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This original edition (issued Chg ece7 15 1936, 
is limited to six hundred and seventy-five copies, 

of which this is Number 40 

After a painting by Charles R. Knight in 1908, under the direction of Henry Fairfield Osborn 

This restoration is based on the classic skeleton known as the Warren Mastodon, which was unearthed in the year 1845 just 
north of the Highlands of the Hudson and presented by the late J. Pierpont Morgan to the Hall of the Age of Man of The American 
Museum of Natural History in the year 1906. The animal took its name from John C. Warren, Professor of Anatomy at Harvard 
University, who described it in detail in his memoirs of 1852 and 18598. 

The painting represents the animal, a bull at least thirty years old, as he appeared in life before he sank into a shell-marl 
basin six miles northwest of Newburgh-on-Hudson, New York; this event, so auspicious to palzontologists of today, occurred in 
the Late Glacial Epoch between 30,000 and 40,000 years ago. In the right background is the north gap in the Hudson River High- 
lands, Storm King Mountain on the right and Break Neck on the left. In the foreground is one of the shallow sheets of water 
covering the shell-marl in which the remains of the animal were so perfectly preserved. The flora of the forest corresponds with the 
description by Asa Gray, in a letter to Doctor Warren, of the contents of the mastodon’s stomach: “An examination with the micro- 
scope showed, from the structure of the woody fibre, that they were boughs of pine or spruce of some sort, and that they minutely 
agreed with the wood of hemlock spruce; so that this is very probably the species they belonged to, but there is no certainty Ofte 





A.B. Princeton, 1877; D.Sc. Princeton, 1880; Honorary LL.D. Trinity, 1901; LL.D. Princeton, 1902: 
Sc.D. Camsrince, 1904; LL.D. Cotumaia, 1907; Px.D. Curistianta, 1911; D.Sc. Yate, 1923: 
D.Sc. Oxrorp, 1926; D.Sc. New York, 1927; LL.D. Union, 1928; Docror' 
oF THE University oF Paris, 1931; Doctor or NATURAL SCIENCE, 
Jouann Wo rGaAnc Goetue University, 1934 

ResEARCH ProressoR OF ZOOLOGY, CotumMBIA University; Honorary Curator-In-Cuter oF VERTEBRATE 
Unirep States Geotocicat Survey; Honorary Prestpent, THE AMERICAN 
Museum or Natura History; Honorary Presipent, 

Tue New York Zoo.ocicat Society 



NEW YORK, 1936 


Volume I has been on the American Museum Press since 1924. In the course 
of the intervening eleven-year period to 1935 the field of research has been greatly 
broadened and many discoveries have been made which alter the Classification of 
the Proboscidea; the consequent alterations appear in the Table of Contents and 
in the Phylogenetic Appendix to the present volume. 

At the time of the author’s lamented death on November 6, 1935, the final 
revision of the manuscript was in progress. This has been carried out along the 
lines laid down by him; since that time no changes have been made in his deter- 
minations and necessary corrections have been inserted as footnotes. 

Volume II, containing Elephantoidea, Appendices, and Index, will be issued 
as soon as possible, probably in 1937. 

Coryriaut, 1936, By 
Tue American Museum or Natura History 

D: B.D, 


DuRING THE YEARS 1907 TO 1936 


1837 — 1913 


Tur AMERICAN Museum or NATURAL Hisrory 


HE author began his researches for this Monograph by devoting the years 1889-1900 to zoogeographical 
| Fea on the origin, migration and distribution of the twenty-two Orders of Mammals known at that 

time. The theoretic conclusion was reached that the Proboscidea originated in Africa. This conclusion was 
confirmed in the Faytim district of northern Africa by Hugh Beadnell (1901-4) and by Charles William Andrews 
(1901-3). In Andrews’ Memoir of 1906 published by the British Museum (see p. 36 of the present monograph) 
he wrote: 

The probability that Africa would be found to be the original home of these animals was pointed out by several writers, 
notably Osborn, Stehlin, and Tullberg. The first of these suggested that probably not only the Proboscidea but also the 
‘Hyracoidea, certain Edentates, the Antelopes, the Giraffes, the Hippopotami, the most specialized Ruminants, and among the 
Rodents the Anomalures, the Dormice, and Jerboas, among Monkeys the Baboons,’ and, as his map suggests, the Sirenia also, 
originated in this region. Osborn also put forward the theory that a succession of migrations from Africa to Europe occurred, 
notably at the end of the Eocene, at the beginning of the Miocene, and again in the earliest Pliocene. It was in the early 
Miocene migration that the Proboscidea passed out of Africa for the first time so far as known. 

The American Museum expedition of 1907 amplified the epoch-making discoveries of Beadnell and Andrews 
and subsequent exploration has demonstrated beyond question that in Africa originated not only the Mceritheres 
and primitive Mastodonts but also the Deinotheres and the Elephants. Not a trace of these proboscideans has 
been found in any other continent earlier than Miocene time, when waves of Deinotheres and Mastodonts, already 
highly specialized, began to enter Eurasia and to migrate to all the continents excepting Australia. 

The actual preparation of this monograph opened in the year 1907 with the author’s expedition to the desert 
bordering the Faytim of northern Egypt, accompanied by Walter Granger of the mammalian paleontology staff 
of the American Museum. In 1908 the author determined to give the Proboscidea a monographie treatment 
similar to that he was then giving to the Titanotheres. Intensive work on the Proboscidea, first entered 
upon in 1920 during the concluding phases of work on the Titanotheres, has continued without interruption for 
fifteen years. Thus the intensive study of the evolution and phylogeny of two dominant and widely different 
types of mammals, the Titanotheres and the Proboscideans, has occupied a total uninterrupted research period of 
thirty-five years, 1900-1935. 

The Titanothere Monograph involved methods of collateral research in geology, comparative anatomy, 
animal mechanics and biology that were largely new to the science of paleontology, chiefly in the science of measur- 
ing the skull (craniometry) and in the science of animal ascent (phylogeny). From the outset the craniometric 
methods and results of the Titanothere researches partially prepared the way for the solution of proboscidean 
problems. In their adaptations the Proboscidea present the widest possible contrasts to the Titanotheres. 
Moreover, there was little conception of the new and rapidly oncoming difficulties in interpreting the classic 
proboscideans as a whole or the entirely unexpected adaptations in numerous new lines of mastodont and ele- 
phantoid ascent. Previously undreamt-of kinds of proboscideans have been discovered. Through the classic 
and the new kinds have been revealed a number of biological principles hitherto unknown. 



In the analysis and synthesis of thousands of observations on the Proboscidea it soon became obvious that 
five main lines of research were essential to a judicial, comprehensive and philosophical treatment: 

First, to survey, for the restatement of historic descriptions and figures, the ancient literature of what 
may be called “the dawn period’ of mammalian paleontology, including Robert Kerr’s naming of the 
American mastodon, Elephas americanus, in 1792 and Blumenbach’s naming of the European mammoth, £. 
primigenius, in 1799; second, to study all the subsequent scientific descriptions of fossil and living proboscideans, 
with republication of essential parts of the original descriptions and reproduction of all the original figures; third, 
to directly or indirectly reéxamine all the fossil proboscidean collections in the world and especially to verify 
and refigure all types of genera and species; fourth, to secure the codperation of mammalian paleontologists all 
over the world in the examination of distant collections inaccessible to the author; fifth and most important, to 
approach this great and difficult subject with an entirely open mind, unbiased by preconceptions, each problem to 
be considered in a fresh and free spirit of observation, induction and generalization. 

Thus approached, the Proboscidea have proved infinite in variety, full of surprises, creative of new ideas and 
demonstrative of new evolutionary principles. 


A research period of unparalleled difficulty proved to be also a period of unparalleled opportunity—oppor- 
tunity such as has never before been afforded to any zoologist, paleontologist or biologist. There has been vast 
expansion of our knowledge since the year 1859 in which appeared Darwin’s ‘‘Origin of Species,” since 1886 when 
Richard Lydekker catalogued the Proboscidea in the great collection of the British Museum (Natural History), 
and even since 1929 when D. M.S. Watson’s article, ‘‘Proboscidea,” appeared in Volume XVIII of the Fourteenth 
Edition of the Encyclopedia Britannica. Even within the year 1935 most surprising discoveries have been made 
bearing on the origin and distribution, affinity and ascent of the Proboscidea, which as far as possible are em- 
bodied in the phylogenetic appendix at the close of the present volume. 

The outstanding result of the expansion of our knowledge of the Proboscidea is the demonstration, partly in 
fulfilment of the prophetic vision of Hugh Falconer, that the single genus Mastodon of Cuvier is the prototype of 
the suborder Mastodontoidea, which comprises all the mastodonts of the world, as fully set forth in the Appendix. 
This suborder embraces thirty-one distinct lines of generic ascent—in brief, of genera; these genera are grouped 
into fifteen distinct subfamilies, which in turn are grouped into four families, out of one of which it is possible 
that the Elephantoidea, the second great suborder of the Proboscidea, evolved. 

In like manner, as fully set forth in Volume II, the single genus Elephas of Georges Cuvier is now known to 
have divided into eight to ten separate lines of generic ascent, which in turn are grouped into four subfamilies, 
actually very wide apart in structure, in geographic distribution, in feeding habits and in adaptation. None of 
these elephantoid lines is known prior to Middle to Upper Pliocene time. The line with the most noble geologic 
and geographic history is that of the genus Archidiskodon, which apparently dated back to the Middle Pliocene of 

South Africa and enjoyed a glorious migration of 15,000 miles to its Late Pleistocene climax in Nebraska, Texas 
and Mexico. 

Thus, taken altogether, the Mastodonts and the Elephants, the Deinotheres and the Mceritheres—the four 

great branches constituting the great order PROBOSCIDEA—are now known to have evolved along no fewer 
than forty-one lines of generic ascent. 



The above revolution, by the discovery of multiple lines of ascent, is first manifested in the zoologie and 
classificatory treatment of the Proboscidea, in which we are impelled to substitute for the purely zoologie pre- 
evolutionary classification of Linnzus an evolutionary or phylogenetic classification based on principles first 
set forth by the author in 1892 and fully exemplified in the Titanothere Monograph (1929). Inherent in phylo- 
genetic classification are entirely new conceptions of the real evolutionary meanings of the old Linnzan terms 
Species and Genus, Family and Order, as well as of the more recent terms Superfamily, Subfamily, Subspecies 
and Mutations. The classic Linnean system based solely on zoologic observation and the creational concept 
is now replaced by a phylogenetic system in which all divisions from the Subspecies to the Order are placed 
vertically as succeeding each other during millions of years of geologic time, rather than horizontally as observed 
in recent or existing time by Linnzeus and all zoologists. 

The Osbornian phylogenetic system that was set forth in the Titanothere Monograph (Vol. I, Fig. 14, p. 16) 
is literally fulfilled in one proboscidean family known as the Serridentide, embracing seven complex lines of ascent 
of the serridentine mastodonts, namely, Serridentinus, Ocalientinus, Serbelodon, Trobelodon, Platybelodon, Toryno- 
belodon, Notiomastodon; also in an elephantoid subfamily known as Mammontinew, embracing three related but 
widely distinct kinds of mammoths—the woolly, the hairy, the southern—Mammonteus, Parelephas and Archi- 
diskodon. Thus for the first time we arrive at the real evolutionary significance of the classificatory terms of 
Linnzeus—Order, Suborder or Superfamily, Family, Subfamily, Genus, Species, Subspecies, Ascending Mutation. 

The facts upon which these phylogenetic results of mastodontoid and elephantoid ascent are established 
may be summarized as follows: 

1) The Proboscidea originated exclusively in Africa, probably in Upper Cretaceous or Lower Eocene time. 

2) Not remote from the stem that gave rise to the manatees and dugongs (Sirenia), proboscideans were distinguished 
from the first by specialization of the second incisor teeth above and below, also by the potentiality of forming transverse 
ridge-crests which, especially in the third superior and inferior grinders, rise from primitive 244-3 ridge-crests to progressive 
30 ridge-crests. 

3) The subfamily radiation into all habitats, from the purely aquatic and amphibious habitat of the Mceritheres to 
swamp, river-border, shallow lake-border, savanna, forest, tundra and desert habitat, from humid to arid climates, from 
equatorial to boreal latitudes and from sea level to elevated mountains (for example, the Andean mastodon), in which thirty- 
one lines of mastodonts and eight to ten lines of elephants have become specialized in every organ. 

4) The conversion of a broad prehensile upper lip of the hippopotamus type in the Meeritheres into the broad prehensile 
upper lip of many of the mastodonts and, finally, into the true proboscis of certain mastedonts and of all Stegodonts and ele- 

5) The genera are primarily distinguishable (a) by the adaptations of the grinding teeth which, when very carefully 
analyzed, as in the Appendix to the present volume, are highly characteristic, and (b) by the respective adaptations of the 
superior and inferior incisive teeth to a great variety of proboscidean floral habitats and modes of feeding; (c) in the Deino- 
theres, in three kinds of shovel-tuskers, also in the Rhynchotheres, the lower incisive tusks are far more important and diversi- 
fied than the upper, while in many mastodonts and in all the Stegodonts and elephantoids the lower tusks disappear entirely, 
the lower jaw being correspondingly abbreviated; (d) the superior tusks function first as offensive weapons of marked 
sexual variation and second as auxiliary feeding tools of great importance, excepting in one division of the elephantoids in 
which the gigantic superior tusks are withdrawn from feeding service and become only of sexual distinction and value. 

6) There is consequently frequent recurrence of adaptive parallelism, namely, of similarly specialized organs, like the 
shortened jaw being independently produced in different phyla, or the rounded and upturned enamel-less superior tusks 
serving as weapons. 

7) With the various functions and forms of the superior and inferior grinders, the superior tusks, the upper lip and the 
proboscis are coérdinated not only the cranium and jaws but also the neck, backbone, and limbs. For example, the three 
types of mammoths—the tundra (Mammonteus), the north temperate (Parelephas) and the southern (Archidiskodon)—alike 
exhibit enormous incurved tusks of use only in combat or for sexual selection purposes and not as auxiliary feeding organs; 
in these three mammoths the posterior half of the backbone is relatively weak because the tusks have relatively little work to 
do as auxiliary feeding organs, whereas in the African and Indian elephants and their relatives the backbone is extremely 
strong, in correlation with active tusks. 



Not revolutionary, but in accord with the clocking of geologic time by similar stages of evolution (homotaxis, 
Huxley) long in use by invertebrate palzontologists, is the evidence yielded by intensive examination which 
the Proboscidea afford for Tertiary geologic correlation of stages in adaptive progression and retrogression in the 
widely distant distribution centers of mastodonts and elephants which migrated into all the continents except 
Australia. The continuous evolution of far-separated adaptations of the grinding teeth, in northern and southern 
Afriea, in Eurasia and in North and South America, can now be coérdinated with unexpected precision, as in the 
outstanding metamorphosis of Archidiskodon proplanifrons of South Africa into the gigantic Archidiskodon 
maibeni of Nebraska (see Volume II). In this correlation the intensive study of the grinding teeth in the masto- 

dontoid and elephantoid divisions plays a leading part. 

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Another result which will prove to be revolutionary in anthropology is the new means afforded of dating 
precisely the main periods of the prehistoric evolution of Man, by intensive measurement of the length, width 
and thickness of the enamel layers in the grinding teeth of the elephantoid division (Osborn 1931.858) in successive 
stages from Upper Pliocene to closing Pleistocene time. Man was a mastodon and elephant hunter from early 
times. Remains of fossil elephants are occasionally found embedded in the same strata with remains of fossil 
man, and the total length of the enamel foldings in proboscidean grinding teeth enables us to date relatively the 
successive phases in the evolution of man. 

From the author’s prolonged thirty-five-year research on Titanotheres and Proboscidea there issue not 
only the principles governing the classic modifying modes of evolution known to Lamarck and Darwin (variation, 
development, degeneration) but also the newly discovered and hitherto unrecognized principle and modes of the 
origin of new characters through aristogenesis or creative biomechanical rectigradation (Osborn 1894.92,1934.922). 


Ascending mutations, species and genera are principally defined by continuously progressive changes in the 
aristogenes and proportions of the grinding teeth and in the upper or lower incisive tusks. Although the rate of 
proportional adaptation varies enormously, for the first time we know approximately how long a period of geo- 
logic time it takes to produce a full-fledged and highly efficient adaptation, as in the metamorphosis of the lower 
incisive tusks of Oligocene Phiomia into the dominant shovel-tusks of Pliocene Amebelodon (see Plate V, pp. 
236-237), or in the metamorphosis of the posterior grinders of T’rilophodon from Lower Miocene into Mio-Pliocene 
time (see Fig. 300, p. 341). 

o——ppecies of Llephas” 
known to Darwin in 1859 

Elephantoidea: Adaptive Radiation from Genus Elephas Linnaeus, 1735 8 generic phyla: Osborn, 1933 

— | 


Through the clear distinction between change of proportion (alloiometry) and the origin (aristogenesis) 
of new parts, also through the newly discovered multiple lines (forty-one or more) of ancestry and ascent technical- 
ly known as phyla, the Proboscidea afford a complete revolution in our biological philosophy and concepts of the 
nature and causes of evolution. The contrast between Darwin’s knowledge of the Proboscidea when he wrote 
the Origin of Species in the year 1859 and the knowledge contained in this monograph is graphically shown by 
comparing the two accompanying charts (1933) with Plate XI (1935). 

Never before has it been possible to follow many lines of phylogenetic ascent over extremely long periods of 
geologic time, noting the progressive adaptive changes in each organ in each phylum to gain the perfection of 
certain mechanisms at the expense of other mechanisms. In general, the specialization of certain organs becomes 
more intense, while clos¢ly contiguous organs remain absolutely stationary. For example, among the shovel- 
tuskers, Phiomia osborni of the Oligocene of Egypt gives rise to the incredibly specialized Amebelodon fricki of 
Nebraska, with a relatively similar and unchanging skeleton and limbs. Amebelodon is paralleled by the flat- 
tusker Platybeledon of the Gobi Desert and of Nebraska in which the whole jaw becomes an enormous shovel, 
the upper jaw and skull being sacrificed and thus greatly reduced in size. 

[This section of the Preface was not completed by the author.—Editor.| 


Coéperation in the preparation of this monograph has been practically worldwide, with the exception of the 
naturalists of Australia, the only continent not invaded by proboscideans. A full list of the names of codperating 
authors and investigators is given in Chapter I, pages 13-15, also in the Bibliography at the close of this volume, 
but I should like to notice here some of the outstanding collaborators and assistants. 

My lifelong and honored friend, Andrew F. West of Princeton University, has greatly added to the value of 
the work by his Latin rendering of the Dedication. 

In the United States I am especially indebted to my friend, Erwin H. Barbour of the University of Nebraska, 
who placed at my service all his materials, his writings and his illustrations. In recent years my research as- 
sociate, Childs Frick, has rendered most timely service in his explorations, researches and astounding discoveries. 
European collaboration that has been invaluable was that of Sir Henry Lyons in supporting the Faytm expedition 
of the year 1907 with all the resources of the Geological Survey of Egypt. In his assistance on the home stretch 
of research on this monograph Arthur T. Hopwood has been particularly helpful in the Calcutta and British 
museums. In both France and Germany I have had the aid of a host of friends. 

CRANIOMETRY. New methods of measuring the skulls of the proboscideans were initiated in the year 1903 
with the aid of William King Gregory in continuation of the new systems of measurement introduced in the 
researches on the craniology of the Titanotheres. In every subsequent year this most helpful and talented col- 
league has responded instantly to every call for codperation and counsel. 

OpontToGRAPHy. The grinding teeth are the remains most frequently found and constitute the chief bases 
of scientific description and classification. The preparation of measurements, calculations and difficult drawings 
of the evolution of enamel foldings in the stegodontoid and elephantoid branches of the Proboscidea has occupied 
two years’ time in the American Museum and the British Museum (Natural History); this work includes the 
excellent aid of Edwin H. Colbert of the American Museum staff in the calculation of indices, of William Dixon 
Lang and William E. Swinton of the British Museum in the new ganometric measurement of the superior and 
inferior grinding teeth of the proboscideans, of Miss Barbara Hopkins of the British Museum in her admirable 
drawings of molar tooth sections and of Sir Henry Lyons of the Science Museum in the preparation of diagrams. 

SysTEMATIC IDENTIFICATION. During the period from 1922 to 1930 Charles Craig Mook assisted in the 
arrangement of the rapidly incoming collections of fossi! mastodonts and elephants. In the year 1930 Doctor 
Mook was succeeded by Doctor Colbert, who has rendered substantial aid in the more recent identification and 
measurements of the material. The proboscidean collection now numbers 1235 specimens. 

InLusTRATION. Especially outstanding is the superb work of illustration rendered by Lindsey Morris 
Sterling from the year 1902 to the time of her death in 1931. Beginning with pen drawings of crania in the 
elephantoid branches, executed in imitation of wood engraving, shown especially in Chapters VI, VIII, X, XIV, 
XX, Mrs. Sterling with rare intelligence, accuracy and constant enthusiasm prepared the greater number of pen 
drawings and diagrams for this monograph and is thus entitled to an enduring rank among the leading scientific 
draughtsmen of the present century. To our deepest regret Mrs. Sterling did not live to see the publication of 
these volumes in which her work will be preserved for all time. 

Reminiscent of the splendid series of illustrations of the extinct reptiles and mammals of North America 
are the frontispiece and five plates reproduced from the masterly oil, water color and pencil sketches of Charles 
R. Knight. All these sketches were executed before the habits and characteristics of these proboscideans were so 
precisely known as at present. In design and artistic conception the Knight restorations are unsurpassed. In 


the year 1928 Margret Flinsch, guided by the most recent studies of the author, entered upon the work of restor- 
ation of the fossil elephants and mastodonts, after prolonged graphic study of the living elephants. Her work 
combines a fine sense of accuracy with rare artistic appreciation of the motions and adaptations of these remarkable 
animals. Beginning with the Moeritheres of the Fay(iim she has vigorously and freshly restored, under the author’s 
direction, all the chief types of proboscideans as members of the forty or more genera and twenty-odd subfamilies 
into which the Order is now divided. A prolonged and very difficult undertaking of the last three years has been 
the outline restorations of most of the known fossil and living types for phylogenetic charts illustrating the adaptive 
radiation of the mastodonts and elephants, also the modeling in plaster of the outstanding mastodonts and 
elephants, aided by the figures of Muybridge. 

The work of illustration also calls for grateful mention of the following artists: Alastair Brown and Roger 
Bullard for their pen and pencil sketches; D. F. Levett Bradley for her mapping and lettering; Abram E. 
Anderson for his fine photographs of skulls and skeletons in the American Museum collection and for his de- 
scription of the microscopic characters of the enamel of the proboscideans in Volume II; Elwin R. Sanborn of 
the New York Zoological Society for his photography. 

Epitine. Beginning in the autumn of the year 1920 and continuing up to the time of publication, Mabel 
Rice Percy has been the editor-in-chief of these volumes. Rarely fitted by intense appreciation of the possibilities 
and results of scientific research, as well as by unflagging energy and patience, exceptional memory and rare 
precision, Miss Percy has rendered imperishable service to the science of paleontology. The taste and beauty of 
the typographic arrangement, the adjustment of the illustrations, the great labor of examining and correcting 
succeeding galley and page proofs to make this monograph a consistent and harmonious statement of fact and 
measurement, are all due to Miss Perey. 

In the closing years Ruth Tyler has assisted in the final proofreading and editorial work of the text as a 
whole and, especially, in the Table of Contents, Bibliography and the Appendix. The introductory pages and 
Preface have been prepared by Florence Milligan. 

Editorial appreciation is also due to Francesca LaMonte and Marcelle Roigneau in the work of translating 
the citations from the French, Spanish, Italian and Russian languages; to the Staff of the American Museum 
Library and to Jannette May Lucas for library assistance; to Miss Percy for the extremely difficult assemblage 
of titles for the Bibliography and to Miss Lucas for assistance in this compilation; under Miss Percy’s direction 
fine work has been done by the American Museum Department of Printing, originally headed by Stephen Klassen 
and including Miss Gerold and Messrs. Warther, Caggana and Klassen, Jr., and on the home stretch headed by 
Edward A. Burns with Messrs. Caggana, Haddon and Lewis and Mrs. Price assisting. 

IT am indebted also to Professor Barbour, Harold J. Cook, Morris Skinner and C. Bertrand Schultz for as- 
sistance in field geology; to James L. Clark and Martin Johnson in comparative zoology; to Nels C. Nelson and 
J. Reid Moir in prehistoric archeology. 

bon Mvefiild Diba 



MSE ey NACH eatin acme eae caskee a “ainal Sad. 'a0 600% ee 0206 06 ap 4 Ad Huiehad wha lar, PIG) op ny 8 hn OEE ele Ouptlelins Coe aS cneetena ete UE eat nne a ie ce ore terete xi 

TLAVSVT (OL TLE GU fe 03 507 Bd 1) 1 Oi nei eee a APU tion i, eens ctr i ea Ia cir aInase Xxvii 



MASTODON; ‘AND PEIOMITA 6. oie cssee tip Scio dashes alate oniae. a1 p Op Pe oe eee ee eee eee 17 

Linnzan zoological classification and Osborn’s phylogenetic classification of the Proboscidea......................... 19 

Osborn’s theory (1900) as to the African origin or center of adaptive radiation of the Proboscidea.................... 34 

Phylogenetic relations and classification of Mwritherium, Paleomastodon, and Phiomia.......... 0.00.00 000 0000s ee 46 

Revision and synopsis of genera, species, and types of the Fayim proboscideans in the order of their original description . 51 

Palzomastodon beadnelle: Andrews; VOOM. 1. i.e ce bos note bone ose ete erase. < «19a eeeps Onin ae ee ees eae 54 

Mieritheriumlyonst ANGLE WA: VOOM soe. aces os tetera ararat 5 omy Seow ew ROUEN eee leg ee 54 

Phiomia'serridens Andrews and Beadnell: 1902 x... .re-sc.255 po Sete entrees ale eo lasgle oe aie ee kee 55 

Merithernim gracile Andrews; 190252 5 hs0. To. 0 oes Seat Spats eet ib res aegis ea tet aychete Meee tee rekere ee eee 56 

Merthertvum trigodon Andrews, 1904) «5s... '.50%-caars oS ise nies wi eve ci oltre vue shat aelece tle io up e ael aiei earn ee eae eee eee 57 

Phiomiaminor Andrews) 19045: cies So cess ss vide oe Stipe baratlens aikae:  yane Seseeesa rata leteiapal? aise te Ltt ieee aaa te een 58 

Palzomastodon partus Andrews, O05 he... 0 2.00.35 sec ote 5 ots noe seis opes aed eee a cae ree 59 

Phiomia wintont Andrews; L905. occ os. ayoc aes 3's 0 20h 0 bbb bin alae toro aap Reade ee el eS een ee 60 

Phiomia barroist Pontier, [90 Mics. cet «a0. sa Ph we a erase ® Sayan tae oak eee ee nee 61 

Merithervum andrews: Schlossersloly © os ws ee oOo nae ed eenais seyereiesa oleae) secgeeks seit eee 61 

Palzomastodon intermedius Matsumoto; 1922: «... <2. de8 as ae = a een eleces SAU tea oe eye eee ee 63 

Phiomia osbornt Maiisumoto, L922 se sacs gcc sacs {tec 5: 308 oho pugen vole) ayaa s res ONS Ne MEAS esate eee ne 64 

Merithervum ancestrale Petronievics, LOQB i. <2 v.. ss5:6 oye + eee ct es ors rove fe eats ele geal a eee eee ee 65 


Meeritherioides, Osborn,1921; superfamily definition): «<5: ¢ << 22-6 aae weeie ee oiercnete terete 69 

Meeritheriids Andrews, 1906, family definition. ..........%.2.-2...200.5 ee 1 fess eaters ietete ol eee ene cna ee 69 

Meeritheriini Winge, 1906; Mceritheriine Osborn, 1923, subfamily definition. ........................00022 000005. (fl 

Menthervum Andrews, 1901-1906; meneric definition... 2. . 05.05) sores oe = « ciel pretest te etait tere 71 

lyonst Andrews; LOO... 5.5.2s5.5 5 ve oe oo aren 2 iy 4d Srnei n @ Oe ale silane ey ns cl een en nee ee 72 

gractle:Andrews, 1902: «2 sss sc: se w)etete wis E55 20% = 2 ove ee 1 splnsclm = lgaee mea 8 ae 73 

andrewst Schlosser, 1911... 5.0 ein vieste cool sea 0 8 5 ee ton aiety ote ances eo Seton lg ate eee 74 

trigodon Andrews; 1904: a... «..55)s Moti dy «ak fm 2 Said « Slits aie eeeecch oe lee tale ls atiia Para ranean gt eat 74 

ancestrale Petronievies; 1923... 6.5.6 :..¢ <= ois oo a cle ware ara, o0a is oho eeieteeigleeueie enka ws ita eee eee 76 

Supposed. Meritheriwm of Baluchistan. . 2.0.0... «005 suis, bescniont sia enere siete eee ele 31s ea hee eee 78 


Deinotheres distinguished from other proboscideans.............:0-.-sceeeeccsv eter ecu deepeewes 5 SW hese 81 

Curtognathids Kaup, 1833, family definitionand synomymiy..3- nero ree te eee 82 

Deinotheriins Bonaparte, 1850; Winge, 19063" Osborn, TO10: 3. Seo eects cee ie er eat ee ee 83 

Deinotherioidea Osborn, 1921, superfamily definition) 02 25 323. ems mers eerste atstetet vel ate an es = eee 83 

Deimothervum Kaup, 1829, generic definition’ and synonymy. <6... 6- << peeteieaye es ates cients te ie steerer S4 

Twenty-six species in order of description, 1715-1085.0 27." 129 Se etre eens os ee er ae ee 84 

History of discovery, 1715-1822, summary by Weinshetmen: css petite ee ool wrens once sce ei ie ets lel ieee 86 

Historical summary, 1883-1934, by Osborn sc. <2. ho ee ree ec ie nie neta ote) nate ta ieee te 95 

Résumé of the. chief generic. and specific characters. <0. <6. s.crrcete be le ate ereratetnee) ore © Wieastonelerc ats ater seteretnatote iste 110 

Conclusions a8 to origin, phylogeny, and adaptive radiations... - eee sec Come ee ales a= oe ee eee 112 

Geologie and geographic distribution of the types of the superfamily Deinotherioidea..........................-2.. 114 

Deratheritum NOWw00ds Sp. DOV... F545 je vine oe beacon wenn one eT re wie a oe co ae tne ce orn eee ey eee eee errs 117 



1h7 (0 ho Py fee en ee ein ry anh A ERC oC OC AAOO Coe cd ho bAaMOCNeEoes ANS So o008 005 119 

@unvierss five classic species” Of Mastodon me cpnciae tori pe teres acre hee eee ee ete ot a 120 

Cuvier’s original conception of his genus ‘Mastodonte,’ 1806: Mastodonte de Ohio, Mastodonte a dents étroites, Petit 
mastodonte de Montabusard, Petit mastodonte de Saxe, Mastodonte des Cordiliéres, Mastodonte humboldien....... .122 

The Zygolophodontine: Separation from the Longirostrine............--- 121156 sees eee eee ee ee eee eee 124 
The Brevirostrine: Separation from the Longirostrine and Zygolophodontine...................--- 22sec eee aeee 125 
Final separation of the Zygolophodontine from the Mastodontinge.............. 0.0... sees cece eee eee eee 126 
Separation of species of ‘Mastodon’ and ‘Elephas’ described in America (1792-1874)... 2.0.00... 00sec eee eee eee ee 127 
Miastodonpinse sresumac of st lo tam ya chiens CCS ete ier tye eee teteteteae ese eee 132 
@hronolosiellistromeeneray an GS pPecles ews sewer ers acer Neen a ee te ete eee eee eee ee el 136 
Progressive characters and generic distinctions of the Mastodontine..................-.. see eee eee eect eee eee 138 
IPaleomastodony ANGrew ss LOO lm eeneric Ge tmitl omar yts wer ter tee epeteet te eclectic oe era et 143 
[MIO Ne spk Wye c eer aE OED ARON OD OOO 5.5 G00 0dsceeuosonddeorco moons padsgbonodsadecagonuaennsao0 6 146 
HGR ESM EAS DUNO Oy! At Pea aIRISamefo UiniG. one d ocd Coad Sone a aERE ROMS Sg oS ARABS AR he cleon sa6 bin ool < 146 
peacdnelio Amare ws sl GO Mss isex tere marcescens Sacer eee teeters ear css cent fac so te ooo AE ce oder ee ae eta eo ee 147 
Maomastodon Osborn 922-5 renenicdetnition\ wera tersrtee clerics eke ene eter oye clea ener eh en eee 151 
GARI OR Wont IP oon amon 5 eae Ot As cane dome ome meee ee omocs cude vc oAaccmh bo oncuuESo ore t- 154 
HADIRAOLS CHGS soa esiayxers, IMPAIRS. 4s op ob osdaceo be PeRe coro lube bou co dcp nedoos cence onacesianans 156 
RiomastodoTA Osborn O26 yeenenie de mnitloniee rere es teres = ore esas taot eee aa oe ee 157 
at Lan IAO 1 oO a0) lke PALS LV eet nee wie ot an dis 6.0 G bo Oo See e eee AMES MED OMe D oO touocosecser arabada. 5 157 
MMEnCANUS PIaeLypicaochlesin yen lO O22 earns meter ere ieysteiste = cis) aiciere © ie lekeree eeictelel keke elect Vet rea Rete ace 159 
suillenesy Simm ICES wom connvsepasnose cab savoa conse SOROS UO SoU OO Uo Coo mOSCOOo oS oumOnOeOS oz: 160 
Arama NERA Oil OBI oaaaie SeAtoene Ge rleclacnn of oc no OOO ROME Cnae amos obo aubuneassooaso obese o occ 161 
Mastodov@uvier 1 S06—1 Sle enerie!Geiinitlonia. «ae cee sisie isla vais 62) c)e. 2 < sichspste faiete alates euotloyeneiai olla vatateleie iste liars teenies 170 
HPT UAMOVIS aR RYAS Shasta d tcaocoe.6 Guat caddce ¢ ae Ree ae OEE MEAD TN PHN Rab oncoaUodcooDOoo AMC SS - 170 
COC DUS FUROR G OU hig INSBIDG Go uoooudd audnodaodh HABE EaRBRRME Se Osa AID oon oohogaedoobonpaocaucucs: 171 
DICE BE Oe eo co. aie hoc o do Sao he Opca cen ane SAS ae ARE AR Opes sods rob dh oon coostodes tabtcco > : 172 
CRT OM GSES BEN) IPA Oakes od oearao sed aa ees Daa ss Oe CO DIEM MOO OLD OCC One ogc coe; onseM ar GoSe oes sec 173 
AUC CHV Ol ooinin I PLN og. ao cade oo He oOo 00 oe oe COMM EBROMMOSs To ppe Ee OvoanO noc ussrancqamodgs sa = 173 
aaa eid oO ERF IBGE Ge say da Se oes o oN Undo 1 CC deb a Lee ene DEAE SoS OR OnonmogeNG oaeatomdaos O42 174 
MOE en Oo OBEN Ss do goas condo de youlso aut 65 lon Open nS on OO morass otadc Coes abamng cous 175 

paisa lms CRBS. oa cone oc a Gee bod Side ao Sec ons nas Sn San eC Bicc ot coh onsonact Ocduobapra cao oc 175 
GRATIS Clim Mipsis fintdlic, WCBRY, 45465505 on daoc st bo HAe A GO GROeRE ODE Coc mae coe soagtonoedsdEsasescoss 176 
Wimmera ginal WWylanir sol brsyerl nse as aces ed dos 00 one atoo 6 hd See eR EEme ar me st dos co noom raonemeodoad os 179 
(Claoravallapni@ Instn Gir eran averea have Sachs oho Sola o oo. don oda a rene eer RRS MnORO ot cts amon cibinibiwd oidicao 0 010% o's 192 
Collateralimelationship tobe sublamilysterodontinet ss. 2- 22.2... ee oe lees or oe wee scree cle le ae aetna ieee 195 
SIUTENOI On HERRON YAN ONO i ionacesin ho op suoo0 4c OOOO RERNRD OSCE oo oMnm aan oiccecatenuoe ooo pe on aa c 198 
ReEsMmMmeo nthe eenenic ang speelic Gist ChLONMS ae qmemees vase eee eles. 2 ec eeble = mists see cta/ seis ence eng oleNeye to teioee ol eletens 198 
Byyaaliay sf ook MAGA MEV Hii POO MOO TOTO 4 os 2 CE RS SOR ES TERED Mn Op OMAR TH oo0ts cor sean aoe dousua2 ace: 203 
DUTENOLCHS MArbe rN ios) (IH al COMET WLS 57) imate uepl ss) eyes aye a1 2). eat ancl ous ganenel eyelet se tele ieloyele etek sce eee 206 

[BOIW RAINES CIA ON TA ESTER OF oon, IO PLO. Soe a pa R CAA PME SORE ERM Re LO Goes DOOtnGoor J ccdsacegssepentons: 207 
Nap B ERTS TCR Waele aes Bia ama Ah 6 Gold ao caio Me enna. AG NCL | 6 Aon AEE Ire arenas at nea rye c 207 
Mastodon vellavus Aymard, 1847 [= Zygolophodon borsoni vellavus]...... 2.0... 0 eee ee tees 209 
Mastodon wialetii Aymard, 1847 [= Zygolophodon borsoni vialetiz].... 22... ee ee eee 209 
Mastodon Buffonis Pomel, 1848 [= Zygolophodon borsoni buffonis]........ 2.62. e cee eee eee 209 
Mastodon Zaddachi Jentzsch, 1883 [= Zygolophodon borsoni zaddachi]... 1... 0.6 eee eee 209 
Rumenis Osborn lo26.eeneric: definition... se notes err eo eeeketene = oe ierenc, sce chicken ub tekeretews ais cusekodete tek Stone ee +.) yates eters 212 
toqmrovdes| Cuvier, 1806518211824" Desmarest, (S20 —US 22 oc ve eta nie iets mo oleiniled ave Qe foe eteeeyelcl = =) adele eee 217 
ioe EE Sse ib iV Aral tse Lae ORR E catty Soa antle Poe ae aon enema OAR AT OA OD Oe ep bods dure fan 219 
UUTECENSTS SUIMOTNETISTS OSWOLI UGA zc. crac peers te ester aeeeetee eeaeree isle ere ene CnC CR rol RAN Act ead ee eee te ee eee 219 
TACT WEY aac) del tol) a a a UE A Sart sien cc o clomid are me ent Aime Scot AO Haka 4 220 

virgatidens von Meyer, 1867 
wahihevmensisil alan 1922 | ieee cks che sence pits share ote wood Pole eases eae eae sete ete 2) al Sa TAT ey = ks she eee eee 223 


Bunomaspodontiaw, family charaeters.:.:..s te-cs cs cee stuma © «= cine. « lly ovene ara Cae oa nee meme ails teaial ica a te 226 
Longirostrins and Petralophodontins, subfamily characters... 272 /).acds «tae eetetereto a= alate tee tha aia es e 231 
Phooma Andrews and Beadnell, 1902, ceneric'definition:... 022... «ar some te nate tel siete sete ans Sent ane ore 236 
MUUTOTANIGTO WE, LOOE: <5 2 icte-aesvaseid «uve pomse e eBouesecky Lieve cw neler tere etd ale la ate ONE eee ee eee nna Tee erent tea 239 
serridens Andrews.and Beadnell, 1902% #5200...) SSOP ed Pe ee cle cle ere e aan eee 239 
ADUTEOT (CE, SETTUMENS) os jo) oes wo. «ns see soareve ooh Fea ote eo atl bik Dalal Seee PD Otero ys ote eee 240 
WWUNLONY ATIOTCWB:, [OOD o.oiscocaacoistslc edwlecc scave sve oe Sup SNS Pinto OREd seen One eee eT et ea a a Ae te ONT vet re tee tote 241 
O8UOrNe MIGLSUMOLO, 1922.5 asec aseindrccasdres alern.d’erele aig 'atave vale ce 'ale pi Poet Oe ee ate SNe eae tate hac 244 
DYGNEUS DICDELEG, USOT <5. oo cn%s ne casd ave rays. o wich oveveo.a gy acest ie-e oc8' alg ia wrote Ba) P w/ele aha Ges anes oa ee es ete 246 
Trilophkodon WV alconer, 1846, 1857, generic definition... <5 f= cc 5:¢ 2 on.s ore ot noche rere Rea ete ee eee oe nT ee ete 249 
Chronologic list of genera and species of the Longirostrinez and Amebelodontine................0...0.. 002.0022 eee 250 
Nyetemanic description Of genera ANG SPECIES... « sicun. ciate + + ouslors ove wins ac apes (a igle ucla settee tet ae me cen 252 
Trilophodon (continued) 
angustidens Cuvier, V8OG: 1SET 5 vc. gosird else ove o ejatarers. ns araceserevexe ee Tea te ohare tec a Me TRU a ee een et Reno 252 
angustidens:gatllard: Osborn, 1929) <16< < sie sear a 5 a: o exes weep vsseoge syste iate's 0. ar oturel sheet at Reset ree Es a oes ee 259 
QNGUSILAENS VAL, QUSITO-GETINONICUS W CLDEL, LO 0S, LOUD es ctor co etate ae 2 fel aie ois tar a dip pare ores tote Rae 259 
angustidens var. ibycus POUTtAU, LOLS... 25 sc dicge s nog + «oes eb elolotena ale poe ears ae ere ie ac oa ae 260 
Choerolophodon Schlesinger; 191'7, subgeneric characters. .....<...-- 2. 0-00. 0- 22s cresle = «01+» wisps vse > aim si aeele a aan 261 
(Choerolophodon) pentelicus Gaudry and Lartet, 1856, 1859, 1862............ 2... cee cee ee ee ee teen ees 263 
DONALONE: HalcONeTs USO E ic. bs: s ctek bp aysie’s «dia ig cue 018.4) 04-b owl ay SI 1a, 5: wields leg tp aoe abate et aioe tee ll eee ete 267 
PUB UNdCUs Livdekker, VEBE. ccc fac ns aie nb ove. wra-tis oo nieve svepin 5) Siege ote) shale lela eie ge ac tole Sts Relat pons pete teeta eee 268 
Tetrabelodon (Hemimastodon) crepusculi Pilgrim, 1908, 1912 [=Suina].....................2 22.0 e eee ee eee 201 
chingrensis Pilerim, 1913: ‘Osborn, L932... «zie. is sis cscs or Foxe he ee = ee ee ee aaa ee 272 
MRALTOONOLNUS EUS § VOUS er acco )a «ars 3 0 2.06 We oie of > @ vyncey WAP es alate ASO eee eee ee 274 
CoopeErd OBDOIN, LBD. 2 oes ccesccenes as cope ses shakes sacar sa near ay outa ofa oases Bs oO ee age ee SONA Sa og 275 
‘nopinalus Borissiak and Beligeva, U928 oo casera e eine oi cre sits nse npn une ne keg aa aig ate esa sa ge ee ee 278 
RASNOLENSTS BP. DOV: 3.0005 cue ss ietvs Sis se wih 0s as vic ese Gc m, aheyere. cashes) pst eae ase eto iat eae 279 
sendarcus Matsumoto, L924 2 og sacs esi e gic en cceis wo cree vit g mbueen oe wile oe eee eet ete 280 
engelsinresensis Wlihn 1922 os hie so ae nso wiv ip, & oie wlave, 3 ere enspehe he agabelie te ne ee eee tele tese eeiee eta tn 281 
esselbornens7s Wain, 1922... — Xe Ae. cos sc 5 ae sin ye oe 5 os he msn tos 9 01, Oe eat or ee 281 
steimhermensts Wlahn ; 1922).0 25. ales o!s gira 0 04.4. + oe aks Aeaeaaae one olin oy ieee eS nee 281 
Other mastodontoids of Klahn 
Puricius wahlheimensis Wiahn, 19223. i. s cies sae oe oo sles aes naa tao pees a eee epee Oe ie eee 282 
Tetralophodon gigantorostris Wiahn; 1922). o.,~ os. sxe poe ce'e © c= lode @ eeisl ele eieie lege te erate eee 282 
Anancus minutoarvernensis Kian, 1922 0. oo. voi ca aces mews sa ete ape ee ele sla mes pain aisle ee ea 283 
Amancus gigantarvernensts Wishn, 1922) 5 6.05 kcesscere wogiayets 3g =n) tay ches eer esheets 283 
Trilophodon (continued) 
pontileniensis Mayet-Fourta, 1918.2... fa. sw 5 acne wns eiacer he coue Spe meter eels sya) sled taleleeaie es iateiate ie oe en 283 
sumplacidens Osborn, 1923... oc. bs ca sone ae ere eo hae ee ace whens tea ney Sine ete ages ogee ets ote 285 
obscurius Leidy, L860... 2... les aus says + eee bin dye 6 eco » wa pe a oo glee nO MIR pti ena nooo: ea 285 
Genomastodon Barbour, 1917; subgeneric definition). <2. ox. wie sis esti euetetele eeteeeee ee e 290 
Megabelodon Barbour, 1914, 1917, subgenerie definition... <0 .)a. ach sere teenie sie ea ee vs eels es ele eae 290 
dinotherioides Andrews, 1909) . sij5 2 a.discacs toile pis elk Pee, octet tere ee Neotel esterase 291 
(Genomasiodon) willistont Barbour, 1904 . - . ..<....¢ «= 00s os) csere eee ies eer noma ae aaa ete Se telnet 293 
(Megabelodon) lulls Barbour, 1914, 1917 3..0).)..... - wis:wpinsesatn cin isso she /eie We iat Sit a es ee eee ee 294 
ligoniferus Cope and Matthew, 1915... «5 s:sacscessavesocle aie) grace ates gies bone alabaster: evahe ee rate ata ee ae 298 
(Genomastadon) osborni Barbour, 1916... 0c¢css xanite« tee swion se see nols stnobws te Os es rae eee eee 298 
giganteus Osborn, 1921 |. ek sie s ws bisseneoasnin eosin eH stasater Oe e0e) er ee) seen ME Ea a ae cafe eee a 304 
Amebelodon (Trilophodon) hickst Cook, 1922:..¢ 2.5 c... <. susiviae eld Ope Wnesioieingd Md <p 7t ac re i etal oie ae 307 
Amebelodon (Prilophodon) paladentatus Cook,.1922 5.))<..y, ©. . og taut Seater iil ieee ats © ae alee ee ene 309 
Trilophodon (continued) 
GbElt Barbour, 1925 « « .. s.2005.<.0. dca nes tuthon OF Cm pte nstenes. Soni AMRIT See RAMOREN Simin eee Ss Ne ik eRe eae ree RE 

Jnicky Peterson) 1928 oo .0. <<< 5.2 5 a's arg > wa nero a RES phat Aut PREM TE TG een sands ere se harass Rees oaks tee 312 
PRIDE: COOK, 19ZB,. .acaas aly Slaros acres 915i adn nae ASD atae rate rtas Ba aya Wife hie cis Ree ys Sets ee ae es at 9 ens > ee 315 


Trilophodon (continued) 
POJOAGUETISLS HITCH; L9QG 205s PS aectere ana Pate areca RNS EID Ree oe Peer cae on ea 320 
enugcensissB rick, L983 5.2 << hose Peo aes iptv ciel coprene ere ays 01 veils eae ees antral aL ev emE Rs ER ee dee te Pepe 323 
(Hatabelodon) meograndensis Hire ke 933 eee ee ere eevee Cee ea ae 324 
(Hatabelodon) gregora Erick, 1983s scien wee ake geet trie chain hae eee entnle eee ee ee eee 324 
aya a hu (olor ES Pee, eee eee tears COREL RA Octane Ain. Conan ree eNO: Galea cuanto ain.natid haloes nimeaeble elo. 326 
Summary of the origin and phylogenetic succession of the Trilophodonts................ 000. ce eee eee ee see e cease 329 
Amebelodontine Barbounw929icsubtannilydeimitlonee reer ene rice cic i ei eerste een en 333 
Descentmtnromyehzomzmon the Olizocenciot NorthrAtiica epee eee eran 334 
Amebelodon Barbour 192 7-¢eneric definition: 4.1... uae aad ee ce hicieron ene eee 335 
‘fide (ABE) 0,0) bop Leaner nen ort ae Se OOPS PRI Rn Sh cia Aiea Ais GAO EG AEs mocks cone coaunanbunbtodoeenc 335 
SUNCLALTUIBAT HOUT, LOGO 2 a Feet cyte dacs Suet co! one Seth aache ANG) PCR RROD RINT ccotetrae c-Met nee ee eee 337 
Torynobelodon Barbour, 1929, of the Platybelodontine, generic definition.................... 00.0 eee cece cee eee 338 
loomist: Barbours 1920) re crs cr cerapacsrs Sree ie cece eae nucle ele Cates aetna ae Meee eee 338 
Mastodon [= Trilophodon] angustidens, cast of Cuvier’s original type................ 00 eee cece eee eee eee eee ee 340 
@entraliconulesidistinetivelotiniloptcd one titre intra errr erie oe eee renee 341 
Distinctions between the Trilophodon phylum and the Tetralophodon phylum, including the typical genus Tetralophodon 
HA! WINS VEAP ORATION MSV naa scouns dodanuad oonU ae AO obos od onuny Ay oeb ag uasanoagooecns 343 
MeinalophodomcontrastedawithysyMCOnNOLa pits er Pee eee eine eer eer ys ee nin ieee 347 
Phylum Tetralophodon Falconer, including the doubtful Lydekkeria, the more progressive and typical Tetralophodon, 
anditheuhichl yaspecializedtMorsclltairn ashes sete io oot evnuss costs ole sang oe eee 348 
Chronologielistoteencrayandispecies encima ate eerste ares casa ooo © ck orcad ans aS aoe 349 
vdekkentoOsbomngO24 subrenenicicharacversmcseyseerei tierra qo cle os riek in ere eee nee ene ee 353 
einalophodom (hydenkerna) i alconenilby.dekicersils iijanvaiemecte: ial -11ce) etenet tenons treat enn meen 354 
Met alopnRodonl (oi CEK kere) ISinens7sNOKET SS Sonn ies seria es ci). oneness 355 
Metralophodontine ivanideniN\laarel: O32 yc cece ci.s,eclee cue lecsce is nc x ios vivetne cin ose ois ssc teel ey nn Ae 356 
MeiralophodonsWalconen, 1847. 8o7, fener charactelsam.o.ce o-. a+... 620 asseiocine adece wae ae dudes oe eee 356 
UDOT OLTOS SAUCY ES PAS ran aes SOE STIS DEAS ON Cat ac an Ce teen a me nee nS nwo as a0 357 
OTIC THUS SON CTT IITs Gao eto. bin bios Guinea aay UR Soe Renae cree res oy Am ae 360 
CHUTKTRORES LEN ma, EPs ats. cdleeaiee Om aol Gas cic blo cea Ach eae emma enon oe oe Ons, SEP ERR AG noid sc 362 
BUI CULE ST SME CHIC CLM SSO ape verse ee Emey WSN Cee ey St Nr sass ves sgovev Se) SHORE At EN OEE Ee as yee 362 
VOGUE Ga CEI EEA OR BI pence acct Cone SO oe Oe CTR ERC Te See oe ore en mindn e nec 365 
COUT DOTIAP Clove ey WOAH ove Scot aul ct eho a HR ceed cea eS iH Cl SCO een RT rT Ce i PO URINATE oe 369 
precampester Osborn, 1923 [= Tetralophodon (Morrillia) barbouri]. .. 0... 000 oc cc cc eee ee 372 
elegamsiklay OI sete cen tanien ele Seraia0 0.8.8 PU OOS Ae eRe aC RET ERCONG Bnd ohio te ha aan Pea a ticle ss 372 
Tmlopnodon wetralophodan) brazosvus Hay. WO23H0n 555-455. ....-. 2 nseee sees oe see eee ese aseeennee 374 
HTH SE SD WO 0.3 05 Sey Bera RE ahs Sl See AR ORCS 30 OO ARC eRe Fete Rie A IE Pi ann Sc 375 
Monmiicc@Ospomelo24. subzeneric detimitionirenim eeeiecee 26 6: fe. ace a + ueipeceenase se ae ciiess ay sen meee 377 
iemampiortin. Uilandina) inten Olson, GP. scoaouesdeaeeegone sbsanne > baceeosnseenseeenosoonncacce 377 
STMT Amey 20 Ke VONDLONTAN Gand pra GLOn ete an ue ne aes en eg eR ect Ly a, a eos 4 he il aa 379 


LNINTID): ISOM AEDT S EE UENIS FB CA ete eee fen hee SRO se IR PRC Ge ee ee Dae 381 
HUTOpeaMiOTeshoneiniand miaration tou ontih Aner Calvan vert se) jee sociale ens sant eee 381 
Serridentine grinders distinguished from T'uricius, Trilophodon............. 00 cece cece ccc cece eee eee ene eeuues 382 
Chronologie order of discovery and description of thirty-four species of Serridentine, four of Platybelodontine, and 

CW OLOMNG HOM As tOd ONIN Ee. .w) =. att.y rt aa P git PSE Pee Mase eat) facs, cia.  aicteah teehee ee eS, cece 384 
Silavo amare Oloxoiaal, IMPAL, hl senroalh Glen. 4.45 ons0oocdsonhonvemes eodonsonepoAsoueduanecsoocheccodaecce 389 
NERrAENicnsIO Sono 23 sp eneric Genmitionee eee eee Mee see en eee eee eee 393 

SEMROeniiiiusisuoLanioracus Schlesinger Ollie eer ane erietarie aici ar eee eee eee 394 

permdentinusmmongoltensis! Osborn 9240 eee eee {lee eae Pay eck ea ed 396 

OcahentimustSemiaentinus) wlonescens Os DORN O20 ete aerate ten aye 397 

Sennidentumisigoviensis| Osborn andi Granger) 1032 sng h ieee nite eae ieee Tae enter 398 


Semudenunus progressus Osborn, 1923)... sac5¢ salen os ss = chai asic ens ea ee eee: ee 401 
EMIUMENLINILS pronuus: COPE; 1813: as hs.da sce ieee 4s Weasels ne eae eRe EG SOR Ee Cee 403 
Enmcenunusiproductus Ope, LB(D. i wari esis sous catn ies < osuue tito ate ne Cet ete eee ee eae anes 404 
Ocalentinusl (Sermdentinus) republicanus Osborn, 1926: .)..........4. suis aciee elie eek eet ee ie nee ee 414 
Ocalientinus (Serridentinus) bifoliatus Osborn, 1929.............. 0c ce ccc cee eee fed ete er cae ee hee 415 
Ovaentimnics (Serridentnus) | ortdants WeiGy 7, 1S80i.~. .:-< 4.1 Abie eine alee ae eee ee 416 
Ocalrentimus\(Sermdentinus) floridanus erate viCk, 1926. > 1s vos siete ene eee 419 
Ocahentnus (Sermaentinus)) oblequidens OSbOrn: 1 O26. aeycs als aici nae ee 419 
Senmaentinus serrzdens Cope; 1884, 2. ccpnvsperm nate cues feats da BET ete ieee ea ee ee ee 423 
meriaentinusanguiriwals Osborn 1928 530. 24.64. 25.3 ue eyo gba ee eee ee 425 
Sepradentinis meorascensts Osborn; VOQ4. oc)... se wee siete peas cesta eee eee 429 
Sarnrdentinis serrzdens CYMUITONTS COPEs LOO... .. « ais <ss6 cds oe. wm asin nicest ee 429 
Senrimeniiviis brewsterensis Osborn, W926 4. os <<. = np <t ui > pig ere eine isthe ee eee 430 
Dibelodon |= Serbelodon(?)| precursor Cope; 1892; 1803). a. 2s «snc nas 2 oe eect ene etree tae ie eee 431 
Sennidentunus.guatemalensis: Osborn, L926. sco s- so ste als bolnass 9 oxo a/ nuetelonseys el nate eee eee 432 

Ocalventinus Hrick,.1933;ceneric. definition’. «ch. <i... caus os wo Ate eee a ie eee ee ee 433 

Ocalrentinusojocaliensis Erick, W938 ac. cc. ciltasrte Mayoe soe eee 's Seis ae ane ee 435 
Serbelodon Brick,,1933; generic definition’... c.:5 2. cterrcule . One Oe oe ere arate oe ee 443 

Serbelodoen barbourensis Mrick. W033): 55... ass ata -nacdia sive Sele oe ee oat eee eae ee ee eee 443 

pervclodon Durnhaims Osborn wl OBB cc ictete.a2e te <,cse/s role sole ote) otek thee le escent Ree eT 444 

Serbelodon:(?) precursor Cope; 18921 893%, 5.0.0 acc an scot pete see et ere ee ie eC on ana 445 
Mrsbelodon Briek,-1933, Zeneric Genmitlows cco ss crac acaeee, ooo ov ace ale ia cc's seco caste aterene See ee aie ee 446 

Trobelodontaoenszs Wrick; L933 st gieeitee setae fate Sette cd entree, ode Se en ee 446 
Serridentinus (continued) 

Serrdentinus barstonis ENICK 9B. coi sce dbs cies Glee & cre Nerve ame ree tele ren ait 447 
iProboseidean life zones, and geologic/borizons of. India: «22... 2 ses se ee nie a cee ee 448 
Serridentinus of India, Japan, and China 

hasnotensts Osborn, 1929s et iors feces ois seve aininse ie wre elv aield ieee he See iee Peeee nae ne ee ee ee 452 

brownt Osborn, 1926........... Pen ed CIN eran Sei ri DSi A SO WAN Conidecoun sake ygnen Ae < 452 

metachinjzenisys Osborn LO2D 5 oii ac iavcseaw arn etn state aoe oe) ate ts tittee ale on oe ener 456 

Chanpiensts| Osborn, LOZ. o.ssc\cascce ie iste svi vi ses e is 0: a/ei ees, 1 Byacd Oe Pare eel alata ate ee ee 456 

prochingrensis Osborn, LOZ scsi 25 aicie fo ahs vous s/ere cies lore ocncataiae ale Spat Sea oe eee rien cnet eee 457 

annectens Matsumoto, LODE E .:. irs tense iaieisi ze 0: 'eie sf +, 7 Eye ile Gi eh cede (alerts ie ee ee eee 457 

lydekkeri Schlosser, 1903) (1906))s.c i... sce. sie ete se aa § clase sleuepelclelsic ie ola aoa elel ee ete Ge Saeed eae 457 

AUMNANE FLOPWOOG, LOBBs.... ise icc sss eee ow olere 0.9/4. n:0 14 win lls ins lss eel ays ONPG ee nee ne 458 
Platybelodontin= Borissiak, 1928, subfamily definition... 5). - w. <1. - )sts eee ee 459 
History of discovery, separation from the Amebelodontineg.................02: 05s eesseseteeet esse eee ep see sae 459 
Platybelodon Borissiak, 1928, 1929) eenerie definition. .... ..-.. <c.s <cie + cies oases eal cheers eee nen 459 

danovi Borissiak, 1928.23.00. 2..008 8 a ee ne ecg ee 461 

gramgert. OSHOrN, 1929 ooo noe eee eee Fatale ls cio ao niet ale Sos Sane Sheng aFetapeaes ote ans Reet een 463 
Porynobelodon barnumbrownt Barbour, LOBY 2.16.5)... 5 an v% «nis Soe eaten e Set alee eee ee 470 
Senndentinus filhols Frick, 1926, V9B8. ..<..... scale... cas vies cle cd ie ie emma Pete ee Teena ate nae ae 473 
Senridentinus (Ocalrentinus?)nebrascensys Osborn, 1924. 5)... 2 11. ieee eet eee eee 473 


GRAPHIC DISTRIBUTION OF THE BEAK-JAWED MASTODONTS.........................00000--- 475 
Rhynchotheriwm distinguished from other bunomastodonts.............0... esse sees eee e essere sues eve eee ees 475 
Rhynchorostrinse Osborn, 1918, 1921, subfamily definition’... ..1. +1 csrne rene crsteeie eters cite inser 477 
Chronologic order of discovery and description of the Rhynchorostrine. ... 2... 00. cece ee eee 479 
Rhynchotherium Falconer, 1856 (MS.), 1863, 1868, generic definition. ......... 0.000000 cece cee cee e veces cess. 480 

SPENCE HOUTCAU, 1918 oe 6: sae: ersicieve agro ates inne soreisge toreieie say tani Faken caye yen ner eRe Le «Te cee Te oe eee gee 485 

DrepidensiGope, USS9 «sa: «95 wis slates sieigTaso0eg SIS) s aie eee bets Micedec te tela eeehee nee Meee ae Temes cont ranean 485 

SHEMET EIA: LBZ 5. 5. ace aby ches cizin ee Sasi eee ree Tere Po eye em ee nr eee to 487 

MECHTONS OSHOID, LODZS.« ..<s.o.cis cs ie 7siels caw Ge IRE Caan aM TST. En rcrureee ce eRe yk IC CEE Sern ee 488 

[=]Skckothervim|)euhypodon Cope, LSSai50. fi iis pene ae es Meee Cette ce. Lai concer Pac eRe es reece 489 

ampacrinale Osborn, LOQG oc... v. 5, secure alte elem herent nent meant eee crc temseene orStsTe ec oe he eR RE Ren Se Oe 491 

Hascae Osborn 19181921 8) iv.cmirs wis heae hie Ch Ee eR IRS Steen Chr td Ree ER ce pene 493 


Rhynchotherium (continued) 
EERE TLL S| O51 00)! eR ee Ee OAS ocd aoe OL coon OU CO Uae oda ann hoe 494 
falconent. OSbOrn, 1923 \a. sais, .ziscces sc ties olka erie hel Rakers) So), Sehona ts yee eee eee ee 494 
Shepardvedense Prick, 192M ic1.65. afeet mcrae aerate ores eer G eae ehagel ee ee) cate eee eee et eae 496 
panedensis Osborn, 1929)\(—Rhynchothemum shepards €denselia. ars) e i/o eye ae sete rela ae 499 
Renynchothervumn (2) fraricisiel ay, VO2G sc yaw cron me Pome terest ourict anal essere eee aR oe tek eee eee 501 
CU Hes oO 3) 0102811 hg A! 2) ets ARE Serr caci ele. Osteo cr i nh cin tn ba tomas cinibtarre oe Rcd wn SoSbid woe ansat 502 
New: Hondurasiphylumi:'Blickothenmumiand Ar belodors nmin metal treats ata eeeist eel eee ee 505 
TUS ROA AROOIT ORO LTG -aLep uo) oul hess ees AR eran cic oe eaecea test aatci cman aor cmn Goyer EA Sem Ogu mots Godab mabe docs: 508 
Aybelodonnondurensis Brick; 1933% ic o.3.2tm = S00 cae os aces es Seer eos Me eee eee eee ee eee 509 
SummanyOniheevolutiomob they Eukiya1CHOrOs press terre eet alent seetetet= eMart et tet eke tener eet ere eee a 511 
IN OPPTOMUANS RO IDIOIN I re eeeees tes iiosecm a: lele kre easier ca cia, Sie Pe ese care ths Oe cats ce Re As Cha ge 515 
RLISCOPICALMM trod WE EL OMS 3c 5c hs: «e's Fi anche, Sores mengs, ase ey Stevan he ted Ses MENtae eeen ee oc aes ach de 515 
Shronoloe1c ord Scion GIS Gove rays 00 KC CS CTL O lore ete eee ete terete eee een 537 
Separaionmombne men ena o7d1ilen.o7y aa | Cccyien OTIC USt pega tet tettet t-te ete tee 540 
INotorosirines OsbormyL92i subtanaily dein tlonieesreererraresci te as cns reels rier ener rene ee en 542 
Gordillerzon Osborn alo 26 sgenericidefinitionmy: setae cies yess ite ere een earca elena eae eee ee 543 
ONLI CUNIETy, VSOG FUSL4 re aealescte a erases ecstatic eke reer hep satan a av cacy « SVUS ea ay ete eg SS ene ome nS 549 
Mastodon) |= Niotomastod on \angencun tsp Arnie SINT Oa Scheie releme re teletets aellelelciele ciaelere eicieiaveeieieiie eee eee ree 550 
Cordillerion (continued) 
tarijenisispAmmec him 190 25 uct tet sycho.0) tenner avo eaten ut eielollo/ Scot6: Sil s/i.a7> A @ ia lersbetet ela Kopee ke ee RRR Pe eter eaten 550 
YALE aT LORRI 2 atti} 0} Oy hole heer ea roma AG Micro eces Ta. cei e e E PRT Reo Gee ae Gh Cocca hea bee: 551 
Teleobunomastodon(?) Revilliod, 1931, bolivianus =Cordillerion (?) bolivianus ref... 2.2.00. eee ee 552 
AG} ACIS Clo) OXSyalkctcy Senet er eereet eee MALMO Rat Lier Up oO eC e ee eet oe Ser ruiesee 6 c.Oon aoc ho ac docket’ « 553 
OLEGOLUTINCOPEAU SOB wis cy aucko Sou eae eee ea ere eae Isao yo bay oySsloley ROIS eave RSNA cae ec oe 554 
GG ODUNISOMIQUISSEN Us BrEUGeN bere OZ 20. e we aera at a.d i - «case, SS CREA RR eee eee 555 
Gordillertons(?olugobunisifelacis Hreudenbergs 19220 rime). sor cj< sc. = - 5c 1 eee nee ieee ere 556 
Gordilleron (“\rolsqobuniswntermedius Breudenberg, 19220... : ..,.....t ss sees ene oo eee re oeeieeee 557 
OG AOTOOS FAO TRASUIS 1 CSO Molesey PPE ino 6 6 ono 6.68 DO OU Rann Meeeraoae 5 amin ohooh oon oUcooeauUnb save cs 558 
ORO ERE NNT 5 3 oe Boon COO oe SOMO Op OO c HOLT ot eens Aas Rete meters snares saree cise eo aoc 559 
GHMSis Os owinin, TOA 50-68 coat ES BOR RO SAC OC eo ERS Can Detector tic ee ees A eRitG SEE es 560 
HRETINS Lele, OAD. clog tio.oce oe Raa Oa ATO S.O Nd Oe eee eee IS OM Te erm Ge es Aine Aa neo BA eed 562 
PLEHOBCOU Gy EL euyeml ODO ncn uae sre RAN cect eRe ert eRS eras) fells v vi gis' shale sansa yoke ee ieee ce cee Oe eae 564 
DETUSOPCETISTS) GG LG yMILO 2G spotter] once eee aapel eeed emp Peg roy Sasi, sic» site ee Seog cee we ge Siero area eee acta ae ae 565 
New knowledge of Cuvieronius afforded by recent discoveries in Bolivia (Tarija) and in Ecuador (near Punin and 
JNTENECCENEID) oe eh ates CaS Geer ART Hea AOL OLE)O octin.o Saxe el nO oiaed eT I Cree enone ge eee hr, oe nett 567 
Type cranium and skeleton of ‘Bunolophodon’ [= Cuvieronius] postremus of Alangasi, Ecuador, and type cranium of ‘B.’ 
lE2@zakeronsusllavorcombunin Biciad Ofer tte ei ry. ha ss © oon eee ee Oe e nee oeeae 571 
lHtnisal s@olinckoriainag WON es atoinde-sonteneton aan acia aotnicle o:o ts aie eee cen een Reena Peete coy sree Mercier TE Ohne ana 575 
EumbaldtinseiOspomeOs4 .culbfamllyad chili Omerryciscre sic. «cr sic oie oey-/e ase raps ieters eexyayn css ces yr ee eee 575 
GrMiexOnis OSHOMMe O23 + FCHETICIGEMMIGLON seycre eye reine sisenesésel oy =y0y9) oi s-auayrs Messed aay spare enttychat nk Sead ecaaeanytetiere ed Mees 575 
Runooldm Guivier pl sO Gi any Desmanest sUSUS) erm my mitsrrecbenc, | sc sr0) o:<0, 0.0.5, 5. c8 ols role ode neon sete cvi cece aude bee caere eae 576 
RGR IUTEN SESSA TIEN Clee Oat (Guat eSSOT 4.2) eee mene Petter 6 ite) Se Sea's soon ahs cane eee Thinacat aes Sn Ree 578 
DOTHLCROTSTE WN LOLEMO LSS Sree tcc ne 1 eee eee TM RCN Sh soo cd  Gcecee sated oe SRP a) de eee 579 
DIGLENSTSPSTOCE MON USS Bic cries ce wom eslerae oe em MNO EE SIO SNS eos. iS Geshe Sy en nes eet LA Oe, Se ci SEE 579 
SUDECROUSCNMEEINITIO LOSS ers osc le rchot in hea NER MME CRS ec RO che in, (c ave ihe a vce SVSPRN ATS eM TH wie meee ly ae 580 
ECULSPANINE CIN O WSS Oi rae cee Scie a's eae RRM OREN POY aed ER dc he Sock Ns BRNO, Late te SR awe ce 580 
ANON EPTONUS NC OP NO USO, ar.. atey ease OPN eR aE ne tsae TPES ETS suet te acche Res Me re AIST oe Pe ere 581 
CUALENSUSADA LO DIsel oO Siete Ae ees css eee ee oem REG = oe le keen A LOA cea cee 581 
(SRO ETDS EMG G7 TONGS, A Ae Ee hol sin aeRO RO IG on Sicha on Clute aes Cee EERE karate nord Ts et Pet BEL ty 07 582 
eile dreyaytlben nol ee /t2 rl to 3) Uae ae eee Ricinorte Cision CpProns.a tre tan tau DCO aR GCG taco aoe MEAL tonic ace o OR OaN Soc c 583 
POStrEMUSSpillmIanM a UOSM eo. 1. 5. « sakes Pies ae ete eho peeres navel pea tease yey ete anos See nee Pa ae ae a a coe Is Se 585 
Synopsis and critique of Dr. Angel Cabrera’s revision of the Argentine Mastodonts, including the new genus and species 
Notiomastodon Orns 2 oir. aon tare ene ek 8 eT EIS es TE Oe a a eo ee 587 
Notiomastodon Cabrera, 1929\ceneric definition 424. .-ar eeer an Cae ee eee 590 

ornatus Cabrera, 1929). os.5.2 bse iiss. 5 ace ao teig cheno te cate oe LOO oee na eT Tin cg oa 590 


Notliomastodon (continued) 

argentinus, Ameghino, 1888... 3.561 3 EAs 9 o Cees acd wine os MO ee es eee ie re eae eae 

Companisonof Cuyeroniue platensis and Cumeronius superba. .... ... se n> veaitente lie siete cieletels oleie st ee 593 

Buvelodon. Barbour; 1914; weneric definition; .c.141cmcs: <1 <.s os oleiea sa 24 eas eee Le ee ie eee ee ee 601 

MROTTIUA BOT POUT: LOLA: 50. oo.s..0: 3lds dictates sph yaperatet tenets «CLIO y, Ses vata distan ah resis a ere re On to 602 


AIMEE GAS, 6 iy afer « ah ave trae wvaiig dears avara, duarre 3% % e699, rei e etace/o1 26 ate\ a) Vente ope age RE eee ee ee 617 
uistonicaliintroductionis:5..045 4G .G eas vec tsi 4 wbaiecie mgs Fls seid ele fale At Oe eee et Se een pene eee ee 617 
Synopsis of the nineteen species of Brevirostrine 2nd eight of the Humboldtine.................. 0.002000. 000-00-- 625 
IBUMOMASLOMOME GR s..5 «5.5: vis eseqess Biafe ess: pneleie Xe valiese Subse le cituslalari Jo,0 pohoZogds cuose couse SOE eae ete ene ere 627 
Brevirostrinss Osborn, 1918; subfamily: definition. 2.7... aw 6 oc. a1. cium <n ale Riera eee een Rete eee 627 
ANONCUS AVINATG, 185), 1859} Ceneric SYNONYIDY ts 08. csc > 2 Go Stas Ys 01n O-atala oe nee ee 630 

MUACTOPUULS BONOCY POs oss. iisieree sale 2 leo a Gia di shwifel oc acpup aaa le eng he gl MPO ty aTR Tae ste a a 631 

arvernensis Croizetiand Jobert, 8285 2. 2. tek cocie aye sie wes Sle 2. pi ee RT RS er ee 632 

arvennensis or evvrosinis Gervais and Geisernes, L84G7. oo 22 sre. c= a ace. cie ee, a nition, oe tote cee teh ela oar nea a ee 634 

falconervOsborns O26 is, cic cresre a sioeere itcia tease, ayctatalie aletate lc Guteabe ekGte bic ETAT A ec OGRE AIO ORS ESP een 636 

WVVTOULLOOTETNENSTS ACs GTO MMLATUELILENSTS EN ALITY Le) acre coerce ates a eusie teres eaeerese pia aioe ene nies ere 637 

LTILEPMEAIUS PACHWAIGs LBSL es cayae ioe aie ccesid Bye berg ciwload iced wat Sot 0a eens Cte cue IE ore 639 

aruermensis progresson Khomenko LOU hos 2 2 iso's oe nin vieretetea ci ck etc pT D HVE A) Ie Er cotati ene eee 639 

SINCNSTS OP WOO, LOSS eb seat Ce Ateiaiaie cocky. eto oe ein cumSta) Geils ALERT ois, xSP aT RIAL eee eee ee 640 

mermmensis Valeonerand Cantley 184722, 2 3.nU jock Tas epee ines om Ae ole eras ve eRe ee eee ee 643 

PUOPETUMENSIS SP. OV ds 2 srerctele Hey siaiw loses Wiviniisr aja \« sasiwiel's olntole Glare, m 9 conte wid hcgutey sy @euete she en Ace eee 647 
Péntalophodon Falconer, 1857, 1865; ceneric:definition™ ... .. g...= 2-0. tas ss amas a -teleere| ate iene apes ene 647 

swalensis Cautley, LSB Gis coe ested see Scesos oes dyad oleae arg cele ehonete Fu 8 aria Pp RS Pe es er 650 

POLCONENT SP sDOW > 5 5.5.<. staysoRers ae reie (eles Bene fa ois Mean asians Sveente OCS Nag. Sie aCe te ap elec 653 
Synconolophus Osborn; 1929 reneric definition: 5 ..i-:<js) susy-1s ic laisls oe = atov ake = els aeeeetetene epee nee eee ne 654 

piychodus: Osborn, L929: ic ote tales teatens och oayecalys are a ah et dh nee Dei See Ee 657 

corrugatus Pilgrim, 1913.......... ble ein- bayer’ 's-315 4) 21a. Siilabin 0h 8 wabbady &. Senate eter oe 658 

fasnott Pilgrim, VOUS sics.0s aie aa e cth Se. tre. ood ele olen ore deeseer trator 4) Sit ere PRESS OLE OE eae eee 659 

dhokpathanenszs: Osborn, 1929)s) os 25 ict cord seo csw we dine ota a ae ee ee 661 

propathanensis Osborn, L929)... fe spe) eve.esc aie, ccs seco toll S wie cpt eye, aye eaten a ENEI e olaaeeeO 665 
Flunboldtidee fam. nOvi. .....s%e aeteis, wi sie dda adja aie bw isp ts weadgle alba Stayeee AURIS SIAL PRUE eRe ne 667 
Humboldtinse Osborn, LOB4: 2% byecis)e/s Wa, c ase. Sia a bis. Te icck aah hanes oe eae eR 667 
istegomastodon Pohlig, 1912) eeneric definition and synonymy .-1. seis: ccleaner eens erent 667 

chapmanz Hays; 1834; 1843). seo. sccsen clear a citys ran nse DONS eae Pel eal ne Tage aes een ee 669 

mirgficus Leidy, 1SB8: «2. Se a Se Nees ein ava oa sein ew ees wate ayes nee reece eee ee 669 

successor: Cope, L892 655 aca We kes cig cl ed o)c.0d sim to sin levee va. wha alle aha te ay teh eoteer ee OCRO a aie ee 671 

temas Osborn, LOZ... 5 isis firedatapele crsaeee i ow iah8) Sn. e a. es olS) oc oye us Beeae ae ean cea dean 673 

arizonz Gidley, 1924). 1926 > csi.c cece 4 eerwiles Pie tevole ww ob ne wre ge ese eee I a een eee a eee 678 

ajtonie: Osborn; 1924... cao 5 40 ie Mecsas ok societal via auelin, wi oth he Ue a ee 682 

priestley? Hay and. Cook; LOBO 5 sce. < dee Lie 8m Bh cs eensena Shee cts ae teeta oe ene pe ese ee 684 

DIANVLLVUS SP: DOV « 5. nieve see ha ale devia oie, vi wle v)ehs ivhalpiole 20g Aiahe calle CAE en er 684 

PAu NIT OV OU UME Deca: ais ine oywiey haere ini Ries «8h no Bowe oes meta at eR Inne Seaman a 685 
Adaptive radiation of the Mastodontoidea as discovered and analyzed up to and including the year 1935............. 685 
Paleomastocontines sublamy DOV)..:.i2'-sisise aie iso's Oe ee GEE ce oe ee eve eo foe 691 
Maomastodon depereti:sp sNOV 22. ais a.c <2crnapsipichs sae eqns 3 oe ese i eT eee as IE ee eminence ee 693 
EAS OR ON PAULO BP. DOV 50.050 <aycdee ie RED yas a CLP oboe ca 8 ot ee 694 
Hastororgrangers Barbour, LOS4ocis...:6 otyys oats na nee #0 Seiad eee eaten n ee ee 695 
VE CSOUOW ACUAMIONS SP. DOV s 2 oe ayu%eiccvegas SAAS apa aoe mk ice Oe Oe Ee ties re os ee Oe eee 696 
Turiaus turicensis from. Upper Bavaria.ciisd een oe Rei ae ee ie RE asic 2s eda eee 697 
UTI CHESS st2ODd CHING «. . 2 wcilstetn dyer eae ee ees ate ee, Oo cede een eee ne 699 
Zyqoiophogon vorsonerel, from Ching &. Sece akties lec ieee tet esas clad cite cies ca ena ce er 699 
pcerolopnodontinss subfam-nov’....kis5 Asha ste tenet e Ruwais outs. c causes OE VR ae Une ee eee 700 

Stegolophodon lydekkeri sp. nov 



Trilophodon connerus Hopwood, 1985... .... 2.6. . eee eee eee een en eee eee tees ened e ees 702 

Trilophodon spectabilis Hopwood, 1935... . 0... --- 0-0 e cece eee ne eet teen nee ett eee e eee nes 702 
Tetralophodon exoletus Hopwood, 1935........ 0... eee e eee etter ee en eee eee n eee eee 704 

Mhe “Spoonbill? Mastodon: «ste cr treyac cao arc ks reeled ge cent tesa ecte ato nent eee een et eae eee het eee 707 
Ynathabelodontine Barbour and Sternberg, 1935, subfamily definition.................... 2.5 eee eee eee eee eee 710 
Ynathabelodon Barbour and Sternberg, 1935, generic definition. .............. 0s eee ee eee cere e eee ee eee eee “lu 

Ynathabelodon thorpei Barbour and Sternberg, 1935. ........... 20:00 e eee eee ee eee eee eae 713 

FMA ond en nde JEM ey Wipandlellipn tn noppodnanocatcaonsanndcmopeumnoseoedoooppasonocGoucHeonfoaasecs 715 
AMANCUSISINENSTS ELOP WOOD AOSD cre crete sim che ee te ete lee cer sectst ene ce 9 ee ere hete le leh fotr allt cts sha eter dons kal =e nearest 721 
Humboldtide:. family definition neice sa rtectarctsis yeti ste ete ee lerene ole vielec eee far= eet tenet tet ayetrse ee toate Men acer 722 

Distinctions between Cuvieronius platensis and Cuvieronius superbus.. 2.0.66. eee eee ee eee 722 
Stegomastodon primitivus Sp. DOV... 20... eee e eee nee ee ee ee eee eee eee eee cee 726 
SErautslamoteoneneinaalaimitils s gg aigwodde eno booscerdoscoan so0sacunusm mn eonuneoTaondgogusocogodoccoupsboo6 729 

AN Waxnvavoaskey cove Cormunhavesicibll ov isn0saa0\o) Aaeeen me en Alor hen ane Goo OM EMA SnD Gro boUTeuongehducosade ammo aoa oM 730 
Notomastodononnatus Cabrera, O20 sr rcnce cise erelete cetera sis oer ecard even o ke cite ere eaeeate eet etete etree et foie =\-P- ott ie ea 731 

Serridentents Wrmand LAOpPwOOd il OB orc cricie eter i eevere iets ss = er cu-V-t ote¥e i cdeyene te et ele etal ete sie) =e) Rea eel T-Vecels 732 
Ocalientinusemmonstielay.. UG3O cer. cise estore ete eaeae ee kell etons ever e nlm svete ote eroded elena che TeN= Fave = IOs N fe kay Kexel cot eel 733 

Osborn’s final (1935) classification of the Mceritherioidea, Deinotherioidea, and Mastodontoidea.................... 735 

BIBLIOGRAPHY OF THE PROBOSCIDEHAS UGO2 NOS aie yea sees se cre acdc eles ope ele ca tatters eta) ett ee deter= =i tet) ties clones 

Frontispiece. Restoration of the Warren Mastodon (Mastodon americanus). After painting by Knight. 
ieee Vigstodonnnss: eiomastoaon, Miomastodon, and Mastodon......02.0. 12.2 tase te sn aeeeree ee tae tee ne 134-135 
meeZyzOlophodontinss: Lygolophodon.”.... ete eee eee See ees clans ns on eee tte ete eran 134-135 
Mitbee ye Clopnodontinws “LUsiCuus: 6. psd.) oo RI ke ee ee en tele seen ae eee ee eee 134-135 
Neen ten Ol ophodontines:. \Stegolophodon. .2.. <4 Fanta. cet e nee elec eat Godse eee eae eee ee een eae ee 134-135 
Vemrarnenclodontinw:. Phiomaand Amebelodon’..-...naate sc ls stat oe een a ee ee oe a net ee ee 236-237 
VI. Amebelodontine: Phiomia osborni and Amebelodon fricki, primitive Lower Oligocene shovel-tuskers. Restorations by 
IMS CHU Se scree cc haps seis nc: oe nate hoyle sere ates ether e naire Ores slate SEA re SR end AEWA CAN TR OO 236-237 
VII. Miocene-Pliocene life zones of the long-jawed, shovel-tusked, and Humboldtine mastodonts of Nebraska, Kansas, 
Colorado; and New Mexico.) . 3 onto Ben Pee re eke ee ee ee Se ete eee eee 328-329 
VIII. Pleistocene of lowa, Nebraska, and Europe. Correlation chart................ce cece eee eee teeter reece eee es 1848-849 
IX. Type jaw of Notiomastodon ornatus Cabrera, also restoration by Flinsch................000 02 eee cece cece cece ee eees 382-383 
X. Phylogeny of the Meritherioidea, Deinotherioidea, and Mastodontoidea, 1934. Folding chart....................... 684-685 
XI. Phylogeny of the Meeritherioidea, Deinotherioidea, Mastodontoidea, and Elephantoidea, 1935. Folding chart... .684-685 
XII. Type jaw and dentition of Notiomastodon ornatus Cabrera. By Bullard................0.000 cece cece cece cee eeens 730-731 
1. Heads of twenty-nine types of proboscideans. Restorations by Flinsch..... ..........02eccceeeecccsceercceceecscces 16 
2. Flood-plain scene of the ancient river Nile in Lower Oligocene time. Restoration by Flinsch.. sia (atacgha a avait Shei eiie at ee 
3. Diagram showing Osborn’s theory (June, 1921) as to the adaptive radiation of the Probosuidaa,, patos or ts nh et 
4. Fundamental arrangement of the cutting teeth, I’-I., in four of the suborders or superfamilies ar the Probesaiden! a Are nr ae 
5. Chief head and dental forms of four of the suborders (I-V) of the Proboscidea.................000cceccecsececeuceees 25 
6. Worldwide distribution of the Proboscidea in past and present time.. oy inti who.) 
7. Osborn’s diagram showing the known and unknown stages (April, 1925) i in ie piiee aiaeoe a Oe Probosaiien ees 32 
8. Osborn’s diagram showing the adaptive radiation of the 37-41 generic phyla of the Proboscidea, as discovered up to 1934 33 
9. Division of the world into three realms (Huxley) and eight main geographical regions. After Osborn.................... 35 
10. Orders of mammals as placed by Osborn in 1900 in their hypothetical chief centers of Tertiary adaptive radiation ......... 35 
11. Section showing lowering levels of the ancient Lake Meeris, of the brackish Birket-el-Qurun, and of the sea. After Osborn... 37 
12. Map of the Fayfim area, formerly covered by Lake Mceris, now reduced to the brackish Birket-el-Qurun ................ 37 
13. Meritheriwm, side view of head with eye and ear in position. Model by Christman...................00ceeeeeeceeees 38 
iemeenomawinion,, side-view or head. Model by Christman. 0.3... s0 02 sie nen aut crete einen iene caer eta ee 38 
15. (Left) Hosiren libyca, the Eocene sea cow; (right) Trichechus americanus, the manatee. Restorations by Knight......... 39 
16. (Left) Meritherium, Phiomia, and Parelephas, evolution of the head, proboscis, nostrils, and tusks. (Right) Meritherium 
andrewsi, Phiomia osborni, and Parelephas jeffersonii. Restorations by Knight................. 0.000 cece eee eee 40 
17. Phiomia osborni, primitive Lower Oligocene shovel-tusker of the Fayim, Egypt. Restoration by Flinsch................ 45 
18. Meritherium lyonsi, first reconstruction of skull and mandible by Charles W. Andrews.............-.00--00eceeceeeeee 46 
LoneiMcntneriwm, tront view of head.. Model’by Ghristman.. oc... issc+'cle tie cy ninale aieieicieie ciel iene aimee eta 48 
Bie Eeniomia wintont, front view of head. “Model by Christman. 7.:..2.-.<. sccm ee cele acters e sien ciereinie ie ieee oie ee 48 
Biomiveritnenvum, Faleomastodon, and Phromia, superior Molars. ..... se cecreieie ie sities sierra eatin et eaten een 50 
po toerthervum, Palzomastodon, and Phiomia, inferior MOlars...... «c+ cs seiee cies eieeie cere ee aieleniela aaieiei ain eee 50 
23. Section through Upper Eocene and Lower Oligocene formations north of Lake Qurun, F hres BgyDl. . 3:5. s0c. coe eee 
24. Map showing geographic distribution of species of the Fayim Proboscidea. . he aig ds Whee ogee WT cee 
25. Complete faunal list (1906) of Qasr-el-Sagha and Fluvio-marine Series. After Andros Sr URE LC Riss 53 
Bom Eateomastoaon oeadnellz. Andrews, type fIPUre:,. 2.0.2 sss sve +a acces in nle ume eke erie eter ateieie im iers eine Meiers Denna eee 54 
miemenioriinerviuniyons: Andrews, type LUNE! «becca = aun «>= na) <0) Ye cveleial eceiese ate attire aie chelates tren r iene char atte e)eh Cacia eet eee eee 55 
Baa ieRntiomiua serriaens Andrews and Beadnell, type figure,.....<.5.sce0 - 2 ve ohiem tems ieic ce caltie ice Me etic career 56 
Boa MURENtUIs gracile ANGrews, tyPe LGULGE, 5... °.-.« sis » vs vele a: since ic. cle wre ste ect alain uses rene enIeic in cee orci enters eae eee 56 
30. Meritherium trigodon Andrews, type figure....................055 i, 018 ats) pee ag clas ices eT uss cal one Lalo (Ara eT a lie Seca ete ee 57 
Bl. ePaleomastoaon (Phiomial minor Andrews, type firure-...<.<-.c< = 00s oe eee cee em a eo ee he ee 95 
De Eoeomasvodan parvus ANGrews; type figure. <ccic nc. cccrns o, wct.e ore Ure Heme tas olsen tae cte ciao otal oes ea 60 
poe (eaLeomastoaon Leniomia| wintont Andrews, type figure. .... .. + as. od2e.dmen aac ns oe ccs satel oo ee eee 60 
34. Palxomastodon beadnelli, Phiomia wintoni, Paleomastodon parvus, Phiomia minor, type second and third lower molars...... 60 
AMEE ieOMasLouon |e nionmialoarrois: Pontier, type tivure..canu cases: nen et ici osisa nots. sock cane aeee Seo eee 61 
Bo, Iverthenum. andrewsi Schlosser, type figure. After Andrews, .....2......2..:ccsces vss cn sees ccccccnweesbbee eas oe 61 
37. Meritherium andrewsi Schlosser, paratype and referred specimens. After Andrews..............-0000eeecceeeeceeuceeee. 62 





