Annals of the Missouri Botanical Garden

ANNALS
OF THE
MISSOURI

by Will H. I

Notes on Hedyotis (Rubiaceae) in North America
by Walter H. Lewis

New and noteworthy woody Rubiaceae of Panama

by John D. Dwyer and Sister M. Victoria Hayden ... 34-47

i II. New species of Eriofheca,

by Andre Robyns .

New species of Lhianthus (Gentk
by Andri Robyns and Thoma

Karyotypes in relation to classification and phylogeny in Clayfonia
by Walter H. Lewis and Yutaka Suda

i of Stelis (Orchidaceae) from Panama 68-69. A new species
ria (Cucurbitaceae) 69-72. Notes on the genus Inga II 72-73.
A new annual" Eriogonum from Utah 74-75. A second series of Cochlospermum
(Cochlospermaceae) from Panama 75-77. Hasseltia rigida Woodson ex A.
Robyns, a new species of Flacourfiaceae from Panama 77-78. Mayna zulhna
(Pittier) A. Robyns, comb. nov. (Flacourfiaceae) 78-79. A new Ormosia
(Leguminosae) from Peru 79.

ANNALS

of the

MISSOURI BOTANICAL GARDEN

VOLUME 55

1968

NUMBER I

A journal containing scientific contributions from the Missouri
Botanical Garden and the Department of Botany of Washington Uni-
versity in affiliation with the Missouri Botanical Garden. Outside
contributions in systematic botany and allied fields will also be con-
sidered. These papers are subject to a charge of $25 per printed page.

Walter H. Lewis, Editor
Missouri Botanical Garden & Washington University
Patricia E. Putman, Assistant to the Editor
Missouri Botanical Garden
Derek Burch, Missouri Botanical Garden & Washington University
John D. Dwyer, Missouri Botanical Garden & St. Louis University
Andre Robyns, Missouri Botanical Garden & Washington University

Beginning with Volume 53, 1966:

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ANNALS VOLUME 55

1968

of the NUMBER '

MISSOURI BOTANICAL GARDEN

Department of Botany, The University of Texas e

European
including

The study of s.

herbaria has led to the i

Introduction

Bouvardia, a rubiaceous genus of primarily small shrubs, is a notable but
poorly understood component of the Mexican (with extensions into the south-
western United States) and northern Central American flora. The habitat differs
but is often a rocky outcrop of steep hillsides or barrancas at elevations above 500
meters, the ambient conditions varying from xeric to very mesic.

No economic significance is attributed to the genus. However, the attractive
flowers of a number of species have led to their adoption as ornamentals, and arti-
ficial hybrids between some of them have been obtained. The nomenclature of
these horticultural subjects is considered outside the scope of this study.

The taxonomy of Bouvardia has received little attention since Standley's treat-
ment in the North American Flora (1921b), in which 30 species were recognized,
a number being newly described. Since Standley's revision, descriptions of 19
additional species have appeared, scattered throughout the literature. The im-
mediate needs in the genus were to evaluate the status of the species described since

1 Based on a dissertation submitted to the Graduate School of The University of
Texas at Austin in partial fulfillment of the requirements for the degree of Doctor of
Philosophy. Assisted by a grant-in-aid, The Society of the Sigma Xi and by National
Science Foundation Grant No. GB-4128 (Principal Investigator, W. Frank Blair). Dr.
Marshall C. Johnston is gratefully acknowledged for supervision of this research.

2 Currently Postdoctoral Fellow, Missouri Botanical Garden and Department of Botany,
Washington University, St. Louis, Missouri. Assisted by National Science Foundation
Grant No. GB-5674 (Principal Investigator, Walter H. Lewis).

Ann. Missouri Bot. Card. 55(1): 1-30, 1968.

The previous issue of the Annals of the Missouri Botanical Garden, Vol. 54,
No. 3, pp. 201-421, was published on March 11, 1968.

[Vol. 55
2 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Standley's revision and re-evaluate those recognized by Standley. The recent aquisi-
tion of material by herbaria throughout the world has emphasized the pertinence
of such a study.

The grouping of species within the genus has also been a matter of concern.
Standley did not formally recognize the three subgenera introduced by Schlech-
tendal (1854), although his organization of species in the key corresponds closely
to them. I have found Schlechtendal's subgenera to be at least a useful contrivance
and consider their retention worthwhile for the sake of convenience alone. How-
ever, as nature is not systematically perfect, it is to be understood that some in-
termediacy exists between them, and that their delimitation is perhaps more prag-
matic than phylogenetic.

The position of Bouvardia within the Rubiaceae has undergone drastic, and
probably justified, alteration. Bouvardia, Heterophyllaea, Hindsia, Manettia,
Danais, Coursiana and Hymenopogon were traditionally treated as members of
the Cinchoneae (Hooker, 1873; Schumann, 1891). However, Bremekamp (1952)
considered winged seeds (the only character uniting the tribe Cinchoneae) to be
of little morphological significance and transfered these genera to the subfamily
Rubioideae and tribe Hedyotideae (having transfered the tribe Hedyotideae from
the subfamily Cinchonoideae) based on the presence of raphides, the absence of
large pits in the basal wall of the testa cells, and the peltate placentas. Breme-
kamp reaffirms this transfer in a later paper (1966) in which he outlines his
revisionary subdivision of the Rubiaceae.

Of the above genera, Bouvardia appears most closely related to Heterophyllaea
and Hindsia from which it is distinguished by characters of the fruit and by flowers
with four rather than five stamens and to Manettia from which it differs in its
shrubby (not scandent) habit, terminal inflorescences and loculicidal capsules.
It is doubtful that the position of placental attachment may be employed as a
character delimiting Bouvardia and Mannetia, as attempted by Standley (1921a,
see key). In specimens of both genera examined, the placenta is attached to the
base of the septum. Bouvardia is distinguished from Hedyotis (sensu Lewis, 1961)
by characters of the placenta, seed, endosperm and stipules.

Systematic Treatment 3
Bouvardia Salisbury, Parad. Lond. pi. 88, 1807.
Aeginetia Cav., Anal. Gi. Nat. 3: 129, 1801; Ic. 6: 51, 1801, not Aeginetia L. (Sp. PI. 632,

1753).

Shrubs, suffrutices or perennial herbs; raphides present. Leaves opposite or
3 to 4(-6)-nate, petiolate or infrequently sessile; stipules interpetiolar, consisting
of a basal sheath and a usually trullate, green (not hyaline or scarious) mucro
or free-portion; blades simple, entire, usually membranous. Inflorescences terminal,
cymose (occasionally reduced to solitary flowers). Flowers actinomorphic, often
showy; calyx-lobes 4, most often lanceolate, typically erect or ascending, persisting

BLACKWELL — BOUVABDIA O

in fruit; corolla tubular or salverform, the tube usually well exceeding the lobes,
the lobes 4 (very rarely 5 on the same plant), valvate; stamens 4, epipetalous,
the anthers linear, dehiscing by longitudinal slits, usually included and subsessile
in pin flowers (but typically positioned well above the middle of the corolla-tube),
included or exserted (the filaments becoming free from the corolla-tube at the
throat and extending beyond) in thrum flowers; ovary ± subglobose, inferior to
Y 3 superior, the style single, slender (to ca 1. mm broad), compressed, elongate,
exserted in pin flowers, included in thrum flowers, terminating in 2 rather short,
linear, often minutely papillose style-branches. Capsules subglobose or slightly
oblate, often somewhat compressed loculicidally, 2-celled (the septum complete),
dehiscing loculicidally then septicidally, with a single ascending, fleshy, peltate
placenta in each cell affixed to the based of the septum; seeds go, brown, vertically
imbricate on the placenta, uniformly and ± evenly winged, the wing neither
dissected nor fringed, the nucleus disciform, flattened or slightly pulvinate in cross-
section, the testa cells thin-walled, lacking large pits in the basal wall, the en-
dosperm fleshly.

Type: Boumrdia triphylla Salisbury [=B. ternifolia (Cav.) Schlecht.].

nflorescence (flower or

arising from each leaf-

l leaf-pairs of a floriferous stem tip) often composed

>f 3 or more flowers (sometimes of only 1); corolla tubular to salver-

brm, white or variously colored, the tube 0.4-3.5 (-4.4) cm long, the lobes

[-7(-8.5) mm long, flared 10-90°; anthers in thrum flowers variously

ided or,

If the corolla is white (often greenish- or yellowish-
white rather than pure white) and salverform and the partial inflores-
cences are solitary flowers, then the floriferous branchlets are often very
short and terminate in nodding, 3 to 7-flowered inflorescences (cf. B.

multiflora) I. Bouvardiastrum (p. 3)

bb. Flowers solitary or if the inflorescences several-flowered, then each partial
inflorescence composed of only a solitary flower; corolla salverform, white,
the tube (1.5-)3.5-8.5 cm long, the lobes (4-)7-28 mm long, flared at 90°;
anthers included in thrum flowers or only the tips protruding

aa. Leaves 3 to 4(-6)-nate at most nodes III. Bouvardia (p. 22)

I. Subg. Bouvardiastrum Schlecht., Linnaea 26: 58, 1854.

Shrubs, suffrutices or rarely perennial herbs, the seasonal branches opposite,
elongate or very short, pubescent or glabrous. Leaves opposite, typically petiolate;
blades glabrous to conspicuously pubescent, the secondaries pinnate, less frequently
subpalmate or plinerved, arcuate, rarely obscure, the reticulation obscure to prom-
inent. Inflorescences few to many-flowered; partial inflorescences often composed
of 3 or more flowers (occasionally 1-flowered). Flowers pedicellate or rarely ses-
sile; calyx-lobes free to the floral cup or joined basally by a calyx-tube; corolla
tubular to salverform, red, orange, yellow, violet, lavender, white, yellowish-white
or greenish-white, externally glabrous or puberulent or more elongate-pubescent,
the tube internally with a villous ring toward the base or more often generally

4 ANNALS OF THE MISSOURI

pilose or villous in the lower half, rarely entirely glabrous within; anthers included
in pin flowers (rarely exserted), in thrum flowers included or often exserted; style
glabrate or rarely pubescent, the branches 0.5-2.5(-4) mm long.

Lectotype: Bouvardia triflora H.B.K. [~B. multiftora (Cav.) Schultes &
Schultes f.].

a. Leaf-blades typically pinnately nerved, less frequently subpalmately nerved
(the majority of the secondaries arising ± together near the base of the mid-
vein but at least 2 arising near the middle) .
b. Corolla-tube with an internal villous ring toward the base.

c. Stipular processes lanceolate, eglandular; inflorescences 7 to 12-flowered;
corolla externally minutely puberulent with trichomes to 0.05 mm long
(sometimes sparsely so), the lobes to 1.3 mm broad 6. B. conzattii

=ir processes often filiform and terminally glandular; inflorescences
7 to 60-flowered; corolla externally glabrous, the lobes 2-4 mm broad

bb. Corolla-tube often pubescent in the lower half bu
organized into an isolated villous ring.

d. Calyx-lobes joined basally by a calyx-tube 0.5-2.5 i
e. Flowers short-pedicellate,

f. Seasonal branches glabrate; petioles "2.5-11.5 mm long; leaf-
blades cuneate to rounded at the base; corolla white to green-
ish- or yellowish-white, the lobes spreading to 90°; anthers

exserted by 2-5 mm in thrum flowers 3. B. loeseneriana

ff. Seasonal branches distally sparsely to densely pubescent (the
trichomes variously 0.05-1 mm long); petioles 0.5-2(-7) mm
long; leaf-blades rounded or cordate basally; corolla red-orange
or apricot, the lobes yellow, spreading to 45°; anthers included

in thrum flowers or only the tips protruding 5. B. cordifolia

ee. Flowers sessile, each with its base surrounded by the stipule-sheath

of a bract-pair, the corolla externally glabrous 4. B. capitata

dd. Calyx-lobes free to the floral cup.

g. Leaf-blades slightly crassulate, the secondaries partially or totally

obscure; corolla red-orange, the lobes yellow 7. B. chrysantha

gg. Leaf-blades membranous, the secondaries readily evident their
whole length; corolla variously colored.

i. Inflorescences obviously terminal on usually well-developed,
leafy branches; corolla externally sparsely or densely
pubescent with trichomes 0.3-1.3 mm long,
j. Seasonal branches distally glabrous or sparsely pilose;
lower leaf-blade surface and distal portion of corolla-
tube sparsely pilose; secondary veins subpalmate ....8. B. suhcordata
jj. Distal portion of seasonal branches, lower leaf-blade-
surface and distal portion of corolla-tube densely villous;

secondaries pinnate 9. B.

ii. Inflorescences pseudoaxillary on greatly reduced, slender,
leafless branchlets; corolla externally minutely papillose-

puberulent (trichomes to 0.05 mm long) 12. B. gr

hh. Corolla externally glabrous.

k. Seasonal branchlets distally glabrous or papillose-puberu-
lent with trichomes to 0.2 mm long; leaf-blades with the

ure or inconspicuous; inflorescences with fewer than 15
>wers; corolla at least sparsely pilose in the lower half.
Floriferous branchlets often suppressed (the vegetative
portion as short as 2 mm); leaf-blades sparsely to

moderately papillose-puberulent above and below; floral
cup minutely hirtellous with trichomes 0.05-0.3 mm
long; calyx-lobes typically lanceolate, usually somewhat
hirtellous; corolla white to greenish- or yellowish-white,
less frequently yellow or red, tubular to salverform;

anthers often exserted in thrum flowers 1. B. multiflora

11. Floriferous branchlets often well-developed; leaf-blades
glabrate but often ciliate marginally; floral cup glab-

long; calyx-lobes linear or linear-lanceolate, often glab-
rate; corolla red or occasionally yellow, tubular; anthers
included in thrum flowers or only the tips protruding

- - 2. B. laevis

kk. Seasonal branchlets distally rather sparsely pilose to densely
arachnose-pubescent with trichomes 02-1 mm long; leaf-
blades with the secondary veins often brochidodrome, the

than 15-flowered; corolla entirely glabrous within

11. B. standleyana

aa. Leaf-blades (3-)5-plinerved.

m. Inflorescences with fewer than 25 flowers; pedicels 6-13 mm long; calyx-
lobes deltoid-acuminate, ca 1 mm long; corolla-tube with an internal

villous ring toward the base 13. B. rekoi

mm. Inflorescences more than 25-flowered; pedicels 0.5-6 mm long; calyx-lobes
lanceolate to linear or subulate, 2-9 mm long; corolla-tube often villous in
the lower half but the pubescence not organized into a villous ring.
l of leaf-blades obscure above, inconspii

1-2 mm broad; corolla tubular,

red, the tube 10-17 mm long; style glabrate 14. B. oaxacana

nn. Reticulation of leaf-blades visible above, prominent below; calyx-lobes
linear-subulate, 0.3-1 mm broad; corolla tubular to salverform, violet or
lavender, the tube 4.5-8.5 mm long; lower half of the style often
pubescent 15. B. quinquenervata

1. Bouvardia multiflora (Cav.) Schultes & Schultes f., Mant. Syst. Veg. 3: 118,

1827.
Aeginetia multiflora Cav., Anal. Ci. Nat. 3: 131, t. 28, fig. 2, 1801; Ic. 6: 52, t. 572, fig. 2,

1801.
Bouvardia versicolor Ker, Bot. Reg. 3: pi. 245, 1817. (Based on a greenhouse-grown plant,

the illustration is taken as the type)
B. triflora H.B.K., Nov. Gen. Sp. PI. 3: 386, 1820. (Type Humboldt & Bonpland s.n. P,

not seen; fragment P)
. lesii DC, Prodr. 4: 366, 1830. (Based on Aeginetia multiflora)
Anotis longiflora Bentham, Pi. Hartw. 23, 1839. (Type Hartweg 206 GH, K; photo MICH)
Bouvardia triflora var. hirsuta Martens & Galeotti, Bull. Acad. Bruxelles 11(1): 236, 1844.

(Type Galeotti 2658 BR)
B. hicolor Kunze, Linnaea 20:24, 1847. (Based on garden plants grown from Mexican

seed, the description is taken as the type)
B. mutabilis Walpers, Ann. Bot. Syst. Lipsiae 5: 127, 1849, as synonym of B. versicolor.
B. schiedeana Schlecht., Linnaea 26: 123, 1854. (Type Schiede s.n. B|)
Houstonia triflora (H.B.K.) A. Gray, Proc. Amer. Acad. Arts Sci. 4: 314, 1860.
Bouvardia gracilis A. Gray, loc. cit. 22: 306, 1887. (Type probably Pringle 1255 G, GH)
B. versicolor var. graciliflora A. Gray in S. Watson, loc. cit. 416. (Syntypes Palmer 154,

369 & 708, all GH)
B. heterophylla Standley, N. Amer. Fl. 32: 107, 1921. (Holotype Heyde & Lux 3137 US

398167; isotype GH)
B. macrantha Standley, loc. cit. (Holotype Purpus 3981 US 841341; isotypes BM, F, GH,

MO, UC)

6 ANNALS OF THE MISSOURI BOTANICAL GARDEN

B. latifolia Standley, loc. cit. 111. (Holotype Langlasse 246 US 385804; isotypes F fragment,

GH,P)
B. salvadorensis Steyermark, Ceiba 4:302, 1955. (Holotype Tucker 1368 F 1492726; iso-
types MICH, UC, US)

Shrubs to 2 m; seasonal branchlets very short (2 mm) or to 10 cm or more
in length, papillose-puberulent with trichomes to 0.2 mm long. Leaves with pe-
tioles to 5 mm long; blades most often ovate-lanceolate (varying from broadly
ovate to narrowly lanceolate or elliptic-lanceolate), 0.7-6.7 cm long, 0.2-3.2 cm
broad, membranous, sparsely or moderately hirtellous with papillose trichomes to
0.3 mm long, the secondaries 4-8, usually pinnate, the reticulation often obscure.
Inflorescences terminal but often seemingly axillary by virtue of the very short
branchlets, 3 to 7 (-11) -flowered, often nodding; partial inflorescences 1 to 3-flow-
ered; Flowers with pedicels 0.5-6(-16) mm long which are papillose-puberulent;
floral cup hirtellous with papillose trichomes to 0.3 mm long; calyx-lobes typically
lanceolate (varying from deltoid-acuminate to subulate), 1-8 mm long, usually
somewhat hirtellous; corolla tubular to virtually salverform, white to greenish- or
yellowish-white, less frequently yellow or red, glabrous, the tube 6-36(-44) mm
long, internally sparsely pilosulous toward the base, the lobes 1-7 (-8.5) mm long,
spreading 10-90; anthers 1.5-2.5 mm long, in thrum flowers either exserted by 1-5
mm or included (tips sometimes protruding), included by 1-6 mm in pin flowers;
style glabrous, the branches 1-4 mm long. Capsules 3-7 mm long, 3.5-9 mm broad;
seeds 2-3.5 mm broad.

Holotype: MA, not seen. Cavanilles gives no information about the locality
or collector in the original description.

Chihuahua to Nicaragua; no specimens known from Vera Cruz, Chiapas or
the Yucatan Peninsula; moist slopes or ravines to dry, rocky hills or buffs; with
pine-oak or thorn-scrub-cactus association; 1000-3000 m; flowering May through
September.

Bouvardia versicolor, B. heterophylla, B. latifolia, B. macrantha and B. salva-
dorensis are reduced to synonymy for the first time. I consider B. multiflora to be
a polymorphic species, the description given encompassing the variability of speci-
mens referred to all the above segregates.

The types of B. heterophylla (Guatemala) and of B. salvadorensis and certain
collections from Temascaltepec, Mexico and Zitacuaro, Michoacan (Hinton 7899,
7911, 7925, 7933, 11996, all US) possess elongate pedicels (often 5-16 mm), short
calyx-lobes (2 mm or less) and seasonal branches often more elongate than those
of the average specimen of B. multiflora. Type specimens of B. latifolia and other
Guerreran specimens such as Crisman & Willis 193 (MICH, TAES) and Cooper
& Rowell 2511 (MICH) possess elongate (to 44 mm), subsalverform (lobes spread-
ing at ca 60°), white corollas which closely approach the type of corolla found
in the subg. Bouvardioides. The type of B. macrantha (Tlacuilotepec, Puebla) and
collections from the vicinity of San Luis Tultitlanapa, Puebla (Purpus 3324, 3328,
3330, 3331, 3332, all MO) are characterized by rather elongate (to 36 mm), tubu-
lar corollas in which the anthers are included (or only the tips protruding) in
thrum flowers. All these specimens are assignable to B. multiflora based on their
total morphology and when all specimens of B. multiflora are considered, com-

BLACKWELL — BOUVARDIA I

plete intergradation is seen to exist. Attempting to formalize components of this
reticulum of variation by the application of different Latin names would lead to
confusion.

2. Bouvardia laevis Martens & Galeotti, Bull. Acad. Bruxelles 11(1): 236, 1844.
B. fiava Decaisne, Fl. Serres 1: 215, pi 38, 1845. (Based on a greenhouse-grown plant,

the illustration is the type)
B. mollis Linden ex Schlecht, Linnaea 26: 85, 1854. (Type LZ|)
B. nubigena Standley & Steyermark, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 382, 1940.

(Holotype Steyermark 31901 F 1043221; photos F, TEX)

Shrubs to 1.5 m; seasonal branches usually well-developed, glabrous or dis-
tally sparsely papillose-puberulent with trichomes to 0.2 mm long. Leaves with
petioles 1-7.5 mm long; blades ovate-lanceolate, subattenuate or subacuminate
at the apex, 1.5-10.5 cm long, 0.7-5 cm broad, membranous, glabrate but often
ciliate marginally with trichomes 0.05-0.8 mm long, less frequently sparsely puberu-
lent above and below (often primarily along the veins), the secondaries 4-10,
pinnate, the reticulation apparent or obscure. Inflorescences 3 to 11-flowered; par-
tial inflorescences 1 to 5-flowered. Flowers with pedicels (l-)3-27 mm long which
are glabrous to moderately papillose-puberulent; floral cup glabrous or sparsely
puberulent with trichomes to 0.1 mm long; calyx-lobes linear-lanceolate or linear,
glabrate or sparsely ciliate, 4-12 mm long; corolla tubular, red or occasionally
yellow, glabrous externally, the tube 16-39 mm long, pilosulous in the lower half,
the lobes 2.5-5.5 mm long, flared up to 40°; anthers 2-4 mm long, included or only
the tips protruding; style glabrate, the branches 1-2.5 mm long. Capsules 4-9 mm
long, 5-10 mm broad; seeds 2-3 mm broad.

Type: Mexico. Vera Cruz: ravines around Zacuapan, 3000 ft, April 1840,
Galeotti 2600 (BR, F fragment ex G-Deless., P not seen; photo of P specimen
at F, TEX, US).

Southwestern Tamaulipas to northwestern Oaxaca, Michoacan and southern
Jalisco; centered in Hidalgo and western Vera Cruz; also found
and extreme southern Chiapas; slopes of barrancas or rocky outcrops of ]
sides; oak-pine woodlands in mesic to semi-arid conditions; 1000-3300 m; flowering
April to August.

Bouvardia laevis and B. multifiora are closely related species and intermediates
are sometimes encountered. Keying out a "problematical" specimen often requires
consideration of a number of characters in combination (see key, p. 5), some of
which are not statable in quantitative or absolute terms, and none of which will
alone serve to identify the plant with certainty. Although it would be easy to
build a case for combining these two species, the majority of specimens are easily
identifiable and the extremes quite divergent. At present it seems unwise to com-
bine them, increasing still further the complexity of the variation coming under
the name B. multifiora.

3. Bouvardia loeseneriana Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 186,
1937. (Type Hinton 8107 BM, F, G, GH, K,

[Vol. 55
8 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Shrubs, seasonal branches short or well-developed, glabrate. Leaves with
petioles 2.5-11.5 mm long; blades ovate to ovate- or elliptic-lanceolate, basally
cuneate to rounded, 2.5-9.8 cm long, 1.3-6.7 cm broad, sparsely pilosulous below
and at the margins with white trichomes 0.2-1 mm long, glabrate above, the sec-
ondaries 4-12, pinnate, the reticulation apparent or obscure. Inflorescences (3-) 7
to 20-nowered. Flowers with pedicels 0.5-5 mm lcng; floral cup glabrous to pru-
inose-puberulent (trichomes to 0.05 mm long) or hirtellous (trichomes to 0.5 mm
long); calyx-lobes deltoid to broadly or narrowly lanceolate, l-3(-4) mm long,
marginally ciliate with trichomes to 0.2(-0.4) mm long, joined basally by a calyx-
tube 0.5-2 mm long; corolla typically salverform, white to greenish- or yellowish-
white, externally densely pruinose- or papillose-puberulent with white trichomes
to 0.05 mm long or glabrous, the tube 24-32 mm long, internally sparsely or mod-
erately pilosulous with trichomes 0.4-1 mm long, the lobes oblong-elliptic or ob-
long-lanceolate, 3.5-7 mm long, spreading at ca 90°, often arcuate; anthers 2-2.5
mm lcng, exserted in thrum flowers by 2-5 mm; style glabrate, the branches 0.5-2
mm long.

Holotype: Mexico. Guerrero: Texquitzin nr Chilapa, 28 Oct 1829, Schultes 89
(Bj) ; isotype F (fragment) .

Guerrero and the southern portion of the State of Mexico to Nayarit; pre-
cipitous mountainsides, wooded slopes and barrancas, steep ravines; typically in
moist habitats but occasionally occurring in more arid barancas; 300-2400 m;
flowering July through December.

In typical specimens of B. loeseneriana the corolla-tubes are minutely but
densely pruinose- or papillose-puberulent externally, the floral cups glabrous or
obscurely puberulent with trichomes to 0.05 mm long, the calyx-lobes deltoid
or broadly lanceolate and not exceeding 3 mm, a calyx-tube to 2 mm long is present
and the seasonal branches are well-developed (see types of B. loeseneriana and B.
hintoni and such collections as Hinton 10521, US; 15003, US; 75979, US; McVaugh
& Koelz 1046, MICH). However, in Colima, southwestern Jalisco and Nayarit,
variable specimens are encountered with glabrous corolla-tubes, glabrous or pru-
inose-puberulent (trichomes to 0.05 mm long) or conspicuously hirtellous (tri-
chomes to 0.5 mm long) floral cups, often slender-lanceolate calyx-lobes to 4 mm
long and short or elongate seasonal branchlets (see McVaugh 15260, 18060, 18964,
19832; Wilbur & Wilbur 2335, 2439, 2440, 2464, 2465; all MICH). Specimens
with glabrous corollas, hirtellous floral cups, slender-lanceolate calyx-lobes and
short branchlets are difficult to distinguish from B. multiflora and are doubtless
closely allied to the latter. They differ from B. multiflora in the presence of a
calyx- tube and usually in the more numerous flowers (more than 7) of the in-

4. Bouvardia capitata Bullock, Hook. Ic. PI. t. 3296, 1935.

Shrubs to 1.5 m; seasonal branchlets rather elongate, to 2 mm broad, woody
or suffrutescent, glabrous or sparsely pilose distally with trichomes to 0.7 mm long.
Leaves with petioles 1-14 mm long; blades ovate to suborbicular or broadly ellip-
tic, 2-10.3 cm long, 1.3-6.3 cm broad, usually sparsely pilose with white trichomes

-BOUVAHDIA 9

to 1 mm long (occasionally restricted to the margins), the secondaries 4-10, pinnate
or the lower 4-6 arising subpalmately near the base of the midvein, the reticula-
tion obscure. Inflorescences capitate, 6 to 36-flowered. Flowers sessile, each sub-
tended by the stipule- sheath of a bract-pair; floral cup typically glabrous; calyx-
lobes linear-lanceolate, 3-8 mm long, pilosulous toward the margins with trichomes
0.3-0.8 mm long, joined basally by a calyx-tube 1-2.5 mm long; corolla ± salver-
form, probably white, glabrous externally, the tube 15-23 mm long, pilosulous in
the lower half; the lobes 3-7 mm long, oblong, spreading to 90°, often arcuate;
anthers 1-2 mm long, exserted by 2-4 mm in thrum flowers; style glabrous. Cap-
sules 4.5-8 mm long, 4.5-11 mm broad; seeds 2.5-4 mm broad.

Type: Mexico. Mexico: Distr Temascaltepec, Palmar, in a barranca, 7 July
1934, Hinfon 63/9 (BM, F, K).

Known only from the District of Temascaltepec in the State of Mexico; bar-
rancas and open woodlands at 1000-1500 m; flowering in July and August.

As pointed out by Bullock (loc. cit.) the congested inflorescence of B. capitata
is distinctive, probably representing a reduction from a compound dichasium. The
base of each sessile flower is surrounded by the often membranous and pluriaristate
stipule-sheath of a bract-pair, the lamina being suppressed. The flowers are grouped
in threes and then in multiples of three, each flower-group or partial inflorescence
being in turn subtended by a stipule-sheath showing an increasing development
of the bract lamina.

Bauvardia capitata is closely related to B. loeseneriana, but in addition to the
sessile flowers subtended by stipule-sheaths, specimens of B. capitata may often
be distinguished by the glabrous corollas with the tube usually less than 23 mm
long, linear-lanceolate calyx-lobes typically exceeding 3 mm in length and possess-
ing a more elongate marginal pubescence, and the always glabrous floral cups.

Shrubs, slender or suffrutices to 1 m; seasonal branches ascending, distally
sparsely to densely papillose-puberulent or villosulous with white trichomes 0.05-1
mm long. Leaves with pubescent petioles 0.5-2(-7) mm long; blades broadly ovate
to ovate-lanceolate, cordate or rounded at the base, 1.5-6.6 cm long, 0.7-5.2 cm
broad, sparsely puberulent with papillose to villous trichomes 0.05-1 mm long,
the secondaries 4-12, pinnate, the reticulation often conspicuous below. Inflores-
cences 7 to 50-flowered, subcapitate. Flowers with pedicels 0.5-2.5(-8) mm long
which are sparsely to densely pubescent; floral cup glabrate to densely white-
pubescent with trichomes 0.05-0.8 mm long; calyx-lobes lanceolate to subulate,
1.5-7.5 mm long, joined basally by a calyx-tube 0.5-2 mm long; corolla tubular,
red-orange or apricot (the lobes and extreme distal part of the tube yellow) mi-
nutely white-papillose-puberulent externally with trichomes to 0.05 mm long, less
frequently glabrous, the tube 12-30 mm long, pilose in the lower half but lacking
a villous ring, the lobes ovate or deltoid to oblong or elliptic-lanceolate, flaring
up to 45°, 1.2-5 mm long; anthers ca 2 mm long, brown, included by 1-6 mm

[Vol. 55
10 ANNALS OF THE MISSOURI BOTANICAL GARDEN

or the tips protruding; style papillose-puberulent to glabrate, white, the branches
1-1.5 mm long. Capsules 3.5-6 mm long, 4-7 mm broad; seeds 1.5-3.5 mm broad.
Type: Mexico: Sesse & Mocino s.n. (MA, not seen); pi 487 of the Caiques des
dessins ... is a drawing of the type.

Mexico, Michoacan, Oaxaca, Guerrero, Colima and Jalisco; steep moist slopes,
shady arroyas or dry rocky cliffs or barrancas; often in oak or pine-oak woodlands;
200-2850 m; flowering June through November.

The majority of specimens of B. cordifolia deposited in herbaria are from the
vicinity of Morelia, Michoacan and Temascaltepec, Mexico and are characterized
by the minute but often dense external puberulence of their corollas (see Black-
well 26 & 27, TEX; Hinton 7950, F; Hitchcock & Stanford 7146, US; Kenoyer
A263, F; Rzedowski 20775, ENCB). A few scattered collections of B. cordifolia
have been made in Colima, southern Jalisco, Guerrero and Oaxaca, but these are
all distinguished by glabrous corollas (see litis et al. 570, McVaugh 15010 & 75873,
Ryan & Floyed 37, all MICH). As I have found no other character correlated
with the difference in pubescence of the corolla which might serve to delimit these
two "groups" of 75. cordifolia, and since their apparent geographic separation may
result from insufficient collecting rather than natural disjunction, I have not rec-
ognized them by formal infraspecific categories. It is perhaps worth mentioning
that an analogous situation involving pubescent and glabrous forms exists in B.
loeseneriana and that the geographic area of interest and pattern of distribution
are roughly similar.
6. Bouvardia conzatto Greenman, Proc. Amer. Acad. Arts Sci. 39: 92, 1903.

Shrubs; seasonal branchlets often distally white-papillose-puberulent with
trichomes 0.05-0.1 mm long. Leaves with petioles 1-3 mm long; stipular processes
lanceolate; blades ovate to lanceolate, 1.5-5 cm long, 0.4-2.8 cm broad, basally
cuneate to broadly rounded, minutely papillose-hispidulous marginally and often
sparsely so above and below (primarily along the main veins below) with tri-
chomes 0.05-0.25 mm long, the secondaries 6-10, pinnate or occasionally the lower
4-6 arising subpalmately near the base of the midvein, the reticulation usually
obscure. Inflorescences 7 to 12-flowered. Flowers with pedicels 0.5-5.5 mm long;
floral cup glabrate; calyx-lobes linear or linear- lanceolate, 2.5-5.5 mm long, 0.3-0.8
mm broad, free to the floral cup; corolla tubular, red or yellowish-red, very min-
utely papillose-puberulent externally with trichomes less than 0.05 mm long (some-
times only sparsely so), the tube 8-17 mm long, 1.5-2 mm broad at the neck,
internally with a villous ring toward the base, the lobes ovate to deltoid, 1-2 mm
long, to 1.3 mm broad; anthers 2-3 mm long, included by 1-4.5 mm; style glabrate.
Capsules 2.5-4.5 mm long, 3-5.5 mm broad; seeds 1.3-2 mm broad.

Holotype: Mexico. Oaxaca: 1750 m, July- Aug 1901, Conzatti & Gonzales
1067 (GH). (sphalm. Conzatti & Gonzales 1076 by Greenman)

Known only from Oaxaca; 1600-2000 m; known to be in flower May through

(not

BLACKWELL— BOUVABJDIA 11

B. myrtifolia Schlecht, Linnaea 26: 121, 1854. (Type Schiede s.n. B|)

B. macilenta Blake, Contr. Gray Herb. 53: 65, 1918. (Holotype Conzatti & Reko 3288 GH)

B. cataphyllaris Bullock, Hook. Ic. PL t. 3297, 1935. (Type Hinton 1131 BM, F, G, GH, K,

MICH, MO, US)

Shrubs or suffrutices to 1 m; seasonal branchlets well-developed or very short,
glabrous or distally papillose-puberulent (trichomes to 0.1 mm long) or pilosulous
(trichomes to 1 mm). Leaves with petioles 0.5-3 mm long; blades ovate to elliptic
or lanceolate, 1-7.8 cm long, 0.4-3 cm broad, often somewhat conduplicate, slightly
crassulate, pale-satiny-green below, glabrous or sparsely puberulent with white
trichomes 0.1-1 mm long (sometimes limited to the margins and midvein), rarely
more densely pubescent, the secondaries usually totally or partially obscure, the
reticulation not visible. Inflorescences 5 to 25-flowered, usually subcapitate. Flowers
with pedicels 0.5-4 mm long; floral cup green, glabrous or occasionally pubescent;
calyx-lobes linear-lanceolate to slender-subulate, 2.5-11 mm long, free to the
floral cup; corolla tubular, red-orange (the lobes and extreme distal part of the
tube yellow), externally glabrous or rarely sparsely pilosulous with white trichomes
to 1 mm long, the tube 15-34 mm long, 2-4 mm broad at the neck, sparsely pilosu-
lous in the lower half (trichomes ca 0.5 mm long), the lobes ovate to oblong, 1.5-4
mm long, 1-3 mm broad, flared up to 40°; anthers white or yellow-white, 1.5-3 mm
long, included by 1-6 mm or the tips protruding; style glabrate, white or yellow-
white. Capsules 3.5-8.5 mm long, 4.5-10 mm broad; seeds 2.5-3.5 mm broad.

Type: "Mexico": nr Sanjaguillo, Karwinsky s.n., not seen.

Oaxaca and extreme southern Peubla to Jalisco; dark loamy soil of volcanic
slopes or rocky bluffs of barrancas; oak zone; 1300-2900 m; flowering mostly from
June through October.

Except for the small size of all structures, the type specimen of B. macilenta
falls within the limits of variation under the name B. chrysantha.

Certain specimens of B. chrysantha such as Hinton 10471 (MO, US) & 10541
(MO, US) and the type of B. cataphyllaris resemble B. multiftora in their short florif-
erous branchlets, hirtellous floral cups and often partially evident secondary veins.
It is thus probable that B. chrysantha intergrades to a limited extent with B. multi-
flora. However, B. chrysantha is retained as a separate species since all but a few
specimens are clearly distinct from B. multiflora. As already indicated, combining
closely related species with B. multiflora would only lead to the creation of one
hopelessly polymorphic species.

8. Bouvardia subcordata Standley, N. Amer. Fl. 32: 105, 1921.

Shrubs, the seasonal branches glabrous or pilose distally and at the nodes.
Leaves with petioles 1-5 mm long; blades ovate to suborbicular, basally broadly
rounded or cordate, 1.8-4.3 cm long, 1.2-3.4 cm broad, membranous, moderately
or rather sparsely (but evenly) pilose with white trichomes 0.2-1 mm long, the
secondaries 6-8, the lower 4-6 often arising subpalmately near the base of the mid-
vein, the reticulation obscure. Inflorescences 3 to 7-flowered. Flowers with pedicels
1-5 mm long which are puberulent with white trichomes ca 0.5 mm long; floral
cup tomentose; calyx-lobes lanceolate or elliptic-lanceolate, 2-7.5 mm long, sparsely
pilose, free to the floral cup; corolla tubular, red (?), externally rather sparsely

12 ANNALS OF THE MISSOURI BOTANICAL GARDEN

white-pilose distally with trichomes 0.3-1 mm long, the tube 15-31 mm long, 2.5-
4.5 mm broad at the neck, pilose in the lower half, the lobes ovate to ovate-
orbicular or rhombic, 2-4 mm long; anthers 2-2.5 mm long, included or the tips
protruding; style glabrate.

Holotype: Mexico. Sinaloa: betw Rosario & Colomas, 13 July 1897, Rose 1628
(US 300476); isotype F.

Known only from the type collection.

9. Bouvardia xylosteoides Hooker & Arnott, Bot. Beechey Voy. 428, 1840.

B. villosa Standley, N. Amer. FL 32: 107, 1921. (Holotype Conzatti 1486 US 764094;

isotype GH)

Shrubs, the seasonal branchlets densely white-villous at least distally with
trichomes to 1. mm long. Leaves with villous petioles 1-6 mm long; blades ovate
to elliptic, basally cuneate to rounded, 1.3-3,3 cm long, 0.8-2 cm broad, villous with
trichomes to 1 mm long (densely villous-tomentose below), the secondaries 4-8,
pinnate, the reticulation obscure. Inflorescences 3 to 7-flowered. Flowers with vil-
losulous pedicels 0.5-2 mm long; floral cup tomentose; calyx-lobes lanceolate or
linear-lanceolate, 2-8 mm long, rather densely pubescent; corolla tubular, densely
white-villous on the upper two-thirds with trichomes 0.4-1.3 mm long (sparsely
pubescent toward the base), the tube 13-26 mm long, sparsely pilose within near
the base, the lobes ovate, 1.5-3,5 mm long; anthers 2-4 mm long; style glabrous.

Type: Mexico. Oaxaca: low mountains around Mitla (sphalm. Mitlam by
Hooker & Arnott), Andrieux 333 (F fragment ex G-Deless., G, K).

Bouvardia xylosteoides has been collected at 1800 m and is known to be in
flower from May through July; all collections known (three, including the type col-
lection) are from central Oaxaca.

10. Bouvardia dictoneura Standley, N. Amer. Fl. 32: 109, 1921.

B. matudai Lundell, Lloydia 2: 105, 1939. (Holotype Matuda 2667 MICH; isotypes F,

MO, US; photos F, TEX)
B. venosissima Lundell, Bull. Torrey Bot. Club 66:602, 1939. (Holotype Matuda 2748

MICH)
B. pachecoana Standley & Steyermark, Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 22, 1943.

(Holotype Standley 86226 F 1093896; isotypes A, US; photo US)

Shrubs, slender, to 2.5 m; seasonal branches often elongate, slender, suffrutes-
cent, bisulcate, sometimes sparsely pilose along the margins of the sulcae with
trichomes 0.1-0.8 mm long, otherwise glabrous. Leaves with petioles 0.5-10 mm
long; stipular processes 1.5-10.5 mm long, often filiform, glandular at the tip;
blades ovate to lanceolate, basally rounded or cordate, 2-10.6 cm long, 1-5.8 cm
broad, usually glabrous but often ciliate marginally with trichomes 0.1-1 mm long,
occasionally sparsely pilose above and below with trichomes to 1 mm, the secon-
daries 4-12, pinnate or rarely subpalmate, the reticulation obscure to prominent.
Inflorescences 7 to 60-flowered. Flowers with pedicels (l-)3-10 mm long; floral
cup glabrous; calyx-lobes linear-lanceolate or linear-subulate, 1.5-6 mm long;
corolla red or orange-red, externally glabrous, the tube 4-13.5(-20) mm long, 2-4.5
mm broad at the neck, a villous ring within toward the base, the lobes broadly
ovate or rhombic to somewhat oblong, 1.5-6 mm long, 2-4 mm broad, flared up to

BLACKWELL— BOUVARDIA 13

60°; anthers 2-3 mm long, included by 1-7 mm or the tips protruding; style glabrate.
Capsules 3-5 mm long, 3.5-6 mm broad; seeds 1-2 mm broad.

Holotype: Mexico. Chiapas: Chicharras (sphalm. Chichorras by Standley),
3000-6000 ft, 6 Febr 1896, Nelson 3757 (US 256554) ; isotype GH.

Southwestern Guatemala and southern Chiapas; volcanic slopes, often in pine
forests; 1000-3800 m; apparently flowering the year round.

A collection from Guerrero (Pilas-Pasion, Distr Montes de Oca), Hinton 10759
(UC, W), has 3-nate leaves (a characteristic of the subg. Bouvardia) but is other-
wise indistinguishable from many specimens of B. dictyoneura. Field work is
needed in the attempt to find additional specimens with ternate leaves and to
assess the reality of the apparent geographic separation from typical opposite-
leaved forms.

11. Bouvardia standleyana Blackwell, sp. nov.

Frutex B. gracilipedi affinis, sed foliorum laminis supra saltern sparse pubes-
centibus, inflorescentiis terminalibus in ramis foliosis elongatis dispositis, corollaque
omnino glabro differt.

Shrubs; seasonal branches 7-30 cm long, suffrutescent, distally sparsely pilose
(often irregularly so) to densely arachnose-pubescent with white trichomes 0.2-1
mm long. Leaves with petioles 1-7.5 mm long; blades ovate to trullate, 1.5-7.5 cm
long, 0.8-4.3 cm broad, membranous, at least sparsely pubescent on both surfaces
with white trichomes 0.2-1 mm long (below primarily concentrated along the
main veins, the trichomes often spreading or curling), occasionally densely arach-
nose-puberulent below, the secondaries 6-12, pinnate, often brochidodrome, the
reticulation apparent above, prominent below. Inflorescences (10-) 15 to 50-flow-
ered, terminal on leafy branches more than 7 cm long (including flowers) and
more than 1 mm broad. Flowers with pedicels 3-14 mm long (at least a few pedi-
cels 8 mm or more) and less than 0.5 mm broad; floral cup sparsely pilosulous to
conspicuously hirsute with trichomes 0.2-0.7 mm long or glabrous; calyx-lobes
lanceolate or elliptic-lanceolate to subulate, 1-3 mm long; corolla tubular, yellow or
light orange, externally and internally glabrous, the tube 7-24 mm long, the lobes
ovate or deltoid to oblong, 2-4 mm long, flared up to 45°; anthers 1-2 mm long,
exserted by 1-3.5 mm in thrum flowers, included by 1-2 mm in pin flowers; ovary
subglobose, ca 1 mm long, the style glabrous, with branches 1-2.5 mm long.

Holotype: Mexico. Michoacan: Distr Coalcoman, S of Naranjillo, 1360 m, 26
Nov 1938, Hinton 12677 (US 2020693); isotypes K, MICH, UC.

Western Jalisco, Michoacan and Guerrero; mountain summits, steep mountain-
sides or rocky slopes; often found in the oak zone but occasionally in sandy soil in
association with more tropical vegetation; 800-1700 m; flowering September through
February.

Bouvardia standleyana is distinguished from its nearest relative, B. gracilipes,
by the glabrous corollas (externally and internally), the well-developed and leafy
floriferous branches, the presence of pubescence on the upper blade surface (though
often sparse) and the more elongate pubescence on the distal portion of the seasonal
branches.

14 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Two collections, Hinton 14901 (US) & 15408 (F), from Mina, Guerrero are
interesting in their apparent intermediacy between B. standleyana and B. bouvar-
dioides (subg. Bouvardia). Although the majority of floral and inflorescence char-
acters, as well as the pilose to arachnose pubescence of the branchlets and leaves
resemble B. standleyana, the leaf size, form, verticillation and the red corollas
strongly suggest specimens of B. bouvardioides from the Western Sierra of Mexico.
However, B. bouvardioides is not known from Guerrero and definitely solving the
problem of the ancestry of such putative hybrids will require intensive field work
followed by experimental studies.

12. Bouvardia gracilipes B. L. Robinson, Proc. Amer. Acad. Arts Sci. 45: 404,

1910.

Shrubs to 1.5 m; seasonal branches glabrous or distally white-papillose-puberu-
lent with trichomes to 0.05 mm long. Leaves with petioles 0.5-3.5 mm long; blades
ovate to ovate-lanceolate, 2-9 cm long, 1-4.4 cm broad, membranous, glabrous above,
often puberulent or pilosulous below on the main veins (and marginally) with
trichomes 0.05-0.5 mm long, otherwise glabrous, the secondaries 6-10, pinnate, often
brochidodrome, the reticulation most conspicuous below, imparting a minutely
varicose appearance to the blade above. Inflorescences 5 to 50-flowered, pseudo-
axillary, the floriferous branchlets less than 7 cm long (including flowers), less
than 1 mm broad, leafless (only a stipule-sheath present at the nodes), with only
1 or 2 very short internodes below the peduncle. Flowers with pedicels 5-22 mm
long (at least a few pedicels 8 mm or more) and less than 0.5 mm broad; floral cup
minutely papillose-puberulent with trichomes to 0.05 mm long or glabrate; calyx-
lobes deltoid to lanceolate, 0.5-2 mm long; corolla tubular, yellow or orange-yellow,
externally minutely papillose-puberulent with trichomes to 0.05 mm long, the tube
10-19 mm long, often somewhat pilosulous within just below the middle (but not
with a dense, villous ring), the lobes broadly ovate to oblong, 2-4.5 mm long;
anthers 1.5-3 mm long, exserted in thrum flowers by 1-2 mm; style glabrate, the
branches 0.5-1 mm long. Capsules 3.5-5.5 mm long, 4-7 mm broad; seeds 1.5-2.5
mm broad.

Holotype: Mexico. Nayarit: Tepic, 5 Jan to 6 Feb 1892, Palmer 1971 (GH);
isotypes F, US.

Nayarit and northwestern Jalisco; steep, heavily forested stream valleys in the
oak zone; growing in cracks and among rocks in stream beds; known to be in
flower in November and December.

13. Bouvardia rekoi Standley, N. Amer. Fl. 32: 108, 1921.

Shrubs; seasonal branches glabrous or distally minutely puberulent with tri-
chomes to 0.05 mm long. Leaves with petioles 1-3 mm long; blades ovate-lanceo-
late, rounded at the base, 3-6.5 cm long, 0.9-2.7 cm broad, glabrous, 3 to 5-plinerved,
the reticulation often prominent below. Inflorescences few-flowered. Flowers with
pedicels 6-13 mm long and 0.2-0.3 mm broad; floral cup sparsely pruinose-puberu-
lent or hispidulous with trichomes to 0.05 mm long; calyx-lobes deltoid-acuminate,
ca 1 mm long; corolla tubular, externally glabrate, the tube 10-19 mm long, a

BLACKWELL — BOUVABJDIA 15

villous ring within near the base, the lobes oblong or oblong-lanceolate, 2.5-4 mm
long; anthers 1.5-3 mm long, exserted in thrum flowers by 1-4 mm; style glabrate,
the branches 1-1.5 mm long.

Holotype: Mexico. Oaxaca: Cafetal Montecristo (Cerro Espino), 1000 m, 7
Dec 1917, Reko3650 (US 867149).

Known only from the type collection.

14. Bouvardia oaxacana Standley, Jour. Wash. Acad. Sci. 13: 7, 1923.

Herbs, perennial or suffrutices; stems simple or sparsely branched, slender,
distally puberulent with white trichomes 0.05-0.3 mm long. Leaves with petioles
0-2.5 mm long; blades broadly ovate to ovate-lanceolate, basally rounded, 3.4-6.7
cm long, 1.3-3.8 cm broad, glabrate or sparsely white-puberulent with trichomes
0.1-0.5 mm long (often primarily along the margin), 5-plinerved, the reticulation
obscure above, visible but inconspicuous below. Inflorescences often more than 30-
flowered, subcapitate. Flowers with pedicels 0.5-4 mm long and 0.5 broad; floral
cup usually densely hirtellous; calyx-lobes lanceolate to elliptic- or linear-lanceo-
late, 2-9 mm long, 1-2 mm broad; corolla tubular, red, externally glabrous or
sparsely hirtellous, the tube 10-17 mm long, villous toward the base within but
lacking a villous ring, the lobes mostly oblong-lanceolate, 3-6 mm long; anthers
2-3 mm long, exserted by 2-6 mm in thrum flowers; style glabrate, the branches
1-2 mm long.

Holotype: Mexico. Oaxaca: Distr Juquila, betw Santa Cruz & El Aguacate,
500 m, 24 Dec 1921, Conzatti 4513 (US 1110842); isotype GH.

Known only from the type collection.

15. Bouvardia quinquenervata Standley, N. Amer. Fl. 32: 108, 1921.

Shrubs; seasonal branches rather elongate, distally hirtellous with trichomes
0.05-0.3 mm long. Leaves with petioles 1-5 mm long; blades broadly ovate to
ovate- or elliptic-lanceolate, basally obtuse to rounded or cordate, 2.6-5.9 cm long,
1.3-3.7 cm broad, glabrate above, obscurely puberulent below on the main veins
with trichomes to 0.1 mm long, 5-plinerved, the reticulation visible above, prom-
inent below. Inflorescences usually more than 50-flowered. Flowers with pedicels
1.5-6 mm long, to 0.5 mm broad; floral cup rather densely hirtellous; calyx-lobes
linear-subulate, 2-6 mm long, 0.3-1 mm broad; corolla tubular to virtually salver-
form, violet or lavender, externally sparsely hirtellous (mainly toward the base)
with trichomes 0.1-0.2 mm long, the tube 4.5-8.5 mm long, rather densely villous in
the lower half (the pubescence not organized into a villous ring), the lobes oblong
to oblong-lanceolate, 2.5-4 mm long; anthers 2-3.5 mm long, exserted in both pin
and thrum flowers by 1-5.5 mm; style somewhat hirtellous or villosulous on the
lower half. Capsules 3-4.5 mm long, 2.4-4 mm broad.

Holotype: Mexico. Chiapas: San Bartolome, 22 March 1904, Goldman 769
(US 470574).

Southern Chiapas; pine forests on volcanic slopes; ca 1400 m; known to flower
December through March.

16 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Based on the many-flowered inflorescences, the short lavender corollas and
stamens exserted in both pin and thrum flowers, B. quinquenervata appears more
closely related to specimens of B. bouvardioides from Chiapas and Guatemala than
to any member of the subg. Bouvardiastrum.

II. Subg. Bouvardioides Schlecht, Linnaea 26: 58, 1854.

Shrubs or suffrutices; seasonal branches opposite, glabrous or pubescent.
Leaves opposite, usually petiolate; blades glabrous or pubescent, the secondaries
pinnate, obscure or rarely subpalmate, the reticulation obscure or inconspicuous.
Inflorescences 1- or several (to 15) -flowered (when several-flowered, each partial
inflorescence composed of only 1 flower). Flowers pedicellate or sessile, ± erect;
calyx-lobes free to the floral cup or rarely connected basally by a calyx-tube; corolla
salverform (lobes spreading at an angle of 90°), white, externally glabrous or
pubescent, the tube internally sparsely pilose throughout or the pubescence limited
to the upper half (never limited to the lower half), less often glabrate internally;
anthers in both pin and thrum flowers included or only the tips protruding; style
glabrate or more often minutely papillose-pubescent (at least on the distal half),
the branches 1-7.5 mm long.

Type: Bouvardia longiflora (Cav.) H.B.K.

a. Flowers sessile or subsessile on a branch terminus (rarely with pedicels to 1 mm
long), usually solitary (if inflorescences several-flowered then the leaf-blades
conspicuously villous beneath toward the base with trichomes to I mm long
and the floral cup villous-tomentose or else the flowers with a calyx-tube ca
2 mm long extending beyond the floral cup).

b. Flowers solitary (if inflorescences 3 or more-flowered then the floral cup
villous-tomentose with trichomes to 1.5 mm long and the leaf-blades con-
spicuously villous below toward the base); calyx-lobes free to the floral

c. Stipular processes lanceolate or slender-subulate, 2.5-10 mm long; blades
glabrate or infrequently puberulent with trichomes to 0.3 mm long (not
concentrated below toward the base); floral cup glabrate or rarely
densely puberulent with trichomes to 0.3 mm long; corolla-tube glabrous

externally or rarely sparsely puberulent with trichomes 1
flowers always solitar)

...16. B. longiflora

cc. Stipular processes often pluriaristate, 1-3.5 mm long; blades conspicu-
ously villous below toward the base with trichomes to 1 mm; floral cup
densely villous with trichomes 0.5-1.5 mm long; corolla-tube externally

; i: l- villous distally with trichomes 0.3-1.5 mm long (if glabrous

then the inflorescences with 3-11 flowers) 17. B. langfassei

bb. Inflorescences 3 to 5 (-7) -flowered (floral cup pruinose-puberulent with
trichomes to 0.05 mm long and the leaf-blades glabrate); calyx-lobes con-
nected basally by a calyx-tube ca 2 mm long 19. B. rosei

Flowers typically at least short-pedicellate; inflorescences often 3- or more-
flowered (if flowers solitary, the leaf-blades less than 9 mm broad and the
plants often low, dense shrubs of xerophytic habit).

d. Petioles (0.5-)3-12 mm long; leaf-blades usually broader than 9 mm, the
secondaries evident; inflorescences 3 to 15-flowered.

e. Plants glabrate; pedicels (0.5-)2.5-19 mm long; calyx-lobes (0.5-) 1.5-5
mm broad, often rather broadly linear-elliptic; corolla-tube 1.5-4.5 (-5.6)
cm long -. 20. B. glabra

BLACKWELL— BOUVARDIA

dd. Petioles 0-3 (-4.5) mm long; leaf-blades usually 1
the secondaries usually obscure; inflorescences 1 to !

f. Xerophytic, low, dense shrubs, the branchlets often becoming pseudo-
spinescent; leaf-blades to 12(-18) mm long, often papillose-puberulent
with trichomes to 0.2 mm long (sometimes very sparsely so); flowers
solitary (inflorescences rarely 3-flowered); calyx-lobes 1.5-5 (-6.5) mm

long - 18. B.

ff. Typically not xerophytes, the branches not at all spinescent; leaf-blades
0.9-7 cm long, glabrate or rarely papillose-puberulent with trichomes
to 0.3 mm long; inflorescences often 3 to 9-flowered, the flowers occasion-
ally solitary; calyx-lobes 2-32 mm long.

g. Leaf-blades having a length/width ratio of 2.4/1 to 18/1; floral
cup typically pruinose- or papillose-puberulent with trichomes to
to 0.05 (-0.3) mm long (sometimes sparsely so); calyx-L '

mm long; ovary ± subglobose, 1-2 mm long 2

gg. Leaf-blades having a length/width ratio of 10/1 to 42/1; floral cup
glabrous; calyx-lobes 12-32 mm long; ovary turbinate or sub-
cylindrical, 2-3.5 mm long .23. B. karwinskyi

: 51, t. 572, fig. 1,
1801.
Bouvardia longiflora var. latifolia Martens & Galeotti, Bull. Acad. Bruxelles 11(1): 236, 1844.

(Holotype Galeotti 2659 BR)
Houstonia longiflora (Cav.) A. Gray, Proc. Amer. Acad. Arts Sci. 4: 314, 1860.

Shrubs to 1.6 m; seasonal branches ascending, glabrous or distally white-
papillose-hirtellous with trichomes to 0.2 mm long, often short and slender and
herbaceous. Leaves with petioles to 10 mm long; stipular processes lanceolate or
slender-subulate, 2.5-10 mm long; blades lanceolate or occasionally ovate- or elliptic-
lanceolate, 1.5-5.7 cm long, 0.5-2.2 cm broad, glabrous or infrequently white-
puberulent with trichomes 0.05-0.3 mm long, the secondaries 4-12, pinnate. Flowers
solitary, sessile or subsessile (pedicels to 1 mm long) ; floral cup glabrate or rarely
densely puberulent with white trichomes 0.1-0.3 mm long; calyx-lobes linear to
linear-lanceolate or linear-oblanceolate, rarely more broadly oblanceolate or ±
elliptic, (2.5-) 6-26 mm long; corolla glabrous externally or rarely sparsely puberu-
lent with trichomes to 0.3 mm long, the tube 3.5-8.5 cm long, internally sparsely
pilose throughout, the lobes 8-28 mm long; anthers 3-5 mm long; ovary 2-5 mm
long, the style glabrate or pruinose-puberulent, the branches 2-6 mm long. Capsules
8-12.5 mm long, 7-12 mm broad; seeds 1.5-4 mm broad.

Holotype: Mexico: betw Queretaro & Guanajuato, Oct (MA, not seen). The
description is based on a specimen in the herbarium of L. Nee without reference
to the collector.

Aguascalientes, Jalisco and San Luis Potosi to Puebla and Oaxaca; occasional
on rhyolitic or basaltic hillsides or steep roadcuts at elevations of 2000-4000 m;
sometimes associated with xerophytic plants in arid or semiarid habitats but more
frequently occurring in mesic woodlands of oak and pine; flowering most commonly
from May through August.

Two collections of B. longiflora from Puebla, Purpus 1249 (GH, MO) from
Tehuacan and Purpus 3327 (MO, UC) from the vicinity of San Luis Tultitlanapa,
are distinguished by their puberulent leaves, densely puberulent floral cups
and shorter calyx-lobes (2.5-6 mm long). At first I thought these formed a distinct

18 ANNALS OF THE MISSOURI BOTANICAL GARDEN

subspecies. However, other collections of "typical" B. longifhra, Purpus 3323 (F)
& 3326 (GH, MO), from the vicinity of San Luis Tultitlanapa were subsequently
seen and cast doubt on the geographical separation of the puberulent forms. Addi-
tional collecting is needed to elucidate their status.

17. Bouvardia langlassei Standley, N. Amer. Fl. 32: 110, 1921.

Shrubs to 1.5 m; seasonal branches distally villous with trichomes 0.2-1 mm
long, often short and herbaceous. Leaves with villous petioles to 7 mm long;
stipular processes 1-3.5 mm long, often pluriaristate; blades obovate or oblanceolate
to ovate or ovate-lanceolate, 1.2-5.2 cm long, 0.5-2,8 cm broad, villous (particularly
beneath toward the base) with white trichomes 0.3-1 mm long, the secondaries
4-10, pinnate or infrequently ± plinerved. Flowers solitary or 3-11 on a seasonal
branch, sessile or subsessile (pedicels to 1 mm); floral cup densely white-villous
with trichomes 0.5-1.5 mm long; calyx-lobes linear or slender-lanceolate, 2.5-9 mm
long; corolla distally sparsely villous externally with slender trichomes 0.3-1.5 mm
long or glabrous (when glabrous, the inflorescence typically several-flowered), the
tube 3.5-6.6 cm long, internally glabrate or sparsely pilose throughout, the lobes
6-14 mm long; anthers 3-4 mm long; ovary 1-3 mm long, the style glabrate, the
branches 1.5-3.5 mm long.

Holotype: Mexico. Guerrero: Testla (Yextla), 45 km W of Chilpancingo,
1500 m, 8 June 1899, Langlasse 1049 (US); isotypes F (fragment), G, GH, P; photo
US.

18. Bouvardia erecta (DC.) Standley, N. Amer. Fl. 32: 110, 1921.

Catesbaea erecta DC, Prodr. 4: 401, 1830.

Hedyotis spinescens Sesse & Mocino, Fl. Mex. 22, 1891. (Type Sesse & Mocino s.n. MA,

Bouvardia flos-joannis Schumann in Loesener, Bull. Herb. Boiss. 3: 621, 1895. (Type Seler

846 GH)
B. flos-joannis var. latifolia Loesener, Repert. Sp. Nov. 18: 358, 1922. (Type Endlicher

1966 W, not seen)

Shrubs to 1.2 m; seasonal branches glabrous or papillose-hispidulous with
white trichomes to 0.2 mm long, ± herbaceous, becoming woody and pseudospines-
cent. Leaves often pseudofasiculate by virtue of crowding and the development of
axillary buds which produce leaves but fail to elongate; petioles to 2.5 mm long;
stipular processes slender-deltoid, 0.5-2 mm long; blades lanceolate or linear to
elliptic-lanceolate or oblanceolate, 3.5-12(-18) mm long, 1-4.5 mm broad, char-
taceous, glabrous or white-papillose-puberulent with trichomes to 0.2 mm long,
the secondaries obscure. Flowers solitary (rarely 3 terminating a branchlet);
pedicels (0-) 1-8 mm long; floral cup glabrous or puberulent with trichomes to 0.2
mm long; calyx-lobes linear to lanceolate or slightly oblanceolate, 1.5-5(-6.5) mm
long; corolla white but often lightly suffused with rose, inconspicuously puberulent
externally with trichomes to 0.05 mm long or glabrate, the tube 2.5-5.5 cm long,
sparsely pilose in the upper half (sometimes densely so at the throat), the lobes

BLACKWELL — BOUVARDIA 19

5-12 mm long; anthers 2-4 mm long; ovary 1-2.5 mm long, the style often minutely
papillose-puberulent, the branches 1.5-5 mm long. Capsules 4-9 mm long, 4-8.5
mm broad; seeds 2-3 mm broad.

Type: Mexico: Sesse & Mociho s.n. (MA, not seen); pi. 460 of the Caiques des
dessin ... is a drawing of the type.

Puebla; limestone slopes with gray or whitish soil and frequent rock outcrops;
often in association with thorn-scrub, cactus and hat palms under arid conditions;
1300-2500 m; flowering from June through early October.

Bouvardia erecta is unique in the subg. Bouvardioides in its adaptation to xeric
conditions. The habit is that of a dense, low shrub of scrubby appearance. The
branchlets are often pseudospinescent and the leaves small and rather densely
crowded.

. rosei Standley, N. Amer. Fl. 32: 109, 1921.

Shrubs, the branches minutely puberulent when young. Leaves with petioles
ca 1 mm long; stipular processes narrowly lanceolate, 3-5 mm long; blades ovate
to elliptic, 2-3.8 cm long, 0.7-2 cm broad, glabrate, the secondaries 4-8, subpalmate
(two arising from the midvein slightly below the midpoint of the leaf and the
remainder originating together near the base or occasionally all originating near
the base). Inflorescences 3 to 5 (-7) -flowered. Flowers typically sessile (occasionally
with pedicels to ca 1 mm); floral cup pruinose-puberulent; calyx-lobes linear-
lanceolate, 4-9 mm long, connected basally by a calyx-tube extending ca 2 mm
above the floral cup; corolla glabrate externally, the tube 4-7 cm long, internally
sparsely pilose throughout, the lobes 11-15 mm long; anthers 3-4 mm long; ovary
ca 1.5 mm long; style moderately papillose-hispid distally, the branches 4-6 mm
long. Capsules 6-8 mm long and broad; seeds 2-3 mm broad.

Holotype: Mexico. Durango: Sierra Madre, 16 Aug 1897, Rose 3516 (US);
isotype F; photo US.

Sierra Madre of Durango and Sinaloa; infrequent; known to be in flower in
August (probably also in September).

20. Bouvardia glabra Polakowsky, Linnaea 41: 565, 1877.

B. glabra var. gracilis Polakowsky, loc. cit. 566. (Type Polakowsky 172 Bf, BM, W; photos

F, G, TEX)
B. glabra var. obtusa Loesener, Verh. Bot. Ver. Brandenburg 65: 106, 1923. (Type Seler

2920 B|)

Shrubs to 1.5 m; branches glabrous. Leaves with petioles (2-) 3- 12 mm long;
stipular processes lanceolate or linear-lanceolate, 1.5-6 mm long; blades ovate- or
elliptic-lanceolate, 2.5-10.5 cm long, 0.9-4.3 cm broad, glabrous or glabrate, the
secondaries 6-12, pinnate. Inflorescences 3 to 7-flowered. Flowers with pedicels
(0.5-)2.5-19 mm long; calyx-lobes elliptic to oblanceolate or ± linear, 3.5-17 mm
long, (0.5-) 1.5-5 mm broad; corolla externally glabrous or rarely minutely puberu-
lent with trichomes to 0.05 mm long, the tube 1.5-4.5(-5.6) cm long, glabrate
within, the lobes 5-16 mm long; anthers 2.5-4 mm long; ovary 1.5-3 mm long;
style pruinose-puberulent, the branches 1.5-7.5 mm long. Capsules 4-10 mm long,
5-10 mm broad; seeds 2-4.5 mm broad.

[Vol. 55
20 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Type: Costa Rica: San Jose, Aug 1875 Polakowsky 337 (B|, F fragment;
photos F, G, TEX, US); possibly from cultivated material.

Guatemala, southern Chiapas and Costa Rica; wooded or shrubby slopes or
barrancas of volcanic mountains in arid to very mesic (cloud forest) conditions;
1300-3500 m; flowering the year round.

I have seen collections from Honduras perhaps assignable to B. glabra but
intermediate between B. glabra and B. dolichantha. Examples of these possibly
hybrid specimens are Hawkes et al. 2095 (C, G, K), Molina 6501 (F) and
Rodriguez 3124 (F). Additional collecting is needed to ascertain more accurately
the distribution of B. glabra, B. dolichantha and B. induta and to look for inter-
mediate forms. These three taxa are a closely related complex in northern Central
America and I view the specific status of B. dolichantha and B. induta as tentative
at present.

21. Bouvardia induta (Robinson) Standley, N. Amer. Fl. 32: 109, 1921.

B. longiflora var. induta B. L. Robinson, Proc. Amer. Acad. Arts Sci. 45: 404, 1910.

B. purpusii Brandegee, Univ. Calif. Publ. Bot. 10: 415, 1924. (Holotype Purpus 9251

UC 223423 p; isotypes F, GH, MO, US)

Shrubs to 1 m; branches distal ly rather densely white-hirsute with trichomes
to 0.8 mm long or glabrous. Leases with petioles (0.5-)3-9 mm long; stipular
processes subulate, 1-3.5 mm long; blades lanceolate to ovate- or elliptic-lanceolate,
(1.5-)2-7.3 cm long, (0.5-) 1-2.5 cm broad, conspicuously white-hirsute with
trichomes to 1 mm long or glabrous, the secondaries 4-10, pinnate. Inflorescences
3 to 15-flowered. Flowers with pedicels 0.5-2.5(-6) mm long which are densely
hirsute to glabrate; floral cup white-hirsute-tomentellous with trichomes to 1 mm
long or minutely puberulent (trichomes to 0.05 mm long) or glabrate; calyx-lobes
linear to linear-lanceolate or linear-elliptic or slightly oblanceolate, 2.5-16 mm
long, 0.5-1 (-2) mm broad; corolla externally villosulous or hirsute with trichomes
0.1-0.7 mm long or pruinose- or papillose-puberulent with trichomes to 0.05 mm
long, rarely glabrate externally, the tube (3.5-) 4.5-8.5 cm long, pilose in the distal
half with trichomes to 1 mm long, the lobes 7-22 mm long; anthers 3-5 mm long;
ovary 1-2.5 mm long, the style distally minutely puberulent and proximally glab-
rous, with branches 1-5 mm long.

Holotype: Mexico. Chiapas: June-July 1864-70, Ghiesbreght s.n. (GH). The
holotype is one of three collections of B. induta on one herbarium sheet, viz. the
one in the middle. The specimens on the left and right are Ghiesbreght 108 and
692 respectively.

Chiapas and adjacent Guatemala; rocky, dry limestone areas in pine forests;
1000-2200 m; flowering March through September.

Some specimens of B. purpusii differ from B. induta in the glabrescence of all
parts (see Purpus 9252 MO, UC, US). However, the holotype is a pubescent plant
similar to the majority of specimens of B. induta. An isotype (F) is a mixed col-
lection, the specimen on the left side of the herbarium sheet being glabrate and
the one on the right conspicuously pubescent. The specimens on the right is
probably a duplicate of the holotype.

BLACKWELL — BOUVARDIA 21

22. Bouvardia dolichantha Loesener, Verh. Bot. Ver. Brandenburg 65: 106, 1923.
B. steyermarkii Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 383, 1940. (Holotype

Steyermark 29671 F 1043386; photos F, TEX)

Shrubs or low suffrutices; branches glabrous or sparsely papillose- or pruinose-
puberulent distally with trichomes to 0.05(-0.3) mm long. Leaves opposite but
occasionally crowded and pseudoverticillate; petioles 0.5-3(-4.5) mm long; stipular
processes deltoid to lanceolate, 0.5-3 mm long; blades linear or lanceolate to
elliptic- or narrowly ovate- lanceolate, 0.9-7 cm long, 1.5-9 mm broad, glabrate or
rarely papillose-puberulent with trichomes to 0.3 mm long, the secondaries obscure
or 4-6 and pinnate. Inflorescences (l-)3 to 9-flowered. Flowers with pedicels 0.5-
5.5 mm long which are glabrous or minutely puberulent; floral cup typically
pruinose- or papillose-puberulent with trichomes to 0.05(-0.3) mm long; calyx-
lobes linear or lanceolate to narrowly elliptic or oblanceolate, 2-11 mm long, 0.5-
1.3(-2) mm broad; corolla externally glabrous or sparsely papillose-puberulent with
trichomes to 0.05(-0.2) mm long, the tube 1.7-8 cm long, internally glabrate or
papillose-puberulent to pilose (trichomes 0.1-1 mm long) in the upper half, the
lobes 4-19 mm long; anthers 2-4 mm long; ovary usually subglobose, 1-2 mm long,
the style minutely puberulent on the distal two-thirds, with branches 1-5 mm long.
Capsules 3.5-8.5 mm long and broad; seeds 2-3.5 mm broad.

Loesener cites Seler 2795 and 2883 without selecting one or the other as the
type. I designate Seler 2883 (US 1205579) as lectotype (Guatemala: Dept of
Huehuetenango, Distr of Nenton, nr Uaxackanal & toward Quen Santo, 1100 m,
9 July 1896); isolectotypes GH, F (fragment); photos F, G, US from B|.

Guatemala and Honduras; occasional on mountainsides along humid banks
in mixed woodlands of oak and pine; 1000-3400 m; flowering May through October.

23. Bouvardia karwinskyi Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 8: 155,
1930.

Shrubs to 1.5 m; seasonal branches glabrate or obscurely hirtellous with tri-
chomes to 0.1 mm long. Leaves opposite but often crowded, sometimes pseudo-
fasciculate by the production of leaves by axillary buds which fail to develop into
branches; petioles 0-2 mm long; stipular processes lanceolate, 1.5-5.5 mm long;
blades linear to linear- lanceolate or narrowly linear-elliptic, 1.6-5.3 cm long, 0.7-3
mm broad, glabrous, the secondaries obscure. Inflorescences 1 to 5-flowered.
Flowers with pedicels 0.5-6 mm long which are glabrous; floral cup glabrous,
calyx lobes linear, 12-32 mm long, 0.5-1.5 mm broad; corolla glabrous externally,
the tube 3.5-6.2 cm long, sparsely pilose in the upper half, the lobes 7-13.5 mm
long; anthers 3-3.5 mm long; ovary turbinate or subcylindrical, 2-3.5 mm long,
the style papillose-puberulent (particularly distally) with trichomes to 0.05 mm
long, with branches 1-2 mm long.

Holotype: Mexico: betw Victoria & Rio Blanco, 1842, Karwinsky 312 (LE);
isotype F (fragment) .

I have been unable to learn the state(s) in which the type and other specimens
of B. karwinskyi were collected; flowering July through October.

22 ANNALS OF THE MISSOURI BOTANICAL GARDEN

III. Subg. Bouvardia Schlecht, Linnaea 26: 58, 1854.

Shrubs, suffrutices or perennial herbs; seasonal branches opposite or verticil-
late, pubescent to glabrous. Leaves 3 to 4(-6)-nate at some or all of the nodes,
petiolate or sessile; blades conspicuously pubescent (mostly below) to glabrous,
the secondaries pinnate, arcuate, less frequently obscure, the reticulation obscure to
prominent. Inflorescences few to many-flowered; partial inflorescences 3- or more-
flowered. Flowers pedicellate or virtually sessile; calyx-lobes free to the floral cup;
corolla tubular to salverform (lobes spreading 10-90°), red, less frequently lavender,
rose, violet-blue or pink, externally pubescent or glabrous, the tube internally
pilose or villous in the lower half (rarely glabrate) or the pubescence organized
into a villous ring near the base; anthers included (rarely exserted) in pin flowers;
included or often exserted in thrum flowers; style usually glabrate, the branches
0.5-3 mm long.

a. Corolla externally pubescent, rarely obscurely pruinose-puberulent or glabrate

but then the leaves obovate and 5 to 6-nate at some of the nodes.

b. Leaves not more than 4-nate, the blades typically neither obovate nor

mucronulate; corolla-tube puberulent externally with trichomes usually 0.1-

0.4 mm long, internally with a villous ring toward the base.

c. Well-branched shrubs or low suffrutices (occasionally the stems simple

and herbaceous but broader than 1.5 mm at the midpoint); leaves

at least short-petiolate, the blades having a length/width ratio of 1.5/1

to 19/1 and most often not linear, the secondaries usually apparent

24. B. ternifolia

cc. Herbaceous or suffrutescent perennials, the stems 1-1.5 mm broad at

the midpoint, simple or sparsely branched distally; leaves often sessile

ally short-petiolate), the blades linear, with a

length/width ratio of 12/1 to 50/1, the secondaries obscure 25. B. tenuifolia

bb. Leaves 5- or 6-nate at one or more nodes, the blades often obovate and
mucronulate; corolla-tube puberulent externally with trichomes to 0.05 mm
long or rarely glabrate, lacking an internal villous ring toward the base

(though occasionally somewhat villosulous) 26. B. ohovata

aa. Corolla externally glabrous.

d. Seasonal branches distally villosulous or hirsute with trichomes 0.2-2 mm
long; leaves short-petiolate or subsessile (petioles 0.5-3 mm long), the
blades often conspicuously pubescent (particularly below) with trichomes
0.2-1.1 mm long (the pubescence either coarse and yellowish or villous
and white); corolla red, the tube 10-27 mm long; inflorescences with fewer
than 60 flowers and often subcapitate.

e. Slender shrubs or suffrutices, the branches angular or terete in cross-
section and less than 3 mm broad at the midpoint; leaf-blades various
(often ovate-lanceolate), white-villous-pubescent below, the secondaries
4-14 (often fewer than 12) ; calyx-lobes usually less than 5.5 mm long;
anthers exserted 1-3 mm in thrum flowers, included in pin flowers,
f. Corolla tubular, the tube typically with a villous ring internally
toward the base, the lobes usually less than 2.2 mm broad, spread-
ing up to 45° 27. B. leiantha

ff. Corolla salverform, the tube villous internally toward the base but
lacking a definite villous ring, the lobes frequently 2.2 mm broad

or more, spreading at 90° 28. B. viminalis

ee. Coarse perennial herbs, often with distinctly tetragonal stems 3-4 mm
broad at the midpoint; leaf-blades usually rhombic-ovate or ovate,
yellowish, with a coarse yellow pubescence beneath concentrated pri-
marily on the main veins, the secondaries (8-) 12-22; calyx-lobes subu-
late (or approaching filiform), (3-) 5.5- 12 mm long; anthers included
in pin and thrum flowers 29. B. scabra

BLACKWELL — BOUVARDIA 23

dd. Seasonal branches distally glabrous or puberulent with trichomes to 0.1
mm long; leaves entirely sessile or conspicuously petiolate (at least some of
the petioles exceeding 4 mm), the blades glabrous or sparse] v
with trichomes to 0.3 mm long; corolla deep-rose, lavender, violet-blue, pink
or red (when red the inflorescences more than 60-flowered and not sub-
capitate), the tube 2-19 mm long,
g. Leaves sessile, the blades linear, having a length/width ratio of 4/1 to

20/1, the secondaries obscure; inflorescences 3 to 20-flowered 30. B. rosea

gg. Leaves distinctly petiolate, the

having a length/width ratio of 2.2/1 t

ous; inflorescences usually more than 60-flowered 31. B. bouvardioides

24. Bouvardia ternifolia (Cav.) Schlecht, Linnaea 26: 98, 1854.

Ixora ternifolia Cav. s Ic. 4: 3, t. 305, 1797.

7. americana Jacquin, Hort. Schonbr. 3: 4, 1798. (Based on I. ternifolia)

Houstonia coccinea Andrews, Bot. Repos., pi. 106, 1800. (Illustration is taken as the type)

triphylla Salisbury, Parad. Lond., pi. 88, 1807. (Based on 1. ternifolia)
B. linearis H.B.K, Nov. Gen. Sp. PL 3: 383, 1819. (Type Humboldt & Bonpland s.n. P,

not seen; photos F, P, TEX, US).
B. angustifolia H.B.K., loc. cit. 384. (Type Humboldt & Bonpland s.n. F, P not seen, US;

photos F, P, TEX, US)

H.B.K., loc. cit. (Type Humboldt & Bonpland s.n. P, not sean; photos F, P,

TEX, US)

ii U.B.K., loc. cit. 385, 1820. (Based on 7. americana)
B. coccinea (Andrews) Link, Enum. Hort. Berol. 1: 139, 1821.
B. quaternifolia DC, Prodr. 4: 365, 1830. (Type Alaman s.n. G-DC, not seen; photo

IDC)
Carphalea pubiflora Sesse & Mociho ex DC., loc. cit. (As synonym of B. quaternifolia)
Bouvardia jacquinii var. B exogyna DC, loc. cit. (Based on a plant grown in Salm-Dyck's

garden; type G-DC, not seen; photo IDC)
B. jacquinii var. y ovata DC, loc. cit. (Based on "Bouvardia triphylla var. B Salisb.," the

supposedly broader-leafed form, the type of which is Salisbury's pi. 88)
B. splendens Graham, Bot. Mag., pi. 3781, 1840. (Based on greenhouse-grown plants; K)
B. triphylla var. splendens (Graham) Lindley, Bot. Reg. 26: t. 37, 1840.
B. tolucana Hooker & Arnott, Bot. Beechey Voy. 427, 1840. (Type Andrieux 332 K, W)
B. quaterniflora Steudel, Nomen. Bot. ed. 2, part 1, 300, 1841, as synonym of Carphalea

pubiflora; probably a mistake for B. quaternifolia DC
B. scabrida Martens 8r Galeotti, Bull. Acad. Bruxelles 11(1): 237, 1844. (Holotype Galeotti

2624 BR)
B. hypoleuca Bentham, PI. Hartw. 288, 1848. (Type Hartweg 1605 K; photo MICH)
B. glaberrima Engelmann in Wislizenus, Tour N. Mex. 106, 1848. (Type Wislizenus 161

GH, MO)
B. ovata A. Gray, PI. Wright. 5: 67, 1853. (Type Wright 1117 GH, MO, US)
Aeginetia hyssopifolia Willd. ex Schlecht., Linnaea 26: 60, 1854. (As synonym of

Bouvardia angustifolia)
Bouvardia tenuiflora Schlecht., loc. cit. 97. (Type Bj)
B. houtteana Schlecht. ex Planchon, Fl. Serres 10: 149, t. 55, 1855. (Illustration is taken

as the type)
B. fcrteVa var. quaternifolia (DC.) Rothrock in Wheeler, Rept. U.S. Geogr. Surv. 6: 137,

1879.
B. triphylla var. angustifolia (H.B.K.) A. Gray, Syn. Fl. N. Amer. 1(2): 24, 1884.
B. ternifolia var. angustifolia (H.B.K.) B. L. Robinson, Proc. Amer. Acad. Arts Sci. 45:

405, 1910.
B. endlichii Loesener, Repert. Sp. Nov. 18: 357, 1922. (Type Endlicher 176a B|; photos

F, G, TEX, US).
B. orizabensis Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 8: 334, 1931. (Holotype

Botteri 604 P; isotypes F fragment, G)

Shrubs or suffrutices or perennial herbs to 1.5 m, the branches sparsely to
rather densely papillose-hispidulous when young with white trichomes 0.1-0.3 mm

[Vol. 55
24 ANNALS OF THE MISSOURI BOTANICAL GARDEN

long. Leaves with petioles 0.5-12 mm long; blades extremely variable (linear,
lanceolate, elliptic, ovate, obovate) though most often elliptic-lanceolate, 1-11 cm
long, 0.1-3.1 cm broad, glabrate to densely papillose-hispidulous or villosulous
with white trichomes variously 0.05-1 mm long, the secondaries 4-12 (often obscure
in narrow, linear leaves), the reticulation apparent or obscure. Cymes 3 to 40-
nowered, sometimes subcapitate. Flowers with pedicels 1-15 mm long; floral cup
sparsely to densely papillose-hispidulous; calyx-lobes lanceolate to linear, 2-10 mm
long; corolla tubular, salmon-red or orange-red or scarlet, externally pubescent with
papillose, white, red-tipped trichomes 0.1-0.4 mm long (rarely obscurely puberulent
with trichomes 0.05 mm long or less or villosulous with trichomes to 0.8 mm long) ,
the tube 9-35 mm long, typically with a villous ring internally toward the base,
the lobes ovate to oblong, 1.5-3.5(-5) mm long, spreading up to 60°, internally
white-pruinose-puberulent; anthers 2-4 mm long, white or red, included by 1-4 mm
or the tips protruding; ovary 1-2.5 mm long, the style-branches red, 1-2.5 mm long.
Capsules 4.5-9 mm long, 5-10.5 mm broad, glabrous or sparsely papillose-hispid;
seeds 2-3.5 mm broad.

The original description of Ixora ternifolia was based on a plant growing in the
Royal Botanical Garden of Madrid, the seed having been sent from Mexico. The
illustration is taken as the type.

Southwestern Texas and New Mexico and southeastern Arizona to Oaxaca and
southern Vera Cruz; the most common species of Bouvardia; frequent in desert and
mesic, montane habitats; 800-3000 m; flowering mostly from late February through
October.

In examining type material of B. scabrida, Galeotti 2624, specimens at P and
W (and photos A, F, MICH, TEX) were found to be B. viminalis. However, the
specimen at BR is B. ternifolia and Galeotti 2624 is thus a mixed collection. As
the original publication clearly describes the BR specimen, it is regarded as the
type of B. scabrida. The "isotypes" and "phototypes" discussed are merely addi-
tional specimens of B. viminalis.

Standley (Publ. Field Mus. Nat. Hist., Bot. Ser. 8: 334, 1931) considered B.
orizabensis to be a "well marked" species "related clearly to B. bouvardimdes."
Examination of type material, however, showed B. orizabensis to be synonymous
with B. ternifolia. Confusion may have arisen because of the obscurity of the
pubescence, particularly of the corolla, on the F and P type specimens of B.
orizabensis. It is worthy of mention, however, that the external pubescence of
the G specimen is similar to that of typical specimens of B. ternifolia and may be
from another plant. Even if Botteri 604 represents a mixed collection, all the speci-
mens involved are still referable to B. ternifolia.

25. Bouvardia tenuifolia Standley, N. Amer. Fl. 32: 104, 1921.

Herbs, often suffrutescent toward the base; stems simple or sparsely branched
distally, 1-1.5 mm broad at the midpoint, glabrous or distally minutely puberulent
to villosulous (white trichomes 0.05-0.7 mm long). Leaves with petioles 0-3 mm
long; blades linear, 2-9.5 cm long, 0.7-4.5 mm broad, glabrous or pubescent with
trichomes similar to those of the distal portion of the stem, the secondaries and

BLACKWELL — BOUVARDIA ZO

reticulation obscure. Cymes 3 to 25-flowered, often subcapitate. Flowers with
pedicels 0.5-6 mm long; calyx-lobes narrowly deltoid to subulate or linear, 1-3 mm
long; corolla tubular, red, externally pubescent with papillose, white, red-tipped
trichomes 0.1-0.4 mm long, the tube 7-26 mm long, a villous ring within near
the base, the lobes ovate or elliptic, 1.5-5 mm long; anthers 2-4 mm long, included
by 1-6 mm or the tips protruding; ovary 1-1.5 mm long, the style-branches 0.5-2
mm long.

Holotype: Mexico. Jalisco: grassy hillsides nr Guadalajara, Oct 1899, Pringle
2292 (US); isotypes A, BM, BR, F, G, LE, M, MICH, MO, MSC, P, UC, W.

Sinaloa and western Durango to northern Jalisco; found in various habitats:
precipitous slopes with pine and oak, open slopes in palm-oak country, grassy hill-
sides, gravelly banks of ravines, dry hills; 150-2700 m; flowering July through De-

Intergradation occurs between B. tenuifolia and B. ternifolia where their ranges
overlap, viz. the Western Sierra Madre from extreme western Durango and eastern
Sinaloa to Nayarit and northwestern Jalisco. The shaky separation of the two spe-
cies by a combination of leaf and habit characters may prove untenable in the
light of future collecting.

26. Bouvardia obovata H.B.K., Nov. Gen. Sp. PI. 3: 385, 1820.
Hedyotis fruticosa Sesse & Mocifio ex DC, Prodr. 4: 365, 1830. (Type Sesse & Mocino s.n.

BM, F)

Herbs or suffrutices to 1.3 m; stems glabrous or sparsely papillose-puberulent
(most noticeably at the nodes and on the extreme distal portion) with white tri-
chomes to 0.1 mm long. Leaves 3 to 6-nate (at least one or two nodes 5- or
6-nate); petioles 0-7 mm long; blades obovate to elliptic, 3.5-13 cm long, 0.9-3.8
cm broad, often mucronulate, glabrate, the secondaries 6-12, the reticulation ap-
parent below, often subobscure above. Cymes 10 to 60-flowered. Flowers with ped-
icels 0.5-12 mm long; calyx-lobes lanceolate, 2-4 mm long; corolla tubular, red,
externally minutely papillose-puberulent with white, red-tipped trichomes to
0.05 mm long, the tube 10-32 mm long, internally glabrate or somewhat villosu-
lous toward the base (not with a villous ring), the lobes ovate to elliptic or oblong,
2-5 mm long, flared up to 30°; anthers 2.5-3.5 mm long, included by 1-8 mm or the
tips protruding; ovary 1-2 mm long, the style-branches 1-3 mm long. Capsules
5.5-9 mm long, 6-10 mm broad, glabrous; seeds 2-3.5 mm broad.

Type: Mexico. Distrito Federal: betw Chapultepec & Tezcoco, "1200 hex.?",
June 1803-04, Bonpland s.n. (P).

Occuring sporadically in Nayarit, Mexico State, the Federal District, Morelos
and Oaxaca; wooded slopes and barrancas of volcanic mountains; rolling grassy
hills in oak forests with a few pines; 1000-2150 m; flowering June through Sep-

Orsted 11039 C)

Shrubs to 1.5 m, the branches hirtellous or villosulous with white trichomes
0.2-1 mm long when young. Leaves with petioles to 3 mm long; blades ovate to

26 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ovate- or elliptic-lanceolate, 1.5-7.5 cm long, 0.7-3.5 cm broad, sparsely to densely
villosulous (particularly below) with white trichomes 0.3-1 mm long, the sec-
ondaries 6-14, the reticulation often prominulous below. Cymes 6 to 45-flowered,
often subcapitate. Flowers with pedicels 1-6 mm long; calyx-lobes lanceolate or
elliptic-lanceolate to linear or subulate, 2-5.5 mm long; corolla tubular, deep red,
glabrous externally, the tube 10-19 mm long, often with an internal villous ring
toward the base, the lobes ovate to elliptic or elliptic-lanceolate, 1.5-3.5 mm long,
0.8-2.2 mm broad, spreading up to 45°; anthers 1.5-2 mm long, exserted 1-3 mm in
thrum flowers; ovary 1-1.5 mm long, the style-branches 1-2 mm long. Capsules
2.5-5.5 mm long, 3.5-6 mm broad, glabrous; seeds 1.5-2 mm broad.

Type: Guatemala: in fields nr Tejar & Chimaltenango, July-Aug 1841, Hartweg
583 (GH, K, P, W, photo US from B|).

Chiapas to Nicaragua; rocky, open or bushy, moist or dry hillsides in oak-pine
forests; sometimes growing in dense tropical forests at lower elevations; 400-4000
m; apparently flowering the year round.

28. Bouvardia viminalis Schlecht, Limiaea 26: 120, 1854.

Shrubs or suffrutices to 1 m; branches villosulous when young with slender
white trichomes 0.2-1 mm long. Leaves spreading, often arcuate and ± condupli-
cate; petioles to 3 mm long; blades ovate-lanceolate, 1.5-5.7 cm long, 0.3-2.6 cm
broad, villosulous (particularly below) with white trichomes 0.2-1 mm long, the
secondaries 4-13, the reticulation prominulous to obscure. Cymes 7 to 35-flowered,
often subcapitate. Flowers with pedicels 0-4 mm long; calyx-lobes lanceolate or
elliptic-lanceolate or linear, 1.5-5.5 mm long, somewhat villosulous or hirtellous;
corolla salverform, externally pinkish-red and glabrous, the tube 10-20 mm long,
internally villosulous toward the base but lacking a villous ring, the lobes typically
4 but occasionally 5 on the same plant, ovate to elliptic or oblong, 2-6.5 mm long, 1-5
mm broad, internally scarlet, flared at ca 90°; anthers 1.5-2.5 mm long, yellowish-
white, exserted 1-2 mm in thrum flowers; ovary 1-2.5 mm long, the style-branches
1-2.5 mm long, pink or white. Capsules 3-6.5 mm long and broad, sparsely hirtel-
lous; seeds 1.5-2 mm broad.

Type: Schiede s.n. (Bf). I designate Pringle 4888 (MO) as neotype (Oaxaca:
Monte Alban, 6000 ft, 4 Sept 1894); isoneotypes BM, BR, F, GH, M, MSC, P, UC,
US, W.

Oaxaca and southern Peubla; rocky, open hillsides in sandy soil; sometimes in
association with Agave, Karwinskia, Croton and cactus; flowering June through

29. Bouvardia scabra Hooker & Arnott, Bot. Beechey Voy. 427, 1840.

Herbs to 1 m, often woody toward the base; stems often simple and tetragonal,
3-4 mm broad at the midpoint, hirsute with yellow trichomes 0.3-2 mm long for
virtually the entire length. Leaves with petioles to 3 mm long; blades ovate or
rhombic-ovate to elliptic-lanceolate, 2.5-9.8 cm long, 0.7-4.7 cm broad, chartaceous
or subcoriaceous, yellowish-hirsute with trichomes 0.2-1.1 mm long (often concen-
trated below along the main veins), the secondaries (8-) 12-22, raised below, the

BLACKWELL — BOUVARDIA 27

reticulation often prominent below. Cymes 15 to 60-flowered, often subcapitate.
Flowers with pedicels 0.5-7 mm long; calyx-lobes subulate, suffused with red, his-
pidulous, (3-) 5.5- 12 mm long, often spreading; corolla salverform, red, glabrous ex-
ternally, the tube 10-27 mm long, rather densely villous internally toward the base
but rarely with a villous ring, the lobes ovate, 2.5-8 mm long, spreading to 90°;
anthers 1-2.5 mm long, included by 2-8 mm; ovary 1-2.5 mm long, the style-
branches 1-2 mm long.

Type: Mexico. Nayarit: betw San Bias & Tepic, Sinclair s.n. (K).

Nayarit and Jalisco; rocky mountainsides, wooded ravines, slopes of barrancas;
in oak zone or sometimes in tropical deciduous forests; 900-1700 m; flowering August
through January.

30. Bouvardia rosea Schlecht, Linnaea 26: 116, 1854.

B. violacea Rzedowski, Ciencia (Mex.) 19: 82, 1959. (Holotype Rzedowski 7680 MEXU,

not seen; isotype ENCB)

Herbs to 0.5 m; stems often simple, green, distally compressed, 1-1.5 mm
broad at the midpoint, glabrate or pruinose- or papillose-puberulent with white
trichomes to 0.1 mm long. Leaves ascending, sessile; blades linear, 2-10 mm long
at the lower nodes, to 40 mm long at the upper nodes (except immediately sub-
tending the inflorescence), 0.5-3 mm broad, chartaceous, glabrous or hirtellous with
white trichomes to 0.3 mm long, the secondaries and reticulation obscure. Cymes
3 to 20-nowered. Flowers with pedicels 0.5-5 mm long; floral cup glabrous; calyx-
lobes lanceolate or elliptic-lanceolate, 1-4 mm long; corolla salverform, rose or
violet-blue or clear pink, glabrous externally, the tube 5-19 mm long, internally
often villous toward the base but lacking a villous ring, the lobes ovate or elliptic-
lanceolate or oblong, 2.5-6 mm long, spreading at 90°; anthers 1-2 mm long, ex-
serted 1-2 mm in thrum flowers; ovary turbinate, 1-1.5 mm long, the style-
branches 0.5-2 mm long.

Type: Mexico. Hidalgo (?): nr San Jose del Oro, Schiede s.n. (B|). I desig-
nate McVaugh 14819 (MICH) as neotype (Guanajuanto: 22 mi W of Xichu, rd
from Xichu to San Luis de la Paz, 2300 m, 14 June 1957).

San Luis Potosi, Guanajuato, Queretaro and Hidalgo; rhyolitic and andesitic
hillsides in pine-oak forests; sometimes in dry habitats; 1800-2800 m; flowering
April through June.

31. Bouvardia bouvardioides (Seemann) Standley, N. Amer. Fl. 32: 102, 1921.
Hedyotis bouvardioides Seemann, Bot. Voy. Herald 296, 1856.

Houstonia bouvardioides (Seemann) Bentham & Hooker, Gen. PI. 2: 60, 1873.
Bouvardia pallida Standley, Jour. Wash. Acad. Sci. 14: 245, 1924. (Holotype Standley

22977 US; isotype GH)

Shrubs to 5 m, often slender and clambering, the branches pruinose-puberulent
or hirtellous when young with white trichomes to 0.1 mm long. Leaves with
petioles (2)4-15 mm long; blades ovate- or elliptic-lanceolate, 2.5-11.5 cm long,
0.7-4.5 cm broad, thinly membranous, glabrate or sparsely appressed-hirtellous with
white trichomes to 0.3 mm long (often confined to the main veins below), the
secondaries 8-14, the reticulation prominulous below. Cymes typically more than

[Vol. 55
28 ANNALS OF THE MISSOURI BOTANICAL GARDEN

60-flowered. Flowers with pedicels 1-7 mm long; calyx-lobes linear to lanceolate,
1.5-6 mm long; corolla tubular to somewhat salverform, red or lavender, glabrous
externally, the tube 2-18 mm long, internally somewhat villous toward the base
(but lacking a definite villous ring), the lobes rather narrowly oblong or elliptic-
lanceolate, 2.5-6 mm long, 1-3 mm broad; anthers 1.5-2.5 mm long, exserted by 1-8
mm in pin and thrum flowers; ovary 1-1.5 mm long, the style branches 1-2 mm
long. Capsules 2-3 mm long, 2.5-3.5 mm broad, hirtellous with white trichomes to
0.2 mm long; seeds 0.4-0.8 mm broad.

Type: Mexico. Durango (?): Sierra Madre, Seemann s.n. (GH).

A disjunct pattern of distribution is evident: one center of dispersal being in
the Western Sierra Madre of Mexico at lower elevations from Durango and Sin-
aloa to northwestern Jalisco, the other from southern Chiapas to El Salvador. This
species occupies diverse habitats at elevations of 300-2000 m and flowers from Oc-
tober to April.

Standley (Jour. Wash. Acad. Sci. 14: 245, 1924) treated the Central American
component of this species as a separate species, B. pallida, reserving the name
bouvarioides for plants of the Western Sierra Madre of Mexico. Although the two
groups are apparently widely disjunct geographically, the characters reputedly
distinguishing them (corolla size and color) completely break down on careful

1. Bouvardia alexcmderae A. Carter, Madrono 13: 142, fig. 1 & 2, 1955 (Holotype Carter

2577 UC 985926, not seen; isotype F). Carter (loc. cit. 144) recognized that
B. alexcmderae has wingless, angular seeds but still felt it to be best placed in
Bouvardia in the section "having large, white, salverform corollas with long
tubes" (subg. Bouvarioides ?). However, based on total morphology, choromosome
number (n = 13) and geographical location, I believe B. alexanderae to have
closest affinity with Baja California species of Hedyotis subg. Edrisia such as H.
saxatalis Lewis and H. brevipes (Rose) Lewis. = Hedyotis alexanderae (A. Carter)
W. H. Lewis.

2. B. chlorantha Bertoloni ex Schultes & Schultes f., Mant. Syst. Veg. 3: 116, 1827, ap-
" l in Bertoloni's herbarium (not seen); the description

3. B. coccinea (Aublet) A. Richard, Mem Soc. Hist. Nat. Paris 5: 272, 1834, not B.

coccinea (Andrews) Link (Enum. Hort. Berol. 1: 139, 1821). = Nacibea coccinea
Aublet = Manettia coccinea (Aublet) Willd., fide Standley, N. Amer. Fl. 32: 97,
1921.

4. B. deamii Donn. Sm., Bot. Gaz 49: 445, 1910 (Holotype Deam 6190 US; isotype

MO). = Rondeletia deamii (Donn. Sm.) Standley, N. Amer. Fl. 32: 60, 1918.

5. "Bouvardia?" discolor Hooker & Arnott, Bot. Beechey Voy. 428, 1840 (Type Andrieux

334 K). = Rondeletia leucophylla H.B.K., fide Standley, N. Amer. Fl. 32: 54, 1918.

6. B. fermginea A. Richard, Mem. Soc. Hist. Nat. Paris 5: 272, 1834, nomen nudum.

7. B. havanensis (H.B.K.) A. Richard, loc cit. = Manettia coccinea (Aublet) Willd.,

fide Standley, N. Amer. Fl. 32: 97, 1921.

8. B. hirsuta (Swartz) A. Richard, loc. cit. = Rondeletia hirsuta Swartz, fide Standley,

N. Amer. H. 32: 75, 1918.

9. B. microphylla Schlecht., Linnaea 26: 112, 1854 (Type Schiede s.n. Bf), the descrip-

tion is not sufficient for determination.
10. B. quinqueflora Dehnhardt, Rivista Napolitana 1, 3: 167. Standley questionably
placed B. quinqueflora in the synonymy of B. chrysantha. However, I have been
unable to trace Dehnhardt's original publication or the specimen (s) on which it

BLACKWELL— BOUVARDIA 29

was based. A description of B. quinqueflora by Walpers (Repert. 2: 507, 1843)
is not sufficiently detailed to permit certainty of disposition.

11. B. racemosa (Ruiz & Pavon) A. Richard, Mem. Soc. Hist. Nat. Paris 5: 272, 1834. = ?

Mannetia racemosa Ruiz & Pavon.

12. B. scandens A. Richard, loc. cit, nomen nudum.
strigillosa Baxter, Loudon's Hort. Brit. Suppl. 3: 502, 1850, the description is too

scanty for determination.
strigosa Bentham, PI. Hartw. 75, 1841 (Type Hartweg 503 K). = Rondeletia
strigosa (Bentham) Hemsley, fide Standley, N. Amer. Fl. 32: 50, 1918.

15. B. uniflora (H.B.K.) A. Richard, Mem. Soc. Hist. Nat. Paris 5: 272, 1834. = Manettia

coccinea (Aublet) Willd, fide Standlt , \ ,u ■. I I '-{2:97, 1921.

16. B. viperalis Schlecht., Linnaea 26: 114, 1854 (Type Schiede s.n. Bf), the description

does not suffice for determination.

17. Cestrum spermacocifolium Willd. ex Roemer & Schultes in L., Syst. Veg. 4: 808, 1819

(Type Humboldt & Bonpland s.n., not ssen), provisionally excluded from the
synonymy of Bouvardia multiflora. Standley incorrectly says that Willdenow's
name was published in synonymy.

18. Hedyotis mexicana Standley, N. Amer. Fl. 32: 104, 1921, as synonym of Bouvardia

ternifolia; possibly an error for Hedyotis fruticosa Sesse & Mocino ex DC. ( =
Bouvardia obovata H.B.K.).

19. Houstonia ochroleuca Raf., Ann. Gen. Sci. Phys. 5: 226, 1820, based on H. coccinea

var. alba Dum. which I have been unable to trace. Merrill (Index Rafinesquianus
226, 1949) states that Houstonia "ochroleuca" Raf. = Bouvardia triphylla Salisb.
= B. coccinea (Andrews) Link. The basis of this synonymy is not known.

Literature Cited

Bremekamp, C. E. B. 1952. The African species of Oldenlandia L. sensu Hiern et K.

Schumann. Verh. Kon. Nederl. Akad. Wet., Afd. Natuurk., sect. 2, 48: 1-297.

. 1966. Remarks on the position, the delimitation and the subdivision of the

Rubiaceae. Acta Bot. Neerl. 15: 1-33.
Hooker, J. D. 1873. Cinchoneae, p. 32-44, In Bentham & Hooker, Genera Plantarum.

Vol. 2.
Lewis, W. H. 1961. Merger of the North American Houstonia and Oldenlandia under

Hedyotis. Rhodora 63: 216-223.
Schlechtendal, D. F. L. von. 1854. Die gattung Bouvardia . . . Linnaea 26: 43-126.
Schumann, K. 1891. Cinchoneae, p. 41-55, In Engler & Prantl, Nat. Pflanzenfam. 4(4).
~ '■oneae.N. Amer. Fl. 32:94.
. N. Amer. Fl. 32:100-111.

INDEX OF LATIN NAMES
New taxa are in boldface type, all other taxa are in roman type; numbers in boldface

type refer to descriptions, numbers in roman type refer to synonyms, numbers with dagger
(t) refer to names incidentally mentioned.

Aeginetia 2; hyssopifolia 23; longiflora lichantha 20t, 21; endlichii 23; erecta

17; multiflora 5 18, 19t; ferruginea 28t; flava 7; flos-

Agave26t joannis 18, var. latifolia 18; glaber-

Anotis longiflora 5 rima 23; glabra 19, var. gracilis 19,

Bouvardia If, 2, 24|, 28f; subg. Bou- var. obtusa 19; gracilipes 13|, 14;

vardia 13t, 14t, 22; subg. Bouvardia- gracilis 5; havanensis 28t; hetero-

strum 3, I6t; subg. Bouvardioides 6f, phylla 5, 6t; hintoni 7; hirsuta 28t;

16, 19t, 28t; alexanderae 28t; angus- hirtella 23, var. quaternifolia 23;

3; bicolor 5; bouvardioides houtteana 23; hypoleuca 23; induta

14t, 16t, 24t, 27, 28t; capitata 8, 20; jacquinii 23, var. fi exogyna 23,

9t; cataphyllaris 11; cavanillesii 5; var. y ovata 23; k

chlorantha 28t; chrysantha 10, lit; 7; langlassei 18; latifolia 6;

coccinea 23, 28 1, 29t; conzatti 10; leiantha 25; linearis 23; loe<

cordifolia 9, lOf; corymbosa 25; 7, 8t, 9t, lOt; longiflora 16t, 17,

i - ira 12, 13t; do- 18t, var. induta 20, var. latifolia 17;

ANNALS OF THE MISSOURI BOTANICAL GARDEN

macilenta 11; macrantha 5, 6|; matu-
tlai 12; miere P lr r Ua 2Si; rowllk 7;
multiflora 4t, 5, 6i, 7t, 8t, lit, 29t;
mutabilis 5; myrtifolia 11; nubigena

obovata 25,

20; quaternifolia 23; quinqueflora
28f, 29t; quinquenervata 15, 16t;
racemosa 29t; rekoi 14; rosea 27;

na 5; splen
It, 13, 14|;

steyermarkii 21;

tenuifolia 24,
olia 3t, 23, 24t, 25t, 29t,
var. angustifolia 23; tolucana 23; tri-
flora 4t. 5, var. hirsuta 5; triphylla
3|, 23, 29|, var. angustifolia 23, var.
P 23, var. splendens 23; uniflora 29|;
venosissima 12; versicolor 5, 6t, var.
graciliflora 5; villosa 12; viminalis
24t, 26; violacea 27; viperalis 29t;
xylosteoides 12

"Bouvardia?" discolor 28t

Catesbaea erecta 18

Carphalea pubiflora 23

Cestrum spermacocifolium 29t

Cinchoneae 2t

Cinchonoideae 2t

Coursiana 2t

Croton 26t

Danais 2t

Hedyotideae 2t

Hedyotis 2t; subg. Edrisia 28t; alexan-

derae 28t; bouvardioides 27; brevipes

28 i: lruticosa 25, 2i»t; mevicana 2})j;

saxatalis 28t; spinescens 18
PHedyotis lutea 9
Heterophyllaea 2t
Hindsia 2t
Houstonia bouvardioides 27; coccinea

23, var. alba 29t; longiflora 17; ochro-

leuca 29t; triflora 5
Hymenopogon 2t

Ixora americana 23; ternifolia 23, 24t
Karwinskia 26t
Manettia 2t; coccinea 28t
PMannetia racemosa 29 1
Nacibea coccinea 28t, 29t
Rondeletia deamii 28t, hirsuta 28t,

leucophylla 28t, strigosa 29t
Rubiaceae It, 2t
Rubioideae 2t

NOTES ON HEDYOTIS (RUBIACEAE) IN NORTH AMERICA 1

by Walter H. Lewis
Missouri Botanical Garden, St. Louis

Abstract
The following rubiaceous taxa are proposed: Hedyotis acerosa var. polypremoides
(Gray) W. H. Lewis, H. alexanderae (A. Carter) W. H. Lewis, H. crassifolia f. albiftora
(Standley) W. H. Lewis, and H. nigricans var. parviflora (Gray) W. H. Lewis. Range
extensions are also given for H. australis Lewis & Moore and H. rosea Raf.

1. Hedyotis acerosa Gray var. polypremoides (Gray) W. H. Lewis, stat. nov.

= Houstonia polypremoides Gray, Proc. Amer. Acad. 21 : 379, 1886. Lectotype selected
Pr ingle 356 (GH) (isolectotype MO), Santa Eulalia Mts, nr Chihuahua, Mexico, Sept
1885 in fruit; several plants are mounted on the same sheet, some of which tend toward
the typical variety . . . therefore the fruiting specimen in the lower right hand corner is
chosen lectotype of the var. polypremoides. Also on the same sheet is the syptype Pringle
16 (GH) with plants varying as for Pringle 356; they were collected at the same locality,
but in_ flower May 1885.

=H. polypremoides var. bigelovii Greenman, Proc. Amer. Acad. 32: 291, 1897. Type
Bigelow 437 (GH).

= Hedyotis polypremoides (Gray) Shinners, Field & Lab. 17: 168, 1949.

Occasional throughout the range of the typical variety, the var. polypremoides
has stems suffruticcse below, opposite leaves, and flowers mostly pedunculate.
Stems of the var. acerosa are usually suffruticose throughout, leaves are commonly
3- or 4-verticillate, and flowers may be sessile or pedunculate. The new variety
is diploid and tetraploid where x = 11 (Lewis, Amer. Jour. Bot. 49: 855-865, 1962).

chihuahua: Santa Eulalia Mts, Pringle 1066 (MO), new Mexico: Lincoln Co, Gray,
Earle & Earle 429 (MO), nr Gray, Skehan 36 (GH, MO), W of Ruidosa, Harrison &
Hicks s.n. (GH), White Mts, Wooton 179 (GH, MO); Otero Co, 1.7 mi W of High Rolls,
Lewis 5525 (MO, SMU), 2.5 mi W of Mescalero, Lei San Miguel Co,

Las Lagunitas, 15 mi S of Las Vegas, Brandegee 11795 (MO), rv.x 'f,i

Mts, Mueller 8170 (GH) ; Culberson Co, Apache Mts, ca 28 mi NW of Kent, Correll 31667
(LL), E of EI Capitan, Waterfall 4492 (GH, MO), nr Frijole, Shreve 10245 (GH), below

1 ^rrell & Johnston 19133 (LL), N McKittrick Canyon, Lewis 5538

(MO), 3 mi E of Nickel Creek, Muller 8254 (LL); Hudspeth Co, Black Mt, Cornudas
Range, Correll & Johnston 24310 (LL), Victoria Canyon, Muller 8220 (LL), id
4787 (GH, MO).

2. Hedyotis t

= Bouvardio
2577 (UC).

Endemic to the Cape region of Baja California, Mexico and known only from
granitic bluffs along the east coast.

baja California sur: Arroyo de Leon (ca 22 mi SE of La Paz), Wiggins et al. 471
(MO), growing on steep granite walls; Arroyo del Salto, E of La Paz, Carter 2577 (UC),
on steep, granite, walls of arroyo; Saltito (ca 30 km E of La Paz), Lewis 5349 (MO, SMU),
occasional in cracks of granite slopes above beach.

By its long corolla tubes and large capsules H. alexanderae superficially
resembles many but not all species of Bouvardia; yet because of a number of well
marked characters shared with Hedyotis, the species clearly belongs to this genus.

Supported by National Science Foundation Grant No. 5042.
Ann. Missouri Bot. Gard. 55(1): 31-33, 1968.

32 ANNALS OF THE MISSOURI BOTANICAL GARDEN

(1) Stipules: scarious in Hedyotis and H. alexanderae, greenish in Bouvardia; (2)
Inflorescences: central flower of cyme shortly pedunculate in many species of
Hedyotis and H. alexanderae, never shorter but peduncles always as long as or
longer than that of adjacent flowers in Bouvardia; (3) Placentae: fused ± medially
to septum in Hedyotis and H. alexanderae, fused basally to septum in Bouvardia;
(4) Seeds: wingless in Hedyotis and H. alexanderae, winged in Bouvardia; (5)
Chromosome number of 2n = 26: known for six species of Hedyotis from Baja
California and H. alexanderae, unknown for Bouvardia; (6) Pollen morphology:
os uniquely flared into crescent shaped extensions on both sides of each colpus
(cf. Lewis, Amer. Jour. Bot. 52: 259, fig. 2, 1965) in all species of Hedyotis from
Baja California and H. alexanderae, unknown for Bouvardia; (7) Distribution:
Hedyotis and H. alexanderae native to Baja California, Bouvardia unknown to the
peninsula. Thus by gross morphology, cytology, pollen morphology and distribu-
tion H. alexanderae differs from Bouvardia, yet is strikingly similar to Hedyotis,
especially H. brevipes (Rose) W. H. Lewis, H. saxatilis W. H. Lewis and other
species also endemic to Baja California.

3. Hedyotis crassifolia Raf. f. albiflora (Standley) W. H. Lewis, comb. nov.

= Houstonia pusilla Schoepf f. albiflora Standley, Rhodora 34: 177, 1932. Holotype
Benke 5191 (F), New Iberia [Iberia Par], Louisiana.

= Hedyotis caerulea (L.) Hooker var. minor (Michx.) Torrey & Gray f. benkei Fos-
berg, Gastanea 19: 31, 1954.

The white flowered form is usually found in the same population as the more
common blue flowered form; the f. albiflora is known from Louisiana, Missouri
(Steyermark, Fl. Missouri 1400, 1963), and Texas (Brazoria Co, 5 mi N of Angleton,
Lewis 6929, MO, SMU).

4. Hedyotis nigricans (Lam.) Fosberg var. parviflora (Gray) W. H. Lewis, comb.

= H. rupicola Greenman, Proc. Amer. Acad. 32: 286, 1897.

= Hedyotis angulata Fosberg in Shinners, Field & Lab. 17: 166, 1949.

= H. nigricans var. angulata (Fosberg) W. H. Lewis, Amer. Jour. Bot. 49: 865, 1962.

A variety close to the var. rigidiuscula (Gray) Shinners but differing by having
capsules about as long as broad, globose, ± y 2 inferior and when mature with
calyx-lobes equalling their length. Infrequent in rocky crevices and hillsides in

5. Hedyotis australis Lewis & Moore.

An extension of this rarely collected though rather common early white flower-
ing species of Hedyotis is reported to include Mississippi and Oklahoma. It is also
known from Arkansas, Georgia (Chambers, Rhodora 65: 271-273, 1963), Louisiana,
and Texas and should be found in Alabama and Tennessee.

Mississippi: Madison Co, Natchez Trace Pkwy, McDougall 1217 (US); Warren Co, 6.3
mi SE of Vicksburg, Lewis 5118 (SMU). Oklahoma: McCurtain Co, Tom, Lewis 6952
(DUKE, MO, NY, OKLA).

LEWIS— HEDYOTIS 33

6. Hedyotis rosea Raf.

= Houstonia patens Elliot var, pusilla Gray, Syn. Fl. 1(2): 25, 1886. Lectotype
selected Hale s.n. (GH), Alexandria [Rapides Par], Louisiana.

= H. pygmaea Mueller & Mueller, Bull. Torrey Bot. Club 63: 33, 1936. Syntypes
Mueller 3 (NY), 4 (NY), DeWitt Co, Texas.

= Hedyotis taylorae Fosberg in Shinners, Field & Lab. 17: 169, 1949.

= H. minima (Beck) Torrey & Gray f. albiflora Lathrop, Rhodora 59: 95, 1957.
Holotype Lathrop & McGregor 35 (KANU), Sec. 32, T25S, RISE, Woodson Co, Kansas.

A very small spring (Febr-early April) annual with large light rose corollas
paling white with age; infrequently collected in the south-central U.S. The species
is known from Arkansas (Moore, Rhodora 58: 331, 1956), Louisiana, Oklahoma
(Waterfall, Rhodora 55: 201, 1953), Texas, and now Mississippi [Simpson Co,
1.4 mi SE of Pinola, Lewis 5124 (SMU, US); Warren Co, 6.3 mi SE of Vicksburg,
Lewis 5117 (SMU)], and, on the basis of the holotype of H. minima f. albiflora,
Kansas. Material in herbaria is often determined as Hedyotis (or Houstonia)
minima (Beck) Torrey & Gray which in fact is H. crassifolia Raf.

NEW AND NOTEWORTHY WOODY RUBIACEAE OF PANAMA

by John D. Dwyer and Sister M. Victoria Hayden
St. Louis University and Missouri Botanical Garden

■ species of Faramea, and four new species and one subspecies of
J diagnoses of Ronabea latifolia Aublet and Uragoga
emetica (L.f.) Baillon are included. Additional notes on Coussarea and Schradera are
presented.

1. Bathysa panamensis Dwyer, sp. nov.

Arbores parvae, ramulis laevibus minute lenticellatis, in sicco opaco-griseis.
Folia subsessilia aut pedicellis ad 0.5 cm longis; lamina obovato-elliptica aut
obovato-elliptico-subtrapeziformis, ad apicem lato-cuneata, ultime gradatim acumi-
nata, acumine ad 1.5 cm longo, versus basim lato-cuneata, basi truncata aut
plerumque brevi-auriculata, rigido-papyracea, laevis, supra glabra, infra in costa
venisque albido-appresso-pilosis, concolor, in sicco brunnea, costa supra plana, infra
prominenti, convexa, venis lateralibus ca 15, arcuatis, 1.5-2 cm distantibus, venis
tertiariis leviter pinnatiformibus; stipulae non visae, caducae. Inflorescentiae
terminales, pyramidali-paniculatae, ad 10 cm longae, pedunculo compresso tortoque,
basi ca 0.5 cm lato, ramis oppositis, divergentibus, usque ad 8 cm longis, jugis
ramorum ca 4, bene distantibus, ca 3 cm longis, cymis terminalibus, patulis,
paucinoribundis, bracteis plerumque persistentibus, patulis, angusto-lanceolatis,
maioribus ad 1.5 cm longis. Flores sessiles aut subsessiles; calyx cupula elliptica,
ca 3.5 mm longa, carnosa, puberula, lobis 4, subtriangularibus, ultime obtusis, ir-
regularibus inaequilateralibusque, ciliolatis, ca 0.8 mm longis, marginibus erosulis;
corolla tubo subcampanulato, basim contracto, tunc dilatato, 3-5 mm longo, carnoso,
extus minute villosuloso, lobis 3-4, ellipticis, 3-4 mm longis, usque ad 3 mm latis,
plerumque quam tubo longioribus, reflexis sed apice involutis; stamina 3-4, antheris
sagittatis, ca 3-4 mm longis, glabris, exsertis, dorsifixis, filamentis ca 3 mm longis,
dense barbatis praeter basim; ovarium disco quattuor lobis rotundis coronato,
uniloculare ovulis co in duabus brevibus placentis intrusis, stylo basi gracili, supra
medium dilatato, usque ad 6.5 mm longo, stigmatibus 2, erectis, ca 1.5 mm longis.
Fructus non visi.

Panama: Rio Ucurganti, Bristan 1187 (holotype MO).

This is the first report of the genus Bathysa north of South America. It is
readily distinguished by its leaf-blades being truncate or briefly auriculate at the
base.

2. Cephaelis bristanii Dwyer & Hayden, sp. nov.

Suffrutices ad 30 cm lati, caulibus in sicco nigris, laevibus, glabris vix nodosis,
plerumque prostratis et radices longos angustos ferentibus. Folia petiolis ad 4 cm
longis, ad 1.5 mm latis; lamina oblonga, apice cuneata, ultime obtusa, basi
contracto-cuneata, 9-16 cm longa, 3.5-6 cm lata, rigido-papyracea, concolor, glaber,

Ann. Missouri Bot. Gard. 55(1): 34-47, 1968.

DWYER AND HAYDEN — RUBIACEAE 35

costa supra gracili, subtus prominula aut plano-convexa, venis lateralibus 15-30,
0.5-0.8 cm distantibus, arcuatis, venis intermediis evanescentibus; stipulae connatae,
depresso-annulares, 1-2 mm longae, utraque minute obtuso-apiculata. Inflores-
centiae axillares, pedunculo solitario, ad 2 cm longo, in sicco nigro, laevi, floribus
crebris in 1-2 capitula rotunda ad 1 cm aggregatis; bracteae multae aggregataeque,
bracteis interioribus gradatim angustioribus, oblongis, acutis, puberulis. Flores
calyce infundibuliformi, vix 2 mm longo, corona cupiformi petaloidea instructo,
lobis indistinctis, fortasse 4, brevibus, truncatis vel obtusis aut saepe redactis tunc
marginibus irregulari-erosis, saepe paucis ciliis rectis ornatis, tubo marginibusque
omnino ca 0.8-1 mm longis; corolla tubo angusto-cylindrico, ad 7 mm longo, in
medio ca 1 mm lato, petaloideo, extus glabro, intus supra medium tubi dense
albido-piloso, lobis 4, usque ad 2 mm longis, obtusis; antherae 5, exsertae, oblongae,
ca 0.8 mm longae, filamentis linearibus, usque ad 2 mm longis, supra medium
tubi affixis; ovarium disco prominenti, coronuliformi, obtuso, 0.5-0.8 mm longo, in
sicco fusco, 2-loculare, stylo lineari, ca 3 mm longo, stigmatibus 2, rectis, ca 1 mm
longis, ovulis 2, basaliter affixis, septo crasso. Fructus rubri (fide Bristan).

costa rica: rain forest nr Rio Toro Amarillo, vie Guapiles, ca al ca 350 m, Godfrey
66248 (MO). Panama: bocas del toro: premontane rain forest, Quebrada Lugron &
Cerro Bonyik nr Rio Teribe, alt 300-900 ft, Kirkbride & Duke 641 (MO), darien: cloud
forest, Cana-Cuasi Trial, Cerro Campaniento nr Tres Bocas, Rio Cuasi headwaters, Kirk-
bride & Duke 1243 (MO); Cerro Pirre, Bristan 575 (holotype MO).

The new species, named in honor of the native Panamanian collector, Narcisco
Bristan, is readily distinguished by its leaf-blades being very obtuse or rounded at
the apex and having lateral veins which are numerous, strict, and parallel; these
in general give a corrugated effect to the blade. The flower heads are reduced and
compact; the flowers have delicately petaloid floral tubes with the anthers exserted
and borne on relatively elongate filaments. The Cuna name (Prov. Darien) for
the species is "hinupichica" (fide Duke).

3. Cephaelis camponutans Dwyer & Hayden, sp. nov.

Suffrutices ad 0.75 m lati, ramulis laevibus, glabris, subnodosis, radicibus
gracilibus, lignosis. Folia valde ascendentia, petiolis gracilibus, ad 3 cm longis;
lamina angusto-elliptica, apice attenuato-acuminata, acumine ad 3 cm longo,
basi vix inaequilaterali, acuta, ad 18 cm longa, ad 5 cm lata, subcoriacea, laevis,
glabra, in vivo subtus purpureo-fusca, costa subtus vix prominula ad plana, proxi-
maliter ca 2 mm lata, venis lateralibus 15-20, in sicco evanescentibus, fortiter
ascendentibus, marginibus leviter crassis; stipulate obovato-rotundae, apice obtusae
(vix bifidae?), ad 1.5 cm longae, ± 1 cm latae, tenues, raphidibus minutis ornatae,
marginibus erosulosis. Inflorescentiae axillares, 1-3 per axillam, terminaliter in
pedunculo dispositae, cernuae capitulataeque, pedunculo ad 1 cm longo, capitulis
compresso-rotundis, ad 2 cm latis, in sicco nigris, bracteis exterioribus solitariis,
subovatis, ad 1.5 cm longis, coriaceis, in dorso mediano-carinatis, bracteis interiori-
bus minoribus angustioribusque, jugo florem solitarium includente, apice eroso-
marginatis. Flores ex parte vidimus; calyx tubularis, lobis 5-6, erectis, inaequila-
teralibus, subulatis quam tubo longioribus, marginibus scariosis; corolla purpurea;
ovarium biloculare, septo integro mediano, ovulo 1 per loculum, basaliter affixo,

36 ANNALS OF THE MISSOURI BOTANICAL GARDEN

stigmatibus 2, erectis, digitiformibus, ca 1 mm longis, stylo fortasse nullo. Fructus
hie immaturi, calyce persistenti coronati, pyrenis 2, ellipticis, seminibus superficie
adaxiale piano, leviter mediporcato, superficie abaxiale convexo, endospermio laevi,
testa modice pigmentifera, fortasse bilamellosa, cellulis macroscopicis parenchymals

bocas del toro: Punta Pena, vie Chiriquicito, ca 2700 ft elev, Lewis et al. 2171 (MO).
Panama: Cerro Jefe, alt 2700 ft, Dwyer & Ganger 7346 (holotype MO).