38. Palzomastodon intermedius Matsumoto, type and paratype figures.............. 0.6 ccc cece eee eee eee eee eens 
39. Palzomastodon intermedius Matsumoto, type jaw and paratype skull.... 0.0... 0... cee cee eee eee eee eee eee 
40. Phiomia osbornt Matsumoto, type figure........<0cc. ccc cet ee ce w o wl Melee aie taal ole) olelels « ebeie ie leleleleislsle ol einei lel ietea le 
41. Primitive Fayam proboscideans: Palzomastodon, Phiomia, and Meritherium. Restorations by Flinsch.......... 
41a. Palzomastodon, Phiomia, and Meritherium. Restorations by Flinsch.......-..-. 1.01. eee e ene e eee eee eee Bbyai tne 
42. Osborn’s reconstruction (1920) of cranium and jaws of Maritheriwm andrewsi-trigodon..... 00.0000 e eee eee eee eens 
42a. Ancient Mceritheres of northern Africa. Restorations by Flinsch............. 2... cece eee eee cece eect eee tneeee 
43. Meritherium lyonsi, axial and appendicular skeleton. After Andrews............-..-. 20sec eee eee ence e eee e eens 
44. Meritherium lyonsi Andrews, second type figure, also paratype figures.......... 06-6. e eee eee eee eee eee eee ee eee 
45. Meritherium gracile and M. lyonsi, referred mandibles. After Matsumoto...............0.. 020s eee e eee eee eee ee eee 
46. Meritherium trigodon and M. andrewsi, American Museum Collection of 1907.............06. eee eee tee eee eee eee eee 
ATE Mcerithenvum ancestrale Petronievacs; Wye LO ULC. eee cr «sete ale lols retel cle ercteye elope! © 2) =i) elete)ie/ ol a¥eNelal nade tono a\eTeve-is\i=/olep=\lelela Felelonei=heltetets els 
48. Meritherium, Palzomastodon, and Phiomia, comparison of superior and inferior molars. ........... +60. se cece eee eens 
49. Comparison of hexabunodont grinders of Palzomastodon with tetrabunodont grinders of Meeritherium................++- 
50. Trilophodon (Meritherium?) pandionis (?) from the Bugti Hills, incomplete fourth superior permanent premolar.......... 
RIMM entimandnewst. Restoratlonibye Huns clecesemace re aiet cette te eiereeters ct ieee ueketeichokaveledetsestets etal ieelefeleketetestelet Neier te tater ke 
52. Deinotherium giganteum type, as originally figured by Kaup (1829) with incisive tusks erroneously upturned.............. 
53. Deinotherium giganteum type, as refigured by Kaup (1835?) with incisive tusks downturned..............-.00+e0eee eee 
54. Deinotheriwm giganteum, referred superior dentition. After Gaudry.................. cece ee eee eee e eee eens 
55. Fundamental arrangement of the cutting teeth in four of the suborders or superfamilies of the Proboscidea................ : 
56. Map showing geographic distribution of types and referred specimens of the Deinotheriine.. a oft anchovces aelens 
57. Deinotherium giganteum in the British Museum, four views of reconstructed original Pppelsheim s minty, “After Andrews.... 
58. Deinotherium giganteum, first Eppelsheim skull and jaws. After cast in Darmstadt Museum. . 
59. Deinotherium giganteum, restored Epplesheim skull and jaws. After Falconer............... 0060s eee eee eee eee teens 
COND inothenumuientenmeavum dey blainvall esa yjoe iO Ule sen avat ayerostete sie chete tees es clcties -\eute e<ct a fale = tale al =)lo\oredoiedal hake eka lotletele i oledsiiaieteteiste 
61. Dinotherium gigantissimum, referred grinding teeth. After Stefainescu......... 0.0.02. 
62. Dinotherium gigantissimum, tibia and pes. After Stefdinescu............- 22:00 eee e eee e cece cece ese reste er er eeces 
63. Deinotherium bavaricum ref., mounted skeleton from Franzensbad, Bohemia, in Vienna Museum..............-+-+++55. 
64. Deinotherium bavaricum of Franzensbad, showing original and restored portions of mounted skeleton in Vienna Museum. . 
65. Deinotherium bavaricum in Vienna Museum. Model by Abel (1982)............... 2. cece eee cee eee tenet teen eeee 
66. Deinotherium, restoration of head. After Gregory and Osborn (1910)................ 2. c cece ee tee e eet eeeceees 
Bip Dei nolnentui wWestoratlones eAt tere Dela (O22) esc cre ratelec cee cis cic ieie e cle = oleae: 1 <ynsntse1cvanobodol bodelnfavoleloie elehe letelelee leit tele talanetee 
Te. [Daniele Wai e 2st as, Uy deuilalinshocosbsobosoccnos booed a PON nO OD BORD AOMOMOOS GON OO CDUGdS TOD DOCDDOUnOUG5O0C 
Bete DE MOEN UAT ROT WOOL UIST MON esr bY [DC MO OLATS i steaicis aie tetete) etoNayeyeesesy oe ote s)016 12s 21 op sts =o) fei eieheler foley oe erin fele/asevein ohayelelene¥ehevoraleyatale 
69: Demothervum, progressive increase in size of mandible........ 2.2... ee cee eee eee ee tees me tert e conan enesepoans 
70. Deinotherium bavaricum, D. giganteum, D. gigantissimum. Restorations (1932) by Flinsch.............. 2.265. ee eee eee eee 
71. Deinotherium, composite figure of deciduous and permanent dentition. After Lartet, eee eeee 
72. Proportionate restorations (1930) of Deinotherium (bavaricum, giganteum, gigantissimum) by Sterling...........6.. 0000+ e0es 
73. Deinotherium gigantissimum Stefainescu of the MAnzati Valley, Rumania. Restoration by Flinsch.....................-- 
74. Prodinotherium [Deinotherium] hungaricum Bhik, type figure............ 0... e cece cece cece eee ne eee eee e tence eeees 
75. Representatives of fourteen genera of the suborder Mastodontoidea up to the year 1935. Restorations by Flinsch... ...... 
76. Mastodonte de VOhio or Le Grand Mastodonte [Mastodon americanus], Cuvier’s typeS.... 0.00.00 eee eect teens 
77. Petit mastodonte |Turicius tapiroides| from Montabusard, Cuvier’s original type figure...........002. se eens 
78. Mastodonte a dents étroites [Trilophodon angustidens], Cuvier’s original type figure........ 0.6... 000 eee ee eee 
79. Mastodonte des Cordiliéres [Cordillerion andium], Cuvier’s original type figure.............. 000s cece ete eee 
80. Mastodonte humboldien |Cuvieronius humboldtii], Cuvier’s original type figure................. at ivaaunis-D Matte orale mare ee pelaene 
81. ‘Mastodon ohioticus’ [=M. (?)pavlowi ref.] molar from Russia. After Pavlow............. 06 cece eee cece eee tet eee e ees 
Soe Vattcnel Vins odonasmmounbedsm 184518465. avec cmecmieea sire vacie ie oie one «cin > cies eavivivsiomerelencuel amie eletiolals fllsheleleioaaalsyalerals 
So ALTCHS ViAStOMOM ASseMOUNTECMINL G40: con 1.08 sence auscy rman need meta pe ercus oi\20)+ a,ccjeta omledeleuelelnieltureynsissitna dedi sns scare ietenie/ stars 
84. ‘Two superior molars figured by Buffon in 1778, referable to Zygolophodon borsoni and Mastodon americanus...........++- 
Soe Vian onbig-boneylick boone County, wen buckys macei tet iitacts ce vatere orencv to 8 <ussid ara, are coor Hetero tatey skeyeuea clea erpialenoney 
86. Map showing geographic distribution of the types of the Mastodontine.................. 0c e cece cece ete cette eee e tees 
Si) Mastodonamericonus jaw andinterior dentition. After Warren sss sos oo5 esse ee accis ae so aeolian cela eeieisisielelerareierd= 
88. Pliomastodon matthewi and Palxomastodon intermedius, comparison of third inferior and superior molars...........-....++++ 
89. Evolution of molar ridge-crests in the Mastodonts, Oligocene to Pleistocene............-20 000. e eee eee eee teens 
90. Superior view of largest known jaws of Phiomia and Palzomastodon superposed for comparisOn..........0.5. 00000 eee rene 

91. Palzomastodon intermedius Matsumoto, type and paratype specimens............ 0.06 e eee cette eet e eee teen eee 
92. Palzomastodon intermedius Matsumoto, type and paratype grinding teeth.............. 50:0 e eee cece eee rete eee eee 












Key to hexabunodont upper and lower molar crowns of Palwomastodon intermedius... .. ce eee eee ees 
Detailed studies of third superior and inferior molars of Palzeomastodon intermedius and Meritherium lyonsi, M. trigodon, and 
AGIs ya) AA ee A eM ee PO Pe ORR Rah ary. iv cae TED Ud 2 ONSITE Qo ey 

Palzomastodon beadnelli Andrews, second type figure............ 0 ccs cece e ence cere eter ne enee er seneeesseeeeseeeee 
Palzomastodon beadnelli, referred limbs and vertebre. After Andrews............6.. 00-0 e eee ete reece eet tee eee 
Palzomastodonas'a forest browser. Restoration (1932) by Flinsch... .... 5.00122. c ee cee ee bo mans s ssleee sisletinsis ees 

Miomastodon and Pliomastodon type molars compared with referred molar of the true Mastodon americanus. After Osborn. . 
Miomastodon merriami Osborn, type figure (superior molars and tusks).. ae 

Miomastodon merriami referred mandible, inferior and superior molars andi tusks, Pawniee Cc mele re sinnadoi. After Frick. . 
Pliomastodon matthewi Osborn, second type and paratype figures........... 0... cece cee cece eee tte ee eee ete eter ences 
Phomastodon matihewi Osborn, third type and paratype figures....... 2... 00.02. c cece ce cae ers e resets eesseedseeeries 
Pliomastodon preereaaas ae Seen tees COty Pe MAOLALAy ss asic HAL taretereteee okie ates nUnt eke Done ey UA MONE Neh oes oe Re ees 

Pliomastodon Saidlersses oe type Skulls! awe obs eS 5 wc) ss oc, 5 Me ales auAea ee tel sear neva oe ot oer are ro 
Pitomasiodon vexillarvus Matthew; type lower Jaws. iocn == 2's) = «210s sielclels olely alale w eleles wilelelieteleliinle + =/aiabeais “1M oie tetatatet=tai eal 
Eeiiomasiodon vexillarius Matthew, type femurs sanvsi ss crete dete ale el cle =) aierete chisel ote ies ele et eve depts ste otete lies te eae ae a 
Mastodon americanus male and female. Restoration (1933) by Flinsch............. 0.6... c cece eee cette eee e eee 
Palzomastodon, Pliomastodon, Miomastodon, Mastodon, Zygolophodon, and Turicius. Restorations by Flinsch............ 
Superior grinding teeth of Zygolophodon and of Mastodon. After Buffon, 1778............ 0.060 e eee eee ee eee eens 
Le Grand Mastodonte, or Mastodonte de l’ Ohio [Mastodon americanus], Cuvier’s types....... 0.6.00. ce eee teens 
Ohio-Incognitum [Mastodon americanus|, Blumenbach’s original type figure... SRR 
Mastodon americanus from Catawba, Ohio, mounted skeleton in Ohio State Univ aire 
Mastodon americanus rugosidens Leidy, type figure (third left inferior molar)................. 00 cece eee e cece eee eee 
Ohio-Incognitum [Mastodon americanus], Blumenbach’s original type figure...... 2.0.0... 0c eee eee eee eee 
WMasiodoniamerrcanus: referred Third NtEriOT MOLAR. « «exe. =e sctsevee aie eie) eels) sv su chahete altel <lnfeteks hele eels felis fet ate N=Rel sUelests = tela = tpanies ie 
Mammut progenium [Mastodon progenius] Hay, type figure (lower jaw)... 2.22... 0. cece cee ete tee ees 
Mammut oregonense {Mastodon oregonensis| Hay, type figure (second left superior molar)... 2.2... 2.56.0 eee eee eee eee 
Mastodon americanus plicatus Osborn, type superior true molar serieS.............0... eee eee e eee ete e eee teen eee aeee 
Mastodon moodiei Barbour, type skull and mandible. After photographs............... 0.0.00 eee e cece e eect eee e eee 
Mastodon raki Frick, type figure (incomplete mandible with dentition)............... eee ee eee eee eee eens 
Mastodon americanus alaskensis Frick. Type figure (incomplete mandible with dentition) ................0. 00 eee eee aee 
Eurasiatic ancestors of the typical Mastodon of America. Theoretic migration lines to North America................ 
Geographic distribution of the typical mastodonts of the United States and Eastern Canada........................0005- 
Warren Mastodon [Mastodon americanus], mounted skeleton in the American Museum. . Gilson 
Whitfield Mastodon [Mastodon americanus], mounted skeleton in the Senckenberg Giscwme Micathtereti ‘Gaunaen 
Warren Mastodon [Mastodon americanus], superior, inferior, and lateral views of the tusks showing annular ees sea 
ATtED OSDOLIY 6 bo. occ eso ccs svee cially  alurbnere esta el Sidea, Sta S G0LG 5 Sach ClR Sta Ce gv SI ae NG RNG IR ene ein fe toot nee 
Elephas indicus (?) sumatranus, referred tusks showing annular growth rings. After photograph.....................-..+-. 
Mastodon americanus ref. from near Fulton, Indiana, palatal view of female skull... ............. 0.0 cee eee eee eee 
Mastodon americanus ref. from Otisville, New York, palatal view............ 6.000 cen scene cee cece esr tesuvuedave cust 
Warren Mastodon [Mastodon americanus], superior view of skull.. 
Mastodon acutidens sp. nov., male skull and jaw from Rochester, ade and shall calf of Ms americanus rah ran Haake 
S110) .00 AN A ne enn Mane fa SAR Se ers ons on Re Acorn vom moe er SS Se She 
Mastodon americanus ref. male and female crania, from Ashley and Fulton, Indiana .......... 
Mastodon americanus ref. from Buffalo, Kansas, crown view of male (?) palate and lower jaw of ‘milie- mee qaiiividuples 
Mastodon americanus ref. from near Fulton, Indiana, middle-aged female palate and lower jaw.. { 
Mastodon acutidens sp. nov. from Rochester, Indiana, showing sharp summits of partly fevelocad ales crowns ieeaeied 
from! a<young male: skull... sis sis.5 sree ce cio Rete ae eel Sete ee ete te eee aER een eS Te cP tat stale oe ice ev cle a etee tens ono 
American Mastodons along the ancient Missouri River of Kansas. After mural by Knight in the American Museum... . 
Map showing geographic distributon of types and referred specimens of Zygolophodon and Turicius. ny 
Miomastodon depereti sp. nov., Trilophodon pontileviensis, and Turicius tapiroides molars of the Lower Mogens ne Mayet 
Turicius turicensis i iapevoidesl and T'rilophodon pontileviensis molars of the Lower Miocene. After Mayet.............- 
Turicius turicensis referred molar, from Murinsel, Croatia (ef. 7. virgatidens). After Vacek...............000 55 ee ee eeee 


141. Turicius turicensis simorrensis Osborn, type molar. After Gaudry... jones, wlOe 
142. Cotypes of Mastodon [Stegolophodon] cautleyi Lydekker equal eee o Oxbars n. Bae. F plconee all Gautley = en pment 197 
143. Middle (=Etage Helvétien) and Upper (=Etage Tortonien) Miocene correlation. After Depéret (1905-1908) and Osborn 
(1910) jen So dahiereecs Gace ts dein atte eee Ee ee ee eee eee ere ee ee 201 
143a. Ascending phylogenetic succession of three species of Twricius and one of Zygolophodon. Restorations by Flinsch......... 201 
144. Head of Turicius turicensis Schinz, exhibiting supplementary superior incisive tusks. Reconstruction by Pontier and Anthony 202 
145. Cranium of Loxodonta africana, exhibiting reduplication of superior incisive tusks. After Anthony and Prouteaux. . ok 202 
146. Chief Lower Miocene localities (= Etage Burdigalien, Sables de l’Orléanais stage) and correlation of France, Suntesanenls foul 
Austria. After! Depéret: (1905-1908) -and@sborns (W910) meserere mera ierekece crcl ees co erehetre oe ree eee eee 204 
1A Zagolophodon pyrenaicus Lartet, type molar, After Gaidny-cacesesee ase eeieier aie ale eile eles eee eee 206 
148. Turicius turicensis simorrensis Osborn and Zygolophodon pyrenaicus Lartet, type molars..............6...0 0.0000 ee eee es 207 
149. Mastodon americanus ref., crown view of inferior dentition and jaw. After Warren................0.000.00 eee eeeeeee 208 — 
150. Zygolophodon borsoni, referred inferior dentition and jaw, from Vialette, France. After Lortet and Chantre............. 208 
151s Zygolonhodon borsonz, referred second/and third inferionprind ers) eee eee eee 208 
152aeZygolophodonborsony.caddochs Jentzsch, type figures (Olan) eerie ete eis eae sian ened etee et ated ees 208 
153. Zygolophodon borsoni, referred third superior molar from Le Petit-Rosey, near Lyons, France. After Lortet and Chantre. 208 
AS sZygolophodon:vorsont, Hays; type figure (molar) a-i.ccmee see eteete eeiiae Sain ele ee eee eee 209 
155. Zygolophodon borsoni [= Pliomastodon praetypica ref.], referred superior molar, from Neudorf, Austria. , After Vacek....... 210 
156. Zygolophodon borsoni, referred right inferior grinder, from Hidvég, Hungary. After Schlesinger........................ 210 
157. Zygolophodon borsoni, referred anterior half of third inferior grinder (erroneously attributed to ?North America). After 
Schlesingerhe ry hcp Cee Ae ae eee errr IS SOA oldies 20S Se A 210 
158. Zygolophodon borsoni and Mastodon americanus. Restorations by Flinsch.. sige S's cateighlacd Bein See Aiea SRS ee 
159. Theoretic migration lines of the forest- and swamp-loving Zygolophodon andl Tires MSc Sete Ne, Leia PAROS NA 215 
160. Petit mastodonte from Montabusard [Turicius tapiroides type]; molar described by Defay (1783), by Cuvier (1806)........ 217 
161. Petit mastodonte Cuvier, 1806 [Turicius tapiroides], new type figure. After Mayet................. 0.0 cece eee see cee eees 217 
162. Turicius turicensis, genotype and homceotype molars from the Upper Miocene of Elgg, Switzerland.................... 218 
G3, UM GOI TPE EIS GOTTA ROSES OS nov dal, viygole\HFabIRS (Gono EN) eo cosnnogcecanwaGedNmonn ooo honMooEoomoonbomoonouvosnonons 220 
DASenUmenisraizcie Warner uy pe wieune: (premolar-and firstimnolsim) ep ern eee oc eyes elise el epaeisircieeleiaicneirs ieee 220 
165. Turicius atticus, referred milk premolars, from Pikermi. Referred by Gaudry to Mastodon turicensis................-.-. 221 
166. Turicius turicensis, referred molar from Murinsel, Croatia (ef. 7. virgatidens). After Vacek....................-0-00-- 221 
167. Turicius virgatidens, referred molar from Terrasse vom Laaerberg, Austria, and Turicius(?) tapiroides, referred molar from 
Halunsdeylaulourainebrances Aditer Sehlesinperteay-m mre cle: cle ie sivtel= © sie\e seelerere hele aoe a tevin saeco eee 221 
IGSMmMICLUS Mi ingaLdens vonMeyer. type eure (Molars)s-eeseeccetar ce ones aei ics ke tatislele © sclale & 4 citreus ile Cckaery ese 222 
169. Long-jawed Trilophodon, the classic ‘Mastodon’ angustidens of Cuvier. After painting by Knight....................... 224 
iO Mee AntGMiCnAN Gn Serntieniuniusn COMPAarison: OliCramianeet seis sites eerie oni > aide «cdl ee dieser beets ns Ce oe eis oe eerie 227 
171. Phiomia, Serridentinus, Trilophodon, and Tetralophodon, evolution of the second and third inferior molars.................. 227 
ij 2a Craniaand incisive tusks inysix bunomastodontisublamiliess).-eee. is. 022 ace a. « «sa sialslecis actus 6 evan cele ae tps stele tiers 229 
173. Dental succession: Bunomastodontide and Elephantide. After Frick, 1926................. 0... c cece eee ee eee eens 230 
174. Dental succession in Mastodon, Rhynchotherium, Serridentinus, and Trilophodon [=Ocalientinus]. After Frick, 1926....... 232 
175. Dental succession in Rhynchotherium, Serridentinus, and Trilophodon [=Ocalientinus]. After Frick, 1926................ 233 
176. Deciduous premolars in Serridentinus, Trilophodon [=Ocalientinus], and Rhynchotherium. After Frick, 1926............. 234 
177. Deciduous and permanent premolars in the Serridentine and Longirostrine. After Frick, 1926.....................--- 235 
178. Palates of the small Phiomia minor and of the large P. wintoni, illustrating differences of facial proportion.............. 237 
ie LLOM eNO Ane ALintont SUpeLiOr denuitioniac elaine incite: cee eins see ete eae eeieie see eee cae meee 240 
180. Phiomia serridens Andrews and Beadnell, second type figure. After Andrews... ” AR etre hiammie tad, i) 
181. Phiomia wintoni (cf. serridens), juvenile dentition, compared with type lower jaw at P. Bestia Be POETICS DATO he 
[Sz eLnoniam@iunonrret, ared skull, and) P: wimion? rel. juvenile jawersas-cs eos. +++ he.sscseces eee se eeciesc se ceeds sc ececse 242 
183. Phiomia minor, skull and tusk, and P. wintoni jaw, composition drawing... .............. ccc e ee eee eee cece cece ee eeee 243 
184. Phiomia wintoni erroneously combined with jaw of Palzomastodon beadnelli. Andrews’ restoration of 1906................. 244 
185. Phiomia osborni Matsumoto, second type figure (mandible and dentition).................0 cee cece eee eee eee eee eens 245 
185a. Five species of Phiomia, in ascending geologic order. Restorations by Flinsch................0. 0000 cece cence ene teens 245 
LSD Pee NOMI DYGMLUSDepereti aby pe yUoUTe (MOLAL) axe tus crete oes ae TOE eee ovis ree APPS ee see eee 247 
187. Phiomia osborni type, P. pygmzus type, and Palzomastodon beadnelli ref., comparison of third inferior molars............. 247 

188. Lower jaws and comparative crania of the hyperlongirostral Trilophodonts, also of the Amebelodonts.................... 248 

189. Map showing geographic distribution of the types and referred specimens of Trilophodon and Amebelodon, also of the type 
DE LENT OT MTT (ETRE SPE EO, oe Pe Ee AS SS tpn ee ieee amet Pa ee eae ge 
190. Mastodonte a dents étroites [= Trilophodon angustidens], Cuvier’s original type figure................ 000000 
191. Petit mastodonte [= Trilophodon angustidens minutus], Cuvier’s original type figure............-....-20000c eee eeeeeeaee 
192. Trilophodon, Turicius, and Deinotherium, deciduous incisive tusks. After Stehlin.............. 000 cece eee eee eee eee 
193. Map of the Lower Miocene, Burdigalian, showing horizons of the Sables de |’Orléanais, ete. After Depéret (1905-1908) 
vere hCO/lefeyn cd (02) (1) eee nn, oe ne as ce a ar EO ena eee Cer ee est aay 
194. Trilophodon angustidens from Simorre, referred second inferior molar. After hee ee 
195. Trilophodon angustidens from Simorre, referred third inferior molar. After Gaudry... a te oe 
196. Superior milk premolars of the left maxilla in the Trilophodonts and Toteslaphadedie “After ede ST ee ery 
197. Trilophodon angustidens ref. from Simorre, succession of milk and permanent teeth. After Gaudry...................... 
198. Trilophodon angustidens gaillardi, type mandible. After pen sketch by Dr. Gaillard.. Ets 
199. Trilophodon and (?)Serridentinus, composition skeleton in Paris Museum; skeletal maeerae od oe: Eanes 
200. Hlephas |= Archidiskodon| meridionalis of Durfort, mounted skeleton, and Trilophodon angustidens, skull of mounted shelter 
mibaris: Museum, - After. pen sketches) by. Eanight 3 orjejsc.'s feted. hg nse nuctelndad spare wis oad ole wiclouel rs felctel a teieisieh eles a ee 
201. Trilophodon angustidens var. austro-germanicus Wegner, type figure (molars)... hs eee 
202. Trilophodon angustidens var. libycus Fourtau, type figure (molars) and referred left Lowen: je JOW.» Cuntangsawcrate teen 
203. Trilophodon and Tetralophodon, milk dentition. After Schlesinger.............. 0.00 cece cee eee eee tee ee ee ee enna 
204. Lower Pliocene (Etage Pontien) of western Eurasia; Middle Pliocene (Etage Plaisancien) in part, as far east as the 
Caspian Sea; Upper Pliocene (Etage Astien), a few important localities.............0..0.0 00000 e cee c ee cece eee eee 
205. Trilophodon (Choerolophodon) pentelicus Gaudry and Lartet, type figure (juvenile skull and teeth). ....................... 
206. Trilophodon (Choerolophodon) pentelicus type (juvenile skull and teeth). After Gaudry...........0..0.00..0 00. eee eee eae 
207. Trilophodon (Choerolophodon) pentelicus, referred superior milk dentition. Brown Collection.. 
208. Trilophodon (Choerolophodon) pentelicus, referred juvenile cranium and milk dentiton. After Dalene sh vray siete ree 
Pm Uropnodon.pandionis EH aleoner, type figure. (nolan) soce. x.0¢-< sn c's aw wie ats yeyerorekete neva aleletepelareaa ereislp pieleviat cele oie eee 
pues Uimiophodon pandionis; type molar:, . After Liydelkkers«.i224.-/...feh)ereerci » aio ofa « mpeyesetene dete sat aru eyeieie lies ee ee ee ee 
211. Trilophodon palxindicus Lydekker, type figure (third inferior molar) and associated second superior molar.............. 
212. Trilophodon palzindicus, referred second inferior molar. After Lydekker................0 0000 cece eee cee eee cee 
213. Trilophodon cooperi, referred third inferior molars from the Bugti Hills, Baluchistan....................000 2.000 c cece eee 
cif el relophoden cooper, referred molar fragment. After Liydekker i. 2:0. 5. 2). os nis eids ole antshay sees ere apy sep ome ie e 
215. Trilophodon cooperi, referred fourth right superior true premolar. After Lydekker... ......... 0.0... 
ibe Hemimastodon crepusculs Pilgrim. [=Sumval], type figure:(molar):..... 2... 0-5-2. > «oe njor -emeniviels tia meen stele ote pie 
217. Trilophodon sp., fourth superior deciduous premolar, referred by Pilgrim to Hemimastodon crepusculi...................- 
218. Trilophodon angustidens gaillardi, type, and Trilophodon chinjiensis, holotype (mandibles)..................022000eee00005 
Pomuriopnodon macrognathus Pilgrim, tyPe)MOlAar. « o,< 22s 05s) 01%,~ e:siece + atae os = ecayetsue fafa pate ehacete tetera ale 
SeUeerilophodon paleindicus, referred third:right inferior molar: ........ < = =:=:=)¢,, 0.1/0 sessions Sel ee ieee eine ia et 
221. Map of Bugti Hills, Lower Miocene-Upper Oligocene (?) horizons west of Dera Bugti, Baluchistan....................... 
222. Evolution of the central conules in Trilophodon, showing T. cooperi type. .......--- 6. eee cc cee ee ee eee tere eens 
Pa Lrziophodon cooper Osborn, paratype—‘ older. palate’ < s.<..\...:. . «10 sx aes sles ale ales oe olene rete cbeieiaysieieet oie ei ee 
224. Trilophodon inopinatus Borissiak and Beliaeva, type figures (cranium, mandible, tusks, and grinding teeth)................ 
252 Classic fossil mammalian exposures of northwestern: India... «2s: «= = dls alec a etemiee Sederel « sleseleie dese ¢ muatnlniel sie ele en 
ene il riophodon sendaicus. Matsumoto, type figure (molar)... <1.a+slotsnelennehe wiensee ee esta eee sine abs eee Lee 
Bowe wumlonhodon esseloornensis Klihn, type figure: Qnolan) «-ssezesscroraysiayorels 21d aototatabataie i aaeieielele le bios aint eis eee eee 
228. Trilophodon steinheimensis Klihn, type figures (molar). After O. Fraas, 1870, and Klahn, 1922......................... 
Peo mucus wahinermensis. Kahn, cotype figures! (mOlATS)2<* 5.5 «10/1 o0a «tess wcoltene abaens tae tesaeetera al tae faire eee 
230. Trilophodon pontileviensis Mayet—Fourtau, cotype figures (molars). Also type molar of Mzomastodon depereti sp. nov. 
BEHOX INT AOS cisssisicrete ict systersiae aia’) Surslalinin ays 9 casera taser eeee © ip lam See SIS chee ete ea ear os ae 
STmEumippnoaon eimplicidens Osborn, type Molar: vss < oras-ajes wis y+ os. vin sie © 1s aioe atele oaks ana ale aria oats 
paeweelreophodon obscurus Leidy, ‘type figure. (MmOlar) 500) ie swiens 0:0 sre cfs aud aveie ales daialer stele lee ehcretatee tee Siete acne eee 
tame lnlonnodon.obecurus, new type figure, «'s-:cicce saan sividiens, pietnceteaye teeere@ Bsa anete everaetebien: Mette = char blaik ieee Aes eee ee 
234. Trilophodonts, primitive and progressive, of Eurasia and North America. Restorations by Flinsch..................... 
235. Trilophodon (Genomastodon) osborni, conjectural restoration. After Barbour.............0..0..0.0 00000 e eee ee 
236. Trilophodon dinotherioides Andrews, type figure (mandible, third molar).................060 0000000 e eee cee eee eees 
237. Trilophodon (Genomastodon) willistoni Barbour, juvenile type skull and jaws ................0...0. 000 cece eee eee eee ees 
238. Trilophodon (Genomastodon) willistont Barbour, type lower jaw. ..... heals fo, Crepe rier aioe teens 
239. Trilophodon (Genomastodon) willistoni Barbour, type lower jaw. Aver sketeh a Barbour: HA Sin catia oben ene res 



240. Trilophodon (Genomastodon) willistoni Barbour, type skull and jaws. After pencil sketch by Osborn.................... 292 
241. Trilophodon (Genomastodon) willistoni, referred humerus and five tusks showing varying widths of enamel bands.......... 293 
242. Lower to Middle Pliocene exposures of Devil’s Gulch on south side of Niobrara River, Brown Co., Nebraska.............. 294 
243. Trilophodon (Megabelodon) lulli Barbour, type skull and jaws............. 00 cece eee eee ee eee eee 295 
244. Trilophodon (Megabelodon) lulli Barbour, type figure (five views of mandible). . ay 296 
245. Trilophodon (Megabelodon) lulli Barbour, type mandible in comparison with mz nandiblest eal costes rocmiie fe re T. 
Aone Ore 18 (VIC) NDT ee meeere coin radio. cta a0 Conran Oe Cone Oon GoaeCnao modo UMC cd sboconymacccoooncod bes 297 

246. Trilophodon ligoniferus Cope and Matthew, type figure (mandible)................. 2000s eee e eee eee eee ee eee eee eee 298 
247. Trilophodon (Genomastodon) osborni Barbour, type mandible................ 00.00 e eee cee ee tect e teeter eeces 299 
248. Trilophodon (Genomastodon) osborni Barbour, type skull and skeleton................ 0600 cece eee e terete eee 300 
249. Trilophodon (Genomastodon) osborni Barbour, type superior and inferior grinding teeth, also mandible. After sketches by 

(01 0,0) 5 Veni ae ee DIO ERI KERIO a Ot Uc hO EM COO a OO On eC OD OA otoegomemonc gsr: 301 
250. Trilophodon (Genomastodon) osborni Barbour, type skull and mandible (skull in plaster ‘cinches’), also conjectural restoration 

OfMhendmAtitersBarbour/sce este tere rere cere sete ore ee eicie io oenl cs dle Staualalcta aie s ts Meggelis elelbucee Gioteie t-te leva tnt eae oan 301 
251. Trilophodon (Genomastodon) osborni Barbour, type ‘‘fifth and sixth” superior molars... ...........0 0.0 eee eee e cece eee 301 
252. Trilophodon (Genomastodon) osborni Barbour, type ‘‘fifth and sixth” inferior molars. . 25 301 
253. Trilophodon (Genomastodon) osborni, type skull, also referred skeletal parts compared oan itose an Elephas indleaus Agee 

IBYid ofc iia Carin OIe ROC ne LOCC eIUES Clade ER rein MOCO ee eT RETO A RITA Aoo oats s00 On: 302 
254. Trilophodon (Genomastodon) osborni, left superior tusk found in pit near skull. After Barbour......................0000+++. 308 
255. Trilophodon (Genomastodon) osborni and Elephas indicus forelimbs compared. After Barbour ..............22....0000 303 
256. Trilophodon (Genomastodon) osborni Barbour, type skull and mandible...............0 6.0 6c te tee ees 304 
Zoe eurilophodon giganteus' Osborn, type mounted skeletomse..-25 «arise «cles «fc ecrcieictelc co ole ee evelele eels) selene cree) chettehe reheat eae 305 
258. Trilophodon giganteus Osborn, type figure (mandible) compared with mandibles of other Longirostrines.................. 305 
259. Trilophodon giganteus Osborn, type skull and jaws, also inferior dentition. ...... 00 ce eee eee eee eee 306 
2602) eA mebelodon (Imlophoden) hicks: Cook, type figure (miolar))) c~. 6s. - 2 2 se ed ee eee oem vee mielererctede tries ae 307 
DGily Amebelodon (lirtlophodon) hicks? Cook, type mieure (mMandible)c cee... -.-< soo. os ye see © bie ee rel steeper 308 
262. Amebelodon (Trilophodon) hicksi, referred superior molar associated with type, presumably of same individual. After Cook. 309 
263. Amebelodon (Trilophodon) paladentatus Cook, type figure (mandible)................0. 00 cece eee eee eee eee eens 310 
264. Amebelodon (Trilophodon) paladentatus Cook, type figure (incomplete inferior molar)................0..0.00 00 ecu eee ee eee 310 
265. Parts of two superior tusks found with the types of Amebelodon (Trilophodon) hicksi and A. (T.) paladentatus. After Cook.. 310 
Bob lmlophodon abvelmbanbour, bype igure: (mandible): s.c- cs onc cc eels v cic ce vee jos cles aie cece olele o-sislelel se late eRe teen ae 311 
267. Trilophodon fricki Peterson, type cranium, mandible, and dentition. After original drawings..................--020000+ 313 
268. Trilophodon fricki Peterson, restoration. After pencil drawing by Dr. A. Avinoff................... 00 cece eee e eee e eens 314 
269. Trilophodon phippsi Cook, type mounted skeleton in Colorado Museum of Natural ee After aes a eae 315 
270. Trilophodon phippsi Cook, type skull and jaws. After photographs. . pdadilend saue 4g ps a dee a 6 alaralenelele Sle Ona eT a eee 
Vi linLopLolon nip pst TeLerreds CLAIM) ANG! JAWS, 2150 cress om cis} cis ie ges 0 sac «-scbse oie dune ov ciel ehouene euelole eis gue e ieee ean 316 
272. Mio-Pliocene beds of Brown County, Nebraska, generalized columnar section. . Sue 317 
273. Key to intermingling of the Longirostrine, Serridentine, Platybelodontine, ead Humboldtiies lene dhe aaciene Nisbrwed 

HSINVOIRS a '9. c:csck o, Gtk Ox 0. cr tr ROAR. age aI CHEE One ace Rn on not c 317 
274. Lower to Middle Pliocene fossil localities in north central Nebraska................. 0c cee e eee e eee cece eee eeeeeeeeraes 318 
Lip me uLO phonon pojoaquensis brick, type figure (SKIL) \oq.acesicyeesie ees vis © lic ccc « c/ele/sichs es seine ere Seley slater One) cierereel one 320 
276. Trilophodon pojoaquensis Frick, type molar, and Ocalientinus ojocaliensis Frick, referred 321 
277. World migration lines of the typical Trilophodon (T. angustidens), of the ‘shovel-tuskers’ (Phiomia, Amebelodon), and of the 

‘prod-tuskers’ (T'rilophodon), from Africa (theoretical), Eurasia, India, and North America...........0-..00eeee0es 321 
278. Ocalientinus ojocaliensis, referred mandible and inferior molar, and Trilophodon pojoaquensis, largest cranium, also superior 

THOME ACE REI CHe  GMISINAl APAWIN ES) shai ffs tease aavs sels Sele tin leiesatwou oe waly wae woaielnes sheen ace 322 
279. Trilophodon cruziensis Frick, type cranium and mandible, also Ocalientinus ojocaliensis referred inferior molar. After Frick 

(onpind dra wines) sree ce acne ave -ceaysoiervact Saw. c oaths inact acgera leh de eee ee eee 323 
280. ‘Prod-tuskers’: Trilophodon (Tatabelodon) riograndensis Frick, genotype mandible; T. (Tatabelodon) gregorii Frick, type 

cranium and mandible. Also palate referred by Frick to Tatabelodon gregorii, by Osborn to Trilophodon osborni. After 

EMC Ka (OMPINAMCGLAWINGS erway tse Mee tek cls coe oe foe buat Pan lee ee ee et ee 325 
281. Trilophodon, (?) Megabelodon, Ocalientinus, Serridentinus, and Trobelodon. Frick’s mandibular types. Original drawings ... 327 
282. ‘Oblique-tuskers’ (T'rilophodon) and ‘uproot-tuskers’ (Serridentinus, Ocalientinus), comparison of mandibles..............- 328 
283. Resemblances and contrasts between the ancestral Oligocene Phiomia, Mio-Pliocene ‘uproot-tusker’ Trilophodon, Plio- 

cene ‘shovel-tusker’ A mebelodon, and tuskless ‘spoon-bill’ Trilophodon (Megabelodon) lulli............ 0.0. ccc eeeeee 330 
284. Amebelodon fricki, ‘shovel-tusker’ of Nebraska. Restoration by Flinsch...............0.cccceccecuccvceccuscueesees 332 
285. Platybelodon grangeri, ‘flat-tusker’ of Mongolia. Restoration by Flinsch...............00-0ccc ccc cecceccecceccseucees 332 

286. Contrasts between the ‘flat-tusker’ Platybelodon, the ‘shovel-tusker’ Amebelodon, the ‘uproot-tuskers’ T'rilophodon chin- 
jiensis and Serridentinus gobiensis—of Mio-Pliocene age, and the probable ancestral Oligocene Phiomia osborni ....... 
Sama mepelonon irc. Barbour, type firure (mandible), .:.is1se« avs ass w nls wdled eiplorew lie ales autatbiwisiates winface wie dalneliiences 6 
PPE ONAISLOGO ITICKE BAL DOLE, TYPO NOIR. «sis s:<.s52 «oe Simian prauatdey ela hinnd As <b ain ret retabootbeeeete ee iene tena neal ee ee ee 
men wAmebelodon fricki, referred inferior molar. After Barbour. «o/s 0 c' »ss <4 » be paises» » pavalelers pislzie wy slenclaed yia@is win ala letaraly 
200: Amebelodon fricki Barbour, second type figure of mandible............+..sseseecsecsesssacceausseessaenssssssssne> 
291. Amebelodon fricki, brush drawing of type mandibular tusks. After Barbour................06 0000000 eecceeececeeeeeees 
292. Amebelodon fricki, conjectural restoration. After Barbour......... 5 o's, cap a9 elendsauk we ¥ PES Oe ire Laces eee 
ea. Amebelodon fricki, revised restoration of head. After Barbour .....,.. + «s+ «+s» dees cs vo sewwis ee oo uivivne bpieb ww pisle'e oes 
204. Amebelodon sinclair: Barbour, type figure (mandibular tusk)............0.cccccseccceccecscsccvcecssavnvvvvtecsoees 
eemaenenacaon imarinative sketch. After Barbour; .<s.<assvs vss bisa bald 9 vis alee 02 viekivy chores DEE Deane eee 
296. Torynobelodon loomisi Barbour, type figure (mandibular tusk)........... 0.00000 cece cece eee eee cc cseees scenes se 
297. Torynobelodon loomisi, conjectural restoration of mandible. ‘After Barbour sss cs.2.+-s°s.k se soca, Sees oa ee eke eee ee 
298. Torynobelodon loomisi, conjectural restoration in supposed marshy habitat. Afte ter Barbour <tavuishe hang Vas FRAG Oa ab 
299. ‘Mastodon’ angustidens Cuvier, type. After one of the original casts in Mantell Collection, presented by the British Mu- 
Beum: to the American Museums : 30:3 sss eress vv: » «vin sis oi¢plelatomtdlongtale stalvisio aps! Derehk beens, eas eee 
300. 14,000,000-year evolution of the Trilophodonts, from the Trilophodon cooperi of Baluchistan to the 7’. macrognathus Mio-Plio- 
mangapare OF Ohinil, Lig... .. ..<:5.s. state ews War enya uvets tasleus-avin astys ley 0s laue /ottdSU ele te PS vets ce teen at i ote le eds a a ete ee aan ot 
301. Tetralophodon punjabiensis and T. campester. Restorations by Flinsch...............00000 00 e ec eens ec eee eee ee euees 
302. Tetralophodonts of Eurasia and North America. Restorations by Flinsch...............0..00ceeeeee cee eeeeeeneeeees 
303. Tetralophodon and Trilophodon, superior intermediate molars. After Falconer.................000000000ee cece eeeee 
304. Tetralophodon, Choerolophodon, and Trilophodon, superior milk premolars of left maxilla.................. ..0.0202005. 
305. Synconolophus dhokpathanensis Osborn, type cranium. After photograph..............0000 00000 c cece eee eee eee eeeeee 
306. Tetralophodon punjabiensis, referred cranium. After photograph.................002ceeecceee cc eeecceeeeteeeceececs 
307. Map showing geographic distribution of the types and referred specimens of the Tetralophodontin. .. ‘ 
308. Anteroposterior evolution and hypsodonty of third superior and inferior molars in Tetralophodon, fron iene to pincer of 
BRIO ENG 2 vn! os ns ces fv 4A adm ehho alk mage wtw\sartisn nina) 5 bv bey ¥. Chapin Boda mo bm hp RRO LRT ye 2 ES Se 
309. Tetralophodon (Lydekkeria) falconeri Lydekker, type figures (jaw and molar)... .......06060 600 e cee eee 
310. Tetralophodon (Lydekkeria) (?) falconeri, referred molars. After Faleoner and Cautley................6.00000-0000000 
Bit. st emalophodon (Lydekkeria) sinensis Koken, type figure (molar)... .0+ 63 ss s)< seu wie Ws slebslale f viele beisiaety eleles ails 
Bie Lvemalophodon longsrostris Kaup, type figure (mandible).......... 4.2... >.ss «ss ps .evs alsin nad eieleldi seis slaiahealals nie Re a 
313. Tetralophodon longirostris. WKaup’s composition of referred skull and type jaws. After Owen, figured as ‘Mastodon’ 
Oy 0 a enn AMP r Se A WIS Sh es 
314. Mastodon grandis Kaup and Scholl [= Tetralophodon longirostris|, type molar. ...... 0.0... 66 00 eee cece eee cence ee 
315. Tetralophodon longirostris, referred palate and superior dentition. After Kaup.............. 00 cee cece e eens 
316. Kaup’s type jaw of Tetralophodon longirostris compared with Phiomia, Serridentinus, and Trilophodon.................. 
317. Tetralophodon grandincisivus Schlesinger, type figure (lower tusk)............00 000 e cece ence ee ee tent eee ee eee enenees 
318. Tetralophodon grandincisivus Schlesinger, superior molar {paratype, Osborn]............. 6.006 cee eee eee eee eee 
319. Tetralophodon longirostris, referred skull of young individual. After Schlesinger. ..............6 0. cece eee ee eee eee eee 
320. Tetralophodon (Trilophodon?) grandincisivus, reconstruction of referred skull and jaws. After seidietin 1 i tensa Ree 
321. Tetralophodon grandincisivus (?), referred superior and inferior grinders. After Schlesinger... eee 
322. Tetralophodon punjabiensis Lydekker, one of the cotypes (Osborn’s lectotype). After Sader » sys natec Benya fat 
323. Tetralophodon punjabiensis Lydekker, new lectotype figure by Osborn...............00 02 c cece eee cece eee eeeeeeeeees 
324. Tetralophodon punjabiensis, referred cranium in Brown Collection.. P 
325. Tetralophodon campester type and T. punjabiensis ref. (molars). Contrast md resemblances, 
326. Tetralophodon bumiajuensis van der Maarel, type figures (cranium, jaw, and grinders) ............... 0.000020 eee eee eee 
327. Map of central Java, showing type horizon of Tetralophodon bumiajuensis..... 0... 0.6 ccc ce eee teen eeees 
328. Serridentinus and Tetralophodon grinding tooth opposition. .............000 00000 c cee eee ee eee eee cee ees 
fo, xtreme Pliocene evolution of T'etralophodon molars. isis. << sistew's wills ols pikes WSF HRI tll Wales UR eS Re 
330. Tetralophodon campester Cope, type. Second figure, after Cope and Matthew. . nee 
331. Tetralophodon campester Cope, type skull and jaws, compared with restored skull and je jaws sof Tr. longivealesn Kanwss s Seca 
332. Tetralophodon elegans Hay, type figure (molar), also referred (?) Trilophodon molar from near Waco, Texas. After Hay... .. 
333. Trilophodon (?Tetralophodon) brazosius Hay, type figure (part of right ramus of lower jaw). ..............-.-......000005 
334. Tetralophodon fricki sp. nov., type palate and dentiton. After Frick (original drawings)...................000000eeeeee 
335. Tetralophodon fricki and Platybelodon grangeri crania compared.................. cece cece ee ence eee eneetsesseces 



















Tetralophodon campester Cope, type palate.............. 00sec ence eect eee etree eee ene tent ne tenet n ees n ee eee 
Tetralophodon (Morrillia) barbouri Osborn, type superior molar........--..- +. +--+. eee eee eee eee 
Tetralophodon punjabiensis and 7’. longirostris, type molars compared.......... 1.60.60. e eects 
Tetralophodon precampester Osborn [= Tetralophodon (Morrillia) barbouri ref.], type superior molar.............-.-.-.+-+4+5 
World migration of Tetralophodon. Restorations by, Flinsch: 9,-2.3... ee ete ee ees ce ele ee fe eee ele eee eee 
Ocalientinus (Serridentinus) .floridanus Leidy. Restoration by Flinsch 
Serridentinus productus Cope. Restoration by Flinsch 
Serridentinus serridens Cope, type figure (molar) 
Serridentunusyprognessis) OSMOLI it ye 11 OLE Tatstereretercteretctcietereneretere onereterer sven reee eeee teens eens ae eee ed eee eee tae ee 
Map showing geographic distribution of the types and referred specimens of the Serridentinz and Platybelodontine......... 
Proboscidean localities of the eastern coast of the United States (map)........... 0... c cece eee eee eee eee eee 
Serridentines of the Florida Pliocene and contemporary Hipparion, Pliomastodon, rhinoceroses, and camels. After 
IME KHIR O10) S) Senaatoce GO aree a due ac GebnI0 F600) cb nocd GBA uraDoDSSOU ME a oad bo ce Oho 70 0.Cod lIn0s ogo c00.00,0.0'6Si010 
Contrast between hyperlongirostral species of Trilophodon and medilongirostral species of Serridentinus and Ocalientinus. 
IRGH IOAN lone) ONS Nee cossactooanc nim adc Odoo UomrgnA CONES QoG AGRGO OOOO OMG od SodoSD ab onOoOCoceDooRDEDOQOCC 
Serridentinus and Tetralophodon grinding teeth superposed..............-. 0200s cee eee cece ee ete e eens 
Contrast of adult superior and inferior molars in Tetralophodon, Morrillia, and Serridentinus 
Serridentinus, Ocalientinus, and Trilophodon inferior grinders. Comparative series........ sec cee cece eee eee 
sau ridentinus suDlapirgiieds pellet, cotype mola) compared with Trilophodon angustidens referred. After Schlesinger 

Serena Pilani Suilesiagee CObyPelmio lar ye eae Rowenta eke Se sl Lo teae Sg, UREN opel Malet Met ed sb altel teas ete ra moyen cs tavarets 
Serridentinus mongoliensis Osborn, type figure (inferior dentition)................. 0.0. c cece ence ence eee eee eee nee enees 
Ocalientinus (Serridentinus) florescens Osborn, type figure (molar) 
Maprotebune Gurstableland wimmer) Mom polistene cn crterreyeertete tet ie teie rete on cis rete teed eke ated Rete cNe leet = =P =aeateene ee e ea 
Senmaentunus qoniensis Osborn) and (Granger; ity Pepe une emye ceva cle =) cfelsyeyas\-\. 1 staljete eteleielielel obchetete efvene cates Newey etal etal iatateleters 
Pliocene and Lower Pleistocene Proboscidea, Perissodactyla, Artiodactyla, and Edentata of the Staked Plains of Texas 
andusanweednro Bedsrot Arizona (TM Ap) ease cistesiscloreverete teas euch eters ciel eve <itia evn ele ove: quatelatetafelvatisvelot ays tleNejoretyatey tats eons e erate 
IMiosPliocenewlertiariesto® Hloriday(map) = Atter SimnapSOMs-y peer cies seers scrote et etieiclel elev eteuste vote eevee otebettst si ielel sielnaetcitaeare 
Serridentinus progressus Osborn, type figure (left ramus of lower jaw).. fi 
Serridentinus progressus Osborn, new figure of type jaw, compared iti refered sieull aad ideeaned af S. ne nee 
diagrammatic figure of type inferior third molar of both species 
Serres (Oro Coie, inane eaiies (CK OIET DS ooo ca 500 Ao OUD B OOO EEEIEn Dae ho nD AO ObEO An cono ooGunmclsa ooo onc 
Senmdenunusiproavusi@©ope, newlimure of typeby Osbormtet-c scence. 26 oc le ee ce fans oe ieee sole 8 epee cseuieless atseieyaiys 
Serridentinus productus and ‘S. serridens, palatal view of cramia..............22 00.002 eee ee eee e etc e wesc ees neee 
SERIO CTIUMUSEPROCUCLUS HIN OGEel DyaBUNSCO Ns yecerse = sete two EE coo axes ©. SMe ees chays le loletereleleverecels Siar stellen aatetalaiemamete 
Serridentinus serridens, referred skull from Clarendon of Texas 
ERPREETIUSETROCUCLES | Gopen by Per Une) (MANIGIDIC) see creretctersteteletetcre o/s) ieee) el «la encheletee tntehe ci ete atefetegeuayer sere atenattararecerneiegs 
Serridentinus productus juvenile and adult (referred and neotype) jaws compared with referred adult jaw of Ocalientinus 
OJ OCUULENETS MMU ALG ETOEUET GK oc e's rele foicgehe Jeeta ote EO a met erat eats hc si-s ais ahs Bushee attsla ceed uaoMb NL Meena ele Yate bey ay cTene Sonata evenevene felt 
Serridentinus productus, referred skull, compared with S. progressus, lower jaws and teeth; also enlarged inferior molar of S. 
(BING IIN ELIS TH Siro 09, aoc RENEE Eso IESE pene RORCH GN SI OVE OO Ch LOO Oe RS OP EAE OO GO Chen a nar OE Cea Cice 
Serridentinus productus, referred skull and jaws from Clarendon beds................0.0 60.00 cece tee teens 
Serridentinus productus, adult skull and restored skeleton in American Museum...............0200 0000 eee eee eee eens 
Trilophodon giganteus type and Serridentinus productus ref., restored skeletons in American Museum; also outline restora- 
TLOTIS OE ge HGLELOSC Lites eeepc eres etc vcrate toe, Sane viernes erent evarcin Ae Sram SRA ORI ces paeeetrecner ee ae cali ay cues Slcle CiaeePene ears 
Primitive Serridentines of Eurasia and North America. Restorations by Flinsch...................0..0. 5.0 eee eee eee 
World migration and adaptive radiation of the Serridentine Mastodonts (map).................00 00 eee e eee eee eens 
Ocalientinus (Serridentinus) republicanus Osborn, type and paratype figures (mandibles) 

Ocalientinus (Serridentinus) republicanus Osborn, type, paratype, and referred dentition. After Osborn................... 
Ocalientinus (Serridentinus) bifoliatus Osborn, type molar and paratype jaw............ 000 cece ee te eee 
Ocalientinus (Serridentinus) floridanus Leidy, type and one of the metatype molars. After Leidy and Lueas............... 5 

Ocalientinus (Serridentinus) floridanus Leidy, type and one of the metatype molars 
Ocalientinus (Serridentinus) floridanus Leidy, type and metatype molars................ 0. eee cee ee eee tee eens 
Ocalientinus (Serridentinus) floridanus Leidy, second type and metatype figures (molars) 
Ocalientinus (Serridentinus) floridanus leidii Frick, new figure of type molars............00. 00000 c cece eee eee eee eee 
Ocalientinus (Serridentinus) obliquidens Osborn, type, paratype, and referred molars................000 000 cece eee eee ae 


383. Ocalientinus (Serridentinus) obliquidens Osborn, type and paratype molars compared with referred molar of Mastodon ameri- 
ATLA sacar ara air wi vierofistal oat do. Buh ar ass cave "avtnn, oh aecMI tea geND| oho abate Ta Mite alarts |S TNg atel Re ee ne ete oral aaa a Re ee 421 
384. Ocalientinus (Serridentinus) obliquidens, referred molar, from Tarboro, North Carolina. After i. PE ae rete ae 421 
385. Serridentinus and Ocalientinus, comparison of inferior grinding teeth.. PORE i eye Seiten, SAL 
386. Map showing exposures of the Clarendon horizon of the ‘Staked Plains’ of mewaas ibe Depa a ie ea ahaa 0216 bie teen ie 
387. Serridentinus serridens Cope, referred young adult skull and jaws, Clarendon of ecann Pye a aioe fond ed a tat De Satine were 425 
Ban mn eIEaentuis: sereaens Cope: type figure (molar) coach Oe. Relea I le eee ne eee 423 
Baum Seucentnus serriaens: Copesnew type sale as oafe aie avdls eis ae Shane AIO tet olin ee caer IE soe ot 423 
Bola SeTAEeNmus serriaens, reterred dentition’ of young: adult... fn: J cigislw zoe w slag vale viele lel oe ere areee rd oes ais ois eetiehclere rete 424 
391. Serridentinus anguirivalis Osborn, type figure (type and paratype molars and referred superior tusk fragments).......... 425 
392. Sheep Creek to Snake Creek exposures, Sioux County, Nebraska...........0 00.0 cece cece cere seers ee eeeeeeeneeceecs 426 
393. Sections through Sheep Creek and Snake Creek quarries, Sioux County, Nebraska... .............00 cece eee eee eee 426 
394. Fauna (Mastodonts, Serridentines, and Rhynchorostrines), Snake Creek B horizon, western Nebraska.................. 428 
Bop SEMOchiInus serridensicymarronie Cope, type Ligure (NOT) =... . sa scsels oss oles Sie cls crataeio le clots eens eae ae eS 429 
Buoresenmzaeninus orewsterensis Osborn, type figure (mOlar) ss .< <-./..0 </.2:s/ieorvein » aes oa s'e 0 se clu lore aye elelaleierars cretion eee 430 
Baie serociodon(?) precursor Cope, type figure (molar): ve soci ses cen See ele ete lee ela lene late G Sieaeeeak Ee eee 431 
Bose seruciodon(?) precursor Cope, new typeveure. <1; Lisi sv’ sis. sve vic ic-«- ablwakale els Blalavecicheleeie: Sat Cfow eles eubieas nv nies oe ele eee 431 
399. Three teeth of Serbelodon(?) precursor, referred by Cope to Dibelodon tropicus.................+-- ere ee ee 431 
Bue senrueniunus quatemalensis Osbornstype figure’ (molar) i2.. wsk.ccoestie « eictela: «elo ole ev ale aie eine biela rs bite Saeed 432 
401. Serridentinus guatemalensis Osborn, after photograph of type molar.................0.ccc cece cence eee e eee e tee e eee eees 432 
402. Ocalientinus ojocaliensis Frick, type and referred specimens. After Frick (original drawings)................-.....2.055 434 
Brae eOcanentnus, progressive evolution of the pentalophid:.:...dscscces © siniel ciciets « o lulbe Vee leie tele fistels & citi ietee) ene aera ays 435 
404. Ocalientinus ojocaliensis, referred third superior and inferior molars.................0 000s ee cece cece cece eect ee eeeeees 437 
405. Ocalientinus ojocaliensis, referred skull and jaws first described by Frick as paratypes of Trilophodon pojoaquensis........... 438 
406. Serbelodon and Trobelodon, blunt ‘shovel-tuskers.’ Restorations by Flinsch...............0000 00 eee eee eens 441 
407. Adaptive radiation of three types of Serridentine crania, jaws, tusks, and grinding teeth.....................0.0000eeeeee. 441 
408. Serbelodon barbourensis and Trobelodon taoensis, genotype and referred specimens..............00 00 cee eee eee ee eee eee ees 442 
Bao a servelodon.and .Platybelodon, shovel-tusks compared: «ir. «.s. 6 2 aise-sele has kok. facie wos otal elaiel lee aos tea else oe eve ee 445 
410. Serbelodon, Platybelodon, Torynobelodon, and Amebelodon tusks superposed. .. 2 MeRie na Mths aes 
411. Trobelodon tacensis Frick, type figure (cranium, mandible, molars), also reine cranium. wiAlPbee Frick i(acvad desedaatal 446 
Bue jeriaentinus barstonis Frick, type figure (premolars)..../655\0 ce 5 Sasha w oaielele © = otee tele Geena Rie en ae ae Sate eee 447 
413. India, proboscidean life zones and geologic horizons, Pilgrim (1910-1927), Pilgrim-Osborn (1927-1932), Osborn-Colbert 
((3 5) eee eee es Pr ren Se ERE ri rah G Paes Gee nC oe ae ioe 448 
414. Proboscidean life zones (Osborn, 1935) within the geologic horizons of India (Pilgrim, 1905-1932) ........................ 449 
415. Molar evolution of the Serridentine in India, compared with Synconolophus and Tetralophodon molars................... 400 
AIG: Serridentinus brownt Osborn, type mandible, maxilla, and tusk... ¢... 0.2.2 s/us10. sew ee nies Ge bie sielalpls i ioteets aieetatsraretsts 453 
aie “Tritophkodon hasnotensis sp. NOV:, type Molar. |). cca sic sise0s es oa! oi bbisPar etal aly ohe wilets Sie ev aMRaermEn le Pe alana a ee TNE PGI Soot ctaeg ta eee ener 454 
418. Serridentinus browni and S. metachinjiensis type molars compared.............0. cece c cence ce cette eee e teens eeeeseee 454 
BaO)s —Sermadentinus chinjiensis Osborn, ‘type MOlar, Po.%ic is 0%0:s es sess wie <ierarn\ahealuly ce peuatelete ePabata etand Pe ar Roiaie open elle Nines ee ae 456 
e20: Serridentinus prochinjiensis Osborn, type molars 2. iissc.i./c ee aia'e Seid sus oce alesa le et alah aPela PIS ose) ole ke sia eet ds oie ay eave 456 
Aon ~Serndentinusiannectens: Matsumoto, type figure (molars) :..;.\ <!em © cam eittmebls cheese lac any seis wheter eee 458 
gee.  Serridentinus lydekkert Schlosser, type figure (molar)..:.....). 22.0. «2s cep es elels eleldele ce wale oie nic nia aklee le ieee eee eee 458 
423. Amebelodont shovel-tuskers and Platybelodont flat-tuskers of Mongolia, North Caucasus, Nebraska, and Colorado. Res- 
torations by Flinsch... 25...) cac bic Pe eS PLT Se ee Re 460 
424. American (Andrews, 1922-1929) and Russian (Khomenko, 1912, Borissiak, 1929) explorations in eastern and western Asia 
(C0512) 0) ee OG ee on ene eins ONRr neh ACRES SrtA tee Oe inter ae Bid UP yt cnt oS 461 
425. Platybelodon danovi Borissiak, type figure (cranium, mandible, and molar)... ..fiinegee Aoapaice seine, 2a; Seaeteeenee enere 461 
eG, 2.F laity belodon danovt Borissiak, type mandibles. ishn@ csi Sass PS oes SO eee ee meee lg, Sea nlc tach e-aie are event fetene teres 462 
Ai.« Pladybelodon:danovt Borissiak, ‘type third inferior molar, .:.....5.ffam 22 se leteactneds soe tos eo us cde Gee eee eee de eee deoiaiate ete 462 
428. Platybelodon, four restorations of head, mouth-parts, ete. After Borissiak............0.00. 000. c ee cee cee eee eee eee 463 
429. Platybelodon grangeri Osborn, type mandible. After Osborn and Granger...............02.cceeee ccc eeee eee eeeneeees 464 
430. Dentinal rod-cones of Platybelodon and hollow incisors of Phiomia. After Osborn. ..... 2.0.2... 0.0.0 465 
Soe sections/of incisors of. Platybelodon and ‘Phromsiasic: 52 ese. a eel ches Slee Sree tS So nero 2) oie ic= nae Se eee 465 
432. Map of region around Iren Dabasu, Inner Mongolia, showing Tung Gur tableland...............0.0.000 0.2 e eee eee eee 466 
433. Deciduous succession in Platybelodon, as displayed in mandible and dentition of Platybelodon grangeri ref. After Osborn.... 466 
434. Platybelodon grangert ref., superior dentition. After Osborn........... ... ccc cece cee cece cee n ec eccnccnteeectecscccee 467 


435. Platybelodon grangeri ref., juvenile fragmentary cranium and nearly complete mandible. After Osborn............. ee 46% 
436. Platybelodon grangeri, referred cranium and jaws (female), also male superior tusk. After Osborn............ ote yee 467 
437. Platybelodon grangeri, referred adult mandible and dentition. After Osborn. .......... 2.26.20. sss eee eee eee eee eee 468 
438. Platybelodon grangeri ref., anterior portion of mandible, showing inferior aspect of lower incisors, placed beside coal shovel 468 
439. Platybelodon grangeri, referred third superior grinder.........-...0-. 650 c cee ee eee ete teen eens 470 
440. Torynobelodon barnumbrowni Barbour, type figure (mandible).........-..-.0:6. 060s cece eect eee eee ete eee eee eee es 470 
441. Platybelodon grangeri, referred second and third inferior grinders... ... 2.0... 0-00-00 eee eee eee ee eee es 471 
442. Torynobelodon barnumbrowni Barbour, type inferior molars. After peneil sketichibys Bar bouter rej cect ieee tte eteaterri nn mera pL 
443. Torynobelodon barnumbrowni Barbour, type figure (mandible).......-.. eee eee eee teen ees 471 
444. Platybelodon grangeri, referred mandible....... eee cts oa en an 288 eA ics Sd rt re eee coer 472 
445. Torynobelodon barnumbrowni Barbour, type mandible............. 6.0.55 e eee eee eet ee 472 
446. Serridentinus nebrascensis Osborn, type figure (molar)............... 02 c cece eee eee eee eee eee eee tee eens 473 
447. Rhynchotherium tlascale, male and female. Restoration by Flinsch............ 2... 5000s eee eee eet eee eee es 474 
448. Falconer’s genotype cast of Rhynchotherium [tlascalx| and Osborn’s type of R. browni sp. nov., both from Mexico (mandi- 

[S1US) Vit ee ee G8 Se erate Sei nr ee Pein eee a3 Oo cried an cee aaiona cits. oO OOD Oona to 012. Gocco." 474 
449. Rhynchotherium brevidens Cope, type and referred third superior molars......... 62-60-6020 020s eee eee eee eee eee es 476 
450. Rhynchorostrine types of lower jaw. After Osborn............... 202: eee cee eee eee ete eee ee eee eee eens 476 
451. Map showing geographic distribution of type and referred specimens of the Rhynchorostrine.......... Mrs occa stl! 
452. Rhynchotherium browni sp. nov., type mandible and dentition (formerly Osborn’s neotype of R. iWascaley, : RUN eel 
453. Type jaws of the iy ae honaahnices comparediwithimlophodoniamoinerzovdess\.na1t hel ee eae eee 484 
454. Rhynchotherium molars from Contra Costa County, California, and Tambla, Honduras..............-.-..-++++0-05005- 484 
455. Rhynchotherium spenceri Fourtau, type figure (mandible, second molar).............. 2600 e eee eee eee eee eee eee ee eee 485 
456. Rhynchotherium brevidens Cope, type figure (molar)............ 600-060 eee tne eee ee ee eee een ees 486 
457. Rhynchotherium brevidens Cope, new type figure..............6. 600 e eee een eee eee een e eee tees 486 
458. Rhynchotherium brevidens, referred molar from state of Washington... ... 6... 26.666 e eee eee eee eee eens 486 
459. Rhynchotherium shepardi Leidy, right and left superior type tusks reconstructed. .........6. 0.6600 e eee eee eee eee 487 
460. Rhynchotherium shepardi Leidy, original type figure (tusk fragment)......... 65.6606 e cee eee eee eee eee 487 
461. Rhynchotherium rectidens Osborn, type right and left upper tusks................-0 6060 e eect e eee ee ees 488 
462. Rhynchotherium [Blickotherium] euhypodon Cope, new type figure. ... 2... 6.6 ee eee eee 490 
463. Rhynchotherium [Blickotherium] euhypodon Cope, type figure (mandible)... 22... 6... eee eee eee es 491 
464. Rhynchotherium [Blickotherium] euhypodon Cope, type figure (palate with superior molars and incisors)................. 491 
465. Rhynchotherium anguirivale Osborn, type figure (molar).........--. 0.00 e eee ee ee eee teen eet eee 492 
466. Rhynchotherium anguirivale, type and referred grinding teeth.........-...... 660s eee ce ee ee ee eee eee 492 
467. Rhynchotherium browni sp. nov., type jaw and dentition................ 666s eee eee ee eee eee eee eee 494 
468. Rhynchotherium falconeri Open type jaw and dentition... ad savansen R405 
469. Rhynchotherium molar from Oak Springs, Contra Costa C aii @ “asa pefeme = ie sie to! Mistatond heer urus. .. 495 
470. Rhynchotherium shepardi edense Frick, type figures (portion of skull, posterior maxillaries, and grinding teeth)............ 497 
471. Cordillerion edensis Osborn, type figure (premaxilla and tusks). After Frick.................. 0222222 e eee eee eee eee 497 
472. Map showing position of San Timoteo and Eden deposits in relation to other Pliocene fossil mammal horizons. After Frick 498 
473. Rhynchotherium shepardi edense, referred superior and inferior grinders. .......- 222-60. c ee eee eet eee eee eee 499 
474. Rhynchotherium shepardi edense, referred immature jaw and maxilla, formerly type of R. paredensis Osborn................ 500 
475. Rhynchothervum francisi Hay, type figure (molar).............. 0.20. .ce eee e tee eee ee ete nee eee cena teens 502 
476. Rhynchotherium chinjiense Osborn, type figure (left ramus, third molar)... .......-.....eceeee ee ee eee ee 502 
477. Rhynchorostrines, comparative restorations by Flinsch................ 06. eee cece eee een ees 503 
478. Rhynchorostrine mandibles and tusks, North Africa to Honduras... ae 3 octane. 79004 
479. Blickotherium blicki Frick, genotype mandible and third molar, and Riaichiotheris mum ee Seas trewRne cranium and 

manoiole, Aer Lids (Grbainall he yyattes) poasnosododbo4ooeco ap adoomooonn ooo Jono ob abodes doco bpoohosanantc 506 
480. Aybelodon hondurensis Frick, genotype mandible and third inferior molar. After Frick (original drawings)................ 509 
481. Rhynchotherium types, as mounted in the American Museum.............. 06.660 s eee eee ee teen eee eens 511 
482. Rhynchorostrine, triphyletic evolution (mandibular series)... .... 2... 2.6... eee eee ee eens 512 
483. Rhynchorostrine, theoretic migration lines (map)........ 0.0.0... e eee cece ener eee tee te ene e ete c ee eeees 513 
484. Cordillerion andium, mastodont of the Cordilleras. Restoration by Flinsch... : sheer hy fap ees Lee! RO 
485. Cuvieronius superbus, giant mastodont of the Argentine plains. Restoration be Flinseh.. Weeks te See ey 
486. Mastodonte des Cordiliéres (‘Mastodon’ andiwm), Cuvier’s original type figure.............0 60.0 cence eee eee ee 517 
487. Mastodonte humboldien (‘Mastodon’ humboldtii), Cuvier’s original type figure... 1.2... 0c. ee eee eee ee 517 


488. ‘Mastodon’ humboldtii [= Cuvieronius (?)platensis| and Cordillerion andium, referred third superior molars. After Gervais. 
BRO CLATeNOniiLe miarensis AMmerhino,, type figure: (TUSK) is etice rates decrees «lech indsbataTevelaie/ Sinker eeatets euetae lake als lehelsialemtisliersit = Siete ial 
AO ECT er Onna rectus Amenhino; type figure (tusk) ics .csds sce % are cele ve. ow otis a anate vielerele Palate Re marae tedden form nro tele a Mt stalaioot ie ys a 
Bolen VOlomastoaon.argenunus Amerhino; type firure’ (GuUskk) iss cctia crn sic. 0 «1 sivis latetetel cudiatataltatin ed viele ie sole bo ieictar elev Gein ee l= 
BoD me CuMnerOnius superous Amechino, type figure (tusk)! oc azencvacrayeraterelele a:2 de Si--epapht tate oteyeNrdees aleve eet otevelen Wiens Wineieva lay 
493. Ameghino’s scheme of the phyletic relations of species of South American proboscideans.........-.-.. 66.00 0ee eee eee eee 
494. Tetrabelodon |Trilophodon| angustidens and Dibelodon (Cordillerion| andium, erania. After Lull.....................2--. 
495. Midcranial sections of ‘M.’ andium, ‘M.’ humboldtii, and E. indicus. After Boule..................00eseeeeeeec estes 
496. Cordillerion andium, single crests and trefoils and enamel foldings of molars. After Boule..................+....-.0005 
Oem GC uperonius Numovoliii, cranium in Paris Museum, After Boulos csccee. ssccmnei eso aesis wee arises isle talate ins ee le 
498. Restorations of Notorostrines and Humboldtines, as known up to 1933, by Flinsch.................0 00 eee eee eee eee eee 
ape bviogenyor the, Mastodontids:: After Bouleie-s:éie:erc bacseres vlecectaeenaid etnias oivd ol atoltcels cleat th oie OREO RESE ele 
uO mC onarlertonecaensis: Brick: «Restoration: by.-Wlinsebs 2 ces io ollie) stacererayat si clovaiaiain sy) ol teres vais talacala mes a anal ees eee ee 
501. Map showing paleogeographic range of Proboscidea in western United States, Central and South America................. 
502. Cordillerion andium in comparison with Cuvieronius superbus. Restoration by Flinsch................00 600 c cece eee 
503. Cordillerion andium, the Andean mastodont of the Cordilleras. Restoration by Flinsch...................00 0000 eee eee 
Mie aC ordinenton ang Oumeronwusigrinding teeth. After Gervais: . cies ¢ «lacs osiiwirse eve cielo ls <televesole\ siatetstal ste) =tets[ott ts). leis staal fetvinate ye 
nm eCoratuenzon andvum, referred: cranium, iafter: BUrMeISbeDs sss siels stsialelev ive dix oe Sake «ie. s Blatssellyel alee sles sis toes eles Wala neyaeiee 
506. Cordillerion andium, referred cranium in Field Museum of Natural History. After pen sketch by Knight................ 
507. Cordillerion andium, Cuvieronius humboldtii, and Ewbelodon morrilli crania............6000 eect eee eens 
508. Mastodonte des Cordiliéres (Cordillerion andium), Cuvier’s original type figure.... 22.0.6... eee eee eee 
SOME OTAteonmanavum, referred jaw; after D?Orbignys i ha. 3 okenaiais sles «ole bboas~ viel telalede «hove :>,eisTaisl id= elara ete ioe ei eimai 
510. Notiomastodon argentinus Ameghino, type figure (tusk), first described as ‘Mastodon’ argentinus..........0000 00 eevee cee 
511. Cordillerion(?) bolivianus, referred molar [genotype of Teleobunomastodon Revilliod].. .... eee eee 
Pio sOorasulerton bolinanus, referred mandible. After: Pompeck]:<....<seseece v cl 2% 0+ 2lely oie sain iam 2 vim see alesis) dela lelate eee eet 
513. Cordillerion tropicus Cope, type figure (right ramus of jaw). After von Meyer’s ‘Mastodon hwmboldti’ ref.................. 
pia, ‘Cordillerion oligobunis Cope, type figure-(mandible).. After Villada.... 2.02.2. sic cc ce 2 oes cesses wieiein ve ows oisinle ate sig gale 
515. Cordillerion oligobunis antiquissimus Freudenberg, type figure (molar). After original photograph........................ 
516. Cordillerion(?) oligobunis felicis Freudenberg, type figure (superior molar)............... 002000 e cee eee eee cette eens 
517. Cordillerion(?) oligobunis felicis, referred inferior molar, after photograph...............06 0: ce eee c erence eee eee eneee 
518. Cordillerion(?) oligobunis intermedius Freudenberg, type figure (molar), after photograph............................000. 
519. Cordillerion oligobunis progressus Freudenberg, type figure (molar), after photograph................6. 0.000 cess eee eeee 
620. Cordillerion gratum Hay, type figure (maxilla, second and third molars).........-.-.-020.++e-+eeees ee ons se reen se sseuee 
521. Map showing position of San Timoteo and Eden deposits in relation to other Pliocene fossil mammal horizons. 
APH er Id sha. ace cee bs ooo o's 0:ce Sunt ld ata bbls ayeepereare ral alain a A is hb Uosleah evga) a leLel le ger a eM U RSE IR iP gn 
522. Cordillerion edensis Osborn, type figure (premaxille and tusks). After Frick...............2..00sescscseececcsereccees 
523. Cordillerion edensis type rostrum and superior tusks compared with those of C. andium. After Osborn................... 
Bei. Cordillercon.orarius Hay; type figure (molars) st. «is sisei:sss,cieie.s: seis bio! esore inn oetaea le alevelel wile) Aietetol staaveleletete te fe gelet= etelet fates arate 
naps (Cordillerion defloccatus Hay, type figure s(mandible),.:. «so... 6.0: .1aje.. isis svehelersctal biele oleie eystopnveletetolainiets arate che relates etal eee 
526. Cordillerion bensonensis Gidley, type figure (fragmentary cranium). ...... 06.6. .626s. cece ce eee ee epee ees vere wees sewers 
527. Localities of mastodont discoveries in Ecuador, Canyon of Chalang, near Punin, Province of Chimborazo................. 
528. Cordillerion andium, referred cranium from Tarija, in Field Museum of Natural History.....................00000005. 
529. Mastodonte des Cordiliéres (‘Mastodon’ andium), Cuvier’s original type figure... é J. ccyenereed 
530. Cuvieronius postremus Spillmann (‘Mastodon of Alangasi’), type cranium, pies resuebeh sae 13) cies ee 
531. Cuvieronius grinding teeth and tusks, type and referred. After Caretite............... 0.0. c cece eee e eee eee e eee eeee 
532. Cordillerion and Cuvieronius crania and jaws. After Nordenskiéld and Carette, also Osborn original.................... 
beei \Cuveronzus postremus Spillmann, skeletons. «2... civ. « avleil rs 3 «ete dooisssiaiesew wo myphe serie) em selbtesielele = oisiphe ual ots tal hale oo 
nes. Cuvieronsus ayore, referred first: left: superior /Da0larsa:s < as 1< 5-6 Js. sts/ais) =fatelolas'levotalale ds sistelate stars falols tele manele inls/alselctsteberatene eas 
535. Cuvieronius ayore, referred third right inferior molar, also mandible, Frick Collection...................22000 eee eee eee 
536. Cuvieronius postremus, supposed flint spear holes in occiput. After Spillmann...................6 005. c cece eee es 
537. Mastodonte humboldien (‘ Mastodon’ [Cuvieronius] humboldtii), Cuvier’s original type figure................00...0 00 eee uee 
588. Mastodon [Cuvieronius] humboldtii, Osborn’s paratype, after Cuvier............. 0... cece c eee e eect eee e eee eeeeeeenes 
539. Cuvieronius humboldtii, referred third superior molar. After Lydekker.................. 0. cece cece eee erent eee eeeeees 
540. Cuvieronius and Cordillerion, third right superior molars. After Gervais............. 00. . cece eee cece eee cece ees 
541. Cuvieronius superbus, mandible referred by Burmeister to Mastodon humboldtit................. ooh, 3 aR ee 



542. Cuvieronius bonaerensis (superbus) Moreno, typecranium. After Cabrera................. 00. - eee eee cee eee teens 579 
Ass Cumeronvus) platensis, Amerhino, type figures (GUSK))io. eit ee eer ieee maitre erent ae cto ete eee 579 
FAAS. Oumeronuis supenbus Ameghino; type: figure’ (tusk:)).jo. acy. rite a aletrusne eieeneteie rs enebere tte teste een chet rtf ya) etc le ete tate Sweet 580 
GM, (Cl ARI OHMS Adena NLP AYo NADER (QHVISN)o oo go acngauimaduccmocuaosnconecmorecHvapappanondaanncdsbiosanaamoe 580 
BAG, Cuvieronius rectus, primitive referred molar. After Ameghino............-..--5.25:02 +50 esse ess setae e eee seen: 581 
547. Cuvieronius chilensis Philippi and Cordillerion bolivianus Philippi, type figures (mandibles), in comparison with Cordillerion 

CCT ee ee aE PEO Pa BO: uc nbs 9.0) fy, AP PCC es GWU. G cin hci dO RATA Hd ow Si lon pay cya ceo che. < 582 
548. Cuvieronius ayore Spiller UMA geruleah hee, (CHPMIDHAN)) Evie OW OAT ON, oo oooapauhosdopenenanamsncnsaddancaucagdcge oc 583 
5AQ) (Cumeronis ayore) and Cordillerconandium,) cramiallySCtlOMs =) is eile = ete oe eee eevee aie ees 583 
550. Cuvieronius postremus Spillmann, type raanciiles Afterjphotograpphitrs, 2.2% S205 Ae. Salo Sk MISS ete ernie roe eo eee 584 
551. Cuneronius postremus Spillmann, type cranium. After photograph. .................... 0.222 eee ee eee eect eee 585 
BEOwOumenonvus) postremus, reterred, molars: Atitersphotopraplin yyy sels otters siete oye) sp eeteeet enero tvnete tenceett rater ete onete tetera 585 
553. Cuvieronius and Stegomastodon, parallelisms and divergences in inferior molars and mandibles........................ 586 
554. Cuvieronius platensis, referred second and third inferior molars.. .......... fh Se a a ne 
555. Cordillerion andium and Notiomastodon argentinus, second cervical ventabras Geese Utter Gabreras ee. Ae ee 587 
556. Cuvieronius superbus referred, tip of tusk artificially carved to convert it into a utensil. After Castellanos............... 589 
557. Notiomastodon ornatus Cabrera, type and referred tusks, type jaw, also type tusk of NV. argentinus. After Cabrera....... 591 
558. Cuvieronius platensis Ameghino and C. rectus Ameghino, type and referred superior tusks. After Cabrera................. 592 
559. Cuvieronius humboldtii Cuvier, paratype third inferior molar, after Cuvier, and C. platensis, referred third superior molar, 

EET Trey Ck 01110) 12): ae a ee a att re re or on 85.0 0 Ost, dinio crt.cht.e.o.c,c 592 
560. Cuvieronius platensis, referred third superior and inferior molars. After Cabrera................... 000 e cece eee eee eee 594 
561. Cuvieronius superbus, referred third superior and inferior molars. After Cabrera...............002202 2 cece eects eee eens 594 
562. Cuvieronius bonaerensis Moreno, type cranium (ef. C. swperbus Ameghino), after Cabrera, also crania and mandibles referred 

bya CalbreraitonStegomastodons | —Cruscenontis|\iphateresissrrarrstcraye)-yeretie aerators 2 < -¥ar-tcbel tells orcas tenia <P aan eevee erent ee 596 
563. Cuvieronius superbus type tusk and mandible; also tusks, cranium, and mandible referred by Cabrera to Stegomastodon 

(@coslisinenbusybyaOsborn: (imjpart) sol = platensismmmrey srr rete reire oicia yen yar= aia rere etre rer tee Fence ee 597 
564. Skeletal remains of Cuvieronius platensis ref. and of C. bonaerensis (superbus) Moreno, type. After Cabrera................ 598 
565. Cuvieronius superbus, referred skeletal parts. After Cabrera........:20.000. 0200 0eecece ces e serene se sess seer eee OOo 
566) Cumeronius superbus and C.platensis. Restorations’ by Elinsch..-..- 22.2... 02. c0ce ees ee ee oe ace re ale alele elelael 599 
567. Eubelodon morrilli Barbour, type skull and jaws (after reconstruction) also referred vertebral series. After Barbour...... 600 
568. Devil’s Gulch section, Brown County, Nebraska, where types of Trilophodon willistoni and Eubelodon morrilli were found.. 601 
569. Hubelodon morrilli Barbour, type jaws and palate before reconstruction. After photograph...................2....005 603 
570. Hubelodon morrilli Barbour, type jaw. After sketch by Barbour:.........0......00000 cece ce sce e cee cece erences eneeen 603 
571. Eubelodon morrilli Barbour, crown pattern of third superior and inferior type molars. After pencil sketch by Osborn..... 603 
572. Eubelodon morrilli, type pelvis, also referred limb skeleton. After photographs.................. 0.0 eee cece eee 604 
573. Eubelodon morrilli skeleton as mounted in the Nebraska State Museum, 1931. Background by Elizabeth Dolan.......... . 605 
574. Eubelodon morrilli, referred cranium and superior dentition, found by Frick expedition. After Frick (original drawings).... 606 
575. Hubelodon morn, type and referred molars and referred tusk... ..........0.00 cee desc weer ets ese e eect eesee reeves ae 607 
576. Hubelodon morrilli and Cuvieronius bonaerensis (superbus) crania and jaws compared.............-..2..0 eee eee eee e eee 608 
577. Eubelodon morrilli, the Humboldtine mastodont of western Nebraska. Restoration by Flinsch........................... 610 
578. Humboldtine: Proversion of inferior ridge-crests in Stegomastodon...........6 000s cece eet ee eee eens 612 
boi Humboldtiinse:) broversionvof inferior ridte-crests im! Crienonvuss. oo oc cee nk ec omens ee ote eee fe elated eislslslc sielelere 612 
580. Humboldtine: Retroversion and centroversion of superior ridge-crests in Cuvieronius and Stegomastodon...........++-+++ 613 
581. Brevirostrine: Proversion of ridge-crests in Anancus and Synconolophus.......... 0060 eee ete eee eee 613 
581a. Notiomastodon ornatus, type first and second inferior molars. New figure................ 00000 e eee ee cee eee teens 614 
582. Anancus arvernensis, the straight-tusked mastodont of Auvergne, southern France, and of northern Italy. Restoration by 

AULA Cy ere tere brtoiettc oes ce nas Sees Senha wats a a ct Bke Syn caveat eS aa. e oud. ctrasnd tied CROPS UB ASUS Geewadascd eh erehe Me Soec ome aaeuete fetes 616 
583. Synconolophus dhokpathanensis, Middle Pliocene mastodont of Dhok Pathan, northern India. Restoration by Flinsch... .. 616 
584. Anancus arvernensis, mounted skeleton in Turin Museum. After Sismonda............ 2.0.00. 0c cece eee eee eee ee 618 
585. Anancus arvernensis, mounted skeleton in Bologna Museum. After photographic reproduction of figure in Capellini’s 

16(0) aXoys7:4 0) (C0) 1 UO 10). Beri Miran ein Pera teteiseetcNts nc -0.clct 4 Solo. ck Sa a Ek a ee Re Pec ah oe ec Rome Geet Ohad co 619 
ps0, William Smiths) Whithneham Tooth’) |[=Anancusfalconern@sboml:ae.- 2 a. seen e cere ieee oe aerosol) el eearabsiet iets 620 
587. Anancus falconeri Osborn, fluted deciduous superior premolar from Norwich Crag. After Lydekker..................-.... 620 
588. Primitive Anancus molars of Europe and of India. After Schlesinger.......... BE a ra ree aon occ obgi eir4! 
589. Anancus perimensis from Perim Island, India, referred second and third ‘ateriGr caches aihtae Falconer and Cautley...... 621 
590. Map showing geographic distribution of eighteen of the types of Brevirostrine of Eurasia and of seven of the types of 

Humboldtine of North America,alsoreferredispecimensemie ste s s nie cee cen sitet aces yal ciaereternninaial 624 


591. Crania of Brevirostrine from Italy and India, and Stegomastodon (Humboldtine) from the United States (Texas)....... 
592. Proversion of inferior ridge-crests in right third inferior molar of Anancus arvernensis brevirostris. .. 2... 0. oo ee ee 
593. Strong proversion of ridge-crests in type third right inferior molar of Synconolophus corrugatus...........00.000-02202 0 
594. Synconolophus dhokpathanensis, restoration of head by Flinsch (right) drawn directly from type cranium and tusks (left) . 
595. Anancus arvernensis and Synconolophus dhokpathanensis, restorations by Flinsch.................. 0000 eee rere eee eee 
596. Anancus arvernensis Croizet and Jobert, cotype figures (mandible, deciduous molars)............6.. 0000 c eee eee eee eee ee 
597. Anancus arvernensis brevirostris, two cotype inferior molars. After Gervais...............02ce eee cece een cee ee tone ne 
598. Anancus falconeri Osborn, type figure (molar), also referred molar of same species. After Faleoner...............+.-+-. 
599. Anancus falconert Osborn, type figure (third inferior molar). After Faleoner..............0...00 200 cece eee ee eee eens 
600. Anancus arvernensis(?), referred mandible, after Sismonda, described by him as ‘ Mastodon’ angustidens.........-.....+++++ 
601. Anancus arvernensis, referred grinding teeth, from Hungary. After Schlesinger............... 0000000 c eee eee eee eee 
602. Anancus arvernensis, referred right inferior molar, from Hungary. After Schlesinger....................0 2.002205 seeeee 
Bias Anancus intermedius Hichwald, paratype jaw. After Michwald...<../0) ci... <1 0025 © 0% 21 © ee wines ole olele siesta iatele siniels ate 
604. Anancus arvernensis progressor Khomenko, type figure (mandible, molars) ...............00 0. cee ce teen eee 
605. Anancus perimensis, paratype and referred skulls, after Falconer and Cautley...............0..0 000 c eee eee eee cee eee 
606. Anancus perimensis, type and referred specimens from Perim Island. After Faleoner and Cautley.............-.....--- 
607. Map showing distribution of Anancus in India: Perim Island, the Punjab, and sub-Himalayan Siwalike.j:7 2.4.08 hceee 
608. Map of Middle and Upper Siwaliks of the Salt Range of the Punjab, India.. 
609. Anancus properimensis sp. nov. and A. perimensis ref., showing evolution of left third ene sales fee dbaonisan of 
REDOOMERL Of SECIMENTAULONL . ..5aid 0 ciccre e creteicne Sys os ahd welt al Aisa hs, cuntele isle lene Slr ot ose, ove PONDS P nS a ety Fopmibee ete eke = Seana tere 
610. Anancus perimensis, referred symphysis. After Lydekker: 547914 0-o eek tm eae ee Eee Se ene eee 
611. Anancus perimensis, referred first superior molar. After Falconer............. 0.0.0 20sec cece eee eee eee eee eens 
612. Anancus perimensis, paratype skull. After Falconer and Cautley.................. cece cece sees ce erence tees neceeeeee 
EI SAyicle NrOPErIMENsis SP. NOV., tVPe MOlAT.. f-....0.0.c00.00 oo deda nahin ddees Weve vey e> Re bere ee oe raee Eee 
614. Pentalophodon falconeri sp. nov., type cranium with superior grinding teeth. After Falconer and Cautley. Third left superior 
molars new. diagrammatic A ULC 36s sie ecrsoreica ensayo, a0, <Stayss » Lares fever Srones SiS ebve aay TmIe) alehe EST Soh ele he dev eRe eat eee ae ete Ree ee 
615. Pentalophodon sivalensis Cautley, type and referred, also Osborn’s ideotype. Synconodont or compacted and proverted in- 
PEELOP TIOLATS!. « sle.cisce oct c cccals oc evsvnels cists wie erace ep a one ere ehareva ea gibrarenere Soc Oe chen mceveraiee arate Mea espe cietstenete tee fee) settee ese 
iinemerenaopnodon svalensis Cautley, type figure: (Molar), tis./. <tsc-e n> vie sts elele ale ols/atelova iol lebaloisley sine re ttete nia ale teleost 
617. Anancus perimensis, referred third left superior molar compared with corresponding molar of A. falconeri. After Falconer 
Piva CH CMT LO Wis ga; atc g'e’inip ayaiace tas cies’ Shoynin obo Bnepebeuece uy Cue wr stews’ e! «cgQIS calscl punve ote: e gche, auetenascerel ee ate eet dette et eee eae 
618. Pentalophodon sivalensis, Osborn’s ideotype molar. After Faleoner and Cautley.................. 0000 eee eee teens 
619. Pentalophodon falconeri, referred cranium in profile. After Gaudry.............. 02.2 cee cee eee eee tee eee eee ete neeee 
620. Pentalophodon sivalensis or P. falconeri, referred molar. After Lydekker.................000 ce cee cee cee eet ence eee aees 
621. Triple phyla of the Eurasiatic Brevirostrines. Restorations by Flinsch................ 2. eee e eee cece cece tee eeeee 
622. Synconolophus, comparison of type and referred specimens (Synconolophus hasnoti, S. corrugatus, types, and S. dhok- 
pathanensis ref.).... ww mus) al chi 9 Shan Salven ergs so ewe aba wipe bog BWR Saha ras SREREN  PaNON ee NSU OR tamer ee 
623. Synconolophus ptychodus and S S. dhokpathanensis, superior dentition. .— 2. .cc2 tere sites etter = eicde atte eee ee 
Gum synconolophus piychodus Osborn, type: MOlar.... ss/cs...5 n.<:+ 2 = ats wie e+ + «letalalevshoteo ebteiayete ebabepevers eels ete SUSIbreN a etetaie) eer nets nae nat 
PEST NCONOLODhus piychodus, referred, MOAT sac 0iz0's corse sr ole\s Rlaiei'vanovoitd ap tahel lu esd foie Wele aheke vlces lay ota otetet okt oi Fede ats tinte saat ere area 
626. Synconolophus corrugatus Pilgrim, type figures (molar and symphysis). After Lydekker....................0022-00eeeees 
Hei Synconolophus hasnot: Pilgrim, type figure (molar). After Liydekkers: 7.54005 ele =~ lien oe l= viele oie ois e o ees dale ta eletalers lols 
628. Synconolophus hasnoti, referred juvenile cranium in three aspects. Brown Collection............ 0.00.00 eee eee eee ee eee 
BuO Synconolophnys ahokpaihanensis Osborn; type: Cranium: cyte cersisaleiere « 6s <b etm eiwin este eee pers eta sto vat intel oveieus tee inta ts et eatin te 
630. Synconolophus dhokpathanensis Osborn, type cranium and superior dentition................ 0062 eee eee cee 
ne unconolophus ahokpatnanensts Osborn, paratype MOAT... 5. wele sees ns stro dete mimpeiel o) sia eae wicker etetafer tee) sie teamn eee 
632. Synconolophus dhokpathanensis, referred superior and inferior molars. ............0.. 000 ee ee eee teens 
Popresynconclophus propathanensts, type mandible and dentition. once oc «/ocle/arele etgiecia sieveieisdel o)-iveler= ele te eva hey use nay ota abs 
634. Stegomastodon arizone Gidley, mounted skeleton of type in National Museum. Photographic reproduction................ : 
635. Stegomastodon arizonex, the curved-tusked Stegomastodont of Arizona. Restoration by Flinsch..............-........+55 
636. Stegomastodon mandibular abbreviation. Ridge-crest internal proversion............600. 000 ee eee 
SE MEESICQOMASLOAOTICHUDMANE LLAYS, type: Lipure (MOLAT)s. osc: nierare stelle sje snvn ete, Sus) EIT Mtonter ave) ots niente o/s © opiate clone rats inde =yerient earners 
638. Stegomastodon mirificus Leidy, type figure (left ramus of jaw)............cc00000 ccc e cee tenet eee e eee setsenssereereees 
aU MmELavomnstonon reeanius OSbOrn, ty Pe SKULL. 35 ..j.insissiccecstaen iavenenstne citoe ele yny eet ebstcumbetereb ate bie sasha tna Statahona: s wthe Or ancte tabale oh neat Fepetetan she 
640. Stegomastodon arizone Gidley, mounted skeleton of type. Restoration and sketch by Knight.................---0+-e0es 
641. Stegomastodon successor Cope, type figure (mandible, molar) 


642. Stegomastodon texanus Osborn, type skull and dentition; also S. mirificus, S. arizonx, S. texanus, and S. aftoniz# restorations 
Dy; aCe ane eee Ann onan icc occudhiwes dco mene sania Innb aeons socnij loo 
643. Stegomastodon texanus Osborn, paratype mandible and dentition compared with S. successor type......... 6.022 eee eee 
644. Stegomastodon, ontogenetic and progressive jaw characters and inferior grinding teeth................66 eee eee eee ee cea ee 
645. Stegomastodon texanus, type and referred superior grinding teeth. .............. 20... c eee eee eee cece eet eens 
646. Stegomastodon arizonzx Gidley, type figure (mandible). After photograph............... 06.0 cece eee eee cece e eee eens 
647. Stegomastodon arizonx Gidley, type figure (cranium). After photographs...............-.. 0.000 e see e eee eee eee eee eens 
648. Physiographic and faunal characters of San Pedro valley of southern Arizona in uppermost Pliocene time. After mural by 
Knight ancAmerican: Mouseumassiie'sccccatcenve sees sons crete torensratete lcneiattenetersls cleats acvirstaviunla for evexeraleienctanoustsRa re letcve aan aoe onr ta aie 
G40 Sienomasindonarizone retermedibbin clin glitter] OTM 0 Leailvarrtyetceraeywieey. fare) oteLeTo)stateiaetetey tote ta ets sare delete ete eae ene 
650. Stegomastodon aftonize Osborn, type figure (superior molars). After original photographs by Samuel Calvin.............. 
651. Stegomastodon priestleyi Hay and Cook, type figure (inferior molar).................. 00. e cece cece e en eeeeees 
652. Molar diagrams showing typical crown pattern of third inferior molar of each of the four families of the Mastodontoidea. . . . . 
653. Miomastodon depereti sp. nov., type molar, compared with type molar of M. merriami.... 0... 0. eee 
Qk, Mikasa lage MEMO; WON an UNA Nenana obKS ((SuhoyeraorewMYEWIS)). 5 oan no ckonnecoboUAoeeedenooonnnepons wubb oasduusoenedonoce 
Gos Mastodon qrangenm banbour bye miguure) (Skul lac etiisls) Peretti ete eee rene olen ts oes tees erence eed eee 
656. Mastodon acutidens'sp. nov., type figure (juvenile skull'and jaws)..................2 22 cc eee eee e eee eee eee eee eee 
657. Turicius turicensis, referred inferior jaws with incisors, also superior second and third molars and superior tusks, from 
[Ojo ef e) oC hy UO Eh an Reg ee ent ae Ree cach hoi ea a ere SORA E OR aot ccna nae ab 6-0 60's ¢ 
658. ‘Mastodon americanus” of Hopwood [= Turicius sp.], from China (left ramus).............-.00 20sec tee tee e eee eee 
659. Zygolophodon borsont, referred third left inferior molar, from China. ........ 0.26.16 eee eects 
660. Stegolophodon lydekkeri sp. nov., type figure (superior molar), after Lydekker.................... 200.0 sce e ee eee eee eee 
661. Structural evolution of the cones, conelets, and ridge-crests in the Stegolophodon phylum, in comparison with Palzomastodon . 
6625 Uimlophodonconnemus Hopwood, type figure (inferior molars)ey sacs see ©. te eie tee cle eee) siete cote estonia teeter 
6635 linlopnodomsnectabilis LLop woody pea vire:(mfeTiON MOLAD) kyr rsis lee =) are) ttre le sleyelelcteteye) ete) laborers ett fe st t=(ee tte =)-teyete tate 
G64eeleinalophodontenoletus Hopwoods type naurel(infertor molar) ce ttre. oe aie ses eee ene eects eee a 
Gb5aee icoavelodonllareterred man diblemn,ecrm <nra-cctnisr tarts Atala ere Aate tua stots): = .bo ah acta uae cpclieverne teva) ai eeletsne keel oie arene tere aiee 
GHG, dlequindiicton ad, MOO URS Cae FOS Hs oan eoeerA Ras 40 ad 8 00000 s ADRDRNRAMRAN So mons apne whoo cdomdo ord patcg ene ot 
667. Megabelodon lulli, referred skeleton mounted in Nebraska State Museum........... 2... 0.0220 
668. Chart showing type localities of various members of the Mastodontoidea, including Gnathabelodon thorpez of northwestern 
TISSUES cs orgs be Al so SRE ea Re eC ee NR er rat aan oe a re ae rere Cer AON R52 
669. Gnathabelodon thorpei Barbour and Sternberg, type figure (mandible, dentition, superior tusk). ...................2.-2... 
670. Migration routes of the three phyla of the Brevirostrine of the family Bunomastodontide from a hypothetie African ances- 
CraloneintOmULOpeandihence tollinGiais.. 1a. eeeeNe ea eter. ol lave alec ae ee es ests aye oeictals Siecle ee eee 
671. Anancus sinensis (Pentalophodon sinensis of Hopwood), type figure (superior molars)... ..........002.00 000 e eee eee 
672. Cuvieronius platensis and C. swperbus, referred jaws and teeth, in the Buenos Aires Museum. ............... 002220000005. 
673. Correlation chart of the Pleistocene of Nebraska, Iowa, and Europe, including type horizon of Stegomastodon primitivus sp. 
TOW s oo 9 ob0-0. 5b O14. tt tae HOES OId rt Ce ET UNE HETE SSCIE a5 cect. cr thee ae eee ey le a ic 0S 
CUAemSLEOONLOSLOROTE DTAITALLUUSISP OV. ty Pe)pAlaue ANG WUSKS =k cietet teste. cen ci bes Ceres Sate eon eos at ay ale w stele efeleichel clpiei= sei aisiie eae 
675. Type palate and paratype third left inferior molar of Stegomastodon primitivus sp. nov., compared with type third left inferior 
TNO ATOMS MTIUUEUTLGLSMOCL Cyan ERIN eee eae ain roe canes os ote risa eee tdels oh Jee letticbenstarerste eae eee 
676. Diagram of Stegomastodon primitivus quarry, northeast of Ainsworth, Nebraska. ............0000 2000 c ee eee eee ee 
677. Serridentinus wimani (Trilophodon wimani of Hopwood), type figure (superior molars)... ........0...002. 000 cece eee ee 
678. Ocalientinus emmonsi (Gomphotheriwm? emmonsi of Hay), type figure (superior molar)... .............000.0 0200 e ee eee 
679. Duplication of columnar section of the Mio-Pliocene Devil’s Gulch and Valentine region of Nebraska, with important cor- 
rections (see Fig. 272, Chap. VIII)...... RENEE Bete Pp sexta or nc ECCLES CRP SEA ROR Ne hh Siar ha Pats wld ot iced ic 
680. Geographic distribution of three of the great primary stocks of the Proboscidea (Mceritherioidea, Deinotherioidea, Masto- 

CTAOGLEN) 4 Bb da hascethn ky dc MRO cok eke nc ge E CSG tae Ro pt ee A tr RO Rr EE a ORY er er I Ar 



1. Principles of revision of generic and specific names. 2. Revision and adoption of generic names of Proboscidea. 
Certain of the names applied to the mammoth and Examples of names of prior generic conception and 
mastodon. definition, with classic orthography. As adopted 
Linnean special creation system (1735-1766). by Osborn up to the year 1923. 
Nomenelatural principles adopted in the present Indeterminate names, collective or section names. 
Memoir. Summary of total generic names adopted up to the 
Principle of soundly established usage and of common end of the year 1933. 

3. Phylogenetic nomenclature and classification adopted in 

Post-1836-selection of the first soundly established gen- the present Memoir. 

eric or specific name. 
Determination of the characters, validity, and names of 4. Acknowledgments of codperation and assistance. 
specific types. 

The proboscideans and the horses divide the honors of age-long association with man, not only in the historic 
period but far back into prehistoric times. Whereas the horses entered Eurasia late in Pliocene time and were 
hunted for food during the closing 100,000-year period, the association of man and the proboscideans appears to 
have been far more remote, certainly extending back into Upper Pliocene time where we discover more or less 
conclusive evidence that man hunted the primitive elephants of the period. During the severe environment of the 
late Paleeolithic age, man had become an extremely clever horse hunter, as shown at Solutré in southern France, 
as well as an extremely clever mammoth hunter, as shown in the ‘‘woolly mammoth pit” of Moravia. Whereas 
many horses were necessary to feed a tribe, a single elephant or mammoth would furnish an entire feast, aside 
from the priceless materials of bone and ivory to be fashioned for economic and artistic purposes. 


From the dawn period of Palzolithic art the mammoth has been looked upon with wonder and curiosity, 
while the horse has compelled admiration by its beauty and adaptation. This introduces us to a further contrast 
in the fabulous, mythical, and scientific history of both these remarkable mammals, namely, that the fossil 
proboscideans of Quaternary age were the first objects in all parts of the world to arouse scientific curiosity, specu- 
lation, and finally research. In the Eastern world there sprang up around the mammoth a distinct ‘earth-burrow- 
ing’ myth; in the Western world there was the natural Biblical interpretation that the fossil elephants were a 
product of the Flood. 

But the present volume is devoted entirely to the succession of discoveries in various parts of the world, in 
fact in all the continents except Australia. These discoveries through the dawn period of scientific research and 
interpretation have been steadily accumulating. They began in 1695 with the first independent observations on 
the ‘ancient elephant’ by Tentzelius (published in 1698). They culminate in the summary (Chap. X XI) of the 
second volume, in which the full tide of experience and of knowledge brings us to the close of the most complete 
and remarkable evolutionary history of mammalian descent thus far revealed. 

Thus, with only occasional allusion to fabulous and mythologic interpretation which in itself would fill 
volumes teeming with interest, we set ourselves to the task of progressive scientific interpretation: 

First, a period of exploration, discovery, and description which virtually began in 1695 when Tentzelius 
victoriously argued with the doctors of Gotha that he had actually found a fossil or extinct elephant rather than 
the remains of a recent elephant as his colleagues hotly contended; continuing (1) with the special creational 



influence of Linneus, Buffon, Blumenbach, and Cuvier, and extending through (2) the intermediate period cul- 
minating in the speculations of Hugh Falconer, a contemporary of Darwin; thence (3) into the post-Darwin 
evolution period, through the bypaths and mazes of erroneous speculation on proboscidean descent, and finally 
(4) into the illumination of recent time resulting from twenty-seven years of continuous research by the present 
author in the entire field of proboscidean exploration and discovery. 

In brief, the history of the discovery and interpretation of the fossil proboscideans—mastodonts as well as 
elephants—is coincident and coextensive with the dawn, rise, and development of the science of Vertebrate Pale- 
ontology, coupled with our slowly deepening and broadening penetration into the mysteries, principles, and laws 
of evolution. First, there had to be waged a century-long battle to prove that these more or less petrified speci- 
mens were actually fossils belonging to the still unknown world of extinct life; second, another century passed of 
more or less accurate scientific description and nomenclature, and, finally, a third century of evolutionary inter- 
pretation and adaptation of the nomenclature to evolutionary knowledge. 

In the various chapters of the present Memoir the history of every great proboscidean type is separately 
traced from the beginning to the very end of its scientific interpretation, with the fullest justice to the actual facts 
observed by the authors of each of the four great periods of proboscidean thought. So far as practicable the obser- 
vations are recorded literally in the language of each author, both as a matter of justice and as a practical matter 
of the dissemination of the widely scattered and at present inaccessible literature in many different languages. 
Also, so far as possible, errors of interpretation are omitted, otherwise this volume would embrace a “comedy of 
errors” and one would lose sight of the main lines of actual fact. 

Immense pains have been taken to reproduce every original description in the language of the author, to 
reproduce every original type figure in facsimile, to retain every original generic name to which the author appears 
to be entitled by originality of observation, as fully explained below in the section on nomenclature of the present 
chapter, and finally to give the fullest credit for priority and originality of thought to the great pioneers in prophecy 
and research, among which are the outstanding names of Tentzelius, Blumenbach, Cuvier, and Falconer. 

As to proboscidean bibliography the series of papers bearing on the subject of this Memoir is arranged alpha- 
betically by authors at the close of the present Volume. A clearer idea, however, of the historic development of 
discovery is shown in the following chronologic arrangement of the writers, greatly renowned, less renowned, or 
obscure, who have enlightened us on this great subject, from Tentzelius (1698) to the present time. 


1. 1698-1836. TrnrzeLius, Burron, Linn=us, BLuMENBAcH, CuvIER PERIOD 

1602 Avalo y Figueroa, Diego de 1767 Baldassari, Giuseppe 1792 Gmelin, Johann Fridrich 

1692 Witsen, Nicolaus 1768 Collinson, Peter 1792 Kerr, Robert 

1693 Ray, John 1769 Hunter, William 1795-1829 Geoffroy Saint-Hilaire, Etienne 

1696 Ludolf, Hiob 1771-1793 Pennant, Thomas 1795-1836 Cuvier, Georges Léopold Chré- 

1698 Tentzelius, Wilhelmus Ernestus 1773 Rozier, Frangois tien Frédéric Dagobert 

1706 Ides, Eberhard Ysbrant 1777 [1780] Pallas, Peter Simon 1799 Corse, John 

1715 Réaumur, René Antoine 1778 Buffon, Georges Louis 1802 Lacépéde, Bernard Germain 
Ferchault de Leclere de Etienne de la Ville 

1729 Sloane, Hans 1779-1810 Blumenbach, Johann Friedrich 1803 Artaud, Soulange 

1734 Beyschlag, Joannes Fridericus 1782-1786 Merck, Johann Heinrich 1807 Adams, Michael 

1734 Seba, Albert 1783 De Hay 1807 Link, Heinrich Friedrich 

1735-1768 Linneus, Carolus 1785 Kennedy, Ildephons 1808-1825 Nesti, Filippo 

1741 Breyne, John Philip 1785 [1788] Joubert de, —— 1808-1837 Fischer de Waldheim, Gotthelf 

1751-1775 ‘Targioni-Tozzetti, Giovanni 1786 Fortis, Giovanni Battista detto 1811 Illiger, Carolus 

1752[1756] Guettard, Jean Etienne Alberto 1814 Rafinesque-Schmaltz, Constan- 

1764 Daubenton, Louis Jean Marie 1788-1803 Camper, Peter _ tine Samuel 



Tilesius von Tilnau, Wilhelm 

Oken, Lorenz 

Smith, William 

1818-1820 Desmarest, Anselme Gaétan 
1818 [1821] Sémmerring, Samuel 


homas von 
Breislak, Scipione 

1820-1823 Borson, Etienne 








2. 1839-1865. 

Gray, John Edward 

Cooper, William 

Harlan, Richard 

Kaup, Johann Jacob 

Lyell, Charles 

Beechey, F. W. 

Brayley, Edward William 

DeKay, James E. 

Meyer, Hermann von 

Hodgson, Bryan Houghton 

Brandt, Johan Friedrich 

Boué, Ami 

Scholl, Johann Baptist 

Hays, Isaac 

Cautley, Proby Thomas 

Klipstein, August von 

Lartet, Edouard 

Caneto, |’ Abbé 

Desor, Pierre Jean Edouard 

Duméril, André Marie Constant 

Geoffroy Saint-Hilaire, Isidore 

Jaecquemin, Emile 

Morren, Charles 

Strauss, F. C. J. von 

Blainville, Henri Marie 
Ducrotay de 








Schlotheim, Ernst Friedrich von 

Krier, Johann Friedrich 

Goldfuss, Georg August 

Cuvier, Frédéric 

Godman, John D. 

Schinz, Heinrich Rudolf 

Fitzinger, Leopold Joseph 
Franz Johann 

HKichwald, Eduard von 

Burmeister, Hermann 

Briggs, C. 

Mather, William Williams 

Bonaparte, Charles Lucien Jules 


Koch, Albert C. 

Foster, J. W. 

Horner, William E. 

Owen, Richard 

Lichtenstein, Martin Heinrich 

Gloger, Constantin W. L. 

Lund, Peter Wilhelm 

D’Orbigny, Alcide 

Grant, [R.] E. 

Lesson, René Primevére 

Nasmyth, Alexander 

Couper, J. Hamilton 

Jackson, J. B.S. 

Falconer, Hugh 

Geinitz, Hans Bruno 

Gray, Asa 

Serres, Pierre Marcel 
Toussaint de 

Giebel, Christoph Gottfried 



Clift, William 

Croizet, l’Abbé Jean Baptiste 

Jobert, Antoine C. G. 

Buckland, William 

Desnoyers, Jules Pierre 
Frangois Stanislas 


Laurillard, Charles Léopold 
Hodgson, William B. 
Gervais, Frangois Louis Paul 
Gay, Claudio 

Temminck, Coenraad Jacob 
Aymard, Auguste 

Jager, Georg Friedrich 
Goppert, Heinrich Robert 
Poppelack, F. 

Pomel, Auguste 

Gibbes, Robert W. 
Quenstedt, Friedrich August von 
Hoernes, Moriz 

Wagner, Andreas 

Agassiz, Jean Louis Rodolphe 
Costa, Oronzio Gabriele 
Sismonda, Eugenio 

Girard, Charles 

Glocker, Ernst Friedrich von 
Hingenau, Otto von 

Pictet, Frangois Jules 
Lockhart, Charles Frangois 
Foetterle, Franz von 
Richardson, John 

Dorlhae, J. 

Reuss, August Emanuel 

3. 1859-1894. DarwiytaAn Preriop TERMINATING witH LeIpy, GAUDRY, AND Copr 

Warren, John C. 
Leidy, Joseph 
Blake, William Phipps 
Hauser, Franz 
Gaudry, Albert 
Rouault, Marie 
Jourdan, Claude 
Abich, Otto Wilhelm 
Hermann von 
Theobald, William 
Gastaldi, Bartolomeo 
Schlegel, Hermann 
Blake, Charles Carter 
Sclater, Philip Lutley 
Logan, William E. 
Suess, Eduard 
Claudius, Matthias 
Le Hon, Henri 
Busk, George 
Murchison, Charles 
Gunn, John 
Marsh, Othniel Charles 
Adams, Andrew Leith 
Cope, Edward Drinker 
Blanford, William T. 
Aradas, Andrea 



Fraas, Oscar Friedrich von 

Hall, James 

Peters, Karl F. 

Dall, William Healey 

Hayden, Ferdinand Vandeveer 

Gill, Theodore 

Chantre, Ernest 

Lortet, Louis 

Dupont, Edouard Frangois 

Marschall, Augusto de 

Biedermann, W. G. A. 

Davies, William 

Yates, Lorenzo G. 

Bachmann, Isidor 

Christy, Henry 

Dana, James Dwight 

Wallace, Alfred Russel 

Major, Charles Immanuel 

Coues, Elliott 

Vacek, Michael 

Stefanescu, Gregoritt 

Ameghino, Florentino 

Fuchs, Theodor 

Acconci, Luigi 




Baretti, Martino 

Grewingk, Constantin Caspar 

Hoernes, Rudolf 

Brauns, David August 

Naumann, Edmund 

Nicolucci, Giustiniano 

Jentzsch, Carl Alfred 

Reiss, Wilhelm 

Weinsheimer, Otto 

Howorth, Henry H. 

Fritsch, K. von 

Thomas, Philippe 

Martin, Johann Karl Ludwig 

Koken, Ernst 

Kittl, Ernst Anton Leopold von 

Verri, Antonio 

Jentink, F. A. 

Carles, Enrico de 

Moreno, Francisco P. 

Riitimeyer, Ludwig 

Weithofer, K. Anton 

Baur, Georg 

Botti, Ulderico 

Cantamessa, Filippo 

Filhol, Henri 










Whitfield, Robert Parr 
Hutchinson, Henry Neville 
Philippi, R. A. 

4. 1900-1933. 

Lydekker, Richard 

Branco [Branca], Wilhelm 

Pohlig, Hans 

Depéret, Charles 

Capellini, Giovanni 

Hofmann, Adolf 

Newton, Edwin Tulley 

Barbour, Erwin Hinckley 

Flower, William Henry 

Lenk, H. 

Zittel, Karl Alfred von 

Felix, Johannes 

Wortman, Jacob L. 

Osborn, Henry Fairfield 

Thomas, Michael Rogers 

Pavlow, Marie 

Lambe, Lawrence Morris 

Tullberg, Tycho Fredrik Hugo 

Boule, Marcellin 

Stehlin, Hans Georg 

Hay, Oliver Perry 

Matthew, William Diller 

Matschie, Paul 

Sclater, William Lutley 

Lucas, Frederick Augustus 

Phillips, P. Lee 

Wiist, Ewald 

Andrews, Charles William 

Andrusov, Nikolai Ivanovich 

Beadnell, Hugh John Llewellyn 

Blake, John Frederick 

Herz, Otto 

Leney, Frank 

Sherborn, Charles Davies 

Bourg de Bozas, Robert 

Clarke, John Mason 

Dawkins, William Boyd 

Nordenskiéld, Erland 

Sternberg, Charles Hazelius 

Stromer, Ernst 

Villada, Manuel Maria 

Zalensky, Vladimir Vladimiro- 

Bate, Dorothea M. A. 

Schlosser, Max 

Gidley, James Williams 

Gregory, William King 

Bartolotti, C. 

Blanckenhorn, Max 

Palmer, Theodore Sherman 

Seguenza, Luigi 

Dubois, Eugen 

De Lorenzo, Giuseppe 

Anderson, Netta C. 

Maddren, Alfred Geddes 

Pompeckj, Joseph Felix 

Udden, Johan August 

Schuchert, Charles 

Pfizenmayer, E. W. 

Pilgrim, Henry Guy Ellcock 




























1893-1896 Portis, Alessandro 
1894-1895 Hayes, Seth 
Graells, D. Mariano de la Paz 

Hornaday, William Temple 

Beck, Richard 

Noack, Th. 

Tokunaga [Yoshiwara], 

Winge, Herluf 

Barbiani, A. 

Fraas, Eberhard 

Merrill, George Perkins 

Sanderson, George P. 

Scott, William Berryman 

Abel, Othenio 

Pontier, G. 

Brown, Barnum 

Gilmore, Charles Whitney 

Wegner, Richard Nikolaus 

Bach, Franz 

Haug, Emile 

Athanasiu, Sava C. 

Lull, Richard Swann 

Mayet, Lucien 

Quackenbush, L. 8. 

Shimek, Bohumil 

Calvin, Samuel 

Wadia, D. N. 

Freudenberg, Wilhelm 

Cook, Harold James 

Evans, George Henry 

Janensch, Werner 

Sierra, Lorenzo 

Stremme, Hermann 

Terra, Paul de 

Schlesinger, Giinther 

Breuil, Henri 

Khomenko, J. 

Dietrich, Wolfgang O. 

Soergel, Wolfgang 

Hilzheimer, Max 

Zuffardi, P. 

Sellards, Elias Howard 

Woodward, Arthur Smith 

Buwalda, John P. 

Heller, Edmund 

Kato, T. 

Lomnicki, M. 

Newton, Richard Bullen 

Niezabitowski, Edward Lubicz 

Oswald, Felix 

Roosevelt, Theodore 

Sato, Denzo 

Schouteden, Henri 

Reck, Hans 

Gez, Juan W. 

Sinclair, William J. 

Cooper, Clive Forster 

Matsumoto, Hikoshichiré 

Cotter, G. deP. 

Kunz, George Frederick 

Merriam, John Campbell 

Airaghi, Carlo 





Trouessart, E. L. 
Heck, L. 
Wagner, George 


Aichel, Otto 

Fourtau, René 

Stefanescu, Sabba 

Bolk, Louis Carles Enrico de 

Brives, Abel 

Carette, Eduardo 

Thevenin, Armand 

Joleaud, Léonce 

Castellanos, Alfredo 

Neuville, Henri 

Frick, Childs 

Reeds, Chester Albert 

Klahn, Hans 

Revilliod, Pierre 

Haughton, Sidney Henry 

Granger, Walter 

Hartnagel, Chris Andrew 

Petronievics, Branislav 

Reyes, Alicia E. 

Roman, Frédéric 

Capitan, Joseph Louis 

Caterini, Francesco 

Majer, Istvan 

Makiyama, Jiré 

Palmer, Rupert William 

Peyrony, D. 

Stauffer, Clinton R. 

Vaughan, Thomas Wayland 

Zukowsky, Ludwig 

Frade, Fernando 

Lang, Herbert 

Plate, Ludwig 

Sandford, Kenneth Stuart 

Furlong, Eustace L. 

Eales, Nellie B. 

Loomis, Frederic Brewster 

Pohle, Hermann 

Schmidtgen, O. 

Wayland, E. J. 

Hopwood, Arthur Tindell 

Anderson, Robert Van Vleck 

Bather, Francis Arthur 

Culver, Harold E. 

Cabrera, Angel 

Dart, Raymond A. 

D’Erasmo, Geremia 

Belizva, E. 

Broom, Robert 

Fabiani, Ramiro 

Peterson, Olof August 

Romer, Alfred Sherwood 

Stock, Chester 

Uhle, Max 

Borissiak, Alexei Alexievich 

Tolmachoff, Innokenty Pavlo- 

Spillmann, Franz 

Absolon, Karel 

Anthony, Raoul Louis 

Hayasaka, Ichiro 


1929 Heurn, F.C. van 1930 Fhik, Julius 1931 Colbert, Edwin Harris 

1929 Prouteaux, M. 1930 Moodie, Roy Lee 1931 Flerov, Constantine C. 

1929 Vaufrey, Raymond 1930 Simionescu, Ioan 1931 Teilhard de Chardin, Pierre 

1929 Yabe, Hisakatsu 1930 Szalai, Tibor 1931-1932 Sunamoto, Etsujiro _ 

1929-1930 Berckhemer, Fritz 1930-1933 Simpson, George Gaylord 1932 Maarel, Frans Henddrik van der 
1933 Koenigswald, Ralph von 

See supplementary Bibliography for 1934 and 1935 at the close of Volume I. 


We should always remember that nomenclature is the tool rather than the master of paleeontologic thought; 
also that no technical principles should override the work of the early discoverers and naturalists. 

In revising the seventeenth to early nineteenth century generic and specific names of Proboscidea, the author 
has found it impossible either to adopt any consistent rules of nomenclature or to adhere to the International 
Code of Zoological Nomenclature of the successive editions of 1901 to 1916, which was formulated by zoologists 
for zoologic practice in which the problems are entirely different from those in palzontologic nomenclature. An 
ex post facto code is difficult in zoology; it is impossible in paleontology until after the time of Cuvier. Zoologic 
types are complete individual specimens, whereas the early and some of the more recent palzontologic types are 
often partial, imperfectly known, and still more imperfectly understood. For example, see the succession of 
names applied to the ‘Mammoth,’ which will be fully set forth in Chapter XVIII, Volume II, and to the ‘Mas- 
todon,’ fully set forth in Chapter VI, as well as the generic names listed more fully in chronologic order below. 

Insertions by the author are in square brackets 

1696 Mammonrtova Kosr Ludloff {Ludolf] (Mammontova, probably derived from the Tatar word mama signifying earth, and 
kost signifying ivory tusks). Original citation inaccessible; cited by Fischer de Waldheim (‘‘Oryctographie,” 
III, Fossiles du Gouvern. de Moscou, 1830-1837, p. 111). 

“T. Elephant. 1. Le Mammont . . . Hlephas mammonteus. m. [Fischer, 1830-1837] . . . Je conserve la 
dénomination la plus ancienne, et je n’écris point Mammouth parce que ce nom n’a pris origine que par corruption 
ou une fausse lecture du mot Mammont [Footnote: ‘Mammouurt en russe; le Yer (+) terminal a été changé par les 
anglais en h et le n a été pris pour un u. D’ailleurs Ludloff est le premier qui en parle (1696) et appelle ces 
ossemens Mammontova Kost et justifie ainsi ma dénomination systématique.’].”’ 

Cited by Cuvier (Ann. Mus., VIII, 1806, p. 45): “‘C’est sous le nom de cornes de mammont, mammontova-kost, 
qu’ils désignent les défenses. ”’ 

1788 MammonteumM Camper (Nova Acta Acad. Sci. Imp. Petropol. Communicanda, II, 1787, p. 251). 

[Type: E. primigenius.] ... “os humanum petrifactum, aut fossile, etiamsi Mammonteorum, Elephan- 
torum, . . . Adserere ex eodem principio audeo Mammonteum animal extinctum non modo esse, sed nullam 
omnino habuisse cum Elephanto similitudinem!”’ 

1806 Mastoponte [= Mastodon] Cuvier (Ann. Mus., VIII, 1806, p. 270). 

[Type: M. americanus.] “Sur le Grand Mastodonte, . . . Animal trés-voisin de l’éléphant, mais 4 macheliéres 
hérissées de gros tubercules, dont on trouve les os en divers endroits des deux continens, et surtout prés des bords 
de l’Ohio, dans l’Amérique Septentrionale, improprement nommé Mammoutu par les Anglais et par les habitans des 

1816 Mastropon Oken (Lehrb. Naturges., Dritter Theil, p. 789). 

“Mastodon” defined, distinguished from Mammoth and Elephant. Cited: 1. [?tapiroides] Montabusard, 
Saxony. 2. M. of Simorre [angustidens]. 3. M. of Chile [humboldtiz]. 4. M. of Cordilleras [andiwm]. 5. M. of 
Ohio [americanus]. 

The full and most confusing chronologic sequence of nomenclature of the Mammoth and the Mastodon 
is set forth in detail in Chapters XVIII, Volume II, and VI, of which the above are examples. 

In surveying the above names the question arises whether Mammonteum Camper, 1788, may be regarded as 
a genus with Elephas primigenius as the type; also, what is the proper orthography of the word Mastodonte G. 


Cuvier, 1806, as compared with the word Loxodonte F. Cuvier, 1825. If Loxodonte is changed to Loxodonta as by 
some authors, why not change Mastodonte to Mastodonta? Similar difficulties arise in connection with every 
generic name proposed for the Proboscidea, excepting only the original Elephas Linnzeus, and even with Elephas 
we are in difficulties, because the Ceylon species, H. indicus Linn., 1754, is not admitted by the Code as antedating 
E. maximus Linn., 1758. Cuvier, the founder of vertebrate paleontology and the first to clearly conceive 
and define the real characters of these extinct mammals, was the last to adopt the Linnean binomial terminology. 
Blumenbach first called the American mastodon Ohio-Incognitum, and then Mammut ohioticum, because he entirely 
lacked the comparative anatomical genius of Cuvier. 


The early editions (I to IX inclusive) of the “Systema Nature”’ of Linnzus are not accepted in the Inter- 
national Code of Zoological Nomenclature. It is important to note that Linnzus used the term Elephas indicus 
in 1754 and called the same animal Elephas maximus in 1758 (“Systema Nature,’ 1758, p. 33). The Editio 
Decima, Reformata, of the ‘Systema Nature”’ by Caroli Linnei, 1758, is accepted in the International Code as the 
starting point of modern systematic zoology. In Chapter XX (the Elephantinz) the specific nomenclature 
under Elephas will be fully discussed. 

(Osborn, “From the Greeks to Darwin,’’ 1894, pp. 128-130): Finally the turning-point to modern ZoGlogy and Botany 
was marked by the great work of Linneus, the Systema Nature. The binary system of nomenclature therein proposed was a 
mere tool for the expression of his broad conceptions of the relation of animals and plants to each other. Species were in his 
mind the units of direct Creation; each species bore the impression of the thought of the Creator, not only in its external form 
but in its anatomical structure, its faculties, its functions; and the end of classification was to consider all these facts and to 
arrange animals in a natural system according to their greater or less likeness . . . he adopted the aphorism of Leibnitz natura 
non facit saltum; to him every species was exactly intermediate between two others: ‘We reckon as many species as issued in 
pairs from the hands of the Creator.’ These were his earlier views in all his writings between 1735 and 1751, in which the 
sentence nullz specie nove recurs, expressing his idea of the absolute fixity of species from the period of their creation as de- 
scribed in Genesis, the only change being that of the extension in numbers, not of variation in kind. Yet Linneus was too 
close an observer to continue to hold this idea of absolute fixity, and in 1762 we find his views had somewhat altered, and 
this is of particular interest because of the hypothesis which he advanced to explain the origin of new species: ‘All the species 
of one genus constituted at first (that is, at the Creation) one species, ab initio wnam constituerint speciem; they were subse- 
quently multiplied by hybrid generation, that is, by intercrossing with other species.’ He was thus inclined to admit a great 
increase of species, more or less recent in origin, arising by hybridity, and losing their perfection of type. He elsewhere sug- 
gested that degeneration was the result of the influences of climate or environment. In the last and thoroughly revised edition 
of the Systema Nature, which appeared in 1766, we no longer find this fundamental proposition of his earlier works, nullz 
Specie nove. 

In the present revision the author has therefore adopted four principles as applicable up to the year 1836. 

1) Recognition of priority of conception and of definition rather than of technical priority of the name. 

2) Principle of soundly established usage and of common sense. 

8) Principle of selection of the first generic or specific name founded on a clearly established character or a clearly designated type. 
4) Well-known principle of ‘elimination’ of species which do not belong, and of ‘restriction’ to species which do belong to the genus. 

The need for such common-sense principles may be illustrated by the case of the names applied to the 
Mammoth and the Mastodon. The early writers on fossil Proboscidea were without the guidance of modern 
rules of nomenclature now embodied in the International Code, either as regards: (1) The orthography or ety- 
mology of names, or (2) the designation of type specimens, or (3) the consideration of priority of publication; nor 
did they realize the value of (4) the Linnean classic orthography, nor (5) the designation of the type locality, nor 
(6) the geologic level or formation in which types occur. 

Consequently to rob Cuvier of his clear conception of grinding tooth structure, which he termed Mastodonte, 
and to substitute the barbaric term Mammut, signifying ‘earth-burrower,’ would be gross injustice to the founder 
of vertebrate paleontology. 




Osborn’s second principle of nomenclature accords in general with the successive revisions by G. Cuvier 
(1799, 1806), by Oken (1816), by G. Cuvier (1817), by Lartet (1836), by Burmeister (1837), by De Blainville 
(1839-1864), by Gloger (1841), by D’Orbigny (1842), by Falconer (1846, 1857, 1865, 1868), by Laurillard (1846), 
by Pictet (1853), by Leidy (1869), by Cope (1889), and by Trouessart (1899). 

BARBARIC AND Ciassic TERMS APPLIED TO ELEPHAS PRIMIGENIUS.—The mammoth (Elephas primigenius 
of Eurasia) received a series of christenings beginning in 1696 when Ludolf termed the mammoth “‘Mammontova 
Kost,” a term cited by Cuvier (1806.1, p. 45), also by Fischer de Waldheim (1830-1837) who insisted that Mammont 
is the proper expression of the Russian vernacular and that Mammouth is a corruption. Camper’s Mammonteum 
(1788) is cited by Leidy (1869, p. 392). This serves to show that in the nomenclature of the seventeenth and 
eighteenth centuries the terms Mammont and Mammouth were repeatedly applied both in vernacular and in 
scientific usage to the animal now known as Elephas primigenius. Camper’s Latin term Mammonteum, by trans- 
position Mammonteus, is the first classic term applied to the mammoth; a full list of the terms subsequently 
applied will be cited in Volume II. 

Mammut Blumenbach (1799), although preoccupied in the vernacular for Elephas primigenius and a synonym 
of Ohio Blum. (1797.2), was revived as a genus for the American mastodon by Hay (1902); so far as we know 
Hay was the first author to recognize Blumenbach’s vernacular term Mammut, a term not employed in the 
designation of the type figures by Blumenbach himself. Our greatest American authority on generic names, 
Palmer (1904), following Hay, has accepted Mammut (op. cit., 1904, p. 397); it has also been adopted by Lull 
(1908). It certainly seems in the interest of paleontology to regard the term Mammut as technically pre- 
occupied by Ohio-Incognitum Blum. and in historic justice to Cuvier to retain firmly in proboscidean nomen- 
clature the appropriate historic name Mastodon Cuvier. 

De Blainville’s citations (1839-1864, p. 245) of the barbaric and classic generic names applied to Mastodon 

americanus are as follows: 
1° Le Granp Mastoponte. 

Ohio incognitum, Blumenbach. Abildungen, n° 19 ,1797- 

Elephas Americanus, Pennant. 

Mammouth Ohioticum, Blumenbach. Manuel d'Hist. nat. 

E. Americanus, molaribus multicuspidibus, lamellis post detri- 
tionem quadri-lobatis, G. Cuvier, sur les esp. d'Eléph. viv. et foss. 

Mastodon giganteum , G. Cuvier. Ann. du Mus. VA, p. 270, pl. 49-56, 


‘MASTODON’ SOUNDLY ESTABLISHED FOR MASTODON AMERICANUS.—Towards the beginning of the eighteenth 
century the American mastodon was discovered. In 1771 Pennant named this animal the American Elephant. In 
1792 Kerr named this animal Elephas americanus; in 1797 Blumenbach (1797.2, Abbildungen, No. 19) first named 
and figured it as Ohio-Incognitum; in 1799 he named it Mammut ohioticum, using the preoccupied word Mam- 
mut, but in 1810 (Abbildungen, No. 19) he again reproduced this figure with the name Ohio-Incognitum on the 
plate. According to Falconer (1846, p. 18) and De Blainville (1839-1864, p. 245) the name Mastodon giganteum 
Cuvier dates 1805; this date is not accepted by Leidy (1869, p. 393) or Palmer (1904, p. 401). In 1806 Cuvier 
described the animal as ‘‘Le Grand Mastodonte,” after defining the term in earlier papers (see De Blainville, 
1839-1864, p. 245). Cuvier did not at the time adhere to the Linnzan classic orthography in naming his genera, 


e.g., Mastodonte (= Mastodon). His name was cited, however, by Oken in 1816 as Mastodon and by Cuvier 
himself as Mastodon in 1817. 

Cuvier’s admirable selection of the diagnostic name Mastodon, which has since been reprinted millions of 
times in all the scientific and popular literature, should not be set aside for any form of the barbaric and pre- 
occupied term Mammut. 


Osborn’s third principle of paleeontologic nomenclature is of an entirely different kind; it applies to the 
rules which should govern us in the selection of generic names founded on two or more species which are now 
known to belong to one or more different genera, e.g., the name Trilophodon Falconer, et seq. 

In the chronologic list of Generic Names and Genotypic Species given below, in each instance the following 
eight rules are observed: 

(1) The original orthography is replaced by the classic orthography. 

(2) The earliest known use of the name is cited in the author’s orthography. 

(3) The words nomen nudum indicate that the author gave no definition by which the ‘species’ could be determined; or that the name 
is @ synonym. 

(4) The earliest type figure is cited and where valid and available is reproduced. 

(5) The genotype species are invariably cited in the order given by the original author. 

(6) The geographic locality is cited where given (= Hab.). 

(7) The geologic age or level is cited also (= Type loc.). 

(8) The name or specific determination adopted by the present author is given. 

Important Note.—Few if any of the eight principles of the selection of generic or specific names can with fair- 
ness be applied retroactively to the 1698-1836 Cuvierian period, as listed above. A notable illustration of the 
perils attached to such procedure is afforded by the first mentioned type specimen of the species Hlephas primigenius 
Blum.; in this classic description Blumenbach first refers to the fossil skeleton described by Tentzelius as un- 
earthed at Burgtonna near Gotha; this skeleton is now positively known to be a specimen of the straight-tusked 
elephant named Elephas antiquus by Falconer in 1847, 1857. The technical type of Hlephas primigenius Blum. 
is the specifie type of Elephas antiquus Fale. Technically, therefore, the species named primigenius antedates the 
species named antiquus. Imagine the dislocation of the entire literature of the Proboscidea if the name antiquus 
were substituted for the name primigenius on technical grounds covering the modern selection of the type, holo- 

type, cotype, ete. 


Beginning with the close of the Cuvierian period in the year 1836, we may with greater justice apply the 
modern principles of paleontologic nomenclature which came into being during the Darwinian period (1859- 
1894) and which have been gradually clarified and intensified in the phylogenetic period (1900-1933). During the 
phylogenetic period two important steps have been taken: First, in revising the work of Cope, Leidy, and Marsh, 
Osborn discovered that the type locality and geologic level were of extreme importance in the study of phyletic 
series of species; second, Schuchert and others sharpened and clarified the conception of the word type, which may 
have an entirely different significance in palzeontology from that conveyed in zoology. Accordingly the modern 
technical procedure may be summarized under the following eleven rules: 

(1) We must first determine the geographic locality and where possible the geologic level as the starting point of the 
determination of the characters of the type, because both these facts have a very important bearing on generic and specific defini- 
tion. Experience teaches that the characters of the original type specimen, the geographic locality, and the geologic level on which 
the specimen was found afford the permanent facts to which all questions of nomenclature must finally be referred in mono- 
graphic investigation. 

(2) The type, where not specifically designated by the original author, is chosen (lectotype) as the first specimen men- 
tioned, the first described, the first figured by the author, consequently great attention has been paid in this Iconography to 
the selection of the particular specimen or specimens which conform with the author’s first description and definition. 


(3) Do not confuse the holotype characters with other characters based on cotype, paratype, or neotype specimens which 
the author may have erroneously considered as belonging to the same species as the type. Much of the early systematic work 
on the Proboscidea was done without discrimination between the permanent facts to be derived from the type specimen and 
the opinions which the original or subsequent authors derived from erroneously associated specimens. Osborn’s rule is that 
specific definition must be based on original types or holotypes only, unless there is absolutely no possibility of doubt that the 
associated specimens are of the same geologic level and belong to the same species. 

Osborn’s usage of the terms type (holotype), cotype, paratype, lectotype, and neotype conforms in general with that of 
Oldfield Thomas (1893), of Cossmann (1904), of Schuchert (1905.1, 1905.2), and of Osborn (1918.473). 

(4) Type=Ho.oryrer. A particular individual specimen “deliberately selected by the author of a species, or it may be 
the only example of a species known at the time of original publication. A holotype therefore is always a single individual, but 
may embrace one or more parts, as the skin, skeleton, or other portions’ (Schuchert, 1905.2, p. 10). The holotype must 
usually be determined from the original description. 

(5) Coryprs=CodrDINATE OR EquivaLent Typrs. This term is applicable when the author’s type description refers 
to two or more individuals without selecting or designating one as the holotype, so that all appear to be identified equally, by the 
original author, with the specific name given. Osborn’s custom in such cases is to designate as the type (lectotype) the 
specimen first mentioned, described, or figured. 

(6) Paratypr=SuBoRDINATE OR SUPPLEMENTARY Typr. Besides the particular individual specimen deliberately 
selected by the author of a species as the type, other specimens mentioned or enumerated in the original description which may 
supplement the characters of the type may be chosen as paratypes. This procedure is especially valuable in paleontology, for 
the type specimen very seldom consists of a complete individual. ‘ 

(7) LecrorypE=Cuosen Type. ‘Where the original diagnosis is without illustrations or is accompanied by figures 
based on two or more specimens, the first subsequent author is at liberty to select from these cotypes a type for the old species, 
adhering, as far as can be ascertained, to the intention of the original author. Such a type specimen is to be designated as a 
Lecrotyrr (=achosen type).”’ (Schuchert, 1905.2, p 12). The lectotype practice of Osborn in paleontology is either (a) to 
select the first individual specimen named by the original author, because it sometimes happens that the second individual 
specimen belongs to a distinct species, or (b) to select the specimen to which the specific name or description obviously refers. 

(8) Nrotrype=New Tyrer. Defined by Schuchert (1905.2, p. 14) as a new ‘supplementary type selected by an author, 
on which a species is to rest because of the loss of the original type or where the original material still extant is so poor 
or fragmentary that from it the characters of the species can not be determined with certainty.” Great care must be taken 
that the neotype comes from the same geologic level as the type. 

(9) Meratypr. Defined by Schuchert, after Thomas (Schuchert, 1905.2, p. 14) as “‘A specimen received from the original 
locality [in paleontology, the exact stratum as well] after the description has been published, but determined as belonging to 
his own species by the original describer himself.’ ”’ 

(10) Tororyrr (Schuchert, 1905.2, p. 14). ‘In paleontology it is further demanded that the topotype should come not 
only from the exact locality but also from the identical stratum that furnished the species.” 

(11) Ipkorypr (Schuchert, 1905.2, p. 15). ‘‘These are the specimens from any place except the original locality, named 
by an author of a species after publication [e.g., M. sivalensis].” 


We now come to the practical test of the above principles of nomenclature in vertebrate paleontology and 
find that step by step the application of these principles enables us to do full justice to the work of the great leaders 
and discoverers in proboscidean paleontology, such as Cuvier and Falconer, only by setting aside in certain cases the 
generic and specific names proposed by revisers and reviewers of less authority and of little or no original thought. 
The student will observe the following five points: 

(1) Great confusion exists in the use of the eighty or more generic names which have been proposed for the Proboscidea. 

(2) Much research has previously been devoted to the revision of this nomenclature by De Blainville, Falconer, Leidy, Cope, 
Trouessart, Ameghino, Palmer, Hay, Matthew, Lull, Schlesinger, and Freudenberg. In the preparation of the present Memoir 
the author has enjoyed the cordial coéperation of the late Joel A. Allen dean of American mammalogists and highest American 
authority on zoological nomenclature, of T. 8. Palmer author of the invaluable ‘Index Generum Mammalium,”’ of the late 
W. D. Matthew who devoted much time and thought to this subject; also of the British Museum authorities, Oldfield Thomas, 
Charles W. Andrews, and F. A. Bather, as well as C. Davies Sherborn author of the “Index Animalium.”’ 

(3) The notes and correspondence relating to this difficult subject are bound and filed in the Osborn Library, including 
the author’s inquiries and replies, under the title: OSsBoRN’s CORRESPONDENCE, NOMENCLATURE OF THE ProposcipnA, 1921-1935. 
As in the case of the revision of species, reference is made to these notes and letters as, for example: (Allen, letter, 1921). 

(4) The reader will observe between 1758 and 1931 a progressive departure from the Linnzan conception of a genus. The 
generic names first proposed are really collective or group names, e.g., Elephas equals all elephants, Mastodon equals all masto- 
donts, T'rilophodon equals all mastodonts with three crests in the intermediate molars. Even so recently as 1886 Lydekker em- 
braced all known elephants, recent or fossil, under the single term Hlephas, and all known mastodonts under the single term 
Mastodon; Freudenberg (1922) has followed this practice. 

(5) Recently certain authors who have gone to the opposite extreme have proposed new generic names based on single 
characters or single stages of evolution. 



The generic names proposed or adopted since 1923 are listed below (p. 11) 

Exeruas Linneus, 1735-1758, for all the elephants of Indian type. Type Elephas indicus =mazimus. 

Mammonteus Camper, 1788. Type Elephas primigenius. This Latinized vernacular name (Mammonteum Camper) pre- 
occupies the long series of generic names subsequently applied to the Mammoth, e.g., Dicyclotherium, Cheirolites, Syno- 
dontherium, and Polydiskodon. 

Mastopon G. Cuvier, 1806, 1817, for all the zygolophodont mastodonts which are actually related to the genotypic species 
M. americanus. 

Loxoponta F. Cuvier, 1825, 1827, for all the elephants of African type. Type Elephas africanus. 

DeEINOTHERIUM Kaup, 1829, for all the Deinotheres. Type Deinotheriwm giganteum. 

Srrcopon Falconer and Cautley, 1846 [1847], 1857, for all pro-elephants which resemble the genotypic species E. cliftii, E 
bombifrons, EL. ganesa, E. insignis. 

Anancus Aymard, 1855, for all the bunolophodont, brevirostrine mastodonts which are closely related to Aymard’s genotypic 
species Anancus macroplus=M. arvernensis. 

TrILOPHODON Falconer, 1846-1857, collective name, restricted to M. angustidens. 

TETRALOPHODON Falconer, 1847-1857, collective name, restricted to M. longirostris. 

PENTALOPHODON Falconer, 1857-1865, collective name, restricted to M. siwalensis. 

RuyncuornertuM Faleoner, 1856-1868, based upon a very distinctive kind of lower jaw from Mexico and including a large 
number of American forms of similar character. Type Rhynchotherium tlascalz. 

ZyGoLoPHopoN Vacek, 1877, an excellent genus based on the species M. borsoni, M. turicensis, M. tapiroides, M. pyrenaicus, 
all of which are probably interrelated and are certainly distinct from the ancestry of M. americanus. Osborn (1926.706) 
removed M. turicensis from Zygolophodon making it the type of the genus Turicius. He also refers M. tapzroides to 
the genus Turicius. 

Arcuipiskopon Pohlig, 1885-1888, founded on the very distinctive specific types Elephas planifrons and E. meridionalis. 

Pataomastopon Andrews, 1901, founded on the very distinctive type P. beadnelli. 

MeritHertum Andrews, 1901, for all the true Meeritheres. Type Meritherium lyonst. 

Pxuromra Andrews and Beadnell, 1902, founded on the very distinctive type Phiomia serridens. 

Unfortunately several of the remaining sixty or more generic names which have been proposed do not rest 
on such indisputable grounds of clear conception and definition as the above. The erroneous or invalid names fall 
into four categories: 

(1) Indeterminate names, that is, names defined without a clear conception of generic characters or without a clear designa- 
tion of the genotypic species, or with an erroneous designation of genotypic species, or other obvious misconceptions. Examples 
of such names are Chio Blum., 1797, Mammut Blum., 1799, Gomphotherium Burm., 1837=Gamphotherium Gloger, Czno- 
basileus Cope, 1877, Hemimastodon Pilgrim, 1912, Promastodon Pohlig, 1912, also Choerolophodon Schlesinger, 1917. 

(2) Collective, Section or Growp Names. Names founded on parallel or convergent dental characters which arise entirely 
independently within different genera and different phyla. The earliest outstanding examples of this kind are Falconer’s collec- 
tive section or group names, T’rilophodon, Tetralophodon, and Pentalophodon, e.g. : 

Trilophodon Fale., 1846-1857, first used as a Section to include Mastodon ohioticus, Deinotherium giganteum, M. tapirotdes, 

M. angustidens, and M. andium, species with three-crested intermediate molars. 
Trilophodon Falc., 1857, subsequently defined as a subgenus, with the genotypic species M. angustidens. 

Tetralophodon Falec., 1847-1857, first used as a Section to include Mastodon perimensis, M. sivalensis, M. arvernensis, M. 

longirostris, and M. latidens, species with four-crested intermediate molars. 
Tetralophodon Warren, 1852, with the genotypic species: M. latidens [=Stegolophodon], M. arvernensis [= Anan- 
cus], and M. sivalensis [= Pentalophodon]. These species obviously belong to several distinct genera or phyla. 
Tetralophodon Falc., 1857, subsequently defined as a subgenus, with M. longirostris as the genotypic species. 

a ee ata Fale., 1857-1865, defined to include species with five-crested intermediate molars; genotypic species M. 


(3) Collective or group names founded upon an erroneous assemblage of species belonging to two or more different genera or 
phyla. Examples of names of this kind are in chronologic order as follows: 

Mastotherium Fischer, 1814. Genotypic species: M. megalodon [= Mastodon], M. leptodon [=Trilophodon], M. microdon 

[= Turicius], M. hyodon [= Cordillerion|. 

Bunolophodon Vacek, 1877. Genotypic species: M. arvernensis [=Anancus], M. pentelicus [= Trilophodon (Choerolopho- 

don)], M. atticus [=Turicius], M. longirostris [= Tetralophodon], M. angustidens [= Trilophodon|. 

Dibelodon Cope, 1884. Genotypic species: D. shepardi [= Rhynchotherium]. Referred: D. tropicus [=Cordillerion], D. 

humboldtii |= Cuvieronius]. 

Tetrabelodon, Cope, 1884. Genotypic species: 7. angustidens [=Trilophodon]. Referred: T. andium [=Cordillerion], T. 

productus = Serridentinus]. 

Polydiskodon Pohlig, 1885-1888. Genotypic species: Elephas primigenius [=Mammonteus], E. indicus |=Elephas], E. 

namadicus {= Palzoloxodon]. 

(4) Obvious synonyms, as listed below, which are also frequently collective, are readily disposed of, e.g.: 

Euelephas Fale., 1857. Genotypic species: EF. indicus [=Elephas], E. armeniacus [=Parelephas], ?E. hysudricus 

[= Elephas (Hypselephas)]. 


SEVERAL DispuTABLE NAmEs.—From the above analysis it appears that there remain several generic names 
improperly defined and as to the validity of which there may be decided differences of opinion, namely, Gampho- 
therium Gloger, 1841, Trilophodon Falc., 1846-1857, Tetralophodon Falc., 1847-1857, Pentalophodon Fale., 1857- 
1865, Mastotherium Fisch., 1814, Bunolophodon Vacek, 1877, Dibelodon Cope, 1884, Tetrabelodon Cope, 1884, and 
Polydiskodon Pohlig, 1885-1888. 

It is in the usage of such names that the widest differences of opinion have been manifested and that the 
greatest confusion as to the true phylogenetic evolution of the Proboscidea has arisen, for the common reason that 
all these names were founded on misconceptions of true relationships. It is in such a group of names also that it 
proves impossible to adopt any uniformly consistent principle of revision. Consequently for reasons more fully 
stated below we have cut the Gordian knot by courtesy to Falconer and have adopted three collective generic 
names, as follows: 

Trilophodon Fale., 1846-1857, collective name, restricted to M. angustidens. 

Tetralophodon Fale., 1847-1857, collective name, restricted to M. longirostris. 
Pentalophodon Falc., 1857-1865, collective name, restricted to M. sivalensis. 

We have eliminated five other collective names, because they are synonyms of collective names previously 
proposed, namely: 

Mastotherium Fischer, 1814, collective name, also a synonym of Mastodon, etc. 

Bunolophodon Vacek, 1877, collective name, also a synonym of Anancus, of Tetralophodon, of Trilophodon. 
Dibelodon Cope, 1884, collective name, also a synonym (in part) of Mastotherium and of Tetrabelodon. 
Tetrabelodon Cope, 1884, collective name, also a synonym of T’rilophodon and of Dibelodon. 

Polydiskodon Pohlig, 1885-1888, collective name, also a synonym of Mammonteus, of Elephas, and of Loxodonta. 


The sixteen generic names adopted up to the year 1923 as representing distinct and separate phyla, namely, 
Elephas, Mammonteus, Mastodon, Loxodonta, Deinotherium, Stegodon, Anancus, Trilophodon, Tetralophodon, 
Pentalophodon, Rhynchotherium, Zygolophodon, Archidiskodon, Palxomastodon, Meritherium, and Phiomia, are 
now amplified by the adoption of Stegomastodon, Eubelodon, Stegolophodon, Miomastodon, Cuvieronius, 
Serridentinus, Ocalientinus, Serbelodon, Parelephas, Pliomastodon, Turicius, Palxoloxodon, Cordillerion, Amebel- 
odon, Platybelodon, Synconolophus, Notiomastodon, Hesperoloxodon, Platelephas, (?) Hypselephas, Trobelodon, 
Blickotherium, and Aybelodon. 

The addition of twenty-three generic names to the sixteen first listed, making thirty-nine’ in all, does not 
signify the discovery of thirty-nine' distinct phyla or branches, because certain of these genera represent geologic 
sections in certain branches; for example, it is probable that Miomastodon, Pliomastodon, and Mastodon were 
actually successive genera; similarly that Phiomia branched into Amebelodon. 

Eliminating such succession of genera, we have thus far discovered no less than TWENTY-SIX GREAT PHYLA in 
which our knowledge includes one or more sections conveniently designated as GENERA. 

SreGomastTopon Pohlig, 1912, founded on the Mastodon mirificus of Leidy 

EvuBELopon Barbour, 1914. Type Eubelodon morrilli, a single individual from Devil’s Gulch, Nebraska. 

SrecoLopHopon Schlesinger, 1917, founded on the distinctive type of Mastodon latidens Clift. 

Mromastopon Osborn, 1922. Type Mastodon merriami, a Middle Miocene mastodont intermediate in character between Palzo- 
mastodon and Mastodon. 

Cuvirrontus Osborn, 1923, based upon the species Mastodon humboldtii Cuvier, characterized by double trefoils and tusks 
without enamel band. 

SERRIDENTINUS Osborn, 1923. Genotypic species: Mastodon productus Cope, M. serridens Cope, M. floridanus Leidy, M. 
obscurus Leidy, and Serridentinus |= Trilophodon] simplicidens Osborn. 

‘Forty-one by the addition of Torynobelodon and Gnathabelodon. 


PARELEPHAS Osborn, 1924. Upper Pliocene to Upper Pleistocene genus based on the species Elephas jeffersonii Osborn, of 


Puromastopon Osborn, 1926, a Pliocene stage based on the species first described by Osborn as Mastodon matthew, subse- 
quently as Pliomastodon matthewt. 

PaLtmoLoxopon Matsumoto, 1924, founded on Hlephas namadicus naumanni Makiyama. 

Turicrus Osborn, 1926, based on Mastodon turicensis Schinz, one of the genotypes of Zygolophodon Vacek, 1877. 

CorpILLERION Osborn, 1926. Type Mastodon andium Cuvier, characterized by single trefoils and spirally twisted tusks with 
enamel band. 

AMEBELODON Barbour, 1927. Type Amebelodon fricki, distinguished especially by its shovel-shaped tusks. 

PLATYBELODON Borissiak, 1928. Type Platybelodon danovi, distinguished especially by its flat tusks. 

SynconoLopuus Osborn, 1929. Type Synconolophus dhokpathanensis, a very progressive species of this phylum from the 
Middle Pliocene of India. 

Norromasropon Cabrera, 1929, founded on the distinctive type Notiomastodon ornatus. 

Hesprerotoxopon! Osborn, 1931, provisionally proposed, with Palzolorodon antiquus italicus as the type, representing a 
phylum quite distinct from that of ‘Elephas namadicus’ of the Siwaliks. 

Burckxotuerium Frick, 1933. Type Blickotherium blicki Frick. A Rhynchorostrine with peculiarly formed symphysis and 
laterally compressed, enamel-banded incisors. 

AyseLopon Frick, 1933. Type Aybelodon hondurensis Frick. A Rhynchorostrine with slender jaws, upcurved rostrum, and 
enamelless tusks. 

Trosetopon Frick, 1933. Type Trobelodon taoensis Frick. Of Serridentine relationship. 

OcauientINUS Frick, 1933. Type Ocalientinus ojocaliensis Frick. Serridentine grinders, abbreviate cranium, and elongate, 
troughlike rostrum. 

SERBELODON Frick, 1933. Type Serbelodonbarbourensis Frick. Short, broad symphysis and laterally flattened incisors. 

PLATELEPHAS gen. noy.!_ Type Elephas platycephalus Osborn, having a relatively elongate, dolichocephalic, and platycephalic 

Hypseepuas gen. noy.! Genotypic species: Elephas hysudricus Falconer, 1845, 1846, and Hlephas platycephalus angustidens 
Osborn, 1929. 


Modern terminology and classification express the discovery of lines of descent. As clearly shown in Chapter 
II of the present Memoir, the Osborn phylogenetic system (1892-1931) adapts the Linnzean ‘special creation’ 
nomenclature to the Darwinian ‘divergent-evolution’ principle of descent. As set forth in a long series of writings 
on this subject, beginning with Osborn, 1892.67, an essential outcome of Darwinism is the substitution of a 
nomenclature conformable to the divergent branching or polyphyletic adaptations of the Proboscidea to diverse 
habitats and different feeding habits. 

EvoLuTionary Stace Tueory. In the 14th edition (1929) of the Encyclopaedia Britannica the articles 
ProposcipEA, ELEPHANT, MAmmotu, and Masropon summarize the general knowledge of these animals as 
prevailing in the decades previous to the year 1929. These divide the Proboscidea into the following nine stages: 

Ist Stage. Mcritheritwm, Upper Eocene of Egypt. Very primitive. 

2d Stage. Palzxomastodon, Lower Oligocene of Egypt. Great advance on Meritherium. 

3d Stage. Tetrabelodon angustidens, Lower Middle Miocene of Europe. Advance on Palzomastodon. 
4th Stage. Tetrabelodon longirostris, Pliocene of Europe. In external appearance entirely elephantine. 
5th Stage. Mastodon, Pleistocene. Terminal stage of the mastodonts. 

6th Stage. Deinotherium, Mio-Pliocene of Eurasia. Side branch. 

7th Stage. Stegodon, Pliocene of India. Transitional to the elephants. 

8th Stage. Hlephas antiquus of Eurasia and other Plio-Pleistocene species. 

9th Stage. EHlephas maximus, E. africanus, of India and Africa. 

The arrangement of the Proboscidea in these nine stages accords with the outworn belief among zoologists 
that one or more of these stages represents a successive or ascending order; for example, that Meritherium gives 
rise to Palzomastodon, as, in turn, Mastodon passes into Stegodon and then into Elephas. This prevailing belief 
is entirely erroneous; it is a vestige of monophyletic theories of animal descent which in the past have been largely 
due to the breaks and gaps in our palzontologic collections. Now that these gaps are in a great measure filled 
by discovery and these animals intensively studied, the older monophyletic theories prove to be illusory; they 

Description in forthcoming Volume II (The Elephantoidea). 


are wholly superseded and replaced by polyphyletic systems in which each of the generic phyla, even of the Oligo- 
cene, passes back into fundamental divisions that occurred during Eocene times. It is proven that the divergence 
or branching of the Proboscidea, like that of all other ungulates, was well advanced in the Eocene. 

In brief, it becomes necessary to replace the old evolutionary stage theory by the new polyphyletic adaptive 
radiation theory of Osborn. 

POLYPHYLETIC ADAPTIVE RapiaTION THEORY.—Each of the successive chapters of this Memoir (Chap. IT, 
“Origin and Classification of the Proboscidea,”’ to Chap. XX, ‘The Subfamily Elephantinz’’) contains a full 
exposition not only of the principles of adaptive radiation but of the necessary modification of the nomenclature 
by polyphyletic discovery. Finally in Chapter X XI (Affinities, Migrations, and Phylogeny’’) the whole subject 
of the adaptive evolution of the Proboscidea along polyphyletic lines is fully set forth. 

Each of the forty-one genera finally established in the present Memoir represents a clearly distinct stage 
of descent; for example, it has very recently been determined that the line of descent terminating in the ‘Elephas 
namadicus’ of India is quite distinct from the line of descent terminating in the ‘Elephas antiquus italicus’ of 
western Europe. 
generic names must be applied to these two lines of descent and to their African relatives, namely: 

It follows that, not only as a matter of convenience but as an expression of truth, different 

Elephas antiquus line of descent = Hesperoloxodon Osborn, 1931. 

Elephas namadicus line of descent = Palxoloxodon Matsumoto, 1924. 

Elephas africanus line of descent = Loxodonta F. Cuvier, 1825, 1827. 

Thus our modern phylogenetic knowledge completely revolutionizes all the older taxonomic systems and 


As recited in the Preface and more fully in Chapter II, the preparation of this Memoir began in the year 1907 
with the author’s expedition to the Fayim of northern Egypt, accompanied by Walter Granger as head of the 
mammalian paleontology staff of the American Museum. This eventful journey verified and amplified the 
remarkable discoveries of Hugh J. L. Beadnell and Charles W. Andrews, and determined the author to give 
the Proboscidea monographic treatment similar to that of the Titanotheres. 

At the time there was little conception of the difficulties of the undertaking. First entered upon in the 
midst of the concluding phases of research on the Titanotheres—the Titanothere Monograph having been sent to 
press in 1920—intensive work on the Proboscidea Memoir has continued with periods of interruption for fourteen 
years—altogether twenty-seven years. 

It soon became obvious that the codperation of practically all the mammalian paleontologists of the world 
was essential to a comprehensive and thorough treatment of the subject. It would be impossible to express to 
each of these collaborators the full measure of the author’s appreciation; it is, moreover, difficult to give a com- 
plete list of those who have helped in many ways. 

OrTHENIO ABEL, Professor of Paleobiology, Universitit, Vienna, 

Karev Apsoton, Curator of Zoology and Paleontology, Moravské 
Zemské Museum, Briinn, Czechoslovakia. 

*CarL ErHan AKELEY, Associate in Mammalogy, The American 
Museum of Natural History, New York City. 

*JoreL, AsApH ALLEN, Curator of Mammalogy, The American Mu- 

seum of Natural History, New York City. 

*CHARLES WILLIAM ANDREWS, Assistant in Department of Geology, 
British Museum (Natural History), London, England. 

Haroup Eimer ANTHONY, Curator of Mammalogy, The American 
Museum of Natural History, New York City. 

Raout ANTHONY, Professor of Comparative Anatomy, Muséum 
National d’Histoire Naturelle, Paris, France. 

Erwin Hincxuey Barsour, Professor of Geology, University of 
Nebraska; Curator, Nebraska State Museum, Lincoln, 

Dorotuea M. A. Bare, Unofficial Scientific Worker, in Depart- 
ment of Geology, British Museum (Natural History), 
London, England. 

*Francis ARTHUR Batuer, formerly Keeper of the Department of 
Geology, British Museum (Natural History), London, 

Fritz BerckHemer, Curator, Geologische Abteilung, Wiirttem- 
bergische Naturaliensammlung, Stiittgart, Germany. 


Wituram Rei Buarr, Director and General Curator, New York 
Zoological Park, New York City. 

Auexer AbexrevicH BortsstaAk, Geological Museum of the 
Academy of Sciences, Leningrad, Union of the Socialist 
Soviet Republics. 

Marce.LLIn Bouse, Professor of Paleontology, Muséum National 
d’Histoire Naturelle; Director, Institut de Paléontologie 
Humaine, Paris, France. 

Epovuarp BourpELLE, Muséum National d’Histoire Naturelle, 
Paris, France. 

Isatan Bowman, Director, American Geographical Society, New 
York City. 

FrerpinAND Brom, Director, Institut fiir Palaiontologie und 
historische Geologie, Bayerische Ludwig-Maximilians- 
Universitit, Munich, Germany. 

Rogert Broom, formerly Professor of Geology and Zoology, Vic- 
toria College, Stellenbosch, South Africa. 

Barnum Brown, Curator of Fossil Reptiles, The American Mu- 
seum of Natural History, New York City. 

Wittram Atanson Bryan, Director, Los Angeles Museum, Los 
Angeles, California. 

Joun P. Buwaupa, University of California, Berkeley, California. 

Ancet Caprera, Professor of Palzeontology, Instituto del Museo y 
Escuela de Ciencias Naturales, Universidad Nacional de 
La Plata; Curator of Paleontology, Museo de La Plata, 
La Plata, Argentina. 

*GIOVANNI CAPELLINI, formerly Professor of Geology, University, 
of Bologna, Italy. 

JorL Ernest Carman, Professor of Geology, Ohio State Univer- 
sity, Columbus, Ohio. 

*Jonn Mason Cuarkb&, formerly Director of the New York State 
Museum, Albany, New York. 

Epwin Harris Coisnrt, Assistant Curator of Vertebrate Palzeon- 
tology, The American Museum of Natural History, New 
York City. 

Harotp JAMES Cook, formerly Curator of the Department of 
Paleontology, Colorado Museum of Natural History, 
Denver, Colorado. 

Curve Forster Coorrr, University Museum of Zoology, Cam- 
bridge, England. 

Raymonp A. Dart, Professor of Anatomy and Dean of the Faculty 
of Medicine, University of the Witwatersrand, Johannes- 
burg, South Africa. 

Giuseppre Dr Lorenzo, Professor of Geology, Universita di Napoli, 
Naples, Italy. 

*CnarLes Drpiret, formerly Dean of the Faculty of Sciences, 
Université de Lyon, France. 

GeremiA D’Erasmo, Professor of Paleontology, Universita di 
Napoli, Naples, Italy. 

Wo.reana O. Dierricn, Geologisches Palaeontologisches Institut 
und Museum der Universitit, Berlin, Germany. 
Raymonp Ler Dirmars, Curator of Mammals and Reptiles, New 

York Zoological Park, New York City. 

EKucen Dusots, Koninklijke Akademie van Wetenschappen, 
Amsterdam, Netherlands. 

Ne ur B. Faxes, Lecturer, Department of Zoology, University of 
Reading, England. 

Jesse Dapp Fiaarns, Director, Colorado Museum of Natural 
History, Denver, Colorado. 

WitHeLtM FRevpENBERG, formerly of Georg August Universitit, 
Gottingen, Germany. 

Cuitps Frick, Honorary Curator of Late Tertiary and Quaternary 
Mammals, also Trustee, The American Museum of 
Natural History, New York City. 

Isustace L, Furtone, Museum of Paleontology, University of 
California, Berkeley, California. 

CLAUDE ene, Director, Musée des Sciences Naturelles, Lyon, 


«James WILLIAM Gip.tay, Assistant Curator of Fossi] Mammals, 
United States National Museum, Washington, D. C. 

CHARLES WHITNEY GILMoRE, Curator, Department of Vertebrate 
Paleontology, United States National Museum, Wash- 
ington, D. C. 

ALLEN H. Gopsey, Professor, Old Testament, Duke University, 
Durham, North Carolina. 

Water GRANGER, Curator of Fossil Mammals, The American 
Museum of Natural History, New York City. 

Maprson Grant, President of the New York Zoological Society, 
New York City. 

Witu1am Kina Grecory, Curator-in-Chief of Living and Extinet 
Fishes, Curator of Comparative and Human Anatomy, 
Research Associate in Paleontology, Associate in Anthro- 
pology, The American Museum of Natural History, 
New York City. 

Herman Gunter, Sate Geologist, Tallahassee, Florida. 

C. R. Hatrer, formerly of The American Museum of Natural 
History, New York City, and of the Department of 
Biology, Southern College, Lakeland, Florida. 

Rosert Torrens Hart, Assistant Curator of Mammalogy, The 
American Museum of Natural History, New York City. 

Wa ter W. Homes, Field Associate in Paleontology, The Ameri- 
can Museum of Natural History, New York City. 

ArtTHuUR TINDELL Hopwoop, Assistant Keeper, Department of 
Geology, British Museum (Natural History), London, 

Wrtu1AM Treme_e Hornapay, formerly Director of the New York 
Zoological Park, New York City. 

*Sir Harry H. Jonnsron, Poling, Arundel, England. 

A. C. pe Jonau, Director, Geological Survey of Dutch East Indies, 
Bandoeng, Java. 

H. Munniks pe Joneau, The Hague, Netherlands. 

GrorGE FrepericK Kay, Head, Department of Geology, Univer- 
sity of Iowa, also State Geologist of lowa, lowa City, Iowa. 

Hans Kuann, Professor of Geology, Universitat, Rostock, Ger- 

Ernst C. Kroun, First Vice-President of the Greater St. Louis 
Museum of Natural History, St. Louis, Missouri. 

*GroRGE FREDERICK Kunz, Research Associate in Gems, The 
American Museum of Natural History, and Vice-Pres- 
ident, Tiffany and Company, New York City. 

Gustav KisTHarpt, Inspector, Zoologische Staatssammlung, 
Munich, Germany. 

Francois ANToINE ALFRED LAcROIX, Secrétaire perpétuel pour 
les sciences physiques de |’ Académie des Sciences, Institut 
de France; Professor of Mineralogy, Muséum National 
d’Histoire Naturelle, Paris, France. 

Hersert LANG, The Transvaal Museum, Pretoria, South Africa. 
Wiiuram Dixon Lana, Keeper, Department of Geology, British 
Museum (Natural History), London, England. 

Louis Laurent, Director-Curator, Muséum d’Histoire Naturelle, 
Marseille, France. 

Frepreric BrewsTER Loomis, Professor of Geology, Amherst 
College, Amherst, Massachusetts. 

Ernar LOnNBERG, Naturhistoriska Riksmuseum, Stockholm, 

*PrepeRIC AuGusrus Lucas, Honorary Director, formerly 
Director, The American Museum of Natural History 
New York City. 

RicHarp Swann LULL, Sterling Professor of Vertebrate Pala- 
ontology, Director of Peabody Museum, Yale University, 
New Haven, Connecticut. 

Frans Henpprik VAN DER Maaret, De Bilt, Netherlands. 

*Paut Marscure, Museum fiir Naturkunde, Berlin, Germany. 

Hikosuicuir6 Marsumoro, Professor of Paleontology, Tohoku 
Imperial University, Sendai, Japan. 


*WittiaM DititerR Marruew, Professor of Palzontology and 
Director of the Museum of Palzontology, University of 
California, Berkeley, California, formerly Curator of 
Geology and Paleontology, The American Museum of 
Natural History, New York City. 

Lucien Mayet, Professor of Anthropology, Université de Lyon, 

JoHn CampBELL Merriam, President, Carnegie Institution of 
Washington, Washington, D. C. 

Eimer Drew Merritu, Director-in-Chief, New York Botanical 
Garden, New York City. 

*GreorGE Perkins Merritt, Head Curator of Geology, United 
States National Museum, Washington, D. C. 

Gerrit SmitH MiItier, Jr., Curator of Mammals, United States 
National Museum, Washington, D. C. 

*Roy Ler Moony, Professor of Paleodontology, College of Den- 
tistry, University of Southern California, Los Angeles, 

CuarLEs CratG Mook, Associate Curator of Geology and Palzon- 
tology, The American Museum of Natural History, 
New York City. 

ALFRED T. Nester, formerly Consul of the United States of Ameri- 
ca at Naples, Italy. 

W. F. F. Oppenoorts, Bandoeng, Java. 

Raymonp C. Ossurn, Department of Geology, Ohio State 
University, Columbus, Ohio. 

THEODORE SHERMAN Paumer, United States Biological Survey, 
Washington, D. C. 

Marie Paviow, Professor, Université de Moscow, Institut 
Géologique, Leningrad, Union of the Socialist Soviet 

*OLtor Aucusr Prrerson, Curator, Mammalian Paleontology, 
Carnegie Museum, Pittsburgh, Pennsylvania. 

Guy Exucock Pinerm, formerly Superintendent of the Geological 
Survey of India, Calcutta, India. 

Hans Pou.iec, formerly of the Universitit, Bonn, Germany. 

CuEsTeR ALBERT ReeEDs, Curator of Geology and Invertebrate 
Paleontology, The American Musum of Natural His- 
tory, New York City. 

GraHAM RENSHAW, Sale, Manchester, England 

Juutes JosepH Repevrn, Curator, Muséum d’Histoire Naturelle, 
Marseille, France. 

Pierre Revituiop, Director, Musée d’Histoire Naturelle de la 
Ville de Genéve, Switzerland. 

Aurcia E. Reyes, Assistant Curator, Museo de Historia Natural, 
Mexico City, Mexico. 

Eimer SamuEt Riaas, Associate Curator of Paleontology, Field 
Museum of Natural History, Chicago, Illinois. 

GUNTHER ScHLESINGER, Director, Niederésterreichische Landes- 
sammlungen, Vienna, Austria. 

*Max Scuuosser, Bayerische Staatssammlung fiir Paliontologie, 
Munich, Germany. 

C. Brrtranp Scuuttz, State Museum, University of Nebraska, 
Lincoln, Nebraska. 

Ex1as Howarp SE.iarps, Associate Director, Bureau Economic 
Geology, Professor of Geology, University of Texas, 
Austin, Texas. 

CHARLES Davies SHERBORN, Bibliographer, British Museum 
(Natural History), London, England. 

GrorRGE GAYLORD Simpson, Associate Curator of Vertebrate Pale- 
ontology, The American Museum of Natural History, 
New York City. 

WoLFGANG SoprGE.L, Professor, Geology and Paleontology, and 
Director, Geologisch-paliont. Institut und Museum, 
Schlesische Friedrich-Wilhelms-Universitéit, Breslau, 

FRANZ SPILLMANN, Director, Cabinete Zool6gico, Museo, Universi- 
dad Central del Ecuador, Quito, Ecuador. 

*GreGORIU STEFANESCU, Professor of Paleontology, University 
of Bucharest, Rumania. 

SaBBa STerANescu, University of Bucharest, Rumania. 

Hans GerorG, President, Naturhistorisches Museum, 
Basel, Switzerland. 