As the axillary inflorescences are cernuous and bell-shaped the name camponu-
tans (L. nutans = bent) was chosen for the specific name. Its narrowly lanceolate
and subcoriaceous leaves with the blades purple beneath in the fresh state and the
lateral veins of the blade evanescent are unique among the Central American
species of Cephaelis. No styles were observed in several flowers dissected, although
the stigmas were evident.

■■ Dwyer & Hay den, sp. nov.— Fig. 1.
, ad 5 m altae, ramulis teretibus, laevibus. Folia petiolis 0.5-8
ad 0.25 cm latis, laevibus, glabris; lamina oblonga, apice
, ad 17 cm longa, ad 8.5 cm lata, membranacea, laevis, glabra,
prominula, ad 2.5 mm lata, subtus prominula, venis lateralibus,
ca 20, 1-1.5 cm distantibus, strictis non nisi prope marginem arcuatis, venis inter-
mediis leviter pinnatinervis; stipulatae connatae, corporibus brevibus inter petiolari-
bus, truncatis, ad 5 mm longis, lobis intrapetiolaribus connatis, ad 1 cm longis,
subscutiformibus, apice bifidis. Inflorescentiae ultime cernuae, solitariae, ad 25 cm
longae, pedunculo ad 20 cm longo, in medio ca 3 mm lato, distaliter ca 4.5 mm
lato, glabro, in sicco porcato, bracteis exterioribus 2, ad 4.8 cm longis, conspicue
nitido-rubris aut purpurascentibus, capitulis interioribus ad 1.5 cm longis rubris
ad purpurascentibus. Flores albi, ca 1.5 cm longi, calyce subquadrato, ad 3 mm
longo, dentibus 5, utroque dente irregulari-marginato; corolla tubo cylindrico, ad
10 mm longo, intus prope basin filamentorum staminum puberulo, lobis 5, tri-
angularibus, ca 3 mm longis; antherae 5, filamentis infra medium tubi affixis.
Fructus nitido-azurei, obovato-oblongi, ca 1.5 cm longi, calyce persistenti, fusco,
pericarpio carnoso, in sicco atrc-fusco>, seminibus funiculo basi loculo conspicue
affixis, subtriangularibus, superficie ventrale plana et in medio breviter sulcata,
testa fuscata.

cocle: El Valle de Anton, foot of Cerro Pilon, alt ca 2000 m, Duke 1247 (MO); id.,
Lewis et al 1759 (MO).

The new species is readily distinguished by its large pendent inflorescences
and conspicuous subscutiform stipules. It is named in honor of Panama's first
woman systematic botanist, Miss Mireya Correa.

5. Cephaelis rigidifolia Dwyer & Hayden, sp. nov.

Suffutices parvi, ramulis nodosis, teretibus sed in sicco internodiis contractis
sic angularibus, glabrescentibus. Folia petiolis 1-2 cm longis, ca 0.2 cm latis,
lignosis; lamina oblonga, apice acuta, ultime acuminata, acumine ad 1.3 cm longo,
basi acuta, ad 13 cm longa, ca 6.5 cm lata, rigido-coriacea, discolor, supra in sicco
fusca, subtus opaco-luteo-viridis, subtus aureo-strigiloso-puberula, costa supra

AND HAYDEN— RUBIACEAE

Fig. 1. Cephaelis correae Dwyer & Hayden: A, habit (X]/ 2 ); B, flower sho\
and corolla (X2); C, longitudinal section of calyx, large ovarian disc and ovary
ovules (X4); D, longitudinal section of corolla tube showing five stamens and j
of tube (X2); E, two stamens: a, axial view (X3); b, abaxial view (X6); F,

style and two stigmas (X2);

face (X3). After Lewis et al. 1759 (MO).

38 ANNALS OF THE MISSOURI BOTANICAL GARDEN

prominula, subtus prominent*, venis lateralibus ca 15, arcuatis, prominentibus,
venulis minoribus intermediis ornatis, venis tertiariis pinnatiformbus; stipulae 2,
connatae, compresso-cupiformes, ca 5 mm longae, coriaceae, utraque stipula denti-
bus 2 erectis rigidis ciliolatis, ca 3 mm longis, instructa. Infiorescentiae (hie in
fructu) ad 10 cm longae, ad 2.5 cm latae, spiciformes, pedunculo compresso-torto,
lignoso, puberulo, ramis paucis, basalibus, distantibus, ad 1.5-2 cm longis, simplici-
bus vel ramulis paucis brevibus lignosis terminatis, fructibus terminali-aggregatis,
plerumque 1-2 in utroque capitulo persistentibus. Fructus sessiles, compresso-
rotundi, didymi, apice cicatrice parvo plerumque excentrico notati, ad 0.8 cm diam
quam longi latiores, puberuli, in sicco virides, pericarpio crasso, seminibus in
X-sect. rotundis, septo crasso.

Panama: Cerro Jefe, alt ca 2700 ft, Dwyer 8075A (holotype MO).

C. rigidifolia has markedly stiff-coriaceous leaves with a golden puberulence
beneath. This vegetative character coupled with the large didynamous fruits borne
on a spike-like inflorescence readily distinguishes the species.

6. Coussarea enneantha Standley, Jour. Wash. Acad. Sci. 18: 282, 1928.

darien: La Boca de Pirre, Bristan 1272 (MO); s. loc, Duke 13781 (MO).

These are the third and fourth collections of this species, all of which are
from the province of Darien, Panama; the other collections are Terry 1476 (MO)
from the Cana-Cuasi Trail (Camp 2) and William 841 (F, photo of type) from

The mature fruits of Duke 13781 are noteworthy as the fruits of C. enneantha
have not been described. Thus: fruits stipitate for 3 mm, the stipe rufo-pubescent,
the pericarp oblong or ovate-oblong, cuneate at the apex, obtuse at the base, 2-2.5
cm long, 0.9-1.1 cm wide, pilose, capped by a cylindrical calycine tube, 5-10 mm
long, ca 2 mm diam, the lobes 4, unequal, ca 3 mm long, truncate or rounded,
the seeds solitary, vertical, the testa membranaceous, the endocarp smooth, fibrous,
the endosperm horny, turning blue-black on drying.

While the membranous seed coat has the cell pattern characteristic of Cous-
sarea, the usual wall thickenings are not apparent throughout. One side of the
seed coat is much more vascular and thicker than the other; the raphides are
abundant.

7. Duroia panamensis Dwyer, sp. nov.

Arbores ad 8 m altae, ramulis subteretibus, in sicco leviter longitudinali-
rimosis, junioribus dense pilosis, nodosis. Folia petiolis 1-2.5 cm longis, lignosis,
pilosis; lamina obovato-elliptica, apice late cuneata, acuminata, acumine ad 1.5 cm
longo, usque ad 26 cm longa, ad 15 cm lata, rigido-papyracea, subconcolor, in sicco
fusca, supra appresso-albido-ciliata, ciliis ca 1-2 mm distantibus, ad tactum scabri-
diuscula, infra pilosa, ciliis aliquibus uncinatis rigidioribusque, costa supra promi-
nula, infra prominenti convexaque, venis lateralibus ca 12, primo a costa divergenti-
bus, tunc ad marginem leviter arcuatis, in foliis maioribus 2-2.5 cm distantibus,
infra prominentibus, venis tertiariis irregulari-pinnatiformibus, marginibus ciliola-
tis; stipulae ovato-ellipticae, deciduae, hie junioribus ad 1.5 cm longis, dense pilosis.

DWYER AND HAYDEN — RUBIACEAE 3V

Flores solitarii, terminales vel axillares sessiles; calyx cupula campanulata, ca 0.8
cm longa, ca 0.5 cm lata, dense barbata, ciliis albido-luteis, adscendentibus, fortasse
4-5 mm longis, lobis ad 3 mm longis, extus dense pilosis, in fructu persistentibus;
corolla tubo tubaeformi, ad 4.5 cm longo, ca 0.6 cm in medio lato, extus dense
albido-piloso, lobis hie 5, ellipticis erectis obtusis, ca 1 cm longis, dense albido-
pilosis, intus glabris. Fructus elliptici, nuciformes, calyce persistenti coronati, usque
ad 3 cm longi, usque ad 1.5 cm lati, 5-7-costati, dense albido-luteo-pilosi, pariete
pericarpii crasso, ad 2 mm lato, placentis 2, intrusis, T-formibus, seminibus multis,
elliptico-oblongis, ca 2 mm longis, trigonis, sub lente minute favosis.

BOCAs del toro: Duwebdulup Peak, N of Rio Teribe at Quebrada Huron, behind
chief's house, alt 300-900 ft, Kirkbride & Duke 571 (holotype MO).

This is the first report of the genus in Panama. Only one other species of
Duroia is known from Central America, D. costaricensis Standley, endemic to
Sierpe, Costa Rica (Pittier 6903, type); this has smaller leaves and presumably
much smaller calycine lobes. Noteworthy is the fact that Duroia is dioecious;
Standley's type description is based on material with $ flowers. With only one
flower available for dissection I am electing to describe only its external morphology.
The flower is ? and all the material deposited at MO is fructiferous. Vegetative
and reproductive features point to Duroia, although the fruits, which seem mature,
are devoid of any gelatinous covering. Superficially the $ flower in the pressed
condition looks like a large-flowered Clitoria. In their field-notes Duke & Kirk-
bride describe the corolla tube as green-white, the anthers yellow, and the stigmas
green. Obviously cf flowers were observed in the field and perhaps were collected.
The collection site of the new species is the home of the few remaining survivors
of the Teribe Indians of Bocas del Toro.

A. Gray, Proc. Amer. Acad. 5: 180, 1861.

, Boy Scout Camp Rd, Dwyer & Elias 7510 (MO); id.,
r8391 (MO).

This is the first report of the genus in Panama. It is well known in Mexico
where it is native. Exostema caribaeum (Jacq.) Roem. & Schult. occurs in Costa
Rica and ranges from Florida to the West Indies and to Mexico.

9. Faramea caput-anguis Dwyer & Hayden, sp. nov. — Fig. 2.

Arbores ad 8 m altae, ramulis in sicco glabris, laevibus. Folia petiolis 2-4 cm
longis, laevibus; lamina oblonga, apice acuta vel acuminata, acumine ad 1.5 cm
longo, basi acuta, ad 20 cm longa, ad 10 cm lata, tenui-coriacea, plerumque subtus
luteo-viridis, minute luteo-puberula, costa supra subplana, infra prominenti, venis
lateralibus ca 20, arcuatis, prominentibus, 1-2 cm distantibus, venulis intermediis
pinnatiformibus; stipulae persistentes, triangulari-subulatae, acutissimae, ad 3 cm
longae, ad 0.8 cm latae, lignosae, glabrae. Infiorescentiae per axillam plures, hie
ad 4, pedunculis divergentibus, laevibus, glabris, ad 9 cm longis, capitulis termi-
nalibus solitariis condenso-cymosis, primo gemminoideis, oblongo-fusiformibus, ad
2 cm longis, ca 1 cm latis, bracteis exterioribus fortasse 5, late oblongo-fusiformibus,
glabris, foliosis, floribus 7-9 per capitulum, jugo utroque bracteolis duabus vel
pluribus (?) instructo. Flores glabri, calyce angusto-campanulato, ad 8 mm longo,

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 2. Faramea caput-anguis Dwyer & Hayden: A, habit, showing leaves, axillary
inflorescences, and erect stipules (ca xy 3 ); B, inflorescence showing diagrammatically in
cross section (Xl'/ 2 ); c, flower, showing calyx and corolla (X2); D, pistil with style,
the ovary and its two ovules in longitudinal section (X7); E, two ovules with basal
septum: a, longitudinal section (X7); b, face view (X7); F, seed: a, face view (XI); b,
endosperm with T-shaped septum, adaxial view (XI); G, fruit in i
A-E, after Dwyer 7248 (MO); F-G after Dwyer et al. 8236 (MO).

DWYER AND HAYDEN — RUBIACEAE 41

lobis 4, inaequilateralibus, subulatis, brevioribus, ca 4 mm longis, longioribus ca
8 mm longis; corolla tubo ca 25 mm longo, angusto ad medium, tunc supra diliatato
lobis 4, lanceolatis, acutis, ad 7 mm longis; antherae 4, oblongo-lineares, ca 5.5 mm
longae, filamentis proxime medium tubi affixis; ovarium ca 1 mm longum, unilocu-
lare, ovulis 2, orbicularibus, geminatis, ca 0.5 mm longis, jugo ovulorum septo
incompleto basali affixo. Fructus glabri, longitudinaliter elongati, oblongi, ca 1.4
cm longi, ca 1 cm lati, calyce persistenti, cylindrico, ca 5 mm longo, pericarpio
carnoso, endocarpio fibroso tenui-friabili, azureo-purpureo, endospermio corneo,
sulco ad aream junctionis seminum duorum ornato, seminibus solitariis magnis,
testa cellulis exterioribus sclerenchymatis bene distantibus, polymorphis, plerumque
oblongis aut triangularibus, 34.5-69 longis X ca 46fi latis, eis stratum unum cel-
lulorum parenchymatorum tegentibus.

Panama: Cerro Jefe, alt ca 2700, Dwyer 7248 (MO); id., Dwyer & Ganger 7375 (holo-
type MO); id., Dwyer et al. 8236 (MO).

The new species is named caput-anguis because of the fancied resemblance of
the floral heads to that of a snake. The pattern of the inflorescence is unmatched
by any Faramea seen. The field notes for Dwyer & Gauger 7375 describe the
external bracts as persistent and black-purple. The pulp of the seed is soft-
cartilaginous. The sclerenchyma elements of the testa are typical of Faramea
seeds; the testa was prepared for microscopic examination by soaking the same for
15 minutes in warm 5% NaOH solution.

10. Psychotria carnosocarpa Dwyer & Hayden, sp. nov.

Frutices, ramulis rigidis, divergentibus, cinereo-fuscis, glabrescentibus praeter
ad apicem dense hirsutis, ciliis pellucidis, subulatis. Folia petiolis ad 1.5 cm longis,
dense villosulis; lamina oblonga, apice cuneata, acumine ca 1 mm longo, basi
cuneata, ad 13 cm longa, ad 5.5 cm lata, rigido-papyracea, discolor, costa prominula,
infra conspicue villosa, ciliis crebris appresso-ascendentibus, venis lateralibus ca
12, primo stricto-ascendentibus, tunc proxime marginem arcuatis, 0.5-1.2 cm dis-
tantibus; stipulae biaristatae, corpore hie indistincto, aristis subulatis, ad 0.9 cm
longis, dense hirsutis. Inflorescentiae terminales, paniculatae, dense puberulae,
breves, ad 6 cm longae, ca 4 cm latae, pedunculo ad 2.2 cm longo, ramis paucis,
inferioribus fortasse quattuor, radiate dispositis, ad 4 cm longis; Flores non vidimus.
Fructus didymi, lobis calycinis brevibus plerumque puberulis coronati, subrotundi
vel compresso-rotundi, ±5 mm longi, 4.5-7 mm lati, in vivo manifeste carnosi, in
sicco nigri, glabrescentes, pyrenis 2, hemisphericis, testa rubro-fusca, cellulis fortasse
uno strato dispositis, subisodiametricis, pigmentis luteis ad aurantiacis coloratis,
endocarpio superficie ventrale plana sed dorsaliter ecostata et sulco alto mediano
instructa, etiam minutum basalem porum gerente, albumine non ruminato, ca 3.5
mm longo, 2.5 mm lato.

bocas del toro: Changuinola to 5 mi S at jet Rios Changuinola & Terebe, alt 100-
200 ft, Lewis, Dwyer, Elias & Robertson 964 (holotype MO).

The new species is readily distinguished from the vast majority of Psychotria
of the New World by its biaristate stipules, radiately branched panicle, and didyna-
mous fruits. The structure of the fruit and the conspicuously acuminate leaf-
blades suggest P. cuspidata Bredem and P. patens Sw. These three species differ

42 ANNALS OF THE MISSOURI BOTANICAL GARDEN

widely in the patterns of the inflorescence. Psychotria carnosocarpa can be easily
distinguished from the pair by its leaf-blades being densely hairy beneath.

11. Psychotria grandicarpa Dwyer & Hayden, sp. nov.

Suffrutices, ramulis teretibus glabris, laevibus. Folia petiolis 1-4 cm longis,
subteretibus lignosis glabris; lamina late elliptica, apice cuneata vel rotunda, basi
acuta inaequilateralique, 10-19 cm longa, 5-10 cm lata, tenui-coriacea, in sicco hie
griseo-viridis, costa supra prominula, infra prominenti et proximaliter ad 2.5 mm
lata, venis lateralibus 10-12, supra planis, late arcuatis, subtus prominulis, ultime in
venam tenuem submarginalem undulatam conjunctis; stipulae semicirculares, ad
1.5 mm longae, brunneae, deciduae. Inflorescentiae fortasse pseudo-terminales (hie
in fructu), pedunculis 3-6, umbellate dispositis, 4-6 cm longis, lignosis, glabris,
simplicibus aut ramulis terminalibus brevibus paucis instructs, floribus in capitula
terminalia, ad 0.8 cm longa, aggregatis. Fructus subsessiles aut pedicellati, pedicel-
lis ad 2 mm longis, in sicco nigri, magni, oblongi aut subfalcato-oblongi (plerumque
semine uno abortivo), apice basique obtusi, apice calyce opaco-luteo persistenti
coronato, lobis calycinis obtusis, omnino ca 5 mm longis, pubescentibus, ad 1.5 cm
longi, distincte 10-costati, pyrenis 2, utraque pyrena utrinque plana, seminibus 2
aut 1 abortu.

Panama: Cerro Jefe, alt ca 2700 ft, Dwyer 7244 (holotype MO); Cerro Jefe nr Rio
Indio, Duke 15228 (MO).

The new species is readily recognized by its coriaceous leaf-blades, its inflores-
cence with several peduncles radiately disposed, and its very large ribbed fruit.
The pyrenes which appear ruminate in cross-section suggest the sect. Mapouria.

12. Psychotria horizontalis Sw. subsp. basicordata Dwyer, subsp. nov.
Suffrutices. Folia superiora sessilia, aliquibus infimis vix petiolatis, lamina basi

plerumque cordata, marginibus in sicco crispis. Fructus puberuli.

canal zone: Fort Sherman, Dwyer 5160 (MO), veraguas: Isla de Coiba (Penal Col-
ony), Dwyer 2348A (holotype MO).

Superficially the two collections appear to be quite different from the numer-
ous Panamanian collections of P. horizontalis. On closer examination the fruits of
the material cited above exhibit the typically persistent calycine lobes, although
admittedly the pyrenes are pubescent. The fact that the leaf-blades are often
subcordate at the base distinguishes them from all other Panamanian Psychotria,
except P. insignis Standley, widely separated phylogentically from P. horizontalis.
The leaves of the new subspecies resemble those of typical P. horizontalis in hav-
ing minute domatia on the undersurface of the lamina. The domatia often have
ostioles from which project minute tufts of hairs.

13. Psychotria olgae Dwyer & Hayden, sp. nov.— Fig. 3.

Frutices ad 7 m alti, ramulis primo teretibus tunc saepe ultime subplano-
compressis laevibus vix nodosis. Folia valde ascendentia sub-sessilia aut vix petio-
lata, petiolis ad 5 cm longis, lignosis; lamina obovato-elliptica aut elliptica, saepe
angusto-elliptica, apice late cuneata, rotunda aut obtusa, basi cuneata, 4.5-11 cm
longa, 1.5-4 cm lata, crasso-coriacea, glabra, marginibus conspicue revolutis, costa
supra prominula, subtus versus basim prominula, proximaliter ad 1.5 mm lata,

DWYER AND HAYDEN— RUBIACEAE

venis lateralibus ca 6, arcuatis, 0.8-1.5 cm distantibus, supra evanescentibus, subtus
vix prominulis, gracilibus; stipulae deciduae (unam juvenilera vidimus) integrae,
triangulares. Inflorescentiae ad 11 cm longae, gemmis oblongo-rotundis, hie ca
3 mm longis; calyx ca 1.5 mm longus, cupula calycina intus jugum glandularum
minutarum nigrarum ferente, utroque jugo lobis calycinis alternanti, lobis 5,
triangularibus, ca 0.5 mm longis; corolla tubo brevi, intus basi puberulo, lobis
5, apice galeatis; anther ae hie immaturae quadratae, ca 1 mm longae, in dorso
raphidibus ornatae; ovarium septo crasso integro instructum, ovulis 2 basaliter
affixis. Fructus in vivo vivido-rubri, obovato-oblongi, ca 2 cm longi, ca 1 cm lati,
apice obtusi, basi acuti, in sicco carnosi, sulcis 12 longitudinalibus ornati, utroque
putamine superficie interiore plana superficie exteriore ruminata seminibus 2, endo-
carpio fibroso infra corpus endospermii, ad 3 mm caudate disposito, ultime acuto,
basi porum minutum ferente, testa opaco-rubra, cellulis epidermidis parenchymatis,
endospermio albo-ruminato.

Fig. 3. Psychotria olgae Dwyer & Hayden: A, habit
view showing calyx and corolla lobes (X3); b, top view
(X3); C, fruit, external view showing costae and apical <

section of fruit, ruminate endosperm shown (X1J4). A-B after Dwyer 7288 "(MO); C-D
after Dwyer et al. 8193 (MO).

); B, floral bud: a, lateral
five valvate corolla lobes
ring (Xli/ 2 );D,

44 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Panama: Cerro Jefe, alt ca 2700 ft, Dwyer 7288 (MO), 8193 (MO); betw Cerro Jefe
& Eneida, alt 2100-2900 ft, Dwyer, Duke & Dressier 8193 (holotype MO).

A combination of characters readily segregates the new species from all New
World Psychotria: the thick coriaceous leaves with the secondary veins evanescent,
the large red fleshy fruits, and the calyx cup with internal glands. Miss Olga
Herrera, in whose honor the species is named, discovered calycine glands in several
species of Psychotria found in Panama: P. horizontals Sw; P. chagrensis Standley;
P. psychotriaefolia (Seem.) Standley, P. carthaginensis Jacq., P. undata Jacq.,
P. fruticetorum Standley, and P. graciliflora Benth. The ruminate endosperm of
P. olgae suggests the sect. Mapouria.

14. Psychotria umbelliformis Dwyer & Hayden, sp. nov.

Frutices, ramulis valde ascendentibus, saepe strictis, teretibus, opaco-cinereis.
Folia petiolis ad 0.7 cm longis, dense aureo-villosulis; lamina ovato-elliptica, apice
acuta ad distincte acuminata, basi obtusa, 6-10 cm longa, 3-4.5 cm lata, rigido-
papyracea, subtus dense villosula, costa gracili, supra prominula, subtus prominenti,
venis lateralibus principalibus 6-10, arcuatis, in sicco luteis, infra prominentibus,
0.7-2 cm distantibus, venis minoribus intermediisque valde irregularibus, aliquibus
a costa stricte orientibus vel aliquibus a costa reflexis, omnibus mox evanescentibus,
venis tertiariis irregulari-pinnatiformibus, patulis; stipulae connatae, puberulae,
corporibus brevibus, truncatis, 1-2 mm longis, utroque duo cornua vix divergentia
lateralia, ad 5 mm longa, ferente. Inflorescentiae terminales, puberulae, pedunculo
3-4 cm longo, rigido, ramis 3, radiate dispositis, ca 2 cm longis, plerumque ad
apicem dilatatis, uno medio strictiore, bracteis inferioribus 2, divergentibus vel vix
reflexis, lineari-lanceolatis, ad 0.4 cm longis, capitulo solitario in utroque ramo et
terminaliter disposito, pauciflora (fortasse ±10 floribus), ad 1 cm longo, quam
longo latiore, bracteis exterioribus paucis, uniseriatim in involucrum aggregatis,
late triangularibus, ad 4 mm latis, ca 3 mm longis, leviter pubescentibus, sub-
anthesi divergentibus, bracteis interioribus angustioribus, paucis ovatis, in gemmis
fortasse jugo bractearum florem solitudinum includente. Flores calycis tubo camp-
anulato, ca 2.5 mm longo, modice crasso, puberulo, lobis 5, ovatis vel triangulari-
bus, apice obtusis vel acutis, ca 0.8 mm longis; corolla in gemmis subfusiformis sed
apice obtusa, albido-puberula sub anthesi ad 20 mm longa, subcarnosa, puberula,
intus infra medium tubum pilosa, lobis 5, sublinearibus, ca 8 mm longis, ±0.8 mm
latis, extus puberulis, patulis; antherae 5, lineari-oblongae, ca 3 mm longae, ca
0.35 mm latae, dorsifixae, filamentis gracilibus, ca 0.8 mm longis, glabris, a basi
tubi ca 8 mm affixis; ovarium vix 1 mm longum, biloculare, septo integro, ovulis
2, modice magnis, basaliter affixis, disco in sicco nigro, ca 0.6 mm alto, stylo ca 16
mm longo, ca 0.35 mm lato, stigmatibus 2, rectis, ca 2 mm longis, subcrassis.
Fructus non visi.

cocle: E slope Cerro Pilon, nr El Valle de Anton, cloud forest, alt 700-900 m, Duke
12154 (MO). Panama: degraded cloud forest, peaks of Cerro Trinidad, Duke & Kirkbride
1641 (holotype MO).

Several characters serve to distinguish the new species: the umbelliform
inflorescences which often assume the form of a Neptunian trident, the densely
villulose undersurface of the leaf-blades with the veins drying yellow, and the few-

DWYER AND HAYDEN — RUBIACEAE

flowered capitate inflorescences subtended by an involucral-like series of bracts.
The flowers are described by Duke & Kirkbride as white and presumably the shrubs

15. Ronabea latifolia Aublet, Hist. PI. Gui. Fr. 154, 1775.

R. erecta Aublet, loc. cit. 156.

Psychotria axillaris Willd. in L., Sp. PL, ed. 4 [i.e. 5] 1 : 962, 1798.

Shrub 0.5-8 m high, the branchlets quadrangular, ridged, often nigrescent,
sericeous or glabrescent. Leaves short-petiolate, the petioles 6-16 mm long, often
stout, sericeous or glabrescent; blade elliptic to oblong, acuminate at the apex,
sometimes abruptly short-acuminate, attenuate or obtuse at the base, 9-15 cm long,
3-12 cm wide, subcoriaceous, olive-black when dry, glabrous above, sparsely
sericeous beneath, sometimes abundantly so along the veins, the midvein prominent
beneath, the lateral veins 6-12, arcuate, clearly anastomosing close to the margin,
the veinlets conspicuous; stipules subpersistent, free, entire, lanceolate or subulate,
acuminate or obtuse at the apex, sericeous. Inflorescences axillary, geminate or soli-
tary, bearing a few sessile flowers, up to 8 mm long, larger in fruit, the peduncle
and bracts densely sericeous; peduncle rather thick, up to 7 mm long, equalling the
petals or smaller; bracts minute, up to 1 mm long, triangular-ovate, acute or obtuse
at the apex. Flowers ca 5 mm long, white or yellow; calyx cupuliform, the tube
ca 2 mm long, subtruncate or minutely and irregularly toothed, glabrous; corolla
with the tube ca 3 mm long, glabrous on the outside, sericeous in the throat, the
lobes suboblong, strongly cucullate, ca 2.5 mm long, glabrous on the outside,
papillose within; stamens attached to the throat of the tube; anthers oblong,
short-acuminate at the apex, ca 1.2 mm long; disc cylindraceous, 0.8-1 mm long;
ovary ca 2 mm long, the style rather thick, ca 2.8 mm long, dilated above, the
stigmas obtuse, ca 0.6 mm long. Fruits (here immature), elliptic, 5.5-7.5 mm long,
2.5-6 mm wide, black, glabrous; pyrenes 2, the ventral face of the seed plane, the
dorsal face ecostate but perhaps muricate.

Known from Panama and Costa Rica, both in lowlands and highlands.

bocas del toro: Fish Creek Mts, von Wedel 2358 (MO); Punta Pena, vie CI
alt ca 1000 ft, Lewis et al. 2187 (GH, MO, US), canal zone: W of Pine Base Camp,
Johnston 1593 (MO), darien: El Real on Rio Pirre, Duke 5425 (MO); s. loc, Duke 8351
(MO); Loma Cuasi behind Manene, Duke 13613; Tumaganti, Duke 14242 (MO). Panama:
Cerro Azul, alt ca 2000 ft, Dwyer 2796; Cerro Jefe, alt 2700-3000 ft, Dwyer & Hay den 4363
(MO); id., Dwyer & Ganger 7344 (MO), san blas: rd betw Mandinga & Cangandi, Duke
14742 (MO); betw Rio Diabolo & Rio Acuati, nr Nargana, Duke 14894 (MO); Rio Diabolo
& Rio Acuati, nr Nargana, Duke 14898 (MO).

In view of the wealth of collections from Panama of this little known species
it is deemed appropriate to present a detailed description. The species obviously
does not belong in Psychotria as is evidenced by the muricate surface of the mature
pyrenes. A less important but obvious character is the few-flowered, axillary
inflorescence, although a few species of Psychotria admittedly have axillary inflo-

16. Schradera blumii Dwyer & Hayden, Phytologia 15: 59, 196'
cocle: Cerro Pilon, El Valle, alt ca 3000 ft, Duke 14996 (MO),
alt 2900 ft, Tyson et al. 3218 (holotype MO).

[Vol. 55
46 ANNALS OF THE MISSOURI BOTANICAL GARDEN

In studying Duke 14996 we encountered the first collection with complete flow-
ers. In describing the species Dwyer & Hayden had only fruit with the persistent
calyx. It seems appropriate, therefore, to describe the flowers: calyx ca 1 mm long, ir-
regularly dentate at the apex, glabrous; corolla-tube cylindrical, surrounded for 1.7
cm of its length by the calyx, the tube and the valvate lobes up to 2.5 cm long,
the upper portion of the tube with white silky hairs within, the lobes thick,
leathery, acute; anthers rectangular, ca 6 mm long, subsessile or on filaments ca
1 mm long, dorsifixed; ovary 2-celled, the septum thick, entire, the axile placentae
bearing numerous ovules, the style ca 7 mm long, the stigmas 2, erect, subulate,
ca 2.5 mm long, papillate on the inner surface.

17. Uragoga emetica (L.f.) Baillon, Hist. PI. 7: 371, 1880.

Psychotria emetica Li., Suppl. PI. 144, 1781.
Syn. PI. 1:203,1805.

C. plagiantha Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 190, 1936.

Shrubs up to 1 m tall, the roots rather thick and gnarled; stems simple, terete,
strigose. Leaves short-petiolate, the petioles 5-16 mm long, flattened, puberulent;
blade elliptic-oblong to oblanceolate, acute to attenuate at the base, acute or short-
acuminate at the apex, sometimes abruptly acuminate, the acumen often pungent,
8-15 cm long, 3-6 cm wide, membranaceous, glabrous above, strigose beneath, more
densely so along the veins, otherwise glabrous, the costa prominent beneath, the
lateral veins 7-11, arcuate, clearly anastomosing close to the margins, elevated
beneath; stipules subpersistent, free, entire, triangular-lanceolate, sometimes subu-
late, acuminate or obtuse, puberulent. Inflorescences axillary, solitary or geminate,
subcapitate, up to 8 mm long, larger in the fruit, strigose; peduncle rather slender,
up to 7 mm long, ca equalling the petioles or shorter; bracts ovate, acuminate or
subulate, up to 2 mm long. Flowers 5-merous, white, subsessile; calyx cupuliform,
the tube 0.8-1.5 mm long, glabrous, often irregularly lobed, the lobes ovate to
oblong, acuminate or obtuse, 0.9-2 mm long; corolla ca 5 mm long, the tube
glabrous, the 5 obtuse lobes slightly shorter than the tube, elongate pilose within;
anthers narrowly oblong, ca 1.5 mm long, elongate-pilose, attached in the middle
of the tube; ovary 2-loculate, with a single basally attached ovule per locule. Fruit
bright-blue, drying black, capped by the persistent calyx, elliptic, 0.5-1 cm long,
0.2-0.25 mm wide, glabrous; pyrenes 2, twisted, each pyrene hemispherical in cross-
sect, the ventral face of the seed plane and with a median ridge, with a small
pore at the base, the dorsal face ecostate, the albumen entire.

Ranging from Guatemala to Bolivia.

bocas del toro: Changuinola Valley, Dunlap 436 (F); vie of Chiriqui Lagoon, von
Wedel 1258 (MO), canal zone: Barro Colorado I, Aviles 10 (F); id., Bailey & Bailey
510 (F); id., Hayden 122A (MO), 1038 (MO); id., Ebinger 549 (MO), darien: vie of El
Real, alt ca 15 m, Allen 955 (MO); La Boca de Pirre, Bristan 1276 (MO); S of El Real,
Duke 5047 (MO); El Real on Rio Pirre, ca 10 mi S, Duke 5445 (MO); s. loc, Duke 8345
(MO); cuipo forest nr Sante Fe, Duke 12264 (MO); forested ridge parallel to Rio Sancanti,
ca 2 mi upstream from Piria, alt ca 120 m, Duke 14386 (MO) ; Puerto St Dorotea, Dwyer
2298 (MO) ; vie of Campamento Buena Vista, Rio Chucunaque & Rio Tuquesa, Stern et al.
931 (MO). Panama: Rio Bayano above confluence with Rio Chepo, Duke 3991 (MO).
san blas: headwaters of Rio Mulatupo, Elias 1773 (MO).

DWYER AND HAYDEN — RUBIACEAE 47

collections cited above permit us to describe this important
species. The twisted pyrenes and pilose anthers immediately segregate this species
from Psychotria and Cephaelis. The axillary inflorescences are striking and are
suggestive of those of Hoffmannia. The roots have a strong and somewhat nauseous
odor and are the source of the drug ipecacuanha, but it is said to be inferior to
Cephaelis ipecacuanha. According to Duke's field notes the plant is known as
"raicillo macho" in Spanish and as "macua" or "moncoa" among the Choco
Indians (Darien, Panama & Colombia) .

We are placing this species in Uragoga with some reluctance as the limits of
Uragoga are not well-defined.

NOTES ON ASCLEPIADACEAE OF PANAMA

by Louis O. Williams
Field Museum of Natural History, Chicago, Illinois

infimieola, Gonolobus

The center of diversity of the Asclepiadaceae in North America is in Mexico
with the greatest concentration of species in Chiapas and in adjacent Guatemala.
The number of asclepiads diminishes rapidly as one proceeds southward from this
center. There are a fair number of species known in Costa Rica and Panama.
Certainly there are more to be expected as botanical exploration continues, es-
pecially in Panama. It would not be surprising if the number now known, 29
native species and one introduced, were doubled in the course of time.

The Asclepiadaceae are considered a difficult family and often, I suspect,
avoided in the field as well as in the herbarium. It is a fascinating group of plants
very much in need of a friend.

Cynanchum infimieola L. Wms., sp. nov.

Lianae parvae, herbaceae. Folia lanceolata vel angusti-ovata, acuminata,
glabra. Inflorescentia umbelliformis, pauci- ad multinora. Flores pedunculis graci-
libus; lobi calycis elliptici vel lanceolati, acuti; corolla campanulata, lobis lanceo-
latis acutis intus barbell ato-puberulentibus; lobi coronae filiformes; gynostegium
stipitatum. Folliculi ignoti.

Small herbaceous vines, the stems twining, sparsely crisped-pubescent in lines,
mostly less than 1 mm in diam, with 1 or more dactyliform glands on the in-
terpetiolar scar. Leaves lanceolate to narrowly ovate, acuminate, with ca 7 pairs
of lateral nerves, glabrous or nearly so; mature blades 1.5-4.5 cm long and 0.8-2
cm broad; petioles slender, crisped-pubescent, 3-9 mm long. Inflorescences um-
belliform, few to many-flowered, axillary or nearly so, the peduncles slender,
crisped-pubescent, mostly 6-10 mm long, the pedicels 2 mm or less long. Flowers
white, ca 2-5 mm long; calyx lobed to the base, lobes elliptic to lanceolate, acute,
sparsely puberulent to glabrous, ca 0.7 mm long; corolla campanulate, ca 2.5 mm
long; lobes twice as long as the tube, lanceolate, acute, slightly spreading at an-
thesis, barbellate-puberulent along the margins within; corona ca as long as the
gynostegium, the lobes filiform, as long as and attached near the base of gyno-
stegium; gynostegium stipitate, ca 2.5 mm long. Follicles unknown.

canal zone: K-2 rd, Dwyer & Hayden 7540 (holotype F; isotype MO), cocle: rd
to El Cope ca 7 mi from Interamerican Hwy, Correct 403 (MO). Panama: Rio Mar, along
rd to beach, Blum & Dwyer 2485 (MO); nr beach at Nueva Gorgona, 8 Oct 1961, Duke
4491 (F, MO); Isla Taboga, alt 0-186 m, 23-24 July 1938, Woodson et al. 1499 (MO).

This is unusual among the species of sect. Metastelma in that it is a lowland
species, occurring not far from the Pacific Ocean; most other species grow in the
mountains. The specific name is derived from this fact. The species is allied to

Ann. Missouri Bot. Gard. 55(1): 48-50, 1968.

WILLIAMS — ASCLEPIADACEAE 4b>

Cynanchum sepicola (Pittier) L. Wms. from which it is easily distinguished by
the filiform corona lobes attached near the base of the gynostegium instead of
lobes nearly as broad as long attached near the apex of the gynostegium.

Gonolobus inaequalis L. Wms., sp. nov.

Liana herbacea, sparse hirsuta vel glabra. Folia ovata vel oblongo-ovata,
cordata, acuminata, sparse hirsuta vel glabrescentia, petiolis gracilibus. Inflores-
centia axillaris vel extra-axillaris, subracemosa. Flores calyce usque ad basim
diviso, lobis lineari-oblongis vel lanceolatis acutis vel obtusis; corolla rotata, pro-
funde lobata, tubo perbrevi, lobis inaequalibus, 3 lobis lineari-oblongis, 2 lobis
lanceolate-ovatis, annulo subnullo; corona carnosa, 5-lobata et undulata; ap-
pendices antherarum lobo suborbiculari munitae.

Herbaceous vines, sparsely hirsute or glabrous, internodes 6-12 cm long. Leaves
ovate or oblong-ovate, cordate, acuminate, glabrous or very sparsely hirsute and
glabrescent, the blade 5-11 cm long and 2-5 cm broad, the petioles slender, ob-
scurely hirsute, shorter than the blade, 3-6 cm long. Inflorescence axillary or extra-
axillary, subracemose, 1-4-flowered, the peduncles hirsute, 1-2 cm long, the pedicels
slender, to 4 cm long. Flowers with the calyx divided to near the base, glabrous,
lobes linear-oblong to lanceolate, acute or obtuse, ca 5 mm long and 2 mm broad;
corolla rotate, the tube short and inconspicuous, deeply lobate, ca 3.5 cm across,
the lobes inequal, 2 smaller than other 3, 3 lobes linear-oblong, acute or acuminate,
15-17 mm long and ca 5 mm broad, 2 lobes lance-ovate, acute, ca 11-13 mm long
and 5 mm broad; annulus of corolla very obscure; gynostegium ca 3 mm high,
corona fleshly, 5-lobate and undulate, ca 1 mm high; dorsal appendage of the
anther with a central suborbicular lobe ca 1 mm long. Follicles unknown.

The specimens have all been determined as G. dubius Pittier but R. E. Wood-
son, Jr. had indicated some of the differences on the type sheet. The specimens
available show the unusual inequality in size of the corolla lobes which would
seem to be a tendency toward zygomorphy.

canal zone: vie of Madden Dam, alt 90 m, 8 Oct 1939, Allen 2012 (MO); Palo Seco,
/ic of Pacora, alt ca 35 m, 5 Nov 1939, Allen

Gonolobus lewisii L. Wms., sp. nov.

Lianae herbaceae vel suffruticosae. Folia ovata vel oblongo-ovata, acuminata,
cordata, utrinque strigosa. Inflorescentia subumbellata vel subracemosa, pauciflora.
Flores calyce sparse strigoso, lobis lanceolatis acutis vel acuminatis; corolla rotata,
praeter annulum ciliatum glabra, lobis ovato-lanceolatis acutis; corona erecta,
carnosa; stigma stellatum; appendices antherarum rotundatae cum utrinque pro-
cesso auriculiformi vel lunato.

Herbaceous or possibly suffrutescent vines, the stems slender and sparsely
crisped-pubescent. Leaves ovate to oblong-ovate, acuminate, cordate to the base,
strigose-pubescent on both surfaces, with 4-6 pairs of secondary veins; blade 4.5-7.5
cm long and 2.5-4 cm broad; petiole slender, sparsely strigose, 1.5-3.5 cm long.
Inflorescence subumbellate or subracemose, few-flowered, shorter than the leaves,

[Vol. 55
BOTANICAL GARDEN

the peduncles extra-axillary 1/3 as large as the stem, 2-3 cm long, the pedicels to
ca 2 cm long. Flowers with the calyx sparsely strigose, divided to near the base,
the lobes lanceolate, acute or acuminate, ca 5 mm long and 2 mm broad; corolla
green, rotate, glabrous (except annulus), 2.5-3 cm across, lobed to near the center,
the lobes ovate- lanceolate, acute, ca 10 mm long and 5 mm broad, the annulus
(outer corona) inconspicuous, barbellate-ciliate; corona erect, fleshy, "scalloped,"
ca 0.5 mm high; anther appendages rounded with a lateral auriculiform or lunate
process on either side; gynostegium ca 2 mm high and 5 mm broad, the stigma
stelliform. Fruits unknown.

Species of the subg. Gonolohus are an interesting lot often quite easily dis-
tinguished by characters in the gynostegium. This is the only one I know with
lateral processes on the dorsal appendages of the anthers. Vegetatively it is like
a dozen others but the umbelliform inflorescence is less compact than in other
Panamanian species.

los santos: cloud forest & disturbed margins, Loma Prieta, Cerro Grande, alt 2400-
2800 ft, 8 June 1967, Lewis, Baker, MacBryde & Oliver 2248 (holotype F; isotype MO).

BOMBACACEAE NEOTROPICAE NOVAE II.
NEW SPECIES OF ERIOTHECA, HAMPEA AND QUARARIBEA

by Andre Robyns

Missouri Botanical Garden, St. Louis, Missouri & National Foundation for

Scientific Research, Belgium

Seven species from the neotropics are described as new to the Bomhacaceae: Eriotheca
peruviana (Peru), Hampea dukei (Panama), H. micrantha (Panama), Quararibea bilobata
(Peru), Q. longitubulosa (Peru), Q. sanblasensis (Panama), and Q. wurdackii (Peru). In

Eriotheca peruviana A. Robyns, sp. nov. [subg. MUlea (Standley) A. Robyns];
a E. discolori (H.B.K.) A. Robyns (Bull. Jard. Bot. £tat Brux 33: 159, 1963) et
E. ruizii (K. Schum.) A. Robyns (loc. cit. 162) floribus longioribus et 3.2-4 cm
longis, staminibus numerosioribus et 75-80 primo visu sat distincta; etiam E.
vargasii (Cuatr.) A. Robyns (loc. cit. 165) affinis, sed floribus leviter longioribus
pedicello dense minuteque stellato-puberulo, receptaculo calyceque extus minute
stellato-tomentello, staminibus tantum 75-80 valde differt. — Fig. 1.

Arbor (?), decidua, ramulis teretibus minute puberulis sed glabrescentibus.
Folia alterna, digitata, 5-foliolata; petiolus teres, basi leviter dilatato-complanatus
apiceque leviter dilatato-subglobosus, usque ad 13 cm longus, sparsissime pilosus
pilisque simplicibus, foliola articulata, sessilia; lamina elliptica ad subobovata,
basi cuneiformis, apice acuta ad breviter acuminata inconspicueque mucronata,
usque ad 10.7 cm longa et 4.5 cm lata, membranaceo-chartacea, marginibus ser-
rulatis, leviter discolor, supra glabra laevisque (ad scabridula et stellato-puberula?),
subtus pallidiora et dense scabridulo-stellato-arachnoidea, nervo mediano infra
prominenti, nervis lateralibus infra vix prominulis. Infiorescentiae praecoces,
cymoso-paniculiformes, laxiflorae, axibus pedicellisque dense minuteque stellato-
puberulis. Flores usque ad 4 cm longi; pedicellus usque ad 8 mm longus, 3-bracteo-
latus bracteolisque caducis; receptaculum 1.5-2 mm longum, minute stellato-
tomentellum, pauciglandulosum; calyx cupuliformis, apice truncatus vel vix sub-
lobatus, 3.5-5 mm longus et apice ca 5 mm diam, extus minute stellato-tomentellus,
intus basi longe sericeo-villosus apiceque breviter adpresseque sericeus, persistens;
petala oblongo-linearia, basi tubi staminei adnata, 3.2-4 cm longa et 4-5 mm lata,
utrinque praeter basin glabram velutina; stamina 75-80, glabra; tubus stamineus
cylindricus, 6.5-7 mm longus, in medio 2-2.3 mm diam, basi leviter dilatatus, apice
disciformi-dilatatus discoque 3.5 mm diam 5-undulato et sulcato; filamenta in
disci sulco inserta, erecta, filiformia, inaequalia, 1.5-2.3 cm longa, apice dilatata;
antherae oblongae, horizontales, extrorsae, ca 0.8-1.5 mm longae, uniloculars,
longitudinaliter dehiscentes; ovarium sessile, plus minusve piriforme, ca 2.5 mm
longum et basi 2 mm latum, tomentellum, 5-loculare, ovulis oo in columella cen-
trali affixis, stylo filiformi usque ad 2.5 cm longo praeter basin stellato-puberulam
glabro, stigmate truncato (vel inconspicue 5-undulato?). Capsula ut videtur

Ann. Missouri Bot. Gard. 55(1): 51-59, 1968.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

saf (Xi/ 2 );

2); D, androecium (Xli/ 2 ); E, anther (X15);
(X13). After Hutchinson et al. 6228.

obovoidea, apice breviter apiculata, valvis ochraceis induratis extus stellato-puberulis
pilisque hyalinis; semina late ovoidea, ca 4 mm longa et 3.5 mm lata, testa fusca
minuteque punctata; lana copiosa ferrugineaque.

Peru, la libertad: Prov Pataz, Canyon of the Rio Maranon, E side of river on rd to
Buldibuyo, 5 km above Chagual, alt 1300 m, 9 Aug 1964, Hutchinson, Wright & Straw
6228 (holotype UC, isotype US).

As the type collection is leafless, the description of the leaves was made from
cultivated plants originating from the seeds of the type collection (Honolulu
Botanical Garden, cultivation number 65.1166, pressed by Hutchinson in July
1967 and deposited at UC under the type collection number).

Hampea dukei A. Robyns, sp. nov.; H. punctulatae Cuatr. (Phytologia 4: 472,
1954) afnnis, sed floribus hermaphroditis multo minoribus et usque ad 14 mm longis
valde distincta.

53

Arbor trunco 15 cm diam, ramulis novellis dense et minute granuloso-tomentel-
lis pilisque ochraceis et stellatis sed glabrescentibus. Folia alterna, simplicia, longe
petiolata petioloque tereti usque ad 14 cm longo stellato-tomentello ad stellato-
puberulo; lamina latissime ovata, basi aperte cordata ad late rotundata, apice
breviter acuminata, usque ad 15 cm longa et lata, tenuiter chartacea, marginibus
integris, discolor, atro-punctulata, supra praecipue secus nervos minute stellato-
puberula glabrescensque, infra pallidiara et stellato-tomentella, e basi distincte
5-7-nervia, nervis principalibus supra prominulis infraque valde prominentibus.
Flores axillares, paucifasciculati fasciculisque usque ad 4-floris, hermaphroditi,
pedicellis longitudinaliter striatis usque ad 15 mm longis dense et minute granuloso-
tomentellis pilisque ochraceis et stellatis; bracteolae non visae; calyx campanulato-
cupuliformis, usque ad 6 mm longus, apice 5-lobatus lobisque transverse anguste
triangularibus et vix 1 mm longis, extus minute denseque ochraceo-stellato-tomen-
tellus, intus glaber et atro-punctatus; petala 5, usque ad 14 mm longa, tubo
ventricoso ca 6 mm longo glabroque, lobis albis contortis inaequilateralibus obovatis
leviter cucullatis extus dense minuteque stellato-tomentellis intusque glabris et
atro-punctatis; tubus stamineus usque ad 3 mm longus, dense barbatus, filamentis
usque ad 2.5 mm longis glabrisque, antheris hippocrepiformibus; ovarium late
oblongum, usque ad 3 mm longum, breviter villosum, 3-loculare loculisque pauci-
ovulatis, stylo filiformi apicem versus sensim dilatato usque ad 12 mm longo apice
declinato parte inferiore glabro parteque superiore papillato. Fructus nondum

above the Cuna-Darien boundary, 21

Hampea micrantha A. Robyns sp. nov.; ab ominibus speciebus generis Hampeae
Schlecht. floribus masculinis brevissime pedicellatis parvibusque facile distincta.

Frutex vel arbuscula 2-4 m altus, ramulis conspicue fusco-punctatis, novellis
stellato-puberulis sed glabrescentibus. Folia alterna, simplicia, longe petiolata
petioloque tereti 3.5-12.5 cm longo conspicue fusco-punctato stellato-puberulo
glabrescentique, stipulis subulatis usque ad 7 mm longis stellato-puberulis cadu-
cisque; lamina aequilateralis ad interdum inaequilateralis, ovata, basi rotundata ad
late obtusa, apice acuminata acumineque obtuso, 10-25 cm longa et 4.5-12 cm lata,
tenuiter ad rigide chartacea, conspicue fusco- ad atro-punctata, marginibus integris
ad leviter sinuatis, supra glabra, infra praecipue secus nervos stellato-puberula
glabrescensque, e basi distincte 3(-5)-nervia, costa supra elevata et infra valde
elevata basique glandulo elongato ornata, nervis secundariis supra prominulis
infraque prominentibus ad prominulis. Flores axillares, paucifasciculati fascicu-
lisque usque ad 4-floris, unisexuales. Flores masculini ca 10 mm longi, breviter
pedicellati pedicelloque crasso usque ad 3 mm longo minute stellato-tomentello;
braeteolae 3, calycis basi insertae, anguste oblongo-ovatae, usque ad 2.5 mm longae,
minute stellato-tomentellae; calyx cupuliformis, apice truncatus et breviter 5-
denticulatus, ca 4.5 mm longus et apice 4 mm diam, fusco-punctatus, extus minute
stellato-tomentellus, intus glaber; petala 5, tubo obconico farcto ca 2.5 mm longo
glabroque, lobis contortis inaequilateralibus obovatis 7.5 mm longis et 4 mm latis

54 ANNALS OF THE MISSOURI BOTANICAL GARDEN

conspicue fusco-punctatis extus praeter partem glabram petalum vicinum tegentem
minute stellato-tomentellis intusque glabris; tubus stamineus anguste conicus,
farctus, apice breviter 5-lobulatus, usque ad 4 mm longus, glaber, parte dimidia
superiore filamenta ferens filamentisque brevissimis et vix 1 mm longis glabrisque,
antheris ca 45 1-2 in quoque filamento hippocrepiformibus; pistillum nullum.
Floras foeminei non visi. Capsula calyce persistent! circumcincta, distincte stipitata,
late obovoidea, apice emarginata et minute mucronulata, usque ad 1.5 cm longa,
coriacea, extus minute fusco-tomentella, in valvis 3 intus glabris loculicida; semina
1-2 in quoque loculo, usque ad 8 mm longa, testa nigro-fusca cum venulis pal-
lidioribus, arillata.

Panama. Panama: betw Cerro Jefe & "School House," NE of Cerro Azul, forest, thicket
at edge of road, Dressier 3227 (staminate flowers, holotype MO); betw Cerro Jefe & La
Eneida, by rd, 16 Febr 1968, Dressier 3383 (capsules, MO).

Only one species of Hampea [H. appendiculata (J. D. Sm.) Standley] was
reported in my revision of the Bombacaceae for the Flora of Panama (Ann. Mis-
souri Bot Gard. 51: 62, 1964). A key which permits separation of the three
species now reported from Panama follows:
a. Leaf blades auriculate-appendaged at the base; flowers 16-18 mm long

H. appendiculata

aa. Leaf blades without auriculate appendages at the base; flowers up to 14 mm

b. Leaf blades very broadly ovate, shallowly cordate to broadly rounded at
the base, short-acuminate at the apex, stellate-tomentellous beneath; flowers
to 14 mm long; pedicel to 15 mm long; staminal tube densely barbate,
indument ochraceous H. dukei

bb. Leaf blades ovate, rounded to broadly obtuse at the base, acuminate at the
apex, stellate-puberulous especially along the veins to glabrescent beneath;
flowers to 10 mm long; pedicel to 3 mm long; staminal tube glabrous;
indument not ochraceous H. micrantha

Quararibea bilobata A. Robyns, sp. nov. — Fig. 2

Arbor 10 m alta, ramulis novellis breviter stellato-ferrugineo-tomentellis sed
glabrescentibus. Folia alterna, simplicia, breviter petiolata petioloque crasso 0.8-1
cm longo breviter stellato-ferrugineo-tomentello sed glabrescenti, stipulis deltoideis
ca 0.3-0.6 mm longis persistentibusque; lamina leviter asymmetrica, elliptica ad
subobovata, basi subrotundata vel obtusa acutave, apice obtusa, 12-25 cm longa
et 5.5-11 cm lata, tenuiter chartacea, utrinque sordida, supra sparse stellato-puberula
glabrescensque, infra stellato-pubescens, basi conspicue triplinervia, nervatura supra
manifesta sed non prominula, costa nervisque secundariis subtus prominentibus,
nervulis venulisque subtus reticulum prominulum formantibus. Flores solitarii,
oppositifolii (?) vel ramis brevibus inserti; pedicellus brevis, usque ad 1 cm longus
sed vulgo brevior, breviter stellato-ferrugineo-tomentellus, bracteolis deltoideis ca
1.5 mm longis nigricantibus persistentibusque; alabastra claviformia; calyx cam-
panulatus, longitudinaliter et conspicue nervatus nervisque prominentibus, usque
ad 2 cm longus, extus scaber et dense breviterque stellato-ferrugineo-tomentellus,
intus dense sericeus, apice 3-lobatus lobisque inaequalibus obtusis usque ad 6 mm
longis; petala 5, alba, anguste obovata, apice rotundata, usque ad 4.5-5 cm longa
et 1.5 cm lata, membranacea, utrinque praeter basin stellato-tomentella ad stellato-

ROBYNS — BOMBACACEAE

Fig. 2. Quararibea bilohata A.
of staminal tube with anthers, anc
section (X6). After Wurdack 2450.

puberula; androecium inclusum, 3-3.7 cm longum, tubo cylindrico apice breviter
5-lobato lobisque rotundatis 2-4 mm longis praeter basin stellato-albido-arach-
noideo, antheris sessilibus tubi apice et lobis insertis ca 1.5-2 mm longis; ovarium
conicum, adpresse stellato-tomentellum, 2-loculare loculisque 2-ovulatis, stylo
androecio longiore 3.5-4.5 cm longo stellato-arachnoideo-tomentello apicem versus
dilatato et manifeste bilobato, stigmatibus flabelliformibus usque ad 4.5-5 mm latis.
Fructus ignotus.

Peru, loreto: Prov Alto Amazonas, rainforest at upper end of Pongo de Manseriche,
Rio Marafion, alt 250 m, 26-28 Oct 1962, Wurdack 2450 (holotype UC, isotype US).

Quararibea bilohata is close to Q. amazonica Ulbr. (Verh. Bot. Ver. Prov.
Brandenburg 50: 91, 1909) from the State of Amazonas in Brazil (type Ule 37b,
probably destroyed in Berlin; photo Field Museum of Natural History 9552 at
MO). These species can be separated as follows:

a. Young branchlets and petioles shortly stellate-ferruginous-tomentellous; leaf
blades sparsely stellate-puberulous to glabrescent above, stellate pubescent
beneath; flowers 4.5-5 cm long; bracteoles deltoid, ca 1.5 mm long; calyx
densely and shortly stellate-ferruginous-tomentellous outside; staminal column
5-lobate, the lobes 2-4 mm long Q. bilohata

> ANNALS OF THE MISSOURI BOTANICAL GARDEN

Young branchlets reddish and glabrous; leaves glabrous except for a few
stellate hairs along the veins on the lower surface; flowers 3.5-3.7 cm long;
bracteoles subulate to lanceolate-subulate; calyx lepidote outside; staminal
column dentate at the apex, the teeth 0.75-1 mm long Q. am,

Quararibea longitubulosa A. Robyns, sp. nov.— Fig. 3.

Arbor 25-35 m alta, ramulis teretibus dense minuteque stellato-puberulis sed
glabrescentibus. Folia alterna ad subopposita, simplicia, petiolata petioloque
tereti 2.5-13 cm longo basi apiceque parum dilatato et dense minuteque stellato
puberulo, stipulis lineari-ovatis acutis obtusisve usque ad 19 mm longis et 4 mm
latis stellato-tomentellis mox deciduis; lamina parum asymmetrica, elliptica ad
late elliptica, interdum leviter obovata, basi cordata, apice obtusa ad breviter ob-
tuseque acuminata, 10-30 cm longa et 5.5-19 cm lata, chartacea, leviter discolor,
praecipue supra subnitida, utrinque sed praecipue secus venas sparse minuteque
stellato-puberula, basi 7-9-nervia, nervatura supra prominula, costa nervisque
secundariis subtus prominentibus, nervulis venulisque subtus reticulum prominulum
formantibus. Flares ramiflori et cauliflori, paucifasciculati, pedicello apicem versus

Fig. 3. Quararibea longitubulosa A. Robyns: A, leaf (X 1 ,
1); C, detail of a lobe of the staminal column with 2 e
ion (X7). After Wurdack 2102.

ROBYNS — BOMBACACEAE 57

sensim dilatato usque ad 2.5 cm longo minute stellato-tomentello, bracteolis prope
pedicelli basin insertis caducisque; alabastra claviformia; calyx infundibuliformis,
ca 2 cm longus, apice 3-lobatus lobisque inaequalibus rotundatis usque ad 8 mm
longis, extus minute stellato-tomentellus, intus dense sericeus; petala 5, alba, an-
guste obovata, apice asymmetrica, ca 3.5 cm longa et 9 mm lata, membranacea,
extus dense stellato-tomentella, intus villosa; androecium longe exsertum, tubo
stamineo cylindrico saltern in sicco obtusiuscule 5-subangulato basi leviter dilatato
usque ad 5.5 cm longo (lobis exclusis) stellato-tomentello indumentoque interdum
apicem versus paucis pilis glandulosis longioribusque intersperso apice in 5 lobos
antheriferos producto, lobis linearibus carnosis usque ad 2 cm longis stellato-
puberulis et sparse glanduloso-pilosis, antheris ca 12-18 in quoque lobo ses-
silibus oblongis et ca 2-3 mm longis; ovarium conicum, 5-obtuso-angulatum, stel-
lato-tomentellum, 5-loculare loculisque 2-ovulatis, stylo androecio parum breviore
dense stellato-tomentello apice parum dilatato curvato et breviter 5-lobato. Fructus
ignotus.

Peru, loreto: Prov Alto Amazonas, high rainforest along Rio Maranon nr Teniente
Pinglo, just above Pongo de Manseriehe, alt 250-300 m, occasional, 4-7 Oct 1962, Wurdack
2102 (holotype US, isotypes F, UC).

Quararibea sanblasensis A. Robyns, sp. nov.; ab affini Q. lep tandra Cuatr. (Lloydia
11: 185, 1948) foliorum lamina angustiore basi obtusa ad subrotunda et mem-
branaceo-chartacea, pedicello 6-8 cm longo, bracteolis longe persistentibus primo
visu sat recedit.— Fig. 4.

Arbor mediocris et 10-20 m alta, ramulis novellis stellato-puberulis. Folia
alterna, petiolo robusto tereti usque ad 2.4 cm longo et stellato-puberulo; lamina
plus minusve inaequilateralis, anguste oblongo-elliptica ad anguste elliptica sub-
obovatave, basi obtusa ad subrotundata, apice plus minusve longe acuminata
acumineque minute mucronulato, 12-40 cm longa et 4-11.5 cm lata, membranaceo-
chartacea, marginibus integris ad leviter sinuatis, utrinque minute puberula, e
basi manifeste 3-5 nervia, costa nervisque secundariis subtus prominulis infraque
valde prominentibus, nervis tertiis plus minusve transversis cum venulis in reti-
culum prominulum anastomosantibus. Flores oppositifolii, solitarii, longe pedicel-
lati pedicelloque tereti 6-8 cm longo stellato-puberulo, bracteolis 3 pedicelli apicem
versus insertis plus minusve distantibus inaequilateralibus subulato-deltoideis ad
anguste deltoideis usque ad 15 mm longis et basi 3.5 mm latis utrinque minute
puberulis longe persistentibusque; calyx tubulosus, usque ad 18 mm longus et 8
mm diam, apice 3(-5)-lobatus lobisque inaequalibus usque ad 4 mm longis, in
vivo viridis, extus dense minuteque puberulus, intus dense sericeus, accrescens;
petala ut androecium nondum visa; ovarium 5-loculare loculisque biovulatis.
Capsula calyce accrescenti late campanulato et usque ad 2 cm diam circumcincta,
drupacea, ovoidea, apice truncato-mamillato et minute apiculato, usque ad 3 cm
longa et 1.5 cm diam, in vivo viridis, in sicco fulva, lepidato-stellato-tomentella,
fibroso-lignosa, 5-locularis loculisque ut videtur uniseminalibus.

Panama, san blas: headwaters of Rio Cuadf, Camp Diablo (Drill Site 22, N 82.2,
E 87.8, alt 273.4 ft, seasonal evergreen forest along river, 18 Dec 1967, Duke, Robyns &
Verhoek 3634 (holotype MO); plain of Sperdi, nr Puerto Obaldfa, nr sea level, PUtier
4353 (US).

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Quararibea sanblasensis can readily be distinguished from the other seven
species of Quararibea occuring in Panama (cf. A. Robyns, Ann. Missouri Bot.
Gard. 51: 54-62, 1964 & 54: 185-186, 1967) by a combination of the following
characters: leaf blades ± inequilateral, distinctly 3-5-nerved from the base; flowers
oppositifolious; pedicels elongated, 6-8 cm long; bracteoles 3, inserted towards the
apex of the pedicel, ± distant, to 15 mm long, persistent; calyx tubular, to 18 mm
long and 8 mm in diam, wingless, broadly campanulate and up to 2 cm in diam
when surrounding the fruit; ovary 5-locular, each locule 2-ovulate; capsule ovoid,
truncate-mamillate and minutely apiculate at the apex, to 3 cm long and 1.5 cm

Quararibea wurdackii A. Robyns, sp. nov.

Arbor 6-8 m alta, ramulis novellis fusco-stellato-tomentellis sed mox glabres-
centibus. Folia alterna, simplicia, petiolo tereti apice leviter pulvinato 2.7-5 cm
longo praecipue apicem versus fusco-stellato-tomentello, stipulis caducis; lamina
anguste elliptica ad elliptica vel subobovata, interdum asymmetrica, basi obtusa
ad rotundata, apice acuminata, 17-35 cm longa et 6-14 cm lata, tenuiter chartacea,

subtus glabra, infra secus costae partem inferiorem stellato-puberula, basi con-
spicue 3(-5)-nervia, costa nervisque secundariis subtus prominentibus, nervulis
venulisque subtus reticulum prominulum formantibus. Flores axillares, ramis brevi-
bus inserti, longe pedicellati pedicelloque tereti gracili usque ad 7 cm longo minute
stellato-puberulo ad stellato-tomentello, bracteolis 3 pedicelli parte superiore in-
sertis lineari-subulatis usque ad 16 mm longis et ca 1 mm latis stellato-tomentellis
caducisque; calyx tubuliformi-campanulatus, apice 3-4-lobatus, usque ad 2.3 cm
longus, extus minute fusco-stellato-tomentellus, intus sericeus, lobis inaequalibus
acutis et usque ad 5 mm longis; petala eburnea, asymmetrica, anguste obovata,
apice rotundata, usque ad 3.5 cm longa et 7-8.5 mm lata, membranacea, extus
praeter basin stellato-puberula indumentoque parte superiore pilis glandulosis in-
termixto, intus fere glabra, marginibus glanduloso-ciliolatis; androecium longe
exsertum, tubo cylindrico saltern in sicco leviter sigmoideo usque ad 6 mm longo,
antheris in quoque lobo 6 biseriatis sessilibus oblongis inaequalibus usque ad 3
mm longis; ovarium depresse conicum minute stellato-tomentellum 5-loculare
loculisque 2-ovulatis, stylo androecium aequanti vel androecio leviter breviore
apice ut videtur obscure 5-lobulato. Fructus ignotus.

Peru: loreto: Prov Alto Amazonas, high rainforest along Rio Maranon nr Teniente
Pinglo, just above Pongo de Manseriche, alt 250-300 m, 4-7 Oct 1962, Wurdack 2129 (holo-
typeUS, isotypeF).

NEW SPECIES OF LISIANTHUS ( GENTIANACEAE )

FROM PANAMA

by Andre Robyns and Thomas S. Elias

Two species of Lisianthus L. from Panama are described as new: L. jefensis and L.

Study of collections of Lisianthus L. from hitherto uncollected areas in Panama,
viz. Cerro Jefe in the Province of Panama and the mouth of Rio Concepcion in the
Province of Veraguas, yielded the following two new species: L. jefensis and L.
scopulinus. During the December 1967 Missouri Botanical Garden's expedition to
Panama under the direction of Walter H. Lewis (in which the senior author took
part), collecting along the mouth of Rio Concepcion and in other inaccessible
areas was made possible through a grant from the U.S. Air Force Office of Scientific
Research (Grant No. F44620-67-C-0055) . This included transportation by heli-
copter from Albrook Air Force Base in the Canal Zone to sites in the Provinces of
Colon, Darien, Los Santos, Panama, Veraguas, and in the Comarca de San Bias.

Lisianthus jefensis A. Robyns & Elias, sp. nov.; a L. seemannii (Griseb.) O. Ktze.
foliorum lamina subcoriacea, inflorescentiis laxis pauciflorisque, pedicellis 1-2.5
cm longis, floribus leviter minoribus, calycis tubo longiore, corolla tubulosa lobisque
valde minoribus et tantum 4-5.5 mm longis, antheris ca duplo longioribus valde
distinctus; a L. soopulino A. Robyns et Elias foliorum lamina subcoriacea, dichasiis
simplicibus vel interdum compositis sed paucifloris, floribus minoribus, calycis tubo
longiore, corollae lobis multo longioribus et 4-5.5 cm longis, antheris majoribus et
ca 3.5-4 mm longis primo visu sat differt— Fig. 1.

Herba vel frutex 1.2-2.4 m altus, omnino glaber, caulibus viridibus teretibus
sed apicem versus subangulatis. Folia opposita, decussata, petiolata petiolisque
amplexicaulibus; lamina anguste obovata ad ovata, basi in petiolum longe attenuata
decurrensque, apice acuminata, usque ad 12 cm longa et 4 cm lata, subcoriacea,
costa subtus prominenti, nervis lateralibus utroque latere 2-3 arcuatis subtus vix
conspicuis ad inconspicuis. Infiorescentiae terminates et axillares, laxae, dichasiales,
dichasiis simplicibus vel interdum compositis sed paucifloris, pedunculis usque ad
4.5 cm longis, pedicellis 1-2.5 cm longis, bracteis oppositis et 2-5 mm longis.
Flores 2.8-3.2 cm longi, 5-meri; calyx tubulosus, 7-9.5 mm longus, lobis anguste
ovatis dorsaliter leviter carinatis apice longe acuminatis 4-6.5 mm longis et ca 2-2.5
mm latis secus margines plus minusve scariosis; corolla vivide flava cum lobis
viridi-flavis, 2.8-3.2 cm longa, tubulosa, parte ! / 3 inferiore constricta, lobis erectis
triangularibus apice acuminatis 4-5.5 mm longis et ca 3 mm latis; stamina plus
minusve corollam aequantia vel manifeste exserta, filamentis tubo corollae ca l / 3
supra basin insertis filiformibus et 19-24 mm longis, anthersis introrsis ^

Ann. Missouri Bot. Gard. 55(1): 60-63, 1968.

ROBYNS AND ELIAS— LISIANTHUS 61

Fig. 1. Lisu
C, id., longitudina"
enlarged). A, after Elias & Hayden

(MO); B-E, after Tyson et al. 3203 (MO).

oblongis ca 3.5-4 mm longis basi bilobatis apice mucronatis mucroneque curvato
et ca 0.3-0.4 mm longo; ovarium anguste ovatum, basi leviter obtuseque 4-angula-
tum, 6-7 mm longum et ca 1.5-2 mm diam, 2-loculare, stylo filiforaii plus minusve
tubum corollae aequanti ad longe exserto, stigmate capitato et ca 1 mm diam.
Capsula calyce persistenti corollaque marcescenti circumcincta, fusiformis, usque
ad 12 mm longa, apice rostrata, 2-septicidali-valvata; semina irregularia, ca 0.5 mm
longa, processibus parvis acutisque tecta.

Panama: Cerro Jefe, Duke 9413 (MO); id., 10-13 mi S of Goofy Lake, Febr 1966,
Duke 8010 (MO); id., alt ca 2900 ft, roadside thicket, Aug 1967, Dwyer & Hayden 8087
(MO, UC); id., cloud forest, Aug 1967, Elias & Hayden 1798 (holotype MO, isotype UC);

ANNALS OF '

MISSOURI BOTANICAL GARDEN

id., summit & forests along rd beyond summit, Aug 1967, Hayden 1008 (COL, DUKE, MO,
in Clusia forest, alt 2700-3000 ft, Jan 1966, Tyson et al. 3203 (MO); id., top,
very common, July 1966, Tyson et al. 4438 (MO).

This new species can readily be distinguished from L. seemannii by the sub-
coriaceous leaf blades, the loose few-flowered inflorescences, the much longer
pedicels (1-2.5 cm long in L. jefensis versus 2-5 mm long in L. seemannii), the
somewhat shorter flowers, the longer calyx tube, the tubular corolla with lobes
only 4-5.5 mm long, and the anthers about twice as long; from L. scopulinus it
can be separated by the subcoriaceous leaves, the simple or sometimes compound
but few-flowered dichasia, the shorter flowers, the longer calyx tube, the much
longer corolla lobes and the longer anthers. Lisianthus jefensis is closely related to
L. peduncularis L. Williams (Fieldiana: Bot. 31: 408, fig. 1, 1968), but differs
mainly in the more coriaceous texture of the leaves and the much smaller flowers
(2.8-3.2 cm long in L. jefensis versus 4.2-6 cm in L. peduncularis).

Lisianthus scopulinus A. Robyns & Elias, sp. nov.; L. skinneri (Hemsl.) O. Ktze.
proximus, sed calycis lobis anguste ovatis apice longe acuminatis longioribus et
5-5.5 mm longis primo visu sat differt; etiam L. jefensi A. Robyns et Elias affinis,
sed dichasiis compositis amplis multiflorisque, floribus majoribus, calycis tubo ca
dimidio breviore, corollae lobis ut antheris etiam brevioribus praecipue recedit.— Fig.
2.

Frutex 1.5 m altus, omnino glaber, caulibus teretibus sed apicem versus
angulatis canaliculatisque. Folia opposita, decussata, petiolata petiolisque amplexi-
caulibus; lamina obovata ad obovato-elliptica, basi in petiolum longe attenuata,
apice longe acuminata, 6.5-18.5 cm longa et 2.8-6.6 cm lata, chartacea, costa et
nervis lateralibus subtus conspicuis, nervis lateralibus utroque latere 2-3 arcuatis.
Infiorescentiae axillares, dichasiales, dichasiis compositis amplis multifloris laxis

ROBYNS AND ELIAS— LISIANTHUS 63

saepeque irregularibus, pedunculis teretibus ad angulatis 1.5-3.5 cm longis, pedicel-
lis 0.8-2.2 cm longis, bracteis oppositis ovatis ca 1.5 mm longis. Flares 3.8-4.6 cm
longi, 5-meri; calyx tubulosus, 6.5-7 mm longus, lobis anguste ovatis dorsaliter
carinatis apice longe acuminatis 5-5.5 mm longis et basi ca 2 mm latis secus
margines membranaceis; corolla flavo-viridis, 3.8-4.6 cm longa, tubulosa, parte y 3
inferiore constricta, lobis erectis ovatis apice acutis 2-3 mm longis et basi latis;
stamina breviter exserta, filamentis tubo corollae ca ! / 3 supra basin insertis filiformi-
bus et 2.6-3.5 mm longis, antheris introrsis versatilibus oblongis 2-3 mm longis basi
bilobatis apice mucronatis mucroneque ca 0.5 mm longo; ovarium anguste ovatum,
basi leviter 4-angulatum, 6-11 mm longum, 2-3 mm latum, 2-loculare, stylo exserto
filiformique, stigmate capitato et ca 1.5 mm diam. Capsula calyce persistent!
corollaque marcescenti circumcincta, fusiformis, 11-14 mm longa et ca 4 mm diam,
longitudinaliter carinata, apice rostrata rostroque usque ad 5 mm longo, 2-septici-
dali-valvata; semina irregularia, ca 0.5 mm longa, processibus parvis acutisque

veraguas: mouth of Rio Conception, cliffs, Dec 1967, Lewis, Croat & Hawker 2799
(holotype MO; isotypes DUKE, F, K).

The new species is related to L. skinneri but differs strongly in having narrowly
ovate calyx lobes which are long-acuminate at the apex and are longer (5-5.5 mm
long, in contrast with 2-3 mm in L. skinneri) ; it is also close to L. jefensis from
which it can easily be separated by the large, compound, many-flowered dichasia,
by the larger flowers, by the calyx tube about half as long, and by the shorter
corolla lobes and anthers; from L. peduncularis it can be distinguished by the large,
compound, many-flowered dichasia, the shorter calyx tube, the shorter corolla lobes
(2-3 mm long in L. scopulinus versus 6-8 mm long in L. peduncularis), and the
smaller anthers.

KARYOTYPES IN RELATION TO CLASSIFICATION
AND PHYLOGENY IN CLAYTONIA

by Walter H. Lewis and Yutaka Suda 1
Center for the Biology of Natural Systems, Washington
University and the Missouri Botanical Garden, St. Louis

Chromosomal morphology in Claytonia involving symmetry, large size and few satel-
lites is correlated with the gross morphologically less specialized species and thei
tion into sections, while karyotypes with asymmetrical and medium- or small-sizi
somes having the most satellites typify the advanced perennials and one annual species
studied. These data further suggest that there are at least two lines of evolul

1 1 i i from sect. Caudicosae to sect. Rhizomatosae to sect. Claytonia, and from a taxon
similar to C. sibirica to sect. Limnia.

In the classification of the genus Claytonia L. (Portulacaceae) proposed by
Gray (1887) and as recently modified (Swanson, 1966; Nilsson, 1966), one annual
and three perennial sections are recognized. Those species in the sect. Caudicosae
with heavy taproots are considered primitive, those with rhizomes or runners in
the sect. Rhizomatosae intermediate between the caudicose perennials and the
specialized geophytic species of the sect. Claytonia, while the annual species of the
sect. Limnia are also thought to be advanced. This subgeneric classification and
proposed phylogeny are based only on morphological characters to which we now
add data from chromosomal morphology.

Species of each section were studied: (1) C. sibirica L., sect. Caudicosae, 2n
= 24 (British Columbia: N of Squamish, Black Tusk recreational area, Lewis
6810, nr Sechelt, Sechelt Peninsula, Lewis 6827; Vancouver I, Cougar Creek at
Hwy 19, Lewis 6822); (2) C. cordifolia S. Watson, sect. Rhizomatosae, In = 10
(Washington: Kittitas Co, Table Mt Rd, Lewis 6736); (3) C. sarmentosa C. A.
Meyer, sect. Rhizomatosae, In = 10 and 15 from one population (Alaska: Hatcher
Pass, Talkeetna Mts, Mitchell 927D1); (4) C. virginica L., sect. Claytonia, 2n =
12 (North Carolina: Buncombe Co, 0.2 mi W of Swannanoa, Lewis 6582) and
2n = 14 (Texas: Bowie Co, Texarkana, Suda 6) ; (5) C. perfoliata Donn ex Willd.,
sect. Limnia, In = 12 (Washington: Kittitas Co, 10 mi W of Cle Elum, Lewis
6728).

Plants were grown in an underground room with 12 hr of light (500 ft
candles) at 24°C and 12 hr of darkness at 18°C. Root tips excised from pot-bound
plants were pretreated with low temperatures (0-2°C) for 16 hr and fixed by modi-
fied Carnoy's solution (4:3:1, chloroform-absolute ethanol-glacial acetic acid) for
30 min. After maceration in N HC1 for 45 min, roots were immersed in 2% acetic-
orcein for 24 hr. Temporary slides were made by the squash method; the best slides
were then made permanent (McClintock, 1929) and are deposited in the Missouri

"We wish to thank Dr. Wm. W. Mitchell, Alaska Agricultural Experiment Station,
Palmer, for sending material of Claytonia sarmentosa, and U. S. Public Health Grant No. 1
P10 ES 00139 ERT for financial assistance.
Ann. Missouri Bot. Gard. 55(1): 64-67, 1968.

LEWIS AND SUDA — CLAYTONIA 65

Botanical Garden Herbarium (MO). Chromosomal measurements were made from
photomicrographs enlarged 5600 X; idiograms (Fig. 1-5) are based on arm ratios
of chromosomes from 3-4 metaphase plates each from different root tips.

Chromosomal symmetry, length, number and satellite frequency for seven
races of five species are summarized in Table 1. Certain trends are striking. For
example, symmetry, expressed by pairs of V (± median and symmetrical) and I
(subterminal and asymmetrical) chromosomes and by percentages of subterminal
chromosomes, is of three kinds. More or less symmetrical karyotypes are character-
istic of C. cordifolia (Fig. 2) and C. sarmentosa (Fig. 3), those of C. virginica (Fig.
4-5) are strongly asymmetrical, while the karyotypes of C. perfoliata (Fig. 1) and
C. sibirica are intermediate between these extremes. This grouping correlates at
least in large part with the subgeneric classification, viz. species in the sect.
Rhizomatosae differ markedly in chromosomal symmetry from C. virginica (sect.
Claytonia), both of which differ from C. perfoliata of the sect. Lirrmia. The latter
is similar to C. sibirica which is, however, an atypical member of the sect.
Caudicosae forming a connecting link with the sect. Limnia (Swanson, 1966) a
conclusion entirely confirmed by the degree of chromosomal symmetry.

On the basis of chromosomal lengths the five species separate into four groups
corresponding exactly to their sectional classification (Table 1). Shortest chromo-
somes averaging 2.2/u. are found for the annual C. perfoliata (sect. Limnia), those
of C. sibirica (sect. Caudicosae) are short to medium, those of C. virginica (sect.
Claytonia) are still longer, while the longest chromosomes averaging 8fi and 8.6/*
are found for C. cordifolia and C. sarmentosa (sect. Rhizomatosae), respectively.
Since phylogeny and chromosomal size may be correlated with plants having larger
chromosomes lacking evolutionary specialization (Davis & Heywood, 1963), the
more primitive C. cordifolia and C. sarmentosa should possess large chromosomes.
They do and, in fact, are the longest in the genus. The annual C. perfoliata with
many specialized features (Swanson, 1966) has in contrast the smallest chromo-
somes; hence small chromosomes and evolutionary advancement are seemingly
correlated as found for example in Crepis (Babcock et al., 1942). Chromosomes
of the annual or perennial C. sibirica are also small and thus parallel C. perfoliata
in this character as well as in symmetry and number. But only the tetraploid race

ngth, symmetry, and number of satellites per plate

2.5-3.7
6.4-9.8
7.5-9.8

4.0-7.3
3.9-5.4

1.6-2.5

W ANNALS OF THE MISSOURI BOTANICAL GARDEN

of C. sibirica was examined and as polyploids within a ploidy series generally have
smaller chromosomes than diploids a direct comparison between this tetraploid
and the other species studied (only diploid races) is not really possible. As noted
the chromosomes of C. virginica are intermediate in size, yet a relationship with
those of C. cordifolia and C. sarmentosa is indicated, i.e. structural alterations and
loss in one arm of each chromosome of C. cordifolia (Fig. 2) or C. sarmentosa (Fig.
3) would give rise to smaller, asymmetrical chromosomes typical of C. virginica
(Fig. 4-5).

We noted also that the number of satellites per karyotype varied by species
and section (Table 1): maximum satellite frequency of species in the morphologi-
cally evolved sect. Claytonia and Limnia is 3.3 per mitotic plate, whereas those
species in the more primitive sect. Rhizomatosae and the tetraploid C. sibirica
average only 1.3 per plate. It appears that a multiplication of satellites is related
to evolutionary advancement of chromosomes and of species per se quite apart
from their level of ploidy.

Finally brief mention should be made of two infraspecific chromosomal differ-
ences illustrated by our results. Claytonia sarmentosa has been examined from only
two localities in Alaska and already plants with 2n = 10, 14, 15, 16, 28 and 32
are known. Further sampling will undoubtedly expand this impressive infra-
specific aneuploidy and polyploidy and hopefully lead to an understanding of
such divergence. These numerical differences, however, are not unique: C. virginica
is known with no fewer than 45 races at many levels of ploidy and including
extensive aneuploidy even at the diploid level (Lewis, 1967; Lewis et al., 1967a;
Lewis et al., 1967b). A hint as to the origin of a diploid race is suggested by the
2n ~ 12 cytotype (Fig. 4) which has a large median pair of chromosomes with
secondary constrictions, a pair clearly absent from the 2n = 14 race (Fig. 5). It
would not be difficult to imagine breakage of the median pair at the centromere to
form two ± similar pairs, such as those observed in the 2n = 14 race, in which
the (now) subterminally positioned secondary constrictions function as centro-
meres. Such an occurrence would give rise to a 2n = 14 race from a presumed
base of x = 6 for the species.

Literature Cited
Babcock, E. B., G. L. Stebbins, Jr. & J. A. Jenkins. 1942. Genetic evolutionary processes

in Crepis. Amer. Nat. 76: 337-363.
Davis, P. H. & V. H. Heywood. 1963. Principles of Angiosperm Taxonomy. D. v.

Nostrand Co., Princeton.
Gray, A. 1887. XV. Contributions to American Botany, Portulacaeeae. Proc. Amer. Acad.

22: 272-285.
Lewis, W. H. 1967. Cytocatalytic evolution in plants. Bot. Rev. 33: 105-115.
, R. L. Oliver & Y. Suda. 1967a. Cytogeography of Claytonia virginica and its

allies. Ann. Missouri Bot. Gard. 54: 153-171.
, Y. Suda & B. MacBryde. 1967b. Chromosome numbers of Claytonia virginica

in the St. Louis, Missouri area. Ann. Missouri Bot. Gard. 54: 147-152.
McClintock, B. 1929. A method for making acetocarmine smears permanent. Stain

Tech. 4:53.
Nilsson, O. 1966. Studies in Montia L. and Claytonia L. and allied genera, III. Pollen

morphology. Grana Palynolog. 7: 279-363.

1966. A synopsis of relationships in Montioideae (Portulacaeeae). Brittonia

NSON, J. R. 1

18:229-241.

I II II li II

I II II II N

I II ii II

I II II II

li .. .. .. .. ..

li ll ll ll li ii

ii ii ii ii ii ii u

Fig. 1-5. Chromosomal idiograms of Claytonia species. Fig. 1. C. perfoliata, 2n = 12
(Lewis 5728). Fn li Fig. 3. C. sarmentosa, In =

10 (Mitchell 927D1). Fig. 4-5. C. virginica, 2n = 12 (Lewis 6582), 2n = 14 (Suda 6).

NOTES

A NEW SPECIES OF STELIS ( ORCHIDACEAE ) FROM PANAMA

Stelis fimbriata R. K. Baker, sp. nov.— Fig. 1.

Herba caespitosa, epiphytica, glabra, usque ad 15 cm alta, caulibus secundariis
brevibus gracilibus ad 5 mm longis, vaginis tubularibus deciduis usque ad 10 mm
longis. Folia lamina coriacea elliptica ad ovata marginata 4-5 cm longa et 2-3 cm
lata apice obtusa minuteque tridentata basi in petiolum 5-8 mm longum breviter
angustata. Inflorescentia usque ad 13 cm longa, folio subtento multo longior, parte
% distali florifer, bracteis infundibuliformibus roseis 1 mm longis pedicellos
persistentes 1.5 mm longos includentibus. Flores rosei et albi; sepala 3-nervia, late
ovata, 3 mm longa, 2.5 mm lata, rosea, basibus marginibusque albis, apicibus
obtusis aut rotundatis, marginibus undulatis dense fimbriatis candidisque; petala
claro-lutea, depresse obovata, undulata, 0.6 mm longa, 0.8 mm lata, secus costam

' $? S I

; fimbriata R. K. Bakar: A, habit (XI); ]

t Bot. Gard. 55(1): 68-79,

NOTES 69

solitariam aliquantum incrassata, aliter ecarnosa; labellum luteum, ovatum, undu-
latum, 1 mm longum, 0.8 mm latum, in unguem brevem attenuatum, costae dimi-
dio basali in callum angustum longitudinalem unguem breviter acuteque bifurcatum
incrassato, venis lateralibus parallelis etiam incrassatis sed leviter brevioribus;
columna lutea, 0.6 mm longa, valde alata, basin versus angustata, alis labelli
margines basales subinvolutos amplectentibus; ovarium gracile, pedicellatum, 1.4
mm longum.

Herb, caespitose, epiphytic, glabrous, up to 15 cm tall; secondary stems short,
slender, up to 5 mm long; sheaths tubular, deciduous, up to 10 mm long. Leaves
4-5 cm long, 2-3 cm wide, coriaceous, elliptic to ovate, marginate, obtuse, the apex
minutely tridentate, the base attenuated into a short petiole up to 5 mm long.
Inflorescence up to 13 cm long, much longer than subtending leaf, upper % flori-
ferous; bracts pink, 1 mm long, infundibuliform, enclosing a persistent 1.5 mm
pedicel. Flowers pink and white, showy; sepals 3-nerved, broadly ovate, 3 mm long,
2.5 mm wide, pink, with clear white bases and margins, the apex obtuse to rounded,
the margin densely white-fimbriate, undulate; petals bright yellow, depressed
obovate, undulate, 0.6 mm long, 0.8 mm wide, somewhat thickened along a solitary
mid-vein, otherwise not fleshy; lip yellow, ovate, undulate, 1 mm long, 0.8 mm
wide, attenuated into a short claw, the basal half of the mid-vein thickened into
a narrow longitudinal callus which is shortly and acutely bifurcate at the claw,
the parallel side-veins also thickened, but somewhat shorter; column yellow, 0.6
mm long, strongly winged above, tapering toward the base, the wings clasping the
upturned basal margins of the lip; ovary slender, pedicellate, 1.4 mm long.

Panama: Cerro Jefe, in mossy forest nr summit, alt 2900 ft, 11 Febr 1967, Baker 249
(holotype MO; living specimen MBG 67-73-137).

This attractive species (subg. Stelis, sect. Concavae Garay) is reminiscent of S.
eublapharis Reichb. f., but is distinctive in the relative simplicity of the lip, and
in the absence of pronounced marginal thickenings of the petals. — R. Kendall
Baker, Department of Botany, Washington University & Missouri Botanical Garden,
St. Louis.

A NEW SPECIES OF AFRICAN LAGENARIA
(CUCURBITACEAE)

While on a botanical expedition to the northern regions of Ghana during the
latter part of December, 1966, I came across a curious Lagenaria at Gambaga near
Wali Wali. The plant is cultivated for shells which are used for manufacturing
musical drums and vessels. The fruits are large with bitter pulp and two-horned
(bicornate) seeds. The description and illustrations are based on plants raised in
the college nursery from seeds gathered during the expedition.

Lagenaria bicornuta Chakravarty, sp. nov.; L. siceraria (Molina) Standley affinis
a quo semine majore et bicorni, fructu giganteo et fere rotundo differt— Fig. 1-2.

Monoecious vines with thick, 5-ridged, hairy stems which harden at maturity.
Leaves with petiole erect, thick, biglandular (glands turbinate 2-3 mm in diam at
the base) at apex, merged with narrow marginal basal veins of the blade; lamina

softly hairy, cordate-orbicular, shortly 7-angled or lobed, apex filiform-i
10-25 cm long and 15-35 cm broad, the margin glandularly serrate except mucronate
midrib-end, the basal sinus deep-semilunar, 3-costate, 2 lateral costa divided at the
base and outer lateral vein again redivided near the base; veins reticulate promi-
nently raised on the lower surface, almost at the same level with the blade on the
upper. Flowers axillary, solitary, male and female usually occurring on different
branches, the female branches few and arising laterally from the longer and more
vigorous male branches. Male flowers with peduncle 5-ridged, longer than the
petiole (10-25 cm long); perianth tube deeply cupular ca 1 cm in diam, the lobes
triangular remote softly hairy; corolla lobes white later brownish-white, 5-6 cm
in diam, ± spathulate, the incurved margin somewhat lobed, the midrib with a
bearded apex, mucronate, the veins whiter than brownish-white blade and distinctly
raised on the lower surface, ca 3 cm long, and 2 cm broad at upper part, the
lower part ± cuneate, unlobed and thicker, thickly 5-nerved; stamens 3, attached
at base of throat of tube, 2 bithecial, 1 monothecial, the filaments of two thicker
than the third, 5-6 mm long, with a glandular cup-shaped disc (metamorphosed

Fig. 1. Lagenaria bicornuta Chakravarty: A, part of one flowering branch showing
male and female flowers (X'/ 2 ); B, male flower split open to show back portion oi ' I
and monothecous stamens, center, and part of bithecous stamen, left & right side (Xy 3 );
C, petal of female flower showing bearded mucronate apex (X|/ 2 ); D, female flower with-
out petals showing hairy ovary and thickened stigma (Xi/ 2 ); E, longitudinal
female flower (Xi/ 2 ); F, cross section of ovary (Xi/ 2 ); G, seeds (Xy 4 ).

72 ANNALS OF THE MISSOURI BOTANICAL GARDEN

pistillodium) forming honey chamber, the anther not glossy, ca 1 cm long, 0.7-0.8
cm broad, loculi long convoluted in an ornamental pattern, the lower surface of
anther (connective) smooth and creamy white, shortly ovate- acuminate, glandular
hairs present between the folds of loculi. Female flowers with peduncle much
shorter than the petiole (2-5 cm long) ; perianth tube very short or almost absent;
corolla lobes much smaller than in the male; ovary oblong, conspicuously softly
hairy, triplacentiferous, the style short, the stigma trilobed each lobe again deeply
bilobed i.e. stigma apparently 6-lobed, the lobes glabrous on upper surface, the
lower hairy. Fruit very large, almost perfectly round, 35-40 cm in diam, almost
white when fully mature; seeds bicornate, ca 3 cm long (including horns), ca 1.1
cm broad, the horns somewhat incurved, ca 4 mm thick, 2-3 mm high, the surface
ornamentally margined, the margin lines having pyramidal apex and open base.

Ghana, cape coast: Univ Farm Akotokyir, 20 Febr 1967, Chakravarty C1001: K, Kl
holotype, K2 (UCCC); C1002 (UCCC); C1003 (K, UGCC).

The species differs from Lagenaria siceraria (Molina) Standley by the presence
of conspicuous two-horned seeds (almost double the size of the seeds of L. siceraria)
and in having gigantic (35-40 cm in diam) and almost perfectly rounded fruits.—
H. L. Chakravarty, Department of Botany, University College, Cape Coast, Ghana.

NOTES ON THE GENUS INGA. II

Two poorly known species of Inga can now be adequately described as to
flowering and fruiting material and one of the species, /. brenesii, placed in its
proper section. Distributional records from recent field work for Darien and San
Bias areas of Panama are also given for several members of the genus.
Inga brenesii Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 495, 1937.

This species previously known only from two collections from the forest of the
central highlands of Costa Rica was placed by Leon (Ann. Missouri Bot. Gard.
53: 330, 1966) in sect. Inga without having seen the fruit. Series Inga is distin-
guished from all other series and sections of Inga by having a rounded, sulcate
legume. Recently, however, a specimen was collected from Costa Rica (Jimenez
2620 F, 2 sheets & 1 bag fruit) with both flowering and fruiting material thus
making possible the accurate placement of this species. The fruit is flattened, 20
cm long, 3-3.4 cm broad, 1.1 cm thick, densely ferrugineous-hirsute, slightly curved
with elevated margins.

Although varying in two minor series characters, gland shape and stipule dura-
tion, this species as supported by other vegetative, floral and especially fruiting
characters should now be placed with sect. Inga ser. Vulpinae sensu Leon. Series
Vulpinae is restricted to South America excepting this species and the closely
related /. tonduzii J. D. Sm. both of which are found in Costa Rica but are not
known from Panama.
Inga saffordiana Pittier, Contr. U.S. Nat. Herb. 18: 176, 1916.

This species previously known only from an incomplete specimen collected in
the forest of Cerro de Garagara in Darien, Panama was believed to be intrageneri-

NOTES 73

cally unique in exhibiting a cauliflorous condition. The type specimen, Pittier
5676 (US) is without flowers and has only immature legumes. One of the distin-
guishing characters used by Pittier was that the inflorescences seem to come from
the old wood of the trunk. Leon (Ann. Missouri Bot. Gard. 53: 354, 1966)
described the flowers apparently from a Colombian collection (Fernandez 267)
which he regarded as matching the type closely. The Colombian specimen differs,
however, by having only 2-3 pairs of leaflets and a long pedunculate inflorescence
which is ramiflorous instead of being cauliflorous. In August, 1967 James Duke and
I discovered on the slopes of Cerro Pirre in Darien, Panama a specimen of /.
saffordiana (Duke & Elias 13865 GH, MO). The tree was small, forming a part of
the understory and its flowers were cauliflorous with some of the inflorescences
issuing from older branchlets. The flowers are long-pedicellate, the calyx conical,
10-12 mm long, with small, acute teeth, the corolla tubular, 14-17 mm long,
expanding slightly toward the apex, the lobes 3-4 mm long, ovate, acute at the
apex. Both the calyx and the corolla are covered with spreading hairs. Since the
flowering material of the Duke & Elias collection matched the description of the
Colombian specimen, I feel that this species can now be adequately described as
to flowering and fruiting material and is probably restricted to the mountain slopes
in Darien, Panama and the neighboring Choco region in Colombia.

Recent collections in Panama in the province of Darien and the Comarca de
San Bias have yielded species which were previously unknown from those areas
and the eastern one-third of the country.

I. thibaudiana DC: nr Rio Canglon, Duke & Bristan 356 (MO).

I. multijuga Benth.: Puerto St. Dorotea, Dwyer 2226 (MO).

/. pauciflora Walp. & Duchass.: Rio Tuira betw Rio Punusa & Rio Mangle, Duke 14609

(MO); Rio Pucro, below village of Pucro, Duke 13127 (MO); along Rio Tuira below

El Real, Stern et al. 968 (MO).
/. hayesii Benth.: 3 mi E of Santa Fe, Tyson et al. 4669 (MO); vie of Campamento Buena

Vista, nr Quebrada Felix, Stern et al. 964 (MO).
1. spectabilis (Vahl) Willd.: vie of El Real, Stern et al. 762 (MO).
/. portobellensis Beurl.: main stream of Rio Cuasi, 0-2.5 m S of Tres Bocas, Kirkbride &

Duke 1132 (MO, NY).

J. multijuga Benth.: opposite Ailigandi, Lewis et al. 149 (GH, K, MO, UC, US), 194 (GH,
MO, UC, US); along headwaters of Rio Mulatupo, Elias 1728 (GH, MO, US).

7. spectabilis (Vahl) Willd.: 139 (GH, K, MO, NY, UC);

Rio Ailigandi, Duke 10838 (MO).

/. goldmanii Pittier: opposite Ailigandi, Lewis et al. 190 (GH, MO, UC, US).

/. mucuna Walp. & Duchass.: Ailigandi, Dwyer 6847 (MO).

1. quaternata Poeppig: Ailigandi, Dwyer 6844 (MO).

— Thomas S. Elias, St. Louis University, St. Louis.

74 ANNALS OF THE MISSOURI BOTANICAL GARDEN

A NEW ANNUAL ERIOGONUM FROM UTAH 1

Soon after the manuscript for the Eriogonum deflexum complex (Brittonia
20: 13-33, 1968) was accepted, a population of dried stems was found which did
not seem to belong to any of the species recognized in that paper. On returning to
this site near Westwater, Grand Co, Utah in September, 1967, the dried stems
were found to represent an undescribed species which may now be known as:
Eriogonum scabrellum Reveal, sp. nov.

A Eriogono deftexo Torrey (subg. Ganysma) differt involucris horizontalibus
sessilibus, perianthiis 1-1.5 mm longis pustulosis basi obtusis, ramis omnibus
scabrellis.

Annual herbs, 1-3 (-5) dm high, with one or rarely two or more stems arising
from thin tan caudices. Stems erect, slender, 5-15 cm long, sparsely tomentose
with tangled white hairs covering the green, scabrellous surface. Leaves subbasal
and sheathing up the stems 2 cm, the leaf-blades 1-3 cm long and wide, cordate,
the apices rounded, the bases cordate, with crispate margins, densely tomentose
below, lightly floccose and green above; petioles as long as or longer than the leaf-
blades, prominently winged. Inflorescences sparsely branched, tending to become
flat-topped, the crowns up to 4 dm long and 15 dm across. Branches dichotomous,
rarely trichotomous, lightly to sparsely floccose with white hairs over the green
scabrellous branches; main branches with 1-4 secondary branches each with a
series of alternating short tertiary branchlets mostly less than 10 cm long at right
angles to it. Bracts scalelike, ternate, 1 mm long, the acute apices widening to the
connate base. Peduncles lacking. Involucres solitary, horizontal, arising from the
bracts at each node, 1.5-2.5 mm long, 1.5-2 mm wide, the 5 acute lobes less than
0.5 mm long, scabrellous. Bractlets linear, 1.5-2.5 mm long, hirtellous. Pedicels
up to 3 mm long, glabrous. Perianth 1-1.5 mm long, white with green midribs
at anthesis, becoming 1.5 mm long and pink to rose or red in fruit, minutely
pustulose over the entire abaxial surface. Tepals dissimilar, the outer whorl of
segments 0.6-0.8 mm wide, obovate, the apices rounded, the bases obtuse, the inner
whorl of segments 0.2-0.5 mm wide, ovate, the apices acute. Stamens excluded, up
to 1.5 mm long, the filaments glabrous except for a few microscopic projections near
the point of attachment, the anthers 0.3 mm long, red, oblong, the pollen grains
pale-yellow, elliptic. Achenes light brown, 2 mm long, the globose base tapering
to a long, sharp, minutely roughened beak. Chromosome number n=20.

Type: 3 mi S of U. S. Hwys 50 & 6 on the dirt road to Westwater, jet 6.8 mi W of
Utah-Colorado state line & ca 17 mi NE of Cisco, along banks of an arroyo on heavy dark
Cretaceous clay soil where it is locally common, Grand Co, Utah, 8 Sept 1967, Reveal &
Davidse 949 (UTC holotype; 34 isotypes will be distributed from UTC).

Additional co >,ys 50 and 6 along the Westwater Rd, 12

Oct 1967, Reveal 958 (BRY, DAOM, NY, UTC).

Eriogonum scabrellum is a member of the sect. Pedunculata Benth. in DC.
of the subg. Ganysma (S. Wats.) Greene and is most closely related to the mem-

1 The field work was supported by the Texas Research Foundation. This paper was
submitted to the Department of Botany, Brigham Young University as partial fulfillment
of three hours of Doctoral Research.

NOTES 75

bers of the E. deflexum complex. The species in this complex have involucres
that are either erect or deflexed. However, E. scabrellum has involucres that are
in a horizontal, or intermediate, position. Other species, such as E. deflexum and
E. hooked S. Wats., which have sessile involucres, have their scalelike bracts ar-
ranged so that the involucres are positioned below the branches. In E. scabrellum
the bracts are arranged so that the involucres are positioned to the side of the
branches and project outwardly although in fruit, the involucres tend to tip down-
wardly slightly. The disposition of the involucres for this new species is unique
to the genus. Unlike any other annual species in the genus, stems and branches
of E. scabrellum are scabrellous, a characteristic known only in E. heermannii
Dur. & Hilg. and its related taxa, a perennial shrub. — James L. Reveal, Department
of Botany, Brigham Young University, Provo, Utah.

A SECOND SPECIES OF COCHLOSPERMUM
(COCHLOSPERMACEAE) FROM PANAMA

Since the publication of the Cochlospermaceae for the Flora of Panama (Ann.
Missouri Bot. Gard. 54: 61-64, 1967) four fruiting collections of Cochlospermum
williamsii Macbr. collected in the Canal Zone, in the Province of Darien, and in
the Comarca de San Bias were received at the Herbarium. This species, originally
described from Amazonian Peru, belongs to the sect. Diporanda and can be readily
distinguished from C. vitifolium (Willd.) Spreng. by the compound-digitate leaves
and by the mature seeds with only a tuft of long, wooly hairs.

Cochlospermum williamsii Macbr., Candollea 5: 388, 1934.— Fig. 1.

Tree 12-23 m tall, the trunk to 15 cm in diam, the wood soft, the branchlets
densely brownish-puberulous, becoming glabrous. Leaves compound-digitate, usu-
ally 5-7 foliolate, the petiole to 25 cm long, longitudinally sulcate, angulate-dilated
at the apex, puberulous when young, glabrescent, the stipules small, deltoid, to
3 mm long, caducous; leaflets inarticulate, subsessile, the petiolule canaliculate
above, the blade narrowly elliptic, sometimes subobovate, attenuate and decurrent
at the base, short-to long-acuminate at the apex, entire-margined, up to 20 cm
long and 6.5-9 cm wide, thin-chartaceous, very brittle when dry, the upper surface
± lustrous and glabrous, the lower surface dull and puberulous especially along
the very prominent costa and prominulous lateral veins, glabrescent. Inflorescences
apparently both terminal and axillary, paniculate, few-flowered, the axes thick,
divaricate, brownish-puberulous. Flowers bright yellow, to 6 cm long, the pedicels
ca 1 cm long, elongated and to 3 cm long in fruit, ± densely brownish-puberulous;
calyx imbricate, the 2 outer sepals smaller than the 3 inner ones, subcoriaceous,
brownish-tomentellous outside, very minutely so inside, caducous; outer sepals
elliptic, obtuse to subacute at the apex, to 1.7 cm long and 1.1 cm wide; inner
sepals elliptic to subobovate, rounded at the apex, thinner towards the margins,
to 2.5 cm long and 1.7 cm wide; petals obovate to broadly obovate, apically deeply
2-lobed, to 6 cm long and 5.8 cm wide, thin-membranous, the lobes rounded, to 2

MISSOURI BOTANICAL GARDEN

r Dressier 3487 (MO).

cm long; filaments oo, unequal, 5-18 mm long, glabrous, the anthers narrowly ob-
long, somewhat arcuate, to 4 mm long, glabrous or sparsely and minutely pilose,
apically 2-porate; ovary subsessile, broadly obovoid, obtusely triquetrous, ca 5 mm
long and 4.5 mm in diam, minutely fulvous-tomentellous; style filiform, to 13 mm
long, densely short-villous at the base, otherwise glabrous, the apex recurved (?).
Capsule obovoid, triquetrous, long-attenuate at the base, emarginate at the apex,
to 7 cm long and 4 cm wide, dark brown, longitudinally striate, and minutely
velvety tomentellous outside, 3-valvate, the exocarp separating from the endocarp
the latter cream-colored to reddish-brown; seeds coiled into a ring, densely lanate,
the hairs tan-colored and 5-10 mm long, the outer seed coat (to which the hairs

NOTES 77

are attached) thin and easily detachable at maturity leaving only a small tuft of
hairs at the hilum, the inner seed coat dark brown to nearly black and ± lustrous.

Reported for the first time in Panama as well as North America; Colombia,
Amazonian Peru, and Amazonian Brazil.

Panama, canal zone: Pifia rd, 21 Apr 1968 (fr), Dressier 3487 (MO), darien: Rio
Uruti (fol, fr), Duke & Bristan 231 (MO) ; woods nr El Real, 4 June 1967 (fol, fr E
for) Duke 11818 (MO), san blas: along the headwaters of the Rio Malatuppu, seasonal
evergreen forest, 17 Aug 1967 (fol, fr) Elias 1757 (MO).

Colombia, amazonas: Trapecio amazonieo, Rio Loretoyacu, alt ca 100 m, Sept-Nov
1944 (fol, fl, fr juv), Schultes 6025 (US), 20-30 Oct 1945 (fol, fr), 6616 (US); Rio Ama-
zonas, nr mouth of Rio Loretoyacu & Puerto Narifio, 13-15 Sept 1966 (fol, fl, fr), Schultes

ma Peru " loreto: P ebas on the Amazon River, forest, July 1929 (fol, fl, fr juv) Williams
1778 (F, US), 1964 (holotype F); Cabollo-Cocha on the Amazon River, forest, Aug 1929
|H A), Williams 2090 (F); Alto Rio Itaya, alt 145 m, Sept-Oct 1929 (fol, fr), Williams

Brazil, amazonas: Basin of Rio Solimoes, Municipality Sao Paulo de Olivenga nr
Palmares, terra firma, high land, Sept-Oct 1936 (fol, fr), Krukoff 8313 (F, MO).

Cochlopsermum williamsii is very closely related to and perhaps conspecific
with C. wentii Pulle (Enum. Vase. PI. Surinam 310, pi. 13, 1906) from Surinam,
from which it seems to differ only by the sepals being minutely tomentellous in-
side, those of C. wentii being glabrous inside. Study of the type of C. wentii is
needed, however, in order to establish the synonomy.— Andre Robyns, Missouri
Botanical Garden, St. Louis, Missouri & National Foundation for Scientific Research
Belgium.

HASSELTIA RIGIDA WOODSON EX A. ROBYNS, A NEW
SPECIES OF FLACOURTIACEAE FROM PANAMA
Hasseltia rigida Woodson ex A. Robyns, sp. nov.; affinis H. monagensi Steyer-
mark (Fieldiana: Bot. 28: 407, 1952), sed arboris magnitudine, foliorum lamina
coriacea marginibusque integerrimis et leviter recurvatis, floribus minoribus et
usque ad 3 mm longis primo visu sat differt.— Fig. 1.

Arbor usque ad 30 m alta, ramulis glabris vel fere glabris. Folia alterna,
petiolo robusto usque ad 2 cm longo supra canaliculate adpresso-puberulo sed
glabrescenti; lamina elliptica ad anguste elliptica vel subcordata, basi cuneata,
apice obtuse acuminata, usque ad 10 cm longa et 4 cm lata, coriacea, marginibus
integerrimis leviterque recurvatis, utrinque sordida, e basi manifeste 3-nervia,
supra glabra basique 2-glandulifera, infra praecipue secus nervos 3 principales
prominentes adpresso-puberula ad glabrescens, nervis secundariis infra prominulis.
Inflorescentiae ut videtur terminales, composito-umbellatae umbellisque iterum
atque interum 3-4-radiatis, axibus adpresso-puberulis ad adpresso-tomentellis,
bracteis parvis deltoideisque. Flores flavido-fusci, hermaphroditi, pedicellis usque
ad 5 mm longis et adpresso-tomentellis; sepala 4, in aestivatione valvata, leviter
inaequalia, ovata, apice acuta, usque ad 3 mm longa et 1.8 mm lata, extus tomen-
tella, intus puberula, persistentia; petala 4, in aestivatione valvata, ovata, apice
acuta, sepalis paululo breviora, extus tomentella, intus puberula, persistentia;
stamina oo, filamentis filiformibus usque ad 2.5 mm longis

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 1. Hasseltia rigida Woodson ex A. Robyns:
base of leaf blade upper surface with 2 glands (X2); CJ
of filament and anther (X30); F, gynoecium (X8); G, b

upper surface (Xj/ 2 ); B,
(X8); D & E, upper part
(X2). After Allen 3733.

antheris minutis subglobosisque; ovarium globosum, glabrum, biloculare, placentis
multiovulatis, stylo simplici usque ad 2.2 mm longo glabroque. Bacca rubra, sub-
globosa, usque 7-8 mm longa, glabra, seminibus 2-4 glabris.

cocle: region N of El Valle de Anton, alt 1000 m, Sept 27, 1946, Allen 3733 (holotype
MO).

—Andre Robyns, Missouri Botanical Garden, St. Louis, Missouri & National Foun-
dation for Scientific Research, Belgium.

MAYNA ZULIANA (PITTIER) A. ROBYNS, COMB. NOV.
(FLACOURTIACEAE)

The revision of the genus Mayna Aublet for the Flora of Panama has turned
up two species for that country, i.e. M. longicuspis (Standley) Standley and another
species, represented by several collections from the provinces of Darien and
Panama, which I was unable to name until I examined the holotype of Carpotroche
zuliana Pittier in the U. S. National Herbarium (Pittier 10513, San Martin on
Rio Palmar, Zulia, Venezuela). As the baccate fruits are bristly (in Carpotroche
the capsular fruits are longitudinally alate) transfer to the genus Mayna is needed:
Mayna zuliana (Pittier) A. Robyns, comb, nov., based on Carpotroche zuliana

Pittier, Bol. Com. Ind. (Venezuela) 4(34) : 38, 1923 (Arboles y Arbustos Nuevos
de Venezuela, secunda y tercera decadas). — Andre Robyns, Missouri Botanical
Garden, St. Louis, Missouri & National Foundation for Scientific Research, Belgium.

A NEW ORMOSIA (LEGUMINOSAE) FROM PERU

Ormosia peruviana Rudd, sp. nov.

Arbor usque ad 18 m alta; ramuli novelli fulvo- vel ferrugineo-subsericei;
stipulae deltoideae, tomentulosae, circiter 1-2 mm longae, basi 1 mm latae, caducae;
folia 7-9-foliolata, axe subtiliter sericeo, glabrata, 6-12 cm longo, petiolo 2-4 cm
longo, jugis inter sese plerumque 2-2.5 cm distanotibus, petiolulis circiter 5 mm
longis, 2 mm diametro, laminis coriaceis vel subcoriaeceis, ovatis, 5-10 cm longis,
3-5 cm latis, apice obtusis, nonnunquam retusis, basi obtusis vel subcordatis, supra
glabris, subtus plus minusve glabris praeter venis maioribus saepe sericeis, venis
secundariis mediocriter elevatis, utrinsecus circiter 10, fere parallels, inter sese
4-10 mm distantibus, angulis venarum costaeque circiter 45°-50°, inflorescentiae
cum axibus fulvido-subsericeis, nee bracteis bracteolisque nee floribus completis
visis, calyce (effracto) fortasse circiter 10 mm longo; fructus dehiscens coriaceus
vel sublignosus, rugulosus, brunneus et fulvo-sericeus, glabrescens, 1-spermus
(fortasse 2- vel 3-spermus), 5-6 cm longus, circiter 3 cm latus, 1-1.2 cm crassus,
valvulis 0.5-1 mm crassis; semina cinnabarina, circiter 17 mm longa, 16 mm lata,
9 mm crassa, hilo elliptic© apicali, 3 mm longo et 1.5 mm lata

cajamarca: open place, mountain slope, alt 2100 m, Colasay, Woytkowski 6964 (holo-
typeMO, isotypeUS).

Vernacular name: huayruru.

This species is readily referable to my Ormosia series Isthmenses (Contr. U. S.
Nat. Herb. 32: 317, 1965). The pods most resemble those of O. colombiana Rudd
but the large seeds are only matched in O. venezolana Rudd.— Velva E. Rudd,
Smithsonian Institution, Washington, D. C.

PUBLISHED BY THE BOARD OF TRUSTEES OF THE MISSOURI BOTANICAL GARDEN
AND WASHINGTON UNIVERSITY PRESS

ANNALS
OF THE

CONTENTS
FLORA OF PANAMA, PART VI

Family 112. Vitaceae

by Thomas S. Elias 81- 92

Family 128. Flacourfiaceae

by Andre Robyns „ 93-144

FLORA OF PANAMA, PART VIII

Family 154. Sapotaceae

by Will H. Blackwell, Jr .145-169

ANNALS VOLUM f 9 S

of the NUMBER 2

MISSOURI BOTANICAL GARDEN

A journal containing scientific contributions from the Missouri
Botanical Garden and the Department of Botany of Washington Uni-
versity in affiliation with the Missouri Botanical Garden. Outside
contributions in systematic botany and allied fields will also be con-
sidered. These papers are subject to a charge of $25 per printed page.

Walter H. Lewis, Editor

Missouri Botanical Garden & Washington University

Sue M. Hulsen, Assistant to the Editor

Missouri Botanical Garden

Derek Burch, Missouri Botanical Garden & Washington University

John D. Dwyer, Missouri Botanical Garden & St. Louis University

Duncan M. Porter, Missouri Botanical Garden & Washington University

Beginning with Volume 53,
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Beginning June 1, 1962 the Stechert-Hafner Service Agency,
10th St., New York 3, N. Y., became sole agent for the Annals of i
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Printed by the Eden Publishing House, St. Louis, Missouri,

ANNALS VOLUME 55

1968

Of the NUMBER 2

MISSOURI BOTANICAL GARDEN

FLORA OF PANAMA

Part VI
Family 112. VITACEAE 1

by Thomas S. Elias
St. Louis University, St. Louis, Missouri
Climbing shrubs, vines or rarely erect shrubs or small trees, unarmed or rarely
armed, stems usually sympodially branched, often with watery juice, bearing
tendrils which may end in discoid suckers; nodes often swollen or jointed, the
stem occasionally winged. Leaves alternate or the lower sometimes opposite, usually
2-ranked, simple, digitately or bipinnately compound, often pellucid-punctate,
usually stipulate. Inflorescences cymose, racemose, spicate or paniculate, usually
well developed and opposite a leaf; axis occasionally flattened and expanded;
peduncles 2 often cirrhose; bracteoles present. Flowers minute, bisexual or unisexual
(the plants then usually monoecious), actinomorphic, numerous; calyx entire or
toothed, the sepals (3-) 4-5 (-7), distinct or basally connate, the petals as many
as the sepals, valvate in bud, minute or obsolete, flat, distinct (connate in Leea) or
as in Vitis apically connate, separating from each other at the base and falling
away as an early deciduous cap; disc evident, annular or lobed; stamens as many
as the petals and opposite them, somewhat perigynous, arising from the base of the
disc, the anthers distinct, rarely connate, introrse, rarely extrorse, 2-thecate, de-
hiscing longitudinally; pistil 1, the ovary superior, 2-(3-8)-loculed and carpelled,
the placentation axile, the ovules 1-2 on each placenta, anatropous, the style 1 and
usually short, the stigma discoid, capitate, or shallowly 2-4-lobed. Fruit a berry,

'Assisted by National Science Foundation Grant No. GB-5674 (Principal Investigator,
Walter H. Lewis).

2 Due to the existing confusion concerning the terrr.
"peduncle" is used in this paper to refer only to the i

Ann. Missouri Bot. Gard. 55 (2): 81-92, '.

[Vol. 55
82 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1-4 (-6) -seeded, the seeds with a small, straight embryo and copious to some-
times ruminate endosperm.

The family contains 1 1 or 12 genera with 600-700 species occurring mainly in
subtropical and tropical areas of the world. The Old World genus Leea has been
placed by some workers in a subfamily of the Vitaceae (Gilg in Engler & Prantl,
Nat. Pflanzenfam. 3(5): 427-456, 1896) or in a family of its own (Suessenguth
in Engler & Prantl, loc. cit. ed. 2., 20d: 372-390, 1953; Cronquist, The Evolution
& Classification of Flowering Plants, p. 261, 1968). Two genera, Cissus and Vitis,
are to be found in Panama.

Useful reference:

Planchon, J. E., Monographie des Ampelidees Vraies. In DC, Monogr. Phaner.
5(2): 305-648, 1887.

a. Inflorescence a well developed panicle; flowers 5-merous; petals fused at the

apex, dropping off as an early deciduous cap; leaves simple; pith brown 1. Vitis

aa. Inflorescence cymose; flowers 4-merous; petals free, expanding, dropping singly;

leaves simple or 3-foliolate; pith white 2. Cissus

1. VITIS
Vitis L., Sp. PL 202, 1753.

Woody vines, deciduous, rarely evergreen, scandent or climbing by tendrils
borne opposite the leaves or arising from the peduncles, the bark often shredding
and falling away, the pith usually interrupted by nodal diaphragms, brown;
tendrils usually branched, rarely simple. Leaves simple or palmately compound,
often lobed, usually rounded and cordate, dentate. Inflorescence a panicle. Flowers
pedicellate, usually umbellate clustered, polygamodioecious, some plants with per-
fect flowers, others staminate with a rudimentary ovary, 5-merous; calyx cupular,
fused with an entire or shallowly lobed margin; petals fused at the apex to form
a deciduous cap; stamens 5, the hypogynous disk of 5 ± free or connate glands
alternate with the stamens and adnate to the base of the ovary, lobed; ovary
bicarpellate, the style short, conical, the stigma usually shallowly bilobed, the
carpels 2-celled. Fruit baccate, usually edible, pulpy, 2-4-seeded, the seeds usually
pyriform and narrowly rostrate basally.

A genus of 60-70 species, mostly of the temperate regions of the northern
hemisphere. Two species are found in Panama, V. vinifera, the common cultivated
grape which has been occasionally planted and the native V. tiliifolia which occurs
throughout Middle America.

1. Vitis tiliifolia Humb. & Bonpl. ex Roem. & Schult. in L., Syst. Veg. ed. 15, 5:

320, 1819.— Fig. 1.
V. caribaea DC, Prodr. 1: 634, 1824.

Vine becoming woody, often climbing, with thick angulate to terete, striate
stems, the pith interrupted by nodal diaphragms, the young branches densely
floccose-tomentose, becoming glabrate, the nodes slightly swollen. Leaves char-
taceous, rounded-ovate to orbicular, frequently with 3 shallow lobes, finely to
coarsely dentate, rarely subentire except at the apex, lateral veins conspicuous

flora of Panama (Family 112. Vitaceae)

Fig. 1. Vitis tiliifolia Humb. & Bonpl. ex Roem. & Schult.: A, habit (Xi/ 2 ); B, mature
I showing apically connate petals (XlO); C, perfect flower (XIO); D, staminate flower
h abortive pistil (XlO); E, seed, ventral surface (X5); F, seed, side view. A-C after
i Wedel 1539 (MO), D after Davidson 567 (MO), E & F after Blum & Tyson 531 (MO).

[Vol. 55
84 ANNALS OF THE MISSOURI BOTANICAL GARDEN

beneath, ending at the apex of the marginal teeth, acute to short acuminate at the
apex, shallowly to deeply cordate at the apex, 5-15 cm long, 4.5-12.5 cm broad,
floccose-tomentose above when young becoming sparsely floccose-tomentose to
glabrate with age, densely floccose-tomentose beneath, the tan to light ferrugineous
tomentum close to lax, usually persistent. Inflorescences of well developed panicles,
the peduncles 4-15 cm long, sparsely to densely floccose-tomentose. Flowers pale
yellow to greenish-yellow, unisexual or bisexual, the calyx undulate, scarcely
toothed, the pedicel 1.5-3 mm long, the corolla 1-1.5 mm long, the lobes oblong
to oblong-obovate, the unisexual flowers with well developed stamens and an
abortive pistil, the perfect flowers with short stamens, the hypogynous glands ±
united, the pistil 1-1.5 mm long, the ovary orbicular, the style short, ending with
a shallowly bilobed stigma. Berries green becoming purplish-black with maturity,
4-6 mm in diam, orbicular, usually 2-seeded; seeds 3-4 mm long, pyriform, slightly
curved, ± stipitate.

Mexico, Central America, the West Indies and northern South America.

bocas del toro: vie of Chiriqui Lagoon, Water Valley, von Wedel 829 (GH, MO),
954 (MO), 1539 (GH, MO, US), canal zone: W slope of Ancon Hill, Woodson et al.
717 (A, MO); s. loc, Blum 555 (MO); Barro Colorado I, Frost 152 (F); Shattuck 736
(F), 7060 (F, US); Woodvj-rth & Vestal 624 (F); Cocolf, Riley 135 (MO), chiriqui:
Boquete, Davidson 567 (F, MO); forest nr Boquete, Pittier 3141 (GH, US), cocle: hills
N of El Valle de Anton, vie of La Mesa, Allen 2486 (GH, MO, US), colon: vie of San
Juan nr Cement Plant Lake, Blum & Tyson 531 (MO), darien: road from El Real to
Pinogana, Duke 5131 (GH, MO); Rio Balsa, betw Manene & Tusijuanda, Duke 13548
of Campamento Buena Vista, nr Quebrada Felix, Stern et al. 952 (GH, MO);
EI Punteadero at bridge crossing over Rio Chucunaque, Stern et al. 162 (GH, MO); Rfo
Uroganti, Bristian 1152 (MO), panama: nr Rfo Tapia, Maxon & Harvey 6619 (GH);
San Jose I, Johnston 591 (GH, MO, US), san blas: Mulatuppu, Duke 8550 (MO).

The original spelling was tiliaefolia, a name widely adopted. Other spellings
like tilifolia and tiliifolia have also been used. According to note 2 of Article 73
and recommendation 73G of the Code (Regnum Veget. 46: 73, 1966) the use of
a wrong connecting vowel or vowels in a name or epithet is considered an ortho-
graphic error. Recommendation 73G states that before a consonant the final vowel
in Latin is reduced to i. Thus in place of tiliaefolia the corrected epithet should
read tiliifolia.

Commonly referred to as "uva" in Panama, the grapes of this species are re-
ported by Duke (personal communication) to be more often used for making wine
and vinegar than for eating. In addition the larger vines contain quantities of
potable water. The tough stems are occasionally used for cordage by the indians.