R. A. Strrton, Department of Geological Sciences, University of 
California, Berkeley, California. 

Cuester Srock, California Institute of Technology, Pasadena, 

Witmer Stone, Director Emeritus, Museum of the Academy of 
Natural Sciences, Philadelphia, Pennsylvania. 

W. E. Swinton, British Museum (Natural History), London, 

Prerre TEILHARD DE CHARDIN, National Geological Survey of 
China, Peiping, China. 

*MicuarL Rocers OLDFIELD THOMAS, British Museum (Natural 
History), London, England. 

INNOKENTY PavLovicH TotmacHorFr. Professor of Paleontology, 
University of Pittsburgh, Pittsburgh, Pennsylvania. 

Epwarp LEFFINGWELL TROXELL, Professor of Geology, Trinity 
College, Hartford, Connecticut; formerly Research 
Associate in Paleontology at Yale University, New Haven, 

G. M. Vevers, Superintendent of the Gardens, Zoological Society 
of London, England. 

Daviy MerepiTH Seares Watson, Jodrell Professor of Zoology 
and Comparative Anatomy, University of London, 

Max Wesker, formerly Director, Zoologisch Institut en Museum, 
Amsterdam, Netherlands. 

CoLEMAN SHALER WiLiiAMs, Department of Vertebrate Pale- 
ontology, The American Museum of Natural History, 
New York City. 

M. Wiiman, Curator, McGregor Museum, Kimberley, South 

ArtTHuR SmitH Woopwarpb, formerly Keeper of Geology, British 
Museum (Natural History), London, England. 

E. D. Van Oort, Director, Rijks-Museum van Natuurlijke His- 
torie, Leiden, Netherlands. 

Hisakatsu YABE, Professor of Geology and Paleontology, Tohoku 
Imperial University, Sendai, Japan. 

NaouipE YATSU, Professor of Zoology, Tokyo Imperial University, 
Tokyo, Japan. 

The outstanding names of the collaborators in the Volume include several great and highly esteemed col- 
leagues who have fallen by the way, such as Charles Depéret of Lyons, William D. Matthew, Joel A. Allen, 
Frederic A. Lucas and Carl E. Akeley of the American Museum, James W. Gidley of the U.S. National Museum, 
Paul Matschie of the Natural History Museum of Berlin, John Mason Clarke of the New York State Museum, 
and finally, but not least, Charles W. Andrews of the British Museum. These names, indicated on the Honor Roll 
by a star, are included in the above list of those who have coéperated more or less extensively with the author 

in the present work. 

All figures one-hundredth natural size, excepting 14 which is one thirty-fourth natural size. 

1,14 Meritherium lyonsi Andrews. 
2  Phiomia osborni Matsumoto. 
3. Trilophodon angustidens Cuvier. 
4 Platybelodon grangert Osborn. 
5  Amebelodon fricki Barbour. 
6  Serridentinus serridens Cope. 
7  Tetralophodon punjabiensis Lydekker. 
8  Palxomastodon beadnelli Andrews. 
9 Mastodon americanus Kerr. 
10 Rhynchotherium tlascale Osborn. 

11  Deinotherium gigantissimum Stefinescu. 
12 Anancus arvernensis Croizet and Jobert. 

13 Synconolophus dhokpathanensis Osborn. 
14 Stegomastodon arizone Gidley. 
15  Notiomastodon argentinus Ameghino. 



won = 


a ns 



Eubelodon morrilli Barbour. 
Cordillerion andium Cuvier. 
Cuvieronius superbus Ameghino. 
Stegodon ganesa Falconer. 

Stegodon trigonocephalus Martin. 
Archidiskodon imperator Leidy. 
Parelephas jeffersonii Osborn. 
Mammonteus primigenius Blumenbach. 
Hesperoloxodon antiquus italicus Osborn. 
Palzoloxodon falconeri Leith Adams. 
Palzoloxodon namadicus Falconer. 
Elephas indicus Linnzus. 

Loxodonta africana Blumenbach. 
Loxodonta africana pumilio Noack. 









. Principles of the Osborn Phylogenetic Classification. 
. Origin of the Proboscidea. 
. The primary stocks established by primary choice of 

food in different habitats. 




. Principle of adaptive radiation expressed in the phylo- 
genetic classification of the Fayim Proboscidea. 
Amphibious habits of Meritheriwm. 

3. Lowland habits of Phiomia, ancestor of the Amebelodont- 


4. Ordinal, subordinal-superfamily, family, subfamily-phy- 
letic, generic-phyletic characters employed in the phylo- 4. Forest-loving habits of Palzomastodon, not ancestral to 
genetic classification of the Proboscidea. Mastodon. 

5. Harmony of Linnean and phylogenetic classification ex- IV. Rrviston AND SyNopsis OF GENERA, SPECIES, AND TYPES OF 

pressing our knowledge up to the close of the year 1933. 

II. Ossporn’s THErory (1900) as TO THE AFRICAN ORIGIN OR 


Andrews confirms the African origin of the Proboscidea, 
Sirenia, and Hyracoidea. 

. Palzomastodon beadnelli Andrews, 1901. 
. Merritherium lyonsi Andrews, 1901. 
. Phiomia serridens Andrews and Beadnell, 1902 
Meritherium gracile Andrews, 1902. 

2. American Museum Faytim Expedition of 1907. Interpre- 
tation and discovery bearing on the phylogeny of the 
Fayfm Proboscidea. . Palzomastodon parvus Andrews, 1905. 

3. Relations of Meritherium, Palzomastodon, and Phiomia . Phiomia wintoni Andrews, 1905. 
to each other and to other mammals. 9. Phiomia barroisi Pontier, 1907. 

4. Matsumoto’s complete separation of Meritherium, Palzxo- 10. Meritherium andrewsi Schlosser, 1911. 
mastodon, and Phiomia. 11. Palzomastodon intermedius Matsumoto, 1922 

5. Africa demonstrated as the homeland of the three families 12. Phiomia osborni Matsumoto, 1922. 
Meeritheriidz, Mastodontidx, and Bunomastodontide. . Meritherium ancestrale Petronievies, 1923. 

. Meritherium trigodon Andrews, 1904. 
. Phiomia minor Andrews, 1904. 

ONOA Pwd eS 


CLASSIFICATION.—The classification of the Proboscidea progresses with our ever increasing knowledge of 
origins, adaptive radiations, and phyletic successions. Consequently classification is progressively modified by 
discovery and research. The classification in the present Memoir attempts to set forth our general knowledge of 
the evolution of the Proboscidea up to the end of the year 1933; the nomenclature, that is, lists of genera, species, 
etc., chiefly terminates at the close of the year 1933, because it is impossible to include all the new specific and 
generic lists of 1934 up to the moment of going to press. The classification of 1935 concludes the present Volume. 

Our rules of nomenclature will be a shock to many zoologists, especially to those who carry the principle of 
priority to the point of adopting misspelled, misapplied, and practically unpublished names. After the most 
honest and painstaking endeavor, we have found it impracticable to apply uniformly all the principles established 
in zoology to nomenclature in paleontology. The fundamental reason is that the imperfect data of many original 



descriptions in paleontology are in wide contrast to the usually perfect data of zoology. The art of nomenclature 
is entirely different in these two branches. Consequently the rules of nomenclature framed by zoologists will 
inevitably be different in certain respects from those which can be adopted by paleontologists. Certain of the 
reasons for this statement will become clear in the following sections. 

OricIn.—Our present knowledge of the origin of the Proboscidea begins with the wonderfully varied and 
interesting forms discovered in the Fayam region of North Africa in 1901 and successively examined by Andrews, 
Schlosser, Matsumoto, Petronievics, and Osborn between 1901 and 1923. In the systematic revision all of these 
Faytim animals will have to be considered together; they are therefore treated in chronological order of descrip- 
tion in the present chapter, although it isnow known that they belong to at least three distinct phyla. After being 
thus separated phyletically, they are again treated in succeeding chapters in their true phylogenetic order, namely: 

Palzomastodon, forest loving, the possible phyletic ancestor of the Mastodontide. 
Phiomia, shore loving, certainly the collective ancestor of the shovel-tusker A mebelodon. 
Meritherium, water loving, standing and ending in a phylum by itself. 

FLoop-PLAIN SCENE OF THE ANCIENT River Nive, Nortx Arrica, Iv Lower OLIGocENE TIME 
ResTORATION BY MarGrer Fiinscu (1932), UNDER THE Direction oF HENRY FarRFIELD OSBORN 

Fig. 2. Theoretic restoration of Palzomastodon beadnelli (upper left), Phiomia osborni (upper right), and Meritherium andrewsi 
(upper center), all one-fiftieth natural size. Meritheriwm lyonsi in foreground one-thirtieth natural size. 


LINN#AN CLASSIFICATION (Linnzus, 1735-1766) aimed to define the apparent relationships and affinities of 
existing animals as they were created and spread contemporaneously on the present surface of the earth, and as 
revealed by the comparative anatomy and zoology of existing animals. The “‘Systema Nature’’ of Linnzeus is 
zoological and creational. 

PHYLOGENETIC CLASSIFICATION (Osborn, 1892-1933) aims to distinguish ties of blood and descent among 
ancestral as well as existing mammals; it also aims to adapt and modify the Linnzan classification to the dis- 
coveries of paleontology, namely, to include animals belonging to phyla or ancestral lines and to connect them with 
existing animals. It expresses the nearness or the remoteness of phyla of all degrees of magnitude, namely, phyla 
of species, genera, subfamilies, families, orders. The term PHyLum has a larger taxonomic significance, e.g., 
Phylum Chordata=animals with a notochord. 

The word phylogenetic signifies the origin or genesis of lines of descent of ancestral animals unknown to Lin- 
neus. The word phylum is derived from the Greek (ef. Liddell and Scott, ““A Greek-English Lexicon,” 1883, 

pp. 1698, 1703): “@vAn, 7 (diw) properly, like @d)\ov, a set of men naturally distinct; . . . In usage @vA7 corre- 
sponded to the Roman tribus, and signified I. a body of men united: 1. by supposed ties of blood and descent, 
aclan or sept, such as those among the Dorians, . . . or 2. by local habitation,” (p. 1698). ‘‘diw, . .. A. trans., 
. . . tobring forth, produce, put forth, ¢b\\a. . . . B. Pass., . . . to grow, wax, spring up or forth, arise, come into 

being,” (p. 1703). 


In 1891, Osborn began to apply a system of phylogenetic classification to the order Perissodactyla (1892.67) 
and developed it in subsequent years especially in treating the family Rhinocerotide (1900.192) and finally 
the superfamily Titanotherioidea (1929). The purpose of this classification of the Proboscidea is to express 
our present knowledge of the more or less profound gaps between the greater and the lesser branches. Most of the 
older classifications were based on the idea that there were but two genera, Elephas and Mastodon. Cope (1889.2) 
divided the Proboscidea into many genera but did not propose a phylogenetic classification. Gaudry (1862-1878), 
Leith Adams (1877-1881), Pohlig (1888-1909), Weithofer (1890), Pilgrim (1905-1926), Soergel (1912-1921), 
Lull (1908), Matthew (1915), Schlesinger (1917-1922), Boule (1920), and Freudenberg (1922) have successively 
discussed and enriched our knowledge of the phylogeny of the Proboscidea but have not expressed this knowledge 
in classification. 

Osborn (1907) began his observations upon the Proboscidea and his survey of the literature covering the years 
1735 to 1934; in 1918 (1918.468), he began the special series of studies on classification as expressive of the 
phylogeny of the Proboscidea which has culminated in the present Memoir. 

Osborn, 1934: The advantage of a phylogenetic classification of the Proboscidea is that the superfamilies or 
suborders expressed with the terminal -orpkEa indicate the four’ larger branches into which the Proboscidea are now 
known to be divided; the family subdivisions corresponding with the terminal -1p# express the five’ known subdivi- 
sions, leaving space for the unknown families; the third system of branching, expressed with the subfamily terminal 
-INE, indicates the seventeen' subfamilies, three of which are now positively known to be as ancient as Lower 

'(Osborn, 1935) Five larger branches (-o1pkE), eight families (-1p22), and twenty subfamilies (1N2) are now known. See Appendix of the present Volume. 



Oligocene time; beneath these branches are the forty-one or more generic phyla, each of which embraces a very 
long line of specific and subspecific descent. This phylogeny of 1933-1934 is shown in figure 8. 

The chief difficulty with a phylogenetic classification of the Proboscidea, as of the Perissodactyla or other 
orders of mammals, is a technical one, namely, to harmonize it, on the one hand, with the Linnzan “special 
creation’”’ system of nomenclature, and, on the other, with the prevailing usage of family, subfamily, and generic 
nomenclature in mammalian zoology, which systems are alike based on affinities displayed between the existing 
terminal twigs of the branches and subbranches rather than on the affinities of the phylogenetic lines which connect 
the existing twigs with their geologically remote and fundamental ancestral branches. By Linnzan and post- 
Linnzean systematists upwards of fifty’ generic names have been applied to the mastodonts and elephants, but there 
is no uniformity in the application of these generic names among paleontologists or zoologists, nor has any other 
principle of revision or of arrangement been worked out by which one may at least begin an advance toward a 

permanent system of nomenclature of this highly important and interesting group, except the present phylogenetic 

African Elepha nt Indian Elephant 
f 5 VA | 
S Z. 
ie hata peti: 3 = PLEISTOCENE 
WN Veo vy 3 ‘ PLIOCENE 
Py Cav ioe 
a7 (O/ lear 
Y ae 
x | eA 
ay 14 MIOCENE 
/ / 
We : i 
/, / 
Mfastodontsl Elepha, 
y, Rete becky yf OLIGOCENE 
\ ved CRAZEPp 7 
Stock ie ay 
é a Forests and Forests Grassy FOCENE 
mph No 22s 

Savannas Plains, Savannas, Steppes. 

& palustNgl 

Fig. 3. Diagram showing Osborn’s theory so far as developed up to June, 1921, as to the Adaptive Radiation of the Proboscidea. 
After Osborn, 1921.532, p. 233, Fig. 1. Compare diagrams of 1925, 1933-1934, and 1935 below. 

Apart from its theoretic value there is an honest difference of opinion about the practicability of a phylo- 
genetic system of classification; some of the leading paleontologists, like W. D. Matthew (letters, February, 1918), 
do not think it is practicable; other colleagues, like W. K. Gregory (letters, February, 1918), do not consider that 
the subfamily division (-rwa) can be properly applied to vertical lines of succession. But to Osborn there does not 

'This estimate was made in 1923; at the close of the year 1933 upwards of eighty genera had been listed by the present author (see Chap. I, p. 9); also 
for complete list of genera and species, see Volume II. 


appear to be any alternative between the continuation of the hopeless confusion which at present prevails in the 
nomenclature and arrangement of the Proboscidea and the absolute illumination and clarity which are thrown 
upon the whole subject by the adoption of the phylogenetic classification. 

In our present phylogenetic revision the only possible way of straightening out the genera is by first straighten- 
ing out the phyla in which they belong. The same is true inregard to the species; the species can be understood only 
in their phyletic relations. Consequently the prelude to our phylogenetic classification has been the prolonged 
and very difficult research on the actual characters of the types of genera and of species, upon which the author 
has been closely engaged for the last sixteen years (1918-1934). During this period the author has learned a great 
deal about the phylogeny of the Proboscidea from other observers and from his own observations, but many 
phyla still remain obscure and imperfectly known; much is to be added to our knowledge by the more intensive 
study of materials already in hand and of the very rich materials still to be discovered, especially in central and 
northern Asia and northern Africa. 

Thus a phylogenetic classification is never final; it is always progressive and adaptive to future discovery. 
Osborn’s classifications of 1921 to 1926 appear to be marked advances upon that of 1918; doubtless it will be 
progressively and perhaps largely modified through future discovery. This is as it should be, because nomenclature 
and classification are the tools of thought, not the masters of thought as some systematists would have it. Other 
phylogenies are those of 1925 (Fig. 7) and of 1933-1934 (Fig. 8) bringing us up to 1935, see the Appendix of 
this Volume. 

With a reasonable rather than a strained regard for priority, nomenclature and classification are the plastic 
vehicles of expression of the actual knowledge of relationships and phylogeny. A reasonable recognition of all the 
great ideas of the past combined with provision for the ideas and the discoveries of the present and of the future 
—these are the watchwords of Phylogenetic Classification. 


As remarked above, phylogenetic classification depends progressively upon the discovery of origins, of 
adaptive radiations, and of phyletic successions. 

The discovery of primary stocks by paleontologists naturally comes last, long after the zoologists have dis- 
covered recent or existing species. Thus the existing elephants have been known by man for thousands of years, 
but the three primary stocks from which they were derived were not discovered until 1901. These primary stocks 
give us the roots and greater branches of our system; the lesser branches have afforded the progressive knowledge 
of adaptive radiation and the still more elusive knowledge of direct phyletic succession. 

The distinguishing feature of this Memoir, which we trust will be of permanent value, is the separation of 
many direct phyletic lines of descent from each other and the recognition of many mastodonts and elephants 
which are very much alike in certain characters, but which are still more unlike in other characters and cannot 
possibly be descended one from the other. 


If we watch ungulates while feeding, we observe that they are very fastidious in the choice of food. Conse- 
quently the beginning of adaptive radiation lies in the choice of food and of feeding habitat. There is a great deal 
of truth in Lamarck’s idea that the wish precedes the organ. Coadaptation and compensation follow the primary 
choice of food. 

The origin and evolution of the three kinds of Fayim proboscideans as described below (p. 34 to p. 50) 
are only part of the evidence for a long continued antecedent adaptive radiation process beginning in the Upper 
Cretaceous and extending throughout Eocene time, by which the Proboscidea, like other hoofed animals, were 
broken up into several great primary stocks, namely: 

An amphibious stock, adapted to rivers and swamps, of limited powers of migration. Represented only 
by the imperfectly known Meritherium of northern Africa and possibly (Pilgrim) of southern 
Asia in Oligocene time =MC@RITHERIOIDEA 

A southern forest stock adapted to forested lowlands. Represented by the Deinotherium of northern 
and central Africa and of southern Eurasia; known from more abundant remains in Miocene 
to Middle Pliocene time = DEINOTHERIOIDEA 

A northern stock, adapted to lowlands, to savannas, and to forests, with better developed limbs and 
powers of wide migration. Represented from the Lower Oligocene of northern Africa to the Mio- 
cene-Upper Pleistocene of Eurasia and North and South America =MASTODONTOIDEA 

A stegodont-elephant stock, adapted to plains, savannas, and steppes; of browsing (forest) or grazing 
(plains) habits, also with powers of wide migration. Represented from the Lower Pliocene of 
southern Asia to the Pleistocene of North America and the Pleistocene and Recent of southern 
Asia and of Africa’ =ELEPHANTOIDEA”’ 

These four? great primary stocks of Eocene time, and probably other stocks still to be discovered, gave off 
from two to six branches each, so that the Proboscidea, as a whole, from Upper Eocene time onward are not two 
branched, or diphyletic, as formerly expressed in the adoption of only two genera, Elephas and Mastodon, but many 
branched or polyphyletic. Species of Mceritheres and Deinotheres remained in Africa and southern Eurasia; 
whereas the mastodonts and elephants, forest and savanna browsers and grazers of the plains, steppes, and for- 
ests, were the long distance travelers and either from an African or from an Asiatic homeland center of origin 
and adaptive radiation in Eocene time reached all the continents of the world, except Australia, in Miocene and 
Pliocene times. 

Always intelligent, resourceful, independent, adaptive, well defended from their enemies of all kinds, we know 
at present of seventeen® entirely distinct main branches of proboscideans several of which persisted into Upper 
Pliocene if not into Pleistocene time, when they were overtaken by the worldwide elimination of the larger 
Mammalia. Only two branches, namely, the Loxodontines and the Elephantines, survive at the present time. 

'(Osborn, 1934) Plio-Pleistocene ancestors of Archidiskodon and Palzoloxodon recently discovered in South Africa (see p. 25). 
*(Osborn, 1935) The STEGODONTOIDEA superfam. nov. separated from the Elephantoidea as a distinet stock. 
8(1935) See footnotes on pages 27 and 30. 



The OrpER PROBOSCIDEA is distinguished by the following characters common to all its members: (1) A 
pair of second upper and lower incisors greatly enlarged and originally opposed as feeding organs, a function 
which resulted in the loss of both the first and third pairs of incisors and in the hypertrophy of the second pair, 
I’-I,; (2) originally tetrabunodont to hexabunodont superior and inferior grinding teeth; (3) originally penta- 
dactyl feet with subequal distribution of weight on all five digits. 

THE ProBoscippa. AFTER OsBorN, 1921.515, ria. 1, Pp. 3. 

I. Meritherium type (MG2RITHERIOIDEA): Median incisors, I'-I,, persisting; 
second incisors, I?-Is, opposed, as in the Rodentia, front encased in enamel. 

II. Deinotherium type (DEINOTHERIOIDEA): Superior incisors entirely 
aborted; inferior incisors (probably I2) downturned and retroverted. 

III. Rhynchotherium type (MASTODONTOIDEA): I-Ie, tusks downturned, 
laterally compressed, partly covered with enamel band (e). 

IV. Elephas type (ELEPHANTOIDEA), including Elephants, Loxodonts, Mam- 
moths, ete., with inferior incisors, Ig, reduced or aborted, superior incisors, I”, gradually 
losing enamel, typically upturned, used both as fighting and as feeding organs. 

On this primary ordinal constitution the Proboscidea radiated adaptively into all habitats known to the her- 
bivorous quadrupeds, namely, (a) fluviatile, lowland, swamp, and forest, (b) forest border, (c) plains and upland 
habitats, with their accompanying vegetation, softer and harder kinds of food, and reactions of the feet and limbs 
to the modes of locomotion corresponding to habitats a, b, c. 


SuPERFAMILY Ranx.—As regards the subordinal separation or rank of thefour’ primary divisions of the Probo- 
scidea which have been discovered up to the present time, these divisions are certainly far more profound than 
those of the four suborders of the Rodentia, namely: I. ScturomorpHA, II. Myomorpua, III. HysrricomorpuHa, 
IV. LacomorpnHa, which are separated from each other not by modifications in the paired incisor, I?-I,, function, 
but by fairly profound muscular adaptations in the mechanism of the jaws. The four primary divisions of the 
Proboscidea are also far more profound than Osborn’s main divisions of the Perissodactyla, namely: I. Trrano- 
THERIOIDEA, IJ. HipporpeEa, III. Tarrromea, IV. Ruryocerororpea, which are distinguished by profound differ- 
ences in the fundamental pattern of the grinding teeth and by adaptive divergence in the original tetradactyl 
manus and pes. 

'See footnotes on pages 19 and 22. 



ELEPHANTOIDEA are separated from each other by adaptive divergences in the function of the upper and 
lower incisor teeth which exceed in their coadaptive results those in the subordinal divisions of any other order 
of ungulates. If the Mceritheres and Deinotheres had lived in the time of Linneus, it is doubtful whether he 
would have united them in the same order with the mastodonts and elephants which are of more obvious affinity 
to each other. 

I. MGRITHERIOIDEA.—Renewed study of Meritherium by Osborn and Matsumoto entirely confirms 
Andrews’ original opinion that Meritherium belongs in the order Proboscidea, as well as Osborn’s opinion that it 
stands very far apart from the other proboscideans and is not directly or indirectly ancestral to either of the other 
three groups. The enlargement of the second upper and lower incisor teeth into mutually abrading tusks, girdled 
with enamel, presents a firm ground of affinity with a still unknown primitive Lower Eocene proboscidean stem 
form. There the resemblance ends. These Meeritheres had no proboscis. The face, of brachyopic type, is 
markedly abbreviated. The cranium is elongated. Thus the facial and cranial proportions are analogous to 
those of the Sirenia. The upper grinding teeth are bilophodont, pointing to a tetrabunodont ancestry, and 
different from the bunomastodont grinders of Palzomastodon, which point to a hexabunodont ancestry. The 
lower grinders exhibit a rudimentary third crest. Andrews’ opinion that the Moeritheres were amphibious is 
probably correct. 

II. DEINOTHERIOIDEA.—All agree that these animals were chiefly browsers and partly amphibious in 
habit, in this respect resembling the Mceritheres but differing in the entire loss of the superior incisor teeth. 
Early loss of the superior tusks released the inferior. In the downturning of the inferior tusks the Deinotheres are 
paralleled by the Rhynchorostrine among the Mastodontoidea. In skull form and in limb and foot structure 
the Deinotheres parallel the true proboscideans. They diverge very widely from proboscideans in the evolution 
of the upper and lower grinding teeth. The primitive Deinotheres present simple, bilophodont grinders, similar to 
those of Meritherium, and are progressively trilophodont. The upper grinders attain a stage which parallels the 
molar pattern of the tapir (T’apirus) among the perissodactyls, but which shows a tendency to the trilophodont 
structure characteristic of the primitive mastodonts and elephants. 

III. MASTODONTOIDEA.—The fundamental character of the front teeth in this superfamily is seen in 
primitive members of the Rhynchorostrine and Longirostrine, namely: (1) Second superior incisors enlarged, 
downturned, divergent, with enamel band on outer side only; (2) second inferior incisors downturned (as 
in Deinotheres) with enamel band on outer side (Rhynchorostrine) or procumbent with no enamel band 

The important functional distinction of the Mastodontoidea is that for a very long period of time the upper 
tusks abraded the outer side of the lower tusks; this probably explains the retention of the superior enamel band. 
In certain lines (Longirostrinz) the procumbent lower incisors persist and the upper incisors retain their primitive 
downcurved position as in Phiomia. In other lines (Mastodontine, Brevirostrine) the lower incisors practically 
cease to function; the upper incisors finally turn upward and inward, but may retain the enamel band for a 
long period (Mastodontine, fide Schlesinger, and Notorostrine, vide Cordillerion andium). 

A distinctive character of the grinding teeth of the Mastodontoidea is evidence of the descent from a 
tetrabunodont ancestral grinder (i. e., without intermediate tubercles or ‘conules’), as is the case with the tetra- 
bunodont ancestral type of Mceritheres and Deinotheres. The rudiments of ancestral conules gave rise to various 


trefoils or paired median outgrowths or crests, so characteristic of all the Bunomastodontide whether beak jawed 
(Rhynchorostrinz), or long jawed (Longirostrinz), or short jawed (Brevirostrine). In each of these subfamilies 
the grinders independently undergo a more or less closely parallel evolution, evolving single trefoils in Upper 
Oligocene and in Miocene time, and double trefoils in Pliocene time. 

Unlike the Mceritheres and Deinotheres, the three intermediate molars (i. e., fourth premolar and first and 
second true molars) invariably become trilophodont, while the third true molars become tetralophodont. At this 
point there is a divergence into (1) Mastodontidx, purely forest living, brachyodont, with simply crested teeth, 
in which the intermediate molars are persistently trilophodont, with arrested trefoils, and into (2) Bunomastodon- 
tide, which pass into tetralophodonty and polylophodonty in some lines, with evolving trefoils. The grinder 
evolution is adapted to a leaf-browsing habit, in distinction to the prevailing grazing habit developed among the 
elephantoids. The development of hypsodonty, and choerodonty (Schlesinger), among these (longirostrine and 
brevirostrine) browsers is analogous to that in the hippopotami and the hypsodont suillines. 

IV. ELEPHANTOIDEA.—One prime distinction in this superfamily is the very early complete loss of the 
lower incisor teeth, accompanied by the early development of the upper incisors into horizontal or upturned tusks 
finally devoid of enamel except at the tips in the young stage. _ Vestigial enamel bands are recorded in early stages 
of the Stegodonts. A second distinctive character is the absence of conule development into trefoils, so character- 
istic of the mastodontoids, and the early tendency to form evenly transverse, more or less mamillate, crests which 
become in the highest degree hypsodont and polylophodont in adaptation to chiefly grazing habits. 

It has been assumed by practically all paleontologists that the Elephants were descended from the Stegodonts. 
This assumption now proves to be erroneous, for neither the Stegodon grinding tooth with enamel valleys closed at 
the bottom, nor the Stegodon cranium with its extremely short face, can give rise to the elephantoid molar or the 
face of the elephantoid cranium. The earliest elephants known (1934) are the primitive species of Archidiskodon 
and of Palxoloxodon of the Plio-Pleistocene of South Africa. The Stegodonts were of independent origin and 
formed an independent parallel branch terminating in the highly specialized Elephas |= Stegodon| aurore Matsu- 
moto from the Upper(?) Pliocene of Mt. Tomuro, Japan, now (1935) separated by Osborn as the STEGODONT- 
OIDEA superfam. nov. 

Fig. 5. (1) Meeritherioidea, one-twelfth natural size. (2) Deinotherioidea, (3) Mastodontoidea, and (4) Elephantoidea, all one-fiftieth natural size 


Only one family up to recent time’ has been proposed in three of the four’ suborders or superfamilies of the 
Proboscidea. In the Mastodontoidea there are two? families, the Mastodontide and the Bunomastodontide. 

I. MGRITHERIOIDEA Osborn, 1921.515, p. 2. Family: Morritheriide Andrews, 1906, p. 99. 

Il. DINOTHERIOIDEA Osborn, 1921.515, p. 2 [=Deinotherioidea of the present Memoir]. Family: 
Curtognati [=Curtognathide] Kaup, 1833.4, p. 516; Dinotheridz Bonaparte, 1845 (fide Palmer, 
1904, p. 738), Dinotheriide Bonaparte, 1850; Dinotheride Zittel, 1891-1893, p. 454; Dino- 
theriidze Osborn, 1918.468, p. 134 [=Curtognathide of the present Memoir]. 

Notr.—The superfamily Deinotherioidea as well as the family Deinotheriide are partly anticipated in the 
term Curtognati Kaup (Kaup, Bull. Soe. Géol. de France, V, 1834, p. 444): “Curtognati—M. Kaup a regu de 
nouveaux ossemens d’Eppelsheim, et a reconnu que le Dinotherium medium était véritablement bien différent du 
D. gigantewum. M. Kaup a pu faire aussi quelques corrections et additions 4 sa description du Dinotherium. Il 
pense que ses deux énormes défenses ne lui servaient pas seulement pour extraire des racines de la terre, mais 
encore, comme 4 la morse, pour l’aider A mouvoir son corps silourd. D’aprés la forme des os intermaxillaires, cet 
animal devait avoir une trompe pour porter la nourriture Asa bouche. I] est 4 placer entre les Mastodontes et les 
Bradypus, et il formerait une famille particuliére 4 laquelle M. Kaup donne le nom de Curtognati. Elle serait 
caractérisée par la machoire inférieure courbée vers le bas, et les deux défenses dirigées vers le bas et en arriére 
(N. Jahrb. f. Mineral. Geognosie, etc., 1833 [1833.4], cah. 5, p. 509 avec 1 pl.).” 

II]. MASTODONTOIDEA Osborn, 1921.515, p.2. Family: Mastodonade Gray, 1821, p. 306 (fide Palmer 
1904, p. 752); Mastodontide Girard, 1852, pp. 326, 328; Mastodontidze Osborn, 1918.468, p. 
134, including the Paleomastodontide of Andrews, 1906, p. 130. 

Osborn has recently added the following family: Bunomastodontide, 1921.515, p. 2, to 
include (1921-1922) all the bunomastodonts which develop the intermediate conules into, first, 
single and, then, double trefoils; also a very important branch with several side branches of mas- 
todonts in which the lophs are transformed into well-defined transverse crests paralleling the 
zygolophodont condition of the Stegodonts, and perhaps giving off a branch into the Stegodon- 
tine. The Bunomastodontide constitute a very distinct family, from which arose several sub- 
families also widely separated from each other. 

IV. ELEPHANTOIDEA Osborn, 1921.515, p. 2. Family: Elephantide Gray, 1821, p. 305 (fide Palmer, 
1904), p. 740 (=Elephasideze Lesson, 1842, p. 156; Elephantide Bonaparte, 1838, p.112, and 
1850; Girard, 1852; Zittel, 1891, p. 458; Osborn, 1910.346, p. 558). 

The above five families of the Proboscidea are distinguished from each other by profound modifications 
in the fundamental structure of the grinding teeth. These differences in fundamental structure are almost as 
profound as those which separate the eight families of the order Perissodactyla, or the seven superfamilies of the 

It is noteworthy that the giant Proboscidea closely parallel the humble Rodentia in the profound modes of 
evolution of the grinding teeth. 

‘See footnotes on pages 19 and 22. 
2(Osborn, 1934) To these two families have been added the Serridentide fam. nov. and the Humboldtide fam. nov. 


The following seventeen‘ subfamily divisions of the Proboscidea have successively been proposed, as ex- 
plained in full chronological detail in Chapters II-X X of the present Memoir: 

I. MGRITHERIOIDEA: 1. Meeritheriini—Herluf Winge, 1906, p. 172; Meoeritheriine Winge- 
: Osborn, 1923.601, p. 1. 
Il. DINOTHERIOIDEA: 2. Dinotherina—Bonaparte, 1841, p. 253, Dinotheriina—Bonaparte, 1850; 

Dinotheriini—Herluf Winge, 1906, p. 172; Dinotheriine—Osborn, 
1910.346, p. 558 [= Deinotheriine of the present Memoir]. 

III. MASTODONTOIDEA: 3. Mastodontina—Brandt, 1869, p. 35; Mastodontinze—Osborn, 1910.346, 
p. 558. 

Bunomastodontinz—Osborn, 1918.468, p. 134. Replaced by Buno- 
mastodontidae—Osborn, 1921.515, p. 2. 

4. Longirostrine—Osborn, 1918.468, p. 136. 
Rhynchorostrine—Osborn, 1918.468, p. 136. 
Brevirostrine—Osborn, 1918.468, p. 136. 

Serridentine (7. serridens phylum, Osborn, 1921.515, p. 8)—Osborn, 
1921.526, p. 330. 

8. Notorostrine—Osborn, 1921.526, p. 330, distinguished and defined. 

9. Zygolophodontinze—Osborn, 1923.601, p. 1, to include the species: 
Zygolophodon tapiroides, Z. turicensis, Z. virgatidens, Z. borsoni, 
and Z. pyrenaicus, as more fully defined below, which were 
mistakenly confused with the Serridentinz by Osborn (1921.515). 

10. Platybelodontinz-—Borissiak, 1928, p. 119. 

11. Amebelodontine—Barbour, 1929.2, p. 139. 

12. Tetralophodontine—van der Maarel, 1932, p. 108. 

13. Humboldtinze—subfam. nov. 

IV. ELEPHANTOIDEA: 14. Stegodontina—Osborn, 1918.468, p. 135, to include the Stegodonts of 
southern Asia. 
15. Loxodontine—Osborn, 1918.468, p. 135, to include the phylum of 
African elephants. 
16. Elephantina—Bonaparte, 1838, p. 112, 1850; Elephantini—Winge, 
1906, p. 172; Elephantinee—Osborn, 1910.346, p. 558, 1918.468, 
p. 1385. 
Euelephantinee—Osborn, 1918.468, p. 136. Abandoned as invalid, 
because the genus Huelephas is invalid. 

17. Mammontinze—Osborn, 1921.515, pp. 1, 14. 

asl ached 

Great Antiquiry.—The above seventeen’ subfamilies have been separated from each other for very long periods 
of geologic time, many of them since Lower and Middle Eocene times, others since Oligocene time. This very 
ancient geologic and adaptive habitudinal radiation is especially characteristic of the subfamilies included within 
the Meeritherioidea and the Deinotherioidea as well as of the eleven’ subfamilies included within the Mastodon- 
toidea. As regards the Elephantoidea, it would appear that the Stegodontine, the Loxodontine, and the 
Elephantine had already widely radiated in Miocene time, but we do not as yet know when the Mammontinze 
were given off from their closest relatives, the Elephantine. 

'(1935) See Appendix of the present Volume I for PaL#omMASTODONTIN® subfam. noy., based on the genus Palzomastodon Andrews, and GNATHABELO- 
_ DONTIN#, based on Gnathabelodon Barbour, also the removal uf Stegolophodon from the Stegodontidw, subfamily Stegodontine, to the new subfamily Sreco- 
LOPHODONTIN®, making twenty subfamilies in all, fourteen of which are included within the superfamily Mastodontoidea. 



SUBFAMILY DIvERGENCE.—Whereas the families of Proboscidea are distinguished from each other by pro- 
found modifications in the fundamental structure of the grinding teeth, the subfamilies of Proboscidea are distin- 
guished mainly by secondary adaptations of the grinding teeth correlated with profound adaptations in the upper 
and lower incisor teeth. For example, compare the lower incisor teeth in the Mastodontine, the Longirostrine, 
the Rhynchorostrine, the Brevirostrine, and the Notorostrine and observe how profoundly different each is from 
the other. This adaptation or loss of the lower incisor teeth is correlated with important adaptations of the upper 
teeth and finally with the structure of the grinding teeth. Here the principle of adaptive compensation is observed ; 
e. g., in the Rhynchorostrine, with their well-developed lower tusks, the grinding teeth remain excessively simple, 
whereas in the Brevirostrine, with the loss of the lower tusks, the grinding teeth become excessively complex. 

The phylogenetic basis of this seventeen-fold' subfamily division is to separate clearly every direct line of 
descent of species and genera from every other direct line of descent; the subfamilies are observed as vertically 
ascending ancestral lines which may be subdivided into one or more generic stages. 

ContTRAST wITH Copr.—This vertical phylogenetic use of the subfamily by Osborn (1918.468, pp. 133, 134) 
presents the most direct contrast to the system of Cope: “Cope’s family classifications were morphological and 
horizontal rather than phylogenetic and geological. Finding one or more single characters possessed in common 
at certain horizontal periods of geologic time by mammals in different lines of evolutionary descent, he seized on 
these common characters as convenient keys to classification. . . . I have reached the opinion that Cope’s method 
of morphological classification is untenable, that the only true and permanent classification is phylogenetic. . . . 
Sometimes the subfamilies proposed by zoologists conform to the phyletic lines discovered by paleontologists; 
sometimes they do not.” 

Inasmuch as the ORDER PROBOSCIDEA is distinguished by profound adaptations of the incisor teeth, the 
SuporpeERs by profound modifications of the incisors, [?-I,, the Famrires by profound adaptations of the grinding 
teeth, the SuBramILiEs by secondary adaptations of the upper and lower incisor teeth and compensatory adapta- 
tions of the grinding teeth, it remains to distinguish the Gmn»RA by successive stages in the evolution of these second- 
ary and compensatory adaptations. 

For example, the four genera of the phylum Mastodontinz in descending order: 
Pleistocene Genus Mastodon: Lower incisor tusks small, variable; upper tusks large, without enamel. 
Pliocene Genus Pliomastodon: Lower incisor tusks intermediate; upper tusks rounded, without enamel. 

Miocene Genus Miomastodon: Lower incisor tusks intermediate; upper tusks upcurved with broad 
enamel band. 

Oligocene Genus Palzomastodon: Lower and upper tusks unknown; upper tusks probably downturned 
with broad enamel band. 

Or, for example, the three genera of the phyla Tetralophodontine, Longirostrine, and Amebelodontine in 
descending order: 

Miocene Gernus Tetralophodon: Lower tusks procumbent; upper tusks downturned. Intermediate 
grinders tetralophodont. 

Miocene Genus Trilophodon: Upper tusks downturned with enamel band. Intermediate molars tri- 

Oligocene Grnus Phiomia: Lower incisors procumbent without enamel; upper incisors fighting weapons 
with enamel band. Intermediate molars trilophodont. 

1See footnotes on pages 19 and 27. 


Thus in the phylogenetic classification of the Proboscidea the Genus embraces and distinguishes the successive 
stages of adaptation and evolution of single or multiple characters, as observed in ascending geologic horizons. 

Among GENERA the phenomena of parallelism, of convergence, and of homoplasy are especially rife. For 
example, the superior incisors of Mastodon, of Anancus, and of certain species of Cordillerion tend to lose their 
enamel and to become upturned and outwardly arched so as to mimic each other closely, and there occurs a 
correlated homomorphy or parallelism in skull structure. Most confusing, however, to the early students (such 
as Falconer) of the Proboscidea was the parallel evolution of the trilophodont intermediate molars, e. g., 
Dp 3 #, M 1 3, M 2, which arose separately in Phiomia, in Palzomastodon (rudimentary), in Anancus, and in 
Stegodon. In fact, it is certain that in many distinct branches of proboscideans trilophodonty was independently 
developed. Similarly the tetralophodont condition of the intermediate molars arose independently in Tetralopho- 
don and in Anancus, also in the ancestral Stegodonts and ancestors of all the true Elephantoidea. The third 
form of generic parallelism is the independent evolution in all the genera of the bunomastodonts of single trefoils 
followed by double trefoils; for example, as in the transition from T'rilophodon angustidens (single trefoils) to Tetra- 
lophodon longirostris (double trefoils), or in the transition from Cordillerion andium (single trefoils) to Cuvieronius 
humboldtii (double trefoils). 

tocene.. Equal-area World Map. 1935. 

ae ae 150 

| Lf 



wy : 3 
a A 
a Ag 


LL aes 


Arrican ELEPHANT: After Blanc, 1897; Bocage, 1890; Buckley, 1876; Chapman, 1868; Chubb, 1909-1919; Claridge, 1915; Cox, 1900; 
Heilprin, 1887; Hippolyte, 1907; Johnston, 1907; Maydon, 1932; Roosevelt and Heller, 1914; Schultze, 1907; Sclater, 1900. 
Inpran Everuant: After Bartholomew, 1911; Blanford, 1888-1891; Blyth, 1872; Bishop, 1921; Champion, 1928; Hornaday, 1885: 
Hunter, 1868; Jerdon, 1874; Laufer, 1925; Lydekker, 1900-1908; Sanderson, 1907; Sclater, 1899. 


Thus by applying the principle of adaptive radiation in geologic time we find it is practicable to develop 
consistently a phylogenetic classification out of the Linnean system which included within the order Proboscidea 
only the single species Hlephas indicus. This phylogenetic system provides a consistent terminology for the main 
stock, its branches and subbranches, parts of which may be Anglicized for convenience as follows: 

Osborn, 1935: This classification of the year 1933 has been recently modified by successive discoveries up to July of the year 1935. 

Accordingly the reader is referred to page 685 et seq. for the classification of 1935 which introduces new forms and new concepts of 
relationship and lines of descent. 

OrpER PROBOSCIDEA (Proboscideans)—Illiger, 1811, p. 96. Upper Eocene to Recent time. 

SuBORDER I. MG@RITHERIOIDEA (Meeritheres)—Osborn, 1921.515, p. 2. A primitive, amphibious, 
swamp-loving stock. 

FamILy M@RITHERIIDA—Andrews, 1906, p. 99; Osborn, 1921.515, p. 1. 

SuBFAMILY Ma@ritHERINa® Winge-Osborn—Osborn, 1923.601, p. 1 (=Mceritheriini Winge, 
1906, p. 172). Upper Eocene, Lower Oligocene of Africa. 

SuporpER II. DEINOTHERIOIDEA (Deinotheres)—Osborn, 1921.515, p. 2. Specialized, fluviatile 
or river border proboscideans. Mio-Pliocene. 

FamILy CURTOGNATHIDA—Kaup, 1833.4, p. 516. 

SuBFAMILY DEINOTHERIINA—Osborn, 1910.346, p. 558 (=Dinotherina Bonaparte, 1841, p. 
253, Dinotheriina Bonaparte, 1850; Dinotheriini Winge, 1906, p. 172). Large, river- 
border, fluviatile forms of southern Eurasia; throughout the Miocene to the close of the 
Pliocene, into the Pleistocene. 

SuBORDER III. MASTODONTOIDEA! (Mastodonts and Bunomastodonts)—Osborn, 1921.515, p. 2. 
Chiefly forest and savanna browsers. 

FamMILy MASTODONTID (“True Mastodonts’’)—Girard, 1852, pp. 326, 328 (=Mastodontide 
Osborn, 1918.468, p. 134), including Paleomastodontide of Andrews, 1906, p. 130. Chiefly 
browsers in the north temperate and boreal forests. 

SuBFAMILY Mastopontinaz—Osborn, 1910.346, p. 558 (=Mastodontina Brandt, 1869, p. 35). 
Springing from unknown African ancestors; Oligocene of North Africa; including M7o- 
mastodon and Pliomastodon (Miocene and Pliocene of Holarctica),and Mastodon americanus 
(Pleistocene forests of North America). Persistent forest browsers. Grinders tetrabun- 
odont, sublophodont, lacking trefoils. 

SUBFAMILY ZYGOLOPHODONTIN (‘‘Yoke-toothed Mastodonts’’)—Osborn, 1923.601, p. 1. 
Browsers of the temperate forests of Europe and Asia, including T'uricius turicensis, T. 
tapiroides, T. virgatidens, Zygolophodon pyrenaicus, terminating with Z. borsoni. Miocene 
to Upper Pliocene. Grinding teeth polybunodont to purely lophodont, lacking trefoils. 

(1935) To the Mastodontoidea have been added two new families, the SERRIDENTID# and the HumBoLpTIp#&, and two new subfamilies, the PaLaomas- 
TODONTIN® and the STEGOLOPHODONTIN® (see Appendix of the present Volume for final classification). 



Famity BUNOMASTODONTID (‘True . Bunomastodonts’”’)—Osborn, 1921.515, p. 2. Arising 
from forms similar to Phiomia of North Africa, probably lowland and swamp browsers, 
radiating into nine! subfamilies. Oligocene to Lower Pleistocene. Grinders provided 
with single or double trefoils—hence ‘‘bunomastodont.” 

SuBFAMILyY LonarrostrIn& (‘‘Long-jawed Bunomastodonts”)—Osborn, 1918.468, p. 136. 
Long-jawed typical bunomastodonts, arising in North Africa (Phiomia*), migrating all over 
North Africa, southern Europe, Asia, and North America north of Mexico; including 
Trilophodon. Grinding teeth originally bunodont, trilophodont. Central conules. 

SuBFAMILY AMEBELODONTIN® (typical “Shovel-tuskers’’)—Barbour, 1929.2, p. 139. Includ- 
ing the shovel-tuskers of Nebraska. Typified by Amebelodon fricki Barbour. 

TETRALOPHODONTIN® (‘‘Four-ridge-crested Bunomastodonts”)—van der Maarel, 1932, 
p. 108. Typified by ‘Mastodon’ longirostris Kaup and Tetralophodon bumiajuensis, van 
der Maarel. Second molars with four ridge-crests, i.e., tetralophodont. 

SuBFAMILY SERRIDENTIN® (‘“‘Medium-jawed Bunomastodonts”)—Osborn, 1921.526, p. 330. 
Modifying the trefoils to the outer walls. Grinding teeth sublophodont; upper and lower 
incisor teeth as in the Longirostrine. Upper Miocene to Upper Pliocene of North America 
(typified by Serridentinus serridens and S. productus); Upper Pliocene of Texas, of Florida, 
and of Guatemala; Miocene of Mongolia (Serridentinus mongoliensis); Upper Miocene 
of India (Serridentinus browni); Pliocene of North China (Serridentinus lydekkert). 

PLATYBELODONTIN”E (“Flat-tuskers”)—Borissiak, 1928, p. 119. Typified by Platybelodon 
danovi Borissiak, including P. grangeri of Mongolia and Torynobelodon barnumbrowni of 
Nebraska. Extremely broad shovel-tusks. 

SuBFAMILY RHYNCHOROSTRIN® (‘‘Beak-jawed Bunomastodonts”’)—Osborn, 1918.468, p. 136. 
With powerful downturned upper and lower tusks. Known only in the southern United 
States and northern Mexico. Typified by Rhynchotherium tlascale of Mexico, R. euhypo- 
don, R. brevidens. 

Supramity Nororostrin (Notorostrines)—Osborn, 1921.526, p. 330. First known in the 
Andean region of South America (Cordillerion andium); subsequently discovered in the 
southern United States and Mexico. Distinguished by the loss of the lower tusks, the 
abbreviation of the lower jaw, the spiral tusks (C. andium). 

HumBo.ptin & (‘“Humboldtines’’)—new subfamily described in the present Memoir. Typified 
by Cuvieronius humboldtii with straight or upturned tusks without enamel; molar enamel 
ptychoid, highly folded. Includes Stegomastodon mirificus. 

SuBFAMILY BREVIROSTRIN® (‘“Short-jawed Bunomastodonts”)—Osborn, 1918.468, p. 136. 
Springing from the same stock as the Longirostrine ; distinguished by the rapid abbrevia- 
tion of the lower jaw, by the early loss of the inferior tusks, by the hypsodonty and laby- 
rinthine multiple trefoil pattern of the grinding teeth. First known in Europe (Anancus 
arvernensis), thence wandering into Asia (Pentalophodon sivalensis, Synconolophus dhok- 
pathanensis); not known to enter North America. 

1(Osborn, 1935) Seven subfamilies by removal of the Serridentine, Platybelodonting, and Humboldting and the addition of the Gnathabelodontine. 
2Phiomia now removed to the Amebelodontine (see Appendix of the present Volume for final classification). 


Suporper IV. ELEPHANTOIDEA! (Elephants and Mammoths of Africa, Eurasia, and North 
America)—Osborn, 1921.515, p. 2. Browsers (Stegodontinz), chiefly tree browsers 
(Loxodontine), general browsers and grazers (Elephantinz), browsers and grazers 

Famity ELEPHANTIDA—Gray, 1821, p. 305 (fide Palmer, 1904, p. 740); Osborn, 1910.346, p. 558. 

SUBFAMILY STEGODONTIN® (Stegodonts)—Osborn, 1918.468, p. 135. Original members closely 
similar to, and perhaps identical with, certain of the Zygolophodontinz of Europe. Dwellers 
in the very warm forest and savanna regions of southern Eurasia and western India. 
Grinders rapidly multiplying crests, but persistently short crowned (brachyodont). Known 
from the Middle Pliocene (Stegolophodon cautleyz) to the Lower Pleistocene (Stegodon 
insignis, S. ganesa) of India. 

SuBFAMILY Loxopontina# (Loxodonts)—Osborn, 1918.468, p. 135. Distinguished by long 
narrow grinding teeth with relatively few crests; embracing Loxodonta africana and all 
the species of African elephants, the cranial varieties of Loxodonta antiqua |= Hespero- 
loxodon|, Upper Pliocene to Upper Pleistocene, Loxodonta namadica [= Palzoloxodon|] 
of India which radiated widely over Africa and wandered all over southern Eurasia, 
attaining the tallest stature known among the elephants; also the dwarfed insular forms 
Loxodonta (Pilgrimia) falconeri, Loxodonta (Pilgrimia) melitensis, Loxodonta (Pilgrimia) 
creticus, Loxodonta (Pilgrimia) cypriotes, ete. 

SuspramMity Mammontin@ (Mammoths)—Osborn, 1921.515, p. 1. Of close original affinity to 
the Elephantine, including (a) the southern mammoths Archidiskodon planifrons and A. 
meridionalis of southern Eurasia, A. imperator of North America, all with broad-plated 
teeth and few crests; also (b) the northern mammoths which apparently include Par- 
elephas trogontheriz of western Europe, P. columbi and P. jeffersonii of North America, and 
the widespread woolly mammoth (Mammonteus primigenius) of the northern steppes. 


- 2 CENE 

A Ale RG Aak 1] 3)" | 22) /~ | (PLIOCENE 



Fig. 7. Diagram showing the known and unknown stages in the Adaptive Radiation of the Proboscidea up to April, 1925. After 
Osborn, 1925.662, p. 21, fig. 2. Compare previous diagram of 1921 (Fig. 3 of the present Chapter IT), also final diagrams of 1933-1934 
(Fig. 8 opposite) and 1935 (Appendix). 

‘See footnote on page 22. 


SuBFAMILY ELepHantin& (‘True Elephants’”’)—Osborn, 1910.346, p. 558 (=Elephantina 
Bonaparte, 1838, p. 112, 1850; Elephantini Winge, 1906, p. 172). Typified by Hlephas 
indicus Linn. of India, which apparently evolved during Tertiary and Lower Pleistocene 
times in northern Eurasia and did not appear in southern Asia (India) and Asia Minor 
until the Upper Pleistocene. 

To those monophyletists who cling to Mastodon-Stegodon-Elephas descent, this seventeen-fold' subfamily branch- 
ing of the proboscideans is very difficult to credit until we examine in detail the anatomy of the known genera and 
species of which these branches are composed and which we soon realize make up a very small proportion of the 
total number of genera and species entering into this titanic order from its origin in Lower Eocene time to its great 
climax in Upper Pliocene and Pleistocene times. The contrast between the known and the unknown is presented 
graphically in the accompanying diagrams (Figs. 7 and 8). 

In the succeeding pages of this Memoir the author will demonstrate that this Adaptive Radiation and Phylo- 
genetic Classification are not imaginary but are based on the solid ground of our present knowledge. This strong 
statement needs several reservations, among them, the ancestry of the Elephantids, now traced to the summit 
of the Pliocene only. 


Heaperoloxedan ~ 



ee OS 
OX Base of Miocene 


' ’ 

_ Base oF Eocene 

ApapTivE RapIATION OF THE PRoposcipEA (1933-1934) 
Fig. 8. Diagram showing the known adaptive radiation of the 37-41 generic phyla of the Proboseidea as discovered up to the year 1934. Compare 
previous diagrams (Figs. 3 and 7), also phylogenetic diagrams of 1935 in the Appendix. —- 

See footnote on page 27. 


In 1899-1900 Osborn, Tullberg, and Stehlin independently advanced the theory that Africa was the con- 
tinent on which the Proboscidea originated. The original homeland of the Proboscidea is no longer in doubt; 
the evidence (1933) favors the geologically ancient continent of Africa. 

OsBorn (1900).—Until the end of the nineteenth century Asia was favored as the homeland of the Proboscidea 
and it is still possible, although not at present probable, that the order originated there. The reasons why Asia is 
not at present favored as the Proboscidea center are threefold, namely: (1) The total absence of Proboscidea in 
the known Upper Eocene and Lower Oligocene exposures; (2) the apparent derivation of the earliest Asiatic 
Proboscidea from ancestral African prototypes; (3) the Miocene appearance of African Proboscidea in Europe and 
North America. If the Proboscidea had originated in Asia in Eocene time they probably would have migrated into 
North America in Eocene or at least in Oligocene time, since the Proboscidea were great travelers. If not the 
original homeland or primary center, Asia certainly became a secondary homeland in which occurred the chief 
evolution and adaptive radiation of the family Elephantide. Current exploration in central Asia will probably 
clear up this whole problem. 

THEORY OF THE AFRICAN CreNTER.—On the contrary, Africa, which remained the “dark continent” of 
paleontology until the beginning of the twentieth century, was seldom regarded as a very important center of 
evolution and adaptive radiation of the Mammalia, despite some excellent suggestions by Riitimeyer (1888) on 
this subject. The first paleontologists and zoologists to make a bold claim for Africa were Osborn (1900), Tullberg 
(1899), and Stehlin (1900). On this subject we may refer to Osborn’s addresses of 1900 (April, 1900.182, ‘“The 
Geological and Faunal Relations of Europe and America during the Tertiary Period and the Theory of the Suc- 
cessive Invasions of an African Fauna,’ and July, 1900.187, ‘Correlation between Tertiary Mammal Horizons of 
Europe and America”). The following citation is from Osborn’s July article (1900.187, pp. 56-58) : 


In Europe there are in the upper Eocene two classes of animals, first, those which have their ancestors in the older rocks; 
second, the class including certain highly specialized animals which have no ancestors in the older rocks—among these, perhaps, 
are the peculiar flying rodents or Anomaluride, now confined to Africa, and secondly the highly specialized even-toed ruminant 
types—the anoplotheres, xiphodonts and others, the discovery of which in the Gypse near Paris Cuvier has made famous. It 
is tempting to imagine that these animals did not evolve in Europe but that they represent what may be called the first inva- 
sion of Europe by African types from the Ethiopian region. 

It is a curious fact that the African continent as a great theater of adaptive radiation of Mammalia has not been sufficiently 
considered. It is true that it is the dark continent of palzeontology for it has practically no fossil mammal history; but it by 
no means follows that the Mammalia did not enjoy there an extensive evolution. 

Although it is quite probable that this idea has been advanced before, most writers speak mainly or exclusively of the 
invasion of Africa by European types. Blanford and Allen it is true have especially dwelt upon the likeness of the Oriental and 
Ethiopian fauna but not in connection with its antecedent cause. This cause I believe to have been mainly an invasion from 
south to north correlated with the northern extension of Ethiopian climate and flora during the Middle Tertiary. It is in a less 
measure due to a migration from north to south. Let us therefore clearly set forth the hypothesis of the Ethiopian region or 
South Africa as a great center of independent evolution and as the source of successive northward migrations of animals, some of 
which ultimately reached even the extremity of South America—I refer to the Mastodons. This hypothesis is clearly implied 
if not stated by Blanford in 1876 in his paper upon the African element in the fauna of India. 



The first of these migrations we may suppose brought in certain highly specialized ruminants of the upper Eocene, the 
anomalures or peculiar flying rodents of Africa; with this invasion may have come the pangolins and aard varks, and possibly 
certain armadillos, Dasypodid, if M. Filhol’s identification of Necrodasypus is correct. A second invasion of great distinct- 
ness may be that which marks the beginning of the Miocene when the mastodons and dinotheres first appear in Europe, also the 
earliest of the antelopes. A third invasion may be represented in the base of the Pliocene by the increasing number of antelopes, 
the great giraffes of the A2gean plateau, and in the upper Pliocene by the hippopotami. With these forms came the rhinoceroses 
with no incisor or cutting teeth, similar to the smaller African rhinoceros, R. bicornis. Another recently discovered African 
immigrant upon the Island of Samos in the A2gean plateau is Pliohyrax or Leptodon, a very large member of the Hyracoidea, 
probably aquatic in its habits, indicating that this order enjoyed an extensive adaptive radiation in Tertiary times. 

It thus appears that the Proboscidia, Hyracoidea, certain edentata, the antelopes, the giraffes, the hippopotami, the 
most specialized ruminants, and among the rodents, the anomalures, dormice, and jerboas, among monkeys the baboons, may all 
have enjoyed their original adaptive radiation in Africa; that they survived after the glacial period, only in the Oriental or 
Indo-Malayan region, and that this accounts for the marked community of fauna between this region and the Ethiopian as 
observed by Blanford and Allen. 

In his April and July, 1900, articles, Osborn printed two world maps (Figs. 9, 10) illustrating his theories 
regarding Africa and Asia; the Africa theory of Osborn was confirmed in 1901; the Asia theory of Osborn was 
confirmed in 1922. 






Insectivora, Cheiroptera, ¢ 
Tillodontia, Rodeptr 
Primates>Mes6donta ..° 
> = INDO-MALAYAN ‘Amblypoda, Condylarthra /Perissodactyls, Aucylopoda . 
os (Oriental) 

(Ethiopian) Archaeoceti 


path t 

Fig. 9. Division of the world into three Realms (Huxley) and 
eight main Geographical Regions. The continental platform is 
raised to the 200-meter line showing the main Tertiary land con- 
nections. After Osborn, 1900.187, Fig. I; compare Osborn, 1900.182, 
Chart II. 

Fig. 10. Orders of mammals as placed by Osborn in 1900 in their 
hypothetical chief centers of adaptive radiation during the Tertiary Period. 
The Prososciwea are placed very near the point of actual discovery in 1901 
of their Fayim ancestors. After Osborn, 1900.187, Fig. III; compare 
Osborn, 1900.182, Chart IV. 

As regards the Proboscidea and Hyracoidea, the above geographic charts of 1900 were confirmed by Charles W. Andrews in 1901. As regards central 
Eurasia as the chief center of radiation of fourteen orders of mammals, the present Central Asiatic Expeditions of the American Museum are strongly confirma- 
tory. As regards the Edentata, Litopterna, Toxodontia, and Typotheria, South America remains the chief center. As regards the Sirenia and Cetacea, it 
now appears probable that they also originated on the African continental shorelines. 

This African or Ethiopian center theory of 1899-1900 was destined to enjoy almost immediate confirmation 
through the discovery in 1901 of the previously unknown Upper Eocene—Oligocene land fauna of North Africa. 
This Fayam region had yielded between 1879-1885 a promising Eocene marine fauna and a Miocene land fauna 
containing one species of proboscidean (found to the north in Moghara, after 1898) related to Trilophodon 


In April, 1901, Charles William Andrews of the British Museum had an opportunity of accompanying Hugh 
Beadnell, the geologist, into the Fayim district. Here a marine (Zeuglodont) fauna had previously been found, 
and on this occasion a considerable number of vertebrate remains, including portions of skeletons of the probo- 
scidean Meritherium and of the sirenian Hosiren, were collected from beds of Upper Eocene age; the discovery 
of fluvio-marine beds of Lower Oligocene age resulted in the finding by Andrews of the first traces of the animal 
named Palzomastodon. Meritherium was the first Eocene proboscidean to be discovered in any part of the world, 
and the type of the second genus, named Palzxomastodon beadnelli, proves to be a possible ancestor’ of the classic 
Mastodon of Cuvier. The expeditions of Beadnell (1901-1904), of Andrews (1901-1903), and collections now 
preserved in the British Museum and the Geological Museum of Cairo, form the subject of a series of papers by 
Andrews, culminating in his great Memoir of 1906, entitled, ‘“A Descriptive Catalogue of the Tertiary Vertebrata 
of the Fayim, Egypt. Based on the Collection of the Egyptian Government in the Geological Museum, Cairo, 
and on the Collection in the British Museum (Natural History), London.” 

After discussing the principal characters of the vertebrate fauna, Andrews (1906) presents the following con- 
clusions regarding the homeland of the Proboscidea and the early history of these hitherto unknown mammals: 

[Andrews, 1906, p. xv] Although Arsinoitherium is certainly the most extraordinary of the Ungulates found in these beds, 
nevertheless the remains of the primitive members of the Proboscidea are perhaps of greater scientific interest, because they 
help to fill, at least to a large extent, one of the most obvious gaps in our knowledge of the extinct Mammalia. Previous to their 
discovery the earliest Proboscideans known were from the Lower Miocene (Burdigalien) of Europe and Northern Africa, and 
although many earlier deposits rich in mammalian remains were known in various parts of the world, in none of them was any 
trace of Proboscidea found, so that their appearance in Europe at the beginning of the Miocene period must be the result of 
their immigration from other regions. The probability that Africa would be found to be the original home of these animals 
was pointed out by several writers, notably by Osborn, Stehlin, and Tullberg. The first of these [Osborn, 1900.182, also 
1900.187] suggested that probably not only the Proboscidea but also the ‘Hyracoidea, certain Edentates, the Antelopes, the 
Giraffes, the Hippopotami, the most specialized Ruminants, and among the Rodents the Anomalures, the Dormice, and Jerboas, 
among Monkeys the Baboons,’ and, as his map [Fig. 10] suggests, the Sirenia also, originated in this region. Osborn also 
put forward the theory that a succession of migrations from Africa to Europe occurred, notably at the end of the Eocene, at the 
beginning of the Miocene, and again in the earliest Pliocene. It was in the early Miocene migration that the Proboscidea 
passed out of Africa for the first time so far as known. Stehlin [1900], who also emphasized the importance of Africa as a 
probable centre of mammalian evolution, expressed much the same views. Tullberg [1899] likewise regarded Africa as a centre 
of mammalian radiation, and pointed to Hystrix (or the whole of the Hystricognathi), the Simiz (Anthropoidea), and the 
Proboscidea as having probably migrated thence in the early Miocene. It is therefore very satisfactory that the earliest traces 
of land-mammals from the Eocene of Africa include remains of primitive Proboscidea, as well as early forms of Hyracoidea, 
Sirenia, and perhaps some of the other groups. 

The earliest-known Proboscidean is Meritherium, which occurs first in the Qasr-el-Sagha beds (Middle Eocene) and 
persisted till the Upper Eocene, its remains having been found in the Fluvio-marine series. This animal was about the size 
of a Tapir, which it must have greatly resembled in general appearance. . . . [p. xvii] The limbs are unfortunately not well 
known. The humerus differs considerably from that of the later Proboscidea, but some of the smaller species of Palzo- 
mastodon (see text-fig. 56) from the Upper Eocene seem to supply intermediate forms: probably the difference arises from 
the fact that Meritheriwm was a more or less amphibious type, while the later Elephants became fitted for progression on 
firm ground. The femur approximates very nearly to the form found in the later Proboscideans. As already mentioned, 
Meritherium was probably an amphibious, shore, or swamp living animal, and it was no doubt owing to the continuation of the 
conditions favourable to its mode of life that it persisted into the Upper Eocene period. . . . 

The largest species of Palzomastodon (P. beadnelli) must have been about the size of a half-grown Indian Elephant: 
in its general appearance it was Elephant-like, but differed in having a longer neck and the symphysial portion of the mandible 
prolonged beyond the skull (see text-fig. 48, p. 131) and covered only with the fleshy snout. Probably it could reach the 
ground with its lower incisors, and the end of the snout may already have been prehensile. . . . [p. xviii] In general appear- 

'(Osborn, 1935) This proves to be an error on the part of both Matsumoto and Osborn; Palzomastodon is not an ancestor of Mastodon, Zygoluphodon, or 
Turicius (ef. p. 43, Matsumoto’s phylogeny). See PALAOMASTODONTIN# subfam. nov., page 65 and Appendix. 


bake Moris Level 


ance Palzomastodon must have resembled a small rather long-necked Elephant, the most notable difference being that the 
trunk, instead of being freely flexible, was supported by, and formed the upper covering of, the elongated mandibular symphy- 
sis; its extremity, however, may have been free and to some degree prehensile. 

The further history of this group can only be briefly summarised here (see Phil. Trans. 1968, 1903, p. 99). The next form, 
Tetrabelodon angustidens from the European Lower Miocene, has the symphysis still more elongated and the narial opening shifted 
further back. At the same time the molars, or at least the posterior ones, are greatly increased in size and possess more trans- 
verse ridges. . . . [p. xix] This animal attained the size of a moderately large Indian Elephant, and except for the inflexibility 
of the mandible-supported trunk must have been very similar in appearance. In the later Miocene the mandibular symphysis 
shortened, leaving the trunk mobile and unsupported, as we now know it; at the same time traces of its original elongated 
condition are retained in the occurrence of deciduous lower incisors in some species of Mastodon and in the peculiar sharp process 
of the symphysis in the Elephants. . . . The above is, of course, only a general summary of the succession of forms which lead 
up to the modern Elephants, and no doubt there have been many checks and side-branches leading only to extinction in the 
course of the vast period that has elapsed since Meritherium existed. . . . [p. xxi] The question of the origin of the Sirenia is 
of great interest, and there seems to be a considerable amount of evidence in favour of the view first put forward by de Blain- 
ville, that they are intimately related to the Proboscidea. In the first place, the occurrence of the most primitive Sirenians 
with which we are acquainted in the same region as the most generalised Proboscidean Meritherium is in favour of such a 
view, and this is further supported by the similarity of the brain-structure and, to some extent, of the pelvis in the earliest- 
known members of the two groups (see pp. 202 & 214). . . . [p. xxvii] Another consideration which adds to the importance 
of Africa as a centre of mammalian evolution has been pointed out by Stromer [1903], namely, that part of it at least has 
probably never been submerged since the Palsozoic period, and formed a portion of a vast Permo-Triassic land-area inhab- 
ited by a great variety of mammal-like Theriodont reptiles from which the Mammalia may have actually arisen. This being 
the case, it is not only the Tertiary, but also the Secondary, deposits of this region that may be expected to yield most im- 
portant data for the history of the Mammalia. 

Map or tHE ReGion EXPLORED BY THE AMERICAN MuseuM ExpepitTion oF 1907 
Route of the Expedition indicated by dotted lines and arrows 

level of 
Libyan Desert 
math — 


Urrcr Bowe Larte Rite Otrosirs 
Containing Arsinoitherium, 
Paleomastoden, Mieritheriam. 



£ Ano 
= Onrosits 

Moore Bone Laver ano EsTuARINE DEPOSITS 
Temp Containing Sirenians,Zeuglodonts , 

le of a 
Ls ics ASK ch SAGHA fa ams Ancestral Elephants( Marithe rium) 


a ee = = 7 = = z 
SSS lomtr Bone Larter Matin Deposirs X 
SS Fentaiming Sirennius and Zeuglordtouts { y 
Rarine Beds A << 
(Marine) = 

5 =) 
Fig. 11 ays 


Fig. 11. Section showing the lowering levels of the ancient Lake Meeris, of the 

brackish Birket-el-Qurun, and of the sea, with the (1) lower marine Zeuglodon beds 
(Middle Eocene), (2) middle Qasr-el-Sagha Meritherium beds (Upper Eocene), 
(3) upper Fluvio-marine Palzomastodon beds (Lower Oligocene). After Osborn, 
1907.302, Century Magazine, p. 827. 

Fig. 12. The horizontal lines indicate the fertile lands of the Nile and of the 
Fayfm. The Faytm area formerly covered by Lake Mceris is now reduced to the 
brackish Birket-el-Qurun. The oblique dots north of Birket-el-Qurun indicate the 
area of the marine Qasr-el-Sagha beds. On the bluffs to the northwest are the 
Fluvio-marine beds, on which was placed the American Museum camp. After 
Osborn, 1907.302, Century Magazine, p. 826. 


Fig. 12 

Andrews thus adopted the Osborn-Stehlin-Tullberg theory that Africa was an important center of adaptive 
radiation of the Mammalia and called attention (p. xxv) to the fact that “the species known must be a mere frac- 
tion of the faunas inhabiting the Ethiopian region during the Middle and Upper Eocene periods’”’; he maintained 
that Osborn, Stehlin, and Tullberg were fully justified “in their assumption that the Ethiopian continent in early 
Tertiary (and perhaps pre-Tertiary) times was a very important centre of mammalian evolution.” 


Finally, the Deinotheres, although first discovered in Europe and Asia, have been found in recent years in 
deposits of Lower Miocene age in central (Deinotherium hobleyt) and northern (D. cuviert) Africa, as fully described 
in Chapter IV. Consequently early members of the Meeritheres, of the Deinotheres, of the true mastodonts, 
and of the bunomastodonts have been discovered in deposits of Upper Eocene to Lower Miocene age in the 
comparatively unexplored continent of Africa. In the Pleistocene of Africa occurs Deinotherium hopwoodi sp.nov. 

The general inference drawn from Andrews’ memoir and popular articles and restorations was that the 
genera Meritherium and Paleomastodon constituted the first and second links in a chain of descent which led 
up to the classic Mastodon angustidens of Europe, thus: 

Mastodon (Tetrabelodon |= Trilophodon]) angustidens of the Miocene of France. 
Palxomastodon, including a variety of species, Upper Eocene to Oligocene of Egypt. 
Meritherium, including at least two species, Middle and Upper Eocene of Egypt. 

Compare front views (Figs. 19 and 20) 

Fig. 13. Side view of the head of M@ritherium with the eye 
and ear in position, the form and position of the nostrils being 

somewhat conjectural. Observe the elongated cranial region and Fig. 14. Side view of the head of Phiomia wintoni, as modeled in 1908 by 
abbreviated facial region; also the direct opposition of the upper Erwin Christman, under the direction of Osborn. Observe the abbreviated cranial 
and lower incisor teeth. Modeled by, Erwin Christman, under the region, the elongated facial region, the upper tusks passing on the outer side of 
direction of Osborn in 1908. After Osborn, 1909.332, p. 140, the lower tusks. Originally described as Palzomastodon. After Osborn, 1909.332, 
fig. 2. p. 189, fig. 1. 

This monophyletic interpretation of Andrews’ conclusions was due rather to his broad inferences than to an 
examination of his actual descriptions in which the divergent adaptations to habit and to habitat, which would 
militate against any monophyletic theory, were clearly pointed out. But despite Andrews’ admirable work, the 
opinion widely prevailed up to the year 1909 that Meritherium gave rise to Palezomastodon and that Palzomastodon 
gave rise to Mastodon angustidens. 


The first to combat this monophyletic interpretation was Osborn who led an expedition into the Fayim 
region early in the year 1907, immediately after the publication of Andrews’ great memoir, assisted by Walter 
Granger and George Olsen. A preliminary study of the extensive collections of Meritheritum, Palzomastodon, and 
Phiomia, followed by comparison and restoration for purposes of exhibition in the American Museum, was entirely 
against the theory that Meritherium gave rise to Palezomastodon; in fact, it was in this process of restoration, as 
shown in the accompanying figures (Figs. 13, 14), that profound divergences between Palzomastodon (= Phio- 
mia) and Meritheriwm were brought out. This led to the publication in “Nature,” July 29, 1909, of Osborn’s 
paper, entitled, “The Feeding Habits of Mceritherium and Palzomastodon.”’ As will be fully set forth in Chapter 
XXI on ‘Affinities, Migrations, and Phylogeny of the Proboscidea,” differences in feeding habits give us 
the key to divergence, divergence gives us the key to phylogenetic separation and succession, and phylogenetic 
separation gives us the key to classification. 

Recent studies by Gregory (1910-1920), Matsumoto (1923), and _ Resroravrion oF THE EocENE AND RECENT SIRENIANS 

Simpson (1932) point towards the existence in Upper Cretaceous and 
Lower Eocene time in Africa of a common ancestral form of mammal 
which by adaptive radiation through ground-loving, shore-loving, and 
water-loving habits, may have given rise to the StRENIANS (Fig. 15), 
the Ma@riTHERES, and the PrRoposcipEANS. Widely 
different and profoundly divergent as the two great 
orders of Sirenians and Proboscideans are today, they 

still exhibit certain common characters in their inter- 

nal anatomy, certain common characters in their 
cranial and labial structure, as well as one unique char- 
acter in their grinding teeth, namely, trilophodonty or 

Fig. 15 (Left). The Eocene sea 
cow, Eosiren libyca, « contemporary 
of M eritherium lyonsi and M. gracile 
in the Qasr-el-Sagha beds, retaining 
the small hind limbs. Restoration 
and drawing by Charles R. Knight, 
under the direction of the author. 
After Osborn, 1907.302, Century 
Magazine, p. 831. 

(Right). The existing Manatee, 
Trichechus americanus, of the rivers 
of Florida, in which the hind limbs 
have entirely disappeared. Restoration and drawing by Charles R. Knight, 
under the direction of the author. After Osborn, 1907.302, Century Magazine, 
p. 831. 

the evolution of three crests on the superior and 
inferior molars. Recent studies by Simpson (1932) 
show that the Pre-Pleistocene forms of America are 
Dugongs rather than Manatees. 


It appears that Osborn in 1909 went too far in separating Meritherium from direct relationship to the Probo- 
scidea; he erred in the opposite direction, as shown in the following quotation (Osborn, 1909.332, p. 139): 
“The conclusion is that Moeritherium was a confirmed and continual river-living animal, feeding mainly under 
water and on the banks, more specialised for aquatic life than the hippopotami, as indicated by its feeble pelvis, 
but less specialised than the Sirenians. It would not be far from the truth to say, from our present knowledge 
of the animal, that Moeritherium is an offshoot of the Proboscideo-Sirenian stock, with slightly nearer kinship to 
the elephants than to the Sirenians.’’ This conclusion should now certainly be greatly modified, because Merv- 
thertwm proves to be of much nearer kinship to the Proboscidea than to the Sirenia; it isin fact one of the primary 
stocks of the Proboscidea. 


To Osborn’s extreme view Andrews replied in 1909, p. 305 (‘‘The Systematic Position of Moeritherium,” 
Nature, LX XXI, p. 305): ‘On the whole, it seems that the weight of evidence is in favour of regarding Meceri- 
therium as a proboscidean, though perhaps not on the direct line of ancestry of Palesomastodon, and retaining 
some characters of the original Proboscideo-Sirenian stock.’”’ Schlosser (1911) considered it doubtful whether 
Meritherium stands in the direct line of ancestry of Palzxomastodon. Gregory (‘“The Orders of Mammals,” 
1910, p. 868) concluded: ‘The genus [Meritherium] represents a very primitive offshoot from the Proboscideo- 

MaeritrHeriuM (Upper Eocene), Porom1A (LOWER OLIGOCENE), AND 

Fig. 16. (Left) Evolution of the head, proboscis, nostrils, and tusks in: 

. (1) Meritherium, (2) Phiomia wintoni, (8) Parelephas jeffersonii, as restored 

by Charles R. Knight in 1907, under the direction of the author. After 
Osborn, 1907.302, Century Magazine, p. 833. i. 

(Right) Comparative restorations to scale of: (1) Meritherium andrewsi, 
(2) Phtomia osborni, (3) Parelephas jeffersonti. Drawn by Charles R. Knight 
in 1907, under the direction of the author. After Osborn, 1907.302, Century 
Magazine; p. 834. Scale approximately one-fiftieth natural size. 

Sirenian stock. Its dentition and certain other characters indicate a nearer alliance with the Proboscidea than 
with the Sirenia, but it is far more primitive than any other known representative of either order.’’ More recently, 
‘Gregory (1920; p. 180)'in his exhaustive study on the lacrymal bone of the vertebrates declares: ‘The orbital 
‘region ‘[of Meritherium]-is much more primitive than that of other Proboscidea, and suggests the sirenian type. 
.. . [p. 245] The Sirenia, although very highly specialized for aquatic life, show special resemblances with 
Meritherium in the skull (including the orbital region) and dentition, and are generally regarded as a derivative 
of the proboscidean stem.”’ 


Matsumoto (1922, 1923, 1924) was the first to make a profound examination of the rich collections of Meri- 
thertum, Palzomastodon, and Phiomia in the American Museum, in comparison with the type and referred speci- 
mens in the British and Cairo Museums. Aided by the new evidence presented in this rich American Museum 
material, he deserves the chief credit: (1) Of clearly distinguishing the genus Palzomastodon from the genus 
Phiomia, (2) of clearly defining and distinguishing three species of Phiomia and three species of Palzomastodon, 
(3) of distinguishing species of Meritherium, (4) of closely comparing these three genera with each other and 
with representatives of the great orders Sirenia and Hyracoidea which occur in the same deposits. 

In the present Memoir Osborn entirely confirms Matsumoto’s principal systematic conclusions. Matsumoto’s 
separation of the genera Palzomastodon and Phiomia is a confirmation of one of Andrews’ observations, namely (An- 
drews, 1905, p. 562): “The species of Palzomastodon fall into two sections, in one of which the posterior end of the 
symphysis of the mandible is situated considerably in front of the level of the anterior premolar, while in the other 
it is only very little in front of that point. The first group, moreover, is distinguished by the comparative simplicity 
of the molars, in which the accessory cusps are scarcely at all developed, and by the small size of the talon of the 
last lower molar; into this subdivision the original species, P. beadnelli, falls, together with a much smaller form 
for which the name P. parvus is now proposed. The type-specimen of this new species is the right ramus of the 
mandible, with the premolars and molars zn situ, though somewhat crushed.” 

The profound structural and adaptive differences between these three genera of proboscideans, Meeritherium, 
Palzxomastodon (sensu strictu), and Phiomia, will be fully set forth in the systematic section below. A significant 
feature of this separation is that it tends to greatly strengthen the theory that Africa was the original homeland 
of the order Proboscidea in which they widely diverged from each other during Lower and Middle Eocene times, so 
that as we find them in Upper Eocene time they are distinctly polyphyletic. This gives a death blow to all the 
monophyletic inferences based on Andrews’ earlier writings. 

The relations of these three great phyla to each other are set forth in the section on phylogeny and classifica- 
tion (Vol. II) and in the present chapter. The relations of these three phyla of Proboscidea to the Sirenia 
and the Hyracoidea respectively are discussed in detail in Matsumoto’s paper of 1923, entitled, “A Contribution 
to the Knowledge of Meritheriwm,” in which he takes up with great fullness the following topics: 

A.—Characters Possibly Indicating Aquatic Adaptation (p. 102). B.—Characters Indicating Terrestrial and 
Non-aquatic Adaptation (p. 104). C.—Characters in Common with the Hyracoids (p. 105). D.—Characters 
Distinctive from the Hyracoids (p. 109). E.—Characters in Common with the Sirenians (p. 111). F.—Char- 
acters Distinctive from the Sirenians (p. 112). G.—Proboscidean Characters (p. 114). H.—Peculiar and 
Pre-Paleomastodont Characters (p. 117). I.—Natural Position of Meeritherium (p. 121). J _—Evolutionary 
Tendencies in the Earlier Proboscideans (p. 121). K.—Phylogenetie Relations Among the Hyracoids, Earlier 
Proboscideans, and Sirenians (p. 123). 

be paraphrased as follows: (1) That the proboscideans and sirenians were very closely related to each other in their 
earlier stages of evolution has been maintained by many eminent authors, such as De Blainville, Andrews, Osborn, 
Gregory, etc.; (2) it is very obvious that in their earlier stages they resembled each other in certain respects, but 



the phylogenetic relationship between the Proboscidea and Sirenia is somewhat less close than is considered by 
those authors (e. g., Osborn) who emphasize chiefly the resemblances between Meritheriwm and the earlier 
sirenians; (3) certain similarities between Merithertum and the earlier sirenians are probably due to their 
original phylogenetic relationship, other resemblances are attributable to convergence, 1. e., similarity of habits, 
still other resemblances are primitive characters common to all three groups of early sirenians, early proboscideans, 
and early hyracoids; (4) as to progressive development, it is obvious that the differences observed between the 
sirenian series of Prorastomus, Eotherium, Prosiren, Eosiren, Miosiren, Halitherium, Halicore, etc. and the progres- 
sive development of the proboscideans Meritherium, Palexomastodon, Trilophodon, and Megabelodon are very 
great; (5) doubtless no known sirenian can be looked upon as an ancestral type of Meritherium, nor can Meri- 
therium be looked upon as an ancestral type of any known sirenian; (6) consequently both the sirenians and the 
proboscideans may have descended from unknown ancestors that stand closer to each other than they do to the 
Hyracoidea stock. Detailed examination and comparison of thirty-four characters in sirenians, hyracoids, and 
early proboscideans led Matsumoto (1923, p. 114) to the following somewhat conflicting conclusion: ‘“‘Among 
these thirty-four characters examined, thirteen, vzz. (2), (9), (10), (12), (13), (15), (16), (17), (18), (19), (21), (25), and 
(26), are common to both the hyracoids and proboscideans in contrast to the [earlier or later] sirenians; ten, v7z. 
(4), (8), (11), (14), (20), (22), (27), (28), (29), and (30), are characteristic of the proboscideans in contrast to both 
the hyracoids and earlier or later sirenians; three, vz. (5), (7) and (23), are characteristic of the proboscideans in 
contrast, at least, to the earlier or later sirenians; three, viz. (32), (33), and (34), are characters vaguely distinctive 
of this genus from the earlier or later sirenians; three, vz. (9), (13), and (23) [these characters are referred to in 
duplicate], are characters of some of the terrestrial mammals in contrast to the aquatic ungulates; and one, v7z. 
(31), is characteristic of this genus in contrast to all the hyracoids, sirenians, and other proboscideans or earlier 
forms of these groups. It may easily be recognized that Merithertum has many characters common to both the 
hyracoids and proboscideans in contrast to the earlier or later sirenians, and many characteristics of the probo- 
scideans in contrast to both the hyracoids and later sirenians or earlier sirenians.”’ 

Osborn, 1924: In our opinion Meritheriwm, a true proboscidean of primitive type, stands much closer to the 
stem forms of the Sirenia than to the stem forms of the Hyracoidea; for the degree of this closeness we must wait 
for fuller evidence from Middle and Lower Eocene deposits in Africa. 

PROBOSCIDEAN CHARACTERS OF MaritHertum (Matsumoto, 1923).—Having dismissed the resemblances 
to the Sirenia and Hyracoidea as principally due to inheritance from primitive mammalian characters in general 
and also to convergence, Matsumoto applies a thirty-five character test and sets forth a very strong argument 
for the dominant relationship of Meritheriwm to the Proboscidea and especially for its peculiar and pre-palzo- 
mastodont characters. After comparing and contrasting Meritheriwm with Palxomastodon, we may paraphrase his 
conclusion as follows (p. 120): (1) In all the twenty-three characters examined, Meritherium is structurally a pre- 
palzomastodont type so far as we admit the conception that Palzomastodon, Trilophodon, and Megabelodon form 
together a fair series of evolutionary stages; (2) it is, of course, beyond doubt that the structural gap between 
Meritherium and Palxomastodon is fairly great, yet the fact should not be neglected that Meritheriwm stands 
structurally before Palzomastodon in the majority of its characters, that is, as a more primitive form; (3) con- 
sequently a presumed ancestral type of Palzomastodon should resemble Meritherium in many characters. 

Osborn, 1924: In our opinion Andrews and Matsumoto have positively established the proboscidean relation- 
ship of Meritherium. Osborn differs from Andrews and Matsumoto in placing Meritherium in an entirely separate 
line of descent, of superfamily rank known as Merritherioidea, distinguished by direct opposition of the upper 
and lower incisor teeth, as in rodents, by many aquatic adaptations in the skull, by profound differences in skull 


proportion, and probably by an amphibious mode of life approaching that of the hippopotami; in contrast to the 
purely terrestrial life of the Palzomastodon and Phiomia phyla. As to the amphibious habits of Mwritherium, 
Osborn and Matsumoto substantially agree. 