2. CISSUS
Cissus L., Sp. PL 117,1753.

Climbing vines, herbs or small shrubs, often woody, occasionally fleshy and
succulent; roots often tuberous; stems terete or angulate, glabrous to densely
pubescent, often ramifying, tendrils always present and opposite the leaves, simple
or bifurcate; stipules paired, always present, generally caducous. Leaves usually
simple or 3-5-foliolate, often lobed, petiolate, the shape, pubescence and texture
highly variable, the venation pinnate or palmate, the margins entire or frequently

floba of panama (Family 112. Vitaceae) 85

toothed. Inflorescences always opposite the leaves, cymes regular or corymbiform
or umbelliform, pedunculate, with bracts and bracteoles. Flowers 4-merous, perfect,
pedicellate, the calyx cupular, entire or with 4 valvate lobes; corolla generally
ovoid or campanulate at the base, the petals valvate, inwardly cucullate, opening
at anthesis and falling away separately; stamens opposite the petals, inserted on
the receptacle at the base of the disc, the filaments erect, the anthers introrse, the
disc surrounding the ovary, the margin 4-lobed; ovary bilocular with 2 anatropous
ovules, the style cylindrical and often persisting after anthesis, the stigma small,
discoid. Fruit a 1 (-2) -seeded berry, the seeds obovoid, attenuate at the base, often
trigonous.

A pantropical genus of ca 400 species of which six are reported from Panama.
This genus exhibits great variation. In C. sicyoides, for example, the amount of
variation expressed within the species is greater than the differences between the
trifoliolate species. The neotropical members of this genus are in need of serious
monographic study. The name Cissus was derived from the Greek kissos meaning

a. Leaves 3-folioIate.

b. Terminal leaflets rhombic; plants densely pubescent with glandular &/or
eglandular hairs; stipules on the younger branches linear-lanceolate to
lanceolate, 3-6 mm long 1. C. rhombifolia

bb. Terminal leaflets elliptic to obovate; plants usually glabrous or with sparse
eglandular hairs; stipules on the younger branches broadly ovate to sub-
rotund, 2-4 mm long.

items 4-gonous, often winged;

cc. Mature peduncles 1-1.8 cm long; stems terete to su
fruits 6-10 mm in diam.

d. Terminal leaflets with a petiole 0.5-2 cm Ion
serrate, mature leaves 4-10 long; flowers rec

fruits 6-8 mm in diam 3. C. microcarpa

dd. Terminal leaflets sessile; leaves sparsely, denticulate, mature leaves

2-4 cm long; flowers pale green; fruits ca 1 cm in diam 4. C. martiniana

aa. Leaves simple.

e. Berries 10-12 mm in diam; buds 3-4 mm long, dark red; leaves strongly

dimorphic, mature leaves coriaceous, glabrous 5. C. biformifolia

ee. Berries 4-6 mm in diam buds 1-1.5 mm long, greenish-white, white to
yellow, rarely red; leaves not strongly dimorphic, mature leaves chartaceous,
glabrous to densely villous 6. C. sicyoides

1. Cissus rhombifolia Vahl, Eclog. Amer. 1: 11, 1796.— Fig. 2A, B.

Vine becoming woody, often climbing, the stems subangulate to terete, striate,
frequently canaliculate, unwinged, slightly swollen at the nodes, densely pubescent
with glandular and/or eglandular hairs, the eglandular hairs to 2 mm long,
septate and transparent except for the darkened junctions of the cells, the glandular
hairs to 0.5 mm long, usually more abundant than the eglandular hairs; stipules on
the younger branches linear-lanceolate to lanceolate, 3-6 mm long, becoming ovate
to broadly ovate on the older stems, 6-12 mm long, subpersistent, usually densely
pubescent. Leaves 3-foliolate, densely villous, especially beneath, the petiole 1.5-8
cm long, striate, densely pubescent, the leaflets chartaceous to subcoriaceous, ser-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 2. A, Cissus rhombifolia Vahl, leaf & inflorescence (xy 2 ); B, C. rhombifolia
Vahl, flower (X10); C, C. martiniana Woodson & Seibert, leaf & inflorescence (Xi/ 2 );
D, C. erosa L. Rich., leaf & inflorescence (X|/ 2 ); E, C. microcarpa Vahl, leaf & inflorescence
(X|/ 2 ). A-B after Lewis et a/. 7544 (MO), C after Seibert 241 (MO holotype); D after
Stern et al. 305 (MO).

flora of Panama (Family 112. Vitaceae) 87

rate, the lateral veins 5-7 pairs, ascending and terminating at the apex of the ser-
rations; tendrils borne opposite the leaves; terminal leaflet 4.8-13 cm long, 2.2-6.5
cm broad, rhombic, the young leaves commonly narrowly ovate, cuneate at the
base, sessile or petiolulate to 5 mm long, acute to short acuminate at the apex;
lateral leaflets 3.6-10 cm long, 1.8-7 cm broad, rhombic to broadly ovate, strongly
oblique at the base, acute to short acuminate at the apex. Inflorescences cymose,
often appearing corymbose with the flowers congested in pseudoumbels, the
peduncles 1-3 cm long, often bright red with exposure to the sun, pubescence of
densely spreading hairs. Flowers pale yellow to greenish-yellow, the pedicel 3-8
mm long, the calyx cupular with four scarsely discernible teeth, densely pubescent,
the corolla of four free petals, oblong to oblong-ovate, acute at the apex, ca 1 mm
long; pistil 1.5-2 mm long, the ovary 2-locular. Berries green becoming black with
maturity, 6-8 mm in diam, orbicular, 1-seeded; seeds broadly pyriform, 5-7 mm
long, 4-5 mm broad.

Mexico, Central America, the West Indies and South America.

canal zone: Barro Colorado I, Shattuck 252 (A), Wetmore & Woodworth 873 (A),
Wetmore & Abbe 139 (A. MO), Bailey & Bailey 337 (GH); vie of Miraflores Lake, White
242 (MO); K-9 road at bridge, Stern et al. 975 (GH, MO); Fort Clayton; Cardenas Creek
area, Tyson 1293 (MO); Farfan beach road, Kirkhride & Ellas 58 (MO); U. S. Army
Tropic Test Center, Albrook, Dwyer 6717 (MO); Fort Kobbe, Duke 3944 (GH, MO);
Curundu, nr Survival School, Tyson 1068 (MO), chiriqui: Cerro Chorcha, Allen 5062
(MO); trail from San Felix to Cerro Flor, Allen 1936 (GH, MO); 12.4 mi N of David,
Lewis et al. 701 (GH, MO, UC, US), cocii: betw Las Margaritas & El Valle de Anton,
Woodson et al. 1762 (A, MO); betw Aguadulce & El Valle de Anton, Woodson et al.
1210 (A, MO); El Valle de Anton, Lewis et al. 2597 (DUKE, K, MO, NY, UC). darien:
vie of El Real, Rio Tuira, Stern et al. 788 (GH, MO, US) ; betw El Real & Rio Canalones,
Duke 4982 (GH, MO); Boca Grande I, Duke 8841 (MO), herrera: Pese, ca 50 mi, Allen
793 (GH, MO, US), los santos: Rio Tonosi, vie of Tonosi, Lewis et al. 1544 •
mi W of Candelaria, Duke 12451 (MO). Panama: vie of El Llano, Duke 5521 (GH, MO);
Gorgona beach, Woodson et al. 1691 (A, MO); thickets & forest nr Arraij'an, Woodson
et al. 1354 (A, MO); betw Pacora & Chepo, Woodson et al. 1653 (A, MO); San Jose I,
Johnston 973 (GH); Rio Tartare, Woodson & Schery 1005 (MO); Piria, Duke 14447
(MO), veraguas: vie of Sona, along highway, Woodson et al. 512 (MO); ca 5 mi NE
of La Mesa, Blum & Tyson 659 (MO), 665 (MO); 12 mi from Santiago toward Divisa,
Dwyer & Kirkhride 7436 (MO); 2-4 mi E of Santiago, roadside savanna, Duke 12359 (MO).
This species is easily, recognized by the rhombic-shaped leaves and dense
pubescence. The glandular hairs characterizing most of the Panamanian material
were not observed on collections from southern Mexico or Guatemala. Although
this condition may warrent subspecific status, I am hesitant about creating another
taxon in this genus until this species has been adequately studied through its range.

2. Cissus erosa L. Rich., Act. Soc. Hist. Nat. Paris 1: 106, 1792.— Fig. 2D.
C. salutaris Kunth in H.B.K, Nov. Gen. Sp. PI. 5: 225, 1821.
Vitis salutaris Baker in Mart, FI. Bras. 14 (2): 211, pi. 52, 1871.

Vine becoming woody, often climbing, the stems angulate, tetragonal, fre-
quently winged on the margins, slightly nodose, glabrous or with a few scattered,
appressed, pilose hairs; stipules broadly ovate to subrotund, subpersistent, 3-4 mm
long, the apex obtuse, glabrous. Leaves 3-foliolate, the petiole 1.8-6.5 cm long,
4-gonal especially at the base, glabrescent, occasionally winged, the wing up to

88 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1.5 mm broad; leaflets chartaceous to subcoriaceous, sharply serrate to sparsely
dentate, occasionally shallowly so, glabrous to sparsely pilose beneath, especially
on the costa, the lateral veins 4-6 pairs, ascending and terminating between the
teeth; terminal leaflet 4.5-15.5 cm long, 1.5-8 cm broad, elliptic to elliptic-ovate
to ovate, cuneate at the base, acute to subobtuse at the apex, sessile to short-
petiolate, rarely to 1.5 cm long; lateral leaflets 3.5-12 cm long, 2-6 cm broad,
inequilateral, ovate to ovate- elliptic, oblique and rounded to subcuneate at the
base, acute to obtuse at the apex. Inflorescences cymose, appearing corymbose with
the flowers congested in pseudoumbels, the mature peduncles (6-) 7- 12 cm long,
glabrous or with a few scattered, appressed, pilose hairs. Flowers bright red to
reddish-orange, the pedicel 2-5 mm long, pilose often sparsely so, the calyx with
four short lobes, often indiscernible, the corolla of four free petals, oblong, acute
at the apex, ca 1 mm long; pistil 1-1.5 mm long. Berries green becoming black
with maturity, 4-6 mm in diam, orbicular to subpyriform, 1-seeded; seeds pyriform,
4-5 mm long.

Southern Mexico, Central America, the West Indies and South America.

canal zone: Ancon Hill, Duke 4587 (MO); Barro Colorado I, Brown 33 (F), Wood-
worth y Vestal 530 (A, F); Gamboa, Tyson et al. 4588 (MO); vie of Miraflores, White
135 (GH, MO): I K-9 road, Stern et al. 36 (GH, MO), 305 (GH, MO);

Frijoles, nr boat dock, Ebinger 70 (MO); Howard Air Force Base, SE of Kobbe Beach,
Oliver & MacBryde 1890 (MO), cocle: vie of El Valle de Anton, Allen 750 (GH, MO,
US); betw Las Margaritas & El Valle de Anton, Woodson et al. 1763 (GH, MO), colon:
vie of Rio Piedras, along road to Puerto Bello, Blum et al. 2536 (MO); vie of Sardinilla
ca 7-8 mi E of Cement Plant, Blum & Tyson 504 (MO); betw Rio Piedras & Puerto
Pilon, Lewis et al. 3215 (DUKE, MO, UC). herrera: 12.5 mi S of Ocu, Lewis et al. 1624
(GH, MO, US), panama: Cerro Azul, Tyson 2151 (MO); Cerro Campana, trail from
Campana to Chica, Allen 2649 (MO); San Jose I, Erlanson 68 (GH), Johnston 429 (GH),
885 (GH), 973 (MO), 976 (GH, MO), san blas: along the headwaters of the Rio Mula-
tupo, Elias 1727 (MO), veraguas: hills W of Sona, Allen 1035 (GH, MO, US).

Cissus erosa occurs frequently throughout Middle and South America and
shows considerable variation in leaf margins and the occasional presence of indu-
mentum on the lower leaf surfaces. Despite Planchon's (in D.C., Monogr. Phan.
5(2) : 548-9, 1887) reduction of C. salutaris to a synonym of C. erosa, some workers
(Publ. Field Mus. Nat. Hist., Bot. Ser. 24: 302, 1949) have maintained the two
as distinct based mainly on the degree of pubescence of the leaves. As specimens
assigned to the two species show an intergradation of characters, the reduction by
Planchon is apparently justifiable.

3. Cissus microcarpa Vahl, Eclog. Amer. 1: 16, 1796.— Fig. 2E.

Vine, usually woody, the stems terete, sulcate, unwinged, lenticels sparse and
inconspicuous, glabrous except for occasional appressed pilose hairs on the younger
stems, the nodes swollen; stipules broadly ovate to subrotund, subpersistent, 3-4
mm long, the apex acute to obtuse, glabrescent. Leaves 3-foliolate, the petiole 1.5-6
cm long, striate, glabrous or occasionally appressed-pilose on the juvenile forms,
the lateral veins conspicuous beneath, chartaceous to subcoriaceous, mucronate-
serrate; terminal leaflet 3-10 cm long, 1.4-6 cm broad, broadly elliptic, elliptic,
ovate-elliptic, cuneate at the base, petiolule 0.5-2 cm long, acuminate at the apex,
mucronate, the lateral veins 5-6 pairs; lateral leaflets 3-8 cm long, 0.6-5 cm broad,

flora of Panama (Family 112. Vitaceae) 89

obliquely elliptic or ovate to rhombic, obliquely rounded at the base, acute to
shortly acuminate at the apex, mucronate, the lateral veins 3-5 pairs. Inflorescences
cymose-umbellate and appearing corymbiform, the peduncles 0.6-1.8 cm long,
glabrous or with a few scattered hairs. Flowers appearing red or reddish-orange,
the pedicel 1-4 mm long, glabrous to sparsely pilose; calyx cupular, the lobes
indiscernible; corolla ca 1 mm long, the lobes ovate to ovate-triangular, acute at
the apex, pistil 1-1.5 mm long. Berries green, turbinate to spherical (?), ca 6-8 mm
in diam, usually 1-seeded, the seeds pyriform, 4-6 mm long.

Mexico, the West Indies, Central and South America.

bocas del toro: vie of Chiriqui Lagoon, von Wedel 1040 (GH, MO, US), 1334 (GH,
MO), 1517 (GH, MO); Almirante, on rd to Bomba, Blum 1335 (MO); Changuinola
Valley, Potrero I, Dunlap 77 (F); Rio Cricamola, betw Finca St. Louis & Konkintoe, Wood-
son et al. 1916 (MO), canal zone: Barro Colorado I, Ehinger 277 (MO); vie of Sala-
manca Hydrographic Station, Rio Pequeri, Woodson et al. 1626 (MO); Coco Solo, U. S.
Army Tropic Test Center, Mine Emplacement Center, Dwyer & Duke 7920 (GH, MO, US),
hills of Cerro Pilon, nr El Valle de Anton, Duke & Correct 14714 (MO); betw
Cerro Pilon & El Valle de Anton, Duke & Dwyer 13951 (MO). Panama: along Pan-Am
Highway, nr Jenine, Duke 3820 (MO); gallery along Rio Terable, nr Pan-Am highway
& El Llano, Duke 5657 (MO); tributary of Rio Chagres, 5 mi SW of Cerro Brewster,
Lewis et al. 3372 (DUKE, K, MO, NY, UC), 3458 (DUKE, K, MO, UC); Cerro Jefe,
10-13 mi beyond Goofy Lake, Duke 8027 (MO).

In Panama this species has been consistently confused with C. rhombifolia.
Cissus microcarpa can be readily identified by long petiolate leaves which are
mucronate-serrate and the generally glabrous condition. Like several other species
of Cissus the upright inflorescences are often showy due to the bright red peduncles.

4. Cissus martiniana Woodson & Seibert, Ann. Missouri Bot. Gard. 24; 191,
1937.-Fig. 2C.

Woody vine or small prostrate shrub, the stems subangulate to terete, un-
winged, conspicuously lenticellate on the older stems, glabrous except for occasional
appressed pilose hairs on the younger stems, the nodes slightly swollen; stipules
broadly ovate to subrotund, persistent, 2-3 mm long, the apex obtuse to subacute,
glabrescent. Leaves 3-foliolate, the petiole 1.5-2.5 cm long, striate, glabrous or
rarely with a scattered, ferrugineous-pilose indumentum, sessile, obtuse at the apex,
minutely mucronate, the lateral veins 4-5 pairs, inconspicuous, subcoriaceous,
sparsely denticulate; terminal leaflet 2-4 cm long, 1-2.2 cm broad, obovate to
elliptic, cuneate at the base; lateral leaflets 1.5-4 cm long, 0.8-2 cm broad, obliquely
obovate to obovate-elliptic, unequally cuneate at the base. Inflorescences cymose,
appearing corymbiform, the peduncles 1-3 cm long with a few scattered tomentose
hairs. Flowers pale green, the pedicel 2-4 mm long, glabrous; calyx with 4 subreni-
form to broadly ovate lobes, the lobes rounded at the apex; corolla 1-1.5 mm long,
the lobes oblong to ovate-triangular, acute at the apex; pistil 1-2 mm long. Berries
green, spherical, ca 1 cm in diam, usually 1-seeded, the seeds pyriform, 6-8 mm
long.

Guatemala southward to Panama and (?) northern South America.

chiriqui: valley of the upper Rio Chiriqui Viejo, vie of Monte Lirio, Seibert 241
(MO holotype, GH isotype); vie of Bajo Chorro, Woodson & Schery 676 (MO), Davidson
248 (A, F, MO, US); valley of the upper Rio Chiriqui Viejo, White 72 (GH, MO, US).

90 ANNALS OF THE MISSOURI BOTANICAL GARDEN

This species is closely related to C. erosa but can be distinguished by its much
smaller leaves, flowers and berries.

5. Cissus biformifolia Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 4: 225,

1929.
C. cardiophylla Standley, loc. cit. 226.

Vine, often climbing and hanging; stems terete to 4-gonal, sulcate to canalicu-
late, glabrous except for the appressed pilose hairs on the young stems, swollen at
the nodes, with sparse minutely elevated lenticels. Leaves dimorphic, the younger
ones narrowly elliptic to narrowly obovate, 4-10 cm long, 1.5-4 cm broad, long
cuneate at the base, short acuminate at the apex, glabrous, chartaceous, the petiole
1.5-4.5 cm long, the older leaves ovate to broadly ovate, 10-18 cm long, 5-13 cm
broad, rounded, cordate or deeply cordate at the base, short acuminate to acute at
the apex, glabrous, coriaceous, the petiole 3.5-7 cm long, often sulcate and canalicu-
late; all leaves with 5-7 pairs of ascending lateral veins and entire at the base
becoming sparsely and shallowly serrate toward the apex; stipules broadly obovate,
3-4 mm long, early caducous, subrotund at the apex. Inflorescences of small
compacted cymes appearing as pseudoumbels, the peduncles up to 2 cm long,
younger ones appressed pilose; buds 3-4 mm long. Flowers dark red, the pedicel
1-2 mm long, with scattered appressed pilose hairs; bracts and bracteoles 2-3 mm
long, ovate, acute at the apex; calyx cupular with four rotund teeth which persist
in fruiting; corolla of four free petals, the petals ovate-oblong, 3-4 mm long, obtuse
at the apex; pistil 2-3 mm long, 2-locular. Berries green becoming purple at
maturity, 10-12 mm long, 8-10 mm broad, suborbicular to obovoid, apex rounded
and usually with a small persistent beak, 1-seeded, the seed 8-10 mm in diam,
obovoid.

Reportedly from British Honduras and ranging southward to Panama. The
dimorphic leaves, larger flowers and fruits and the glabrous condition distinguish
this species from the common C. sicyoides.

bocas del toro: Changuinola valley, Dunlap 323 (US holotype, F isotype); 10-15
mi inland from mouth of Changuinola River, Lewis et al. 861 (GH, K, MO, UC, US);
vie of Chiriqui Lagoon, von Wedel 2983 (GH, MO); Duwebdulup Peak, N of Rfo Teribe
across from Quebrada Huron, Kirkbride & Duke 576 (MO, NY); Santa Catalina, river
bank & beach. Blackwell et al. 2692 (MO).

6. Cissus sicyoides L., Syst. Nat. ed. 10, 2: 897, 1759.
C. obtusata Bentham, Bot. Voy. Sulphur, 77, 1844.

Vitis sicyoides (L.) Morales in Poey 5 Repert. Fis-Nat. Isla Cuba, 1 : 206, 1866.

Cissus brevipes Morton & Standley, Publ. Field Mus. Nat. Hist, Bot. Ser. 18: 653, 1937.

Vine, occasionally shrubby, climbing or sprawling; stems terete, striate, ±
canaliculate, glabrous, tomentose to villous, lenticels usually sparse, inconspicuous,
± swollen at the nodes, the older stems often woody with the epidermis becoming
loose to form a sheath around the stem and peeling off. Leaves variable, broadly
ovate to ovate-elliptic to oblong to subrotund, 2.5-15 cm long, 1.7-12.5 cm broad,
truncate to rounded to cordate at the base, acute to short acuminate at the apex,
the margins shallowly serrate, the teeth short tipped, the lateral veins 4-6 pairs,
ascending, the costa and veins ± conspicuous beneath, chartaceous; petiole

flora of Panama (Family 112. Vitaceae) 91

l-6.5(-8.5) cm long, terete, ± swollen at the base, glabrous to densely villous;
stipules narrowly ovate to ovate-oblong, acute to obtuse at the apex, 2-4 mm long,
caducous. Inflorescences of small congested cymes, often appearing as pseudo-
umbels, the peduncles 1-5 cm long, terete to angulate, ± sulcate, densely villous
to glabrous; buds ca 1 mm long, ovoid. Flowers greenish-white, white or yellow,
becoming red when growing in bright, exposed areas, the pedicel 2-3 mm long,
densely villous to glabrous, the calyx cupular with 4 shallow rotund lobes, the
lobes often persisting in fruit, the corolla of 4 free, oblong petals, 1.8-3 mm long,
pistil 2-3 cm long, bilocular; the 4-lobed disc conspicuous, persisting in fruit.
Berries 4-6 mm in diam, obovoid, 1-seeded, often with the style persisting as a
beak, green turning dark red to black at maturity; seeds 3.5-5 mm in diam,
obovoid.

Southern United States, the West Indies, Central and northern South America.
bocas del toro: vie of Chiriqui Lagoon, von Wedel 1367 (GH, MO); id., Old Bank
I, von Wedel 1891 (MO), 7958 (MO), 2070 (MO); id., Water Valley, von Wedel 838
(GH, MO), 1811 (MO); id., Isla Colon, von Wedel 2470 (GH, MO), 2952 (GH, MO);
region of Almirante, Cooper 183 (F), Blum 1334 (MO); Chiriquicito to 5 mi S along
Rio Guarumo, Lewis et al. 2015 (GH, MO); vie of Nievecito, Woodson et al. 1854 (MO);
10-15 mi inland, S from mouth of Changuinola River, Lewis et al. 981 (GH, MO, US);
s. loc., von Wedel 309 (MO); Santa Catalina, Blackwell et al. 2751 (COL, MO, UC).
canal zone: Ancon Hill, Duke 4588 (GH, MO); Barro Colorado I, Shattuck 268 (MO),
588 (F), 589 (F), 810 (F), Woodivorth & Vestal 38 (F), 154 (F, GH), 553 (F), Wilson
3 (F); Fort Amador on causeway & I, Tyson 2009 (MO), 2010 (MO), 2020 (MO), Chagres,
Fendler 52 (MO); 5 mi N of Cocoli Tyson 3874 (MO); Farfan beach, from Thatcher
Hwy to Palso Seco, Lewis et al. 54 (GH, K, MO, UC, US); Farfan Beach road, Kirk-
bride & Elias 59 (MO); 5 mi N of Gamboa on pipeline rd, Blum & Loftin 2305 (MO);
in government forest along Las Cruces Trail, Hunter & Allen 693 (MO), 700 (MO);
vie of Rio Chilibre bridge, Blum & Dwyer 2133 (MO); Fort San Lorenzo, Tyson & Blum
3674 (MO); Trans Isthmian Highway ca 19 mi from Colon, Burch et al. 1001 (DUKE,
F, GH, K, MO, NY. UC, US), chiriqui: vie of San Bartolome, Peninsula de Burica,
Woodson & Schery 913 (MO); vie of Puerto Armuelles, Woodson & Schery 832 (MO);
Boquete, Davidson 675 (F, MO), cocle: N rim of El Valle de Anton, nr Cerro Turega,
Woodson & Schery 182 (MO), colon: vie of San Juan at Cement Plant Lake, Blum &
Tyson 526 (MO), darien: Cana, Rio San Jose below former goldmine headquarters, Stern
et al 635 (GH, MO); forest nr Sante Fe, Duke 8431 (MO); 3 mi E of Santa Fe, Tyson
et al. 4659 (MO); 0-4 mi up Rio Sabana from Santa Fe, Duke 4138 (GH, MO); Rio
Chico, vie of Yavisa, Allen 5093 (MO); Rio Lara, Duke 8853 (MO); vie of El Real, along
trail to Rio Pirre, Stem et al. 291 (GH, MO), 295 (GH, MO); Tucuti, Terry & Terry
1400 (F. GH, MO); Rio Balsa, betw Manene & Tusijuandra, Duke 13575 (MO); Patiho,
Duke 10507 (MO); Rio Pinas, Duke 10559 (MO), herrera: rd from La Avena to out-
skirts of Pese, Burch et al. 1310 (GH, K, MO, UC, US); Pese, ca 50 mi, Allen 792 (F,
GH, MO), los santos: 1-2 mi W of Candelaria, Duke 12448 (MO); 5-9 mi from Chitre
on rd to Las Tablas, Burch et al. 1221 (GH, MO); Monagre beach rd, 0.3-1. „ , : . ■
beach, ca 5 mi SE of Los Santos, Lewis et al 1681 (COL, MO, UC); Tone-
bank, Tyson et al 2996 (MO). Panama: Rio Canasas, gravel bar, Duke 14514 (MO);
1 mi N of Rio Chagres along Roosevelt Boyd Hwy, Blum & Tyson 1985 (MO), 1986
(MO); Chiman, Lewis et al. 3324 (MO), 3336 (MO); ca 6 mi E of Chepo on Pan-Am
Highway, Duke 4031 (MO), 4041 (MO), 4062 (GH, MO); Gorgona Beach, Woodson et
al 1697 (GH, MO); Las Sabanas, E of Panama City, Maurice 811 (MO); Las Sabanas,
nr Chepo, Hunter & Allen 96 (MO); savanna nr Playa Rio Mar, Duke 11763 (MO);
vie of Pacora, Allen 1007 (MO); Pedro Gonzales I, Dwyer 680A (MO), Allen 2584 (MO);
San Jose I, Johnston 875 (GH, MO), Duke 12518 (MO); Taboguilla I, Duke 5876 (MO);
Tocumen, Dwyer 1848 (MO), san blas: Soskatupu, Elias 1687 (MO).

Cissus sicyoides, a highly polymorphic species, is one of the more common

92 ANNALS OF THE MISSOURI BOTANICAL GARDEN

vines in the neotropics. Readily adapting to disturbed areas, it is one of the few
plants to invade successfully the very rocky beaches along lakes and the seacoasts.
In this habitat, the leaves are usually small, subrotund and are glabrous. An
alternate form is encountered in more protected, less exposed habitats where the
leaves are usually ovate, truncate at the base and usually quite pubescent. I hesitate
to attempt assigning varietal names since Planchon (loc. cit. 521-531) listed 16
varieties in 1887 and since then no adequate work exists on the neotropical species
of Cissus. Although some Panamanian collections of C. sicyoides can be assigned
to a variety, e.g. C. sicyoides var. ovata Planchon, the majority of the material
seems to defy placement.

The tough stems are often used for cordage. In addition, the leaves when
macerated in water yield a soapy solution which may be utilized for washing
clothes. Practically every specimen I examined from Panama showed varying de-
grees of infection by the smut, Mycosyrinx cissi (DC.) Beck. The intemodes of
severely infected plants are reduced, the leaves fail to develop properly and the
plant has the appearance of "witches broom." The diseased plant is so strange that
Presley (Rel. Haenk. 2: 35, t. 53, 1834) described a new genus (Spondylantha)
of flowering plants from such a specimen.

> incidentally mentioned.

Cissus 82t, 84, 85t, 89t, 92t Leea 82f

biformifolia 90 Mycosyrinx cissi 92

brevipes 90 Spondylantha 92f

cardiophylla 90 Vitaceae 81, 82t

erosa 87, 88t, 90t Vitus 82
martiniana 89 caribaea 82

rhombifolia 85, 89t
salutaris 87, 88t

85t, 90, 91t. 92t

FLORA OF PANAMA

by Robert E. Woodson, Jr. and Robert W. Schery
and Collaborators

Family 128. FLACOURTIACEAE 1

by Andre Robyns 2

Missouri Botanical Garden and Department of Botany, Washington University,

St. Louis, Missouri

Trees or shrubs, the branchlets sometimes spine-tipped or axillary spinose.
Leaves usually alternate and/or distichous, rarely opposite or verticillate, persistent,
the petiole short to long, sometimes 2-glandular at the apex, the stipules usually
small and caducous, sometimes large, foliaceous and persisent, rarely absent; blade
simple, entire or not, sometimes 2-glandular at or near the base, penninerved or
3-5-nerved from the base, sometimes with pellucid dots or lines. Inflorescences
axillary or terminal, fasciculate, racemose, spicate, corymbose or paniculate, some-
times flowers axillary and solitary, the peduncles or pedicels sometimes adnate to the
petioles of the subtending leaves, the bracts and bracteoles minute. Flowers actino-
morphic, 5 or $ 9, sometimes dioecious or polygamous; sepals 2-several, contorted
or imbricate, rarely valvate, sometimes undifferentiated from the petals, usually
distinct, sometimes more or less united into a tube, sometimes persistent or even
accrescent; petals, when present, hypogynous or ± perigynous, rarely epigynous,
equal in number to the sepals and alternating with them, or sometimes more
numerous than the sepals, contorted or imbricate, sometimes with a scale within
the base; torus often glandular or sometimes expanded into a glandular disk (be-
tween androecium and gynoecium); stamens usually more numerous than the
petals, often oo, 1- or many-seriate, or isomerous with the petals and opposite to
them; filaments free or in fascicles alternating with glands, rarely united into a
tube; anthers 2-thecate, sometimes appendaged, usually longitudinally dehiscent,
rarely opening by terminal pores; pollen grains usually 3-colporate; ovary superior,
semi-inferior to rarely inferior, 1-locular with several (2-10) parietal placentas, the
later sometimes ± deeply protruding into the middle of the ovary, infrequently
3-5-locular; ovules usually oo on each placenta, anatropous or amphitropous; styles
isomerous with the placentas, free to completely united, rarely absent. Fruit a
valvately dehicent capsule, or fleshy or dry and indehiscent, the pericarp some-
times alate or prickly; seeds few to numerous, sometimes conspicuously arillate,
the endosperm usually copious and fleshy, the embryo straight or curved; cotyledons
usually broad, often cordate. — x = 10, 11, 12.

A family of about 85 genera and 1300 species, nearly all woody, chiefly of
tropical distribution, with some extensions into the temperate zone; 15 genera, at

1 Assisted by National Science Foundation Grant No. GB-5674 (Princioal Investigator,
Walter H. Lewis).

2 Visiting Curator of the Flora of Panama, Missouri Botanical Garden; Charge de
Recherches of the National Foundation for Scientific Research, Belgium.

Ann. Missouri Bot. Gard. 55(2): 93-144, 1968.

[Vol. 55
94 ANNALS OF THE MISSOURI BOTANICAL GARDEN

present, reported from Panama, several additional genera occurring in Central
America, namely: Bartholomaea Standley & Steyermark, Macrohasseltia L. Wms.
(Honduras, Costa Rica), Olmediella Baillon, and Pleuranthodendron L. Wms.
(from Mexico, through Central America, to northern South America).

Economically the family is of little importance; Oncoba spinosa Forsk. from
Africa, cultivated at the Plant Introduction Garden at Summit, Canal Zone
(Steyermark s.n., MO, in 1935) has edible fruits.

The Flacourtiaceae are characterized by a combination of several characters:
numerous stamens, receptacle often glandular or often expanded into a glandular
disc, ovary superior and 1-locular with parietal placentation, copious endosperm,
and often undifferentiated perianth. In some genera the placentas are deeply pro-
truding into the middle of the ovary and the ovary is sometimes even plurilocular,
e.g. in Prockia with a 3-5-locular ovary and Hasseltia with a 2(-3)-locular ovary;
these genera are perhaps better placed in the Tiliaceae (see Hutchinson, The Genera
of Flowering Plants 2: 476, 478, 1967). The family is somewhat indeterminate
and, as noted by Sleumer (in Van Steenis, Fl. Males., ser. I, 5: 2, 1954), "no single
character exists wherewith to distinguish Flacourtiaceae from other families or to
recognize them in the field."

The following key to the genera is based only on Panamanian collections.

a. Petals present.

b. Petals more numerous than the sepals,
c. Fruits echinate.

d. Styles simple; stigma shortly 3-lobulate; flowers 5 or $ by abortion

1. Lindackeria

dd. Styles 3, each one 2-lobed, the lobes laciniate; flowers dioecious

2. Mayna

cc. Fruits with broad, vertical wings 3. Carpotroche

11, in large, compound, repeatedly 3- to 4-radiate

lers small, subglobose; ovary 2(-3) -celled 4. Hasseltia

large, in few-flowered racemes; anthers elongated-linear;
incompletely 5-9-locular by the intrusion of the placentas
5. Neosprucea

g. Sepals 3 (-4); stamens pluriseriate, inserted on a densely villosulous
disc; ovary superior, with 5-8 intruding, lamelliform placentas

-6. Banara

gg. Sepals (5-) 6-7; stamens in fascicles of 3, rarely 2, the fascicles
alternating with minutely tomentellous glands; ovary half inferior,

with 2-6 (-8) parietal placentas 7. Homalium

aa. Petals absent.

h. Flowers 5, branchlets unarmed (except in Casearia stjohnii).
i. Stipules large, foliaceous, and persistent or not.

leaves slender and rather long-petiolate; flowers pedicellate, in
racemes; calyx 5.5 to 6.5 mm long; stamens <x ; ovary 3-5 Jocular ....8. Prockia
jj. Stipules somewhat falcate, entire-margined, caducous; leaves short-
petiolate; flowers sessile, paniculate-spicate, densely crowded or not
along the secondary rachises; calyx to 3.5 mm long; stamens 4;

ovary 1-locular - 9. Tetrathylacium

ii. Stipules small to usually very small, caducous, or absent; ovary 1-locular.

flora of Panama (Family 128. Flacourtkvceae)

mm. Style a
11. Staminodes absent.

n. Indumentum of simple hairs; leaf blades pellucid-punctate;

sepals to 9 mm long; anthers not appendaged ...12. Laetia

nn. Indumentum mostly stellate; leaf blades epunctate; sepals

1.3-5 cm long; anthers mucronate 13. Ryania

kk. Inflorescences racemose; leaves estipulate, 3- (5-) nerved from the

base; calyx, at length, split into 2-3 lobes 14. Lunania

owers $ ? (or polygamous) ; branchlets often with axillary spines; leaves

1. LINDACKERIA
Lindackeria C. Presl, Reliq. Haenk. 2: 89, pi. 65, 1835.

Shrubs or trees, unarmed. Leaves alternate, petiolate, the petiole often elongate
and pulvinate at the apex, the stipules none or early caducous, the blade rather
large, entire-margined or the margins dentate, glabrous or pilose below, the hairs
simple or stellate, penninerved. Inflorescences axillary or terminal, short to elongate,
racemose or paniculate, usually many-flowered, infrequently flowers solitary. Flowers
rather small, g or c? by abortion, pedicellate, the bracts early caducous; sepals
3, imbricate; petals 6-12, imbricate, equalling ± the sepals; stamens oo, the fila-
ments filiform, free or rarely united into a tube, the anthers basifixed, linear,
longitudinally dehiscent; ovary shortly stipitate, smooth, muricate, or echinulate,
usually hairy, 1-locular, with 3 parietal placentas, the placentas pauci- or multi-
ovulate, the style simple, the stigma simple or obscurely 3-lobulate. Fruits capsular,
globose, coriaceous to ligneous, tuberculate or echinate, tardily 3-valvate; seeds
few, ovoid, the endosperm copious, the embryo large, the cotyledons large, cordate,
foliaceous.

About 12 species in the tropics of Africa and America; only the following
species reported from Panama and Central America.

1. Lindackeria laurina C. Presl, Reliq. Haenk. 2: 89, pi. 65, 1835.— Fig. I.
Mayna laurina (C. Presl) Bentham, Jour. Proc. Linn. Soc, Bot. 5 (Suppl. 2): 87, 1861.
Oncoha laurina (C. Presl) Warb. in Engler & Prantl, Nat. Pflanzenfam. 3 (6a): 18, 1893.

Tree 3-8 m high, sometimes to 15(-35) m high, the trunk 5-35 cm in diam,
the branchlets angulate, striate, glabrous, often grayish. Leaves glabrous, the
petioles 2.5-11 cm long, slightly pulvinate at the apex; blade narrowly elliptic,
elliptic or ± ovate, obtuse to rounded at the base, long-acuminate at the apex, the
margins entire to usually undulate, 10-31 cm long and 5-11 cm wide, papyraceous,
± shiny above, paler beneath, the costa prominent beneath, the secondary veins
slightly prominent beneath. Inflorescences axillary and terminal, paniculate, to
22 cm long, the rachis glabrous. Flowers white, the pedicels angulate, to 1.5 cm
long, glabrous, ± shining; sepals ± obovate or largely obovate, cucullate, reflexed
at anthesis, 6-9 mm long and 4.5-5.5 mm wide, glabrous or inconspicuously

ANNALS OF THE MISSOUBI BOTANICAL GARDEN

Fig. 1. Lindach
(X5); D, gynoecium (X5); E. dehiscent
E after Stem et al. 516 (MO).

puberulous and shining outside, caducous; petals elliptic to narrowly elliptic, obtuse
to ± rounded at the apex, ± undulated along the margins, 9-12 mm long and
2.5-5 mm wide, glabrous, shining outside; stamens 26-40, the filaments scarcely
connate at the base, unequal, 2-4.5 mm long, glabrous or sparsely and inconspicu-
ously puberulous, the anthers ± oblong, emarginate at the base, slightly bifid at
the apex, ca 3.5-4.2 mm long, glabrous or sparsely and inconspicuously puberulous

1968]

flora of Panama (Family 128. Flacourtia\ceae) 97

at both ends; ovary very shortly stipitate, the stipe ca 0.5 mm long or less, ovoid,
densely appressed-soft-echinate, the bristles minutely puberulous; style 5-6 mm
long, minutely puberulous especially on the lower half, the stigma shorty 3-lobulate.
Capsule long surmounted by the persistent style, echinate, to 1.5 cm in diam
(without bristles), the pericarp rugose, shiny, and inconspicuously puberulous, the
bristles to 8 mm long and inconspicuously puberulous; seeds few - 1, ± angulate,
to 7 mm long.

Southern Mexico, Guatemala, British Honduras, Costa Rica, Panama, and per-
haps Colombia and Venezuela; known colloquially in Panama as uvre (cf. Cooper
& Slater 167) and chopo cucullo (cf. Duke & Bristan 394).

This species is generally known in Panama under the name Oncoba lamina.
According to Duke & Bristan the fruits have a bad flavor and the leaves are used in
the Province of Darien to cure snakebites.

canal zone: Mojinga swamp nr mouth of Rio Chagres, alt below 1 m, Allen 907 (F,
MO, NY, US); Barro Colorado I, Aviles 110 (F), Bangham 406 (F, US), Frost 137 (F),
Shattuck 687 (F, MO), 1088 (F, MO, US), Starry 296 (F); vie of Cerro Viejo on K 16 C,
Blum 1271 (MO); drowned forest of Rio Puente nr jet with Rio Chagres, alt 66 m,
Dodge et al. 16828 (MO); Thatcher Ferry Rd, Duke 5755 (MO); Madden Dam,
Boy Scout Camp Rd, roadside woods, Dwyer & Elias 7472 (MO); Gatun Station,
Hayes 15 (MO, US), 92 (NY); s. loc, Hayes 18 (US); rd W of Pina Base Cam
1589 (MO), 1770 (MO); vie of Frijoles, Piper 5794 (NY, US); Agua Clara, on il.
River, alt 10-40 m, Pittier 3991 (NY, US); betw Corozal & Ancon,

(NY, US); 5 mi NW of Cocoli, alt 500 ft, Tyson 1616 (MO), chiriqui: vie of Remedios,
alt 0-150 m, on edge of mangrove swamp, Allen 3665 (F, MO, NY, US) ; Progreso, Cooper
& Slater 167 (F, NY, US); Tole, vie of Santa Ana Well, alt ca 1000 ft, thicker ,
paralleling brook, Dwyer & Kirkhride 7464 (MO): vie

5228 (F, NY, US), colon: vie of Camp Pina, alt 25 m, Allen 3675 (MO, NY, US), darien:
Rio Tuira, betw Rio Punusa & Rio Mangle, Duke 14581 (MO); San Jose River, Duke &
Bristan 394 (MO); vie of Cana, summit of Cerro Cana, alt 2500 ft, Stern et al. 516 (MO,
US). Panama: vie of La Chorrera, alt 30 m, Allen 2751 (MO, US); Rio Tocumen down
road from La Siesta, Dwyer 4338 (MO); Chiman, rain forest, Lewis et al. 3236 (MO); La
Chorrera, Bro. Paul 501 (MO, US).

2. MAYNA

Mayna Aublet, Hist. PI. Gui. Fr. 921, pi. 352, 1775.
Dendrostylis Karsten & Triana, Linnaea 28: 431, 18563.

Shrubs or small to medium-sized trees. Leaves alternate, petiolate, the petiole
pulvinate at the apex, the stipules early caducous, the blade entire-margined or
the margins dentate. Flowers dioecious; staminate flowers axillary, in small fascicles
or sometimes solitary; pistillate flowers axillary and usually solitary; sepals 3,
imbricate; petals 6-9, imbricate, longer than the sepals; staminate flowers with oo
stamens, the filaments inserted on a scarcely thickened torus, free or almost so,
pilose, the anthers linear, longitudinally dehiscent; pistillate flowers with the
ovary echinulate, 1-locular, with 3 parietal placentas, the placentas multi-ovulate,
the styles 3 or 4, each one 2-lobed, the lobes laciniate. Fruits baccate, globose,

■ding to a note by A. Dugand in the U. S. National Herbarium the correct
Dendrostylis should read: Dendrostylis Karsten & Triana in Triana, Nuevos
Species di Plantas para la Flora Neo-Granadina 27, 1854 (Bogota).

[Vol. 55
98 ANNALS OF THE MISSOURI BOTANICAL GARDEN

bristly, indehiscent; seeds oo, the testa red or orange, fleshy; endosperm copious;
embryo straight; cotyledons cordate.

A small genus much in need of revision; about 15 species in tropical America,
mainly in northern . South America; two species reported from Panama.

In the absence of fruits it is very difficult, if not impossible, to distinguish this
genus from Carpotroche Endl.

a. Leaf blades glabrous; staminate flowers with the sepals 5-7 mm long and with
the petals 10-11.5 mm long; fruit to 2.5 cm in diam 1

aa. Leaf blades with the upper surface sparsely hirtellous especially along the veins
and with the lower surface softly hirsute; staminate flowers with the sepals to
8 mm long and with the petals 12-14.5 mm long; fruit to 1.5-2 mm in diam

1. Mayna longicuspis (Standley) Standley, Trop. Woods 52: 27, 1937.
Sloanea longicuspis Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 4: 229, 1929.
Carpotroche subintegra Standley, loc. cit. 236.

Shrub or small tree 2-5 m high, the trunk 2.5-5 cm in diam, the branchlets
terete, ochraceous, glabrous or almost so. Leaves clustered towards the tip of the
branchlets, the petiole 1-4 cm long, thickened at the apex, canaliculate above,
sparsely puberulous to glabrous; blade elliptic to obovate, usually narrowly so,
attenuate and acute to obtuse at the base, acuminate to long-acuminate at the apex,
the margins entire to remotely and obscurely serrulate, sometimes coarsely serrate
towards the apex, the serrations often mucronulate, 15-23 cm long and 4-8.5 cm
wide, chartaceous, glabrous, the costa prominent beneath, the lateral veins prominu-
lous beneath. Staminate flowers white, in few-flowered fascicles, the pedicels 1-3
mm long, shortly appressed-puberulous, the bracteoles triangular-filiform; sepals
± rotund, conchiform, 5-7 mm long and 6-6.5 mm wide, shortly appressed-puberu-
lous; petals 7, obovate, rounded at the apex, 10-11.5 mm long and 5.5-6 mm wide,
glabrous; stamens oo, the filaments ca 0.5-0.6 mm long, strigillose, the anthers
oblong-linear, ca 1.8 mm long, strigillose along the margins. Pistillate flowers not
seen. Fruit yellow or green, globose, to 2.5 cm in diam (without bristles), sur-
mounted by the 3 persistent styles, densely bristly, the bristles to 7 mm long and
hirtellous as the pericarp; seeds orange, to 1 cm long, the testa striate, shining.

Only reported from Panama. This species is close to and perhaps conspecific
with the Colombian M. suaveolens (Karsten & Triana) Warburg (in Engler &
Prantl, Nat. Pflanzenfam. 3(6a): 19, 1893).

canal zone: Quebrada Lopez, alt 30 m, Allen 2127 (F, MO, US), chiriqui: Progreso,
Cooper & Slater 234 (type US); vie of San Bartolome, Peninsula de Burica, alt 0-50 m,
Woodson & Schery 870 (F, MO), cocle: N rim of El Valle de Anton, alt 600-1000 m, Allen
1646 (F, MO), darien: Quebrada Bidoto (Peccary Creek) off Rio Areti, Duke 13591
(MO). Panama: Cerro Trinidad, alt 800-1000 m, Allen 3771 (F, MO), san blas: Perme,
Cooper 638 (type Carpotroche subintegra F, NY); along headwaters of Rio Malatupu,
seasonal evergreen forest, Elias 1763 (MO); forest around Puerto Obaldia, alt 0-50 m,
Pittier 4296 (F, US).

2. Mayna zuliana (Pittier) A. Robyns, Ann. Missouri Bot. Gard. 55: 78,

1968.— Fig. 2.
Carpotroche zuliana Pittier, Bol. Com. Ind. (Venezuela) 4(34): 38, 1923 (Arboles y
Arbustos Nuevos de Venezuela, secunda y tercera decadas).

flora or panama (Family 128. Flacourtkiceae) 99

100 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Shrub or small tree, 4.5-7.5 m tall, the trunk 7.5-15 cm in diam, the branchlets
hirsute when young. Leaves with the petioles slender, 0.8-3 cm long, canaliculate
above, hirsute, the stipules subulate, 7.5-10 mm long, hirsute; blade narrowly
obovate to elliptic-obovate, long- attenuate towards the base, ± obtuse at the base,
acuminate at the apex, remotely serrulate-ciliate along the margins, 11-30 cm
long and 4-11.5 cm wide, chartaceous, the upper surface ± shiny and sparsely
hirtellous especially along the veins, the lower surface dull and softly hirsute, the
costa prominent beneath, the secondary veins prominulous. Staminate flowers
solitary or in small fascicles towards the end of the branchlets, the pedicels to
10 mm long, densely short-sericeous, the bracts subulate, 4-6 mm long, densely
short-sericeous; calyx valvate, the sepals rotund-conchiform, to 8 mm long and
wide, densely short-sericeous outside except on the margin covered in prefloration,
glabrous inside; petals 7-8, white, obovate, rounded, 12-14.5 mm long and 5-7 mm
wide, appressed-silvery-sericeous in the middle outside, glabrous inside and
marginally outside; stamens ca 50, the filaments almost free, ca as long as the
anthers, silvery-pilose, the anthers oblong, ca 2 mm long, very sparsely pilose,
longitudinally dehiscent; rudimentary ovary lacking. Pistillate flowers not seen.
Fruit with the pedicel to 8 mm long, articulate below the middle (?), densely
short-sericeous, the bracts and sepals as in the staminate flowers, long-persistent;
bacca globose, surmounted by the persistent styles, 1.5-2 cm in diam (without
bristles), densely bristly, the bristles ca 3-5 mm long, hirtellous, the styles 3 or 4,
hirtellous below, each style deeply bifid, the lobes laciniate; seeds few, broadly
ovate-angulate, to 7.5 mm long, striate, glabrous.

Panama and Venezuela (Dept Zulia).

darien: headwaters of Rio Chico, alt 500-750 ft, fairly frequent in heavy forest, Allen
4630 (MO); betw Quebrada Venado & Peje swamp on the headwaters of Rio Tuqueza,
Bristan 1060 (MO); km 16 from Yaviza along Quebrada Uvital off Rio Chucunaque,
Duke 4994 (MO); Rio Balsa, betw Manene & Tusijuanda, Duke 13570 (MO); InterAmer
Hwy, ca 1 mi SE of Rio Tuira, Duke 14561 (MO) ; betw Rio Punusa & Rio Pucro, Duke
14648 (MO); in pasture belonging to Mr. Pablo Othon, Duke & Bristan 290 (MO);
Patifio, on cliffs along the beach, Pittier 6673 (US); vie of Campamento Buena Vista, Rio
Chucunaque above confluence with Rio Tuquesa, growing on muddy bank, Stern et al.
908 (MO, US). Panama: woods along PanAmer Hwy ca halfway betw El Llano & Rio
Mamoni, Duke 5629 (MO); Tocumen, Dwyer 1179 (MO).

3. CARPOTROCHE
Carpotroche Endl., Gen. PI. 918, 1839.

Shrubs or trees. Leaves alternate, short- to long-petiolate, the stipules early
caducous, the blade entire-margined or the margins mostly serrate. Flowers odori-
ferous, rather large, monoecious, dioecious or rarely polygamo-dioecious, the stami-
nate flowers axillary, in short racemes or almost fasciculate, the pistillate ones
axillary and solitary or few-fasciculate; sepals 2-3, imbricate, persistent; petals
4-12, ± biseriate, imbricate; stamens oo (pistillate flowers lacking any rudiments
of stamens), the filaments short, inserted on a slightly thickened torus, pubescent,
the anthers linear, basifixed, longitudinally dehiscent; ovary superior (no rudimen-
tary ovary in staminate flowers), longitudinally costate, 1-locular, the placentas
4-8, many-ovulate; styles isomerous with the placentas, short, persistent, the stigmas

1968]

flora of Panama (Family 128. Flacourtiaceae) 101

scarcely capitellate. Fruits capsular, rather large, coriaceous to ligneous, longitudi-
nally alate, the wings up to 16, large and undulated, crowned by the persistent
styles, indehiscent; seeds numerous, smooth, immersed in a fleshy pulp; albumen
copious; embryo straight; cotyledons foliaceous.

A neotropical genus of about 20 species; one species reported from Panama.

1. Carpotroche platyptera Pittier, Contr. U. S. Nat. Herb. 12: 178, pi. 19, figs.
15, 16, 1909.-F:g. 3.

Shrub 1.50-2.40 m high, or small tree up to 6 m high, the trunk to ca 7.5 cm
in diam, the young branchlets inconspicuously puberulous. Leaves on petioles to
5 cm long, thick, and inconspicuously puberulous; blade obovate to narrowly
obovate, long-attenuate towards the base, obtuse at the base, acuminate at the
apex, coarsely serrate to serrulate along the margins, to 50 cm long and 18 cm
wide, chartaceous, inconspicuously puberulous mainly along the costa and lateral
veins beneath, the costa very prominent and the lateral veins prominent below.
Staminate flowers fasciculate in the leaf-axils (or on the trunk?), the pedicels
short, puberulous; sepals 2, concave, to 6.5 mm long, puberulous outside, glabrous
inside; petals 4 or more (5 or 6 in Panamanian material), oblong-elliptic to nar-
rowly oblong-elliptic, the apex involute at least in bud, to 6.5 mm long and 1.7-2.7
mm wide, sparsely appressed-pilose in the middle especially outside; stamens oo
(21-26 counted in Panamanian material), the filaments ca 0.5 mm long, densely
barbate, the anthers to 3.5 mm long, appressed-pilose. Pistillate flowers (not seen;
description ace. to Pittier, loc. cit. 179), solitary, to 30 mm in diam; sepals as in
staminate flowers but larger; petals 8, elliptic-obovate, ± obtuse; ovary ovoid,
longitudinally 8(- 10) -ribbed, pilose, with 4(-5) placentas, the styles 4(-5), distinct,
very short. Capsule ± globose, reddish or purplish, 2.5-4.5 cm long (wings in-
cluded), ligneous, with 8-10 broad, vertical, undulate-margined wings, these 0.8-2
cm broad in the middle, rigid-chartaceous, inconspicuously puberulous; seeds ovoid,
angulate, to 0.8 cm long.

Guatemala to Panama, along the Atlantic coast, in forest.

Fig. 3. Carpotroche platyptera Pittier: Capsule (XI), after von Wedel 1453 (MO).

102 ANNALS OF THE MISSOURI BOTANICAL GARDEN

It is difficult to say whether the plants are monoecious or dioecious. On
several herbarium sheets, however, staminate flower buds and capsules are present
(although the latter are never attached to the specimens) and this suggests that the
flowers are monoecious. I did not see pistillate flowers!

Carpotroche crassiramea Pittier (loc. cit. 180) from Costa Rica is probably
synonymous with our species, the only significant difference being the number of
wings on the capsule: 10 wings in C. crassiramea and 8 wings in C. platyptera.

This species is called colloquially rabo de gallo in Panama (cf. von Wedel
2186). According to Seibert (1571) the bark and wood smell much like potassium

eocas del toro: region of Almirante, Daytonia Farm, Cooper 367 (F, US), 377 (F, NY,
US); Changuinola Valley, Dunlap 129 (F); 1045 mi S from mouth of Changuinola River,
Lewis et al. 879 (MO); Nievecito, United Fruit Co., Seibert 1571 (US); lower Changuinola
River, Stork 129 [ui Lagoon, von Wedel 1029 (MO, US); Water Valley,

vie of Chiriquf Lagoon, von Wedel 1453 (MO), 1471 (MO), 1650 (MO); Fish Creek, vie of
Chiriqui Lagoon, von Wedel 2186 (MO, US).

4. HASSELTIA
Hasseltia H.B.K., Nov. Gen. Sp. PI. 7: 231, pi 651, 1825.

Shrubs or trees. Leaves alternate, petiolate, the stipules soon caducous; blade
entire or serrate along the margins, 3-nerved from the base, with 2 glands on the
upper side at the base. Inflorescences terminal or axillary, in large, compound,
repeatedly 3- to 4-radiate umbels. Flowers small, g , pedicellate; sepals (3)4,
valvate, persistent; petals (3)4, valvate, persistent; stamens oo, free, inserted on a
glandular or eglandular disc, the filaments filiform, the anthers small, subglobose,
2-thecate, basifixed, longitudinally dehiscent; ovary free, superior, 2- (3-) celled by
complete union of the 2(3) opposite parietal placentas, the ovules oo, the style
simple, entire. Fruits baccate, each cell usually 1-seeded; seeds pendulous, the
testa coriaceous, the endosperm present; embryo straight; cotyledons foliaceous.

A neotropical genus of about five to eight species; three species reported from

a. Leaf blades chartaceous, coarsely and irregularlj
aa. Leaf blades subentire- to entire-margined.

puberulous 2. H. cf.

bb. Leaf blades coriaceous, the margins entire and slightly recurved; sepals

to 3 mm long; ovary glabrous 3. H. rigida

1. Hasseltia floribunda H.B.K., Nov. Gen. Sp. PL 1: 232, pi. 651, 1825.— Fig. 4.

Shrub or small tree 3-10 m high, the trunk to 30 cm in diam, the branchlets
glabrous or appressed-puberulous. Leaves dark green, with the petioles to 4 cm
long, canaliculate above, glabrous or appressed-puberulous; blade from narrowly
to broadly-elliptic, sometimes oblong-elliptic, acute at the base, obtusely acuminate
at the apex, coarsely and irregularly serrate along the margins, to 22.5 cm long
and 11 cm wide, chartaceous, glabrous above, glabrous or appressed-puberulous
especially along the veins below, the main veins and secondary veins prominent

flora of panama (Family 128. Flacourtiaeeae) 103

Fig. 4. Hasseltia fioribunda H.B.K.: A, habit (Xi/ 2 ); B, flower (X9); C & D, stai
(X30); E, gynoecium (X8); F, bacca surmounted by the persistent style (X5). A-E i
Erlanson 380 (US); F after Shattuck 855a (MO)

[Vol. 55
104 ANNALS OF THE MISSOURI BOTANICAL GARDEN

beneath, the network of the veins prominulous below. Inflorescences usually large
and many-flowered, the axes puberulous. Flowers greenish, yellowish-white or
white, the pedicels slender, to 1 cm long, puberulous to tomentellous; sepals 4,
reflexed, ovate, acute, to 2.8 mm long and 1.3-1.7 mm wide, puberulous to tomentel-
lous; petals 4, reflexed, obovate, acute, to 2.7 mm long and 1.2-1.4 mm wide,
puberulous to tomentellous; stamens 30-40, inserted on a glandular disc (apparently
8 separate glands), the filaments 3-4 mm long, puberulous to sparsely puberulous,
the anthers ca 0.25-0.3 mm long; ovary substipitate, globose-ellipsoid, ca 1.5-1.8 mm
long and 1.3-1.5 mm wide, puberulous to subglabrous, the style ca 1.5-2 mm long.
Bacca subglobose, surmounted by the persistent style, 4-6 mm long, the pericarp
succulent, red to black, puberulous, becoming glabrous; seeds 1 or 2, 3.5-4.5 mm
long, the testa glabrous.

From Honduras to Panama, and in northern South America.

eocas del toro: nr Bocas del Toro, Carleton 198 (F, US); Ghanguinola Valley,
Dunlap 550 (F). canal zone: Barro Colorado I, Bailey & Bailey 534 (F), Bangham 539
(F), Brown 39 (F), 57 (F), Dwyer 1450 (F), Shattuck 674 (MO), 855a (MO, US),
855b (MO), Standby 31441 (US), 40929 (US), Wetmore & Abbe 196 (F), Wilson 58 (F),
703 (F), Woodworth & Vestal 448 (F), 646 (F), Zetek 5071 (F); on Las Cruces Trail,
thmian Ilwy. Blum & Miller 2283 (MO); 12 m S of Colon, vie Rio Providencia,
Blum & Tyson 2318 (MO); nr Quebracho on Panama Railroad, Christopher son 122 (US);
lerson 129 (US); vie of Hill C-6, Ft. Sherman, shaded areas, Duke
4389 (MO); Madden Dam, Dwyer 2183 (MO); Empire Station, Hayes 36 (MO); Bojio
Station, Hayes 946 (NY); common in many places along Panama Railroad, Hayes 1016
i illo, Piper 5604 (NY, US), 5655 (US); hills N of Frijoles, Standley
27428 (US); Darien Station, moist thicket, Standley 31614 (US); Obispo, Standley 31686
(US); rd from Ft. Sherman to Gatun Locks, Tyson & Blum 3805 (MO), colon: Portobello,
La Cruzio Trail, Ebinger 100 (MO); betw France Field, Canal Zone, and Catival, wooded
swamp, Standley 30266 (US), 30297 (US), darien: trail betw Pinogana and Yavisa, alt
ca 15 m, Allen 246 (F), 289 (F, MO, NY); headwaters Rio Chico, alt 500-750 ft, Allen
4604 (F, MO); Rio Pirre, 2-5 mi above El Real, Duke 5089 (MO); hills nr Pidiaque,
Duke 8039 (MO); Rio Pirre, Duke & Bristan 8272 (MO), 8278 (MO); Rio Sabana, King
Leopold III 104 (MO); forest around Pinogana, Pittier 6528 (US); El Real de Sta. Maria,
Pittier 6967 (NY, US); vie of Paya, Rio Paya, Stem et al. 186 (MO, US); Rfo Tuira, S
of El Real, Don Pablo Othon's pasture, foothills of Cerro Pirre, Stern et al. 748 (MO, US);
Cana & vie, alt 2000-6500 ft, Williams 984 (NY, US). Panama: along Rio Juan Diaz above
Juan Diaz, alt ca 30 m, Allen 940 (F, MO, NY, US); Bohio Soldado, Cowell 243 (NY, US);
nr confluence of Rio Pacora & Rio Corso, alt ca 450 m, Duke 11968; San Jose I, Erlanson
380 (NY, US), Johnston 372 (MO, US); nr Chepo, Kluge 32 (F), s.n. (US); Rio Tapfa,
Standley 26139 (US), 28231 (US); Rio Tocumen, moist forest, Standley 29330 (US), 29360
(US); Juan Diaz, Standley 31597 (US), province unknown: s. loc, Seemann s.n. (NY).

guatemalensis Warb. in Engler & Prantl, Nat.
3(6a):32,fe72,D-E, 1893.

Tree to 30 m tall, the trunk to 55 cm in diam, the branchlets puberulous when
young. Leaves with the petioles to 2 cm long, canaliculate above, appressed-puber-
ulous when young; blade elliptic, obtuse to acute at the base, obtuse to obtusely
short-acuminate at the apex, subentire-margined, to 12.5 cm long and 5.5 cm wide,
chartaceous, somewhat shiny and glabrous above, dull and sparsely appressed-
puberulous when young especially along the veins beneath, the main veins and
prominulous beneath. Inflorescences large and many-flowered, the
; puberulous to tomentellous. Flowers with the pedicels slender, to 1 cm long,

flora of Panama (Family 128. Flacourtiaicme) 105

tomentellous; sepals (3-) 4, ovate, acute, ca 4 mm long and 3 mm wide, tomentel-
lous; petals (3-) 4, ovate, acute, ca as long as the sepals but somewhat narrower,
tomentellous; stamens inserted on a glandular disc, with glabrous filaments to
3 mm long; ovary puberulous, 2- or 3-celled, the style ca 1.5 mm long. Bacca (im-
mature) subglobose, sparsely puberulous.

chiriqui: nr sawmill on Rio Chiriqui Viejo, 3 km N of Camp EI Volcan, alt 4300 ft,
evergreen rain forest, Little 6063 (F, MO, NY).

The above described collection is very close to H. guatemalensis (see Standley
& L. Williams, Flora of Guatemala, Fieldiana: Bot. 24(7): 94, 1961) (Mexico to
Nicaragua), but differs by the tomentellous pedicels and sepals, and by the sparsely
puberulous young fruits. Furthermore, some flowers are 3-merous (3 sepals and
3 petals), and the dissection of one ovary showed clearly a 3-celled condition.
More material is needed, however, to evaluate these characters.

3. Hasseltia rigida Woodson ex A. Robyns, Ann. Missouri Bot. Gard. 55: 77, 1968.
Tree to 30 m tall, the branchlets glabrous or nearly so. Leaves with the petioles
robust, to 2 cm long, canaliculate above, appressed-puberulous, becoming glabrous;
blade elliptic to narrowly elliptic, or subobovate, cuneate at the base, obtusely
acuminate at the apex, the margins entire and slightly recurved, to 10 cm long
and 4 cm wide, coriaceous, dull on both sides, glabrous and with the venation in-
conspicuous above, appressed-puberulous especially along the 3 prominent main
veins to glabrescent beneath, the secondary veins prominulous and the network
of the venation inconspicuous below. Inflorescences apparently terminal, large,
many-flowered, the axes appressed-puberulous to appressed-tomentellous, the bracts
small, deltoid. Flowers tan-colored, the pedicels to 5 mm long, appressed-tomentel-
lous; sepals 4, slightly unequal, ovate, acute, to 3 mm long and 1.8 mm wide, tomen-
tellous outside, puberulous inside; petals 4, ovate, acute, slightly shorter than the
sepals, tomentellous outside, puberulous inside; stamens with the filaments to 2.5
mm long, sparsely puberulous, the anthers minute; ovary globose, glabrous, the
style to 2.2 mm long and glabrous. Bacca red, subglobose, to 7-8 mm long, glabrous;
seeds 2-4, glabrous.

Known only from Panama.

cocle: region N of El Valle de Anton, alt 1000 m, Allen 3733 (holotype MO).

5. NEOSPRUCEA
Neosprucea Sleumer, Notizbl. Bot. Gart. Mus. Berlin 14: 47, 1938.
Spruceanthus Sleumer, loc. cit. 13: 362, 1936, non Verdoorn (Ann. Bryol. Suppl. 4: 151,

Small trees. Leaves alternate, petiolate, the stipules minute and caducous;
blade 3-5-nerved from the base, distinctly 2-glandular at the base on the upper
surface or eglandular, the margins remotely and obtusely glandular-serrate. In-
florescences axillary or terminal, few-flowered, racemose. Flowers large, £ , the
pedicels articulated; sepals (3)4, valvate, persistent; petals (3)4, similar to the
sepals, valvate, persistent; stamens oo, subperigynous, multiseriate, the filaments
filiform, distinct, short, the anthers elongated-linear, basifixed, 2-thecate, dehiscing

106 ANNALS OF THE MISSOURI BOTANICAL GARDEN

by an oblique introrse slit at the apex; receptacle (or disc?) densely pilose be-
tween the filaments; ovary free, superior, incompletely 5-9-locular by the intrusion
of the placentas, the ovules oo; style simple. Fruits baccate, dry, indehiscent, sur-
rounded by the persistent perianth and surmounted by the persistent style; seeds
numerous, small, angulate, the testa smooth.

A genus of six species occuring in tropical South America, but with one species
extending northwards into Panama.
1. Neosprucea aff. sararensis Cuatr., Trop. Woods 101: 21, 1955.

Small tree (?), the branches glabrous. Leaves with the petioles to 15 mm
long, canaliculate above, rounded beneath, short-strigose; blade narrowly elliptic
to narrowly oblong-elliptic, ± rounded to attenuate (and somewhat oblique?) at
the base, long-acuminate at the apex, remotely crenate along the margins, the
crenations distinctly glandular below, 14-20.5 cm long and 4.5-7.5 cm wide, char-
taceous, distinctly 3-nerved from the base, sparsely short-strigose beneath especially
along the prominent main nerves, the secondary nerves transversely subparallel
and prominulous. Flowers light green, in short, axillary racemes, the (fruiting)
pedicels to 12 mm long, terete, rather thick, articulated close to the base, densely
and rather stiff -puberulous; sepals (surrounding the fruit) unequal, broadly
Dvate, apparently shortly united at the base, obtuse at the apex, 11-13 mm long
and 9-13 mm wide, rather fleshy, tomentellous outside, glabrous inside in the
middle but tomentellous towards the margins; petals (surrounding the fruit) ovate,
obtuse, 11 mm long and 5-6 mm wide, rather fleshy, tomentellous outside, puberu-
lous inside; stamens not seen; receptacle densely short-hirsute (the hairs ca 1 mm
long). Fruit globose, longitudinally several-lobed, ca 1 cm in diam, apiculate
(base of persistent style), the pericarp thin, brittle, blackish, glabrous, ± shiny,
incompletely several-locular; seeds to 4.5 mm long, the testa brown, shiny, thin.

Panama and Colombia.

cocle: El Valle de Anton at the foot of Cerro Pilon, cloud forest, alt ca 2000 ft,
Dwyer & Coma 7997 (GH, MO, US); El Valle de Anton & vie, alt 500-700 m, Seibert
465 (F, MO).

In absence of flowering material, especially of stamens, it is impossible to
name the above collection with certainty. In the key to the genus published by
Guatrecasas in 1955 (Trop. Woods 101: 24-25) it is closest to N. sararensis de-
scribed from the Dept Norte de Santander, Colombia. This collection was deter-
mined: N. grandiflora (Spruce) Sleumer by Williams in 1962 (Fieldiana: Bot.
29: 376). According to Cuatrecasas (loc. cit), in N. grandiflora the leaves are
glabrous, the flowers are sessile or subsessile, the calyx is ca 7 mm long, the fila-
ments 2 mm long, and the anthers 5 mm long, while in N. sararensis the leaves are
subglabrous beneath, the pedicels 12-14 mm long, the calyx 9-10 mm long, the
filaments 3-4 mm long, and the anthers 2 mm long.

6. BANARA

Banara Aublet, Hist. PI. Gui. Fr. 547, pi. 217, 1775.

Trees or shrubs. Leaves alternate, distichous, short-petiolate, the petiole some-
times 1- or 2-glandular at the apex, the stipules small and fugacious; blade simple,

(Family 128. Flacourtiaceae) 107

± inequilateral and oblique at the base, the margins glandular-serrate (the glands
on the lower side of the serrations), penninerved. Inflorescences usually terminal,
infrequently oppositifolious, paniculate or racemose, rarely corymbose. Flowers
small, g , rarely polygamous or dioecious, the pedicels articulated above the base,
the bracts and bracteoles deltoid, minute; sepals 3(-4), valvate, shortly united at
the base, persistent; petals isomerous with the sepals and similar to them, ± imbri-
cate in bud, persistent; stamens oo, hypogynous or the outer ones slightly epigynous,
pluriseriate, inserted on a glabrous or villosulous disc, distinct; filaments filiform;
anthers small, introrse, didymous-subglobose, basifixed, 2-thecate, longitudinally
dehiscent; ovary sessile, 1-locular or incompletely pseudo-3-8-locular, the placentas
3-8, parietal, filiform or lamelliform and intruding into the ovary, the ovules oo,
small and pluriseriate; style simple, the stigma minutely capitate. Fruits baccate,
fleshy or coriaceous, surmounted by the persistent style, indehiscent; seeds numerous,
embedded in a fleshy pulp, the testa crustaceous; endosperm copious, carnose;
embryo small; cotyledons thick.

A neotropical genus of about 30 species; one species reported from Panama.

i Aublet, Hist. PI. Gui. Fr. 548, pi. 217, 1775.— Fig. 5.

Shrub or small tree 3-15 m high, the trunk up to 20 cm in diam, loosely
branched, the branchlets whitish-gray-appressed-tomentellous. Leaves with the
petioles (3-) 6- 12 mm long, canaliculate above, whitish-gray-appressed-tomentellous,
the apex (junction of blade and petiole) often provided with 1 or 2 somewhat
stalked cupulate glands; stipules narrowly triangular, to 4 mm long, whitish-gray-
appressed-tomentellous; blade ovate or oblong, sometimes narrowly oblong, rarely
oblong-obovate, equilateral to markedly inequilateral, rounded or truncate or sub-
cordate at the base, bluntly caudate-acuminate and usually inconspicuously
mucronate at the apex, coarsely serrate along the margins, 7-16 cm long and 3-7
cm wide, chartaceous, discolor, the upper surface dark grayish-green when fresh,
± shiny and usually dark brown when dry, and sparsely appressed-pubescent,
the lower surface dull, pale, densely to sparsely appressed-puberulous, the costa
very prominent and the secondary veins prominent beneath. Inflorescences termi-
nal or rarely oppositifolious towards the end of the branchlets, paniculate, some-
times ± racemose, the axes whitish-gray-appressed-tomentellous. Flowers with
the pedicels 3-5 mm long, whitish-gray-appressed-tomentellous; sepals 3, rarely 4,
oblong-ovate, obtuse to acute at the apex, ca 4.5 mm long and 3 mm wide, ap-
pressed-tomentellous on both sides; petals ± obovate, rounded at the apex, ±
crispate along the margins, slightly longer than the sepals; stamens inserted on a
densely villosulous disc; ovary ovoid, glabrous, with 5-8 lamelliform placentas, the
stigma minutely capitate and obsoletely 5-8-lobulate. Bacca subgldbose, 6-10 mm
in diam, the pericarp thin-coriaceous, green when fresh, blackish when dry; seeds
ovate, ca 1-1.5 mm long, the testa black, shining, longitudinally sulcate.

A rather variable species (especially regarding leaf shape) distributed in Costa
Rica, Panama, and northern South America. According to field notes, the flowers
are said to be white, yellow-green, cream tinged with pale green, or pale purple.

canal zone: Barro Colorado I, Aviles 894 (F), Bailey 225 (F), 308 (F), 618 (F),
Bangham 379 (F), 402 (F), Kenoyer 449 (US), 663 (US), Shattuck 894 (US), Starry 226

ANNALS OF THE MISSOURI BOTANICAL GARDEN

(F); Albrook, U. S. Army Tropic Test Center, Dwyer 6712 (MO); betw Chagres Batteries &
Fort San Lorenzo, Fort Sherman Military Reservation, Maxon & Valentine 6981 (US) ; Cerro

Gordo, nr Culebi M , - [ , - Trail nr Miraflores

Lake, Tyson 1361 (MO); Curundu Survival School Area, Tyson & Dwyer 4453 (MO).
chiriqui: Tote, vie of Santa Ana Well, alt ca 1000 ft, Dwyer & Kirkbride 7464 (MO).
gocle: Rio Hato Airstrip, Blum & Dwyer 2472 (MO); El Valle de Anton, on slopes of
Cerro Pilon, alt 600-1000 m, cloud forest, Duke 12077 (MO), 12128 (MO), 73207 (MO),
Dwyer & Correa 7969 (MO), 7987 (MO); bstw Las Margaritas & El Valle, Woodson et al.
1242 (F, MO, NY), colon: vie of Sardinilla, ca 7-8 mi E of cement plant, Blum & Tyson
484 (MO). Panama: Cerro Campana, elfin forest beyond Motel Su-Lin, alt 2700-3000 ft,
Duke 8652 (MO) ; San Jose I, along rd betw Bodega Bay & Rio Mata Puerco, common even
in abandoned road bed, Duke 12564 (MO), vie of Naval Station, Erlanson 498 (NY, US);
Chilibre, Dwyer h*V lM<>>, 11 ]0 (M(U; fhktm a f-,usK .it Aimljan, alt ca 15 m,
Woodson et al. 1351 (F, MO, NY), veraguas: ca 5 mi N of Santiago by the Santa Maria
River, Blum & Tyson 609 (MO); Ida de Coiba, Penal Colony, Dwyer 2349 (MO); Cerro
de Tute, Dwyer 4319 (MO).

1968]

flora of panama (Family 128. Ffocourtiaeeae) 109

7. HOMALIUM
Homalium Jacquin, Enum. Syst. PI. Ins. Carib. 5, 24, 1760.

Trees or shrubs. Leaves usually alternate, petiolate, the stipules minute and
caducous, rarely absent, the blade entire to usually serrate-crenate, the teeth
glandular beneath, penninerved, without pellucid glands. Inflorescences axillary,
racemose or paniculate. Flowers g , sessile or pedicellate, the pedicels articulated,
the bracts usually minute, caducous or ± persistent; calyx tube (or receptacle?)
turbinate, adnate to the base of the ovary; calyx lobes 5-8(-12), usually narrow,
persistent or accrescent; petals isomerous with the calyx lobes, inserted in the
throat of the calyx, alternating with the calyx lobes, persistent or accrescent;
stamens epipetalous, isomerous with the petals or in fascicles of 2-8(or more), the
filaments filiform, the anthers small, extrorse, dorsifixed, longitudinally dehiscent;
disk represented by a gland opposite each sepal and alternating with the filaments
or fascicles of filaments; ovary with the lower half adnate to the calyx tube (or
half inferior?), 1-locular, the placentas parietal, 2-6(-8), each with (l-)2-7 ovules
near the apex, the styles isomerous with the placentas, the stigmas simple or
capitellate. Fruits capsular, half inferior, ± coriaceous, indehiscent or incompletely
2-8-valvate from the apex; seeds few or solitary, small, angulate, the testa crustace-
ous; endosperm copious.

A large pantropical genus of over 200 species (cf. Hutchinson, The Genera
of Flowering Plants 2: 216, 1967).

All the neotropical species of Homalium belong to the subg. Myriantheia
Warb. (stamens in fascicles) sect. Racoubea Aublet; at present, only one species
is reported from Panama.

Useful references:

Williams, L. O., Homalium Jacq., In Tropical American plants, II Fieldiana-
Bot. 29: 262-263, 1961.

1. Homalium racemosum Jacquin, Enum. Syst. PI. Ins. Carib. 24, 1760.— Fig. 6.
H. stenosepalum Blake, Contr. U. S. Nat. Herb. 20: 234, 1919.

Tree to 30 m tall, the trunk to 90 cm in diam, the bark grayish, the branchlets
glabrous to finely puberulous, lenticellate. Leaves with the petioles 3-13 mm long,
glabrous or finely puberulous; blade elliptic to oblong-elliptic, sometimes narrowly
so, acute to obtuse at the base, acuminate at the apex, the acumen often blunt, the
margins undulate-crenulate to crenate, 8-15 cm long and 3-6 cm wide, thin-
chartaceous to chartaceous, ± shiny on both sides, glabrous or almost so, the costa
prominent below, the lateral veins slightly prominent below. Inflorescences usually
racemose, sometimes paniculate, to 12 cm long, the rachis minutely puberulous.
Flowers white, the pedicels to 10 mm long, articulated above, near or below the
middle, puberulous, the bracteoles narrowly triangular, to 1.5 mm long, minutely
puberulous, caducous or ± long persistent; calyx tube turbinate, minutely puberu-
lous; calyx lobes (5)6-7, ovate to narrowly oblong-ovate, subobtuse to acute at the

110 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 6. Homaliun
(X5); D, petal (X5); E,
After Stern et a!. 917 (MO)

apex, 2.5-5 mm long and 1-1.5 mm wide, minutely puberulous, persistent; petals
ovate to broadly ovate, subobtuse at the apex, to 5-6 mm long and 3.5 mm wide,
minutely puberulous, persistent; stamens in fascicles of 3, rarely 2, the filaments to
4 mm long, glabrous, the anthers ca 0.5 mm long; glands ± transversely elliptic,
to 1.3 mm X 0.8-1 mm, minutely tomentellous; ovary conical, tomentellous, the
styles 3 or 4, distinct from the base, to 1.2 mm long, glabrous or puberulous beneath,
the stigmas simple. Mature fruit not seen.

Mexico, Central America, northern South America, and the West Indies;
called colloquially guayabillo in Panama (cf. Duke & Bristan 234).

canal zone: s. loc, Christopherson 139 (US); around Frijoles, alt 10-30 m, on river
bank, Pittier 2693 (NY, US) ; along Rio Chagres, below Gatun, nr sea level, Pittier 2804

1968]

flora of panama (Family 128. Flacourtiaceae) 111

(holotype H. stenosepalum US; isotypes F, MO, NY), darien: Rio Ucurganti, Bristan
1180 (MO); Rfo Chucunaque betw Rio Membrillo & Rio Subcuti, Duke 8586 (M
frequently found nr rivers, Duke & Bristan 234 (MO); Rio Tuira betw Rio Paya & Rio
Pucro, Duke & Kirkbride 14059 (MO); vie of Campamento Buena Vista, Rio Chucunaque
above confluence with Rio Tuquesa, Stern et al. 917 (MO, US).

Analysis of immature fruits (Christopherson 139 & Pittier 2804) shows that
the capsules are only 1-seeded at maturity. The inner walls of the pericarp are
hairy.

8. PROCKIA
Prockia P. Br. ex L., Syst. Nat. ed. 10, 1074, 1759.

Shrubs or small trees. Leaves alternate, slender-petiolate, the stipules usually
large, foliaceous, persistent; blade membranous, 5-7-nerved from the base, glandular-
serrate along the margins (the glands on the lower side of the serrations), 2-
glandular above at the base. Inflorescences terminal, racemose or fasciculate, rarely
flowers solitary. Flowers g , the pedicels articulated above the base, the bracteoles
2, narrowly triangular; sepals 3, rarely 4, valvate, persistent; petals isomerous with
and smaller than the sepals, persistent, or none; stamens oo, pluriseriate, free,
inserted on the slightly elevated receptacle, the filaments filiform, the anthers small,
didymous-subglobose, basifixed, 2-thecate, longitudinally dehiscent; ovary free,
3-5-locular, the ovules co; style simple, the stigma scarcely enlarged. Fruits bac-
cate, dry, surmounted by the persistent style, indehiscent, 3-5-locular; seeds
numerous, small, imbedded in a white pulp, the testa brittle; endosperm copious;
embryo straight; cotyledons thick.

A neotropical genus of about 10 species; one species known, at present, from
Panama.

1. Prockia crucis P. Br. ex L., Syst. Nat. ed. 10, 1074, 1759.— Fig. 7.

Shrub or small tree 1.2-10 m high, the trunk to 10 cm in diam, the branchlets
grayish-puberulous or glabrous. Leaves with the petioles to 20 mm long, glabrous
or softly hirtellous, the stipules conspicuous, sessile or nearly so, extremely inequi-
lateral, oblique at the base, acuminate at the apex, glandular-crenate along the
margins, up to 17 mm long, usually ca 5-8 mm long, glabrous or softly hirtellous;
blade broadly to narrowly ovate, usually rounded or sometimes subcordate at the
base, acuminate at the apex, glandular-crenate or -serrate along the margins, 4-11
cm long and 2-7 cm wide, thin-chartaceous, slightly discolor, glabrous or shortly
puberulous above, glabrous or softly hirsute below, 5-7-nerved from the base, the
main nerves slightly prominent. Inflorescences racemose. Flowers with the pedicels
terete, slender, 7-17 mm long, glabrous or softly hirtellous, the bracteoles narrowly
triangular, 3-5 mm long; sepals ovate, attenuate and obtuse to acute at the apex,
5.5-6.5 mm long and 3.5-4 mm wide, green, short-hirsute outside, glabrous below
to tomentellous towards the apex inside; petals none; filaments unequal, 4-6 mm
long, yellow, glabrous, the anthers ca 0.3 mm long; ovary subglobose, ca 2-2.5 mm
long and 2.5-3 mm in diam, shortly appressed-hirsute, the style to 4 mm long,
short-hirsute below to glabrous above, the stigma slightly thickened. Bacca ±
globose, 6-7 mm in diam, the pericarp thin, green when young, becoming light red

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 7. Prockia cruris P. Br. ex L.: A, habit (Xi/ 2 ); B, base of leaf blade, upper s
face, with 2 glands (XI); C, flower (X5); D & E, stamen, upper part of filament i
anther (ca X20); F, gynoecium (X5); G ; fruit surrounded by the persistent calyx t
topped by the persistent style (X5). A-F after Stern & Chambers 35 (MO); G after Ty.
& Blum 4104 (MO).

flora of Panama (Family 128. Flacourtiaceae) 113

to very deep red, almost black when completely ripe, shortly pilose; seeds ± ellip-
soid, ca 1.5 mm long and 1 mm in diam, the testa black, longitudinally striate.

A polymorphic species, especially regarding the indument, ranging from
southern Mexico, through Central America, to South America; also in the West
Indies.

canal zone: vie Tropic Test Center Albrook tower, out C-15, Blum 2174 (MO); Rd
K-10, 1-2 mi from Kobbe-Arraijan Hwy, Duke 11709 (MO); Madden Dam, Boy Scout
Camp Road, roadside woods, Dwyer & Elias 7493 (MO); s. loc, Johansen 20 (US); Naval
Ammunition Depot, Group 300 Rd, Stern & Chambers 35 (MO, NY, US); Madden Dam
area nr road, Transisthmian Hwy, edge of woods, Stern et al. 44 (MO, US) ; Fort Clayton
nr old hospital building #519, Tyson & Blum 3913 (MO); old town site of Red Tank,
Tyson & Blum 4015 (MO); several mi off main road S of the island (Barro Colorado I?),
damp shady habit, White 111 (F, MO). Panama: along Chiva-Chiva trail to Search Light
Station beyond Chiva-Chiva, alt 10-100 m, Allen 952 (F, MO, US); nr Rfo Tapia, Juan
Diaz region, edge of forest, Maxon & Harvey 6633 (US); area just NE of Madden Dam,
Stimson 5179 (MO); 1 mi W of Tocumen, in frequently burned areas, Tyson & Blum
4104 (MO) ; boggy grasslands and marginal thickets, betw Pacora & Chepo, alt ca 25 m.
Woodson et al. 1666 (F, MO, NY).

9. TETRATHYLACIUM
Tetrathylacium Poeppig in Poeppig & Endl., Nov. Gen Sp. PI. 3: pi. 240, 184 1 4 ,

p. 34, 1843*.
Edmonstonia Sesmann, Bot. Voy. Herald pi. 18, 1852 4 , p. 98, 1853 4 .

Trees, infrequently shrubs, unarmed. Leaves alternate, short-petiolate, the
stipules large, foliaceous, caducous, the blade rather large, rounded to cordate at
the base, the margins entire to remotely repand-serrate, epunctate, penninerved.
Inflorescences axillary, paniculate-spicate, the flowers sessile and densely crowded or
not along the secondary rachises (spikes), elongated in fruit. Flowers small, g,
each one subtended by a bract and 2 bracteoles, these small and closely appressed
along the rachis; calyx urceolate, thickened, 4-gibbous or 4(-5)-angulate (com-
pressed by the crowding of the flowers along the rachis), 4-lobed, persistent, the
lobes small, rounded, membranous, biseriately imbricate; petals none; stamens 4,
inserted within the calyx tube, the filaments alternate with the calyx lobes, the
anthers introrse, longitudinally dehiscent, exserted or not; ovary sessile, 1-locular,
with 4 parietal, many-ovulate placentas; style very short, the stigma obscurely 4-
lobed. Fruits baccate, coriaceous, the seeds numerous, exarillate, each seed sur-
rounded by a thin membrane (small compartment, of placentary origin?), the
testa foveolate-alveolate.

A genus of three species, ranging from Costa Rica to Peru; two species reported
from Panama.

a. Flowers very densely crowded along the secondary rachises (spikes) of the

inflorescences, calvx 4(-5) angled by the crowding of the flowers I. T johansenii

aa. Flowers not crowded along the secondary rachises (spikes) of the inflorescences;

calyx 4-gibbous 2. T. macrophyllum

4 For dates of publication see Stafleu, Regnum Veget. 52: 362, 441, 1967.

[Vol. 55
114 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1. Tetrathylacium johansenii Standley, Jour. Wash. Acad. Sci. 15: 479, 1925. —

Fig. 8.

Tree to 30 m high, the trunk sometimes buttressed, infrequently shrub, the
branchlets inconspicuously puberulous to glabrous. Leaves with petioles to 7 mm
long, the foliaceous stipules narrowly oblong-ovate, somewhat falcate, to 17 mm
long and 3 mm wide, minutely puberulous; blade oblong, often narrowly so, or
obovate-oblong, or elliptic-oblong, slightly unequal and rounded to subcordate at
the base, long-caudate-acuminate at the apex, remotely repand-serrate to subentire
along the margins, to 25 cm long and 9 cm wide, subcoriaceous, lustrous and
glabrous above, dull and glabrous to minutely puberulous especially along the
prominent costa beneath, the secondary veins prominulous below. Inflorescences
to 8 cm long, the spikes to 2.5 cm long, much enlarged in fruit, the flowers very
densely crowded along the secondary rachises (spikes), the primary and secondary
rachises densely and minutely puberulous, the bracts and bracteoles minute, decidu-
ous. Flowers white, the calyx 4(-5) -angled (by crowding of the flowers along the
rachis), not gibbous, to 2 mm long, glabrous outside, pubescent inside, the lobes
ca 0.5 mm long; stamens with the filaments ca 0.5 mm long, the anthers 0.5 mm
long; ovary apparently glabrous. Bacca globose to obovoid, to 2.5 cm long, glabrous,
the seeds ovoid, to 2 mm long.

Costa Rica(?), Panama, and probably Colombia.

The crowding of the flowers along the axes of the spikes makes them look like
a tiny mosaic and gives the inflorescences a very distinctive look. The fruits are
(ace. to Duke 13115) in clusters to 30 cm in diam and are loaded with ants.

canal zone: Mojinga swamp nr mouth of Rio Chagres, alt below 1 m, Allen 904
(F, NY, US); Barro Colorado I, Carpenter 76 (F); nr Gatun, Goldman 1863 [holotype US,
isotype NY (fragment of inflorescence)]; forest along the Rio Indio de Gatun, nr sea level,
v 2 (US); Naval Ammunition Depot, Group 300 Rd, Stern & Chambers 34 (F,
MO, NY, US), darien: vie of El Real, alt ca 15 m, Allen 959 (F, MO, NY, US); Rio Pucro,
below village of Pucro, Duke 13115 (MO); trail betw El Real & Pinogana, Stern et al.
280 (MO, US), san blas: forests around Puerto Obaldia, alt 0-50 m, Pittier 4300 (US).

The following sterile collections belong probably to this species: canal zone: s. loc,
lohansen 4 (US); Mount Hope Cemetery, Standley 28767 (US). Panama: Rio Tocumen,
moist thicket, Standley 29408 (US).

2. Tetrathylacium macrophyllum Poeppig in Poeppig & Endl, Nov. Gen. Sp.

PL 3: pi 240, 1841, p. 34, 1843.
Edmonstonia pacifica Seemann, Bot. Voy. Herald pi. 18, 1852, p. 98, 1853.

Hum macrophyllum var. pacificum (Seemann) Tr. & PL, Ann. Sci. Nat., Bot.,
17:106,1862.
T. pacificum (Seemann) Standley, Jour. Wash. Acad. Sci. 15: 479, 1925.
T. costaricense Standley, loc. cit. 480.

Tree to 15 m tall, the branchlets inconspicously puberulous to glabrous.
Leaves with robust petioles 0.5-1.5 cm long, the stipules not seen; blade from nar-
rowly to broadly (in Panamanian collection) oblong, somewhat asymmetrical and
deeply cordate to ± truncate at the base, long-acuminate to caudate-acuminate at
the apex (shortly caudate-acuminate in Panamanian collection), entire to remotely
and rather inconspicuously serrate along the margins, to 26 cm long and 14 cm
wide (in Panamanian collection, usually narrower in extra-Panamanian collec-

flora of panama (Family 128. Flacourtiaceae) 115

Fig. 8. Tetrathylacium johansenii Standley: A, habit (Xi/ 2 ); B, flower (X10)j C, id.,
longitudinal section, the gynoecium removed (X10); D, stamen (X20); E, gynoecium
(X20); F 5 bacca (XI). A-E after Allen 959 (MO); F after Stern et al. 280 (MO).

[Vol. 55
116 ANNALS OF THE MISSOURI BOTANICAL GARDEN

tions), coriaceous, glabrous above, glabrous or puberulous beneath, the costa and
secondary veins prominent below. Inflorescences up to 16 cm long, much elongated
in fruit, flowers not crowded along the secondary rachises (spikes), the primary
and secondary rachises densely and minutely puberulous, each flower subtended
by a bract and 2 connate bracteoles, the bract and bracteoles less than 1 mm long,
ciliolate and persistent. Flowers reddish or maroon; calyx 4-gibbous, to 3.5 mm
long, glabrous outside, pubescent inside, the lobes to 1.5 mm long; stamens with
the filaments to 2.2 mm long, glabrous, the anthers exserted, sagittate, to 1.5 mm
long; ovary minutely puberulous, the style thick, less than 1 mm long. Bacca red,
subglobose, to 15 mm long, minutely puberulous; seeds ± ovoid, to 1.5 mm long.

Costa Rica to Peru.

cocle: hills N of El Valle de Anton, alt 1000 m, Allen 2147 (F).

10. CASEARIA
Casearia Jacquin, Enum. Syst. PI. Ins. Carib. 4, 21, 1760; Select. Stirp. Amer. Hist.

132, 1763.

Trees or shrubs. Leaves alternate, distichous, petiolate, the stipules usually very
small and early caducous, rarely ± persistent, the blade entire-margined or the
margins crenate or serrate, often pellucid-punctate and/or striate. Flowers axillary,
mostly in fascicles or glomerules, rarely solitary, sometimes in corymbose or panicu-
late inflorescences, £ , small, usually greenish or yellowish; pedicels articulated,
bracteate at the base, the bracts usually numerous, scale-like and often forming a
cushion in the leaf axils; sepals 4-6(-9), imbricate, ± united at the base into a
short or rather elongated calyx tube, persistent or sometimes slightly accrescent;
petals absent; stamens (5-) 6- 15 (-22), 1-seriate, inserted on the tube of the calyx
or at its base, alternating with as many staminodes, these inter- or intrastaminal,
clavate or flattended and often pilose especially at the apex; filaments equal or
alternately unequal, free or united among themselves and with the staminodes at
the base into a ± perigynous tube; anthers small, elliptic or subglobose, sometimes
apically and dorsally glandulose; ovary free, 1-locular, the ovules usually numerous
on 3-4 parietal placentas; style simple or divided at the apex into 3 branches, the
stigmas 1 or 3, capitate. Fruit a dry or succulent capsule, 3-4-valved (3-angled
when fresh, often 6-ribbed when dry); seeds few to numerous, with a fleshy aril,
the testa coriaceous or crustaceous; endosperm fleshy, embryo straight, cotyledons

About 250 species in the tropics and subtropics of both hemispheres.
The genus is badly in need of a comprehensive revision, "for some of the
common species have been repeated by various authors under several different
names" (cf. Bentham, Jour. Proc. Linn. Soc, Bot. 5 (Suppl. 2): 88, 1861). Eight
species are reported here from Panama, while a few other species occur in Central
America, north of Panama,
a. Stigma 1; staminodes interstaminal (sect. Pitumha).

b. Inflorescences fasciculate; seeds without resinous glands.

c. Fascicles distinctly pedunculate, the peduncles to 4 mm long 1. C. arborea

flora of Panama (Family 128. Viacom-

'.. Branchlets densely tawny-tomentellous; le

2. C. grandiflora

y pubescent to inconspicuously puberuloi

picuously c

. Branchlets minutely pubescent to inconspicuously puberulous; leaves
cuously discolorous.

d at the base into a tuba 1-1.5 mm

long; leaf blades with the margins sharply serrate —.3. C. arguta

ee. Sepals 4.5-5.5 mm long, scarcely united at the base.

f. Pedicels ca 2-3 mm long, articulated below the apex; branches
often with stout, spreading spinescent twigs; stipules deltoid,

ca 1.5 mm long; leaf blades ± distinctly crenate 4. C. stjohnii

ff. Pedicels to 6 mm long, articulated ca 2-2.5 mm above the
base; twigs without spines; stipules subulate, 2-5 mm long;

leaf blades serrulate-denticulate 5. C. guianensis

bb. Inflorescences cymose-corymbiform; seeds with the testa covered with

numerous, dark, resinous glands 6. C. nitida

g. Style simple; leaves copiously pellucid-punctate; sepals 2-3 mm long; anthers
didymous, subglobose, with a dorsal, apical gland; staminodes ± intra-
staminal (sect. Crateria) 7. C. sylvestris

gg. Style divided at the apex into 3 branches; leaves without pellucid dots;
didymous, subglobose, with a dorsal, apical gland; staminodes ± intra-
staminal (sect. Piparea) 8. C. commersoniana

1. Casearia arborea (L. Rich.) Urban, Symb. Ant. 4: 421, 1910 & 8: 447, 1920.
Samyda arborea L. Rich., Act. Soc. Hist. Nat. Paris 1: 109, 1792.

Shrub to 5 m high, the branchlets puberulous when young, soon glabrous.
Leaves shortly petiolate, the petiole up to 4-5 mm long, the stipules narrowly ovate,
to 1 cm long, puberulous, early caducous; blade narrowly oblong, somewhat
inequilateral, slightly oblique and acute to rounded at the base, long-caudate-
acuminate at the apex, closely crenulate-serrulate at the margins, to 12 cm long
and 4.3 cm wide, chartaceous, pellucid-punctate or -lineate, the upper surface
shining, glabrous or with the slightly prominent costa puberulous, the lower sur-
face somewhat paler, dull, puberulous along the venation, the costa and lateral
veins prominent. Inflorescences fasciculate, the fascicles up to 25-flowered, distinctly
pedunculate, the peduncles to 4 mm long, minutely puberulous, the bracts minute.
Flowers yellow, the pedicels up to 4 mm long, articulated at or slightly below the
middle, minutely puberulous especially below the articulation; calyx ca 4-4.5 mm
long, the lobes 5, oblong, united at the base into a short, 5-angulate tube less than
1 mm long, rounded and slightly cucullate at the apex, ca 2-2.5 mm wide, minutely
puberulous to nearly glabrous on both sides; stamens 10, inserted near the apex
of the calyx tube, slightly united at the base among themselves and with the
staminodes, the filaments unequal, ca 2-2.3 mm long, glabrous, the anthers
didymous, subglobose, ca 0.5 mm long, with a dorsal, apical gland, the latter
slightly pilose or not; staminodes 10, interstaminal, to 1.5 mm long, barbate at the
apex; gynoecium ca 3.7 mm long, the ovary 3-angular, ca 1-1.2 mm in diam,
glabrous but pilose at the apex, gradually attenuate into the style, the latter pilose
at the base, otherwise glabrous, the stigma capitate. Capsule ellipsoid-subglobose,
angulate, apiculate, ca 4-5 mm long, pilose at the apex, splitting into 3 valves at
maturity; seeds up to 6 per capsule, ellipsoid, ca 2 mm long, the testa minutely
alveolate, the aril fimbriate-lacerate.

[Vol. 55
118 ANNALS OF THE MISSOURI BOTANICAL GARDEN

West Indies, British Honduras to Panama, Guianas, and Brazil.
canal zone: Barro Colorado I, clearing, Shattuck 670 (F); Panama: top of Cerro Jefe,
alt 2700-3000 ft, Tyson et al. 4398 (MO).

2. Casearia grandiflora Camb. in St.-Hil., Juss. & Camb., Fl. Bras. Merid. 2:

239, pi. 126, 1830.

Small tree (?), the branchlets densely tawny-tomentellous. Leaves shortly
petiolate, the petiole to 4 mm long, densely tawny-tomentellous, the stipules linear,
fugaceous; blade oblong, obtuse or ± acute and sometimes slightly inequilateral at
the base, rather long- acuminate at the apex, crenulate-serrulate at the margins, up
to 10 cm long and 3 cm wide, chartaceous, obscurely pellucid-punctate, markedly
discolorous, the upper surface dark green when fresh, dark brown when dry, ±
shining and glabrous or with the costa puberulous, the lower surface dull, densely
tawny-tomentellous and with the costa and lateral veins prominent. Inflorescences
sessile or nearly so, fasciculate, 10-20-nowered, the bracts rather large, subcircular,
tawny-tomentellous outside, glabrous inside. Flowers shortly pedicellate, the pedicel
articulated close to the base, tawny-tomentellous; calyx ca 7 mm long, the tube ±
5-costate and ca 2-2.5 mm long, the lobes 5, rather broadly elliptic, rounded at the
apex, ca 3.5 mm wide, tawny-tomentellous without, puberulous within; stamens
10, inserted near the apex of the calyx tube, scarsely united at the base among
themselves and with the staminodes, the filaments unequal, up to 2.5 mm long and
glabrous, the anthers didymous, subglobose, ca 0.5 mm long, with a dorsal, apical,
barbate gland; staminodes 10, interstaminal, ca 1.8-2 mm long, tomentose-
barbate; gynoecium 5.5-6 mm long, the ovary ca 1.5-1.8 mm in diam, glabrous at
the base, densely whitish-pilose at the apex, gradually attenuate into the style, the
latter densely whitish-pilose below to glabrous apically, the stigma capitate and
densely papillate. Capsule (ace. to Eichler in Mart., Fl. Bras. 13(1): 479, 1871)
turbinate-ellipsoid, 6-angulate, apiculate, 7-8 mm long, glabrous; seeds 3-4, oblong,
ca 2.5 mm long, the testa glabrous, minutely reticulate-foveolate, the aril small,
fimbriate-lacerate.

Southern Darien in Panama, Colombia, Venezuela, the Guianas and Brazil
(Para, Ceara, Maranhao).

darien: vie of La Palma, alt 0-50 m, Pittier 5709 (F, US).

3. Casearia arguta H.B.K., Nov. Gen. Sp. PL 5: 364, 1823.

Shrub or small tree 1.5-12 m tall, the trunk to 12 cm in diam, unarmed, the
branchlets minutely pubescent. Leaves short-petiolate, the petiole up to 1 cm long,
canaliculate above, minutely pubescent; blade narrowly oblong or narrowly ovate-
oblong, obtuse to acute at the base, long- acuminate at the apex, to 17 cm long and
6 cm wide, chartaceous to rigid-chartaceous, pellucid-punctate or not, slightly dis-
color, the margins sharply serrate, more or less shining and minutely pubescent
to glabrous above, somewhat paler, dull and minutely pubescent below, the costa
and lateral veins prominent beneath. Inflorescences sessile, fasciculate, the fascicles
dense, the bracts numerous and forming a cushion in the leaf axils. Flowers
greenish-white or white, the pedicels up to 8 mm long, usually shorter and to 4-5

flora of panama (Family 128. Flacourtiaiceae) 119

mm long, articulated close to the base, minutely pubescent; calyx 5-7 mm long, the
lobes rarely 6, narrowly ovate-oblong, united at the base into a tube 1-1.5 mm long,
acute to obtuse at the apex, 3-nerved, minutely puberulous without, minutely
puberulous to nearly glabrous within; stamens (9)10(11), the filaments filiform,
slightly unequal, 1.5-3.5 mm long, glabrous, united at the base among themselves
and with the staminodes into a very short tube less than 0.5 mm long, the anthers
ca 0.5-0.7 mm long, usually inconspicuously mucronate and often with 1 or few
whitish setae at the apex; staminodes (9)10(11), interstaminal, narrowly obovate,
1-2 mm long, densely whitish-pilose; gynoecium 3.5-5.5 mm long, the ovary nar-
rowly ovoid, ca 0.8-1.2 mm in diam at the base, densely whitish-pilose, gradually
attenuate into the style, the latter pilose at base to glabrous at apex, the stigma
capitate and densely papillate. Capsule ± globose, mucronate (base of style), 1-2
cm in diam, yellow at maturity, blackish when dry, glabrous except the slightly
pubescent apex; seeds conglutinate, to 5 mm long, the aril reddish or pinkish.

Southern Mexico, Central America, and tropical South America; known in
Panama as pica lengua (fide Pittier 5142) and raspa lengua (fide Standtey 28149).

bocas del toro: vie of Chiriqui Lagoon, von Wedel 1052 (MO, US), 1322 (MO, US);
Changuinola Valley, Dunlap 216 (F, US); banks of Changuinola River, Dunlap 518 (F,
US), canal zone: vie W end Gatun Lake dam, Blum & Tyson 1992 (MO); along drowned
Rio Axote Caballo, alt 6-70 m, in clumps in savanna, Dodge et al. 16847 (MO); Cruces,
Hayes 1019 (NY); Gamboa, moist thicket, Standley 28483 (US); vie of Miraflores, in
shaded swamps, White & Wh kite 96 (F, MO);

Balboa Heights, Greenman & Greenman 5023 (MO), chiriqui: vie of San Felix, alt 0-120
m, Pittier 5142 (F, NY, US), cocle: Aguadulce, nr sea level, in savannas, Pittier 4939
(US); 4 mi W of Anton on 1 bf Blum 2594 (MO); El Valle, valley floor

& lower slopes along highway, Miller 1836 (US); vie of El Valle, alt 600-1000 m, Allen
1178 (F, MO, US), darien: Rio Tuira, betw Rio Punusa & Rio Mangle, Duke 14596
(MO), herrera: rd from La Avena to outskirts of Pese, alt ca 200 ft, Burch et al. 1319
(MO); vie of Ocu, hill above the cantera of Sr. Joaquin Carrizo, limestone area, much cut
over & browsed by animals, Stern et al. 1714 (MO), panama: Punta Paitilla, Piper 5422
(US); nr Punta Paitilla, moist thicket, common, Standley 26244 (US); vie of Bella Vista,
Piper 5381 (NY, US); vie of Juan Franco Race Track, nr Panama City, moist thicket,
common, Standley 27768 (US); Sabanas, N of Panama City, Bro. Paul 579 (US); nr Old
Panama, Bro. Heriberto 290 (F, NY, US) ; Nuevo San Francisco, in thicket, Standley 30727
ndley 26201 (US), 28149 (US), 30681 (US); nr the big
swamp E of the Rio Tocumen, wet forest, common, Standley 26556 (US); along rd betw
Panama City & Chepo, Dodge et al. 16649 (MO); Laguna de Portala, nr Chepo, alt 50 m,
Pittier 4770 (NY, US), veraguas: Cafiazas, Tyson 3645 (MO); 2 mi S of Canazas, Tyson
3730 (MO); hills W of Sona, alt ca 500 m, Allen 1039 (MO), province unknot, [< ,
Hayes 350 (NY), 603 (NY).

4. Casearia stjohnii Johnston, Sargentia 8: 213, 1949.

Tree or large shrub 3-12 m tall; branches often with stout, spreading, spinescent
twigs 1-3.5 cm long; branchlets minutely pubescent. Leaves with the petioles
slender, 5-15 mm long; blade obovate, obovate- elliptic or ± elliptic, acute at the
base, rounded-obtuse and acuminate at the apex, 6-12 cm long and 3-6.5 cm wide,
the acumen 4-7 mm long, chartaceous, the margins ± distinctly crenate, conspicu-
ously pellucid-punctate, the upper surface lustrous and glabrous, the lower surface
dull and inconspicuously appressed-pubescent mainly along the prominulous costa
and the 5-7 pairs of lateral veins. Inflorescences axillary, densely fasciculate, the
fascicles sessile, 8-15-flowered. Flowers greenish, the pedicels articulated below the

120

apex, 1-2 mm long, the bracts numerous, crowded, often longer than the pedicels,
ca 2-3 mm long, pallid-strigulose towards the apex, persistent; buds subglobose;
calyx 4.5-5.5 mm long, the lobes ovate-oblong, obtuse, 1.8-2.3 mm wide, scarcely
united at the base into a tube 0.5 mm long or less, minutely appressed-pubescent
without, glabrous or sparsely strigulose within; stamens 8(9), the filaments filiform,
1.5-2.5 mm long, glabrous, united at the base among themselves and with the
staminodes into a very short tube, the anthers oblong, ca 0.5-0.7 mm long; stami-
nodes 8(9), elongated-oblong, ca 1-2 mm long and 0.4-0.5 mm wide, conspicuously
white-pilose; gynoecium 3-3.5 mm long, pilose, the ovary ovoid, 1 mm in diam at
the base, gradually attenuate into a short style, the stigma conical-capitate. Fruit
unknown.

Known only from San Jose Island.

Panama: San Jose I, northern end of island, Erlanson 369 (holotype US, isotype NY),
274 (US).

5. Casearia guianensis (Aublet) Urban, Symb. Ant. 3: 322, 1902.

Shrub or small tree 1.5-6 m high, deciduous, without spines, the branches few,
elongate, the branchlets inconspicuously puberulous, soon glabrous. Leaves with
the petioles to 10 mm long, the stipules subulate, 2-5 mm long, minutely ferrugi-
nous-puberulous, soon caducous; blade obovate, acute at the base, acuminate at
the apex, the acumen rather blunt, serrulate-denticulate at the margins, to 18 cm
long and 7 cm broad, chartaceous, with numerous pellucid dots and lines, minutely
puberulous on both sides when young, the costa prominent and the lateral veins
prominulous below. Inflorescences sessile, densely fasciculate, axillary (but usually
on defoliated branchlets). Flowers white, the pedicels to 6 mm long, articulated
ca 2-2.5 mm above the base, minutely ferruginous-puberulous especially below the
articulation; calyx ca 5 mm long, the lobes ± oblong, scarcely united at the base,
obtuse and subcucullate at the apex, 1.5-1.8 mm broad, minutely puberulous;
stamens (7)8, united at the base with the staminodes into a very short tube, the
filaments subequal, ca 2.5 mm long, glabrous, the anthers oblong, ca 0.7-0.8 mm
long (sometimes inconspicuously apiculate?); staminodes (7)8, linear-spathulate,
ca y 2 as long as the filaments, villous; gynoecium ca 4-4.5 mm long, the ovary
pyriform, ca 1 mm in diam at the base, villous, gradually attenuate into the
glabrous style, the stigma conical-capitate. Capsule subglobose, with 6 longitudinal,
blunt ridges, 8-13 mm long, white or green outside when fresh, glabrous or nearly
so, the pericarp ± reticulated, splitting into 3 valves at maturity, these reddish
inside; seeds up to 13 per capsule, subovoid, 3-3.5 mm long, very minutely foveolate,
the aril orange-colored.

West Indies, Costa Rica, Panama, and northern South America; known col-
loquially in Panama as palo de la cruz (cf. Pittier 6551).

canal zone: Chagres, Fendler 192 (US); Barro Colorado I, forest along shore of Gatun
Lake, E of Laboratories, Killip 40020 (US); Balboa, thicket, Standley 32105 (US), darien:

J mogana, f
on hillside,

6557 (US), herrera: 12.5 mi S of Ocu,

ndary woods on hillside, Lewis et al. 1665 (MO). Panama: Bella Vista, Bro. Heriherto

flora of Panama (Family 128. Flacourtiaiceae) 121

224 (US); id., in thicket, Standley 25305 (US); low woods E of Bella Vista, along beach,
Maxon & Valentine 6929 (US); Sabanas, N of Panama City, Bro. Paul 606 (US); nr Tapia
River, Juan Diaz region, thicket at edge of forest, Maxon & Harvey 6717 (US); San Jose I,
Erlanson 94 (NY, US), 261 (NY, US), Johnston 1139 (MO, US).

This species is very closely related and perhaps even conspecific with C.
aculeata Jacq.

6. Casearia nitida (L.) Jacquin, Enum. Syst. PL Ins. Carib. 21, 1760; L. Wms.,

Fieldiana: Bot. 29: 359, 1961.
Samyda nitida L., Syst. Nat. ed. 10, 1025, 1759.
Casearia corymhosa H.B.K., Nov. Gen. Sp. PI. 5: 366, 1823.
C. banquitana Krause, Beih. Bot. Centralbl. 32, Abt. 2: 345, 1914.
C. laevis Standley, Contr. U. S. Nat. Herb. 23: 845, 1923.
C. banquitana var. laevis (Standley) Johnston, Sargentia 8: 211, 1949.

Shrub or small tree 1.5-7.5 m tall, the branchlets inconspicuously puberulous
to glabrous. Leaves short-petiolate, the petiole to 5(6) mm long, canaliculate
above, inconspicuously puberulous to glabrous; blade oblong-elliptic to oblong-
obovate, often narrowly so, the base oblique or not, ± rounded to subcordate or
sometimes obtuse, the apex usually bluntly short- acuminate, rarely long-acuminate
or obtuse or rounded (and subemarginate?) , the margins minutely serrulate to
crenulate-serrulate, to 16 cm long and 5.5 cm wide, membranous to chartaceous,
copiously provided with pellucid dots or lines, glabrous above, glabrous or incon-
spicuously puberulous beneath, the costa and lateral veins prominent beneath.
Inflorescences axillary, cymose-corymbiform, the axes glabrous to inconspicuously
puberulous. Flowers white, the pedicels to 6 mm long, articulated below the
middle, the bracteoles small and deltoid; calyx to 5 mm long, the lobes 5, united
at the very base, elliptic, rounded, erect-patent at anthesis, inconspicuously puberu-
lous; stamens 8, inserted at the apex of the calyx tube (or even above?), the
filaments subequal, 2.5-3 mm long, glabrous, slightly united at the base among
themselves and with the staminodes, the anthers oblong-ellipsoid, ca 0.7 mm long;
staminodes 8, interstaminal, linear-spathulate, to 1.5 mm long, pilose; gynoecium
ca 4 mm long, the ovary ovoid, ca 1 mm in diam at the base, rather sparsely pilose
as the style, attenuate into the style, the stigma capitate. Capsule globose to
ellipsoid, with 3, rarely 4, prominent longitudinal ridges, to 15 mm long and 11
mm in diam, the pericarp glabrous, yellowish-orange or reddish when mature and
fresh, blackish when dry; seeds 1-3, ovoid to angulate-ovoid, to 7(8) mm long,
the testa glabrous but covered with numerous dark resinous glands, the aril
incomplete.

Widely distributed in tropical America; known in Panama as comida de low
(fide Cooper cV Slater 14), mamar and maho (fide Duke 14461).

bocas del toro: Almirante region, Cooper & Slater 14 (US), canal zone: edge of
forest at roadside just beyond Fort Sherman Research Forest Site, Stimson 5224 (MO) ; rd
to Devils Beach, Johnston 1573 (MO); S of island (Barro Colorado I?), White 113 (F,
MO); Madden Dam, Dwyer & Hayden 3 (MO); vie of Miraflores Lake, White 143 (F,
MO); nr junction of Miraflores & Cocoli roads, White 87 (F, MO); Sosa Hill, Balboa
brushy slope, Standley 25257 (US); Farfan Beach area, Tyson 1828 (MO), chiriqui:
Progreso, Cooper & Slater 199 (F, NY, US), colon: Fato (Nombre de Dios), at sea level,
"""i (US), darien: Rio Ucurganti, Bristan 1186 (MO); Rio Chucunaque, betw Rio
& Rio Subuti, Duke 8585 (MO); Rio Chucunaque, betw Rio Membrillo & Rio

122 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Yaviza, Duke 8617 (MO), los santos: 1-2 mi W of Candelaria, Duke 12444 (MO); nr
Santo Domingo, Dwyer 2499 (MO); Monagre Beach thicket, Dwyer 4137 (MO); several
km SE of Pedasi, beside dirt rd just S of Rio Galdera nr Punta Mala, Stimson 5289 (MO).
Panama: Alhajuela, Dwyer 1141 (MO); around Alhajuela, Chagres Valley, forest on dry
alt 30-100 m, Pittier 3497 (US); nr Matias Hernandez, moist thicket, Standley
28922 (US); Agricultural Experiment Station at Matias Hernandez, Pittier 6724 (US);
betw Las Sabanas and Matias Hernandez, moist thicket, Standley 31908 (US); nr Tapia
River, Juan Diaz region, edge of forest, Maxon & Harvey 6625 (US) ; vie of Pacora, alt 0-20
m, Allen 3448 (F, MO, NY, US); betw Cafiazas & Sabalo, alt ca 100 m, Duke 14461
(MO); Taboga I, alt 0-250 m, Pittier 3540 (US), Woodson et al. 1522 (F, MO, NY); San
Jose I, in shade of forest, Erlanson 151 (US), rocky bluffs, Erlanson 159 (NY, US), 255
(US), 448 (US), 588 (NY, US), san blas: Mulatuppu, Duke 8531 (MO), veraguas:
Santiago, 12 mi from Santiago towards Divisa on Transisthmian Hwy, roadside thicket,
Dwyer & Kirkbride 7408 (MO).

7. Casearia sylvestris Sw., Fl. Ind. Occ. 752, 1798.

Samyda sylvestris (Sw.) Poiret in Lamarck, Encycl. Meth., Rot. 6: 492, 1805.

Shrub or small tree 2-10 m high, the trunk to 12 cm in diam, the branchlets
puberulous but soon glabrous. Leaves with the petioles to 10 mm long, slender,
ite above, minutely puberulous when young but soon glabrous, the
stipules minute, 1-1.5 mm long, caducous; blade narrowly oblong-ovate or narrowly
oblong-elliptic, sometimes narrowly ovate or elliptic, the base symmetrical or often
- asymmetrical and usually acute, the apex long-acuminate, the acumen
slender and up to 2.5 cm long, the margins entire or nearly so, to 14 cm long and
5 cm wide, membranous to thin-coriaceous, dark green, lustrous especially above,
pellucid-punctate, glabrous, the costa prominulous below. Inflorescences axillary,
sessile, densely fasciculate, the fascicules many-flowered, the bracts minute and
forming a cushion in the leaf axils. Flowers pale yellow or white, the pedicels
slender, to 5 mm long, articulated near or below the middle, puberulous to densely
puberulous below the articulation; calyx 2-3 mm long, the lobes 5, broadly ovate,
jnited at the base into an obtusely angulate tube 0.5-0.8 mm long, rounded, 1-1.5
mm wide, erect-patent at anthesis, minutely ciliolate, glabrous or sparsely puberu-
lous outside, sparsely puberulous to puberulous inside; stamens 10, inserted at the
apex of the calyx tube, the filaments slightly unequal, to 1.5 mm long, almost
glabrous to sparsely puberulous, slightly united at the base among themselves and
with the staminodes, the anthers didymous, subglobose, ca 0.3 mm long, with a
dorsal, apical gland; staminodes 10, ± intrastaminal and alternating with the
stamens, to 1-1.2 mm long, whitish-puberulous; gynoecium 2-2.2 mm long, the
ovary obtusely angulate, ca 0.7-1 mm in diam, glabrous or sparsely pilose at the
apex, attenuate into the style, the stigmas 3, capitellate, papillate. Capsule sur-
rounded by the persistent, patent calyx, ovoid-globose, 3-4 mm long, obtusely
angulate, apiculate, glabrous or nearly so, reddish or purple at maturity, splitting
into 3 valves; seeds 2-6, ± ovoid, to 2 mm long, the testa light brown, glabrous,
foveolate, the aril incomplete, reddish.

Throughout tropical America; known locally as corta lengua (Cooper & Slater
213).

bocas del toro: vie of Chiriquf Lagoon, Water Valley, von Wedel 1747 (MO, US);
Isla Colon, von Wedel 64 (MO), 2970 (MO, NY, US); region of Almirante, Buena Vista
Camp on Chiriqui Trail, alt 1250 ft, Cooper 587 (F, US), canal zone: Rio Chagres,

1968]

flora of panama (Family 128. Flacourtiaceae) 123

Fendler 186 (MO, US) ; forest along banks of Quebrada La Palma & Canon of Rio Chagres,
alt 70-80 m, Dodge bf Allen 17359 (MO); nr Fort Randolph, brushy slope, Standley 28701
(US); nr Coco Solo Weather Station, Duke 4286 (MO);Barro Colorado I, Bangham 408 pro
parte (F), 436 (F) . MO), 749 (MO), 1097 (F, MO), Woodworth & Vestal

429 (F); Cerro Gordo, nr Culebra, alt 50-290 m, Pitt,: 21 i (US) -'neon Hill, woods,
aui: Progreso, Cooper & Slater
213 (F, NY, US), cocle: Bismark above Penenome, Williams 557 (NY, US), c<
from Puerto Pilon on side of dirt rd to Maria Chiquita Beach, Correa 137 (MO); betw
France Field, Canal Zone, & Catival, wooded swamp, common, Standley 30169 (US); along
trail to triangulation station on top of Tumba Vieja, alt 90-400 m, Dodge 16759 (MO).
darien: El Real Quebrada Trapiche, Duke & Bristan 323 (MO), panama: La Chorrera,
Bro. Paul 498 (US); Juan Diaz, moist woods, common, Standley 30589 (US); I
moist woods, Standley 30679 (US); Rio Tocumen, wet forest, Standley 26733 (US); San
Jose I, in forest or edge of forest, Johnston 232 (MO, US), 345 (MO, US), 1334 (MO, US).
iAN blas: along the headwaters of the Rio Mulatupo, seasonal evergreen forest, Elias 1744
(MO), veraguas: Isla de Coiba (Colonia Penal), Dwyer 2316 (MO).

8. Casearia commersoniana Camb. in St.-Hil., Juss. & Camb., Fl. Bras. Merid.
2:235, 1830.— Fig. 9. ■

C. myriantha Turcz., Bull. Soc. Imp. Nat. Moscou 36(1, fasc. 2) : 609, 1863; Johnston,

Sargentia8:212, 1949.
C. javitensis Auct. non H.B.K.; Standley, Contr. U. S. Nat. Herb. 27: 274, 1928.

Shrub or small tree 3-10(-15) m tall, the trunk to 10-20 cm in diam, the bark
smooth, the branchlets glabrous or almost so. Leaves glabrous, short-petiolate, the
petiole to 5-8 mm long, the stipules linear-subulate, early caducous; blade narrowly
elliptic to narrowly ovate-elliptic to narrowly oblong, sometimes elliptic, acute at
the base, acuminate at the apex, the acumen usually long and rather blunt, re-
motely crenulate-serrulate to almost entire along the margins, to 25 cm long and
9 cm wide, rigid-chartaceous, lustrous especially above, dark green, without pellucid
dots, the costa and 5-8(-10) pairs of lateral veins prominent beneath. Inflorescences
sessile, fasciculate, the fascicles few- to many-flowered, the bracts small and form-
ing a cushion in the leaf axils. Flowers white or greenish, the pedicels 3-10 mm
long, articulated below the middle, puberulous; sepals 4-5, scarcely united at the
base, ovate, acute, unequal, ca 3-4 mm long, reflexed to erect-patent at anthesis,
puberulous especially outside, persistent or slightly accrescent; stamens 9-15, slightly
longer than the sepals, the filaments to 4.5 mm long, slightly unequal, glabrous,
the anthers ellipsoid, ca 0.5-0.7 mm long, eglandular; staminodes intrastaminal,
usually isomerous with the stamens, linear, ca 1.5-2 mm long, puberulous, especially
towards the apex; gynoecium ca 3.5-4.2 mm long, the ovary subglobose, ca 1-1.5
mm in diam, sparsely to ± densely hirsute, the style glabrous or nearly so, divided
at the apex into 3 branches to 0.8 mm long, the stigmas 3, capitate. Capsule sub-
globose, 3-angulate, apiculate, 5-10(-15) mm long, red or brownish outside when
fresh, blackish outside and yellowish brown inside when dry, sparsely appressed-
pilosc, thin-walled, splitting into 3 valves at mat rii I , subglobose, 4-5

mm long, the testa brown, whitish-pilose, the aril pale, incomplete, subentire,

Southern Mexico, Central America, and northern South America.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

' 2 )\ B, flower (X8);
amerC upper part of filament & anther (X20); E, gynoecium (X8); F, dehiscinj
X3). A-E after Tyson & Blum 3911 (MO); F after Dwyer 6837 (MO).

flora of panama (Family 128. Flacourticmeae) 125

bocas del toro: Almirante, just N of Dos Milla, hillside, McDaniel 5123 (MO).
canal zone: Chagres, Isthmus of Panama, Fendler 185 (type C. myriantha, MO, US);
headwaters Rio Arenal, Johnston 1501 (MO); Camp Pina, vie of Hill C-6, Ft. Sherman,
Duke 4406 (MO); Mount Hope Cemetery, moist thicket, Standley 28834 (US); Gatun,
Hayes 98 (NY), 606 (NY), 613 (NY); Barro Colorado I, Bangham 443 (F), 587 (F, US),
Shattuck 1151 (F, MO), Starry 289 (F); Gamboa, Stevens 1058 (US); Obispo, moist woods,
Standley 31759 (US); Ft. Clayton nr old hospital building #519, Tyson & Blum 3911
(MO); Cerro Galero, Rd K6, alt 1000 ft, Stern & Chambers 29 (F, MO, NY, US); s. loc;
Epplesheimer s.n. (F). chiriqui: vie of David, alt 30-80 m, along streamlet. ,'

(US); vie of San Felix, alt 0-120 m, Pittier 5256 (F, N betw France Field,

Canal Zone, & Catival, brushy slope, Standley 30322 (US); Loma de la Gloria, nr Fato

(Nombre de Dios), in forests, alt 10-104 m, Pittier 3854 (F, NY, US). DARitn , I 1 i,

Duke & Bristan 8295 (MO); Camp Esloganti, helipad in premontane rain fori

ft, Duke 15502 (MO). Panama: hills above Campana, alt 600-800 m, Allen & Alston 1863

(F, MO); Rio Tapfa, moist forest, Standley 28259 (US

US), 1347 (MO, US), san blas: Ailigandi, Duke 9318 (MO), Dwyer 6837 (MO).

Because of insufficient material, a few collections of Casearia remain unnamed:

(1) Hayes 3 (MO, US) = 899 (NY) (Canal Zone: nr Gatun): tree 10 m high,
the branchlets rusty-tomentose; leaves short-petiolate, the blade ovate to obovate,
discolorous, shiny and glabrous above, rusty-pubescent below; capsule ± globose,
apiculate, with 3 longitudinal, prominulous ridges, to 2.5 cm long, tawny-tomentel-
lous; this collection can perhaps be referred to the West Indian species Casearia
hirsuta Sw.

(2) Woodson et al. 987 (MO) (Chiriqui: vie of Casita Alto, Volcan de Chiriqui,
alt ca 1500-2000 m) ; tree 6 m high; leaves like in C. arguta; capsule to 3 cm long,
glabrous, the seeds conglutinate, to 12 mm long; this collection is related to
C. arguta, but the capsule and seeds are unusually large.

(3) Williams 646 (NY, US) (Darien: nr Marraganti): according to the field notes,
this is a plant 5 m high, with the stem ca 6.5 cm in diam, with thorns 2.5 cm
long; it is perhaps related to C. aculeata Jacquin, but the absence of stipules and
flowers makes positive determination difficult.

11. ZUELANIA
Zuelania A. Richard in Sagra, Hist. Fis. Pol. Nat. Isla Cuba 10: 33, 1841.

Trees or shrubs. Leaves alternate, petiolate, the stipules fugacious, the blade
entire to serrulate, penninerved, pellucid-punctate. Inflorescences axillary, fascicu-
late towards the apex of the branchlets. Flowers often precocious, g , the pedicels
bracteate at the base and articulated; sepals 4-5, imbricate, shortly united at the
base, persistent; petals wanting; stamens oo (20-40), subperigynous, the filaments
filiform, alternating and united basally with many clavate staminodes, the anthers
subbasifixed, introrse, longitudinally dehiscent; ovary free, 1-locular, the ovules oo
on 3 parietal placentas, the stigma sessile or subsessile, thick, peltate. Fruit a fleshy
capsule, globose, opening at length by 3-valves; seeds numerous, arillate; endosperm

Zuelania guidonia (Sw.) Britton & Millsp., Bahama Fl. 295, U
%etia guidonia Sw., Nov. Gen. Sp. PI. Prodr. 83, 1788.
<xelania roussoviae Pittier, Contr. U. S. Nat. Herb. 18: 163, pi 79, 1916.

OF THE MISSOURI BOTANICAL GARDEN

Fig. 10. Zuelania guidonia (Sw.) Britton & Millsp.: A, habit (Xi/ 2 )
(Xi/ 2 ); C, flower (X5); D, sepal, 4 stamens, and 5 staminodes (X5); E, stamen (XlO);
F, gynoecium (X6); G, fruit (XI). A-F after Johnston 613 (MO); G after Pittier 2710
(MO).

flora of panama (Family 128. Flacourtiaceae) 127

Tree, deciduous, 10-25 m high, the trunk 30-50 cm in diam at the base, the
crown rounded, the bark grayish, the branches sparsely verruculose, the branchlets
brownish-hirsute. Leaves often fasciculate towards the apex of the branchlets, the
petiole to 2 cm long, brownish-hirsute, the stipules narrowly ovate, 3-6 mm long,
appressed-pubescent; blade narrowly oblong to oblong to oblong-elliptical, oblique
and rounded or obtuse at the base, obtuse to acute to shortly obtusely acuminate,
serrulate to subentire along the margins, 6-25 cm long and 3-9 cm wide, chartace-
ous, discolorous, green above and grayish or slightly brownish beneath when fresh,
pellucid-punctate, pubescent on both sides but the indumentum denser beneath,
the costa and lateral veins prominent beneath. Inflorescences in globose fascicles
in generally defoliate axils at the apex of the branchlets, the fascicles to 15-
flowered. Flowers yellowish, the pedicels to 18 mm long, shortly fulvous-hirsute,
articulated below the middle, the bracts thin-scarious, to 4 mm long, short-hirsute
without; calyx to 6-7 mm long, the sepals ovate, rounded at the apex, deflexed at
anthesis, tomentose without except on the covered margins; stamens with the fila-
ments to 4 mm long, glabrous or sparsely pilosulose, the anthers ± ovate-oblong,
subemarginate at the base, obtuse at the apex, to 1.8 mm long; staminodes about
Y 2 as long as the filaments, glabrous or sparsely pilosulose; gynoecium to 4 mm
long, the ovary ovoid, ca 2.5-3 mm broad, densely tomentellous, the stigma sub-
sessile. Capsule baccate, ± globose to depressed-globose, shallowly 5-sulcate, up to

5 cm broad (5-10 cm broad ace. to Zetek Z-3343) and 3 cm high, yellowish-green
to dark green when fresh, pilose; seeds numerous, obovoid, angular, ca 4 mm long,
the aril orange.

Southern Mexico, Central America, and the West Indies; known colloquially
in Panama as cagajon (fide Aviles X-ll & Zetek Z-3343) or caranon (fide Pittier
6605).

canal zone: Chagres, Fendler 318 (MO); nr Fort Randolph, swampy woods, Standley
28665 (US); Barro Colorado I, Aviles X-ll (F), Shattuck 796 (F), Wilson 91 (F), Wood-
worth & Vestal 719 (F), Zetek Z-3343 (F), 3463 (F), belw Rio Grande

6 Pedro Vidal, on rd to Arraijan, alt 50-150 m, Pittier 2710 (holotype US, isotypes F, MO,
NY); rd K9D, alt 260 ft, Stern & Chambers 44 (F, ^ c:le: Penonome &
vie, alt 50-1000 ft, Williams 235 (US), darien: vie of La Palma, alt 0-50 m, Pittier 6605
(NY, US). Panama: Rio Tocumen, moist forest, Standley 26750 (US); San Jose I, Johnston
611 (MO, US), 613 (MO, US).

On San Jose Island, Johnston (Sargentia 8: 215, 1949) describes this tree as
follows: "The tall pallid trunk and the shallow wide crown give it a distinctive
appearance, especially so during the dry season, when it is leafless . . . When
freshly cut the thick bark is tan and marked with dark streaks, but it soon changes
to mustard color and begins oozing a clear, yellowish exudation. The twigs are
slightly reddish and roughened with somewhat protruding transverse lenticels . . .
The foliage is shed early in February and generally not renewed until late in the
dry season ... It [the fruit] is firm but somewhat juicy in texture, produces honey-
like exudations when bruised, and eventually splits into three broad diverging parts
to expose the seeds and the conspicuous orange placentas."

128 ANNALS OF THE MISSOURI BOTANICAL GARDEN

12. LAETIA
Laetia Loefl. ex L., Syst. Nat. ed. 10, 1074, 1373, 1759.

Shrubs or trees. Leaves alternate, distichous, petiolate, the stipules early cadu-
cous, the blade crenate or serrate, rarely entire-margined or nearly so, usually
pellucid-punctate, sometimes coriaceous and epunctate. Inflorescences axillary or
terminal, fasciculate or cymose. Flowers rather small, g , the pedicels articulated
above the base, the bracteoles small, the bracts and bracteoles sometimes united
into a cupule; sepals 4-5, imbricate, free or scarcely united at the base, somewhat
petaloid, reflexed or not at anthesis, deciduous; petals absent; stamens 10-20 or
oo, inserted on an obsolete disc, the filaments free, equal or not, the anthers small,
elliptic or linear, not appendaged, introrse, attached at or above the base, longitudi-
nally dehiscent; staminodes none; ovary free, 1-locular, with 3 parietal placentas,
the placentas few- to many-ovulate; style short, simple or shortly but distinctly 3-
lobed, the stigmas capitate or lobed. Fruit a berry-like capsule, tardily valvately
dehiscent, sometimes resinous within; seeds few to numerous, arillate, the testa
coriaceous; endosperm copious; embryo straight; cotyledons broad and foliaceous.

A neotropical genus of about 20 species; three species, at present, reported from
Panama and North America.

bb. Pedicels to 12 mm long, glabrous; sepals to 4 mm long, reflexed at anthesis,
glabrous; stamens 12-20, the anthers to 1.5 mm long; styel undivided; leaf
blades with the margins minutely appressed-serrulate to almost entire ...2. L. procera
aa. Inflorescences dichasial, few-flowered; sepals 4, 8-9 mm long; stamens co; style

undivided; leaf blades with the margins entire to indistinctly crenulate ....3. L. thamnia

1. Laetia micrantha A. Robyns, Ann. Missouri Bot. Gard. 54: 190, 1967.— Fig. 11.
Tree 18-20 m tall, the trunk 12.5-25 cm in diam, the wood hard, the branchlets
puberulous. Leaves pendent on branches, the petioles 5 mm long, puberulous-
villous, the stipules narrowly triangular, ca 4 mm long and 1 mm wide at the
base, fugacious; blade narrowly ovate-oblong to narrowly oblong, the base sub-
oblique to markedly oblique, obtuse to rounded on one side and acute to obtuse
on the other side, the apex long-acuminate, the acumen slender, the margins ser-
rulate, 6-13 cm long and 1.5-4.5 cm wide, membranous to thin-chartaceous, densely
pellucid-punctate, ± shining especially above, minutely puberulous on both sides,
the costa prominent and the lateral veins prominulous beneath. Inflorescences
axillary, densely fasciculate, the bracts numerous, small, ca 1 mm long, persistent,
forming cushions in the axils of the leaves. Flowers white, the pedicels filiform, to
4 mm long, articulated ca 1 mm above the base, minutely puberulous; sepals 5,
free or nearly so, ± obovate, slightly cucullate, rounded at the apex, ca 1.8-2 mm
long and 1-1.3 mm wide, membranous, erect at anthesis, minutely puberulous out-
side; stamens 10, exserted, uniseriate, the filaments filiform, ± equal, 2-2.5 mm
long, glabrous, the anthers ca 0.3 mm long; ovary subglobose, ca 0.7-0.8 mm in

(Family 128. Flacourtiaceae) 129

diam, glabrous, each placenta 2-ovulate, the style ca 1.3 mm long, shortly but
distinctly 3-fid at the apex, the branches each 0.2 mm long, the stigmas 3, slightly
enlarged. Capsule unknown.

Endemic in the Province of Darien; known colloquially as raspa lengua (fide
Duke & Bristan 238).

darien: Perrecenico River, Duke & Bristan 238 (MO); trail betw Paya & Palo de las
Letras, Stern el at de, in dense wood, Stern et al.

513 (holotype MO, isotype US).

Duke & Bristan (238) report that the fruit is edible and that the wood may be
used for firewood when dry.

i A. Robyns: A, habit (Xi/ 2 ); B, flower (X15); C, s
anther (ca X40); D, gynoecium (X15). After Stern

[Vol. 55
130 ANNALS OF THE MISSOURI BOTANICAL GARDEN

2. Laetia procera (Poeppig) Eich. in Mart, Fl. Bras. 13 (1): 453, 1871.
Samyda procera Poeppig in Poeppig & Endl., Nov. Gen. Sp. PI. 3: 67, 1845.
Guidona procera (Poeppig) O. Ktze., Rev. Gen. PI. 1 : 44, 1891.

Slender tree to 12 m tall or more, the trunk terete, the branchlets glabrous,
sometimes pruinose. Leaves with the petioles to 1 cm long, canaliculate above,
glabrous and sometimes pruinose, the stipules deltoid, 0.5-0.8 mm long, caducous;
blade narrowly oblong, sometimes slightly inequilateral, rounded to subcordate at
the base, long-acuminate at the apex, the acumen mostly blunt and to 12 mm long,
minutely appressed-serrulate to aJmost entire along the margins, to 16 cm long and
4.5 cm wide, chartaceous, densely pellucid-punctate, ± lustrous on both sides,
slightly paler beneath, glabrous, the costa and lateral veins prominent beneath.
Inflorescences supra-axillary, in sessile 15-30-flowered fascicles, the bracts numerous,
small, forming a cushion. Flowers white, glabrous, the pedicels slender, to 12 mm
long, articulated close to the base; sepals 5, obovate, rounded at the apex, to 4 mm
long and 2.5 mm wide, reflexed at anthesis, persistent; stamens 12-20, scarcely
perigynous, the filaments filiform, ± crispate, to 3 mm long, the anthers oblong,
to 1.5 mm long; ovary ovoid, attenuate at the apex into a short, undivided style,
the stigma capitate. Capsule globose-ellipsoid, 1.5-2 cm long, splitting into 3 valves
stellate-coherent at the base, the pericarp often wrinkled; seeds numerous, ovoid,
to 3.5 mm long, reticulate-foveolate, the aril white and fleshy.

Reported for the first time from Panama and North America; Guyanas; north-
ern Brazil.

canal zone: River rd nr Congo bridge, Johnston 1527 (MO); rd W of Pina Base
Camp, Johnston 1791 (MO).

3. Laetia thamnia L., Amoen. Acad. 5: 379, 1760.

Tree to 10 m high, the branchlets glabrous. Leaves with the petioles to 1 cm
long, caniliculate above, glabrous, the stipules broadly triangular, ca 1 mm long,
caducous; blade narrowly elliptic to elliptic, slightly inequilateral, the base oblique,
acute to obtuse on one side, ± rounded on the other side, the apex obtuse to
acuminate, the margins entire to indistinctly crenulate, to 12.5 cm long and 4 cm
wide, membranous to chartaceous, pellucid-punctate, glabrous or inconspicuously
puberulous when young, the costa prominent beneath. Inflorescences axillary,
dichasial, few-flowered, the peduncles inconspicuously puberulous to glabrous, the
bracts broadly triangular, ca 1 mm long; buds globose, minutely to inconspicuously
puberulous. Flowers white, the pedicels slender, 9- 13 mm long, articulated 2-3 mm
above the base, inconspicuously puberulous especially below the articulation to
glabrous; sepals 4, broadly elliptic or obovate, rounded at the apex, 8-9 mm long
and 5-7 mm wide, reflexed at anthesis, minutely puberulous to glabrescent, cadu-
cous; stamens oo, the filaments filiform, unequal, 2.5-4.5 mm long, finely and softly
puberulous, the anthers broadly elliptic, ca 0.5-0.7 mm long; gynoecium ca 4-5 mm
long, the ovary broadly ovoid, to 3 mm in diam, tomentellous, attenuate at the
apex into a short, undivided style, the stigma capitate. Capsule globose or nearly
so, to 3 cm in diam, densely and minutely ferruginous-tomentellous, the pericarp
to 5 mm thick and fleshy, many-scaled.

flora of panama (Family 128. Flacourtiaceae) 131

Southern Mexico, Central America, Colombia, and the West Indies; known
locally as conejo (fide Pittier 3514).

canal zone: Chagres, Fendler 106 (MO, US); betw Chagres Batteries & Fort San
Lorenzo, Fort Sherman Military Reservation, Maxon & Valentine 6969 (F, NY); around
Alhajuela, Chagres Valley, alt 30-100 m, ,r 3514 (F, NY);

Barro Colorado I, Bailey & Bailey 504 (F), Shattuck 1096 (F, MO, US). Panama: along
Rio Juan Diaz, alt ca 30 m, Allen 932 (MO).

The following sterile collections probably belong to this species: Zetek 3687 (F, MO),
3906 (F, MO), 4999 (F), all from Barro Colorado I, Canal Zone.

13. RYANIA
Ryania Vahl, Eclog. Amer. 1: 51, pi. 9, 1796, nom. gen. conserv.
Patrisia L. C. Richard, Act. Soc. Hist. Nat. Paris 1: 110, 1792.

Trees or shrubs, the wood hard, the indumentum mostly stellate. Leaves alter-
nate, distichous, shortly petiolate, the stipules 2, glandular near the base within,
deciduous; blade equilateral or subasymmetrical, the margins entire to irregularly
denticulate-serrate, membranous to coriaceous, epunctate. Flowers often showy,
axillary, solitary to 4-fasciculate, £ , heterostylous, pedicellate, the pedicels articu-
lated and bracteolate at the base; sepals 5, quincuncial, nearly free, petaloid,
spreading at maturity, erect after anthesis, deciduous or persistent; petals none;
stamens 30-70, inserted in 2-3 series at the apex of the very short calyx tube, free
or nearly so, subequal, the anthers attached above the base, oblong to linear, to
9 mm long, often conspicuously mucronate at the apex, introrse, longitudinally
dehiscent; pollen smooth, with a single longitudinal colpus; disc coroniform,
urceolate; ovary superior, sessile or manifestly stipitate, 1-locular, with 3-9 parietal
placentas; ovules oo, many-seriate, anatropous; style short to long, entire or 3-9-fid
at apex, the stigmas capitellate. Fruits capsular, ultimately valvately dehiscent, 1-
locular, many-seeded; seeds small, arillate, hispidulous with scattered stellate hairs;
endosperm copious; embryo straight; cotyledons flat and thin.

A neotropical genus of 9 species distributed in Panama, Trinidad and South
America; one variety of the polymorphic R. speciosa reported from Panama.

Useful reference:

Monachino, J., A revision of Ryania (Flacourtiaceae). Lloydia 12: 1-29, 1949.

Probably all the species of Ryania are poisonous. All parts of the plant are to
a greater or lesser extent deadly, the concentrated alkaloid acting as a violent
stomach poison on both warm and cold blooded animals. According to Monachino
(loc. cit. p. 5) the Indians of the Amazon have used R. angustifolia to poison
alligators. Derivatives of Ryania (e.g. R. speciosa) are used as insecticides and "it
is said to be as good and possibly better than DDT against the European Corn
Borer, promising against the sugar cane borer and the Oriental fruit moth, and
effective control of the soybean caterpillar." (Monachino, loc. cit).

1. Ryania speciosa Vahl var. panamensis Monachino, Lloydia 12: 18, 1949. —

Fig. 12.

Shrub or tree 1-12 m tall, the branches elongated, the branchlets minutely
stellate-tomentose. Leaves with the petioles to 0.5(1) cm long, canaliculate above,
densely and minutely stellate-tomentose, the stipules small, subulate, ca 3-7 mm

ANNALS OF THE MISSOURI

. Ryania speciosa Vahl var. panamensis Monachino: A, habit (X'/ 2 );
flower (XI); C, section of sepal showing the stellate indument (X2); "~
part of filament and anther (X5); E, id., lateral view (X5); F, gynoecium (X2); G,
dehiscent capsule surrounded by the persistent calyx (Xl). A-F after Stevens 1251 (US);
G, after von Wedel 1947 (MO).

long, deciduous; blade narrowly elliptic to slightly obovate, slightly oblique or not
and mostly rounded to sometimes obtuse at the base, caudate to long-caudate-
acuminate at the apex, to 21 cm long and 7 cm wide, chartaceous, the acumen to
2.5 cm long, the margins entire, the upper surface ± shining and glabrous or nearly
so (midvein often stellate-puberulous near the base), the lower surface sparsely
and minutely stellate-puberulous to glabrous except for the stellate-tomentellous to

1968]

flora of panama (Family 128. Flacourtiaceae) 133

stellate-puberulous prominent costa and lateral nerves. Flowers white or light
green, solitary to 2 (-3) -fasciculate, the pedicels to 8 mm long, disarticulating at the
base, minutely stellate-tomentose, the bracteoles 3.5-5 mm long, subulate, caducous;
buds angulate-ovate; sepals narrowly ovate, acute, scarcely united at the base, 1.3-5
cm long and 0.4-0.8 cm wide, minutely stellate-tomentose to stellate-puberulous
outside, minutely and softly stellate-tomentellous inside, persistent; stamens 2-seri-
ate, the filaments filiform, free, subequal, to 2.5 cm long, glabrous or the lower-
most part pilose, the anthers 2.5-4 mm long, mucronate; disc ca 2 mm high (teeth
included), the teeth up to 1 mm long, barbate-villose; ovary sessile or nearly so,
ovoid to ellipsoid, densely appressed-fulvous-sericeous, the style up to 1.7 cm long,
appressed-sericeous at the base to glabrescent or glabrate at the apex. Capsule bac-
cate, surrounded by the persistent calyx, ± globose, apiculate (persistent base of
style) , to 2.5 cm or more in diam, the exocarp thick, suberose, scabridulous and fer-
rugineous-stellate-tomentellous outside, the endocarp crustaceous; seeds ovoid, ±
bluntly angulate, ca 5 mm long, the testa with scattered stellate hairs, the aril uni-
lateral, membranous.

Known at present only from the Province of Bocas del Toro, the Comarca de
San Bias, and from the Canal Zone.

bocas del toro: vie of Chiriqui Lagoon, Big Bight, von Wedel 2886 (holotype NY;
isotypes MO, US); id., Old Bank I, von Wedel 1928 (MO, US), 1947 (MO), 2119 A (MO,
US); Laguna de Chiriqui & vie, Hart 31 on Wedel 306 (MO), canal

zone: Fort Lorenzo Trail, Stevens 1251 (US); along Pavon Road, forest understory,
Johnston 1537 (MO); Santa Rita hills, cut-over secondary forest, Smith & Smith 3438 (F).
colon: E Santa Rita Ridge, Correa & Dressier 647 (MO), san blas: 2-5 mi S of Mandinga
airport, trail E of Cangandi-Mandinga airport road, Duke 14766 (MO).

The only significant distinction between the above variety and the var. speciosa
resides in the length of the anthers. In var. speciosa they are 5-7 mm long, while
in var. panamensis they are only 2.5-4 mm long. Study of additional material is
needed in order to evaluate this characteristic properly.

14. LUNANIA
Lunania W. J. Hooker, Lond. Jour. Bot. 3: 317, 1844, nom. gen. conserv.

Shrubs or trees. Leaves alternate, petiolate, estipulate, the blade entire-
margined or nearly so, 3- (5-) nerved from the base, with or without scattered pel-
lucid dots. Inflorescences axillary or terminal, in simple or branched racemes, the
pedicels short and articulated at the base, the bracts and bracteoles minute.
Flowers £ , small; calyx subglobose, at length valvately split to the base into
2-3(-5) spreading or reflexed membranous lobes; petals none; stamens 6-12, inserted
around a cupular hypogynous disk and alternating with its lobes, the filaments
shorter to longer than the anthers, the anthers basifixed, 2-thecate, longitudinally
and extrorsely dehiscent; ovary free, 1-locular, the ovules oo, multiseriate on 3
broad, parietal placentas; styles 3, free to the base or not, short to elongate, the
stigmas terminal. Capsules coriaceous, 3-valvate, few- to many-seeded; seeds small,
the testa usually punctate, arillate; endosperm present.

A neotropical genus of probably less than 20 species; two species, represented
by only four collections, reported at present from Panama.

[Vol. 55
!4 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Calyx splitting into 2-3 lobes, these reflexed and curled after anthesis; filaments
very short and ca 0.2-0.3 mm long; anthers to 1.5 mm long; disc and ovary
tomentellous - - 1. L. parviflora

.2. L. piperoides

1. Lunania parviflora Spruce ex Bentham, Jour. Proc. Linn. Soc., Bot. 5 (Suppl.

2): 90, 1861.— Fig. 13.
L. cuspidata Warburg in Engler & Prantl, Nat. Pflanzenfam. 3 (6a): 47, 1893.
Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 722, 1937.

Tree to 20 m tall, the branchlets glabrous or inconspicuously puberulous.
Leaves with the petioles to 2.2 cm long; blade inequilateral, ovate to elliptic, or
ovate-oblong, sometimes narrowly so, oblique and acute to rounded at the base,
long-caudate-acuminate at the apex, the acumen to 3 cm long and blunt, the
margins entire or nearly so, to 19 cm long and 7.5 cm wide, thin-chartaceous to
chartaceous, somewhat lustrous above, dull beneath, 3- to 4- to 5-nerved from the
base, glabrous above, inconspicuously puberulous and hirtellous along the main
veins to glabrous below, the main veins somewhat impressed above and prominent
below, the veinlets scarcely prominulous and loosely reticulate on both sides.
Racemes pendulous, to 40 cm long, the rachis inconspicuously puberulous. Flowers
yellowish-green to cream-colored, the pedicels 1.5-2 mm long; buds subglobose,
inconspicuously puberulous; calyx splitting into 2-3 lobes, these 1.5-2 mm long,
reflexed and curled after anthesis, long persistent; stamens 10-12, the filaments
very short and ca 0.2-0.3 mm long, glabrous, the anthers narrowly deltoid, to 1.5
mm long, apiculate; disc ca 0.5 mm high, tomentellous; ovary ca 1 mm long,
tomentellous, the styles 3, to 1 mm long, at first close together, later on spreading,
glabrous, the stigmas papillate. Capsule when young puberulous, mature one not

Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia, and Amazonian
Brazil.

san blas: Ailigandi River, Duke & Bristan 353 (MO) ; headwaters of Rio Cuadi, Camp
Diablo (Drill Site 22), N. 82.2, E. 87.8, alt 273.4 ft, seasonal evergreen forest along river,
Duke et al. 3609 (MO).

2. Lunania piperoides Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 4: 317,

Shrub to medium-sized tree to 20 m tall, the branchlets minutely puberulous
and hirtellous, glabrescent; blade somewhat inequilateral, ovate to oblong-ovate,
sometimes narrowly so, rounded to ± cuneate at the base, long-caudate-acuminate
at the apex, the acumen to 2.5 cm long and blunt, the margins entire to somewhat
undulate, to 13.5 cm long and 5.5 cm wide, thin-chartaceous to chartaceous, 3-
nerved from the base, somewhat shining on both sides, glabrous above, minutely
puberulous and hirtellous along the main veins to glabrous beneath, the main veins
slightly impressed above and prominent beneath, the veinlets slightly prominulous
and loosely reticulate on both sides. Inflorescences of branched racemes, the racemes
to 10 cm long, the rachis densely and minutely puberulous. Flowers greenish, the
pedicels to 1.5 mm long, the bracts ca 0.8 mm long, the bracteoles ca 0.5 mm long,

flora of Panama (Family 128. Flaeourtiaceae) 135

arviflora Spruce
[., later stage ot development, th
(X6). After Duke et al. 3609 (MO).

A, habit (Xi/ 2 ); B, flower (ca X10);
'""" "" (ca XlO); D, young capsule

[Vol. 55
136 ANNALS OF THE MISSOURI BOTANICAL GARDEN

persistent; calyx splitting into 2 spreading, cucullate, glabrous lobes to 2 mm long;
stamens 10, the filaments to 1.5 (-3) mm long, the anthers subglobose, ca 0.5 mm
long, the disc ca 0.5 mm high, glabrous; ovary glabrous, the styles 3, somewhat
flabellate towards the apex, glabrous. Fruit (not seen, ace. to original description)
subglobose, 5 mm long, the pericarp smooth, pale red, and glabrous; seeds numer-
ous, embedded in pulp, ovoid, 1 mm long.

Honduras and Panama.

bocas del toro: vie of Chiriqui Lagoon, Fish Creek, mountains, von Wedel 2396 (MO,
US), cocle: El Valle de Anton, forest behind Club Campestre, alt ca 700 m, Duke 13248
(MO).

15. XYLOSMA
Xylosma G. Foster, Fl. Ins. Austr. Prodr. 72, 1786, nom. gen. conserv.
Myroxylon J. R. & G. Foster, Char. Gen. PI. 125, 1776, non L. f. (Suppl. 34, 233, 1781).

Shrubs or small trees, often with axillary spines, the branchlets commonly
lenticellate. Leaves alternate, sometimes borne in fascicles, usually short-petiolate,
estipulate, the blade often ± coriaceous, usually glandular-dentate, penninerved,
rarely entire-margined, without pellucid-glands. Inflorescences axillary, fasciculate
or contracted-racemose, rarely racemose. Flowers small, dioecious or rarely polyga-
mous; pedicels articulated above the base, the bracts minute; sepals 4-5(-6), imbri-
cate, usually scale-like, slightly connate at the base, often ciliolate along the mar-
gins, usually persistent; petals none; stamens oo (8-35 in Panamanian spp.), usually
surrounded by an annular or glandular, fleshy disc, the filaments free, filiform,
short- to usually long-exserted, the anthers minute, basifixed, extrose, longitudinally
dehiscent; ovary sessile, inserted on an annular disc, 1-locular, with 2-3, rarely
4-6, parietal placentas, each placenta with 2, sometimes 4-6, ovules, the style entire
or ± divided, sometimes very short, the stigmas scarcely dilated to dilated; rudi-
mentary ovary wanting in c? flowers. Fruits baccate, rather dry, indehiscent,
surmounted by the persistent style, the pericarp rather thin-coriaceous, the seeds
2-8, ± angular by mutual pressure, the testa thin; endosperm copious; embryo
large, the coyledons broad.

About 100 species throughout the tropics, but absent in Africa.

In the absence of a comprehensive revision of the genus as a whole, it is
impossible to give a satisfactory treatment of the genus in a regional flora. The
specific delimitations within the genus are extremely difficult and neither the
flowers nor the fruits yield good distinctive characters. All the keys in the
regional tropical American floras are based mainly on vegetative characters, and I
have been forced, alas, to do the same in this flora. It is consequently obvious that
my treatment of Xylosma is only provisional.

Nine species are listed in the following keys, four species without specific name.
In view of the existing confusion within the genus I am reluctant to describe any
new taxon and I leave the evaluation of the given characters of the unnamed
species to the monographer-to-be of the genus!

The bark contains tannin and the fruit is sometimes the source of dyestuff;
the wood is moderately hard and rather heavy.

flora of Panama (Family 128. Flacourtiaceae)

Inflorescences fasciculate or contracted-racemose,
b. Leaves, at least most of them, borne in fascicles ;
pseudo-branches

closely.

rounded serrations, theii

of the blade, the network of the

a 20-25; along the Atlantic Coast
2. X.

Xfie,

gg. Leaves long-acuminate at the apex, the i

venation prominulous on both sides; stamens ca 35; style

4-5 lobed; Prov of Panama, lowlands 6. Xylosma sp. 1

ee. Stamens 8-12; Prov of Code, at ca 500 m alt 7. Xylosma sp. 2

dd. Flowers polygamous; Prov of Code, at sea level 8. Xylosma sp. 3

, Leaves hispidulous or minutely hirtellous below,
h. Branchlets and petioles densely short-hispid; leaf blades hispidulous
below; petioles 2-4 mm long; Prov of Bocas del Toro, at sea level

X. hispidula

1. Xylosma anisophylla Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 135,
1932.

Shrub or small tree 2-7 m tall, glabrous, the branchlets often armed with
spines, these to 4.5 cm long. Leaves, at least most of them, borne in fascicles at the
end of very short pseudo-branches, subsessile to very shortly petiolate, the petiole
to 4 mm long; blade usually narrowly obovate to obovate, infrequently elliptic,
cuneate-attenuate at the base, obtuse to rounded at the apex, remotely and obtusely
glandular-serrulate along the margins, the serrations inflexed towards the margin
and with their tips covering slightly the margin, to 6 cm long and 3.5 cm wide,
chartaceous to rigid-chartaceous, slightly lustrous above, dull beneath, the costa
prominulous beneath, the venation reticulate and inconspicuous on both sides.
Flowers fasciculate, dioecious, yellow, with slender, glabrous pedicels to 8 mm long;
sepals 4, triangular-ovate, acute, to 1.5 mm long, ciliolate along the margins,
glabrous; stamens ca 20, the filaments slender, to 2.5 mm long before dehiscence,
to 4 mm long after dehiscence. Bacca subglobose, 5-6 mm long, glabrous, sur-
mounted by the persistent, 2-3-lobed style, the stigmatic lobes spreading; seeds
(ace. to Standley, loc. cit. 136) 5, ferrugineous.

Mexico, British Honduras, and Panama.

[Vol. 55
138 ANNALS OF THE MISSOURI BOTANICAL GARDEN

moist forest, Standley 28256 (US); nr Matias Hernandez, moist thicket, Standley 28873
(US); Nuevo San ket, Standley 30739 (US); betw Pacora & Chepo, boggy

grasslands & marginal thickets, alt ca 25 m, Woodson et al. 1667 (F, MO).

2. Xylosma panamensis Turcz., Bull. Soc. Imp. Nat. Moscou 36 (1, fasc. 2): 554,
1863.

i 44, 1891.

Shrub or small tree 2-10 m high, the trunk up to 20 cm in diam, often spiny,
the axillary spines on twigs sharp and up to 4 cm long, the branchlets inconspicu-
ously puberulous to glabrous. Leaves subsessile to short-petiolate, the petiole to
6 mm long; blade polymorphous, typically elliptic to broadly elliptic (sometimes
± subrotund) and rounded on both ends, sometimes ovate or even narrowly ovate
and obtuse to acute at the apex, the margins remotely appressed-glandular-serrulate,
the serrations rounded, inflexed towards the margin and with their tips slightly
overlapping the margin, to 12 cm long and 8 cm wide, rigid-chartaceous to sub-
coriaceous, ± lustrous above and dull beneath, the costa prominent beneath, the
reticulated venation inconspicuous on either side, glabrous. Flowers in fascicles,
dioecious, yellowish, the pedicels slender, to 6 mm long, minutely puberulous,
bracteate at the base; sepals 4-5, subrotund to ovate, ca 1.2-2 mm long, ciliolate
along the margins, minutely puberulous externally; stamens ca 20-25, the filaments
short- to long-exserted and to 2-3 mm long. Bacca subglobose, to 6 mm long, at
first green, turning red, and finally black at maturity, glabrous, surmounted by the
persistent, 2-3-lobed style, the stigmatic lobes spreading; seeds 2-5, angulate-ovoid,
the testa reddish-brown, glabrous.

Collected only along the Atlantic Coast in Panama; called colloquially
canirico (cf. Duke 8480) and jobo de lagarto (cf. Pittier 4111).

The species is allied to the X. flexuosa complex, but can be distinguished by
the leaves usually broadly rounded at the apex, by the small, sometimes even
inconspicuous, closely appressed, rounded serrations (small auricles) with their
tips slightly overlapping the blade, and with the network of the venation incon-
spicuous on the upper surface.

bocas del toro: region of Almirante, Flat Rock, along beach & in second growth,
Cooper 548 (F, NY, US); Changuinola Valley, Bar Mouth, Dunlap 512 (F, US), 532
J i '\ Ma Colon, alt 0-120 m, von Wedel 499 (MO), 2479 (MO, NY, US), 2823 (MO,
NY, US), 2850 (MO, NY, US); vie of Chiriquf Lagoon, von Wedel 1361 (MO); id., Water
Valley, von Wedel 1516 (MO, US), canal zone: Fort Sherman, front of army barracks,
on beach, Dwyer & Robyns 153 (MO); Chagres, Fendler 194 (type F, MO, US); rd S
of Fort Sherman, edge of mangrove swamp, Johnston 1650 (MO); vie of Fort Sherman,
thicket along beach, Standley 31142 (US); Fort Sherman on main post, Tyson 2238 (MO),
Tyson & Blum 3771. colon: Aspinwall, Hayes 885 (NY); Colon, Kuntze 1884 (F, NY);
mouth of Rio Piedras, beach & adjacent area, Lewis et al. 3175 (MO); vie of Viento Frio,
along beach & nr sea level, Pittier 4111 (MO); betw France Field, Canal Zone, & Catival,
brushy slope, Standley 30430 (US), san blas: Mulatuppu, Duke 8480 (MO), 8546 (MO);
Isla Soskatupo, Duke 8512 (MO); Nargana I, nr Airport, Tyson & Dwyer 1172 (MO).
veraguas: mouth of Rio Concepcion, beach, cliffs & adjacent swamp, Lewis et al. 2823
(MO), 2828 (MO), province unknown (Canal Zone or Colon) : s. loc, Hayes 644 (NY).

flora of Panama (Family 128. Flucourtiaceae)

Fig. 14. Xylosma panamensis Turcz.: A, habit (Xi/ 2 ); B, leaf margin, upper surf;
(much enlarged); C, flower (X8); D, bacca surmounted by the persistent style (X6).
B & D after Tyson & Dwyer 1172 (MO); C after von Wedel 2479 (MO).

140 ANNALS OF THE MISSOURI BOTANICAL GARDEN

3. Xylosma flexuosa (H.B.K.) Hemsley, Biol. Centr.-Amer. 1: 57, 1879, sensu

lato.
Flacourtia flexuosa H.B.K, Nov. Gen. Sp. PI. 7: 239, 1825.
Hisingera nitida sensu Seemann, Bot. Voy. Herald 249, 1854.
H. elliptica Clos, Ann. Sci. Nat., Bot., ser. 4, 8: 226, 1857.
Myroxylon ellipticum (Clos) O. Ktze., Rev. Gen. PL 1: 44, 1891.
Xylosma hemsleyana Standley, Jour. Wash. Acad. Sci. 17: 169, 1927.

Shrub or small tree, 2-12 m high, the trunk to 10-12.5 cm in diam, armed or
not, the spines on the trunk often branched, the spines on the branches simple,
the branchlets inconspicuously puberulous to glabrous. Leaves short-petiolate, the
petiole to 5(-7) mm long; blade polymorphic, glabrous, commonly elliptic to
broadly elliptic, infrequently ovate (or even narrowly ovate), acute at the base,
obtuse to acute at the apex, seldom obtusely acuminate, distinctly glandular-
crenate-serrate along the margins, to 6 (-8) cm long and 4.5 cm wide, rigid-
chartaceous, ± lustrous and with the network of the veins conspicuous on the
upper surface, rather dull and with the costa prominent below, the network of the
veins less conspicuous on the lower surface. Flowers in fascicles or contracted
racemes, dioecious, greenish to yellowish, the pedicels to 5 mm long, minutely
puberulous or glabrous, bracteate at the base; sepals 4 or usually 5, broadly ovate
or deltoid, acute, ca 1.5-2 mm long, spreading in tf flowers, ciliolate along the
margins, usually minutely puberulous on both sides; stamens ca 16-20, the
filaments to 2.5 mm long, glabrous; ovary oblong-ovoid, glabrous, the placentas 2-3,
each one 2-ovulate. Bacca subglobose, 5-6 mm in diam, at first green, turning
bright red at maturity, surmounted by the persistent, 2-3-lobed style, the fruiting
pedicel elongated and to 8 mm long, usually distinctly articulate ca 2 mm above
the base; seeds 4-6, angulate-ovoid, to 4 mm long, the testa reddish-brown, glabrous.

Mexico, Central America, and northern South America; in Panama, this species
is found only in the Province of Chiriqui at 900-2250 m altitude; known in Panama
as cachito (cf. Stern et al. 2029) and roseto (cf. Pittier 3330).

Xylosma flexuosa, treated here in a rather broad sense, can be easily distin-
guished from X. panamensis by the leaf blades distinctly crenate-serrate and by the
network of the veins clearly noticeable when dry on the upper surface. Further-
more, the mature berries are said to be bright red in X. flexuosa, while they are
black in X. panamensis. Finally, the habitat of the two species is quite different.

chiriqui: llanos on slopes of Volcan de Chiriqui Viejo & along Rio Chiriqui Viejo,
alt 1200 m, Allen 969 (F. MO); Boquete, Boquete Distr, alt 3800-5500 ft, Davidson 615
(F, MO, US), 736 (F, MO, US), 779 (F, MO, US), savanna, 787 (F); vie of Boquete,
from Boquete to 3 mi N, alt 3300-4200 ft, second growth, cultivated areas & roadsides,
Lewis et al. 577 (MO); Bajo Boquete, alt 900-1300 m 3 Bro. Maurice 736 (MO), Pittier 3330
(US); vie of El Boquete, Maxon 5141 (US); vie of Boquete, Finca Collins, alt 5800-6700
ft, Stern et al. 2029 (MO); Camiseta, Volcan de Chiriqui, alt 7500 ft, Terry 1360 (MO);
Rio Chiriqui Viejo Valley, on llanos, White 56 (MO), 109 (MO); Finca Lerida to Boquete,
alt ca 1300-1700 m, Woodson et al 1104 (F, MO), 1168 (F, MO).

4. Xylosma hispidula Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 724,

1937.

Shrub or small tree, the trunk provided with numerous, long, slender, compound
spines, the branchlets slender, densely short-hispid, the hairs brownish, the axillary

1968]

flora of Panama (Family 128. Flacourtiaiceae) 141

spines to 2.5 cm long. Leaves very short-petiolate, the petiole 2-4 mm long, densely
short-hispid; blade narrowly elliptic to elliptic, obtuse at the base, acuminate to
caudate-acuminate at the apex, the acumen from acute to obtuse, the margins
obtusely serrulate, the serrations glandular on the lower side, to 9 cm long and
3.5 cm wide, rigid-chartaceous, glabrous or nearly so above, hispidulous below
especially along the slightly prominent costa and prominulous lateral- veins.
Inflorescences axillary, fasciculate, up to 10-flowered, bracteate at the base. Flowers
dioecious (or polygamous?). Male flowers not seen. Female flowers (young fruits)
(or hermaphrodite flowers?) with slender hispidulous pedicels to 5 mm long; sepals

4. subcircular, rounded apically, 1-1.5 mm long, the margins densely ciliolate,
glabrous or nearly so dorsally, pilose inside. Bacca (juvenile) globose-ovoid, ca 4
mm long, glabrous, surmounted by the 2 persistent, subsessile, spreading stigmas.

Known only from one collection from Panama.

bocas del toro: Changuinola Valley, Spur 2, Dunlap 561 (holotype, F, isotype US).

There is some doubt whether the flowers are dioecious since one occasionally
observes what appears to be a filament of a stamen around juvenile berries and
within the glandular disc.

5. Xylosma intermedia (Seemann) Tr. & PL, Ann. Sci. Nat., Bot. ser. 4, 17: 100,

Hisingera intermedia J
Myroxylon intermediw

Shrub ca 3 m high, unarmed, the branchlets lenticellate. Leaves short-
petiolate; blade oblong-elliptic, narrowed towards the base, acuminate at the apex,
obtusely serrate along the margins, 10-12.5 cm long and 3.5-5 cm wide, coriaceous,
lustrous and glabrous on both surfaces. Inflorescences racemose, 8- 10-flowered, ca
3.5 cm long, the peduncles and pedicels pubescent. Flowers polygamous (originally
described as hermaphrodite), the calyx 4-merous, the sepals oblong, obtuse; stamens
25-30 or more, the filaments slender, somewhat longer than the sepals, glabrous,
the anthers globose; ovary ovoid, attenuate at the apex into a short, 3-lobed style,
the stigmatic lobes cuneate, the ovules 6. Bacca with 1-2 oblong seeds.

Known only from Panama.

chiriqui: nr village of San Lorenzo, Seemann 1623 (type K, not seen; photo NY).

As I have not seen the type, which is the only known collection of this species,
the description given here has been taken from the original publication by Seemann
and from the description given by Triana & Planchon. The species is remarkable
on account of the racemose inflorescences and the polygamous flowers.

6. Xylosma sp. 1

Shrub or small tree to 5 m tall, the trunk unarmed or with branched spines,
the branchlets unarmed or with small, axillary spines, glabrous. Leaves glabrous,
the petiole to 5 mm long; blade narrowly ovate, obtuse to acute at the base,
long-acuminate at the apex, obtusely glandular-crenate to -serrate along the mar-
gins, to 11.5 cm long and 4 cm wide, chartaceous to rigid-chartaceous, ± shiny on
both surfaces, the costa slightly prominent below, the network of the venation

[Vol. 55
142 ANNALS OF THE MISSOURI BOTANICAL GARDEN

prominulous on both sides. Flowers yellow, dioecious, in axillary fascicles, the
pedicels to 7 mm long, articulated in the lower i/ 3 (in <? flowers) or ± in the
middle or slightly above the middle (fruiting pedicels), inconspicuously puberu-
lous; sepals (in cf flowers) usually 4, very broadly ovate, obtuse, ca 2-2.5 mm long
and 2 mm wide, inconspicuously ciliolate or not along the margins, puberulous
especially at the apex externally; stamens ca 35, the filaments slender, long-exserted,
to 6 mm long, glabrous. Bacca ellipsoid to subglobose, ca 6 mm long, not sur-
rounded by the calyx (calyx apparently caducous), orange-pink, glabrous, sur-
mounted by the short, persistent, 4-5 lobed style, the lobes short and spreading;
seeds angulate-ovoid, to 4.5 mm long, the testa reddish-brown, glabrous.

Costa Rica and Panama.

canal zone: Barro Colorado I, Bailey & Bailey 484 (F), Woodworth & Vestal 419
(F); on rd from Chepo to El Llano, Tyson & Smith 4125 (MO).

This species has been erroneously determined as X. intermedia (Seemann) Tr.
& PL (cf. Standley, Contr. U. S. Nat. Herb. 27: 273, 1928). In the latter species
the inflorescences are racemose and the flowers are polygamous while in our species
the inflorescences are fasciculate and the flowers are dioecious.

The above species is close to X. excelsa Standley & L. Williams (Ceiba 1 : 245,
1951) from Costa Rica. The leaves of both species are very similar, but X. excelsa
differs from the above by the large size of the tree (to 25 m) and by the polygamous
flowers (own observation of type collection: Allen 5650 in F, MO, US).

7. Xylosma sp. 2

Small tree to 4 m tall, the branchlets unarmed, glabrous or inconspicuously
puberulous. Leaves glabrous, short-petiolate, the petiole to 2-3 mm long; blade
elliptic to broadly elliptic, acute at the base, short-acuminate at the apex, glandular-
serrulate along the margins, the serrations sometimes very small and ± obtuse,
to 6 cm long and 4 cm wide, chartaceous, the costa slightly prominent beneath,
the network of the venations slightly prominulous above. Flowers fasciculate,
dioecious. Male flowers yellowish, the pedicel to 3.5 mm long, inconspicuously
puberulous; sepals 4, ovate to oblong-ovate, acute or rounded apically, 1.5-2 mm
long and ca 1 mm wide, finely ciliolate along the margins, glabrous outside,
puberulous inside; glandular disc ca 0.5 mm high; stamens 8-12, the filaments to
2 mm long, glabrous. Female flowers and fruit not seen.

cocle: N rim of El Valle, Allen & Alston 1856 (F, MO, US).

8. Xylosma sp. 3

Shrub or tree (?), the branchlets glabrous, armed, the axillary spines 1-4 cm
long. Leaves glabrous, subsessile or the petiole to 2 mm long; blade elliptic, obtuse
to acute at the base, obtuse to rounded-obtuse at the apex, appressed-glandular-
serrulate along the margins, the serrations obtuse, inflexed towards and slightly
overlapping the margin, to 4 cm long and 2.5 cm wide, rigid-chartaceous, some-
what shiny especially above, the costa prominulous beneath, the network of
the venation slightly prominulous on both sides. Flowers fasciculate, polygamous.
Male flowers not seen. Bacca subglobose, 5-6 mm in diam, glabrous, surmounted

flora of panama (Family 128. Flacourtiaceae) 143

by the persistent, deeply 2-lobed style, the fruiting pedicel to 8 mm long, glabrous,
the persistent sepals 4, deltoid, to 2 mm long, ca 1 mm wide, glabrous, the
persistent filaments of the stamens to 3.5 cm long; seeds 1 or 2, orbicular or semi-
orbicular, 3.5-4 mm long, the testa reddish-brown, glabrous.
cocle: Aguadulce, along outskirts of tidal belt, Pittier 4991 (US).

9. Xylosma sp. 4

Shrub 2.4 m tall, the branchlets with stout, axillary spines to 7 mm long,
inconspicuously puberulous. Leaves with the petiole to 1 cm long or slightly
longer, ± canaliculate above, inconspicuously puberulous; blade elliptic to ovate-
elliptic, acute and somewhat decurrent at the base, long-acuminate at the apex,
rather coarsely and obtusely glandular-serrate along the margins, to 13.5 cm long
and 6.5 cm wide, rigid-chartaceous to subcoriaceous, dull on both sides, inconspicu-
ously puberulous along the veins to glabrescent above, minutely hirtellous below,
the network of the venation inconspicuous above, the costa prominent and the
secondary veins and veinlets prominulous beneath. Inflorescences contracted-race-
mose. Flowers dioecious. Male flowers pale yellow, the pedicels to 6 mm long,
minutely puberulous, bracteolate at the base; calyx 4- or 5-merous, the sepals ±
subrotund, somewhat acute at the apex, to 2.8 mm long and 2.3 mm wide,
eciliolate, glabrous outside, short-pilose inside; glandular disc around the stamens
ca 1 mm long; stamens 16-17, the filaments slender, to 3-3.5 mm long, glabrous.
Female flowers and fruit not seen.

bocas del toro: N slopes of Cerro Horqueta, Robalo Trail, alt 6000-7000 ft, Allen
4941 (F, MO).

Latin Names

Numbers in boldface type refer to descriptions; numbers in roman type ref

numbers with dagger (t) refer 1

to names incidentally mentioned.

Banara 106

laevis 121

guianensis 107

myriantha 123

Bartholomaea 94t

nitida 121

Carpotroche 98f, 100

ramiflora 120

crassiramea 102f

stjohnii 119

platyptera 101, 102f

sylvestris 122

subintegra 98

Dendrostylis 97

Edmonstonia 113

Casearia 116, 125f

pacifica 114

aculeata 121 f, 125 1

Flacourtia flexuosa 140

arborea 117

Flacourtiaceae 93, 94 1, 131t

arguta 118, 125f

Guidona procera 130

banquitana 121

Hasseltia 94f, 102

— var. laevis 121

floribunda 102

OF THE MISSOURI BOTANICAL GARDEN

Lunania 133
cuspidata 134
parviflora 134

cruris 111
Ryania 131
angustifolia 131f
speciosa 131 1

— var. panamensis 131, 133t
— var. speciosa 133f

arborea 117
nitida 121

sylvestris 122
Sloanea longicuspis
Spruceanthus 105

Tiltaceae94t

intermedium 141

Neosprucea 105
grandiflora 106t
sararensis 106t

FLORA OF PANAMA

Part VIII
Family 154. SAPOTACEAE 1 ' 2

by Will H. Blackwell, Jr.

Missouri Botanical Garden and Department of Botany, Washington

University, St. Louis, Missouri

Trees or shrubs with milky latex (lactiferous sacs present in the pith and
cortex and the leaves and fruit), glabrous or pubescent (the trichomes unicellular
and 2-armed). Leaves alternate or rarely opposite, petiolate, estipulate; blades
simple, entire, often leathery, penninerved. Flowers usually $ , regular, axillary
to leaves and/or leaf-scars, fasciculate or occasionally solitary, the pedicels ses-
sile; sepals 4-12, imbricate or spiralled, in one or two whorls, free or united only
at the base; corolla gamopetalous, the lobes 4-8, shorter or longer than the tube,
as many as or fewer than (rarely more than) the sepals, with or without basal
petaloid appendages; stamens epipetalous, usually equal in number to and op-
posite the corolla-lobes, an outer antisepalous whorl of staminodes often present
(typically equal in number to and appearing to be in the same whorl with the
stamens); ovary superior, (l-)4 to 14-carpelled and -loculed, completely septate,
with 1 ascending, lateral or basal, anatropous, bitegmic ovule in each locule, the
style single, entire or lobed at the summit. Fruit a berry, often fleshy, the exocarp
frequently becoming corky or sclerotic; seeds 1-several, large, with or without
endosperm, the testa hard and shiny, the scar lateral to basal, large or small.

A family of about 40 genera and 600 species, widespread in the tropics of the
New and Old World; six genera occur in Panama. Several representatives are
economically important.

Key to Genera 3
a. Each corolla-lobe with a pair of petaloid appendages arising from the base
(the appendages dorsal or lateral), if these lacking then the sepals in two dis-
tinct whorls of three each; staminodes present and petaloid.
b. Sepals biseriate (3 + 3); corolla-lobe appendages dorsal or absent; ovules
attached laterally; seed-scar lateral or essentially so, equalling or extend-
ing beyond the middle of the seed . ..1. Manilkara

Assisted by National Science Foundation Grant No. GB-5674 (Principal Investigator,
Walter H. Lewis).

2 Synonyms presented in this paper included, in addition to names used in a previous
flora dealing with a portion of Panama (e.g. Standley, Flora of the Panama Canal Zone,
Contr. U. S. Nat. Herb. 27:299-301, 1928), only those based on Panamanian types or
those relegated to synonymy for the first time. For additional information on nomeclature
and synonymy, reference should be made to Cronquist's excellent publications (enumerated
after the generic descriptions) and to Moore & Stearn (The identity of Achras zapota L.
and the names for the sapodilla and the sapote. Taxon 16: 382-395, 1967).

3 The key to genera is intended for identifkatioi
Broader application may lead to misidentification.
Ann. Missouri Bot. Gard. 55(2): 145-169, 1968.

16 ANNALS OF THE MISSOURI BOTANICAL GARDEN

bb. Sepals not in two distinct whorls though sometimes quincuncial; corolla-
lobe appendages lateral; ovules attached basilaterally; seed-scar basal or
basilateral, not reaching the middle of the seed.

c. Plants unarmed; flowers borne at defoliated nodes only; ovary glabrous,
the style 1.5-2 mm long; apex of fruit abruptly tapering to the persistent

style; seeds with endosperm 2. Dipholis

cc. Plants often with spines; flowers axillary to leaves and occasionally
also leaf-scars; ovary pubescent, the style 3-7 mm long; apex of fruit
rounded or subtruncate, not tapering to the style; seeds lacking en-
dosperm -3. Bumelia

. Corolla-lobes lacking appendages; sepals usually uniseriate (though often
strongly imbricate or spiralled), if distinctly biseriate then 2 + 2; staminodes
present (often not petaloid) or i '
d. Secondary lateral veins ±

absant 4. Chrysophyllur

dd. Secondary laterals not paralleling (often essentially perpendicular to)
the primary laterals; staminodes present (frequently scale-like or linear-
lanceolate or filiform, occasionally petaloid).

e. Leaf-blades usually canaliculate above the midvein, the channel
terminating in a small pouch at the bass of the blade; petiole usually
Y 2 the length of the blade or more; ovary glabrate; seed-scar often
elongate but not reaching the middle of the seed; endosperm present

5. Mastichodendron

ee. Leaf-blades n n I ;: mole less than ]/ 2 the blade length; ovary
conspicuously pubescent; seed-scar excaeding the middle of the seed;
endosperm lacking 6. Pouteria

1. MANILKARA
Manilkara Adanson, Fam. 2: 166, 1763.

Trees, unarmed. Leaves alternate; blades coriaceous, the primary laterals
numerous, fine, rather closely spaced, often subobscure, straight (not arcuate near
the margin). Flowers fasciculate in the axils of leaves or leaf-scars or solitary;
sepals usually 6, infrequently 4 or 8, in 2 whorls of (2)3(4); corolla-lobes as
many as the sepals, each with a pair of dorsal ± petaloid appendages arising
from the base, occasionally these fused to the lobes or lacking; staminodes usually
petaloid, rarely minute or replaced by functional stamens; stamens as many as
corolla-lobes or rarely twice as many (morphologically 2 whorls), typically at-
tached to the corolla at the juncture of the tube and lobes; ovary often pubescent,
6 to 14-celled, the ovules affixed laterally. Fruit 1 to several-seeded, fleshy; seed
compressed, the scar rather elongate, lateral or basilateral (but equalling or sur-
passing the middle of the seed), ± linear, the endosperm copious.

Thirteen species are known in North America, four occurring in Panama.

Useful reference:

Cronquist, A. Studies in the Sapotaceae, IV. The North American species
of Manilkara. Bull. Torrey Bot. Club 72: 550-562, 1945.

a. Flowers 2-12 per axillary fascicle; corolla-tube usually shorter than the corolla-
lobes; ovary glabrate or pubescent only on the upper portion,
b. Pedicels glabrous; outer sepals glabrate by anthesis; corolla-lobes equalled
in length by dorsal appendages arising from their base (each appendage
deeply bifid or trifid); ovary glabrate; fruit smooth or but slightly rough-

flora of panama (Family 154. Sapotaceae)

Corolla-lobes narrowly oblanceolate or spatulate, 0.5-1 mm broad, equalled

in length by simple dorsal appendages .3. M. meridionalis

, Corolla-lobes oblong to ovate, 1.5-3 mm broad, lacking dorsal appendages

bidentata (DC.) Chevalier, Revue Bot. Appl. & Arg. Trop.

12: 270, 1932.
Mimusops darienensis Pittier, Contr. U. S. Nat. Herb. 18: 249, 1917.
Manilkara darienensis (Pittier) Standley, Trop. Woods 31 : 45, 1932.

Tree to 35 m. Leaves with petioles 1.3-4.2 cm long; blades narrowly obovate
or sometimes elliptic, obtuse or rounded to shallowly emarginate or short-acu-
minate, 6-30 cm long, 3-12 cm broad, glabrous, the primary laterals fine and
rather closely spaced, occasionally subobscure. Flowers 3-12 per fascicle, the fas-
cicles axillary to both leaves and leaf-scars; pedicels glabrous, 1.2-3 cm long;
sepals ovate or ovate-oblong, 4-6 mm long, the inner minutely appressed-sericeous,
the outer soon glabrate; corolla 5-7 mm long, subrotate, the tube ca ] / 5 the total
length, the lobes narrowly elliptic-lanceolate, the dorsal appendages as broad as
and usually slightly longer than the lobes, free from the lobes to the tube, distally
divided for ca % of their length into 2 or 3 slender lanceolate segments; staminodes
1.2-3 mm long, ovate-lanceolate, erose; staminal filaments equalling or slightly
exceeding the staminodes; ovary glabrate, the style glabrous, 4-5 mm long. Fruit
smooth or slightly roughened, globose or ellipsoid-globose, 2-3.5 cm long.

Hispaniola, Puerto Rico, Lesser Antilles, northern South America and Panama.

bocas del toro: s. loc, Cox s.u. (US), canal zone: hills aroud Gatun, Pittier 2699
(US), panama: nr Chepo, Kluge 55 (US), san blas: hills nr Puerto Obaldfa, Pittier 4318
(holotype of Mimusops darienensis US), 4384 (US).

Manilkara bidentata is important as a source of balata, a nonelastic rubber
obtained from the latex which is a nonconductor, resistant to water. Balata and
a similar product, gutta-percha (obtained from another member of the Sapotaceae),
are used in the construction of marine cable, machine belting, telephone receivers,
golf balls, waterproofing, adhesives and a number of other items.

chicle (Pittier) Gilly, Trop. Woods 73: 14, 1943.— Fig. 1.
Achras calcicola Pittier, Jour. Wash. Acad. Sci. 9: 438, 1919.
Manilkara calcicola (Pittier) Gilly, Trop. Woods 73: 15, 1943.

Tree to 40 m. Leaves with petioles 1-3.5 cm long; blades 8-24(-29) cm long,
3-7 (-10) cm broad, oblanceolate or narrowly obovate to elliptic or elliptic-oblong,
obtuse or rounded to acute or acuminate, paler and more yellowish beneath, ap-
pressed-sericeous-strigulose below when young but eventually glabrate. Flowers yel-
lowish- or grenish-white, 2-5 per fascicle; pedicels strigulose, 0.5-3 cm long; sepals
5-9 mm long, ovate to ovate- lanceolate or ovate-oblong, the inner and outer sericous-
tomentulose; corolla 5.5-9 mm long, the tube y 4 - l / 2 the total length, the lobes
ovate, often erose or erose-dentate at the apex, lacking dorsal appendages; sta-

148 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Njll II II <l A i\ (\

Fig. 1. Manilkara chicle (Pittier) Gilly: A, habit (Xi/ 2 ); B, corolla c
chment of stamens and staminodes (X5); C, flower with corolla removed (X5 ] / 2 ); D,
t (X2/ s ); E, seed (X%). A-C after Sttmson 5294 (MO); D-E after Johnston 753 (GH).

flora of panama (Family 154. Sapotaceae) 149

minodes 2.5-4.5 mm long, distally erose-1 acini ate, often shallowly bifid; stamens
equalling the staminodes; ovary minutely sericeous on top, glabrous elsewhere,
the style 6-9 mm long, glabrous except at the base. Fruit mealy-roughened, 2-4
cm in diam, subglobose or obovoid, several-seeded; seeds 14-19 mm long, flattened,
the basilateral scar barely reaching the middle.

Oaxaca, Mexico through Central America to Colombia.

canal zone: area W of Limon Bay, Gatun Locks & Gatun Lake Johnston 1535 (A) ;
Ancon Hill, Standley 26384 (US), darien: vie of Pifias, Duke 10653 (MO); Patino, Pittier
5698 (GH, US), los santos: i Vimson 5294 (MO), panama:

San Jose I, Johnston 336 (GH), 342 (MO, GH), 646 (GH), 753 (GH, MO, US); around
Alhajuela, Chagres Valley, Pittier 3457 (holotype of Arenas calcicola US).

Although bearing the specific epithet chicle, this species is not the principal
source of chicle gum. The latex is difficult to coagulate and inferior to that of
M. zapota.

3. Manilkara meridionalis Gilly, Trop. Woods 73: 12, 1943.
M. tabogaensis Gilly, loc. cit. 10.

Tree to 30 m. Leaves clustered toward the branch tips; petioles 1-2.7 cm
long; blades elliptic or oblong-elliptic to narrowly obovate, 5.5-13 cm long, 1.7-4.5
cm broad, glabrous or sparsely rufescent along the midrib beneath. Flowers soli-
tary in the leaf-axils; pedicels 1-2.3 cm long, tomentulose or eventually glabrate;
sepals ovate, 6-9 mm long, minutely tomentulose externally; corolla cylindric,
glabrous or pubescent, ca 9 mm long, the tube usually comprising y 2 - 2 / 3 of the
length, the lobes narrowly oblanceolate or spatulate, 0.5-1 mm broad, the dorsal
appendages ovate-lanceolate to elliptic or obovate, broader than but ca equal in
length to the lobes, simple (not divided into elongate segments distally), basally
united with the lobes for ca l / 3 of their length; staminodes ±: ovate, erose-fimbriate,
2.5-5 mm long, exceeding the staminal filaments; ovary tomentose, the style 5.5-8.5
mm long, glabrous except at the base. Fruit brown, mealy-roughened, to 3.5 cm
long; seed strongly compressed, ca 2 cm long, the linear scar extending from near
the base to well beyond the middle of the seed.

Guerrero, Mexico to Colombia and Venezuela; introduced in the West Indies.

panama: Taboga I, Standley 27099 (US), Woodson et al. 1455 (holotype of M. tabo-
gaensis NY, not seen; isotypes A, MO) .

4. Manilkara zapota (L.) van Royan, Blumea 7: 410, 1953*.
Achras zapota L, Sp. PI. 1190, 1753.

Tree to 40 m. Leaves clustered toward the branch tips; petioles 0.8-3 cm
long; blades elliptic or oblong-elliptic to somewhat obovate or oblanceolate, 4-15
cm long, 1.5-6 cm broad, subconcolorous, glabrate at maturity, the reticulation
usually evident below. Flowers solitary in the leaf-axils; pedicels 1.2-2.5 cm long,
rufous-tomentulose (some of the pubescence deciduous with age); sepals 6-10
mm long, ovate or occasionally oblong, tomentulose, the outer often losing some

4 For a discussion of nomenclature and complete list of synonyms of M. zapota see
Moore & Stearn, Taxon 16: 382-395, 1967.

150 ANNALS OF THE MISSOURI BOTANICAL GARDEN

(rarely all) their pubescence with age; corolla 6-11 mm long, the tube usually
comprising J^-% the total length, the lobes oblong to ovate, 1.5-3 mm broad, entire
or erose or dentate at the apex, lacking dorsal appendages; staminodes petaloid,
rather narrowly ovate-lanceolate, 3-4.5 mm long, erose; stamens %-% as long
as the staminodes; ovary densely sericeous, the style 4.5-8 mm long, glabrous
except at the base, the apex often irregularly toothed or lobed. Fruit brown, mealy-
roughened, ellipsoid or ovoid or subglobose, to 10 cm in diam; seeds brown, com-
pressed, 16-24 mm long, the linear scar extending from near the base to beyond
the middle.

Mexico (as far north as San Luis Potosi and Nayarit) to northern South
America and from southern Florida through the West Indies; probably native only
from Mexico to Costa Rica.

canal zone: Balboa, Standley 27121 (US), 30860 (US).

The latex of the sapodilla or chicle-tree is the commercial source of the gum
base of the chewing gum industry. It contains 20-25% of the gutta-percha-like
gum and is obtained by tapping the trunk by a series of connected, half-encircl-
ing, zig-zag gashes. To prevent death of the trees, they are tapped only once every
two or three years. The United States is the leading consumer of chicle, the bulk
being imported from British Honduras. The pear-shaped fruit is esteemed by
people of tropical America and is eaten uncooked.

2. DIPHOLIS
Dipholis DC, Prodr. 8: 188, 1844.

Trees or shrubs, lacking spines or thorns. Leaves alternate, the blades cori-
aceous or subcoriaceous. Flowers small, white or green, numerous or few per axil-
lary fascicle, occasionally solitary, the clusters axillary to leaves, leaf-scars or
both; sepals (4)5(-9), uniseriate; corolla rotate or funnelform, 5(6) -lobed, each
lobe with a pair of lateral appendages arising from the base; staminodes petaloid,
these and the appendages of the corolla-lobes often erose, fimbriate or laciniate;
anthers often exserted; ovary glabrous or rarely appressed-puberulent, 5-celled,
the ovules attached basilaterally. Fruit fleshly, usually 1-seeded at maturity,
abruptly tapering to the short, persistent style; seed with a small, virtually basal
scar and copious endosperm, the cotyledons thin.

Approximately 15 species; heavily centered in the West Indies (principally
the Greater Antilles) but also occurring in Central America, Mexico and southern
Florida; one species is known from Panama.

Although Steam (Jour. Arnold Arb. 49: 282-283, 1968) argues impressively
against the recognition of Dipholis as an entity generically distinct from Bumelia,
I favor its retention at present. I have found the delimiting characters listed by
Cronquist (Jour. Arnold Arb. 26: 445, 1945) to hold up rather well in the species
I have examined personally and there can be no doubt of their validity with re-
gard to Panamanian representatives.

Useful reference:

Cronquist, A. Studies in the Sapotaceae, III. Dipholis and Bumelia. Jour.
Arnold Arb. 26: 435-471, 1945 (Dipholis p. 435-445).

flora of fanama (Family 154. Sapotaceae)

151

1. Dipholis minutiflora Pittier, Contr. U. S. Nat. Herb. 13: 464, 1912.— Fig. 2.

Tree to 35 m. Leaves with petioles 0.5-1.8 cm long; blades elliptic-oblanceolate

or narrowly elliptic-obovate, rounded to acute or subacuminate, 5-20 cm long,

2-10 cm broad, glabrous or finely white-strigulose (particularly below). Flowers

Fig. 2. Dipholis min
tachment of stamens and
fruit (XU/z). A after Woodson
Hinton7600 (MO), Mexico.

:ier: A, habit (X'/ 2 ); B,
(X8); C, flower with <
995 (MO); B-C after

corolla opened to s
corolla removed (X
Allen 1564 (MO);

[Vol. 55
152 ANNALS OF THE MISSOURI BOTANICAL GARDEN

often numerous per fascicle, the fascicles occurring only at defoliated nodes, each
borne on a small woody cushion; pedicels 3-10 mm long, very sparsely to moder-
ately appressed-sericeous; sepals 5(-8), closely imbricate, sometimes unequal, sub-
orbicular or broadly ovate, glabrate, carnosulous, ca 2 mm long; corolla 4-4.5 mm
long, the 5 lobes erose, ca 3 mm long, ± ovate, narrowed toward the base, the
appendages lance-ovate, laciniate, V 2 -Va the length of the lobes; staminodes 2-3
mm long; narrowly ovate, erose; staminal filaments ca 2 mm long; ovary glabrous,
1 mm long, the style 1.5-2 mm long, glabrous, conical, evenly tapered from the
ovary, virtually as broad as the ovary at the base. Fruit broadly ovoid, 1.5-2.5 cm
long, 1-2 cm broad, conical-apiculate, the flesh scant; seed ellipsoid, slightly com-
pressed, 14-18 mm long, the scar 3-5 mm long.

State of Mexico to Panama.

CHiRiQui: vie of Cerro Punta, Allen 1564 (F, MO); Quebrada Velo, vie of Finca
Lerida, Allen 4676 (MO); valley of the upper Rio Chiriqui Viejo, White 109b (F, MO),
White & White 1 (MO); Bajo Mona, mouth of Quebrada Chiquero, along Rio Caldera,
Woodson et at 995 (F, MO), panama: hills above Campana, Allen 1314 (F, MO).

3. BUMELIA
Bumelia Swartz, Nov. Gen. Sp. PI. Prodr. 49, 1788.

Shrubs or trees, usually spiny or thorny. Leaves alternate or casually opposite,
the blades firm. Flowers few to numerous per axillary fascicle, rarely solitary,
green or white, (4)5(6)-merous; sepals uniseriate; corolla-lobes well exceeding
the tube, each with a pair of lateral appendages arising from the base or these
rarely lacking; staminodes petaloid, entire to erose or laciniate; anthers often ex-
serted; ovary pubescent or less frequently glabrous, usually 5-locular, the ovules
affixed basilaterally, the style slender to virtually filiform. Fruit fleshy, 1-seeded,
broadly rounded or subtruncate or retuse at the apex (not tapering to the persistent
style); seed with a small subbasal scar, lacking endosperm, the cotyledons fleshy.

A New World genus of about 23 species, the majority in the United States,
Mexico and the West Indies but several extending into Central and South Amer-
ica; one species occurs in Panama.

Useful reference:

Cronquist, A. Studies in the Sapotaceae, III. Dipholis and Bumelia. Jour.
Arnold Arb. 26: 435-471, 1945 (Bumelia p. 445-471).

1. Bumelia persimilis Hemsley subsp. persimilis, Biol. Centr.-Amer., Bot.

2:298, 1882.-Fig.3.
B. panamensis Standley, Trop. Woods 4: 9, 1925.

Shrub or tree to 18 m, possessing spines. Leaves with petioles 2-10(-12) mm
long; blades elliptic- or somewhat ovate-lanceolate to elliptic or elliptic-oblanceo-
late, acute or acuminate to obtuse, 3.5-12.5 cm long, 1.3-5 cm broad, glabrous and
lustrous above, at first sparsely appressed-rufous- or cinereous-sericeous below,
soon glabrate, with 10-30 pairs of primary lateral veins, these brochidodrome.
Flowers white, several to numerous per fascicle, the fascicles axillary to leaves and
occasionally also leaf-scars; pedicels 3-6 mm long; sepals ovate, 1.8-3.7 mm long,
unequal, quincuncial, glabrate to moderately appressed-sericeous; corolla rotate-

. of Panama (Family 154. Sapataceae) 153

Fig. 3. Bumelia persimilis Hemsley subsp. persimilis: A, habit (Xi/ 2 ); B, portioi
corolla showing attachment of stamens and staminodes (X7'/ 2 ); C, flower with coi
removed (X7'/ 2 ); D, fruit (X2). A after Williams 16585 (MO), Costa Rica; B-C ;
Schipp 1077 (MO), British Honduras; D after Hunter & Allen 298 (F).

154 ANNALS OF THE MISSOURI BOTANICAL GARDEN

funnelform, the tube ca 1.5 mm long, the lobes 3-4.5 mm long, oblong-ovate,
subtruncate and erose at the apex, narrowed at the base, the appendages lance-
filiform, ca y 4 the length of the lobes; staminodes 2.5-3 mm long, lanceolate to
obovate, ± laciniate; staminal filaments 3-3.5 mm long, the anthers 0.7-2 mm
long; ovary appressed-sericeous, the style 3-7 mm long, a stigma not apparent.
Fruit spheroid or ellipsoid, smooth, 1.2-2.5 cm long, the apex rounded.

Vera Cruz and Oaxaca, Mexico through Central America to Venezuela.

cocle: El Valle de Anton, along Rio Indio trail, Hunter & Allen 298 (F). Panama:
nr Chepo, Kluge 12 (holotype of B. panamensis US, not seen).

The other subspecies of B. persimilis, subsp. subsessiliflora (Hemsley) Cron-
quist, is exclusively Mexican (Chihuahua and Durango to Michoacan and Oaxaca)
and is characterized, among other features, by a dense, rufous, spreading pubescence
of the young twigs and lower leaf -blade surface.

4. CHRYSOPHYLLUM
Chrysophyllum L., Sp. PI. 192, 1753.

Shrubs to medium-sized trees. Leaves alternate; blades with the secondary
laterals and areoles ± paralleling the primary laterals. Flowers several to nu-
merous in each axillary cluster, occasionally solitary; sepals (4)5(6), uniseriate,
often quincuncial when 5, united at the base; corolla not exceeding 6 mm in
length, the lobes (4)5(6), lacking appendages; staminodes absent; staminal fila-
ments attached to the top of the corolla-tube or base of the corolla-lobes; ovary
appressed-pubescent, 4 to 12-celled, the ovules affixed laterally or basilaterally,
the style short (less than 1.75 mm in Panama taxa) , columnar, the stigma discoid,
marginal lobes often evident. Fruit fleshy; seeds 1-several, the scar large, broadly
elliptic to subcordate, at least 5 mm long, lateral or basilateral, the endosperm
copious.

Approximately 40 species occur in the New World and a number of Old
World species are known; two species are found in Panama.

Useful references:

Cronquist, A. Studies in the Sapotaceae, I. The North American species of
Chrysophyllum. Bull. Torrey Bot. Club 72: 191-204, 1945.

. Studies in the Sapotaceae, V. The South American species of

Chrysophyllum. loc. cit 73: 286-311, 1946.

a. Blades sparsely white-strigulose below or glabrate; corolla-lobes less than i/ 2 as

long as the tube; stigma 5-lobed; fruit to 2 cm broad - 1. C. panamense

aa. Blades densely rufous-sericeous below (with a coppery sheen, even on dried
specimens); corolla-lobes equalling or slightly exceeding the tube; stigma 7 to
12-lobed; fruit 3 cm broad or more 2. C. cainito

1. Chrysophyllum panamense Pittier, Contr. U.S. Nat. Herb. 18: 165, 1916.—

Fig. 4.
C. panamense var. macrophyllum Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 366,

Tree to 15 m. Leaves with petioles 0.8-2.5 cm long; blades elliptic or broadly
oblong to elliptic-obovate, acuminate, to 25(-33) cm long and 11 (-14) cm broad,

flora of panama (Family 154. Sapotaceae) 155

Fig. 4. Chrysophyllum panamense Pittier: A, habit (X%); B, corolla opened to
show attachment of stamens (X7); C, flower with corolla removed (X7); U, fruit (><!%)„
A after Starry 82 (F) and Shattuck 1024 (F); B-C after Zetek 3810 (F); D after Zetek
Z4327 (F).

156 ANNALS OF THE MISSOURI BOTANICAL GARDEN

glabrous above, glabrate below or sparsely and finely white-strigulose. Flowers
several-numerous in each axillary cluster, the pedicels 3-7 mm long, appressed-
gray-pubescent; sepals to 2 mm long, ovate or suborbicular, strigulose externally
(occasionally losing some of the pubescence); corolla subcylindric, pale green to
yellow or white, 3.5-5.5 mm long, externally often appressed-rufous-sericeous on
the lobes and adjacent portions of the tube, the lobes deltoid-ovate, ± erect, less
than y 2 the length of the tube; anthers 0.4-0.6 mm long, the filaments attached
at the juncture of the corolla-tube and lobes; style 1-1.5 mm long, the stigma with

5 small marginal lobes. Fruit several-seeded, to 2 cm broad, oblate-spheroid to
ovoid, often lightly ribbed; seeds pale brown, ca 1 cm long, with the basilateral scar
extending to the middle or beyond.

Panama and Costa Rica, chiefly on the Atlantic slope.

bocas del toro: region of Almirante, Cooper 353 (F); s. loc, von Wedel 230 (F, MO).
canal zone: Barro Colorado I, Aviles 969 (F), Bailey & Bailey 397 (F), Bangham 591
(F), Ebinger 210 (MO), Shattuck 778 (F), 969 (MO), 7024 (F, MO), Starry 82 (F),
718 (F). Wetmore & Abbe 169 (F), Zetek 3810 (holotype of var. macrophyllum F, iso-
type MO), Z-4327 (F). Z-4330 (F); s. loc, Hayes s.n. (F, MO); along the Trinidad River,
Pittier4W5 (holotype US).

2. Chrysophyllum cainito L., Sp. Pi. 192, 1753.

Tree to 30 m. Leaves with petioles 0.8-2.3 cm long; blades elliptic or oblong,
occasionally ovate or somewhat obovate, short-acuminate, to 15(-19) cm long
and 8.3 cm broad, shiny- glabrous above, appressed-rufous-tomentulose below.
Flowers often numerous per axillary cluster, the pedicels 5-16 mm long, appressed-
rufous-sericeous; sepals ca 1 mm long, suborbicular or broadly ovate, closely
rufescent; corolla green, yellow or purplish-white, 3-5 mm long, rotate-funnelform,
the tube glabrous, the lobes appressed-sericeous externally except at the margins,
ovate or lance-ovate, equalling or slightly exceeding the tube; anthers 0.6-0 9 mm
long, the short filaments attached to the corolla-lobes near the base; style 0.5 mm
long or less, the stigma with 7-12 small marginal lobes. Fruit 3-10 cm broad,
subglobose, several-seeded; seeds oblique-obovate, flattened, 1-2.5 cm long, the
scar lateral, extending nearly the length of the seed.

Probably native to the West Indies; cultivated and often naturalized in Cen-
tral America and Mexico and occasionally in northern South America and southern
Florida.

bocas del toro: Changuinola Valley, Dunlap 24 (F); vie of Chiriqui Lagoon, von

Wedel 2523 (MO), canal zone: nr Madden Dam & along Azote Caballo Rd nr Alajuela,

Dodge 16573 (MO); upper Chilibre River, i/ 2 -l mi below Chilibre, Seibert 1505 (MO).

•astures & forested river banks E of Gaulaca, Allen 5033 (MO) ; Progreso, Cooper

6 Slater 247 (F), 264 (F). cocle: floor of El Valle de Anton, Allen 2747 (F). darien:
Rio Sabana above Sante Fe, Duke 14104 (MO); vie of Campamento Buena Vista, Rio
Chucunaque above confluence with Rio Tuquesa, Stern et al. 853 (MO), herrera: vie of
Ocu, Allen 3647 (MO), los santos: from 1 mi S to 10 mi N of Tonosi, Duke 12488 (MO).
Panama: woods along Pan-Am Hwy ca halfway betw El Llaon & Rio Momoni, Duke 5527
(MO); Chepo, Kluge 49 (F); Taboga I, Woodson et al. 1537 (MO), veraguas: dry slopes
of Cerro Tute, region W of Santa Fe, Allen 4441 (MO); Coiba I, Dwyer 2331 (MO).

Chrysophyllum cainito is the best known species of the genus and is widely
planted as a shade tree and for its succulent, edible fruit which resembles a small

flora of Panama ( Family 154. Sapotaceae ) 157

apple. When the fruit is cut in cross-section, the several compressed, brown seeds
are seen to radiate in a star-like fashion around the central axis — hence the common
name "star apple." The foliage is bright blue-green above and coppery or golden
beneath and offers a striking contrast when stirred by the wind.

5. MASTICHODENDRON
Mastichodendron Cronquist, Lloydia 9: 245, 1946.

Trees, unarmed. Leaves alternate or subopposite; blades typically exceeding
6 cm, ± canaliculate above the midrib, the channel often terminating in a small
pouch at the base of the blade. Flowers in clusters at defoliated nodes; sepals 5,
uniseriate; corolla subrotate to ± cylindric, the lobes 5, exceeding the tube, lack-
ing appendages; staminodes not petaloid; staminal filaments attached to the corolla
near the level of the sinuses; ovary glabrate, usually 5-locular, the ovules at-
tached basilaterally. Fruit fleshy, 1-seeded at maturity; seed 1-2.7 cm long, the
scar basilateral, as much as 9 mm long but not extending to the middle of the
seed, the endosperm copious.

Five species are known in North America, one reaching Panama; the status
of several Chinese species is open to question (Cronquist, Lloydia 9: 245, 1946).

Useful reference:

Cronquist, A. Studies in the Sapotaceae, II. Survey of the North American
genera. Lloydia 9: 241-292, 1946 (Mastichodendron p, 244-252).

1. Mastichodendron capiri (DC.) Cronquist var. tempisque (Pittier) Cron-
quist, Lloydia 9: 250, 1946.— Fig. 5.

Tree to 25 m. Leaves often clustered toward the branch tips; petioles 3-9 cm
long, 1-1.5 mm broad, usually l / 2 - 2 / 3 the blade length; blades elliptic or elliptic-
ovate to elliptic-obovate, acuminate or acute to rounded or emarginate, 6.5-15 cm
long, 3-7 cm broad, glabrate, the secondary laterals angled at ca 45° with respect
to the primary laterals or irregularly anastomosing, the reticulation prominulous
and rather coarse. Flowers several-numerous per cluster, the clusters occasionally
closely spaced (seemingly one large cluster below the leaves of the season) ; pedi-
cels 3-8 mm long, glabrate or pubescent; sepals 1.5-3.2 mm long, rounded or
broadly ovate; corolla yellow or greenish-yellow, 5-8.5 mm long, the lobes broadly
oblong to obovate, slightly cucullate, imbricate; staminodes squamiform, ovate
or ovate-lanceolate, erose, 0.75-1.5 mm long, not more than ! / 2 the length of the
staminal filaments; anthers 2.3-3.8 mm long, sagittate at the base; style 2-3 mm
long, thickening gradually toward a vitually imperceptible juncture with the
ovary, with several longitudinal creases, the stigma inconspicuous, the lobing
scarcely evident. Fruit ovoid or ellipsoid, yellowish, apiculate, to 4 cm long and
2.5 cm thick, the flesh scant; seed ellipsoid, 1.5-2.5 cm long, only slightly com-
pressed, the scar ovate-ellipsoid, 6-9 mm long.

Panama to Oaxaca, Mexico and perhaps occasionally north-westward.
cocle: Penonome & vie, Williams 421 (F, US).

Mastichodendron capiri var. capiri, characterized by pubescent leaves, occurs
primarily from Oaxaca to Nayarit, Mexico.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 5. Mastichodendron capiri (DC.) Cronquist
A, habit (Xi/ 2 ); B, corolla opened to show attachmen
C, flower with corolla removed (X6); D, fruit (X%). A-C after Godfi
Costa Rica; D after Smith 2508 (MO), ~

(Pittier) Cronquist:

' (X5);

(MO),

flora of panama (Family 154. Sapotaceae) 159

6. POUTERIA 5
Pouteria Aublet, Hist. PI. Gui. Fr. 85, pi. 33, 1775.

Trees or occasionally shrubs, lacking spines. Leaves alternate or sometimes
subopposite; blades membranous or coriaceous, the primary lateral veins strongly
arcuate near the margin, not crowded, the secondary laterals often ± perpendicu-
lar to the primary laterals (at least toward the margin), occasionally irregular.
Flowers white to yellow or green, pedicellate or sessile, the fascicles axillary to
leaves or leaf-scars; sepals 4-12, uniseriate or biseriate (always 2 + 2); corolla
cylindric to subrotate, the lobes 4-6(-7), lacking appendages; staminodes petaloid
or more often not petaloid (lance-filiform, linear or small and inconspicuous);
staminal filaments attached at or near the top of the corolla-tube, less frequently
at the middle or near the base (virtually free); ovary pubescent, (l-)4 to 10-
loculed, the ovules attached laterally. Fruit fleshy or woody or gall-like; seeds
1-several, lacking endosperm, the scar lateral, exceeding the middle of the seed,
often broad.

About 40 species occur in North America; 15 species, as recognized in this
treatment, are found in Panama.

Useful reference:

Cronquist, A. Studies in the Sapotaceae, II. Survey of the North American
genera. Lloydia 9: 241-292, 1946 (Pouteria p. 257-292).

Sepals 4-7 [unknown in P. lucentifolia but presumably so],
b. Flowers obviously pedicellate (the majority of the pe
mm long),
c. Leaf-blades sericeous beneath.

d. Pubescence pale; corolla-lobes fringed-ciliolate;
attached at the middle of the corolla-
rufous; corolla-lobes nor < !: I i

!af-blades glabrous or pubescent only c
Sepals 2-3 mm long,
f. Reticulation i " " "

1. P. euryphylla

1 filaments

...2. P. calistophylla

3. P. lucentifolia

iNiamui or icar-Diaaes nne; iruit I-seeded, less than 2

d; seed scar 8-10 mm broad 4

ee. Sepals 4-12 mm long.

ff. Reticulation of leaf-blades fine; fruit 1 -seeded, less than 2

8-10 mm broad 4. P. stipitata

;ins without specific orientation (irregularly

Is 4; corola glabrous externally ~ ~

Secondary laterals ± perpendicular to the primary laterals and
parallel to each other; sepals (4)5-6; corolla sparsely sericeous-
strigose externally 6. P. campechiana

Pouteria is a careful and invaluable revision, the

rial available precludes, even now, the po ifail I

auaining a reany satisractory classification. A number of species are known only from

the type collection and the complete morphology of the flower and/or fruit re

known for several, e.g. P. cooperi, P. sambuensis, P. lucentifolia. The key to species should

[Vol. 55
ANNALS OF THE MISSOURI BOTANICAL GARDEN

h. Sepals 5; corolla-tube shorter than the corolla-lobes.

i. Petioles 6-17 mm long; secondary lateral veins irregularly an-
er coarse; outer sepals ovate, the inner oblong-

; corolla tubular or campanulate; ovary 4-locular 7. P. chiricana

ii. Petioles 15-33 mm long; secondary laterals fine, close, regularly
disposed (though sinuous); all sepals ovate; corolla subrotate;

ovary 5-locular - 8. P. subrotata

hh. Sepals 4, 6 or 7; corolla-tube exceeding the corolla-lobes [unknown
in P. sambuensis],
j. Sepals 6-7; corolla-lobes 5; staminal filaments attached at the top

of the corolla-tube; fruit pyriform or obovoid 9. P. sclerocarpa

j]'. Sepals and corolla-lobes 4; staminal filaments attached at the
middle or base of the corolla-tube; fruit oblate-spheroid to ellipsoid
■Jiaracters known for P. sambuensis].

k. Fruit conspicuously hairy 10. P. sambuensis

kk. Fruit glabrous.

I. Leaf-blades finely appressed-sericeous below (sometimes
glabrate in age); outer sepals ovate and finely rufescent, the
inner rather narrowly oblong-spatulate; corolla 2.5-3 mm
long; staminal filaments attached at the base of the corolla-
tube 11. P- stylosa

II. Leaf-blades glabrate; sepals all ± ovate, the outer soon
glabrate; corolla 4.5-8 mm long; staminal filaments at-
tached at the middle of the corolla-tube or slightly below
12. P. caimito

aa. Sepals 8-12 or more.

m. Secondary lateral veins mostly perpendicular to the primary laterals,
decidedly more prominent than the fine reticulum; sepals conspicuously
appressed-pubescent on the back; fruit fleshy.

n. Primary lateral veins 20-50 pair; sepals (at least some) emarginate
or more deeply bilobed at the apex; fruit ellipsoid to ovoid or sub-
globose, brown, mealy-roughened 13. P. sapota

nn. Primary laterals 12-20 pair; sepals entire or scarcely emarginate;
fruit obovoid, yellowish-green, the surface irregularly wrinkled (at

least in herbarium specimens) and also mealy roughened 14. P. fossicola

mm. Secondary laterals rather irregularly disposed, scarcely more prominent

than the coarse reticulum; sepals glabrous; fruit woody 15. P. cooperi

1. Pouteria euryphylla (Standley) Baehni, Candollea 9: 249, 1942. 6

"Tree about 15 m high; leaves elliptic or elliptic-obovate, rather abruptly
and narrowly acuminate, about 10-22 cm long, 4-12 cm wide, glabrous above, very
closely and finely sericeous-strigose with pale hairs beneath; primary lateral veins
about 9-13 pair, not becoming crowded and obscure near the base of the blade,
as they do in many species; secondary lateral veins, except those near the midrib,
nearly perpendicular to the primary ones, but also somewhat sinuous and often
branched; petioles about 1.5-6 cm long; flowers few in the axils, the closely seri-
ceous-strigose pedicels about 4-5 mm long; sepals 4, about 3.5 mm long, broadly
elliptic-ovate, the outer closely hairy on the back, the inner glabrous except for
a strip down the middle of the back; corolla about 3.7-4.5 mm long, the lobes
a little shorter than the tube, evidently fringed-ciliolate; filaments attached about

"Quotation mi
available for study
9:257-292, 1946).

flora of panama (Family 154. Sapotaceae) 161

at the middle of the corolla-tube, and trace raised and conspicuous to the base;
anthers about 1 mm long, short-cuspidate; staminodes about 1 mm long, triangular-
lanceolate, ciliolate like the corolla-lobes, and alternating with them, nearly or
quite in the same series; ovary densely hairy, 4-loculate, the style a little over 2
mm long; fruit unknown."

Known only from the type collection.

bocas del toro: Buena Vista Camp on Chiriqui Trail at 1250 ft, Almirante region,
Cooper 611 (holotype F, isotype NY; not seen).

2. Pouteria calistophylla (Standley) Baehni, Candollea 9: 419, 1942.

Tree ca 20 m high. Leaves with petioles 1-3 cm long; blades obovate or ellip-
tic-obovate, acuminate, 10-22 cm long, 4.5-10 cm broad, glabrous above, rufous-
sericeous-tomentulose beneath (the pubescence mostly appressed), the primary
laterals 12-22 pair, prominently raised below, the secondary laterals (except the
2-3 most medial ones) subperpendicular to the primary laterals (although some-
what sinuous), sometimes obscured beneath by pubescence, the reticulation sub-
obscure. "Well-developed flowers unknown, but a very young flower has a stout
pedicel 5 mm long, with 5 sepals and a 5-lobed corolla, the filaments attached
at the level of the sinuses, the staminodes narrow, approximately triangular-
lanceolate; an overmature flower has 5 deltoid-ovate very firm sepals about 2 mm
long, the ovary parasitized, probably originally 2-loculate; fruit unknown."

Known only from the type collection.

bocas del toro: region of Almirante, Cricamola Valley, Cooper 481 (holotype F, not
seen; isotype US).

3. Pouteria lucentifolia (Standley) Baehni, Candollea 9: 424, 1942.
ley, Publ. Field Mus. Nat. Hist, Bot. Ser. 4: 251, 1929.

I (Standley) Baehni, Candollea 9: 353, 1942.

Tree ca 11 m high. Leaves with petioles 8-24 mm long; blades elliptic-obovate,
acuminate or occasionally obtuse or acute, 7-25 cm long, 3.3-10 cm broad, lustrous,
glabrous or occasionally sparsely hirsutulous on the midrib beneath, the primary
lateral veins 8-15 pair, the secondary laterals prominent below and often slightly
raised, becoming ± perpendicular to the primary laterals toward the margin al-
though somewhat irregularly anastomosing, the reticulation coarse, often prominent
below. "Flowers unknown, but a young developing fruit has a thick pedicel 5
mm long and nearly glabrous ovate sepals 3 mm long; pedicels evidently not
elongating in fruit." Fruit golden-brown, mealy-roughened, globose, 4-5 cm in
diam, the fruiting pedicel 5-6 mm broad; seeds 5, ellipsoidal, strongly flattened,
2.5-3.5 cm long, 1.3-1.6 cm broad, lustrous-brown, the seed-coat 0.4-0.6 mm thick,
the scar linear, 1.5-2 mm broad, extending the length of the seed.

Honduras, Costa Rica and Panama.

bocas del toro: region of Almirante, Cooper 369 (holotype of Lucuma pentasperma
F, not seen; isotype A); Changuinola Valley, Dunlap 591 (F).

4. Pouteria stipitata Cronquist, Lloydia 9: 265, 1946.

Tree to 10 m. Leaves with petioles 4-17 mm long; blades narrowly obovate
to elliptic, acuminate, 5.5-13 cm long, 2-4.5 cm broad, glabrate or appressed-

162 ANNALS OF THE MISSOURI BOTANICAL GARDEN

pubescent along the main veins beneath, the primary laterals 7-11 pair, the sec-
ondary laterals conspicuous but often irregularly anastomosing, sometimes be-
coming ± perpendicular to the primary laterals toward the margin, the reticula-
tion fine. Flowers fasciculate at defoliated nodes; pedicels 2-5 mm long; sepals
4-5, suborbicular or broadly ovate, 2-2.5 mm long, finely strigulose to glabrate;
corolla thin, 2.4-3.2 mm long, the lobes 4, subrectangular, 0.7-1.2 mm long, fringed-
hairy on the margin; staminodes oblong-spatulate, 0.3 mm long, marginally
fringed-hairy (especially near the tip); staminal filaments attached at the middle
of the corolla-tube; ovary 4-locular, densely ascending-hirsute externally with
pale trichomes, the style glabrous, 1-1.5 mm long. Fruit yellow, finely warty-
roughened, the body ellipsoid or obliquely-ellipsoid, 2-2.5 cm long, ca 1.6 cm
broad, constricted basally to a stipe ca 7 mm long and 3 mm broad; seed 1, ellipsoid,
ca 1.5 cm long, not compressed, the scar elliptic-obovate, 8-10 mm broad, extend-
ing the length of the seed.

Panama; known only from Barro Colorado Island and the Perlas Archipelago.

canal zone: Barro Colorado I, Shattuck 1125 (F), Zetek 4693 (holotype F). panama:
San Jose I, Erhnson 324 (GH, US), Johnston 634 (GH, MO, US), 1291 (GH).

5. Pouteria dominigensis (Gaertner f.) Baehni var. dominigensis, Lloydia

9: 278, 1946.
Lucuma serpentaria H.B.K., Nov. Gen. Sp. PI. 3: 242, 1819.

Tree to 10 m. Leaves with petioles 0.5-2 cm long; blades obovate-oblanceolate,
rounded to subacute, 4-13 cm long, 1.8-6 cm broad, glabrate except when very
young, lustrous, usually conspicuously reticulate-veiny on both sides, the primary
laterals less than 1 cm apart, the secondary laterals without specific orientation,
sometimes subequalling the primary laterals. Flowers 1-several per axil; pedicels
3-8 mm long; sepals 4 (2 + 2), 4-9 mm long, subequal, ovate, the outer finely
appressed-rufous-sericeous, the inner similarly pubescent down the middle, other-
wise glabrous; corolla cylindric, yellow or white, minutely papillose but glabrous,
8-16 mm long, the tube comprising ca % of the total length, the lobes (5)6, ±
oblong; staminodes linear-subulate, ca 3 mm long; staminal filaments 1-2 mm
long, attached at the top of the tube, the anthers 1.3-2 mm long; ovary densely
rufous-pubescent, (5)6(-8)-loculed, the style 6-11 mm long, glabrous except at
the base. Fruit yellow, fleshy, oblate, 2-5 cm long, 3-6 cm broad, the base of the
persistent style surrounded by a conspicuous light-colored areola 1-2 cm broad;
seeds 1-several, 1-3 cm long, the scar 3-10 mm broad.

West Indies and southern Florida; cultivated in the Canal Zone.

canal zone: s. loc, Johansen 36 (F); Ancon, Mell s.n. (F); Balboa, Standley 26894
(F), 30859 (F).

The other variety of P. dominigensis, var. cuprea (Urban & Ekman) Cron-
quist, is apparently limited to Haiti and the Dominican Republic and is distin*
guished by the leaves which are strongly rufous-strigose beneath or only tardily
glabrate.

flora of panama (Family 154. Sapotaceae) 163

a '\mmi

:A.j

Fig. 6. Pouteria campechiana (H.B.K.) Baehni: A, habit (Xi/ 2 ); B, corolla opened
to show attachment of stamens and staminodes (X4); C, flower with corolla removed
(X3i/ 2 ); D, fruit (xy 4 ); E, seed (Xi/ 2 ). A after Piper 6027 (F); B-C after Palmer 386
(MO), Mexico; D after Hinton 9185 (MO), Mexico; E after Allen 2609 (F).

164 ANNALS OF THE MISSOURI BOTANICAL GARDEN

L. glabrifolia Piitier, Contr. U. S. Nat. Herb. 20: 481, 1922.
Pouteria glabrifolia (Pittier) Cronquist, Lloydia 9: 282, 1946.

Tree to 25 m. Leaves with petioles 6-33 mm long; blades oblanceolate or
narrowly obovate to elliptic, often acuminate but sometimes acute or broadly
obtuse or virtually rounded, 6.5-30 cm long, 2-8.5 cm broad, glabrous, the sec-
ondary laterals subparallel, ± perpendicular to the primary laterals (the 3-4
most medial secondary laterals rather strongly arcuate and joining the midvein),
the reticulation fine, prominent to subobscure. Flowers 1-9 per leaf-axil; pedicels
5-17 mm long; sepals (4)5-6, ovate (sometimes very broadly so), appressed-
puberulent externally, 4-12 mm long, the innermost the longest; corolla cylindric,
7-16 mm long, sparsely to moderately sericeous-strigose externally (glabrous toward
the lobe-margins and on the lower portion of the tube), the tube comprising
Y 2 - 2 /z of the total length, the lobes (4)5(-7), oblong; staminodes narrowly linear-
lanceolate, often minutely papillose, 2.5-5 mm long; staminal filaments attached
slightly below the top of the tube, ca 1.5 mm long, the anthers ca 2 mm long;
ovary densely rufous- pubescent, usually 5-locular, the style 5-12 mm long, glabrous
except at the base but minutely papillose, the stigma subcapitate, usually 5-lobed.
Fruit edible, subglobose or pyriform, apiculate, brown to orange or yellow at
maturity, to 7 cm long and broad, the pulp orange or yellow; seeds 1-4, brown,
polished, 2-4 cm long, the scar 1-2 cm broad, extending the entire length.

Mexico (as far north as Colima and San Luis Potosi) to Panama; also in
Cuba and the Florida Keys but probably introduced.

canal zone: vie of Ancon, Piper 6027 (F, US), cocle: Penonome 8c vie,
56 (US), colon: vie of Donoso, Holdridge 6202 (MO), darien: Casaya I, Duke 10374
(MO); vie of Pifias, Duke 10589 (MO); forests around Pinogana, Pittier 6542 (holotype
of P. glabrifolia US; isotypes F, GH); 2 mi E of Santa Fe, Tyson et al. 4839 (MO).
Panama: Trapiche I, Allen 2609 (F), 2627 Duke 10320 (MO); Saboga I,

lanson 201 (US), 235 (US), 396 (US), Johnston 526
(MO, US), 733 (MO, US), 1171 (MO, US).

Though pointing to a close relationship, Cronquist (Lloydia 9: 282, 1946)
considered P. glabrifolia distinct from P. campechiana by virtue of its larger flowers
with corollas more loosely hairy on the back. However, I find nothing distinctive
in the corolla pubescence of the type of P. glabrifolia and, save for the slightly
larger flowers, it would seem to be a typical specimen of P. campechiana.

7. Pouteria chiricana (Standley) Baehni, Gandollea 9: 421, 1942.

Tree to 30 m. Leaves with petioles 6-17 mm long; blades elliptic or oblong-
elliptic to somewhat obovate, acuminate, 8-16 cm long, 2.5-6.2 cm broad, glabrous
or sparsely sericeous below along the midrib, usually conspicuously reticulate-
veiny beneath, the secondary laterals irregularly anastomosing, often slightly
raised below. Flowers subsessile in small axillary clusters; sepals 5, 2.2-2.6 mm
long, closely pubescent, the outer ovate, the inner oblong-spatulate; corolla tubular
or campanulate, 3-3.2 mm long, the lobes 5, ca 2 mm long, rounded; staminodes
small and inconspicuous; ovary 4-locular. Fruit ca 3.5 cm long, broadly ellipsoid,
smooth, 1 -seeded, narrowed to a ± definite, short and broad, stipitate base; seed
ellipsoid, ca 2.4 cm long and 1.8 cm broad, the seed-coat dull, with a fine raised

flora of panama (Family 154. Sapotaceae) 165

honeycomb pattern on the external surface, 0.5-0.7 mm thick, the scar oblanceolate,
ca 8 mm broad, extending the length of the seed.

Panama.

bocas del toro: region of Almirante, Cooper 457 (F). chiriqui: Progreso, Cooper &
Slater 230 (A, US), 254 (holotype F, isotype Y; not seen).

8.Pouteria subrotata Cronquist, Lloydia 9: 277, 1946.

Tree 10-15 m high, the limbs and branchlets erect. Leaves with petioles 1.5-
3.3 cm long; blades obovate to elliptic-obovate, acuminate, 10-23 cm long, 5-11.5
cm broad, glabrous above, glabrous or sparsely strigulose below (the trichomes
scattered and not concentrated along the main veins), not conspicuously reticulate-
veiny, the primary lateral veins prominent, 8-11 pair, the secondary laterals nu-
merous, fine, close, sinuous but regularly disposed and subperpendicular to the
primary laterals except near the midrib (there strongly curved). Flowers subsessile
(pedicels to 2 mm), clustered in leaf-axils and at defoliated nodes; sepals 5, ovate,
1.7-2 mm long, densely appressed-puberulent externally (the margins of the inner
sepals glabrous), more sparsely pubescent within; corolla white, subrotate, 3.5-3.8
mm long, the lobes 5, 2.5-3.1 mm long, ovate-oblong, sparsely to moderately ap-
pressed-sericeous externally except near the margins; staminodes linear-filiform,
ca 2 mm long; staminal filaments 1.7-1.9 mm long, attached at the base of the
corolla-lobes, the anthers ca 1 mm long; ovary densely appressed-white-sericeous,
5-locular, the style stout, 1.5-2.5 mm long, glabrous except at the base. Fruit
green or yellow-green, turning red, oblong-ellipsoid, 2-2,5 cm long, ca 1.2 cm broad,
1-seeded, the flesh scant; seed ellipsoid, ca 2 cm long, filling the fruit, the scar
8-9 mm broad, extending the length of the seed.

Panama (Darien).

darien: Rio Paya, Duke & Kirkbride 14073 (MO); forests around Pinogana, Pittier
6548 (holotype US).

9. Pouteria sclerocarpa (Pittier) Cronquist, Lloydia 9: 287, 1946.

Tree to 25 m. Leaves with petioles 1-2.2 cm long; blades rather narrowly
elliptic-obovate, acuminate, 10-25 cm long, 4-8 cm broad, glabrous, the primary
laterals 13-20 pair, the more marginal secondary laterals ± perpendicular to the
primary laterals, the reticulum slightly raised beneath. Flowers subsessile (pedi-
cels to 2 mm long), several per fascicle, the fascicles occurring at defoliated nodes;
sepals 6-7, ovate, the 2 outermost sepals appressed-pubescent externally, 3-4 mm
long, the inner sepals glabrate externally and 5-6.5 mm long, all sepals pubescent
within; corolla 6-8 mm long, the tube comprising ca % of the total length, the
lobes 5, ovate; staminodes ca 2 mm long, petaloid, narrowly ovate, subtruncate;
staminal filaments attached at the top of the corolla-tube, the anthers ca 1 mm
long; ovary 5-locular, densely ascending-sericeous externally, the style glabrous,
5-7 mm long, the stigma 5-lobed. Fruit yellow, sclerous, pyriform to obovoid,
somewhat narrowed toward the base, 5-7.5 cm long, 3-4.5 cm broad; seed solitary,
to 4 cm long and 2.5 cm broad, the large scar extending its full length.

Known only from the type collection.

san blas: plain of Sperdi, nr Puerto Obaldia, Pittier 4357 (holotype US, isotype MO).

166 ANNALS OF THE MISSOURI BOTANICAL GARDEN

10. Pouteria sambuensis (Pittier) Baehni, Candollea 9: 250, 1942.
Labatia sambuensis (Pittier) Pittier, Contr. U. S. Nat. Herb. 20: 481, 1922.

"Tree 10 m high; leaves glabrous, oblanceolate to obovate, acute at the base,
acuminate at the tip, mostly 20-25 cm long and 5-9 cm wide; primary lateral
veins about 13-15 pair; secondary lateral veins prominent, anastomosing, some-
what irregular; petioles short, about 1-1.5 cm long; sepals 4, presumably large;
other flower parts unknown; fruit sessile or subsessile, fleshy, densely hairy, ovoid,
about 5.5 cm long, 4 cm thick, obscurely sulcate from base to top."

Known only from the type collection.

darien: foothills of the Garagara Mts, Sambu Valley, Pittier 5621 (holotype US,

11. Pouteria stylosa (Pierre) Dubard, Not. Syst. 1: 381, 1909.
Lucuma standleyana Pittier, Contr. U. S. Nat. Herb. 18: 166, 1916.
Labatia standleyana (1 :. cit. 20: 481, 1922.

Pouteria standleyana (Pittier) Baehni, Candollea 9: 255, 1942.

Tree to 7 m. Leaves with petioles 3-10 mm long; blades elliptic to narrowly
obovate, acuminate, 7-20 cm long, 2.5-7 cm broad, glabrous above, finely and
closely appressed-sericeous below (the trichomes white to slightly fulvous) or
sometimes glabrate in age, the primary laterals 10-20 pair, the more marginal
secondary laterals perpendicular to the primary laterals, the more medial skewed
at about 45°, the reticulation more apparent above. Flowers axillary to leaves
and leaf-scars, 1-several per axil, sessile (often several borne on a low, woody
cushion); sepals 4 (2 + 2), 2.5-3.6 mm long, the outer broadly ovate and finely
rufescent, the inner rather narrowly oblong-spatulate and glabrous except down
the middle; corolla cylindric, glabrous, 2.5-3 mm long, the tube 2 / 3 of the total
length or more, the lobes 4, broad, subtruncate or slightly emarginate; staminodes
lanceolate, 0.7-0.8 mm long; staminal filaments attached at the base of the corolla-
tube; ovary densely ascending-rufescent, 4 to 6-locular, the style usually glabrous,
1-2 mm long, the stigma 4 to 6-lobed, the lobes readily evident. Fruit woody,
glabrous, ca 3 cm long and 3.5 cm broad; seeds 2, to 2 cm long and 1.6 cm broad,
somewhat compressed, the seed-coat 1-2 mm thick, firmly grown to the pericarp.

Panama; perhaps also Honduras, Costa Rica and Colombia.

bocas del toro: region of Almirante, Konkintoe 10 mi above Holstein, Cooper 509
(F). canal zone: area W of Limon Bay, Gatiin Locks & Gatiin Lake, Johnston 1820 (A,
MO); Mamei Hill, Pittier 3807 (holotype of Lucuma standelyana US, not seen; isotype
F). Panama: vic of El Cermeno, Allen 2572 (F), Zetek 4804 (F); Vista Alegre, Rio
Aguacate, 2 mi beyond Arraijan, Zetek 5511 (F, US), province unknown: s. loc, Hayes
67 (type, not seen).

Cronquist (Lloydia 9: 273, 1946) states that several sterile specimens from
Costa Rica and Honduras (Standley 40129, 54634, 55606 & 55816, all F) resemble
this species but have much larger (to 50 cm long and 20 cm broad) and less
markedly acuminate leaves possesing 20-40 pairs of primary lateral veins and
petioles 2-7 cm long. He indicates that these plants may prove to represent a
distinct species when better known. I have seen a similar specimen from Colombia
(Killipetal. 38307 US).

flora of panama (Family 154. Sapotaceae) 167

12. Pouteria caimito (Ruiz & Pavon) Radlkofer, Stizungsb. Akad. Wissens.

Miinchenl2:333, 1882.

Tree to 30 m. Leaves with petioles 5-17 mm long; blades obovate-oblanceolate
to elliptic, acute or acuminate, 5-24 cm long, 2-11 cm broad, glabrate, the sec-
ondary laterals without particular orientation, the reticulation close. Flowers
1-several per axil, sessile or subsessile (pedicels to 1.5 mm); sepals 4 (2 + 2),
3-4.5 mm long, ovate to oblong-ovate, the inner appressed-sericeous down the
middle, otherwise glabrous, the outer soon glabrate; corolla cylindric-urceolate,
glabrous, 4.5-8 mm long, the tube 2 / 3 of the total length, the lobes 4; staminodes
linear, flattened, ca 1.5 mm long; staminal filaments ca 2 mm long, attached at
the middle of the corolla-tube or slightly below, the anthers ca 1 mm long; ovary
4-locular, densely ascending-sericeous externally (the trichomes 1-2 mm long),
the style glabrous, 2-4.5 mm long. Fruit yellow, fleshy, globose to ellipsoid, 5-10
cm long, 4-8 cm broad; seeds 1-4, the scar extending the entire length.

Brazil and Peru to Trinidad and southern Panama.

damen: Sambu River, Pittier 5555 (F, US).

sapota (Jacquin) Moore & Stearn, Taxon 16: 383, 1967 7 .

(L.) Pierre in Urban, Symb. Antill. 5:97, 1904, auct. quoad

Tree to 30 m. Leaves clustered toward the branch ends, the branch ends often
fulvous-tomentose; petioles 1-4.5 cm long; blades oblanceolate or obovate, often
acuminate, 10-40 cm long, 4-14 cm broad, glabrate or tawny-sericeous below along
the midvein and primary laterals, the primary laterals 20-50 pair, the secondary
laterals essentially perpendicular to and connecting the primary laterals, more
prominent than the close reticulum. Flowers subsessile (pedicels to 2 mm long),
clustered at defoliated nodes, each cluster often borne on a small woody cushion;
sepals 8-12, spirally imbricate, ± orbicular, densely appressed-sericeous externally
except at the margins (there entirely glabrous), often emarginate or more deeply
bilobed at the apex, 2-6 mm long, the innermost the longest; corolla ± cylindric,
6-10 mm long, the tube comprising 2 / s - l / 2 of the length, the lobes 4-5, oblong-
obovate, appressed-sericeous except at the margins; staminodes linear-lanceolate,
2-3 mm long; staminal filaments attached at the top of the tube, 2-3.5 mm long,
the anthers ca 2 mm long; ovary densely ascending-sericeous, 5-locular, the style
3.5-7 mm long, often pubescent on the basal V3-V2, the stigma 5-lobed. Fruit
fleshy, ellipsoid or ovoid or subglobose, mealy-roughened, 8-20 cm long, brown,
the pulp yellow to red or pink, often milky; seed 1, ellipsoid, not compressed,
often 5-6 cm long, lustrous brown, the elliptic or obovate scar extending the entire
length, ca 2-2.5 cm broad.

Southern Mexico to northern South America and the West Indies; widely
cultivated; original natural range uncertain.

: below Chilibre, Seibert 1517 (MO, US); Balboa, Standley 26080
of Rio Tabasara, on hwy, Woodson et al. 441 (MO).

[Vol. 55
168 ANNALS OF THE MISSOURI BOTANICAL GARDEN

vie of Ola, POtter 5089 (US), herrera: rd betw Las Minas & Pese, Duke 12307 (MO).
panama: Laguna de Portala, nr Chepo, Pittier 4628 (US); Taboga I, Standley 27916 (US).
san blas: Ailigandi, Dwyer 6849 (MO); mainland opposite Ailigandi, from mouth of
Ailigandi River to 2.5 mi inland, Lewis et al. 163 (MO), province unknown: s. Inc.,
Duchassaing s.n. (F-fragment) .

Pouteria sapota, like several other members of the Sapotaceae valued for their
edible fruits or other uses, has numerous common names, the most frequent prob-
ably being "sapote." The fruit is a favorite among natives throughout tropical
America and is eaten raw or used to make sherbets, preserves and beverages.

14. Pouteria fossicola Cronquist, Lloydia 9: 289, 1946.

Tree 10 m high. Leaves ± clustered toward the branch tips; petioles 1.2-4
cm long; blades obovate, often acuminate, 8-30 cm long, 4-13 cm broad, finely
white-strigulose on the main veins beneath, sometimes also with a few similar
hairs scattered over the lower and occasionally the upper surface, the primary
laterals 12-20 pair, the secondary laterals regularly disposed, mostly perpendicular
to the primary laterals, decidedly more conspicuous than the fine reticulum. Flow-
ers clustered at defoliated nodes; pedicels ca 5 mm long; sepals ca 8, broad, densely
appressed-grayish-sericeous dorsally except for thin glabrous margins, rounded
or scarcely emarginate, spirally arranged, increasing the size centripetally, the
largest ca 6 mm long; other flower parts unknown. Fruit obovoid, fleshy, 8.5 cm
long, 4.5 cm broad, yellow-green, the surface irregularly wrinkled (at least in
herbarium specimens) and also mealy-roughened; seed 1, obovoid, not compressed,
7 cm long, 3.5 cm broad, the scar obovate, basally acuminate, 3 cm broad, extend-
ing the length of the seed, the seed-coat firm, shiny, yellowish-tan.

Known only from the type locality.

canal zone: Barro Colorado I, Bangham 583 (A, F), Salvoza 999 (holotype A),
Zetek 3870 (F).

15. Pouteria cooperi Cronquist, Lloydia 9: 291, 1946.

Tree ca 15 m high. Leaves with petioles 0.7-3.1 cm long; blades elliptic or
narrowly obovate, acuminate, 8-17 cm long, 3-7 cm broad, loosely hirsutulous
on the main veins beneath, in addition with a few scattered hairs on the lower
surface, the primary laterals 8-13 pair, the secondary laterals irregularly anas-
tomosing or occasionally ± perpendicular to the primary laterals, scarcely more
prominent than the coarse reticulum. "Flowers (separately preserved) apparently
sessile or nearly so; calyx of about 12 or perhaps more thick and fleshy broad
sepals, increasing in size centripetally, the inner about 4-5 mm long when dry,
all finely warty-wrinkled when dry, otherwise apparently glabrous, with narrow
thin margins; other flower-parts unknown; fruit reputedly woody."

Known only from the type collection.

bocas del toro: region of Almirante, Cricamola Valley, Holstein farm, Cooper 499
(holotype NY, not seen; isotypes F, US).

flora of panama ( Family 154. Sapotaceae )

Index of Latin Names
to descriptions; numbers ii

1 type refer to synonyms;

calcicola 147, 149t
zapota 145t, 149
Bumelia 150t, 152
panamensis 152, 154f
persimilis

Manilkara 146
bidentata 147
calcicola 147
chicle 147

Mastichodendron 157
. capiri 157t

meridionalis 149

[ BOTANICAL C

ANNALS
OF THE
MISSOURI BOTANICAL GARDEN

VOLUME 55

1968

NUMBER 3

CONTENTS

The Flora of Panama by Walter H. Lewis, The Flora of the Galapagos
Islands by Duncan M. Porter, The Illustrated Flora of Illinois by
Robert H. Mohlenbrock, Flora North America Project by Stanwyn
G. Shetler 1 7 ! - 1 78

Experimental studies of the species concept

by Edgar Anderson 179-192

Interspecific relationships in the Polypodium pectinatum-plumula
complex
by A. Murray Evans 193-293

Palynotaxonomic study of the Phytolaccaceae

by Joan W. Nowicke 294-364

Two new species of Polygala endemic to Panama

by Walter H. Lewis .365-367

Psidium (Myrfaceae) in the Galapagos Islands

by Duncan M. Porter 368-371

A revision of the Panamanian species of Rondeletia (Ruhiaceae)

by Joseph H. Kirkbride, Jr 372-391

A new species of Cucurbifa from Ecuador

by Hugh C. Cutler and Thomas W. Whitaker 392-396

N&TES 397-400

Hedyoth acerosa var. bigelovii, comb. nov. (Rubiaceae) 397. Shyrinchum
(Iridaceae), a new species from Texas and Mexico 397. Notes on
" an species of Voyria (Gentianaceae) 398-400. The First Jesse M.
ward 400.
Jme 55 „. 401-402

ANNALS VOLUME 55

1968

of the NUMBER 3

MISSOURI BOTANICAL GARDEN

A journal containing scientific contributions from the Missouri
Botanical Garden and the Department of Botany of Washington Uni-
versity in affiliation with the Missouri Botanical Garden. Outside
contributions in systematic botany and allied fields will also be con-
sidered. These papers are subject to a charge of $25 per printed page.

Editorial Committee
Walter H. Lewis, Editor

z Burch, Missouri Botanical Garden & Washington University
j D. Dwyer, Missouri Botanical Garden & St. Louis University
r M. Porter, Missouri Botanical Garden & Washington University

Contents of previous issues of the Annals of the Missouri Botanical
Garden are listed in the Agricultural Index, published by the H. W. Wilson
Company.

Beginning June 1, 1962 the Stechert-Hafner Service Agency, Inc., 31 East
10th St., New York 3, N. Y., became sole agent for the Annals of the Missouri
Botanical Garden. The Agency handles all subscriptions, all claims begin-
ning with volume 50 (1963) and all requests for back issues. Out-of-print
numbers will be reprinted as may be required.

Under the arrangement with the Stechert-Hafner Service Agency, the
lies also all purchases of any portion of the Annals subseries
the "Flora of Panama."

Printed by the Eden Publishing House, St. Louis, Missouri,

ANNALS VOLUME 55

1968

of the NUMBER 3

MISSOURI BOTANICAL GARDEN

THE FLORA OF PANAMA 1

by Walter H. Lewis

A pleasurable function Dr. Hugh Cutler gave me today is to introduce to you
our first speaker this afternoon, Dr. Duncan M. Porter, our new staff member in the
herbarium. But before we hear from him on the Galapagos Island project I want
to give you a little propaganda on our own floral project, the Flora of Panama. In
general style and approach the two Floras have much in common so I won't spend
time on an aspect Dr. Porter will cover, but what I did want to do is to give you
a brief history of the Panamanian project, its good and bad points and what value
it has been in stimulating related systematic programs.

Panama is an exceedingly rich floristic area with large elements of both the
Colombian and northern cordilleran floras represented. In its 47,000 square miles
we estimate no fewer than 8,000 species. How did the Garden become interested
in this piece of the tropics and what are some of the factors relating to the Flora
of Panama today?

The Garden founded a tropical station in the Canal Zone in 1926, and began
a series of expeditions from here in 1934. The first "Contributions toward a
Flora . . ." appeared in 1937 with Dr. Robert E. Woodson, Jr. as editor and assisted
first by Dr. Russell Seibert and later by Dr. Robert W. Schery. Seven Contributions
were published through 1943, chiefly from collections made from 1934 till 1941 by
members of the Garden staff (cf. Dwyer, Ann. Missouri Bot. Gard. 51 1 109-117,
1964). Earlier, collectors had concentrated their work in the Canal Zone or on
off shore islands so what we had in fact was the initiation of the Flora of Panama
in 1943, largely based on collections from the Canal Zone plus those from seven
years of general collecting, some of it continuous, such as that by Paul H. Allen,
but mostly by just periodic visits to Panama. Except for western Chiriqui and the
El Valle region, no mountainous area north and west of the Zone was then
conveniently accessible so of course little material was collected. The whole area

^he first of four papers and informal remarks on floristics given at the Missouri
Botanical Garden's 15th annual symposium on systematics during the afternoon session of
October 19, 1968. The symposium on "The Practical Values of Systematics" was chaired
by Dr. Donovan S. Correll, Texas Research Foundation, Renner, and organized by Dr. Hugh
C. Cutler of the Missouri Botanical Garden.
Ann. Missouri Bot. Gard. 55(3): 171-178, 1969.

172 ANNALS OF THE MISSOURI BOTANICAL GARDEN

south and east of the Zone to the Colombian border had hardly been visited
excepting by Pittier. My intention here is not to go into a long discourse on col-
lecting or the lack of it, but rather to give an illustration of when not to start a
flora; in this case with well over half of the country not yet visited by a botanist
and when the richest area, the mountains, remained virtually untouched. But the
Flora was started, ill-conceived in my opinion as a project for 1943, yet what is
really worse is that by virtue of its beginning, collecting essentially stopped and
was not reactivated until this decade, a lapse of nearly 20 years. It has become
quite clear to us what a tragic mistake this was as we now uncover dozens of new
species and dozens of generic extensions into Panama, particularly from South
America. I think the lesson was learned, albeit the hard way, for apart from the
quality of the Flora, which obviously suffered, we ignored the great stimulants to
all aspects of an herbarium's operation, namely collecting and exploration. Yet on
the basis of our collections since 1959 material has been almost doubled for families
still to be treated in the Flora, With them we obviously have a better understand-
ing of distribution and ecology of taxa, and a better opportunity for making correct
taxonomic judgments.

One very positive effect of the Flora both during the non-collecting and current
phase has been its role in graduate education. Too few floras, particularly those
involving the tropics, have been a part of the academic world where stimulus to
graduate training and graduate exposure to floristics is maximumly possible.
Fortunately, because of our close association with Washington University, we are
currently managing direct graduate participation by insisting that students join one
collecting expedition to the tropics, and in our advanced course in taxonomy, that
students choose to study a small family as it occurs in Panama. Such exposure
may not be very great but we think it sufficient for a student to develop an ap-
preciation for what is involved in writing a flora, often his only direct experience
with floristics during a graduate career.

Apart from collecting and training as significant activities interwoven with
our floral project, we have several others I want to mention only briefly. One has
been our small help to the University of Panama in establishing their new her-
barium (PAN). Miss Mireya Correa, its first curator who was trained at Duke
University, arrived in St. Louis yesterday for about two months of study and de-
termination of her collections. This perhaps best illustrates our desire to stimulate
taxonomic resarch within Panama and to have a long-ranged program based at the
University of Panama, now a reality of just a few months.

We have for several years discussed the feasibility of including all green ter-
restial plants in the Flora and thereby include the bryophytes. Our new Curator
of Cryptogams, Dr. Marshall Crosby, will assume this responsibility as soon as our
bryophytes are recurated and reorganized. Dr. Crosby comes to us from Duke Uni-
versity where he worked under Dr. Louis Anderson.