PALMOMASTODON ANCESTRAL TO THE TRUE Masropon (Matsumoto, 1924).—In Matsumoto’s succeeding 
paper (1924.1), “A Revision of Palawomastodon,” ete., he concludes by completely separating Mwritherium and 
placing it in an independent line, as shown in the accompanying diagram (op. cit., p. 58): 

In my opinion, the phylogenetic relationship of the genera just referred to, can be diagrammatically shown as follows: 

Mastodon Tetralophodon Megabelodon 
Zygolophodon Trilophodon Cherolophodon 
Palzomastodon Phiomia 


In discussing the phylogeny of the earlier Proboscidea, Matsumoto (1924.1, pp. 55-57) makes a final comparison 
between the skull and dentition of Ma@ritherium, Paleomastodon, and Phiomia; he rightly places Meritherium 
in aside-line of its own, but wrongly determines that the true Palzomastodon points towards Zygolophodon and the 
true Mastodon of the Mastodon americanus type for the following reasons. (1) The skull of Palzomastodon is not 
clearly known; judging from the shape of the palate it is probably shorter skulled than Phiomia which is distinctly 
long skulled; the skull of Zygolophodon of the European Miocene is not yet clearly known; the true Mastodon 
is distinctly short skulled. (2) The palate of the true Palzomastodon is very wide in proportion to the length of the 
cheek teeth, while that of Phiomza is rather narrow in the same proportion; the palate of the true Mastodon is 
very wide. (3) The mandibular symphysis of Palzomastodon appears to be rather short, while that of Phiomia is 
very long; the symphysis of the true Mastodon is short. (4) The largest and most conspicuous of the anterior mental 
foramina lies just below the first cheek tooth, P;; in the true Mastodon also the largest and most conspicuous of 
these foramina lies just below the anterior cheek tooth; in many short-jawed mastodonts and elephants the largest 
and most conspicuous of the anterior mental foramina lies just below the anterior cheek tooth also; the position 
of this foramen may be correlated with the development of the relatively short symphyseal region and the rela- 
tively small lower tusks, in contrast with the long symphyseal region and large lower tusks in the Phiomia-Trilopho- 
don series. (5) The cheek teeth of Palzomastodon are proportionately shorter and wider than those of Phiomia; 
the cheek teeth of Zygolophodon and Mastodon are also proportionately shorter and wider than those of Trilophodon. 


(6) The cheek teeth of Palezomastodon show a lower ridge formula than those of Phiomza; the potentiality of getting 
a higher ridge formula was lower in the Zygolophodon-Mastodon phylum than in the Phiomia-Trilophodon phylum. 
(7) The cheek teeth of Palzxomastodon are bunolophodont attaining a typically lophodont character when mode- 
rately worn, while those of Phiomia are typically bunodont; the cheek teeth of Zygolophodon and Mastodon are 
lophodont, while those of Phiomia and Trilophodon are bunodont. (8) In the cheek teeth of Me@ritherium, Palzo- 
mastodon, Zygolophodon, and Mastodon no trefoil pattern of cusps is developed, while in the Phiomia-Trilophodon- 
Tetralophodon phylum the trefoil pattern of cusps is well developed. (9) In the cheek teeth of Meritherium, 
Palzxomastodon, Zygolophodon, and Mastodon the crests are not very thick anteroposteriorly, the valleys are widely 
open and even the walls and bottoms of the valleys are worn, while in those of Phiomia, Trilophodon, and Tetra- 
lophodon the crests are very thick anteroposteriorly, the valleys are not so widely open, the worn surfaces are 
almost even. In the cheek teeth of Maritherium, Palxomastodon, Zygolophodon, and Mastodon the surface of the 
enamel is rather smooth, while in those of Phiomia, Trilophodon, and Tetralophodon the same is very rough. 

For all these reasons, Matsumoto continues, Palzomastodon appears nearly to correspond to a theoretical 
ancestral type of the Zygolophodon-Mastodon phylum; Palzomastodon appears to correspond to the beginning of a 
very great phylum, namely, the Zygolophodon-Mastodon phylum. 

Matsumoto thus announces a very important phylogenetic discovery, i. e., that Palzomastodon stands 
near the ancestry of the true Mastodontide. This discovery was immediately approved by Osborn, before the 
publication by Matsumoto of his complete results, with the following comments (see Matsumoto, 1922, p. 6, 
“Note by Henry Fairfield Osborn, August, 1922’’): ‘The significance of the above revision [Matsumoto’s] is 
that the true Palzomastodon beadnelli has bilophodont intermediate molars and a relatively broad skull; it isa 
rare animal both in the British Museum and American Museum collections; according to Andrews ((letter], 1922) 
the genotype (Palxomastodon beadnelli) was found at the very base of the Fluvio-marine Beds, Lower Oligocene, 
50 or 100 feet below the Phiomia level. . . . Associated with the type is a very large femur and humerus.”’ Palzo- 
mastodon is a relatively short-jawed animal, with bilophodont intermediate molars, whereas Phiomia is a very long- 
jawed animal with trilophodont intermediate molars. 

Matsumoto’s theory is now (1935) rendered improbable by Osborn’s closer studies in the present Memoir 
of the limited material of the true Palzomastodon from the Faytm (see footnote on p. 36 above). 

PuiomiA ANCESTRAL TO THE TRUE TRILOPHODON.'—From the first Andrews recognized the kinship of Phiomia 
to Trilophodon angustidens; Matsumoto goes further and proves that Phiomia is directly ancestral to what he 
terms the Trilophodon-Tetralophodon-Megabelodon phylum, whereas it has nothing to do with the ancestry of the 
Zygolophodon-Mastodon phylum. The kinship of Phiomia to Trilophodon is clearly indicated in the following 
characters: (1) Phiomia, Trilophodon, and Megabelodon are distinctly long skulled; (2) the palate of Phiomia is 
rather narrow in proportion to the length of the cheek teeth, the palates of Trilophodon and Megabelodon are also 
distinctly narrow; (3) the mandibular symphysis of Phiomia is very long, while the mandibular symphysis of 
Trilophodon and of Megabelodon is extremely long; (4) the anterior mental foramen in Phiomia lies on either side 
of the symphyseal region far anterior to the front cheek tooth, P;, and to the posterior end of the symphysis, in 
both Trilophodon and Megabelodon the position of the anterior mental foramen being quite similar to that observed 
in Phiomia; (5) the cheek teeth of Phiomia show a trilophodont ridge formula almost similar to that of Trilophodon, 
the potentiality of a multiple ridge formula is thus high in the Phiomia-Trilophodon-Megabelodon phylum, whereas 
it is low in the Palzomastodon phylum and still lower in the Meritherium phylum; (6) the cheek teeth of Phiomia 
are typically bunodont, those of Trilophodon, Tetralophodon, and Megabelodon are also bunodont; (7) in Phiomia, 

\(Osborn, 1935) The three or four known species of Phiomia are now recognized as directly ancestral only to the shovel-tusker Amebelodon of Nebraska. 
See Plate v between pages 235 and 236. 


Trilophodon, Tetralophodon, and Megabelodon the trefoil pattern of cusps is well developed; (8) the transverse 
crests in Phiomia, Trilophodon, Tetralophodon, and Megabelodon are very thick anteroposteriorly, the valleys are 
not so widely open, and the worn surfaces are almost even; (9) the surface of the enamel is relatively rough in 
Phiomia, Trilophodon, Tetralophodon, and Megabelodon; (10) the basal cingula are very rough and strong in 
Phiomia, Trilophodon, Tetralophodon, and Megabelodon, whereas they are rather feeble in Palzomastodon and 
Mastodon and more or less strong in Meritherium; (11) thus Phiomia appears to correspond to the beginning of a 
very great phylum, namely, the Trilophodon-Tetralophodon phylum of the Old World. 

Osborn, 1924: Quite independently of Matsumoto, Osborn had confirmed and extended Andrews’ opinion 
as to the Phiomia-Trilophodon phylum, and as early as 1917 (Osborn, 1918.468) placed Phiomia (= Palxomastodon) 
in the true longirostral line which he termed the Longirostrinez; but Matsumoto deserves the priority not only for 
clearly distinguishing Palzomastodon from Phiomia, but for suggesting that Palzomastodon was related to the 
subfamily which Osborn designates as Mastodontine. 


Thus Andrews, Osborn, and Matsumoto have successively prepared the way for a true understanding of the 
three great and widely distinct phyla of Proboscidea in northern Africa in Upper Eocene and Lower Oligocene times. 
As more fully pointed out in the phylogenetic classification of the forthcoming Volume II, the wide divergence 
of these phyla tends to confirm the theory that Africa was the homeland of the stem forms of all the Proboscidea, 
although it does not as yet finally demonstrate it. 

Osborn’s present classification of the thirteen species of the primitive North African Proboscidea is shown on 
page 65. 

Fig. 17. Paromta ossornt, Primitive Lower OxiGoceNne Swovet-Tusker OF THE Faybm, Eoypr 
Restorep (1932) By MARGRET FLINSCH, UNDER THE Direction oF Henry FarrrieLD OSBORN 
All figures one thirty-sixth natural size 

Phiomia osborni frequented the flood-plain region of the Nile and the present restoration shows a group of these animals in their 
natural habitat in Lower Oligocene time. The three specimens are based on a single lower jaw, which Matsumoto in 1922 dedicated to the 
present author; their estimated shoulder height is 1345 mm. or 4 ft. 5 in. 


The systematic descriptions and the nomenclature of the Faytim Proboscidea will be clearly reviewed below. 
Let us first examine their structure more closely with regard to adaptations to certain habits and modes of life 
and to progressive tendencies towards increasing perfection and specialization in adaptation. The Proboscidea 
follow the principle which we have clearly observed in many other ungulates, especially the families Brontotheri- 
de, Rhinocerotide, and Equide, which the author has been studying and comparing for the past thirty-four 
years, namely, a phylum having once started in a certain adaptive direction, if unchecked, will proceed to the extreme 
of adaptation. 

For example, elongation of the jaws, of the skull, and of the teeth begins in Phiomia and reaches an extreme 
in certain species of trilophodonts in which the progressive development is known as longirostral. 

Text-fig. 40. 


Fig. 18. First reconstruction by Charles W. Andrews of skull and mandible of Meritherium lyonsi. A, from above; B, from left side. 
After Andrews, 1906, text figure 40. About one-fourth natural size. 

The Osborn-Matsumoto reconstruction (Fig. 42) of the skull of M. andrewsi-trigodon differs widely in the relations of the upper and lower incisor 
teeth, I?-Is. These teeth are actually less tusklike and more gliriform than as restored by Andrews (Fig. 18). 

Classification should be expressive of phylogeny as brought about through adaptive radiation into diverse 
habits and habitats which influence, first, the entire dental and cranial structure, second, the entire foot, limb, and 
body structure, third, the adaptation of dental, cranial, limb, and body structure to defense and offense, and, f ourth, 
coadaptation and compensation through gain and loss of other organs. 

The analysis of the feeding and locomotor habits is the basis of the phylogeny and therefore of the classifica- 
tion of the Proboscidea, from the first appearance of these animals in the Upper Eocene up to the surviving recent 

forms in Asia and Africa. 


RésuMé OF PrREcEDING SEecTion.—Through the discoveries of Andrews (1901-1906) in the Eocene-Oligo- 
cene of the Fayim of Egypt of three very distinct types, namely, of Palzomastodon (very rare), of Meritherium 
(numerous), of Phiomia (very numerous), these three animals have become famous as the earliest known stages in 
the ancestry of the elephants. Since the wish is always father to the thought, and nothing was more eagerly 
sought for prior to 1901 than the primitive progenitors of the Proboscidea, it was altogether natural to place 
Meritherium in or near the direct line of ancestry of Hlephas, and consequently, with this idea, to provide Meri- 
therium, Paleomastodon, and Phiomia with a proboscis of less or greater length such as would befit more or less 
remote direct ancestors of the elephant. 

As such direct ancestors these three animals have gone forth into the general literature, and have therefore 
taken an entirely false and unnatural position in popular treatises on phylogeny. Andrews tried to correct this 
error in his great Memoir of 1906, and Osborn (1909.332) in his article “The Feeding Habits of Mceritherium and 
Palzomastodon,” followed by Matsumoto (1923), pointed out that since the dental and cranial structure of 
these three animals was profoundly different, it followed that the feeding habits were profoundly different and 
that they fed in three different habitats, consequently that their limb and body structure must have been different , 
finally, that none was ancestral to the other or to the genus Elephas. In brief, we were all so eager to welcome 
these animals into the order Proboscidea, that we too hastily assumed their ancestral position. 

We shall see that Meritherium, Phiomia, and Palxomastodon are widely separated both by habits and by 
structure from each other, as well as from the still to be discovered direct ancestors of the true elephants. 


The first more cautious note against the true-Elephas ancestry theory was sounded by Andrews in his Memoir 
of 1906 (1906, p. xvii) from which Osborn (1909.332, p. 139) made the following citation and comments: 

‘As already mentioned, Mceritherium was probably an amphibious, shore, or swamp living animal, and it was no 
doubt owing to the continuation of the conditions favourable to its mode of life that it persisted into the Upper Eocene 
period. In the meantime, however, either from this or some closely allied type, there had arisen another animal more 
adapted to terrestrial life and showing a great advance in the direction of the typical Proboscidea: to this creature the 
name Palzomastodon has been given.’ Elsewhere (p. xxi) Dr. Andrews notes that Mceritherium favours the view, first put 
forward by de Blainville, of an original relationship between the Proboscidea and Sirenia. Later on in the same work (p. 119) 
the same author, in commenting on the similarity between the pelvis of Mceritheri'um and that of the Eocene sirenian 
EKotherium, observes:—‘Then it may fairly be suggested that Mceritherium and Eotherium, both occurring in the same region 
(one the most primitive Proboscidean, the other occupying the same position with regard to the Sirenia), are, in fact, closely 
related, and had a common ancestor in early Tertiary times, probably in the Lower Eocene.’ On page 105 we find a comment 
on the remarkable likeness between the brains of Mceritherium and the Sirenia. 

CRANIAL AND DrenTAL ADAPTATIONS.—After these suggestive comments by Andrews were published, other 
specimens were secured by the American Museum Expedition of 1907 to the Fayim, including two fairly well pre- 
served skulls of Meritherium (Fig. 42), which gave witness afresh to the general analogy of the skull of Meritheriwm 
to that of the Sirenia. This material proves that Meritherium not only had no proboscis, but that it was a totally 
different animal from Phiomia both in its appearance and habits and only very remotely related to it; this material 
proves further that in czenotelic’adaptation the Meritherium skull is closer to that of the sirenians and of the 
hippopotami than to that of any of the proboscideans. The cutting teeth, lips, and mouth parts of Meritherium 
oppose each other in an entirely different manner than do those of Phiomia, so that it may be truly said that there 


was not the least resemblance between either the mouth parts or the feeding habits of these two animals. In 
Meritherium the nasal bones do not greatly recede, there was consequently little or no free retractile power of the 
upper lip, which is always the first step in the evolution of the proboscis as witnessed in living species of Tapirus. 
Comparison of the muzzle of Meritherium with that of Hyraz, of Castor, of Hippopotamus, and of other mammals 
with an enlarged pair of front teeth, tends to show that the upper and lower lips were heavy and fleshy, and some- 
what similar in form, in function, and in prehensile power; that the blunt tusks wore directly against each 
other and were entirely sheathed in enamel; that the lips were capable of closing over the tusks when the mouth 
was shut, somewhat as in the hippopotamus. The tusks were adapted as feeding organs rather than as fighting 
weapons, probably because Meritherium was protected from attack by its partly aquatic habitat. 

Compare side views (Figs. 13 and 14) 

Fig. 19. Front view of the head of Maritherium with the eye 
and ear in position; the form and position of the nostrils somewhat 
conjectural. The eyes are seen to be very far forward, well raised 
toward the top of the face; the ears also are raised high on the side 

of the head; both these peculiarities are adaptations to aquatic 
life to bring these sense organs near to the surface of the water 

in swimming, so that they will emerge first and disappear last. As Fig. 20. Front view of the head of Phiomia wintoni, as 
modeled by Erwin Christman, under the direction of Osborn, 1908. modeled by Erwin Christman, under the direction of Osborn, 1908. 

The above method of comparison of Phiomia and Meritherium is that of making life-size models of the skulls in which the sense organs, 

such as the eye and the ear, are placed exactly in position, while the contours of the lips, nostrils, mouth parts, and external ears are largely 

conjectural. In Osborn’s opinion (1926) the ear of Phiomia is erroneously restored, for it is much too elephantine. After Osborn, 

1909.332, pp. 139, 140. 

The conclusions drawn from the front teeth, I?-I,, from the simple bunodont grinding teeth, from the very 
short face (brachyopy), from the long cranium (dolichocrany), and from the extremely small bony eye sockets, 
are that Meritherium was a confirmed and continual river-living animal, feeding mainly under water and along 
the banks of rivers, more specialized for aquatic life than the hippopotamus, as proven by its feeble pelvic bones, 
but far less specialized for aquatic life in its limb structure than the Sirenia. 

This does not prove that Meritherium is of the order Sirenia, as Osborn suggested in 1909; it is certainly an 
independent member of the Proboscidea, as Andrews originally maintained and as Matsumoto has stoutly 
contended. Its cranial analogies are with the sirenian skull; its cranial and dental homologies are with the Probo- 
scidea. Its limbs and skeleton relate it to the primitive Proboscidea. 


PHIoMIA WITHOUT A Proposcis.—The restoration-model of the head of Phiomia (=the Palzomastodon of 
the previous literature) also is without a proboscis but is provided with a stout upper lip which was capable of 
pressing closely against the procumbent paired lower incisor teeth. As Osborn observed (1909.332), Phiomia in all 
probability had not developed a proboscis. A profound difference between Phiomia and Meeritherium is brought 
out in comparing the front (Figs. 19 and 20) and side (Figs. 13 and 14) views of the head, in which it is seen that 
whereas the eyes of Phiomia are in the typical mammalian position above the first permanent grinder, those of 
Meritherium are very far forward, well raised in the front part of the head, and of very diminutive size, as is 
shown by the shallowness of the eye sockets. 

The distinctive peculiarity of Phiomia is that its eyes are in the position typical among mammals, that is, 
above the first true grinders, M'-M,. The reason that the eyes appear to be so far back is that the lower jaws are 
extended unusually far forward. The upper jaws recede, terminating at the sides in the sockets of the very sharp, 
laterally compressed tusks which at this stage of evolution were chiefly developed as fighting or defensive weapons, 
while only of indirect value as feeding organs. When the upper and lower lips are restored in such a manner 
as to enable the animal to close its mouth, the upper tusks are so largely covered by the lips that they are not 
especially prominent. In contrast with Meritherium the nasal bones and narial openings in Phiomia deeply 
recede; thus a very wide space is left to be filled by the large retractile, prehensile upper lip which could un- 
doubtedly be raised or lowered. 

EvIpENCE AGAINsT A Proposcis.—The question now arises (Osborn, 1909.332, p. 140): ‘How far had this 
lip begun to transform into a proboscis? Was there a free projecting proboscis as represented in several previous 
restorations? A negative answer appears to be furnished by the structure and mode of wear of the lower incisors. 
Together these form a broad, protrusive, spoon-shaped feeding organ, which is invariably greatly worn on the 
upper surface and somewhat less at the ends. This worn upper surface seems to prove that in the prehension of 
food the edge of the upper lip was constantly pressed downward against these teeth, thus, with the aid of fine 
particles of grit and sand, which were occasionally taken in, causing wear. In brief, the food appears to have been 
seized between the upper lip and the spoon-shaped lower teeth. Paleeomastodon was a browser, and this lip could 
be turned up and retracted effectively to pull down smaller branches, but there is no reason to suppose that it had 
the free curling and independent prehensile power which characterises a true proboscis. If we critically consider 
the theory of the animal possessing a proboscis of considerable length, we find it rests upon the idea of kinship 
with the elephant rather than upon careful study of the mouth parts themselves.” 

If Phiomia possessed an independent prehensile proboscis extending beyond the line of the mouth for the 
seizing of food, we cannot assign any function to the large and much worn paired lower incisor teeth; the most 
probable theory of their function, therefore, seems to be that shown in the model (Fig. 14), namely, the prehension 
of food by pressing the upper lip against the lower incisors, rather than the prehension of food by grasping with 
a long and flexible proboscis; this function is progressively developed in the Amebelodont successors of Phiomia, 
also without a proboscis. 

The first rule in the restoration of adaptations is not to be too much influenced by kinship, but to adhere to 
the evidence afforded by the hard parts themselves. According to this rule Phiomia had no proboscis; it had a 
powerful upper lip which pressed against the tips of the lower incisor teeth. Osborn (1933) believes that this 
function was continued into the subfamily of the Amebelodontine (Longirostrines) to which Phiomia directly 
gave rise. 


One proof of the forest habits of Palzomastodon is the rarity of its fossil remains; forest-living animals are 

relatively rare. 

The true Palzxomastodon, typified by Palzomastodon beadnelli Andrews, 1901, and supplemented by the more 
recently described Palzomastodon intermedius Matsumoto, 1922, is only partly known, namely, the palate, the 
grinding teeth, and portions of the zygomatic arch; therefore we can only form a conjecture as to its habits from 
our knowledge of the animals which appear to be analogous to it, namely, the true mastodonts, known to be of 
forest-living habitat and of leaf-browsing habits. Consequently analysis of the habits of the Fayim proboscideans 

Phiomia. wintoni. 
Amer. Mus. 13450 Ref, 



Palazomastodon intermedius 
B Amer Mus. 13449" 

: ( 
Amer, Mus, 13431 Ref. OSs Eek 

A Mosritherium 

All 1/3 nat. size 

Fig. 21. 

“ Meritherium, Palzxomastodon, and Phiomia. 
teeth. A, Meritherium, Amer. Mus. No. 13481; B, Palzomastodon, 
Amer. Mus. No. 138449; C, Phiomia, Amer. Mus. No. 13450.”’ All 

figures one-third natural size. 

is incomplete in respect to this genus. 

animals as follows: 

Rivers and swamps 

1. Face abbreviated; cranium elongated. 

2. Upper and lower second incisor tusks 
of equal size, directly opposed, sheathed 
in enamel; feeding, not fighting, organs. 

3. Eye sockets extremely small, placed 
anteriorly; eyes and auditory openings 

4. Upper and lower lips opposing each 

5. Molars short and broad, tetra- 

6. Amphibious or aquatic in habit, 
hence rarely fossilized. 

Superior molars, compare Matsumoto, 1923, fig. 10: 


Fig. 22. 

Left upper grinding teeth. 

G Phiomia osborni 
Amer. Mus. 13468 

B _ Palzzomastodon intermedius 
Amer. Mus. 14547 

Amer. Mus. 13437 Ref. (rev.) 

All 1/3 nat. size 

Inferior molars, compare Matsumoto, 1923, fig. 11: 
“ Meritherium, Palezomastodon, and Phiomia. Left lower grinding 
A, Meritherium, Amer. Mus. No. 13437 (reversed), second 
and third lower molars; B, Paleomastodon, Amer. Mus. No. 14547, 
first, second and third lower molars; C, Phiomia, Amer. Mus. No. 

13468.” All figures one-third natural size. 

It is certain that the habits and habitat were different from those of 
Phiomia and profoundly different from those of Meritherium. We may therefore contrast these three Fayim 

Lowlands and savannas 

1. Face elongated; cranium normal. 

2. Lower tusks horizontal, no enam- 
el; upper tusks sharply pointed, re- 
curved, enamel band on outer surface; 
fighting, not feeding, organs. 

3. Eye sockets and auditory open- 
ings in normal position. 

4. Upper lip opposing lower incisors. 

5. Molars narrow and elongate, tri- 
lophodont, large central conules 

6. Shore or lowland habitat, hence 
frequently fossilized. 



Forests and savannas 

1. Facial and cranial proportions un- 

2. Tusks unknown. Upper tusks prob- 
ably circular in section, with lateral enamel 

3. Eye sockets and auditory openings 
in normal position. 

4. Upper lip probably forming a pro- 

5. Molars relatively broad, hexabuno- 
dont, subtrilophodont, central conules rudi- 

6. Probably of forest habitat, 
rarely fossilized. 



Revistons (1901-1923).—The Faydm proboscideans described by Andrews (1901-1906), and added to by 
Pontier (1907), Schlosser (1911), Matsumoto (1922-1924), and Petronievics (1923), are so fundamentally im- 
portant and so confused in the present literature that it is necessary to give them a revision by themselves prior 
to the phylogenetic consideration which they will receive in Chapters III, VI, and VIII of this Memoir. This 



Rhagatherum falaeomastodon 
Merunerum —Megalohyrax 
Ancodon Plotemaia 
Saghatherlam Phiomys 
Prerodon Aplerodon 

Palzomastodon intermedius Type 
Palxomastodon parvus Type 
Meritherium andrewsi Type 
Merritherium trigodon Type 

ee F051) Mammal boner TS LJ 
(e) FLuvi0-MARINE BEps 
fee ss =| _j| Phiomia osborni Type 
rained O| Phiomia serridens Type 
jaa: Phiomia wintoni Type 
oe seen ae =| |Q¢| Phiomia minor Type 
ARS! — See a $ 
“Arsinoltherium y = Palzomastodon beadnelli Type 

Upper Mokattam Level fee 

Martane : a Meritherium lyonsi Type 
Poatrals Maritherium gracile Type 

[Meritherium ancestrale Type] 

Zeuglodon Focetus 


Present Surface of Lake Qirun 



Fig. 23. Section through the Upper Eocene and Lower Oligocene formations north of Lake Qurun, Fayim, Egypt 
Arrows indicate levels richest in remains of fossil mammals. Approximate geologic distribution of proboscidean types shown 
in right-hand column. After Andrews, Beadnell, Granger, and Osborn. Reproduced from Osborn, 1910.346, p. 199, fig. 89 

revision is chiefly based on the authority of Dr. H. Matsumoto (1922, 1923, 1924), who reviewed the entire Fayam 
collection of the American Museum and the original type collection of the British Museum for the purpose, 
inasmuch as neither Osborn nor Andrews had been able to verify these observations in detail. In 1905 Andrews 
(1905, p. 562) observed that the species of the genus Palzomastodon fall into two groups; in 1922 Matsumoto (1922, 
p. 1) divided these groups into two genera, viz.: Palzomastodon Andrews and Phiomia Andrews and Beadnell. 
In general Osborn accepts Matsumoto’s main conclusions, which have a very important bearing on the 
phylogeny of the Proboscidea. 

The geologic levels of the type and genotype specimens are indicated in the above diagram. The Qasr-el- 

Sagha beds are now regarded as Upper Eocene; the Fluvio-marine beds as Lower Oligocene. 




1. Fluvio-marine beds Palzomastodon Beadnelli Andrews, 1901 Genotype of Palzomastodon 
2. Qasr-el-Sagha beds Meritherium lyonsi Andrews, 1901 Genotype of Meritherium 
3. Fluvio-marine beds Phiomia serridens Andrews and Beadnell, 1902 Genotype of Phiomia 
& 4. Qasr-el-Sagha beds Meritherium gracile Andrews, 1902 = Meritherium gracile 
E 5. Fluvio-marine beds Meritherium trigodon Andrews, 1904 = Meritherium trigodon 
© 6. Fluvio-marine beds Palzomastodon minor Andrews, 1904 = Phiomia minor 
g 7. Fluvio-marine beds Palzomastodon parvus Andrews, 1905 = Palzomastodon parvus 
E 8. Fluvio-marine beds Palzomastodon wintoni Andrews, 1905 = Phiomia wintoni 
z 9. Fluvio-marine beds Paleomastodon Barroisi Pontier, 1907 Bees: See Gece 
fe Phiomia minor (?) 
4 10. Fluvio-marine beds Meritherium Andrewsi Schlosser, 1911 = Meritherium andrewsi 
11. Fluvio-marine beds Palzomastodon intermedius Matsumoto, 1922 = Palxomastodon intermedius 
12. Fluvio-marine beds Phiomia osborni Matsumoto, 1922 = Phiomia osborni 
13. Qasr-el-Sagha(?) beds Mceritherium ancestrale Petronievics, 1923 = Meritherium ancestrale 

Fig. 24. Geographic distribution of species of the Fayiim Proboscidea according to the heavy face numerals in preceding list. The white dots within the 
black areas represent the approximate localities where the types of these thirteen species were found; the figures in the circles represent the species. 
Mastodon [ = Phiomia| pygmxus Depéret, 1897, from Algeria, omitted in this map (see Fig. 189). 

GroLocic LevELs oF M@rITHERIUM SpeEciES.—The above chronologic list of species was corrected by 
Andrews himself as to geologic level; thus the genotype (Meritheriwm lyons?) as well as the type of Meritherium 
gracile are both from the Qasr-el-Sagha beds, whereas a referred specimen of Meritheriwm lyonsi(?) as well as 
the type of Meritherium trigodon are from the overlying Fluvio-marine series. It is this referred specimen of 
Meritherium lyonsi(?) which Schlosser selected as the type of Meritherium andrewst. 

It is not known whether these smaller, intermediate, and larger species constitute two geologic ascending 
series; itis probable that they do. 

QuARRY DisTRIBUTION REcorpDs.—The shifting river-channel sands and gravels in which the species of 
Meritherium, Phiomia, and Palzomastodon were deposited are exposed in different quarries; the numerous speci- 
mens referable to these three genera and six species found by the American Museum Expedition of 1907 were 
recorded as occurring in Quarry A, Quarry B, and Quarry C, or at various distances from these chief quarries. 


Perhaps future examination of these quarry (A, B, C) records will enable us to determine the relative geologic 
levels. So far as examined the American Museum quarry records do not indicate the separation of the species by 
the quarries, e. g., from Quarry B are recorded the following species: Palzomastodon parvus,a single specimen; P. 
intermedius, two specimens; P. beadnelli, one specimen; Phiomia minor, seven or more specimens; P. wintoni, thirty 
or more specimens. Thus the same Quarry B of the American Museum records contains three species of Palzo- 
mastodon and two species of Phiomia. This demonstrates that Quarry B represents a very long period of deposition. 
The very progressive species Phiomia osborni was found eight miles west of Quarry A; it undoubtedly represents 
a higher geologic level. See full quarry records in American Museum card catalogue, also as printed by Matsu- 
moto (1923, 1924). 

GroLogic LEVELS OF PALHZOMASTODON AND PHIOMIA Species.—Andrews informs us that the type of Palzo- 
mastodon beadnelli came from the very base of the Fluvio-marine series, 50 feet below the typical level of Phiomia; 
yet P. beadnelli is the largest and most progressive species of the true Palzomastodon. Unfortunately we have as 
yet no record of the geologic levels on which were found the type specimens into which six species of Palzo- 
mastodon and Phiomia are now divided, namely: 

Large, progressive specific stages Phiomia osborni Palzomastodon beadnelli 
Intermediate specific stages Phiomia wintoni Palzomastodon intermedius 
Small, primitive specific stages Phiomia minor Palzomastodon parvus 

The associated mammalian and reptilian fauna with these species of proboscideans is shown in the Andrews 
table (1906, p. ix) reproduced herewith. 

Fluvio-marine Series (4 in Section).—Variegated sands, sandstones, clays, and 
warls, with limestone-grits and thin bands of limestone. In the lower beds of 
this Series are large numbers of silicified trees associated with vertebrate remains, 
including :—Arsinoitherium zitteh, A. andrewsi, Saghatherium antiquum, S. minus, 
S. magnum, S. majus, Megalohyrax eocanus, M. minor, Paleomastodon beadnelli, 
250 P. wintoni, P. parvus, P. minor, Meritherium (?) lyonsi, M. trigonodon, Phiomia 
a serridens, Ancodon gorringei, A. parvus, A. sp., Rhagatherium cegyptiacum, 
Geniohyus mirus, G. fajumensis, G. major, Hyenodon sp., Pterodon africanus, Fig. 25. Complete faunal list 
Apterodon macrognathus, Sinopa ethiopica, Eremopezus eoceenus, Crocodilus articeps, | (1906) of the Upper Eocene (155) 
O. megarhinus, Tomistoma gavialoides, Testudo ammon, T. beadnelli, T. isis, Qasr-el-Sagha Series, also of the 
Stereogenys libyca, Podocnemis fajumensis, P. blanckenhorni with var. ovata, Pelo- | yor Oligocene (250) ‘Sinwig: 

medusa progaleata, Aetobatis sp. simrinw Hesiod of Muni kane 

; Andrews, 1906, p. ix. Compare 

Qasr-el-Sagha Series (Carolia Beds) (5 in Section).—Alternating limestones, marls, | figures 11, 12, and 23. 
clays, and sandstones. The vertebrate remains include :—Meritherium lyonsi, | 
M. gracile, Barytherium grave, Eosiren libyca, Zeuglodon osiris, Crocodilus sp., 
155 Tomistoma africanum, Psephophorus eocenus, Thalassochelys libyca, Podocnemis 
antiqua, P. stromeri and var. major, Stereogenys cromert, S. podocnemioides, 
Gigantophis garstini, Pterosphenus schweinfurthi, Fajumia schweinfurthi, Socnopea 
grandis, Pristis fajumensis, P. ingens; Eopristis reinachi, Propristis schweinfurthi, 
Myliobatis sp., Carcharodon sp. 


| Upprr MoxKartram. 

ORIGINAL DESCRIPTIONS AND PRESENT REFERENCE.—The remaining pages of the present Chapter II are 
devoted to the firm establishment of Charles W. Andrews’ types, type figures, type descriptions, and type geologic 
levels, in accord with the monographic system which prevails throughout this Memoir. It is'‘a token of Andrews’ 
signal ability as a paleontologist that he interpreted so accurately the chief characters of this proboscidean fauna 
from materials invariably scattered and more or less fractured in the course of fluvio-marine deposition. In 
italics is given Andrews’ original reference, e.g., Palzomastodon minor Andrews, 1904; in heavy face is given 
the present generic and specific reference, e.g., Phiomia minor Andrews, 1904. It is earnestly hoped that these 
references, representing years of labor on the part of Andrews, Matsumoto, and the present writer, may stand 
and be accepted as a basis of future monographic research. 



Palzomastodon beadnelli Andrews, 1901 
Figures 2, 26, 34, 41, 41a, 90-95, 97; see also page 147 

Fluvio-marine formation of the Fayaim, Egypt=Upper Eocene of 
Andrews = Lower Oligocene of the present Memoir. 

This was the first proboscidean to be described from the 
Fay(im and was properly given the name Palzomastodon, namely, 
the ancient Mastodon. 

(Andrews, letter, 1922): The type was found at the very base 
of the Fluvio-marine beds, Lower Oligocene, 50 or 100 feet below 
the Phiomia level; this genotype specimen, with a very large 
femur and humerus, was the only true Palzomastodon material 

Palzomastodon Beadnelli Andrews, 1901. Zoologist, Vol. V 
(4), August 15, 1901, pp. 318, 319 (Andrews, 1901.1); name only. 
Tageblatt des V Intern. Zoologen-Congresses, Berlin, No. 6, 
August 16, 1901, p. 4 (Andrews, 1901.2), notice; published volume 
(Verhandlungen) dated 1902, see especially p. 528. Geol. Mag., 
Dec. IV, N.S., Vol. VIII, pp. 400-409, September, 1901 (Andrews, 
1901.3); description and figure. Typr.—(Op. cit., 1901.3, 
p.401): . . . ‘‘nearly complete left ramus of the mandible of a 
Proboscidean.” In Andrews’ supplementary description “A 
Descriptive Catalogue of the Tertiary Vertebrata of the Faytim, 



Fic. 1.—Left ramus of mandible of Paleomastodon Beadnelli. natural 
size. (A) From above; (B) outer surface. 

Fig. 26. Original type figure of Palzomastodon beadnelli Andrews, 1901. 
After Andrews, 1901.3, p. 401, fig. 1, A, B. (Op. cit., p. 401): .. . ‘nearly 
complete left ramus of the mandible of a Proboscidean.” Original (Geol. 
Mus., Cairo, C. 10014) destroyed; cast Brit. Mus. M. 8059; cast Amer. 
Mus. 9984. One-sixth natural size. 

Egypt” (1906, p. 150, Pls. x11-xvi, text figs. 50, A, 51) the type 
specimen is given as “‘ Left ramus of mandible with pm. 4 (broken) 
and m. 1-3 (Pl. xv, figs. 1, 14); Geological Museum, Cairo.” 
Original (Geol. Mus., Cairo, C.10014) destroyed in Custom House 
at Cairo; cast in British Museum (M.8059), also cast in American 
Museum (Amer. Mus. 9984). Horizon anp LocaLiry.— 


Fluvio-marine beds, Lower Oligocene, Fayfim, Egypt. TYPE 
Ficgure.—Andrews, 1901 3, p. 401, fig. 1, A, B. 

Andrews (1901.3, p. 401) observes: ‘‘One of the most impor- 
tant specimens found in the higher beds (probably Lower Oligocene) 
is the nearly complete left ramus of the mandible of a Proboscidean, 
which is in many respects similar to that of Mastodon angustidens, 
but belonged to a much smaller and in several respects more gen- 
eralized form.” Andrews (1904, p. 115) added the following meas- 
urements of the type and supposed female specimen of P. bead- 
nelli in comparison with P. minor: “This species may be called 
Palzomastodon minor; its dimensions compared to those of P. 
beadnelli are shown in the following table. . . 

P.[=Phiomia] minor P. beadnelli _ P. beadnelli 

(? female) (type) 
m.3 47 mm. 65 mm. 78 mm. 
m. 2 AD pe 5D) Gone 
m. 1 oot Aliortt 48> 
pm. 4 Do ee 39. 48‘ 
pm. 3 280 30 ? 

Matsumoto (1922, p. 3) confirms Andrews’ description of 1901 
and adds Amer. Mus. 13481 as a referred specimen. In 1924 
(p. 4) he distinguishes Palxomastodon beadnelli from P. interme- 
dius and P. parvus as follows: 

“3 Length of lower molar series measuring 194 mm.; 
that of lower premolar and molar series, 285mm. (Andrews’ 
type) beadnelli 

2.—Length of lower molar series measuring 159 mm. (type 
[Amer. Mus.] 14547); that of upper molar series, 150-152 
mm. (paratype [Amer. Mus.] 13449); that of upper premolar 
and molar series, 250 mm. (ditto) intermedius. 

1.—Length of lower molar series measuring 130 mm.; 
that of lower premolar and molar series, 197 mm. 
(Amd rews tyne) ae cette eile oleae est etsy 

Meritherium lyonsi Andrews, 1901 
Figures 2, 27, 41a, 42, 42a, 43, 44, 49; see also page 72 
Qasr-el-Sagha formation of the Fayim, Egypt=Middle Eocene of 
Andrews= Upper Eocene of the present Memoir. 

This was the second proboscidean to be described from the 
Faytim and was very appropriately given the name Meritherium, 
in reference to the ancient Lake Merris of the Greeks. 

Meritherium lyonsi Andrews, 1901. Tageblatt des V Intern. 
Zoologen-Congresses, Berlin, No. 6, August 16, 1901, p. 4 
(Andrews, 1901.2), notice; published volume (Verhandlungen) 
dated 1902, see especially p. 528. Geol. Mag., Dec. IV, N. S., 
Vol. VIII, pp. 400-409, September, 1901 (Andrews, 1901.3), 
description and figure; also “A Descriptive Catalogue of 
the Tertiary Vertebrata of the Fayim, Egypt,” 1906, 
pe 120) Piss vin. DX, ox Typer.—(Andrews, 1906, p. 120): 


“Mandible associated with upper molars and a dorsal vertebra.” 

Geol. Mus., Cairo, C.10000; cast Amer. Mus. 9977. Horr- 
ZON AND Locauiry.—Qasr-el-Sagha beds, Upper Eocene, 
Fayfim, Egypt. Tyre Ficgure.—Andrews, 1901.3, p. 404, 
fig. 2. 

One-fourth natural size. (A) Ups 

Fig. 27. Original type figure of Meritherium lyonsi Andrews, 1901. 
After Andrews, 1901.3, p. 404, fig. 2. (Andrews, 1906, p. 120): ‘Mandible 
associated with upper molars and a dorsal vertebra.’’ Geol. Mus., Cairo 
(C. 10000); cast Amer. Mus. 9977. One-fourth natural size. (Andrews, 
1901.3, p. 404, fig. 2): A, Upper teeth (C.10001) =paratype; A, B, portions of 
large skull, B showing front of snout with tusklike second incisors (C.10002, 
cast Amer. Mus. 9978)=paratype; C, left ramus of mandible from outer side 
(C. 10000)=type. (See Fig. 44.) 

The referred Meritherium lyonsi (?) recorded by Andrews in 
the overlying Fluvio-marine beds was chosen as the type of 
Meritherium andrewsi by Schlosser. Matsumoto (1923, p. 124) 
distinguishes the two species of the Qasr-el-Sagha formation as 

“(1.) Larger form of the Qasr-el-Sagha Formation. Lower 
premolars very short: P*4 [P24], ca. 69 mm. (Andrews); 
lower molars very long, M:-s, ca. 104 mm. (Andrews). All the 
lower cheek-teeth very wide. Po, triangular, its widest part 
corresponding to the posterior lobe. P*4, 67-78 mm.; 
MLSS amma (ANGLO WS) / a. ajs.0.5.-s0e6 « sae salle O M. lyonsi. 

(2.) Smaller form of the Qasr-el-Sagha Formation. Lower 
premolars not very short in comparison with the length of 
lower molars: Ps-;, ca. 62 mm. (specimens in the American 
Museum); lower molars very short: M;-;, 83 mm. (specimen 
in the American Museum). All the lower cheek-teeth are 
narrow. P24, 62 mm. (Andrews); M!*%, 75—ca. 79 mm. 
AUC WS) Reet Petri is Nett Src ces « sce nicurtets M. gracile.” 

Matsumoto (op. cit., p. 125) adds the following information 
from the American Museum collection: 

Amer. Mus. 13444; “two of the three fragments of mandibular 
rami of this specimen number appear to belong to this species. 

They are very peculiarly weathered, as a characteristic of the 
weathered specimens from the Qasr-el-Sagha Formation, with 
much-weathered and badly preserved molars in situ. Qasr-el- 
Sagha Formation of the Fayim.” 

“The dimensions of the teeth of these fragments, in compari- 
son with those of Andrews’ specimens, are tabulated as follows 
(in mm.): 

| Lower Teeth Upper Teeth 
{Amer. Mus.] 
13444 ditto (Andrews) 
fv ss De 
length AY Se” 22 7 (Pb eae tA 
P 2 \ width ae aig 23? 
P3 length Peery en 38 26-5. «ts <n Phar 
width ne ek SOND Ae beh eee 
P4 length io eee | Deee Meet w Scie 
width oS ih eo Di Dery hs Sea 
length Sue Mateen MONO ZO Me. Beater 
M1) width ee a ee 
M2 length 29 28 35 BEN aa OO. crore 
width 25. 251" 39 2aV ar t28 a. A Sectnee 
M3 length 40 39 42 sete OO) oul oletst 
width 28 28 30 Lee eee 30+! 
Length of P 24 69+! 68 Sr a Dears 
Length of M 1-3 104+! datro EOD ernie 

‘These dimensions are estimated from Andrews’ figures.”’ 

Phiomia serridens Andrews and Beadnell, 1902 
Figures 28, 180, 181; see also page 239 

Fluvio-marine formation of the Fayim, Egypt=Upper Eocene of 
Andrews = Lower Oligocene of the present Memoir. 

This was the third proboscidean to be described from the 
Faydm and was appropriately given the name Phiomia, the Greek 
equivalent of the province now known as the Fay(im. 

Phiomia serridens Andrews and Beadnell, 1902. “A Pre- 
liminary Note on Some New Mammals from the Upper Eocene 
of Egypt,” Survey Dept., Pub. Works Ministry, Cairo, 1902, pp. 
1-9 (original description); “Descriptive Catalogue of the Terti- 
ary Vertebrata of the Fayim, Egypt,”’ Andrews, 1906, pp. 169-171 
(supplementary description), Pl. xvm, figs. 4, 4a. TyPE.— 
Anterior portion of a left mandibular ramus, bearing Di, and Dp>-s 
in situ; Geological Museum, Cairo (C.10007); cast Amer. Mus. 
9981. Horizon AND LocaLity.—Beds of the Fluvio-marine forma- 
tion (Lower Oligocene), Fayim, Egypt, north of Birket-el-Qurun. 
Type Figure.—Andrews and Beadnell, 1902, figs. 1, 2. Matsu- 
moto (letter, 1921) remarks of the referred specimen shown in fig- 
ure 3 (op. cit., 1902) that it is not the type and not a proboscidean. 

Matsumoto (letter, 1921): Schlosser first pointed out that 
this genus and species might be merely a juvenile form of a 
Palzxomastodon [{i. e., Phiomia] (Neues Jahrb. f. Min., Vol. I, Pt. I, 
1905, p. 157, Referate); Andrews accepted Schlosser’s view, after 
securing a beautifully preserved mandible of a very young Palzo- 


mastodon {i.e., Phiomia] probably referable to Ph. wintoni (Andrews, 
Geol. Mag., Dec. V, N.S., Vol. IV, 1907, p. 97); though the type 
specimen is merely a very young individual, it shows certain indica- 
tions that it might belong to the wintoni-minor type of Palzomas- 
todon [i. e., Phiomial, as distinguished from the beadnelli-parvus 
type; moreover, the specific name Phiomia serridens antedates any 
of the specific names of the wintoni-minor type. 

Matsumoto (1924.1, p. 14) remarks that the genotype of 
Phiomia serridens Andrews is a species probably identical with 
Palzomastodon [=Phiomia] wintoni Andrews, 1905; or possibly 
with Palzomastodon [=Phiomia] minor Andrews, 1904. 

Osborn, 1924: The type (genotype) cast of Phiomia serridens 
is close in size to P. wintoni; this observation is validated by 
another but slightly larger milk tooth specimen, namely, Amer. 
Mus. 13458. 

of the mand/ble 

Anterior portion of left ramus 

Dorsal aspect 

of the mandible. 

Anterior portion of left ramus 

Lateral aspect 

Fig. 28. Original type figure of Phiomia serridens Andrews and Bead- 
nell, 1902. After Andrews and Beadnell, 1902, figs. 1 and 2. Anterior portion 
of a left mandibular ramus, bearing Diz and Dp»-3 in situ; Geol. Mus., Cairo 
(C. 10007), cast Amer. Mus. 9981. One-half natural size. 

Meritherium gracile Andrews, 1902 
Figures 29, 41, 42a, and 45; see also page 73 

Qasr-el-Sagha formation of the Fayim, Egypt=Middle Eocene of 
Andrews =Upper Eocene of the present Memoir. 

Meritherium gracile Andrews, 1902. “Preliminary Note on 
some Recently Discovered Extinct Vertebrates from Egypt,” 
Geol. Mag., Dec. IV, N.S., Vol. [X, pp. 291-295 (original descrip- 
tion); ‘‘Descriptive Catalogue of the Tertiary Vertebrata of the 
Faytim, Egypt,” 1906, pp. 127, 128 (supplementary description), 
Pl. xvu, figs. 1, 2. Typr.—(Andrews, 1906, p. 127): ‘An im- 
perfect skull . . . including the palatal region, associated with 
cervical, dorsal, and lumbar vertebre; Geological Museum, Cairo.” 


Geol. Mus. Cairo [C.10003]; cast Amer. Mus. 9979. Horr 
ZON AND Locatiry.—(Andrews, 1906, p. 127): “Qasr-el-Sagha beds 
(Middle [Upper] Eocene): north of Birket-el-Qurun,” Fay(im, 
Egypt. Typs Ficurn.—Andrews, 1906, Pl. xvu, figs. 1, 2. 
Matsumoto (1922, p. 5, 1923, p. 125) rightly characterizes 
Meritherium gracile as the small species of the Qasr-el-Sagha beds. 
His definition of this species is as follows (op. c7t., 1923, p. 125): 

““(2.) Smaller form of the Qasr-el-Sagha Formation. Lower 
premolars not very short in comparison with the length of 
lower molars: P:-s, ca. 62 mm. (specimens in the American 
Museum); lower molars very short: M;-;, 83 mm. (specimen 
in the American Museum). All the lower cheek-teeth are 
narrow. P*4, 62 mm. (Andrews); M', 75—ca. 79 mm. 

(Annidtews))a:csca2 ss oeisievceeme clei caeracraciotie M. gracile.” 

The additional information based on the American Museum 
collection from the Qasr-el-Sagha formation of the Faytim is 
quoted below (Matsumoto, 1923, p. 128): 


Fig. 29. Original type figure of Meritheriwm gracile Andrews, 1902. 
After Andrews, 1906, Pl. xvu, figs. 1, 2. (Op. cit., p. 127): ‘‘An imperfect 

skull . . . including the palatal region, associated with cervical, dorsal, and 
lumbar vertebre;” Geol. Mus., Cairo (C. 10003), cast Amer. Mus. 9979. 
(Fig. 1) . . . “anterior portion of skull, palatal view: one-third nat. size. 

Qasr-el-Sagha beds (Middle Eocene).” (Fig. 2) . . . “occipital region of 
same skull, posterior view; one-third nat. size. Same horizon.” 

Amer. Mus. 13443; ‘mandible, with P;-M; of the left side 
and M,, of the right side in situ. [Amer. Mus.] 13444; one of the 
three fragments of mandibular rami of this specimen number, with 
badly preserved molars in situ. [Amer. Mus.] 13445; fragment of 
a right mandibular ramus of a young individual, with the teeth 
broken away. [Amer. Mus.] 13446; fragment of a left mandibular 
ramus, with the crowns of the teeth broken away. All from the 
Qasr-el-Sagha Formation of the Fayam.” 

“The mandible of [Amer. Mus.] 13443 measures 305 mm. in 
length without incisors, 8.5 mm. in length of symphysis, 55 mm. 
and 50 mm. in the distance between the two first molars and the 
two last molars respectively, 225 mm. in the bicondylar width and 


76 mm. in the height of the mandibular ramus at M, without 
the teeth. In this specimen the symphysial depression already 
cited is observed to be present. In the fragmentary mandible of 
{Amer. Mus.] 13446, the same depression is clearly observed, also.”’ 

“The dimensions of the cheek-teeth of the specimens at hand, 
in comparison with those of Andrews, are tabulated as follows (in 

Lower Teeth Teeth 
[Amer. Mus.] [Amer. 
13443 Mus.] | (Andrews) | 
right-left 13444 
{length uate; secs ae 22 
P 2 ) width ie ie ee 
length Se 22 oats 20 
P3 \ width Lee 15 .. | 23 
length res 21 acre 20 
e4 width roe 18 swe PAY ere 
M1 length 23 23 22 23% 25 
width 20 19 19? 23 «21 
oJ length 28 28 27 a 27 | 
width 22.5 23 22 25 23° | 
length 34 33 aan 28 28 
width 24 24 ile! | aa 55 | 
Length of P24 | 63+ Gis. wos MietnOMe TL | 
(alveoli) (ditto) 
Length of M 1-3 84 84 cee mea) ¢ 923-4 


This dimension is estimated from Andrews’ figure.” 

“Thealveoliof each I, and I?of themandible of [Amer. Mus.] 13443 
measure 10 mm. and 20 mm. in transverse diameter, respectively ; 
the lateral extension of, and the minimum distance between, the 
two alveoli of lower tusks are 50 mm. and 9 mm. respectively. 
The two alveoli of first incisors are situated just below and anterior 
to the part corresponding to minimum distance between the two 
alveoli of tusks. Judging from these alveoli, the lower first incisors 
might be located not strictly inside, but inside and below and 
anterior to, the pair of lower tusks, which might be rather closely 
set to each other. These lower tusks appear to be distinctly smaller 
than those of M. lyonsi and andrews?.” 

Meritherium trigodon Andrews, 1904 
Figures 30, 42, 42a, 46, 49; see also page 74 

Fluvio-marine formation of the Fayim, Egypt=Upper Eocene of 
Andrews = Lower Oligocene of the present Memoir. 

Meritherium trigodon Andrews, 1904. “Further Notes on the 
Mammals of the Eocene of Egypt,” Geol. Mag., 1904, Dec. V, 
N. S., Vol. I, pp. 109-115 (original description); ‘‘Descriptive 
Catalogue of the Tertiary Vertebrata of the Fayim, Egypt,” 
1906, pp. 128, 129 (supplementary description, including change of 
name, i. e., trigodon to trigonodon). Typr.—(Andrews, 1904, 
p. 112):... “portion of the right ramus of a mandible containing 

the three molars: of these m.3 is in perfect and unworn condition, 
while m.2 and m.1 have lost portions of their outer sides.” Brit. 
Mus. M.8499; cast Amer. Mus. 9980. HoRIzON AND 
Locauiry.—(Andrews, 1906, p. 128): ‘“‘Fluvio-marine beds 
(Upper Eocene {Lower Oligocene]); north of Birket-el-Qurun,” 
Fayim, Egypt. Type Ficure.—Andrews, 1906, Pl. rx, fig. 5. 


Fig. 30. Original type figure of M@ritherium trigodon Andrews, 1904. 
After Andrews, 1906, Pl. rx, fig. 5. (Andrews, 1904, p. 112): ... “portion 
of the right ramus of a mandible containing the three molars: of these m.3 
is in perfect and unworn condition, while m.2 and m.1 have lost portions of 
their outer sides.” Brit. Mus. M.8499; cast Amer. Mus. 9980. Natural size. 

Marsumoro, 1923, p. 134.—Meeritherium trigodon is a smaller 
form of the Fluvio-marine formation. In revising the American 
Museum collection, Matsumoto refers the following specimens to 
this species (p. 134): 

Amer. Mus. 13430; ‘greater part of a full-grown skull, bear- 
ing all the upper cheek-teeth in situ.” 

Amer. Mus. 13431; “fragment of a skull, including a greater 
part of a right half of palate, bearing P*-M$ in situ.” 

Amer. Mus. 13433; “left M? attached to a fragment of upper 

Amer. Mus. 13435; “fragment of a right ramus of mandible, 
with all the cheek-teeth in situ.” 

Amer. Mus. 13436; “fragment of a left ramus of mandible, 
with all the cheek-teeth in situ.”’ 

Amer. Mus. 13439; “right P, and M,, with their roots 
broken away.” 

On this material M. trigodon is redefined as follows (Matsu- 
moto, 1923, p. 125): 

“(4.) Smaller form of the Fluvio-marine Formation. 
Lower premolars not very short in comparison with the length 
of lower molars: P24, ca. 63 (specimen in the American Mu- 
seum)—70 mm. (ditto, as well as Andrews); lower molars 
rather short: My-3, 93 (specimens in the American Museum)— 
98 mm. (Andrews). All the lower cheek-teeth very narrow: 
P., fusiform in upper view, its widest part corresponding to 
the middle part. P?4, 60-63 mm. (specimen in the American 
Museum); M'?, 83-85 mm. (ditto). Skull, small and lightly 


9982. Horizon AND Locauity.—(Andrews, 1906, p. 168): 
“‘Fluvio-marine beds (Upper Eocene [Lower Oligocene]): north 
of Birket-el-Qurun,’ Fayim, Egypt. Tyre Ficgure.— 
Andrews, 1906, Pl. xtv, figs. 1, 1a, and text fig. 50 D, p. 143. 

Tyree Description.—(Andrews, 1904, p. 115): ‘‘A portion of 
the right ramus of a mandible shows that there existed in the Upper 
Eocene beds a species of Palzomastodon considerably smaller than 
P. beadnelli, even allowing for a very wide range of individual varia- 
tion in size in that species. . . . This species may be called Palzo- 
mastodon minor; its dimensions compared to those of P. beadnelli 
are shown in the following table . . . 

built. Sagittal and occipital crests very strong. Zygomatic 
width very large in comparison with the length of skull. P.[=Phiomia] minor P. beadnelli __ P. beadnelli 
Distance between the external auditory openings, as well as (?female) (type) 
the width of occiput, very large.............. M. trigodon.” m. 3 47 mm. 65 mm. 78 mm. 
Matsumoto (op. cit., p. 134) gives a detailed description of the m. 45" 48 tae 65re 
six specimens in the American Museum listed above as referred m. 1 oot able ace AS. 
to this species, concluding (p. 137) with the detailed measure- pm. 4 250 Sy) 48“ 
ments of the grinding teeth as follows (in mm.): pm. 3 Pah S10 ? 
Lower Teeth Upper Teeth 
[Amer. Mus.] (Andrews) [Amer. Mus.] 
13435 13436 13439 13430 13431 13433 
right left 
P2 length 19.5 23? 23 19 22? 
width 10 11 11 21 18? eae 
P38 length 22 24? 25 23 23.5 24 
width 15 14? Neh We s5 |) 2s 24? 25? 
P4 length 23 23.5 25 24 22) 20:5 20 22 
width 18 18.5 19 1 20h ee eee 23? 25.5 
M1 length 27 24 27.5 26 27 | 25 22 24 
width 21 19 21 oe 21 | 25 26.5? 24 een 
M2 length 29 31 32 29 28 30 29 
width 24 24 ne 28 27 25? 24 
M3 length 37 37 40 31 32 30 
\ width 26.5 25 NE re: 26.5 27 26 
Length of P 2-4 63 70 on th) 60 63 aes 
Length of M 1-3 93 93 98 85 83 84 

Phiomia minor Andrews, 1904 
Figures 31, 34, 178, 179, 182, 183; see also page 239 

Fluvio-marine formation of the Fayfim, Egypt=Upper Eocene of 
Andrews = Lower Oligocene of the present Memoir. 

Palzomastodon minor Andrews, 1904. “Further Notes on 
the Mammals of the Eocene of Egypt,’’ Geol. Mag., 1904, Dec. V, 
N.S., Vol. I, p. 115 (original description); ‘Descriptive Catalogue 
of the Tertiary Vertebrata of the Faytim, Egypt,’ 1906, pp. 168, 
169 (supplementary description), Pl. xtv, figs. 1, 1a, and text fig. 
50D, p. 148. Typr.—(Andrews, 1904, p. 115): ... “part 
of the ramus [right] and the coronoid process of an immature 
mandible, in which m.3 has not yet been cut, although it is 
completely developed.” Brit. Mus. M.8479b; cast Amer. Mus. 

(Andrews, notes, 1922): ‘These measurements [of P. bead- 
nelli] are no doubt from a specimen of P. wintoni.”’ 

Matsumoto (1924.1, p. 16) reviews the type of Paleomastodon 
minor Andrews and transfers it to the genus Phiomia. He then 
compares it very carefully with specimens referred by Andrews to 
Palzomastodon beadnelli and to P. [Phiomia] wintoni, also with 
the type of Palzomastodon barroisi Pontier, and concludes that 
Phiomia minor is a valid species which may be clearly. distin- 
guished from both Phiomia wintoni and P. osbornt. 

He refers to Phiomia minor (1924, pp. 18, 19) eighteen speci- 
mens in the American Museum collection, of which the principal 
numbers are the following (also Matsumoto, 1922, p. 3): Ameri- 
can Museum referred specimens Nos. 13469, 13471, 13475, 13483, 


13486, 13448, 13455, 13461, 13464, 13465, 13467. Of the total 
number of eighteen specimens it is important to note that Quarry B 
and vicinity yielded ten specimens; Quarry B also yielded Phiomia 
wintoni in abundance. Quarry A yielded three specimens of P. 
minor, also P. wintoni; one important specimen, namely, Amer. 
Mus. 13469, was found eight miles west of Quarry A. 


Fig. 31. Original type figure of Palzomastodon minor Andrews, 1904. 
After Andrews, 1906, Pl. x1v, figs. 1, 14, and text fig. 50 D (see present Memoir 
Fig. 34D), p. 143. (Andrews, 1904, p. 115): . . . ‘‘part of the ramus [right] 
and the coronoid process of an immature mandible, in which m.3 has not yet 
been cut, although it is completely developed.” Brit. Mus. M.8479b, cast 
Amer. Mus. 9982. One-half natural size. 

The following (Matsumoto, 1924.1, p. 20) are the detailed 
measurements of the mandibles: 

“The mandibles of the specimens Nos. 13469, 13471 and 13475, 
in comparison with one described by Andrews, measure as follows 
(in mm.). 


suMoTO, 1924.1, pp. 19-21).—(1) Amer. Mus. 13469 bears an extra 
pair of incisors, I; or C, just behind the enlarged I, or C; these 
extra teeth are either third incisors or canines. (2) Mandible of 
Amer. Mus. 13471 very large in its dimensions [referable to 
Phiomia wintoni]. (3) Skull (Amer. Mus. 13448) bearing all the 
upper cheek teeth, P’-M’*, measured and described by Matsumoto 
in detail. 

Palwomastodon parvus Andrews, 1905 
Figures 32 and 34; see also page 146 

Fluvio-marine formation of the Fayim, Egypt=Upper Eocene of 
Andrews = Lower Oligocene of the present Memoir. 

Palxomastodon parvus Andrews, 1905. “Note on the Species of 
Palxomastodon,’’ Geol. Mag., 1905, Dec. V, N. S., Vol. II, pp. 
562, 563 (original description); “Descriptive Catalogue of the 
Tertiary Vertebrata of the Fayfim, Egypt,” 1906, pp. 162-168 
(supplementary description), text figs. 50C, p. 143, and 55, p. 163. 
Typr.—(Andrews, 1905, p. 562): . “right ramus of the 
mandible, with the premolars and molars in situ, though somewhat 
crushed.”’ Brit. Mus. M.8479a; cast Amer. Mus. 9976. Horr- 
zON AND Locatiry.—(Andrews, 1906, p. 163): ‘‘Fluvio-marine 
beds (Upper Eocene [Lower Oligocene]): north of Birket-el- 
Qurun,” Fayim, Egypt. Type Ficure.—Andrews, 1906, text figs. 
50C, p. 143, and 55, p. 163. 

Type Description (Andrews, 1905, p. 562).—In his paper 
Andrews observes the two sections of Palzomastodon, subsequently 
separated by Matsumoto into Palzomastodon and Phiomia: 

“The species of Palxomastodon fall into two sections, in one of 
which the posterior end of the symphysis of the mandible is situated 
considerably in front of the level of the anterior premolar, while in 
the other it is only very little in front of that point. The first 
group, moreover, is distinguished by the comparative simplicity 
of the molars, in which the accessory cusps are scarcely at all 
developed, and by the small size of the talon of the last lower molar; 
into this subdivision the original species, P. beadnelli, falls, 


Amer. Mus. 13471=PHIOMIA WINTONI 

Length from Tip of Symphysis to Posterior Side of 

Length of Symphysis 

Length from Tip of Symphysis to Posterior Side of Mz 

Minimum Antero-posterior Width of Ascending Bar 

Maximum Width of Anterior Half of Symphysial 

Minimum Width at the Constriction of Symphysial 

Height of Ramus at Ps 

Ditto at My; 

Ditto at Anterior Lobe of Ms 

Height of Ascending Bar at Condyle 

[Amer. Mus.] |[Amer. Mus.][Amer. Mus.] 
13469 13471 13475 Andrews 
Young; Young; 
prob. 9 prob. o prob. 2 prob. 
435+ 590+ 600 
137 215 226 
ae 435 475 
122 155 
60 5. bes 55 | 
61 83 70 
59 90 72 
we 82 
112+ 190 


together with a much smaller form for which the name P. parvus 
is now proposed. The type-specimen of this new species is the 
right ramus of the mandible, with the premolars and molars in 
situ, though somewhat crushed.” 

Matsumoto (1922, p. 2) confirms Andrews’ type description 
of 1905 and type figure of 1906 and adds as a single Ameri- 
can Museum specimen No. 13497 from the upper Fluvio-marine 
formation of the Fayfiim. (Matsumoto, 1924.1, p. 4): “Length 
of lower molar series measuring 130 mm.; that of lower pre- 
molar and molar series, 197 mm. (Andrews’ type)... . [P.] 
parvus.” (Op. cit., p. 4): [Referred] Specimen.—{Amer. Mus.] 
No. 13497; a left lower third molar; Amer. Mus. Exp. 1907, 
Quarry B, Fluvio-marine formation, Fayim, Egypt. This tooth 
measures 52 mm. in length and 32 mm. in width. It is longer 
than and as wide as the lower third molar of Andrews’ type, 
which is stated by Andrews to be 46 mm. long and about 32 
mm. wide.” 

Osborn, 1924: Osborn confirms Andrews’ and Matsumoto’s 
opinion as to the specimens of P. parvus, namely, the unique type 
and the referred specimen, and gives a description also measure- 
ments in comparison with Palzomastodon beadnelli and P. 
intermedius (Chap. VI, pp. 148, 146, 147). 

Text-fig. 55. 

Right ramus of mandible of Paleomastodon parvus. Type specimen. 

Fig. 32. Original type of Palzomastodon parvus Andrews, 1905. After 
Andrews, 1906, text fig. 55, p. 163 (see also our Fig. 34C). (Andrews, 1905, 
p. 562): . . . “right ramus of the mandible, with the premolars and molars 
in situ, though somewhat crushed.’’ Brit. Mus. M.8479a; cast Amer. Mus. 
9976. One-fourth natural size. 

Phiomia wintoni Andrews, 1905 
Figures 33, 34, 178, 179, 182, 183; see also page 241 

Fluvio-marine formation of the Fayim, Egypt=Upper Eocene of 
Andrews= Lower Oligocene of the present Memoir. 

Palzomastodon wintoni Andrews, 1905. “‘Note on the Species 
of Palxomastodon,” Geol. Mag., 1905, Dec. V, N.8., Vol. II, pp. 
562, 563 (original description); ‘Descriptive Catalogue of the 
Tertiary Vertebrata of the Fayim, Egypt,’ 1906, pp. 156-162 
(supplementary description), text fig. 538, p. 157. TyPE.— 
(Andrews, 1905, p. 563): . . . ‘a mandible with the incisors and 
posterior molars in situ.” (Andrews, 1906, p. 156): ‘‘A mandible 
wanting the angular region and the anterior cheek-teeth. . . ; 
British Museum.” Brit. Mus. M.8414. Cast (Amer. Mus. 14557) 
of two teeth from type mandible, r. Mo-s. Horizon AND 
Locauiry. — (Andrews, 1906, p. 157): ‘Fluvio-marine beds 
(Upper Eocene [Lower Oligocene]): north of Birket-el-Qurun,” 
Faytim, Egypt. Type Fieurn.—Andrews, 1906, p. 157, 
text fig. 53. 


Text-fig. 53. 

Mandible of Palaomastodon wintoni, type specimen, from above. 

cond., condyle; cor., coronoid process ; d.c., dental canal; ¢., incisor. The premolars and first molar 
on the left side have been restored from another specimen. 

Fig. 33. Original type figure of Palzomastodon wintoni Andrews, 1905. 
After Andrews, 1906, text fig. 53, p. 157. (Andrews, 1905, p. 563): ... “a 
mandible with the incisors and posterior molars in situ.”” (Andrews, 1906, p. 
156): ‘‘A mandible wanting the angular region and the anterior cheek-teeth 
... 3 British Museum.” Brit. Mus. M.8414; cast Amer. Mus. 14557. One- 
eighth natural size. 

Tyre DescripTion.—(Andrews, 1905, p. 563): . . . “the 
other is a much larger form, and is important as_ being 
probably the commonest of all the species; for this the 
name P. wintoni is suggested, in honour of Mr. W. E. de 
Winton, by whom the expenses of my third collecting trip to 
the Faytim were defrayed. It is considerably larger than 
P. parvus and P. minor, but smaller than P. beadnelli. From this 
last species it is also distinguished by (1) the extension back of 
the symphysis so that its posterior end is only just in front of the 
anterior premolar; (2) the position of the mental foramen on 
the side of the symphysis instead of behind it; (3) the greater 
complication of the molars and their greater length in proportion 

The second and third lower molars of: A, Paleomastodon beaduelli (left side); B, P. wintont 

(left side); C, P. parvus (right side); D, P. minor (right side). All % nat. size. 

Fig. 34. After Andrews, 1906, text fig. 50, p. 148, all reduced from 
two-thirds to one-half natural size. Type second and third lower molars, as 
named in the present Memoir. 

A, Type: Palzomastodon beadnelli (left side), type specimen, Cairo 
Museum, My and M3. B, Type: Phiomia wintoni (left side), type specimen, 
British Museum, My and M3. C, Type: Palzomastodon parvus (right side), 
type specimen, British Museum, Mz and M3. D, Type: Phiomia minor 
(right side), type specimen, British Museum, Mz and M3. 

i Sa 


to their width; the last lower molar consists of three transverse 
crests and a distinct talon. The type-specimen is a mandible with 
the incisors and posterior molars in situ.” 

P. winton1 Revision.—Matsumoto (1922, p. 3) points out 
that the immature type of Phiomia serridens {see also P. minor], 
referred specimens of Palzomastodon beadnelli Andrews, 1901, and 
cotypes of Paleomastodon barroisi Pontier, 1907, may all belong 
to the same animal, namely, Phiomia wintoni Andrews, 1905. He 
also adds twenty-seven referred specimens from the American Mu- 
seum collection of 1907. Matsumoto (1924.1, pp. 24-40) gives a 
clear and very able review of the characters of this dominant 
species, referring to it fifty-five specimens in all. 

Phiomia barroisi Pontier, 1907 
Figure 35 

Fluvio-marine formation of the Fayim, Egypt=Upper Eocene of 
Andrews = Lower Oligocene of the present Memoir. 

Synonym of Phiomia minor and of P. wintoni. 

Paleomastodon Barroisi Pontier, 1907. “Sur une espéce 
nouvelle de Paleomastodon (Paleomastodon Barroisi).”’ Ann. Soc. 
géol. du Nord, XXXVI, pp. 150-154. Typr.—Last left 
superior and inferior molars. Horizon AND LOCALITY. 
Bartonien of the Fayim, Egypt. Type FicurEs.—Pontier, 
1907, pp. 150, 151, text figs. 1, 2. 

Fic. 4 
Paleomastudon Barroisi Pontier 
Arriére molaire superieure gauche 
Echelle « 3/5. 
Fig. 35. Type figures of Paleomastodon Barroisi Pontier, 1907. 

Fic. 2 
Paleomastodon Barroisi Pontier. 
Arriére molaire infétieure gauche. 
Echelle : 3/5. 

After Pontier, 1907, pp. 150 and 151, text figs. 1 and 2. 
superior and inferior molars. Three-fifths natural size. 

Last left 

Matsumoto (1922, p. 3) considers that Paleomastodon barroisi 
is a synonym in part of Phiomia minor Andrews, 1904, and in part 
of Phiomia wintoni Andrews, 1905. He observes (1924.1, p. 16, foot- 
note): ‘The last lower molar of Pontier’s two cotypes falls within 
the limit of variation of P. minor; it may belong to the supposed 
male types of the same species.’’ (Op. cit., p. 24, footnote) ‘The 
last upper molar of Pontier’s two cotypes appears to me to belong 
to the presumed female type of the species [P. winton?].”’ 

Osborn, 1924: Osborn confirms Matsumoto’s reference and 
considers that the two cotypes of Palzomastodon barroisi Pontier 
may come within the limits of male and female specimens of 
Phiomia minor and of P. wintoni. 

Meritherium andrewsi Schlosser, 1911 
Figures 2, 36, 37, 41, 42, 42a, 46, 49, 51; see also page 74 
Fluvio-marine formation of the Fayim, Egypt=Upper Eocene of 
Andrews = Lower Oligocene of the present Memoir. 
Meritherium Andrewsi Schlosser, 1911. ““Beitriige zur Kennt- 
nis der Oligoziinen Landsiiugetiere aus dem Faytim: Agypten.’’ 

Fig. 36. Type figure of Meritherium Andrewsi Schlosser, 1911. After 
Andrews, 1906, Pls. v1 and rx; specimens provisionally referred to Meri- 
therium lyonsi by Andrews. Geological Museum, Cairo (C. 7867); cast Amer. 
Mus. 9983. One-fourth natural size. 

Beitr. Pal. Geol. Osterr.-Ung., XXIV, p. 130. Typr.— A 
young, nearly complete skull, Geological Museum, Cairo (C.7867), 
sast Amer. Mus. 9983; paratypes, also based on specimens 
referred by Andrews to Meritherium lyonsi, namely, right and 
left series of lower premolars and molars in almost perfect preserva- 
tion (Brit. Mus. M.8501), right lower premolars of another individ- 
ual (Cairo Mus. C.8127), and right upper premolars of a large 
individual (Brit. Mus. M.8500). Horizon AND LOCALITY.— 
(Andrews, 1906, Pl. rx): “Fluvio-marine beds (Upper Eocene 
[Lower Oligocene]),” Fayfim, Egypt. Tyre Ficure.— 
Andrews, “Descriptive Catalogue of the Tertiary Vertebrata of 
the Fayim, Egypt,” 1906, Pls. vit, 1x (skull and teeth from the 
Fluvio-marine formation provisionally referred to M. lyonsi by 


Matsumoto (1922, p. 5) confirms Schlosser’s definition of 
Meritherium andrewsi based on the larger form from the Fluvio- 
marine (skull, Geol. Mus., Cairo, C.7867) which Andrews had 
referred *to Meritherium lyonsi, type figure Andrews, 1906, Pls. 
vit and 1x. In 1923 (p. 125) he describes MW. andrews as follows: 

Speciric Cuaracters.—“(3.) Larger form of the Fluvio- 
marine Formation. Lower premolars not very short in comparison 
with the length of lower molars; Ps-4, 70 (specimen in the American 
Museum)—73 mm. (Andrews); lower molars rather long: M}-3, 99 
(Andrews)—100 mm. (specimen in the American Museum). Allthe 
lower cheek-teeth are narrow. Ps, fusiform in upper view, its widest 

Andrewsi Schlosser, 1911. 
Oligocene) Fluvio-marine beds. 

Fig. 2. Paratype, Brit. Mus., M.8501. Figs. 3,3a. Para- 
type, Geol. Mus., Cairo (C.8127). Fig. 4. Paratype, Brit. Mus., 
M.8500. Natural size. 

Observe the primitive tetrabunodont form of the Meritheri- 
um andrewsi molars 


Fig. 37. Paratype and referred specimens of Meritherium 

All from ‘‘Upper Eocene” (= Lower 
After Andrews, 1906, Pl. 1x; 
provisionally referred to Meritherium lyonsi by 

(See also our Fig. 36.) 

part corresponding to the middle part. P?*,70-75 mm. (Andrews) ; 
M'®, yet unknown, but M!*, ca. 61 mm. (British Museum, east in 
the American Museum). Skull large and heavily built. Sagittal 
crest rather weak. Zygomatic width rather small in comparison 
with the length of skull. Distance between the two external audi- 
tory openings, as well as the width of occiput, small, being smaller 
even than in the next form [M. trigodon]...........M.andrewsi.” 

REFERRED SPECIMENS (Matsumoto, 1923, p. 129).—‘‘[Amer. 
Mus.] No. 13432; right half of a full-grown skull, without teeth, 
well preserved in the limit of the parts represented. [Amer. Mus.] 
No. 13434; right upper tusk, vz. I?, with well-worn crown. Extra 


no.; left first upper incisor, with imperfectly preserved crown. 
{Amer. Mus.] No. 13437, greater part of a mandible, with the right 
series of cheek-teeth, except P, which might have been acciden- 
tally lost in the life of the animal, and with left Mo; in situ. All 
from the Fluvio-marine Formation of the Fay(im.”’ 

Matsumoto (op. cit., pp. 129,132) observes that while Schlosser’s 
type and the specific name Meeritherium andrewsi are valid, 
Schlosser erred in referring to this species other specimens which 
belonged to Merithertum trigodon. We gives detailed descrip- 
tion and measurements of referred skull (Amer. Mus. 13432) in 
comparison with M. trigodon, as well as detailed comparative 
measurements with the Andrews-Schlosser type; also char- 
acters and measurements of the first upper incisors, I', of the second 
upper incisors, I’, which constitute the tusks, also of the third upper 
incisors, I’, which are rarely found. A referred I* has a triangular 
crown, it measures 14 mm. in length, 8 mm. in width, and 13 mm. 
in height of crown. All three upper incisors of Meritheriwm 
andrewsi appear to have had the crown and root well differentiated. 
In the lower tusks, Iz, one measures 28 mm. in length of crown, 
as compared with 38 mm. in length of crown in I’. 

Osborn, 1924: Osborn confirms Matsumoto’s reference of this 
large Fluvio-marine species, and with the codperation of Doctor 
Matsumoto restores the skull (Fig. 42) of Merithertum andrewsi- 
trigodon partly with the aid of M. lyons materials. 

Paleomastodon minus Andrews, 1905=Phiomia minor 

Palzomastodon minus Andrews, 1905. “Note on the Species 
of Palzomastodon,”’ Geol. Mag., 1905, Dec. V, N. 8., Vol. II, p. 

(Andrews, letter, 1922): “This was only a slip, later in the 
same paper P. minor was used.” 

Palwzomastodon intermedius Matsumoto, 1922 
Figures 38, 39, 49, 88, 91-94, 300; see also page 146 

Fluvio-marine formation of the Fayim, Egypt=Upper Eocene of 
Andrews = Lower Oligocene of the present Memoir. 

Palxomastodon intermedius Matsumoto, 1922. ‘Revision of 
Palzomastodon and Meritherium. Palxomastodon intermedius, 
and Phiomia osborni, New Species,” Amer. Mus. Novitates No. 51, 
November 21, 1922, pp 1-6. iypu;—(Opi cit. pr2)ie 
“fragment of left mandibular ramus bearing all three molars in situ, 
with parts of alveoli of penultimate and last premolars.” Amer. 
Mus. 14547. Paratypes: Amer. Mus. 13480, ‘‘a fragment of left 
mandibular ramus bearing last molar and posterior root of penulti- 
mate molar zm situ.” Amer. Mus. 13449, palate. Amer. Mus. 
14548, maxilla, P*M?. Horizon and Locauity.—Fluvio- 
marine formation (Lower Oligocene), north of Lake Qurun, Fayim, 
Egypt. Paratypes: Amer. Mus. 13480, southwest of Quarry B; 
Amer. Mus. 13449, Quarry A; Amer. Mus. 14548, north of Lake 
Qurun. Type Ficurr.—Op. cit., p. 2, fig. 1. 

Tyre Description.—Matsumoto’s original description is as 
follows (op. cit., 1922, p. 2): 

“ Palzomastodon intermedius, new species. Type: American 
Museum No. 14547; fragment of left mandibular ramus, bearing 

all three molars in situ, with parts of alveo’: of penultimate and 
last premolars.” 

“Paratype: American Museum No. 13480; a fragment of left 
mandibular ramus bearing last molar and posterior root of penulti- 
mate molar in situ.” 

“American Museum referred [paratype] specimens: 
13449, 14548.” 

“All from Fluvio-marine formation.” 


Fig. 38. Palzomastodon intermedius Matsumoto, 1922. Type and 

paratype specimens. After Matsumoto, 1922, p. 2, fig. 1, p. 3, fig. 2. A, 
Type specimen, fragment of left mandibular ramus (Amer. Mus. 14547). 
One-third natural size. External view. B, Paratype specimen, fragment 
of left mandibular ramus (Amer. Mus. 13480). One-third natural size. 
Superior view. 

“The palate of paratype specimen of Palzomastodon inter- 
medius, No. 13499 [13449], measures as follows: ”’ 

Length from the frontal plane tangential to the 
anterior limits of the crowns of the two P? 
to the tip of the posteriorly directed process 
at the posterior limit of the median suture 

between the two palatines.................250 mm. 
Distance between the two P?... . ..... ...... . 53 mm, 
Distance between the two M!.................. 77mm. 
Distance between the two M®.................. 75 mm.” 

“All the upper molars are distinctly bilophodont, as a generic 
character, the rudiment of the third ridge being much feebler 
and much less conspicuous than that of the lower molars. The 
mode of wearing corresponds well to what is stated of the lower 
molars. Besides, all the generic characters of all the cheek teeth 
of this species are the same as those stated in the diagnosis of the 

1924.1, ep. 10).—In this Bulletin Palxomastodon intermedius is 
described and measured in great detail from four specimens. (1) 



laeomastodon intermedius 
Amer. Mus. 13449 Paratype 

1/4 nat. size 

laeomastodon intermedius 
Amer. Mus. 14547 Type 

Fig. 39. Type jaw and paratype skull of Palzomastodon intermedius 
Matsumoto assembled and refigured in lateral view. One-fourth natural size. 

B, B1, Type jaw of P. intermedius (Amer. Mus. 14547). The cavity may 
represent a hollow space within the jaw or a continuation of the alveoli of the 
lower incisors. B1, Crown view of same type jaw. 

A, Al, Paratype skull fragment of P. intermedius (Amer. Mus. 13449), 
lateral view. A1, palatal view of same specimen. 

Compare figure 93 of Chapter VI, enlarged diagram of hexabunodont third 
superior and inferior molars. 

The type (Amer. Mus. 14547); the paratypes (Amer. Mus. 13480), 
a fragment of left mandibular ramus from near Quarry B, also 
“Quarry B [A]. . . [Amer. Mus.] No. 13449; a large fragment of 
skull consisting chiefly of the palate, bearing anterior premolars 
(P?) and all molars of both sides in situ, and with alveoli of penulti- 
mate and last premolars of both sides; Amer. Mus. Exp. 1907. 
[Amer. Mus.] No. 14548; a fragment of skull and palate, bearing 
penultimate premolar to penultimate molar of left side in situ; 
purchase, 1909. All the specimens, Fluvio-marine formation of the 
Fayim, Egypt.’ (2) ‘In all the lower molars [M;-s] the third 
ridge is rather poorly developed, being distinctly narrower than the 


first and second ridges; the posterior valley is distinctly narrower 
antero-posteriorly and shallower than the anterior valley; so that 
the third ridge looks simply like a talon. As a generic character, 
the very bottoms as well as the walls of the valleys were worn, 
even in the very earlier stages of wearing.”’ (3) “All the upper 
molars are distinctly bilophodont, as a generic character, the rudi- 
ment of the third ridge being much feebler and much less conspicu- 
ous than that of the lower molars. The mode of wearing 
corresponds well to what is stated of the lower molars. Besides, all 
the generic characters of all the cheek teeth of this species are the 
same as those stated in the diagnosis of the genus.” 

Phiomia osborni Matsumoto, 1922 
Figures 2, 17, 40, 41, 41a, 90, 171, 185, 185a, 187, 222, 258, 283, 286, 300, 316, 
430; see also page 244 

Fluvio-marine formation, eight miles west of Quarry A, Alexandria 
Trail, Fayam, Egypt=Upper Eocene of Andrews =Lower Oligocene of the 
present Memoir. 

Phiomia osborni Matsumoto, 1922. “Revision of Palzo- 
mastodon and Meritherium. Palzxomastodon intermedius, and 
Phiomia osborni, New Species,’ Amer. Mus. Novitates No. 51, 
November 21, 1922, pp. 1-6. Typr.—(Op. cit., p.3): ... “a 
nearly complete mandible, bearing all the teeth zn sztu.”” Amer. 
Mus. 13468. Type Locatiry.—Eight miles west of Quarry A, 
Alexandria Trail, Fayfiim, Egypt. Type Ficure.—Op. cit., p. 

Fig. 40. Original type figure of Phiomia osborni Matsumoto, 1922. After 
Matsumoto, 1922, fig. 3. Type specimen, mandible (Amer. Mus. 13468). 
One-seventh natural size. Lateral view, right side. 

Typr Drscription.—Matsumoto’s original description (1922, 
p. 5) is as follows: “This species, Phiomia osborni, appears to 
be more progressive than Phiomia minor and Phiomia wintoni in 
the better developed posterior ridge of the first and second lower 
molars and in the better developed posterior talon of the last lower 
molar; and to be more archetypal than the same in the more 
gradual increase in size posteriorly of the lower cheek teeth.” 

CHARACTERS OF PHIOMIA OSBORNI (Matsumoto, 1924.1, pp. 
40-49).—The type (Amer. Mus. 13468) is regarded as a male and 
detailed measurements of mandible and teeth are given in compari- 
son with a supposed female of Phiomia wintoni (Amer. Mus. 13476). 
(1) Judging from the large last molars, Ms, as well as from the 
large symphyseal region and tusks, this mandible is regarded as 
a male with the following principal measurements: 

Length from tip of symphysis to posterior 

SIGeOManple wee. cence 1s ae ee eka 
ene thvofsymphysisee iit sire rare 
Length of premolar-molar series.......... . .250-255 
Length of true molar series..............4.-- 177-180 “ 

640 mm. 



(2) The cheek teeth occupy relatively more space than in the 
supposed female of P. wintont (Amer. Mus. 13476); mandible 
shorter behind symphysis than in P. wintoni female (Amer. Mus. 
13476), see also Andrews, 1906, text figure 54 of P. wintoni. 
(3) Thus this supposed male type of P. osborni appears to have a 
shorter jaw behind the mandibular symphysis than the supposed 
female of P. wintonz; this indicates a post-symphyseal reduction 
of the jaw in P. osbornt. (4) The left and right lower tusks protrude 
58 mm. and 83 mm. respectively from the jaw; each has a distinct 
notch at the tips which may be the result of wear in digging 
and uprooting plants. (5) Symphysis extends back as far as the 
middle part of P;. (6) Lower premolars comparatively large and 
elongate. (7) The last lower molar, M;, extremely elongate and 
comparatively narrow although not beyond the limit of variation 
of M; in P. wintoni. (8) Posterior cheek teeth, Mj-;, increasing in 
size more gradually than in P. wintoni; posterior lobe of P, 
distinctly wider than anterior lobe; differences of proportion are 
observed in the premolar teeth in both P. minor and P. wintoni. 
(9) Posterior talon of M; constitutes an imperfect fourth lobe con- 
sisting of two prominent cusps, besides a few smaller cuspules, an 
important progressive character. (10) Basal cingula of all molars 
very strong. (11) Intermediate cusps (trefoils) well developed. 


Summary (Matsumoto).—This species appears to be more 
progressive than P. minor and P. wintoni in the more strongly 
developed posterior ridge of the first and second lower molars, M;-2, 
and in the better developed posterior talon of the last lower molar, 
M;; and to be more primitive than the same in the more gradual 
increase in size posteriorly of the lower cheek teeth, My-. 
Nothing is yet known about the skull and upper teeth; it 
is of course possible that some of the fragments of the skulls 
and some of the upper teeth referred to P. wintoni may really 
belong to P. osborni, although it is impossible at present to 
be certain of this. 

Osborn, 1924: Osborn confirms Matsumoto’s separation of 
this animal and adds drawings and diagrams showing its distinct- 
ness from Palzomastodon beadnelli and also illustrating the progres- 
sively distinct trilophodont structure of M; and Mb, and the tetra- 
lophodont structure of Ms. 

Meritherium ancestrale Petronievics, 1923 
Figure 47; see also page 76 
Qasr-el-Sagha(?) formation of the Fayam, Egypt=Middle Eocene of 
Andrews = Upper Eocene of the present Memoir. 
A full account and figure of this recently described species of 
Meritherium are given in Chapter III of the present Memoir. 