The last example I shall mention is the project of Dr. Thomas Croat who,
in collaboration with the Smithsonian Tropical Research Institute in the Canal
Zone, is doing a new concise manual of Barro Colorado Island. Dr. Croat has
spent about five months on the Island during the past year collecting and making

FLORISTICS 173

valuable notes on this typical rainforest community (he returned just two days
ago brandishing a major leg wound inflicted by a charging peccary!). The manual
he hopes to produce will include much needed biological data such as times of
flowering and fruiting backed by complete herbarium specimens. I think it is a
sad commentary of our manuals from the area that often one cannot associate
fruiting material with the flowering specimens; flowers of course are needed for
most keys and all too often fruiting specimens just remain undetermined. Dr. Croat
plans to make a series of keys for the dominant taxa: to mature fruits, to vegetative
parts alone and so forth. These ought to be very helpful for anyone working in
the area including, for example, the mammologist who wants to know what his
animals are eating . . . and as a matter of record, to help him find an answer
provided the initial incentive for this manual. What the program is doing for the
Flora of Panama is obvious, but what it will do as a practical guide to the rain-
forest plants of Central Panama is even more important.

Here then are our activities associated with the Flora of Panama, but the hard
work of putting many of the familial treatments together rests with our new
Curator of the Project, Dr. Duncan M. Porter. Dr. Porter comes to us from the
Galapagos Islands project under Dr. Ira L. Wiggins, he taught at the University
of San Francisco and has his doctorate from Harvard University. It is my distinct
pleasure to introduce to you our new Curator of the Flora of Panama and Assistant
Professor of Botany at Washington University, Dr. Porter.

THE FLORA OF THE GALAPAGOS ISLANDS 1

The Galapagos Islands lie astride the equator in the eastern Pacific Ocean,
about 500 miles west of Ecuador. This archipelago consists of some 15 larger
islands and numerous islets and rocks, covering a land area of approximately 3,000
square miles. The largest island is 75 miles long and 45 miles wide at its broadest.
Several of the mountains of the group reach to a height of over 5,000 feet, although
most of the land is relatively low.

The islands in reality are immense piles of lava projecting out of the ocean,
covered with craters, fumaroles, vents, and other volcanic phenomena. Great lava
flows extend from the crater rims to the sea, and the area is one of the most vol-
canically active in the world.

About 1,200 people inhabit the Galapagos. They are concentrated in the few
spots where there is either permanent water, or there is sufficient and constant
enough rainfall to collect and store in barrels. Water is the great limiting factor

174 ANNALS OF THE MISSOURI BOTANICAL GARDEN

on the islands for all forms of life. Permanent water holes are few and far between.
The plants depend mainly upon the sporadic rain that usually falls from January
to April or May, although plants at higher elevations are bathed in fog almost
daily. Most animals obtain their water by eating the plants. Both this scanty rain-
fall and the relatively cool climate (the thermometer rarely rises above 85°F) are
due to the Humboldt Current, sweeping up from the south along the western edge
of South America and bathing the Galapagos shores with the cool waters of the
Antarctic Ocean.

Scanty and sporadic rainfall, a relatively cool climate, and drying winds have
here combined to produce literature's classic desert islands. Their vegetation is
much sparser than one would guess from their geographical position, and the
number of species, subspecies, and varieties of vascular plants known to occur here
is roughly 700.

Two so-called "floras" have been published for the Galapagos Islands in this
century: that of B. L. Robinson (Flora of the Galapagos Islands, Proc. Amer. Acad.
38: 77-270, 1902) and that of Alban Stewart (A botanical survey of the Galapagos
Islands, Proc. Calif. Acad., ser. 4, 1: 7-288, 1911). In reality both of these works
are merely annotated checklists, giving only some synonymy, locality data, and in
only a few cases any plant descriptions, and these minimal. At present there is no
single publication to which one with an interest in the plants of the Galapagos
Islands may refer in order to key out specimens from the archipelago or to determine
► be applied to the plants growing there.
\ William Beebe wrote several sentences in his book Galapagos: World's
End (Putnam, N. Y.) that still apply to the situation in 1968: "A very thorough
list of plants has been published, giving exact distribution, elevation, ecological
summary and relative abundance, but, except for the technical names, no hint as
to the growths themselves. Professor Wheeler and I were very anxious to identify
certain plants because of their relationships to animal organisms, but found it
impossible from published works on the Galapagos. ... If explorers would give
just a thought to others who might come after, and take the time to write out the
simplest kind of a key, it would be a kindness beyond gratitude."

Beebe is about to have his wish. Since July 1966 I have been privileged to be
associated with Professor Ira L. Wiggins of Stanford University in the writing of
a complete flora for the vascular plants of the Galapagos Islands.

The Flora of the Galapagos Islands project was instigated by the late Dr. E.
Yale Dawson, who was very much interested in the cacti of the archipelago. Upon
Dawson's leaving the San Diego Natural History Museum in late 1964, and the
consequent loss of institutional backing for his proposed governmental grant,
Wiggins took charge of the Flora. Research on the bulk of the project was carried
out after Wiggins had taken it over. The project was funded by the National
Science Foundation and administered by the California Academy of Sciences in
San Francisco. I was employed by the project for 13 months as a Postdoctoral
Assistant.

The primary function of a flora is to enable a person unfamiliar with the
plants of an area to identify them and to determine the names to be applied to

FLORISTICS 175

them. This we have attempted to do. However, this should not be taken to imply
that The Flora of the Galapagos Islands is only a handbook or manual. It is a
comprehensive study of the vascular plants occurring in the archipelago. Not only
does the Flora include a detailed description of each taxon of vascular plant known
to occur on the islands, it also includes an illustration of at least one representative
of each genus, keys to all taxa, complete synonymy (or a reference to where it may
be found), citations of herbarium specimens examined, geographical distributions,
notes on variation and relationships, and ecological data for every species, sub-
species, and variety treated.

Our chief concern has been with the plants as they occur in the archipelago.
In the main, descriptions are based only upon specimens actually collected in the
Galapagos Islands. The overwhelming majority of Galapagos collections are in
the herbaria of the California Academy of Sciences, Harvard University, and
Stanford University. Specimens in these herbaria and several others have been
utilized in the preparation of the Flora. Field studies in the archipelago in 1964
and 1967 have added materially to previous collections and have helped us to
gain a first-hand knowledge of the plants.

Plants known to occur in the Galapagos Islands are not the only ones included
in the Flora. Notes are provided on species reported from the islands, but which
are not represented by any known collections. However, such species are not in-
cluded in the keys.

A number of cultivated plants also are included in the Flora, Some of these,
such as Psidium guajava (the cultivated guava), have escaped from cultivation and
become established in the native plant communities; others, like Tropaeolum
majus (the garden nasturtium), have the capacity to do so; and still others, like
Mangifera indica (the mango), may persist after cultivation has ceased and the
native vegetation has moved back into the previously disturbed area. Such informa-
tion is not usually to be found in a flora, unless an escape is able to reproduce
itself in the wild. However, the non-taxonomist in particular is invariably inter-
ested in every plant that catches his eye, and the spectacular flowers to be found
on many cultivated plants make them the first noticed by many.

Thus, The Flora of the Galapagos Islands is written not only for the specialist.
A conscious effort has been made to provide a useful guide, not merely for the
professional taxonomist, but for anyone interested in the Galapagos flora, whether
he be an ecologist, a geologist, a herpetologist, an ornithologist, or a plant taxono-
mist; whether he be in the laboratory or in the field.

1/b ANNALS OF THE MISSOURI BOTANICAL GARDEN

THE ILLUSTRATED FLORA OF ILLINOIS

by Robert H. Mohlenbrock
Southern Illinois University, Carbondale

A multi-volumed work on the complete flora of Illinois is under preparation
at Southern Illinois University; I am editor and chief contributor to the series.
The work will provide an account of every kind of plant known to occur in Illinois,
except bacteria and certain groups of fungi. Keys for identification will be provided,
along with the descriptions, synonymy, discussions of ecology and economic uses,
chromosome numbers, and distribution maps for each species. In addition, line
illustrations of each species, showing diagnostic features, are being prepared by a
staff of illustrators. Publication will be by the Southern Illinois University Press.
Financing for the project, thus far, has been by the University Press and the
Graduate School of Southern Illinois University. A board of advisers has been
created to screen the manuscripts and serve as consultants for the project. The
board is composed of Dr. Gerald W. Prescott, Michigan State University; Dr. C. J.
Alexopoulos, University of Texas; Dr. A. J. Sharp, University of Tennessee; Dr.
Rolla M. Tryon, Jr., The Gray Herbarium; and Dr. Robert F. Thorne, Rancho
Santa Ana Botanic Garden. The first volume dealing with the ferns of Illinois
appeared in April, 1967. Two volumes on monocots (excluding grasses and sedges)
are scheduled for release around August 1, 1969. The storage of the entire flora
on magnetic tape is being investigated; computing techniques are already being
employed in the recording of distributional data.

FLORA NORTH AMERICA PROJECT

by Stanwyn G. Shetler
Smithsonian Institution, Washington, D.C.

Plant taxonomy serves an eminently practical function through floristics.
Without the synoptical guides and floras that are the products of floristic research,
neither the biologist nor the layman would be able to identify the plants of the
earth and to know where each grows. Such diagnostic information is basic on the
one hand to further scientific research and on the other hand to wise exploitation
of our plant resources to meet human needs.

North American taxonomists have just undertaken an immense, new, co-
operative effort to compile and integrate our knowledge of the plants of this
continent. This effort, estimated to take about 15 years, is called the Flora North
America Project. Taxonomists hope that it will become their "Project Synthesis"
of the 20th century, providing the organizing theme and the intellectual rationale
for the broader, more cooperative approach to floristic-taxonomic research that is
necessary to meet the challenges of our ecological age, when society is becoming
increasingly more conscious of its natural environment and of the need to control
the quality of this environment. Public officials and laymen alike are depending

FLORISTICS 177

on the biologists more than ever for instant facts and correlations concerning our
plant and animal resources. Flora North America should fill a great void by provid-
ing for the first time in history a concise encyclopedia of all vascular plants in the
whole of North America north of Mexico. The preparation of this flora will be
accomplished through the cooperative labors of many taxonomists, organized and
guided by several committees.

Apart from the preparation of the flora itself, a second, supporting objective
is to place floristic-taxonomic research on a modern, computerized, information-
system basis. FN A must be a computer-age treatise— a taxonomically-structured
data bank with an associated package of computer programs ("flora software") to
serve as a "kit of tools" for use in accessing and manipulating the data.

The third, intangible goal is to stimulate new interest in taxonomy and the
study of the higher levels of biological integration— organisms, species, populations,
communities, floras — especially among the coming generations of students who
should find in the FNA Project the philosophical rationale, unifying and organizing
genius, and advanced technical resources to conceive and carry out taxonomic
researches on a scale hitherto impossible.

The time is overdue for plant taxonomists, who in a sense have always been
the keepers of the botanical data bank, to develop a modern storage and retrieval
system using the latest methods of electronic data processing and communication,
so that they can regain their historic control of the data bank. Publication has
exploded such that it no longer is possible to integrate new information rapidly
and concisely by conventional methods and media.

Floras and systematic monographs are time-honored, if non-automated, storage
and retrieval systems, which through the years the taxonomist has produced as
both the means and the ends of his research. In the FNA Project we propose to
computerize such systems and thereby to update and extend the concept of the
flora or monograph to cover not only the traditional printed book but also today's
dynamic electronic data bank. Output from a data bank can take many forms
and present many different kinds of data. FNA output, in addition to the basic
identification manual, may include distribution maps, illustrations, glossaries,
indices, gazeteers, descriptions, ecological summaries, statistical calculations, phe-
netic diagrams, and research reports. Thus, FNA will not be a single publication,
embalming the state of knowledge. Instant update, revision, and cumulation will
be cardinal attributes of the automated FNA information system.

It is expected that the FNA data will be organized into a series of matrices
which lend themselves to easy manipulation by computer. Computer programs
are being developed to be used as a package or "black box" by taxonomists for
performing automatically many of the purely mechanical chores of compiling,
correlating, and editing floristic-taxonomic data. Some of the programs will enable
the initiated user to construct by machine his own, up-to-the-minute identification
keys, descriptions, maps, etc., from the data matrices. Time-share use of the
computer will be exploited for on-line identification of plants from remote termi-
nals. Data matrices and flora software will be constructed to serve not only the
scientific research purposes of the taxonomists but also the practical needs of other

178 ANNALS OF THE MISSOURI BOTANICAL GARDEN

scientists and laymen. FNA should help greatly to bridge the gap between
scientist and layman. The electronic methods also should eliminate or simplify
many of the routine labors which dilute the basic intellectual content of taxonomy.
Then the taxonomist can be more scholar and less curator than in the past.

EXPERIMENTAL STUDIES OF THE SPECIES CONCEPT

This review is dedicated to Dr. J. M. Greenman, former Curator of the Mis-
souri Botanical Garden Herbarium and my colleague during much of the time
these studies got underway. In answering my naive questions about the species
problem, he frequently quoted to me the statement "Species are but judgments."
It was due to him that I became fascinated with the problem of finding factual
evidence as to the ways in which such judgments are formed.

Figure 1 illustrates two extreme uses of the word SPECIES in discussions of
the species problem. Such discussions are often at cross purposes because those
participating are unaware of these differences.

Fig. 1. Diagram of two extreme uses of the word Species.

The diagram shows three taxa of one genus, they may be species, subspecies,
or formae but each of the three is represented by individual plants or animals
which have certain features in common. This is symbolized in the diagram by
5 individuals of taxon A with no markings, 5 of taxon B with dots and 5 of taxon
C with crosses. At the top of the figure are symbolic representations of two gentle-
men, Mr. X and Mr. Z, each of whom has been studying the 15 specimens. Above
Mr. X's head is symbolized his concept of the taxonomic relationship of A, B, and
C. Mr. Z's concept is diagrammed in a similar manner. The diagram would be
exactly the same if the two men were in perfect agreement as to the classification
of the 15 specimens or if they disagreed in part, or completely. Mr. X, for example,
might feel that A with no markings should be species A, and that B and C were

Ann. Missouri Bot. Gard. 55(3): 179-192, 1969.

1BU ANNALS OF THE MISSOURI BOTANICAL GARDEN

differing forms of that species. Mr. Z might think that all the differences among
the 15 specimens were too minor for taxonomic recognition. In any such example,
however, in discussions between Mr. X and Mr. Z, the word "species" might (1)
be referring to groupings of the individual specimens of A, B, and C which merited
recognition as species or (2) the word "species" might be equated to the concepts
X and Z which had matured in the minds of the two men and in which examina-
tions of A, B and C might or might not have played a part.

These matters are well known to most authorities on the species problem.
Few of them have made experimental analyses of the species as a concept, but
these prejudices are now breaking down rapidly.

Eight such papers were published between 1954 and 1966. They are reviewed
below in the order in which they were published. In quoting from those papers
of which I was one of the authors, the original text has been shortened but in no
case has the sense been changed, though slight differences in shades of meaning
are unavoidable in radical condensations.

(1) Speciation in Uvularia, by Edgar Anderson & T. W. Whitaker (1934).
This paper was (pp. 30-31) "an attempt to present objectively in a codified form
the essential facts as to resemblances and differences within and between two
similar but distinct species; to reproduce in a concise manner for non-taxonomists
the kind of data which are consciously or unconsciously used by taxonomists in
the delimitation of species. If a species cannot as yet be defined in terms that
are meaningful to workers in other fields, one can present the range of variation
within and between two closely-related species in a summarized form for non-

"It is not a taxonomic thesis. The two species were chosen for study because
there was general agreement that they were specifically distinct from one another
and were closely related species of one genus." By diagrams to scale, objective
evidence was presented that Uvularia perfoliata and U. grandiflora had four meas-
urable differences in the leaves, five in the nodes of the stem and inflorescence
and four in the flowers. Of all these, only three were discontinuous; the acknowl-
edged discontinuity between the two species is "a discontinuity of combinations
reinforced by a few truly discontinuous differences."

It does not seem to be generally realized that species may be, and customarily
are, thought of in different ways by different groups of biologists. Biologists en-
gaged in taxonomic work will think of species in terms of the precise differences
that permit their ready classification; that make it possible to arrange species in
an herbarium or to construct a key to a genus. To them the essential differences
between these two species of Uvularia will be those few discontinuous ones that
are ordinarily used in identifying the species.

"With this attitude we have no quarrel, recognizing systematics as a difficult
and necessary business and that those who have it in hand must be allowed to
develop their own methods. Yet there are those who are interested in the biologi-
cal makeup of the units which are being classified. This group will include some
taxonomists; the separation of the two kinds of thought is not absolute. To us

ANDERSON — EXPERIMENTAL STUDIES OF SPECffiS CONCEPT 181

the difference between two species is the difference between one kind of germplasm
and another. As geneticists we have gained an impression of two different life-
stuffs, each of which reacts variously with the environment. Individual plants
produced cooperatively by the germplasm and the environment will show only
one facet of that germplasm. For the full expression of a particular species there
will be required a series of individuals produced under various environments."

In presenting evidence for differences between the two species of Uvularia
in their branching patterns and leaf shapes and in the internode patterns of their
stems, a combination of measurements and standardized glyphs was used (p. 32).
These were developed into a series of charts for graduate classes and lectures at
symposia and for the general public. They are discussed below in reviewing the
fifth paper.

The first paper in the field of this review presents little objective evidence
about the concepts in the minds of systematists, but the diagrams (Fig. 1) do
analyze the kinds of evidence that are before ones eyes when working in an her-
barium, a museum, or an experimental plot. Taxonomists certainly differ greatly
in their ability to form useful "judgments" from such subtle data as trends in
shape. Greater skill in that direction is probably responsible for the "taxonomic
intuition" of such taxonomists as George Engelmann. His judgments about the
classification of western American plants were derived mostly from tiny scraps of
material, but they have stood the test of time.

(2) An experimental study of hybridization in the genus Apocynum, by Edgar
Anderson (1936).

In his Doctor's thesis my colleague Dr. Robert E. Woodson published an
interpretation of the phylogeny of the genus Apocynum which was decades ahead
of its time. He envisaged it as a genus in which inter-specific hybridization has
been so important that its evolutionary pattern is more like an anastomosing net-
work than a branching tree. To me his ideas "though stimulating and interesting,
seemed rather in need of experimental confirmation by other than purely mor-
phological criteria. After much friendly argument an experiment was planned,"
a simple progeny t?st of two common American species, Apocynum cannabinum
and A. androsaemifolium, strikingly different plants, and their putative hybrid,
Apocynum medium. Woodson gathered seed from several different plants at a
site in Indiana where all three species grew near one another, and I made supple-
mental collections in New England. I supervised raising the seedlings to flower-
ing age, scoring pollen fertility, photographing the flowers and making herbarium
specimens of each mature flowering plant (Fig. 2).

Since I was then working in Boston and Woodson in St. Louis, it was easy
to get precise, objective data on his concepts of speciation in Apocynum from his
labels on the 40 reticulately-related Apocynum specimens I sent him. They had
all been collected the same day and were tagged with a randomized set of num-
bers, the only key to which was in my record book. Woodson had no way of know-
ing how the specimens fitted together to make 8 progeny tests until I sent him a

ANNALS OF THE MISSOURI BOTANICAL GARDEN

A. androsaemifolium

A. androsaemifolium

A. androsaemifolium

A. androsaemifolium

A. androsaemifolium

A. androsaemifolium

very typical
fairly typical

androsaemifoliu

before their determinations were recorded

glabrous leaves
like a small androsaemifoli
glabrous

sparsely pubescent
glabrous
glabrous
unusually small flowers

Probably a glabrous A.
A. medium

medium

A. medium

A. medium

A. androsaemifolium

A. medium

but very small flowe

quite typical
but nearly glabrous

androsaemifolium?

androsaem ifolium
androsaemifolium?

corolla rather small

ANDERSON — EXPERIMENTAL STUDIES OF

Fig. 2. Progeny tests of three species of Apocynum. Each flower was chosen i
representative for one plant, all X y 2 . Left, A. androsaemifolium; Center, A. mediun
Right, A. cannabinum.

rough copy of the information in Table 1. It confirmed all his hypotheses, in-
cluding some I had been skeptical about.

Table 1 demonstrates that all the 6 aberrant specimens that he labelled with
questions marks, as well as all the 13 which were so variant that he added in-
formal comments to the label, were seedlings of A. medium, the supposed hybrid,
thus indicating it to be of hybrid ancestry. Table 1 also presents evidence for a
phenomenon that I do not remember having talked, read or thought about up to
that time, the various restrictions to free recombination of multiple-factor char-
acters which operate in hybrid germ plasms. The total effect of these restrictions
in Apocynum is so strong that two A. medium seedlings, 449-4 and 504-4 were
labelled by Woodson as unquestioned A. androsaemifolium and one, 446-11 as
unquestioned A. cannabinum var. glaberrimum! The paper concluded by suggest-
ing that "the chief effect of hybridization in this genus in eastern North America
at the present time is to increase variability in the parental species."

It was these experiments with Apocynum which lead me to examine the gen-
eral restrictions to recombination in multiple factor characters and eventually to
describe, define and diagram introgressive hybridization.

(3) The Concept of the genus II. A survey of modern opinion, by Edgar
Anderson (1940).

From discussions with my colleague Jesse M. Greenman and our students,
I became increasingly interested in species as a concept. I sensed that there might
be a "Genus Problem" as well as a species problem. Accordingly, when a na-
tional symposium on Genera was organized in 1938, I sent out a questionnaire
asking in two different ways a basic question about genera and species. The ques-
tionnaire was set up to allow any one of five different answers to the first question
and four to the second one and was mailed to 43 taxonomists.

[VOI
184 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Question 1 No. of re\

Genus the more natural unit 26

Species the more natural unit 8

! other 11

No opinion
Question meaningless

Question 2
Genera originate in the same way ;
Genera may originate in a different
Genera may originate in same or i

Forty-two taxonomists responded, so many of them with additional <
that a negative correlation between age of respondent and interest in the symposium
could be demonstrated in tabular form (see below).

The replies were more uniform than some of the respondents had expected.
Twenty-one of them were identical. This orthodox opinion was that genera, on
the average, are more natural groups than species, that they originate in the same
way, and that generic differences could be compounded from specific differences.

By dividing the respondent into two groups: one, taxonomists whose experi-
ence had been mostly in monographic work, and group two, taxonomists who
were not monographers or who had extensive experience in other biological dis-
ciplines, this correlation could also be tabulated. Of the monographers, two-thirds
were "orthodox"; of the non-monographers less than one-third. However, though
I was still under 40 when the questionnaires were mailed out, I felt that the "genus
problem" required extensive experience with more than one group of organisms,
to answer such questions intelligently. I agreed with Liberty Hyde Bailey when
he answered "A fair agreement has been reached as to the limits of genera and
the limits of species, without much reference to philosophical considerations.
Discussion is likely to be made by persons who have no taxonomic training and
the conclusions would be of little practical value." C. W. Weatherby, then the
assistant curator of the Gray Herbarium, appended to his answer a short distillation
out of his own experiences that has important c

INTEREST OF RESPONDENT Und

Not in sympathy 2 2

Replied without comment 2 7 7 16

Replied with discussion 6 6 5 17

Replied and expressed interest 12 1 1 14

Total 20 14 15 49

"Taxonomy is only a glorified guess — an attempt to construct a cross-section of the
lines of descent in a form intelligible to the human mind. It always contains two
variable quantities — the plasticity of organic nature and the differing points of
view of the people who work at it. You can generalize successfully, if at all, only

185

by keeping these facts constantly in mind. The only general rule is that there is
no general rule. Therein lines the fascination of taxonomy for those who like it.
Each group one tackles presents a fresh and original problem: for each, one has
to work out anew the method by which he may best achieve that transforming of
confusion into order which is the great satisfaction of pure taxonomy."*

When the replies to the questionnaire arrived they were of such interest to
taxonomists and other biologists that they were bound and filed in the Missouri
Botanical Garden Library. Unfortunately this volume was not put in the locked
cases which house the Pre-Linnaean Library and other treasures. It was read so
frequently that it became dog-eared. Twenty years later it disappeared from the
shelf.

(4) Concordant vs. discordant. Variation in relation to introgressive hybridiza-
tion, by Edgar Anderson (1951).

The above title is misleading for the purposes of this review. The extensive
experimental studies of species concepts reported in the preceding review demon-
strated great variation between able biologists in perceiving leaf shape and inter-
node-pattern differences. This paper first presents evidence for the taxonomic
validity of these characters. With mathematical precision it then constructs a
demonstration of the differences in appearance to be expected between populations
of plants in which extensive introgression has or has not taken place. The paper
belongs in this review because it presents evidence about the importance of the
complex differences it refers to as "trends", as measures of species differences in
plants and animals.

All taxonomists tested were aware of leaf-shape differences, though most non-
taxonomists were more conscious of size than of shape differences. Correlated
changes in size and shape, however, had proved so difficult for them to grasp* that
a large half of this paper is a demonstration of that relationship with tracings,
diagrams, and discussions.

The first evidence presented is leaf shapes of two common wild cherries of
the eastern United States, the choke cherry, Prunus virginiana and the rum cherry,
P. serotina. "The outstanding difference is in the trend in proportion. As the
leaves of Prunus virginiana get longer, they get correspondingly broader. As the
leaves of P. serotina get longer they get only slightly wider. In P. serotina, there-
fore, the largest leaves are the narrowest, the smallest the broadest; in P. virginiana
the largest are the broadest, the smallest the narrowest. Such differences in trend of
related parts seem to be generally characteristic of specific differences. I found such
differences in trend in all the species (of several genera) for which I had made
pressed population samples."

Trends in internode patterns became easy to demonstrate and to record
precisely by a method originated by a botany major, Dorothy Schregardus. She
used it to analyze the growth and development of the common sunflower and it

l apprehended them, i

186 ANNALS OF THE MISSOURI BOTANICAL GARDEN

was published as Anderson & Schregardus (c), a method for recording and analyzing
variations of internode pattern. The internodes of the stem were diagrammed along
the horizontal axis at regular intervals from left to right, 1, 2, 3, 4, etc. as in a
scatter diagram of correlation. The length of each internode was then diagrammed
vertically in centimeters to an appropriate scale, with a dot. The successive dots
were connected with straight lines whose slope was a precise record of the rate of
increase or decrease from one internode to the next for the internodes of the entire
stem. This simple device has been effective in various studies of plant develop-
ment and in making exact measures of plant-to-plant differences when interpreting
the variation of introgressing populations.

Its most extensive use has been in classifying Zea mays. Additional symbols
were added to these maize-internode diagrams. A solid black circle marked the
position of the last internode below the male inflorescence (the "tassel") and a
solid black triangle the position of the corn ear, or of each ear in multiple ear
varieties and races. Such diagrams, with minor variations, have been used in more
graphs on the races of maize in Brazil, Cuba, Colombia, Central America, the
West Indies, Chile, Peru, and Bolivia* sponsored by the National Academy of
Sciences and the National Research Council. They are proving useful in develop-
ing hybrid corn for Latin America and in the tropics and sub-tropics generally.
They demonstrate how a concern for understanding the concepts by which taxono-
mists evaluate differences between species grew into the use of basic physiological
differences in classifying economic plants.

(5) Efficient and inefficient methods of measuring specific differences, by Edgar
Anderson, (1954).

This reprint from Kempthorne, Bancroft, Gowen and Lush is included be-
cause, though it was prepared for an audience of statisticians and mathematicians,
it contains the only published evidence of my most extensive experimental investi-
gations of the species as a concept. As in the first paper reviewed, Uvularia fur-
nished the research material. "To get at the fundamentals of the species problem,
one needs to choose the simplest possible species. Uvularia grandiflora and U.
perfoliata were chosen for demonstration because there has been universal agree-
ment among those who have dealt with them that they are both good species, that
they belong to the same genus, and that they present no special problems of
classification, distribution or ecology. Both are known to be diploids; there are
no indications of other complicating factors. These two species differ in the
presence or absence of hairs on the underside of the leaves and by curious glandular
outgrowths on the inner face of the perianth in U. perfoliata that have no counter-
part in U. grandiflora." They also differ by many minor, more-or-less-correlated
differences in the size, proportion, texture, number, and arrangement of the various
internodes, leaves, and scales that make up the vegetative parts of the plant that

* See for example, Races of maize in Bolivia, Ramirez et al., in collaboration with G.
Edward Nicholson Calle, Edgar Anderson, William L. Brown. 1960. Publication 747: Nat.
Acad. Sci.— Nat. Res. Council, vii + 159 pp.

ANDERSON EXPERIMENTAL

OF SPECIES CONCEPT

187

are above ground. To analyze the nature of these differences it was necessary to
choose a few from among the hundreds of sense impressions coming to us from
each plant. The length of every leaf and the length of every internode was
recorded, as well as the position of every flower and every scale-leaf (cataphyll).
From these measurements and scores, an "ideograph", a sort of diagrammatic skele-
ton of each specimen, was reconstructed (Fig. 3). The width of the leaf, the angle
of branching, the sizes of the flowers, and of the cataphylls at the base of the
stem, were all conventionalized. Flowers are represented as black disks, leaves as
wide lines but to scale, cataphylls as short narrow lines. The length and number
of internodes is indicated by the positions of the leaves and cataphylls. The varia-
tion of all these variables was analyzed mathematically. Uvularia grandifiora was
shown to be "represented by a coherent group of individuals." Uvularia perfoliata
was not so coherent. Some of the specimens vary in the direction of U. grandifiora,
that introgression from that species, previously unsuspected, may be
These conclusions were confirmed and extended by
Dietz(1952).

Experimental studies of the perception of these species-differences were made
with reproductions of the 20 ideographs on plain white cards, a little larger than
playing cards, or with poster-size enlargements for public lectures. People being
tested were usually not told how many species were represented. It was found that
biologists differ greatly in ability to separate the two species correctly from these
cards and in the speed with which they can do it. At one extreme was a graduate
; anatomy. With no previous experience, he sorted all twenty

V

V \ :

vvv

v/

y

species of Uvularia, above U.

188 ANNALS OF THE MISSOURI BOTANICAL GARDEN

out correctly in less than ten minutes. At the other extreme was an able monogra-
pher and museum curator. He studied the cards carefully and laid them out in
little groups of almost identical ideographs. One or two of these he was able to
unite, but most of his units he placed halfway between two others until the whole
table-top was covered with a complex web-like arrangement of little groups of
cards. He was intrigued by the challenge and kept at the problem intermittently
for hours with no success. I suspect that he had more of the basic facts about each
specimen in his mind than anyone else I ever tested, but that he had no instinctive
way of ignoring maturity differences when looking for specific differences. I suspect
also that he was not facile in apprehending differences in internode patterns. When
the method of diagramming internode patterns described in the following review
was adapted to the Uvularia ideographs, it produced diagrams which were easier
for students to classify correctly.

From testing students with these ideographs, and from observing taxonomists
at work, I believe that there are innate differences between people in their ability
to apprehend significant clusters of variables as indicators for malnutrition, ma-
turity, specific differences, etc. Some of those most highly endowed are unaware
that they have exceptional gifts in that direction. Able scientists with little of this
ability can be helped by a good teacher. Some able taxonomists with little of it,
and that little not well-developed, are prejudiced in appraising the work of those
who are highly endowed.

(6) An analysis of variation in a variable population of Cladonia, by Edgar
Anderson and E. D. Rudolph (1956).

This analysis of introgression between species of two lichens is included in
these reviews because the same specimens studied by the authors were sent to an
authority on Cladonia, identified only by numbers, for precise identification. His
groupings of the specimens and the authors' are in close agreement. The authors'
groupings confirm the generalization of the second review that one of the important
results of hybridization is enrichment of variation in the parental species. The
contrast between his concept of speciation and their approach to the problem is
shown in detail in Figs. 7 and 8. They demonstrate his concepts as well adapted
to cataloguing such a hybrid swarm, and the authors' as adapted to analyzing its
evolutionary dynamics. This also justifies the paper's introductory remark "that
populations which suggest hybridization to the experienced eye have much the
same variation pattern as back-crosses of experimentally produced hybrids."

More than any other study of introgression this paper shows step by step how
precise measures were worked out for analyzing the variation. The methods adapted
for measuring potential variation, demonstrated in figures one to four, could prob-
ably be adapted to various kinds of non-vascular plants.

(7) An experimental investigation of judgments concerning genera and species,
by Edgar Anderson (1957).

Having learned (No. 3 above) that taxonomists agree in their judgments of
species and genera more than they realize, I gathered objective evidence about con-

cepts of species and genera by a simple method. I compared the assignments to
species and genera made in two rival floras dealing with the vegetation of the
eastern United States. From Fernald's last revision of Gray's Manual and Gleason's
New Britton and Brown Illustrated Flora, I made extensive spot checks of their
agreement or disagreement in various families and genera. For areas which they
both cover they agree at least eight or nine times out of ten. "This is of course
a rather limiting case. The flora of eastern North America is relatively stable
and it has been well and intensively studied for over a century. In California
speciation is much more intricate for all kinds of organisms. Microclimates are
much more highly developed. Tertiary and Pleistocene sea-level, rainfall-pattern,
and mountain-height changes have been extreme and have led to complicated
speciation patterns. The flora furthermore has not been intensively studied for so
long a time, yet when one takes two of the standard floras of the state and deals
with comparable areas he finds complete agreement 4 to 6 times out of 10 and
differences only in detail 2 to 3 times out of 10." Acrimonious disputes between
taxonomists have mostly been over the small percentages of instances in which
they did not agree.

These concurrences of opinion convinced me that there is a GENUS PROB-
LEM worth studying as well as a SPECIES PROBLEM. Through the kind offices
of Dr. Robert Cooper, 16 specimens of Uvularia perfoliata and U. grandiflora were
studied in turn by three New Zealand systematists who were unacquainted with the
North American Flora. None of them had specialized on the Liliaceae. All three
placed all 16 in the same genus, proving that there is a Genus Problem worthy of
extensive study in its own right.

"The central core of the instructions accompanying the sheets was as follows:
What I should like is your judgment after examining the specimens as to how
many species, varieties, and genera are involved and (by the numbers) how these
16 plants are to be classified. I am not at all interested in having you identify
them as to family or genus. Quite the opposite; I hope you will not refer to the
literature until after you have dealt with the specimens (if then). This much I
can assure you. They were chosen because their classification presents no problems
and has never been under dispute. If, for instance, they belong to five species in
two genera, then the species are all clear cut and the genera are universally recog-
nized as coherent, distinct genera. If there are any varieties or sub-species present
in the material then they too are well-marked variants but not so distinct as to
merit specific recognition in the opinion of anyone who has dealt with them."

Note that the recipients were given no clue as to the number of genera, specier,
and varieties included in the sample; they were, if anything, encouraged to believe
that more than one genus was involved. They were assured that no problem
genera, nor hybrid swarms, nor apomictic groups of doubtful relationship had been
included in the sample.

Of the three systematists, one was a distinguished monographer of another
family of plants, and he worked independently of the other two. They discussed
the specimens with each other but made their final judgments independently. One
had carried on extensive field work in biosystematics, the third was an algologist.

190 ANNALS OF THE MISSOURI BOTANICAL GARDEN

The latter separated U. grandiftora from U. perfoliata but divided each of them
into two separate species. It was concluded that there is close agreement among
taxonomists on the relation of species to genera even for specialists of different back-
grounds and different kinds of training. Their judgments are apparently intuitive
and inarticulate. The species-genus relationship deserves coordinated research with
biological and psychological techniques.

(8) Folk taxonomies and biological classification, by Brent Berlin, Dennis E.
Breedlove and Peter H. Raven (1966).

This short report summarizes the advancing understandings emerging from a
remarkable project. It is Science's first accurate, comprehensive attempt to analyze
differing concepts of classification in studying the flora of an entire community.

It begins with a refreshingly realistic appraisal of all recent work in this field,
and the authors make an exception only for Harold Conklin's Yale dissertation on
the Hanuoo culture. "Unfortunately most of the data contributing to our under-
standing of folk taxonomies are casually collected, non systematic, incomplete and
anecdotal."

They concentrated on an area of about 160 square kilometers, the municipio
of Tenejapa in Chiapas, Mexico. It ranges from 2700 feet above sea level with
legume-evergreen forest in more moist areas, such as along rivers, to mixed pine-
oak-sweet-gum forest in the more temperate, middle regions, rising to cloud forest
at 9000 feet. "More than 1500 species of vascular plants probably occur in the
municipio." From repeated interviews with native informants they believe their
sample of more than 1100 plant names in Tzeltal, the native tongue, "is nearing
completion."

All their analysis of Tzeltal plant names is based on the "Tzeltal specific" as
a unit. They define it as "any taxon which includes no other taxa" and continue
"For the purposes of this report we have taken a sample of about 20 percent of our
data by including the first 200 Tzeltal-specific names in our alphabetical files. We
have no reason to believe that such a procedure biases the results in any significant

For exact comparisons between the taxonomies of Botanical Scientists and the
Tzeltal specific names, they divided the latter into three categories, 1) under-differ-
entiation, Tzeltal specifics which include two or more botanical species, 2) a one-
to-one correspondence and 3) over-differentiation, when more than one Tzeltal
specific corresponds to a botanical species. Approximately 41% of the 200 were
under-differentiated, 25% were the same and 35% were over-differentiated.

The cultural significance of these Tzeltal specifics to the Tzeltal was graded
high, moderate, and low for (1) plants of little or no utility for them, (2) plants
of moderate cultural significance, as plants used for fuel or firewood but not
cultivated, (3) plants intensively cultivated such as maize, beans, chili peppers
and squash, primarily food or cash crops. From these scorings, a strong positive
correlation between cultural significance and degree of differentiation was revealed.
Forty of the 50 over-differentiated species were judged to be highly significant.

One unexpected result of the analysis was that 40 out of 68 of the plants with

ANDERSON — EXPERIMENTAL STUDIES OF SPECIES CONCEPT 191

a one-to-one correspondence were introduced to Tenejapa after the Spanish con-
quest. They are invariably used today for the same purposes and in many instances
retain their Spanish names.

The authors conclude on a philosophical note, telling us that when we put
together such entities as species with the highest proportion of shared attributes, we
cannot logically insist that these entities share any one particular attribute. This
may tell us little about the structure of nature itself but a great deal about our

Conclusions

During the period that I published the first seven papers of this review, I was
increasingly impressed with the importance of joint taxonomic-psychological in-
vestigations of the two aspects of the species problem discussed and diagrammed in
(1). For none of that time, however, did I have a colleague well-grounded in the
fields of psychology pertinent to these studies. The 8th review demonstrates the
ways in which such problems could have opened up fruitful new fields of investiga-
tion had such collaboration been possible.

Within the last decade I have become aware that experimental investigations
of taxonomists' conceptions of taxa above the species might produce more effective
collaboration of taxonomists with physiologists and biochemists.

When a taxonomic monograph of a group of plants has achieved its goal and
put like ones close together that tend to share many common features, it can be
widely useful. Taxonomists of the higher plants could be as helpful to biochemistry
as were those medical school pharmacologists whose discoveries led to such useful
concepts as "psychedelic drugs", in grouping whole sets of compounds together.
A very little has already been done. The association of dangerous cyanogens with
the Rosaceae had long been recognized. See Kingsbury (F) for discussion of this
and many other examples.

Various taxonomists are already using biochemical information in classifying
plants. The relationship between biochemists and plant taxonomists could become
genuine collaboration, profitable to both sciences. Sorbitol, for instance, is common
to closely related Asiatic and American species of Larix (larch) but is not produced
in less closely-related species (reviewer's unpublished information). Such facts are
useful not only to monographers of the genus Larix but also to chemical manu-
facturers looking for cheap sources of sorbitol.

Before such a two-way association could become facile, there would have to be
a wider understanding among taxonomists as to which sub-genera, genera, sub
families and families of higher plants are based on a wide variety of characteristics,
each of which is shared by most of the group, in other words is a "natural" group
and not an "artificial" one. Various genera, families and higher categories in plant
taxonomy are widely recognized as "natural" or "artificial". Taxonomists frequently
discuss these matters informally among themselves, but I know of no exposition of
the problem except the elementary one cited in (5).

It is becoming widely understood, for instance, that the rose family is a natural
group and less widely that one of its sub-divisions, the Pomoideae, (the pome

192 ANNALS OF THE MISSOURI BOTANICAL GARDEN

fruits — apples, pears, quinces, hawthorns, etc., fruits with a core) are an introgres-
sive, polyploid, apomictic complex whose genera, species and sub-species are arti-
ficial. They form such a complex, interwoven network of relationships that there
have been wide disagreements between experts in cataloguing them.

No one or two taxonomists could yet compile a working list of outstanding
natural and artificial taxa, complete enough to be generally useful. On the other
hand, a modern taxonomist, familiar with computer techniques, could organize a
survey of present day concepts that could be mechanically sorted. It would, at the
very least, be a beginning that would call attention to the problem. It might grow
into a catalogue that would be widely helpful.

The closing three paragraphs of the paper by Berlin et al. (8) with a discussion
of "general" versus "special" classification, indicate the advances of understanding
and of utility to be expected from expert conceptual classifications.

Bibliography
Numbers refer to Papers reviewed; letters to backgrou
(1) Anderson, Edgar & Thomas W. Whitaker. 1934.
Arnold Arb. 15:28-42.

1936. An experimental study of hybridization
Ann. Missouri Bot. Gard. 23: 159-168.

i & H. C. Cutler. 1942. Races of Zea Mays )
. Missouri Bot. Gard. 29: 69-89.
Edgar. 1940. The concept of the genus II. a survey of modern i
Bull. Torrey Bot. Club 67: 363-369.

(C) Anderson, Edgar & Dorothy Schregardus. 1944. A method for recording and
analyzing variation in internode pattern. Ann. Missouri Bot. Gard. 31 : 239-247.

(4) Anderson, Edgar. 1951. Concordant vs. discordant variation in relation to mtrogressive
hybridization. Evolution 5: 133-141.

(D) Dietz, Robert A. 1952. Variation in the perfoliate Uvularias. Ann. Missouri Bot.
Gard. 39:219-254.

(5) Anderson, Edgar. 1954. Efficient and inefficient methods of measuring specific differ-
ences, in Kempthorne, Bancroft, Gowen and Lush. Iowa State College Press, 93-106.

(6) Anderson, Edgar & E. D. Rudolph. 1956. An analysis of variation in a variable
population of Claudonia. Evolution 10: 147-156.

(E) Roberts, L. M. et al. 1957. Races of maize in Colombia. Natl. Acad. Sci-Natl. Res.
Council Publ. No. 510.

(7) Anderson, Edgar. 1957. An experimental investigation of judgments concerning genera
and species. Evolution 11 : 260-262.

(8) Berlin, Brent, Dennis E. Breedlove & Peter H. Raven. 1966. Folk taxonomies and
biological classification. Science 154: 273-275.

(F) Kingsbury, John M. 1965. Deadly harvest; a guide to common poisonous plants.
Holt, Rinehart & Winston, N. Y. 128 pp.

INTERSPECIFIC RELATIONSHIPS IN THE
POLYPODIUM PECTINATUM-PLUMULA COMPLEX 1

by A. Murray Evans
Department of Botany, The University of Tennessee, Knoxville

Studies of the life cycle, anatomy, morphology, cytology, ecology, distribution pat-
terns and an analysis of the probable evolutionary relationships of taxa in the Polypodium
pectinatum-plumala complex, compose the basis of a systematic study of this complex.
Field studies in Florida, Jamaica, and Costa Rica, plus use of greenhouse grown plants
South America, have aided in the understanding of the biological interrelation-
ships of the tax;! 'tiding types, from 20 major American and European
herbaria have been utilized in this study, which has led to the recognition of 35 taxa,
including five new species, three new varieties and six new combinations. Keys, descriptions,
typification, synonomy and specimen citations are included for each taxon.

Introduction

In attempting to understand the interrelationships of the very closely related
species of the Polypodium pectinatum-plumula complex, studies of the ecology,
variation in certain populations, and the natural history of the several species, as
well as traditional morphological and anatomical studies, were of considerable
help. Because the group has a wide tropical distribution, extensive field studies
were not possible, but field studies were made in Florida, Jamaica and Costa Rica,
and I obtained living rhizomes of several species for culture from collectors in
South America. Although it was necessary to rely heavily on data obtained from
the species in Florida and in greenhouse culture, several aspects of the life cycles
of these species were examined, and this information was useful in interpreting
the remaining species.

Historically, the major preoccupation of taxonomists has been in reducing the
enormous and polyphyletic genus Polypodium to its present restricted sense. No
systematic coverage of this whole group as a unit has ever been attempted, and
the taxonomy has consisted mainly of the addition of new taxa one by one.
Lindman (1903) treated eight of them in an overall study of tropical American

Contribution from the Botanical Laboratory, The University of '

e Graduate School of r .
ints for the degree of Dot
! G-10846 (to Warren H. '
GB-3966 travel funds from the Rackham Graduate School and the Botanical Gardens, both
of The University of Michigan, and publication funds from the Faculty Research Fund of
The University of Tennessee. Acknowledgement is gratefully made to Dr. Warren H.
Wagner, Jr., for supervision of the research, to Mr. Conrad V. Morton for extensive nomen-
clatural aid, and to Mr. G. R. Proctor of the Institute of Jamaica and the Director and
staff of the Organization for Tropical Studies for aid and support of field work. Specimens
were borrowed from the following herbaria (abbreviations follow those of Lanjouw &
Stafleu, Index Herbariorum, 3rd. ed., 1956) : B, BR, C, FI, FLAS, G, GH, HB, K, L, LE, M,
MICH, MO, NY, P, R, S, S-PA, US.
Ann. Missouri Bot. Gard. 55(3): 193-293, 1969.

194 ANNALS OF THE MISSOURI BOTANICAL GARDEN

species. De la Sota (1960), in his research on the Polypodiaceae of Argentina,
grouped Polypodium into several complexes and treated the eight species of the
present group from that country.

Morphology and Ecology

Microscopic morphological observations were facilitated by several techniques.
The living material was first killed in FAA (formalin, acetic acid, alcohol) or dried
material was relaxed in 5% NaOH for 2-3 days before sectioning,
stipes, and blade segments were sectioned at 2S(j. on a freezing
mounted unstained in diaphane; or they were imbedded in paraffin, sectioned on
a rotary microtome, stained in safranin and fast green, and mounted in diaphane.
For studies of venation and epidermal patterns, leaf segments were cleared in 5%
NaOH, bleached in 10% Clorox, stained by the tannic acid-ferric chloride method
(Foster, 1934) and mounted in diaphane or piccolite. Spore preparations were
made either by the acetolysis technique (Erdtman, 1943) and mounted in silicone
oil (20,000 viscosity grade), or teased from the frond in 95% EtOH and mounted
directly in diaphane.

Gametophytes were grown on a variety of growth media. For young stages,
spores were sown on either distilled water or liquid Biejerinck's Solution (Brown,
1920) . For later stages and mature gametophytes, spores were sown on agar fortified
with Knop's Solution (Brown, loc. cit.) inpetri dishes, or sown in 3" porous clay
pots with either pulverized sphagnum or hardwood humus. The spores were
sterilized by treating them with varying concentrations of Clorox, 10% for 10
minutes being optimal. However, good results were usually obtained in nonsterile
conditions. The pots containing soil media were autoclaved, but the petri dishes
with agar were not, because it was found that fungi were inhibited when the agar
was not autoclaved. Spores, treated either by sterilization or from washed fronds
which had been dried in sterilized envelopes, were sown in a sterile transfer

Gametophytes were observed microscopically either in water mounts, mounted
in Hoyer's Solution without being killed, or were killed in 70% EtOH and cleared
and stained as the leaf segments above. The small gametophytes were handled
in quantity by transferring them in short upright sections of wide glass tubing with
the bottom covered with cheese cloth.

Voucher specimens of all experimental collections are deposited in the Univer-
sity of Michigan Herbarium.

Rhizome
The stem (Fig. 17-19) is a long-creeping to short-creeping or sub-erect rhizome,
with the fronds short-spaced (never more than 1.5 cm apart) or approximate to
fasciculate and articulate on short pseudopodia 1 to 2 mm long. The plants may
be epiphytic, terrestrial, or epipetric, but the rhizomes are always exposed. They
vary in diameter from 1 to 3 mm in small species such as P. truncorum to 10 to
13 mm in the larger species such as P. ptilodon or P. recurvation. The internal
anatomy is quite uniform. The vascular system is dictyostelic with 3 (in P. trun-

EVANS— POLYPODIUM 195

coram) to 17 small exarch bundles in a single ring (Fig. 1, A-C). The endodermis
is thin-walled, but the inner wall of the ground parenchyma cells adjacent to the
endodermis is always strongly thickened. Polypodium singer i and P. truncorum,
two small epiphytic ferns, show a good deal of dark staining, thick-walled col-
lenchyma tissue in all but the outer portion of the rhizome (Fig. 1, C). No true
sclerenchyma tissue is present.

The rhizomes are clothed with paleae throughout, which fall basically into
two groups of scales— (a) the most common type, associated with the P. pectinatum
allies, and (b) those found in the groups of P. cupreolepis and P. truncorum.

The former group (a) has narrow- to linear-triangular, dark red-brown lustrous
paleae, basifixed and with an acuminate or filiform apex. The paleae are either
broadly expanded at the basal attachment or have a narrow base at the apex of the
rhizome. They are dark colored along their margins, usually with a paler area at
the base. The cells are narrow, except at the base of the scale, where they usually
become shorter and nearly isodiametric. They are non-clathrate. This is in con-
trast to several groups in Polypodium (such as the P. vulgare or the P. squamatum
groups) in which the paleae are usually basally peltate, at least sub-clathrate, and
are usually darker in the center rather than along the margin.

These paleae all have a dorsal cluster of unicellular hairs (and are therefore
termed "comose") toward the base of the paleae, which are usually visible under
low magnification (Fig. 1, G, J-L). In certain species, such as P. pectinatiforme,
P. atrum, or P. dispersum, the hairs are uniformly present, although they may not
be conspicuous. In P. plumula, on the other hand, the comose hairs are sometimes
absent. In P. bolivianum the hairs of the paleae are long and dense, and they
mostly obscure the scales themselves except at the rhizome apex (Fig. 1, K, L), and
in P. consimile var. pastazense they totally obscure the paleae.

In the second group (b) the paleae are cordate or ovate, basifixed at a single
point, not so polished or lustrous as in the previous group, and vary in color from
golden to dark red-brown. They are non-comose and either concolorous or some-
times paler toward the base (Fig. 1, I).

The roots are approximately 0.5 mm in diameter with an outer thin-walled
cortex of 3-9 layers of parenchyma cells, and a sclerified darkly colored inner cortex
usually 4-6 layers deep occupying one-fourth to one-half of the diameter of the
root. The endodermis is a thin-walled and inconspicuous layer lining the inner
cortex. The bundle is diarch with few (4-8) large tracheids and clusters of small
tracheids at the two protoxylem points.

Root Proliferations
Although root proliferation leading to sporophytic reproduction is known in
pteridophytes, it is infrequent. Root proliferations are known in Platycerium,
Amphoradenium (syn. Adenophorus) , and Asplenium, and in the present study
they have been found to occur in Polypodium dispersum, P. ferrugineum, P.filicula,
P. pectinatum, P. plumula, P. siccum, and P. singeri (Fig. 7).

196 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Although root proliferations have occasionally been thought to be new plants
arising from slender elongated stems or rhizomes (similar to the stolons of Nephro-
lepis), this is not the case in the present complex. As seen in cross-section, a single
vascular trace of the root type crosses the cortex of the parent root. The new
rhizome is usually first noticed as a small scaly bump approximately 1 mm in
diameter on the side of the root. New roots and leaves develop in typical fashion
from this small rhizome.

Juvenile Leaves

Within the Polypodium pectinatum-plumula complex, sporophyte leaves of a
"juvenile type" can develop from any of three sources— from root proliferations in
various species (as discussed above), from apogamous outgrowths in P. dispersum
and from sexual embryos.

Leaves produced from root proliferations and from gametophytes have the
same morphological stages. Figure 2, D-F, shows the developmental leaf series of
P. dispersum, P. plumula, and P. ptilodon var. caespitosum. The first leaves are
invariably simple and elliptic with a single midvein of spirally thickened tracheids.
The first leaves of P. ptilodon var. caespitosum are larger and more broadly elliptic
than those of either P. dispersum or P. plumula. The three hair types, discussed
below, appear on the first leaves of all the species listed (Fig. 15, C, D, F, G),
except that ctenoid hairs are lacking on P. plumula. After the second or third
leaf, the veins become forked and the blade becomes lobed. The leaves which fol-
low are alternately lobed. Subsequent to these are leaves which are sub-oppositely
lobed as in the mature leaf. The venation then develops the mature condition of
being 2 or 3 times forked (Fig. 2). No young stages of any of the species with
anastomosing venation were available for study, but presumably they follow the
same pattern modified only by vein fusions.

Mature Leaves

The fronds may be upright, curved or pendent. Most of the species have up-
right and rather stiff fronds, but certain of the epiphytic ones with upright rhizomes
attached to vertical substrates tend to have more flexuous or pendent fronds, such
as P. plumula (Fig. 3, F), P. curvans (Fig. 3, C), P. choquetangense, P. siccum,
or P. truncorum.

The frond outline is narrow-elliptic, -ovate, or -deltoid (Fig. 3). In those
species that have narrowly elliptic fronds (e.g. P. plumula, P. alfredii, or P. cupreo-
lepis) and those with narrow-ovate fronds (e.g., P. ptilodon, P. dispersum, P. atrum,
or P. camptophyllarium) the stipe may be long or short. Several of the species have
the lower segments contracted quite abruptly, as in P. plumula (Fig. 3, F), and a
long stipe; in others, such as P. consimile or P. filicula, the stipe is reduced or
almost absent, and the basal segments are very gradually reduced to lobes. The
deltoid or subdeltoid fronds, such as those of P. recurvatum (Fig. 3, G) and P.
bolivianum, have an abrupt demarcation between stipe and rachis at the base of
the blade.

The fronds are articulate, abscissing from a short pseudopodium. The tip of

EVANS— POLYPODIUM 197

the frond and the segments curl tightly adaxially in response to drought, either
from atmospheric dryness or water loss after collection. The whole group possesses
the ability to respond in this manner, but it is most strongly expressed in such
species as P. plumula, P. dispersum, P. filieula, and P. cupreolepis. Both this
curling ability and the articulation of the fronds reduce the exposed leaf surface,
and appear to be adaptations to the epiphytic or epipetric mode of life and to
periods or seasons of little atmospheric humidity.

Stipe and Rachis

The stipe and rachis are distinctive. In the mature leaf the main axes are
either black or of various shades of red-brown; they always have a highly polished
appearance. In juvenile fronds the main axis may be lighter or even stramineous,
but it turns dark and lustrous as the blade matures. The only exception is that
certain of the northern plants of P. hygrometricum have a light axis even at
maturity, although the majority of specimens of this species have dark axes. The
color may also vary between living and dried plants. The axes of P. dispersum
and P. plumula are black in both states. In P. recurvatum, P. trunoorum, and P.
ptilodon var. ptilodon they are red-brown in both the living and dried state.
Polypodium ptilodon var. caespitosum has a consistently red-brown axis and costa
when dry, but the axis is quite blackish in living material. A wide sclerified layer
in the outer cortex gives the axis its hard and lustrous external appearance. This
layer also gives the rachis the characteristic springiness or elasticity readily recogniz-
able even on herbarium sheets, particularly in the strongly pendent epiphytes, such
as P. plumula. In P. curvans and P. choquetangense the main axis is usually
sinuous or contorted, and the apex of mature fronds often remains circinate.

The stipe and rachis are terete, and the stipe has either narrow wings or nar-
row lateral green stripes along its entire length.

Trichomes

Three types of hairs are present on the fronds of these ferns, as illustrated in
Fig. 4. (1) The first is a small, simple, clavate hair composed usually of only two
or three cells, the terminal one being somewhat larger and blunt-tipped (Fig. 4,
I 3 ). These hairs are usually no longer than 150^. (2) The second type is ap-
parently derived from the first. In its most characteristic form, this hair consists
of an arched axis of approximately four cells, with lateral branches usually of only
one, sometimes two, cells. There are usually three or four of these lateral branches
all arising from one side producing a "ctenoid" hair. However, modifications of
this type of hair are frequent in which the number of branches is reduced, some-
times to one, thus suggesting intergradation with the simple clavate hair (Fig. 4,
I). (3) The third hair type is a long, simple, acicular, multicellular hair (Fig. 4,
J 3 , K 4 ). These hairs are usually pale on the young frond and turn a golden color
at maturity. The acicular hairs vary considerably in length, from 0.2 to 2.0 mm.
They are of diagnostic importance in several species, as indicated below.

The three hair types are found on the stipe, rachis, costa, and lamina. The
ctenoid hair is apparently present on all juvenile fronds, but is not present on the

198 ANNALS OF THE MISSOURI BOTANICAL GARDEN

mature fronds of P. plumula, P. cupreolepis, P. truncorum and their immediate
allies. The small, clavate hairs are universally present, primarily on the lamina.
They are usually so short as to be overlooked, except in P. sursumcurrens, in which
they are long, blackish, and visible to the naked eye. The acicular hairs have not
only considerable variation in size, but vary also in density and distribution. They
are absent in a few species, such as P. bolivUmum, and are only scattered on the
lamina in most species, but are quite abundant and dense in P. pectinatum, and
P. camptophyllarium. In all four varieties of P. ptilodon the acicular hairs have
a curious pattern of distribution such that the lamina is essentially glabrous except
for an oval area of dense pilosity around the sorus, often visible to the naked eye
or with a hand lens (Fig. 1, E). These are not to be confused with the receptacular

The most frequent type of rachis scales or paleae are those of the P. pectinatum
allies (Fig. 5, H, K) which are long, filiform, inconspicuous, and few in number.
They are uniformly dark red-brown and of long, thin cells, except that they have
more lightly colored and small isodiametric cells at the base. They occur on both
the upper and lower surfaces of the main axis, although usually only on the stipe
and lower portion of the rachis. At the base of the stipe they tend to have broader
bases and approach the shape of the rhizome paleae, which, in the P. pectinatum
assemblage, are invariably linear-triangular. The margins are entire except in P.
bolivianum, where they have simple or forked fimbriae on the margin, and in P.
eurybasis, which occasionally shows slightly fimbriate margins. In both of the
latter species this is an amplified condition as regards stipe paleae, and approaches
the more fimbriate condition of the rhizome paleae of those species. Several other
species, such as P. pectinatum or P. camptophyllarium, have inconspicuously fim-
briate rhizome paleae, but have entire rachis paleae. This is thus apparently a
relative change: those species with quite heavily fimbriate rhizome paleae show
some indication of this on the rachis paleae, but those species with few or no
fimbriae on the rhizome paleae lack any at all on the paleae of the rachis.

The second group of species (P. atrum, P. dispersum, P. bermudianum, P.
plumula, P. filicula, P. ferrugineum, P. cupreolepis, P. alfredii and P. sursumcur-
rens) have rachis paleae which are non-filiform (Fig. 5). They may be narrow-
elliptic (P. alfredii, P. sursumcurrens and P. ferrugineum) (Fig. 5, F), narrow-
triangular with irregular hastate bases (P. dispersum, P. atrum, P. bermudianum,
and, in part, P. plumula) (Fig. 5, A 4 _ 7 , B, C, D, J) or they may be essentially
cordate, as in P. plumula, P. filicula, and P. cupreolepis (Fig. 5, A t _ 3 , G, I, L). In
all but the narrow-elliptic paleae, the margins may be inconspicuously and ir-
regularly fimbriate (Fig. 5). Polypodium filicula is the only species with cordate
or triangular paleae in which most paleae have an entire margin. Except for
overall shape, all of the paleae of this group of species are similar to those of
the rhizome in coloration and cell structure. Those species with hard, lustrous,
red-brown rhizome paleae have similar rachis paleae. Species such as P. cupreo-
lepis (Fig. 1, I and Fig. 5, G) and P. ferrugineum, with pale and dull rhizone
paleae, have similar rachis paleae.

EVANS — POLYPODIUM 199

The narrow-elliptic paleae on the rachis are sparse like the filiform paleae
in the previous group. However, those species with hastate-triangular or cordate
paleae possess them on the rachis in greater quantity, so that they are readily
visible to the unaided eye.

In all taxa except P. plumula and P. filicula the paleae are flat at the base.
In the two exceptions the scales appear inflated or "bull ate" at the base. This
character appears occasionally also in the paleae of P. curpreolepis.

The species with conspicuous and numerous rachis paleae (P. atrum, P. ber-
mudianum, P. dispersum, P. plumula, P. filicula, and P. cupreolepis) also have
paleae on the costae. In all cases, these are reduced paleae similar to those of the
rachis. In some, as shown in P. plumula (Fig. 4, H), the homologies between paleae
and hairs are apparent. Although there are basically only three hair types, as
described above, in those taxa with costal paleae an additional large, many celled,
clavate, simple hair is also present. It is evident that this is the ultimate reduc-
tion of the paleae.

Segments

Figure 6 illustrates outlines and venation patterns of the segments of rep-
resentative species. Figure 3 shows the orientation of the segments in selected
species. All the species have simple and entire segments except P. eurybasis, P.
curvans and P. paradiseae, which have crenate or crenulate segments, and P.
choquetangense, which has pinnatifid segments. All species have inconspicuously
ciliate segment margins.

Whereas the segments are typically perpendicular to the rachis and straight
or slightly falcate, several of the species (P. absidatum, P. curvans, P. truncorum,
and P. choquetangense) have noticeably ascending segments (Fig. 3, C, D). In
most species the segments have a symmetrical base in which both the lower and
upper edges are dilated. However, in thosa species with ascending segments the
upper edge meets the rachis abruptly at a nearly right angle. Also, several species,
such as P. atrum, P. paradiseae, P. pectinatiforme, and P. filicula, have segments
of the lower portion of the frond deflexed, and the lower edge of the segment
correspondingly meets the rachis at a right angle or less (Fig. 3, B & Fig. 18). In
P. alfredii, several plants from the northern part of its range have basal segments
deflexed so that the lowermost segments are nearly or quite parallel to the stipe.
There are several species, however, with all segments essentially at right angles
to the rachis, but in which the lower edge of the segment meets the rachis at a
right angle, or in which it may even be slightly incised to form a small sinus be-
tween rachis and basal margin (Fig. 3, A).

Figure 4 shows variations in the shape of the abaxial epidermal cells and
the arrangement of the stomates. The cells over the lateral veins are character-
istically narrower and more attenuated than those away from the veins. The
stomates occur only on the abaxial surface. The guard cells are slightly elevated
above the epidermal cells and are without modified subsidiary cells. Guard cell
sizes, as they relate to the ploidal level, are discussed below.

The internal tissue of the segments is compact and quite uniform from one

ZVV ANNALS OF THE MISSOURI BOTANICAL GARDEN

species to another. They average approximately four layers of cells in the mesophyll
(two to three in P. truncorum and five or six in P. pectinatiforme), two layers in
a somewhat irregular palisade above and two (three, four) slightly less compact
and more irregular layers in the spongy mesophyll (Fig. 1, D).

Figure 6 shows the representative range of venation patterns in the segments.
Although the group typically has free-forked venation, three species (P. filicula,
P. siccum, and P. truncorum) have simple veins. Entirely simple veins are not
common in the Polypodiaceae, although the condition is frequent in the Gram-
mitidaceae, a reason why P. trucorum has been considered as a ctenopteroid fern
by Copeland (1956). However, on the basis of the spores, sporangia, trichomes,
and the characters of the rachis, all three of these simple-veined species are properly
members of the Polypodium pectinatum-plumula complex (de la Sota, 1963).
Anastomosing venation, while not the rule in the present complex, occurs all
through the Polypodiaceae. In the group under consideration here, anastomosing
occurs in varying degrees from casual, irregular junctures in each segment, as in
P. pectinatum, to very regular and complete patterns in P. camptophyllarium
var. macedoi. Polypodium camptophyllarium var. camptophyllarium has consider-
able variation, although it usually has at least a sporadic distribution of areoles.
In some plants in southern South America the anastomosing in the latter variety
becomes extensive and may even exceed that which is expected in var. macedoi
(Fig. 6, Q. S). When anastomoses occur in this complex, they are always with
a free included veinlet. This is opposed to the areolate condition in the Gram-
mitidaceae, in which there are no included veinlets.

The costa, in all cases, is sclerified, sometimes only in the abaxial portion
of the bundle sheath (e.g. P. dispersum and P. pectinatiforme), and sometimes
throughout the bundle sheath, but then usually more so on the abaxial side, as
in P. truncorum and P. ptilodon (Fig. 1, D). In several of the species with large
fronds, the bases of the secondary veins are also sclerified. Except in P. ferrugineum,
in which the costae are black and the rachis brown, the costae and rachis are
the same color.

Sori
The sorus is terminal on the first acropetal veinlet in all species (Fig. 6).
In those with anastomosing venation this veinlet becomes the free included vein-
let. Whereas the sorus is typically medially or extra-medially placed on the seg-
ment, it may vary all the way from infra-medial (in P. consimile) to submarginal
(in P. sursumcurrens, P. alfredii, and P paradiseae).

Wilson (1959 a, b) has described in detail the sporangia of P. plumula and
P. ptilodon var. caespitosum (as "P. pectinatum") . I have found these structures
to be similar and with little deviation throughout the group. Although the annulus
usually has 13 or 14 bow cells, P. camptophyllarium var. lachniferum has only
12, and P. sursumcurrens and P. curvans have 16.

Wagner (1964) discusses and summarizes the known types of paraphyses
in the ferns. Figure 4 illustrates the types found in this complex. Sori of all species
produce receptacular paraphyses, and in all but two these are simple or occasionally

EVANS— POLYPODIUM ^»1

forked, multicellular, clavate hairs. In all but one species they are similar to
the clavate laminar hairs, although they are often longer, and vary from 150-
250^ long. In P. camptophy liar turn var. macedoi the receptacular paraphyses are
a curious mixture of the clavate and the acicular hair types (Fig. 4, M), being
forked hairs with one clavate and one acicular branch. However, the same type
of hair is also characteristic of the lamina. In P. sursumcurrens the paraphyses
are unusual in differing considerably from the laminar trichomes (Fig. 4, P).
They range up to 4(% long. Their outline is contorted, and a multicellular,
expanded portion is developed near the apex. In most species the receptacular
paraphyses are few and hard to observe except by dissection and microscopic ex-
amination, but in P. sursumcurrens they are numerous and visible even with
low magnification. Unlike the elaborate "sporangiasters" of P. virginianum, ap-
parently derived from modified sporangia (Martens & Pirard, 1943), the para-
physes of P. sursumcurrens are evidently related to the simple, clavate, laminar
hairs, as in the remainder of the complex.

Capsular paraphyses (Fig. 4) also occur in approximately half of the species
of the complex. They are always short, simple, 1- to 3-celled hairs arising near
the top of the capsule near the annulus. They vary in length from 45^ in P.
pectinatum to lOfye in P. hygrometricum. They may be of quite regular number
and occurrence— one per capsule in P. pectinatum; two in P. hygrometricum and
P. ferrugineum; or of quite variable number, as in P. dispersum or P. plumula, in
which they range from one to five per capsule. In several species they are present
on young capsules but may be shed as the capsule matures.

With two exceptions, spores in this complex are essentially alike. One ex-
ception, Polypodium dispersum, has only 32 mitotically produced, globose spores
per sporangium. The other, Polypodium camptophyllarium var. camptophyllarium,
has, for the most part, normal, reniform, tuberculate spores, but a few collec-
tions from Jamaica have quite irregular spores, which suggests that apogamous
reproduction may be present in scattered populations of this species (see the tax-
onomic description of this variety).

The remaining taxa all have 64 spores per sporangium and are therefore
presumed to have a normal sexual life cycle. The spores are uniformly reniform
and monolete (Fig. 12, H), although they vary in shape from narrowly reniform
and curved to globose-reniform. They are all without perispore and the surface
is tuberculate, although the sculpturing varies from almost smooth to strongly
tuberculate. The spore sizes, as they relate to the ploidal level, are discussed in
the section on cytology and evolution.

Game

The gametophytes of Polypodium dispersum are so unusual that a separate
section has been devoted to them in the section on sporogenesis. The gametophytes
of the Polypodiaceae and Grammitidaceae have been treated by Stokey & Atkin-
son (1958), and the gametophytes of P. plumula and P. ptilodon var. caespitosum

ZVZ ANNALS OF THE MISSOURI BOTANICAL GARDEN

(as "P. pectinatum") were reported by Stokey (1959). All my observations agree
with those previously described.

The sex organs are typical of the Polypodiaceae, as described by Stokey.
After fertilization the sporophyte plant is first seen as a tiny elliptic leaf (Fig. 2)
and a root that appears either at the same time or slightly before the leaf. The
leaves then develop as described above in the section on juvenile leaves.

Ecology and Geography
These ferns extend from Bermuda, peninsular Florida, northern Mexico and
the Galapagos Islands south to northern Argentina and southern Brazil (Fig. 8, B).
I consider them to occupy five distributional zones (Fig. 8, D): (1) Central Amer-
ica, (2) western South America, (3) southern Brazil, (4) northern and north-
western South America, and (5) the West Indies, considered a sub-zone of the
last. As the distribution maps show (Fig. 8-11), several species occur in more
than one zone, although a particularly large number of species are confined to
the Central American and southern Brazilian zones. I have included the West
Indies as a sub-zone of the northern South American zone because, out of the
nine species present in the West Indies, six are known also in northern South
America and do not occur in Central America. The remaining three species (P.
dispersum, P. plumula, and P. pectinatum) are wide-ranging and cross several
zones (Fig. 9, 10).

Although as a whole the genus Polypodium is primarily epiphytic, some
species are terrestrial or epipetric. The same is true of the P. pectinatum-plumula
complex. No species is strictly terrestrial, although P. ptilodon (including all
its varieties) and P. consimile var. consimile are primarily terrestrial or occur on
the bases of trees (Fig. 7, E). Polypodium bermudianum, P. dispersum (Fig. 7,
A, G) and P. ferrugineum are primarily epipetric, although the two latter are
recorded as occurring also epiphytically or terrestrially. Polypodium choquetan-
gense, P. curvans, and P. cupreolepis are either epipetric or epiphytic. Polypodium
sursumcurrens, P. siccum, P. singeri, and P. truncorum are evidently strictly epiphy-
tic. Polypodium singeri is curious in that it apparently grows with its roots reach-
ing the ground, so that it perhaps is only hemi-epiphytic. Polypodium truncorum
has a particularly narrowly defined substrate, and is found only among the old
stipe bases of tree ferns, especially Alsophila and Hemitelia. The remaining species
are primarily epiphytic, but also occur occasionally as epipetric or terrestrial plants.
On the whole, this is primarily a group of tropical and sub-tropical montane
plants. Elevation data indicates that they grow from 1500-3500 m in altitude,
but a few species are found at low altitudes. Polypodium dispersum, P. plumula,
and P. ptilodon var. caespitosum occur in peninsular Florida at nowhere over 100
m in elevation, although they are known elsewhere at much higher elevations.
Polypodium hygrometricum and P. recurvatum are known only from sea level to
1800 m and P. bermudianum is confined to the low lying Bermuda Islands.

Certain of the epipetric species are recorded from exposed rocky areas or cliffs.
No species grow in the lowland rain forests of the Amazon basin. Judging from
the generally montane distribution, the response to drying (desiccation and curl-

EVANS— POLYPODIUM 203

ing of the frond without the death of the plant) and the articulate stipes, this
group is not adapted to areas of constant high humidity, such as in the Amazon

Ecology of Root Proliferations

Several species in this complex reproduce partially by root proliferations;
this appears to be related to their distribution and ecological requirements. Pro-
liferations in P. ferrugineum, P. filicula, P. siccum, and P. singed were found
only on herbarium specimens, so it is difficult to note more than their mere pres-
ence. However, P. singeri normally grows as an epiphyte on small woody twigs.
Several herbarium sheets show that the individual rhizomes are often short (ca
2 to 3 cm long) and grow in a series along the twigs (Fig. 7, D). The rhizomes
and roots are exposed and the rhizomes arise as root proliferations.

Polypodium plumula and P. pectinatwn grow either on the trunks or hori-
zontal branches of trees, and are true epiphytes (Fig. 7, B). Root proliferations
occur infrequently as small plantlets formed from the roots of larger plants. I
have also observed small plantlets arising from the superficial roots of a green-
house culture of the former species.

Although P. dispersum is capable of reproducing by spores (as discussed be-
low in the treatment of sporogenesis), its primary mode of local dispersal is evi-
dently by root proliferations. In Florida, for example, this species is usually seen
as dense mats of juvenile plants completely covering limestone rocks (Fig. 7, G).
Often none of these plants bear sporangia. I saw only three soriferous colonies
of this species in the course of field trips in Florida, although I studied numerous
populations.

The root proliferations in nature usually occur only a few centimeters apart
when the plants are growing on rock surfaces. However, at "Blechnum Ravine"
in Annutalaga Hammock, Florida, P. dispersum was growing in sand and the
plants were widely spaced. In one case proliferations were spaced approximately
0.5 m apart on a very long root.

The rhizomes of these proliferations are always short and superficial on the
substrate with no intermediate structure between parent root and rhizome, so
that proliferations occur only on those roots that are close to the surface. As these
seven species in which proliferations are known are all either epipetric or epiphytic,
they normally grow where the substrate is thin and many of the roots are exposed
or near the surface. These proliferations may be produced in response to reduced
moisture or greater aeration around exposed roots. Increased light also might be
a factor. However, the occurrence of proliferations in the dark at the bottom
of the pot in greenhouse culture suggests that this is not so, unless a minimal
exposure to light is sufficient.

Sporogenesis and the Life Cycle of Polypodium dispersum

I have carried out life cycle studies primarily on P. dispersum and P. ptilodon

var. caespitosum. Experimental collections of P. dispersum have included a plant

from Jamaica (Proctor 22896) and collections from Florida (Citrus Co, Wagner

62061, Evans 2008). Collections of P. ptilodon var. caespitosum have been from

204 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Florida (Highlands Co, Evans 1157; Citrus Co, Evans 2005; Sumter Co, Evans
2004).

Sporogenesis studies were carried out using a modified acetocarmine squash
technique. Segments with sporangia of a suitable age for meiotic figures were
killed and fixed in Newcomer's Solution (Newcomer, 1953) and stored in a deep
freeze until used. To prepare microscope slides, sporangia from a single sorus
were placed in a drop of acetocarmine on a slide. No iron mordant was added,
thereby avoiding overstating of the cells. The sporangia were spread apart to
avoid clumping, and a cover slip was added. The slide was heated slightly, but
tapped only lightly with a dissecting needle in order not to spread the individual
cells too far from the mother sporangium. Firm pressure was then applied with
the thumb. The combination of few and wide-spaced sporangia, light tapping,
and firm pressure with the thumb, resulted in preparations in which the cells were
well squashed, but the individual cells of a particular sporangium were still in
close proximity to each other. In this way numerous sporangia could be easily
and quickly prepared and observed. The slides were made permanent by inverting
them in 2:8 acetic acid: alcohol solution until the cover slips dropped off. After
rinsing with the acetic acid-alcohol solution, the cover slips were replaced with
a drop of diaphane.

In the P. pectinatum-plumula complex, the sexual life cycle (as determined
by the number of spores per sporangium) is present throughout except in P. dis-
persum. In this common and widespread species, a pattern of apogamous reproduc-
tion thus far unique among pteridophytes has been found.

Sexual Life Cycle
The sexual life cycle of the higher ferns is well known. In comparing this
with apogamous reproduction, it is only necessary to summarize the process of
sporogenesis in a sexual species, as exemplified by P. ptilodon var. caespitosum
(Fig. 12). In the developing sporangium, the archesporial cell divides mitotically
four times to form 16 2 X spore mother cells (Fig. 12, B). The spore mother cells
are readily recognized by their size and form, and also by the large, darkly stain-
ing nuclei. They undergo one meiotic division (Fig. 12, C, D) to form a tetrad
of young spores (Fig. 12, E-G), resulting in 64 monolete spores per sporangium
(Fig. 12, H). Upon spore germination and maturation of the bisexual gametophyte,
fertilization may occur; the sporophyte plant develops from the fertilized egg, thus
completing the cycle.

Obligate Apogamous (Meiotic Apogamous) Life Cycle
Although not so widely known as the sexual life cycle, this process is well

documented, as in several papers by Dopp (c.f. 1939) and later by Manton (1950)

and others.

Using herbarium specimens, apogamy can usually be detected by making

preparations of mature but unopened sporangia. Some of the sporangia, if the

plant is apogamous, will have 64 aborted spores, whereas others will show 32

normal-appearing spores.

EVANS— POLYPODIUM 205

Apogamous reproduction is normally a process in which serogenesis is modi-
fied and sexual fusion is by-passed. It always involves plants with an unbalanced
genotype that could not, in the usual sexual cycle, form viable spores. The proc-
ess is known in many species of ferns and is considered to be quite widespread.
Wagner (1963) estimates that approximately 7% of the 72 pteriodphytes of Giles
Co, Virginia, U. S. A., reproduce apogamously. Mehra (1961) found 30 apogamous
ferns among 350 cytologically known ferns from the Himalayan region, or about
8.6%. The highest frequency of obligate apogamy is among triploids, although
apogamy is known at other ploidal levels, including the diploid (Mehra he. cit).

In order to compare the apogamous situation in P. dispersum with the more
usual apogamous sporogenesis, the following brief outline of the latter will pro-
vide a comparison. In obligate apogamy the 2X archesporial cell in certain de-
veloping sporangia divides mitotically three times forming 8 cells. There is then
a further chromosomal division without cytokinesis, and a reorganization of the
genetic material into single nuclei ("restitution nuclei"). This results in eight
4X spore mother cells from the initial 2X archesporial cell. Meiosis in these
doubled cells is then normal, but produces only 32 viable 2X spores per sporangium.

It should be added that these meiotic apogamous ferns usually also undergo
spore formation in another way. In some sporangia, the spores develop without
the formation of restitution nuclei. This is essentially the process of sporogenesis
in sexual species. However, because of the hybrid-like behavior of the chromo-
somes in apogamous ferns, in which the genomes do not pair normally except
in the restitution nuclei, this process results in irregular and aborted spores.

According to Manton (1950) these two processes can occur in different spo-
rangia on the same apogamous plant, in the same or different sori, and in quite
varying proportions, from only a few 32-spored sporangia per sorus to many
sporangia of this type per sorus.

The spores produced from 32-spored sporangia develop into gametophytes
which usually form sex organs. The archegonia are aborted, or may be absent;
but the antheridia and sperms may be functional. The new sporophyte develops
not from the sexual apparatus but directly from thallus tissue.

Ameiotic Alternation of Generations in Polypodium disperum
Obligate meiotic apogamy has been considered the only successful or func-
tional apogamous life cycle in ferns, and Manton (1950) termed the process as
"monotonously uniform." However, in the P. pectinatum-plumula complex, P.
dispersum displays a unique alternative in apogamous reproduction in ferns (Evans,
1964).

A study of sporogenesis in P. dispersum (Fig. 13) shows four complete mitotic
divisions of the archesporial cells to form 16 spore mother cells in all sporangia,
without restitution nuclei. The 16 spore mother cells are readily visible, for they
separate, enlarge, and stain more darkly than other cells in the sporangium (Fig.
13, C). All sporangia observed contained only 32 spores. This involved the ob-
servation of several hundred sporangia both in sporogenesis studies and on herb-
arium specimens. In metaphase figures Polypodium dispersum has 111 unpaired

206 ANNALS OF THE MISSOURI BOTANICAL GARDEN

chromosomes (Fig. 13, E) . Squashes of metaphase figures in the eight-celled stage
of the archesporium and in the 16 spore mother cells show that the chromosome
configuration is essentially identical, both stages showing 111 unpaired chromo-
somes (Fig. 13, A, D). The mode of cytokinesis of the 16 spore mother cells (Fig.
13, F, H) indicates that each mother cell produces only 2 spores.

The spores are thus formed by one additional mitotic division of the spore
mother cells, and meiosis, characteristic of all other known apogamous ferns, is
rbsent (hence my distinction of the two types of apogamy as "meiotic" and
'ameiotic"). This is an example of diplospory. Mehra (1961) reports that "di-
plospory which is so widespread in apomictic angiosperms has so far not been
observed as a functional mechanism in apomictic ferns, although there are stray
records of its occurrence." Reports of Trichomanes proliferum forma /? (Bell, 1960;
Mehra & Singh, 1957) and in Ophioglossum and Isoetes (Verma, 1956, 1960) sug-
gest that this process may also occur in pteridophytes other than P. dispersum.
However, none of the latter reports contain information pertaining to the gameto-
phyte generation or whether or not these plants are capable of completing their
life cycle. Braithwaite (1964) indicates a similar situation in Asplenium aethio-
picum to that observed here. He describes the formation of 32 spores from 16
spore mother cells as being due to a partial meiotic division and a restitution
nucleus. These would appear to be very similar processes in these two different

The irregularities in this process are few. There are occasional sporangia
which contain a lower number than 32 spores and some spore fragments. Occa-
sionally, mature spores are contracted medially or submedially as though the
cell had begun to divide again. I do not yet know whether these "pinched" spores
are viable. In the examination of several hundred divisions only one cell was
found that may have doubled its chromosomes. As I obtained over 60% germina-
tion of spores, I am quite sure that these few irregularities contribute little or
nothing to this germination percentage, and that the viable spores are formed as
outlined above.

The morphology of the Polypodium dispersum spore is a further departure
from the norm. All other species in this complex have the usual reniform, mono-
lete, polypodioid spores, which are formed in tetrads.

The P. dispersum spores, in contrast, are distinctly globose, and the scar is
exceedingly variable, being either incomplete or suggesting a monolete or trilete
pattern (Fig. 14). Occasionally the scar is absent. Spores of apogamous ferns
normally resemble the spores of closely related species. In the surface tubercula-
tion, spores of P. dispersum are like those of related species, but in the spore and
scar shape they are readily distinguishable microscopically.

In agar plates fortified with Knop's Solution, and grown under constant light
at 20°C, spores germinated in about 10 days, and development was rapid; young
apogamous sporophytes forming in about two and a half months.

The first rhizoid appears with, or even before, the first prothallial cell (Fig. 2,
A), and subsequent rhizoids develop with each additional cell, often more than

EVANS — POLYPODIUM 207

one rhizoid per cell. After one to three filamentous cells develop, an elliptical
plate is formed, and rhizoid initiation is reduced.

The cell plate of P. dispersum is not cordate, as are most polypodioid gameto-
phytes. At the time of sporophyte proliferation the prothallus is either small and
elliptical (Fig. 15, G), or it may be strap-shaped and several times branched (Fig.
2, D). The latter is typical of artificial cultures. In nature, both small elliptical
thalli, not much larger than the bases of the sporophytic proliferations, as well as
longer, branched thalli are found. Rhizoids are either marginal or near the
margin. If the thallus is branched, rhizoids are usually more dense at the conflu-
ence of the branches. They are never wholly basal and ventral, as is normal in
other sexual species of Polypodium. The shape of the thallus and the position of
the rhizoids suggest that P. dispersum has gametophytes like those of the Gram-
mitidaceae. However, the early development of both the rhizoids and the cell
plate are characteristically polypodioid. Also, the rhizoids of P. dispersum are a
light brown, not dark brown and stiff, as is typical of the Grammitidaceae (Stokey
& Atkinson, 1958).

Two features of Polypodium dispersum gametophytes are unique: the absence
of gametangia and the ability to produce stomata on a thallus one cell thick.
Previously known apogamous ferns produce sex organs even though they are not
functional in self-reproduction. No sex organs were observed in either wild or
cultured collections of P. dispersum gametophytes.

Stomates were observed on several of the gametophytes that had already
proliferated sporophytes. As the thallus is everywhere only one cell thick, the
stomatal apparatus merely consists of two typical guard cells over an opening
between thallus cells (Fig. 2, B & Fig. 15, D, E). Its function, if any, is unknown.

Stomates have previously been observed on apogamous gametophytes (Steil,
1919), but always at the base of the sporophyte and associated with cells already
modified in morphology toward sporophytic epidermal cells (elongate and with
undulate lateral walls). However, in P. dispersum, these stomates occur on lobes
without sporophytes (although always on gametophytes containing proliferations)
and in tissue with cells still strictly of gametophytic structure.

The sporophytic proliferation in Polypodium dispersum is first noticeable as
a small swelling on the dorsal surface, usually near the margin of the thallus (Fig.
2, 15). In the very small, elliptic gametophytes it arises from the center and
quickly occupies all but a small fringe of gametophyte tissue around the base of
the small sporophyte plant. This swelling very early produces both the simple and
ctenoid, clavate hairs of the juvenile sporophyte (Fig. 15, D, F, G). The swelling
may be only a few cells high or it may become 1 or 2 mm long before differentia-
tion takes place at the apex. A fine strand of two or three partially over-lapping,
spirally thickened tracheids appears in a central core of narrow cells in the swell-
ing. One or two tiny strap-shaped leaves with a delicate midrib, marginal acicular
hairs and simple and ctenoid laminar hairs arise, followed by a series of leaves like
those of sexually produced and root proliferated sporophytes. The first leaf (and
sometimes the second) appears before the roots. This is in contrast to the sexually

208 ANNALS OF THE MISSOURI BOTANICAL GARDEN

produced sporophyte, in which the first root appears with or usually before the
first leaf.

As in any obligate apogamous fern, the ameiotic apogamous cycle of P.
dispersum allows dispersion and colonization without the necessity of fertilization.
It is unique, however, in that the cycle has entirely the character of sexual repro-
duction with neither meiosis nor syngamy and the usual associated opportunities
for genetic assortment and recombination. At the same time that meiosis and
syngamy of the sexual cycle are missing, syndiploidy and meiosis of the meiotic
apogamous cycle are also missing. The only way to inject changes into the genome
of P. dispersum would be by random mutation and selection for or against this
mutation. But any mutations selected would then be carried by all spores of the
plant. Consequently, one might expect that this species would have a limited
ecological and geographical potential. Yet this species is frequently collected and
is widespread in the New World tropics (Fig. 9). Morphological variation from
one individual to another is about as great as in most of its relatives. There is even
one distinctive collection of numerous specimens (Wright 1051) with incised seg-
ments from Cuba, where the typical form is common. In spite of its seemingly
limited genetic potential, its somatic life cycle has evidently been quite successful
in adapting this species to its environment.

Cytology and Evolution

One of the major efforts in recent systematic studies of the Filicineae has been
to subdivide the classical all-encompassing and undoubtedly polyphyletic family
Polypodiaceae. Serious attempts at a more natural arrangement have led to such
revisions as those of Bower (1923), Christensen (1938), Ching (1940), Dickason
(1946), Copeland (1947), Holttum (1947, 1949), Alston (1956) and Pichi-Sermolli
(1959). Within the Polypodiaceae sensu stricto there has been, at the same time,
a subdivision of the classical genus Polypodium into presumably more natural
genera. Today there is a general agreement as to the extent of the genus Poly-
podium, although Copeland (1947) segregated out Goniophlehium Presl, a step I
do not think warranted on present evidence.

The Polypodium pectinatum-plumula group is more similar in certain gross
appearances to Ctenopteris in the Grammitidaceae than to the rest of the genus
Polypodium as a whole— so close, indeed, that Copeland (1956) placed P. trun-
corum.ia Ctenopteris and suggested that P. pectinatum and P. plumula might also
belong there. Copeland's paper inspired critical studies of the sporangia (Wilson,
1959 a, b) and gametophytes (Stokey, 1959) of the latter two species. Table 1
lists character differences between this complex and the Grammitidaceae.

I believe that these families, based primarily on the differences in spores,
sporangia, paleae, and gametophytes, should be maintained, although there still
remain vexing problems as to their origin. There are, basically, two lines of
thought— one that the Grammitidaceae arose from the Polypodiaceae (Pichi-
Sermolli, 1959; Christensen, 1938), and the other that the Grammitidaceae arose
independently from gleichenioid stock (Holttum, 1947, 1949; Stokey, 1951; Cope-
land, 1947). There are also dual propositions as to the origin of the Polypodiaceae,

EVANS — POLYPODIUM ZU9

Comparison of the Polypodium pectinatum-plumula complex with the

P. pectinatum-plumula complex Grammitidaceae

Paleae non-clathrate (rarely sub-clathrate), Paleae clathrate or not, often conspicu-
entire or partially and inconspicuously ously setose
papillate or fimbriate

Stipe rarely articulate
Short-spaced or approximate to fasciculate Mostly fasciculate

Sporangial stalk with 2-3 columns of cells Sporangial stalk with 1 column of c

Rhizoids appearing with first prothallial Rhizoids appearing after 10-15 prothallial

cell, or later cells have appeared

Plate early forming (after 1-2 months) Plate late forming (after 2 months- 1 year)

Simple papillate hairs common Simple papillate hai

one that they arose from dipteroid stock (Holttum, 1947, 1949; Pichi-Sermolli,
1959), the other that the Polypodiaceae also arose from gleichenioid stock (Cope-
land, 1947; Stokey, 1951). The origin of the Grammitidaceae from gleichenioid
ancestors appears to have much support in actual resemblances, i.e. the trilete
spores, free forked or simple venation, and the rather stiff pectinate blades. It has
been argued that the Polypodiaceae arose from dipteroid stock through forms like

210 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Dipteris, Matonia, and Cheiropleuria, which have complicated stelar anatomy and
anastomosing venation. However, one can also read the opposite sequence from
gleichenioid stock to the free- veined polypodia, perhaps much like the present
members of the P. pectinatum-plumula complex. One of several difficulties in
tracing evolutionary patterns in Polypodium is that of the free vs. anastomosing
venation. Christensen (1928) builds a case for the origin of free venation in the

Table 2. Chromosome observations.

Taxa, with collection & location Number

-Mexico, Oaxaca, Zacatepec, Michel 1613; Costa Rica, n-37

P. dispersum Evans— Florida, Citrus Co, Evans 2007, 2008, Wagner 62061, "2n"=l
Hillsborough Co, Evans 1189, 2019; Jamaica, Portland Parish, Evans
2400, 2402, 2403, Proctor 22896.

P. ferrugineum Mart. & Gal.— Mexico, Nayarit, Ceboruco volcano, Michel n=3

, hygrometricum Splitg.— Costa Rica, Puntarenas Prov, Evans 2733.

Rio Grande do Sul, Sao Leopoldo,

Parish, Evans 2588; Costa Rica, Alajuela Prov, Evans 2984-1.

rtumula Humb. & Bonpl. ex Willd.— Florida, Hernando Co, Wood,
s.n., Hillsborough Co, Evans 1186, Sumter Co, Evans 1190, 1192,
1193; Jamaica, Portland Parish, Evans 2398.

■ Co, Evans .
ica, Portland" Parish, Evans 2411, 2412, 2414; Mexico, San
Luis Potosi, Xilitla, Michel 602.

vtilodon Kunze var. ptilodon— Peru, Prov Bagua, Dept Amazonas,

P. recurvation Kaulf— Brazil, Rio Grande do Sul, Sao Leopoldo, Sehnen
P. siccum Lindm.— Brazil, Rio Grande do Sul, Sao Leopoldo, Sehnen s.n.

EVANS — POLYPODIUM 211

P. vulgare group from the goniophlebioid venation. In the group under present
study there are several species with partial to almost complete reticulation, species
which are also pubescent, as are most of the truly goniophlebioid species. These
reticulate venation patterns could be considered as either generalized or specialized
character states, but are here treated as examples of a condition derived from free
venation. The interpretation of this character is involved with the i
of the origin of the family, either from free-veined gleichenioid stock c
ing dipteroid stock. The answer is not immediately self-evident.

Cytology

Fourteen taxa were available for chromosome counts (Table 2). These are too
few to be of application in interpreting the whole picture of speciation in the
complex; however, all counts are useful. It was initially assumed that Polypodium
dispersum was a hybrid between P. plumula and P. ptilodon var. caespitosum, but
chromosome counts indicate that this is not likely. In the case of P. ptilodon var.
ptilodon and var. caespitosum, a difference in their chromosome counts helps in
interpreting them as at least separate varieties. Polypodium ferrugineum is quite
similar to P. cupreolepis, except that the former has larger spores. It might have
been considered a tetraploid derivative of P. cupreolepis, were it not known to be
a diploid.

Table 3 indicates mean spore and guard cell lengths (based on 10 measure-
ments per specimen and 3-10 specimens of each species or variety). Those cyto-
logically known taxa have their ploidal level also indicated. On the assumption
that spore and guard cell length are usually affected by polyploidy, this can give
some indication of possible ploidal levels in certain of the species. Unfortunately,
there are not enough cytologically known taxa to give real indications of such
levels, but certain suggestions can be made. The taxa are here arranged in descend-
ing order of length of spores.

In this table there is some indication that those taxa with guard cell lengths
of ca 45/u. or greater, correlated with a spore length of 50fi or greater, may be
polyploids. Conversely, those taxa with guard cells less than 40/u. long, correlated
with spore lengths of less than 47 y. could be expected to be diploids. On this
basis it is predicted that those taxa marked with an (*) are probable polyploids,
and those marked with an (o) are expected diploids.

Divergence Index and Common Ground Plan
The relationships of the P. pectinatum-plumula complex are diagrammed in
Fig. 16. This is a method for expressing evolutionary relationships developed by
Wagner (1961, 1962), at the University of Michigan and used by such previous
authors as Hardin (1957), Mickel (1962), and Scora (1964).

A list of as many characters as possible that show contrasting character states
within the complex is first compiled. The generalized and the specialized state
of each character is indicated. This is ascertained by noting the frequency and
distribution of a certain character state. The assumption is that the more common
the character state the more likely it is to be the generalized state. It is necessary

I ANNALS OF THE MISSOURI BOTANICAL GARDEN

Table 3. Comparison of spore and guard cell size

var. hchniferum
var. camptophyllarium
pectinatiforme

dispersum
camp tophyllar ium
var. abbreviatun

ptilodon var. ptilodon
eurybasis var. eurybasis
consimile var. consimde

54

n=74

57

54

r=74

52

52

50
50
50

n-37

39

65
41
40

50

rc=37

37

50

n=74

34

49

47
47

n=37

38
39

45

n=37

47

45
45

n=37

41
37

43

n=37

39

43

"2n"=lll

38

43
42

n=37

43

42
41

,=37

50
40

i by taking into account related taxa or groups of taxa outside
the one in question. In the analysis of a genus one must thus examine other
closely related genera. In the present case, members of the genus Polypodium out-
side the complex were considered. In this way, it is possible to evaluate with strong
probability a character state which appears frequently in a given group, but which
may actually be a derived character state when taking the overall group into
consideration. 2

EVANS POLYPODIUM 213

An initial list of characters, each showing paired character states, was selected.
However, certain of these characters did not appear to have direct evolutionary
significance; they had no obvious correlations. To simplify the diagram and reduce
the information to the most definite characters and trends, these fluctuating char-
acters were eliminated from the ground plan.

Each of the remaining 23 characters was then evaluated as to whether it is a
generalized or specialized character state, the generalized state being given the
value (0), and the derived state (1). This is based on the principle that, as we
do not really know any actual quantitative value for a specific character, it is best
to give them all equal values,
entire segments, and i
few or absent — however, I have given a value of 1.5 to these isolated and exag-
gerated expressions of an already specialized character state. Also, in evaluating
the character of basal segments perpendicular to the rachis (0) or deflexed (1), I
have added an intermediate level (0.5) for those cases which have a sporadic
expression of the specialized state. Such situations could be avoided by establishing
additional characters for these conditions, e.g. segments pinnatifid (1) versus seg-
ments crenate (0). But considering that the "ultra" specialized character state 1.5
in each case occurred in species obviously related to other species exhibiting the
specialized character state 1, this seemed to be the clearest path to take. The same
is true of the intermediate 0.5 state.

The total values of the character states for each entity are totaled, giving each
a quantitative index showing its approximate divergence from the hypothetical
ancestor, which had all the characters in the generalized state. This value then
indicates at which level, i.e. on which semicircle on the divergence figure, the
species will be placed. The position depends on the correlation of as many derived
character states as possible from one species to another. Where it is necessary to
cause a line to fork and there is no extant plant which "fits" this position, a
"hypothetical ancestor," or "prototype" to those following it on the lines (indicated
by open dots in Fig. 16) is inserted. This brings out an interesting point regarding
the use of these "hypothetical ancestors" within a very closely related group of
plants. Mickel (1962), in constructing a ground plan for the species of the genus
Anemia, subgenus Coptophyllum, had to use 19 "hypothetical ancestors" in
evaluating 41 taxa. Scora (1964) had to use 13 "hypothetical ancestors" in placing
22 taxa in the genus Monarda. Yet in the present study, in placing 35 species, only
8 were necessary. This may be due to the wider spread of morphological diversity
that usually occurs in the species of a whole genus or subgenus, as compared to
a single species complex within a genus. In the present study the species are

not so elsewhere in Polypodium and is therefore considered a derived character state. In
the same way, the black polished rachis is considered a derived character state, as it occurs
with less frequency than the red-brown polished rachis. In the case of the poli
non-polished rachis, this character is not used at all in the ground plan analysis because
this character state is present throughout the group and therefore has no evolutionary
significance within the complex, though it might well have significance in an evaluation
of the whole genus or family.

214 ANNALS OF THE MISSOURI BOTANICAL GARDEN

recognized to be closely related, and therefore the divergence pattern reflects this
closeness in that many species appear to be directly related in a linear or diverging

The following are the evaluations of the characters used in formulating the
common ground plan:

A. Rhizome: Throughout most of the genus, as well as the family, the rhizome
is long-creeping. In this complex several of the species of smaller stature, particu-
larly those allied with P. plumula, have a more specialized, short-creeping, sub-
erect or erect rhizome.

B.-C. Rhizome Paleae: The generalized shape of the paleae is narrow- or
linear- triangular. However, in several of the species, e.g. P. cupreolepis, P. alfredii,
P. ferrugineum, P. sursumcurrens, and P. filicula, the paleae are ovate or cordate
(B). One of the specialized features of practically the whole complex is the
comose rhizome paleae (C) (Fig. 1). Inconspicuously comose paleae also occur in
several species not closely allied to this complex as judged by other characters,
which have been excluded from the present treatment. There are a few of the
included taxa allied with P. plumula which characteristically lack this comose
condition. Since palear comosity is probably specialized, those species which have
not developed it are considered generalized in this respect.

D. Rachis Color: The rachis is lustrous or polished, and though this is un-
doubtedly a specialized condition, it is a null character as far as divergence within
the complex is concerned. However, there are essentially two color phases: red-
brown in the P. pectinatum allies and either red-brown or black in the P. plumula
allies. As stated above, the black condition is considered specialized.

E.-F. Rachis Firmness: Though most of the group have a stiff and erect rachis,
as do most species of Polypodium in general, several members have specialized
springy elastic rachises, readily recognizable on herbarium sheets (E). There are
also two species, P. curvans and P. choquetangense, that show an additional
specialization in the twisting or curling of the rachis under normal growth condi-
tions (F). (This is not to be confused with the curling of the rachis and costa
causing the frond to curl up under dry conditions.)

G. Rachis Pubescense: Although the whole group is characterized by a variety
of types of rachis trichomes, and the rachises vary from nearly glabrous to pilose,
P. eurybasis var. villosum has a striking densely-villose rachis distinguishing it
from all other species.

H. Ctenoid Hairs of the Rachis: Although inconspicuous ctenoid hairs can
probably be found on any species of Polypodium, at least on juvenile fronds, the
species of the P. pectinatum group are noteworthy because of the numerous, large,
conspicuous ctenoid hairs developed also on the mature rachis (Fig. 1, F). This
is considered a specialized condition.

I. Rachis Paleae (Fig. 5) : Though the rachis paleae vary considerably through-
out Polypodium, in this complex and in those polypods most similar to those of
this complex, the inconspicuous, long, thin, filiform type of palea is most frequent
and therefore considered primitive. However, a number of species allied with P.
plumula exhibit numerous hastate-triangular or cordate, conspicuous paleae on the

215

rachis, which are probably specialized. They are either concolorous or have a dark
margin and paler base (Fig. 1, H) and are thus different from the clathrate paleae
with pale margins in the P. squamatum and P. vulgare groups.

J. Frond Size: Though there is considerable size variation in Polypodium (Fig.
3), there are a few species which are significantly reduced in size. These are
represented here by P. ferrugineum, P. filicula, and P. camptophyllarium var.
abhreviatum, which have fronds generally less than 25 cm long.

K. Length of the Stipe: Whereas the majority of the species of Polypodium
have a well-defined stipe that is readily disguishable from the rachis, several species
have the specialized condition of sessile or nearly sessile blades, as represented in
the P. pectinatum-plumula group by P. consimile, P. filicula, and P. ptilodon var.

L. Blade Base: The most general blade outline in Polypodium is elliptic-ovate,
or linear with a cuneate or subtruncate base. However, several species, here repre-
sented by P. bolivianum and P. recurvatum, have a specialized, very abruptly
truncate blade base (Fig. 3, G).

M. Basal Segments: Most polypodioid ferns have segments that are all essen-
tially perpendicular to the rachis, and this is considered a generalized character.
Certain species, however, show a distinctly different orientation at the base of the
blade, with the basal segments becoming increasingly deflexed (Fig. 3). This
usually creates an asymmetry at the segment base in which the upper edge meets
the rachis at a broadly confluent angle, whereas the lower edge meets the rachis
at a much sharper angle (see character 0). Species with basal segments irregularly
deflexed are given an intermediate value of 0.5.

N. Segment Orientation: Pinnatifid polypodioid ferns commonly have either
straight or slightly falcate segments which are essentially perpendicular to the
rachis or slightly ascending (usually not more than 20° above the horizontal).
The group of P. absidatum, P. curvans, and P. choquetangense have distinctly
ascending segments exceeding ca 30° above the horizontal (Fig. 3, C).

O. Segment Base Symmetry: The segments of pinnatifid polypodioid ferns
usually have an essentially symmetrical base, expanded and confluent on the
rachis. In several species of this complex the lower edge of the segment meets
the rachis at a right angle or may even be partially incised on the lower side,
creating a semi-pinnate condition. This character has only been considered as
specialized, however, when the segments are at right angles to the rachis (Fig. 3,
A). When the segments are either ascending or deflexed this is automatically re-
flected in an asymmetry at the base.

P. Segment Margin: Whereas the majority of these species have the generalized
condition of segments with entire margins, the species allied to P. eurybasis and
the P. curvans — P. choquetangense line have crenate margins, as do P. paradiseae
and P. sursumcurrens (Fig. 3, Fig. 6). In P. choquetangense (Fig. 6, T) the seg-
ments are deeply pinnatifid, and I have assigned this character state the "ultra"
value of 1.5.

Q.-R. Laminar Pubescense: This refers only to the relatively conspicuous
acicular, non-appressed hairs, since all members of this group have inconspicuous

[Vol. 55
216 ANNALS OF THE MISSOURI BOTANICAL GARDEN

clavate hairs. The presence of a conspicuous pubescence is considered derived from
the essentially glabrous condition (Q), and the presence of definite patterns of
distribution of these hairs, as in the P. ptilodon group (Fig. 1, E), is considered
an additional derived character state (R).

S. Lateral Veinlets: The great majority of polypodioid ferns have either
anastomosing or free forked venation patterns. However, the specialized state of
simple venation is present in this group in P. truncorum, P. siccum, and P. fdicula
(Fig. 6).

T. Sorus Position: In the free veined species of Polypodium the sorus is always
terminal on the first acropetal veinlet and is usually medial between the costa and
the laminar margin (Fig. 6). However, in several instances, the fertile veinlet
typically extends almost to the margin, thus reflecting the condition of comparable
veinlets in a sterile segment. In these instances the sori are then marginal or sub-
marginal, and this is considered a derived character state.

U. Receptacular Paraphyses: In almost all taxa the receptacular paraphyses
are composed of uniform cells and are similar to the clavate hairs of the lamina,
though they are occasionally branched. They are universally present in this com-
plex, and in most of the species examined in the rest of the genus and the family
Polypodiaceae, and so it is believed that this simple paraphysis type is generalized.
However, P. sursumcurrens exhibits an unusual paraphysis with an irregular sub-
terminal multicellular clavate head (Fig. 4, P). As the head is composed of un-
differentiated, though contorted, cells, it is considered to be derived not from re-
duced sporangia ("sporangiasters" as in P. virginianum) , but from coalesced cells
of a strongly twisted simple type of paraphysis. Polypodium camptophyllarium var.
macedoi exhibits another distinctive paraphysis type; this is regularly forked, but
one fork is tipped by a clavate cell, the other by a sharply setose cell (Fig. 4, M).
The small appressed hairs of the lamina of this species are similar to the paraphyses.

V. Spore Length: While great length may be a simple morphological specializa-
tion by itself or reflect polyploidy, in either case it represents a divergence from
the ground plan and is thus considered specialized.

W. Number of Spores Per Sporangium: In the higher ferns, 64 spores per
sporanguim is a regular occurrence and has become a standard for indicating the
normal sexual life cycle. Similarly, 32 spores per sporangium indicates an apoga-
mous asexual reproduction, as is the case in P. dispersum. Apogamy is considered
a specialized type of life cycle.

Taxonomic Revision of the Polypodium pectinatum-plumula Complex 3

In treating this group of closely related species of Polypodium, there are no
clear-cut limits in inclusion and exclusion. The species here included are circum-
scribed by the characters of blades with pectinate segments, essentially free veins,
approximate to fasciculate fronds, rigid or elastic, lustrous, red-brown or black
stipe and rachis, triangular and comose or cordate and non-comose, non-clathrate

3 Until a broad

and thorough <
'isest to withhc

EVANS— POLYPODIUM 217

rhizome scales, 2-3 rowed sporangial stalks, and reniform spores. The species of
closest superficial resemblance in Grammitidaceae are excluded on the basis of the
sporangial stalk, spores, and other characters (see Table 1 above). Certain species
of Polypodium have been excluded that might, in a wider treatment of the whole
genus, be included here in the future. Polypodium hartwegianwn, P. chnoophorum,
P. moritzianum, and others have inconspicuously comose rhizome scales but have
been excluded on the basis of the lack of the lustrous rachis, the different texture
of the lamina, or the more crisped silvery laminar trichomes. I have, on the
contrary, included P. hygrometricum and P. recurvatum, which perhaps are more
properly placed with these excluded species. However, the latter two have been
particularly confused with the included species both as to taxonomy and nomen-
clature. Future work should be directed at delimiting the different groups (possibly
as subgenera) within Polypodium sensu stricto so that all the species can eventually
be grouped and treated systematically.

The measurements of rhizomes, fronds, blades, and segments include a mean
size bracketed by an expected maximum and minimum limit. These were compiled
by measuring ten random specimens of each recognized entity. "Ctenoid hairs"
refer to small, appressed, multiple-branched hairs with all the branches arising
from one side of the main axis (Fig. 1, F & Fig. 4, I). "Comose" rhizome paleae
refer to those with tufts of unicellular hairs arising from the dorsal surface (Fig.
1 ) . The costa is "decurrent on the rachis" when it joins the rachis in a downward
arc to form a short wing along the rachis and a visible curvature of the costa.
When the costa meets the rachis at a right angle, there is also a slight expansion
on the rachis but no visible curvature of the costa. In defining plane shapes (e.g.
fronds, segments, scales), I have attempted to follow the Systematics Association,
Descriptive Terminology, Chart 1 (Taxon 11: 145-156, 1962), with the addition
of the term "cordate," as it so usefully applies to the overall shape of many paleae
(Fig. 5). All observations have been made from mature fertile fronds, and all
characters of rachis, costa and lamina, unless otherwise noted, refer to the abaxial
surface only.

The colors of the paleae, rachis, and costa are treated as black or one of the
following three shades of red-brown. Because there is no satisfactory equivalent
for the intermediate color, I have used the terms "light red-brown" (ferrugineous),
"red-brown", and "dark red-brown" (castaneous).

As a result of the close morphological similarity of many of these species, it
has been necessary to rely heavily on what may appear to be quite minute and
technical characters. In evaluating the shape or comosity of paleae, or the size,
shape, and distribution of hairs, it has often been necessary to use optical magnifi-
cations beyond the hand lens and up to 20 to 30 X or more.

The laminar trichomes consist basically of two types. One is the appressed,
clavate, simple, forked, or ctenoid hair. These are rarely over 250/* long and would
be generally overlooked in routine identifications, except when they are conspicuous
on the stipe base in the larger members of the P. pectinatum group. The second
are conspicuous, acicular, simple hairs from 0.25 mm to 2.0 mm long. These
! the generally visible pubescence of the frond. However, in the descrip-

218 ANNALS OF THE MISSOURI BOTANICAL GARDEN

tions both the conspicuous acicular hairs and the inconspicuous clavate hairs are
referred to. Because the clavate hairs would often be overlooked, certain species
may appear superficially glabrous, when in fact small clavate hairs are undoubtedly
present. In these cases, the description may refer to the lamina as "essentially
glabrous."

Where the soral size is important, it is indicated by the approximate portion
of the width of the segment that two sori occupy, e.g. "occupying half the width
of the segment." Only the mature sori are referred to in this :

• THE POLYPODIUM

Rachis paleae relatively conspicuous, persistent, other than filiform; rachis

or black with lateral brown stripes; rachis paleae narrow-elliptic or cordate
with acute apex, entire; sporangia with c
3. Rachis paleae narrow-elliptic, few an

basal segments of the blade strongly i

rachis red-brown or black with lateral brown stripes 5. P. alfredii

3. Rachis paleae cordate, numerc

segments perpendicular to the rachis, with symmetrical bases; sporangia

4. Rhizome paleae light golden color; rachis red-brown but costa
black or darker brown than the rachis; sporangia with 2 long setae
6. P. ferruginei

4. 3wn; rachis and costa both red-brown; spor-

Rhizome paleae narrow-triangular with acuminate apex, comose; rachis
red-brown or black without lateral stripes; rachis paleae cordate,
row-triangular with acuminate apex and hastate base, inconspicuously
toothed or fimbriate; sporangia always setose, sometimes inconspic
5. Fronds 10 (6-17) cm long; rachis paleae cordate, dark brown with dark

base, inconspicuously toothed; veins unforked 10. P. jilicula

5. Fronds 15-65 cm long; rachis paleae cordate or narrow-triangular and
hastate, dark red-brown, lighter at the base, fimbriate; veins 1-2-forked.
6. Fronds narrow-ovate or linear; segments linear, the basal segments

shorter, not denexed, reduced to mere auricles; veins 1-forked ....9. P. plumula
6. Fronds narrow-ovate; segments narrow-ovate, the basal segments
shorter, denexed, sometimes reduced to auricles; veins 2-forked.

7. Rachis and costa brown; Bermuda 8. P. hermudianum

7. Rachis and costa black; not from Bermuda.

8. Rachis paleae subulate with hastate base; lower segments
only slightly shortened, strongly deflexed; 64 reniform spores

i reduced to auricles, slightly
32 globose spores per sporangium; wide-ranging
11. P. dis

• and inconspicuous; rachis

; as long as the 1

; 1-forked 26. P.

>r 3-forked.

tminate, gray-green, puberulent with short silvery

-. - 25. P. recurvatum

. Segments acute to obtuse, green, glabrous 16. P. bolivianum

EVANS — POLYPODIUM

Basal segments less than half as long as the longest segments,

to wings, lobes or auricles.

12. Veins simple; rhizome paleae glabrous.

13. Segments ascending 15°-25° above the horizontal;
(2.5-4.5) mm wide; rhizc

13. Segments perpendicular
iow seg ™ n r ts , 1_ ? mm wide » rhizome paleae light , ^- u

l em L'V7 k f ; r i izome p / leae ™r se (except ;

14. Segments strongly ascending 25°-50° above the horizontal, crenulate
to pinnatihd.

15. Segments pinnatifid almost to the costa .... 15 p choqueta

15. Segments merely crenulate or crenate.

16. Blades herbaceous; segments 2(1.5-3) mm wide; hairs of the
rachis ca 1 mm long; apex of the blade usually circinate

' .' n > ^i I ti 1 , ,

16. Blades coriaceous to herfc

wide; hairs of the rachis ca 0.5 mm long; apex of the blade

"•■wily straight at maturity 13. P

14. Segments perpendicular to 1

patch around the sorus, other-
hairs (then these
gradually re-
23. P. p

19. Lower edge of the segments perpendicular t
at least in ' '
20. Spaces between si

segments; clavate hairs on the lar
short, blackish; sori submarginal to r
receptacular paraphyses twisted an<
terminal clavate multicellular head;

ginal; receptacular paraphyses simple and straight.
2L ?o a( S 2 " 15 times as lon S as the s tipe; fronds

28-130 cm long; blade 4-26 cm wide 17. P. eurybasis

21. Rachis 4-14 times as long as the stipe; fronds "
19-55 cm long; blade 3-7 cm wide.
22. Rachis paleae linear; sori medial; rhizome
paleae narrow-triangular, red-brown; north-

00 ^ rn , Ar g en 1 tina and southern Brazil 3. P. singer i

22. Rachis paleae narrow-elliptic; sori medial to
submarginal; rhizome paleae ovate to cordate,

brown; Mexico and Central America 5. P alfredii

. Upper and lower edge of the segments similar, equally
expanded and adnate on the rachis; veins 2-forked
23. Rachis 13 (6-20) times as long as the stipe; segments
reduced at blade base to obtuse or rounded lobes,

23. Rachis 2-5 tn '" , Jn/re- 00 ™"^
duced at the blade base but not to mere lobes, nar-
row-triangular, acute at the apex, crenulate or
crenat8 17. P. eurybasis

ANNALS OF THE MISSOURI BOTANICAL GARDEN

24. Lamina thinly

ovate, red-brown with dark bro
. Lamina pilose with pale or go!
sori medial to nearly marginal;
or partially anastomosing; rhiz
linear-triangular, red-brown, lig
25. Costa meeting the rachis ai

27. Fronds less than 30 em long; veins 1-forked, free

throughout ....22d. P. camptophyllarium var. abbr

27. Fronds more than 30 em long; veins 2(1) -forked,

always at least partially anastomosing; spor-

29. Lamina ' puberulent with hairs usually
much shorter than 0.2 mm long; spor-
angial setae ca 65> long; segments
usually abruptly reduced to 2 or 3 pairs
of lobes at the blade base, perpendicular
to the rachis throughout, the basal
lobes expanded and symmetrical

...22. P. camptophyllarium

1. Polypodium truncorum Lindm., Hedwigia 43: 309, 1904.

P elasticum Rich var. glazhvii Baker in Mart., Fl. Bras. 1(2): 517, t. 64, fig. 2 1870.
' (Svntvpes- B irina, Muller s.n. not seen; Rio de Janeiro, Glaziou

1723 ! 2467 not seen; Minas Gerais, Caldas, Regnell II 319* MO)
P. baked Lindm., Ark. Bot. 1:240, t. 11, fig. 9, 1903 non Luerss 1882. (Lectotype: Brazil,

Minas Gerais, Caldas, Pedra Branca, Regnell II 379*a S; isolectotype S-PA)
Ctenopteris glaziovii sensu Copel., Phil. Jour. Sci. 84: 416, 1956, p.p., not as to type.
i (Lindm.) Copel., loc. cit. 450.

EVANS — POLYPODIUM 221

Rhizomes very short-creeping or erect, 1.5 (1-2) mm in diam; rhizome paleae
narrow-triangular, brown to dark brown, lustrous, basifixed, non-comose, non-
clathrate, the margins entire; fronds 25 (8-42) cm long, caespitose on the rhizome;
rachis 9 (6-20) times as long as the stipe; stipe and rachis red-brown, with scat-
tered whitish or golden acicular hairs, ctenoid hairs absent; rachis paleae incon-
spicuous or absent, if present then linear to short-filiform, entire; blades narrow-
to linear-ovate or -obovate, 23 (7-38) cm long, 3.5 (2.5-4.5) cm wide, subtruncate
to cuneate at the base; segments 19 (13-23) mm long, 3.5 (2.5-4.5) mm wide,
narrow-triangular, acute, expanded and symmetrical at the base, chartaceous to
membranaceous, slightly toothed, ascending ca 65°-75° from the rachis, slightly
reduced but not deflexed at the blade base, the basal pair usually reduced to auricles,
or occasionally blades gradually reduced to a cuneate base; lamina essentially
glabrous except for a few long, clavate hairs; costae perpendicular to or slightly
ascending on the rachis, with few long acicular hairs; veins simple, free; guard
cells ca 41^ long; sori medial to extramedial, round, with long simple clavate
paraphyses; sporangia without setae; spores globose-reniform, smooth, ca 45/x long.

Type: Based on P. bakeri Lindm., 1903, not Luerss., 1882. Regnell's numbering
system was complex; 7/ 319a, II 319*, II 319*c and 77 319*d have been examined and
they represent other species, the two last named numbers being P. singeri.

Habitat: epiphytic, mostly on tree-fern trunks, occasionally terrestrial, in wet
forests, 500-1800 malt

Distribution: southern Brazil and northern Argentina.— Fig. 11.

Bbazil: minas gerais: Canjerana, Regnell II 319* p.p. (MO, non US); Caldas, Regnell
II 319*b (S-PA); Carangola, N of Serra da Grama, Mexia 4272 (GH); Vicosa Fazenda de
Aguada, Mexia 5105 (GH). mo de Janeiro: Tijuca, Dusen 5022 (MO); nr Rio de Janeiro
& Bahia, Webb s.n., in 1867-1868 (MICH), sao paulo: Alto da Ssrra, Estagao Biologica,
Smith 1844 (MICH); Campos do Jordao, Leite 3562 (GH, MO); Serra da Bocaina, Brade
20799 (MO), 20801 (MO); Serra da Cantareira, Eiten & de la Sota 2170 (US), parana:
Serra do Mar, Ypiranga, Dusen 3651 (GH, MO); Desiro Ypiranga, Dusen 6776 (MO).
Santa catarina: Ararangua, Serra da Pedra, Reitz C271 (US); Azambuja, Brusque, Reitz
1795 (US), Smith & Reitz 6136 (GH, MO); Brusque, Mata Hoffmann, Reitz 3071 (US);
Ibirama, Horto Florestal, Smith & Klein 7554 (US); Itacorobi, Santa Catarina I, Rohr 320
(US); Luis Alves, Itajai, Reitz 159 (US), rio grande do sul: Sao Leopoldo, Reitz 1314

Argentina: misiones: Yerbal Viejo, Burkart 1584 (GH); Cainguas, Qae Mayo, Montes
27.218 (US).

I have not seen the type of P. glaziovii Bak. (Glaziou 9062, presumably at BM
or P), which I believe to be a Ctenopteris. Copeland (1956) transferred P. glaziovii
Bak. to Ctenopteris, using Baker's original description and citing the type specimen,
but in a way indicating that he had not seen it. He then stated, "I find the veins
simple; the spores tetrahedral, the stipes glabrescent," and cited eight collections.
I have examined five of these collections. The spores of these are not tetrahedral.
The veins are simple, as Copeland stated, but this alone is insufficient for separation
of Ctenopteris and Polypodium in all cases. The stipes are glabrescent, but with
polypodioid articulate hairs and not ctenopteroid unicellular setiform hairs. These
five sheets are all P. truncorum. It is evident thus that Copeland had a confused
concept of his Ctenopteris glaziovii (Baker) Copel.; the type may be a true

222 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Ctenopteris, but at least five of the collections cited by Copeland belong to
Polypodium.

This species is very similar to P. siccum, from which it can be distinguished
by the very small rhizome with crowded stipes, the dark chestnut rhizome scales,
membranous frond texture, and the narrow ascending segments with the basal
pair reduced to mere auricles and the next pair only slightly reduced. That it is
a true Polypodium and not a Ctenopteris was shown by de la Sota ( 1963) .

2. Polypodium siccum Lindm., Ark. Bot. 1:234, t. 11, fig. 4, 1903.

P. heteroclitum Fee, Crypt. Vase. Bras. 1:93, t. 26, fig. 4, 1869, non Desv., 1811 (Type:

Brazil, Rio de Janeiro, Glaziou 2410 BR, C, P)

Rhizomes short-creeping to erect, 2 (1-3) mm in diam; rhizome paleae nar-
row-triangular, light red-brown, basifixed, non-comose, non-clathrate, entire;
fronds 27 (10-50) cm long, caespitose on the rhizome; rachis 8 (4.5-11) times
as long as the stipe; stipe and rachis red-brown, pilose with long acicular simple
hairs, and also with long clavate hairs, ctenoid hairs absent; rachis paleae in-
conspicuous or absent, if present then linear to filiform, light red-brown, entire;
blades linear to narrow-ovate, 23 (8-36) cm long, 3 (2-5) cm wide, cuneate to
narrow-cuneate, occasionally subtruncate at the base; segments 15 (9-22) mm
long, 1.5 (1-2) mm wide, perpendicular to the rachis, reduced to a narrow wing
at the blade base, linear-triangular, chartaceous to herbaceous, acute, slightly
toothed, expanded and symmetrical at the base, or with the lower edge perpendicu-
lar to the rachis in the lower part of the blade, the sinuses approximately equal
to the width of the segment; lamina with numerous long golden clavate hairs;
costa decurrent on the rachis, with scattered long acicular hairs; veins simple,
free; guard cells ca 37^ long; sori submarginal, round, with simple clavate para-
physes, ca 0.3 mm long; sporangia without paraphyses; spores globose-reniform,
smooth, ca 45,u long; n=37.

Lectotype: Brazil, Rio de Janeiro, Glaziou 3339 S; isolectotypes BR, C, S-PA.
Syntypes: Brazil, Cafioas, Rio Grande do Sul, Regnell I A391 G, GH, NY, S,
S-PA, US; Brazil, Hamberger Berg, Rio Grande do Sul, Regnell I A501 S, S-PA;
Paraguay, Cordillera of Villa Rica, Balansa 667 B, BR, G, S-PA.

Habitat: epiphytic, often on tree ferns, in forests, 450-1200 m alt.

Distribution: southeastern Brazil, Paraguay and northern Argentina.— Fig. 10.

Brazil: rio de Janeiro: Novo Friburgo, Dusen 1899 (US), Leite 4104 (MO); Rio
de Janeiro & Teresopolis, Clarke s.n. (US) ; Santo Antonio, Serro dos Orgaos, Brade 16317
(MO) sao paulo: Iguape, Rio Peronpaua, Brade 7737 (US); Morro das Pedras, Brade
s.n., in 1918 (US). Parana: Lapa, Braga 1041 (US); Serra do Mar, Ypiranga, Dusen 14433
(GH, MO); Villa Velha, Dusen 14813 (US); Volta Grande, Dusen 14141 (US), santa
catarina: Mun. Curitibanos & Campos Novos, Smith & Klein 8287 (US)

Will (MO); Lages, Spannagel 67 (US); Mun. Porto Uniao, Smith & Reitz
8672 (US) rio grande do sul: Caiioas, Regnell IA391 (G, GH, NY, S, S-PA, US); Neu-
Wurstemberg, Bornmuller 120 (GH); Sao Leopoldo, Anchieta 2664 (US), Chacara Meyer,
Reitz 161 (US); Regnell 1A501 (S, S-PA).