Palzomastodon beadnelli 
MASTODONTIDA Feet at intermedius 

Palzomastodon parvus 

Phiomia barroisi 
Phiomia osborni 

BUNOMASTODONTIDA® Phiomia serridens 
Phiomia wintoni 
Phiomia minor 

M@RITHERIID® Meritherium trigodon 

Meritherium andrewsi 

Upper Eocene QASR-EL-SAGHA BEps 
IM eritherium lyonst 

M@RITHERIIDA Meeritherium gracile 

\Meritherium ancestrale 


Palzomastodon beadnelli 
Palxomastodon intermedius 
Palzomastodon parvus 

Paleomastodon barroisi 
Phiomia osborni 
Phiomia serridens 
Palzxomastodon wintoni 
Palzomastodon minor 

Meeritherium trigodon 
Meritherium andrewsi 

Meritherium lyonsi 
Meritherium gracile 
Meritherium ancestrale 

According to their present reference these Faytim species, treated in chronological order of description above, 
will be treated in subsequent chapters in the order of their relationships and phylogenetic succession, as follows: 



Palzomastodon: Fam. MASTODONTID®, Subfam. nov. PALAOMASTODONTIN (see Appendix). 


Fam. MG@ERITHERIID®, Subfam. MarirHeriina (Chapter ITI). 

To the above members of the Bunomastodontide should be added the Upper(?) Oligocene species Phiomia 
pygmzus originally described as Mastodon angustidens Cuy. mut. asc. pygmeus Depéret, 1897, and as occur- 

ring in the more recent formation of Kabylie, Algeria. 

ResToRATIONS (1932) OF THE Primitive FaytmM PRoBoscIDEANS 

Uniform reduction to one thirty-sixth natural size 

Fig 41. When compared with the imaginary scene (Fig. 2, p. 18) on the ‘Ur-Nile’ or ‘ Ancestral- 
River-Nile’ of the Faydm region of North Africa, these reconstructions are designed to dispel the wholly 
erroncous notion that Meritherium, Phiomia, and Palzomastodon are in any way ancestral to each other. 
On the contrary, their cranial, dental, and skeletal characters show them to be profoundly different and 
divergent both in habit and habitat. The relative rarity of remains of Palzomastodon is consistent with 
its interpretation as a forest-loving form, in contrast to the more abundantly represented shore-loving 
Phiomia and the still more frequent water-loving Meritherium. 

(Upper) Palzomastodon beadnelli Andrews, after material described by Andrews, Matsumoto, and 
Osborn. Cranium, ears, proboscis, and tusks largely conjectural. 

(Left lower) Phiomia osborni Matsumoto, after cranial and skeletal parts described by Andrews, 
Matsumoto, and Osborn. 

(Right lower) Meritherium andrewsi Schlosser. After materials described by Schlosser, Andrews, 
and Matsumoto, and cranial reconstruction by Osborn (Fig. 42, p. 68). 

Of the three restorations, Palzomastodon is the most hypothetical; as shown in the figures of Chapter 
VI, “The Subfamily Mastodontine,’, besides the skeleton, only the jaw and palate are known, while the 
upper and lower incisive tusks, cranium and mouth parts, and cranio-aural parts are still unknown. 
Phiomia, on the other hand (as shown in Chap. VIII, pp. 236-247), is relatively well known in its cranial 
structure. Mcritherium is also relatively well known (Chap. III, pp. 67-79) both in cranial and limb 
structure. The mouth parts and feeding habits of Phiomia are deductible from our recent knowledge of its 
shovel-tusker descendants (Amebelodon), as conceived and restored by Osborn and Borissiak. 

ac \! , 







1. Subordinal, family, subfamily, and generic characters. Type and characters of Meritherium andrewsi 
Reconstructions of Meritherium by Osborn. Schlosser, 1911. 

Type and characters of Meritherium trigodon 

Andrews, 1904. 

ial’ : Type and characters of Meritherium ancestrale 
Type and characters of Meritherium lyonsi An- Petronievies, 1923. 

drews, 1901. Meritherium molars compared in detail with those 
Type and characters of Meritherium gracile An- of Palzomastodon and Phiomia. 

drews, 1902. Supposed Meritherium of Baluchistan. 

2. Revised descriptions and definitions of the species 
of Meritherium in phylogenetic order. 


Meritherium is very fully treated in the preceding Chapter II, in comparison with the contemporary probo- 
scideans, by a complete review of the opinions and theories of Charles W. Andrews, who first recognized and 
described the genus (1901-1906) and the majority of the species embraced within it. Also see the opinions and 
theories of Hikoshichir6 Matsumoto and those of the present writer on the habits, habitat, and relationships of 
this genus, which, while standing apart from and not directly ancestral to any of the other Proboscidea, displays 
many primitive ancestral characters of the order Proboscidea that are lost in other phyla. 

In the present Chapter III it remains to complete the systematic description of the species of this genus, 
to define clearly the suborder Mceritherioidea and the family characters, to establish the types and type geologic 
levels of the species, to quote Matsumoto and Petronievies in their detailed studies of the cranial structure of 
Meritherium, and to make a very detailed comparison of the grinding teeth of M@ritherium and of Palzomastodon. 


Ref@ A.14./5432 


Reconstruction (1906) or M. trons (Fra. 18). 

Fig. 42. Second reconstruction of the skull of Maritherium (M. andrewsi, M. trigodon), 1920, which is to be compared with the first 
reconstruction of M. lyonsi by Andrews (Fig. 18). 

This reconstruction by Henry Fairfield Osborn with the codperation of Dr. Hikoshichiré Matsumoto assigns chiefly a gliriform 
feeding function to the superior and inferior incisor teeth. There is no evidence that the upper incisor teeth were chiefly weapons or 
tusks, as indicated in the British Museum reconstruction and the model of the skull by Andrews (see Fig. 18). 

A, Al, Top and side views of skull chiefly based on Amer. Mus. 13432 (M. andrewsi), 13430 (M. trigodon). 

B, B1, Top and side views of jaw chiefly based on Amer. Mus. 13437 (M. andrewsi), 13436, 13435 (M. trigodon). 

The American Museum specimens were used as follows: For size and general characters No. 13432; for the first incisor No. 13442; 
for the second incisor No. 13434; the faint lines indicate position of the root of I?-Ip; for the structure of the grinding teeth Nos. 15898 
and 13430; for the structure of the pterygoids and occiput No. 13430. 

_ pr Ere we RP 

500mm.,1'73/4"e 500mm.,1’734"e 699MM., 2’3Ye"e 625mmM.,2’/e"e 650MmM.,2/1v2"e 

Fig. 42a. Ancient Mceritheres of northern Africa. One-hundredth natural size. Restorations by Margret Flinsch, under the direction of 
Henry Fairfield Osborn. 



Original reference: Amer. Mus. Novitates, No. 1, 1921, p. 2 (Osborn, 1921.515). 

SUBORDINAL CHARACTERS.—(1) Palustral, amphibious, and semi-aquatic quadrupeds, with ancestral 
genetic affinities to the Proboscidea in the dentition, and analogies to the Sirenia and Hyracoidea in the 
skull. (2) Vertebre estimated: Cervicals 7, dorsals 19-20, lumbars 4, sacrals 4; vertebral structure indi- 
cating an ambulatory and amphibious habit (fide descriptions of Andrews, 1906, pp. 112-117). (3) 
Girdles, scapule, and pelvis very primitive with analogies to the sirenian type, not expanded as in the 
proboscidean graviportal type. (4) Proportions of humerus and forearm unknown; humerus distally of 
primitive locomotor type. Femur with ligamentum teres pit, greater and lesser trochanter, absence of 
inner trochanter; straight, not unlike Proboscidea. (5) Probably pentadactyl. (6) Cranium primitive, 
greatly elongated, tubular brain cavity, small brain case of archaic type, zygomatic arches slender. (7) 
Face abbreviated, orbits small, nasals abbreviated. (8) Somewhat reduced eutherian dentition: 14, Cs, 
P%, M#. Second incisors, I’-I,, enlarged, curved, gliriform, partly surrounded by enamel. (?) Motion 
of jaw vertical-orthal. 

The subordinal characters are derived from the detailed observations, descriptions, and definitions of Andrews 
(1906, pp. 99-120), as confirmed by Osborn (1907-1923), and as extended by Matsumoto (1922-1923). The most 
recent review is that of Petronievics (1923) cited below under the description of Meritherium ancestrale. 

The numerous archaic and primitive characters in the trunk and limbs of these animals and the 
feeble muscular areas of the scapule and pelvic girdles appear to indicate that they had never passed through a 
long period of terrestrial locomotion either mediportal or graviportal, much less cursorial. Until the complete 
adaptation of the fore and hind limbs and the feet is known we cannot determine how far they were terrestrial or how 
far amphibious. The girdles and limbs are certainly more aquatic in adaptation than those of the hippopotami. 
On the other hand, the Meerithere limbs and girdles are not to be compared for a moment in aquatic adaptation 
with those of the contemporary sirenians. (See Fig. 43.) 

Conc.usion as TO Hasirs.—In the absence of a more complete knowledge of the limbs, we may conclude 
that the Mceritheres were chiefly shore-living or marsh-living animals, amphibious in shallow waters, adapted in 
dentition to a succulent vegetation which could be seized by the opposing upper and lower tusks and by the broad 
lips, which are sirenoid and hippopotamoid rather than elephantoid. 


Original Reference: “A Descriptive Catalogue of the Tertiary Vertebrata of the Fayim, Egypt” (Andrews, 1906.1, p. 99). 

DEFINn1TION.—The family Mceritheriidae Andrews may be redefined as follows: (1) Nares small 
terminal; (2) orbits opening upwards and outwards; (3) auditory meatus elevated, an aquatic adap- 
tation; (4) superior premolars, P**, tritubercular (quadritubercular in Phiomia and Palzomastodon), pre- 
molar tetartocones rudimentary or absent; (5) molars tetrabunodont (quadritubercular), bilophodont, 
with incipient trilophodonty, quadrate or slightly elongate in proportion; (6) superior molars, M' 
strictly bilophodont, M? with rudimentary third crest, M* with enlarged third crest; (7) inferior molars, 
M, trilophodont, rudimentary, M, trilophodont, trilophodonty not pronounced; (8) all three pairs of 
premolars and of molars of both jaws functioning at same time; (9) no trace of trefoil pattern or of in- 
termediate conules in any of the Meritheriwm molars. 

Of the eight characters assigned by Andrews (1906, p. 99) to his family Mceritheriide, five are included above 
as subordinal characters. We do not agree with Andrews when he remarks: . . . “‘probably a small proboscis 
was present,”’ because the structure of the muzzle proves that the animal had a square upper lip. 

The family Mceritheriide stands well apart in its dental adaptations from its contemporaries, either Palzo- 

mastodon or Phiomia. The food was seized by the gliriform incisors and there was an up-and-down or chopping 
motion (vertical-orthal) of the bilophodont, subtrilophodont grinders, as compared with the lateral grinding 


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‘S[ENPIAIPU JUSIOYIP OUIN, *OZIS [BIN}eU paTYyy-ou0 []B ‘G-T “SBY ‘IX [Tq ‘9O6T ‘SMoIPUY 10}j8 UOTONposdar ofIuIsORY “Ep “SIT 


‘dunt weumayy ‘3s2y4 YT 48 Top puempoom Wo 



motion of the molars in Palzomastodon and in Phiomia. The delicate zygomatic arches indicate feeble develop- 
ment of the masseter and temporal muscles. 

These family characters probably represent the climax in a long period of evolution. Close comparison of 
the grinding teeth of Maritherium (Fig. 49A, B, C) with those of Palzomastodon intermedius (Fig. 49D, E) 
indicates that they are tetrabunodont and sublophodont, and it also shows that Mcrritherium differs from 
Palxomastodon in the total absence of any trace of the ‘central conules’ in the molars. 

SUBFAMILY: MarirHeriNni Winge, 1906; MariraertNnas Osborn, 1923 
Original reference: “Jordfundne og nulevende Hovdyr (Ungulata) fra Lagoa Santa, Minas Geraes, Brasilien.” E. Museo Lundii, 
1906, III, p. 172 (Winge, 1906.1); also Amer. Mus. Novitates, No. 99, 1923, p. 1 (Osborn, 1923.601). 

SUBFAMILY CHARACTERS.—(1) First superior and inferior incisors, I'-I,, partly functional. (2) 

Second superior and inferior incisors, I’-I,, greatly enlarged. (3) Third superior incisors, I’, and su- 

perior canines, C, greatly reduced. (4) Third inferior incisors, I;, and inferior canines, C, entirely 

wanting. (5) Inferior molars, M,;, elongated, with rudimentary third lobe (tritolophid) behind the 
protolophid and metalophid. (6) Incisors sheathed in enamel. 

Winge’s definition of the subfamily Mceritheriini, which he included within the family Elephantide, corre- 
sponds with Osborn’s redefinition of the family Meeritheriide above. Inasmuch as Meritherium in its five 
included species, M. lyonsi, M. gracile, M. trigodon, M. andrewsi, and M. ancestrale, are the only Mceritheres 
known, the definition of this subfamily is merely a formal matter. 

It is not unlikely that other subfamilies of Mceritheriide will be discovered, especially if the Mceritheres should 
be traced into southern Asia, as suggested by Pilgrim, and should be found in other parts of the African continent. 

Genus: MC@ERITHERIUM Andrews, 1901-1906 

Original reference: Tageblatt des V Internat. Zool.-Cong., Berlin, No. 6, Aug. 16, p. 4 (Andrews, 1901.2). (Published Volume 
Verhandlungen, 1902, p. 528.) 
Genotypic species: Mcritheriwm lyonsi. 

GrenerRIC Cnaracters.—(Andrews, 1906, p. 99): Dental formula:—i.2; ¢.0; pm.3; m.3. The second 
incisors in both jaws greatly enlarged and tusklike: The’ last premolar not bilophodont; molars bilo- 
phodont. This genus is represented by several species, of which Meritherium lyonsi is the type. It is 
found in both the Middle and Upper Eocene deposits of the Fayim [=Upper Eocene and Lower 
Oligocene of the present Memoir. 

Matsumoto, 1923, p. 121.—After very detailed comparisons, not summed up in a generic definition, 
Matsumoto reaches the following conclusion: “I.—Natural Position of Meeritherium. It may be evident from 
the preceding statements that Meritherium differs strongly from either the hyracoids or the sirenians in many 
characters, on the one hand, and has many proboscidean and pre-paleeomastodont characters on the other hand. 
If much weight should be given to the sirenian resemblances of Meritherium, then more weight should be given 
to its hyracoid resemblances. In my opinion Meritherium is to be treated as a very archetypal member of 
the proboscideans as correctly stated by Andrews at the first.’’ Also: (1) Skull short in comparison with 
zygomatic width; (2) long nasofronto-parietal region in proportion to length of skull; (3) zygomatic arches 
relatively long; (4) sagittal and lambdoidal crests well developed; (5) orbits situated just above P** in Meritherium, 
just above M'* in Palxomastodon; (6) mandible relatively short; (7) diastemata relatively short; (8) ascending 
bars of mandible incline slightly forwards; (9) lower border of mandible ascending; (10) symphysis of mandible 
rather short; (11) upper [and lower] tusks short in comparison with those of the other proboscideans; (12) premolars 
large in proportion to the molars, the series of cheek teeth increasing gradually in size backwards; (13) ridge 
formula simpler than in Palzomastodon, much simpler than in Phiomia; (14) Meritheriwm structurally a pre- 
palzomastodont type, that is, Meritherium stands structurally as more primitive than Palzomastodon in twenty- 
three characters which are peculiar to it and also which indicate its remote relationship to Palzomastodon. 


Compare original descriptions of the same species in Chapter II, wherein the chronological order of description is clearly set forth. 
Consequently the reader is referred to both Chapters II and III and to the more recent paper of Petronievics for our present knowl- 

edge of the species of the genus Meritherium. 

Meritherium lyonsi Andrews, 1901 
See Chap. II, p. 54 
Qasr-el-Sagha formation of the Fayim, Egypt=Middle Eocene of An- 
drews= Upper Eocene of the present Memoir. 
The establishment of this species by Andrews and its descrip- 
tion, based on forty-six specimens contained in the British and 
Cairo museums from the Qasr-el-Sagha beds and one specimen in 

short: P24 [P24], ca. 69 mm. (Andrews); lower molars very long, 
M,3, ca. 104 mm. (Andrews). All the lower cheek-teeth very 
wide. P», triangular, its widest part corresponding to the posterior 
lobe. P?4,67-78 mm.; M'?,85mm. (Andrews)..... M. lyonsi.” 
It is this genotypie species M. lyonsi of the Upper Eocene 
Qasr-el-Sagha formation which furnishes the chief characters of the 

genus Meritherium as described and figured in great fullness by 

Fig. 44. Second type figure of Meritherium lyonsi Andrews, 1901. 
Cairo (C. 10000); cast Amer. Mus. 9977. One-third natural size. 

After Andrews, 1906, Pl. x, figs. 1, 14, 2, 8. Geol. Mus., 

Three individuals: (1, 14) Type specimen (C. 10000), mandible associated with upper molars and a dorsal vertebra (op. cit., 
p. 120); (2) “Upper molars and premolars found close to type” (p. 123), of the right side = paratype (C. 10001); (3) front portion 

of skull, side view (C. 10002, cast Amer. Mus. 9978) = paratype. 

the American Museum collection, are in part set forth in the 
systematic revision of this Memoir above, pages 54, 55. 

Speciric Cuaracters.—(Matsumoto, 1923, p. 124): ‘‘(1.) 
Larger form of the Qasr-el-Sagha Formation. Lower premolars very 


Same as figure 27. 

Andrews (1906, pp. 99-126, Pls. v1, rx, and x). The detailed 
measurements and indices of the superior and inferior teeth in com- 
parison with those of M. gracile, M. andrewsi, and M. trigodon are 
transcribed and estimated from Matsumoto as follows: 

Lower Teeth M. lyonsi M. gracile M. andrewsi M. trigodon 

l. Ww. Ip I Ww. iy l. Ww. | eal l. Ww. I. 
P, 22 16 73 24-26 11 -14 54 | 19.5-23 10 -11 48 
P; 23 21 90 22 15 68 | 27-29 17.5-19 65 | 22 -25 14? -17 68 
P, 25 23 90 21 18 86 | 24-25 21 87 22 -25 18 -20 80 
M, 26.5 24.5 92 22-23 19-20 87 | 28-30 21 -22 73 24 -27.5 19 —-21 76 
M, 28 -35 25 -30 86 27-28 22-23 82 32-34 26 -27 79 | 29 -382 24 83 
M; 39 -42 28 -30 71 33-34 24 72 37-40 26 -28 70 | 37 -40 24 -26.5 66 
P2-5 69+ 63+ 70-73 63+--70 
Mi. 104+ 84 | 99-100 93 -98 
I, width 20+ 28 | 20 

Upper Teeth 

iE? 27 23? 85 22 18 82 25-27 20 -22 81 | 19 -22? 18? -21 95 
Ps 26.5 29.5 111 20 23 115 25-26 28 -32 123 23 -24 24? -26 108 
P 23 27.5 119 20 21? =105 21-22 27 -29 132 | 20 -22 23? -25.5 116 
M' 29 27 93 23-25 21-23 92 29-32 25 -27 84 22 -25 24 -26.5 106 
M? 26? -30 23.5-28 93 24-27 23-25 92 | 32-33 28 -29 88 28 -30 24 -28 93 
M* 32 -37+ 28 -30+ 81 28 24-25 86 | | 380 -32 26 -27 84 
PA 67 —68 62 70-75 | 60 -63 
M* 85 | 75-79 | 83 -85 
I? width 28 26 21 

American Museum Couiection.—Matsumoto 
refers (1923, p. 125) to the species Meritherium lyonsi 
two of the three fragments of mandibular rami 
included in Amer. Mus. 13444; the other frag- 
ment of a ramus he refers (op. cit., p. 128) to M. 
gracile. his descriptions being included above in 
Chapter II, pages 54 and 56 respectively. 

Meritherium gracile Andrews, 1902 
See Chap. II, p. 56 
Qasr-el-Sagha formation of the Fayim, Egypt = Middle 
Eocene of Andrews= Upper Eocene of the present Memoir. 

This small form of the Qasr-el-Sagha beds 
Upper Eocene, is relatively rare, four specimens 
being described by Andrews from the British and 
Cairo museum collections while four are described by 
Matsumoto and Osborn from the American Museum collection. 

Speciric CuHaracrers.—(Andrews, 1906, p. 127): “This 
species is distinguished from M. lyonsi by its comparative lightness 
of structure, the narrowness of the palate, the smaller size of the 
upper molars and premolars, particularly of m. 3, the strong de- 
velopment of the cingulum in these teeth, and by the considerable 
inflation of the cranial region of the squamosal, which apparently 
contains extensive air-sinuses.”’ 

(Matsumoto, 1923, p. 125): ‘(2.) Smaller form of the Qasr- 
el-Sagha Formation. Lower premolars not very short in compari- 
son with the length of lower molars: Pe-s, ca. 62 mm. (specimens in 
the American Museum); lower molars very short: M3, 83 mm. 
(specimen in the American Museum). All the lower cheek-teeth 
are narrow. P*4, 62 mm. (Andrews); M'*, 75—ca. 79 mm. 
RRTELEGS CUE) MRED aah Rates ANS olor, aya wien Sees M. gracile.” 

Am.Mus.No 13443 


Fig. 45. Meritherium gracile and M. lyonsi from the Qasr-el-Sagha for- 
mation, Upper Eocene of the Fayim. American Museum collection of 1907. 
After Matsumoto, 1928, figs. 3 and 4. 

Amer. Mus. 13443, M. gracile; mandible with P3-Mg of the left side 
and M;-3 of the right side in situ. 

Amer. Mus. 13444, M. lyonsi; one of the three fragments of the man- 
dibular rami, with badly preserved molars in situ. 

Andrews (op. cit., 1906, pp. 127, 128) gives a detailed descrip- 
tion with measurements of the four specimens in the British Mu- 
seum; Matsumoto (op. cit., 1923, p. 128) also gives a detailed 
description with measurements of the specimens in the American 
Museum collection. In his comparison of the four species, Mat- 
sumoto’s detailed dental measurements are as above. 

PuyLoGENetic SummMAry.—From the limited materials at 
hand neither Andrews, Matsumoto, nor Osborn has been able to 
establish the phylogenetic or ancestral relationship of the Qasr-el- 
Sagha species (M. lyonsi and M. gracile) to the Lower Oligocene 
Fluvio-marine species (M. andrewsi and M. trigodon). 

We might suppose that the larger form M. lyonsi gave rise to 
the larger form M. andrews, but Matsumoto does not believe so. 
(1) Matsumoto observes that the larger Upper Eocene species M. 
lyonsi seems to be less closely related to its smaller contemporary 
M. gracile than the larger Lower Oligocene form M. andrewsz is to 
the smaller Lower Oligocene form M. trigodon; this inference is 
drawn from the absolute and relative dimensions of the grinding 
teeth. (2) M. andrewsi and M. trigodon appear to represent two 
entirely distinct species and not male and female representatives 
of the same species, as Schlosser has suggested. (3) So far as can be 
judged from the relative dimensions of the lower cheek teeth, M. 
gracile appears tq be more closely allied to the two geologically 
succeeding species M. andrewsi and M. trigodon than to its con- 
temporary M. lyonsi; in other words, it appears that M. gracile 
might be the ancestral type of both M. andrewsi and M. trigodon. 

Meritherium andrewsi Schlosser, 1911 
See Chap. II, p. 61 

Fluvio-marine formation of the Fayim, Egypt=Upper Eocene of An- 
drews = Lower Oligocene of the present Memoir. 

This palustral animal, based on specimens referred to M. 
lyonsi by Andrews, occurring in the Fluvio-marine, beds with a 
rich terrestrial fauna, is naturally much less abundant than its 
predecessors in the Qasr-el-Sagha marine beds; it is represented by 
twelve specimens in the British Museum (all referred by Andrews 
to M. lyonsz) and by three specimens of M. andrewsz in the Ameri- 
can Museum, two of which (Amer. Mus. 13432 and 13437) have 
been largely used in the second reconstruction of the skull of 
Meritheritum shown in figure 42. As set forth in the systematic 
revision above (p. 61), Schlosser and Matsumoto have clearly 
separated and defined this larger form as a distinct species, M. 
andrewsi, named in honor of Charles W. Andrews. 

Speciric CHARAcTERS.—Matsumoto’s characterization (Mat- 
sumoto, 1923, p. 125) of this species is cited in full on pages 62, 63 
of this Memoir. These characters agree with the reconstruction of 
the skull by Osborn and Matsumoto shown in figure 42, in which it 
appears that there are considerable differences in size and propor- 
tions between the larger M. andrewsi (Fig. 36) and the smaller 
M. trigodon (Fig. 30). 

Meritherium trigodon Andrews, 1904 
See Chap. IT, p. 57 

Fluvio-marine formation of the Fayim, Egypt=Upper Eocene of An- 
drews = Lower Oligocene of the present Memoir. 

American Musnum Couiection.—Meritherium trigodon is 
represented by the single type specimen (Brit. Mus. M. 8499) 


described by Andrews in 1904, and by six specimens in the American 
Museum collection, described, figured, and measured by Matsumoto, 
namely, Amer. Mus. 13430, 13431, 13433, 13435, 13436, 13439. 

The cranium is inferior in all its dimensions to the larger and 
more massive M. andrews? and there appear to be minor differences 
in the characters of the median and lateral incisors, I*, I. A well- 
preserved second right upper incisor, I’, with its casing of enamel 
(Fig. 42), illustrates the profound difference between the gliriform 
second superior incisor of M. andrewsi and the superior tusk of 
Phiomia (Fig. 183); observe especially the posterior surface 
beveled by the action of the second inferior incisor in contrast to 
the tusk of Phiomia which is worn on the internal side. Observe 
also the profound difference between the lower jaw in superior 
aspect (Fig. 42) and the lower dentition of Phiomia and Palzo- 
mastodon (Fig. 48) as seen in superior view; also the uniformly 
worn grinding series, P2-M;, of Amer. Mus. 13485 (Fig. 46), as 
compared with the successional wear in Phiomia. 

Speciric CHaracters.—(Andrews, 1906, pp. 128, 129—see our 
Fig. 30): “Type Specimen.—Posterior portion of right ramus of 
mandible (Pl. 1x, fig. 5); British Museum. This species is dis- 
tinguished by the form of the posterior lower molar, which nar- 
rows posteriorly, the talon consisting almost entirely of one large 
tubercle; also by the rapidity with which the teeth decrease in 
size from behind forwards. . . . This difference in the talons 
appears to justify the separation of the present form as a distinct 
species at least, and not improbably further material will show 
that a new genus must be established. The enamel of the whole 
tooth is raised into irregular ridges and small tuberosities. The 
dimensions (in centimetres) of the teeth are:— 

Length. Width. 
1s li, AOE eee 2.6 im 
1 EER RO EER has, Sines with 3.2 ? 
LOS ya RR Oe ec Bie RENE ost fOr 4 2.4 

The length of the molar series is 9.8 em.” 

Matsumoto’s definition (1923, p. 125) is cited above (pp. 57, 
58) under the systematic revision. He considers that Andrews lays 
too much weight on the shape and structure of the talonid or 
tritolophid of M; in distinguishing this species from M. andrewsi, 
remarking that the posterior talon of M; appears to be one of the 
most variable features in the teeth of Meritherium, and that its 
surface varies according to age and the degree of wearing action 
of the tooth. 

CRANIAL CHARACTERS.—Matsumoto observes (1923, p. 134) 
in his description of the skull: ‘The skull of [Amer. Mus.] No. 
13430 measures 240 mm. in the length from the point at which the 
vertical plane tangential to the anterior borders of both P? meets the 
median longitudinal line on the palate, to the basion, 145 mm. in 
the length from the same point to the median point of the posterior 
border of palate, about 180 mm. in the length from the anterior 
limit of the temporal vacuity in palatal view to the posterior lower 
border of squamosal, 42 mm. in the minimum width of the mid- 
cranial region (this dimension might be somewhat less than it 
ought to be in primary condition, as this part of this specimen 
appears to be crushed secondarily from side to side), 280 mm. in the 
zygomatic width, about 190 mm. in the distance between the upper 
borders of external auditory openings (including reliably restored 


Am.Mus.Na 13430 

Fig. 46. American Museum 
Collection of 1907. After Matsu- 
moto, 1923, figs. 1, 2, 6, 7, 8, and 9. 

A, B, Amer. Mus. 13430, M. 
trigodon, a full-grown skull, lateral 
and superior views; partly restored 
occipital and sagittal crests, nasal 
and symphyseal region, and anterior 
teeth. This skull is described and measured in detail hy Matsumoto (1923, p. 134) in his 
description of M. trigodon. About one-fourth natural size. (1) Observe the excessively 
slender, widely arching zygomata, the tubular cranial region, the slender sagittal and 
lambdoidal crests. (2) Zygomatic width 280 mm. 

C, Amer. Mus. 13435, M. trigodon, fragment of the right ramus of the mandible, all 

the cheek teeth, Po-M3 in situ. One-fourth natural size. 
- Am,Mus.No.13435 D, Amer. Mus. 13434, Meritherium andrewsi Schlosser, second upper right incisor 
tooth, I2; upper, internal view, lower, external view. One-half natural size. Through error 
this appeared in the original legend (Matsumoto, 1923, p. 126) as M. lyonsi. 

E, Amer. Mus. 13436, Meritherium trigodon, “ramus of mandible, with all the cheek 
teeth in situ’’ After Matsumoto, 1923, p. 126, fig. 1, through error referred to M. lyonsi 
in the legend. 

F, Amer. Mus. 13431, M. trigodon, fragment of skull including greater part of right half 
of palate bearing P°-M® in situ. One-fourth natural size. 


Am. Mus No!3431 


parts to a slight extent), 220 mm. in the width of occiput, 91 mm. 
in the lateral extension of the two occipital condyles, 52 mm. in the 
width of palate between the two P?, 34 mm. in the same between 
the two M!, and about 150 mm. in the maximum height of the 
skull, including the upper cheek-teeth (including a restored part of 
sagittal crest to a slight extent). In this skull the sagittal and 
occipital crests are extraordinarily well developed; and the occiput 
is strongly concave, so that its upper half, including the lambdoid 
crest, inclines distinctly backwards.” 

Meritherium ancestrale Petronievics, 1923 

Qasr-el-Sagha(?) formation of the Fayim, Egypt=Middle Eocene of 
Andrews = Upper Eocene of the present Memoir. 

Meritherium ancestrale is regarded by Petronievics as nearer 
to Palzomastodon than any previously described species of 

Meritherium ancestrale Petronievics, 1923. “Remarks upon 
the Skulls of Meritherium and Palzomastodon,” Ann. Mag. Nat. 
Hist., Vol. XII (9), pp. 55-61. Typr.—Skull and mandible 
(Brit. Mus. M. 9225). Horizon anp Locautity.—Fayim 
district of Egypt, Qasr-el-Sagha (?) formation= Middle Eocene of 
Andrews= Upper Eocene of the present Memoir. Type Fic- 
URE.—Petronievics, 1923, p. 56, fig. 1. 

Fig. 1. 

Part of the left side of the skull of Maritherium ancestrale, 3 nat. size. 
as., alisphenoid; pt., pterygoid; sg., squamosal. 


Fig. 47. Type of Meritherium ancestrale Petronievics, 1923, p. 
56, fig. 1. Reproduced same size as the author’s original figure and 
including the author’s legend. Brit. Mus. M.9225. 

Type Description.—(Op. cit., 1923, p. 57): “There are two 
main characters of our specimen which justify the supposition that 
it represents a distinct species:—(1) the elongated palatine vacuity 
between anterior maxillary prolongations forming the inner border 
of the sockets for the tusks; (2) the backward direction of the 
occipital plate. As the first of these two characters brings our 
specimen nearer to Palzomastodon than any other, we will give to 
the new species the name Meritherium ancestrale, sp. n.”’ 

The following is a direct quotation, with some omissions, 
from the description of Doctor Petronievics (op. cit., pp. 55-57): 

“{1] There is in the British Museum [M.9225] a skull and 
mandible of Meritherium not yet described, which is in several 
respects important for deciding the question of the ancestral 

relationship of Meritherium to the Proboscidia. This skull was 
found by Baron Nopesa several years ago (1905) in Middle Eocene 
beds in the Faytim district of Egypt [Footnote: ‘Comp. C. W. 
Andrews, ‘A Descriptive Catalogue of the Tertiary Vertebrata of 
the Fayim, Egypt’: London, 1906, p. 121 seg. (specimen M. 9225 
in the British Museum).’]. I propose to describe the features of 
this skull, which either give some supplementary data as to the 
morphology of the skull of the genus Meritheriwm or are peculiar 
to the species. I propose also to give some additional evidence on 
the skull of Palzomastodon. Finally, I will add a list of the re- 
semblances and differences in the skull, mandible, and dentition 
of the two genera. [2] In the skull in question the basioccipital is 
separated from the basisphenoid by a suture situated somewhat 
obliquely on a ridge at the level of the glenoid surface. The suture 
between basioccipital and exoccipitals is not plain. The exoccipi- 
tals with the condyles are as in Meritherium lyonsi. [3] The same 
is to be said about the supraoccipital bone; but the whole occipital 
plate in our specimen is directed backwards, while in Meeritherium 
lyons? itis directed forwards, and the lateral depressions accompany- 
ing the median ridge are deeper in our specimen than in the old 
one. [4] The squamosal and its relation to the parietal and supra- 
occipital are the same as in Meritherium lyonsi. [5] The jugal 
seems to be more slender than in the latter. [6] The tympanic in 
this specimen is better preserved than in the skull of Meritherium 
lyonsi. [7] The parietals are as in the latter. It is uncertain if 
they meet the alisphenoids—probably not. [8] The frontals are 
as in Meritherium lyonsi, with no postorbital process. The exact 
relation of the frontals to alisphenoids and parietals is uncertain, 
but they reach the former. [9] The existence of a separate lachry- 
mal cannot be established. [10] While in Meritherium lyonsi the 
position of the alisphenoid on the side-wall of the skull could not 
be determined, in our specimen the limit of the alisphenoid in this 
respect is quite clear (text-fig. 1). Above the alisphenoid canal 
the alisphenoid forms a triangular bone. The backward side of 
this triangle is constituted by the suture separating the alisphenoid 
from the squamosal, while the upperside is marked by a suture 
separating the alisphenoid from the frontal (perhaps also from the 
parietal). It is to be remarked that the position of this part of the 
alisphenoid bone in our specimen is, perhaps, exactly the same as in 
Palzxomastodon. [11] As the premaxille are crushed in front, their 
relation to the nasals and maxill# cannot be made out. [12] The 
maxille are very large and greatly elongated as in Merithercum 
lyonsi; but, while in this latter there is only a small palatine vacuity 
between the slightly divergent anterior prolongations of the 
maxille, this vacuity is larger in our specimen, and the anterior 
prolongations of the maxillz undoubtedly form part of the inner 
border of the sockets for the tusks, although they are somewhat 
crushed anteriorly. In this respect our specimen is nearer to 
Palzxomastodon than any other species of Meritherium. Meri- 
therium gracile has these prolongations longer than Meritherium 
lyonsi, but they do not reach the inner border of the tusks (comp. 
Andrews, l. c. pl. xvii. fig. 1). [13] The exact shape of the ptery- 
goid is not to be established, but it is certain that its vertical plate 
projects beyond the end of the descending plate of the alisphenoid 
(fig. 1). [14] The upper teeth are so worn that their exact structure 
is not to be made out; but they seem to be the same as in Meri- 
therium lyonst.” 


The author follows with a very detailed comparison of the 
cranial characters of the type of Meritherium ancestrale with those 
of Palzxomastodon beadnelli and P. wintoni Andrews, and concludes 
(pp. 58, 59): “The ancestry of Me@ritherium to Palxomastodon 
has become very probable from the new additional features of the 
skull in the above-described specimen of Meritherium. But, in 
order to make it possible to judge better on this question [Foot- 
note: ‘On this point three main opinions have been advanced :— 
(1) Meritherium lies in the direct line of ancestry to the later 
Proboscidea; (2) Meritherium is a primitive offshoot of the Probo- 
scidean stock; and (3) Meritherium is an offshoot of the Sirenian 
stock allied to Proboscidea. The first opinion has been maintained 
by Andrews in his first fundamental memoir on the question (comp. 
C. W. Andrews, ‘The Evolution of the Proboscidea,’ in Phil. 
Trans. 19038, p. 101), but later on (comp. his Catalogue, Introduc- 
tion, p. xvii) he was rather inclined to adopt the second view. The 
third opinion was expressed for the first time by H. F. Osborn (in 
his article ‘The Feeding-habits of Meritherium and Palzomastodon,’ 
in Nature, July 29, 1909), but abandoned (comp. his work ‘The 
Age of Mammals,’ 1910, p. 203 seq.) after the vigorous refutation of 
Andrews (in his article ‘The Systematic Position of Meritherium,’ 
in Nature, Sept. 9, 1909). In a recent paper, however, the 
eminent American palzontologist seems to have returned to his 
former wrong opinion on the subject (comp. H. F. Osborn, ‘ Palzxo- 
mastodon, the Ancestor of the Long-jawed Mastodons only,’ in 
Proc. Nat. Acad. Sci. vol. v. 1919, pp. 265-266).’], we give here a 
list of the resemblances and differences in the skull, mandible, and 
dentition of the two genera [Footnote: “A similar but much shorter 
list is to be found in W. K. Gregory, The Orders of Mammals, 
1910, p. 367 seq.’], which are mainly founded on the characters of 
Meritherium ancestrale on the one hand, and of Palxomastodon 
wintoni on the other.”’ : 

Osborn, 1925: In Chapter II of the present Memoir many 
reasons are given by both Matsumoto and Osborn why we should 
exclude Meritherium entirely from the ancestry of Palzomastodon. 


Figures 42, 45, 46, 48, 49 

The five new figures and reconstructions of the grinding teeth 
assembled in the present Memoir, namely, figure 48, a comparison 
of the grinding teeth of three genera, figure 45, a comparison of 
Meritherium lyonsi and M. gracile, figure 46, a comparison of M. 
trigodon and M. andrewsi, figure 42, second reconstruction of the 
skull of Meritherium andrewsi and M. trigodon, and figure 49, 
life-size detailed studies of Meritherium and Palzxomastodon 
grinding teeth, exhibit very clearly the fundamental resemblances 
and the progressive differences between the dental and cranial 
structure of these three Fayim genera. Figure 49 beautifully 
displays the superior grinders of Meritheriwm trigodon in contrast 
with those of Palzomastodon intermedius, demonstrating that 
Palzomastodon cannot in any way be considered as a descendant of 

(1) In both Meritherium and Palzomastodon Ms; is trilophodont 
(Fig. 49C, E); (2) in both Meritherium and Palzomastodon the 
second and third superior molars, M**, are in an incipient bilopho- 
dont condition; (3) in Palzomastodon M,-; have a rudiment of a 


Amer. Mus. 13449 


AQ Mosritnerium . 
. Amer. Mus. 13431 Ret. (rev.) 

All 1/3 nat size 

Amer. Mus. 13468 

B Paixomastodon 
Amer. Mus. 14547 

A Moeritherium 
Amer. Mus. 13437 Ret. (rev. 

Fig. 48. Comparison of upper and lower molars to the same 
scale of: 

A, True Meritherium, (lower) Amer. Mus, 13437, (upper) 
Amer. Mus. 13431. 

B, True Palzomastodon, (lower) Amer. Mus. 14547, (upper) 
Amer. Mus. 13449. 

C, True Phiomia, (lower) Amer. Mus. 13468, (upper) Amer. 
Mus. 13450. 

All 1/3 nat size 

tritolophid not observed in Meritherium; (4) Palzomastodon 
is hexabunodont or sexitubercular, conules persisting in M**, while 
Meritherium is tetrabunodont or quadritubercular, the conules 
having disappeared; (5) the inner and outer cones are united (Fig. 
49D, E) by a transverse crest in Palzomastodon, M**, Mo-s; 
they are separated by a longitudinal cleft in Meritherium. (6) 
As to adaptation, this demonstrates that Maritherium had more 
of the omnivorous, cheerodont habit, while Palzomastodon had 
more of the browsing, lophodont habit. The evolution of the 
Palzomastodon molars is discussed in full below under Palzomas- 
todon (Chap. VI). 

ConcLusion.—This detailed comparison serves to demonstrate 
that both Meritherium and Palzomastodon may have descended 
from an original primary tetrabunodont stock; that the inter- 
mediate tubercles or ‘central conules’ never developed (Fig. 37) in 
Meritherium, while they did develop in Palzomastodon inter- 
medius, M*8, Mos. These conules, moreover, are in process of 
development in Palzomastodon, the lophs being composed of six 
cones (Figs. 93, 94). 



In 1912 Pilgrim erroneously referred to Meritherium a frag- 
mentary tooth (Ind. Mus. A 452) contained within “The Verte- 
brate Fauna of the Gaj Series in the Bugti Hills and the Punjab,” 
found near Khajuri, Bugti Hills, Baluchistan, which he described 
as follows (p. 15): “ Mceritherium (?), sp. Plate mu, fig. 3. Amongst 
the Bugtispecimensis . . . a fragmentary tooth, possessing char- 
acters which lead one, with no uncertainty, to diagnose it as Probo- 

AM. 15898 €ast) 

AM. 343) 



to render it possible to recognize its presence. The incompleteness 
of the transverse crests entirely precludes the possibility of its 
belonging to Dinotherium. It is clearly distinguished from premolars 
of Tetrabelodon, not only by the rectangular instead of rounded 
corners, but also by the perfectly symmetrical arrangement of the 
cusps, and the breadth and absolutely transverse situation of the 
valley between the crests. The surface of the enamel also possesses 
irregular strie, which are not seen in Tetrabelodon. In all these 



Fig. 49. Meritherium (A, B, C) and Palzomastodon (D, E) grinders figured and compared in close detail, demonstrating 
that the hexabunodont grinders of Palexomastodon contrast with the tetrabunodont grinders of Meritherium. See same figure in 

Chapter VI (Fig. 94), see also key figure 93. Natural size. 

A, Maritheriwm lyonsi ref. (Amer. Mus. 15898 cast). First and second right superior molars, M!-M?. 

B, Meritherium trigodon ref. (Amer. Mus. 13431). Right superior grinding teeth, P?-M°. 

C, Meritherium andrewsi ref. (Amer. Mus. 13437). Second and third inferior grinders, Mo-M3. 

D, Palzomastodon intermedius paratype (Amer. Mus. 13449). Third right superior molar, r.M°. 

E, Palzomastodon intermedius type (Amer. Mus. 14547). Third left inferior molar, 1.M3 (reversed in drawing). 

scidean. Although only three quarters of it remain there is no 
doubt as to its being a rectangular, oblong tooth, possessing two 
transverse crests, separated by a broad straight transverse valley. 
Each crest consists of an inner and outer pointed cusp, perfectly 
distinct from one another. The presumed postero-internal cusp is 
continued into a small hinder lobe. A beaded cingulum is well 
developed on that side of the tooth, whose preservation is such, as 

particulars, it agrees exactly with the hinder upper molars of Merv- 
therium lyonsi, Andrews. It is, however, rather larger than this spe- 
cies, and the relative height of the cusps differs. It is by no means 
unlikely that further material of the Bugti species will show that it 
belongs to a different genus from Meritheriwm. At present its 
interest lies in the proof it affords of the existence in Baluchistan in 
Gaj times of a primitive Proboscidean allied to Meritherium. 


3. (7) 


Fig. 50. Incomplete fourth superior permanent premolar, P‘, possibly 
referable to T'rilophodon pandionis. Originally referred by Pilgrim to Meri- 
therium. After Pilgrim, 1912, Pl. m1, figs. 3, 3a: “Meritherium (?) sp. Pilg. 
Upper molar, 3, surface view, 3a, side view. From the base of the Gaj near 
Khajuri, Bugti Hills. (Ind. Mus., No. A 452.) Page 15.” 

Provisionally, therefore, it may be regarded as an indeterminate 
species of the Egyptian genus. The measurements of the tooth 
are approximately as follows:— 


Vy, ie 

.42 mm. 
i)! We 

Length. . 
Breadth.... . 

Locality—The specimen was found at Khajuri, near the very 
base of the series, and hence, is probably lowermost aquitanian.”’ 

Osborn, 1926: The supposed Meritherium superior molar of 
Baluchistan proves to be a fourth superior premolar, P‘, probably 
referable to Trilophodon pandionis, a characteristic species of 
Sind, India. The original comparison by Pilgrim of this tooth 
with Meritherium lyonsi, as quoted above, was qualified by his 
remark: “It is by no means unlikely that further material of the 
Bugti species will show that it belongs to a different genus from 
Maritherium.’’ Similar tetrabunodont superior premolar teeth 
occur as P4-P, both in Trilophodon and in Serridentinus; such a 
tooth was selected by Cope as the type of his ‘Mastodon’ proavus 
(=Serridentinus proavus of the present Memoir), as illustrated 
in figures 362 and 363. 

Unfortunately Pilgrim’s original reference of this premolar 
tooth to Meritherium(?) has been widely quoted as demonstrating 
the presence of the genus Meritherium in India. 

Fig. 51. Restoration (September, 1932) of M@ritherium andrewsi, after materials described and figured by Andrews (1906), Schlosser 

(1911), and Matsumoto (1922-1923), and reproduced in figures 41 and 42. 

One-sixteenth natural size. 

This restoration embodies the Andrews-Matsumoto-Osborn conception of Meritherium as of a swamp- and river-loving stock. As 
detailed above, the conclusion is, that it was a confirmed and continual river-living animal, feeding mainly under water and on the banks, 
as indicated by its feeble pelvis, more specialized for aquatic life than Hippopotamus, but less specialized than the Sirenians. The resem- 
blances to the Sirenians are largely parallelisms. Mcritherium is one of the primary stocks of the Proboscidea. The marked resemblances 
to the river-living Hippopotamus are also parallelisms, namely, the small, elevated orbits, with small eyes and probably small ears, the 
elevation of the eye and ear openings to the upper level of the face, the prominent anterior tusks, the relatively short, stout limbs, probably 
encased in fat. The habitat was on a river-bed meandering over a low-grade flood-plain, as indicated in the restoration. 



\ AY 

Koupad nat oe. 

Fig. 52. Type of Deinotherium giganteum as originally figured by Kaup, with the incisive tusks erroneously upturned, 
1829, XXII, Heft IV, Taf. 1, one-ninth natural size. 

Fig. 53. Type of Deinotherium giganteum as refigured by Kaup, 1835.1, Add. Tab. 1, fig. 5, 
with incisive tusks downturned. One-fifth natural size. Original in Museum of Darmstadt. 
Cast Amer. Mus. 14286. 

The date of publication of this figure is somewhat doubtful, but it would appear (see 
Kaup, 1841.1, p. 18) that this was prepared to supplement Kaup’s Cahier IV of the ““Ossements 

Fig. 54. Superior dentition of Deinotherium giganteum ref., one- 
fourth natural size. Copied from Gaudry, 1878, p. 187, fig. 247. 






1. Chief distinctions of the Deinotheres from other proboscideans. 4. Conclusions as to origin, phylogeny, and adaptive radiation. 
2. History of discovery of the Deinotheres between 1715 and 5. Geologic distribution of Deinotherium, Trilophodon, Anancus, 

1934. Rhynchotherium, Zygolophodon, Turicius, Miomastodon, 
Historical summary by Weinsheimer, 1715-1882. Serridentinus, and Tetralophodon. Causes of extinction. 
Historical summary by Osborn, 1883-1934. Appendix: Deinotherium hungaricum Bhik and Deinotherium hop- 

3. Résumé of the chief generic and specific characters. woodi sp. nov. 


It is not possible for the author to treat the Deinotheres in the same critical or exhaustive manner as the 
species of proboscideans belonging to other families are treated in the present Memoir; nor is it possible to give 
the type descriptions or reproductions of the type figures as in other chapters. Such exhaustive revision and 
monographic treatment can only be made after a tour of the museums of Europe and of India where the actual 
type specimens are to be found and the precise records of the type localities are given. 

In the succeeding historical summaries by Weinsheimer and by Osborn there is recited the wide range of 
opinion as to the structure, habits, and ordinal relationships of these remarkable mammals between the date of 
the first description in 1715 and the present time; the author’s personal opinions on these points are summarized 
at the end (pp. 112-115). 

Following the ruling of the International Commission on Zoological Nomenclature, the generic name must 
be spelled Deinotherium throughout, in accordance with Kaup’s original orthography of 1829 derived from the 
Greek dewvds and Onpiov. The same principle applies to the family name Curtognati, proposed by Kaup in 1833, 
which may be given the modern family terminal and orthography CURTOGNATHID®, although as in all 
matters of proboscidean nomenclature the early authors naturally did not follow modern nomenclatural canons. 
The history of the term ‘Curtognati,’ equivalent in German to ‘Krummkiefer,’ in reference to the sharp downward 
curvature of the lower jaw, is given as follows in the Bulletin of the Geological Society of France for 1834 (Boué, 
Bull. Soe. géol. de France, 1834, V, p. 444): 



Curtognati—M. Kaup a regu de nouveaux ossemens d’Eppelsheim, et a reconnu que le Dinotherium medium était véri- 
tablement bien différent du D. giganteum. M. Kaup a pu faire aussi quelques corrections et additions 4 sa description du 
Dinotherium. Il pense que ses deux énormes défenses ne lui servaient pas seulement pour extraire des racines de la terre, 
mais encore, comme A la morse, pour l’aider 4 mouvoir son corps si lourd. D’aprés la forme des os intermaxillaires, cet 
animal devait avoir une trompe pour porter la nourriture 4 sa bouche. Il est 4 placer entre les Mastodontes et les Bradypus, 
et il formerait une famille particuliére 4 laquelle M. Kaup donne le nom de Curtognati. Elle serait caractérisée par la 
mAchoire inférieure courbée vers le bas, et les deux défenses dirigées vers le bas et en arriére (NV. Jahrb. f. Mineral., Geognosie, 
etc., 1833, cah. 5, p. 509 [516] avec 1 pl.). 

The following is from Kaup’s original description, 1833.4, p. 516: 

Nach meiner Ansicht steht es am besten zwischen Mastodon und Bradypus als eigene Familie, die ich Curtognati 
(Krummkiefer) nenne, und durch den nach Unten gekriimmten Unterkiefer mit den zwei Stoss-Zahnen ebenfalls nach Unten und 
Hinten gerichtet, charakterisire. Die Gattungs-Diagnose wiirde seyn $ Stoss-Zihne. Der ausfiihrlichere Charakter: Keine 
Stoss-Zihne im Oberkiefer, zwei nach Unten gekritmmte Stoss-Zihne im Unterkiefer,  Backen-Zihne, wovon der dritte 
dreihiigelig, die iibrigen zweihiigelig, und nur der erste des Unterkiefers mit einem schneidenden Rand auf der vorderen 
Halfte versehen ist. In der Jugend ist vor dem ersten Backen-Zahn noch ein tiberzihliger, und der Milchzahn des zweiten des 
Oberkiefers ist in die Liinge gestreckt und dreihiigelig. Riissel, Krallen zum Aufscharren, Gang auf den Hand-Randern, wie beim 

In 1868 Falconer erroneously assigned the name A ntoletheriwm to specimens from Attock belonging to the genus 
Deinotherium; this error was corrected by Lydekker in 1885. 

ANTOLETHERIUM FALCONER, 1868.—(Falconer, 1868, I, p. 416): “‘Among the fossils discovered by Lieut. 
Garnett, near Attock, is a portion of the lower jaw of a tapiroid animal containing what appear to be the last 
premolar and the first and second true molars. Of this specimen I have received an excellent drawing executed by 
Col. Baker, who regarded the species as allied to Tapir. (Plate xxxrv. figs. 1 and 2.) The teeth certainly differ 
generically from those of Dinotheriwm in the massive connecting bridge between the two ridges, speedily running 
them into one confluent disc. The bend of the first ridge is very tapiroid. The specimen appears to me to be of an 
undescribed genus for which the provisional name Antoletherium (avrod\n the east, and @npiov) would be 

(Lydekker, 1885, p. 105): “A. 20. Fragment of the right ramus of the mandible, containing jaa 
and m2; from Kushdlgarh. Figured in ‘Faleconer’s Paleontological Memoirs,’ vol.i., pl. xxxiv., figs. 1 and 2 (as 
Antoletherium), and in the ‘Pal. Ind.’ op. cit. [Lydekker, 1880] pl. xxix., figs. 2,3. Presented by Lieuts. Garnett 
and Trotter.’’ No. A. 20 is listed by Lydekker (1885, p. 105) under specimens of Dinotheriwm pentapotamiz in 
the Indian Museum, Calcutta. See also Lydekker, 1880, p. 184, in which he states: ‘“‘As I have already observed 
in the above-quoted passage in the ‘Records’ [Geol. Surv. India, 1877, Vol. X, p. 33], the figure of this specimen 
given by Dr. Murchison in the ‘Palzeontological Memoirs’ [Footnote: ‘Vol. I, Plate xxxrv, figs. 1-2.’] under the 
name of Antoletherium is entirely incorrect. It appears probable that Dr. Falconer never saw the original specimen, 
but made his new genus Antoletherium solely on the evidence of the imperfect drawing from which the figure was 
taken; a copy (or the original) of this drawing was sent by Dr. Falconer to Professor Owen, whose note on the 
subject will be found on page 416 of the first volume of the ‘Palzeontological Memoirs.’”’ 


Original and subsequent nomenclatural references and modifications for this Memoir are as follows: 



Deinotherium Kaup, 1829, p. 401 = Deinotherium 
Dinotherium Kaup, 1832. All subsequent authors adopted the abbreviated 

spelling = Deinotherium 
Curtognati Kaup, 1833 = CURTOGNATHIDAL 
Dinotheridz Bonaparte, 1845, in his “Catalogo metodico dei Mammiferi 

Europei,” cited by Palmer, 1904, p. 738 = CURTOGNATHID 
Dinotheriidz Bonaparte, 1850, cited by Osborn, 1918.468, p. 134 =CURTOGNATHID 
Dinotherina Bonaparte, 1841 (1837, fide Graells, 1897, p. 573); Dino- 

theriina Bonaparte, 1850, cited as a subfamily = DEINOTHERIIN] 
Antoletherium Falconer, 1868 = Deinotherium 
Dinotheriini Winge, 1906, p. 172 = DEINOTHERIIN 
Dinotheriinz Osborn, 1910.346, p. 558 = DEINOTHERIIN & 
Dinotherioidea Osborn, 1921.515, p. 2, ‘typified by the Miocene and 

Pliocene Dinotheres of Eurasia”’ = DEINOTHERIOIDEA 

The following is Osborn’s definition of the suborder or superfamily DEINOTHERIOIDEA (Osborn, 
121.515, p. 3): 
II].—Dinotherioidea. All agree that these animals were chiefly fluviatile and amphibious in habit, in this 
respect resembling Mceritheres but differing in the entire loss of the superior incisor teeth. Early loss of upper tusks released 

the inferior. In the downturning of the inferior tusks the Dinotheres are paralleled by the Rhynchorostrine among the Masto- 

rl? rf! U! UP 

rle rh; U; Up 

rle Up 
A B 

Fig. 55. Fundamental arrangement of the cutting teeth in the four super- 
families of the Proboscidea. For lateral aspect of the lower cutting teeth, compare 
fig. 1, p. 16. See also fig. 4, p. 23. 

A.—Mearitherium, B.—Deinotherium. C.—Rhynchotherium. Composition of two species. 
D.—Stegodon. 8S. ganesa stage. Scale not uniform. 

dontoidea. In skull form and in limb and foot structure the Dinotheres parallel the true proboscideans. They diverge very 
widely from proboscideans in the evolution of the upper and lower grinding teeth. The primitive Dinotheres present 
simple bilophodont grinders, similar to those of Meritherium, and are persistently bilophodont. The upper grinders attain a 
stage which parallels the molar pattern of the tapir (Tapiroides[Tapirus]) among the perissodactyls, but show no ten- 
dency to the trilophodont, tetralophodont, or polylophodont structure characteristic of the mastodons and elephants. 

Thus the technical range of classification and nomenclature for these animals stands as on the following page. 



FamILy: CURTOGNATHID Kaup, 1833 
SuBFAMILY: DEINOTHERIIN® Bonaparte, 1850, Winge, 1906, Osborn, 1910 

Genus: DEINOTHERIUM Kaup, 1829 

So far as known the Deinotheres are certainly monophyletic. There is a single phylum of successive ascending 
mutations and species extending from the base of the Miocene, where they suddenly appear in Europe (D. cuvier7), 
to the Lower or Middle Pliocene of Rumania, where the pre-final stage is attained in D. gigantissimum. The final 

stage is D. hopwoodi of the Pleistocene of Africa. 


Bole = 2-2 160 


Fig. 56. Geographic distribution (according to the numbers in the list below) of the types of Deinotherium. The approximate localities where these 
twenty-six' types were found are represented by the white dots within the black areas; these dots each carry a number in a circle representing the chronologic 

seqnence of type description. The white crosses represent referred specimens. 

See Figure 56 

ORIGINAL NAME IN Present Memoir GroLocic AGE 

1715 Réaumur publishes Antoine de Jussieu’s drawing of a grinding 
tooth discovered near Lyons, France, in the previous 

1773 Rozier describes and figures a tooth discovered near Vienne, 
Dauphiny, France 

1798-1822 Cuvier ascribes grinding teeth found in France to ‘“‘Tapirs 

1. 1828-1829 Kaup describes a lower jaw from near Eppelsheim, 
Germany, now in the Museum of Darmstadt, assigning 

= Deinotherium sp.(?) 
= Deinotherium sp.(?) 

= Deinotherium sp.(?) 

in 1829 the name Deinotherium giganteum = Deinotherium gigantewm Lower Pliocene 
2. 1831 Gmiind, Bavaria Deinotherium Bavaricum 
von Meyer = Deinotherium bavaricum Upper Miocene 
3. 1832 Eppelsheim, Germany Dinotherium maximum Kaup 
(MS.), fide von Meyer, 
1832.2, p. 78 = Deinotherium giganteum Lower Pliocene 

ISee No. 6a, p. 85, and footnote 1. 

4. 1832 
6. 1833 
6. 1835 
7. 1835 
8. 1836 
9. 1839 
10. 1839 
11. 1839 
12. 1841 
13. 1841 
14. 1845 
15. 1845 
16. 1845 
17. 1861 
18. 1868 
19. 1876 
20. 1880 
21. 1892 
22. 1911 
23. 1912 
24. 1930 
25. 1935 

Comminge, Carlat-le-Comte, 
and Chevilly, France 


Dinotherium Cuvieri Kaup 

Kaup proposes the family name Curtroenarti for the Deinotheres 

Eppelsheim, Germany 
Podolia, Russia 

. 1835-1837 Podolia, Russia 

Ural Mountains 
Simorre, France 

Locality (?) 
Locality (?) 
Locality (?) 

Eppelsheim, Germany 
Mosskirch, Germany 

Darling Downs, Australia 

Compubay [Cambay, cf. p. 90], 


Perim Island, India 

Grive Saint-Alban, Isére, France 
Perim Island, India 

Attock, Indus Valley, India 
Sind, India, Salt Range 
Gaiceana, Tecuciti, Rumania 

Bugti beds, Bugti Hills, 

Karungu, British East Africa 

Bugti beds, Bugti Hills, 

Kotyhaza (Dep. Négrad, 

Olduvai, Tanganyika Territory, 
British East Africa 

Kavirondo Gulf, Africa 

Dinotherium medium Kaup 

Dinotherium proavum (Dinotherii 
proavi Eichwald) 

Mastodon podolicum' Eichwald 

Dinotherium uralense Eichwald 

Dinotherium secundarium Kaup 
(teste Lartet, 1836, p. 218) 

Dinotherium [gig.] var. majus de 
Blainville (fide Weinsheimer, 1883, 
p. 210) 

Dinotherium [gig.] var. medium de 
Blainville (fide Weinsheimer, 1883, 
p. 210) 

Dinotherium [gig.| var. minus de 
Blainville (fide Weinsheimer, 1883, 
p. 210) 

Dinotherium Kénigit Kaup 

Dinotherium minutum von Meyer 
(in Bronn) 

Dinotherium Australe Owen 

Dinotherium angustidens Koch 
Dinotherium intermedium de Blainville 
Dinotherium indicum Falconer 

Dinotherium levius Jourdan 

Dinotherium Perimense Falconer, cited 
in Murchison (name only) 

Dinotherium pentapotamix Lydekker 

Dinotheriwm sindiense Lydekker 

D. [Dinotherium|] gigantissimum 

Dinotherium ndricum Pilgrim (name 
abandoned by author and sub- 
specific term gajense substituted— 
see below) 

Dinotherium hobleyi Andrews 

Dinotherium indicum Lyd. var. 
gajense Pilgrim 

Prodinotherium hungaricum 

Deinotherium hopwoodi sp. nov. 

Deinotherium? sp.? Leakey 


= Deinotherium cuvieri Lower Miocene 

= Deinotherium medium Lower Pliocene 
= Deinotherium proavum 
= Deinotherium podolicum 
= Deinotherium uralense 
= Deinotherium secunda- 

rium (?) 

Upper Miocene(?) 
Lower Pliocene 

Middle Miocene 
= Deinotherium giganteum Lower Pliocene 
Lower Pliocene 

= Deinotherium giganteum 

Lower Pliocene 
Lower Pliocene 

= Deinotherium giganteum 

Upper Miocene 
=Diprotodon australis (not 
a proboscidean) 

= Deinotherium sp.(?) Middle Pliocene 

= Deinotherium intermedium Upper Miocene(?) 

= Deinotherium indicum (fide 

= Deinotherium levius 

Middle Pliocene 
Upper Miocene 

= Deinotherium indicum Middle Pliocene 

= Deinotherium indicum Upper Miocene 

= Deinotherium sindiense (?) Middle Miocene 

=Deinotherium gigantissi- 

Middle Pliocene 

= Deinotherium indicum ga- 

= Deinotherium hobleyi Lower Miocene 
= Deinotheriwm indicum 

gajense Lower Miocene 

= Deinotherium hungaricum Lower Miocene 

= Deinotherium hopwoodi 
sp. nov. Middle Pleistocene 
= Deinotherium hopwoodi (?) Pleistocene 


The history of discovery is most picturesque and even romantic because of the wonder and astonishment 
caused by the discovery of these pre-Diluvian mammals and the many and often grotesque theories as to their 
habits and external appearance. 

For the early history of discovery up to the year 1883 we may depend chiefly on the extremely thorough and 
scholarly article published by Dr. O. Weinsheimer in the Palzeontologische Abhandlungen, Band I, Heft 3, Berlin, 
1883, entitled, “Uber Dinotherium giganteum Kaup,” pp. 208-216. This history we have translated with 

1Type mandibular symphysis. 
Jahrb., 1837, p. 44) referred by Eichwald to Deinotherium. 

physis of the lower jaw of Deinotherium giganteum. 
*An undescribed specimen, of Pleistocene age, associated with the type remains of Homo kanamensis (see Leakey, 1934, pp. 204, 206, 207, also p. 105 of 
the present Memoir). 

First named Mastodon podolicum by Eichwald in 1835 (p. 736, Tab. v1, Lym, see also p. 734); subsequently (Neues 
Falconer (1846, p. 4) states in a footnote that the specimen appears to be a fragment of the sym- 


marginal inserts anda few comments of our own together with illustrations of some of the important specimens 
alluded to by Doctor Weinsheimer in his interesting narrative of the progress of discovery. For the bibliography of 
the emouhercs, the reader is referred not only to Weinsheimer and to the authors mentioned by him in the 
following historical summary but to the Bibliography at the close of the present Volume I. The more recent 
history of discovery is compiled by Osborn (p. 95). 

Translated by C. D. Matthew from the original of Weinsheimer, with notes, modifications, and additions by Osborn, 1925 

Réaumur, 1715.—In 1715 Réaumur (1715)! published an illustration from a drawing by Antoine de Jussieu of 
a tooth of Deinotherium found near Lyons in the beginning of the previous century, without, however, ascribing it to 
any particular animal. From that time no further remains of this remarkable animal of the Miocene Tertiary 
were found until Rozier (1773) described and figured another—a molar tooth discovered in Dauphiny in the 
neighborhood of Vienne by Gaillard—without knowing of the tooth previously figured by Réaumur. 

The third and most remarkable discovery of Deinotherium remains was of two lower jaw fragments, unearthed 
near Comminge, France, and described and figured by de Joubert (1785). In the same year Ildephons Kennedy 
(1785) published his ““Abhandlung von einigen in Baiern gefundenen Beinen,” in which he describes five teeth and 
an astragalus as the best preserved parts of a complete skeleton of Deznotherium found April 6, 1762, near Reichen- 
berg in Lower Bavaria. Kennedy says that these remains show striking resemblance to remains [of mastodonts] 
from the Ohio River in North America, from Peru in South America, from Asia, and from the small town of Fiirth 
in Lower Bavaria (1785, t.2 [IV], f.6), and remarks at the close of his article that these remains belonged to an 
animal that had long been extinct, and that could not be distinguished from the Siberian mammoth, the 
“Ochsenvater der Louisianer.”’ 

Cuvier, 1798—-1822.—These teeth, described by de Joubert and Kennedy, were more carefully studied by 
G. Cuvier (1798) who, on account of their similarity to the teeth of the tapir, ascribed them to some gigantic 
extinct individual of that genus, which was contemporary with Mastodon and Elephas. Some of the teeth which 
Kennedy described were better figured by S6mmering (1818) who ascribed them to Cuvier’s giant tapir. Other 
teeth and jaws of Deinotherium, or fragments of such, were found later at various sites in France, such as St. 
Lary, Arbeichan, Grenoble, Carlat-le-Comte, Chevilly, and others, and—together with the specimen already 
described by Cuvier (1822)—ascribed to a new genus; these Cuvier named ‘‘Tapirs gigantesques,” for he already 
suspected, in spite of their similarity of form, that on account of their great variation in size they did not belong 
to one single species, and therefore he used the plural. 

Kaup, 1828-1829.—Somewhat later Kaup (1829) made a report to the Naturforscherversammlung of Berlin, 
through Berthold, describing the left half of a large lower jaw, found in 1828 near Eppelsheim in Hesse-Darmstadt 
and sent to the Museum of Darmstadt where it remains to the present day. Much as the molar teeth in this jaw 
resembled those of the tapir, the teeth in the anterior part of the jaw were so widely different from the correspond- 
ing onesin the tapir that Kaup felt justified in distinguishing a new genus and species, Deinotherium giganteum, which 

he believed to be related to Hippopotamus. The tusks of the lower jaw he erroneously placed [Fig 52] pointing 

Since that time—particularly during the 30’s—a great many remains of Deinotherium have been found at 
various sites in Europe, that is, in France and Germany, as well as in Asia, so that the material and the literature 

"Dates in parentheses after authors’ names refer to the bibliographic list given on pp. 214-216 of Weinsheimer'’s article. 


respecting this genus are now very extensive. Moreover, the various authors who have occupied themselves with 
the specimens have reached widely divergent conclusions in regard to the dentition, the various species, and the 
classification of Deinotheriwm. 

THEORIES AS TO Hasits AND Hasirat.—Thus, according to Kaup (1832), Deinothertwm apparently had four 
to six tusks in the upper jaw and two in the lower jaw, which were used to stir up the earth when seeking the 
roots of which its food consisted. There were apparently six molars in the upper jaw, and to all appearances yet 
a seventh, anterior to these, which, however, as in Hippopotamus, was lost early. In the lower jaw, on the 
contrary, there were only five molars. He distinguishes two species—one large, Deinotheriwm giganteum Kaup, 
and one smaller, Dinothertwm Cuviert Kaup. 

According to H. von Meyer (1833 [1832.1]) De:notherium is most nearly related to Rhinoceros incisivus; he 
recognizes the same species as Kaup, but—on account of its frequent occurrence in Bavaria—he calls the Bavarian 
animal Deinotherium bavaricum, which is one quarter smaller than Deinotheriwm giganteum. He dissents from 
Cuvier’s view that Deinotheriwm was contemporary with Mastodon and Elephas because their remains have never 
been found associated, claiming that it was contemporary with ‘Mastodon’ angustidens ‘and Rhinoceros incisivus, 
and became extinct earlier than the mastodon of Cuvier, and that it belongs to the latest Tertiary deposits. In 
the plates given by Kaup (1832) and H. von Meyer (1833 [1832]) the tusks of the lower jaw are pictured point- 
ing upward. 

Kaup, 1833.—In 1833 the Museum of Darmstadt (Kaup, 1833, p. 419) acquired an almost perfect left half 
of a lower jaw from Eppelsheim, from which it could be seen that the anterior part of the jaw was inclined down- 
ward, whereupon Kaup (1833, [1833.4, Taf. vi], p. 509) corrected his earlier mistake (1829—see our figure 52). He 
now states that Deinotherium lived only on land, on which—like the sloth Bradypus—it dragged itself along. Its 
principal food consisted of roots and tubers which it dug out of the earth with the two tusks of its lower jaw, and 
stuffed into its mouth with its trunk (Kaup, 1833 [1833.2], p. 172); but the tusks also served, like those of the 
walrus, to help its progress on land. It forms a separate family, intermediate between Mastodon and Bradypus, 
which he named Curtognati (i.e., crooked jaw: Kaup, 1833 [1833.4], p. 509 [516]). 

Pauuas (1777)—ErcHwatp (1835).—Eichwald (1835) describes two molars from Podolia found associated 
with teeth of Mastodon podolicum,' and assigns them to a new species which he names Dinotherium proavum. In 
it he also includes a molar from the Ural described by Pallas (1777), subsequently (1835) named Dinotherium 

Larter (1835 [1836]).—Lartet (1835) ascribed a few teeth found in the sands of the freshwater deposit 
of Sansan, Gers, to the species D. secundariwm Kaup. 

Buckianp, 1835.—From the existence and downturned position of the tusks Buckland (1835) draws the 
following conclusions in regard to the life and habits of Deinotherium. For a land animal the monstrous lower jaw 
and the consequent length of body would be very cumbrous and wearisome. With an aquatic animal these would 
not be disadvantages. He considers, therefore, disagreeing with Kaup (1833 [1833.2], p. 172), that Deinotherium 
frequented freshwater lakes and rivers, and was most nearly related to the tapirs. The tusks were used for up- 
rooting and plucking roots and large aquatic plants. From their construction it is seen that for this purpose they 
united the mechanical strength of the pickaxe with that of a very heavy harrow. The weight of the head, pressing 
on the downwardly pointing teeth, would increase the effectiveness of such a use, just as a heavy weight is fastened 
to a harrow to increase its effect. Possibly Deinotherium also used its tusks to lay hold of the bank and so to 

1See footnote 1 on page 85. 


anchor itself, Thus the nostrils would come above the level of the water, so that the creature might sleep without 
danger and could breathe, while its body, floating in the water, remained concealed with the muscles relaxed and 
resting; for the weight of head and body would also serve to keep the tusks fast hooked in the same place, like 
the weight of a sleeping bird whose claws hold it fast to the branches on which it sits. It is also most probable that 
Deinotherium, like the walrus, used its tusks to drag itself on the land, or in case of danger or of attack as a means 
of defense. From the construction of the shoulder-blade it is clear that the fore feet were used in conjunction with 
the tusks for uprooting and dragging out roots and large aquatic plants. Hind limbs it did not have. 

Fig. 57. Four views of the reconstructed original Eppelsheim skull of Deinotherium giganteum in the British Museum (Natural History). After 
Andrews, 1921, pp. 525-534, figs. 1-4. 

This is the first skull found at Eppelsheim and referred to by Weinsheimer (1883, p. 210) as having been exhibited in Paris, March 15 and 16, 1837, 

to the members of the Academy of Sciences and other savants for their consideration, resulting in widely divergent views, and afterward sold to the 
British Museum in London. 

According to Andrews (1921, pp. 525, 526), this skull was discovered in 1835 by von Klipstein in the well-known ossiferous sands of Eppelsheim (Hesse- 
Darmstadt); the specimen was described by von Klipstein and Kaup in 1836 under the title ‘‘ Beschreibung und Abbildungen von dem in Rheinhessen auf- 
gefundenen colossalen Schadel des Dinotherii gigantei”’; while in Paris the skull was examined by de Blainville, who has given one of the best descriptions 
as yet published; it was then intended to send the specimen to London, but whether this was done at that time is uncertain; some years later, in 1849, 
it was certainly in London, and was offered to the British Museum for purchase; it was then examined and reported on by Buckland and Owen; the pur- 
chase, however, was not completed, and the skull seems to have been sent back to Darmstadt. About 1866, von Klipstein sold his collection to Dr. Old- 

ham, Director of the Geological Survey of India, and from him the British Museum acquired the specimen, which is in an excellent state of preserva- 
tion, despite the widespread idea that it was injured on its journey to London. 

Kaup AND von Kuipstern, 1836.—Shortly after, one of the most interesting paleontological discoveries was 
made in the Lower Pliocene of Eppelsheim, Germany, namely, a complete skull of Deinotheriwum which was 
described by Kaup (von Klipstein and Kaup, 1836). Kaup holds to his original view, and thinks that the walrus-like 
structure of the posterior half of the skull—on account of which Buckland considered Deinotherium an aquatic 
animal—offers no difficulties, and that such a flattened occipital region would permit the insertion of more 
powerful muscles than an abruptly ending occiput. Beside, these muscles would have to be excessively powerful 
if Deinotherium used its tusks for tearing and rooting up the earth. This Eppelsheim skull—later sold to the British 
Museum in London, uninjured in transport—was exhibited by von Klipstein and Kaup in Paris, March 15 and 16, 

1837, to the members of the Academy of Sciences and other savants for their consideration, and again there 
resulted widely divergent views. [See Fig. 57 of the present Memoir.] 


Der BuaAINviLLE, 1837.—Thus de Blainville (1837.1) gives a minute description of the Eppelsheim skull, and 
classes Deinotherium as a genus belonging to the family of aquatic pachyderms |i. e., Sirenia] together with Halicore 
and Manatus, while Elephas, Mastodon, and Tetracaulodon belong to the terrestrial pachyderms [i. e., Proboscidea]. 
In regard to the habits of Deinotherium he practically agrees with Buckland. Later (1839) de Blainville recognizes 
two species, Dinotheriwm Cuviert Kaup (1832) and Deinotherium giganteum Kaup (1829), and of the latter he 
distinguishes three separate varieties—Dinotherium majus de Blain., Dinotherium medium de Blain., and Dino- 
thertum minus de Blain.—based on the size of the teeth. 

Dumeéril (1837, p. 427) and Isidore Geoffroy Saint-Hilaire (1837, p. 429) agree with de Blainville, while Strauss 
(1837, p. 529) considers Deinotherium exclusively aquatic and not able, like Hippopotamus and Manatus, to come 
on land. From the large narial openings he concludes that it had a trunk—the existence of which de Blainville 

1/16 nat. size 

D. giganteum 

D. giganteum 
Falc. 1847, Pl. XLIV, Fig. 1 (rev.) 

Falc. 1847, Pl. XLil, Fig. 1 

Fig. 58 

Fig. 58. First Eppelsheim skull and jaws of Deinotherium giganteum, after the cast and restoration of Kaup, about one- 
twelfth natural size. Copied from Gaudry, 1878, p. 188, fig. 248. Type jaw in Darmstadt Museum; skull in British 
Museum (see Fig. 57). 

Fig. 59. Restored Eppelsheim skull and jaws of Deinotherium giganteum Kaup. After Falconer, 1846 [1847], from 
original in Museum of Darmstadt. 

Al, Op. cit., Pl. xin, fig. 1. A, Op. cit., Pl. xxiv, fig. 1 (rev.). 

doubted—used like that of the elephant for seizing food. The tusks were not used for uprooting plants, because 
they showed no signs of wearing down, but were used like those of the elephant for self-defense only. It formed a 
separate family, transitional between pachyderms and cetaceans. Jacquemin (1837) placed it between Rhinoceros 
and Elasmotherium, and Owen (1840) between the tapir and the pachyderms with trunks. Owen [1840-1868] 
gives the dental formula of Deinotherium as follows: 

1. Deciduous teeth; Dir=t; Des=s; Dp3=s=16. 

2. Permanent teeth; I+; Ci=s; Pi=3; M%=$=22. 
He claims that of the three milk molars in the young animal, the anterior has no corresponding permanent tooth, 
that the tusks of the male are twice as large as those of the female, and that in both cases they were used for 
uprooting aquatic plants, since, like Hippopotamus, it was an aquatic pachyderm. 

Kaup, 1841.—After the discovery of the first great [Eppelsheim] skull [now in the British Museum (Natural 
History)], the Museum of Darmstadt received a great number of additional Deinotherium remains from Eppelsheim 


and Westhofen, which Kaup studied and described in the ‘‘Akten der Urwelt”’ in 1841. By that time he already 
had extensive material at his disposal, including the complete [Eppelsheim] skull, three skull fragments, 13 lower 
jaws, and 115 molar teeth. He now changes his former views, and considers that the adult Deinothertum had only 
five molars in both upper and lower jaw, while in the young six were present, anterior to the first being a “super- 
fluous”’ one which, however, soon fell out. 

All the teeth of Deinotherium previously known belonged to only one species [i. e., D. giganteum Kaup] with a 
wide range of variation in size. Dinotherium proavum Eichwald [1835], Dinotherium medium Kaup [1833.3] (see 
below), Dinotherium Cuviert Kaup [1832] sive bavaricum H. von Meyer [1831] are therefore not admitted by Kaup as 
species. These teeth do not justify the recognition even of large or small varieties. Nevertheless Kaup introduces 
(1841, p. 49) in addition to his Deinotherium giganteum a new species which he calls Dinothertum Konigit Kaup. 

‘Dinotherium giganteum varies greatly in size, so that individuals of this species may have attained a length 
of 18 Parisian feet, while others again may have been barely 11 feet long.’ On the other hand, the smaller species, 
Dinotherium Konigii, is supposed not to have exceeded the Indian Rhinoceros in length. In regard to the life and 
habits of Deinotherium Kaup reiterates his former opinions. 


Fig. 60. De Blainville’s original type figure of Dinotherium intermedium, 1839-1864 [1845], Pl. ur. 
Possibly from the Upper Miocene of Simorre (cf. p. 111). Right mandibular ramus. One-third natural size. 

Owen’s Error, 1843.—Up to 1848 remains of Deinotherium had been found only in Europe, but now it was 
claimed that remains of the creature had also been found in Australia. Certain of these were sent to London 
and studied by Owen (1843) who ascribed them to a separate species which he named Dinotheriwm australe 
Owen. Later, however, these remains were recognized as belonging to [the giant marsupial] Diprotodon. 

Kocu, 1843.—In 1843 Koch (1845, p. 41) saw in the British Museum in London a part of a lower jaw, 
discovered in Compubay,' western India, and labeled “Mastodon angustidens.” By a detailed examination he 
found that it did not belong to the genus Mastodon but to Deinotherium, and, if not to a very old specimen, at 
least to one fully adult, which—on account of the comparatively small molars—he named Dinotherium angusti- 
dens Koch. He completely agrees with Buckland (1835) in regard to its life and habits, and classes it with the 
Proboscidea as intermediate between T'etracaulodon Godman and Missourium Koch. Geinitz (1846) agrees with 

Fauconer, 1845.—Shortly after this remains of Deinotherium were also found in western India. Falconer 
(1846|1845]) describes the posterior half of a lower left molar of Deinotherium from a chalky ferruginous conglomerate 

(‘Im Jahre 1843 sah ich im britischen Museum zu London eine in Compubay in Ostindien aufgefundene, mit der Ueberschrift: ‘Mastodon angustidens’ 
versehene untere Kinnlade.” (Koch, 1845, p. 41.) For “Compubay,” a locality not on any existing map, probably read Cambay, western India.| 


from Perim, a small island at the mouth of the Narbada [Nerbudda River] in the Gulf of Cambay on the west coast 
of India, which differs from the corresponding parts of an Eppelsheim tooth in having the transverse crest and 
the enamel thicker. He therefore ascribes it to a new species which he names Dinotherium indicum Falconer. 
He considers Deinotherium very nearly related to Mastodon Ohioticus (Falconer, 1868, 1, p. 85) and belonging 
likewise to the Proboscidea. 

Gervais, 1848.—Gervais (1848) classes Deinotherium with Mastodon in the order Proboscidea, and recog- 
nizes three species—Deinotherium giganteum, Dinothertum intermedium de Blainville (see fig. 60), and Dinotherium 
Cuvieri—while Wagner (1850), Girard (1851 [1852]), and Pictet (1853) continue to place it in the order Sirenia. 
Burmeister (1851) holds it for an herbivorous sea monster, 15-20 feet long, with short, thick neck, powerful, 
slender rump, and broad, effective flippers for creeping like the walrus, which like all the Sirenia frequented the 
mouths of large rivers, and used its tusks for assisting its movements on shore and for uprooting aquatic plants. 

DISCOVERIES IN WESTERN Europe, 1850-1858.—Again in the 50’s a great number of remains of Deinothertum 
were found in a variety of places, and announced chiefly in brief reports. Opinions in regard to species, dentition, 
and classification continued divergent. Jiiger (1850) reports the occurrence of Dinotherium Cuvieri Kaup sive 
bavaricum H. von Meyer at Mésskirch [Upper Miocene], and of Deinotherium giganteum Kaup near Salmendingen 
in the pisolitic iron ores of the Swabian Alb. Quenstedt (1851 [1850]) reports the occurrence of the latter in the 
pisolitic iron ores of Melchingen south from Tiibingen, and later (1853) in those of Frohnstetten, and says that 
Deinotherium had not five, but six, molars in both jaws. Von Hingenau (1852) found molars near Nikolsburg and 
Keltschau; Glocker (1852) a tusk and several molars in the blue marl near Abtsdorf; Fétterle (1853) the right half 
of a lower jaw with tusk near Fiinfkirchen and a molar near Keltschau; and Merian (1854) a molar in the Delsberg 
valley in the Bernese Jura. Pomel (1854 [1853]) classes Deinothertum with Elephas and Mastodon among the 
Proboscidea; Bronn (Lethaea, III, p. 803) as intermediate between Sirenia and Pachydermata; and Agassiz 
(Bronn, Lethaea, III, p. 805) among the herbivorous Cetacea. 

In 1853, in consequence of the collapse of an embankment on the Prague-Briinn railway, a skeleton of Deino- 
thertum giganteum was exposed in the blue marl near Abtsdorf, the bones of which lay together, covering a space 
about 24 feet long and 18 feet wide. Many of them fell to dust after being for a time exposed to the air; others, 
before word of the discovery was noised abroad, were shattered or cleared away. All that was preserved was the 
complete jaws and dentition, the first and second cervical vertebra, a few dorsal and caudal vertebre, a large part 
of the tarsus and metacarpals, and a few large fragments of the limb bones. These skeletal parts of a juvenile 
specimen, which are now in the Bohemian Museum of Prague, prove incontestably that Deinotheriwm should be 
classed not with the Cetacea, but with the Pachydermata as nearly allied to Mastodon (Reuss, 1855). Neverthe- 
less Giebel (1855) again classes it among the Sirenia as nearly allied to Halitherium. 

Deinotherium remains were found in France at this time by Lockhart (1853) in the Department of Loiret: 
by Bayle (1855) near La Chaux-de-Fonds in the Miocene Molasse which is identical with the deposit of Simorre 
and Sansan [Miocene] and the Faluns of Touraine; and by Rouault (1858) near Rennes, his discovery consisting 
of a large molar of Dinotherium Cuviert. Hauser (1856) discovered remains of a Deinotherium skull in the Leitha 
chalk near Breitenbrunn on the Neusiedler See, Hungary. Gaudry and Lartet (1858) found a tibia 95 em. long, 
and Wagner (1857) askull fragment with five permanent teeth anda forearm of a young individual at Pikermi [Lower 
Pliocene]. Wagner believes that all Deinotherium remains known up to this time belong to one and the same 
species. Aichhorn (1857) found teeth of Deinotherium gigantewm near Wies in Steiermark [Miocene]. Fdétterle 
(1857) found a fragment of the right side of the chin with a molar and several bones of the torso embedded in fine 


sand intermixed with rubble (Schotter) lying above the upper brackish blue marl. At other sites in Austria Poppe- 
lack (1858), Hoérnes (1858), and von Hingenau (1858) found various more or less important remains of 


Larter, 1858 [1859].—Lartet (1858) holds the opinion that the first dentition in the upper jaw of the young 
Deinotherium consists of three milk teeth on either side, followed by the first molar of the second dentition, and 
then by the second molar. The first milk tooth falls out but is not replaced; the second and the last are replaced 
by premolars which are simpler than the corresponding milk teeth. Accordingly he gives the following dental 


1. First dentition: Dit=t; Dp 2-43=3=14. 
2. Second dentition: It=+; P 3-423; Ms=3s=22. 

Lartet recognizes four species: 

1. Dinotherium giganteum Kaup, syn. Dinotherium proavum? Kichwald. 

2. Dinotherium sp.? Lartet. 

3. Dinotherium bavaricum? H. v. Meyer, syn. Dinotherium intermedium? de Blainville. 
4. Dinothertum Cuviert Kaup. 

Gaupry, 1860-1878.—Gaudry (1860) in his excavations at Pikermi, Greece [Lower Pliocene], collected a 
great number of skeletal fragments of Deinotherium giganteum which he considers as an ambulatory terrestrial 
animal related to Mastodon and Elephas, which was also his opinion in 1861 and 1862 when (p. 162) he gives a 
detailed description of the Greek remains and feels obliged to consider the various species of Deinotheriwm merely 
as varieties of one species (Il. ¢. p.171). Miller (1861) feels obliged to doubt whether Deinotherium was exclusively 
an aquatic animal, and Hensel (1862 [1863]), on account of its skeletal structure, classes it among the pachyderms 
with trunks. Suess (1863) reports a small Deinotherium (Deinotherium bavaricum?) from the [Upper Miocene] coal 
seams of Parschlug, Eibiswald, Turnau, Wies, etc., in Steiermark, and a large one, similar to the unnamed species 
of Lartet, in purely marine deposits. Rachoy (1863) and Stur (1864) found remains of Deinotherium bavaricum in 
the overlying sandstone of the coal bearing Tertiary basin of Leoben in Steiermark. 