Paraguay: E of the Vilarica Mts, Balansa 667 (B, BR, G, S-PA).

Argentina: corrientes: General Paz, Paraje Angostura, Ibarrola 3819 (MO) ; Mburu-
cuya, Santa Teresa, Petersen 1758 (MO, US), misiones: Loreto, Burkart 1551 (GH);
Posadas, Bonpland, Ekman 66 (MO).

EVANS— POLYPODIUM 223

This species is best recognized by its long, linear, delicate blades, submarginal
sori, and bright ferruginous rhizome paleae. Within its range, the only similar
species is P. truncorum, which has, however, much wider segments and darker
paleae. Polypodium siccum (like P. truncorum) strongly resembles the ctenopteroid
ferns, but in the critical characters of spores, sporangial stalks, and paleae, it is
properly placed in Polypodium.

One specimen indicates that this species reproduces in part by root prolifera-
tions, though apparently not so commonly as in P. dispersum or P. filicula.

3. Polypodium singeri de la Sota, Opera Lilloana 5: 181, 1960.

P. pectinatum L. var. awita Rosenst., Hedwigia 46: 138, 1906, non P. auritum Lowe, 1858.

(Holotype: Brazil, Lages, Spannagel 116a S-PA)
P. pectinatiforme Lindm., loc. cit. 43: 309, 1904, p.p., not as to lectotype.

Rhizomes short-creeping, 3 (2-4) mm in diam; rhizome paleae narrow-tri-
angular, red-brown, lustrous, basifixed, acuminate, non-comose, non-clathrate,
entire; fronds 25 (19-30) cm long, caespitose; rachis 6 (3-8) times as long as the
stipe; stipe and rachis red-brown, with scattered golden, acicular hairs, ctenoid
hairs absent; rachis paleae inconspicuous, red-brown, linear, entire; blades narrow-
ovate, 22 (15-27) cm long, 4.3 (3-6) cm wide, cuneate at the base; segments 22
(15-25) mm long, 3 (2.5-3.5) mm wide, perpendicular to the rachis, strongly de-
flexed at the blade base or reduced to auricles, linear-ovate, obtuse at the apex,
asymmetrical at the base with the upper edge widely adnate on the rachis and the
lower edge perpendicular to the rachis or incised, occasionally auriculate, charta-
ceous, entire, the sinuses equal to or narrower than the segments; lamina essentially
glabrous except for scattered clavate, simple hairs; costa perpendicular to the rachis,
with scattered, golden, acicular hairs; veins simple or once-forked, free; sori medial,
round, with simple clavate paraphyses; sporangia without setae or with one capsular
seta ca 115^ long; spores reniform, slightly tuberculate, ca 32^ long.

Holotype: Argentina, Misiones, San Ignacio, Loreto, 2 Febr 1958, Cristobal,
Ahumada, & de la Sota 53 LIL, not seen; isotypes BR, S, US.

Habitat: epiphytic on very small tree stems and tree-fern trunks, occasionally
terrestrial, in humid forest, 700-1200 m alt.

Distribution: southeastern Brazil to northern Argentina. — Fig. 8.

Brazil: minas gerais: Caldas, Regnell II 319* p.p. (US, non MO), 77 319*c (S), 77
319*d (S); Passa-quatro, Brade 19006 (MO), sao paulo: Campinas, Heiner 504 (S), 616
(S-PA). Parana: Jaguariahyva, Dusen 14880 (GH, MO); Rio Negro, Annies (Rosenstock
113) (US); Villa Nova, Annies (Rosenstock 113) (S, S-PA, US); Vierdel 1* (S-PA). santa
catarina: Sao Carlos, Chapeco, Reitz 3769 (US), rio grande do sul: Silveira Martins,
Santa Maria, Pivetta 158 (US); Serra Leitao, Jurgens 105 (S); Santa Cruz, J
(Rosenstock 113) (US), (Rosenstock 113a) (S-PA); Regnell 750 (S).

Argentina: mtsiones: Candelaria, La Pastora, Montes 2340 (MO); Loreto, Moreau
), San Ignacio, Montes 2249 (GH); Posados, Bonpland,
s.n. (S), Ekman 67 (S, S-PA), 81 (S).

This species has a curious habit due to its vegetative reproduction by root
proliferation. It is usually found growing on small upright twigs as a series of
short interconnected rhizomes. The whole appears as a layer of roots surrounding
small stems with a series of short rhizomes imbedded in the root mass. The species

224 ANNALS OF THE MISSOURI BOTANICAL GARDEN

is well characterized by this curious serial arrangement of rhizome, the conspicu-
ously asymmetrical and glabrous segments, the small size of the frond, and the
non-comose rhizome scales. It is apparently closely related to P. alfredii of Central
America, although well separated by the broader blade, more asymmetrical seg-
ments, and the longer rachis paleae.

Within this species there appear to be two variations. One form has the seg-
ments with smooth entire margins and the sporangia generally without setae, while
the other tends to have slightly undulate margins and one seta on most of the
capsules. The plants are alike in all other respects, however, so that unless careful
population studies in the future indicate otherwise, I believe them to be taxonomi-
cally inconsequential. Rosenstock based his P. pectinatum var. aurita on a leaf
form with narrow excurrent auricles on the lower edge of the segment.

4. Polypodium cupreolepis A. M. Evans, sp. nov. — Fig. 17.

Rhizoma breviter repens, paleis ovatis vel cordatis, pallide vel fusco-brunneis,
non comosis, integris, basi affixis, apice acutis, cellulis non clathratis; frondes ap-
proximatae articulatae, ca 30 cm longae; stipites et rhachides rigidae nitentes brun-
neae, pilose dissitos ferentes; paleae rhachidis numerosae consipcuae cordatae,
aureae vel ferrugineae, integrae, apice acutae; lamina pectinata anguste ovata, ca
22 cm longa et 4 cm lata, basi abrupte cuneata vel subtruncata; segmenta linearia
integra, rhachide perpendicularia vel interdum basalia paullo deflexa; lamina solum
cum pilis inconspicuis clavatis; venae liberae, 1-furcatae; sporangia sine setis;
sporae late reniformes laeves, ca 36fi longae.

Rhizomes short-creeping 4.5 (3-7) mm in diam; rhizome paleae ovate to
cordate, pale to dark brown, non-lustrous, basifixed or basally cordate, acute, non-
comose or rarely inconspicuously comose, entire; fronds 30 (10-60) cm long, ap-
proximate on the rhizome; rachis 3 (1.5-5) times as long as the stipe; stipe and
rachis brown, with long, scattered, acicular hairs; rachis paleae numerous, conspicu-
ous; broadly cordate, bullate, acute, basifixed, golden to light red-brown, non-
lustrous, entire; blades narrow-ovate, 22 (9-47) cm long, 4 (2.5-6.5) cm wide,
abruptly cuneate to subtruncate at the base; segments 20 (12-32) mm long, 2.5
(1.5-3) mm wide, perpendicular to the rachis, or occasionally slightly ascending
in the upper part of the blade, strongly reduced and sometimes deflexed at the
blade base, herbaceous, obtuse to acute at the apex, subsymmetrical at the base or
the lower edge perpendicular to the rachis, entire; lamina essentially glabrous, but
with inconspicuous clavate hairs; costa decurrent on the rachis, with scattered
acicular hairs; veins once-forked, free; guard cells ca 31^ long; sori medial, round,
with numerous, long, clavate paraphyses; sporangia without setae; spores broadly
reniform, smooth, ca 36^ long.

Holotype: Mexico, Michoacan, hills of Patzcuaro, 8 Nov 1890, Pringle 3353 US;
isotypes, B, GH, MICH, MO, US.

Habitat: epiphytic or epipetric, occasionally terrestrial, in montane forests,
1100-2900 malt.

EVANS — POLYPODIUM 225

Distribution: western and southern Mexico to Costa Rica. — Fig. 10.

Mexico: sinaloa: betw Durango & Villa Union, Ownbey 1939 (US); Ocurahui, Sierra
Surutato, Gentry 6337 (GH, MO), durango: 22 mi E of Santa Luc.

to Durango, Reeder 2487 (US), jalisco: 15 mi SSE of Autlan, Wilbur 1927 (US); below
Canoa de Leoncito, McVaugh 13486 (US); Santa Monica, McVaugh 14042 (US); 7 mi
SSW of Tecalitlan, McVaugh 16254 (US), hidalgo: Trinidad, Pringle 13475 (US), puebla:
Ixtaccihuatl, Purpus 1828 (GH, MO, US), morelos: Valle del Tepeite, Lyonnet 779 (MO).
distrito federal: Eslava, Lyonnet 3315 (US), 3376 (US), michoacan: Morelia, Campanario,
Arsene 8464 (GH, MO, US); Cerro Azul, nr Morelia, Arsene, 3 May 1910 (GH); Tancitaro,
Uruapan, Hinton et al. 15478 (GH, MO, US); Tancitaro, Leavenworth 336 (GH), 516a
(GH), Mt San Miguel, Leavenworth & Hoogstraal 1097 (MO);

Cacique, Hinton 13178 (GH, MO, US). Golima, Goldsmith 40 (GH);

Revillagigedo Islands, Socorro I, Mason 1633 (GH). Guerrero: Omilteme, Rowell 3049 A
(US); 2 mi W of Omilteme, Hamilton & Rowell 3245 (US), oaxaca: Totontepec, Nelson
796 (US), chiapas: Pico de Loro, nr Escuintla, Matuda 4268 (GH).

Guatemala: alta verapaz: Coban, von Tuerckheim 645 (US), quezaltenango: Cerro
Quemado, Kellerman 5940 (US), chimaltenango: Las Calderas, Standley 60040 (US).
escuintla: Volcan de Fuego, Salvin s.n. (GH).

El Salvador: santa ana: NE of Cerro del Aguila, Tucker 1249 (US), san vicente:
Volcan de San Vicente, Standley 21609 (US).

Nicaragua: jinotega: Jinotega, Howard 79 (US).

Costa Rica: san jose: Las Nubes, Scamman 7228 (GH).

This species has generally been identified as "P. pulchrum" the type of which
has, however, a black rachis and costa, triangular, lustrous, red-brown rhizome
scales, setose sporangia, and somewhat different rachis scales. I consider P.
pulchrum a synonym of P. plumula Humb. & Bonpl. Polypodium ferrugineum
Mart. & Gal., a smaller plant with two setae per sporangium, a black costa, a brown
rachis, and much paler golden-tan rhizome scales, appears to be its nearest relative.

The isotype at Berlin has been annotated by Hieronymus as "P. patzcuarense"
which seems to be an unpublished name.

5. Polypodium alfredii Rosenst., Repert. Sp. Nov. 22: 15, 1925.

P. alfredii var. curtii Rosenst., loc. cit. (Holotype: Costa Rica, region of Rio Chris, nr Juan

Vinas, 1200 m, Brade 695 S-PA; isotype US)
P. tablazianum Rosenst., loc. cit. 14. (Lectotype: Costa Rica, Tablazo, 1800 m, 4 Mar. 1908,
Brade & Brade 14 S-PA. Syntypes: Costa Rica, Tablazo, 1900 m, 20 July 1909, Brade
& Brade 696 NY; Costa Rica, Carpintera, 1500 m, 14 Dec 1909, Brade & Brade 149
NY, S-PA, US)
P. cyathicola Copel, Univ. Calif. Publ. Bot. 19:292, t. 43, 1941. (Holotype: Mexico, Vera-
cruz, Cuantlancillo (Orizaba) alt 1600 m, on Cyathea in wet ravine, 1 Mar 1938,
Copeland, Mex. Ferns 127 MICH; isotype US)

Rhizomes short-creeping, 4 (3-5) mm in diam; rhizome paleae brown, non-
lustrous, ovate to cordate, acute to rounded, basifixed, entire or rarely irregularly
fimbriate toward the apex; fronds 36 (20-55) cm long, fasciculate on the rhizome;
rachis 7.5 (4-14) times as long as the stipe; stipe and rachis red-brown, or black
with lateral brown stripes, with scattered, long, acicular hairs, ctenoid hairs absent
or few and inconspicuous; rachis paleae brown, non-lustrous, acute, basifixed, entire;
bullate-cordate to narrow-ovate and conspicuous, or narrow-ovate to narrow-
obovate, inconspicuous and scattered, blades 31 (20-50) cm long, 4.8 (3.3-7) cm
wide, narrowly oblong, cuneate at the base or subtruncate with the lower segments
strongly deflexed, sometimes parallel to the stipe; segments linear-triangular to

226 ANNALS OF THE MISSOURI BOTANICAL GARDEN

linear, acute, 25 (15-35) mm long, 3 (2-3.5) mm wide, symmetrical at the base
in the upper half of the frond, the lower edge perpendicular to the rachis or deflexed
in the lower half of the frond, ciliate, entire; lamina chartaceous, with scattered,
short, clavate hairs; costa brown, occasionally darker than the rachis, with scattered,
long, acicular hairs, slightly decurrent on, or perpendicular to, the rachis; veins
once-forked, free; guard cells ca 35^ long; sori submarginal to extramedial, round,
with numerous long, clavate paraphyses; sporangia without setae; spores reniform,
smooth, ca 49^ long; n = 37.

Holotype: Costa Rica, Turrialba, 650 m, 5 Aug 1909, Brade & Brade 697 S-PA;
isotype US.

Habitat: epipetric or epiphytic on tree trunks, 1220-2500 m alt in ravines and
mountain forests.

Distribution: southeastern Mexico to Costa Rica.— Fig. 10.

Mexico: hidalgo: Chapulhuacan, Sharp 441768 (TENN, US); nr Jacala, Knobloch
718 (US), veracruz: Cuantlancillo, Copeland 127 (GIT rro de Ejecatepetl,

Santos 2990 (MICH, US); nr El Puerto, above Acultzingo, Sharp 44822 (US); Jalapa, Orcutt
2819 (MO, US), Smith 2219 (GH, MO); Orizaba, Mohr s.n., in 1857 (US), Mohr & Botteri
s.n., in Sept, 1857 (US); Orizaba to Tzhuatlanchilla, Bourgeau 2609 (GH, US), puebla:
El Cerro de Cuhuatepetl Tehuacan, Santos 3678 (MICH, US), morelos: Valle
. Lyonnet 779 (GH). oaxaca: Cuicatlan, Cuyamecalco, Conzatti & Gomer 3499
(US), chiapas: Los Lagos, SE of Comitan, Carlson 1777 (US); Pico de Loro, Matuda 4268
(MO); San Felipe, nr Ciudad las Casas, Carlson 1615 (US); San Pablo, Miinch 121 (US);
Ghiesbreght 262 (GH).

Guatemala: alta verapaz: Santa Cruz, Johnson 980 (US), chimaltenango: Tecpan,
Skutch 572 (US).

Honduras: comayagua: El Achote, above Siguatepeque, Yuncker et al. 6191 (US);
nr El Rincon, Yuncker et al. 6076 (GH, MO, US); Barranco El Socorro, Williams & Wil-
liams 18377 (US), francisco morozan: Cerro de Uyuca, Morton 6914 (US), 6975 (US);
betw La Branza & Las Flores, Molina R. 1107 (US), 1298 (US); betw Pena Blanca & Lo de
Ponce, Williams & Molina R. 17128 (US).

Costa Rica: guanacaste: nr Tilaran, Standley & Valeria 44666, 44713, 44757 (US).
alajuela: Alajuela, Alfaro 6056 (GH, US), san jose: Tablazo, Brade & Brade 696 (NY),
14a (S-PA). cartago: Cervantes & Padayas, Biolley 95 (US); El Muneco, Stork 4708 (US);
Juan Vifias, Reventazon Valley, Cook & Doyle 204 (US), 242 (US); Orosi, Scamman 6117
i 4625 (GH, MO); Peralta, Rowlee & Rowlee 60 (US); Tuis, Tonduz
11309 (US); Turrialba, Scamman 6116 (GH), 6121 (GH, US), 7230 (GH, US), 7765
(GH); Rio Turrialba, Smith 5091 (US).

There are two extreme types. Typical northern material (represented by Cope-
land's P. cyathicola) has either a long-cuneate frond base and a short stipe, or
subtruncate and strongly deflexed segments at the blade base, a black rachis with
lateral brown stripes, and inconspicuous narrow-ovate rachis scales. The typical
southern material represented by P. tablazianum and P. alfredii has a brown rachis
and conspicuous, cordate rachis scales. However, these characters are less consistent
than the entire, dark brown rhizome scales, the linear-triangular chartaceous seg-
ments, and the extra-medial sorus position. The rachis scales throughout the P.
pectinatum-plumula complex show patterns of consistency in some and inconsist-
ency in other species. Polypodium plumula (see Fig. 5) has a clinal variation of
rachis scales. The rachis scales of P. dispersum and P. atrum are very similar
(more similar than the range of variation in P. plumula) but because of other

EVANS — POLYPODIUM 227

e species warrant recognition. In P. alfredii (as here
construed) both the cordate and lanceolate scales can be found on any one frond
except on those plants from Mexico where the broader scales are completely lost.

6. Polypodium ferrugineum Mart. & Gal., Mem. Acad. Bruxelles 15: 36, t. 6, fig. 2,
1842.

Rhizomes short-creeping, 3.5 (2-5) mm in diam; rhizome paleae broad-ovate
to cordate, golden-tan, basifixed, acute, non-clathrate, non-comose, the margins
entire; fronds 20 (8-40) cm long, approximate to fasciculate on the rhizome; rachis
light red-brown, thinly pilose with long, harsh, acicular hairs, and also with clavate
hairs, ctenoid hairs absent; rachis paleae scattered, inconspicuous, linear-ovate to
-obovate, light red-brown, obtuse to acute, entire; blades narrow-ovate, 17 (9-35)
cm long, 4 (2-6.5) cm wide, cuneate to subtruncate at the base; segments 20
(10-32) mm long, 2.5 (1.5-3) mm wide, linear to narrow-ovate, perpendicular
to the rachis except the strongly deflexed ones at the blade base, herbaceous, acute,
the base expanded and symmetrical in the top half of the frond, subsymmetrical
or with the lower edge perpendicular to the rachis in the lower half of the blade,
entire; lamina with scattered clavate hairs, and occasional acicular hairs; costa
decurrent on the rachis, dark red-brown or blackish, with scattered, long, acicular
hairs, without paleae; veins once-forked, free; guard cells ca 40^ long; sori medial,
round, with simple, clavate paraphyses; sporangia with 3 (2-4) capsular setae, 70-
\20ju, long; spores globose reniform, smooth, ca 50^ long; n = 37.

Holotype: Mexico, Oaxaca, forests of Zacatepeque and Juquila, Galeotti 6354
BR not seen, fragment US.

Habitat: epipetric, rarely epiphytic, from canyons, cliffs and river banks, 500-
2100 malt.

Distribution: western Mexico. — Fig. 8.

Mexico: sinaloa: Colomas, Rose 3201 (US); Panuco, Pennell 20010 (US); Paraje del
Zapote, Salazar 393 (US); Sierra Monterey, Gentry 5883 (GH, MO, US), nayarit: Ceboruco
,a Alarjea, N of Yxtlan, Mexia 880 (GH, MO, US);
Santa Maria del Oro, Pennell 19868 (US); Cerro de San Juan, Mexia 690 (GH); E of
Jalcocotan, rd to Tepic, McVaugh 13342 (US). Jalisco: E of Mamantlan, S-SE of Autlan,
Wilbur 1969 (US); Pihuamo, Jones 511 (MO, US), hidalgo: Hidalgo, km 330, Herb.
Copeland 16056 (MICH), morelos: Cuernavaca, Kenoyer 40 (US), Rose & Painter 6870
(GH, US); Teposteco, Lyonnet 2583 (US); Ramanoa Teatzalan, nr Tepoztlan, Seler 4516
(GH); Tepoztlzan, Herb. Copeland 125 (( /alle del Tepeite, Lyonnet 779

(US). Mexico: Temasealtepec, Ypericones, Hinton 4161 (GH). colima: Revillagigedo I,
Socorro I, Barkelew 236 p.p. (US). Guerrero: Mina, Manchon, Hinton 9607 (MICH);
Mazatlan, Correll 14391 (US), oaxaca: Cerro de San Felipe, Conzatti & Gonzalez 334
(GH). chiapas: Ixtape, Munch 12 (US).

This central and southern Mexican species is distinctive in its small size,
light brown rachis and darker costa, long-setose sporangia, light brown rachis
scales, and golden, ovate rhizome scales. Although this species has usually been
identified as P. plumula, its nearest relatives are P. cupreolepis and P. alfredii, and
it combines certain of the characters of each. It has normal appearing spores and
is a diploid, so it is probably not of hybrid origin.

[Vol. 55
228 ANNALS OF THE MISSOURI BOTANICAL GARDEN

7. Polypodium sursumcurrens Copel., Univ. Calif. Publ. Bot. 19: 291, t. 42, 1941.

Rhizomes short-creeping, 4 (3-5) mm in diam; rhizome paleae ovate to broad-
ovate, red-brown at the margins with dark brown center, lustrous, basifixed or
cordate, acute to obtuse, occasionally inconspicuously comose, non-clathrate, entire;
fronds 30 (12-65) cm long, fasciculate on the rhizome; rachis 4 (2.5-6) times as
long as the stipe; stipe and rachis red-brown, thinly pilose with blackish hairs up
to 1 mm long, without ctenoid hairs; rachis paleae absent; blades narrow-ovate,
25 (10-56) cm long, 5.5 (3.5-8.5) cm wide, cuneate at the base; segments 28 (18-
44) mm long, 3 (2-4) mm wide, perpendicular to the rachis, reduced and slightly
deflexed at the blade base with the lowest 2 or 3 pairs reduced to auricles, linear-
triangular, acute, asymmetrical at the base with the lower edge perpendicular to
the rachis throughout the blade, herbaceous, entire to crenulate, the spaces between
the segments 2 to 3 times the width of the segments; lamina with numerous
blackish, simple, clavate hairs up to 175^ long; costa perpendicular to, or slightly
decurrent on, the rachis, with scattered, long, acicular hairs; veins once-forked, free,
guard cells ca 38^ long; sori marginal to submarginal, round, with numerous
twisted paraphyses with a multicellular irregular clavate head, ca 375^ long;
sporangia without setae; spores globose-reniform, smooth, thick-walled, ca 49/t long.

Holotype: Mexico, Veracruz, mountain tops at head of Orizaba Valley, 2400 m,
epiphyte in mossy woods, Copeland, Mexican Ferns 128 MICH; isotypes GH, US.

Habitat: epiphyte in montane forests, 2400-2600 m alt.

Distribution (Fig. 9): Mexico: Veracruz: El Puerto, Sharp 44822 (US), puebla: Honey
Station, Pringle 13475 (MICH) .-Fig. 9.

This species is a narrow endemic in eastern Mexico. I have seen only two
collections in addition to the type. Copeland mentions the distinctive feature of
the blackish hairs of the lamina. However, they are the usual golden color but
are uniformly attacked by a pyrenomycete fungus, which gives them this character-
istic blackish appearance.

The relationships of this species are not obvious, though it is clearly in the P.
plumula group. It is probably closest to P. alfredii, P. ferrugineum, and P. cupreo-
lepis, from which it differs in the larger size, more widely spaced segments, espe-
cially in the larger fronds, the gray-green color, long rachis hairs, dark brown,
ovate rhizome paleae, the lack of rachis paleae, and especially the unique
paraphyses.

8. Polypodium bermudiamim A. M. Evans, sp. nov. — Fig. 17.

P. pectinatum var. squamosum Lindm., Ark. Bot. 1:238, 1903, p.p., as to Bermuda, Herb.

Farlow (GH), not as to lectotype. (See P. dispersum).

Rhizoma breviter repens, paleis anguste ovatis, castaneis, non comosis, incon-
spicue dentatis, basi affixis, apice acuminatis; frondes approximatae articulatae, ca
35 cm longae; stipites et rhachides rigidae nitentes castaneae, sparse pilosae; paleae
rhachidis numerosae parvae, anguste triangulares, castaneae, basi hastatae, apice
acuminatae; lamina pectinata, anguste ovata, ca 25 cm longa et 7 cm lata, basi
abrupte cuneata; segmenta anguste ovata, ca 35 mm longa et 4-5 mm lata, rhachide
apice obtusa, basi symmetrica; lamina solum cum pilos brevibus

EVANS — POLYPODIUM 229

clavatis; venae liberae 1-furcatae; sporangia cum saepe 2-capsularibus setis; sporae
globoso-reniformes, paullo tuberculatae, ca 43/u longae, 64 per sporangium.

Rhizome short-creeping, 5 (4-7) mm in diam; rhizome paleae narrow-ovate,
red-brown, acuminate, non-comose, basifixed, non-clathrate, inconspicuously
toothed; fronds 35 (15-55) cm long, approximate on the rhizome; rachis 3 (2.5-4)
times as long as the stipe; stipe and rachis red-brown, pilose with long, simple,
acicular hairs, ctenoid hairs absent; rachis paleae conspicuous, red-brown, paler
at the base, narrowly triangular, hastate at the base, acuminate, occasionally
toothed; blades narrow-ovate, 25 (14-33) cm long, 6.8 (5.7-8.5) cm wide, abruptly
cuneate at the base; segments 35 (28-44) mm long, 4-5 mm wide, perpendicular
to the rachis, slightly reduced or auriculate at the base of the blade, herbaceous,
obtuse, symmetrical at the base, entire, the spaces between the segments equaling
the width of the segments; lamina essentially glabrous, with short, clavate hairs;
costa decurrent on the rachis, pilose with acicular hairs, with reduced paleae similar
to those of the rachis; veins once-forked, free; guard cells ca 43^ long; sori medial,
round, with simple clavate paraphyses; sporangia mostly with 2 two-celled capsular
setae, 130-175^ long; spores globose-reniform, slightly tuberculate, ca 43^ long.

Holotype: Bermuda, Tuckers Town, on rocks, 10 Febr-9 Mar 1908, Brown
464 US 848332; isotype GH.

Habitat: on limestone rocks in lime-sink areas.

Distribution: Bermuda: Goode s.n., Mar, 1877 (MO), Apr, 1877 (US); Goode s.n.
(US); Reinwardt s.n. (GH). Hamilton island: Walsingham, Harshberger s.n., June 1905
(GH, US); Walsingham Caves, Moore 3118a (GH), Taylor 49-1163 (MICH); Paynter
Vale, Farlow s.n., June 1881 (GH), Britton & Brown 267 (US).

This species can be distinguished from P. plumula by the brown rachis and
the broad segments, and from P. ptilodon var. caespitosum by the once-forked veins
and the rachis scales. It is distinguished from P. dispersum by the number of
spores per sporangium and the brown rachis. Of the sexual species, it is probably
closest to P. atrum, from which it can be distinguished by its brown rachis, once-
forked veins, basal segments not deflexed, and the costa being decurrent on the
rachis. Its close similarity to P. atrum and P. dispersum indicates its affinities to
these two species. The problem of specific and varietal rank in these three species
is complicated by the unique life cycle of P. dispersum, which demands its separa-
tion. I have, therefore, accorded specific rank to this narrowly confined endemic.

9. Polypodium plumula Humb. & Bonpl. ex Willd. in L., Sp. PL, ed. 4, 5: 178,
1810.

P. schkuhrii Raddi, Opusc. Sci. Bol. 3:287, 1819. (as "Schkuhri"). PI. Bras. 1: 19, t. 27,
fig. 2, 1825. (Holotype: Brazil, Corcovado, Raddi FI not seen; isotype B)

P. elasticum L. C. Rich, ex Desv., Mem. Soc. Linn. Paris 6:233, 1827, non P. elasticum
Bory ex Willd., 1810. (Type: P. elasticum L. C. Richard, 1792, nom nud., was
validated by Desvaux by accepting this name and citing P. elegans Poir. and P.
plumula Humb. & Bonpl. ex Willd. as synonyms; the name is thus to be considered
an illegitimate renaming. The name P. elastic tly accepted by Baker

in Hooker & Baker, Syn. Fil., ed. 2, 332, 1874, who cited P. plumula as a synonym
but not P. elegans, thus effectively fixing P. plumula as lectotype. Thus, P. elasticum
is to be considered a superfluous renaming of P. plumula Humb. & Bonpl.)

[Vol. 55
230 ANNALS OF THE MISSOURI BOTANICAL GARDEN

P. pulchrum Mart. & Gal, Mem. Acad. Bruxelles 15: 41, 1842. (Holotype: Mexico, Veracruz,

Jalapa, ad auercos, 4000', Galeotti 6332 BR)
P. pectinatum var. schkuhrii (Raddi) Baker in Hooker & Baker, Syn. Fil., 333, 1867.
P. pulchrum Mart. & Gal.

Rhizome short- to long-creeping, 5 (3-7) mm in diam; rhizome paleae narrow-
triangular, red-brown, lustrous, basifixed, acuminate, non-comose or occasionally
comose, non-clathrate, the margins regularly papillose; fronds 35 (20-60) cm long,
approximate on the rhizome; rachis 4.5 (2-7.5) times as long as the stipe; stipe
and rachis black, with scattered, long, acicular hairs, ctenoid hairs absent; rachis
paleae broadly cordate with short-acuminate apex (in Central America), cordate-
or hastate-acuminate (in Central America and the West Indies), or hastate and
narrowly triangular (South America), basifixed or cordate, bullate or not, golden
to red-brown, the margins toothed or papillate and occasionally fimbriate; blades
narrow- to linear-elliptic, 28 (15-52) cm long, 4.7 (3-7.5) cm wide, cuneate
or occasionally subtruncate at the base; segments 24 (16-40) mm long, 2.5 (2-3)
mm wide, perpendicular to the rachis or slightly ascending, abruptly reduced to
lobes at the blade base, linear, straight, obtuse, symmetrical and expanded at the
base, herbaceous, entire, the spaces between the segments } / 2 to 1 times tne widtn
of the segments; lamina with scattered acicular and clavate hairs; costa decurrent
on the rachis, black, occasionally brownish near the rachis in Central America,
with scattered, long, acicular hairs; costal paleae similar to those of the rachis
but reduced; veins once (or twice) -forked, free; guard cells 34^ long; sori medial,
round, with long, simple, clavate paraphyses; sporangia mostly with 3 or 4 two-
celled capsular setae; spores reniform, slightly tuberculate, ca 50^ long; n=74.

Lectotype: Venezuela, Caracas, Bredemeyer Herb. Willd. no. 19655-1, photo-
graph B, fragm. NY. Syntype: Venezuela, Caripe, Cumana, Humboldt & Bon-
pland 429 Herb. Willd. no. 19655-2, photograph B. The choice of lectotype is
discussed below.

Habitat: epiphytic on tree trunks and branches, occasionally terrestrial or
epipetric on wet swampy or montane forests, from sea level to 2600 m alt.

Distribution: Florida, West Indies, eastern Mexico, south to southern Brazil.—
Fig. 9.

U. S. A.: Florida: brevard co: Indian River, Whitney 1875 (GH). citrus go: Pineola
Noble s.n. (RLAS) ; Pineola grottoes, Evans 2006 (MICH), dade co: Whiskey Creek,
Everglades Ntl Pk, Craighead s.n., Jan 1960 (FLAS); SW of Royal Palm Pk, Cuthbert
& Small s.n., 14 May 1919 (FLAS). hernando co: Brooksville, Jones 24 (US), hills-
borough co- Hillsborough River St Pk, R21E, T27S, Sect 8, Evans 1186 (MICH); 10
mi NE of Tampa, Correll 5868 (GH, US), marion co: Ocala, Smith s.n., Apr 1879 (GH).
monroe co: Aiken, Key Largo, Pollard et al. 203 (US), polk co: 12 mi E of Lake Marion,
vie of Winter Haven, McFarlin 4112 (MICH), seminole co: Oviedo, Eaton 1023 (GH);
Sanford Rapp s.n., Mar 1911 (FLAS). ST. John's co: 14 mi W of St. Augustine, Reynolds
sn in 1877 (US), sumter co: Drake Point, Lake Astachula, Smith s.n., 29 Mar 1879
(US); Indian Field Ledges, Evans 1143 (MICH); Rutland Creek, St. John s.n., 22 Dec
1934 (FLAS); Wonders Ham >1 Mar 1883 (GH, MICH, US), volusia

co: Enterprise, Faxon s.n., Apr 1873 (GH); Ormond Beach, Freeman 59323 (US).

l as: NW of El Progreso, Stanford et al. 985 (GH, MO, US); Gomez

Farias, Palmer 300 (GH, MICH, MO, US); S of Hi- et al. 2055 (US).

anos, Palmer 263 (GH, MO, US); 4 mi W of Pendencia, Graber

233 (US); Tamasopo Canyon, Pennell 17983 (US), Pr ingle 3392 (GH, MO); Tamaz-

EVANS— POLYPODIUM 231

unchale, Lundell 7104 (MICH), Copeland s.n., 27 Dec 1938 (MICH, US); E of Xilitla,
King 4354 (US); Xilitla Road 9 mi, Kenoyer & Crum
3993 (GH, MICH). Veracruz: E of Chavarillo, Weatherwax 181 (MO); Coatepec, Sanchez
S. 245 (US); El Mirador, Purpus 16642 (U<

Seaton s.n., 29 Aug 1891 (GH); Orizaba, Ojo de Agua, Copeland s.n., 4 Febr 1938 (MICH,
US); Potrero Viejo, Copeland 126 (GH, MICH); Teocello Falls, Rhoads s.n., Mar 1899
(US), hidalgo: Jacala, Chase 7426 (MI( • <>er, 15 Nov 1937

(MO), Lyonnet 1299 (US); 41 mi N of Jacala, Barkley 17MOW (MICH); Molango, Moore
1970 (GH). oaxaca: Santiago de Jocotepec, Santos 3378 (MICH, US), chiapas: Mimic.
Pichucalco, Gilly & Hernandez X. 175 (MICH), campeche: Hacienda San Pablo, nr
Champotoh, Collins 50 (US).

Guatemala: el peten: San Clemente to Dos Arroyos, Bartlett 12820 (MICH, US);
Tikal National Pk, Lundell 16748 (US); Uaxactum, Bartlett 12754 (MICH);
to San Clemente, Bartlett 12810 (MICH), quezaltenango: Colahuache, Rojas 549 (US);
San Felipe, Kelb . suchitepequez: Cuyotenango, Rojas 151 (US); Las

Animas, Shannon 276 (US), retalhuleu: San Felipe, Kellerman 5584 (US), santa
rosa: Ojo de Agua, Heyde & Lux 4083 (US).

British Honduras: corozal: Honey Camp, Lundell 537 (GH, MICH, MO, US),
•., 22 July 1930 (US), belize: Maskall, Gentle 1240 (US), el cayo: Cohune
Ridge, Lundell 6460 (MICH); Mountain Pine Ridge, San Agustin, Lundell 6748 (MICH);
Valentin, Lundell 6276 (MICH, US).

Honduras: comayagua: Agua Salada, Williams & Molina R. 11451 (GH); Siguatepe-
que, Standley & Chacon P. 6684 (GH). atlantida: La Fragua, vie of Tela, Ames 7 (US).

El Salvador: ahuachapan: Sierra de Apaneca, Finca Colima, Standley 20182 (US).
sonsonate: Izalco, Standley 22189 (US), san Salvador: Calderon 1274 (US), cuscatlan:
Colina de Santa Tecla, Calderon 1786 (US), san vicente: Volcan de San Vicent
27609 (GH, US).

Costa Rica: alajuela: Alajuela, Scamman 6122 (GH), Alfaro 6056 (US), san jose:
Tablazo, La Uruca, Beyer 1 (US); Hitchcock s.n., 22-24 Oct 1911 (US); Pittier 1231 (US).

Panama: veraguas: Bahia Honda, Taylor 1390 (MICH, US).

Cuba: oriente: Finca Playucla, Ekman 4451 (US); Baracoa, Shafer 3927 (US); Gran
Piedra, Clement 7156 (US); Loma Mensir S); Monte Verde, Shafer 8696

(US); El Cobre, Pollard & Palmer 394 (US); Loma del Gato, Hioram & Clement 6486
(US), Leon, Clement & Roca 9960 (US), Clement 400 (US).

Dominican Republic: Santiago: Vallecito, San Jose de las Matas, Valeur 465 (GH).

Jamaica: Abbey Green, Maxon 10096 (GH, US), Arntully, Orcutt 3105 (MICH, MO);
Chester Vale, Philipson 758 (US); Mt Horeb, Taylor s.n., 27 April 1956; Guys Hill
P. O., Proctor 57/9 (MO); nr Kingston, Lehmann 972

1891; Marshalls Pen, Proctor 22898 (MICH); Silver Hill Gap, Maxon & Killip 1234 (GH,
US).

Puerto Rico: Las Mesas, nr Mayagiiez, Holm 253 (GH, US).

Guadeloupe: Camp Jacob, Duss 4093 (GH); St. Lelande (Grand-Val), Questel 1163
(US).

Surinam: Avanavero Falls, Stahel 4600 (MO); Augustus Falls, Tafelberg, Maguire
24754 (US); Grassi Falls, Saramacca River headwaters, Maguire 24945 (GH, MO, US),
24005 (GH, MO, US).

British Guiana: Kanuku Mts, Takutu River, Smith 3289 (GH, MO, US).

Venezuela: delta amacuro: Lower Orinoco, Santa Catalina, Rusby & Squires 367
(US), miranda: Guatopo Ntl Pk, Steyermark 90029 (US), carabobo: Maracai, Vogl s.n.,
(GH).

Colombia: magdalena: Santa Marta, Campo Alegre, Smith 1026 (GH); La Vuelta
del Tigre, Minca Rd, Bennett 36 (US), santander: Rio Surata valley, betw E Jaboncillo
Killip & Smith 16404 (GH, US), antioquia: Frontino, West Andes of Antioquia,
Lehmann 7408 (US), valle del cauco: Cauca Valley, E of Zarzal, Pennell et al. 8407
(GH, US), meta: Los Llanos, Villavicencio, Cuatrecasas 4690 (US); nr Rio
Pennell 1559 (US); Puerto Lopez, Little 8358 (US); mouth of Rio Atacuarf, i
Black 46-226 (GH).

Peru: san martin: Rio Negro, Woytkowski 6198 (US); Tarapoto, Woytkowski 35134
(MO); Tingo Maria, Allard 20470 (US), loreto: Maynas, Lupuna Cocha, Tryon 5186
(US); Mishuyacu, nr Iquitos, Klug 1382 (US); Santa Rosa, Urubamba Valley, Cook &

[Vol. 55
232 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Gilbert 1727 (US), huanuco: Chinchao to Puerte Durend, Coronado 90 (GH, US);
Yanayacu, flues 2000 (US), junin: Colonia Perene, Killip & Smith 25047 (US), along
Rio Perene, nr "Hacienda 3," Killip & Smith 25169 (US); La Merced, Killip & Smith
23941 (US); Rio Paucartambo Valley nr Perene Bridge, Killip & Smith 25382 (US);
Schunke Hacienda, above San Ramon, Schunke A158 (US) ; nr San Ramon, Coronado 261
(GH); Manto, n US), 6479 (US);

Yunguy, Woytkowski 6598 (US), cusco: Convencion, Herrera 147 (US); Convencion,
Sahuayaco, Vargas C. 1825 (GH); Convencion, Mission nr Quillabamba, Mexia 8090
(GH, MICH, MO, US), Coronado 112 (GH, US), Soukup 162 (GH); Potrero, Tryon
5390 (US); Playa Rosalina Rio Alta Urubamba, Biles 1720 (US).

Bolivia: la paz: Apolo, Williams 1120 (GH, MO, US); La Floride, South Yungas,
Buchtien 483 (US juaya, North Yungas, Buchtien

5004 (US); Polo-Polo, nr Coroico, North Yungas, Buchtien 3507 (GH, MO, US), 3507a
(US); San Antonio de Mapiri, Buchtien 1074 (US); San Bartolome (near Calisaya), South
Yungas, Krukoff 1 S). santa cruz: Buena Vista,

Prov Sara, Steinbach 5410 (GH); Espina, Rusby 136 (US); Motacucito, Sara, Steinbach
2506 (GH).

Brazil: amazonas: Rosariulio, Kuhlmann 218 (MO), ceara: Sitio Uruguaniana, 4 km
W of Guaramiranga, Cutler 8318 (US), bahia: Monte Cruzeiro, Rose 20041 (GH, US);
Rio Grongogy Basin, Curran 282 (GH, US); Toca de Onca, Rose 20076 (US), minas
gerais: Ilhen, Fazenda da Tabunha, Mexia 4969-a (GH); Mariana, Vanthier 591 (GH).
rio de Janeiro: betw Alto da Serra & Meio da Serra, Sr «,) ; Corcovado Mt,

W 149 (GH); Estrada Velha de Petropolis, Smith 6466 (US); Ilha Grande, Rose

20363 (US) ; Itaguai, Rio Mazomba, Brade 20163e (MO) ; Organ Mts, Wagner s.

1902 (MICH); Petropolis, Smith 6466 (GH, MO); Rio de Janeiro, Lindman A159 (US),
Regnell 250 (US), Gaudichaud s.n., (GH).

This species has historically been broadly interpreted and frequently mis-
construed. I have segregated out those entities which are clearly distinct, i.e. P.
bermudianum, P. dispersum, P. cupreolepis, P. ferrugineum and P. alfredii, thus
leaving a fairly uniform species but with some variation still to be more thoroughly
explored. Figure 5 shows outline drawings of representative rachis paleae of
specimens from different geographical areas. The broad cordate type from Central
America appears to be the basic type and from that the narrower paleae of the
West Indian plants and the extremely narrow paleae with large median marginal
fimbriae of the South American plants have arisen as clinal modifications. The
Central American plants with extremely cordate and golden rachis paleae also
needs further study. These plants exhibit costae with brown bases and some brown
streaking on the sides of the rachis. Although they are otherwise like the typical
material, in these characters they show tendencies toward P. alfredii and P.
cupreolepis. Their spores appear normal but no cytological information is known
other than that P. plumula has been counted as a tetraploid in Florida and
Jamaica.

A further question concerns a few collections from Brazil with only 32 spores
in each sporangium. These spores are identical to the spores of the West Indian P.
dispersum; and I have, therefore, included the specimens in that species. However,
their morphology is closer to that of P. plumula than that of the typical West
Indian P. dispersum. As I believe that P. plumula is probably one of the original
parents of P. dispersum, it may be that the Brazilian P. dispersum is of different
origin — possibly a hybrid between the southern P. plumula and P. filicula. There
is, however, little evidence on which to base such an hypothesis and none of this
material is known cytologically.

EVANS — POLYPODIUM 233

Polypodium elegans Poir. (1804), (non Cav. ex Swartz, 1806) has been referred
to P. plumula (e.g. by C. Chr., 1906; de la Sota, 1960). However, the type (St.
Dominique, Nectoux s.n., Herb. Webbianum, ex Herb. Desfontaines (FI) can be
referred to P. otites L. (P. tenuifolium Humb. & Bonpl. ex Willd.). I have included
P. schkuhrii Raddi in synonymy. Though no type was designated, I have seen an
illustration by Raddi (1825) and a Raddi collection (B) which fit his description
(1819) and which I believe to represent an isotype. The type is probably at

The choice of lectotype for P. plumula may be explained as follows. In the
original description in 1810 of P. plumula, Willdenow attributed the name to
Humboldt & Bonpland and cited the localities Cumana and Caracas, both in
Venezuela. I have seen photos of the two syntypes in the Willdenow Herbarium
in Berlin: Sheet 19655-2, collected in Cumana, Caripe, Humboldt 429, and Sheet
19655-1, collected in Caracas by Bredemeyer of which there is also a type fragment
in NY. These two specimens represent different species in my opinion. In 1816,
Humboldt, Bonpland, & Kunth described P. plumula again and a second species
that they called P. molle, which are both represented in Paris by authentic speci-
mens in the Humboldt & Bonpland Herbarium. The specimen called P. plumula
and which agrees with their description corresponds with the Bredemeyer specimen
in Berlin and the one called P. molle agrees with the Humboldt 429 collection in
Berlin. Thus, Humboldt & Bonpland did collect two species both of which are
represented in Paris but only one in Berlin. Since the original P. plumula was
based on a mixture, one element has to be selected as lectotype. It should be
considered that Humboldt, Bonpland, & Kunth did this in 1816 by keeping one
element under the epithet plumula and describing the other as a different species,
P. molle. Thus the Humboldt 429 specimen in Berlin was removed as P. molle
from the concept of P. plumula, and inferentially the latter was left based on the
Bredemeyer specimen, which is thus the lectotype. This specimen is conspecific
with the Humboldt & Bonpland specimen in Paris which very likely originally
provided the name. However, the Paris specimen can hardly be taken as lectotype
since it may never have been seen by Willdenow at the time he drew up the
original description. This Bredemeyer specimen represents a widespread species
that has commonly been identified as P. plumula. The specimen Humboldt 429
at Berlin, the isotype of P. molle H.B.K. (which is a later homonym thrice over),
represents a species that has also not usually been separated from P. plumula but
which is different, as noted elsewhere, and which I am calling P. dispersum.

10. Polypodium jilicula Kaulf., Enum. Fil., 275, 1824.

P. elasticum L. C. Rich. var. filicula (Kaulf.) Baker in Mart., Fl. Bras. 1(2): 517, 1870.

Rhizome short-creeping, 1-2 mm in diam; rhizome paleae narrow-ovate to
narrow-triangular, brown, non-lustrous, basifixed, acute, non-comose, non-clathrate,
finely serrate; fronds 10 (6-17) cm long, fasciculate on the rhizome; rachis 5.4
(3.5-11) times as long as the stipe; stipe and rachis red-brown, with scattered
acicular hairs ca 0.3 mm long; rachis paleae numerous, conspicuous, cordate, bul-

234 ANNALS OF THE MISSOURI BOTANICAL GARDEN

late, acute, dark brown, finely serrate; blades narrow-ovate to narrow-obovate,
9 (5.5-15.5) cm long, 2.5 (1.5-3) cm wide, cuneate at the base; segments 15 (10-
20) mm long, 1.5 (1-2) mm wide, perpendicular to the rachis or slightly ascend-
ing, not deflexed but reduced to lobes at the blade base, linear, herbaceous, obtuse
or rounded, symmetrical and expanded at the base, entire or undulate, the spaces
between the segments ca 1/2 to 1 times the width of the segment; lamina glabrous
or with scattered, short, clavate hairs; costa decurrent on the rachis, with scattered
acicular hairs ca 0.25 mm long, without paleae; veins simple, free; guard cells
ca 41^ long; sorus supra-medial, round, with short, simple, clavate paraphyses;
sporangia without setae; spores globose-reniform, smooth, ca 50/* long.

Type: Brazil, Chamisso s.n. not seen, B(?) or LE(?).

Habitat: epiphytic, occasionally epipetric, in humid forests at 500-2200 m alt.

Distribution: western and central South America from Colombia to southern
Brazil and northern Argentina.— Fig. 9.

Colombia: north de santander: Culaga Valley, nr Tapata, Killip & Smith 20162
(GH, US).

Ecuador: carchi: km no 78, Rio Blanco, Ibana to San Lorenzo, Dodson & Thien
1563 (US).

Peru: amazonas: Bagua, Rio Utcubamba, 40 km S of Bagua Grande, Hutchison 1472
(US), san martin: Tarapoto, Woytkowski 35236 (MO), cusco: Biies 1589 (US); Cusco
to Santa Ana, Herrera 874 (US) ; Convention, Potrero, 8 km W of Quillabamba, Tryon
5373 (GH, US); Urubamba Machupicchu, Vargas C. 3346 (US); Urubamba Valley, Herrera
3297a (US); nr Urubamba River, Heller 2203 (US).

Bolivia: la paz: Guanai to Tipuani, Britton & Rushy 1448 (MICH); Tigre Pata,
1125 (GH, US), cochabamba: Canamina, Rushy 82 (US), santa cruz: El
Fuerte, Jamaipata, Steinbach 8272 (GH); Tres Cruces, Herzog 1538 (GH, US).

Brazil: minas gerais: Serra da Mutuca, nr Vargem de Ouro Podre, Williams & Vicente
6196 (GH, US); Sao Sebastio do Paraiso, Brade 17968 (MO), goias: Jatai, Queixada, Rio
Corrente, Macedo 21554 (MO, US). Rio de Janeiro: Estrada Velha de Petropolis, Smith
6468 (US); Rio de Janeiro & Bahia, Webb s.n., in 1868 (US). Parana: Jaguariaiva, Dusen
15925 (GH, MO, US). Rio grande do sul: Pareci Novo, nr Monte Negro, Beetle 1753
(US); Sao Luis, Latto Pirabo, Jurgens & Stier (Rosenstock 256) (US).

Paraguay: Guarapi, Balansa 2874 (US); Cerros de Tobaty, Hassler 6172 (GH).

Argentina: jujuy: Ledesma, Dinelli 16726 (GH). misiones: Loreto, Moreau s.n.,
26 July 1931 (GH); San Ignacio, Hunziker 764 (MO), Vattuone & Bianchi 169 (US);
Gobernador Roca, Schwarz 6297 (GH); San Ignacio, Cristobal, Ahumada & de la Sota 51
(US); Yabebiry, Moreau 48162 (US).

Although I have not seen the type, there can be little doubt about the ap-
plication of filicula to this very distinctive species. It is easily recognized by its
small size, cuneate blade base, dark cordate rachis scales, unforked veins, and its geo-
graphical isolation from any other closely similar species except P. plumula. It
probably arose initially from the stock of the latter species, but it is presently
quite distinct from it in a number of characters.

This species, like P. dispersum and P. plumula, reproduces readily by root
proliferations. Several herbarium sheets show this character well, and a few sheets
show dense mats of juvenile plants similar to the dense populations of small sterile
plants of P. dispersum. It is presumed that this habit is correlated with dryness and
exposure of the roots through a thin substrate.

EVANS— POLYPODIUM 2

11. Polypodium dispersum A. M. Evans, Amer. Fern Jour. 58: 173, pi. 27, 1968.
P. molle H.B.K., Nov. Gen. Sp. PI. 1:8, 1816, non Schreb., 1771. (Type:

Cumana, Humboldt & Bonpland s.n. P not seen, fragment B)
P. pectinatum var. squamosum Lindm. Ark. Bot. 1:238, 1903. (Lectotype: Brazil, Mato
Grosso, Fazenda Sao Jose, Regnell A2671 S. Syntypes: Jamaica, Herb. Alstroemer
S-PA, and Herb. Casstroem S-PA; Bermuda, Herb. Farlow GH; Brazil, Rio de Janeiro,
Mosen 113 B, S, S-PA)
P. microsorum Lindm., Ark. Bot. 1:239, 1903, p.p. as to Cuba, Wright 1051, not as to
lectotype. (See P. pectinatiforme, which is P. microsorum Lindm., not Mett.)
Rhizome short-creeping, 5 (4-8) mm in diam; rhizome paleae narrow-triangu-
lar, red-brown, lustrous, basifixed, acuminate, slightly comose, non-clathrate, in-
conspicuously toothed; fronds 43 (27-63) cm long, approximate on the rhizome;
rachis 3 (2.5-4) times as long as the stipe; stipe and rachis black, thinly pilose with
acicular hairs, occasionally with ctenoid hairs; rachis paleae conspicuous, narrow-
triangular, hastate, non-bullate, basifixed, dark red-brown with pale base, lustrous,
acuminate, inconspicuously toothed, fimbriate at the base; blades narrow-triangu-
lar, 32 (17-48) cm long, 6.8 (4.5-9) cm wide, subtruncate to abruptly cuneate
at the base; segments 35 (23-47) mm long, 4 (3-6) mm wide, perpendicular to
the rachis, occasionally deflexed at the blade base, reduced (sometimes to mere
lobes) at the blade base, herbaceous, obtuse, symmetrical at the base, entire, the
spaces between the segments approximately equal to the width of the segments;
lamina with few, long, acicular and clavate hairs; costa decurrent on the rachis,
thinly pilose with acicular hairs, with reduced paleae similar to those of the rachis;
veins twice-forked, free; guard cells ca 38^ long; sori medial, round or occasionally
oblong, with a few, simple, clavate, paraphyses; sporangia mostly with 2 (1-4)
two-celled capsular setae; spores globose to ovoid, slightly tuberculate, ca 43^
long, with irregular, incomplete or interrupted variable scar, 32 per sporangium;
n=lll (apogamous); gametophyte strap-shaped, branched, with marginal rhizoids,
often with stomates, but without sex organs, proliferating 1-3 apogamous sporophyte
proliferations.

Holotype: Florida.. Citrus Co, R20E, T21S, Sect 1, Pineola Grottoes, 21 Sept
1963, Evans 2008 MICH; isotypes TENN, US.

Habitat: epipetric (mainly limestone), occasionally terrestrial or epiphytic,
on walls, in reeky hillsides, open woods, relatively dry areas, 100-2000 m alt.

Distribution: Florida, West Indies, eastern Mexico, through western South
America to southern Brazil. — Fig. 9.

U. S. A.: flordia: alachua co: Buzzard's Roost, Gainesville, Weber & West sn

18 Nov 1927 (FLAS), Knoppen s.n., 19 Mar 1928 (US); Devil's Mill Hopper, C

West s.n., 11 Apr 1926 (FLAS). brevard go: Indian , j n 1899 (MO)

citrus co: Pineola, West & Arnold s.n., 13 Nov 1932 (FLAS); Lecanto, St. John 325
(FLAS). hernando co: Annutalaga Hammock, R19E, T21S, Sect 20 & 29, Evans 2012
(MICH); on left bank of Withlacoochee River, 13 mi NE from Brooksville, Smith sn
22 Mar 1883 (US); Brooksville, Garrett s.n., 10 Oct 1953 (FLAS); Istachatta Underwood
287 (GH); nr Nobleton, Knoppen s.n., Mar 1928 (US). Hillsborough co: Hillsborough
River St Pk, R21E, T27S, Sect 8, Evans 1189 (MICH), marion co: Anthony S-ct 1
T14S, R21E, Ward 1871 (FLAS), Blake et al. s.n., 2 Apr 1950 (FLAS); Ocala.
Apr 1879 (GH), Shockley s.n., Mar 1878 (GH). martin go: Sewell's Point, Curtis 5861
(GH, FLAS, MO, US), monroe co: Key Largo, Small 7294 (US); Pumpkin Key, Small
s.n., 9 Mar 1915 (FLAS), R40E, T59S, Sect 12, Evans 1165 (MICH), pasco go- Blandton

236 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Underwood s n (GH). sumter co: Drake's Point, Lake Astachula, Smith s.n., 29 Mar
1879 (GH); Indian Field Ledges, R21E, T20S, Sect 34, Evans 1142 (MICH).

Mexico: tamaulipas: Gomez Farias, Palmer 562 (GH, US), san luis potosi: Tama-
sopo, Pringle 3999 (GH, US), hidalgo: km 350, coll. unknown, 28 Mar 1938 (MICH).
Veracruz: Rio Seco, nr Cordoba, Woronow 3002 (US), oaxaca: Cerro Concordia, Con-
zatti3043 (US).

Guatemala: alta verapaz: Chilaseo, Salvin (GH); Coban, von Tuerckheim s.n.,
July 1885 (GH, US).

British Honduras: el cayo: Camp Six-Vaca Road, Lundell 6545 (GH), Mt Pine
Ridge, Rio On, Lundell 6798 (MICH), toledo: Toledo, Peck 820 (GH).

Honduras: comayagua: Agua Salada, Williams & Molina R. 11451 (US); Siguatepe-
que, Yuncker et al 5729 (GH, MICH, MO, US), francisco morazan: Las Mesea, Standley
28656a (US), Williams & Molina R. 10096 (US), colon: Cuyamel, Carleton 476 (US).
el paraiso: 5 km from Yuscaran, Molina R. 10053 (US).

Cuba: Wright 1051 (B, BR, G, GH, MO, L, NY, S-PA, US), pinar del mo: Arroyo
del Sumidero, Shafer & Leon 13699 (US); Bahia Honda to El Rosario, Shafer 12009 (US);
El Guama, Palmer & Riley 161 (US); Taco-Taco, Baker 3803 (US), las villas: Cien-
fuegos, Combs 349 (GH, MO); Hanabanilla Falls, Trinidad Mts, Britton et al. 4856 (US);
Las Vegas de Mataqua, Buenos Aires, Jack 6524 (MO); Mina Carlota, SE of Cumanayagua,
Sierra de San Juan, Howard 5656 (GH); Sierra Gavilan, above San Bias, Morton 3992
(US), 4142 (US); Trinidad Mts, power plant at San Bias, Howard 5388 (GH, MO), cama-
guay: La Gloria, Shafer 102 (US), oriente: El Gobre, Pollard & Palmer 412 (US);
Arroyo Jimenez, Pico Turquino, Sierra Maestra, Ekman 14794 (US).

Haiti: Bayeus, Loomis s.n., May 1927 (US); Diquini, Miller s.n., Mar 1925 (US);
Furcy Leonard 4757 (GH, US); Hinche, Massif des Cahos, M. Vaillecife, Ekman 6121
(US),' Hinche, Massif du Nord, M. Pinquois, Ekman 6160 (US); Jean Rabel, Leonard
13048 (US); La Vallee, Tortue I, Leonard 11681 (GH, US), 77580 (US); Mission, Fonds
Varettes, Leonard 3646 (US); Petionville, Leonard 4928 (GH, US), Petionville, Massif
de la Selle, Momence River, Ekman 1514 (US); Riviere Soleilhet, Holdridge 2047 (MICH);
St. Michel de l'Atalaye, Mt la Cidre, Leonard 7610 (US), La Lomas, Leonard 7499 (US).

Dominican Republic: azua: Sierra de Ocoa, San Jose de Ocoa, Bejucal, Ekman 11887
(US), barahona: Aceitillar, Sierra de Bahoruco, Jimenez (Marcano 3120) (US), la vega:
Cotuy, Abbott 761 (US); Gajo de Constanza, Jimenez 2111 (US), monte cristo: Arroyo
Asiento Frio, Dist Moncion, Valeur 214 (MO). Santiago: San Jose de las Matas, Jimenez
926 (US), 861 (US), santo domingo: Barrabas, Raunkiaer 72 (US); San Pedro de
Macoris', Rose 4189 (US).

Jamaica: Herb. Alstroemer (S-PA); Herb. Casstroem (S-PA). Castleton, Maxon
824 (US), 758 (US); Flamstead, Port Royal Mts, Maxon 8647 (US); Greenwood, 5 mi
ESE of Little River P. O., Proctor 3931 (US); Kingston, Long Mt, Wilson & Webster
436 (US); Lydford P. O.; mine area, Howard & Proctor 13501 (GH); Mandeville, Maxon
2556 (US), Orcutt 5021 (MO); Montego Bay, nr Salt Spring, Maxon & Killip 1658 (GH);
Rio Bueno, Proctor 16619 (MO); 1 mi S of Rudds Corner, Procior 22896 (MICH); Schwal-
lenburgh, Mt Diablo, Wilson & Webster 489 (US); Silver Hill Gap, Maxon 1144 (US);
St. Andrews, Mt James, Maxon 8602 (US).

Puerto Rico: 10 mi SW of Carolina, Wagner s.n., in 1944 (US); Maricao, Hess 243
(US).

Venezuela: monaras: betw Caripe & San Agustfn, Steyermark 61778 (US), distrito
federal: Caracas, Bailey 464 (US); Chacafto Gorge, Pittier 9479 (US); betw Caracas
& La Guaira, Rose 21727 (US); betw Cotiza & Los Venados, Allart s.n., Oct 1924 (US),
Vogl s.n., (GH).

Colombia: magdalena: Santa Marta, Smith 1026 (MO, US), meta: Puerto Lopez,
Little 8290 (US), norte de santander: Region del Sarare, La Cabuya, Cuatrecasas,
Schultes & Smith 12135 (GH, US).

Ecuador: manabai: hills S of Olmedo, Haught 3492 (GH, US).

Galapagos Islands: Albemarle island: Iguana Cove, Snodgrass & Heller 133 (GH,
US), 14 (GH). charles island: Stewart 967 (US), 968 (US). Chatham island: SW
End, Middle region, Baur 358 (GH); Stewart 966 (US); NW Ansel region, Schimpff 155
(GH, MO, US), indefatigable island: Academy Bay, Stewart 962 (GH, US), Svenson
74 (US); SE side, Stewart 964 (GH, US); Fortuna, Howell 9278 (GH).

EVANS— POLYPODIUM

Bolivia: Porango, Herzog 1496 (S). la paz: Guanai fc, ^
Brazil: cearA: Sitio Urguaniana, 4 km W of Guaramiranga,
MO), minas gerais: Ouro Preto, Macedo 3074 (MO, US), mat* grosse: JJuriti, nr Santa
Mo^n m P p7<£pA fSs). (Regnrf ' ' i776) (S) ' Regne " 7/ 2482 (S) - M ° D * JANEIRO:
This species was noted as distinct by Maxon, and numerous herbarium sheets
at US from the West Indies and Florida are annotated with the epithet "dis-
persum" in his handwriting. Although Maxon never published it, I have adopted
his epithet.

This species has a type of apogamous life cycle thus far unique among ferns,
in which the spores number only 32 per sporangium and are formed by mitosis
rather than meiosis (Evans, 1964). It reproduces commonly by root proliferations,
often forming dense populations of sterile plants in this manner. A further dis-
cussion of the life cycle and reproduction of this species may be found above.

The similarity of this species to P. atrum is striking. They differ in that P.
dispersum has broader rachis scales and the basal segments are only slightly or
not at all deflexed. Of more fundamental importance, however, is that these two
species reproduce very differently. Polypodium atrum has 64 spores per sporangium
and therefore can be assumed to reproduce by the normal sexual process. Poly-
podium dispersum reproduces apogamously and has no sex organs on the gameto-
phyte. This means that there is no possibility for interaction between these two
species. Polypodium dispersum is a triploid and, on morphological grounds, I
suspect it arose as a hybrid between P. atrum and P. plumula. Polypodium plumula
is a tetraploid, so I predict that P. atrum is a diploid. As P. atrum occurs only
in Mexico and northern Central America, P. dispersum perhaps arose there and
then migrated to the other parts of its range.

The relationships between P. dispersum and P. bermudianum are discussed
under the latter species. See also the discussion of P. plumula for comments on
the typification of P. molle H.B.K.

12. Polypodium atrum A. M. Evans, sp. nov.— Fig. 18.

Rhizoma breviter repens, paleis lineari-triangularibus, basi affixis, castaneis,
comosis, integris; frondes approximatae articulatae, ca 40 cm longae; stipites et
rhachides rigidae nitentes nigrae, pilos dissitos aciculares gerentes; paleae rhachidis
numerosae parvae anguste triangulares vel lineari-triangulares, castaneae integrae
basi saepe hastatae; lamina pectinata, anguste oblonga, ca 33 cm longa et 8 cm
lata, basi subtruncata vel interdum abrupte cuneata; segmenta anguste ovata, ca
40 mm longa et 4 mm lata, apice obtusa, basalia paullo reducta et valde deflexa-
lamina pilos dissitos aciculares ferens; venae liberae 2-furcatae; sporangia saepe
cum 2-4 capsularibus paraphysibus; sporae globoso-reniformes, tuberculatae ca
34^ longae, 64 per sporangium. P. disperso simile, segmentis basalibus deflexis,
paleis rhachidis angustioribus, et numero sporae differt.

Rhizome short-creeping, 4-6 mm in diam; rhizome paleae linear-triangular
expanded at the base, red-brown with lighter base, basifixed, lustrous, comose,'
entire; fronds 42 (30-65) cm long, approximate on the rhizome; rachis 3.5 (2 5-5)'
times as long as the stipe; stipe and rachis black, with scattered, long, simple

[Vol. 55
238 ANNALS OF THE MISSOUKI BOTANICAL GARDEN

acicular hairs; rachis paleae small, narrow- to linear-triangular, often hastate
at the base, red-brown, entire; blades narrow-oblong, 33 (21-50) cm long, 8 (6-
11.5) cm wide, herbaceous, subtruncate or occasionally abruptly cuneate at the
base; segments 42 (32-60) mm long, 4 (3-7) mm wide, obtuse, symmetrical at
the base in the upper half of the frond, with the lower edge perpendicular to the
rachis in the lower half of the frond, entire, the basal segments slightly reduced
and strongly deflexed, or occasionally abruptly reduced to auricles, the spaces be-
tween the segments equaling or exceeding the width of the segments; lamina
with scattered acicular hairs and short, simple or occasionally forked clavate hairs;
costa perpendicular to the rachis, black throughout or dark red-brown at the base,
with scattered, long, acicular hairs; veins twice-forked, all free; guard cells ca
35^ long; sori medial, round, with long, simple, clavate paraphyses; sporangia
mostly with 3 or 4 two-celled capsular setae 35-95^a long; spores globose-reniform,
tuberculate, ca 34^ long.

Holotype: British Honduras, El Cayo District, Mountain Pine Ridge, San
Augustin, on boulders on bank of Rio Frio, July-Aug 1936, Lundell 6639 US
1638286; isotypes GH, MICH.

Habitat: terrestrial or epipetric, woods, cliffs, river banks, 60-900 m alt.

Distribution: southeastern Mexico to British Honduras.— Fig. 9.

Mexico: veracruz: Atayac River, Copeland 16058 (MICH); Cordoba, La Luz, Kerher
97 (US); Cordoba, Metlac River, Copeland 124 (MICH, US); C6rdoba, Mt Orizaba, Seaton
424 (GH US) ; Cordoba, San Alejo Mts, Couch M26 (US) ; Cordoba, Monte de San Pablo,
Woronow 2996 (US); Valle de Cordoba, Bourgeau 1432 (GH); Playa Azul, nr Catemaco,
Stoutamire 3563 (TENN); Potrero Viejo, Copeland 123, 126 (MICH); San Andres Tuxtla,
Laguna Encantada, Dressier & Jones 77 (GH, MO, US); Zacualpan, Purpus 2165 (GH,
MO US) oaxaca: Tuxtepec, Chiltepec, Martinez-Calderon 693 (MICH, US); Mogone,
Orcutt 5211 (MO), tabasco: Balancan, San Isidro, Matuda 3372 (MICH, US), chiapas:
San Fernando de Tuxtla, Collins & Doyle 167 (US).

Guatemala: el peten: Chicbul, La Libertad, Lundell 3378 (US), 3378-A (MICH),
La Libertad, Lundell 2980 (MI" '! 1 i :VuV i 1 l.s •; Tikal Ntl Pk, Dos Aguadas, Lundell 15588
(US), Remate rd S of Tikal, Lundell 15878 (US). IZABEL: Cacao, betw Panzos &
Senahu, Barber 197 (US).

British Honduras: el cayo: Camp Six-Vaca Rd, Lundell 6545 (MICH, US); Cohune
Ridge, Lundell 6494 (MICH, US); San Antonio, Bartlett 13062 (MICH, US).

Honduras: santa Barbara: San Pedro Sula, Thieme 5692 (GH, US), comayagua:
Tiquitapa River, Los Dragos, Seguatepequi area, Steeves & Ray 435 (GH, US); Jutiapa,
Saguatepaqui area, Steeves & Ray 414 (GH). yoro: Aguan River valley, Coyoles, Los
Flores Yuncker et al 8150 (MO, MICH); Rio Pelo, Cordillera de Mico Quemada, Ames
25 (US), atlantida: La Ceiba, nr Cangrajal River, Yuncker et al. 8792 (MO, MICH, US).

The relationships of this species are discussed under P. dispersum and P. bermudianum.

13. Polypodium absidatum A. M. Evans, sp. nov.— Fig. 20.

Rhizoma longe repens, paleis anguste triangularibus, basi dilatatis, atro rubro-
brunneis basi pallidoribus, acuminatis, comosis, integris vel inconspicue fimbriatis;
frondes approximatae, stipitibus et rhachibus atro rubro-brunneis pilos aureis
acicularibus praeditis; paleae rhachis inconspicuae filiformes integrae; laminae
anguste ovatae, basi anguste cuneatae; segmentis ascendentibus acutis integris vel
crenulatis, basi asymmetricis, infimis ad auriculas reductis; laminae subtus pilis
dissitis acicularibus et pilis numerosis longis clavatis praeditae; costae valde decur-

EVANS — POLYPODIUM 239

rentes; venae unifurcatae; sori mediales, paraphysibus simplicibus clavatis praediti;
sporangia plerumque setifera.

Rhizome long-creeping, 5 (3-7) mm in diam; rhizome paleae narrow-tri-
angular, expanded at the base, dark red-brown, lighter at the base, lustrous, basi-
fixed, acuminate, comose, non-clathrate, entire or inconspicuously fimbriate; fronds
38 (15-57) cm long, approximate on the rhizome; rachis 3.5 (2-5) times as long
as the stipe; stipe and rachis dark red-brown, with golden, acicular hairs ca 0.5 mm
long, ctenoid hairs present; rachis paleae inconspicuous, filiform, entire; blades
narrow-ovate, 30 (12-46) cm long, 6.5 (3.5-9) cm wide,
base; segments 33 (20-50) mm long, 3.5 (2-4.5) mm wide, i
from the horizontal, reduced to auricles at the blade base, linear-triangular,
coriaceous to herbaceous, acute, asymmetrical at the base, entire to crenulate;
lamina with scattered acicular hairs ca 0.3 mm long, and with numerous, long,
clavate hairs; costa strongly decurrent, with scattered acicular hairs ca 0.5 mm
long; veins once-forked, free; guard cells ca 66fi long; sori medial, round, with
simple, clavate paraphyses ca 0.25 mm long; sporangia mostly with 2 capsular setae
up to 0.25 mm long; spores reniform, tuberculate, ca 59^ long.

Holotype: Colombia, Santander, Paramo de Romeral, Killip & Smith 18518
US 1353919; isotype GH.

Habitat: montane forest, epiphytic, occasionally epipetric or terrestrial, 2800-
4100 m alt.

Distribution: greater Antilles and western South America south to Bolivia. —
Fig. 11.

Cuba: oriente: Gran Piedra, Clement 7155 (US).

Dominican Republic: azua: top of La Pelona, Ekman 13643 (US), la vega: Sabana
Aha, Ekman 13794 (US).

Jamaica: Hart 45 (US), 771 (MO).
Venezuela: merida: Laguna Negra, Gines 1748 (US).

Colombia: magdalena: Paramos, Sierra Nevada de Santa Marta, Seifriz 463 (US).
santander: Paramo de Romeral, Killip & Smith 18518 (GH, US) ; Paramo de Santurban,
Vetas, Killip & Smith 17925 (GH). boyaca: Valle de las Playas, Grubb & Guymer P40
(US); Valle del Corallitos, Grubb & Guymer P103 (US), valle: Paramo de Bavaya, Rio
Bugalagrande, Corrales, Cuatrecasas 20551 (US).

Ecuador: ca: opos of Volcan de Chile, Wiggins 10605 (US), cusco:

Quito, Holdridge 1579 (US), pinchincha: Mt Pichincha, Holmgren 289 (Gil, U
15 (US), Mille s.n. (MO); Sodiro s.n., in 1905 (US), napo-pastaza: Antisanilla, Anthony
& Tate 318, 319 (US); Mt Atazco, Mille 899 (US).

Peru: la libertad: Otuzco, Agallpampa, Lopez M. 1024 (US).
Bolivia: tarija: Padcaya, Fiebrig 2875 (GH).

Although often identified as "P. curvans" or "P. curvatum" by various authors,
the types of these taxa do not agree with this species. The affinities of P. absidatum
are discussed below under P. curvans Mett.

14. Polypodium curvans Mett., Ann. Sci. Nat. (Paris) V, 2: 253, 1864.

P. curvatum sensu Mett., Abhandl. Senckenb. Naturf. Ges. Frankfort 2(1): 58, 1856, non

Swartz, 1801.
P. circinatum Sodiro, Crypt. Vase. Quit. 333, 1893. (Type: Ecuador, Azuay, nr Cuenca,

Rimbach 35 Q?)

Rhizome short-creeping, 3 (2-5) mm in diam; rhizome paleae narrow-tri-
angular, dark red-brown with a lighter base, lustrous, basifixed, acute to acuminate,

240 ANNALS OF THE MISSOURI BOTANICAL GARDEN

comose, non-clathrate, entire or inconspicuously fimbriate; fronds 43 (18-90) cm
long, approximate to fasciculate on the rhizome; rachis 8 (344) times as long as
the stipe; stipe and rachis dark red-brown, with scattered, golden acicular hairs
up to 1 mm long, and with scattered, short, clavate hairs, ctenoid hairs inconspicu-
ous or absent; rachis paleae inconspicuous, filiform, dark red-brown, entire; blades
linear, 38 (15-80) cm long, 4.5 (2.5-10) cm wide, linear-cuneate at the base,
often circinate at the apex at maturity; segments linear, 30 (13-53) mm long, 2
(1.5-3) mm wide, strongly ascending ca 45° above the horizontal, herbaceous,
acute, asymmetrical at the base, confluent, widely-spaced and gradually reduced
to mere auricles at the blade base, crenulate to crenate; lamina with numerous,
long, clavate hairs up to 200^ long, without acicular hairs; costa decurrent on the
rachis, with scattered, acicular, golden hairs ca 0.75 mm long; veins once-forked,
free; guard cells ca 43fi long; sori medial, round, occupying most of the lamina,
with simple, clavate paraphyses ca 200/z long; sporangia with 1 or 2 capsular setae
up to 200jU long, the sporangial stalk with 2 columns of cells; spores reniform,
tuberculate, ca 44^ long.

Type: based on P. cwvatum sensu Mett. Mettenius' description was based on
a Lechler specimen from Peru. The locality and number are given in Mett. Fil.
Lechl. 1: 7, 1856, as Agapata, Peru, Lechler 2006, which is thus the holotype (B?;
isotype L, Morton photo 1859).

Habitat: in montane forests, epiphytic or epipetric, 1600-4000 m alt.

Distribution: Ecuador to western Bolivia.— Fig. 11.

Ecuador: Quitensian Andes, Couthouy s.n. in 1855 (GH).
s n in 1848 (GH) chimborazo: W slope of E Cordillera of Riobamba, Rimhach 27 (GH,
US), napo-pastaza: Antisanilla, Anthony & Tate 322 (US); Cubillan, W slope, Rimhach

12 ^eru: Agapata, Hohenacker 2006 (GH); Chasqui, MacBride & Featherstone 27 July-

13 Aug, 1922 (US); Mayuyoc, Biles 1027 (US), junin: Huancayo, Killip & Si

(GH, US), 23364 (US); Quebrado de Occopilla, Soukup 3642 (GH, US) apurimac: Prov
Abancay, Ampay, Stork et al. 10626 (MO); Bosques de Ampay, Vargas 364 (GH, MO, US);
sTcllaccasa Pass, Abancay-Cuzco trail, West 3820 (GH, MICH, MO), cusc
al Ampay (Apurimac), Santander & Vargas, Aug 1937 (GH); Prov Conversion, Santa
Ana, Hacienda Potrero, Herrera 877 (US); Prov Paucartambo, Valle del Paucartambo,
Hacienda Churu, Herrera 271 (GH, US), 1107, 1605, 1655 (US); Prov Urubamba, Chupani
Vargas C. 11130 (GH). puno: Carabaya, Ollachea to Puente Ackopampa, Vargas C. 3346

Bolivia: la paz: Pelechuco, Williams 2583 (GH, US).

Polypodium curvans is similar to P. absidatum, from which the former differs
by having much smaller and more delicate fronds, strongly circinate at maturity,
and longer hairs on the rachis and sporangia. The spore and guard cell sizes of
P. curvans suggest that it may be a diploid, whereas the spores and guard cells of
P. absidatum (see Table 3) are considerably larger and in the polyploid range.
Polypodium curvans, P. absidatum, and P. choquetangense appear to be closely
related species.

In superficial appearance, P. curvans strongly suggests ctenopteroid affinities.
However, this is not confirmed in any of the critical criteria of sporangial stalks
or spore shape, and it is probably placed in the P. pectinatum complex.

241

15. Polypodium choquetangense Rosenst., Meded. Rijks Herb. Leiden 19: 18, 1913.

Rhizome long-creeping, 4-5 mm in diam; rhizome scales narrow-triangular,
red-brown, lustrous, basifixed, acuminate, comose, non-clathrate, entire; fronds 55
(36-90) cm long, approximate on the rhizome; rachis 7 (3-10) times as long as
the stipe; stipe and rachis dark red-brown, with very long, scattered, simple acicular
hairs, and with long, irregular, ctenoid, clavate hairs, without paleae; blades
narrow-ovate, cuneate at the base; segments linear, strongly ascending ca 25°-50°
above the horizontal, reduced to auricles at the blade base, herbaceous, acute, with
constricted attachments to the rachis, pinnatifid to a narrow wing along the costa;
lamina essentially glabrous, with scattered clavate hairs; costa decurrent on the
rachis, with scattered, long, acicular hairs; veins once-forked, free; guard cells ca
50^ long; sori inframedial, round, with simple clavate paraphyses; sporangia with-
out setae; spores reniform, tuberculate, ca 42ft long.

Type: Bolivia, Hab. Choquetanga grande in silvis montanis ad arborum
truncos, 3300 m alt, Oct 1911, Herzog 2387 L, S.

Habitat: wet rocks and tree trunks, montane forest, 2700-3500 m alt.

Distribution: known only from the province of Cochabamba, Bolivia (Fig. 11).

Bolivia: cochabamba: Ayopaya, Sailapata, Cardenas 3085a (US), 3167 (US); Choro,
Brooke 6090 (G, S); Jatun Pino to Carrasco, Cardenas 5939 (US).

This species is unique because of its deeply pinnate-pinnatifid blades. It is
the only pinnate member of the pectinatum complex. The base of the frond
matures and sheds spores while the apical portion is still juvenile and unfurling,
showing an almost indeterminate growth unknown elsewhere in the group. The
very sinuous, almost circinate rachis, the strongly ascending segments, and the
very long hairs of the stipe and rachis are found only in this species and P. curvans,
which is clearly its nearest relative.

16. Polypodium bolivianum Rosenst., Repert. Sp. Nov. 5: 236, 1908.

P. carpinterae Rosenst., Repert. Sp. Nov. 22 : 16, 1925. (Holotype: Costa Rica, La Carpin-

tera, 1850 m 10 Apr 1908, Brade & Brade 14, p.p., S-PA)

Rhizome long-creeping, 8 (6-10) mm in diam; rhizome paleae narrow- to
linear-triangular, expanded at the base, dark red-brown with pale base, lustrous,
basifixed, acuminate, comose, with hairs obscuring most of the paleae, densely and
conspicuously fimbriate; fronds 75 (35-135) cm long, spaced ca 1 cm apart to
approximate on the rhizome; rachis 3 (2-5) times as long as the stipe; stipe and
rachis dark red-brown, with numerous, conspicuous, ctenoid hairs, occasionally with
short, acicular hairs; rachis paleae inconspicuous, small, filiform, with fimbriate
margins; blades narrowly triangular, 55 (28-107) cm long, 15 (10-27) cm wide,
truncate or rarely the lower segments slightly shortened; segments 80 (45-145) mm
long, 6 (4-9) mm wide, perpendicular to the rachis or the lower segments occasion-
ally deflexed, coriaceous, obtuse to acute, entire, symmetrical at the base, often
slightly constricted near the base; lamina with few, short clavate hairs ca 125^
long; costae perpendicular to the rachis, with a few clavate and ctenoid hairs be-
low, without paleae; veins 2 or 3 forked, free or occasionally partially anastomosing,
obscure; guard cells ca 43^ long; sori supramedial, round, with a few clavate

242 ANNALS OF THE MISSOURI BOTANICAL GARDEN

paraphyses ca 150^ long, and with a cluster of setiform hairs around the receptacle,
these obscured in the mature sorus; sporangia without setae; spores reniform,
tuberculate, ca 43^ long; n = 37.

Holotype: Bolivia, South Yungas, Sirupaya, nr Yanacachi, ca 16° lat, 2300 m,
humid forests, 29 Nov 1906, Buehtien 481 S-PA; isotype US.

Habitat: in montane forests, epiphytic on tree bases and trunks, occasionally
terrestrial or epipetric on mossy rocks, (200) 1000-2700 m alt.

Distribution: southern Mexico, through western South America to western
Bolivia.-Fig. 8.

Mexico: chiapas: Ghiesbreght 394 (GH).

Guatemala: alta verapaz: Finca Sepacuite, Cook & Griggs 462, 555 (US) ; Pansamala,
von Tuerckheim 644 (US).

Honduras: comayagua: Siguatepeque, Standley 56350 (US); Fl Achote, hills above
Siguatepeque, Yuncker et al. 5973 (US) ; Barranco de Trincheras, nr Siguatepeque, Morton
7582 (US).

Costa Rica: alajuela: Palmira, Zarcero region, Smith 4332 (MO), heredia: Vara
Blanca, betw Poas & Barba volcanoes, Maxon 8329 (US), san jose: Rio del Convento,
valle de Diquis, Pittier 12112 (US); San Ramon, Tonduz 17604 (US), Brenes 14221 (US).
cartago: Pejivalle, Standley & Valerio 47053 (US); Santa Clara de Cartago, Maxon 8208
(US).

Panama: chiriqui: Casita Alta, Volcan de Chiriqui, Woodson et al. 887 (MO, US);
Cerro de la Horqueta, Maxon 5405, 5409, 5423 (US); Cerro de Lino, above El Boquete,
Maxon 5216, 5216a (US); El Boquete, Cornman 839, 1276, 1327 (US), Maxon 5248, 5249
(US), darien: Cana, Williams 847 (US).

Venezuela: aragua: Colonia Tovar, Pittier 9358 (GH, US).

Colombia: magdelena: Santa Marta, Sierra de Onaco, Smith 1027 (GH, MO, US);
Mt San Lorenzo, Santa Man, I. US), santander: Cerro Armas, H aught

1958 (US); Las Vegas, Killip & Smith 21153 (US); Mt Pefia Blanca, Charta, Killip &
Smith 19284 (CI- ti Smith 16569, 19284 (US).

cundinamarca: Macizo de Bogota, Quebrada Chica, Schultes 18579 (US), Cuatrecasas
5242 (US), Black 46-424 (GH); W of Salto de Tequendama, nr Ermitano, Uribe 3352,
3354 (US), antioquia: Boqueron, betw Medellin & Palmitas, Hodge 6600 (GH). caldas:
Rio Boquia, Salento, Killip & Hazen 8843 (GH, US); Rio Santa Rita, Salento, Killip &
Hazen 8995 (US), valle del cauca: Pavas, Killip 11660 (US); Rio Cali, Pichinde, Alto
de las Brisas, Cuairecasas 18280 (US).

Ecuador: imbabura: Rio Chaluayaco, below Magnolia, lower Intag Valley, Drew
E-604 (US), leon: Hacienda Solento, nr Santa Rosa, Canton, Pajii, Mexia 6684, 6684a
(US), pichincha: Mt Pichincha, Mille 16 (MO, US), galapagos islands: Albemarle I,
Stewart 963 (GH, US).

Peru: junin: Chanchosmayo valley, Schunke 118 (US); Huacapistana, Killip & Smith
24146 (US), Coronado 272 (GH); Oxapampa, Soukup 2347 (GH); E of Quimiri Bridge, nr
La Merced, Killip & Smith 23958 (US); Vitoc, Soukup 4432 (US), cusco: San Miguel,
Urubamba Valley, Cook & Gilbert 1752, 1762 (US).

Bolivia: Bang 2228 (US); Bang s.n. (MO), la paz: Mapiri, Rusby 356 (US); Polo-
Polo, Coroico, N Yungas, Buehtien 3510 (GH, US); Hacienda Simaco at Tipuani, Buehtien
5269 (GH, US); Yungas, Rusby 357 (US), santa cruz: Yungas de San Mateo, Comarapa,
Steinbach 8430 (GH).

This very large fern with coriaceous, almost waxy, fronds, strongly truncate
blade bases, and densely fimbriate rhizome scales is one of the most distinctive
members of the P. pectinatum group. Its affinities are not clear, although the
generally glabrous lamina and the obscure tuft of setose hairs around the sorus
suggest a relationship to P. ptihdon. In northern South America the rhizome scales
are less densely fimbriate, and it becomes more difficult to separate the species from

EVANS — POLYPODIUM 243

P. eurybasis, which generally has a more pubescent lamina, subtruncate blade with
more acute and crenulate segments, and not so harsh a texture.

I have included here two collections from the Galapagos Islands that are sub-
truncate and have a few ctenoid hairs and a few long hairs on the brown rachis.
These may represent an endemic species, but because the specimens are juvenile,
I am tentatively placing them here where they fit most closely.

17. Polypodium eurybasis C. Chr., Svensk. Vet. Akad. Handl. Ill, 16(2): 71, t. 16,
fig. 12, 13, 1937.

Rhizome long-creeping, 5-9 mm in diam; rhizome paleae narrow-triangular,
light to dark red-brown, lustrous, basifixed, acute to acuminate, comose, non-
clathrate, entire to inconspicuously fimbriate; fronds 28-130 cm long, approximate
on the rhizome; rachis 3 (2-5) times as long as the stipe; stipe and rachis red-
brown to dark red-brown, subglabrous to densely villose with acicular hairs, ctenoid
hairs also present; rachis paleae inconspicuous, filiform or linear, entire; blades
narrow-triangular to narrow-ovate, 20-88 cm long, 4-26 cm wide, subtruncate to
abruptly cuneate at the base; segments 22-135 mm long, 3-9 mm wide, linear-
triangular, narrow- triangular at the blade base, perpendicular to the rachis,
straight or slightly falcate, herbaceous to subcoriaceous, acute, symmetrical at the
base or with the lower edge approaching a right angle with the rachis, crenulate
to crenate; lamina essentially glabrous, with simple and forked clavate hairs up to
200^ long, and occasionally with scattered acicular hairs; costa perpendicular to,
or slightly decurrent on, the rachis, with a few reduced ctenoid hairs and with
scattered acicular hairs up to 0.5 mm long; veins once- or twice-forked, free; guard
cells 44-50^ long; sori medial, round, with simple or branched, clavate paraphyses;
sporangia without setae; spores reniform, tuberculate, 40-44^ long.

This is a widespread and variable species distinguished primarily by its
subtruncate blades, sharply acute and crenate segments, prominent veins, and
glabrous lamina. It is probably most closely related to P. bolivianum, from which
some of the northern South American material is difficult to distinguish. It can be
subdivided into three varieties.

1. Rachis densely villose; rhizome paleae entire 17c. P. eurybasis var. villosum

1. Rachis subglabrous to thinly pilose; rhizome paleae entire to inconspicuously
fimbriate.
2. Veins once (twice) -forked; segments straight, crenate or crenulate; :

paleae red-brown, inconspicuously fimbriate 17b. P.

2. Veins twice-forked, segments straight or drooping at the apex,

rhizome paleae dark red-brown, entire to inconspicuously fimbriate!
17a. P. eurybasis var. eurybasis

17a. Polypodium eurybasis var. eurybasis.

Rhizome paleae dark red-brown, attenuate, fronds 74 (30-114) cm long; stipa
and rachis dark red-brown, with scattered acicular hairs 0.5 to 0.75 mm long, and
also with conspicuous ctenoid hairs; blades narrow-ovate, 55 (21-87) cm long, 11
(4.5-18) cm wide; segments 58 (22-95) mm long, 6 (3-8) mm wide, straight or
slightly drooping near the apex, herbaceous, symmetrical and expanded at the base,
crenulate; veins twice-forked; guard cells ca 50^ long; spores ca 40^ long.

244 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Holotype: Haiti, Massif de la Selle, Marigot, Jardins Bois-Pin, ca 1900 m, 9
June 1928, Ekman H10062 S; isotypes B, US.

Habitat: terrestrial, epipetric or epiphytic on trunks or branches, in montane
forests, 1000-2600 m alt.

Distribution: Cuba, Haiti, Venezuela.— Fig. 11.

Cuba: oriente: Pico Turquino, Ekman 5455 (US), Loma Regino, Ekman 14605 (US).

Haiti: Massif de la Selle, Croix-des-Bouquets, Ekman 3088 (US); Mission, Fonds
Varettes, Leonard 3990 (GH, US).

Venezuela: monagas: Cerro de Turumiquire, Tate 120 (US); Gerro Negro, above
La Sabana de Las Piedras, NW of Caripe, Steyermark 62080 (MO, US), distrito federal:
El Junquito, Killip & Rohl 37168, 37190 (US), aragua: Colonia Tovar, Fendler 221 B (GH,
MO). Bolivar: Ptari-tepui, Steyermark 59595 (US); Mt Roraima, im Thurn 104 (US),
Steyermark 58741 (US).

See P. eurybasis var. glabrescens for discussion.

17b. Polypodium eurybasis var. glabrescens (Rosenst.) A. M. Evans, comb. nov.
P. lachniferum var. glabrescens Rosenst., Repert. Sp. Nov. 11:57, 1912. (Holotype. Bolivia,

N Yungas, Unduavi, 3400 m, Buchtien 2770 S-PA; isotype US)
P. lachniferum var. glabrescens f. incurvatum Rosenst., 1 i >da"). (Holotype:

Bolivia, N Yungas, Unduavi, 3400 m, Buchtien 2769 S-PA; isotype US)

Rhizome paleae usually not expanded at the base, light red-brown, basifixed,
inconspicuously fimbriate; fronds 80 (28-120) cm long, rachis 2.5 (2-3.5) times as
long as the stipe; stipe and rachis red-brown, thinly pilose to subglabrous with
short acicular hairs, ctenoid hairs present; blades narrow-triangular to narrow-
ovate, 56 (20-88) cm long, 12 (4-26) cm wide, subtruncate; segments 65 (22-135)
mm long, 6 (3-9) mm wide; lamina essentially glabrous, with scattered, simple,
clavate hairs ca 0.25 mm long; costae perpendicular to the rachis, with scattered
acicular hairs up to 0.35 mm long; veins once (twice) -forked, prominent; guard
cells ca 44/* long; sori large, covering one-third to one-fourth of the width of the
lamina, with simple, clavate paraphyses; spores ca 44/u long.

Habitat: pendent epiphyte in wet, shady, montane forests, 1500-3600 m alt.

Distribution: Costa Rica to western Bolivia.— Fig. 11.

Costa Rica: alajuela: Zarcero, Smith H 127 (MO). Heredia: Cerros de Zurquf, NE
of San Isidro, Standley & Valeria 50346 (US), san jose: Laguna de la Chonta, NE of
Santa Maria de Dota, Standley 42128 (US); Las Nubes, Standley 38846 (US).

Venezuela: aragua: Colonia Tovar, Fendler 221 (US), merida: La Venta, ]ahn 851
(GH, US), monac .miquire, Tate 148 (US).

Colombia: M S). santander: California, Killip & Smith 16902 (GH,

US); La Baja, Killip & Smith 18135 (GH, US), cundinamarca: Bogota, Ariste-Joseph s.n.
(US); Guadalupe, Bogota, Haught 5040 (US); Macizo de Bogota, Quebrada de las Delicias,
Cuatrecasas 5463, 5608, 5675 (US) ; Bogota, Rio San Francisco Valley, Montserrate, Hawkes
& Garcia-Barriga 94 (US), Ewan 16901 (US); Bogota Plateau, San Cristobal, Niemeyer
175 (US); Bogota, old San Cristobal to Ubaque rd, Schiefer 322 (GH); Sabana de Bogota,
Pring 133 (MO); Mun Chipaque, Gutierrez V. 50 (GH). caldas: Magana, Old Quindio
p & Hazen 9431 (US); Paramo de Buena Vista, Pittier 1147 (US); Cabeceras
del Rio Palo, Quebrada de Santo Domingo, Cuatrecasas 19265 (GH, US), narino: Cordoba,
Ewan 16235 (US); headwaters of Rio Guapuscal, Ewan 16553 (US); San Jose, below La
Victoria, Ewan 16218 (US); Soledad, Ewan 16510 (US).

Ecuador: pichincha: Mt Pichincha, Mille s.n. (GH). napo-pastaza: Pifo, Mille s.n.,
in 1898 (US).

Bolivia: la paz: Unduavi, N Yungas, Buchtien 2771, 2772 (US); Sirupayo, nr

EVANS — POLYPODIUM

Yanacachi, S Yungas, Buchtien 485 (US).

Steinbach 9235 (GH, MO); Rio Tocorani, Herzog 2275 (US); Prov Sacaba, Steinbach 5868

(MO).

Of the three varieties of P. eurybasis, this one appears to be the most general-
ized element. It is distinguished from var. eurybasis by a more glabrous lamina,
lighter green foliage, paler and broader rhizome scales, usually more crenate seg-
ments, and usually only once-forked veins. It is closer to var. villosum, as discussed

17c. Polypodium eurybasis var. villosum A. M. Evans, var. nov.— Fig. 19.

P. eurybasi var. glabrescente simile, paleis rhizomatis semper integris et
rhachidibus dense villosis differt.

Similar to var. glabrescens except rhizome scales with entire margins; stipe
and rachis densely villose with acicular hairs ca 0.5 mm long; range of dimensions
not so great; fronds 75 (40-130) cm long; blades 56 (30-84) cm long, 12 (8.5-21)
cm wide; segments 62 (45-104) mm long, 5.5 (4-7) mm wide.

Holotype: Colombia, Cundinamarca, collected on rocky ridge in brush on foot-
hills above Bogota, just N of mouth of Quebrada El Obispo, 830 m alt, 10 Jan
1943, Fosberg 19688 US 2290492; isotype US.

Habitat: epiphyte, occasionally epipetric or terrestrial in rich humus, in
montane forest, 220-330 (830) m alt.

Distribution: Panama to western Bolivia. — Fig. 11.

Panama: chiriqui: Camp Aguacatal, Maxon 5300 (US).

Venezuela: distrito federal: Avila de Caracas, Vareschi s.n., Feb 1963 (TENN).

Colombia: Mutis 3235 (US); Ariste-Joseph s.n. (US), magdalena: Santa Marta,
Carriker 16 (US); Sierra de Perija, Casacara Valley, 23 km E of Codazzi, Grant 10982
(US), cundinamarca: Boca de Monte, W margin, Sabana de Bogota, Cuatrecasas, Jaramillo
M. & Huertas 25809 (US); Bogota, Ariste-Joseph 1919, 1920 (GH, US); Montserrate, above
Bogota, Ewan 16920 (US); Paramo de Cruz Verde, Apollinaire & Arthur 4 (US),
Cuatrescasas 419 (US); Sibate, Pennell 2494 (US), caldas: Pinares, above Salento, Pennell
9258a (US), cauca: Canaan, Mt Purace, Pennell & Killip 6691 (GH, US).

Ecuador: pichincha: Corazon Peak & Corazon Pass, Ewan 16421 (US) ; Mt Pichincha,
Mills 16 (US), canar: San Marcos, 5-8 km NE of Azogues, Camp E-2614 (US), azuay:
N of Sevilla de Oro, Camp E-5209 (GH, MO, US), loja: Zamora-Huaico, Espinosa 2271
(US).

Peru: huanuco: Mito, MacBride & Featherstone 1621, 1731 (US); Muna, trail to
Tambo de Vaca, MacBride 4280 (US).

Bolivia: la paz: Sorata, Williams 2575 (GH, US), cochabamba: Incachaca-San
Antonio, Prov Chapare, Steinbach 8988 (GH, MO); Prov Sacaba, Cochabamba, Steinbach
5868 (GH, MO).

This variety is differentiated from var. glabrescens primarily on the basis of
the more densely villose rachis and the entire rhizome scales. As they both cover
essentially the same range, these apparently are not clinal variations, and appear
sufficiently distinct to warrant formal recognition. The peculiar distribution and
relationships of this variety to the other two suggest that, if this is truly a distinct
variety, there may be an ecological isolation not evident from the data available.

246 ANNALS OF THE MISSOURI BOTANICAL GARDEN

18. Polypodium pectinatum L., Sp. PL 1085, 1753.— Fig. 20.

P. otites sensu Willd. in L, Sp. PL ed. 4, 5: 177, 1810, non L, 1753 (Based on Vahl s.n.,

Herb. Willd. no. 19653, "Habitat in America," B-photo)
Goniophlebium pectinatum (L.) J. Smith, Lond. Jour. Bot. 4: 57, 1842.
P. pectinatum L. var. majus Kuhn ex Krug, Bot. Jahrb. Engler 24: 128, 1897, nomen

nudum. (Based on: Santo Domingo, Eggers 2595b, c, d B; Dominica, Eggers 12 B?;

Martinique, Hahn 55 P)
P. pectinatum L. var. wagneri sensu Jenm., Bull. Bot. Dept. Jamaica 4: 125, 1897. (Not

as to basionym Polypodium wagneri Mett, a species of sect. Goniophlebium. Based

on: Jamaica, Jenman s.n. NY)

Rhizome long-creeping, 6.5 (5.9) mm in diam; rhizome paleae linear-triangu-
lar, dark red-brown with pale base, basifixed, filiform at the apex, inconspicuously
comose, non-clathrate, inconspicuously and sparsely fimbriate; fronds 60 (30-126)
cm long, approximate on the rhizome; rachis 6 (3.5-9) times as long as the stipe;
stipe and rachis light red-brown, thinly pilose with acicular hairs up to 0.5 mm
long, usually much shorter, ctenoid hairs inconspicuous to very conspicuous;
rachis paleae usually absent, when present inconspicuous, filiform and entire;
blades linear-elliptic, 50 (24-113) cm long, 7.5 (4.5-10.5) cm wide, abruptly
cuneate at the base; segments 37 (23-53) mm long, 4.5 (3-7) mm wide, perpendic-
ular to the rachis, abruptly reduced to a few pairs of lobes at the blade base,
straight or slightly falcate, obtuse, the base expanded on the rachis and symmetrical,
herbaceous, entire; lamina puberulous with small fine acicular hairs up to 0.25
mm long, usually shorter, also with inconspicuous, clavate hairs up to lOO/x long;
costa slightly decurrent on the rachis, the trichomes as on the lamina and with
reduced ctenoid hairs; veins twice (once) -forked, partially to almost completely
anastomosing except at the apex of the segments; guard cells ca 47^ long; sori
medial, round, small, with a few simple, clavate paraphyses; sporangia ca 180^
in diam, usually with one inconspicuous clavate capsular seta ca 65^ long; spores
reniform, tuberculate, ca 45/* long; n = 37.

Type: There is no specimen labeled P. pectinatum in the Linnaean Herbarium,
nor any specimen agreeing with the diagnosis; therefore, the species was evidently
based only on the references cited. There were two of these: Polypodium lonchi-
tidis folio, Pet. fil. 31. t.7.f. 14 and Polypodium nigrum tenerius sectum. Plum,
amer. 26. t. 37 (The "tenerius" is an error of Linnaeus for "tenuius") The Petiver
figure is copied from Plumier, and therefore there is essentially only the Plumier
citation left, automatically making the Plumier plate the lectotype. This ques-
tion was put to Mr. J. E. Dandy of the British Museum (Natural History) and
this is his conclusion also. The Plumier citation refers to his "Description des
plantes de l'Amerique," 1693. The text indicates that his species came from "nos
Antilles," i.e. from the French Antilles. In a later work, "Tractatus de filicibus
Americanis," (p. 64, t. 83, 1705) Plumier repeated the same description and figure;
he amplified the text by replacing "nos Antilles" by Martinique and by adding
the locality Jamaica, on the basis of a publication by Sloane on Jamaica. Plumier's
original description was thus based on one of his own collections from Martinique
which is thus the type locality. In publishing P. pectinatum, Linnaeus picked
up only the locality Jamaica in error. The only species of this group occurring in

EVANS— POLYPODIUM 247

Martinique is the species as I have delimited it here, which also does occur in
Jamaica. Linnaeus' description of the rhizome as "nudo" was evidently taken
from Plumier's illustration, in which the artist has omitted showing the rhizome
scales, probably in order to show more clearly the leaf-scars (which are drawn
in a very exaggerated way) ; no species of this alliance has or could be expected
to have a naked (i.e. scaleless) rhizome.

Habitat: on limestone rocks or epiphytic on tree trunks and horizontal
branches, occasionally terrestrial, in lowland or montane humid forest, from sea
level to 1100 malt.

Distribution: West Indies and Costa Rica to northwestern South America —
Fig. 10.

Tonduz 11309 (US).

Panama: San Pablo, Blake s.n., 8 July 1872 (GH). bocas del toro: Chiriquf Lagoon
von Wedel 2611 (MO), cocii: El Valle, Allen 228 (US); Penonome, Williams 495 (US)!
canal zone: Frijoles, Standley 27571 (MO, US) ; Barro Colorado I, Standley 31410, 40973
(US); Rio Indio de Gatun, Maxon 4855 (US); Rio Medio, Miller 1729 (US).

Cuba: pinar del rio: source of Rio Taco-Taco, Sierra de Los Organos, Morton 4295,
4300, 4315, 4485 (US), orients: El Yunque, nr Baracoa, Underwood & Earle 1292 (US),
Pollard & Palmer 183 (US); Las Ninfas, Guantanamo, Hioram 1458 (US); W of Santiago
Graves 1482 (US) ; 20 km SW of Companfa de Moa Mill, Sierra de Moa, Howard 5973
(MO); N spur of Sierra Maestra W of Rio Yao, Morton & Acuna 3352 (US); Sierra Nipe
Loma Mensura, Ekman 3149 (US), Woodfred & Shafer 3119, 3210 (US), Rio Piloto!
Ekman 2100 (US); Yateras, Josephina, N of Jagiiey, Maxon 4103 (GH, US); Yateras, S of
Jagiiey, Maxon 4219 (GH); Yateras, Monte Verde, Maxon 4325 (GH, US), Shafer 8701

Haiti: Marmelade, Leonard 7289a (US); Pilbury Hill, Miller 188 (US).
Dominican Republic: barahona: Fuertes 1013 (GH); Paradis, Abbott 1580a (US).
duarte: San Francisco de Macon's, La Bracito, Abbott 2033 (US), la vega: Piedra Blanca
Allard 13814, 13990, 13974, 13988 (US), samana: Laguna, Abbott 367, 369 (US); Rojo
Cabo, Abbott 164, 1164 (US); Samana, Abbott 490 (US); Sanchez, Abbott 112, 113 (US).
santo domingo : La Cumbra, Raunkiaer s.n., 4 Aug 1906 (US); Mt Isabel de la Torre,
Eggers 2595c, 2595d (B). seibo: Samana Bay, Miller 1032 (US); 3-4 mi W of San Lorenzo
Bay, Abbott 1273 (US).

Jamaica: Roper s.n., in 1895 (MO). Cinchona, Clute s.n., 21 Febr 1900 (GH) ; Hartford,
Priestman's River, Maxon 2525, 2538 (US); Hollymount, Mt Diablo, Maxon 2319 (US) ■
John Crow Mts, E of Seamen's Valley, Portland, Maxon & Killip 239 (GH, US); Port
Antonio, Millspaugh, 28 Jan-6 Feb 1899 (GH) .

Puerto Rico: Aibonito, Hioram 269 (US); betw Arecibo & Utado, Britton & Cowell
2048 (US) ; 8 mi SW of Carolina (US) ; El Yunque, Scamman 6560 (GH) ; Wagner s n
16 Apr 1944 (US); El Yunque, Rio Piedras, Johnston 750 (US); Loma Icaco Sierra de
Naguabo, Shafer 3393 (US); Mariaco, Kuhn 426 (US); Mayagiiez, Heller 4492 (GH, MO,
US); rd from Ponce to Adjuntas, Underwood & Griggs 765 (US); Adjuntas Rd, 7 mi from
Ponce, Heller s.n., 2 Dec 1902 (US); Sierra de Luquillo, Dale s.n., Febr 1926 (MICH);
Utuado, Underwood & Griggs 21 (US); Vega Alta, Wagner s.n., 30 Jan 1944 (US)

Lesser Antilles: st. kitt's: Nine Turn Gut, Box 267 (US); Phillips Level, Proctor
!S ( S H) " M0NT * ERRAT: Turner '■"■ (US); Shafer 748 (US); Centre Hills, Salem, Proctor
18837 (GH, US). Guadeloupe: Camp Jacob, Duss 4094 (US); Bains Jaunes, Questel 3165
(US); Matouba, Scamman 8168 (GH). dominica: Roseau Valley, Hodge 56 (GH) ; Morne
Brule, Portsmouth, Hodge 57 (MO, US); Sylvania, Hodge 1151 (US); Morne Diablotin
Hodge 2703 (GH, US). Martinique: Fontaines Didier, Duss 1676 (GH MO US) Hahn
55 (P). st. lucia: 2.5 mi SSE of Soufriere, Proctor 17869 (GH, U
249 (GH). Grenada: St George Parish, Grenville Vale, Hunnewell 19435 (GH"
42 (US); Grand Etang, Beard 1260 (MO, US); Mt Pleasant, Milton, 19 Mar 1924 (US)
st. eustatius: Fairchild s.n., 22 Jan 1932 (US) . ' '

248 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Venezuela: bolivar: Cerro Pijiguao, N end of Serrania Suapure, above Pijiguao,
Wurdack & Monachino 41300 (US).

Colombia: santander: Rfo Surata valley, above Surata, Killip & Smith 16569 (GH).
meta: Sierra de la Macarena, Carlo, Entrada, Philipson & Idroho 1755 (US), vaupes: Rio
Guaviare, San Jose del Guaviare, Cuatrecasas 7468 (US), choco: Negria, Rfo San Juan
Killip 35027 (GH, US), Primavera, Scolnik 1653 (US), cauca: Gorgona I, Taylor 1227
(US), narino: Rio Oretopungo, nr Rio Putumayo, above Puerto Asis, Ewan 26764 (Ub).

Ecuador: manabi: Salango, Haught 3372 (US); Quininde, Holdridge 1678 (US).

Peru: san martln: Tingo Maria, Allard 21442 (US); Boqueron Pass Allard 21734
(US), loreto: Pongo de Manseriche, Rio Santiago, Alexia 6212 (GH, MICH, MO, US),
6368a (GH, US).

Polypodium pectinatum, as interpreted here on the basis of the type illustration
mentioned above, agrees with the usual concept of the species as it occurs in the
West Indies. This interpretation goes back at least to Swartz, who correctly under-
stood the Plumier description and illustration. I have seen three specimens from
the Swartz Herbarium in Stockholm which are labeled pectinatum by Swartz and
which agree with my concept of the species. A number of specimens from the
West Indies and elsewhere have been wrongly identified as P. pectinatum; these
are now referred to P. dispersum, P. ptilodon, and other species.

This species is best characterized by the very short hairs of the lamina, the
short sporangial setae, and the abrupt reduction in the segments to several pairs
of mere lobes at the base of the blade. Morphologically, it is most similar to P.
camptophyllarium var. camptophyllarium, which has much longer and more dense
hairs and is a larger and coarser plant. Specimens of P. pectinatum from the Lesser
Antilles tend to have somewhat longer, though sparse, hairs and have the basal
segments more gradually reduced than is typical. However, these plants are clearly
related to the other West Indian material of P. pectinatum and not to P.
camptophyllarium var. camptophyllarium.

19. Polypodium venturii de la Sota, Opera Lilloana 5: 186, fig. 31, 1960.

Rhizome long-creeping, 5 (4-6) mm in diam; rhizome paleae narrow-tri-
angular, dark red-brown, lustrous, basifixed, acute, conspicuously comose, sub-
clathrate with lighter non-clathrate base, entire; fronds 55 (45-60) cm long, ap-
proximate on the rhizome; rachis 3 (2-5) times as long as the stipe; stipe and
rachis red-brown, pilose with acicular, golden hairs, ctenoid hairs conspicuous;
rachis paleae inconspicuous, filiform, entire; blades narrow-ovate, 40 (28-47) cm
long, 8 (6.5-9) cm wide, cuneate to subtruncate at the base; segments 40 (33-45)
mm long, 5 (4.5-5) mm wide, perpendicular to the rachis or slightly deflexed at
the blade base, linear-triangular, straight, obtuse, subsymmetrical at the base,
chartaceous-herbaceous, entire, spaces between the segments 3 / 4 to 1 times the width
of the segments; lamina thinly pilose with clear, acicular hairs ca 0.35 mm long
and with scattered clavate hairs ca 150^ long; costa decurrent on the rachis, the
trichomes as on the lamina; veins once- or twice-forked, free or occasionally
partially anastomosing; guard cells ca 56^ long; sori inframedial to medial, round,
with simple clavate paraphyses ca 0.2 mm long; sporangia without setae; spores
reniform, tuberculate, ca 50/* long.

EVANS— POLYPODIUM

Holotype: Argentina, Tucuman, Tafi, Yerba Buena, 800 m, 17 Apr 1921,
Venturi 1232 LIL. Not seen, but de la Sota has verified my material.
Habitat: epiphytic or on fallen logs, in forests, 600-2700 m alt.

Colasay, Woytkowski 7013 (US), loreto: Pampayacu, Kanehira

Argentina: tucuman: Chichigasta, Las Pavas, Venturi 2974 (GH, US).

This species is similar to P. camptophyllarium, from which it can be distin-
guished by the usually completely free venation, smaller sori, non-setose sporangia,
and the softer, more delicate foliage.

Although the two collections from Peru are less pilose and the hairs longer,
in other characters they agree well with the Argentinian plants.

20. Polypodium pectinatiforme Lindm., Hedwigia 43: 309, 1904.

Lindm., Ark. Bot. 1:239, t. 11, fig. 2, 1903, non Mett., 1855. (Lectotype:
Brazil, Rio Grande do Sul, Porto Alegra, Regnell I A381 S. Syntypes: Brazil, Rio de
Janeiro, Mosen 114 S, S-PA; Minas Gerais, Caldas, Mosen 2195 C, M, S, S-PA; Rio
Grande do Sul, Hamberger Berg & Colon, Silvcira Marlins, Regnell 1 A493 S, S-PA,
US, 493'/ 2 S; Cuba, Wright 1051 B, BR, G, L, NY, S-PA).

Santa Catarina, Joinville, Schmalz I
P. pectinatiforme Lindm. var. hirsutum

(Holotype: Brazil, Rio Grande do Si

Rhizome long- to short-creeping, 5 (2-10) mm in diam; rhizome paleae linear-
triangular, dark red-brown with pale base, lustrous, basifixed, acute, inconspicuously
comose, non-clathrate, entire; fronds 40 (12-65) cm long, approximate on the
rhizome; rachis 7 (5-11) times as long as the stipe; stipe and rachis red-brown,
thinly pilose with golden, acicular hairs up to 1 mm long, sometimes also villose
with short acicular hairs, ctenoid hairs conspicuous; rachis paleae inconspicuous,
filiform, entire; blades narrow-ovate, 35 (10-60) cm long, 9 (3.5-14) cm wide,
cuneate at the base; segments linear to linear-triangular, straight, 40 (15-65) mm
long, 3 (2-6) mm wide, perpendicular to the rachis, the basal segments strongly
deflexed and shortened or reduced to auricles at the blade base, asymmetrical at
the base, the lower edge perpendicular to the rachis, acute, herbaceous, entire to
crenulate; lamina with numerous acicular hairs up to 0.3 mm long, also with
occasional clavate hairs; costa perpendicular to the rachis, thinly pilose with
long, acicular hairs up to 0.5 mm long, sometimes with ctenoid hairs; veins once
(twice) -forked, free; guard cells ca 39^u long; sori supramedial to sub-marginal,
round, with very long, simple, clavate paraphyses overtopping the young sorus,
these obscure in the mature sori; sporangia without setae; spores reniform, tubercu-
late, ca 47^ long.

Type: based on P. microsorum Lindm., non Mett. The syntypes do not consti-
tute a homogenous sample of the species, in that as presently construed Wright
1051 is P. dispersum, Mosen 2195 is P. singeri, and Mosen 114 in S is P. ptilodon
var. robustum, but in S-PA is P. pectinatiforme.

Habitat: epiphytic, occasionally epipetric or terrestrial, in forests, 1400-1700 m

250 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Distribution: southern Brazil, northern Argentina and Paraguay.— Fig. 10.

Brazil: minas gerais: Caldas, Regnell 319 (MO, US); Serra de Caldas, Mosen 2198
(C, S, S-PA); Serra de Mutuca, beyond Barreiro, Williams 6642 (GH, US), 6644 (GH),
Vargem de Ouro Podre, Williams 6196a, 6197 (GH). Rio de Janeiro: Rio de Janeiro,
Glaziou 396 p.p. (BR, non P); Mosen 113 p.p. (S, non S-PA); Mosen 114 p.p. (S-PA, non
S); Serra dos Orgaos, Liietzelburg 6915 (MICH), sao paulo: Campos do JordSo, Harsh-
berger 930 (US), Leite 388 (GH). santa catarina: Sombrio, Reitz CI 159 (US); Lages,
Spannagel (Rosenstock 396) (US), Spannagel s.n., in 1920 (US), rio grande do sul:
Hamburger Berg, Lindman LA. 493 (S, S-PA, US); Sao Leopoldo, Rick s.n. (GH); Silveira
L n Lindman A. 493'A (S).

Paraguay: Caaguazu, Hassler 9084 (GH).

Argentina: misiones: Yerbel Viejo, Burkart 1584 (GH).

Superficially, P. pectinatiforme and P. paradiseae are very similar. The latter
is considerably larger, however, with segments more gradually reduced at the base
of the blade, and with characteristically more strongly crenate and falcate segments
with marginal sori. The rhizome scales of P. pectinatiforme are larger and only
inconspicuously comose.

Rosenstock's P. pectinatiforme var. hirsutum applies to specimens with a
more villose rachis. This may be a distinct taxon, but I do not have sufficient
material at hand to evaluate it satisfactorily.

21. Polypodium paradiseae Langsd. & Fisch., Icon. Fil. 11, t. 11, 1810.

P. pectinatum L. var. paradiseae (Langsd. & Fisch.) Baker in Mart., Fl. Bras. 1(2): 517,

1870.

Rhizome long-creeping, 8 (6-10) mm in diam; rhizome paleae narrow-tri-
angular, red-brown, lustrous, basifixed, acuminate, comose, non-clathrate, entire
or with a few inconspicuous fimbriae; fronds 120 (70-170) cm long, spaced ca 2 cm
apart on the rhizome; rachis 7 (4.5-11.5) times as long as the stipe; stipe and
rachis red-brown, pilose with long, golden, acicular hairs, ctenoid hairs conspicu-
ous; rachis paleae inconspicuous, filiform, entire; blades 100 (60-150) cm long,
16 (10-20) cm wide, narrow-rhombic, narrow-cuneate at the base; segments 80
(55-105) mm long, 6.5 (4-9) mm wide, linear-triangular, falcate, gradually
shortened and deflexed to mere auricles at the blade base, coriaceous, acute, unequal
at the base with the lower edge perpendicular to the rachis or incised, 24 (18-30)
pairs of segments in the middle V 4 of the frond, crenate; lamina thinly pilose with
golden, acicular hairs, ca 0.3 mm long and with scattered clavate hairs ca 0.25
mm long; costa perpendicular to or curved downward from the rachis, pilose with
long, golden, acicular hairs ca 0.4 mm long; veins 2-3-forked, free; guard cells ca
43^ long; sori marginal to submarginal, round, with simple clavate paraphyses
ca 0.25 mm long; sporangia without setae; spores reniform, tuberculate, ca 43p long;
n = 37.

Holotype: Brazil, Santa Catarina, Santa Catarina I, Langsdorff s.n. Herb.
Fisch. LE; isotype Herb. Willd. B, photograph.

Habitat: terrestrial or occasionally epiphytic, in wet forests, up to 1600 m alt.

Distribution: southeastern Brazil. — Fig. 10.

rio de Janeiro: Matto, Macieiras, Mt Itatiaya, Smith 1764 (US); Monte Serrat, Mt
Itatiaya, Smith 2296 (GH); Serra do Itatiaio, Dusen 467 (US), sao paulo: Alto da Serra,
Wacket (US); Campos do Jordao, Bailey 889 (US); Guaruja, Dusen 14213 (US). paranA:

EVANS — POLYPODIUM 251

Alexandra, Dusen 17128 (GH); Serra do Mar, Desiro Ypiranga, Dusen 14471 (GH);

Tacarachy, Dusen 6602 (GH, MO, US), 14743 (MO), 14743a (GH, MO), santa catarina-

.rina I, Itacorobi, Rohr 1040 (US); Itajaf, Praia Braba, Smith & Reitz 6093

ville, Muller (US); Mun. Port I by new airport E of Porto

ith y Reitz 8845 (US); Sombrio, Reitz 1750 (US). Rio Grande do sul- Sao

Leopoldo, Leite 1462 (211), 1716, 1855 (GH), Reitz C679?; Sao Francisco do Paula* Vila

Oliva, Rambo 31204 (MO); Sao Salvador, Leite 1934 (212) (GH).

This species appears to intergrade with P. ptilodon var. robustum. However,
the marginal sori and the crenate, narrow segments, gradually reduced and deflexed
at the base of the blade are characteristic of P. paradiseae. Also, if the total length
of the blade is measured and then the numbers of pairs of segments counted in the
middle fourth of the blade length, these two species are strikingly separated. In
my test of this character Polypodium paradiseae had 24 (18-30) pairs of segments,
whereas P. ptilodon (including all four varieties) had 16 (10-21) pairs. Applying
a Mann Whitney-U test to samples of 10 fronds of each species, the two groups
were found to be significantly different at the .002 level.

22. Polypodium camptophyllarium Fee, Mem. Foug. 8: 86, 1857.

Rhizome short- to long-creeping, 5-10 mm in diam; rhizome paleae linear-
triangular, dark red-brown, basifixed, filiform at the apex, comose, with linear non-
clathrate cells, entire to slightly and inconspicuously fimbriate; fronds 1.6-16 dm
long, closely spaced to approximate on the rhizome; rachis 1.5-7 times as long as
the stipe; stipe and rachis light to dark red-brown, pilose with acicular hairs 0.4-1
mm long, ctenoid hairs present; rachis paleae inconspicuous, filiform, entire; blades
narrow- to linear-ovate, 13-138 cm long, 4-17 cm wide, cuneate at the base; seg-
ments 20-90 mm long, 3-8 mm wide, perpendicular to the rachis, or slightly deflexed
at the blade base, reduced to lobes or auricles at the blade base, slightly falcate to
straight, herbaceous to coriaceous, obtuse to acute, the base expanded and symmetri-
cal or the lower edge approaching a right angle with the rachis toward the base
of the blade, or incised, entire; lamina pilose with pale acicular hairs up to 0.5 mm
long, and with scattered, clavate, simple or forked hairs ca 0.2 mm long; costa
decurrent on the rachis, with fine, pale, acicular hairs like those on the lamina or
with coarser, longer and yellower hairs than those of the lamina; veins once- or
twice-forked, free to completely anastamosing with a free included veinlet; guard
cells 52-57^ long; sori medial to supramedial, with long, simple or forked, clavate
paraphyses; sporangia with 0-3 multicellular capsular setae up to 250^ long; spores
reniform, smoothish to tuberculate, 43-60^ long.

Polypodium camptophyllarium is a widespread and variable species, distin-
guished primarily by the long-pilose pubescence of the rachis and lamina. It is
subdivided into four varieties showing varying degrees of similarity to P. pectinatum.
I. Fronds less than 30 cm long; veins once-forked, free throughout

22d. P. camptophyllarium var. abbreviatum

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Veins only partially anastomosing; receptacular paraphyses and laminar

s simple ' l > 4.5- (1.5-7) times as long as the stipe; basal segments

reduced to auricles or strongly deflexed and only partially reduced; plants

Rachis and costa pilose with soft hairs up to 0.5 mm long, the lamina
the same; sori occupying half the width of the segment or less

22a. P. camptophyllarium var. camptophyllarium

thinly pilose with scattered, harsh acicular hairs ca
0.75 mm long, the lamina thinly pilose with finer hairs ca 0.35
mm long; sori occupying most of the width of the segment
22c. P. camptophyllarium var. lachniferum

22a. Polypodium camptophyllarium var. camptophyllarium.

P cinerascens Lindm, Ark. Bot. 1: 238, t. 11, fig. 6, 1903. (Lectotype: Brazil, Rio de Janeiro,

Regnell I 474 S, isolectotypes BR, C, S, S-PA, US. Syntype: Regnell I A99 S, S-PA)
P. robustum var. cinerascens (Lindm.) Hassl., Trab. Inst. Bot. Farm. Buenos Aires 45:69,

P. lachniferum Hieron., Bot. Jahrb. Engler 34: 515, 1904, p.p., not as to lectotype. (for

Fronds 70 (25-165) cm long, spaced ca 1 cm apart on the rhizome; rachis 4
(1.5-5.5) times as long as the stipe; stipe and rachis light red-brown, pilose with
pale, acicular hairs up to 0.75 mm long; blades 58 (23-138) cm long, 9 (3.5-17)
cm wide; segments 45 (15-90) mm long, 5 (4-8) mm wide, perpendicular to the
rachis, reduced to mere lobes at the blade base, slightly falcate to straight,
herbaceous; lamina pilose with pale, acicular hairs, all ca 0.5 mm long; costa with
trichomes like those on the lamina; veins twice (once) -forked, usually partially
anastomosing; guard cells ca 52^ long; sori occupying half the width of the seg-
ment or less, with long simple clavate paraphyses; sporangia with 2 or 3 capsular
setae ca 250/4 long; spores reniform, tuberculate, ca 54 (47-60)^ long, mostly with
64 normal appearing spores per sporangium; n = 74.

Holotype: Colombia, Norte de Santander, Environs of Ocafla, in 1850, Schlim
128, p.p. P; isotypes BR, G, K photograph, not L.

Habitat: terrestrial, occasionally epiphytic, in rocky soil or on rocks in forests,
often in openings or at the edge of the forest, 350-2100 m alt.

Distribution: Costa Rica, Cuba, Jamaica, northwestern South America to
southern Brazil.— Fig. 10.

Costa Rica: limon: Guapiles, Standley 37291 (US).

Panama: Panama: 5 mi N of Cerro Azu, rd to Cerro Jefe, Blum et al. 1692 (FSU).

Cuba- oriente: El Cristo, Ekman 1468 (US); Sierra Maestra, Termino del Cobre,
Clement 1129 (US), Finca Reunion to Loma del Gato, Ekman 6925 (US); El Cobre,
Pollard y Palmer 412 (GH, MO, US).

Dominican Republic: barahona: Montiada Nueva, SE of Polo, Howard 8431 (GH).
san juan: Juan Santiago, Howard 9288 (GH, US); Hondo Valle, Howard 8753 (GH, US).
Santiago: Arroyo Juan Fino, Jicome, Valeur 655 (GH, US), la vega: Salto de Constanza,
1215 (US); Salto de Jimenoa, Jarabacoa, Jimenez 1533 (US).