CRANIAL CHARACTERS, CLauptus, 1865.—Claudius (1864 [1865]), from the structure of the auditory laby- 
rinth of Deinotherium, shows positively that this genus, like the elephant, belongs to the family of the Proboscidea 
and among the pachyderms. According to his researches both Elephas and Deinotherium have a depressed auditory 
cochlea with less than three convolutions, but notably broad at the base. The planes of the superior and posterior 
convolutions of the semicircular canals which intersect in the crus commune, form an angle less than 90°, while in 
Rhinoceros and Tapirus the cochlea is higher, the angle formed by the superior and posterior convolutions is at 
least 90°, and the cochlea has hardly more than two convolutions. From the size of the canalis facialis the exist- 
ence of a trunk may be inferred with certainty. In Deinotherium the facialis nerve is led from the inner ear through 
an elliptical foramen 5’”’ long and 2’” wide. The latter figure, which remains the same for the passage through the 
tympanic cavity, gives the diameter of the nerve which, from the inner ear outward, penetrates the thin bone in a 
diagonal direction and thus produces an elliptical opening. The nervusfacialis of Deinotherium was about the size 
of the medianus nerve in man, and served like it for the innervation of an enormous number of muscle fibers. In 
the elephant this canal has the same dimensions: in the tapir it is proportionally much narrower. The stapes is 
similar in structure to that of the elephant; the incus somewhat smaller (6’”"; 4’); in its morphological rela- 
tions the labyrinth corresponds perfectly to that of the elephant; similarly the canalis ganglionaris of the cochlea 



agrees perfectly with that of the elephant, and consequently shows marked differences from all other mammals in 
this respect. 

GerRvAIS (1864), FaLconEeR (1868).—In Hérault, between Narbonne and St. Chinian, in the freshwater 
Molasse of Montredon, Gervais (1864) discovered teeth of Deinotherium giganteum in the Upper Miocene [Lower 
Pliocene], as well as fragments of tusks, several bones of manus and pes, a metacarpal, and a phalanx, also an in- 
ner ear. In Haute-Garonne, Salaro (1864)["| discovered a pelvis of Deinotheriwm in which he believed he recognized 
marsupial epipubic bones, and therefore concluded that it was a monstrous pachyderm-like marsupial related to 
Megatherium, a view also supported by H. von Meyer (1865, p. 1 et seg.). Quenstedt (1867) has doubts whether 
Deinotherium should be classed with the pachyderms or the cetaceans, while Carus (1868) would place it between 
the Sireniaand the Proboscidea. According to Falconer (1868) Deinotherium had only two milk molars in both 
upper and lower jaws—the posterior with three, and the middle one with two transverse crests. The anterior or 
third remained undeveloped. The two milk teeth were replaced vertically by an equal number of premolars, 
both of which have two crests. Following these come the three true molars, of which the first has three transverse 
crests, and the otherseach two. He divides the Proboscidea into Mastodon and Elephas, and holds Deinotherium for 
a divergent form of the latter. 

OwEN (1840) ro LypEKKER (1876).—Among the more recent works on Deinotherium is that of Brandt (1869). 
He agrees with Owen (1840) and Lartet (1858) that of the three milk molars [Dp 2-4] the anterior one [Dp 2] 
has no corresponding replacing tooth [P 2], while both the others when they fall out are replaced vertically by two 
premolars [P 3-4]. Thereconstruction of Deinothercwm which he figures is very similar in form to the elephant, 
and he also says that Deinotheriwm was a true member (genus) of the family of elephant-like animals, more nearly 
allied to Mastodon than to the elephants, more nearly allied than either Mastodon or Elephas to many other 
pachyderms, and related—though slightly—to the Sirenia. Its life and habits resembled those of the elephant. 
-Inaseparate chapter he lists the various species of Deinotherium, with the remark that he will not increase the 
confusion and divergence of opinions by offering new and unproven views. 

Peters (1871) describes a number of Deinotheriwm remains from the highest Miocene deposits of southern 
Steiermark, which include Deinotherium gigantewm Kaup, Dinotherium medium Kaup, Dinotherium Cuviert Kaup, 
Deinotherium bavaricum H. von Meyer, the latter being only a variety of Deinotherium giganteum Kaup. He considers 
Deinotheriumas a creature of fluviatile habitat, spending more than half its time immersed in river water. Bach- 
mann (1875) describes and pictures a lower jaw from the Delsberg valley in the Bernese Jura, which he identifies 
with Deinotherium bavaricum H. v. M., and considers as belonging to the pachyderms and not nearly allied to 
Manatus. Zittel (1875) ranks it with the Proboscidea, between Mastodon and Elephas. Holliinder (1877) also 
classes it with the Proboscidea, but says that the structure of the skull is not unlike that of the Sirenia, and that 
it was of aquatic habit. In regard to the dental succession in Deinotherium, Vacek (1877) fully agrees with Lartet 
(1858). He considers Deinotheriwm as the direct ancestor of Mastodon in the family Proboscidea, and so also 
does K6llner (1882). 

Lydekker (1876) describes a molar from India which he identifies with ‘Dinotherium pentapotamicum’ 
Falconer and later (1880) recognizes a third Indian species, beside Dinotherium pentapotamicum and Dinotherium 
indicum Falconer, which he founds on a fragment of lower jaw from Sind and names Dinotherium sindiense Lyd. 
He classes Deinotherium with the Proboscidea. 

‘See Sanna-Solaro in Bibliography at close of present Volume I. 


The above historical summary is on the authority of Dr. O. Weinsheimer (1883), and the reader is referred to 
his monograph “Ueber Dinotherium giganteum Kaup,” in which he also gives an invaluable Literatur-Verzeich- 
niss (pp. 214-216) of one hundred and fifteen titles, from Réaumur (1715) to Vacek (1882). Weinsheimer then 
reviews (1883, pp. 217-238) the whole subject of the dentition as observed chiefly in the collections of the mu- 
seums of Germany. He describes the newly found jaw in the Darmstadt Museum (pp. 238-244) also the cranial 
and skeletal material in the same Museum (pp. 244-247), and then considers the nomenclature (pp. 247-264) 
of the following sixteen species described or named up to the year 1880: 

Synonymy Monospecific 
fide Weinsheimer fide Weinsheimer 
Deinotherium giganteum Kaup type, Eppelsheim, 1829 = Deinotherium giganteum Deinotherium giganteum 
Dinotherium cwieri Kaup type, Eppelsheim, 1832 = Deinotherium cuvieri Deinotherium giganteum 
Dinotherium medium Kaup type, Eppelsheim, 1833 = Deinotherium giganteum Deinotherium giganteum 
Dinotherium maximum Kaup type, Eppelsheim, 1831, 1832 = Deinotherium giganteum Deinotherium giganteum 
Deinotherium bavaricum von Meyer type, Upper Miocene, 1831 = Deinotherium cuvieri Deinotherium giganteum 
Weinsheimer gives a very full discussion with measure- 
ments relating to this synonymy 
Dinotherium proavum Hichwald type, 1835, Podolia, Russia = Deinotherium proavum Deinotherium giganteum 
Dinotherium secundarium Kaup (ref. Lartet), 1835 =Deinotherium giganteum (fide 
Lartet-Weinsheimer) Deinotherium giganteum 
Dinotherium kénigui Kaup type, 1841 =Deinotherium cuvieri Kaup sive 
D. bavaricum von Meyer Deinotherium giganteum 
Dinotherium minutum von Meyer type (in Bronn, 1841), 
Upper Miocene, Mésskirch = Deinotherium bavaricum (?) Deinotherium giganteum 
Dinotherium australe Owen type, 1843 = Diprotodon australe 
Dinotherium angustidens Koch type, Compubay, western India 
[Cambay (?), western India], 1845 = Deinotherium angustidens Deinotherium giganteum 
Dinotherium intermedium de Blainville type, 1845 =Deinotherium bavaricum (fide 
Lartet, 1858) Deinotherium giganteum 
Dinotherium indicum Falconer type, 1845, compare Dinotherium 
perimense, Falconer, 1868, p. 415, Pliocene, Perim Island = Deinotherium indicum Deinotherium giganteum 
Dinotherium levius Jourdan type, 1861 =Deinotherium giganteum (fide 
Gaudry-Lartet) Deinotherium giganteum 
Dinotherium pentapotamiz Falconer type (MS.), 1868 = Deinotherium pentapotamiz Deinotherium giganteum 

Dinotherium sindiense Lydekker type, Middle Miocene, Sind, 1880 =Deinotherium sindiense Deinotherium giganteum 

Monospeciric Error.—The above synonymy is partly based on the erroneous treatment by Weinsheimer 
of Deinotherium giganteum |Lower Pliocene] as a collective species which embraces many of the Miocene stages 
also. He closes, accordingly, with the following ‘“‘Schlussbemerkungen”’ (pp. 280, 281), that in view of the 
great variation in size of the teeth of Deinotherium, and despite the various geologic horizons or layers in which 
the remains of Deinotherium occur, we have to do with one single European species, namely, Deinotherium 

Hiernach sind also alle Species, welche nach Zihnen oder Kiefern aufgestellt sind, die in Grésse und Gestalt von den dem 
Dinotherium gigantewm Kaup zugerechneten nicht mehr abweichen, als von den betreffenden Autoren angegeben wurde, zu einer 
einzigen Species zusammenzufassen, fiir welche die von Kaup zuerst gebrauchte Bezeichnung Dinotherium giganteum beizube- 
halten ist. Der Grund der mehr oder weniger grossen Abinderungen in Grésse und Gestalt der Zihne und Kiefer ist, wie wir 
gesehen, theils auf individuelle, theils auf sexuelle und besonders auch auf Altersverschiedenheiten zuriickzufiihren. . . . Um 
dies zu erkliiren, miissen wir annehmen, dass das Dinotherium verschiedener Zeitriume und Landschaften die bedeutenden 
physischen Veriinderungen, welche allenfalls zwischen die Ablagerungszeiten der verschiedenen Schichten der jiingeren Tertiiir- 
periode fallen, iiberdauert hat, ohne in merklicher Weise von denselben beeinflusst worden zu sein und ohne wiihrend der sehr 
langen Zeitdauer, welche zwischen diesen Veriinderungen liegt, die geringste Sechwankung seiner Artenmerkmale zu erfahren. 


Weinsheimer’s Memoir concludes with an invaluable review (pp. 269-280) of the geographie distribution of 
Deinotherium in the provinces of Germany, Switzerland, France, Austria-Hungary, Russia, Greece, and India. 

The historical summary below by the present author relates to some of the chief and more important papers 
which have appeared since Dr. O. Weinsheimer’s review (1883), namely, between 1883 and 1934, but also includes 
citations from some of the earlier authors, as follows: 

Richard Owen, 1840-1868 Charles W. Andrews, 1911 C. Forster Cooper, 1922 

Gregoriti Stefainescu, 1892-1899 R. Fourtau, 1918 Giinther Schlesinger (letter), 1923 
E. Kittl, 1908 A. Brives, 1919 Othenio Abel, 1924 

Lucien Mayet, 1908 Charles W. Andrews, 1921 R. W. Palmer, 1924 

RicHaRD OwEn (1840—1868).—First we may revert to Richard Owen (1840-1868), who records (1840, p. 609, 
1868, III, pp. 358, 359) the permanent dentition in Deinotherium as, I73; C33; Pi¢; M?3=22, and adds three 
superior and inferior deciduous premolars and one deciduous incisor, namely, Dp 2-43, Dir. 

As regards the tusks, Owen observes (1868, III, p. 359): “The generic peculiarity of the Dinotherium is most 
strongly manifested in its tusks. These, fig. 288, 7, are two in number, implanted in the prolonged and deflected 
symphysis of the lower jaw, in close contiguity with each other, and having their exserted crown directed down- 
ward and bent backward, gradually decreasing to the pointed extremity. In jaws with molar teeth of equal size, 
the symphysis and its tusks offer two sizes; the larger ones, which have been found four feet in length, with tusks 
of two feet, may be attributed to the male Dinothere; the smaller specimens, with tusks of half size, to the female. 
The ivory of these tusks presents the fine concentric structure of those of the Hippopotamus, not the decussating 
curvilinear character which characterises the ivory of the Elephant and Mastodon. No corresponding tusks, nor 
the germs of such, have yet been discovered in the upper jaw of the Dinotheriwm.”’ 

STEFANESCU, 1892.—Professor Gregoriti Stefiinescu of the University of Bucharest discovered remains of 
Deinotherium far exceeding in size any previously described, to which he gave the name Dinotherium gigantissimum, 
in allusion to Kaup’s term Deinotherium giganteum of 1829. 

There was first discovered in 1878, at Gaiceana, district of Tecuciti, a molar tooth, probably an Mz, of 120 
mm. in length and 120 mm. in width, thus exceeding in size the corresponding tooth of Deinotheriwm giganteum. 
This molar becomes the type grinding tooth of Stefainescu’s new species Dinotherium gigantissimum, 1892. It was 
embedded in yellowish-gray sand (op. cit., 1892, fig. 24) belonging to the Middle Miocene [Lower or Middle Pliocene]. 
An abbreviated summary of this very important discovery taken from his paper published in 1892 is as follows: 

(Stefanescu, 1892, pp. 81, 82): “I received in 1878 a fossil molar tooth found at Gidiceana, in the judet 
(district) of Tecuciti. It was the last but one molar of a Dinotherium, but it was so large that it could not have 
belonged to the usual D. giganteuwm, as you may judge from the following dimensions: 

Antero-posterior diameter meters 0.12 
Transverse diameter " 0.12 
Height of the crown bi: 0.08 
Height of the root ss 0.14 
Distance between the hills of the crown . 0.05 

Thickness of the hills at their basis sf 0.05 


These uncommon dimensions should lead us to look at these remains as belonging to another species than the 

usual D. gigantewm, which may be named D. gigantissimum.” 

In the same paper Stefiinescu describes a second discovery at Manzati of which the following is an 
abstract (p. 82): 

MAnzati Vattey.—In 1890, in the valley of MAnzati, district of Tutova, was discovered a head of Deinotherium, 
partly removed by visitors, also other remains, preserved as follows: 

1. Right branch of lower jaw with five molars, P;-Ms3, referred to D. gigantissimum [Stefanescu, 
1896 (1899), Tab. 1]. 

2. Portion of left branch of lower jaw with two posterior molars, referred to D. giganteum 
[Stefanescu, 1895, p. 175]. 

3. Fragment of right branch of upper jaw with portion of palatal bone and three molars, 
referred to D. gigantissimum |Stefanescu, 1896 (1899), Tab. m1]. 

4. Tenribs, one of which measures 1200 mm., of doubtful reference, Stefanescu, 1892, p. 83. 

5. Scapula, measured zn situ, transverse diameter 1150 mm., Stefanescu, 1892, p. 83. 

Fig. 61. Outline of referred grinding teeth of Deinotherium gigantissimum, after Stefiinescu, 1895, Tab. tv and y. Reduced to two-thirds 
natural size. 
A, Second left inferior molar, 1.Mo, from MAnzati. 
B, Third left superior molar, 1.M®, from Giiceana. 

According to the above descriptions of Stefiinescu of 1892, 1895, 1896 [1899], remains referred to Deinotherium 
giganteum and to D. gigantissimum occur in the same horizon in the Manzati Valley (see Stefinescu, 1892, fig. 24, 
section through Manzati Valley). 

Hisroricat Review.—In the Introduction to his paper of 1895 (pp. 173-177) Stefinescu gives a full historical 
review, ‘Historique du Dinothérium,”’ of the successive discoveries of remains of Deinotherium together with 
discussions as to the anatomy and affinities of this animal, beginning with Cuvier (1822) and ending with 

Hollander (1877) and Bieber (1884), namely, from the Tapir gigantesque of Cuvier to the Franzensbad 


Stefainescu’s original description of the type molar of Dinotherium gigantissimum, “Découverte d’une molaire 
de Dinothérium en Roumanie,”’ 1878, is not accessible for the present Memoir. The accompanying outlines (Fig. 
61) copied from Stefinescu’s paper of 1895, Tab. 1v and v, exhibit a second inferior molar and a third superior 
molar referred by him to this species. 

In 1895, however, Stefiinescu gave a full description of the discovery of Deinotherium in Rumania, under the 
subtitle ‘“Le Dinothérium en Roumanie,’ 
Boghitii, Giiceana, district of Tecuciti, referring them in part to Deinotheriwm giganteum and in part to the 
largest species which he had named Dinotherium gigantissimum. 

’ in which he mentions the discovery in 1878 of remains at Fundul 

After reviewing (op. cit., 1895, pp. 173-177) the 
chief materials found in Rumania referable to D. gigan- 
teum and to D. gigantissimum, Stefinescu describes and 
figures in detail (pp. 177-195, Tab. 1-v) the skeleton 
and teeth from MaAnzati referred to the giant species D. 
gigantissimum, including the successive stages of dis- 
covery of: (a) Inferior tusk (1889), (b) left inferior 
maxilla and large part of right inferior maxilla, also 
part of left superior maxilla (1890), two iliz, two 
scapulz, vertebree, four limbs, eighteen ribs, one tusk, 
fragments of cranium, débris of teeth and of different 
bones (1890). He concludes (p. 183): ‘‘Comme on le 
voit, j’ai eu le bonheur de trouver 4 Manzati le squelette 
presque complet de ce colossal Dinothérium; je n’ai 
trouvé cependant que deux vertébres, mais il est probable 
qu’on trouvera aussi les autres.”’ The bones were 
found within a space 10 m. in length and 3 m. in 
breadth, 30 sq. m. in all. Stefiinescu indicates the fol- 
lowing conditions regarding depth (op. cit., p. 183): 
“Ces faits prouvent ou que le Dinothérium de Manzati 
est mort envasé dans un endroit bourbeux, en cherchant 


iy boire ou A s’y rafraichir, ou que, aprés sa mort, 

Fig. 62. Front and side views of tibia and pes of Dinotherium gigan- 

: ie ; 
il a été presque aussitot recouvert d’une couche de tissimum, after Stefinescu, 1899, Tab. rv. 
limon, qui a empéché le corps de se décomposer 4 ‘®lfthnaturalsize. 

This retouched drawing is one- 

Pair, ou d’étre la proie des animaux féroces qui eussent dispersé les os et y eussent laissé la trace de leurs dents 

The following details are important: (1) In this excavation all the parts belong incontestably to one individual; 
(2) nearly all the bones were found in normal connection; (3) no doubt remains that Deinotheriwm had but three 

digits [D.II, III, 1V—Fig. 62], with a vestigial D.I in the pes; the tarsal and metatarsal bones are more or less 



From the detailed measurements of all these parts, we may extract the following: Found with the skeleton 
from MAnzati, a second left inferior molar, 1. M:, diameters, ap. 111 mm., tr. at protolophid 90 mm., index 81. 
Found at Giiceana, a left third superior molar, |. M*, diameters, ap. 107 mm., tr. 105 mm., index 98. 


Anteroposterior diameter of M2 

Anteroposterior diameter of M; 

Space occupied by five inferior 
molars, P3-M3 

Length of inferior tusks 

Length of tibia (ef. Fig. 62) 

Height of tarsus (ef. Fig. 62) 

Total length of median Mts. III 

Tapir Deinotherium 
gigantesque giganteum 
Cuvier Kaup 
85 81 

The author concludes (1895, p. 197): 

Deuxiéme molaire de la mAchoire 
inférieure [second inferior molar, 


Diamétre antéropostérieur 

Diamétre transversal de la colline 
postérieure (4 la base) 

Diamétre transversal de Ja colline 
antérieure (4 la base) 

Derniére molaire de la mAchoire 
supérieure [third superior 
molar, M*] 

Diamétre antéropostérieur 

Diamétre transversal de la colline 
postérieure (base) 

Diamétre antéropostérieur de la 
colline antérieure 

Tapirus gigan- Dinoth. gigan- 
teus de France teum d’Eppels- 
Cuvier heim Kaup 

0,085 0,081 

0,073 0,072 

0,075 0,076 




Dinothertum Dinotherium 
indicum Fale.  gigantissimum 
and Pilgrim Stefanescu 
99 111 
Dinothérium Dinothérium 
Indicum de Périm _ gigantissim. 
Falconer de Manzati 
0,099 0,111 
0,088 0,097 

de Manzati 




de Gdiceana 



The author observes in his paper of 1896 [1899], p. 127, as follows: ‘‘Si, d’une part, nous considérons les 
dimensions de beaucoup plus grandes des dents du Dinothérium Gigantissimum, en comparaison avec celles des 
autres Dinothériums connus, et surtout avec celles du Dinothérium Giganteum, si, d’autre part, nous prenons en 
considération l’existence d’un bourrelet crénelé continu A la base de toutes les molaires de la mAchoire supérieure, 
et surtout de la premiére molaire, ce qu’on ne trouve pas de la sorte chez les autres espéces, nous pouvons, 4 bon 
droit, regarder le Dinothérium de Manzati comme appartenant A une autre espéce, et conséquemment, regarder 
comme étant assez justifiée, la creation de l’espéce Gigantissimum.” 

‘Limb measurements of D. bavaricum of Franzensbad inserted below (pp. 100, 102). 



Upper Miocene of Franzensbad, Bohemia 

E. Kirrr, 1908.—The first description of this skeleton is that of Kittl in his article ‘Das Dinotheriumskelett 
von Franzensbad im k. k. naturhistorischen Hofmuseum”’ (1908). In describing the skeleton, Kittl remarks that 
this almost completely restored skeleton of Deinotherium arouses wide attention in the Vienna Museum because 
the only other skeleton which appears to be nearly as complete is in the University Museum of Bucharest. He 
speaks of the abundance of scattered skeletal materials in sand and gravel deposits of western Europe, chiefly of 
fluviatile and flood-plain origin, in contrast to the rarity of entire skeletons, in fact, an entirely complete skeleton 
had never been discovered up to 1908. The two most complete discoveries of this animal are those made near 
Franzensbad in Bohemia and near MAnzatiin Rumania, the former exhibited in Vienna, the latter in Bucharest. 

These animals belong to different species and are widely dis- 
tinguished by their size. The Franzensbad Deinotherium attains 
the dimensions of a half-grown elephant, whereas the Rumanian 
skeleton |D. gigantissimum] equals in size the largest of the living 
elephants. Its specific distinctions, however, rest less upon size 
than upon the structure of the grinding teeth; the condition of 
wear of the teeth in both skeletons indicates prolonged use. 

Comparing the [Upper Miocene] Franzensbad Deinothere with 
other skeletons, it appears to belong to a small, perhaps to the 
smallest, species of the genus Deinotherium, whereas the Rumanian 
skeleton [D. gigantissimum|] belongs to the largest variety of this 
genus. Between these two extremes we can range a series of inter- 
mediate size. It is the middle-sized Deinotherium which was first 
observed; as early as 1812 Cuvier designated them as large tapirs 
owing to the resemblance of the back teeth to those of the tapir; in 
1833 [1829] they received the generic name Dezinotherium with 
reference to their very large size. The discovery of a complete 
skull of Deinotherium in 1836 in a sand pit near Eppelsheim south 
of Mainz enabled us to understand these animals more clearly. 
It was, however, through the skeletal structure that the Deinotheres 

were first compared with the trunk-bearing pachyderms, i. e., Fig. 63. Mounted skeleton of the Upper Miocene 
. Deinotherium of Franzensbad, Bohemia, in the Royal 
elephants and mastodons, to which they have the greatest number Museum of Natural History, Vienna, referred to Deino- 

of resemblances. The most striking differences of the genus Deino- — terium bavaricum. After photograph kindly furnished 
E : . ee the present author by Prof. Othenio Abel of Vienna. 

therium, as contrasted with other proboscideans, lie in the structure 

of the skull and especially of the teeth; above all in the structure 

of the lower jaw which is strongly elongated and sharply bent downwards, the great incisor tusks bending back- 
wards and not forwards. Their second distinction is in the structure of the grinding teeth in which two sharp 
transverse crests appear, except in the middle back tooth, M?-M,, which is trilophodont. In the permanent 
dentition of the Deinotheres there are five grinding teeth in each jaw, namely, P*-P;, P*-P,, M'-M,, M?-M., M®-Msg, 
in all twenty grinders. 


The Franzensbad skeleton, attaining a length of 3.2 m. (=10 ft. 6 in.) and a height of 2.5 m. (=8 ft. 214 in.), 
is referred to the smallest species Deinotherium bavaricum. It was found in a cleft of calcareous rock near Franzens- 
bad in the year 1883. The bones belonging to it were partly complete, partly broken into numerous fragments, 
the assembling of which occupied a long time. The complete skull with a lower jaw, several of the large bones of 
the extremities, numerous vertebre and foot bones, and a large number of ribs were modeled. For this purpose 
fossil bones and models were employed from the Vienna and other museums, also the skeleton of an elephant as an 
object of comparison. 

Referring to the stature of the species of Deinothertum: 

Deinotherium bavaricum (or D. cuvierz) includes animals of the smallest size, Middle Miocene, second Mediter- 
ranean layer. 

Deinotherium levius (or D. medium) includes animals of middle size, generally distributed; D. giganteum, 
animals of the larger size, Pliocene, Congerienschichten and Belvedereschotter. 


ScHLESINGER, 1923.—The original materials included in this mount are shown in figure 64B. The mount 
as it appears in the Vienna Museum of Natural History, with the iron-work removed, is shown in figure 64A. 

From notes and observations kindly forwarded by Dr. Giinther Schlesinger (letter, Vienna, December 17, 1923) a modified description and figure of the 
skeleton of Deinotherium bavaricum are presented herewith. 
A, Photograph, side view of skeleton as mounted, with the iron uprights removed by retouching of the photographic plate. 
B, Sketch furnished by Doctor Schlesinger showing the original parts of the skeleton preserved. 
(1) OrtatnaL Parrs.—Lower jaw without incisor teeth, atlas, axis, cervicals 2, 4, dorsals 5, 9, 10, lumbars 17, 18, 22, parts of sacrum, caudal vertebra. 
Of left forelimb: Left scapula, left humerus without articular end, ulna, distal end of radius and upper proximal end, unciform, and metacarpals II, III. Of 
right forelimb: Scapula, ulna, radiale of the carpals, ulnare, trapezoid, magnum, metacarpals I, III, IV. Of the hindlimb: Pelvis, femur of right and left 
side, left and right patelle, right fibula, left caleaneum, right and left astragali, left navicular. (2) Parrs ResroreD rv PLasrer.—Sacrum partly restored. 
Cervicals, dorsals, and lumbars apparently too abbreviate. Skull and dorsal spines partly restored. (3) Errors 1n Mountine.—Scapula and pelvis appar- 
ently too upright; possibly the restoration of the extremities elongates them too much. 

e MEASUREMENTS BY PRoressor ABEL (LETTER, JANUARY 13, 1926).—Length of humerus 81.5 em., of ulna 80em., of femur 105.5 em., and of mandible 
3 cm. 


It was at the request of the present author that further information regarding the original materials and the 
restored portions of this skeleton was kindly sent by Dr. Giinther Schlesinger in a letter dated Vienna, December 
17, 1923, as follows: 

Ihren Brief vom 15 November 1923 mit Dank bestitigend beeile ich mich, die verlangten Photographien zu iibermitteln. 
Durch die Liebenswiirdigkeit unseres Kollegen Prof. Schaffer vom Naturhistorischen Museum ist es mir méglich, Ihnen zwei 
Aufnahmen in 3 Kopien zu tibermitteln [Fig. 64A, B]....Meine niheren Angaben iiber die Verteilung von Originalstiicken 
und Erginzungen in Gips werden Sie allerdings iiber das Skelett sehr enttiuschen. Wirkliche Fossilreste sind nur: Mandibel 
(ohne Incisoren), Atlas, Epistropheus, 2, 4. Halswirbel, 5, 9, 10-17, 18, & 22, Rumpfwirbel, das Sacrum und der Schwanz, 
ferner von der linken Vorder-Extremitiit Scapula, Humerus (ohne Gelenkskopf), Ulna, distales Ende und oberstes proximales 
Ende des Radius, Unciforme und Metacarpale 2 & 3. Von der rechten Vorder-Extremitit sind original: Scapula, Ulna, 
Radiale, Ulnare, Trapezoid, Magnum, und Metacarpale 1,3,&4. Vonden Hinter-Extremititen ist das Becken original ferner: 
Femur sin. et dext., Patella sin. et dext., Fibula dext., Caleaneus sin., Astragalus sin. et dext., Naviculare sin. Alles andere ist 

Die Montierung des Skelettes halte ich mit Ihnen fiir zu hoch. Der Fehler diirfte daher gekommen sein, dass das Sacrum 
nur zum Teil vorhanden ist, und die Ergiinzung der Wirbelkérper anscheinend zu kurz ausgefallen ist. Auch scheinen mir 
Schulterblatt und Becken zu steil zu stehen. Vielleicht ist auch bei der Gipsergiinzung der Extremitaitenknochen etwas zu 
viel an Liinge herausgekommen. Allerdings diirfte dies kaum erhebliche Werte ausmachen. 

ABEL, 1924.—At the request of the present author, Prof. Othenio Abel in a letter dated Vienna, January 8, 
1924, kindly supplemented in somewhat more detail the information conveyed by Doctor Schlesinger, accom- 
panying it with a new and more direct frontal photograph (see Fig. 63): 

[1. Original and restored parts] Vor allem diirfte es fiir Sie wichtig sein, zu erfahren, welche Knochen erhalten und 
welche ergiinzt sind. Ich erfuhr gestern, dass Dr. Schlesinger Ihnen bereits geschrieben hat, wovom er mich nicht verstindigt 

hat; so weiss ich nicht, ob er Ihnen diese Liste mitteilte. Ich habe sie heute Vormittag aufgenommen und zwar sind von diesem 
Skelette erhalten: 

Unterkiefer (ohne Incisivem) 10 Brustwirbel (mit Neurapophyse) 
Atlas lve ne ( nur Zentrum) 
Epistropheus 18 as SF 
4 Halswirbel 4 (vorletzter) Lendenwirbel (nur Zentrum) 
6 “a Sacrum 
5 Brustwirbel (nur Zentrum) 1 bis 5 Caudalwirbel 
Linke Kérperseite: Rechte K6rperseite: 
2 Rippe 1 Rippe 
Be 3 “(nur Oberende) 
6 ae 4 ae 
7 “ce 5 ae 
Sas 7 ‘ (nur Oberende) 
17 “e 10 ae 
ll ae 
12 oe 
Scapula Scapula 
Radius Radius (nur untere Epiphyse) 
Ulna Ulna 
Unciforme Unciforme 
Metacarp. II, III Metacarp. I, III, IV 
Becken (sehr fragmentarisch) 
Femur Femur 
Patella Patella 
Astragalus Astragalus Fig. 65. Deinotherium bavaricum in the Vienna 
Calcaneus Museum. Model by Othenio Abel (1932), about one- 

Naviculare fiftieth natural size. 




[2] Dimensionen: Distanz vom Unterende des linken Unterkiefers (Angulus) bis zum 
Vorderrande der Stosszahnalveole: 66 cm. [=2 ft. 2 in. approx.] 
Lange des linken Humerus: 75 cm. [=2 ft. 5% in. approx.] 
Linge des linken Femur: ca. 100 em. [=3 ft. 314 in. approx.] idl 

[3] Was die Rekonstruktion des Skelettes anbetrifft, so scheint mir vor allem ein Fehler darin zu liegen, dass die Richtung 
der Neurapophysenachsen ficherférmig divergiert, was unmoglich ist. Ferner steht das Sacrum viel zu hoch tiber der Stand- 
flache. Es muss tiefer herabgesenkt werden. Die Stellung ware etwa so wie auf meiner Rekonstruktion des Dinotherium 
giganteum, von der ich Ihnen eine Photographie beischliesse (vgl. ‘ Lebensbilder aus der Tierwelt der Vorzeit,’ Fig. 82, pag. 91). 
Sodann steht die linke Scapula ganz unrichtig, weil zu weit nach einwarts gedreht und ausserdem ist die Hohenlage zum 
Thorax unrichtig angenommen, wie aus dem Vergleiche mit dem rezenten Elefanten hervorgeht. Wenn Sie meine Pause der 
Photographie mit dem Skelette eines Elefanten in den ‘Recherches sur les Ossemens fossiles,’ TPS Tek, a. pag. 204, Zur 
Deckung zu bringen versuchen, mit dem die Photographie des Dinotherium bavaricum in der Grésse gut tbereinstimmt, so 
werden Sie sehen, dass sich Dinotherium bavaricum in tiberraschender weise als in den allgemeinen Proportionen durchaus wie 
ein rezenter Elefant verhilt und sich von Bunolophodon ete. unterscheidet. Darum habe ich auch in meiner Rekonstruktion 
des D. giganteum, das sich von D. bavaricum nur unwesentlich, abgesehen von der Grésse und einigen anderen unbedeutenderen 
Merkmalen unterscheidet, den lebenden Elefanten zum Vorbilde genommen. 

[4. Hapits] Ich halte Dinotherium fiir einen Laubfresser, der seine Stosszihne zur Nahrungsaufnahme nur insoweit 
verwendete, als er mit ihrer Hilfe Baumiiste abbrach, wie ich dies in meiner Rekonstruktion andeutete. Ausserdem hat Dino- 
therium seine Stossziihne wohl auch als Waffen verwendet, hat aber damit kaum den Boden erreichen kénnen, ebensowenig wie 
ein Elefant in der Standstellung dies zu tun vermag (ich denke an den indischen Elefanten). Dinotherium ist selbst verstindlich 
etwas ganz anderes als Mastodon oder Bunolophodon hinsichtlich seines Korperbaues. Unter den fossilen Proboscidien der 
Tertiirzeit kommt es meines Erachtens dem Elefanten der Gegenwart am néchsten. 

Fig. 66 Fig. 67 

Fig. 66. Restoration of the head of Deinotherium modified from Gregory and Osborn (Osborn, 
1910.346, p. 247). It will be observed that the proboscis is drawn much shorter than that in the 
restoration of Abel (Fig. 67); this short proboscis is probably erroneous. 

Fig. 67. Restoration of Deinotherium, after Othenio Abel, “‘Lebensbilder aus der Tierwelt der 
Vorzeit,”’ 1922, p. 91, fig. 82, showing the animal with a hairy coating. It willbe observed that the pro- 
boscis is drawn much longer than in the Osborn-Gregory restoration (Fig. 66); this is probably correct. 


British Museum. W. D. Marruew, SepTemBer, 1920.—The skull of Deinotherium restored and figured by 
Kaup is complete and very little crushed. The occiput pitches forward almost horizontally, with a great nuchal 
pit. The upper tusks are certainly absent. The breadth and the overhang of the nasal region are mostly natural. 

AnpreEws, 1921.—The original skull purchased from Kaup by the British Museum was carefully described by 

Charles W. Andrews in the year 1921 (p. 532) as follows: 

The skull of Dinotherium is, in many respects, one of the most remarkable known. Although fundamentally its 
characters are clearly Proboscidean, nevertheless it differs widely from the skulls of the other members of the group and, 
indeed, in some respects from that of any other mammal. In the true Elephants and Mastodons the peculiar form of the skull 
is mainly due to the enormous development of cellular bone in the occipital region, increasing the area available for the 
attachment of the muscles necessary for the support of the heavy trunk and tusks. 


In Dinotherium, although the upper tusks are wanting, the trunk, judging from the large size of the nasal opening, must have 
been enormous, and the weight of the head was further increased by the great deflected mandibular symphysis with its large 
tusks. In this case, however, the area for the attachment of the supporting muscles was supplied by the widening out of the 
occipital surface, which was further increased laterally by the extension outwards of the squamosals. Little or no cellular bone 
seems to have been developed, the occipital surface above the post-tympanic flanges being nearly flat except for the depression 
for the nuchal ligament. ‘This flattening of the occiput, combined with its forward inclination, must have made it possible 
for the animal to move its head up and down through a large are, a movement perhaps connected with the use of the downwardly 
directed lower tusks. The great width of the proboscis, probably rendered possible by the absence of upper tusks, led to the 
widening out of the skull in the orbital region, producing the shelf-like projection of the maxille# above noticed. The great width 
of the glenoid surface for the mandible is a peculiarity for which it is difficult to account, unless it is correlated with the general 
widening of this region of the skull. 

The skeleton of Dinotherium is still very imperfectly known, but such bones as have been described show that the animal 
must have been quite Elephantine in structure and appearance except as to its head, the legs being pillar-like and the neck 
short. A femur probably associated with the skull above described measured 150 cm. in length. The numerous speculations 
as to the appearance and habits of Dinotherium have been summarized by de Blainville and Stefanescu in their works referred 
to above. Most writers seem to suppose that the animal was chiefly fluviatile and aquatic in its habits, but there appears to 
be no good reason for believing that it was more so than the Elephants. 

InprA. Patmer, 1924.—In the Memoirs of the Geological Survey of India there appears an account by 
R. W. Palmer of an incomplete skull of Deinotherium which was published after his death in October, 1922, and 
the paper of 1921 by Charles W. Andrews, cited above, and that of 1922 by C. Forster Cooper, cited below, were 
therefore not consulted (see Palmer, Nature, Oct. 25, 1924, p. 624): 

The specimen described is an incomplete fragment of a skull showing the basal surface from the condyles to the pterygoid 
region. The anatomical features shown do not differ markedly from those described by the late Dr. Andrews for the celebrated 
skull of Dinotherium giganteum in the British Museum, but the new specimen being in better condition, certain points are 
more clearly established. With regard to the question of the validity of the Indian species D. indicum and D. pentapotamiz, 
the author concludes that they cannot be upheld and that they are both to be referred to the European form D. giganteum. 
This agrees with the views already published by Forster Cooper which were the outcome of astudy of material from Baluchistan. 

(Palmer, 1924.1) Found in the Lower Chinji Sandstone, Lower Pliocene (fide Pilgrim, 1913). 


Deinotherium hobleyi is the first species of the genus Deinotherium described from the continent of Africa; 
no mention is made by Fourtau (1918) of the presence of Deinotherium in the Lower Miocene of Moghara, although 
the Moghara locality (Fourtau, op. cit., p. 91) is described as certainly the richest in northern Africa in the remains 
of mastodonts, containing the types of ‘Mastodon’ | =Rhynchotherium(?)| spenceri and of ‘Mastodon’ {= Trilophodon| 
angustidens var. libyca, as described by Fourtau. In fact, Fourtau remarks (op. cit., p. 91): 

Mais il est un fait assez bizarre A constater, c’est l’absence totale de restes de Dinotherium, le compagnon habituel des 
mastodontes miocénes. I] semblerait done que ces animaux ne sont pas d'origine africaine, et que les restes signalés dans 
lOuganda et en Ethiopie [Footnote: ‘E. Haug.—Traité de Géologie, tome II (2), p. 1727.’], 4 des horizons bien supérieurs, 
proviendraient d’une migration postérieure 4 l’apparition des Dinotherium en Asie et en Europe. 

Sir Harry Johnston independently supports the African origin of the Deinotheres and in 1925 writes to Osborn 
as follows: 

Jounston (Lerrer, Marcu 16, 1925).—You assign to the Dinotheria a ‘European and Asiatic’ habitat, but unless I 
have been grossly misled I rather fancy that the discoveries of Hobley, Fraas and one or two other Germans investigating 
Upper Egypt have shown the Dinothere group of Proboscideans to have originated in Egypt or Equatorial East Africa. 
Hobley discovered remains of pygmy Dinotheria in the Miocene formations lying east of the Victoria Nyanza—some such 
year as 1908. When I was in Germany just prior to the War I was shown specimens collected by German palzontologists 


from Stuttgart and Munich, which, though fragmentary, certainly seemed to point to a primitive Dinotherium. These, 
I understood, were discovered in Upper Egypt [i. e., Tébessa]. T hey or I may have been wrong, however, and it is possible that 
Hobley’s East African Proboscideans may not have been Dinotheres. But think theirstatements are worth your investigation. 

AnpREws, 1911.—Seven years previous to Fourtau’s paper and twenty-eight years after the publication of 
Weinsheimer’s history, Charles W. Andrews recorded (1911.1, p. 35, Abstract, Proc. Zool. Soc. London) the dis- 
covery near Karungu, British East Africa, of a species which he named Dinotherium hobleyi, after C. W. Hobley, 
Commissioner of Mines in British East Africa. 

(Andrews, op. cit., p. 35): The specimens described were sent to the British Museum by Mr. C. W. Hobley, Commissioner of 
Mines for British East Africa. They included portions of the mandible with teeth, acaleaneum, and a patella of asmallspecies 
of Dinotherium nearly allied to D. cuvieri, from the Lower and perhaps Middle Miocene beds of France. The new species, 

which he [Andrews] proposed to call Dinotherium hobleyi, differed from D. cuviert in several particulars—e. g., the inner anterior 
column of pm 3 was more distinctly developed, and the talon of M; had a distinct tubercle on its inner side. 


Amer. Mus, 27005 

Fig. 68a. Type third superior and inferior molars (casts Amer. Mus. 27006, 
27005) of Deinotherium hopwoodi sp. noy.,from Olduvai, near the southeast shore 
: gy. { ; of Lake Victoria, regarded as of Middle Pleistocene age. One-half natural size. 
ae ig. 68. ete type figure of Dinotherium hobleyi Andrews, 1911.2, Collected by Mr. Arthur T. Hopwood of the British Museum. Observe especially 
Sasa gs. 1, la, one-third natural size. From near Karungu, British the subdivision of the summits of the crests into 14 to 16 conelets. Originals (Brit. 

rica. Mus. M. 14118, M. 14119 respectively). See page 117 below for description. 

The same author published (1911.2, p. 943) a full description of Deinotherium hobleyt, from which the following 
has been extracted: 

. .. from the neighbourhood of Karungu on the east side of Lake Victoria Nyanza. Most of the specimens are indetermin- 
able fragments, probably picked up on the surface, but in addition to these there are some beautifully preserved teeth with a 
portion of the mandible (Pl. xivmu, figs. 1, 1a) of a small species of Dinotherium: a small imperfect Proboscidean caleaneum 
(fig. 5), a patella, and some other fragments probably belong to the same animal. tt OSes size and, on the whole, in the pattern 


of the teeth this Dinotherium is very similar to D. cuvieri Kaup, which is from the lower and middle Miocene of France, being 
apparently especially characteristic of the Burdigalien horizon. Detailed comparison however shows some differences. . . . 
These differences, coupled with the remoteness of the localities in which the two forms are found, seem to justify the establish- 
ment of a new species for this East African animal and I propose that it shall be called Dinotherium hobleyi Andrews. 

BriveEs, 1919.—A. Brives notes in 1919 (p. 90) the discovery of a single tooth of Deinotherium in the sands 
of Djebel Kouif near Tébessa, which he refers to the Lower Miocene Deinotherium cuvieri and describes as follows: 

Le Djebel Kouif est un vaste plateau de 16 4 18 kilométres de tour d’une altitude moyenne de 1150 4 1200 ™, qui est situé 
prés de la frontiére tunisienne 4 27 kilométres au Nord-Est de Tébessa. . . . C’est dans ces sables qu’a été trouvée une belle 
dent de Dinotherium en parfait état de conservation. C’est la premiére fois que cet animal monstrueux est signalé dans |’ Afrique 
du Nord et ce fait a une importance capitale, car il permet de fixer d’une maniére précise l’Age des sables qui le renferme. . . . 
Les seuls débris organiques trouvés dans les assises consistaient en fragments de troncs d’arbres silicifiés dépourvus d’écorce et de 
racines. . . . Au col de Beccaria, Ph. Thomas a recueilli et cité: Araucariorylon xgyptiacum Krauss, Bambusites Thomasi 
Fliche, Palmoxylon Cossoni Fliche, Ficorylon cretaceum Schenk, Acaciorylon antiquum Schenk, Jordanta tunetana Fliche, Nicolia 
zxgyptiaca (?) Unger. . . . La présence du Dinotherium ne laisse plus aucun doute sur l’Age miocéne de ces dépdts; il reste A 
préciser maintenant si ces assises sont du Tortonien ou du Pontien. . . . La dent que je présente est identique comme dimen- 
sions 4 un exemplaire de Dinoth. Cuvieri des faluns de Maine et Loire . . . La dent du Kouif est une 2° molaire supérieure; 
elle est presque carrée (5 centimétres 4 au collet), les crétes sont légérement usées, les racines manquent. 

Leakey, 1932-1934.—To the north of Olduvai, about 300 km., is Karungu, the type locality of Deinotherium 
hobleyt (Fig. 56, 22), on the east shore of Lake Victoria. About 30 km. distant from this was found by Dr. L. S. B. 
Leakey, on the islands in the Kavirondo Gulf, a tooth of a Deinothere probably referable to the Pleistocene species 
D, hopwoodi, associated with remains of Homo kanamensis of Kanam, East Africa (see footnote, p. 85, of present 


ForstER Cooper, 1922.—In the Proceedings of the Zoological Society for the year 1922, C. Forster Cooper 
published the following observations on the Deinotheres discovered in the Lower Miocene Bugti beds of Baluchis- 
tan, also in higher geologic levels. 

Middle Pliocene, Perim Island, is the type geologic level of Falconer’s Dinotherium indicum, 1845; its specific 
characters lie in the shape of the jaw, there being no difference from D. giganteum in the teeth excepting a greater 
thickness of the enamel. Falconer (1868, p. 415) apparently refers to the same species as D. Perimense, a name 
nowhere defined. 

Upper Pliocene, Attock. Falconer mentioned (MS. labels) a smaller variety from Attock but did not 
definitely name it; this smaller variety was subsequently named Dinotherium pentapotamiz by Lydekker in 1876, 
but in 1886 he united this species as a synonym with D. indicum, regarding the differences as individual variations. 

Middle Miocene, Salt Range, Sind, India, is the type geologic horizon of Dinotherium sindiense Lydekker, 
1880; regarded by Lydekker (1880, p. 196) as distinct from D. pentapotamix; the type (Lydekker, op. cit., Pl. 
XXXI, fig. 4) regarded by Forster Cooper as too fragmentary to afford adequate evidence of the specific distinctions. 

Lower Miocene, Bugti beds, Bugti Hills, Baluchistan, is the type geologic level of Dinotherium ndricum 
Pilgrim, 1908. (Pilgrim, 1908, p. 157): “It differs very markedly from the other known species.” Subsequently, 
however (1912, p. 16), Pilgrim withdrew the name D. ndricum and substituted the name D. pentapotamiz 
[indicum] var. gajense. 

Forster Cooper (1922, p. 621, 622) considers that the characters assigned by Falconer and Pilgrim, and the 
published figures, lie within the limits of variation of D. giganteum of Darmstadt, e. g., Brit. Mus. collection No. 
M.3494. In this Cooper agrees with Weinsheimer (see p. 94 of the present Memoir). 


Buett Beps (Forster Coorrr, 1922, pp. 622-624).—The material of the present collection from the Bugti beds is, as 
is the common experience, both scanty and fragmentary, and consists only of part of a lower jaw and some separate upper and 
lower teeth. The lower jaw (text-figs. 8 & 9) belongs to a small form, and in size and structure compares closely with D. hobleyi 
[Footnote: ‘Andrews, P. Z.S.1911.’]. The third ridge of the last molar is not parallel to the front ones, but slopes at an angle 
similar to that found in D. hobleyi and D. levius. Of the separate teeth, none shows any great difference from specimens of 
European forms with which it has been compared. An upper molar, probably a second, is here figured (text-fig. 10). There is 
nothing in it which can be construed as a ‘longitudinal ridge,’ nor is there anything in the nature of a ‘cusp blocking the valley’ 
which has not been found within the range of variation of European specimens. A third upper premolar (PI. rv, fig. 6) is also 
figured to show the separation of the inner cusps said to be characteristic of the Indian forms, but which, as has just been 
stated above, occurs also in D. giganteum. The remaining teeth in the collection are not distinguishable from the smaller forms 

from Europe, and call for no comment except for three specimens. 
Comparative measurements (pp. 625 and 626) of the lower and upper grinding teeth of Deznotherium sp. and 
of Trilophodon angustidens from the Bugti beds of Baluchistan, from Chevilly (Lower Miocene), and from 
Sansan and Simorre (Middle Miocene) of France are given, and the following conclusions are reached. 
Summary.—The Proboscidea of the Bugti beds of Baluchistan include Trilophodon angustidens | =cooperi 
Osb.] and a small Deinotherium. The Indian 7. angustidens is a small animal, more primitive than the Middle 
Miocene forms of Sansan, for which the name 7’. angustidens paleindicus may be retained. 
The Deinotherium resembles the smaller European Deinotheres, 
also the D. hobleyi of Africa. It is doubtful in our present state of 
knowledge whether the Indian Deinotheres are separable from the 






So \. All 1/12 nat. size 

5 Procressive INcREASE IN Size of MANDIBLE or DeINoTHERIUM Fig. 70. Restorations (1932) by Margret 
Fig. 69. Outline to same scale, namely, one-twelfth natural size, of: Flinsch under the direstion of H a Fairf Id 
ee Adult lower jaw of Deinotherium gigantissimum as partly restored by Stefiinescu, Oca One-hundredth natural a Y : 
after Stefainese 399. T's : 
e Adult lower ase Dane gigantewm, after Kaup, 18: ly i was Fae hace ee ee 
: : aM se aup, 1835.1, Add Tab. 1, fig. 5. Deinotherium bavaricum and the incomplete 
A, Juvenile lower jaw of Deinotherium cwieri ref., after Lartet, 1859, Pl. xi, fig, 4. skeleton of D. gigantissimum 




Mayet, 1908, pp. 199—-206.—This valuable specific revision is the first to treat the ‘mutations’ and species of 
Deinotherium in ascending geologic order, beginning with the Lower Miocene Deinotheriwm cuvieri of the Sables de 
VOrléanais and continuing with the Middle Miocene ‘D. bavaricum,’ with the Upper Miocene D. levius, and with 
the Lower Pliocene D. giganteum, as follows: 

D. cuviert 

Dinotherium de taille relative- 
ment petite, atteignant 4 
peine la moitié de celle du 
D. giganteum. 

Superior Dentition 

(1) Coneavity of crests fac- 
ing posteriorly, M!' three 
crested, M?, M* two crested, 
crowns, with the exception of 
M', less broad than long. M!, 
crests gently concave, trito- 
loph diminishing in breadth, 
total length 72 mm. M?, 
transverse crests incompletely 
reunited internally, tritoloph 
narrower than  protoloph, 
length 60 mm. M$, length 
62 mm. 

Inferior Dentition 

(2) Concavity of crests 
turned forward, molars longer 
than broad. Ps, protolophid 
thick and divided into two 
points, length 43 mm. P,, 
length 48 mm. Mj, atrophy 
and narrowing of tritolophid, 
length 60 mm. Mbp, thickening 
of cingulum representing the 
talon, length 59 mm. Ms, 
talon thick, narrow, triangular, 
detached from the crown, re- 
flected backward, length 71 

(3) Tusks somewhat short 
and obtuse. 

D. bavaricum 
(Burdigalien?) Helvétien 

Taille  sensiblement plus 
grande que celle du D. 
Cuvieri, mais moindre que 
celle du D. levius et surtout 
du D. giganteum. 

Superior Dentition 

(1) Same characters as in 
D. cuviert. M', same char- 
acters as in D. cuvieri. M?’, 
same characters as in D. 
cuviert, excepting length 72 
mm., breadth of protoloph 69 
mm., of metaloph 66 mm. 

Inferior Dentition 
(2) Same observations as in 

D. cwiert. Ps, protolophid 
thick and bifid. My, trito- 
lophid narrower than the 

anterior crests, length 73 mm. 
M2, talon greatly reduced, 
length 63 mm. Ms, talon 
triangular, contracted as in 
D. levius, well detached from 
the crown, and thrown back- 
ward, length 72 mm. 

D. levius 


Taille un peu moindre que 
celle du D. giganteum. 

Superior Dentition 
(1) Same observations as in 
D. cuviert. P*, narrow in- 
ternally and somewhat. tri- 
angular, length 66 mm. P%, 
length 70 mm. M?', marked 
abbreviation in breadth of 
tritoloph, other characters 
more pronounced, length 84 
mm. M?, metaloph narrower 
than protoloph, length 73 mm. 
M?, two lobes more unequal 
than those of M?’, metaloph 

smaller, length 77 mm. 

Inferior Dentition 

(2) Same observations as in 
D. cwviert. Ps, anterior lobe 
compressed into a single tri- 
angular point, etc., small pos- 
terior talon, length 55 mm. 
Ps, proto- and metalophid 
gently concave anteriorly and 
somewhat united on outer side. 
Pre- and postcingula little de- 
veloped, length 70 mm. My, 
reduction of tritolophid in 
breadth, length 78 mm. Mb, 
postcingulum smaller, more 
detached than in Me of D. 
giganteum, length 73 mm. 
Ms, posteingulum narrow, tri- 
angular, detached, length 80 

(3) Tusks shorter, thicker, 
less sharp than in D. gigan- 

(4) Mental foramen placed 
more anteriorly below Ps. 

D. giganteum 

Trés grande taille. Dimen- 
sions des dents fort 


Superior Dentition 

(1) Same observations as in 
D. cuvieri. P*, of subquadrate 
form, length 80 mm. P‘, 
length 78 mm. M!, three 
transverse crests, i. e., proto-, 
meta-, and tritoloph, gently 
concave posteriorly, tritoloph, 
less strong and less broad, 
length 95 mm. M?, two trans- 
verse crests, i. e., proto- and 
metaloph, and crenulated cin- 
gulum, length 91 mm. M*, 
proto- and metaloph approxi- 
mately equal, length 91 mm. 

Inferior Dentition 

(2) Same observations as in 
D. cuviert. Ps, anterior lobe 
simple, narrow, single pointed, 
length 63 mm. P4, two trans- 
verse crests, 1. e., proto- and 
metalophid, length 70 mm. 
M,, three transverse crests, 
subequal, tritolophid slightly 
more narrow than metalophid, 
length 104 mm. Mb, two 
transverse crests, length 83 
mm. Ms, talon nearly as 
broad as metalophid, crenu- 
lated, closely applied to base of 
metalophid, length 95 mm. 

(3) Inferior tusks long, point- 
ed at extremity. 

(4) Mental foramen at level 
of Ps or at interval between 
P3 and Pa 


Fig. 71. Proportions of Deinotherium giganteum, after Lartet, 1859, Pl. x11, figs. 1-4, reduced to one-third natural size. Lartet remarks (op. cit., pp. 
506, 507): ‘Les piéces en nature qui ont servi de modéle au dessinateur pour recomposer, dans les figures 1 et 2 de cette planche, les séries théoriques de pre- 
miére et de seconde dentition, sont de provenances diverses et probablement aussi d’espéces distinctes; mais on s’est attaché 4 ramener uniformément toutes 
ces dents aux proportions du Dinotherium gigantewm, avec réduction 4 demi-grandeur de nature.’’ Compare Palmer, 1924, Pl. 111, Deinotherium sp. (?). 

Fig. 1, a, b, c, Superior milk premolars, Dp”, Dp®, Dp’ (upper row). 

a’, b’, e’, Inferior milk premolars, Dp2, Dps, Dps (lower row). 
Fig. 2, A-E. Permanent dentition, A=P*, B=P*, C=M!, D=M?, E=M? (upper row). 
A’-E’, Permanent dentition, A’ = P3, B’ = Ps, C’=My, D’ = Mp, E’ = Ms (lower row). 
Fig. 3, Fragment of jaw, c= Dp, being replaced by B= P4; C = My [H.F.O.]. 
Fig. 4, Juvenile jaw showing below A and B=P3 and Py, D= Mbp, above a-c = Dp2, Dps, Dps, C = My. 


The above synthetic illustration contains materials from Lower Miocene to Lower Pliocene horizons, as follows: 
Fig. 1, a, b, = Deinotheriwm cwvieri, Lower Miocene of Pontlevoy (Loir-et-Cher). 

c, Middle Miocene of the sub-Pyrénées. 

a’, Upper Miocene of Simorre (Gers). 

b’, Miocene of Gers. 

ce’, = Deinotherium bavaricum (?) ref., Upper Miocene. 
g. 2, A=Deinotherium cuvieri (?), Miocene of Gers. 

E, M®, = Deinotherium giganteum, Lower Pliocene of Eppelsheim. 
Fig. 3, Fragment of jaw from Ile-en-Dodon (Haute-Garonne), Middle Miocene. 
Fig. 4. Jaw, Ile-en-Dodon (Haute-Garonne), Middle Miocene. 
For further details, see Lartet’s minute description of Plate x1, 1859, pp. 506-510. 




1908. SyNoNyMy AND CITATIONS (FIDE Mayet, 1908) 

Dinotherium Cuvieri Kaup, Deinotherium bavaricum von Meyer, Deinotherium giganteum Kaup, 
1831 [1832] 1831 1829 

1812 Tapir gigantesque Cuvier 1831 Dinotherium medium et secon- 1861 Déinotherium levius Jourdan 
darium Kaup 

1825 Tapir gigantesque Cuvier 1832 Dinothertum bavaricum von Meyer 

1831 [1832] Dinotherium Cuvieri Kaup 1850 ee intermedium de Blain- 

1850 Dinotherium Cuvieri de Blainville 1859 Dinotherium bavaricum Lartet 

1858 [1859] Dinother‘um Cuviert Lartet 1875 Dinothertum bavaricum Bachmann 

DEINOTHERIUM CUVIERI (OBSERVATIONS OF Mayet, 1908, pp. 199-206).—Remains very abundant in the 
Lower Miocene Orléanais, especially at Chevilly, chiefly isolated bones and teeth, type mandible from Chevilly. 
Entire dentary series represented by remains from Chevilly, Bricy, Boulay, ete., species clearly distinct from the 
Middle Miocene D. bavaricum, the Lower Pliocene D. giganteum, or the Upper Miocene D. levius. 

Dinotherium cuvieri is the sole species found in the Sables de |’Orléanais; certain molars approach those of D. 
bavaricum, a form of much larger size. Characters of D. cuviert notably different from those of D. giganteum or D. 
levius, the latter type coming from the Upper Miocene of Grive-St.-Alban, Rhone basin. Thus D. cwvieri appears 
suddenly at the beginning of the formation of the Sables de |’Orléanais, survives into the Faluns du Blésois, and 
then disappears, to be succeeded by D. bavaricum, an ascending mutation of larger size. 

DEINOTHERIUM BAVARICUM (OBSERVATIONS BY MAyYET, 1908, pp. 205, 206).—Von Meyer’s type of Deino- 
therium bavaricum from Gmiind, Bavaria, is a Deinothere of medium size closely intermediate between the Lower 
Miocene D. cuwviert and the Lower Pliocene D. giganteum. The dental characters of D. bavaricum resemble those 
of D. cuviert more closely than those of D. giganteum. From the Sables de |’Orléanais, Chevilly, one molar only 
is mentioned [?] as representing D. bavaricum. Although D. bavaricum of the Upper Miocene approaches in 
size D. levius of the Lower Pliocene [Upper Miocene] the dental characters of D. levius show much stronger 
affinity to those of D. giganteum. Consequently the larger teeth of the Sables de l’Orléanais probably belong to 
large examples of D. cuviert. 


Anteroposterior Measurements Be PY OM ME MP Pe PM Meee es 

Deinotherium gigantissimum 111 348-357e 
¢ indicum 100 99 
wi giganteum 80 78 95 91 91 63 70 = 104 83 95 282 
H levius 66 70 84 73 77 55 70 78 73 80 231 
¢ bavaricum 72 73 63 72 208 
. cuviert 72 60 62 43 48 60 59 71 190 

Osborn, 1925: From the above observations of Mayet it becomes perfectly clear that Deinotherium, like all 
other large herbivorous quadrupeds, passed through a series of ascending mutations, witnessed (a) in every single 
character of the superior and inferior grinding teeth, also (6) in the steady increase in size. The net anteroposterior 
diameter of the three lower grinding teeth, obtained by adding the length in millimeters, and including a 
comparison with D. giganteum, is as above. 


1. ProGresstvE DouBLING IN Sizy.—In the above table the measurements given by Mayet of Deinotherium 
cuvieri, D. bavaricum, D. levius, and D. giganteum are amplified by the Falconer (1868, p. 407) and Lydekker 
(1880, p. 189) measurements of D. indicum and by the Stefanescu (1896 [1899], Tab. 1, 111) measurements of D. 
gigantissimum; they demonstrate a progressive increase in size, from the Lower Miocene stage (D. cuviert) to the 
Pliocene stage (D. gigantissimum). 

The second inferior molar, Mz, of D. gigantissimum (=111 mm.) is about 95% longer than the second inferior 
molar of D. cuvieri (=59 mm.). The inferior molar series, M,3, of D. gigantissimum (=348-357e mm.) is also 
about 95% greater than the inferior molar series of D. cuviert (=190 mm.). 

Consequently, corresponding to the doubling in size of the dental series, we may roughly estimate that the 
Mid-Pliocene D. gigantissimum was an animal about double the size of the Lower Miocene D. cwviert. 

These proportions shown in the lower jaws of Deinotheriwm cuvieri, D. giganteum, and D. gigantissimum 
(Fig. 69) are also expressed in the diagrammatic restorations (Fig. 72) in which D. gigantissimum is represented 
as nearly double the size of D. bavaricum, with D. giganteum as intermediate in its skeletal proportions. 

Pwr GAN @ © 




Fig. 72. Proportionate restorations of the year 1930, in profile, of Middle to Upper Miocene and Lower to Middle Pliocene specific stages of Deinotherium, 
with an estimated shoulder height of: 
Deinotherium bavaricum 8 ft. 24 in. 
Deinotherium giganteum OSG 
Deinotherium gigantissimum 12 

2. GroLocic Succession or Species.—The Eurasiatic distribution of the Deinotheres, as carefully 
worked out for the first time in the accompanying correlation table, displays the ascending geologic succession of the 
species in western and eastern Europe, in southern Asia, and in Africa. When we compare the small Deinotherium 
indicum gajense from the Lower Miocene of India with the D. pentapotamix {[=D. indicum] of the Upper 
Miocene of India, we observe that there is a closely corresponding progressive increase in size of the 
Deinotheres in all parts of Eurasia where they have been discovered. The approximate geologic parallel between 
the Deinotheres of western Europe and those of corresponding horizons in India is displayed in Table I (Geologic 
Correlation) and Table II (Specific Distribution) of the present chapter. 

3. SpEctric CHARACTERS IN Successton.—As explained in the introductory paragraph of this chapter, 
“It is not possible for the author to treat the Deinotheres in the same critical or exhaustive manner as the species 
of proboscideans belonging to other families are treated in the present Memoir; nor is it possible to give the type 

descriptions or reproductions of the type figures as in other chapters.” The reader is referred to the original type 


descriptions and figures wherever published, as indicated in the ‘‘Order of Discovery and Description of Twenty- 
six Species of Deinotherium described or named, 1715-1935” (p. 84 above) and in the corresponding bibliographic 
references below to the works of Kaup, von Meyer, Eichwald, Koch, de Blainville, Falconer, Jourdan, Lydekker, 
Stefinescu, Pilgrim, Andrews, and Ehik, the only paleontologists to our knowledge who have described type species 
and specimens of Deinothertum. See also Haug, Joleaud, and Leakey in Bibliography. 

4. Dentition RELATIVELY NON-PROGRESSIVE.—As compared with other proboscideans both mastodontoid 
and elephantoid the dentition of the Deinotheres is relatively non-progressive; the fundamental pattern of the 
grinding teeth was established extremely early in geologic time certainly during the unknown Oligocene stages, 
because in the Lower Miocene specific stages it is fully established, especially the number of ridges and the char- 
acter of the crests of the upper and lower true molars, M1-M 3. 

Mayet (1908, pp. 199-206) in his valuable table of specific distinctions (translated in full above) pointed out 
that the chief progressive characters between Deinotheriwm cuviert, D. bavaricum, D. levius, and D. giganteum 
consist in (a) progressive increase in size, (b) detailed changes of proportion between the anterior and posterior 
crests, protoloph, metaloph, and tritoloph, (c) in the greater or lesser union of the crests externally and internally, 
(d) in the shape and size of the incisive tusks, and (e) in the relative position of the mental foramen of the jaw. 

Mip. PLiocENE D.  gigantissimum| D. indicum Type 
Type D. perimense Fale. 
D. giganteum ref. (=D. indicum) 
D. angustidens 
Ty Pieiicer yee oe 
Low. PLioceNE D. gig. majus D. giganteum Type | D. uralense Type 
D. gig. medium (=D. maximum T., D. 
D. gig. minus medium T., D. secun- 
darium T.(?), D. 
kénigit T.) 
Up. Miocene D. levius Type D. bavaricum Type D. proavum Type | D. Bate co Be 
D. intermedium Type | D. minutum Type D. bavaricum ref. Type 
Min. Miocene D. giganteum(?) ref. | D. bavaricum ref. D. sindiense Type 
D. secundarium (?) ref. 
Low. Miocene | D. cuvieri ref. | D. cuvieri Type D.ndricum Type( = 
D. hobleyi D. indicum gajense)| 

Compare pages 84, 85, 114, and 115. 


These specific changes, as pointed out by Mayet, are of minor grade as compared with the profound changes 
which occur in the grinding teeth of other proboscideans by the addition of crests, by progressive hypsodonty, etc. 

The conclusion is therefore justified that Deinotherium is relatively conservative in the structure of its dentition, 

while sharing with all other proboscideans progressive increase in size. 


GEroLocic ORIGIN AND RanNGE.—Deinotherium appears simultaneously in the Lower Miocene of Africa in D. 
hobleyi type and D. cuvieri referred, which apparently are in a stage of evolution similar to that of D. cwviert type 
of the Lower Miocene Sables de l’Orléanais of France, which, in turn, are similar in size to the D. indicum gajense 
type of the Lower Miocene Bugti horizon of Baluchistan. These three specific types, which agree approximately in 
size, require very close examination and comparison before it can be determined which is the most primitive. 

The evidence seems to point to an African origin of Deinotherium, since there is no indication of ancestral 
forms of this genus in any Eocene or Oligocene horizon of Eurasia at present known. 

Mrcration.—Well defended by its tusks, well nourished by its precociously lophodont grinding teeth, well 
transported by its long limbs, Deinotheriwm spread rapidly through northern Africa, southern France, and far 
eastward into the peninsula of Baluchistan and India. Its traveling companions were the Trilophodon angustidens 
libycus of the Moghara desert, northern Africa, the T'rilophodon angustidens of the Loire, Sables de |’Orléanais, and 
the Trilophodon coopert Osb. of Dera Bugti, Baluchistan. 

While ranging in the same geographic region, Deinotherium was very divergent in habit and local habitat from 
Trilophodon, as indicated by the following adaptive and generic contrasts: 


Cranium and jaws abbreviate. Proportions 
brachycephalic. A very elongate proboscis capable 
of reaching higher branches of trees also the ground. 

Superior tusks early aborted. Inferior tusks 
rounded, downcurved, recurved, aiding the probos- 
cis in the prehension of branches and leafage. 

Grinding teeth sharply crested; Dp4 and M1 
trilophodont, P4, M2, and M3 bilophodont; total 
number of complete transverse crests not exceed- 
ing seven; all grinders in use at same time; per- 
sistently brachyodont, never hypsodont; prob- 
ably resembling Tapirus in browsing habit on leaf- 
age. Superior grinders tending to form ectolophs, 
like those of primitive tapirs. 

Limbs elongate, increasingly elephantoid, raising 
body well off the ground. Feet reduced to three 
short functional digits in manus and pes. Dorso- 
lumbar vertebr and trunk abbreviate. 


Cranium elongate; lower jaws excessively 
elongate. Limited development of the proboscis, 
probably a prehensile upper lip, compensated for by 
great elongation of the lower jaw. 

Superior tusks persistently progressive in size, 
downcurved, aiding the prehensile upper lip in the 
prehension of food. Inferior tusks horizontal, spatu- 
late, aiding the proboscis in the prehension of food, 
also employed in the digging and uprooting of plants. 

Grinders persistently bunomastodont in pattern; 
transverse crests increasing to four or five, adapted 
to the crushing of food, tending to hypsodonty; 
finally only the posterior grinders, M3-Ms, in use in 
advanced age. Browsing and partly grazing habit 
on low leafage and uprooted plants. 

Limbs mastodontoid. Feet tetradactyl to pen- 
tadactyl; digits abbreviate. Trunk elongate; low 
bodied; pelvis broad; mastodontoid in proportions. 

The above contrasts of structure and habit prove that from the very first the Deinotherioidea were sharply 
divergent from the Mastodontoidea. While the geographic range is very similar, as far at least as southern Asia, 
Deinotherium has been found only once in the north (D. uralense), whereas Trilophodon was a bold northern 
migrant reaching the 40th parallel of the United States in Middle Miocene time. 


Consistent with their divergence of structure is the fact that, while fossil remains of Deinotherium are some- 
times found in the same horizon with fossil remains of Trilophodon, more frequently they are found separate. 

ProBABLE Hapits AND Hasirat.—There is little in the limb, foot, trunk, or skeletal structure of Deino- 
therium, now that it is quite fully known from the wonderful Franzensbad specimen (Fig. 64), to support the 
original theory that this was a fluviatile or water-loving animal which frequented streams and used its lower tusks 
for the prehension of food from the banks. Yet during warm seasons it doubtless bathed (Fig. 73) in shallow 
waters, like the modern Indian elephant. 

Rather we incline to conclude that it was a forest-living animal, subsisting upon leafage and tree boughs, to 
which its sharply crested grinding teeth were adapted like those of the tapir and of the tree-or shrub-browsing types 
of rhinoceroses. The very powerful trunk, indicated in the Eppelsheim cranium (Figs. 57, 58, 59), was eminently 
adapted to the collecting of tree boughs and leafage, as in the case of the existing African and Indian elephants; 
the open supranarial space is enormous. The absence of large superior tusks, which are seen in all the Mastodonti- 
de and Elephantide, explains the relatively flattened form of the top of the cranium and the forwardly inclined 
occiput, in wide contrast to the elevated, hypsicephalic and brachycephalic cranium, with vertically placed occiput, 
for the attachment of large neck muscles to counteract the strain of the greatly elongated tusks. 

Consequently Deinotherium was neither mastodontoid nor elephantoid in profile but relatively flattened and 
depressed. The profile aspect of the skull shown in our restorations (Figs. 72, 73) is totally different from that of 
either Mastodon or Elephas, while the proboscis was quite as large as that of these animals, and the body height 
and length of limb approximated that of the largest Proboscidea, as also shown in the restoration (Fig. 70) in 
comparison with other proboscideans drawn to the same one-hundredth scale. 


Species of eight mastodont genera, namely, Trilophodon, Anancus, Rhynchotherium, Zygolophodon, Turicius 
Miomastodon, Serridentinus, and Tetralophodon, were contemporaneous with Deznotherium, as shown in Table 
II below (Geologic and Geographic Distribution of the Types of the Superfamily Deinotherioidea, as compared 
with the Types of the Contemporary Mastodontoidea and Stegodontoidea’). 

The student should turn immediately to Table II below, in which the geologic and geographic distribution 
of most of the types of these deinotherioid and mastodontoid species are carefully set forth, also to the geographic 
distribution charts showing exactly where these types were found; by this means the zoologic and geographic 
environment of the Deinotheres is clearly brought out. 

It will be observed that while the Deinotheres belong chiefly in competition with animals of the Masto- 
dontoid period, they are now known to survive into the period of Elephantoid distribution. 

Aided by different local adaptive radiation, probably in the forested regions of southern Eurasia, the Deino- 
theres with their brachyodont, tapiroid grinding teeth were able to compete with the bunomastodont genera, 
Trilophodon, Tetralophodon, and Anancus. It is probable that they enjoyed an entirely different local geographic 
range from the purely zygolophodont genera, Zygolophodon and Turicius. 

It is not an unreasonable hypothesis of extinction that the brachyodont Deinotheres were unable to compete 
with the incoming Stegodonts with their very numerous transverse crests and tendency to hypsodonty. At all 
events, the climax of the series (Deinotherium gigantissimum) in southern Eurasia becomes extinct in Middle Plio- 
cene time, just prior to the appearance of numerous species of Stegodon. In Africa the Deinotheres survived. 

This hypothesis of the failure of the brachyodont, tapiroid types of Deinothere grinding teeth to compete 
with the subhypsodont and hypsodont types of the rising superfamily Elephantoidea' will afford a parallel in the 
Proboscidea of what we observe in the ungulates all over the world, because in Pliocene time hypsodont ungulates 
began to prevail over the brachyodont ungulates, as first observed in Kowalewsky’s great Memoir of 1873. 

‘See footnote 2 on page 22. 


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Fig. 73. 

climax in size, to which the specific name gigantissimum appropriately refers. 



The gigantic Deinotheres represented in this drawing are based on the dimensions of the type lower jaw and five grinding teeth 
described in the year 1899 by Gregoriti Stefinescu of the University of Bucharest, discovered in the Manzati Valley, central Rumania. 
As stated on pages 95 to 98, the skeletal material described and figured is not from the type locality; the tibia and pes are figured on 
page 97 (Fig. 62). The restoration scene is in the MAnzati Valley during the Middle Pliocene period when these animals reached their 

Warm midsummer season. 

The limb proportions of the skeleton are certainly mastodontoid rather than elephantoid; they are based on proportions of the 
Deinotherium bavaricum of Franzensbad (Fig. 63, p. 99). The three different feeding poses and attitudes represent a theoretic interpreta- 
tion of the adaptations of the lower tusks as well as of the proboscis in these remarkable animals. The tusks were used along river banks for 
uprooting function; also in combat between the males and in warding off enemies, while the proboscis was used in the conventional probo- 

scidean way in reaching for and drawing down foliage. 


Deinotherium hungaricum Fhik, 1930 
Figure 74 

Type: Kotyh4za (Dep. Négrid), Hungary. Lower Miocene (?), Burdi- 
galian age. Paratype: Kirdld (Dep. Borsod), Hungary. 

Prodinotherium hungaricum Bhik, 1930. ‘‘Prodinotherium 
hungaricum N.G., N.Sp.”’ with an Appendix by T. Szalai “On the 
Geological Occurrence of Prodinotherium hungaricum Bhik,”’ 
Geol. Hungarica, Ser. Palaeont., Fasc. 6, pp. 3-21. TYPE. 
Lower mandible containing |.Ps—Ms, r.Ps, Mz, Ms; also fragment 
of scapula and limb fragments (humerus, ulna, radius, carpals, and 
portions of metacarpals). Locatiry AND Horizon.— 
Kotyhiza (Dep. Nograd), Hungary. Embedded in a blue clay 

belonging to the Aquitanian (?) period. PARATYPE.— 
Molar and tusk fragments, from Kirald (Dep. Borsod), 
Hungary. Typ Ficgurn.—Op. cit., Pl. 1, figs. 1-3, Pls. 1, 

lt, and rv. ParatyPr Ficgures.—Op. cit., Pl. 1, figs. 4-7. 
Speciric Cuaracters.—(Cf. Bhik, p. 14) Third premolar. 
. “in the case of D. Cuviert and D. Hobleyi the hypoconid and 

the entoconid [of Pm;] are connected by a posterior cingulum, while 

with the specimen of Kirdld the entoconid of pm3 lies inside of the 

cingulum and is thus in no connection with it. This peculiar forma- 
tion of the pm3 distinguishes the Hungarian specimen from all 
Dinotheria hitherto described, so as from the larger species too.”’ 
Mental foramen. . “in the case of the Hungarian animal it 
{posterior mental foramen] is to be found in the vertical separating 
the pm3 from the pm4. The place of the anterior mental foramen 
is unknown with D. Hobleyi; in the case of D. Cuvieri it falls below 
the middle of pm3; in the Hungarian specimen it is situated below 
the anterior edge of the pm3. In consideration of all these cir- 
cumstances I give anew name viz: D. hungaricwm to the specimen 
found at Kotyhaza and Kirald. . . . From the systematic point 
of view the mesatipody is a much more primitive character, than 
the systematically highly progressed dolichopody. This very 
interesting and important difference in the structure of the foot 
induces me to class the D. hungaricum into a new genus, under the 
genetic name of Prodinotherium.” 

“The Prodinotherium hungaricum must be by all means one 
of the eldest Dinotheria hitherto known from an exactly determined 
geological age.”’ 

GroLocy.—(Szalai, op. cit., p. 19): “On both find places of 
Prodinotherium hungaricum Bhik, namely at Kotyh4za, in the 


neighborhood of Salgétarjan, and at Kirald not very far from the 
former locality, the bed layers of the coal-measures are generally 
formed by a tough clay. Its material is sometimes of a swelling 
kind, sometimes of a sandy one, and its colour is bluish gray if wet, 
and greenish gray when dry. Embedded vegetal remains are to 
be found rather abundantly in this clay, among which Calamus 

. and Cinnamomum ... are the most frequent, whilst the 
Vertebrate fauna is represented by Mastodon angustidens Cuv., 
. . . Aceratherium tetradactylum Lart., . . . Prodinotherium hun- 
garicum Bhik, ... Testudo Fejérvdryi n. sp... . and some 
Trionyx . . . remains.” 

Osborn, 1932: The presence of ‘Mastodon’ angustidens in- 
dicates that these coal measures are of Burdigalian (Lower Mio- 
cene) age rather than of Aquitanian (Upper Oligocene) age; in 
fact, Szalai remarks (op. cit., p. 21) “The Prodinotheriwm remains 
described by Ehik are especially important from the stratigraphical 
point of view: for they are the only Dinotherian fossils which un- 
doubtedly belong to the transition beds connecting the Upper 
Oligocene with the Lower Miocene, precisely this set of strata 
representing the Aquitanian. The biohistorical importance of 
such statement lies, moreover, in the fact that no Dinotherian 
remains are known, up to now, from strata older than those be- 
longing to the Aquitanian.”’ 



Fig. 74. 

U, figs. 1 and 2, in oblique (upper) and right lateral (lower) aspects. 
one-eighth natural size. 

Type mandible of Prodinotherium hungaricum Bhik, 1930, Pl. 


In the opinion of the present author, the characters of the 
paratype premolar of Kirald (PI. 1, fig. 7) serve to distinguish a 
new specific rather than a new generic stage. Consequently we 
are inclined to consider Prodinotherium as a synonym of Deino- 
therium until further evidence is adduced to the contrary. 

Deinotherium hopwoodi sp. nov. 
Figure 68a 

From Olduvai, near the southeast shore of Lake Victoria, Tanganyika 
Territory, Africa. Middle Pleistocene. 

The most surprising discovery of recent years is the survival 
of Deinotherium into the Middle Pleistocene of central Africa. 
The beautifully preserved type molars (Fig. 68a, p. 104) were 
regarded by Hopwood (letter, Feb. 5, 1934) as referable to the Lower 
Pliocene D. giganteum stage, from which they are quite distinct. 
For reasons given below they are made the type of Deinotherium 

These grinders (Fig. 68a), an 1.M; (Brit. Mus. M. 14119, cast 
Amer. Mus. 27005), ap. 97 mm., tr. 87 mm., and an |.M® (Brit. 
Mus. M. 14118, cast Amer. Mus. 27006), ap. 91 mm., tr. 96 mm., 
compare in size with those of the Lower Pliocene Deinotherium 
giganteum Kaup (1.Ms;, ap. 95 mm., tr. 83 mm.; 1.M’, ap. 91 mm., 
tr. 93 mm.), while inferior to those of the Middle Pliocene D. 
gigantissimum (1.M3, ap. 114 mm., tr.?; 1.M°, ap. 114 mm., tr. 
117 mm.). 

In the same pit with the above type grinders were found 
three other molars,’ namely, third inferior premolar, |.P3, ap. 
81 mm., tr. 63 mm., index 78, fourth superior premolar, r.P‘, 
ap. 89 mm., tr. 90 mm., index 101, and finally a second su- 
perior molar, |.M?, ap. 101 mm., tr. 100 mm., index 99. 

The teeth found (Fig. 56+) in the Lower Pleistocene of 
Abyssinia (Haug, 1911; Joleaud, 1928), referred to D. gigan- 
teum, probably belong to this species. 

The type locality (Fig. 56, 25), Olduvai, Tanganyika 
Territory, Lat. 2° 58’ S, Long. 3° 24’ E, is a most inaccessible 
semi-desert place, a day’s march from the nearest good water 
supply. Hopwood (letter, Feb. 5, 1934) reports that the type 
teeth were found in the lowest of the four horizons referable 
to the Middle Pleistocene. They were mixed up with a partial 
skeleton of a very young Elephas antiquus recki, and on the 
other side of the dry water-course were found other remains 
of the same elephant at the same level; also that teeth belong- 
ing to Deinotherium have been found in the Lower Pleistocene 
of Abyssinia (Haug, 1911, Joleaud, 1928), as well as in the 
Pleistocene of Kanam, Kenya Colony. 

ConeLets.— Beautifully preserved and fitting closely in 
apposition, as if belonging to a single individual, these type 
grinders display the conelets of which the crests were formed, 
namely: protolophid of 1.Ms (16 conelets), metalophid of 
1.Ms (14e conelets); protoloph of 1.M® (14 conelets), metaloph 
of 1.M8 (12 conelets). On this character, which to Osborn’s 
knowledge has not been mentioned before, also on the superior 
size, the species Deinotheriwm hopwoodi has been based by 
Osborn, in recognition of Doctor Hopwood’s never-failing 
coéperation with the author in the preparation of this Memoir, 
in addition to his distinguished work in the African field. 

'See also Leakey’s discovery of Deinotherium sp.?, of Pleistocene age, in central Africa, Kavirondo Gulf (pp. 85 and 105 of the present Memoir). 
°The originals of these referred molars are in the British Museum, namely, Brit. Mus. M.14116, M.14115, and M.14117, respectively. 





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SERBELODON BURNHAM!  2050Mm™m, 6’834’e 







1. Cuvier’s five classic ‘species’ of Mastodon represent four 5. The Zygolophodontine (=Mastodon borsoni): Final 

[five] subfamilies. separation (Russia, 1894) from the Mastodontine 
2. Cuvier’s original conception of his genus ‘Mastodonte,’ (= Mastodon americanus). 

1806. Types and definitions of five species. 6. Separation of species of ‘Mastodon’ and ‘Elephas’ 
3. The Zygolophodontine: Separation by Schinz and Vacek described in America (1792-1874). 

from the Longirostrine. 7. Osborn’s classification (1927) of the superfamily Masto- 
4. The Brevirostrinw: Separation by Lortet and Chantre dontoidea. 

from the Longirostrine and Zygolophodontine. 8. Cuvier and the Revolutions of the Globe. 

(January, 1934) Many discoveries since 1927 have altered or modified the statements made in this chapter. During the seven years 
(1927-1934) four additional subfamilies were discovered or defined, namely, the Amebelodontine, the Platybelodontinz, the Tetralo- 
phodontinz, and the Humboldtinz subfam. nov.; also the families Serridentidze fam. nov. and Humboldtidz fam. nov. (1935) 
Three new subfamilies are added, namely, Paleomastodontinz, Gnathabelodontinz, and Stegolophodontinz. The final classification, 
therefore, will be found in the Appendix at the close of the present Volume. 

Cuvier, 1806.—Cuvier, the founder of vertebrate palzontology, fell heir to all the collections in the Muséum 
d’Histoire Naturelle of Paris and was the first to distinguish specifically the true mastodont form of molar without 
trefoils from the bunomastodont form of molar with trefoils. Cuvier was more impressed with the resemblances 
between these two forms of molars than with the differences; consequently he included both forms within his 
single genus ‘Mastodonte’ (= Mastodon), which he divided into five classic species that became the starting point 
of all future descriptions and which survive to the present day; these species also are subdivisible according to 
modern discovery, as below; he did not adopt the binomial system of Linnzeus until 1817. 

OsBorn, 1927.—We now know that these five Cuvierian ‘species’ belong in two’ distinct families, namely, the 
Mastodontide and the Bunomastodontide, and in four' distinct subfamilies, namely, the Mastodontine (including 
Mastodon giganteum Cuv.), the Zygolophodontine (including M. tapiroides Cuv.), the Longirostrine (including 
M. angustidens Cuv.), and the Notorostrine (including M. andiwm Cuv. and M. humboldtii? Cuv.). 

It has taken more than a century of research, concluding with the present Memoir, to discover that five entirely 
different lines of descent actually separated these five Cuvierian ‘species’ in geologic time and divided them up 
among five generic and four’ subfamily phyla, as follows: 

Mastodon giganteum Cuv....Mastodon americanus. ...Subfam. Mastodontine. ...Fam. Mastodontide. . . .Superfam. Mastodontoidea 

Mastodon angustidens Cuv..Trilophodon angustidens... “  Longirostrine.... “ Bunomastodontide  “ a 
Mastodon tapiroides Cuv.....Turicius tapiroides.......  “  Zygolophodontine “ Mastodontide.....  “ 5 
Mastodon andium Cuv...... Cordillerion andium...... “  Notorostrine...... “ Bunomastodontide  “ % 
Mastodon humboldtii Cuv...Cuvieronius humbolatii?...  “‘ Notorostrine ...... “ Bunomastodontide  “ ‘ 

\(Osborn, 1934) These Cuvierian ‘species’ now belong in three families and five subfamilies through the addition of the Humboldtide fam. nov. and the 
Humboldtine subfam. nov. 

2(Osborn, 1934) now referred to the Humboldtine subfam. noy., of the Humboldtid® fam. nov. 