Haiti- Bombardopolis, Leonard 13457 (US) ; Dept l'Artibonite, Ennery, Leonard 9576

cy, Leonard 4432 (GH, US); Massif du Nord, Hinche, Ekman 6166 (US); Marc

Sal, Holdridge 1380 (US); Massif de la Selle, Petionville, Ekman 10024 (US); ' lissic i

Fonds Varettes, Leonard 3958 (US); St. Louis du Nord, Leonard 14469 (GH, US); Dspt

du Nord, St. Michel de 1'Atalaye, Leonard 7627 (GH, US).

Jamaica- Abbey Green, Maxon 10060 (GH, US); above Cedar Valley, Silver Hill Gap,
Maxon 10272 (GH, US); Cinchona, Clute s.n., 21 Febr 1900 (US); Cinchona to Morce's

EVANS POLYPODIUM 253

Gap, Killip 204 (US); Port Royal Mrs, Flamstead, Maxon 8683a (US); Mandeville, Gilbert
s.n., Febr 1895 (MO), Maxon 2581 (US); Mocho, Catadupa, Maxon & Killip 1551 (GH
US); Moneague, St. Ann, Hunnewell 15217 (GH); Mt Diabolo, Maxon & Killip 469 (GH
US); St. Helen's Gap, St. Andrew, Maxon & Killip 631 (GH, US); Troy, Underwood 2930
(US), 3296 (US).

Venezuela: mongas: Quebrada Colorado Grande, SW of Caripe, Steyermark 61690a
(US), distrito federal: Quebrada de San Lazaro, nr Caracas, Pittier 9760 (US).

Colombia: magdalena: Haught 3942 (US); betw Pueblo Viejo & San Miguel, Seifriz
359x (US), bolivar: El Patico, Rio Paez, Lehmann K. 110 (US), santander: Pica-Pica
Valley, above Tapata, N of Toledo, Killip & Smith 20191 (GH, US), valle: nr Con-ales
Lehmann 4424 (B); El Naranjo, Rio Dagua, Lehmann 5049 (B, US).

Peru: Cedrobamba, Herrera 1580 (US), huanuco: Chinchao to Puerte Durand,
Coronado 91 (US), apurimac: Quebrada Nataza, Vargas C. 2299 (US), cusco: Urubamba
River, Machu Picchu, Heller 2206 (US), Herrera 3479 (US).

Bolivia: la paz: Apolo, Williams 1124 (US); N Yungas, Milluhuaya, Buchtien 4235
(MO, US); Sirupaya, Yanucachi, S Yungas, Buchtien 482 (US); Tres Crucas, Herzog 1616

Brazil: minas gerais: Mun. Ouro Preto, Andorinhas, Macedo 2864 (MO); Mun.
Ituiutaba, Aroeira, Macedo 2383 (MO); Mun. Nova Lima, Serra do Curral, Williams 6505
(GH);RegnellIA99 (S).

This is the central and widespread variety which ties the four together. The
other three were not, however, necessarily derived from this variety, though var.
macedoi very likely has been. The var. abbreviatum has considerably smaller
spores, suggesting that it is a diploid and may have arisen by divergence. The
var. lachniferum can, in most cases, be distinguished by the rather coarse golden
scattered hairs on the rachis and costa, the slightly to strongly deflexed lower seg-
ments often reduced and auriculate, and the sori, which occupy the major portion
of the segment. However, the collections which I have range from the West Indies
to Peru and show little variation between range limits and considerable intergrada-
tion throughout its range with var. camp tophy liar ium. The range of this and
related varieties is complicated by its extension over what I consider to be four
geographical areas (see Figs. 8 & 10). At the overlapping of three of these distri-
butional areas in northern South America, the representatives of the P. pectinatum-
plwnula complex in Colombia and Venezuela are often difficult to interpret.

The material of var. camptophy liar ium from the West Indies and from
Colombia is somewhat different. In the West Indies the segments tend to be
more obtuse and the plants somewhat smaller, the spores more irregular, and a
few evidences of apogamy are suggested, as discussed below. The material from
Colombia tends to be larger with straighter and more acute segments. The
laminar trichomes are essentially identical throughout the range. In Brazil the
plants again begin to exhibit segments which are more obtuse and falcate as in
the West Indian material, except that the stipe is considerably longer, and the
anastomosing of the venation is more complete, as in var. macedoi. Because few
collections from this area have been examined and because there does appear to be
sufficient difference between the long-stiped materials which I have called var.
camptophyllarium and var. macedoi, I am maintaining the latter as a distinct

Throughout the range of this species the spores seem normal, though large
and somewhat irregular. The spores are much like the large and somewhat irregu-

254 ANNALS OF THE MISSOURI BOTANICAL GARDEN

lar spores of P. ptildon var. caespitosum. A few of the specimens from Cuba and
Jamaica have quite irregular reniform spores. They appear viable but are attached
in pairs by the terminal ends; sometimes only by one end, thus making a double
spore in a horseshoe shape; sometimes by both ends, making a deformed, dough-
nut-shaped, double spore. These may be apogamous hybrids. Further cytological
and populational studies of P. camptophyllarium are in progress, but it is sug-
gested that these few plants may be isolated cases of apogamous reproduction.
I have included these specimens with var. camptophyllarium as they represent
only a small fraction of the total material, and they cannot at present be dis-
tinguished other than by a critical observation of the spores.

22b. Polypodium camptophyllarium var. macedoi (Brade) A. M. Evans, comb.

P. macedoi Brade, Arq. Jard. Bot. Rio de Janeiro lis 30, t. 10, 11, fig. 3, 1951. (Holotype:
Brazil, Estado de Minas Gerais, Ituiutaba, Macedo 1098 RB-63398 not saen, isotypes

Similar to var. camptophyllarium, except: rachis only 2.5 (1.5-3) times as
long as the stipe; veins completely anastomosing or free at the apex only; lamina
with forked, clavate-setose hairs; receptacular paraphyses consisting of forked hairs
with one clavate and one setose branch; spores reniform, smoothish, ca AAft long.

Habitat: terrestrial in humid forests and along river banks.

Distribution (Fig. 10) : Bolivia: la paz: Polo-Polo, Corioco, N Yungas, Buchtien 3505
(US): Yungas, Rusby 357 (US).

Brazil: minas gerais: San Vicente, Mun. Ituiutaba, Macedo 1894 (S, US), 2302 (MO,
US), 2383 (US), 2864 (US).

This variety is distinguished from var. camptophyllarium on the basis of
the completely anastomosing venation, longer stipes, and the presence of forked
hairs on the lamina, as well as receptacular paraphyses consisting of one clavate
and one setose branch.

Although in the original description Macedo considered this a goniophlebioid
Polypodium because of its venation, in all other respects this variety is typical
of the P. pectinatum-plumula complex. The extent of the anastomosing of the
venation is only an extreme expression of what occurs to a lesser extent in var.
camptophyllarium and in P. pectinatum s.s.

22c. Polypodium camptophyllarium var. lachniferum (Hieron.) A. M. Evans,

P lachniferum Hieron., Bot. Jahrb. Engler 34:515, 1904. (Lectotype: Ecuador, Mt Tun-
guragua, 1500-2500 m, Lehmann 458 B, isolectotypes LE, US. Syntypes: Colombia,
Cauca, Central Cordillera, nr Corrales, 2000-2800 m, Lehmann 4424 B; Colombia, El
Valle, nr El Naranjo, on Rio Dagua, 300-1000 m, Lehmann 5049 B, US) The Colom-
bian collections are referable to var. camptophyllarium.

Rhizome as in var. camptophyllarium except the rhizome paleae with entire
margins; fronds 46 (33-70) cm long, approximate on the rhizome; rachis 5 (1.7-7)
times as long as the stipe; stipe and rachis light to dark red-brown, thinly pilose
with coarse, golden, acicular hairs ca 1 mm long, ctenoid hairs conspicuous; rachis
paleae inconspicuous, filiform, entire; blades narrow-ovate, 35 (25-60) cm long,

EVANS— POLYPODIUM 255

7.5 (5-11.5) cm wide, broad-cuneate at the base; segments 40 (25-60) mm long,
4.5 (3.5-5) mm wide, perpendicular to the rachis, slightly deflexed and reduced
to auricles at the blade base, slightly falcate, herbaceous to coriaceous, acute, the
lower edge perpendicular to the rachis or incised, entire; lamina thinly pilose
with acicular hairs up to 0.35 mm long and with scattered clavate hairs; costa
decurrent on the rachis, with scattered long hairs up to 0.75 mm long; veins twice-
forked, free or occasionally partially anastomosing; guard cells ca Slfj. long; sori
medial, round, occupying most of the width of the lamina, with long, simple,
clavate paraphyses; sporangia occasionally with one capsular seta ca 200 ft long;
spores reniform, smoothish, 54 (41-69)^ long; n=74.

Habitat: epipetric or terrestrial, montane, on and about rocks on wooded
mountain sides, 600-2300 m alt.

Distribution: Cuba, Jamaica, northwestern South America. — Fig. 10.

Cuba: oriente: La Perla, Shafer 8474 (US); Loma del Gato, Cobre Range of Sierra
Maestra, Leon et al. 10530 (US).

Jamaica: Arntully, Orcutt 3105a (US); Blue Mt Peak, Hitchcock s.n., 13 Dec 1890
(MO); Chestervale, Underwood 1148 (US), Cinchona, Maxon 2596 (US), Underwood
473 (US), Cinchona Plantation, Underwood 1127 (US); Farm Hill, Orcutt 3402 (US);
Port Royal Mts, Flamstead, Maxon 8683 (GH, US); Blue Mt range, Helen's Gap to Morce's
Gap, Chrysler 1741 (US); Mandeville, Churchill s.n., 16 Mar 1897 (MO); Moody's Gap,
Maxon 1207 (US) ; Cedar Valley, rd to Silver Hill Gap, Maxon 10306 (GH, US)
s.n., 26 Mar 1897 (GH).

: Mucuruba, Gehriger 195 (US); Paramo de la Sal, John 632 (US).
: betw Pueblo Viejo & San Miguel, Seifriz 378 (US), 384 (US).
> Killip & Smith 18488 (GH, US); betw Piedecuesta & Las Vegas,
Killip & Smith 15508 (GH, US), cundinamarca: Bogota, Quebrada, Ewan 15593 (US);
Suba Hill, Sabana of Bogota, Schiefer 596 (US); Facatativa, Ariste-Joseph A190 (US);
Pandi, Perez s.n. (Col 552) (US); mts W of Salto de Tequendama, Uribe Uribe 3382 (US).
antioquia: Bello, Archer 161 (US); Boqueron, Daniel 1634 (US); Copacabana, Henri-
Stanislas 1606 (US).

Ecuador: Grubi, Mille 14 (US), tungurahua: Banos, Popenoe s.n., 6 Mar 1921
(US), azuay: Cuenca, nr union of Rios Tarqui & Yanuncay, Prieto, (Camp E-2658) (US);
Rio Milchichic, 5 km N of Cuenca, Prieto (Camp E-27 Jan Lucas, Dodson

& Thien 564 (US), napo-pastaza: betw Rio Blanco & Rio Verde, rd from Barios to Puyo,
Dodson & Thien 1970 (US).

Peru: cajamarca: Las Juntas, Rose 23221 (US), junin: Huacapistana, Coronado
275 (GH); Manto to Yaupi, Woytkowski 6518 (US), Manto, km 20, Woytk,
(US); Prov Tarma, 5 km SW of Huacapistana, 25 km NE of Tarma, Tryon 5426 (GH,
US); nr Yaupi, Woytkowski 6414 (US), ayacucho: Ccarrapa, betw Huanta & Rio Apuri-
mac Killip & Smith 22473 (US).

This variety is closely related to var. camptophyllarium, the best differentiating
characters being that var. lachniferum is generally a shorter plant, with distinctly
deflexed, reduced and auriculate basal segments; and that there are two distinctly
different sizes of acicular hairs on the frond. There are scattered but long and
harsh hairs on the costa and rachis and shorter, finer hairs (not as dense as in
var. camptophyllarium) on the limina. The sori cover almost the entire width
of the segment, whereas in var. camptophyllarium, the sori, in most cases, cover
only a half or less of the width of the segment.

256 ANNALS OF THE MISSOURI BOTANICAL GARDEN

22d. Polypodium camptophyllarium var. abbreviatum A. M. Evans, var. nov. —

Fig. 18.

A var. camptophyllario simile, frondibus ca 23 cm latis, approximate, laminis
ca 18 cm longis et 6 cm latis, basi late cuneatis, segmentis ca 30 mm longis et
3 mm latis, rhachide perpendicularibus basalibus paullo deflexis exceptis, venis
1-furcatis, liberis, soris medialibus, laminam subtus fere tegentibus, sporangii cum
setis 2-capsularibus differt.

Rhizome as in var. camptophyllarium; fronds 23 (15-30) cm long, approximate
on the rhizome; rachis 4.5 (3.5-5) times as long as the stipe; stipe and rachis
red-brown, pilose with clear, acicular hairs ca 0.4 mm long, ctenoid hairs present;
rachis paleae inconspicuous, filiform, entire; blades narrow-ovate, 18 (13-25)
cm long, 5.5 (4-7) cm wide, broadly cuneate at the base; segments 30 (20-40)
mm long, 3 mm wide, perpendicular to the rachis except the lower few slightly
deflexed, mostly expanded and symmetrical at the base except the lower few pairs
of segments incised on the lower edge and gradually reduced to auricles, herbaceous,
acute, entire, the laminar and costal trichomes as in var. camptophyllarium ex-
cept shorter; costa decurrent; veins once-forked, free; sori medial, round, covering
most of the width of the frond; sporangia with two capsular setae ca 0.14 mm
long; spores reniform, slightly tuberculate, ca 43/* long.

Holotype: Peru, Cusco, Prov La Convenci6n, Distr of Santa Ana, Hacienda
Echarabi, 1300 m, Jan 1926, Herrera 872 US 1342112.

Habitat: terrestrial on rocky hillsides, 1300-1800 m alt.

Distribution (Fig. 10): Peru: cusco: 20 km N of Ollontaytombo, Hitchcock 22516
(US); Pampakjahua, Coronado 105 (US).

This is a local endemic. It is smaller than either var. camptophyllarium or
var. lachniferum. It is related to the former by the trichomes of the lamina and
rachis and to the latter by the reduction and constriction of the basal segments.
Except for the overall small size and the smaller, regular spores, it could easily
be considered as a hybrid of the other two varieties.

23. Polypodium ptilodon Kunze, Linnaea 9: 42, 1834 (as "ptiloton," in error).

Rhizome long-creeping, 8 (5-12) mm in diam; rhizome paleae narrow-triangu-
lar, dark red-brown, lustrous, basifixed, acuminate, comose, non-clathrate, en-
tire; fronds 27-170 cm long, closely-spaced to fasciculate on the rhizome; stipes
1-36 cm long; rachis 5-42 times as long as the stipe; stipe and rachis red-brown,
with scattered acicular hairs up to 2 mm long, ctenoid hairs conspicuous; rachis
paleae inconspicuous, filiform, entire to inconspicuously fimbriate; blades narrow-
to linear-ovate, 25-130 cm long, 3.5-22 cm wide, narrow-cuneate at the base; seg-
ments 2-11 cm long, 3-10 mm wide, perpendicular to the rachis or slightly as-
cending, gradually reduced to lobes or auricles at the blade base, straight to sub-
falcate, acute to rounded, expanded at the base and symmetrical or with the lower
edge perpendicular to the rachis, herbaceous to coriaceous, entire (crenulate),
the spaces between the segments ]/ 2 to 1 times the width of the segments; 10-21
pairs of segments in the middle l / 4 of the blade; lamina with scattered silvery

EVANS — POLYPODIUM 257

acicular hairs up to 0.35 mm long, more densely pilose in an oblong area around
the sorus and with scattered clavate hairs; costa decurrent on, or perpendicular
to, the rachis, with ctenoid hairs, and with scattered, golden, acicular hairs up
to 2 mm long; veins 1-3(4) -forked, free; guard cells 44/t long; sori medial, round
or occasionally oblong, with simple and branched, clavate paraphyses up to 165/*
long; sporangia with 1-3 capsular setae 45-140^ long; spores reniform, tuberculate,
41-52^ long.

This species is subdivided into four closely related varieties. Polypodium
ptilodon var. ptilodon is known to be a diploid, with small spores. The var.
caespitosum is a tetraploid, with larger spores; var. robustum, with similar spores
is presumed to be a tetraploid. The var. pilosum has medium-sized spores, and is
a smaller plant than the other varieties, with longer hairs on the rachis and costae.
1. Stipe 2 (1-5) cm long; segments rounded to obtuse; hairs of the rachis and

costa 1.5-2 mm long 23d. P. ptilodon var. pilosum

1. Stipe 13 (3-36) cm long; segments obtuse to acute; hairs of the rachis and
costa less than 0.75 mm long.

2. Lower edge of the segments perpendicular to the rachis in the lower half
of the blade; fronds 110 (50-170) cm long; spores ca 52,u long; Bolivia,
Argentina, Paraguay, Uruguay, and southern Brazil ....23c. P. ptilodon var. robustum
2. Upper and lower edges of the segments similar and equally expanded
and adnate on the rachis; fronds 85 (30-115) cm long.
3. Stipe 14 (12-17) cm long; fronds 95 (55-115) cm long; spores ca 41ft

long; Peru and Bolivia 23a. P. ptilodon var. ptilodon

3. Stipe 9 (343) cm long; fronds 70 (30-105) cm long; spores ca

23a. Polypodium ptilodon var. ptilodon.

Fronds 97 (55-115) cm long, spaced ca 1 cm apart on the rhizome; rachis
6 (4-7) times as long as the stipe; stipe and rachis pilose with golden acicular
hairs ca 0.75 mm long; rachis paleae entire; blades narrow-ovate, 80 (45-100)
cm long, 15 (10-18) cm wide; segments 70 (55-90) mm long, 9 (8-10) mm wide,
gradually reduced to mere wings at the blade base, obtuse, expanded at the base
and symmetrical, the spaces between segments approximately equal to the width
of the segments, 14 (13-15) pairs of segments in the middle one-fourth of the
blade; costa perpendicular to the rachis, with scattered acicular hairs up to 0.3
mm long; veins 2-or 3-forked, 41^ long; n = 37.

Type: Peru, Dept San Martin, in woods at Pampayacu, July 1829, Poeppig
s.n. Since the original in the Kunze Herbarium in Leipzig is presumed to have
been destroyed, the duplicate at B is designated lectotype.

Habitat: dense forests, 1100-1700 m alt.

Distribution (Fig. 11): peru: amazonas: Rio Utcubamba, Cerro Tapur, 40 km S
of Bagua Grande, Hutchison 1470 (MICH), junin: Chanchosmayo Valley, Schunke 126
(US); San Ramon, Schunke Hacienda, Schunke A151 (US).

Bolivia: la paz: Polo-Polo, Corioco, N Yungas, Buchtien 3508 (MO, US), 3510 (US);
Mapiri, Rusby 356 (US); Simaco, Tipuani Valley, Buchtien 5269 (GH).

This variety is a diploid with spores averaging ca 11^ shorter than either of
the two other similar varieties. Inasmuch as var. caespitosum is known to be a
tetraploid, one may presume that var. robustum, with the same spore size, is also

[Vol. 55
258 ANNALS OF THE MISSOURI BOTANICAL GARDEN

a tetraploid. It would appear that var. ptilodon was the original parental stock
in the species. The var. caespitosum is so similar to var. ptilodon that it could
be an autotetraploid, although its smaller size indicates that it could also be of
hybrid origin with any of several other Caribbean species.

23b. Polypodium ptilodon var. caespitosum (Jenman) A. M. Evans, Amer. Fern

Jour. 58: 170, 1968.
P. pectinatum auct., non L., 1753.
P. pectinatum var. caespitosum Jenman, Bull. Bot. Dept. Jamaica 4: 125, 1897. (Holotype:

Jamaica, Old Enfiknd, 4000 ft, jennum. s.n. Ml, not > which is P. comunik

As in var. ptilodon except: fronds 70 (30-105) cm long; rachis 9 (6-14) times
as long as the stipe; stipe and rachis red-brown to blackish; blades 60 (27-90) cm
long, 12 (6.5-18) cm wide; segments 65 (35-85) mm long, 7 (4-10) wide, obtuse
to acute; guard cells 50/* long; sporangia with 1 or 2 two-celled capsular setae
ca 140 p long; spores ca 56^ long; n=74.

Type: Based on P. pectinatium var. caespitosum Jenman.

Habitat: terrestrial or on mossy tree bases or logs, occasionally epipetric,
usually in damp woods, mountain slopes or ravines, from sea level to 1500 m alt.

Distribution: Florida, Greater Antilles, eastern Mexico to Honduras.— Fig. 11.

Florida: brevard co: N of Titusville, Turnbull Hammock, Small & DeW inkier 10799
(US), citrus co: Homosassa Springs, Correll 5798 (MO); 3 mi NE of Lake Lindsay,
Cooley 5488 (FLAS, US); Pineola grottoes, Evans 2005 (MICH), collier co: Fakahatchee,
Eaton 1133 (GH); Deep Lake, Royal Palm Hammock, Scull s.n. (FLAS). dade co: Hattia
Bauer Hammock, Small 7421 (US), hernando co: Annutalagga Hammock, NW of Brooks-

.. ^ . ' - - 0"-'= '^> > '00 " '-aa Oaaa ' .i ■ -■;»:;,:->■ va. I-' aept 1953 (FLAS);

Brooksville, Choochahattie Hammock, St. John et al. s.n., 13 Jan 1935 (FLAS); Istachatta, Cur-
tiss s.n., Aug 1897 (US); Pineola, McFarlin 3618 (MICH); Brooksville, Spring Lake, Perkins

<>: Avon Pk Wildlife Management Area, N end of Bill
7758 (MICH); Sebring, Highlands Hammock, Correll & McFarlin 6247 (US); Parker
I, 4 mi E of Childs, Ward et al. 2420 (FLAS), Correll 6174 (GH). Hillsborough co:
Hillsborough River St Pk, R21E, T27S, Sect 8, Evans 2018 (MICH), lake co: Eustis,
, 15 Oct 1929 (FLAS). mana-
tee co: Manatee, Garber 15 (FLAS, US), marion co: Stevens s.n., May 1883 (MICH).
orange co: Rock Spring, Harper s.n., 11 Febr 1915 (US), pasco co: Blanton, Small et al.
10858 (US), polk co: Fort Meade, Smith s.n., Mar 1880 (US); 7 mi S of Lakeland,
Scott Lake, Sheridan s.n., 28 Nov 1958 (FLAS); Peace Creek, Smith s.n., Apr 1880 (US);
Winter Haven, Faulkner Hammock, McFarlin 3547 (FLAS, MICH), putnam co: 4 mi
W of Interlachen, West & Arnold s.n., 29 May 1942 (FLAS). seminole co: Sanford,
Orange Co, Rapp s.n., Mar 1910 (FLAS). sumter co: Indian Field Ledges, R21E, T20S,
Sect 34, Evans 2004 (MICH, TENN); Camp's Grove, St. John s.n., 4 Dec 1933 (FLAS).
volusia co: Halifax R, Reynolds s.n., in 1877 (US); 5 mi SW of Port Orange, Freeman
59327 (US).

Mexico: tamulipas: Gomez Farias, Palmer 306 (US), san luis potosi: Hacienda
de Tamasopo, Prinze 3974 (GH, MO. I el 602 (MICH).

Veracruz: Atoyac River, Copeland s.n., 3 Mar 1938 (US); Cordoba, Conzatti & Gonzalez
575 (GH), Fink 74a (US), Woronow 3093 (US); Zacualpan, Purpus 15728 (MICH),
3878 (US).

Honduras: cortes: Rio Lindo, N of Lake Yojoa, Morton 7665 (US).

Cuba: Loma San Juan, Hioram 7064 (US), las villas: Buenos Aires, Howard 5246
(GH, MO), Morton 4154 (US); Arroyo de Manaca, Herradura, Britton 4997 (US).
oriente: La Perla, Leon 3849 (US); Loma del Gato, Clement 1172 (GH), Hioram &
Clement 6490 (US); Santa Ana, 6 mi N of Jaguey, Yateras, Maxon 4212 (GH, US), pinar
del rio: source of Rfo Taco-Taco, Morton 4304 (US).

EVANS— POLYPODIUM

Haiti: Plaisance, Leonard 9383 (US); St. Louis du Nord, Leonard 14314 (US).
Jamaica: Cinchona, Clute 321 (MO); Morce's Gap, Hatch 30 (US); Mount James,
n "-" (US), 8608 (US); Silver Hill Gap, Maxon 1143 (US), Maxon 1145 (US);
"--« 933 (US).

The affinities of the variety are discussed above under var. ptilodon.
23c. Polypodium ptilodon var. robustum (Fee) A. M. Evans, comb. nov.
P. robustum Fee, Crypt. Vase. Braz. 1 : 92, 1869. (Type: Brazil, Rio de Janeiro, Angra dos
Reis, 16 June 1868, Glaziou 2407 BR, C, K photo, P, S, US)

cens f. majus Hassl., Trab. Inst. Bot. Farm. Buenos Aires 45: 70,
(Holotype: Paraguay, Sierra de Amambay, Hassler 11327 G)
. cinerascens f. minus Hassl., loc. cit. (As "minor"). (Lectotype: Paraguay,
Hassler 624 G. Syntype: Paraguay, nr Tobaty, Hassler 6281 G)
As in var. ptilodon except: fronds 110 (50-170) cm long; blades 90 (45-130)
cm long, 14 (9-22) cm wide; segments 70 (40-110) mm long, 8 (5-11) mm wide,
gradually reduced to auricles at the blade base, expanded and symmetrical at the
base in the upper part of the blade, with the lower edge perpendicular to the
rachis in the lower half of the blade, 16 (10-21) pairs of segments in the middle
one-fourth of the blade; costa decurrent on the rachis, thinly pilose with golden,
acicular hairs ca 0.5 mm long; spores ca 52^ long.

Habitat: terrestrial, occasionally epiphytic or epipetric, in moist woods and
along streams, up to 1400 m alt.

Distribution: Bolivia and eastern Brazil to northern Argentina.— Fig. 11.
Bolivia: la paz: Tipuani, Hacienda Simaco, Buchtien 5249 (US).
Brazil: ceara: Sitio Pe de Ladeira, 3 km E of Guaramiranga, Cutler 8324 (GH).
minas gerais: Fazenda de Aguada, Vicosa, Mexia 5180 (GH, MO, US), mo de Janeiro:
Meio de Serra, Smith & Brade 2290 (GH) ; Monte Serrat, Mt Itatiaya, Smith 2296 (GH) ;
Nova-Friburgo, Leite 4816 (MO); Petropolis, Drogo 324 (US); Rio de Janeiro, Wilkes
Exped. in 1838-1842 (US); Tijuca, Ball s.n., 22-23 July 1882 (GH); Glaziou 1725 (BR,
C); Mosen 114 p.p (S, non S-PA). sao paulo: Igaupe, Brade 7735 (US); Mun. Moji-
Guacu, Fazenda Campininha, 6 km NNW of Padua Sales, Eiten & de la Sota 2123 (US),
Parana: Tibagy, Reiss 67 (GH). rio grande do sul: Mun. Rio Pardo, Jurgens s.n. (Rosen-
stock 422) (US); Sao Leopoldo, Reitz 168 (US).
Uruguay: Montevideo, no data (MO).

Paraguay: betw Rio Apa & Rio Aquidaban, San Luis, Fiebrig 4417 (GH); Sierra
de Maracaju, Igatimi, Hassler 5511 (GH); Villarica, Jorgensen 4605 (MO).

Argentina: misiones: Iguazu, Puento Iguazii, de la Sota & Cuezzo 1556 (US) ; Posadas,
Ekmans.n., in 1907-1908 (MO).

See under P. paradiseae and P. ptilodon var. ptilodon for comments regarding
the affinities with var. robustum.

23d. Polypodium ptilodon var. pilosum A. M. Evans, var. nov.— Fig. 20.

A var. ptilodonte simile, frondibus approximatis vel fasciculatis, stipitibus
brevibus pilis numerosis longis acicularibus (interdum pilis brevioribus) praeditis,
rhachibus cum pilis eis stipitium similibus et paleis inconspicue fimbriatis praeditis,
lamina lineari-elliptica, segmentis apice rotundatis, basi symmetricis, infimis
numerosis reductis triangularibus, costis horizintalibus, pilis dissitis longis acicular-
ibus praeditis, venis 1-vel 2-furcatis, sporangiis cum 2 vel 3 inconspiuis setis
praeditis, sporis 47^ longis differt.

As in var. ptilodon except: rhizome long-creeping to suberect; fronds 49 (27-73)

260 ANNALS OF THE MISSOURI BOTANICAL GARDEN

cm long, approximate to fasciculate on the rhizome; stipe 2 (1-5) cm long; stipe
and rachis with numerous long acicular hairs 1.5-2 mm long, occasionally with
shorter, acicular hairs; rachis paleae with inconspicuously fimbriate margins;
blades linear-elliptic, 47 (26-70) cm long, 6.5 (3.5-9) cm wide; segments 32 (18-
48) mm long, 5.5 (3-9) mm wide, reduced to numerous pairs of triangular lobes at
the blade base, rounded at the apex, symmetrical at the base, the spaces between
segments less than one-half the width of the segment; costa perpendicular to the
rachis, with scattered long acicular hairs 1.5-2 mm long; veins once- or twice-
forked; sporangia with 2 or 3 short, inconspicuous, capsular setae up to 45^ long;
spores ca 47 ft long.

Holotype: British Guiana, Demerara, Essequibo River, Jenman s.n. NY.

Habitat: epiphytic or occasionally epipetric in humid forests, 400-1000 m alt.

Distribution: Trinidad: Blanchisseuse Rd, Hombersley 306 (US) ; Maracas, Hombersley
30 (US), 88 (US), Morne Bleu, Britton et al. (US).

British Guiana: Essequibo River, nr mouth of Onoro Creek, Smith 2781 (GH);
Shodikar Creek (Essequibo tributary), Smith 3010 (GH).

Venezuela: neuva esparta: Copey, Gines 3799 (US), miranda: Guinand Estate
(Cardenas), Siqui ■ 5949 (US), amazonas: Cerro Duida, Cano Negro,

Steyermark 57993 (US).

Peru: san martin: Tingo Maria, Allard 21541 (US), huanuco: Tingo Maria, Rio
Huallaga, Tryon 5334 (GH, US), junin: Santa Rosa, Pichis Trail, Killip & Smith 26201
(US).

Bolivia: la paz: San Carlos, Mapiri region, Buchtien 208 (US), 7072 (US), cocha-
bamba: Antahuacana, Buchtir, icrto Polonia, Rfo Coni, 14 km E of San

GH, MO).

Brazil: ceara: Sitio Pe da Ladeira, 3 km E of Guaramirango, Cutler 8324 (GH, MO).
rio de Janeiro: Ilha Grande, Rose 20350 (US).

This is the most distinct of the varieties of P. ptilodon because of the rounded
segments, the conspicuously longer hairs of the rachis and costa, and the very
short stipe. It is like the other varieties in the curious distribution of acicular hairs
only around the sorus (a character found only in this species) and in the gradual
reduction of the basal segments. Superficially it resembles P. consimile, but the
harsher texture, the trichomes of the lamina and costa, and the rhizome scales are
different.

24. Polypodium consimile Mett., Ann. Sci. Nat. (Paris) V, 2: 253, 1864.

Rhizome short- to long-creeping, 5 (3-7) mm in diam; rhizome paleae linear-
triangular to triangular, dark red-brown to blackish, lustrous, basifixed, attenuate
to acute, non-clathrate to sub-clathrate, slightly comose to comose with hairs
completely obscuring the paleae throughout the rhizome, entire; fronds 25-65 cm
long, close-spaced to approximate on the rhizome; stipe short to almost lacking;
rachis 6-20 times as long as the stipe; stipe and rachis light to dark red-brown,
with few acicular hairs up to 0.75 mm long and with inconspicuous ctenoid and
clavate hairs; rachis paleae inconspicuous, linear, entire; blades narrow-ovate, 23-
60 cm long, 4-15 cm wide, narrow-cuneate at the base; segments 20-75 cm long,
5-11 mm wide, perpendicular to the rachis, reduced to numerous short lobes at
the blade base, straight, herbaceous, obtuse to rounded, the base symmetrical,
expanded and confluent throughout, entire, the spaces between segments narrower

EVANS — POLYPODIUM 261

than the segments; lamina essentially glabrous, with occasional long, acicular
hairs and small scattered, clavate hairs; costa decurrent on the rachis, with
trichomes like those of the lamina; veins twice-forked, free; guard cells ca 43^ long;
sori inframedial, round, small, with simple and branched, clavate paraphyses up
to 160^ long; sporangia golden or brownish, with or without capsular setae; spores
reniform, tuberculate, ca 40^ long.

As here construed, this species consists of the two varieties discussed below.
1. Sporangia setose; rhizome paleae narrow-triangular, not obscured by dorsal

hairs; blade 6.5 (4-11) cm wide ...24a. P mil mimil

I. Sporangia without setae; rhizome paleae broad-triangular, completely obscured

by dorsal hairs; blade 12 (10-15) cm v, ide P consimik

24a. Polypodium consimile var. consimile.

P. consimile Mett. ex D. C. Eaton, Mem. Amer. Acad., n.s., 8: 198, 1860, nomen nudum.
(Based on: Venezuela, propre colonia Tovar, Fendler 220 B, GH, MO,' US)

P. consimile Mett. var. minus Hieron., Bot. Jahrb. Engler 34:519, 1904 (as "minor")
(Lectotype: Colombia, Tolima, Rio Ambica, 2000 m, Lehmann 2353 B fragment,'
isolectotypes LE, US. Syntype: Venezuela, Aragua, Colonia Tovar, Moritz 255 p p
These two collections were separated by Hieronymus as "Forma 1" and "Forma 2,"
without assigning formal names. The specimen of "Forma 1" is chosen lectotype)

P. pityrolepis Rosenst, Repert. Sp. Nov. 22: 16, 1925. (Holotype: Costa Rica, Rio Chris
Juan Vinas, 1200 m, 30 Mar 1910, Brade & Brade 694 S-PA; isotypes NY, US)
Rhizome paleae linear- to narrow-triangular, comose hairs not obscuring the

paleae; fronds 40 (25-60) cm long; rachis 12 (6-17) times as long as the stipe;

blades 36 (23-50) cm long, 6.5 (4-11) cm wide; segments 34 (20-55) mm long,

5 (4-9) mm wide; sporangia mostly with 2 capsular setae up to 80^ long.

Holotype: Colombia, Ocana, Norte de Santander, 6000-7000 ft. Schlim 633 B;

isotypes BR, G, L.

Habitat: mountain slopes, montane forest and ravines, terrestrial or on mossy

tree bases, 300-2300 malt.

Distribution: Hispaniola, Jamaica, Guatemala south to Colombia and Vene-
zuela.— Fig. 8.

Guatemala: el quiche: Chama, Johnson 206 (US), alta verapaz: Cubilquitz von
Tuerckheim s.n., Nov 1903 (US). '

Costa Rica: Cabeceras del Bkfs, Pittier 10572 (US); Cooper 6057 (MO), Cooper sn
Nov 1886 (GH). alajuela: Zarcero, Smith 48/32, 48/137, 48/195, 48/242, 48/247 (US)"
F72 (MO), san jose: La Hondura, Standley 37801 (US); La Palma, rd to La Hondura'
Scamman 7763 (GH); Las Nubes, Scamman 7229 (GH). cartago: Cartago, Cooper sn'
Rarre^R (GH ' M °' ^ Cen "° ^ 1& Carpintera> Standle V m ™> 35537 (US); Navar^

Panama:' chiriqui: Bajo Mona, mouth of Quebrada Chiquero, Woodson et al 1028
(US); El Boquete, Cornman 893, 1177, 1198 (US); "Camp I," Holcombs Trail, 10 mi 'above
El Boquete, Killip 5228, 5253 (US); above El Boquete, Rfo Panduro, Killip 5419 (US)
Roballo Trail, Rfo Piarnasta toward the Cordillera, Killip 5424 (US).

Dominican Republic: monte cristo: Las Rosas, Ekman s.n., 6 June 1926 (US)

Haiti: Furcy, Leonard 4611, 4728, 4776 (US); Massif de la Selle, Croix-des-Bouquets,
Ekman s.n., 2 Apr 1925 (US); Mission, Fonds Varettes, Leonard 4022 (US)

Jamaica: Clyde River, below Cinchona, Underwood 435 (US).

Venezuela: monagas: Cerro de Turumiquire, Tate 103, 104 (US) distrito federal-
Cordillera del Avila, above Caracas, betw Los Venados & Papelon, Steyermark 55070 (GH
MO, US), aragua: Colonia Tovar, Fendler 220 (B, GH, MO, US), 220B (GH US)

: santander: Rio Surata, above Surata, Killip & Smith 16647 (GH US)

Bogota, Ariste-Joseph (GH, US).

[Vol. 55
262 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Although superficially similar to P. ptilodon var. pilosum, this species differs
in the shorter rachis hairs, the lack of the stiff hairs around the sori, and the
segments with symmetrical bases.

There may be valid varieties in this species other than the var. pastazense
discussed below. The plants from Central America, represented by P. pityrolepis
Rosenst., tend to have a lighter brown, glabrous or subglabrous, narrower blade,
shorter stipes, and darker brown sporangia than the material from elsewhere in
the range. Venezuelan plants tend to have shorter, more clathrate, and less comose
rhizome scales, almost golden sporangia, and darker brown rachises and longer
stipes.

24b. Polypodium consimile var. pastazense (Hieron.) A. M. Evans, comb. nov.
P. pastazense Hieron., Hedwigia 48:257, t. 13, fig. 22, 1909. (Holotype: Ecuador, betw

Banos & Jivarfa de Pintuc in Pastaza Valley, Stuhel 1011 B)

Rhizome paleae short, triangular, dark red-brown, acute, non-clathrate, comose
with hairs completely obscuring the scales throughout the rhizome; fronds 54 (42-
65) cm long, spaced ca 1 cm apart on the rhizome; rachis 15 (12-20) times as long
as the stipe; stipe and rachis red-brown, glabrescent with short, acicular hairs;
blades 50 (40-60) cm long, 12 (10-15) cm wide; segments 60 (45-75) mm long,
9 (6-11) mm wide, obtuse at the apex; sporangia without setae.

Habitat: epiphyte in montane forests, 325-2300 m alt.

Distribution: Colombia and Ecuador. — Fig. 8.

Colombia: narino: betw Rio Miraflores & Rio San Martin, Volcan de Cumbal region,
Ewan 16157 (US).

Ecuador: napo-pastaza: Andes, Spruce 5260 (US) ; betw Puyo & Canelos, Mexia 6832
(US); Hacienda La Mascota, Canton Mera, Mexia 7016 (US), zamora-chinchepe: Cordil-
lera Cutucu, Camp E-1393 (US).

In the original description, Hieronymus noted the similarity between his species
and P. consimile. I believe that it is sufficiently distinct to maintain varietally,
because of the more robust aspect, the more nearly glabrous lamina, non-setose
sporangia, and particularly the complete obscuring of the rhizome scales by the
dorsal hairs. The spore and guard cell sizes do not suggest hybridity or polyploidy,
and it is probable that this is a local endemic variety just south of the broader
range of var. consimile and undergoing allopatric speciation.

25. Polypodium recurvatum Kaulf., Enum. Fil. 106, 1824.

P. externum Fee, Crypt. Vase. Bras. 1 : 85, t. 28, fig. 3, 1869, non Forst., 1786. (Holotype:
Brazil, Rio de Janeiro, Glaziou 2403 B)

P. paradisiastrum Fee, Crypt. Vase. Bras. 1: 90, t. 29, fig. 2, 1869. (Lectotype: Brazil, Rio de
Janeiro, Corcovado, Glaziou 974 P, isolectotype BR. Syntypes: Brazil, Rio de Janeiro,
Corcovado, Glaziou 396 P, at BR is P. pectinatiforme, Brazil, Therezopolis, Glaziou
1725 P. Glaziou 1725 at BR and C represent P. ptilodon var. robustum)

P paraguayense Baker, Jour. Bot. 310, 1878. (Type: Paraguay, forests nr base of Cerro
Telado, nr Villa Rica, Balansa 388 G)

P. pectinatum L. var. recurvatum (Kaulf.) Sodiro, Crypt. Vase. Quit. 335, 1893.

P. paradisiastrum f. crenulatum Rosenst., Hedwigia 46: 140, 1906 (as "crenulata"). (Holo-
type: Brazil, Santa Catarina, Lages, Spannagel 145 S-PA)

P paradisiastrum forma pectinatum Rosenst., loc. cit. (as "pectinata"). (Holotype: Brazil,
Rio Grande do Sul, Serra Joao, Rodriguez, liirgens 159 S-PA)

EVANS— POLYPODIUM 263

P. recurvatum var. minus Rosenst. ex HassL, Trab. Inst. Bot. Farm. Buenos Aires 45:69,

1928. (Holotype: Paraguay, Hassler 3990 G; isotype B)
P. recurvatum var. paraguayense (Baker) HassL, loc. cit.

P. recurvatum var. subbipinnatifidum Rosenst. ex Hassl., loc. cit. (Holotype: Paraguay,
Hassler 12241b G)

Rhizome long-creeping, 6 (5-7) mm in diam; rhizome paleae narrow-tri-
angular, light red-brown, non-lustrous, basifixed or basally cordate, acute, incon-
spicuously comose, non-clathrate, the margins inconspicuously papillate; fronds
53 (22-90) cm long, spaced ca 1 cm or less apart on the rhizome; rachis 2 (1.5-3.5)
times as long as the stipe; stipe and rachis red-brown, with scattered, short acicular
silvery hairs, ctenoid hairs absent; rachis paleae inconspicuous, filiform, entire;
blades triangular to narrow-triangular, 36 (16-65) cm long, 14 (10-24) cm wide,
truncate at the base; segments 72 (30-120) mm long, 4.5 (3-6) mm wide, perpen-
dicular to the rachis, occasionally slightly ascending, sometimes slightly shortened
at the blade base, straight or occasionally slightly falcate, often slightly constricted
toward the rachis, herbaceous or coriaceous, acuminate, expanded and symmetrical
at the base, entire (rarely crenate), the spaces between segments 1-3 times as wide
as the segments; lamina glabrous; costa perpendicular to the rachis, with scattered,
short, acicular hairs; veins twice-forked, free; guard cells ca 39^ long; sori medial,
round, with simple, clavate paraphyses; sporangia with 2 (1-5) short capsular setae;
spores reniform, tuberculate, ca 43^ long; n = 37.

Type: Brazil, Santa Catarina, Santa Catarina I, Chamisso s.n. B, LE.
Habitat: epiphytic, occasionally epipetric or terrestrial, in forest, 50-1000 m alt.
Distribution: southeastern Brazil and Paraguay.— Fig. 8.

Brazil: Campo da Lanca, Annies s.n. (Rosenstock 113b) (US), bahia: Toca de Onca,
% OSe , 2 ° n l°J (US) - RI ° DE J ANEIRO: Itatia y a > R °™ 20591 (US); Meio da Serra, Smith &
Brade 2289 (GH); Nova-Friburgo, Leite 4232 (MO); Organ Mts, Rose 20792 (US), Wilkes
U. S. South Pacific Exp. no. 15 (GH); Sumare, Brade 20607e (MO); Rio de Janeiro,
Monroe s.n., in 1864-70 (GH). sao paulo: Campos do Jordao, Leite 3570 (GH), Leite
3498 (MICH), Porto 3089 (MO). Parana: Desvio Ypiranga, Dusen 8337 (GH, MO, US).
santa catarina: Mun. Blumenau, Smith & Reitz 6287 (US), Hausa, Liidenvaldt 1.860 (US);
Horto Florestal I. N. P. Ibirama, Reitz & Klein 3090 hmalz 87 (MO);

Lages, Spannagel 5, 69, 118 (US); Marata, Porto Uniao, Reitz 4709 (US); Sao Bento,
Dutsch s.n. (Rosenstock 113a) (US); Sierra da Pedra, Ararangua, Reitz C279 (US) rio
grande do sul: Sao Salvador, Leite 2639 (209) (GH).

Paraguay: Villarrica, Jorgensen 4385, 4605 (MO), Hassler 8781 (B, G); Colonia
Presidente Gonzalez, Regnell I, A. 1813 (US).

This is a distinctive species because of its gray-green foliage, long-acuminate
segments, strongly truncate blade, and bright, inconspicuously comose, rust-colored
rhizome paleae. Its only really close relative is P. hygrametricum, which is smaller
in all respects and has only once-forked veins and more obtuse segments.

Many of the specimens from Paraguay have a more herbaceous texture and
wider and often deeply lobed segments. They are clearly P. recurvatum, although
several specific, varietal, and formal names for these have been created. Even
though the range of the Paraguayan materials might suggest it, these do not
represent intermediates between P. recurvatum and P. hygrometricum, so far as I

[Vol. 55
264 ANNALS OF THE MISSOURI BOTANICAL GARDEN

26. Polypodium hygrometricum Splitg., Tijdschr. Nat. Gesch. 7: 409, 1840.

P. pectinatum L. var. caliense Hieron., Bot. Jahrb. Engler 34:517, 1904. (Holotype:

Colombia, El Valle, Cordillera Occidental de Cali, nr Las Juntas del Dagua, Lehmann

7668 B; isotypes LE, US)
P. truncatulum Rosenst., Repert. Sp. Nov. 9:343, 1911. (Holotype: Bolivia, Antahuacana,

valley of Rio Espiritu Santo, Buchtien 2168 S-PA; isotype US)

Rhizome long- creeping, 4 (3-5) mm in diam; rhizome paleae narrow-tri-
angular, sometimes expanded and cordate at the bass, light red-brown, basifixed,
acuminate, inconspicuously comose, non-clathrate, the margins inconspicuously
papillate; fronds 27 ( 10-50) cm long, close-spaced to approximate on the rhizome;
rachis 7 (3-15) times as long as the stipe; stipe and rachis red-brown to dark
red-brown, pilose with short to long, simple, acicular, silvery hairs, ctenoid hairs
absent; rachis paleae inconspicuous, filiform, entire; blades 23 (8-40) cm long, 5
(2-9) cm wide, narrow-ovate to narrow-triangular, subtruncate to truncate at the
base; segments 25 (10-45) mm long, 4 (2-5) mm wide, perpendicular to the rachis
or slightly ascending throughout, narrow-ovate to linear-triangular, herbaceous,
obtuse to acute at the apex, expanded and symmetrical at the base, entire; lamina
pilose with simple, silvery, acicular hairs, clavate hairs absent; costa decurrent on
the rachis, brown or occasionally stramineous or blackish, pilose as on the lamina;
veins once-forked, free; guard cells ca 43^ long; sori medial, round, with short,
simple, clavate paraphyses; sporangia with 2 (1-5) acicular capsular setae; spores
reniform, tuberculate, ca 48^ long; n = 37.

Holotype: Surinam, Para, Splitgerber 1069 L.

Habitat: epiphytic, occasionally epipetric or terrestrial, in humid forests, 30-
1800 m alt.

Distribution: southern Mexico to western Bolivia.— Fig. 8.

Mexico: chiapas: Escuintla, Col. Cintalapa, Matuda 18392 (US), Esperanza, Matuda
18116 (US); Huixtla, Purpus 7226 (GH, US).

Guatemala: Xiiiana, Finca San Jose Nil, Hatch & Wilson 403 (US) ;

Pueblo Nusvo, Rojas 548 (US).

Honduras: colon: Claura, Spinden 38 (US).

Nicaragua: Managua: Sierras de Managua, Chaves 7 (US).

Costa Rica: Rio Poas, Pittier 2426 (US), limon: La Columbiana Farm, Standley
36896 (GH, US), heredia: La Selva, Rio Puerto Viejo, Scamman & Holdridge 8098 (GH),
Scamman 7518 (GH). san jose: El General, Skutch 2667 (MICH, MO, US); San Isidro
del General, Chrysler & Roever 5318 (MICH, US), cartago: Jesus Maria, Lankester 617
(US).

Panama: Aquarubia, Killip 2800 (US); Pecora, Killip 2703 (US), canal zone: Barro
Colorado I, Bailey 525 (US), Barbour Lathrop Trail, Wetmore & Woodworth 150 (GH);
betw Frijoles & Monte Lirio, I US); N of Frijoles, Standley 27449 (US);

Rio Indio de Gatun, Maxon 4815, 4849 (US), darien: Cana, Williams 931 (US). Panama:
Juan Diaz, Killip 2755, 2765 p.p. (US); Charave River, Chepo, Pittier 4721 (US); Pearl
Archipelago, Canyon Rd, Erlanson 414, 420 (US); Rio Merino, Erlanson 541 (US); below
pumping station, Johnston 383 (US), Area M, Johnston 71 (US).

British Guiana: Kanaku Mts, Mt Iramaikpang, Smith 3670 (GH).

Venezuela: nueva esparta: EI Valle, Margarita I, Miller & Johnston 163 (Gil, MO,
US), miranda: Parque Nacional de Guatopo, Steyermark 90178 (VEN). bolivar: El Mor-
rocoy, Guayapo, Bajo Caura, Williams 11807 (US); La Union, Medio Caura, Williams
11263 (US).

Colombia: magdalena: Codazzi, Haught 3760 (GH, US) ; Quebrada Soraria, 6 km E
of La Jagua, Haught 3615 (GH, US) ; Santa Marta, Rio Piedras, Smith 1028 (GH, MICH,
MO, US), santander: Barranca Bermeja, Magdalena Valley, betw Sogamoso & Colorado

EVANS — POLYPODIUM 265

Rivers, Haught 1424 (MICH, US), cundinamarca: Sasaima, Gonzalez & Daniel 1885 (US).
meta: jet of Guejar & Zanza Rivers, Smith & Idrobo 1499, 1506 (US), choco: Schott
1216 (MO); Lloro, 50 km S of Quibdo, jet of Rio Atrato & Rio Andagueda, Archer 2066
(US); S of Rio Condoto, betw Quebrada Guarapo & Mandinga, Killip 35138 (US) valle
del cauca: Las Juntas del Dagua, Lehmann 7668 (US); Rio Dagua, Quebrada del Rio
Blanco, Cuatrecasas 13671 (US); Rio Dagua, betw La Elsa & Rio Blanco, Killip 34734 (US).
"'► Santiago, above Pongo de Manseriche, Mexia 6216 (MICH); betw
■~ *- (lower Rio Huallaga basin), Killip & Smith 28164 (US).
" ', Killip & Smith 23608 (US).

'ana, ca 160 km NE of Cochabamba, Buchtien 2168

This species is easily recognized by its bright- or gray-green foliage, bright-
ferruginous rhizome scales, truncate blades, and silvery pilose pubescence. It
varies somewhat from north to south. Material from Central America is smaller,
pale- or lime-green, sometimes with a stramineous rachis or costa and more blunt
segments. The southern form is larger, more gray-green, and with more acute
segments. The hairs of the blade vary in length throughout the range but are
always characteristically silvery. Except for pubescence, which is most strongly
expressed in the middle of the range (i.e. northern South America), the variation
is evidently a cline ranging from smaller, more obtusely lobed plants in Central
America to taller, acutely lobed ones in Peru.

The allied species Polypodium recurvatum from Paraguay and Brazil differs in
being more glabrous and more robust and in having very long, almost acuminate
segments, but it is otherwise quite similar. It is isolated geographically from the
range of P. hygrometricum, and the differences between the two probably result
from allopatric speciation.

Species Dubiae et Excludendae
Polypodium bolivianum Rosenst. var. brevipes Rosenst, Repert. Sp. Nov. 12: 473,
1913 (Type: Bolivia, North Yungas, Polo-Polo, nr Corioco, 900 m alt, Buchtien
3497 S-PA?). This is probably a small specimen of P. bolivianum Rosenst.
P. inversum Veil., Fl. Flum. 11: 11, t. 72, 1827, nomen nudum; Arch. Mus. Nac.
Rio 5: 448, 1881. No Velloso types exist; the name must be based on the
description and illustration of a plant from Rio de Janeiro, Brazil. It is
usually considered a synonym of P. plumula (Morton, pers. comm.), but other
similar species in that region include P. dispersion, P. siccum and P. filicula,
P. lomariiforme Kunze, Linnaea 9: 42, 1834 (as "lomariaeforme") (Type: "in flor.
Peruv. montibus aridioribus as Cassapi 1829, Lect Diar. 1152, Herb. Poepp.,
Kze., etc."). I have seen a specimen at K labeled as having been sent to KunzJ
from Poeppig. It is dated 1830, but Mr. Morton (pers. comm.) indicates that
Poeppig's Peruvian collections are often misdated. Kunze's herbarium at
Leipzig is destroyed. The Kew specimen may be an isotype although Dr. Jar-
rett questions this (pers. comm.). There may be authentic material at Berlin.
The specimen agrees with the protologue except that the frond is pinnatisect
rather than pinnate, and it agrees with my P. camptophyllarium Fee var.
lachniferum (Hier.) Evans. If a satisfactory determination of type material
supports this, P. lomariiforme would take precedence.

[Vol. 55
266 ANNALS OF THE MISSOURI BOTANICAL GARDEN

P. maenurum Link, Hort Reg. Bot. Berol. Descr. 2: 96, 1833 (Type: none cited,
probably at B). The protologue compares it to P. paradiseae Langsd. & Fisch.,
and it has been considered a synonym of P. recurvatum.

P. otites L., Sp. PI. 1085, 1753 (Type: none in LINN; based on the illustration in
Petiver, Pteridographia 32, t. 1, fig. 16). Although this name has been as-
sociated with P. pectinatum L„ it is not a member of this complex, and is more
nearly allied with P. tenuifolium Humb. & Bonpl. ex Willd. [in L., Sp. PL,
ed. 4, 5: 185, 1810 (Type: Venezuela, Cumanacoa, Humboldt 437 B-photo)].

P. pectinatiforme var. brevipes Rosenst. in Buchtien, Contrib. Fl. Bolivia 1:38,
1910, nomen nudum.

P. pectinatiforme f. parvum Sehnen, Pesquisas, Bot. 10: 33, 1960. No type is cited
and the name is, therefore, invalidly published.

P. pectinatum var. acuminatum Baker, Jour. Bot. Brit. For. 25:25, 1887 (Type:
Costa Rica, Cooper K). The specimen is incomplete and cannot be determined
with confidence. It appears not to belong to this complex.

P. pectinatum L. var. baezanum Mille, Revista Col. Nac. Vic. Rocofuerte 9: 202,
1927 (Type: none cited; authentic material may be at B or in Ecuador). It
is not possible to place this name with confidence; it may be P. consimile Mett.

P. pectinatum L. var. bourgaeanum Fourn., Mex. PI. 76, 1872. (Syntypes: "In valle
Cordobensi, dec. sp. Bourgeau 1431, 1436; Guadalupa, Nova-Grenada Lindig
45; Brasilia."). No syntype has been seen, but the protologue indicates that
the plant has ellipsoid sori and anastomosing veins, neither of which would
place it in P. pectinatum L., and would tend to exclude it from the group

P. pectinatum L. var. brachypus Sodiro, Crypt. Vase. Quit. 334, 1893 (Type: "Crece
en los bosques de Gualea colectado por el Sr. D. Rodolfo Riofrio."). No type
has been seen; the description is inconclusive.

P. pectinatum L. var. hispidum Christ in Pittier, Prim. Fl. Cost. 3: 15, 1901 (as
"hispida") (Type: "El Paramo, 3000 m, versant E du massif de Buena
Vista, Jan 1897, Pittier 10474 P? or BR?). = Ctenopteris semihirsuta (Klotzch)
Copeland, Phil. Jour. Sci. 84: 450, 1955).

P. pectinatum L. var. jurgensii Rosenst, Hedwigia 43: 229, 1904 (Syntypes: Brazil,
Santa Cruz, auf Berg Joao Rodriquez, 200 m, Jurgens & Stier 82 S-PA?; Join-
ville, Santa Catharina, Schmalz 56 S-PA?). Schmah 56 at MO is not a mem-
ber of this complex, and other material has not been seen.

P. pectinatum. L. var. paradisiae Sodiro, Crypt. Vase. Quit. 334. 1893, non Baker,
1870 (Type: "Crece en la region tropical y subtropical, en lugares secos y
pedregosos; en la orilla del rio Pilaton."). This is presumed to be a new
variety rather than a transfer as P. paradiseae Langsd. & Fisch. is cited only
with a query, and does not occur in Ecuador. From the description, P.
bolivianum suggests itself.

P. venustum Desv., Gesell. Naturf. Freund. Berlin Mag. 5:315, 1811 (Type:
"Habitat in America calidiore Antillisque", Herb. Desv. P-photo). Although
Christensen (1906) referred this name to P. taxifolium L., the photo of the

EVANS — POLYPODIUM 267

type indicates that it is a eupolypodium. It would appear to be P. camptophyl-
larium Fee, over which it would take precedence, but I would have to see the
the specimen to be sure that it was not P. eurybasis or P. consimile.

Literature Cited
Alston, A. H. G. 1956. The subdivisi
Bell, P. R. 1959. The experimental h

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Bower, F. O. 1923. The Ferns (Filicales). Vol. I.

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Braithwaite, A. F. 1964. A new type of apogamy in ferns. New Phytol. 63: 293-305.
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Torreya 20: 76-83.
Chtarugi, A. 1960. Tavole cromosomiche delle Pteridophyta. Caryologia 13:27-150.
Ching, R. C. 1940. On natural classification of the family Polypodiaceae. Sunyatsenia

5:201-268.
Christensen, C. 1906. Index filicum. Copenhagen.

. 1928. On the systematic position of Polypodium vulgare. Dansk Bot. Ark. 5: 1-10.

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Copeland, E. B. 1947. Genera Filicum. Ronald Press, New York.

. 1956. Ctenopteris in America. Phil. Jour. Sci. 84: 381-475.

Dickason, F. G. 1946. A phylogenetic study of the ferns of Burma. Ohio Jour. Sci. 46: 73-

Dopp, W. 1939. Cytologische u. genetische Untersuchungen innerhalb der Gattung Druop-

teris. Planta 29: 481-533.
Erdtman, G. 1943. An introduction to pollen analysis. Chronica Botanica, Waltham.
Evans, A. M. 1964. Ameiotic alternation of generations: A new life cycle in the ferns.

Science 143: 261-263.
Fabbri, F. 1963. Primo supplemento alle "Tavole Cromosomiche delle Pteridophyta" di

Alberto Chiarugi. Caryologia 16: 237-335.
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meristematic tissue. Stain Technol. 9: 91, 92.
Hardin, J. W. 1957. A revision of the American Hippocastanaceae. Brittonia 9: 145-195.
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268 ANNALS OF THE MISSOURI BOTANICAL GARDEN

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EVANS — POLYPODIUM

tion of P. truncorum, de la Sota 21 (EVANS), showing only t
thick-walled cells of the inner ground parenchyma, X5.5. (]
de segment of P. ptilodon var. robustum, &
ing the mesophyll and the scarified bundle sheath, X80.
laminar hairs around the sorus in P. ptilodon var. ptilodc, ...
X6.5. (F) Ctenoid hairs of the rachis of P. ptilodon \ar j0 n 1470 (MICH)'

X6.5. (G) Rhizome palea of P. dispersum, Evans 2007 (MICH), X18. (H) Rachis palea
°1£ ^P^ 5 "^ Evans 2007 (MICH), X18. (I) Rhizome palea of P. cupreolepis, Pringle-
3353 (GH), X18. (J) Rhizome palea of P. plumula, Copeland 16059 (MICH), XI8.
(K) Palea from the rhizome apex of P. bolivianum, (US 1007896), X18. (L) Palea from
the side of the rhizome of P. bolivianum, (US 675781), X18

mates on the gametophyte of P. dispersum, Evans 2008 (MICH). (C) Stages of develop-
ment of the young gametophyte of P. ptilodon var. caespitosum, Evans 2002 (MICH).
(D) Mature gametophytes and heteroplastic leaf series of juvenile leaves of P dispersum
Evans 2008 (MICH). (E) Heteroblastic series of juvenile leaves of P. plumula, Evans
1187 (MICH). (F) Mature gametophytes and heteroblastic leaf seriss of juvenile leaves
of P. ptilodon var. caespitosum, Evans 2005 (MICH).

/«x ? g> R e P resent ative frond shapes. (A) P. ptilodon var. robustum, Pabst s.n. (MICH).
(B) P. paradiseae, Sehnen 2 (MICH). (C) P. curvans, Stork, et al. 10626 (MO) (D)
P. truncorum, Pabst s.n. (MICH). (E) P. dispersum, Evans 2008 (MICH). (F) P plumula
Evans 1187 (MICH). (G) P. recurvatum, Sehnen 4 (MICH).

Fig. 4. Epidermal cells, hairs and paraphyses of the lamina. Fig. A-G, Epidermal
cells. (A) P. dispersum (US 1287974). (B) P. filicula (US 1280298) (C) P siccum
(US 1516124) (D) P. truncorum, Mexia 5105 (GH). (E) P. hygrometricum, Chrysler
y Roever 5318 (MICH). (F) P. absidatum (US 1833997). (G) P. camptophyllarium
var. macedoi (US 2081770). (H) Transition of costal hairs to costal paleae in P. plumula
(US 1049810). (I) Variation in the ctenoid hairs of P. ptilodon var. caespitosum Mc
Nov. 4, 1930 (MICH): 1. Rachis; 2. Costa; 3. Lamina; 4. Receptacular para-
physes. Fig. J-P, Laminar hairs and paraphyses. (J) P. dispersum: 1. Capsular and re-
ceptacular paraphyses, Wagner 62061 (MICH); 2. Clavate hairs of the costa and lamina
(US 1287974); 3. Acicular hairs of the abaxial surface of the costa (US 1287974). (K)
P. filicula: 1. Receptacular paraphyses, Ahmuda & de la Sota 51 (EVANS); 2 Clavate
hairs of the abaxial surface of the lamina (US 1280298) ; 3. Acicular hairs of the segment
margin (US 1280298); 4. Acicular hairs of the abaxial surface of the costa (US 1280298).
(L) P. absidatum (US 1833997) : 1. Capsular setae; 2. Clavate hairs of the lamina- 3
Hairs of the abaxial surface of the costa; (M) P. camptophyllarium var. macedoi (US
2081770): 1. Clavate, short and long acicular and setose-clavate forked hairs of the lamina-
2. Acicular hairs of the segment margin; 3. Acicular hairs of the abaxial surface of the
costa; 4. Capsular setae and receptacular paraphyses. (N) P. venturii, Venturi 2970 (GH) ■
1. (Receptacular paraphyses; 2. Clavate and acicular laminar hairs. (O) P ptilodon var
robustum, de la Sota & Cuezzo 1556 (EVANS), Capsular seta and simple and branched

, Copeland 128 (MICH): 1. Clavate 1

- - > showing the range from the simple hair to thoss with
a multicellular subterminal expansion.

Fig. 5. Representative types of rachis paleae. (A) P. plumula from: 1. Jamaica,
Maxon et al. 1234 (GH); 2. Mexico Copeland 16055 (MICH); 3. Mexico (US 1745606);
4. Panama, Taylor 1390 (MICH); 5. Colombia (US 1692356); 6. Peru (US 2356738)-
7. Brazil (US 762365). (B) P. dispersum, Evans 2007 (MICH). (Q P. atrum Lundell
6639 (US). (D) P. bermudianum, Brown 464 (GH). (E) P. singer i Ahmuda & de la
Sota 53 (EVANS). (F) P. ferrugineum, Copeland 125 (MICH). (G) P cupreolepis
Pringle 3353 (GH). (H) P. ptilodon var. caespitosum (FLAS-P1718). (I) P filicula
Ahmuda & de la Sota 51 (EVANS). (J) P. dispersum (FLAS-P1155, FLAS-P4879). (K)
P. ptilodon var. caespitosum (FLAS-P4881). (L) P. plumula (FLAS-PI153, FLAS-P5166).

[Vol. 55
270 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 6. Venation patterns and segment shapes. Most of the segments lack the ex-
panded basal portion, M and N lack medial portions, and O and S x lack terminal portions.
(A) P. truncorum, Mexia 5105 (GH). (B) P. siccus (US 1516124). (C) P. filicula (US
1280298). (D) P. cupreolepis, Pringle 3353 (GH). (E) P. sursumcurrens, Copeland 128
(GH). (F) P. bermudianum (sterile segment with portion of a fertile segment) (US
848332). (G) P. dispersum, Evans 2007 (MICH). (H) P. hygrometricum, Chrysler &
Roever 5318 (MICH). (I) P. plumula, Maguire 25005 (GH). (J) P. atrum, Stoutamire
3563 (MICH). (K) P. ptilodon var. caespitosum, McFarlin s.n., 4 Nov. 1930 (MICH).
(L) P. pectinatiforme, Copeland 13601 (MICH). (M) P. bolivianum (US 675781). (N)
e var. pastazense (US 2254236). (O) P. recurvatum, Copeland 21561 (MICH).
(P) P. venturii, Venturi 2974 (GH). (Q) P. camptophyllarium var. maceodi (US 1081770).

Fig. 7. Habitat studies. (A) P. dispersum, on mossy sandy soil, Evans 1189 (MICH),
xy 12 . (B) P. plumula, on Quercus virginianum trunk, Evans 1187 (MICH), xy 12 . (C)
P. dispersum, small proliferations from an exposed root mass, Evans 1189 (MICH), X%
(D) P. singeri, short rhizomes probably originating as root proliferations along a small
twig, arrows mark the individual rhizomes, Gerdez 54 (S-PA), xy 6 . (E) P. ptilodon
var. caespitosum, growing on a rotten log, Evans 2004 (MICH), X% 2 . (F) P. dispersum,
root growing through the hole in the bottom of the pot, the
narked with an arrow, Evans 1130 (MICH), X'/,,.

: small non-soriferous plants, presumed to have orig-
inated entirely from root proliferations, Evans 2016 (MICH), X% 4 .

Fig. 8. Geographic distribution of species in the Polypodium pectinatum-plumula
complex, (as represented by herbarium specimens). (A) Geographic distribution of P.
ferrugineum, P. hygrometricum, P. recurvatum. (B) Overall distribution of the species
of the complex; major and marginal areas are distinguished according to the key on the
map. (C) Geographic distribution of P. bolivianum, P. consimile var. consimile & var.
pastazense, P. singeri. (D) The five areas of distribution of the individual species of the
complex as indicated by the key accompanying the map.

Fig. 9. Geographic distribution of species in the Polypodium pectinatum-plumula
complex, (as represented by herbarium specimens). (A) P. plumula, P. sursumcurrens.
(B) P. atrum, P. dispersum, P. filicula.

Fig. 10. Geographic distribution of species in the Polypodium pectinatum-plumula
complex, (as represented by herbarium specimens). (A) P. alfredii, P. paradiseae, P.
r , ime , P. pectinatum, P. venturii. (B) P. camptophyllarium var. abbreviation,
var. camptophyllarium, var. lachniferum & var. macedoi, P. cupreolepis, P. siccum.

Fig. 11. Geographic distribution of species in the Polypodium pectinatum-plumula
complex, (as represented by herbarium specimens). (A) P. ptilodoi

var. pilosum, var. ptilodon, var. robustum. (B) P. absidatum, P. ch *

P. eurybasis var. eurybasis, var. glabrescens & var. villosum, P.

Fig. 12. Stages of sporogenesis in Polypodium ptilodon var. caespitosum, Evans 1157
(MICH). (A) Cytokinesis in the 8 pre-mother cells, X160. (B) 16 spore mother cells,
X160. (C) Meiosis in the 16 spore mother cells, X130. (D) 74 pairs of chromosomes at
meiotic metaphase in a spore mother cell, X760. (E) Young spore tetrads with four nuclei,
before cytokinesis, X160. (F) Young tetrads during cytokinesis, X240. (G) Young tetra-
spores X240. (H) Mature spores, showing long-reniform shape and tuberculate surface,
X760.

Fig. 13. Stages in sporogenesis in Polypodium dispersum, Wagner 62061 (MICH).
(A) Mitosis in the eight pre-mother cells, X160. (B) Cytokinesis in the eight pre-mother
cells, X160. (C) 16 spore mother cells, X160. (D) Mitosis in the 16 spore mother cells,
X160. (E) 111 univalent chromosomes at metaphase in the spore mother cells, X760.
(F) Cytokinesis in the 16 diads of young diplospores, X160. (G) Mature spores, one
without a scar, the other with a monolete scar, X760. (H) 32 young diplospores, X160.

EVANS POLYPODIUM

Fig. 15. Gametophyte of Polypodium dispersum, Evans 2008 (MICH). (A) Ger-
minating spore, X120. (B) Formation of the cell plate and branching of the thallus, X120
(C) Young apogamous sporophytes proliferating from gamstophytes on agar plan
sporophytes are indicated by arrows, X4. (D) Two-and-one-half -month-old gameto-
phytes showing a portion of an apogamous sporophyte and the position of stomates on
the gametophyte (marked with an arrow), X24. (E) Three stomates from gametophyte
in D, X 120. (F) Young apogamous sporophyte showing simple and branched clavate
hairs, X50. (G) Young apogamous sporophyte from small elliptical gametophyte, X20.

Fig. 16. Specialization in the Polypodium pectinatum-plumula complex. (A) Rhizome
long-creeping (0) to short-creeping or erect (1); (B) Rhizome paleae narrowly triangular
(0) to cordate (1); (C) Rhizome paleae non-comose (0) to comose (1) to comose hairs
completely obscuring the paleae (2.5); (D) Rachis brown (0) to black (1); (E) Rachis
stiff (0) to elastic (1); (F) Rachis straight (0) to spiral , (G) Rachis gla-

brous or thinly pilose (0) to densely villose (1); (H) Ctenoid hairs absent (0) to present
(1); (I) Rachis paleae filiform (0) to non-filiform (1); ally 25 cm to 150

cm (0) to less than 25 cm (1); (K) Frond stipitate (0) to sessile (1); (t) Base of the
blade subtruncate or cuneate (0) to definitely truncate (1); (M) Basal segments per-
pendicular to the rachis (0) to occasionally deflexed (0.5) to regularly deflexed (1); (N)
Segments perpendicular to the rachis (0) to ascending (1); (O) Perpendicular segment
base symmetrical (0) to asymmetrical when the segment is perpendicular to the rachis
(1); (P) Segment margin entire (0) to crenate (1) to pinnatifid (1.5); (Q) Lamina glabrous
(0) to pubescent (1); (R) Pubescence evenly distributed (0) to localized (1); (S) Lateral
veinlets forked (0) to simple (1); (T) Sori medial (0) to marginal (1); (U) Receptacular
paraphyses simple (occasionally forked) and with uniform cells (0) to: U 1 paraphyses
with a subterminal irregular multicellular expansion (1) or: U 2 paraphys. i I

branched— one branch tipped with a clavate cell and the other tipped with a setose cell
(1); (V) Spores up to 50 mu long (0) to over 50 mu long (1); (W) Spores 64 per spo-
rangium (0) to 32 per sporangium (1).

Solid dots represent extant taxa; hollow dots represent hypothetical ancestral inter-
mediates. The character letter is given the first time the specialized condition appears

(0.5); capital

Fig. 18. Type specimens of new taxa. Left. P. atrum A.
(US 1638286). Right. P. camptophyllarium var. abbreviatum i
(US 1342112), xy 4 .

Fig. 19. Type specimens of new taxa. Left. P. eurybasis var. villosum A. M. Evans,
Fosberg 19688 (US 2290492). Right. P. ptilodon var. pilosum A. M. Evans, Jenman s.n.,
in 1897 (NY), xy 4 .

Fig. 20. Type and representative specimens. Left. Type of P. absidatum A. M. Evans,
Killip & Smith 18518 (US 1353919). Right. "
L. s.s., Evans 2587 (TENN), xy 4 .

272 ANNALS OF THE MISSOURI BOTANICAL GARDEN

,*/; Z

< # u

Fig. 1. Morphology of the sporophyte (legend p. 2

-^W fwlf*'

274 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 3. Representative frond shapes (legend p. '<

EVANS — POLYPODIUM

Fig. 4. Epidermal cells, hairs and paraphyses of the lamina (legend

276 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 5. Representative types of rachis paleae (legend p. 269),

EVANS — POLYPODIUM

278 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 7. Habitat studies (legend p. 270).

Fig. 8. Geographic distributions (legend p. 270).

280 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 9. Geographic distributions (legend p. 270).

■fXH

t^t

[

riA

- v^

'WL-^m-

■ \

•I

J

<kJ

y

r'W-^

isMs^y

<

£- — -

lyyT^^.

Fig. 10. Geographic distributions (legend p. 270).

Fig. 11. Geographic distributions (legend p. 270).

EVANS POLYPODIUM

' ■'•■ -**-*

k J ~** B

3 H

Fig. 12. Sporogenesis (legend p. 270) .

ANNALS OF THE MISSOURI BOTANICAL GARDEN

EVANS — POLYPODIUM

o

i>

Fig. 14. Spores (legend p. 271).

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 15. Gametophyte (legend p. 271).

Fig. 16. Legend p. 271.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 17. Type specimens (legend p. 271).

Fig. 18. Type specimens (legend p. 271).

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 19. Type specimens (legend p. 271).

EVANS— POLYPODIUM

Fig. 20. Type and representative specimens (legend p. 271)

29Z ANNALS OF THE MISSOURI BOTANICAL GARDEN

INDEX OF LATIN NAMES
New taxa are in boldface type, all other taxa are in roman type;
boldface type refer to descriptions, numbers in roman type refer
numbers with dagger (t) refer 1

Adenophorus 195t

Alsophila 2021

Amphoradenium 195t

Anemia subg. Coptophyllum 213t

Asplenium 195t; aethiopicum 206t

Cheiropleuria 210t

Ctenopteris 208t, 221 1, 222; glaziovii
220, 221; plumula 230; semihirsuta
266; truncorum 220

Cyathea 225t

Dipteris 210t

Goniophlebium 208t; pectinatum 246

Hemitelia 202t

Isoetes 206t

Matonia 210t

Monarda 213t

Nephrolepis 196t

Ophioglossum 206f

Platycerium 195t

Polypodium absidatum 199t, 212t,
215t, 2191, 238-239, 2401; alfredii
196t, 198t, 199t, 200t, 210t, 212t,
214t, 218t, 219t, 224f, 225-227, 227t,
228t, 232t, var. curtii 225; atrum
1951, 196t, 198t, 199t, 212f, 218t, 226t,
229t, 237t, 237-238; auritum 223;
bakeri 220, 221; bermudianum 198t,
199t, 202t, 212t, 218t, 228-229, 232t,
237t, 238f; bolivianum 195t, 196f,
198t, 210t, 212t, 215t 3 218t, 241-
243, 243t, 265, 266, var. brevipes
265; camptophyllarium 196t, 198t,
220t, 249t, 251-256, 265, 267, var. ab-
breviatum 212t, 215f, 220t, 251t,
253f, 256, var. camptophyllarium
200t, 201t, 210t , 212f, 248f, 252t, 252-

254, 254t, 255t, 256t, var. lachnife-
rum 200t, 210t, 212t, 252t, 253t, 254-

255, 256t, 265, var. macedoi 200t,
201t, 212t, 216t, 251t, 253t, 254;
carpinterae 241; chnoophorum 217f;
choquetangense 196t, 197t, 199t,
202t, 212t, 214t, 215t, 219t, 240t,
241; cinerascens 252; circinatum 239;

h M I96t, 2001, 215t, 219t,
260f, 260-262, 266, 267, var. consi-
mile 202t, 212t, 26 It, 261-262, 262t,

195t, 196t, 197t, 198t, 199t,202t,221t,
212t, 214t, 218t, 224-225, 227t,
228t, 232f; curvans 196t, 197t, 199t,
200t, 202f, 212t, 214t, 215t, 219t,
239f, 239-240, 241t; curvatum 239t,
239, 240; cyathicola 225, 226t; dis-
persum 195t, 196t, 197t, 198t, 199t,
200t, 201t, 202t, 203t, 203-208t, 210t,
21 If, 212f, 216f, 218f, 223f, 226t,
2281, 229t, 232t, 233t, 234t, 235-
237, 238t, 248t, 249f, 265; elasticum
229, var. filicula 233, var. glaziovii
220; elegans 229, 233, 235t, eurybasis
198t, 199f, 215t, 219t, 243t, 243-245,
267, var. eurybasis 212t, 243t, 243-
244, 245f, var. glabrescens 212t,
243f, 244f, 244-245, 245f, var. vil-
losum 212f, 214t, 243t, 245t, 245;

21 It, 212t, 214t, 215f, 218t, 225t,
227, 228t, 232t; filicula 195t, 196t,
197t, 198t, 199t, 200f, 203t, 212t, 214t,
215t, 216t, 218t, 223t, 232f, 233-
234, 265; glaziovii 221; hartwegianum
217t; heteroclitum 222; hygrometri-
cum 197t, 201f, 202t, 210f, 212f,
217t, 218t, 263f, 264-265; inversum
265; lachniferum 252, 254, var. gla-
brescens 244, var. glabrescens f. in-
curvatum 244; lomariiforme 265;
lonchitidis folio 246; macedoi 254;
naenurum 266; microsorium 235, 249,
249t; molle 233t, 235, 237t; morit-
zianum 217t; nigrum tenerius sectum
246; otites 233f, 246, 266; paradiseae
199t, 200t, 2 lOt, 212t, 215t, 220t,
250t, 250-251, 2f "

262,

212t, 261f,

i 261, i

pectinatum 2
pastazense 262; patzcuarense 225 1";
mm tii «mc 195t, 199t, 200t, 212t,
220t, 223, 235, 249-250, 262, var.
brevipes 266, var. hirsutum 249, 250t,
f. parvum 266; pectinatum 1951",
198t, 200t, 201f, 202t, 203t, 208f,
210t, 212t, 217t, 220t, 240t, 242t,
246-248, 251t, 254f, 258, 266, var.
i, var. aurita 223,

EVANS — POLYPODIUM

Ml.Ul.ni

230,

truncatum 249, var. wagneri 246;
pityrolepis 261, 262t; plumula 195t,
196t, 197t, I98f, 199t, 200t, 20 It,
202t, 203t, 208t, 210t, 21 It, 212f,
214t, 218t, 225f, 226t, 227t, 228t,
229t, 229-233, 234t, 237 1, 265;
ptilodon 194t, 196t, 198t, 200t, 202t,
216t, 219t, 242f, 248f, 25 It, 256-
260, var. caespitosum 196t, 197f,
200t, 201t, 202f, 203t, 204t, 210t,
21 If, 2121, 229t, 254t, 257t, 258,
var. pilosum 212t, 215f, 257t, 259-
260, 262t, var. ptilodon 197t, 210t,
211t, 212t, 257t, 257-258, 2591, var.
robustum 212f, 249t, 25 It, 257t,
259, 262; pulchrum 225, 230, var.

minus 230; recurvatum 194t, 196t,

197t, 202t, 210t, 212t, 215t, 217t,

218t, 262-263, 265t, 266, var. minus

paraguayense 263, sub-

263; robustum 259,

cinerascens 252, 259, var. cin-

f. minus 259; schkuhrii 229, \
siccum 195t, 196t, 200t, 202t, :
210t, 212t, 216t, 219t, 222f, ;

223, 265;

212t, 2191

squamatum 195t, 215t;
198t, 200t, 201t, 202t, 212t, 214f,
215t, 216t, 219t, 220t, 228; tabla-
zianum 225, 226t, taxifolium 266;
tenuifolium 233t, 266; truncatulum
264; truncorum 194t, 195t, 196t, 197t,
198t, 199t, 200t, 202t, 208t, 212t,
216t, 219t, 220-222, 223t, venturii
212t, 220t, 248-249; venustum 266;
virginianum 201t, 216t; vulgare 195t,
21 It, 215t; wagneri 246
Trichomanes proiferum forma B 206t

PALYNOTAXONOMIC STUDY OF THE PHYTOLACCACEAE 1

by Joan W. Nowicke 2
Department of Botany, Washington University, St. Louis, Missouri

The Phytolaccaceae, a largely tropical and subtropical
utilizing pollen morphology in addition to floral
species, including the novelty Ercilla syncarpellata Nowicke,
of pollen and gross morphology; keys are provided to all major taxi
genera recognized (Anisomeria, Ercilla, Phytolacca, Gallesia, Segui
stigma, Schindleria, Hilleria, Petiveria, Ledenbergia, Monococcus, Agdestis, Microtea,
Lophiocarpus, Stegnosperma, and Barbeuia) are placed in six subfamilies (Phytolaccoideae,
Rivinoideae, Agdestioideae, Microteoideae, Stegnospermoideae, and Barbeuioideae) three
of which are newl all subfamilies except the Rivinoideae (with Seguierieae

and Rivineae) being monotribic. Four major pollen types are recognized: 3-colpate with
variations of minor os formation or polar exine thickening; pantoporate; 12-colpate with
four colpi forming a square at each pole and four perpendicular to the equator; and 15-
colpate with five colpi forming a pentagon at each pole and five perpendicular to the
equator. In terms of gross morphology the family is considered relatively primitive with
only occasional examples of characters considered advanced. In terms of pollen morphology,
the family has advanced types (those other than 3-colpate) well represented.

Introduction

The Phytolaccaceae comprise a weedy family of largely tropical and subtropical
plants which have been placed, almost without exception, in the order Centro-
spermae (Chenopodiales of Hutchinson, 1959; Caryophyllales of Bessey, 1915).
It is a relatively natural order, best characterized by its uniform placentation, ovule
structure, perisperm storage tissue and the unique presence of betacynanins.

Heimerl (1889), in his survey of the Phytolaccaceae for the Pflanzenfamilien,
recognized six tribes: Rivineae [Gallesia Casar., Seguieria Loefl., Monococcus F.
Muell., Phaulothamnus A. Gray, Ledenbergia Klotzsch, Rivina L., Petiveria L.,
Microtea Swartz, Hilleria Veil, (as Mohlana Mart.), Adenogramma Reichb.],
Limeae (Polpoda Presl, Limeum L., Barbeuia Thouars), Stegnospermeae (Stegno-
sperma Benth., Psammotropha Eckl. & Zey.), Phytolacceae (Phytolacca L., Aniso-
meria D. Don, Giesekia L.), Gyrostemoneae (Didymotheca Hook, f., Gyrostemon
Desf., Tersonia Moq.), Agdestideae (Agdestis Moc. & Sesse). He did not recognize
Trichostigma A. Rich., Ercilla Juss. (Ercilia), or Schindleria H. Walter as distinct
genera, and Lophiocarpus Turcz. was not treated but rather included in the
Chenopodiaceae.

Walter's (1909) monograph was the first comprehensive treatment of the
family and he recognized two subfamilies, Phytolaccoidei

1 Based on a dissertation submitted to the Graduate School of Washington University
in partial fulfillment of the requirements for the degree of Doctor of Philosophy. Assisted
by a Junior Fellowship from the Center for the Biology of Natural Systems and by National
Science Foundation Grant No. GB-5042 (Principle Investigator, Walter H. Lewis). Dr.
Lewis is gratefully acknowledged for supervision of this research.

2 Currently Postdoctoral Fellow, Missouri Botanical Garden and Department of Botany,
Washington University, St. Louis, Mo.

Ann. Missouri Box. Gard. 55(3): 294-363, 1969.

NOWICKE— PHYTOLACCACEAE 295

(Anisomeria, Ercilla, Phytolacca, Barbeuia, Didymotheca, Tersonia, Gyrostemon,
Codonocarpus A. Cunn., Hilleria, Seguieria, Gallesia, Rivina, Trichostigma, Leden-
bergia, Schindleria, Petiveria, Monococcus) , Stegnospermoideae with only Stegno-
sperma, Agdestis which is not placed in a subfamily or tribe, as well as three other
anomalous genera (Achatocarpus Triana, formerly placed in the Amaranthaceae
by Bentham & Hooker, 1883; Microtea, Phaulothamnus) with affinities to the
Chenopodiaceae for a total of 22 genera. He removed six genera (Limewn, Giesekia,
Adenogramma, Psammotropha, Polpoda, Semonvillea Gay, the last genus Heimerl
had treated as a subgenus of Limeum) to the subfamily Ficoideae of the Aizoaceae.
In addition to this major change, he recognized Ercilla as distinct from Phytolacca,
Trichostigma as distinct from Rivina, and included his recently established genus,
Schindleria (Walter, 1906).

In the second edition of Pflanzenfamilien, Heimerl (1934) divided the Phyto-
laccaceae into five tribes: Rivineae, Phytolacceae, Agdestideae, Stegnospermeae,
and Barbeuieae, the last three being monogeneric; in addition he cited two genera,
Microtea and Lophiocarpus, as connecting links to the Chenopodiaceae. He re-
duced the total number of genera to 17 by removing six previously included by
Walter (1909) and adding Lophiocarpus, a genus formerly placed in the Cheno-
podiaceae (Bentham & Hooker, 1883), and placed by N. E. Brown (1909) in the
Phytolaccaceae as congeneric with Microtea. Four of the genera which Heimerl
separated from the Phytolaccaceae comprised the Gyrostemonaceae (Gyrostemon,
Codonocarpus, Didymotheca, Tersonia) all of which have unisexual flowers and
a high carpel frequency (rarely two or one). The Achatocarpaceae was constructed
for two dioecious genera, Achatocarpus and Phaulothamnus, found in the American
tropics and subtropics.

Hutchinson (1959) included the Phytolaccaceae in the Chenopodiales; the
latter consists of 10 families of which four resulted from a further division of the
Phytolaccaceae. The family was consequently reduced to three genera: Phytolacca,
Anisomeria, and Ercilla. Agdestis and Barbeuia were placed in monotypic families
and the remaining genera, with the exception of Stegnosperma which he placed as
a monogeneric family in the Pittosporales, comprise the Petiveriaceae (Gallesia,
Hilleria, Ledenbergia, Lophiocarpus, Monococcus, Microtea, Petiveria, Rivina,
Schindleria, Seguieria, Trichostigma).

Eckardt (1964) made some changes in the larger taxa, recognizing three sub-
families, the Phytolaccoideae with four tribes, and the Stegnospermatoideae and
the Microteoideae each with a single tribe. He does not cite all genera, but his
treatment appears to follow closely that of Heimerl (1934).

The present work is the first major treatment and revision of the Phytolaccaceae
in the generic concepts of Heimerl (1934). Although Walter's (1909) monograph
is a competent treatment of the family, several serious flaws exist: the familial
limits, i.e. inclusion of genera whch have since, justifiably, been removed; and the
fact that in many instances, generic as well as specific, the exsiccatae listed could
not possibly have permitted an adequate consideration of the wide variation which
the family, with the result that many of the taxa established are

296 ANNALS OF THE MISSOURI BOTANICAL GARDEN

invalid. This study, although incomplete in parts, attempts to coordinate morpho-
logical descriptions, including pollen, floral and vegetative, in order to revise the
generic and specific limits.

Synopsis of the Phytolaccaceae
Phytolaccaceae Lindl., Nat. Syst. ed. 2, 210, 1836.
Petiveriaceae Link, Handb. 1: 392, 1829 3 .

Herbs, shrubs, or trees. Leaves simple, alternate, petiolate to =t sessile, entire,
generally estipulate (stipules thorny in Seguieria). Inflorescences racemes, spikes,
or irregular panicles rarely cymules. Flowers small, perfect or rarely unisexual
(plants then dioecious), ± actinomorphic (weakly zygomorphic in Hilleria and
Anisomeria) ; calyx composed of 4-5 free or slightly connate segments, dry and
inconspicuous or occasionally corolla-like; corolla absent (staminodia petaloid in
Stegnosperma) ; stamens 3-oo (number variable within a species or even an
inflorescence), frequently arranged in one or two whorls, sometimes deposited on
a hypogynous disc but placed irregularly in relation to sepals or rarely alternate,
the filaments linear, or awl-shaped, the anthers mostly linear, tetrasporangiate,
introrse (extrorse in Hilleria), dehiscing longitudinally; ovary superior (semi-
inferior in Agdestis), composed of 1-16 free or united carpels, each carpel with
one basal, campylotropous ovule, the styles usually equal to the carpel number
or absent, the stigmas capitate or ± penicellate or not apparent. Fruit a berry,
capsule, drupe, utricle, achene or samara; seed one per carpel; embryo curved
around a mealy perisperm.

A family of 17 genera and ca 70-80 species mostly in the New World tropics
and subtropics, but also found in Africa, Australia and Hawaii.

Economically the family is of little importance, but some species contain
partially toxic substances which are used medicinally. The roots and fruits of some
contain Saponin, which can be utilized as a soap. Phytolacca dioica L. is frequently
planted as a shade tree in the tropics because it is fast growing. The berries of
some Phytolacca spp. have been utilized as an adulterant of red wine, and the
young sprouts and leaves can be made into a "poke salad".

Pollen

Pollen grains single, prolate, subprolate or prolate spheroidal, ca 16-35/* (E)
X ca 18-39^ (P), 3-colpate, 3-colporoidate, 12-colpate in a 4-4-4 pattern, 15-colpate
in a 5-5-5 pattern, or pantoporate, exine ca 1.5-3^ in thickness, sometimes thickened
at the poles to 5//, sexine ± equal to or slightly thicker than nexine and sparsely
small spinulose to ± smooth (see Appendix).

Cytology

Relatively few chromosome counts are available for the family, but all evidence
points to a base of x = 9 (18).

Hilleria latifolia 2n = 36 (Mangenot & Mangenot, 1958).

H. Walter

Petiveria alliacea L. 2n = 72 (Sugiura, 1937).

Phytolacca acinosa 2n = 36 (Sugiura 1936b).

Roxb.

P. americana L. 2n - 36 (Bostick, 1965, N. G;

Lewis et al., 1962, Texas).

P. australis Phil. n = 18 (Heiser, 1963).

P. dioica L. 2n = 36 (Schnack & Covas, 1947).

P. octandra L. 2n = 36 (Sugiura, 1936a).

Rivina humilis L. 2n =•- 108 (Nowicke, 1967; Sugiura, 1936a).

Anatomy

The entire order Centrospermae is characterized by its distinctive stem struc-
ture (essentially anomalous secondary thickening) and according to Metcalfe &
Chalk (1950) certain genera in the Phytolaccaceae (Agdestis, Anisomeria, Barbeuia,
Gallesia, Petiveria, Phytolacca, Seguieria) have successive rings of vascular bundles
in the inner parenchymatous portion of the pericycle. Concentric rings of xylem
and phloem occur in sufficiently thick stems of Ercilla, Gallesia, Phytolacca, Rivina
and Seguieria.

Another distinctive feature of the Phytolaccaceae, as well as the entire order,
is found in the character of its pigments. It is one of the "beta-cyanin families"
(Dreiding, 1961), which are closely related and characterized by their inability to
produce anthocyanins which is replaced by the ability to synthesize betacyanins
(and betaxanthins).

Taxonomy

In this treatment I recognize six subfamilies, all monotribic with the exception
of Rivinoideae which is divided into two tribes based primarily on the striking
differences in fruits and types of inflorescences.

I. Phytolaccoideae H. Walter— ovary of 3-16 carpels, free or united; fruit a
drupe, achene or berry. 3 genera: Anisomeria, Ercilla, Phytolacca.

II. Rivinoideae Nowicke— ovary of one carpel and one seed; fruit an achene,
drupe, utricle or samara. 9 genera: Gallesia, Seguieria, Rivina, Trichostigma,
Schindleria, Hilleria, Petiveria, Ledenbergia, Monococcus.

III. Microteoideae Eckardt ex Nowicke — ovary of one carpel with 2-4 stigmas
and one seed; fruit an achene. 2 genera: Microtea, Lophiocarpus.

IV. Agdestioideae Nowicke— ovary of 3-4 carpels, 3-4 stigmas, semi-inferior,
and one seed. One genus: Agdestis.

V. Stegnospermoideae H. Walter — ovary of 3-5 united carpels, 3-5 seeds,
petaloid staminoidia; fruit a capsule. One genus: Stegnosperma.

VI. Barbeuioideae Nowicke — ovary of 2 united carpels, 2 seeds; fruit a capsule.
One genus: Barbeuia.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Key to
a. Fruit(s) a capsule, 2 or 3-5 locular.

b. Capsule 2-locular; inflorescences axillary cymules or fascicles; staminodia
absent; plants drying black; endemic to the Malagasy Republic

bb. Capsule 3-5 locular; inflorescences racemes, sometimes cymules; staminodia
petaloid; plants drying green; Central America and the West Indies

subf. V Stegnospermoideae (p. 356)

aa. Fruit (s) a berry, drupe, samara, achene, or utricle.

subf. VI Barbeuioideae (p. 358)

c. Ovary of 3-16 carpels, free or united.

d. Ovary semi-inferior, with 3-4 united carpels and 3-4 stigmas; seed one;

± woody vines; leaves cordate subf. IV Agdestioideae (p. 355)

dd. Ovary superior, 3-16 free or united carpels and one stigma per carpel; seeds
as many as the carpels; herbs, shrubs or trees; leaves variable, but not

cordate - subf. I Phytolaccoideae (p. 298)

cc. Ovary of one carpel, with one seed.

e. Ovary with one stigma; fruit a samara, drupe or utricle, or if an
achene then not globose, and conspicuously 4-6 hooked or covered with
recurved spines subf. II Rivinoideae (p. 320)

ee. Ovary with (2-) 3-4 stigmas; fruit an achene, globose with pericarp

wrinkled, glochidiate, warty or ridged subf. Ill Microteoideae (p. 346)

PHYTOLACCOIDEAE
I. Subf. Phytolaccoideae H. Walter, Pflanzenr. IV, 83 (Heft 39) : 29, 1909. (Type

Phytolacca L.)
Tribe Phytolacceae Reichb., Fl. Exc. 586, 1832. (Type Phytolacca L.)
a. Carpels distinctly free; sepals unequal and ± fleshy; leaves succulent-leathery;

inflorescences mostly terminal 1- Anisomeria

aa. Carpels free or united; sepals equal or only weakly unequal and thin; leaves
not succulent-leathery; inflorescences axillary or terminal,
b. Carpels usually free, rarely united; inflorescences dense, short, axillary

Chile .

bb. Carpels free or united, usually united; inflorescences racemes or ± long

spikes; mostly herbs; cosmopolitan 3. Phytolacca

1. ANISOMERIA
Anisomeria D. Don, Edinb. New Phil. Jour. 13: 238, 1832. (Type A. coriacea D.

Don)
Pircunia Bertero, Mercurio Chileno 744, 1829; Amer. Jour. Sci. 23:264, 1833, non Moq.

(in DC, Prodr. 13(2) :29, 1849).

Herbs or shrubs, sometimes succulent, calcium oxalate crystals present. Leaves
alternate or in fascicles of ca 3, ovate, ovate-elliptic, or ± spatulate, mucronate,
retuse or rounded, entire or slightly undulate, the bases rounded to attenuated,
glabrous, ± leathery; sessile to petiolate, petioles sometimes thickened at the base
in fascicular arrangement and appearing stipular. Inflorescences spikes or spike-like
racemes, mostly terminal. Flowers perfect, ± zygomorphic; sessile or pedicellate;
bract single or absent; bracteoles 2 and fleshy, or absent; sepals 5, unequal, ±
united at the base, orbicular, fleshy; stamens 10-20, appearing in two whorls, the
filaments ± thickened; ovary 5-8 carpellate, free, the styles as many as the carpels,
the stigmas inconspicuous to slightly thickened. Fruit a loose collection of drupelets
(?), red to brown; seed one (Fig. 1).

This genus of three species has been described as restricted to Chile; however,

some locations cited on herbarium specimens are from the Chilean-Argentinean

Pollen grains single, prolate, ca 26[i (E) X ca 34-39^ (P), 3-colpate, colpi
ca 23-32/* long, the exine ca 2-3 fi in thickness, sexine ± equal to or slightly thicker
than nexine and finely reticulated (Fig. 4).

The publication of Bertero (1829) does predate that of Don, but the descrip-
tions of the former author are seminude and until I see collections of Bertero
which have definitely been determined by him as Pircunia drastica, and thus leave
no doubt as to the plant described in the Mercuric- Chileno publication, I think
the generic name Anisomeria should remain.

Two names have deliberately been omitted from the following synonomy,
namely, Anisomeria coriacea var. petalifera H. Walter (Pflanzenr. IV, 83 (Heft
39) : 32, 1909) which may well be a variety of A. fruticosa Phil, judging from the
thickened filaments of the latter; however, since I have not seen petaloid specimens,
I withhold judgment. The other name, A. densiflora H. Walter (loc. cit.), does
not apply in my opinion to a distinct species, but could, because of the immaturity
of the inflorescence, be placed in synonomy under either A. coriacea D. Don or A.
fruticosa. I tend to favor the former reduction because the leaves of the type
(Lechler s.n. photo F, from Bj") resemble those of A. coriacea, which agrees with
Walter's (1909) description.

Fig. 1. Inflorescences of Anisomeria D. Don. A, A. littoralis (Poepp. & Endl.) Moq.
(XI); B, A. coriacea G Don (Xi/ 2 ); C, A. fruticosa Phil, mature and immature inflores-
cences. A after Grandjot s.n. (MO); B after Grandjot s.n. (MO); G after Werdermann

300 ANNALS OF THE MISSOURI BOTANICAL GARDEN

The genus is clearly related to Phytolacca and is distinguished by its rather
weak zygomorphic condition, fruit type and, to a greater or lesser extent, its general
habit.

1. Anisomeria coriacea D. Don, Edinb. New Phil. Jour. 13: 238, 1832. (Type

Cuming s.n. G?)
Pircunia drastica Bertero, Mercurio Chileno 744, 1829; Amer. Jour. Sci. 23: 264, 1833.

, rrtcro) Poeppig & End!., Nov. Gen. Sp. PI. 1: 26, pi 43, 44, 1835.

Shrubs, weak, or succulent herbs with woody bases. Leaves mostly alternate,
ovate, rarely lanceolate or spatulate, mucronate, undulate, the bases attenuate, up
to 6 cm long and 3 cm wide, succulent to leathery; petiole indistinct. Inflorescences
spikes, up to 25 cm long, terminal. Flowers sessile; bract single, ca 2-2.5 mm long,
lanceolate; bracteoles absent (?); sepals 5(-6), ± unequal, rounded, ca 3 mm long
and 3-4 mm wide; stamens ca 20, in two irregular whorls, the filaments linear,
ca 2.5-2.9 mm long, the anthers ca 1.7-1.9 mm long; ovary 5-6 carpellate, free, the
styles as many as the carpels and ca 1.5-2 mm long, the stigma on the upper
surface. Drupelets 5-6, red-brown, ca 8 mm long, style ± persistent (Fig. IB).

Chile 4 : coquimbo: s. lot, Gay s.n., (G, NY). Santiago: nr Juncal, Elliot 631 (K); vie
of Santiago, Grandjot s.n., in 1932 (MO), without province: Reed s.n. (K); Bridges
526 (K).

Pollen grains ca 39^ (P), colpi ca 32^ long, exine ca 3[i in thickness, sexine
somewhat thicker than nexine.

Pollen examined: Grandjot s.n. (MO).

2. Anisomeria littoralis (Poepp. & Endl.) Moq. in DC, Prodr. 13(2): 25, 1849.
Phytolacca chilensis Miers, Trav. 2 : 532, 1826, nom. nud.

Pocpp. & Endl., Nov. Gen. Sp. PL 1 : 27, pi. 45, 1835. (The plate is taken as

someria chilensis Miers ex H. Walter, Pflanzenr. IV, 83 (Heft 39) : 33, 1909, non Phy-
tolacca chilensis (Miers ex Moq.) H. Walter [Pflanzenr. IV, 83 (Heft 39): 45, 1909].
Shrubs. Leaves mostly fasciculate, ovate-elliptic, obtuse, mucronate or retuse,
the bases obtuse or rarely attenuate, up to 5 cm long and 2 cm wide,
succulent to leathery; petioles to 1.5 cm long, becoming swollen and woody at
the base. Inflorescences racemes (appearing as a spike in bud), mostly terminal,
up to 7 cm long. Flowers with pedicels up to 6 mm long at maturity; bract ab-
sent; bracteoles two, ca 0.6-0.7 mm long, located ca midway on pedicel; sepals
5, unequal, somewhat united at the base, ca 2 mm long, and ca 0.6-2 mm wide,

4 The geographical citation procedure is: large countries of S. America, the province
or state, the locat" . . ' .- - • ■ ■ - tically by collector; Mexico and the United States, the
state, then alpha I- > « III .-III tor; large countries of Africa, Australia, and Asia, the

location or province, then alphabetically by collector; all others, alphabetically by col-

NOWICKE— PHYTOLACCACEAE 301

± orbicular; stamens 10-20, the filaments ca 2 mm long, stout, the anthers ca
1.2 mm long; ovary 5-8 carpellate, free, the styles as many as the carpels, thick,
short, strongly recurved, the stigma on the upper surface. Drupelets 3-5, green-
brown, ca 8-9 mm long (Fig 1A).

Chile: coquimbo: vie of lower Choros River, Reed s.n. (K): estate of Frai loree.
Munoz B161 (GH), B218 (GH); Skottsberg & Skottsberg 763 (F, NY). Santiago- vie of
Santiago Grand/ot s.n., in 1935 (MO). Valparaiso: Algarrobo, Kausel 4339 (F); Quillota,
tiertero 1263 (MO).

Pollen grains ca 34^ (P), colpi ca 23^ long, exine ca 2-2.5^ in thickness,
sexine ± equal to nexine.

Pollen examined: Grandjot s.n. (MO); Skottsberg & Skottsberg 763 (NY)
(Fig. 4).

It is unfortunate that Walter (1909) applied the nude name Phytolacca
chilensis Miers to two different taxa, a species of Anisomeria and one of Phytolacca.
Since it is impossible as yet to determine to which entity Miers referred, even
though I have seen Miers collections of Anisomeria, I have applied it to a bona
fide species of Phytolacca.

3. Anisomeria fruticosa Phil., Linnaea 29: 38, 1857-1858. (Type Philippi 873

photo F, from Bf)

Shrubs. Leaves mostly alternate, ovate, rounded to retuse, entire, the bases
rounded to obtuse, up to 6 cm long and 4 cm wide, ± leathery; petioles to 12
mm long, the bases ± thickened and woody. Inflorescences spike-like racemes,
up to 20 cm long, mostly terminal. Flowers with pedicels ca 2-2.5 mm long at
maturity; bract absent; bracteoles two, ca 1 mm long; sepals 5, slightly united
at the base, unequal, ± orbicular, ca 2.5-3 mm long and 0.6-2.2 mm wide; stamens
10-20, the filaments ca 2 mm long and ca 0.8 mm wide, fleshy, the anthers ca
1 mm long; ovary 6-7 carpellate, free, the styles as many as the carpels, short,
thick and recurved, the stigmas on the upper surface. Drupelets 6-7, up to 1 cm
long (Fig. 1C).

yet Z^LT?™?^ Vic ° f Tdta1 ' 1° hnston 519 5 (GH), 5611 (GH); Werdermann
785 (F, K, MO, NY).

Pollen grains ca 34^ (P), colpi ca 24-25^ long, exine ca 2-2.5^ in thickness,
sexine ± equal to nexine.

Pollen examined: Werdermann 785 (MO).

2. ERCILLA
Ercilla A. Juss., Ann. Sci. Nat. 25: 11, 1832; Edinb. New Phil. Jour. 14: 261, 1833.

[Type E. spicata (Bert.) Moq.]
Suriana Domb. & Cav. ex D. Don, loc. cit. 13: 238, 1832.
Bridgesia Hooker & Arnott in Hooker, Bot Misc. 3: 168, pi. 102, 1833, pro parte, non B

incisifolia Bertero, (Bertero 1361 photo NY, from G).
Ercilia Endl., Gen. Sp. PI. 977, 1840.
Apodostachys Turcz., Bull. Soc. Nat. Hist. Moscou 21(1): 577, 1848.

Shrubs. Leaves ovate to ovate-orbicular to ovate-elliptic, acute to retuse-
mucronate, entire to very finely undulate, the bases slightly cordate to obtuse, ±

302 ANNALS OF THE MISSOURI BOTANICAL GARDEN

papery to coriaceous, glabrous; petiolate. Inflorescences spikes or spike-like racemes,
mostly axillary, rarely terminal, densely flowered. Flowers sessile to shortly pedicel-
late; bract single; bracteoles 2, closely appressed to sepals; sepals 5, ± equal
free, ovate to ± elliptic, turning black on dessication; stamens 8-12, free, inserted
on a hypogynous disc; ovary 4-8 carpellate, free or united, the styles as many as
the carpels and free, slender, the stigmas not apparent. Fruit a loose collection
of drupelets (?) or berry (?); seeds 1-7.

Chile.

Pollen grains single, prolate, ca 19-20/* (E) X ca Zip (P), 3-colpate, colpi
ca 20-22/1 long, exine ca 2p in thickness, sometimes thickened at the poles to ca
2.5-3^, sexine ± equal to nexine and sparsely small spinulose.

Heimerl (1934) stated that Ercilla spicata (Bertero) Moq. is the sole species
of the genus, for E. volubilis Juss. scarcely differs from it. Walter (1909) recognized
both and separated the two by leaf texture, viz. coriaceous in E. spicata and papery
in E. volubilis, and by the sunken condition of the midvein in the former. Both
leaf texture as well as shape vary widely in the genus, although there are, ad-
mittedly, some very leathery-leaved specimens. Walter's description of E. volubilis
is based' on a single collection, Domhey 944, which he cites as being from Peru,
and which Harms in a footnote (Heimerl, 1934) states as being a doubtful loca-
tion for the genus, a conclusion with which I agree, especially since the NY speci-
men of this collection is marked Chile. This particular sheet of Dombey 944 is
unusual in that it may represent a mixed collection since the inflorescences of
the two specimens are somewhat different, the one on the right having spike-
like racemes, the left one having shorter spikes. For the present I am recognizing
two species based on carpel condition, i.e. free or united, and am treating all free
carpellate specimens as E. spicata.

aa. Carpels united, the styles free 2. E. syncarpellata

1. Ercilla spicata (Bertero) Moq. in DC, Prodr. 13(2): 35, 1849. (Type Dombey

944 G, NY)
Galvezia spicata Bertero, Mercurio Chileno 642, 1829.

nlis A. Juss., Ann. Sci. Nat. 25 : 11, pi. 3, fig, 1, 1832.
Suriana volubilis (A. Juss.) Domb. & Cav. ex D. Don, Edinb. New Phil. Jour. 13:238,

1832.
Bridgesia spicata Hooker & Arnott in Hooker, Bot Misc. 3: 169, pi. 102, 1833. (Syntypes:

Bridges s.n. BM; Cuming 349 K; Mathews 244, not seen)
Apodostachys densiflora Turcz., Bull. Soc. Nat. Moscou 21(1): 577, 1848. (Type Bridges

s.n. BM)
Phytolacca volubilis (A. Juss.) Heimerl, Pflanzenfam. 3(lb): 11, 1889.

Shrubs. Leaves up to 8 cm long and 5.5 cm wide; petioles to 1.5 cm long.
Inflorescences mostly spikes, rarely spike-like racemes, up to 10 cm long. Flowers
mostly sessile, rarely with pedicels to 1 mm long; bract single, ca 2.2-2.8 mm
long, ± elliptic; bracteoles two, ca 1.7-2.4 mm long; sepals ovate, ca 3.5-5 mm
long; stamens 8-10(-12), the filaments 5-6 mm long, the anthers ca 1.6 mm long;
ovary 4-8 carpellate, free, the styles ca 1.2-1.6 mm long, recurved. Drupelets 4-8,
reniform, ca 4-5 mm long; seed ca 3.5-4 mm long, testa shiny black.

NOWICKE — PHYTOLACCACEAE 303

Chile: concepcion: vie of Concepion, Macrae s.n. (K). maule: Cauquenes, Joseph
1822 (US), o'higgins: Donihue, Bertero 289 (GH). valdivia: Chinguil, Hollermayer 1922
(NY, US); Coral, Buchtien s.n. (US); Panguipulli Lake, Joseph 2405 (US); s. loc, Philippi
s.n. (US). Valparaiso: s. loc, Cuming 349 (K); Bertero 289 (GH). without province:
Gay 362 (NY, US), s.n. (K, NY, US); Dombey 944 (NY); Joseph 4398 (US).

Pollen grains ca 20/* (E), colpi ca 20^ long, exine thickened at the poles to
ca 2.5-3^ in thickness.

Pollen examined: Gay 362 (NY).

2. Ercilla syncarpellata Nowicke, sp. nov.

Inflorescentiae spicato-racemosae; flores pedicellis ca 1.5 mm longis; antherae
ca 1.5-1.6 mm longae; ovarium brevi gynophoro, 5-7 carpellatum, carpellisque
connatis.

SJvubs. Leaves ovate-elliptic, acute-mucronate, entire, the base obtuse to
slightly cordate or oblique, up to 7 cm long and 3 cm wide, ± coriaceous, petioles
up to 6 mm long. Inflorescences spike-like racemes, up to 8.5 cm long, mostly
axillary. Flowers with pedicels ca 1.5 mm long; bract single, ca 2-2.8 mm long,
awl-shaped; bracteoles two, ca 1-1.3 mm long; sepals 3-4 mm long; stamens ca
10, the filaments ca 5-6 mm long, the anthers ca 1 mm long; ovary on short gyno-
phore, ca 0.5 mm long, 5-7 carpellate, united, the styles separate or closely ap-
pressed at the bases, recurved, the stigmas on the upper surface. Fruit unknown.

Holotype: Chile. Valdivia: La Aguade, Gunckel 1837 (MO), 15 Nov 1930;
isotype GH.

Chile: valdivia: Coral, Gunckel 62 (BM); Krause s.n. (US).

Pollen grains ca 19^ (E), colpi ca 22^ long, exine not noticeably thickened
at the poles.

Pollen examined: Krause s.n. (US).

3. PHYTOLACCA
Phytolacca L., Sp. PI. 441, 1753. (Type P. americana L.)
Phytholacca Brot., Fl. Lusit. 2: 224, 1804.
Sarcoca Rat, FI. Tellur. 3: 55, 1836.

Pircunia Moq. in D.C, Prodr. 13(2): 29, 1849, non Bertero (Mercuric chileno 744, 1829).
Trees, shrubs or herbs; calcium oxalate crystals ± conspicuous. Leaves lanceo-
late to ovate or rarely ± deltoid, acute, acuminate, mucronate, retuse-mucronate,
entire, rarely finely undulate, the bases rounded, obtuse, or attenuate, rarely ±
oblique, ± glabrous; subsessile or petiolate. Inflorescences racemes or spikes,
rarely racemes with thyrsiform bases, axillary or terminal, ± scurfy to pubescent.
Flowers perfect, or unisexual and plants dioecious, mostly actinomorphic; sessile
or pedicellate; bract single, rarely absent, narrowly-lanceolate to awl-shaped;
bracteoles two, rarely absent, lanceolate to awl-shaped; sepals 5, rarely 4 (-9),
narrow-oblong, oblong, elliptic, oblanceolate or ovate, white to red; stamens func-
tional or rudimentary, 8-25, deposited irregularly in one or two whorls, generally
on a hypogynous disc, the filaments sometimes widened at the bases; ovary ab-
sent, rudimentary or functional, sometimes on a short gynophore, 5-16 carpellate,
free or united, the styles as many as the carpels, the stigmas on the upper surface.

304 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fruit a berry, ribbed, or a loose collection of drupelets (or achenes ?); seeds 5-16
for berry, one for drupelet, reniform, testa shiny black.

Cosmopolitan in distribution but principally in the warmer regions.

Pollen grains single, prolate spheroidal, subprolate, or prolate, ca 17-29/* (E)
X ca 25-36> (P), 3-colpate, colpi ca 17-26// long, exine ca 2-2.5^ in thickness,
sometimes slightly thickened at the poles, sexine ± equal to nexine, rarely 2X as
thick, and finely reticulated to sparsely small spinulose (Fig. 7).

Without exception, Phytolacca is the most difficult genus in the Phytolac-
caceae in which species can be accurately defined or classified. The problem lies
principally with the fact that many taxa hybridize readily (Fassett & Sauer, 1950;
Sauer, 1951), thus obscuring the characters by which they are recognized. In ad-
dition infraspecific variability and even extensive variation in the same specimen,
e.g. inflorescences in which the flowers may have either one or two staminal whorls,
contribute to the difficulty. An illustration of the type of problem encountered
in species differentiation is the occurence of inflorescences which are thyrsiform
at the base— a character supposedly best delimiting two South American species,
P. thyrsiflora Fenzl. and P. sanguinea H. Walter. However this same character
can be observed to a disturbing extent in P. americana L., to a lesser degree in
P. heterotepah H. Walter and even in specimens collected in Asia.

The best characters to delimit taxa in this problematical genus should of
course be qualitative, but it appears that many of these parameters are under weak
genetic control. Heimerl (1934) noted that the approximately 35 species described
are in part very closely related and are difficult to distinguish. In my opinion
there are far less than 35 distinct species of Phytolacca; many names have un-
doubtedly been assigned to specimens of hybrid origin.

A discussion of all species which have been described is not feasible here, and
I am, for the most part, revising and amending the species in the sense of Walter's
(1909) treatment.

Walter (1909) placed 26 species of Phytolacca into three subgenera based
on the degree of connation of the carpels: free, connate at the base with the apices
free, or completely united carpels. Subgenus 1 Pircunia (Moq.) H. Walter con-
tains P. heptandra Retz, P. esculenta van Houtte, P. acinosa Roxb., P. latbenia
(Buch.-Ham.) H. Walter, and P. cyclopetala H. Walter in the sect. 1 Pircuni-
astrum Moq., characterized by hermaphroditic flowers, and P. dodecandm L'Herit,
P. goudotii Briq., and P. nutans H. Walter in the sect. 2 Pircunioides H. Walter,
characterized by dioecious plants. Subgenus 2 Pircuniopsis H. Walter, characterized
by carpels connate at the base with the apices free, contains a hermaphroditic
group, the sect. 1 Pircuniophorum H. Walter, with three species, P. chilensis
(Miers ex Moq.) H. Walter, P. sanguinea H. Walter, and P. rugosa A. Br. &
Bouche, as well as the sect. 2 Pseudolacca Moq., with two dioecious species P.
dioica L. and P. weberbaueri H. Walter. Subgenus 3 Euphytolacca Moq., the
largest group and characterized by carpels completely united contains a very large
hermaphroditic sect. 1 Phytolaccastrum H. Walter with P. thyrsiflora Fenzl, P.
heterotepala H. Walter, P. brachystachys Moq., P. americana L., P. polyandra
Batalin, P. rivinoides Kunth & Bouche, P. meziana (J. D. Smith) H. Walter, P,

NOWICKE — PHYTOLACCACEAE 305

micruntha H. Walter, P. australis Phil, P. octandra L., P. pwpurascens A. Br.
& Bouche, and P. icosandra L., and a monotypic dioecious sect. 2 Phytolaccoides
H. Walter containing P. pruinosa Fenzl.

The degree of carpel connation upon which Walter (1909) based the three
subgenera functions well for recognition of the subg. Pircunia, with its completely
separate carpels, a condition which is unmistakable in fruit, but for distinguish-
ing the subg. Pircuniopsis, with carpels connate at the base, from subg. Euphyto-
lacca, with carpels completely united, it is a difficult character. However, in view
of the fact that I am not monographing the genus and do not wish to add further
to the taxonomic and nomenclature! confusion by making an unmeaningful
change, I think it best at this time to maintain the subgenera as outlined by Walter.
The dioecious species are a unique problem, not only in subgeneric classifica-
tion of male specimens, but also because some species have female flowers with
normal appearing stamens. These may be mistaken for functionally perfect flow-
ered species.

A useful, if not essential, criterion for classifying material of Phytolacca is
geographic location. It is unfortunate, however, that so many binomials seem to
be based primarily on this feature; in some instances location is the key to rapid
and accurate identification, e.g. Cyprus and P. pruinosa. A knowledge of the
species commonly found in a particular region can also aid in recognition of hy-
brids. Specimens from the northern regions of South America are the most per-
plexing, due to the large number of species found there and to the almost con-
tinuous variability of their characteristics.

In this treatment I am recognizing 20 species, but in some instances do so
with misgivings. Most of the latter involve specimens where geographic location
is apparently the only feature which distinguishes them as species. Examples
are P. brachystachys from the Hawaiian Islands, which is very similar to P. rugosa
of Central and South America, and P. purpwascens from Haiti, which may very
likely be a hybrid of P. icosandra and P. americana or P. rivinoides.

Following Walter's (1909) infrageneric classification in his subg. 1 sect. 1,
I recognize P. heptandra, but have reduced P. esculenta to P. acinosa. To my
knowledge, I have not seen P. latbenia or P. cyclopetala, but this does not deny
their existence; they are unfortunately omitted. In his dioecious sect. 2 I recognize
P. dodecandra, yet doubt the validity of P. goudotii and P. nutans.

In his subg. 2 sect. 1, I recognize P. sanguinea, P. rugosa, and P. chilensis.
In the dioecious sect. 2, I recognize both species cited by Walter, P. dioica and P.
weberbaueri, even though I have not included a description of the latter because
I have seen only a male specimen. I am also including here P. tetramera Hauman-
Merck, which Walter neglected.

In the subg. 3 sect. 1, I follow Walter's treatment rather closely, with the
following exceptions: P. polyandra I have not seen and omit; P. heterotepala and
P. meziana I include, but with reservations that they may be of hybrid origin;
and P. micrantha and P. australis are reduced to P. bogotensis H.B.K. In the
monotypic dioecious sect. 2, I recognize P. pruinosa.

306 ANNALS OF THE MISSOURI BOTANICAL GARDEN

In accordance with the Code, I have changed the subg. 3 Euphytolacca sect.
1 Phytolaccastrum, which contains the type species, to the subg. Phytolacca, sect.
Phytolacca.

aa. Carpsls ± united.

d. Carpels with apices ± free, styles not conni\, bg. 2 Pircuniopsis

e. Flowers perfect sect. 1 Pircuniophorum

f. Inflorescences 4. P. sanguined

mm long; carpels ca 9-10 16. P. purpurascens

nn. Flowers mostly with one staminal whorl.

r. Plants of Hawaii; carpels mostly 5-6 17. P. brachystachys

rr. Plants of North or South America; carpels mostly 7-10.

, Inflorescences ± open racemes; plants of North

America 19. P. am

ate sect. 2 Phytolo

NOWICKE— PHYTOLACCACEAE 307

Subg. 1 Pircunia (Moq.) H. Walter, Pflanzenr. IV, 83 (Heft 39): 38, 1909. (Type

P. acinosa Roxb.)
Pircunia Moq. in DC., Prodr. 13(2): 29, 1849, non Bertero (Mercuric, chileno 744, 1829).

Sect. 1 Pircunia
sect. Pircuniastrum Moq. in D.C., Prodr. 13 (2): 29, 1849.

1. Phytolacca heptandra Retz., Obs. 6: 29, 1791.
P. striata Hoffm., Comm. Getting. 12: 27, 1796.

" " i DC., Prodr. 13(2): 30, 1849, nom. nud. pro syn. Pircunia striata

var. residiformis.

iud. pro syn. Pircunia stricta var. residiformis.
i Ecklon & 7 "

Pircunia stricta Moq., loc. cit. (Type Ecklon & Zeyher s
P. stricta var. resediformis Moq., loc. cit. (Type Ecklon & Zeyher s.n. P)
P. stricta var. latifolia Moq., loc. cit. (Type Drege s.n. G, not seen, in IDC Micro-Edition,
Candolle Prodromi Herbarium)

Herbs, slender, to ca 1 m. Leaves lanceolate-elliptic, rounded-mucronate,
entire, the bases obtuse, up to 10 cm long and 2.5 cm wide; ± sessile to petioles
1 cm long. Inflorescences spikes or spike-like racemes, up to 13 cm long, sparsely
flowered. Flowers ± sessile or with pedicels to ca 3-4 mm long; bract single, ca
1.4-1.6 mm long, linear-lanceolate, bracteoles two, ca 1 mm long, lanceolate; sepals
5, somewhat united at the bases, rounded, ca 2.5-3 mm long and 1.3-1.5 mm wide;
stamens 6-7, in one whorl, the filaments ca 1.5-2 mm long, widened at the bases,
the anthers ca 0.8 mm long; ovary 5-7 carpellate, free, the styles ca 0.8 mm long,
± straight. Fruit a collection of 5-7 drupelets, each 2-3 mm long.

South Africa: Albert Dist, Cooper 1358 (K); 8 mi from Greytown, Wylie 28018
(MO); Somerset East, MacOwen 1453 (F, NY); s. loc,, Cooper 366 (K, NY), Ecklon &
Zeyher s.n. (MO).

Pollen grains prolate spheroidal, ca 26^ (E) X ca 27^ (P), colpi ca 17^ long,
exine ca 2fi in thickness, sexine ± equal to nexine and sparsely small spinulose.

Pollen examined: MacOwan 1453 (NY).

2. Phytolacca acinosa Roxb., Hort. Bengal. 35, 1814.

P. esculenta Van Houtte, Fl. Serres 4: 398 B, 1848.

Pircuinia esculenta (Van Houtte 236, 1853-54.

Phytolacca kaempferi A. Gray, Mem. Amer. Acad. n.s. 6: 404, 1858. (Type Small s.n. GH)

P. pekinensis Hance, Jour. Bot. 7: 166, 1869. (Type Williams 12648, location unknown)

Herbs, stout, ca 2-3 m. Leaves ovate, ovate-elliptic, or rarely ovate- lanceolate,
acute to ± mucronate, entire, the bases attenuate to slightly rounded, up to 35 cm
long and 16 cm wide, ± glabrous; petioles to 6 cm long. Inflorescences racemes, up
to 30 cm long, mostly axillary, rarely terminal, the peduncle ± smooth to scurfy.
Flowers with pedicels 5-13 mm long, slightly winged at the base in some; bract
single, (l-)2-4 mm long, lanceolate to awl-shaped; bracteoles two, ca 1.5 mm
long; sepals 5, ± unequal, 3-4 mm long and 1.8-2.3 mm wide, ovate to rounded;
stamens 7-15, sometimes in ± two whorls, the filaments ca 1.8-2 mm long,
widened at the bases, the anthers ca 0.8-1 mm long; ovary sessile or on short
gynophore, ca 0.5 mm long, 6-9 carpellate, somewhat united at the base in flower,

[Vol. 55
308 ANNALS OF THE MISSOURI BOTANICAL GARDEN

mostly free in fruit, the styles ca 0.6-1 mm long, mostly straight, slightly recurved
at the tip. Fruit an assemblage of drupelets, each up to 4 mm long.

Asia.

China: changyang: Wilson 873a (K). chihli: Wang 20194 (NY), hupeh: Chow 532
(NY); Henry 4351 (MO), kansu: Potanin s.n. (K). kiangsu: Tsu 255 (K, MO), kwang-
tung: Tso 20891 (NY), lungchow: Morse 576 (NY), shantung: Chiao 2609 (F).
szechan: Henry 5511 A (K). yunnan: Forrest 5989 (K); Henry 10705 (NY), 10705A (MO).
without province: Henry 2045 (K); Licent 1467 (K).

Japan: Albrecht s.n. (GH, NY); Arimoto s.n. (MO); Dickens s.n. (K, NY) s.n. (NY);
Higg s.n. (NY) ; Oldham 671 (GH, K); Wright s.n. (NY).

S. Korea: Yongso/c 8063 (F).

India: Assam: Ward 8453 (K). himachal pradesh: Gammie 18587 (K). Kashmir:
Stewart 7873 (MO), punjab: Jain & Bharadwaja s.n. (NY), sikkim: Watt 5709 (K).
without province: Griffith 4360 (K).

Pakistan: Stewart s.n. (NY).

Pollen grains subprolate, ca 23^ (E) X ca 28^ (P), colpi ca 23^ long, exine
ca 2.5fi in thickness, sexine ca 2X as thick as nexine or ± equal to nexine and
finely reticulated.

Pollen examined: Henry 10705 (MO).

This Asian group of very robust herbs with conspicuously free-carpelled ovaries,
is treated as an "aggregate species". Walter (1909) distinguished P. acinosa from
P. esculenta primarily by the condition of the peduncle and inflorescence axis,
scabrous in the former and glabrous in the latter, coloration of floral parts, and
shape of sepals, all of which are not necessarily correlated in the manner described
in his key. The individual characters, particularly coloration, are unsound bases
for specific recognition.

Sect. 2 Pircunioides H. Walter, Pflanzenr. IV, 83 (Heft 39) : 42, 1909.
3. Phytolacca dodecandra L'Her., Stirp. Nov. 143, pi. 69, 1789. (Type Bruce s.n.

K)
P. ahyssinica Hoffm., Coram. GStting. 12: 27, 1796.
P. elongata Salisb., Prodr. 345, 1796.
P. lutea Marsigl. ex Steud., Norn. ed. 1, 618, 1821.

byssinica (Hoffm.) Moq. in DC, 13(2): 30, 1849.
Phytolacca scandens Hilsenb. & Boj. ex Moq., loc. cit., nom. nud. pro syn. Pircunia

Shrubs or herbs, dioecious, ± scandent. Leaves ovate or rarely ovate-lanceolate
or ± deltoid, acute-mucronate to retuse-mucronate, entire, the bases rounded,
oblique-rounded, or ± obtuse, up to 14 cm long and 8.5 cm wide; petioles up to
3.5 cm long. Inflorescences racemes, pistillate or staminate, up to 30 cm long,
axillary or terminal, pubescent. Staminate flowers with pedicels ca 4-6 mm long;
bract single, ca 1-1.2 mm long, very narrow; two bracteoles, ca 0.6 mm long, very
narrow; sepals 5, ± equal, narrow-oblong, ca 2-2.3 mm long and 0.7-0.9 mm wide;
stamens 13-15, in two whorls, the filaments ca 2.5 mm long, widened at the base,
the anthers ca 1 mm long; ovary rudimentary, 3-5 carpellate. Pistillate flowers
with pedicels 5-7 mm long at maturity; bract single, ca 1 mm long, awl-shaped;
two bracteoles, ca 0.6-0.8 mm long, ± awl-shaped; sepals 5, ± equal, oblong-ovate,
ca 2.5 mm long and 1.5 mm wide; stamens rudimentary, ca 8-12, deposited in two

NOWICKE — PHYTOLACCACEAE 309

(rarely one) whorls, the filaments ca 2 mm long, widened at the bases, the anthers
ca 0.8 mm long; ovary 5 carpellate, ± free, the styles widened at the bases, the
stigmas weakly penicellate. Fruit a collection of 5 or fewer drupelets, ca 3.5-4.5
mm long; seed 1, reniform, ca 3.5-4 mm long, testa shiny black.

Central and southern Africa and reported from Madagascar (Walter, 1909).

Nigeria: Oban, Talbot 1381 (MO).

Republic of Congo: Lake Kivu, Under 2032 (A); betw Sileko & Basoko, Louis 11404
(MO); Yangambe, Louis 264 (MO), 8638 (MO), 13890 (MO).

Ethiopia: s. loc, Loccardo s.n. (GH) ; Schimper 131 (GH).

Ruanda: Biumba Terr, Troupin 11819 (MO).

Tanzania: Bezirk Bagamogo, Schlieben 4136 (MO); N of Lake Nyasa, Stolz 2265 (A).

Uganda: s. loc, Dummer 5405 (A).

Rhodesia: Headlands Dist, Greenlow 88 (MO); Inyanga, Whellan 672 (MO).

Pollen grains subprolate, ca 23/u (E) X ca 28-29^t (P), colpi ca 17-18// long;
exine ca 2p in thickness, slightly thickened at the poles, sexine sparsely small
spinulose.

Pollen examined: Troupin 11819 (MO).
Subg. 2 Pircuniopsis H. Walter Pflanzenr. IV, 83 (Heft 39): 45, 1909. (Type P.

diaica L.)

Sect. 1 Pircuniophorum H. Walter, loc. cit.
4. Phytolacca sunguinea H. Walter, Pflanzenr. IV, 83 (Heft 39) : 46, 1909. (Lecto-

type selected: Lehmann 4479 US, isolectotypes F, K. Syntypes: Humboldt &

Bonpland 822; Karsten s.n.; Linden 852; all not seen)

Herbs, somewhat succulent, to 2 m. Leaves lanceolate-elliptic to elliptic to
rarely ± ovate, acute, entire to very finely undulate, the bases obtuse to attenuate,
up to 18 cm long and 6 cm wide, glabrous; subpetiolate to petioles ca 2.5 cm long.
Inflorescences racemes, thyrsiform at the base, up to 15 cm long, the peduncle ! / 3 to
]/ 2 this length, axillary or terminal, flowers crowded at maturity, scurfy. Flowers
with pedicels to 1 cm long; bract single, ca 7-9 mm long, lanceolate; bracteoles two,
sometimes appearing to be absent, ca 0.8-1 mm long, lanceolate; sepals 5, ± equal,
oblong-elliptic, ca 4-5 mm long and 2-3 mm wide, ± red; stamens 8-11 (-15),
deposited irregularly in one whorl, rarely two whorls, the filaments ca 2-2.5 mm
long, the anthers ca 0.8-0.9 mm long; ovary ca 9 carpellate, almost completely
united, the styles ca 1.5-2 mm long, recurved near the tip. Fruit a berry, ca 9 ribbed,
black, ca 6 mm in diam.

Mostly northern South America.

Costa Rica: Hatheway 1358 (US).

Colombia: cauca: El Tambo, von Sneidern 1146 (F, NY, US); Popayan, Lehmann
4479 (F, K, US); Mt Purace, Pennell & Killip 6510 (US); Quebrada del Rio San Marcos,
Cuatrecasas 14776 (F, US), putumayo: Mts above Laguna de La Cocha, Fosberg 20439
(NY, US); Valle de Sibundoy, Cuatrecasas 11597 (F, US), santander: mts E of Las Vegas,
Killip & Smith 15783 (NY); Paramo Rico, Killip & Smith 17703 (NY), tolima: betw
Cajamarca & summit of Divide, Killip & Varela 34558 (US).

Venezuela: merida: Paramo de las Lajas, Hamburg-Tracy 141 (K).

Pollen grains prolate spheroidal, ca 25-26^ (E) X ca 28^ (P), colpi ca 22-23^
long, sexine ± equal to nexine and finely reticulated.
: Killip & Smith 15783 (NY).

[Vol. 55
310 ANNALS OF THE MISSOURI BOTANICAL GARDEN

5. Phytolacca rugosa Br. & Bouche, Ind. Sem. Hort. Berol. 13, 1851. (Type

Warszewicz s.n., location unknown)

Herbs, woody at the bases, branches erect, angled, to 2 m. Leaves elliptic to
elliptic-lanceolate, acute to long acuminate, entire, the bases attenuated or obtuse,
up to 19 cm long and 6.5 cm wide; petioles to 5 cm long. Inflorescences racemes,
to 15 cm long, mostly axillary. Flowers with pedicels, 3-7 mm long; bract single,
ca 3-4 mm long, lanceolate; bracteoles two, < 1 mm long; sepals 5, ± equal,
elliptic, 2-3 mm long, reflexed in fruit, white to pink; stamens 6-12, deposited on
a hypogynous disc, the filaments ca 2 mm long, the anthers < 1 mm; carpels 4-9,
united at the bases, styles not connivent and straight. Fruit a berry, sharply ridged,
green-black, ca 6 mm in diam.

Mexico south to Colombia.

Mexico: guerrero: Hinton 11082 (MO), jalisco: Mexia 1661 (MO). michoacAn:
Hinton 11896 (MO).

Costa Rica: Dodge & Thomas 4551 (MO), 5320 (MO).

Panama: Allen 311 (MO), 7577 (MO); Blum et al. 2403 (MO); Dwyer et al. 469
AO); Siehert 302 (MO); Woodson & Schery 660 (MO); Woodson

Pollen grains prolate, ca 24-26^ (E) X ca 34^ (P), colpi ca 24^ long, exine
slightly thickened at the poles, sexine finely reticulated.
Pollen examined: Hinton 118% (MO).

6. Phytolacca chilensis (Miers ex Moq.) H. Walter, Pflanzeur. IV, 83 (Heft 39):

45, 1909, non H. Walter, loc. cit. 33.
P. chilensis Miers, Trav. 2: 532, 1826, nom. nud.

chilensis Miers ex Moq. in DC, Prodr. 13(2): 29, 1849. (Type Bridges s.n: K)

Herbs, with woody bases. Leaves ovate-elliptic, mucronate, entire, the bases
obtuse, up to 13 cm long and 5 cm wide; petioles to 3.5 cm long. Inflorescences
spikes, to 20 cm long, axillary or terminal. Flowers sessile or with pedicels ca 1
mm long; bract single, ca 4-5 mm long, lanceolate; bracteoles two, ca 2 mm long,
lanceolate; sepals 5, ± equal, ovate, rounded, ca 3-4 mm long and ca 2.5 mm
wide; stamens 9-12, in 2 whorls, the filaments ca 2 mm long, the anthers ca 0.7 mm
long; carpels 5-8, united only at the bases, the styles not connivent and straight.
Fruit a weak berry, 5-8 ridged, ca 6-7 mm in diameter.

Chile: s. loc. Bridges s.n. (K).

No pollen examined due to paucity of flowering material.

Known only from the type collection, it nonetheless separates very easily from
P.sanguinea and P. rugosa by its long spike. The possibility of hybrid origin from
P. icosandra L. x P. rugosa (which could give the general characters of P. chilensis)
are remote because of the geographical location. See also the discussion of
Anisomeria littoralis.

Sect. 2 Pircuniopsis

in DC, Prodr. 13(2): 30, 1849.

NOWICKE — PHYTOLACCACEAE 311

7. Phytolacca tetramera Hau. Mer., Apuntes Hist. Nat. 1: 108, 1909. (Type
Hauman-Merck s.n., photo US, from B|)

Herbs, dioecious, to ca 0.5 m. Leaves spatulate to elongate lanceolate, mucro-
nate, entire and finely undulate, the bases attenuate, up to 19 cm long and 5 cm
wide, midrib prominent, up to 2 mm wide; subpetiolate to petioles 1-3 cm long.
Inflorescences spikes, rarely spike-like racemes, staminate ca 4-5 cm long, the pistil-
late up to 10 cm long, mostly axillary. Staminate flowers sessile or with pedicels
ca 2-3 mm long; bract single, present or absent, ca 2 mm long, closely appressed
to sepals; bracteoles absent; sepals 4, unequal, ovate-elliptic, ca 2-2.5 mm long
and 1.8-2.3 mm wide; stamens 12-15, irregularly deposited, the filaments ca 2.5-3.5
mm long, the anthers ca 1-1.5 mm long; ovary absent or of 2-3 carpels, thin and
abortive. Pistillate flowers sessile or rarely with pedicels to 1 mm long; bract single,
ca 2-2.5 mm long, sepal-like and keeled at the base, closely appressed to calyx,'
bracteoles absent; sepals 4, unequal, ovate, ca 2-2.5 mm long and 1.6-2.2 mm wide-
stamens absent; ovary 6-9 carpellate, ± united, the styles somewhat connivent
Berry 6-9 ribbed, ca 4-6 mm in diam.

Argentina:
Plata & Magdak^
Cabrera 626 (NY).

Pollen grains subprolate, ca 23p (E) X ca 28-29,/ (P), colpi ca 17-18* long,
sexme ± equal to nexine and finely reticulated.
Pollen examined: Cabrera 626 (NY).

An easily identified species, not only because of the unique sepal number, but
also because of the distinct leaf shape.

8. Phytolacca dioica L., Sp. PI. ed. 2, 632, 1762. (Type Alstroemer 129 LINN not

seen; from IDC Micro-Edition 607.5)
P. populifolia Salisb., Prodr. 345, 1796.

lioica Moq. in DC, Prodr. 13(2) : 30, 1849.
Phytolacca arborea Moq., loc. cit., 31, nom. nud. pro syn Pircunia dioica.

Trees, dioecious, to 25 m. Leaves ovate, acute, entire to finely undulate the
bases rounded and sometimes decurrent, up to 12 cm long and 6 cm wide; petioles
up to 7 cm long. Inflorescences racemes, up to 15 cm long, axillary or terminal
Staminate flowers with pedicels to 4 mm long; bract single, ca 1 mm long- bracteoles
two, ca 1 mm long; sepals 5, ca 3 mm long; stamens 20-25, in two whorls the
filaments ca 4-5 mm long, the anthers ca 1.5-2 mm long; ovary occasionally
present, 2-4 abortive carpels. Pistillate flowers with pedicels ca 3 mm long ± stout-
bract single ca 0.5 mm long; bracteoles two, ca 0.5 mm long; sepals 5, ± equal,'
elliptic, ca 2-3 mm long, persistent in fruit; stamens ca 10, rudimentary; ovary 8-12
carpellate, incompletely united, the styles not connivent. Fruit a weak berry, ca

South America.

Ecuador: S Naranjapata, Schimpff 534 (MO)

tJirz^SftSft&s&'ig g$ NY> - •"" ™ : ,oinvlIk -

Uruguay: Montevideo, Herter 220 (MO).

312 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Argentina: chaco: s. loc, Jorgensen 1995 (MO); Venturi 9825 (MO), corrientes:
Santa Maria, Pedersen 455 (MO, NY), misiones: Santo Pipo, Schwarz 4844 (MO), tucu-
man: Quinta Lillo, Descole-Borsini 35934 (NY).

Paraguay: guaira: Villarrico, Jorgensen 3903 (MO), without province: Hassler 447a
(MO, NY), 3379 (NY), 3380 (NY).

Pollen grains prolate, ca 17 ft (E) X ca 25ft (P), colpi ca 17^ long, exine ca
2fi in thickness, sexine ± equal to nexine and finely reticulated.

Pollen examined: Venturi 9825 (MO).

9. Phytolacca weberbaueri H. Walter, Pflanzenr. IV, 83 (Heft 39) : 49, 1909. (Type

Weberbauer 4817, photo F from Bf).

Peru: cajamarca: vie of Casa Hacienda, Hutchison & von Bismarck 6349 (F, MO,
US).

Pollen grains prolate spheroidal, ca 21-22^ (E) X ca 24ft (P), colpi ca 17 fi
long, sexine ± equal to nexine and finely reticulated.

Pollen examined: Hutchison & von Bismarck 6349 (MO).

The description is omitted because of a lack of female specimens.

Subg. 3 Phytolacca.

Subg. Euphytolacca Moq. in DC, Prodr. 13(2): 31, 1849.

Sect. 1 Phytolacca.
Sect. Phytolaccastrum H. Walter, Pflanzenr. IV, 83, (Heft 39): 50, 1909.

10. Phytolacca icosandra L., Sp. PI. 631, 1753. (Type LINN, not seen; from IDG

Micro-Edition 607.4)
P. malabarica Crantz, Inst. 2: 484, 1769.

P. mexicana Gaerto. 77, t. 8, 1788, non Crantz (Inst. 2: 484, 1769)

P. triquetra Moench, Meth. Suppl. 107, 1802.

P. mexicana Swec : 337, 1827, non Crantz (Inst. 2 : 484, 1769)

■i K unth & Bouche, Ind. Sem. Hort. Berol. 15, 1848.
P. acuminata Moq. in DC, Prodr. 13 (2): 33, 1849, nom. nud. pro syn. P. icosandra.
P. longespica Moq., loc. cit. (Type Bates s.n. P)
P icosandra L. var. fraseri Moq. in DC, Prodr. 13(2): 34, 1849. (Type Fraser s.n. G, not

seen in IDC Micro-Edition, Candolle Prodromi Herbarium)
P nova-hispania Millsp., Publ. Field Mus. Nat. Hist, Bot. Ser. 2: 41, 1900. (Type Mill-

svauzh 1413 F)
P icosandra var angustitepala H. Walter, Pflanzenr. IV, 83 (Heft 39) : 61, 1909. (Type

Kerber 216a, location unknown)
P. icosandra var. sessiliflora (Kunth & Bouche) H. Walter, loc. cit.

Herbs, sometimes woody at the bases, stems angled or grooved, to ca 2 m.
Leaves variable, elliptic, to obovate or rarely lanceolate, acute, entire, the bases
obtuse to attenuate, up to 16 cm long and 8 cm wide, ± glabrous, subsessile to
petioles ca 4 cm long. Inflorescences spikes or spike-like racemes, up to 30 cm long,
mostly axillary. Flowers mostly sessile or rarely with pedicels to 4 mm long; bract
single, lanceolate, ca 4 mm long; bracteoles two, ca 1 mm long; sepals 5, ± equal,
broadly elliptic to ± lanceolate, ca 3 mm long and ca 2 mm wide, persistent in
fruit, pink tinged; stamens ca 8-20, usually in two whorls, on a hypogynous disc,

NOWICKE — PHYTOLACCACEAE 313

the filaments ca 2-2.5 mm long, widened at the bases, the anthers ca 1 mm long-
ovary 6-9 carpellate, united, the styles connivent in flower and recurved Fruit a'
berry, dark green to brown, rarely purple, ca 5-7 mm in diam.

Mexico, Central America, the West Indies and northern South America.
o«**S™ : DURANGO: JH almer 157 (MO), guerrero: Hinton 9240 (MO), jalisco- Prinele
9525 (MO), mexico: Ortenburger 16M650 (MO), michoacan: Bro Arsene8703 (Mm

SS- X^rS? (M0); Sc Z ery 724 (MO) - —lonT2ml mS:
^^Tatf^S SINALOA: Gentry 59n (M0) - SONORA: Gen »y 1422 cmd>-

Costa Rica: Godfrey 66075 (MO).

Panama: Leu; is et al. 321 (MO).

Cuba: Wng/ii 1392 (MO).

Haiti: Holdridge 1630 (MO).

Colombia: cauca: El Tambo, Hulten 9 (GH).

Venezuela: miranda: Los Mariches, Pittier 11965 (MO).

Pollen grains prolate, ca 26^ (E) X ca 36u (P), colpi ca 26a long, exine ca
2^ in thickness, thickened at poles to ca 2.5>, sexine ± equal to nexine and finely
reticulated. J

Pollen examined: Gentry 1423 (MO).

This is one of the more distinct species of Phytolacca perhaps best recognized
by a combination of long inflorescences, almost sessile flowers and two staminal
whorls, although any one of these three features alone is not sufficient.

1L IDctlSS: 607 S 1 P ) P1 - ^ * m ' 1763 ' (T - UNN ' "* ~* f -

p ^TiST^^l^i l J&T Gaertn - (Fruct l! 377 ' pl 77 ' * * 1788) nec

P. decandra Descourt Fl Antill. 5: 32, pl. 312, 1827, non L. (Sp. PI. ed. 2: 631, 1763)
P. octandravar. grandifolia Moq. in DC, Prodr. 13(2): 32, 1849. (Type Galea ti 372 ^
P. avnosa Pope, Wayside Pl. Hawaii 61, pl. 25, 1929, non Roxb. (Hart Bengal 3 I^8I4)

Herbs, ± succulent, to 2 m. Leaves lanceolate, lanceolate-ovate, acute to
acute-acuminate, sometimes mucronate, entire, the bases obtuse to attenuate up to
22 cm long and 7.5 cm wide; subpetiolate to petioles ca 2.5 cm long. Inflorescences
spikes or rarely spike-like racemes, up to 14 cm long, axillary or terminal, ± scurfy.
Flowers sessile or with pedicels to 1 mm long; bract single, ca 2 2-25 mm lone
anceolate; bracteoles two, ca 1 mm long, narrow-lanceolate; sepals 5, subequat
oblong to ovate, ca 2-3 mm long and 1.2-2 mm wide; stamens 8-10, in one whorl,

united! ! T I ^ l ° n i ^ SntherS < ' mm l0ng; ™* 7 " 10 <*P^
ir! diam conniven *- Fruit a berry, 7-10 ribbed, green-black, ca 4.5-6 mm

A ± cosmopolitan species.

Mexico: federal district: Lyonnet 220 (K, MO), morelos: Clark 7293 (MO^ nit™

Guatemala: Greenman & Greenman 5698 (MO). g W -

Colombia: norte de santander: Mutiscua, Killip & Smrt/i 7965/ (NY)

f J ( ^0). mir ^da: upper Pico de Naigmata, Pittier 6272 (NY).
Bolivia: cochabamba: Pocona, Steinbach R6R2 (Y Mn ^Tv^ V '

India: Kodaikanal,

314 ANNALS OF THE MISSOURI BOTANICAL GARDEN

kInyaT^STs of Eldort, Bogdan 1813 (K); Kakamega Forest, Lucas 100 (K); Ngong
Hills, Kokwaro 3/2 (K); vie of Norfolk Hotel, Kiwika 249 (K).

Rhodesia: Hondi View, Noel 2341 (MO).

South Africa: Utrecht Natal, Pole-Evans 3894 (K).

Australia: new south wales: Constable 19105 (MO).

New Zealand: Rangitoto I, Walker 4288 (MO).

Hawaii: oahu: Degener 8892 (MO), 8893 (NY). Kauai: Forbes 507 (K, MO).

Pollen grains prolate, ca 26^ (E) X ca 36^ (P), colpi ca 21-22^ long, exine ca
2.5ft in thickness and finely reticulated.

Pollen examined: Gaumer 674a (MO).

There are suggestions of characteristics of this species in many specimens;
bona fide collections of P. octandra may be rather easily identified by the ± sessile
flowers, inflorescences which are not conspicuously long (as in P. icosandra L.),
and which have a relatively short peduncle and a single whorl of stamens.

12. Phytolacca thyrsiflora Fenzl ex J. A. Schmidt in Mart., Fl. Brazil. 14(2) : 343,

pi 80, 1872.

Herbs to 3 m. Leaves ovate-elliptic to elliptic, ± acute, entire or very finely
undulate, the bases obtuse to attenuate, up to 14 cm long and 7 cm wide, glabrous;
subpetiolate to petioles ca 3 cm long. Inflorescences raceme-like, thyrsiform near
the base or often ± completely to tip, up to 30 cm long, axillary or terminal, the
flowers not crowded at maturity, ± scurfy. Flowers with pedicels to 1.4 cm long;
bract single, ca 2.5-4 mm long, lanceolate; bracteoles two, sometimes obsolete, ca
1.5-1.8 mm long, lanceolate; sepals 5, ± equal, oblong-ovate, ca 2-3 mm long and
1.5-2 mm wide; stamens 9-12, in one whorl, rarely a partial second whorl, the
filaments ca 1.8-2 mm long, the anthers ca 0.8 mm long; ovary 7-9 carpellate and
united. Fruit a berry, 7-9 ribbed, green to black, ca 5-7 mm in diam.

Brazil, French Guiana and Paraguay.

Brazil: esphuto santo: Espirito Santo, Robert s.n. (K). goias: S of Corumba de Goias,
Irwin et al 10883 (MO); W of Veadeiros, Irwin et al. 12793 (MO), maranhao: Maracas-
sume River, Froes 1997a (K). minas geraes: betw Itamuri & Realeza, Duarte & Castellanos
33176 (F);'4 km SE of Vicosa, Irwin 2171 (US), pernambuco: Recife, Tavares 985 (US).

15354 (K). santa catarina: anth
Caruru de Cacho, Reitz 1889 (NY).

French Guiana: Broadway 637 (US).

Paraguay: amambay: Sierra de Amambay, Hassler 9909 (K, NY).

Pollen grains subprolate, ca 25/i (E) X ca 30^ (P), colpi ca 22-23^ long,
sexine ± equal to nexine and finely reticulated.

Pollen examined: Reitz 1889 (NY).

13. Phytolacca heterotepala H. Walter, Pflanzenr. IV, 83 (Heft 39) : 51, 1909.

(Syntypes: Bourgeau 199 pro parte; Ehrenberg s.n.; Hahn s.n.; Schiede s.n.;

Schumann 1185 pro parte; all not seen).

Herbs to ca 2 m. Leaves ± lanceolate to ovate, acute to acute-mucronate,
entire, the bases obtuse, up to 13 cm long and 6 cm wide; petioles up to 5 cm long.
Inflorescences spike-like racemes, up to 25 cm long, mostly axillary, slightly scurfy.

NOWICKE— PHYTOLACCACEAE 315

Flowers with pedicels 1-4 mm long; bract single, ca 2.5-3 mm long, lanceolate;
bracteoles two, ca 0.8-1 mm long, ± awl-shaped; sepals 5(-6-8), unequal, ±
oblong, ca 3-4 mm long and 1.5-2.2 mm wide, stamens 15-22, in two whorls,' the
filaments ca 1.8 mm long, the anthers ca 0.9-1 mm long; ovary 8-9 carpellate'and
united. Fruit a berry, ca 8-9 ribbed, purple-black, ca 6-7 mm in diam.
Western United States and Mexico.

United States: California: Howell 34602 (US), 34604 (US), 35095 (US) 35121 (US)
Mexico: tamaulipas: Viereck 333 (US). "w^.awip),

Pollen grains subprolate, ca 28^ (E) X ca 35^ (P), colpi ca 24p long, exine
slightly thickend at the poles, sexine finely reticulated.
Pollen examined: Howell 34602 (US).

The only collection listed in Walter's (1909) original description which I
have seen is Schumann 1185 (US), and I believe this to be P. octandra L. Never-
theless, the specimens from California possess the distinctive characters which
Walter (1909) cited, i.e. unequal sepal size and two staminal whorls. Specimens
collected by Howell in the vicinity of San Francisco, exhibit hybrid characters such
as 8-9 sepals, sterility, and thyrsiform inflorescences, particularly Howell 35122
(US), which because of the above anomalies, I do not cite (see Howell, 1960).
14. Phytolacca meziana H. Walter, Pflanzenr. IV, 83 (Heft 39): 57, 1909. (based
on P. icosandra var. octogyna)

R ISn^ifiSffi/; a Smith ' Bot - Gaz - 18: 210 ' 1893 - ™* »** * ^

Herbs, somewhat suffrutescent at the base. Leaves lanceolate-elliptic acute
entire, the bases obtuse, up to 14 cm long and 4.5 cm wide, glabrous; petioles up
to 5 cm long. Inflorescences racemes, up to 28 cm long, mostly axillary, ± scurfy.
Flowers with pedicels to 1 cm long; bract single, ca 7-9 mm long, lanceolate;
bracteoles two, ca 2 mm long, lanceolate; sepals 5, ± equal, oblanceolate to oblong
ca 4 mm long and 2 mm wide; stamens 15-20, in two whorls, the filaments ca 2-2 2
mm long the anthers ca 0.9-1 mm long; ovary 7-8(-9) carpellate, united, the
styles ca 1.5 mm long, slightly recurved at the tip, the stigma on the upper surface.
Fruit a berry, 7-8 ribbed, black, ca 5-6 mm in diam.
Central America.

Guatemala: Heyde & Lux 3031 (US).
Costa Rica: Standley 42697 (US).

Pollen grains subprolate, ca 2^ (E) X ca 30-31,, (P), colpi ca 19-21,. long,
exine shghtly thickened at the poles, sexine ± equal to nexine and finely reticu
lated. '

Pollen examined: Heyde & Lux 3031 (US).

15. Phytolacca rivinoides Kunth & Bouche, Ind. Sem. Hort. Berol. 15, 1848 (Type

Moritz s.n., location unknown)
P. icosandra Wright, Mem. 268, 1828, non L. (Sp. PL 631, 1753).
P Z r?Z^ q W Se ,U EX °\ P / 6 - 3 ' 18 5 M RB - K - < Nov - Gen - SP- PL 2: 183, 1817)

ZsZ7ra ya ^"^ " MOq " M DC Pr ° dr - 13(2): 33 ' I849 ' ™. nud! pro syn P.
P. polystigma Benth. ex Moq., loc. cit.
P. polystyla Schomb. ex Moq., loc. cit., 460.

[Vol. 55
316 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Herbs, succulent, robust, erect to 5 m. Leaves ovate-elliptic, rarely lanceolate,
acute, acuminate, or rarely mucronate entire, the bases obtuse or rarely oblique,
up to 21 cm long and 9 cm wide; petioles to 7 cm long. Inflorescences racemes,
ca 40-50(-70) cm long, axillary or terminal, robust, the peduncle red-purple. Flow-
ers with pedicels 7-13 mm long, filiform in some; bract single, ca 2 mm long,
awl-shaped; bracteoles two, ca 0.5-0.7 mm long; sepals 5, ± equal, ovate, ca 2
mm long and 1.5 mm wide, white to pink, absent in fruit; stamens 10-17, in two
whorls on a hypogynous disc, the filaments ca 1.5-2 mm long, the anthers ±
globose, < 1 mm long; ovary 10-16 carpellate, united, the styles connivent at
the base, free and recurved above. Fruit a berry, juicy, ribbed when dry, purple-
black, 2-3 mm in diam and delicate appearing to 5-6 mm in diam and robust.

Mexico, south to Bolivia and Brazil, and the West Indies.

Mexico: nuevo le6n: Frye & Frye 2548 (MO).

British Honduras: Schipp 125 (MO).

Honduras: Yuncker et al. 6166 (MO), 8459 (MO).

SSJtet- M^Tl^hMO), 6406 (MO); Godfrey 67290 (MO); Rajas
205 &^%WS£Sl (MO), 8104 (MO); FUnger 49 (MO); Hunter
fr A1l£ m I (MOV Lewi et al. 874 (MO); Tyson 2186 (MO), 3364 (MO); Tyson et al
2922 (MO); von Wed d833 (MO), 2078 (MO), 2079 (MO), 2631 (MO); Woodson et al.

Dominican Republic: Valeur 11 (MO).
Guadeloupe: Duss 2400 (MO).
Jamaica: Hespenheide 709 (MO).
Puerto Rico: Otero 645 (MO) ; Stimson 1684 (MO).
Trinidad: Broadway 6778 (MO) .

Colombia- putumayo: Rio San Miguel, Schultes 3646 (MO), santa marta: s. loc,
Smith 1161 (MO), 2664 (MO), valle: Jamundi, vonSne^n 4558 (MO).
Venezuela: merida: Colonia Tovar, Fendler 1084 (MO).

G 7 Y r E D^C^ 2X^2/32 (MO), 322, (MO), 3639 (MO) 4481 (MO).
Brazil: acre: NW of Cruzeiro do Sul, Prance 2796 (MO.) amapa: vie of Mt Bruyers,

^"bolwu: beni: Werdermann 2165 (MO), la paz: S Yungas % Krukoff^ \
nordyungas: nr Coroico, Buchtien 4117 (MO), s/
(MO), without province: Buchtien 5403 (MO).

Peru- cuzco: Quincemil, Vargas 7779 (MO), loreto: vie of Pongo de Mansenche,
Mexia 6231 (MO); Boqueron Padre Abad, Woytkowski 34358 (MO) puno: trail bctw
Santo Domingo & Maehu, Metcalf 30655 (MO), san martin: vie of Moyobamba, King
3494 (MO).

Pollen grains subprolate, ca 27^ (E) X ca 35^ (P), colpi ca 22-23,* long,
sexine ± equal to nexine and finely reticulated (Fig. 7).

Pollen examined: Frye & Frye 2548 (MO).

An easily recognized species because of the very robust racemes, long pedicels
and high carpel number.

16. Phytolacca purpurascens A. Br. & Bouche, Ind. Sem. Hort. Berol., app. 13,

1851. (Type Warszewicz s.n., location unknown)

Herbs. Leaves elliptic, lanceolate-elliptic, or rarely ± ovate, acute or mucro-
nate, entire, the bases obtuse, up to 22 cm long and 8 cm wide; petioles to 7 cm

NOWICKE— PHYTOLACCACEAE 317

long. Inflorescences spike-like racemes, up to 28 cm long, mostly axillary, lightly
pubescent. Flowers with pedicels 3-6 mm long; bract single, ca 2.3-2.8 mm long,
lanceolate; bracteoles two, ca 1.5-1.8 mm long, lanceolate; sepals 5, subequal, ±
ovate, ca 2.5 mm long and 1.8-2 mm wide; stamens 13-17(-20), in two whorls,
the filaments ca 1.5-1.7 mm long, the anthers ca 0.7-0.8 mm long; ovary (7-8-)
9-10 carpellate, ± completely united, the styles ± connivent. Fruit a berry 7-10
ribbed, ca 6-7 mm in diam.

Sparsely distributed in the West Indies and reported from Central America
(Walter, 1909).

Haiti: Leonard & Leonard 11631 (MO, US), 13394 (US), 739/2 (US), 15454 (US).

Grand Bahama I.: Lewis 7166 (MO).

Pollen grains prolate spheroidal, ca 27^ (E) X ca 30^ (P), colpi ca 23^
long, sexine ± equal to nexine and finely reticulated.

Pollen examined: Leonard & Leonard 13912 (US).

Phytolacca purpurascens is closely related to P. icosandra but can usually be
distinguished from the latter by its longer pedicel, a somewhat unreliable character
because of continuous graduation. This taxon may well be of hybrid origin from
P. icosandra X P. rivinoides or P. americana.

17. Phytolacca brachystachys Moq. in DC, Prodr. 13(2): 31, 1849. (Type Beechey
P. abyssmica Hook. & Am., Bot. Beech. Voy. 94, 1832, non Hoffm. (Comm. Gotting. 12: 27,
P. bogotensis Mann, Proc. Amer. Acad. 7: 198, 1867, non H.B.K. (Nov. Gen. Sp. PI. 2: 183,

Herbs. Leaves ± elliptic, acute or mucronate, entire, the bases obtuse, up to
17 cm long and 9 cm wide; petioles to 3.5 cm long. Inflorescences racemes, up to
16 cm long, mostly axillary, ± scurfy. Flowers with pedicels to 5 mm long; bract
single, ca 2-2.5 mm long, lanceolate; bracteoles two, ca 1-1.5 mm long, lanceolate;
sepals 5, ± equal, oblong, ca 3-4 mm long and ca 1.8-2 mm wide; stamens 7-9,
in one whorl, the filaments ca 1.5 mm long, the anthers ca 0.8 mm long; ovary
5-7(-8) carpellate, united the styles connivent or sometimes free. Fruit a berry
5-7 ribbed, ca 4-6 mm in diam.

Hawaii: Degener 8891 (MO), 8896 (MO), 8897 (MO); Degener & Wieblse 3302
(MO); Forbes 184 (MO), 208 (MO), 7036 (F, MO); Forbes & Rock s.n. (MO); Fosberg
10946 (F); Heller 2772 (F, MO); Mann & Brigham 426 (F, MO).

Pollen grains subprolate, ca 29^ (E) X ca 32^ (P), colpi ca 22-23^ long,
exine ca 2.5^ in thickness, sexine ± equal to nexine and finely reticulated.

Pollen examined: Heller 2772 (MO).

18. Phytolacca bogotensis H.B.K., Nov. Gen. Sp. PI. 2: 183, 1817.

P. austmlis Phil., Anal. Univ. Chil. 43: 536, 1873. (Type Philippi s.n. photo NY, from

P. rnbmntha H. Walter, Pflanzenr. IV, 83 (Heft 39) : 57, 1909. (Type Loretz & Hiewnymus

P. parviflora Hau.-Mer., Apuntes Hist. Nat. 1 : 107, 1909. (Type Hauman-Merck s n
location unknown)

318 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Herbs, ± succulent, to 2 m. Leaves lanceolate to elliptic, acute to ± acum-
inate, entire, the bases obtuse to attenuate, up to 18 cm long and 7 cm wide; sub-
sessile to petioles ca 3.5 cm long. Inflorescences spike-like racemes, up to 20 cm
long, axillary or terminal, scurfy. Flowers with pedicels ca 2-4 mm long; bract
single, 2-3 mm long, lanceolate; bracteoles two, ca 1-1.2 mm long, narrow-lanceo-
late; sepals 5, subequal, oblong to ± ovate, ca 2-3 mm long and 1.2-2 mm wide;
stamens 7-10, in one whorl, rarely a partial second whorl, the filaments ca 2 mm
long, the anthers ca 0.8-0.9 mm long; ovary 7-10 carpellate, united, the styles thin.
Fruit a berry, 7-10 ribbed, green-brown, ca 5-6 mm in diam.

South America.

Colombia: cauca: Popayan, Lehmann 616 (GH).

Ecuador: E of Loja, Espinosa 485 (NY); eastern Rio Bamba, Schimpff 791 (MO).

Chile: valdivia: S of Jose de La Mariquina, Hollermayer 1205 (MO).

Bolivia: la paz: vie of Sorata, Mandon 1030 (NY); Williams 828 (K, NY), without
province: Yungas Mts, Scolnik & Luti 502 (NY).

Eyerdam & Beetle 22227 (MO); Rio Yala r
de Hules, Venturi 9130 (MO).

Pollen grains prolate ca 25^ (E) X ca 34^ (P), colpi ca 23-24^ long, exine
ca 2-2.5^* in thickness and finely reticulated.

Pollen examined: Eyerdam & Beetle 22227 (MO) .

Phytolacca bogotensis is very difficult to separate from P. octandra; the pedicel
length, ± sessile in the latter species, appears to be the only consistent character.

19. Phytolacca americana L., Sp. PI. 441, 1753.

P decandra L., Sp. PI. ed. 2, 631, 1763, non Descourt. (Flor. Med. Antilles 5: pi. 312, 1827).

(Type LINN, not seen; from IDC Micro-Edition 607.3)
P vulgaris Crantz, Instit. 2: 484, 1766.
P. rigida Small, Bull. N.Y. Bot. Gard. 3: 422, 1904. (Type Small & Wilson 1893 NY)

Herbs, ± succulent, to ca 3 m. Leaves lanceolate-elliptic, elliptic or rarely
± ovate, acute or sometimes mucronate, entire, the bases obtuse, up to 30 cm long
and 12 cm wide (occasionally much larger) ; petioles to ca 6 cm long. Inflorescences
racemes, sometimes thyrsiform at the bases, up to 30 cm long, mostly axillary,
glabrous to lightly scurry. Flowers with pedicels to 12 mm long; bract single,
ca 2-3 mm long, lanceolate; bracteoles two, ca 1-1.5 mm long, lanceolate; sepals
5, ± equal ovate and rounded, ca 2.2 mm long and 2 mm wide; stamens ca 10,
in one whorl, the filaments ca 2 mm long, the anthers ca 0.8 mm long; ovary
ca 10 carpellate, united or sometimes the apices free, the styles mostly connivent
in flower. Fruit a berry, ca 10 ribbed, ca 6-8 (-10) mm in diam.

Widely distributed in eastern North America and introduced into Europe,
Africa and Asia; selected exsiccatae listed.

Canada: Ontario: Marie-V ictorin et al. 56838 (MO); Soper & Shields 377 (MO).

United States: Florida: Hitchcock 301 (NY); Moldenke 250 (MO); O'Neill s.n.
(MO); Small & Small 4120 (NY), 4682 (NY), 5439 (NY); Small & Wilson 2010 (NY);
Tracy 7531 (NY). Indiana: Beam 18 (MO). Louisiana: Ball 542 (MO). Massachusetts:
Churchill sn (MO); Seymour 265 (MO). Mississippi: Pollard 1156 (MO). Missouri:
Eggert s.n. (MO) ^O); Kellogg 1713 (MO). Oklahoma: Houghton 3936

(MO). Tennessee: Ruth 191 (MO), west Virginia: Berkley 840 (MO). Wisconsin: Fassett
27388 (MO).

NOWICKE — PHYTOLACCACEAE 319

Pollen grains ca 25^ (E) X ca 34^ (P), colpi ca 23-24^ long, exine ca 2fi
in thickness, slightly thickened at the poles to ca 2.5ft, sexine equal to or greater
than nexine and finely reticulated.

Pollen examined: Soper & Shields 377 (MO).

Although P. rigida Small has been considered conspeeific with P. americana
by many authors, including Sauer (1952), there does seem to be a higher frequency
of certain characteristics in specimens from Florida, the type locality for P. rigida.
These features include a shorter raceme, hence the specific epithet which refers
to the erectness of the inflorescence, and pedicels which are, on the average, a
little shorter and stouter than for most individuals of P. americana. Hardin (1964)
in a study of the two taxa cited the above observation plus minor differences in
leaves, bracteole width, number of fruits, etc., all of which, as his Table 1 (p. 163)
indicates, have overlapping measurements. The most striking discovery was the
difference in the xylem of the peduncle, in which P. rigida averages a 70% thicker
cylinder. In spite of Hardin's results, and he makes no definite designation of
the status of the Florida collections, I tend to favor a reduction of P. rigida to P.
americana in view of the widespread morphological variation in the entire genus.
Hardin's tentative conclusions and ideas are worthy of mention because of their
applicability to the problem of speciation in Phytolacca in general (Hardin, 1964,
p. 162) : "P. rigida may represent an ecological variant of P. americana which is
not yet distinct; i.e., an early stage in sympatric speciation. On the other hand,
two distinct species may have existed at one time and with forced migration and
sympatry during Pleistocene or before, the old barriers and discontinuities have
been eliminated. A third possibility is that P. americana has been "contaminated"
by introgression from a tropical or subtropical species such as P. octandra which
has an erect raceme, by way of Mexico or the Caribbean."

Sect. 2 Phytolaccoides H. Walter, Pflanzenr. IV, 83 (Heft 39): 61, 1909.

20. Phytolacca pruinosa Fenzl, Del. Sem. Hort. Vindob 8, 1855. (Type Kotschy
s.n. W, not seen)

Herbs, dioecious, stems ± succulent, mostly unbranched, to 1 m. Leaves
ovate-lanceolate, acute, rarely mucronate, entire, the bases obtuse-attenuate, up
to 10 cm long and 4 cm wide; petioles indistinct. Inflorescences racemes, the
staminate up to 12 cm long, the pistillate up to 7 cm long and densely flowered.
Staminate flowers with pedicels up to 6 mm long; bract single, ca 2.5-3 mm long,
lanceolate; bracteoles two, ca 1 mm long, located ca midway on pedicel; sepals 5^
± equal, ovate ca 3.5-4 mm long and ca. 2.5 mm wide, green to pink; stamens
15-20, in two whorls, the filaments ca 2-2.5 mm long, widened at the base, the
anthers ca 1-1.2 mm long; ovary rudimentary, 5-7 carpellate. Pistillate flowers
with pedicels up to 3.5 mm long; single bract, ca 2.2-3 mm long; two bracteoles,
ca 1.8-2 mm long; sepals 5, ± equal, ovate-triangular, ca 3-3.5 mm long and 2.5-3
mm wide, green to pink; stamens rudimentary and much reduced, 12-15, appear-

[Vol. 55
320 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ing to be deposited in two whorls; ovary 6-9 carpellate, united, the styles recurved.
Fruit a berry, 6-9 ribbed, ca 5-6 mm in diam; seeds reniform, ca 3.5 mm long.

Endemic to Cyprus.

Cyprus: Haradjian 487 (MO); Oswald 147 (K); Young 7808 (K).

Pollen grains ca 27^ (E) X ca Up (P), colpi 20-23^ long, exine slightly
thickened at the poles, sexine medium finely reticulated.

Pollen examined: Young 7808 (K).

Excluded Specimens:

Horticultural specimens from Kew, the type collections of P. clavigera W. W.
Smith (Bot. Mag. 149: pi. 8978, 1923). At first glance this specimen appears to
be the robust Asian species, P. acinosa; however, the carpels are definitely united.
Since it is of horticultural origin, I hesitate to treat it as a distinct species until
I have seen more collections.

Pittier 6272 (US). This is the type collection of P. venezuelensis O. C. Schmidt
(Notizbl. Ber. 8: 312, 1923), which in my opinion is a stunted specimen of P.
octandra. The carpel connation is not characteristic of subg. Pircuniopsis, to
which Schmidt assigned it.

RIVINOIDEAE
II. Subf. Rivinoideae Nowicke, subf. nov. Ovarium unicarpellatum et uniovula-

tum; achenium, drupa, samara, vel utriculus. (Type Rivina L.)

A. Tribe Seguierieae Nowicke, trib. nov. Inflorescentiae paniculiformes;
samara. (Type Seguieria Loefl.)

a. Calyx of 4 sepals, becoming woody and remaining erect in fruit 4. Gallesia

aa. Calyx of 5 sepals, herbaceous and becoming reflexed in fruit 5. Seguieria

4. GALLESIA
Gallesia Casar., Stirp. Bras. Dec. 5: 43, 1843.
Monotypic.

1. Gallesia integrifolia (Spreng.) Harms in Engler & Prantl, Nat. Pflanzenfam.

2(16): 144, 1934. (Neotype selected: Ducke 24211 US, isoneotypes G, S)
Thouinia integrifolia Spreng., Neue Entdeck. 2: 155, 1821.
Crataeva gorarema Veil., Fl. Flum. 1: 191; 5: pi. 4, 1825.
Gallesia scorododendrum Casar., Nov. Stirp. Bras. Dec. 5: 44, 1843.
G. gorazema (Veil.) Moq. in DC, Prodr. 13(2): 8, 1849. (Type Vauthier 146 G, P)

Trees to 30 m. Leaves ovate to ovate-elliptic, acute to acuminate, entire, the
bases obtuse or slightly cordate, up to 12 cm long and 6.5 cm wide, glabrous, ±
leathery; petioles ca 2-3 cm long. Inflorescences irregular panicles, ca 10-20 cm
long, axillary or terminal, softly pubescent. Flowers perfect, ± actinomorphic,
± sessile; bract single, ca 1.2 mm long; bracteoles two, ca 1.2 mm long; sepals
4, somewhat unequal, ovate, ca 4 mm long and 1.4-2.5 mm wide, green-brown,
leathery, pubescent, thickened at the base, erect in fruit and becoming enlarged
and the margins split; stamens numerous, irregularly deposited, the filaments ca

NOWICKE — PHYTOLACCACEAE 321

1.5 mm long, the anthers ca 1.5 mm long; ovary 1-carpellate, compressed laterally,
the style flattened, the stigma papillose on one edge. Fruit a samara, ca 2.5-3 cm
long, brown, the non-stigmatic edge gradually curved or ± constricted at the base;
seed one, ovoid to elliptic, compressed laterally.

Central South America: Bolivia, Brazil, Ecuador and Peru.

Pollen grains single, subprolate, ca 21-22^ (E) X ca 24-25/* (P), 3-colporoi-
date, colpi ca 9-lO^u long, os irregular, ca 4p wide and ± as long as colpi, exine
ca 1.7 fi in thickness, ca 2fi at the poles, sexine ± equal to nexine and ± smooth.

Although clearly related to the large genus Seguieria Loefl. by its almost identi-
cal flower plan and samara fruit, Gallesia is nevertheless easily identifiable by its
4-merous calyx, which has a woody texture and remains erect in fruit. Gallesia
ovata O.C. Schmidt is here reduced to a variety of G integrifolia (Spreng.) Harms
for the following reasons: while there appears to be some variation in the shape
of the samara wing, the condition is somewhat indeterminate, and the wider leaves
cited for G ovata do not constitute a distinguishing character.

la. Gallesia integrifolia (Spreng.) Harms var. integrifolia.

Bolivia: la paz: San Bartolome, Krukoff 10118 (G, S, US).

Brazil: acre: upper Rio Jurupary, Krukoff 5216 (G, S, US); Seringal, Ducke 24212
(S, US), amazonas: Rio Purus, Ducke 24211 (G, S, US), bahia: basin of Rio Grongogy,
Curran 7 (US), mato grosso: Recreio, lute 7427 (US), minas geraes: Caldas, Regnell
III 1014 (S, US). Parana: Patrimonio, Dusen 16786 (S). rio de Janeiro: Copacabana,
Glaziou4753 (S).

Ecuador: el oro: vie of Piedras, Little 6621 (US).

Peru: junin: Satipo, Vasquez 16 (G).

Pollen examined: Ducke 24211 (US).

lb. Gallesia integrifolia (Spreng.) Harms var. ovata (O.C. Schmidt) Nowicke,

G. ovata O. C. Schmidt, Repert. Sp. Nov. 32: 97, 1933. (Type Raimondi 11606 B|)
Brazil: acre: nr mouth of Rio Macauhan, Krukoff 5405 (G, S, US).
The Krukoff collection is cited by Macbride in the Flora of Peru (1936) as
Gallesia integrifolia even though he recognizes both "species." It differs, however,
from all other collections of Gallesia in the constriction of the samara wing near
the base.

5. SEGUIERIA
Seguieria Loefl, Iter Hispan. 191, 1758. (Type S. americana L.)
Seguiera Adans., Fam. PI. 2: 443, 1763.
Albertokuntzea O. Ktze., Rev. Gen. PI. 2: 550, 1891.

Trees, shrubs, or lianas. Leaves lanceolate, lanceolate-elliptic to ovate, acute
to mucronulate, entire, the bases obtuse to slightly rounded, papery to ± leathery,
mostly glabrous; petiolate or subpetiolate; stipular thorns present or absent on
flowering branches, sometimes conspicuous on older branches. Inflorescences ir-
regular panicles, rarely ± racemes, axillary or terminal, mostly pubescent. Flowers
perfect, ± actinomorphic, pedicellate; bract single, awl-shaped, sometimes keeled

322 ANNALS OF THE MISSOURI BOTANICAL GARDEN

at the base; bracteoles two, or rarely absent, mostly awl-shaped; sepals 5, ± equal,
oblong to oblanceolate, somewhat veined, reflexed in fruit; stamens co, generally
in 2-3 irregular whorls, the filaments ± filiform, the anthers linear; ovary 1-carpel-
late, compressed laterally, the styles flattened, the stigma lightly papillose on one
edge. Fruit a samara, the base or ovary with or without a tubercle and ridges or
winglets, green to black on drying, the stigmatic edge convex, straight, or concave;
seed one, ovoid (Fig. 2).

South America.

Pollen grains single, prolate, ca 15-22// (E) X ca 25-30^ (P), 3-colpate, colpi
ca 18-25^ long, exine ca 1.5-2^ in thickness, sexine ± equal to nexine and almost
smooth.

In 1909 Walter described the genus Seguieria with 23 species in two sections.
The sect. 1 Euseguieria H. Walter, is characterized by a small tubercle and ridges
at the base of the samara and includes 13 species. The sect. 2 Seguieriella H.
Walter has samaras lacking the above characters and includes the remaining
species. Specific alignments within the sections are very difficult and the flowers,
and in most instances the fruits, do not yield good distinctive characters. My treat-
ment of the genus is admittedly based on insufficient material, even though all

Fig. 2. Seguieria L. samara types. A, S. coriacea Benth.; B, S. foliosa Benth.; C, S.
vauthieri Moq; D, S. floribunda Benth.; E, S. parvifolia Benth. A after Regnell III 1013
(US); B after Schomburgk 661 (K); C after Vauthier 29 (K); D after Gardner 722 (US);
E after Schwarz 4256 (MO).

NOWICKE— PHYTOLACCACEAE 323

loans from the world's major herbaria included a request for specimens. In view
of Walter's high number of species, I think two conclusions are possible; firstly,
that the group has not been well collected and distributed, or secondly, that as so'
many species are based on only one collection, they may represent extreme forms
of polymorphic species. Undoubtedly both factors plus the lack of fruiting material
have contributed to great difficulty with the genus. It is badly in need of mono-
graphic revision.

The original treatment of sect. 1 was severely limited by lack of material for
many of the species which Walter authored. However, after an examination of
type material at Kew and subsequent borrowing of many of these collections, I
have recognized and described nine of the 13 species and transferred one, S.
brevithyrsa H. Walter, to sect. 2. To my knowledge, I have not seen specimens
of S. inermis H. Walter and S. wangerinii H. Walter and omit their treatment;
and based on a photo of the type, S. pachycarpa H. Walter has been reduced to S.
foliosa Benth. Even though most descriptions are from type material, I wish to
emphasize that the entire treatment of the section is highly provisional due to lack
of additional collections to reinforce the validity of species.

The sect. 2 Seguieriella, which included 10 species according to Walter (1909),
is here reduced to six or possibly seven species, not including the transfer of S.
brevithyrsa. Seguieria paraguayensis Morong, S. macrophylla Benth. and S. glaziovii
Briq. are all rather distinct species. Seguieria affinis Heimerl (including S. rigida
H. Walter) is very close to S. glaziovii and may prove to be conspecific; the former
has very long stipules on the older stems, a character not specifically listed for S.
glaziovii, which may be lacking on the latter specimens due simply to poor collect-
ing techniques. Two wide ranging and very variable species, S. parvifolia Benth
[including S. votsahii H. Walter and S. guaranitica Speg.; the latter reduction
agrees with Heimerl (1934)] and S. americana are very difficult to define, and the
determiner must almost resort to utilizing these two complexes as "catch-alls" for
specimens which are not readily identifiable as one of the more distinct species.
Even so, I think in most instances the specimens would be correctly classified.
Seguieria mammifera H. Walter is not treated due to lack of material. The photo
of the type (Riedel s.n.) does very little to elucidate the species characteristics.

Following the Code, the sect. Seguieriella, which contains the type S. ameri-
cana, must become sect. Seguieria, and sect. Euseguieria I have renamed sect
Walteria.

a. Samaras with ridges or winglets and/or a tubercle at the base Sect. 1. Walteria

b. Leaves leathery.

c. Leaves ± lanceolate, or if lanceolate-elliptic then at least some 10 cm
long.

d. Leaves crowded, the largest < 10 cm long; inflorescences mostly
axillary and < 10 cm long 1. S langsdorffil

dd. Leaves not crowded, at least some > 10 cm long; inflorescences

mostly > 10 cm long and axillary or terminal 2. S lonsifolia

cc. Leaves ± ovate, elliptic-ovate, or if rarely then < 10

e. The stigmatic edge of the samara wing convex, or if not distinctly

ANNALS OF THE MISSOURI BOTANICAL GARDEN

f. Non-stigmatic edge of samara wing constricted near the top of

the ovary, and wings < 15 mm at its widest point 3. S. floril

ff. Non-stigmatic edge of samara wing arising conspicuously from

ee. The stigmatic edge ± straight or concave, the non-stigmatic edge

not noticeably arising from the base of the ovary 5. S. c

. Leaves not distinctly leathery,
g. At least some leaves with retuse tips.

h. Stipules small, < 2 mm long, straight; leaves ovate to ovate-^
rounded, the I

hh. Stipules 2-3 mm long, recurved; leaves ovate-elliptic to ovate, the

bases obtuse 7. S. emarginata

gg. Leaves without retuse tips.

i. Leaves lanceolate-elliptic to ovate-lanceolate, at least some 10-11 cm
long; flowering branches without stipules; inflorescences mostly

8. S. laurifolia

ii. Leaves ovate, the largest < 10 cm long; flowering branches with

recurved stipules; inflorescences mostly axillary 9. S. foliosa

. Samaras without protuberances at the base Sect. 2 Seguieria

± rounded, pale green on drying

i combination of characters,
aves ± lanceolate-elliptic, conspicuously mucronate.
Samaras black on drying, the stigmatic edge ± straight, a small

protuberance near the distal end 11. S. glaziovii

. Samaras brown on drvine. the stigmatic edee convex and gradually

12. S. affinis

Leaves not lanceolate and not conspicuously ]

h. Climbers; leaves ovate to ± elliptic, the stipules thick and recurved;

samaras black or dark brown on drying, distal protuberance on wing

drying 13. S. macrophylla

ii. Leaves small : bases ± obtuse; samaras black

on drying 14. S. hrevithyrsa

hh. Without the above combination of characters.

j. Samara wing with protuberance; leaves generally elliptic or

ovate-elliptic, the stipules ± straight; widespread 15. S. parvifolia

j]". Samara wing without protuberance; leaves more ovate, or ovate-
rounded, the stipules recurved; mostly from Colombia ....16. S. americana

Sect. 1 Walteria Nowicke, nom. nov. (Type selected: S. floribunda Benth.)
sect. Euseguieria H. Walter, Pflanzenr. IV 83 (Heft 39) : 87, 1909.

1. Seguieria langsdorffii Moq. in DC, Prodr. 13(2): 6, 1849. (Type Langsdorff s.n.

G, K)
Albertokuntzea langsdorffii (Moq.) O. Ktze., Rev. Gen. PI. 2: 550, 1891.

Trees to ca 15 m. Leaves crowded, lanceolate or lanceolate-elliptic, acute to
mucronate, entire, the bases obtuse or attenuate, up to 10 cm long and 4 cm wide,
± leathery; petioles to ca 1 cm long; stipules to 5 mm long, erect, slender. In-
florescences ± racemes or weak panicles, up to 10 cm long, mostly axillary, sparsely
pubescent. Flowers with pedicels to 5 mm long; bract single, ca 1-1.5 mm long,
awl-shaped and keeled at the base; bracteoles two or none, ca 0.5 mm long; sepals

NOWICKE— PHYTOLACCACEAE 325

oblong, ca 2.5-3 mm long and 1.8-2.3 mm wide; filaments ca 2 mm long, the
anthers ca 1.5-1.8 mm long. Samara unknown.

: s loc., Riedel s.n. (GH). minas geraes: s. Ioc, Langsdorff s.n. (K).
— J ~, Glaziou 8259, (K); s. loc, Filho 98 (F, MO), with-

Pollen grains ca 15p (E) X ca 25p (P), colpi ca 18-lfyi long.
Pollen examined: Riedel s.n. (GH).

2. Seguieria longifolia Benth., Trans. Linn. Soc. London 18: 235, 1839. (Type Pohl

Albertokuntzea longifolia (Benth.) O. Ktze., Rev. Gen. PI. 2:550, 1891.

Trees or shrubs? Leaves lanceolate to lanceolate-elliptic, acute, entire, the
bases obtuse to slightly rounded, up to 13 cm long and 4 cm wide, ± leathery;
petioles ca 4-7 mm long, stipules small, ca 1-2 mm on flowering branches and re-
curved. Inflorescences panicles, irregular, up to 17 cm long, axillary or terminal,
sparsely to densely villous. Flowers with pedicels to 5 mm long; bract single, ca
0.5-0.7 mm long, awl-shaped; bracteoles two, ca 0.5 mm long, awl-shaped; sepals
oblong, ca 3-4 mm long and 1.5-2.5 mm wide; filaments ca 2 mm long, the anthers
ca 1.5-1.8 mm long; ovary with winged and/or ridged base. Samara unknown.
SdZo*TttO BRASILIA: MathGa barb0S ° J PoM S - n - (K) - WITHOUT STATE = Pohl 3747 (F);

No pollen examined due to lack of suitable flowering material.

The specimens cited have minor variation in their leaf shape, i.e. the tip and
the base. While I include the Sella collection with some reservation, I believe this
to be the best decision until further collections are observed.

3. Seguieria floribunda Benth., Trans, Linn. Soc. London 18: 235, 1839 (Type

Gardner 722 BM, F, K, P, US)
Albertokuntzea floribunda (Benth.) O. Ktze., Rev. Gen. PI. 2:550, 1891.

Shrubs or woody vines. Leaves elliptic-ovate to elliptic-lanceolate, acute, entire,
the bases obtuse, up to 10 cm long and 6 cm wide; petioles ca 2-4 mm long;'
stipules 0.5-4 mm long, recurved. Inflorescences panicles, irregular, ca 18-22 cm
long, axillary or terminal, pubescent. Flowers with pedicels to 6 mm long; bract
single, < 1 mm long, awl-shaped; bracteoles two, < 1 mm long, awl-shaped;
sepals oblong, ca 4.5-6 mm long and 2-3 mm wide; filaments ca 1.5-1.8 mm long,'
the anthers ca 1.5-1.8 mm long. Samara ca 3.5-4 cm long, brown, the base with
prominent winglets, the stigmatic edge convex (Fig. 2D).

Brazil and Peru.

722 ( bTf, ™ uT : " ^ Remdl '" ' m (VS - * 1845> - mm " """ C -*~

Peru: madre de dios: Rio Acre, Ule 9487 (K, US).

No pollen examined due to lack of suitable flowering material.

The Ule specimens with the numbers 9487 and 9486 on the same sheet (K)
are cited by Macbride in the Flora of Peru (1936) as S. foliosa Benth. I have seen
the types of both species, and while the Ule collections are not identical to either
they are much closer to S. floribunda than to S. foliosa.

[Vol. 55
326 ANNALS OF THE MISSOUKI BOTANICAL GARDEN

4. Seguieria vauthieri Moq. in DC, Prodr. 13(2): 7, 1849. (Type Vauthier 29 K)
Alhertokuntzea vauthieri (Moq.) O. Ktze., Rev. Gen. PI. 2:550, 1891.

Trees or shrubs? Leaves ovate, obtuse, entire and slightly inrolled, the bases
obtuse, up to 3 cm long and 4 cm wide, somewhat leathery; petioles ca 3-4 mm
long, the stipules very small, ca 1 mm long or absent. Inflorescences panicles,
weakly so, ca 13 cm long, terminal, ± glabrous. Flowers with pedicels to ca 7 mm
long; bract single, ca 1 mm long; bracteoles two, ca 0.8-1 mm long; sepals and
stamens unknown. Samara ca 3.5-4 cm long and 1.6-2 cm wide, brown, the
stigmatic edge ± convex, the non-stigmatic edge arising from the base of the ovary
(Fig.2C).

Known only from a single fruiting collection.

Brazil: Brasilia, Vauthier 29 (K).

No pollen examined due to lack of suitable flowering material.

Walters (1909) concept of this species is erroneous. His description is based
solely on one of the confusing Regnell III 1013 collections (1864) of which there
appear to be three, all collected on different dates, 1845, 1855 and 1864. The 1864
collection bears Walter's determination as well as the location of Caldas. This
specimen differs markedly in samaral characters from that of Vauthier 29 (K), the
type collection of S. vauthieri. I have included Regnell III 1013 (1864 & 1855), as
well as an unnumbered Regnell collection from 1866, under S. coriacea Benth.
because of their similarity in leaf shape, stipule size and direction, and inflorescence
size and position, to that of the type for this species, Blanchet 2908.

5. Seguieria coriacea Benth., Trans. Linn. Soc. London 18: 235, 1839. (Type

Blanchet 2908 BM; photo F, from G)
Alhertokuntzea coriacea (Benth.) O. Ktze., Rev. Gen. PI. 2:550, 1891.

Shrubs or small trees. Leaves elliptic-lanceolate, acute to slightly mucronulate,
entire, the bases obtuse, up to 8 cm long and 3.5 cm wide, ± leathery; petioles ca
3-5 mm long; stipules up to 13 mm long, ± straight to slightly recurved. In-
florescences panicles, irregular, up to 40 cm long, axillary or terminal, lightly
pubescent, ± woody. Flowers with pedicels to 6 mm long; bract single, ca 1 mm
long, awl-shaped; bracteoles two, ca 0.8 mm long, awl-shaped; sepals oblanceolate,
up to 4 mm long and 2-2.6 mm wide; filaments ca 1.8-2 mm long, the anthers ca
1.5 mm long. Samara ca 3.5-4 cm long and ca 1.7 cm wide, brown-green, the base
with prominent winglets, the stigmatic edge concave (Fig. 2A).

Brazil: bahia: Acurua, Blanchet 2908 (BM; photo F, from G). minas geraes: s. loc,
Regnell III 1013 (K, in 1855; US, in 1864), s.n. (US, in 1866).

Pollen grains ca 18/* (E) X ca 27^ (P), colpi ca 19^ long.

Pollen examined: Regnell 1013 (K).

The specific epithet which Bentham (1839) applied to this species is perplex-
ing with regard to the BM collection of Gardner 2908. Of the six species which he
described, this specimen has perhaps the least coriaceous leaves. A photo (F from
G) of a more mature portion of the type does appear to have the coriaceous leaves
characteristic of the Regnell collections. For the present I am assuming, based on

NOWICKE— PHYTOLACCACEAE 327

l of the leaves (Bentham, 1839, p. 235) "foliis subsessilibus
oblongis obtusissimis coriaceis" and choice of names in light of the remaining
collections on which he based five other species, that he saw extensive material of
Gardner 2908 and that the BM specimen represents an immature leaf stage. See
also the discussion of S. vauthieri.

6. Seguieria alberti H. Walter, Repert. Sp. Nov. 8: 79, 1910. (based on S. elliptica)
S. elliptica H. Walter, Pflanzenr. IV, 83 (Heft 39): 89, 1909, non Fries (Ark. Bot. Stockh.

8 : 20, 1909) . (Type Glaziou 8260 K)

Trees or shrubs? Leaves ovate to ovate-orbicular, entire, obtuse and ± retuse,
the bases rounded or slightly obtuse in immature leaves, up to 6 cm long and 3.5
cm wide; petioles ca 3-6 mm long, stipules on flowering branches ca 2 mm long,
horizontal to ± ascending. Inflorescences panicles, irregular, axillary or terminal,
up to 10 cm long, ± villous. Flowers with pedicels to 5 mm long; bract single,
ca 1 mm long, awl-shaped; bracteoles two, ca 1 mm long, awl-shaped; sepals
oblong, ca 4-5 mm long and 2-2.5 mm wide; filaments 2.5-3 mm long, the anthers
ca 1-1.2 mm long; ovary with ridges in mature flowers. Samara unknown.

iB&G).

7. Seguieria emarginata H. Walter, Pflanzenr. IV, 83 (Heft 39) : 89, 1909. (Type

Glaziou 5730 K)

Trees or shrubs? Leaves ovate-elliptic to ovate, entire, blunt-acute and retuse,
the bases obtuse, up to 8 cm long and 5 cm wide; petioles 5-7 mm long, stipules
on flowering branches ca 2-3 mm long and recurved. Inflorescences panicles, ir-
regular, axillary or terminal, up to 8 cm long; ± glabrous. Flowers with pedicels
to 5 mm long; bract single, ca 1 mm long, awl shaped; bracteoles two, ca 0.5 mm
long, awl-shaped; sepals oblong, ca 4-5 mm long; filaments ca 3-4 mm long, the
anthers ca 1.5 mm long; ovary with ridges in mature flowers. Samara unknown.

Brazil: rio de Janeiro: s. loc, Glaziou 5730 (K; photo F, from B).

No pollen examined due to paucity of flowering material.

When the type specimens of S. emarginata (Glaziou 5730) and S. alberti
(Glaziou 8260), both from Kew, are compared, they are markedly different in leaf
shape, S. emarginata being conspicuously more elongate. However, a photo (F) of
the Berlin specimen of Glaziou 5730 has very broad leaves, and at first glance is
almost more similar to Glaziou 8260 than it is to the Kew specimen of Glaziou
5730. Nevertheless I have treated the two as separate taxa based on leaf shape, and
the stipule size and direction, but I admit that the distinction is somewhat obscure.

8. Seguieria laurifolia H. Walter, Pflanzenr. IV, 83 (Heft 39): 92, 1909. (Type

Glaziou 2488 K; photo F, from C)

Trees or shrubs? Leaves lanceolate-elliptic to ± ovate-lanceolate, acute, entire,
the margins ± inrolled, the bases obtuse, up to 11 cm long and 4 cm wide;
petioles up to 5 mm long, stipules absent on flowering branches. Inflorescences
panicles, irregular, rarely ± racemes, up to 15 cm long, axillary or terminal,

[Vol. 55
328 ANNALS OF THE MISSOURI BOTANICAL GARDEN

lightly villous. Flowers with pedicels to 6 mm long; bract single, < 1 mm long,
awl-shaped; bracteoles two, < 1 mm long, awl-shaped; sepals oblong, ca 4 mm
long and 2.2-2.5 mm wide; filaments ca 3 mm long, the anthers ca 1.8 mm long;
ovary appearing to have prominent winglets. Samara unknown.

Known only from the type collection.

Brazil: rio de Janeiro: Juiz de Fora, Glaziou 2488 (K; photo F, from C).

No pollen examined due to lack of suitable flowering material.

9. Seguieria foliosa Benth., Trans. Linn. Soc. London 18: 236, 1839. (Type Schom-

burgk 661 BM, F, K)
S. pachycarpa H. Walter, Pflanzenr. IV, 83 (Heft 39): 93, 1909. (Type Riedel s.n., photo

F, from BT)

Trees or shrubs to ca 3 m. Leaves ovate-elliptic, entire, acute to slightly
mucronulate, the bases obtuse to somewhat rounded, up to 12 cm long and 5 cm
wide, mostly 6-8 cm long and ca 4 cm wide; petioles ca 3-5 mm long; stipules to
10 mm long, recurved. Inflorescences panicles, irregular, or ± racemes, up to 12 cm
long, mostly axillary, lightly pubescent. Flowers with pedicels to 1 cm long; bract
single, ca 1-1.5 mm long, awl-shaped; bracteoles two, ca 0.6-0.8 mm long, awl-
shaped; sepals oblanceolate, ca 3-4 mm long and 1.5-2 mm wide; filaments ca 2
mm long, somewhat thickened at the base, the anthers ca 1 mm long. Samara
up to 3.5 cm long, brown, the base ridged, the stigmatic edge ± straight to slightly
convex, the non-stigmatic edge ± straight (Fig. 2B).

Brazil and Guyana.

Brazil: ceara: Maracanau, Ducke 2576 (NY).

Guyana: Rupununi Dist, Irwin 797 (US); s. lot, Schomburgk 661 (BM, F, K).

Pollen grains ca 25fi (E) X ca 30-3 lp (P), colpi ca 22-23^ long.

Pollen examined: Ducke 2576 (NY).

Sect. 2 Seguieria.

sect. Seguieriella H. Walter, Pflanzenr. IV, 83 (Heft 39) : 94, 1909.

10. Seguieria paraguayensis Morong, Ann. N. Y. Acad. Sci. 7:210, 1893. (Type

Morong 690 MO)

Trees or shrubs. Leaves ovate, mucronate, entire, the bases obtuse to slightly
rounded, up to 8.5 cm long and 6.5 cm wide, glabrous, ± leathery; petioles up to
1.5 cm long; stipules up to 3 mm long, ± straight to erect, mostly on older
branches. Inflorescences panicles, ca 6 cm long, mostly axillary, few-flowered,
lightly pubescent. Flowers with pedicel length variable, up to 7 mm long; bract
single, ca 2.5-2.8 mm long, lanceolate; bracteoles absent; sepals ± equal, oblong,
ca 4 mm long and ca 2 mm wide; filaments ca 2.5-3 mm long, the anthers ca 2
mm long. Samara up to 3 cm long, yellow-brown, the stigmatic edge convex.

Paraguay: San Bernardino, Hassler 3712 (F, NY), 3887 (F, NY); vie of Ypacaray,
Bossier 12400 (MO, NY); s. loc, Fiebrig 869 (A, F, US), Hassler 1764 (NY), s.n. (F),
Morong 690 (MO).

Pollen grains ca 17^ (E) X ca 26/* (P), colpi ca lfyi long.

Pollen examined: Hassler 12400 (MO).

NOWICKE— PHYTOLACCACEAE 329

11. Seguieria glaziovii Briq., Ann. Conserv. Jard. Bot. Geneve 4: 214, 1900. (Type
Glaziou 13126 G, K)

Trees to 30 m. Leaves elliptic-lanceolate, mucronate, entire, the bases obtuse,
up to 11 cm long and 4.5 cm wide, ± leathery; petioles ca 0.5-1 cm long; stipules
ca 0.5 cm long, slender. Inflorescences irregular panicles, up to 18 cm long, mostly
axillary, rarely terminal, rachis pubescent. Flowers with pedicels up to 8 mm
long; bract single, up to 1.5 mm long, awl-shaped, keeled at the base; bracteoles
two, ca 0.8 mm long, awl-shaped; sepals subequal, ca 4-5 mm long and 2-2.5 mm
wide, veined, prominently so in fruit; filaments ca 3 mm long, the anthers ca 2.2
mm long. Samara ca 2.5-3 cm long, black on drying, the stigmatic edge ± straight.

Brazil: santa catarina: Brusque, Klein 288 (NY), 290 (NY); Reitz 3464 (US);
Ibirama, Reitz & Klein 1563 (NY); Itajai, Klein 1183 (I ' • , n 2257 (NY, US),

2409 (NY, US).

Pollen grains ca 18-lfyi (E) X ca 29-30/* (P), colpi ca 25fi long.

Pollen examined: Reitz & Klein 2409 (NY).

12. Seguieria affinis Heimerl in von Wettstein, Ergebn. Exped. Sudbrasil 1901 1: 6,

1908. (Lectotype selected: Novaes 1027 US; syntype Novaes 1026 WU, not

S. rigida H. Walter, Pflanzenr. IV, 83 (Heft 39) : 98, 1909. (Syntypes de Moura 985 LE,

B|; Riedel s.n. LE, B|, all not seen)

Trees, to 18 m. Leaves lanceolate-elliptic, mucronulate, entire, the bases obtuse,
up to 11 cm long and 5 cm wide, glabrous to slightly pubescent, ± leathery;
petioles to 6-7 mm long; stipules to 4.5-5 cm long on older stems, slender. In-
florescences panicles or racemose, to 20-25 cm long, axillary or terminal, pubescent.
Flowers with pedicels ca 6-7 mm long; bract single, ca 1-1.5 mm long, awl-shaped;
bracteoles two, ca 0.6 mm long, ± triangular; sepals subequal, oblong and nar-
rowed at the base, ca 4.5 mm long and 3-3.5 mm wide, veined; filaments ca 3.5
mm long, the anthers ca 1.7 mm long. Samara ca 3.5 cm long, brown, the base
darkened, the stigmatic edge convex.

Brazil.

Brazil: minas geraes: rd to Barroso, Mexia 4444 (BM, F, MO); rd to Sao Miguel,
Mexia 4358 (BM, F, K, MO), sao paulo: Campinas, Novaes 1027 (US).

Pollen grains ca 18-% (E) X ca 28-29^ (P), colpi ca 21-22^ long.

Pollen examined: Mexia 4358 (F).

13. Seguieria macrophylla Benth., Trans. Linn. Soc. London 18: 235, 1839. (Type
Schomburgk 348 K)

Albertokuntzea macrophylla (Benth.) O. Ktze., Rev. Gen. PI. 2:550, 1891.

Lianas? Leaves ovate-elliptic, acute, entire, the bases rounded-obtuse, up to 17
cm long and 9 cm wide, leathery; petioles ca 0.5-1 cm long; stipules up to 6 mm
long, recurved, stout. Inflorescences panicles, up to 40 cm long, axillary or terminal,
pubescent. Flowers with pedicels ca 6 mm long; bract single, ca 1.2 mm long,
awl-shaped; bracteoles two, ca 0.8 mm long, awl-shaped; sepals subequal, ca 4 mm
long and 1.6-2.4 mm wide; filaments ca 2.5 mm long, the anthers ca 1.7 mm long.

330 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Samara up to 3.1 cm long, brown-black on drying, the stigmatic edge ± straight.

Brazil, Guyana, Peru and Venezuela.

Brazil: amazonas: nr mouth of Rio Embira, Krukoff 5206 (A), maranhao: Maracas-
sume River Basin, Krukoff 1924 (A, BM, MO), para: Belem, Archer 7936 (F, MO); Bouche
de lac de Faro, Ducke 8657 (US).

Guyana: Essequibo River, Schomburgk 348 (K); Kanuku Mts, Smith 3650 (A).

Peru: loreto: Yurimaguas, Poeppig 2176 (F).

Venezuela: delta: Cano del Gorisal, Bond et al. 208 (GH). without province:
Sacupana, Rushy & Squires 57 (F, MO).

No pollen examined due to lack of suitable flowering material.

14. Seguieria brevithyrsa H. Walter, Pflanzenr. IV 83 (Heft 39) : 87, 1909. (Type

Rusby 1353 BM, GH, NY)
S. inerensis Britton, Bull. Torrey Bot. Club 48:331, 1921. (Type Britton, Freeman &

Nowell 2527, probably NY)

Shrubs, decumbent, or woody vines. Leaves elliptic-lanceolate, acute-acum-
inate, entire, the bases obtuse, up to 11 cm long and 4 cm wide, glabrous, ±
leathery; petioles up to 1 cm long; stipules ca 4-6 mm long, recurved. Inflores-
cences racemes or weak panicles, up to 10 cm long, axillary, glabrous, black on
drying. Flowers with pedicels up to 6 mm long; bract single, 3-4 mm long, lanceo-
late; bracteoles absent; sepals subequal, oblong, ca 5 mm long and 3.5 mm wide;
filaments ca 3 mm long, the anthers ca 2 mm long. Samara up to 4.5 cm long,
black on drying, ± slightly ridged at the base, the stigmatic edge straight or con-
Reported only from Trinidad and Bolivia.

Trinidad: Smith 2706 (US).

Bolivia: s. yungas: San Bartolome, Krukoff 10166 (F, MO, US), without province:
Guanai, Rushy 1353 (GH, NY).

No pollen examined due to lack of suitable flowering material.

Walter (1909) included this species in his sect. 1, Euseguieria, characterized
by the extra tubercle at the base of the fruit, a condition difficult to detect in flow-
ers, and certainly not visible in the single collection from which he described
the species. The collection of Krukoff 10166, in fruit only, which agrees with
Rusby 1353 in leaf shape, stipule size and direction, inflorescence characters, and
blackening on dessication, as well as in geographic location, has two slight ridges
on the flattened sides of the ovary but no evidence of the tubercle. For this reason
I have transferred this species to the sect. Seguieria.

(Type

Seguieria guaranitiea Speg., Anal. Soc. Cient. Argent 16: 88, 1883.

Albertokuntzea parvifolia (Benth.) O. Ktze., Rev. Gen. PL 2: 550, 1891.

Seguieria elliptica R. E. Fries, Ark. Bot. Stockh. 8:20, 1909, non H. Walter [Pflanzenr.

IV, 83 (Heft 39) : 89, 1909] (Type Fries 313 US) .

Shrubs, ± scandent. Leaves ovate-elliptic to ovate, mucronulate, entire, the
bases obtuse, up to 11 cm long and 5 cm wide, ± leathery; petioles to 8-9 mm
long; stipules to 1 cm long, ± straight, mostly on older branches. Inflorescences

NOWICKE— PHYTOLACCACEAE 331

panicles or rarely racemose, up to 20 cm long, axillary or terminal, pubescent.
Flowers with pedicels up to 6 mm long; bract single, ca 1-1.2 mm long, awl-shaped,
membranaceous; bracteoles two, ca 0.8 mm long, awl-shaped, membranaceous;
sepals subequal, ca 4.5 mm long and up to 3 mm wide, veined, conspicuously
so at the base in fruit; filaments ca 3.5 mm long, the anthers ca 1.7 mm long.
Samara up to 4 cm long, green-brown, the stigmatic edge convex, rarely ± straight,
a small protuberance near the tip (Fig. 2E) .

A variable species widely distributed in South America.

Brazil: parana: Iguacu Ntl Pk, Pereira 5314 (F). rio grande do stit • Vila VU*
Rambo 41919 (F, MO). K J ° SUL " Vlla EIsa '

Argentina: misiones: Acaragua, Bertoni 3172 (F) ; Campo Grande. Schwarz 4419 f MC» •

s/sTmov s tri 425 \ ^°^r^r^ er ™ 4 <?>•• p«Xffl

1 1 c ^ ? ; S mS/JK* S r CkW ^ 4267 (MO) ' Santa Rita Schwarz 4204 (F). salta: Bosque
8 8 S ( f)'. ( )J La CaUera ' Pier0tU 2 ° 3 (F) ' betw EI Pi( * uete & Palmero, Ragonese

„ pAI ^H, AY: c*™ 11 - DEp r: Villa Elisa, Pedersen 3152 (MO, US) gaira- Villarrico
^ S (MO) (F ' M ° ); " l0C " HOSSler 1849e (NY) ' 3?86 (F > ™V> ^(Mo"m)Mlwn g

Pollen grains ca 21^ (E) X ca 28-29^ (P), colpi ca Up long, exine ca I5u
in thickness.

Pollen examined: Hassler 7055 (MO).

16. Seguieria americana L., Syst. Nat. ed. 10, 1074, 1759. (Neotype selected: Smith

342 MO, isoneotypes F, US)
S. aculeata Jacq., Select. Stirp. Amer. 170, 1763.
Albertokuntzea americana (L.) O. Ktze., Rev. Gen. PI. 2: 550, 1891.

Shrubs, scandent or climbers. Leaves ovate or ovate-elliptic, acute to slightly
accuminate and emarginate, the bases obtuse, up to 10 cm long and 7 cm wide
glabrous, papery to slightly leathery; ± sessile to petioles 3 mm long; stipules up
to 1 cm long, recurved. Inflorescences panicles, up to 15 cm long, axillary or
terminal, pubescent. Flowers with pedicels 6-7 mm long, slender; bract single,
ca 05 mm long, membranaceous; bracteoles two, ca 0.4 mm long, membranaceous;
sepals ± unequal, oblong, ca 3.5-4 mm long and 1.2-2.5 mm wide; filaments ca
2-2.5 mm long, the anthers ca 1.2 mm long. Samara ca 3.5 cm long, green-brown
the stigmatic edge ± straight.

South America.

Colombia: atlantico: Barranquilla, Dugand 106 (F), 474 (F) 1112 (F)- b P tw r*U
& Baranca, Dugand 272 (F US); Puerto Colombia, Bra. ElialV f (f/^S); bltw tn

W \?9? a ^ Rlta ' Dumnd 68 M F); S ° Iedad ' Du ^ d lm (F). magdalL Ta Paz
Haught 2330 (F). santa marta: s. be, Smith 342 (F, MO, US) '

Venezuela: aragua: Tuy Valley, Pittier 12201 (US)

Bolivia: s. loc, Williams 249 (BM, NY).

Paraguay: without province: Fiebrig 776 (F), Hassler 8393 (F), 11502 (F)

Pollen grains ca 21-22^ (E) X ca 30^ (P), colpi ca 20-2U long.

Pollen examined: Dugand 683 (F).

Excluded collection: Broadway s.n. (K). This is the type collection for S
cordata Britton (Bull. Torrey Bot. Club 48: 331, 1921), which has neither flowers
nor fruit and is impossible to describe or identify.

[Vol. 55
332 ANNALS OF THE MISSOURI BOTANICAL GARDEN

B. Tribe Rivineae Agardh, Aphor. 218, 1825. Inflorescences spikes or racemes;
fruits not a samara. (Type Rivina L.)

a Flowers unisexual; fruits covered with hooked spines; Australia 12. Monococcus

aa. Flowers perfect, rarely unisexual; fruits not covered with hooked spines; South
and Central America,
b. Fruit a 4-6 hooked achene; inflorescences elongate spikes; calyx of 4 narrow-

lanceolate, erect sepals U PeUvena

bb. Fruit an utricle or drupe not hooked; inflorescences racemes; calyx not or 4
narrow-lanceolate, erect sepals,
c. Flowers ± zygomorphic; sepals connate at the base m late flower and

£ ru j t y. tiulena

j I x _ a tin m iplu , s pals nui V r,mn„ c nwi" in lat flower

d" Leaves ± deltoid; stamens 4; fruit a red, orange, or purple drupe ....6. Rivina

dd. Without the above combination of characters.

e. Racemes from woody stems, not pendulous; fruit a purple^ r-

black drupe

ee. Racemes not from woody stems or if so,

f. Racemes pendulous; flowers perfect or functionally unisexual,
the pistillate with 4-6 rudimentary stamens, the sepals net-
ve i ne d 10. Ledenhergia

ff. Racemes ± erec- the stamens 12-25, the

sepals with ca 3 parallel veins 8. Schindleria

6. RIVINA
Rivina L., Sp. PL 121, 1753. (Type R. humilis L.)
Rivinia L., Gen. PL ed. 5, 57, 1754.
Piercea Mill., Gard. Diet. ed. 7, 1759.
Solanoides Moench, Meth. 307, 1794.
Tithonia L. ex. O. Ktze., Rev. Gen. PI. 2: 552, 1891.
Monotypic.

1. Rivinia humilis L., Sp. PL 122, 1753. (Type LINN, not seen; from IDC Micro-
Edition 163.1)
R. laevis L., Mant. 41, 1767.

R. purpurascens Schra ,. 17, pi. 5, 1808.

R. portulaccoides Nutt, Trans. Amer. Phil. Soc. II, 5: 167, 1837. (Type Nuttall s.n. BM)
For a complete synonomy list, see: Ann. Missouri Bot. Gard. 48: 76, 1961, By K. Raeder.

Herbs or subshrubs, woody at the base, to 70 cm. Leaves deltoid to ovate,
acuminate, entire, the bases truncate to rounded, rarely oblique, up to 12 cm
long and 6 cm wide, glabrous to finely pubescent; petioles up to 6 cm long. In-
florescences racemes, up to 15 cm long at maturity, terminal or axillary. Flowers
perfect, actinomorphic, with pedicels to 8 mm long in fruit; bract single, ca 1.3-2
mm long, awl-shaped, ± deciduous; bracteoles two, ca 0.2-0.3 mm long, closely
appressed to the sepals; sepals 4, subequal, oblong, ca 2-3 mm long, white or
pink; stamens 4, alternate, the filaments ca 1.2-2 mm long, the anthers ca 0.8-1
mm long; ovary globose to elliptic, compressed laterally, 1-carpellate, the style
short but distinct, ca 0.3-0.5 mm long, the stigma capitate. Fruit a drupe, up to
4.5 mm in diam, ± globose, orange, red, or purple; seed one, lens shaped, ca
2.5-3.5 mm in diam, the testa pubescent.

NOWICKE — PHYTOLACCACEAE 333

Southwestern United States to Florida, south through Central and South
America to Argentina; also introduced into Africa and reported from Asia and
Australia (Heimerl, 1934).

„^P N ^ D States: *«z°na: Gilman 53 (MO); Jones s.n. (MO). Florida: Blanton 6426
(MO); Curtiss 2340 (MO), 5383 (MO); Hitchcock 302 (MO); Hood s.n. (MO)- Janish
& Janish 454 (MO) ; Moldenke 325 (MO) ;Murrill s.n. (MO); Nash 1273 (MO)- O'Nc!
fwSJ 5 « fl/m ,f 458 (MO)j ^ (MO) > 27344 < MO )' R eWolds s.n., 11565 (MO) ;' S
(MO); SifioB & SmoU 4162 (MO); Tract/ 6429 (MO), 9349 (MO); iWer 376 (MO).
Louisiana: Riid-eH sa (MO); STiort s.n. (MO). Oklahoma: Houghton 4049 (MO).
Texas: Bush 1128 (MO); C/ian^er 7040 (MO); Daws s.n. (MO); Eggert sn (MO)-
Perm & Duncan 2632 (MO), 2890 (MO), 3/57 (MO); Heller 1422 (MO);']ermysn
MO); Jootsju (MO); ^f™n 423 (MO); lihifceimer 295 (MO), 374 (MO), 7/73
(MO), s.n. (MO); Macfcenne 48 (MO); Moore & Steyermark 3014 (MO); MueZZer 8094
^°>,w^v (MO): ° rcu " 58S6 (MO > : Palmer U7 ° ( MO )> 98 & (MO), 70770 (MO)
70794 (MO), 70357 (MO), 74357 (MO), 26782 (MO), 30527 (MO); Pa L 7359 (MO)
Reverchon 812 (MO) 7588 (MO); Ku* 229 (MO), s.n. (MO); T^y 9357 (MO)^/'
(MO); Traverse 1086 (MO); Pre/ease s.n. (MO); Ward s.n. (MO); W&foson JSS (MO);
Young s.n. (MO) . '

Mexico: baja California: Gentry 4748 (MO); Jones 24020 (MO), chihuahua-
Gentry 1846 (MO); PaZmer 254 (MO), coahuila: PaZmer 729 (MO) Stewart sn (MO)-"
Wamocfc & Bar/cZey 74720 (MO); Wynd & MueZZer 292 (MO), colima: West 3529 (MO)
^ D nt LG ^J: ingle 7428 (MO) - NUEV0 le6n: Bro - Ars ™ 61 38 (MO), sinaloa: Gentry
J? 5 r. (M S2n /°i^ RA: Geritry 7078A (M0 >' 1606 ( M °)J W fes*w & «°«frw 340 (MO);
W^w^n? 1 " 1 "™ ^ 6622 o (M0); Meyer & Rogers 2484 (MO); Palmer
127 (MO), 736 (MO), yucatan: Gaumer 322 (MO), 7599 (MO); Gaumer et al 23419
(MO), 23561 (MO); LuneZeZZ 858 (MO), 942 (MO), without state I Pwy77t J MO)
British Honduras: LundeZZ 380 (MO) ; Scnipp 469 (MO) .
Honduras: Williams & Molina 10510 (MO).
Guatemala: Steyermark 47631 (MO).
™ ^o^,?: 6 ™™ & Greenman 5423 (MO) ; Ro/as 472 (MO) ; Smith A667 (MO) ;
l nomas olod (MO).

^ P ,TnS : u Uen \ 36 * ( y°l'J 89 (MO); Carlet ° n 52 < M °)' Co °P er 77 (MO); Duke

™> ( * k H r*f%^ /e "5V MO); «" ^«W 609 (MO), 256 (MO), 2882 MO);

WTute & M* 88 (MO); Woodson & Scnery 7028 (MO); Woodson et al. 1807 (MO)

Bahamas: Wilson 7981 (MO), 8076 (MO), 8738 (MO). K '

et aZ. 790 (MO) ; Van Hermann 335 (MO) .

Dominican Republic: Valeur 486 (MO).

Grenada: Broadway s.n. (MO).

Guadeloupe: Grisebach s.n. (MO).

Jamaica: Crosby et al. 90 (MO); Hitchcock s.n. (Kingston) (MO) sn (Constant
Springs) (MO); LZoyrf 7072 (MO); Nichols 189 (MO); YuJklr 17074 (MO)

Martinique : Sieber 48 (MO) . '

7655 (MO) 5527° : (MO) er ^ (MO) ' ^ ^^ ^^ ^ (M ° )j ^^ 306 (M ° )s
. (MO); Ricksecker 134 (MO).

: Rincon Hondo, Allen 236 (MO), 375 (MO); Santa Marta,
^ of Palmira, Garcia 6394 (MO).

Lower Orinoco, Rusby & Squires 80 (MO), merida- Colonia
Tovar, Fendler 1088 (MO). Margarita i, Miller & /onnston 263 (MO).

DusenX(NYT : BatU " te ' ^^ ^ (MO) ' "" ™ JANEIR ° : ^ ° f Ri ° de Jandr °'

Uruguay: Herter 70078 (MO).

Argentina: buenos aires: Zorate, Krapovickas 3020 (MO), chaco- CamDo Bnnp/znla
ffS ^ (NY) ' 26?9 ( T ); S - SV**~« '*» (MO). c6^ I HfcSSfe
fe f^,^?- ENTRE MOS: Tezanos Pin do, Huidofcro 3647 (MO). Formosa: Espinilloa
Morel 7196 (MO); La Frontera, Morel 8380 (MO); Siete Palmas, Morel 8448 ! (MO)
misiones: Oro Verde, Schwarz 7826 (MO); Posadas, Scnu>arz 5649 (MO), salta: Balboa,'

334 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Cabrera 3135 (NY); Coronel Moldes, Meyer 3748 (NY). Santiago del estero: Yutuyacu,

^"Saguay: Villarica, Jorgensen 3904 (MO, NY). Ypacaray, Hassler 12198 (MO);

S " ° C B0LmA? S LA paz: Coripata, Bang 2083 (MO), Buchtien 8257 (NY); Coroico, Buchtien
s.n. (MO), santa cruz: Yorochito, Steinbach 8146 (MO), without province: Bang 574
(MO).

Ecuador: Albermarle I, (Galapagos I) Stewart 1434 (MO).

Siam: Zimmerman 146 (MO).

Hawaii: oahu: Fosberg 9388a (MO).

Pollen grains single, prolate, spheroidal, ca 35^ (E) X ca 35> (P), 15 colpate,
5 at each pole and 5 perpendicular to the equator, colpi ca 11/z long, exine ca
2-2.5 n in thickness, sexine ± equal to nexine and very finely reticulated.

Pollen examined: Orcutt 5886 (MO); Pringle 7428 (MO); Ruth 229 (MO).

Rivina humilis is one of the most variable species in the Phytolaccaceae.
Walter (1909) recognized three species, R. humilis, R. portulaccoides Nutt., and
R. purpurascens Schrad., based on sepal color and size, erectness of the inflores-
cence, and relationship of leaf length to inflorescence length, all of which are
overlapping characters. I have reduced all to synonomy under R. humilis. Some
of the variability may result from the diverse weedy habitats in which Rivina
is frequently found— some specimens are almost "stunted" in appearance, wiry,
small leaves, short internodes, etc., and present a marked contrast to the more
robust collections. However, to give each of these variants species rank is unsound.

7. TRICHOSTIGMA
Trichostigma A. Rich, in Sagra, Hist. Fis. Pol. Nat. Cuba, Part 2, Hist. Nat.

10: 306, 1845. [Type T. octandrum (L.) H. Walter]
Rivinia Mill, Gard. Diet. Abridg. ed. 4, 3, 1754.
Villamilla Ruiz & Pavon ex Moq. in DC., Prodr. 13(2): 10, 1849.

Shrubs or trailing vines. Leaves ovate to elliptic, acute to long acuminate,
entire, the bases cordate to obtuse, punctate, ± glabrous to sparsely hairy on veins
beneath; petiolate. Inflorescences racemes, axillary or terminal. Flowers perfect,
± actinomorphic, pedicellate; bract single, awl-shaped to lanceolate; bracteoles
two, minute; sepals 4, ± equal, oblong, green to white; stamens 8-25, separate,
irregularly deposited in two whorls, ± sessile or filaments filiform, the anthers
linear; ovary 1-carpellate, cylindrical to ± globose, the style short or absent, the
stigma generally penicellate. Fruit a drupe, ± globose, black to red-purple; seed
1, lens shaped, the testa red-brown.

Northern South America, Central America and the West Indies; a genus of
three species.

Pollen grains subprolate or prolate spheroidal, ca 27-37^ (E) X ca 27-35^
(P), 3-colpate with colpi 22-23^ long, or 15 colpate, 5 at each pole and 5 per-
pendicular to the equator with colpi ca 4.5-6^ long, exine ca 2-2.3^ in thickness,
sexine ± equal to nexine or slightly thicker than nexine, and ± smooth.

The results of pollen analysis, 3-colpate grains in Trichostigma octandrum L.,
and 15-colpate in a 5-5-5 pattern for T. polyandrum (Loes.) H. Walter and T.
peruvianum (Moq.) H. Walter, are perplexing in view of the very similar floral

NOWICKE — PHYTOLACCACEAE 335

and vegetative morphology, especially of T. octandrum and T. polyandrum. Al-
though the two pollen types are very different in terms of aperture structure, I
make no recommendation for division of the genus based solely on this feature.

a. Leaf bases cordate, petioles pubescent _ 1. T. peruvianum

aa. Leaf bases obtuse, petioles glabrous.

b. Stamens 20-25; inflorescences few and more than 10 cm long, the pedicels

10 mm or longer ...2. T. polyandrum

bb. Stamens 8-10; inflorescences many and up to 10 cm long; the pedicels up

to 10 mm long 3. T. octandrum

1. Trichostigma peruvianum (Moq.) H. Walter, Pflanzenr. IV, 83 (Heft 39): 111,

1909.
Rivina peruviana Moq. in DC, Prodr. 13(2): 10, 1849. (Type Matthews 1455 G, K)
Villamilla tinctoria Ruiz & Pavon Fl >l 402) ex Moq., loc. cit.

Ledenbergia roseo-aenea Lem., Ulustr. Hort. 16: pi. 591, 1869.
Villamilla peruviana Hook. f. in Benth. & Hook, f., Gen. PI. 3 : 81, 1880.
Rivina roseoaenea (Lem.) O. Ktze., Rev. Gen. PL 2: 551, 1891.
Villamilla roseo-oenia (Lem.) Rusby, Mem. Torrey Bot. Club 6: 110, 1896.

Shrubs to 2 m. Leaves ovate, acuminate, entire, the bases cordate, up to 27
cm long and 11 cm wide, sparsely hairy on veins beneath; petioles 2-3 cm long,
conspicuously pubescent. Inflorescences mostly terminal, 25-30 cm long, rachis
pubescent. Flowers with pedicels up to 8 mm long; bract single, 1.5 mm long;
bracteoles two, ca 0.3-0.4 mm long, triangular; sepals ca 4 mm long; stamens 10-12
usually in two whorls, the filaments ca 1.5-2 mm long, the anthers ca 1.5-1.8 mm
long; style short and thick or absent. Drupe ca 4-6 mm in diam, black.
Peru and rarely Ecuador.
Ecuador: s. loc, Mexia 7221 (US).

Peru: loreto: Balsapuerto, Killip & Smith 28690 (F); Klug 2976 (F, MO); Soledad,
Killip & Smith 29696 (F); betw Yuramaguas & Balsapuerto, Killip & Smith 28344 (F).
san martin: San Roque, Williams 6935 (F).

Pollen grains prolate spheroidal, ca 30-32^ (E) X ca 30-32^ (P), 15-colpate,
5 at each pole and 5 perpendicular to the equator, colpi ca 4.5-5.5)/ long, exine ca
2fi in thickness, sexine ± equal to nexine and ± smooth.
Pollen examined: Klug 2976 (MO); Williams 6935 (F).

2. Trichostigma polyandrum (Loes.) H. Walter, Pflanzenr. IV, 83 (Heft 39): 112

1909.
Rivina polyandra Loes., Bot. Jahrb. 23: 123, 1896. (Type Rothschuh 114 B|)
Villamilla polyandra (Loes.) H. Walter, loc. cit. 37(Beibl. 85): 24, 1906.

Shrubs, ± erect or trailing vines. Leaves ovate to elliptic, acute to long
acuminate, the bases obtuse, up to 16 cm long and 8 cm wide, ± glabrous; petioles
2-3 cm long. Inflorescences mostly terminal, occasionally axillary, up to 25 cm long.
Flowers with pedicels to 15 mm long; bract single, ca 1.5-2 mm long; bracteoles
two, ca 0.5 mm long; sepals 4-5 mm long, enlarging in fruit to ca 8-9 mm long;
stamens 20-25, deciduous, sessile to filaments 0.6 mm long, the anthers ca 2 mm
long; the style short, the stigma sparsely penicellate. Drupe ca 4.5-5.5 mm in diam,

336 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Central America.

Costa Rica: Rojas 441 (MO); Standley & Valedio 44442 (US), 46272 (US).

panama: von Wedel 714 (MO), 752 (MO), 926 (MO), 940 (MO), 1439 (MO), 7547
(MO), 2577 MO); Woodson et al. 1832 (MO).

Pollen grains prolate spheroidal, ca 27-29/* (E) X ca 27-29^ (P), 15 colpate,
5 at each pole and 5 perpendicular to the equator, colpi ca 5-6// long, exine ca
2.3fi in thickness, sexine ± equal to nexine or slightly thicker than nexine, and
± smooth.

Pollen examined: Standley & Valeria 44442 (US), 46275 (US).

3. Trichostigma octandrum (L.) H. Walter, Pflanzenr. IV, 83 (Heft 39) : 109, 1909.

Rivina humilis var. scandens L., Sp. PI. 122, 1753.

R. octandra L, Cent. PI. 2:9, 1756. (Type LINN, not seen; from IDC Micro-Edition

163.3)
7?. dodecandra Jacq. Obs. Bot. 1 : 6, 1764.
R. scandens Mill., Gard. Diet. ed. 8, Rivinia no. 2, 1768.
R. mutisii Willd. ex. Schultes, Mant. 3: 305, 1827.
R. americana Rat, FI. Tellur. 3: 56, 1837.

Trichostigma rivinoides A. Rich, in Sagra, Hist. Cuba 10: 306, 1845.
Rivina octandra L. var. obtusifolia Moq. in DC, Prodr. 13(2) : 11, 1849.
R. ehrmbergiana Klotzsch ex Moq., loc. cit, nom. nud. pro syn. R. octandra.
R. moritziana Klotzsch ex Moq., loc. cit., nom. nud. pro syn. R. octandra.
VillamiUa octandra Hook. f. in Benth. & Hook, f., Gen. PI. 3: 81, 1880.

Shrubs, decumbent or woody vines. Leaves elliptic, acuminate, entire, the bases
obtuse, up to 13 cm long and 6 cm wide, glabrous or pubescent; petioles ca 2-3 cm
long. Inflorescences ca 10 cm long, mostly axillary. Flowers crowded, with pedicels
ca 6-7(-10) mm long at maturity; bract single, ca 2-3 mm long, awl-shaped;
bracteoles two, ca 0.4-0.5 mm long; sepals 2.5-4 mm long and 1.5-2.5 mm wide,
white to green; stamens 8-10, the filaments ca 2-3 mm long, the anthers ca 1.5-2
mm long; style absent, the stigma penicellate. Drupe ca 4-5 mm in diam, black.

Widely distributed in Central America, the West Indies and South America.

United States: Florida: Small & Matthaus 9904 (MO).

Mexico: Guerrero: Hinton 14154 (MO), morelos: Pringle 8491 (MO), sinaloa:
Lamb 418 (MO), tamaulipas: Palmer 349 (MO), without state: Rose 14677 (US).

British Honduras: Gentle 4028 (MO).

Costa Rica: Smith 1672 (MO).

Panama: Allen 944 (MO), 17279 (MO); Terry & Terry 1396 (MO).
ght 471 (MO).

Guadeloupe: Duss 2399 (MO); Bertero s.n. (MO).

mard 9873 (MO) ; Leonard & Leonard 13623 (MO).

Jamaica: Harris 11966 (MO); Yuncker 18185 (MO), 78345 (MO).

Puerto Rico: Sintenis 3931 (MO), 70766 (MO).

St. Croix: Ricksecker 343 (MO); Ricksecker 325 (MO).

Tobago: Broadway 2967 (MO).

Trinidad: Broadway 5118 (MO).

Colombia: magdalena: Santa Marta, Smith 1718 (MO, NY).

Venezuela: merida: Colonia Tovar, Fendler 1087 (MO, NY), monagas: W of Santa
Barbara, Steyermark 61768 (MO), without state: Punta Predra I, Rusby & Squires 419
(MO).

Brazil: amazonas: mouth of Rio Embira, Krukoff 4872 (MO); Sao Paulo de Olivenca,
Ducke 404 (MO) ; Rio Solimoes, Krukoff 4507 (MO).

Argentina: sa Iver, Eyerdam & Beetle 22800 (MO), misiones: Posadas,

Ekman 1884 (GH).

NOWICKE — PHYTOLACCACEAE 337

Bolivia: pando: jet of Beni & Madre de Dios Rivers, Rusby 741 (MO), santa cruz:
Missiones Guarayos, Werdermann 2648 (MO).

Peru: san mart i 4185 (MO), 4364 (MO).

Pollen grains subprolate, ca 28-29^ (E) X ca 34-35^ (P), 3-colpate, colpi ca
22-23/t long, exine ca 2fi in thickness, sexine ± equal to nexine and ± smooth.
Pollen examined: Allen 944 (MO), 17279 (MO).

Excluded collection: Pollard & Palmer 340 (MO, NY). This is a fruiting speci-
men in which the robustness of the inflorescence, i.e. length of pedicel, size of sepals
and overall inflorescence length, is characteristic of T. polyandrum, whereas the
geographical location and intermediate stamen number, ca 13-14, are typical of
T. octandrum.

8. SCHINDLERIA
Schindleria H. Walter, Bot. Jahrb. 37(Beibl. 85): 24, 1906. [Lectotype selected:
S. racemosa (Britton) H. Walter]

Shrubs, rarely herb-like with woody bases, drying black or yellow-green. Leaves
elliptic or elliptic to ovate-elliptic, acuminate or acute-acuminate, entire, the bases
obtuse, slightly cordate, or rarely ± oblique, glabrous, or sparsely to densely pubes-
cent; petiolate. Inflorescences racemes, in some appearing as a delicate spike in bud,
mostly axillary, or axillary and terminal. Flowers perfect, actinomorphic, pedicel-
late; bract single, lanceolate and keeled at the base or awl-shaped; bracteoles two,
minute, closely appressed to sepals; sepals 4, ± equal, oblong and ± rounded, veins
barely discernible or ± prominent; stamens 12-25, irregularly deposited, the fila-
ments filiform, the anthers linear; ovary 1-carpellate, ± cylindrical to ovoid, ±
compressed laterally, style absent, stigma penicellate. Fruit an utricle, compressed
laterally, green-brown; seed 1, lens shaped, testa shiny black.
Two species in Peru and Bolivia.

Pollen grains single, subspheroidal, ca 17-21^ (E) X ca 17-21^ (P), 12-17
pantoporate, ora ca 3-3.5^ in diam, exine ca 2-2.5^ in thickness, sexine ± equal to
nexine and finely reticulated.

I have reduced the six species listed by Heimerl ( 1934) to two, which are best
separated by the striking color differences in the dried specimens. Schindleria
racemosa (Britton) H. Walter has inflorescences and leaves drying black or brown-
black and includes S. glabra H. Walter and S. mollis H. Walter, both of which
were based on single collections.

Schindleria densiflora (O. Ktze) Monachino, retaining its green color or be-
coming yellow on drying, comprises a variable group of specimens in terms of leaf
base and texture. In flower structure, however, the group is much more uniform
and no character or set of characters serves to separate this complex into distinctive
alliances. Monachino (1952) first proposed this treatment, although admittedly on
insufficient material, and elucidated the nomenclatural problems involved with it.
With the exception of Bang 2607, the type of S. rivinoides H. Walter and about
which I maintain some reservations concerning conspecificity with S. densiflora,
members of the group are coarse, weedy semi-shrubs found in southern Peru and
northern Bolivia.

a. Plants turning black or brown-black

aa. Plants remaining green or turning yellow upon dessication 2. S.

[Vol. 55
338 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1. Schindleria racemosa (Britton) H. Walter, Bot. Jahrb. 37(Beibl. 85): 24, 1906.

(Lectotype selected: Bang 414 MO; isolectotypes F, NY, US; syntypes Rusby

743 K, MO, NY)
Villamilla racemosa Britton in Rusby, Mem. Torrey Bot. Club 4:251, 1895. (Syntypes

Bang 414 NY; Rushy 743 NY)
Schindleria glabra H. Walter, Bot. Jahrb. 37(Beibl. 85): 24, 1906. (Type Weherbauer 1168

Bt, not seen)
S. mollis H. Walter, Pflanzenr. IV, 83 (Heft 39): 116, 1909. (Type Lobb 691, photo F,

from W)

Shrubs, or herbs with woody bases, black on drying. Leaves elliptic, acute-
acuminate, entire, the bases obtuse, up to 15 cm long and 6 cm wide, slightly to
very pubescent, especially on the veins beneath; petioles ca 1.5-4 cm long. In-
florescences racemes, up to 28 cm long, axillary or terminal. Flowers with pedicels
ca 6-8 mm long; bract single, ca 1.2-1.5 mm long, lanceolate and keeled at the
base; bracteoles two, minute, ca 0.1-0.2 mm long, triangular and closely appressed
to sepals; sepals ca 2-3.5 mm long and 1.5-1.7 mm wide, veins barely discernible;
stamens ca 12-15, the filaments ca 1.2-2 mm long, the anthers ca 1.2-1.5 mm long;
ovary slightly elongate. Utricle ca 2 mm in diam.

Bolivia: la paz: Colaya, Mexia 4302 (GH); Coroico, Buchtien 3772 (NY, US); Yungas,
Bang 414 (F, MO, NY, US); Rusby 743 (K, MO, NY, US), santa cruz: Bella Vista,
Buchtien 6291 (US).

Pollen grains ca 2\(x (E) X ca 21^ (P), 12-15 pantoporate, ora ca 3^ in diam,
exine ca 2.5^u in thickness.

Pollen examined: Rusby 743 (MO).

2. Schindleria densiflora (O. Ktze.) Monachino, Phytologia 4: 39-41, 1952.
Rivina densiflora O. Ktze., Rev. Gen. PI. 3(3): 268, 1898. (Type Kuntze s.n. NY, not seen)

'alter, Bot. Jahrb. 37(Beibl. 85): 24, 1906. (Type Bang 1292 MO)
S. rivinoides H. Walter, loc. cit. (Type Bang 2607 MO)

S. weberbaueri O. C. Schmidt, Notizbl. Berlin Bot. Gart. 8:313, 1923. (Type Weherbauer
6758 F, GH, MO, NY)

Shrubs, or herbs with woody bases, to 3 m, green to yellow on drying. Leaves
elliptic to ovate-elliptic, acuminate, entire, the bases obtuse, slightly cordate or
rarely ± oblique, size variable, up to 20 cm long and 7 cm wide, mostly glabrous,
some sparsely pubescent on veins beneath; petioles to 8.5 cm long. Inflorescences
racemes (appearing as delicate spikes in bud), up to 35 cm long (in fruit), mostly
axillary, rarely terminal. Flowers with pedicels ca 7-9 mm long at maturity; bract
single, ca 1 mm long, awl-shaped; bracteoles two, ca 0.2 mm long, closely appressed
to sepals; sepals 2-3 mm long, some prominently veined; stamens ca 15-25, the
filaments ca 1.5 mm long, the anthers ca 1 mm long. Utricle ovoid, compressed
laterally and ridged on the edges, ca 1.8-1.9 mm in diam, testa wrinkled, yellow-
Bolivia and Peru.

Bolivia: cochabamba: Chimore, Cardenas 5482 (US); s. loc, Steinbach 9361 (GH,
NY), la paz: Mapiri, Buchtien 1694 (GH, US), santa cruz: Samaipata, Steinbach 8201
(GH). without province: N Yungas, Buchtien 4327 (US); s. loc, Bang 2607 (F, GH,
MO).

Peru: ayacucho: betw Huanta & Rfo Apurimac, Killip & Smith 23092 (NY, US).

NOWICKE — PHYTOLACCACEAE 66V

cuzco: Lares Valley, Weberbauer 7924 (F, GH, MO, NY); Manto-Lares, Marin 2164 (F);
Hacia Pilcopata, Vargas 13282 (US); Quincemil, Vargas 7746 (MO); betw Santo Isabel
& Mistiana, Scolnik 906 (NY), huanuc. kowski 34509 (MO); Tal des

Mayro, Weberbauer 6758 (F); Tingo Maria, Asplund 12488 (US), madre de dios: s. loc,
Vargas 16931 (US), puno: vie of Santo Domingo, McCarroll 54 (NY).

Pollen grains ca 2Lu (E) X ca 2\p (P) s 12-17 pantoporate, ora ca 3-3.5^ in

Pollen examined: Woytkowski 34509 (MO).

9. HILLERIA
Hilleria Veil., Fl. Flumin. 47, 1: pi 122, 1825. [Type H. latifolia (Lam.) H.

Walter]
Mohlana Mart., Nov. Gen. Sp. PI. 3: 171, pi. 290, 1829.

Herbs, robust, some species suffrutescent at the base, to 1.5 m. Leaves ovate to
elliptic, acute, acuminate, or long acuminate, entire, the bases obtuse, ± glabrous
above to sparsely hairy on the veins beneath; petiolate. Inflorescences racemes,
axillary or terminal, ± black on drying. Flowers perfect, ± zygomorphic, pedicel-
late; bract single, awl-shaped; bracteoles two, minute, visible only in bud; sepals
4, unequal, the lowermost enlarging conspicuously in fruit and somewhat uniting
at the base with the 2 laterals; stamens 4 or 8-13, alternate or deposited irregularly,
the filaments filiform, the anthers 2X longer than broad; ovary spherical or slightly
compressed laterally, 1-carpellate, the styles present or absent, the stigma feathery
or capitate. Fruit an utricle, lens-shaped, ridged on the edges, pericarp wrinkled;
seed one, testa shiny black.

Northern South America and introduced into Africa; a genus of 3 species.

Pollen grains single, subspheroidal, ca 23-29^ (E) X ca 23-29^a (P), 12-
colpate, 4 at each pole and 4 perpendicular to the equator, colpi ca 6-9^ long,
sometimes ± indistinct, exine ca 2-2.5^ in thickness, sexine ± equal to nexine
and finely to very finely reticulated (Fig. 5).

Young specimens of Hilleria, i.e. those not in late flowering or fruit, can be
confused with Schindleria H. Walter, especially that section of the latter genus
which turns black upon dessication. However, the pollen of the two genera, 12-
colpate in Hilleria, and polyporate in Schindleria, provides a means for unmistak-
able identification.

a. Stamens 8-13; stigmas feathery, ± sessile 1. H. longifolia

aa. Stamens ± 4; stigmas capitate.
Style absent, stigma

bb. Style short, ca 0.2-0.3 mm long, i

1. Hilleria longifolia (H. Walter) Heimerl, Oesterr. Bot. Zeitschr. 61: 10, 1911.
H. latifolia (Lam.) H. Walter var. longifolia H. Walter, Pflanzenr. IV, 83 (Heft 39) : 82,

1909. (Type Poeppig 1541 W)

Herbs, erect, to 1-2.5 m. Leaves elliptic, ± long acuminate, entire, the bases
obtuse, up to 23 cm long and 6 cm wide, glabrous above, coarsely hairy on veins
beneath; petioles up to 6 cm long. Inflorescences up to 40 cm long, terminal or
axillary. Flowers with pedicels to 6 mm long; bract single, up to 4 mm long;

340 ANNALS OF THE MISSOURI BOTANICAL GARDEN

bracteoles two, ca 0.2 mm long; sepals at maturity ca 3.5 mm long for lowermost,
ca 3 mm long for remaining 3; stamens 8-13, irregularly deposited, the filaments ca
0.8-0.9 mm long, the anthers ca 1 mm; style absent, the stigma feathery. Utricle
ca 2.5 mm in diam.

A rare species reported only from Peru and Bolivia.

Bolivia: suryungas: Colaya, Mexia4302 (MO).

Peru: cuzco: Quincemil, Marin 1536 (US), san martin: Margarita, Ferreyra 1017
(US); vie of Tingo Maria, Allard 22267 (US).

Pollen grains ca 23^ (E) X ca 23^ (P), colpi ca 8-9^ long, sexine medium
finely reticulated.

Pollen examined: Allard 22267 (US).

2. Hillerm latifolia (Lam.) H. Walter, Pflanzenr. IV, 83 (Heft 39) : 81, 1909.
Rivina latifolia Lam., Encycl. Meth. Bot. 1: 324, 1791. (Type Martin s.n. P)
R. affinis Nees & Mart, Nov. Acta Acad. Nat. Cur. 11 : 30, 1823.

Flumin. 47, 1 : pi 122, 1825.
Rivina apetala Schum. & Thonn., Beskr. Guin. PI. 84, 1827.
Mohlana nemoralis Mart., Nov. Gen. Sp. PI. 3: 171, pi 290, 1829.
M. gumeensis Moq. in DC, Prodr. 13(2): 15, 1849. (Type Schumann s.n., location un-

M. latifolia (Lam.) Moq, loc. cit. 16. (Type Martin s.n. P)

M. apetala (Schum. & Thonn.) Engler, Pflanzenwelt Ost-Afr. 5: 175, 1895.

Herbs, slightly suffrutescent at the base, to ca 1 m. Leaves ovate to elliptic,
acuminate to long acuminate, entire, the bases obtuse, up to 20 cm long and 6 cm
wide, ± glabrous above, and coarsely hairy on the veins beneath; petioles to 8 cm
long. Inflorescences up to 30 cm long, mostly axillary, rarely terminal. Flowers
with pedicels up to 5 mm long; bract single, ca 2 mm long; bracteoles two, ca 0.2
mm long; sepals at maturity ca 4 mm long for lowermost, ca 3 mm long for re-
maining 3; stamens 4, alternate with the sepals, the filaments ca 0.8 mm long, the
anthers ca 0.8 mm long; the style absent, the stigma capitate. Utricle ca 2 mm in

South America and introduced into Africa.

Colombia: cundinamarca: Quebrada Cabana, Killip et al. 38373 (US).

Ecuador: guayas: Hacienda Barcelona Trail, Gilmartin 555 (US).

Peru: cuzco: betw Victoria & Echarate, Vargas 7555 (MO, US), huanuco: Pozuzo,
Machride 4625 (US); Tingo Maria, Asplund 12039 (US), junin: La Merced, Soukup 3373
(US), loreto: lower Rfo Huallaga, Killip & Smith 28904 (US), 29079 (US), san martin:
Rioja, Woytkowski 6127 (US); Zepelacio nr Moyobamba, Klug 3438 (MO, US).

Bolivia: beni: vie of Rurrenabaque, Cardenas 1771 (US), sara: Rio Palometillas,
Steinhach 6799 (MO).

Paraguay: Hassler 8287 (MO).

Argentina: salta: Rio Blanco, Venturi 7634 (US); vie of Tartagal, West 8417 (MO).

Ivory Coast: Leeuwenherg 4142 (MO); Roherty 12376 (MO).

Ghana: Darko 631 (MO); Oldeman 757 (MO).

Nigeria: Ross 153 (MO).

Cameroons: Bates 673 (MO); Staudt 922 (US); Zenker 316 (MO, US), 4628 (MO);
Zenker & Staudt 15 (US).

Republic of Congo: Corbisier 745 (MO); Germain 229 (MO); Louis 924 (MO), 7318
(MO, US), 77087 (MO), 77095 (MO).

Uganda: Dummer 327 (US), 459 (MO, US).

NOWICKE— PHYTOLACCACEAE 341

Pollen grains ca 29 fi (E) X ca 2fy (P), colpi 6-7 fi long, sexine very finely
reticulated (Fig. 5).

Pollen examined: Bates 673 (MO); Dummer 459 (US); Louis 924 (MO);
Woytkowski 6127 (US).

Hilleria latifolia and H. secunda (Ruiz & Pavon) H. Walter are very difficult
to distinguish. Frequently the styles are broken and what appears at first glance to
be H. latifolia may be classifiable as H. secunda when a sufficient number of flowers
is examined. Also, some specimens are almost undeterminable because of the inter-
mediate character of the style length, which is difficult at best to observe because of
its small size.

3. Hilleria secunda (Ruiz & Pavon) H. Walter, Pflanzenr. IV, 83 (Heft 39) : 82,

Rivina secunda Ruiz & Pavon, Fl. Peru Chile 1 : 65, pi. 102, 1794. (Type Ruiz & Pavon s.n.

R. acuminata H.B.K., Nov. Gen. Sp. PI. 2: 184, 1817.

Mohlana secunda (Ruiz & Pavon) Mart., Nov. Gen. Sp. PI. 3: 172, 1829.

Rivina inaequalis Hook., Icon. PI., pi. 130, 1837. (Type Mathews 1604, location unknown)

Mohlana secunda var. acuminata (H.B.K.) Moq. in DC, Prodr. 13(2): 15, 1849.

Hillera secunda (Ruiz & Pavon) O. Ktze., Rev. Gen. PI. 2: 551, 1891.

Mohlana meziana H. Walter, Bot. Jahrb. 37(Beibl. 85): 25, 1906. (Type Ule 6500 Bt, K)

Hilleria meziana H. Walter, Pflanzenr. IV, 83 (Heft 39): 83, 1909. (Type Ule 6500 B|, K)

Herbs, slightly suffrutescent at the base, erect, to ca 1 m. Leaves ovate to
elliptic, acute to acuminate, entire, the bases obuse, up to 15 cm long and 6 cm
wide, coarsely hairy on the veins beneath; petioles to 6 cm long. Inflorescences up
to 20 cm long, mostly axillary. Flowers with pedicels ca 5 mm long; bract single,
ca 1.5 mm long, much overtopping the flowers in bud; bracteoles two, ca 0.2 mm
long; sepals at maturity ca 3.5 mm long for lowermost, ca 2.25 mm long for re-
maining 3; stamens 4-7, alternate with the sepals, or ± deposited irregularly, the
filaments ca 0.8-1 mm long, the anthers ca 1 mm long; style ca 0.2-0.3 mm long,
the stigma capitate. Utricle ca 2 mm in diam.

Northern South America.

Colombia: cundinamarca: Icononzo, Pennell 2762 (MO, US) ; La Mesa, Fernandez &
Mora 1386 (US), tolima: vie of Totare River, Haught 2395 (A, US).

Venezuela: without state: Quebrada de Chacaito, Pittier 12996 (MO, US).

Ecuador: guayas: Manglar Alto, Anthony & Tate 14 (US), manabi: El Recreo, Eggers
15513 (US) ; N of La Tuna, Haught 3364 (US).

Peru: cajamarca: Monte Seco: Soukup 3866 (US), huanuco: Tingo Maria, Asplund
12129 (US), junin: La Merced, Sandeman 5037 (US), loreto: Tarapoto, Ule 6500 (K).
san martin: Alto Rio Huallaga, Williams 5757 (US), 6884 (US); Rio Mayo, Ferreyra 7813
(US); San Roque, Williams 7611 (US); N of Tingo Maria, Allard 20923 (US), 21822

Bolivia: beni: San Buenaventura, Williams 343 (US), la cruz: s. loc, Kuntze s.n
(US).

Pollen grains ca 25^ (E) X ca 25^ (P), colpi ca 8-9^ long, exine ca 2fi in
thickness, sexine finely reticulated.

Pollen examined: Ferreyra 7813 (US); Haught 2395 (US); Williams 7611
(US).

ANNALS OF THE MISSOURI BOTANICAL (

10. LEDENBERGIA

in DC, Prodr. 13(2): 14, 1849. (Type L. s

Trees or shrubs. Leaves alternate, elliptic to ± ovate, acuminate to acute,
entire, the bases obtuse, glabrous to slightly pubescent; petiolate. Inflorescences
racemes, mostly axillary, pendulous. Flowers perfect, or uni