It was characteristic of Cuvier that he was very reluctant to admit the existence of more than one genus 
of either mastodonts or elephants, namely, Mastodon and’ Elephas; or to admit the existence of more kinds of 
fossil proboscideans than he himself had seen or described, This reluctant spirit has been manifested by many 

of his followers down to the present time. 


See Chapter VI for detailed descriptions 

Cuvier was not in the habit of clearly designating his type specimens; consequently we select for the types 
of the Mastodon the teeth from the Ohio River first discovered, described, and figured as the types of Masto- 
donte de l’ Ohio [= Mastodon americanus Kerr—Fig. 76]. 


Fig. 76. Cuvier’s types of Mastodonte de l’Ohio or Le Grand Mastodonte, 1806.2, 
Pl. 49 [1]. [=Mastodon americanus Kerr of the present Memoir.] See figure 112. 

See Chapter VII for detailed descriptions 

Petit Mastodonte (= Mastodon |Turicius| tapiroides Cuvier, 1806—see Figs. 77, 160, and 161), from the 

Fig. 77. Cuvier’s 
type of Petit masto- 
donte, 1806, from 
Montabusard =T'urici- 
us tapiroides. 

Lower Miocene of the Caleaire de Montabusard, with lophoid crests, without trefoils, has 
become the prototype of the zygolophodont phylum of the Miocene, including M. turicensis 
Schinz, M. affinis Jourdan, M. borsoni Hays, as shown in the conspectus of Mayet (1909, 
p. 45, B). This phylum is treated by Schlesinger (1917, pp. 146-162) under the heading 
“Mastodon (Zygolophodon) tapiroides Cuvier.’’ Mastodon tapiroides Cuv. is treated by 
Osborn in the present Memoir as Turicius tapiroides. The reasons for the separation by 
Osborn in 1926 (Osborn, 1926.706) of the genus Turicius from the genus Zygolophodon are 
given in Chapter VII, pp. 198-202, of the present Memoir. See also Pls. 1 and ut, between 
pp. 134-135. 

'(Osborn, 1934) Five subfamilies (see footnote on page 119). 


See Chapter VIII for detailed description (p. 252) and new figure (Fig. 299) of Cuvier’s type 

Cuvier’s type (Figs. 78, 190, 299) from Simorre (Mastodonte a dents étroites = Mastodon {Trilophodon| 
angustidens Cuv.), with narrow molars and single trefoils, appears in the modern 
literature (e. g., Schlesinger, 1917) as a collective species of all the Miocene Lon- 
girostrines of Europe; it includes, as synonyms or trinomials, Mastodon minutus 
Cuv., 1824, of Saxony, M. simorrense Lartet, 1851, M. gaujaci Lartet, 1851, and 
M. cuviert Pomel, 1854 [1848], as revised and listed by Mayet (Mam. Fos., 
1909, p. 44, A). This phylum is treated most fully by Schlesinger (Denk. Natur- 
hist. Hofmus., I, 1917, pp. 5-63), under the heading ‘Mastodon (Bunolophodon) 
angustidens Cuv.” A new figure of the type has been prepared (see Chap. VIII, 
Fig. 299) from a cast presented to the American Museum in 1927 by the British 
Museum. This cast was one of two sent to Gideon Mantell by Cuvier, both 
of which were acquired by the British Museum with the purchase of the Mantell Fig. 78. Cuvier’s type of Mastodonte a 
Collection about the middle of the last century. dents: Grontes, 1806 [= Triaphoton areas 

dens of the present Memoir]. See new figure 
from cast (Fig. 299) in Chapter VIII. 


See Chapter XII for detailed descriptions 

Regarding Cuvier’s South American types: (1) It is important to note that the type (Fig. 79) of Mastodonte 
des Cordiliéres [=Cordillerion andium of the present Memoir], distinguished by single trefoils, is from near a 

Fig. 80. Cuvier’s type 


Fig. 79. Cuvier’s type of Mastodonte des of Mastodonte humboldien, 
Cordilitres, 1806 |=Cordillerion andium of the 1806 [= Cuvieronius humboldtii 
present Memoir]. of the present Memoir). 

voleano in the vicinity of Quito, Ecuador; (2) the type (Fig. 80) of M. humboldien, distinguished by double 
trefoils (now referred to a distinct genus, Cuvieronius, in honor of Cuvier), isfrom near Concepcion, Chile. Further 
characterization of these South American species is given in the phylogenetic section below (Chap. XII). 


(January, 1935) Written by Osborn in the year 1927, Sections 4-7 are now somewhat out of date (see Appendix at close of the present Volume). 

Cuvier conceived his genus Mastodonte as embracing animals possessing the five different types of grinding 
teeth shown in his five type figures above; this is a ‘collective’ genus in the same sense that most of the original 
genera of Linnzus were collective, that is, embracing several ‘species’ and ‘genera’ in the modern usage of these 

According to all authors, Cuvier’s first references to his genus Mastodonte are in the Annales du Muséum 
d’Histoire Naturelle of 1806 (1806.2, p. 270). In the same year, in a subsequent paper, he observed (1806.3, p. 
413): ‘“‘Ainsi le genre se trouvera composé de cing espéces, toutes également inconnues aujourd’hui sur la terre.” 

His original descriptions, references, and figures of these five species are as follows (Cuvier, 1806.2, pp. 270, 
293; 1806.3, p. 412): 

“Je nommerai donc la grande espéce [op. 
cit., pp. 412, 413], 
Mastodonte del’ Ohio; [Mastodon giganteum Cuv.= Mastodon Ann. Mus., VIII, p. 293, Pl. 49 [1], Figs. 
americanus] 1-5; also Pls. 50 [11] to 56 [vu], fide 
de Blainville. (Type confirmed by de 
Blainville in his revision of 1839-64, 

p. 245.) 
Celle de Simorre et d’ailleurs, [Mastodon angustidens Cuv.=Trilopho- Ann. Mus., VIII, p. 405, Pl. 66 [1], 
Mastodonte a dents étrovtes; don angustidens| Fig. 4; Pl. 67 [i], Figs. 2, 4, 6, 11, 

and 13 (fide de Blainville). (Type 
confirmed by de Blainville in his 
revision of 1839-64, p. 246.) 

Celle des petites dents, [Mastodon tapiroides Cuv.=Turicius Ann. Mus., VIII, p. 411, Pl. 68 [1], 
Petit mastodonte;: tapiroides| Fig. 6. (Type confirmed by de 
[de Montabusard] Blainville in his revision of 1839-64, 
p. 251.) 
[de Saxe] [Mastodon minutus aut. minor=(?) Tri- Ann. Mus., VIII, p. 411, Pl. 67 [1], 
lophodon angustidens minutus] Fig. 11. (Type confirmed by de 
Blainville in his revision of 1839-64, 
p. 250.) 
La grande 4 dents carrées, [Mastodon andium Cuv., 1824,? Ann. Mus., VIII, pp. 411, 413, Pl. 67 
Mastodonte des Cordiliéres = Mastodon cordillerarum Desm., 1820- {], Fig. 1. (Type confirmed by de 
{prés du volean d’Imbaburra, 1822, =Cordillerion andium| Blainville in his revision of 1839-64, 
Quito, Ecuador] p. 249.) 
Et la plus petite, [Mastodon humboldtii Cuvier in Des- Ann. Mus., VIII, pp. 412, 413, Pl. 67 
Mastodonte humboldien marest, 1818, p. 447 =Cuwvieronius {1], Fig. 5. (Type confirmed by de 
[de la Conception du Chili]’’ humboldtii| Blainville in his revision of 1839-64, 
p. 249.) 
Le Grand Mastodonte Cuy. (Cuvier, 1806.2, Ann. Mus., VIII, pp. 270, 293) or Mastodonte de l’Ohio Cuv. 
(Cuvier, 1806.3, p. 412) = Mastodon giganteum Cuv., 1817 (“Le Régne Animal,” p. 233). Typr.—Cuvier, 
1806.2, Pl. 49 [1], figs. 1-5; confirmed by de Blainville (1839-1864, p. 245). Type DescripTion.—Cuvier, 

1806.2, pp. 270, 273, 293; 1806.3, p. 412. 
= Mastodon americanus of the present Memoir. 

“Mastodonte & dents étroites”’ Cuy., 1806 (Ann. Mus., VIII, 1806, pp. 405, 412) = Mastodon angustidens Cuv., 

1817 (‘Le Régne Animal,”’ p. 233). Type.—Cuvier, 1806.3, Pl. 66 [1], fig. 4; confirmed by de Blainville 
'In 1806 Cuvier regarded the little teeth of Montabusard (=M. tapiroide Cuv., 1821) and of Saxony (=M. minutus Cuv., 1824) as belonging to the 
same species, namely, Pelit mastodonte. See Cuvier, 1806.3, p. 411: ‘La dent de Montabusard, pl. m1, fig. 6, correspond si bien & celle de Saxe pour sa 
largeur, que je ne doute pas que ce ne soit un germe de l'une des postérieures de la méme espéce, cassé en avant.” 
*The specific names M. andium Cuv. and M. cordillerarum Desm. refer to the same type description (Cuvier, 1806.3) of the Mastodonte des Cor- 
dilitres. We may follow de Blainville and Falconer in choosing the specific name andium. 


(“Ostéographie des Mammiféres,” 1839-1864, p. 246). Type Descriprion.—(Cuvier, 1806.3, p. 405): “Je 
commence par une dent de Simorre, pl. 1, fig. 4.” (Detailed description and measurements of this tooth and com- 
parison with other teeth from Simorre also of M. giganteum, pp. 401 to 420). 

= Trilophodon angustidens of the present Memoir. 

Petit mastodonte, celle des petites dents, Cuvier, 1806.3, p. 413 = Mastodonte tapiroide Cuv., 1821 (Cuvier, 
1821-1824, Vol. I, p. 268), subsequently written Mastodon tapiroides (see Desmarest, 1820-1822, p. 386, also 

Cuvier, 1821-1824, Vol. V, Pt. 2, p. 527). Typre.—Cuvier, 1806.3, Pl. 68 [11], fig. 6; confirmed by de Blain- 
ville (1839-1864, p. 251). Tyre Locauity.—Caleaire de Montabusard. (See below Petit mastodonte 
= Mastodon minutus.) TypE Description.—(Cuvier, 1806.3, p. 411): ‘La dent de Montabusard, pl. 1, 

fig. 6, correspond si bien 4 celle de Saxe pour sa largeur, que je ne doute pas que ce ne soit un germe de |’une 
des postérieures de la méme espéce, cassé en avant.” 
(de Montabusard, L. Miocene) = Turicius tapiroides of the present Memoir. 

Petit mastodonte, celle des petites dents, Cuvier, 1806.3, p. 413 = Mastodon minutus Cuv., 1824 (Cuvier, 1821- 
1824, Vol. V, Pt. 2, p. 527). TypEe.—Cuvier, 1806.3, Pl. 67 [uu], fig. 11; confirmed by de Blainville (1839- 
1864, p. 250). Type Locauiry.—Saxony. (See above Petit mastodonte= Mastodon tapiroides.) . TYPE 
Description.—(Cuvier, 1806.3, p. 411): ‘‘Telle est la dent de Saxe, envoyée autrefois par le professeur de Got- 
tingue, Hugo, A Bernard de Jussieu, et que l’illustre neveu de celui-ci a bien voulu me communiquer, PI. 11 [11, 67], 
fig. 11, entiérement semblable en figure et en proportions 4 celle de la fig. 4, pl. 1. Elle est exactement d’un tiers 

(de Saxe, Miocene) = Trilophodon angustidens minutus of the present Memoir. 

Mastodonte des Cordiliéres, la grande 4 dents carrées, Cuvier, 1806.3, p. 413= Mastodon Andium Cuv., 

1824 (Cuvier, 1821-1824, Vol. V, Pt. 2, p. 527), and M. cordillerarum Desmarest, 1820-1822, p. 385. TyYPE.— 
Cuvier, 1806.3, Pl. 67 [1], fig. 1; confirmed by de Blainville (1839-1864, p. 249). Type Locauiry.— 
Voleano near Quito, Ecuador. Type Description.—(Cuvier, 1806.3, p. 411): “Les plus grandes ont les 

mémes dimensions que leurs correspondantes de l’Ohio. M. de Humbold en arapporté une qu'il a trouvée prés du 
volean d’Imbaburra, au royaume de Quito, 4 1200 toises de hauteur. Elle est assez décomposée et encore enduite 
de cendres voleaniques. Son émail est teint en roussdtre; elle est longue de 0,12, et large de 0,085. Voyez pl. 
11 [67], fig. 1.”” (De Blainville, 1839-1864, p. 249): ‘‘Cette espéce a été proposée, pour la premiére fois, par M. G. 
Cuvier, en 1806, dans son mémoire sur différentes dents de Mastodontes. Ann. du Mus., tome VIII, p. 411, et 
dans les mémoires réunis tome I, p. 11 [tome II, 1812.1, p. 11], et enfin dans la seconde édition de ses Recherches 
sur les Ossements fossiles de Quadrupédes (tome I, p. 266, 1821 4 1825). Elle ne reposait et ne repose encore que 
sur un petit nombre de piéces, trois dents molaires. (Loc. cit., pl. 2, fig. 1 et fig. 12.)” 

= Cordillerion andium of the present Memoir. 

Mastodonte humboldien, la plus petite, Cuvier, 1806.3, p. 413 = Mastodon Humboldtii Cuv. (see Desmarest, 

1818, p. 447); M. humboldii Cuvier, 1821-1824, Vol. V, Pt. 2, p. 527). Type.—Cuvier, 1806.3, Pl. 
67 [11], fig. 5; confirmed by de Blainville (1839-1864, p. 249). Type Locauiry.—Near Concepcion, 
Chile. Type Description.—(Cuvier, 1806.3, p. 412): “M. de Humbold est encore celui qui les a découvertes. 

Je lui en dois une qu’il a rapportée de la Conception du Chili; elle est fort usée, mais bien conservée, teinte 
en noir, longue de 0,08, et large de 0,06. Voyez pl. 1 [67], fig. 5.” 
=Cuvieronius humboldtii of the present Memoir. 


(1) One of the earliest revisions of the mastodonts, in the broad sense, following Cuvier, is that of Desmarest 
(1820-1822) in which all of Cuvier’s five species are reviewed and the name Mastodon cordillerarum applied to 
Cuvier’s Mastodonte des Cordiliéres. In this revision six species of ‘Mastodon’ are recognized, namely, 
Mastodon giganteum, M. angustidens, M. cordillerarum Desm. (=M. andium Cuv.), M. Humboldtii, M. minus 
(=M. angustidens), and M. tapiroides (pp. 384-386). M. minus (= minutus) is a synonym of M. angustidens. 

(2) The next revision is that of Schinz (1824, p. 278), in which five of the same species are recognized and 
the zygolophodont species ‘Mastodon turicense’ from the lignites of Elgg, Canton Zurich, is added. 

(3) There followed successive revisions in which it was recognized that M. tapiroides Cuv. and M. turicensis 
Schinz, animals possessing lophodont grinders without trefoils (=Zygolophodontine), should be separated off 
from animals possessing bunomastodont grinders with trefoils (= Bunomastodontide), namely, M. angustidens 
and M. longirostris. 

(4) Finally, in 1879, Lortet and Chantre distinguished two groups of species, separating in group II (= Zygo- 
lophodontinze Osborn), ‘“‘Mastodontes 4 dents présentant des collines larges et tapiroides,” the following (p. 304): 

II Lorret AND CHANTRE (1879) 
Mastodon Borsoni Hays, 1834 
Mastodon Turicensis Meyer, 1839 
Mastodon Tapiroides de Blainville, 1839 
Mastodon Buffonis Pomel 
Mastodon Vellavus Aymard, 1846 
Mastodon Vialetti Aymard, 1846 

(PRESENT Menmorr, 1927) 
= Zygolophodon borsont 
= Turicius turicensis 
= Turicius tapiroides 
=Zygolophodon borsoni buffonis 
= Zygolophodon borsoni vellavus 
= Zygolophodon borsoni vialetiz 

(5) Practically the same zygolophodont phylum is adopted by Mayet (1909, p. 45) who, with Depéret, 
enjoys the advantage of direct examination and comparison of the type and referred specimens included within 
Mayet’s Section B “Mastodontes 4 Mamelons Disposes en Crétes Transversales (Type Lophodonte).” These 
species, namely, Mastodon borsoni, M. turicensis, M. tapiroides, [and M. pyrenaicus|, are those chosen by the 
Austrian paleontologist M. Vacek in 1877 as typical of the genus Zygolophodon. 

BuNOLOPHODONT AND ZyGoLOPHODONT MiocENE AND PuLioceNe Specips oF Europe AFrrer Mayet, 1909 

Mastodon angustidens Cuvier. 

1825, Mastodon angustidens, Cuvier, Ossements fossiles, t. 1, p. 250. 

1825. Mastodon minutus. Cuvier, Id., p. 267, pl, Il, fig. 11. 

1851. Mastodon Simorrense. Lartet, Notice sur la colline de Sansan, 
p- 24. 

1851. Mastodon Gaujaci. Lartet, Id., p. 24. 

1854. Mastodon Cuvieri. Pomel, Calalogue méthodique, p. 76. 

1859. Mastodon angustidens. Lartet, Note sur la dentition des pro- 
boscidiens vivants et fossiles (Bulletin de la Société géolo- 
on de France, 1859, p. 469, pl. XVI, fig. 1-4; pl. XX, 

ig. 6). 
1859. Mastodon angustidens. P. Gervais, Zoologie et Paléontologie 
_frangaises, 2° édit. 

1870. Mastodon arvernensis. Fraas, Die Fauna von Steinheim, Stutt- 
gart, 1870, 

1887. Mastodon angustidens. Depéret, Vertébrés miocénes de Ja vallée 
a (Archives du Muséum d'histoire naturelle de Lyon, 
t. {), 

1898. Mastodon angustidens. Mayet, Mammniféres miocénes des sables 
de lOrléanais et des faluns de la Touraine (Annales 
de U'Universilé de Lyon, fasc. 24); == des sables de l'Orléa- 
nais, p. 140, pl. VII, fig. 3, 4, 5,6; =e des faluns du Blésois, 
Pp. 297, pl. XI, fig. 2 et 3), 

Species, including types and referred specimens, related to 
the Mastodon angustidens phylum. After Mayet, 1909, p. 44. 
[=Longirostrinw and Brevirostrine of the present Memoir] 



Mastodon turicensis Schinz. 

1825. Mastodon tapirojdes. Cuvier, Ossements fossiles, t. 1, p. 267, 
pl. Ill, fig. 6. Cf. aussi Guettard, Mémoires, t. V1, 10° m., 
pl. VII, fig. 4. 

1827. Mastodon turicensis. Schinz, Naturg. und Abbild. d. Sauge- 
thiere, 1827, p. 243. 

1842, Mastodon turicensis. H. von Meyer, Palzontologica zu* Geschi- 
chte der Erde und ihrer Geschopfe, Francfort. 

1850, Mastodon tapirojdes. Blainville, Ostéographic, G. Eléphant. 

1859. Mastodon Borsonii. G. Gervais (p. part.), Zoologie et Paléonto- 
logie .frangaises, 2° éd., p. 68. 

1878. Mastodon affinis. Jourdan, Archives du Muséum de Lyon, t. II. 

1878. Mastodon tapiroides. Lortet et Chantre, Recherches sur les 
Mastodontes (Archives du Muséum de Lyon, t. II, p. 285. 
pl. VII et IX), 

1887. Mastodon turicensis. Depéret, Vertébrés miocénes de la vallée 
du Rhéne (Archives du Muséum de Lyon, t. IV, p. 131). 

1908. Mastodon turicensis. L. Mayet, Mammiféres miocénes des sables 
de l’Orléanais et des faluns de la Touraine (Annales de 
CUniversité de Lyon, fasc. 24); == des sables de l'Orléanais, 
p. 194, pl. VIII, fig. 1 et 2; == des faluns du Blésois 
p. 298, pl. XI, fig. 4 et 5. 

Type and referred species related to the Mastodon turi- 
censis phylum. After Mayet, 1909, p. 45. 

{=Zygolophodontinz of the present Memoir] 


(6) In Vacek’s invaluable Memoir of 1877, “Uber Osterreichische Mastodonten,”’ we not only find a clear 
separation of the Zygolophodontine from the Longirostrine (T'rilophodon, Tetralophodon, etc.) , but a clear separa- 
tion of the Brevirostrine (Anancus, etc.) from the Longirostrine. Vacek’s classic memoir is fully treated in 
Chapter VII (the Zygolophodontine). 


Following Cuvier, in a long period of exploration, discovery, and research, representatives were found of a 
new and important phylum, unknown to Cuvier, namely, the Brevirostrine, including ‘Mastodon’ arvernensis of 
Auvergne and related forms, while many additional members of the phyla known to Cuvier were discovered and 
traced into genera by Falconer. 

The state of knowledge and opinion in 1878 is reflected in the splendid memoir ‘‘Recherches sur Les Masto- 

dontes”’ of Lortet and Chantre, 1879, pp. 285-311, Pls. 1-xvi (bis). In this invaluable memoir the mastodonts 
are grouped as follows: 

Masrtopontes A Dents Errorres Masropontes A DENTS PRESENTANT DES COLLINES 
[p. 297] [p. 304] 
Mastodon dissimilis....... Jourdan, 1840. Mastodon Borsoni........ Hays, 1834. 
2 Arvernensis....Croizet et Jobert, 1828. 3 Turicensis......H. v. Meyer, 1839. 
- Blainville, 1839. 4 Blainville, 1839....(pro parte) 
: angustidens....Cuvier, 1836........ (pro parte). Borsoni........ de Blainville, 1839. Blainville, 1839. .(pro parte). 7. Buffonis........Pomel. 
* angustidens....Laurillard, 1846.....(pro parte). ws Vellavus........Aymard, 1846. 
“ brevirostris..... P. Gervais, 1859. Z: Vialettiv sn 2-2 Aymard, [1846]. 
Anancus macroplus.......Aymard, 1846. S Borsont..< 2.4. Vacek, 1877. 
Borsoni........ Lartet, 1859. 

{=Brevirostrine and Longirostrine of the present Memoir] 
| =Zygolophodontine of the present Memoir] 

[p. 303] 
Mastodon longirostris..... Kaup, 1835. 

4: angustidens. Blainville, 1839... .(pro parte). 
angustidens....P. Gervais, 1859...... (pro parte). 
longirostris..... Owen, 1861. 
longirostris..... H. v. Meyer, 1867. 
longirostris..... Falconer, 1868. 
longirostris..... Vacek, 1877. 

{=Longirostrine and Tetralophodontine of the present Memoir] 

It will be observed that while Lortet and Chantre (1879) follow Cuvier in uniting the brevirostrine, longi- 
rostrine, and zygolophodontine mastodonts into the single genus ‘Mastodon,’ they practically separate the species 
into three groups which correspond respectively with the subfamilies Brevirostrine, Longirostrine, and Zygolo- 
phodontinz of the present Memoir. 

Consequently the grouping of species by Lortet and Chantre (1879) is nearer the truth than that of Mayet 
(1909), because Mayet includes under his Group A, “‘Mastodontes 4 Mamelons Arrondis (Type Bunodonte),” 
species belonging in both the Longirostrine and Brevirostrine phyla, while he rightly includes in his Group B, 


‘““Mastodontes 4 Mamelons Disposes en Crétes Transversales (Type Lophodonte),” the entire Zygolophodontine 

In the Miocene of Europe no representatives of the Brevirostrinz have been found, but this phylum is appar- 
ently represented in the Pliocene of India, e.g., Anancus perimensis. The author has not yet had an opportunity 
of personally examining the classic collections of the Proboscidea of Miocene Europe, which must be very carefully 
reéxamined before their true phyletic, generic, and specific characters can be determined. We have positive evi- 
dence of the existence of four entirely separate and distinct proboscidean lines of descent, and we see some evidence 
of the existence of a fifth line in Miocene Europe, although the fifth phylum has not been positively demonstrated 
as yet. These theoretical lines' are as follows: 

descended from perhaps a branch of descended from perhaps a branch of a branch of 
Miomastodon Zygolophodon Phiomia Phiomia unknown ancestors 
Pliomastodon Turicius Trilophodon Serridentinus (?) [| Stegolophodon|] 

These four to five subfamilies are represented by species of Miocene and Pliocene age as follows: 

1. Mastodontinsze by species of Miomastodon and Pliomastodon, e.g., Miomastodon tapiroides americanus and 
Pliomastodon americanus praetypica. 

2. Zygolophodontine by the species T'uricius tapiroides, T. turicensis, and Zygolophodon borsoni. 

Longirostrine by the species T'rilophodon angustidens; all longirostral types. 

4. Serridentinz, medilongirostral Serridentines with medium length of jaw, specifically defined by Serri- 
dentinus filholi in Europe; represented in Asia by S. mongoliensis; in America by S. productus, 
S. serridens, and other species. 

5. Stegodontine (i.e., ancestors of) possibly represented by a species, Zygolophodon |= Stegolophodon|] sub- 

latidens, recently described by Schlesinger from eastern Europe. 


It appears, from Marie Pavlow’s revision (1894) cited below, that all 
Russian students of the Proboscidea worked under the spell of Cuvier and 
conservatively adhered to his generic name Mastodon even though they 
clearly recognized the wide distinctions which exist between the genotype 
species Mastodon americanus and many other species which he included within 
this genus. As detailed above, the last phylum to be split off from the 
Mastodontine of the true Mastodon americanus type is that embracing a large 

number of European and Asiatic species which may be grouped under the Big 6. gh ootacgeternt toe 
P . La Se ; ” : ohioticus’ by Pavlow, from Pestchana, 
name Zygolophodontine. Beginning in 1824 with the discovery of Mastodon Podolia, Russia. (Pavlow, 1894, PI. 1): 
. . . . . dh i 2 i i i 9 
hinicendis and coumoun of oF s F Fig. 4, une m~ [Mbp] inférieure droite... 
| ul ag in 1834 with the recognition of M. borsoni, this Univ de Bosco. |= me inadnneen 
important and entirely distinct phylum, the Zygolophodontine, was grad- That this is a true species of the true 
2 aor aaite ‘ eer, A 5 genus Mastodon appears froin the two 
ually recognized, until in 1877 Vacek proposed the generic name Zygolophodon. principal ‘mamelons’ composing each crest. 

(Osborn, 1934). These theoretical lines of generic descent are considerably modified in the Appendix of the present Volume. 

AMERICA (1792-1874) 

The American palzontologists up to the year 1878 were no less conservative than their European confréres 
in retaining the two generic names Mastodon and Elephas for all kinds and formsof proboscideans. The new generic 
names Euelephas, Tetralophodon, and Trilophodon of Falconer were reluctantly admitted only by certain authors. 
Between 1884 and 1922, however, the tendency to split the mastodonts into a number of genera did not assert 
itself generally until Osborn began the present revision, as shown in the following chronological list of names as 
originally used and as revised in the present Memoir. 

1792 LHlephas americanus Kerr...................------d Mastodon americanus MASTODONTIN 
isa Mastodon borsons Hays: .... 22... 2c ccc ce ee nteee Zygolophodon borsoni ZYGOLOPHODONTIN-£ 
1834 Mastodon chapmani Hays..........................Stegomastodon chapmani {[HcuMBoLDTIN£ subfam. nov.] 
1838 Elephas jacksoni Mather......... ................Parelephas jacksoni MAMMONTIN 
1842 Elephas americanus DeKay.. weeeeeeees....--Mammonteus primigenius americanus MaMMONTIN 
1857-1868 Elephas columbi Fiuieoaer: .........Parelephas columbi MAMMONTIN 
1858 Mastodon (Tetralophodon) miri nels tie ee Stegomastodon mirificus (HuMBOLDTIN® subfam. nov.] 
1858 Elephas imperator Leidy.. . .Archidiskodon imperator MAMMONTIN-E 
1859-1861 Elephas texrianus Owen n 1859, ead Blake, 1861. ..Parelephas columbi MAMMONTIN& 
1868 Mastodon americanus Leidy....................-.4 Mastodon americanus MasToDONTIN © 
1869 Mastodon obscurus Leidy..............-.------.- Trilophodon obscurus LONGIROSTRIN-E 
aaah | Mastodon shepardt Leidy...: ... 202.22 2d35. 2 0300282 Rhynchotherium shepardi RHYNCHOROSTRIN © 
1873 Mastodon proavus Cope...........................Serridentinus proavus SERRIDENTIN 
iis Mastodon productus Cope..........<..---is++-++--- Serridentinus productus SERRIDENTINE® 

A full list of the generic and specific synonyms of Mastodon americanus will be found in Chapter VI (Sub- 
family Mastodontine), under the heading of this species. 


From the foregoing sections it appears that the collective genus ‘Mastodonte’ of Cuvier, 1806, has become, 
through discovery, analysis, and research, equivalent to the superfamily Mastodontoidea. This superfamily, 
as it comprises all the mastodonts in all parts of the world, is far more comprehensive than Girard imagined 
in his Mastodontide, because it includes two families, Mastodontide and Bunomastodontide, and not less than 
seven subfamilies and seventeen genera, the number of which is constantly being enlarged." 

Through discoveries by Schinz (1824), by Vacek (1877), by Lortet and Chantre (1879), by Pavlow (1894). 
by Mayet (1909), and by Osborn (1918-1925), the seven distinct subfamily phyla, Mastodontine, Zygolopho- 

dontine, Longirostrine, Serridentine, Rhynchorostrine, Notorostrine, and Brevirostrine, have been separated 

Thus the superfamily Mastodontoidea requires a definition broad enough to embrace the characters common 
to these seven subfamilies. 

'This, like all other classifications of the 1927 period, is now replaced by the 1935 classification set forth in the Appendix at the close of the present Volume. 
See also page 11 and footnotes on pages 27, 30, and 31 above. At the present time (July, 1935) the genera included within the Mastodontoidea number 





Families:! Mastodontidae Girard, 1852, Osborn, 1918; Bunomastodontidz Osborn, 1921 
Original reference: Mastodontoidea Osborn, Amer. Mus. Novitates, No. 1, p. 1 (Osborn, 1921.515); Masto- 
dontide Girard, Proc. Amer. Assoc. Adv. Sci., 1852, pp. 326, 328; Osborn. Bull. Geol. Soc. Amer., XXIX, 1918, 
Bunomastodontide Osborn, Amer. Mus. Novitates, No. 1, 1921, p. 2 (Osborn, 

p. 134 (Osborn, 1918.468). 



Derinition.—Chiefly forest and savanna living proboscideans; Sideirs migrating in the north and 
Browsers on tree and shrub leafage and on twigs; uprootersin part. Superior 
and inferior incisive tusks adapted to varied food habits, branching and uprooting, and to varied modes 

south temper ate regions. 

of defense; 
premolar S. 

retaining or losing enamel bands. 

Deciduous premolar series prevailing over permanent 
Intermediate molars, P4-M2, early becoming trilophodont; third molars, M’-M; progres- 

sive to 4-7 crests; cones, conules or conelets progressive to lophodont, bunolophodont, or ptychodont 



Nipple-toothed mastodonts 

Tusk enamel disappearing. 
Grinders sublophodont to 


Yoke-toothed mastodonts 

Incisive tusks oval, (?) re- 
taining enamel band. Grind- 
ers becoming truly crested, 
with numerous conelets. 


Body prevailingly elongate, broad, massive, short limbed. 



Long-jawed mastodonts 

Upper and lower tusks 
functional in feeding, superi- 
or tusks retaining enamel 
band, inferior tusks spatu- 
late, losing enamel band. 
Grinders with central con- 
ules or conelets in valleys. 


Serrate-toothed mastodonts 

Superior and inferior inci- 
sive tusks functional in feed- 
ing, retaining enamel band. 
Rostrum moderately elon- 
gate. Grinders without 
central conules or conelets, 
with external and internal 
serrated spurs. 


1. =Mastodon gigantewm Cuvier = Mastodon tapiroides Cuvier 3. =Mastodon angustidens Cuvier 4. Unknown to Cuvier — 


Beak-jawed mastodonts South American mastodonts Short-jawed mastodonts 

Superior and inferior tusks down- 
turned, functional in feeding, retain- 
ing enamel band. Grinders with inter- 
mediate conules or conelets. 

Superior tusks offensive weapons, 
losing enamel band; inferior incisive 
tusks disappearing. Grinders with 
alternating external and internal 
lobes, i.e., cones, lacking imterme- 
diate conules or conelets, developing 
double or quadruple trefoils. 

Jaws gradually abbreviating. Su- 
perior incisive tusks purely offensive 
(without enamel) or used in feeding 
(with enamel). Grinders with inter- 
mediate conules or conelets. 

Cuwieronius Anancus 
Rhynchotherium Eubelodon Pentalophodon, [Synconolophus] 
5. Unknown to Cuvier 6. =Mastodon andium and 7. Unknown to Cuvier 

M. humboldtii Cuvier 
The above synopsis shows that in 1927 Cuvier’s ‘Mastodon’ was known to embrace five classic species and 

seven subfamilies. To these are now (1935) added seven subfamilies, making fourteen subfamilies in all (see 
Appendix of the present Volume I), 

(Osborn, 1934) The suborder Mastodontoidea now includes the additional families Serridentide fam. noy. and Humboldtide fam. noy. 

*(Osborn, 1934) The subfamily relationships of [Palzomastodon] are now uncertain. (1935) See pages 143-149 below, also Paleomastodontine subfam. 
nov. in Appendix of the present Volume. 


This historic chapter may appropriately conclude with quotations from ‘‘Les Transformations du Monde 
animal,” a work published in the year 1907 by Charles Depéret, who next to Cuvier and Gaudry ranks among the 
leaders of the paleontology of France. Depéret ardently sets forth and defends Cuvier as the creator of com- 
parative anatomy and of paleontology. (Depéret, 1907.2, pp. 7, 10, 13, 14, 19, and 22): 

L’admiration générale suscitée dans le monde savant par la série de mémoires que G. Cuvier publia 4 partir de 1798 et qui 
furent réunis, en 1812, sous le titre de: Recherches sur les ossements fossiles, n’a pas diminué [sic] de nos jours, malgré un recul de plus 
d’un siécle. Tout naturaliste qui veut se familiariser avec l’organisation des animaux supérieurs vivants ou fossiles doit encore 
aujourd’hui commencer ses études par la lecture de cet ouvrage magistral, ot se trouvent exposées, avec une clartéet une précision 
lumineuses, les notions fondamentales des deux sciences sceurs: |’ Anatomie comparée et la Paléontologie des vertébrés. . . . 

Ainsi, Cuvier a non seulement démontré la présence dans les couches sédimentaires d’une série de faunes terrestres super- 
posées et distinctes, mais il a eu le premier, et trés nettement, l’idée du perfectionnement organique graduel de ces faunes depuis 
les plus anciennes jusqu’aux plus modernes. C’est 1A une notion fondamentale, dont on oublie trop souvent d’attribuer le 
mérite 4 Cuvier, dans les jugements d’une sévérité excessive et souvent injuste, que les transformistes ont porté et portent encore 
sur les idées cuviériennes en matiére de paléontologie philosophique. . . . 

L’illustre naturaliste expose ses idées, avec sa clarté habituelle, dans l’admirable Discours sur les révolutions du globe, qui 
forme l’introduction de son grand ouvrage sur les ossements fossiles. Pour lui, les extinctions de faunes ont été A la fois 
completes et brusques, provoquées par des événements géologiques violents ou révolutions du globe, d’un caractére de généralité 
assez grand, mais non cependant absolu. En faveur de cette hypothése, Cuvier invoque de nombreux faits, d’ordre géologique, 
qui, pris isolément, et en tenant compte des documents connus 4 cette époque, sont d’une rigoureuse exactitude, et dont les 
relations réciproques seules deviennent discutables ou méme inexactes. . . . 

Quant au procédé de ce renouvellement, le reproche a été bien souvent adressé 4 Cuvier d’avoir admis 4 son tour une autre 
hypothése tout aussi peu démontrable scientifiquement, celle des créations successives. Mais c’est encore lA une critique tout 
4 fait injustifiée. Nulle part, le mot de création ne se rencontre dans |’ceuvre de Cuvier et il suffit de lire avec attention le 
Discours sur les révolutions du globe pour voir que dans |’esprit de l’illustre savant, il s’agit seulement d’invasions de nouvelles 
formes animales venues brusquement de contrées lointaines inconnues. Ici, l’idée est assez fondamentale pour qu’il vaille la 
peine de citer: 

‘Au reste, lorsque je soutiens,’ dit Cuvier, ‘que les bancs pierreux contiennent les os de plusieurs genres et les couches 
meubles ceux de plusieurs espéces qui n’existent plus, je ne prétends pas qu’il ait fallu une création nouvelle pour produire les 
espéces existantes, je dis seulement qu’elles n’existaient pas dans les mémes lieux et qu’elles ont dd y venir d’ailleurs.’ . . . 

Le lecteur voudra bien excuser sans doute la longueur de cette citation, dont l’intérét est considérable; elle démontre 
jusqu’a l’évidence qu’il faut reporter 4 Cuvier tout |’honneur d’avoir posé, avec une netteté et une exactitude admirables, 
Vhypothése si importante et si féconde du renouvellement des faunes par voie de migrations. . . . 

Ainsi, deux points paraissent absolument certains pour d’Orbigny: d’une part, la création en bloc de faunes entiéres— 
opinion que Cuvier n’avait jamais formulée—de |’autre, la disparition brusque de chacune de ces faunes. Pour le premier de ces 
grands faits, le savant paléontologiste ne tente méme pas, on |’a vu plus haut, le moindre essai d’explication scientifique. . . . 
C’est, on le voit, la reproduction 4 peu prés intégrale des révolutions du globe de Cuvier, avec une généralisation encore plus 
grande. ... 

Si l’explication de ce grand fait par des créations successives ne saurait nous satisfaire au point de vue scientifique, nous 
aurons plus loin & en chercher une interprétation rationnelle par des phénoménes de migration de faunes ou de migration de 
milieux, analogues & celles que Cuvier avait déja si bien mis en évidence pour les animaux terrestres. 

we : > 

Pas ee = = 


Tur WARREN Mastropon As IT wAs First MounTep (1845-1846), sHoRTLY AFTER ITS DISCOVERY 

Fig. 82. The figure is reproduced from the American Journal of Agriculture and Science, Volume II, Number 2, conducted by E. Emmons, 
Albany, and A. J. Prime, Newburgh. In their article, “The Great American Mastodon,” Messrs. Emmons and Prime remark: ‘‘The skeleton has 
since been arranged and set up, and this has been done with great care and the strictest attention to the articulating surfaces of all the bones, which 
we believe has not been the case with others which have been put together.” 

THe WarrREN Mastodon 
As photographed in the 

interior of the Warren Mu- 

seum, 92 Chestnut Street, 


Fig. 83. For fifty-seven 
years, 1849-1906, the War- 
ren Mastodon, mounted as 
shown herewith, was ex- 
hibited in the Warren Mu- 
seum in Boston. In 1906 it 
was acquired, thanks to the 
generosity of the late J. Pier- 
pont Morgan, by the Ameri- 
can Museum. 

The skeleton, as here de- 
picted, is covered with heavy 
black varnish. The imitation 
tusks are made of papier- 
mAché and were so length- 
ened as to sweep the ground 
and curve outwards at the 
extremities. The chest and 
backbone were raised two 
feet above the top of the 
shoulder blade, or scapula, 
and as a result the natural 
height of the animal was 
increased from nine feet to 
twelve feet. Beneath the 
Warren Mastodon are tusks 
and grinding teeth of other 
specimens. Around the base 
of the walls are many verte; 
bre of the giant Zeuglodon, 
the archaic fossil whale of 
the southern United States. 

THE WarrREN Mastopon 
as remounted for The Ameri- 
can Museum of Natural 
History, under the direction 
of Curator Henry Fairfield 
Osborn, July, 1907, by chief 
preparator Adam Hermann 
and assistants, is shown in 
figure 124, page 178. 






1. History of the subfamily Mastodontine, 1705-1934. 8. Characters of Pliomastodon and its included species. 

2. Résumé of the subfamily characters. 9. Historical review of the discovery of the genus Mastodon 

3. Separation of mastodonts of the Mastodon americanus (1705-1934) and of the species Mastodon americanus 
and Zygolophodon borsoni phyla in Russia. (1792-1869). 

4. Résumé of discovery and description of the Mastodon- 10. Characters of Mastodon and its included species and 
tine. varieties. 

5. Progressive characters and generic distinctions of the 11. Skeletal characters of Warren and Whitfield mastodons. 
Mastodontine. 12. Age and progressive cranial and dental characters of 

6. Characters of Palzomastodon and its included species. Mastodon americanus, revealed in the American 

7. Characters of Miomastodon and its included species. Museum collections. 

In Chapter IT it is shown how the Mastodontoidea may be clearly distinguished from the Mceritherioidea, 
the Deinotherioidea, and the Elephantoidea. In Chapter V it is shown how Cuvier’s original genus ‘Mastodonte’ 
(=Mastodon) came to embrace two' distinct families, seven' distinct subfamily phyla, and seventeen’ distinct genera. 
In the present Chapter VI Cuvier’s genus Mastodon, based on the type Mastodon americanus, becomes the 
terminal member of the subfamily Mastodontine, otherwise known as the ‘true mastodonts,’ with a geologically 
prolonged lineage of its own certainly traceable into Asia and Europe and possibly back to the Lower Oligocene 
Palzomastodon of North Africa. Accordingly Palzomastodon is systematically treated in this chapter, and the 
reader is referred to Chapter II where Palzxomastodon is described in its North African environment of Oligocene 
time in competition with Meritherium and Phiomia. The discovery of a complete cranium and jaws may 
compel us to remove Palzxomastodon from the subfamily Mastodontinz’, in which it now rests on the evidence 
at present available. It is certainly not a direct ancestor of Mastodon. 

(October, 1934) The very important addition of the Lower Miocene Miomastodon depereti sp. nov., from the Sables de l’Orléanais 
of France, of the Mastodon pavlowi sp. nov., from Pestchana, Podolia, Russia, has recently been made to the subfamily Mastodon- 
tine, as described in the phylogenetic appendix of the present Volume; see fig. 230D, p. 284, of Chap. VIII, and Pl. I, p. 134; also of 
the Mastodon acutidens sp. nov. of the Pleistocene of North America (see Figs. 131 Al-A4, 135, and Pl. I, also Appendix). 


Original reference: Mastodontina Brandt, 1869, Mém. Acad. Imp. Sci. St. Petersb., (VII), XIV, No. 1, p. 35. Mastodontine 
Osborn, 1910, “The Age of Mammals,” p. 558 (Osborn, 1910.346). 

Although the first fossil remains of a member of this great subfamily were discovered on the banks of the 
Hudson River as early as 1705, also near the banks of the Ohio River (Big-Bone Lick, Kentucky) by the Indians 
who gave them in 1739 to a French officer named Longueil, the origin, migrations, and early geologic history 
of the Mastodontinz are still comparatively obscure. 

This we believe is because the Mastodontine have included forest-living animals from the very beginning of 
their history, for fossil remains of forest-living animals are always rare in Tertiary deposits. 

Only by such a forest-loving hypothesis can we explain the unparalleled abundance of the remains of 
Mastodon americanus in the ancient forest, bog, and swamp deposits of the eastern and middle United States, as 

‘See page 128. 
*See pages 143-149 below, also Palwomastodontine in Appendix. 



summarized in the nation-wide records collected by Dr. J. M. Clarke, Dr. O. P. Hay, and others. And only by this 
hypothesis also can we explain the abundance and remarkable preservation of the more or less complete skeletons 
of our American Mastodon, which we believe outnumber those of all other extinct proboscideans put together. 

Masropon.—In the northern portions of the American continent these remains become very rare, and as we 
pass westward into the plains region and northward into Eurasia not a single fossil record is found until we reach 
Podolia, Russia, from which locality Marie Pavlow in 1894 described a single tooth (Fig. 81) which she referred 
to ‘Mastodon ohioticus.’ This central Eurasiatic region also yields teeth of the species belonging to the related 
subfamily Zygolophodontinw. When the grinders of Mastodon americanus are placed side by side with those of 
Zygolophodon borsoni we observe that they are really profoundly different; we verify this observation as we 
proceed westward into Austria, Germany, and France and discover that the bilobed grinding tooth of the true 
Mastodon may be traced back into its simple ancestral forms, which are quite distinct from the quadrilobed 
grinding tooth of the true Zygolophodon. 

Mromastopon.—This new stage of ancestral Mastodon grinder, as it appears in the Miocene of Holarctica, is 
placed in the distinct genus Miomastodon Osborn, because with these simple grinders are associated upper incisive 
tusks with broad enamel band; no Lower Pleistocene Mastodon thus far discovered has the enamel band. Thus the 
Pleistocene Mastodon americanus certainly leads back into the Miocene Miomastodon of western Eurasia and 
of North America. An intermediate stage is the Pliocene Pliomastodon lacking the enamel band. 

PALZOMASTODON.—Somewhat doubtful at present is the theory advanced by Matsumoto, and more recently 
supported by Osborn, that the original ancestor’ of the Mastodontine is to be found in the true Palzomastodon 
(as distinguished from the longirostral Phiomia) which occurs with relative rarity in the Oligocene river sands of 
the Faytim of northern Africa. Fossil remains of Palzomastodon are relatively rare, the skull is little known, but 
so far as our present knowledge goes the Palxomastodon grinding teeth are not ancestral in type to those of the 
true American Mastodon. 


SUBFAMILY CHaracters.—Skull brachycephalic, brachyopic; cranium relatively broad and low. 
Mandibular rami rapidly abbreviating, with persistently rounded inferior tusks becoming variable. 
Superior tusks rounded, with enamel band in the Palzomastodon' and Miomastodon stages disappearing 
in the Upper Pliocene (Pliomastodon) and Pleistocene (true Mastodon) stages; superior tusks upturned 
and out-turned, adapted to browsing and uprooting purposes, powerful offensive and defensive weapons. 
Grinding teeth hexabunodont in Palxomastodon, primitively tetrabunodont in Miomastodon and 
Mastodon by the absence of the intermediate conules, gradually acquiring a bunolophodont form; 
intermediate grinders, Dp*-Dp,, M'-M,, M?-M,, becoming trilophodont; third superior grinders, M°, 
progressive to tetralophodont stage only; third inferior grinders progressive to tetralophodont + 
or four and a half ridge-crests only. 

Warning the reader of the present limitations of our knowledge, which chiefly arise from the very great rarity 
of fossil materials in the Eastern Hemisphere, we may give a theoretic summary of the ancestral history of the 

GEOGRAPHIC AND GroLoaic Rance.—Springing from unknown ancestors of the Eocene-Oligocene of North 
Africa, the true Mastodontin appear in the Lower Miocene Miomastodon of western Eurasia and soon find their 
way eastward into North America, appearing in the Middle Miocene species of Nevada, Miomastodon merriami, 

‘See footnotes on pages 36 and 131. 


and in the Lower Pliocene species of western Nebraska, Pliomastodon matthewi, meanwhile leaving behind 
in Austria-Hungary their relatives, Miomastodon tapiroides americanus and Pliomastodon americanus praetypica, 
which give rise to the rare true Mastodon of southern Russia, referred to ‘Mastodon ohioticus’ by Pavlow. Rarely 
found in northern and western North America, they multiply rapidly in the favorable forests of the middle and 
eastern United States in the typical form Mastodon americanus. 

SuBFAMILY HaBiraL ADAPTATION.—These broad, massive, low-bodied, low-headed, well-defended Mastodon- 
tine probably evolved chiefly in the north temperate forests of Eurasia and of North America avoiding the ex- 
tremes of warm climate in both hemispheres. The American Mastodon is known to have lived on the foliage of 
trees and shrubs aided by the vertical chopping motion of its jaws and the free interlocking action of its superior 
and inferior ridge-crests, the valleys not being filled or blocked with the trefoils, as in the valleys of the Longi- 
rostrine molar. That these true mastodonts were temperate and cold-loving animals, enjoying coniferous herbage, 
is shown by their avoidance of southern Eurasia, where no trace of the subfamily is found, also by their early 
disappearance in northern Africa. 


The reader is referred to Chapter V for the history of the separation of the Zygolophodonts in Europe as 
distinguished from their separation in Russia. See also Pls. 1, 11, and m1, between pp. 134-135 of the present 

We owe to the learned Russian paleontologist Marie Pavlow the full historic record, from the time of 
Buffon, of the occurrence of the true Mastodon (ohioticus) americanus in Russia and its clear distinction from the 
true Zygolophodon borsoni of Russia, a molar of which was presented to the great French naturalist Buffon in 1770 
and beautifully deesribed and figured by him in 1778 (Fig. 84A). 

Paviow, 1894, pp. 1, 2, 36.—Avant d’aborder la description de tous ces restes fossiles, il me semble nécessaire de faire une 
courte revue de la littérature sur ce qui est connu jusqu’a présent comme restes de Mastodon en Russie . . . [1] Buffon. en 
1775 ['], a été le premier 4 décrire [Footnote: ‘Buffon. Epoques de la Nature. 1775. PI. 1-11.’] deux dents provenant de la 
Russie, et rapportées plus tard par Lartet au Mastodon Borsoni. Une d’elles que le comte de Vergennes lui avait donné en 1770 
(Il. c. Pl. ret 1) a été indiquée comme trouvée dans la Petite-Tartarie [Footnote: ‘On désignait par ce nom a la fin du XVIII 
siécle toute la Nouvelle Russie d’aujourd’hui.’]._ Une autre dent, apportée par l’abbé Chappe de Sibérie (Pl. 111), a éveillé un 
grand doute sur sa provenance. 

[2] Buffon, sans indiquer le genre auquel ont appartenu ces dents, les croit identiques 4 celles trouvées prés de l’Ohio et ne 
doute pas qu’il y avait un animal inconnu jusqu’alors et commun aux deux continents. Ces échantillons types se trouvent 
maintenant dans la Grande Galerie du Museum A Paris. * 

[3] Presque en méme temps Pallas a décrit, en 1770-77 [Footnote: ‘Pallas. Acta Acad. Petropolitanae. 1777. Pl. 9.’] une 
dent trés usée et trouvée dans les sables ferrugineux, prés de Belaja, affluent de la Kama. Pallas rapprochait cette dent de 
celles de l’animal de l’Ohio et plus tard Blainville et Eichwald [Footnote: ‘Blainville. Ostéographie. Eichwald. Paléontologie 
de la Russie.’] l’ont rapportée au Mastodon tapiroides. Le dernier de ces savants indique que cette dent se trouve a |’Institut 
de Mines 4 St. Pétersbourg. 

[4] Il me semble que, autant qu’on peut en juger d’aprés le dessin de ce débris trés mal conservé, cette dent a du appartenir 
& un Mastodon du type ‘Zigolophodon’, c’est 4 dire possédant des crétes absolument nettes, dépourvues de mamelons inter- 
médiaires. Ce type est le mieux représenté par Mastodon Borsoni. 

[5] Soixante ans a peu prés se sont écoulés aprés cette description de Pallas, sans que personne mentionne des mastodontes 
trouvés en Russie, et ce n’est que depuis 1835, que des nouvelles trouvailles et des descriptions ont été faites. 

‘In the bibliography Pavlow gives the reference as follows: ‘Buffon. Supplément Al’histoire naturelle. Epoques de la nature. Tome V. Pl. 1-5, 1778.” 



[6] Ainsi Fischer de Waldheim en 1835, en déterminant d’aprés les dessins les fossiles trouvés prés de Rzazan, indique dans 
une courte notice [Footnote: ‘Fischer de Waldheim. Bull. Natur. Moscou p. 394, Pl. x.’] une jeune dent d’un Mastodon qu’il 
considére comme la premiére trouvaille de ce genre faite en Russie. Le dessin n’est pas assez bien fait, pour pouvoir déterminer 
l’espéce de ce Mastodon. L’échantillon est indiqué comme se trouvant dans la collection du Lycée de Riazan. ... 

(7. Op. cit., p. 36]: Age géologique et répartition géografique du groupe Zygolophodon. L’age géologique de ce groupe des 
Mastodon est bien prolongé. On rencontre leurs différents représentants depuis le mzocéne, durant le pliocéne, en Europe et le 
pleistocéne en Amérique. Les formes les plus anciennes ont été indiquées en Espagne (Mast. tapiroides) dans les lignites de 
Brihuega dans le miocéne moyen (ou inférieur—Lartet p. 4751. cit.). | Dans le miocéne moyen elles abondent: la France en a 
deux représentants: Mast. tapiroides Lartet et Mast. turicensis Schinz (af. Borsoni), dans les faluns de Touraine, graviers 
d’Orléanais, lignites de Soblay (Ain). 

[8] Mast. turicensis Vacek (af. Borsoni) a été trouvé en Silésie. Le miocéne en Suisse (Elgg) et la molasse de Winterthur 
sont trés riches en Mast. turicensis Schinz (af. Borsonz), ainsi que le mio-pliocéne d’Europe qui débute 4 Oeningen (Suisse). 

[9] Mast. Borsoni Lartet, et Borsoni Hays provient du Pliocéne inférieur d’Asti, d’Auray. Celui de Bravard, Lortet et 
Chantre du Pliocéne supérieur du Puy-de-Déme, Auvergne. Celui de Mr. Forsyth Major du Pliocéne supérieur du Val d’Arno. 

[10] Le Mast. virgatidens de v. Meyer n’est désigné que comme provenant des dépdts tertiaires de Foulda (Allemagne) 
sans que l’dge soit précisé. 

[11] Les différentes dents de M. Borsoni Vacek proviennent de divers dépéts; ainsi, Pl. v1, f. 3 indiquée comme provenant du 
miocéne supérieur de Neidorf éveille les doutes de l’auteur sur |’exactitude de cette indication, 4 cause de l’ancienneté des dépéts. 
Pourtant cela ne nous parait pas impossible, prenant en considération la trouvaille de M. twricensis Schinz dans ces dépéts et la 
parenté, presque l’identité de ces 2 formes. 

[12. Op. cit., p. 37] Quant au rapport génétique de ces formes il nous semble possible d’exprimer les suppositions suivantes: 
a) que Mast. tapiroides Lartet (non Schinz) de Simorre est l’espéce la plus ancienne dans ce groupe, et qui a donné naissance 
a b) Mast. turicensis et Borsoni de Touraine, de Soblay, de Zurich et d’ Asti, laquelle 4 son tour a précédé c) Mast. aff. Borsoni de 
|’Auvergne, de l’Allemagne, de |’Autriche et de la Russie; et qu’une branche, qui a dt se détacher de ce dernier a la fin du 
Miocéne (aff. ohioticus), a donné les formes de d) Mast. ohioticus de la Russie, développées dans le Pliocéne. 

Burron, 1778. Superiork Grinpinc TEETH or ZyGOLOPHODON (LEFT) AND OF MasTopoNn (RIGHT). ORIGINALS SAID TO BE IN THE 
Paris Museum (FIDE Paviow, 1894) 

Fig. 84. Grinding teeth of Zygolophodon borsoni referred (left) and of Mastodon americanus referred (right), after Buffon, 1778, 
Pls. 1 and ry, one-fourth natural size. [Inverted by H. F. O.] 

A, Buffon’s figure of the molar of an animal which we now know to be related to Mastodon [ = Zygolophodon| borsoni, found in Russia 
(“la petite Tartarie”’), presented to Buffon in 1770 by M. le Comte de Vergennes. Observe three to four lobes, i.e., cones or conelets, in 
each transverse crest. 

; B, Buffon’s figure of the molar of an animal now known as Mastodon americanus, from Big-Bone Lick, Kentucky, near the Ohio 
River, sent to Buffon by M. Collinson. Observe two lobes, i.e., cones, in each transverse crest, the true Mastodon type. 

Both of these beautifully engraved molar teeth appear to be third superior grinders, M°, although they lack the rudimentary fifth 

crest or pentaloph characteristic of the third superior molars of Mastodon americanus (F ig. 133). 




MastopontiIn&: Persistent LoNairupiNAL Sutcus, Two Cones AND Four Coneets tn Eacu Rince-crest, Rupimentary Ecro- (INFERIOR) 

Lower Oligocene A, Palzxomastodon beadnelli ref., r.M3. No suleus. Complete transverse crests and prominent central conules (C, C) prove that 
Palzomastodon is Nov ancestral to Mastodon. Amer. Mus. 13481, Faydm of Egypt (ef. Fig. 187c). 
Pleistocene L, Mastodon acutidens sp. nov., 1.M’. Longitudinal suleus, progressively sharpened ridge-crests. See figures 131 Al-A4 and 

135. Amer. Mus. 17727, Rochester, Indiana. 
K-J, Mastodon americanus ref., r. and 1.M?*, Mo.3. Sulcus persistent, 4 conelets, superior entotrefoils (KK), inferior ectotrefoils (J) 
(=losanges of Cuvier). See figures 133 and 134. Amer. Mus. 12464, Buffalo, Kansas, and Amer. Mus. 14294, near 
Fulton, Indiana. 
I, G, Mastodon pavlowi sp. nov., type, |.M**, 1.M3. Primitive. Longitudinal suleus, 4 conelets, no trefoils. After Pavlow, 1894, 
Pl. 1, figs. 2 and 3, Pestchana, Podolia, Russia. 
I’, ‘ Mastodon ohioticus’ ref. |= M. pavlowi ref.], r.Mz. Rounded conelets 4-5, entotrefoils. After Pavlow, 1894, Pl. 1, fig. 4 
(see Fig. 81 of present Memoir), Pestchana, Podolia, Russia. 
Pliocene C, D, E, H, Pliomastodon praetypica cotypes and ref. Primitive. Rounded conelets 4, persistent longitudinal! sulcus, rudimentary 
C, D, Cotypes r.M**, 1.Mo.s, after Schlesinger, 1922, Taf. xv, figs. 2 and 4, Szabadka and Batta-Prd, and 1.Ms, Taf. xrx, fig. 2, 
Batta-frd, Hungary. 
E, H, Referred 1.M! (rev.), after Vaeek, 1877, Taf. v1, fig. 3, Neudorf, Austria, and 1.M®, Taf. v1, figs. 1, la, near Theresiopel, 

Lower Miocene B, Miomastodon depereti sp. nov., 1.M*. [Erroncously determined as an 1.Ms, Fig. 230, p. 284, of the present Memoir.] The 
most primitive stage thus far known. Deep longitudinal sulcus, 3-4 eonelets, 4 ridge-crests and rudimentary penta- 

loph. After Mayet, 1908, Pl. vu, fig. 3, Chevilly, Sables de l’Orléanais, France. 


Molars of the true Mastodon phylum are readily distinguished: (1) By the persistence of the median sulcus even into the progressive stage (L) 
of Mastodon acutidens; (2) by the persistent four conelets: (3)by the rudimentary ectotrefoil and entotrefoil spurs. 

i il i ee ae — 






External CG 

| 4 External & 
(E, D), ConreLets 4 (B) Progressive To 6 (D, EB, F, G), No Ecro- or ENToTREFOILS. RIDGE-CRESTS 
Lower Oligocene A, Palzomastodon intermedius type and paratype, Mj.3, M' (see Fig. 92). Longitudinal suleus blocked by proto- and meta- 
conules. Central conules (C, C) in inferior molars. Nor ancestral to Mastodon or Zygolophodon. 

Middle Miocene — B, Zygolophodon pyrenaicus type, r.My (see Fig. 148). Primitive longitudinal sulcus, 4 conelets on three posterior crests. 

Middle Pliocene ©, Zygolophodon borsoni type, r.M® (see Fig. 154). 4-6 conelets on three anterior ridge-crests. 

Upper Pliocene D, Zygolophodon borsoni ref., r.Mo.s, 1.Mo.3 (see Fig. 150). Progressive 4-5-6 conelets on perfected inferior molar ridge-ecrests. 
Vestigial sulcus. 

Middle Pliocene E, Zygolophodon borsoni ref., 1.M?, ‘intermediate molars’ 4-5 conelets. After Schlesinger, 1922, Taf. xrv, fig. 5 (praetypica), 
Rikoskeresztur, Hungary. Ms, after Schlesinger, 1922, Taf. xv, fig. 5, Bics-Bodrog, Hungary. 

Upper Pliocene I, Zygolophodon borsoni ref., 1.M® (see Fig. 153). Primitive irregular longitudinal sulcus, 5—6 conelets. 

G, Zygolophodon borsoni ref., 1.M® (see Fig. 111). Primitive sulcus, 5 conelets on proto-, meta-, tritolophs, 4 conelets on tetarto- 

loph. After Buffon, 1778, Pl. 1. 

Palxomastodon intermedius (A) is distinguished from Zygolophodon as follows: (1) Median sulcus lacking; (2) central conules (C, C) present in 
anterior valleys; (3) entotrefoils sometimes present. 
Molars of the relatively rare representatives of the Zygolophodon phylum are readily distinguished by the following three outstanding characters: 
First, the median sulcus separating the four conelets in Zygolophodon pyrenaicus (B) is not very distinct; it is reduced or vestigial in Z. borsoni 
4 (D-G). 
Second, by the subdivision of the four transverse conelets of Zygolophodon pyrenaicus (B) into five conelets in Z. borsoni superior molars (D-G). 

Third, by the entire absence of the ecto- and entotrefoil spurs, which are more or less prominent on prolonged usage in specimens of Pliomastodon 
and Mastodon (Pl. 1, C-K). 

oe a 


ho as 

Ant. C 




ie dz! 


A J 


ZYGOLOPHODONTIN®, Turicius: LONGITUDINAL Sutcus (A), VESTIGIAL OR ABSENT (A—J), RipGE-crests Constant 4 (A-I), 5 (J), RupIMeNnTARY 
Ecro- aNp Enrorreroit Spurs (A, F, H), Conpiers 4-5-6 (A, B), Rapipty Muvrietyina, 7 (C)-10 (E)-11-13 (F)-10-25 (H-H3), 

Lower Miocene A, Turicius tapiroides type, r.M3. Primitive longitudinal suleus, 4-6 conelets. After Mayet, 1908, fig. 66, Calcaire de 
Montabusard, France. 

B, Turicius tapiroides ref.,1.M3. After Mayet, 1908, Pl. vi, fig. 2, Sables de l’Orléanais du Blésois, Pontlevoy, France. 

C, Turicius tapiroides ref.,r.My. Conelets 5-7, ridge-crests 4. After Mayet, 1908, Pl. x1, figs. 4and 5, Pontlevoy, France. 

Upper Miocene D, Turicius turicensis type, 1.M*. After Schinz, 1833, Taf. 1, fig. 1, Elgg, Canton Zurich, Switzerland. 
E, Turicius turicensis homceotype, r.My. Conelets 5-10. After von Meyer, 1867, Taf. m1, fig. 1, Elgg, Canton Zurich, 

F, Turicius turicensis ref., r.My. Conelets 5-13, trefoil rudiment. After von Meyer, 1867, Taf. 11, fig. 2. 
G, Turicius turicensis ref., r.Ms, r.M?. Conelets 4-7. After casts of originals in the Munich Museum (1899 IX 1>= 
Amer. Mus. 22492, from Freising, Upper Bavaria, and 1891 I 50=Amer. Mus. 22491, from Moosburg, Bavaria, 
Middle (?) Pliocene H-H3, Turicius virgatidens types, r.Ms (H), r.My (H 1), 1.M® (H 2, 3). Conelets 6-25. After von Meyer, 1867, Taf. rv, 
figs. 1-5, ‘Gelben Lehme’ bei Fulda, northeast of Frankfort, Germany. 
Lower Pliocene I, Turicius atticus type, 1.P*, M'. Conelets 4-5. After Wagner, 1857, Tab. vu, fig. 16, Pikermi, Greece. 
Pliocene J, Turicius wahlheimensis cotype, 1.M’. Conelets compressed, 4-9, 5 ridge-crests. After Klihn, 1922, p. 77, fig. 17, 
Esselborn, Rheinhessen, Germany. 

Molars of Turicius are readily distinguished from those of the Mastodon and Zygolophodon phyla by the following characters: 
First, by the accelerated sharpening and elevation of the transverse crests (A-F). 
Second, by the accelerated multiplication of the conelets from six (A) to seven (C) to thirteen (F) to twenty-five (H), thus producing a lofty, 
sharpened crown parallel with that of Stegodon, with minute conelets more numerous than in Stegolophodon or primitive species of Stegodon. 
Third, by the early reduction of the median sulcus, more apparent in Miocene species (A-F), vestigial in Pliocene species (H-H3), absent in 
specialized T'uricius wahlheimensis (J). 


Sc 0 
2. ws 





FPN ee, 


ZA \ B 


STEGOLOPHODONTIN £, STEGOLOPHODON: Primitive Lonerrupinat Sutcus, SHapep (A, B), Persistrxe ry I-IV Anrertorn Ripce-crests (E, El), 
tw L-III Anrertor Rince-crests (F), Vesticrat iv Ripce-crests I-III (G, H, 1). Coneters Rounpep, IncreasivG From 4-5 (A, D), 
5-7 (C, H, 1, G). Posrerion Rmce-crests IV-VI Procressive, with Congtets 5-7 (G), ConeLets not Exceepine 5 (H, I) 
Mio-Pliocene __B, Stegolophodon cautleyi progressus type, r.M'. Amer. Mus. 19446, near Chinji Bungalow, India. Summit of Lower Chinji 
horizon, 2,000 feet above base of Lower Siwaliks. 

C, Stegolophodon nathotensis type,r-Mz. Conelets5-6. Amer. Mus. 19455, near Nathot, Lower Middle Siwaliks, India. Lower 
Chinji horizon. 

Lower Pliocene D, Stegolophodon latidenstype,r.Mz. Ridge-crests I-VII, conelets 4, suleus on ridge-crests I-IV. After Clift, 1828, PL xxxvm, 
fig. 1, near Yenangyaung, Burma. Lowest levels of Irrawaddy Series (fluviatile). 

E, Stegolophodon latidens type, r.M*. Ridge-crests I-VI, sulcus on ridge-crests I-IV. E 1 (section), ridge-crests coalescent at 
base (I-III). After cast (Amer. Mus. 21978) of Clift’s type, 1828, Pl. xxxvn, fig. 1, near Yenangyaung, Burma, lowest 
levels of Irrawaddy Series (fluviatile). 

F, Stegolophodon latidens ref., r.M'. Sulcus on ridge-crests I-III, conelets 4-6, ridge-crests 4%. After Matsumoto, 1926.1, PI. 
y, figs. 1 and 3 (Prostegodon), Shiwogama, Miyagi District, Province of Rikuzen, Japan. 

Pliocene (?) G, Stegolophodon lydekkeri sp. nov.,1.M®. Ridge-crests I-VI, conelets 4-7, sulcus on ridge-crests I and II only. After Lydekker, 
1886.2, fig. 19 (as M. latidens), Borneo. 
Middle(?) Pliocene A, Stegolophodon sublatidens type, r.M°®. Conelets 4-5. After Schlesinger, 1917, Taf. xvu, fig. 2, Teschen (Schlesien), Austria. 
Middle Pliocene H, Stegolophodon cautleyi type, r.M®. Ridge-crests I-V, sulcus on ridge-crests I and II, conelets 5. After Lydekker, 1886.1, 
p- xv, fig. 6, Perim Island, India. 
Upper(?) Pliocene I, Stegolophodon stegodontoides Pilgrim, type, r.M*. Ridge-crests I-VI, sulcus on ridge-crests I-III, conelets 5. After Lydek- 
ker, 1880, Pl. xxxrx, Lehri, Punjab, India, possibly Upper Siwaliks. 
Molars of Stegolophodon are readily distinguished from those of the Mastodon, Zygolophodon, and Turicius phyla by the following characters: 
First, by the persistence of the median sulcus separating the inner and outer pairs of cones of all the crests (A, B, D), of the three to four an- 
terior crests (E, F, I), of the two anterior crests (G, H). 
Second, by the rounded, bunoid conelets separated by median suleus (A—H). 
Third, by the closure of the enamel in the base of the transverse valleys, as seen in section (El), very characteristic of Stegodon. 
The second and third of the Stegolophodon characters enumerated below link this genus with the genus Stegodon. But we must remember that 
Stegolophodon cautleyi is of Middle Pliocene age (Perim Island), contemporary with the true Middle Pliocene Stegodon bombifrons (Dhok Pathan). 
See Volume II, Chapter XV, Stegodontoidea. 



TruE MASTODON AMERICANUS Type IN EurasiA.—It is interesting to note that in addition to the lophodont 
and zygolophodont forms related to Mastodon borsoni and M. turicensis, described by Pavlow and beautifully 
figured by her (1894), there also occurs in several localities, e.g., Pestchana and Krasnoie, Podolia, Russia, 
the molar type which she rightly compares closely with that of M. ohioticus, i.e., M. americanus (see Fig. 81 of 
the present Memoir). Of these Russian specimens Marie Pavlow (1901, p. 12) speaks as follows: 

Je veux profiter de l’ocecasion, puisque je parle des mastodontes, pour compléter la description des restes fossiles du 
Mastodon Ohioticus, trouvé 4 Pestchana en 1892, et décrit par moi en 1894 (1. cit.). En parlant de la partie antérieure de la 
mandibule de cet animal, je me suis bornée a dire (p. 10), qu’on ne trouve sur ce morceau de mandibule aucune trace d’alvéole 
pour la défense. Cela me semblait suffisant pour faire croire, que cet animal adulte n’avait pas de défenses inférieures et qu’il 
pourrait étre rapporté par ce caractére au Mast. Ohioticus, les autres caractéres de dents venant appuyer cette détermination. 
Mais ayant rencontré dans la littérature quelques objections 4 ce sujet, je me crois obligée de donner une description plus 
détaillée de cette partie de mandibule, en l’accompagnant d’un dessin (PI. 1, fig. 3). 

This description, with photographic figures (1894, Pl. 1, figs. 2 and 3), convinces us that we have from Pest- 
chana a form closely resembling Mastodon americanus but which may not be specifically identical’; the Pestchana 
specimens are certainly closer to M. americanus than to Zygolophodon borsont. 

The fact that this recent review by Pavlow of the Eurasiatic Proboscidea yields examples of only four 
individual specimens of Mastodon americanus from two localities, Pestchana and Krasnoie, as compared with the 
relative abundance of Zygolophodon borsoni, is another proof of the great scarcity of members of the true Mastodon 
americanus phylum up to the period of its abundant revelation in the eastern forests of North America. As shown 
in Pl. 1, pp. 134-135, other specimens referred by Vacek to Zygolophodon prove to belong to Mastodon. 


The earliest discoveries of the American Mastodon on the Hudson River in 1705, and those in Kentucky near 
the Ohio River in 1739, antedated by thirty years the dissemination of the various editions of the “Systema 
Nature” (1735-1766) of Linneus. From 1752 to 1778 Buffon described the ‘‘os d’éléphans,”’ as cited in full below, 
of this Kentucky animal without assigning to them a specific name; subsequently Kerr’s description of Elephas 
americanus appeared (1792). Nevertheless these fossils attracted little attention, in fact, even as late as 1797 
Blumenbach affixed the clumsy name Ohio-Incognitum to one of the fossil teeth found on the Ohio River, and as 
late as 1806 Cuvier named the same animal Le Grand Mastodonte and Mastodonte de l’Ohio. 

Consequently the history of proper nomenclature lags far behind the history of discovery, and the very last 
stage in the long progress of paleontology is the ancestral history or phylogeny. 

Tyre Locauity, Bie-Bonr Lick, Ky.—The type locality of the first discoveries of Mastodon in America is 
certainly the Big-bone-swamp (fide Kerr, 1792, p. 116), subsequently known as the ‘Big-Bone Lick,’ a few miles 
southeast of the Ohio River, in Kentucky. Thus the animal took its second name Ohio-Incognitum Blumenbach, 
1797, from the Ohio River, and became known as Mammut ohioticum Blumenbach in 1799, Mastodonte de l’Ohio 
Cuvier in 1806. 

Chronologie notes respecting early explorers of Big-Bone Lick were published by William Cooper in 1831 
{Monthly American Journal of Geology and Natural Science, Vol. I, 1831, p. 159]. This list contains the state- 
ment that a French officer named Longueil secured fossil bones from a morass near the Ohio River in 1739, and 
that they were collected by some Indians and not by a Frenchman. The list goes on to say that Col. George 
Croghan visited Big-Bone Lick in 1765. 

(1935) Made the type of Mastodon pavlowi (see Appendix). 


Cuvier, 1806.2, p. 280, ry PART, AND 1834,Vol. II, p. 266.—Le plus célébre de ces dépdts, celui qu’ont visité Longueil, Croghan 
et tant d’autres, celui qui a fait donner au mastodonte le nom d’animal de |’Ohio, porte lui-méme celui de Big-Bone-Strick ou 
Great-Bone-Lick. Il est dans l’état de Kentucky, 4 la gauche et au sud-est de |’Ohio, 4 quatre milles du fleuve, trente-six 
milles au-dessus de l’embouchure de la riviére de Kentucky [Footnote: ‘Volney, Tableau du climat et du sol des Etats-Unis 
d’ Amérique, I, page 100.’], presque vis-A-vis celle de la riviére dite la Grande Miamis. C’est un lieu enfoncé entre des collines, 
occupé par un marais qu’entretient un filet d’eau salée, et dont le fond est d’une vase noire et puante. Les os se trouvent dans 
la vase et dans les bords du marais, au plus A quatre pieds de profondeur, suivant le rapport que nous en a fait feu le général 
Collaud, qui avait été sur les lieux, et qui, en fouillant pendant trois jours seulement, avait recueilli vingt-quatre morceaux. 
Leur abondance y est étonnante. DéjA Croghan croyait y avoir vu des restes de plus de trente individus; mais on en a recueilli 
depuis un bien plus grand nombre. 

Fig. 85. Big-Bone Lick, Boone County, Kentucky, 
from map in Anthony Finley’s ‘A new American atlas,”’ 


{5 weimdionZAY ¥) etc., folio, Philadelphia, 1826, No. 10 (fide P. Lee 


Phillips, “A List of Maps of America in the Library of 
Congress,” 1901, p. 838). 

This map is of great historic interest as showing one 

of the birthplaces of vertebrate paleontology in America. 

f Visited by the French officer Longueil, mentioned and 

Jaa ous >) yA / < also fully described by Buffon, Blumenbach, and 

/ \ especially Cuvier, founder of vertebrate paleontology. 

The Mastodon in the New World took the part played 

by the Mammoth in the Old World, in fact, it is re- 

markable that Hlephas americanus Kerr (1792) antici- 
pates Elephas primigenius Blumenbach (1799). 

Bia-Bone Lick, 

), \ an ? 
rf 2 e 7 A 
tw” ES So AR 

: 8 \ 

In the following generic list only the chief specific names applied to the American Mastodon are included; a 
full list of this polynomial species is given below under the heading “Mastodon americanus” (p. 165). 

See Figure 86 

1705 New York First mention of fossil remains near Albany, New York. 
1714 First published account of two teeth and a thigh bone found 
thirty miles south of Albany, New York, on the Hudson 
1739 Big-Bone Lick, First fossil remains found by Indians in the Big-Bone Lick, 
Ohio River Ky., near the Ohio River, and given to a French officer 

by the name of Longueil; subsequently mentioned by 
Buffon, Blumenbach, and Cuvier. 

1778 = io A single molar tooth from the Ohio River figured by Buffon 

(Fig. 84 B). 
1. 1792 tf S JHMATHOS CRAIGS INSU G5 No pa heouen ob ae aaseapoadansoos = Mastodon americanus 
2. 1797 ue Ee Onveaincogminmeblimenbach na neeeneeie ene eee eee oS 4 
2. 1799 os if Mammunonioicun Blumenbacht,) sees... 0se eee eee tf 2) 
3. 1803 if of BIZ DhOSMmocrocennalus7 As Ampere. easiness nee eetes i o 
4. 1806 e “ NGronduVvlastodontei Cuvier acencccnscr ore ee - 
4. 1806 f 3 Mastodanterde UOnto\@uvier-twen- sete Lee en eee o i 
6. 1808(?) “ « Harpagmotherium canadense Fischer de Waldheim (cf. Sherborn, 

1924, p. 1022, “Harpagonotherium canadense, Anim. foss. 

SSIDETION AE cue dussse ys Sida uet Rica m amen aa sn eee = “ a 

ba. 1814 No. America Mastodon Macrodon Rafinesque = HY as 





6. 1814 Ohio River Mastotherium megalodon Fischer de Waldheim............... = Mastodon americanus 

4. 1817 Ly Mastodon giganteum Cuvier. . = rs as 

4. 1824 aS = Mastodon maximus Cuvier . : = di a 

7. 1854 Canada Elephas Rupertianus Richardson, “Swan River, Lake Winni- 

Pegivadin CANARY ci ererd eta eee Rels yale sea chess bus vane = Mastodon americanus rupertianus 
[1835 Russia Mastodon podolicum HWichwald.........................-.. =Deinotherium podolicum (see Chap. IV, 
p. 85)] 
4, 1868 Ohio River Mastodon americanus Kerr [Leidy]. First use of Kerr’s 
specific name in America.. . = Mastodon americanus 

8. 1890 §. Carolina Mastodon rugosidens Leidy, Beaufort C ounty.. = Mastodon americanus rugosidens 

9. 1894 Russia Mastodon ohioticus ref., Pavlow, Podolia.. ; Ae = Mastodon sp. (?). See No. 26 below 
10. 1901 Faytim Palxomastodon Beadnelli Andrews, Egypt.. eceeeeeaesse.. =Palxomastodon beadnelli' 
11. 1905 o Palzomastodon parvus Andrews, Egypt...................=Paleomastodon parvus' 
12. 1914 Iowa Mammut progenium Hay, Harrison County................= Mastodon progenius 
13. 1919 Hungary Mastodon (Mammut) americanus Penn. forma praetypica 

PCHLeSINGer ane meme ae a Nhe costes wages aem Speen = Pliomastodon americanus praety pica 
14. 1921 Nebraska Mastodon matthewt Osborn, Sioux County... . = Pliomastodon matthewt 
15. 1921 Nevada Mastodon merriami Osborn, Thousand Creek, ‘Humboldt 
COM noes rraaae thrash cuise erely nce mn aie sigs, and Gena = Miomastodon merriami 

16. 1921 Hungary Mastodon tapiroides americanus (Schlesinger in Osborn)... = Miomastodon tapiroides americanus 
17. 1922 Faytim Palxomastodon intermedius Matsumoto, Egypt............=Palzomastodon intermedius' 
18. 1926 Illinois Mastodon americanus plicatus Osborn, Walnut.............. = Mastodon americanus plicatus 
19. 1926 Oregon Mammut oregonense Hay, Baker County...................=Mastodon oregonensis 
20. 1930 Florida Pliomastodon sellardsi Simpson, Brewster................ = Pliomastodon sellardsi 
21. 1930 California Pliomastodon verillarius Matthew, Fresno iene! teh Adna's = Pliomastodon verillarius 
22. 1931 Nebraska Mastodon moodiei Barbour, Milford. . = Mastodon moodiei 
23. 1933 New Mexico Mastodon raki Frick, Hot Springs... = Mastodon raki 
24. 1933 Alaska Mastodon americanus alaskensis Frick, 1 near Fairbanks. . = Mastodon americanus alaskensis 
24a. 1934 Nebraska Mastodon grangeri Barbour, near Pender, Thurston Co. .. = Mastodon grangeri 
25. 1935 France Miomastodon depereti sp. nov.,? @hevillyanivan. - ee Groots = Miomastodon depereti sp. nov. 
26. 1935 Russia Mastodon pavlowi sp. nov.,? Pestchana...............-... = Mastodon pavlow? sp. nov. 
27. 1935 Indiana Mastodon acutidens sp. nov.,? Rochester... ........ ....=Mastodon acutidens sp. nov. 

The white dots within the 
The white crosses represent referred specimens. Nos. 24a and 27 
See also geographic distribution map forming the inside cover, or 

Fig. 86. Geographic distribution (according to the numbers in the list above) of the principal species of the Mastodontinae. 

black areas represent the approximate localities where the types were discovered. 
omitted on this map. Nos. 25 and 26 are omitted here but appear on map (Fig. 123a). 
front end-paper, of the present Volume. 

The above list of genera and species shows that there are many gaps in this great Mastodontine phylum to 
be filled by future discovery; also that while the Pleistocene Mastodon stage is well known, the Pliocene and Mio- 
cene stages of Pliomastodon and Miomastodon are known only from a few grinding teeth and tusks. The Oligo- 
cene stage, Palxomastodon, is still very imperfectly known.’ 

'See pages 143-149 below, also Palwomastodontine in Appendix at close of the present Volume. 
*See Appendix at close of the present Volume for description of Miomastodon depereti sp. nov., of Mastodon pavlowi sp. nov., and of Mastodon acutidens sp. nov. 


GENERIC AND SpEciFIc Succession.—As explained in the preceding section, the inclusion of the Lower 
Oligocene genus Palzxomastodon' as a form related to the phylum Mastodontine is provisional and depends 
upon our further knowledge of this relatively rare animal. While we are certain that the true Palzxomastodon 
possesses a longer jaw but a broader and shorter skull than its Lower Oligocene contemporary Phiomia, while the 
superior grinding teeth are comparatively short and broad, and while in the crowns of the superior and inferior 
grinding teeth the proto- and metaconules forbid the ancestral relationship of Palzomastodon to Pliomastodon, 
Miomastodon, and Mastodon, we must await further knowledge of the cranium and of the cutting teeth, also of the 
incisors of Palzomastodon, before we can form a positive opinion on this very important and interesting question. 

Meanwhile we may contrast the characters of these four genera so far as they are known: 

or Nortu AMERICA 

Europe AND oF NortH 


Palzomastodon Andrews, 1901 Miomastodon, Pliomastodon Osborn, Mastodon Cuvier, 1817 

1922, 1926 
Palate and grinding teeth relatively Palate and grinding teeth somewhat Palate and grinding teeth relatively 
short and broad. elongated. elongated. 
Dental formula: I$=3=$ Dp} Ps=4 Dental formula: I $=3=3 Dp} P} Dental formula: I $=3=¢ Dp $+ 
Mitzi. Mo=0=s- P9=9=4 (vestigial) M +3. 
Ridge formula: P2¢ P32 P43 Ridge formula: Probably P 43 M 13 Ridge formula: Dp 22 Dp 32 Dp 48 

M 1222 M 2227 M 3224. 

Grinding teeth brachyodont, buno- 
lophodont, with proto- and meta- 
conules (see Fig. 93). No median 

Grinding teeth with more elevated 
lophs. A median suleus; no conules. 

M13 M23 M3 4+ 

Grinding teeth subhypsodont. A 
median sulcus; no conules. 

Rudimentary third crests in ‘inter- 
mediate molars,’ P 2-M 1. 

Lower jaw and symphysis greatly 

Incisive tusks oval in section, with 
broad enamel band. 

Palzomastodon parvus. 
Palzomastodon intermedius. 
Palzomastodon beadnelli. 

Fully formed third erests in ‘inter- 
mediate molars.’ 

Lower jaw and symphysis mod- 
erately elongated. 

Incisive tusks rounded, with enamel 
band (Miomastodon); without enamel 
band (Pliomastodon). 

Pliomastodon matthew. 

Pliomastodon americanus praetypica. 

Miomastodon tapiroides americanus. 

Miomastodon merriami. 

Third crests fully formed in ‘inter- 
mediate molars.’ 

Lower jaw and symphysis  ab- 

Incisive tusks rounded, without 
enamel band. 

Mastodon americanus. 

Mastodon americanus rugosidens. 
Mastodon americanus plicatus. 
Mastodon progenius. 

Dentau Succession.—The relations of the deciduous premolars, Dp*(?), Dp;(?), of Palzomastodon in dis- 
tinction from the permanent premolars, P*, P,(?), have not been observed. Nor do we know the relations of the 
deciduous premolars to the permanent premolars in Miomastodon or in Pliomastodon. 

According to the observations of Hays (1834) and of Warren (1852, 1855) cited below, the true Pleistocene 
Mastodon americanus agrees with Pleistocene and recent species of the Elephantide in the suppression of the 
permanent premolar teeth; the fourth true superior and inferior premolars, P*-P;, form in the jaw but they do not 
erupt. In Elephas the corresponding permanent teeth, P*-P,, are suppressed entirely. 

‘See pages 143-149 below, also Palwzomastodontine in Appendix. 



The succession and relationship of members of the true Mastodontine phylum depend upon our very close 
examination and comparison of the structure of the grinding teeth, as displayed in the accompanying comparative 
figures. As first observed by Lartet, the grinding teeth of the true Mastodon americanus are fundamentally differ- 
ent from those of Mastodon borsoni which Vacek has rightly referred to Zygolophodon. 

Mep1iAn Sutcus.—In the carefully drawn Plates 1, 1, 11, and tv, illustrating the evolution of the grinding 
teeth in Mastodon, Zygolophodon, Turicius, and Stegolophodon, a median longitudinal sulcus separates the exter- 
nal and internal cones both in the superior and inferior molars. This demonstrates that the ancestral probo- 
scidean molar was tetrabunodont, as in Meritherium, not hexabunodont, as in Palzomastodon. 

RipGcE Formut#.—In the carefully drawn figures of the superior and inferior grinders of Palzomastodon the 
construction of the ridge-crests of the grinding teeth should be examined with great care, because it is difficult to 
express the rudimentary condition of the third crest or tritoloph in a formula. It will be seen (Figs. 90B, 91, 92, 
93, 94, 95) that the tritoloph and tritolophid are in a formative stage; the tritoloph is rudimentary even in M® (Fig. 
94D); it consists of a single cone in M', M? (Fig. 91); the éritolophid is completely formed in M, (Fig. 92B); it 
is less fully formed in M, (Fig. 92B). Consequently Palzomastodon is far more primitive than Mastodon in the for- 
mation of the tritoloph and tritolophid, yet it exhibits the ancestral stages in the development of this third crest so 
characteristic of all the Proboscidea. 

By close comparison of all the figures of the upper and lower grinding teeth of Palzomastodon, Miomastodon, 
Pliomastodon, and Mastodon, it is observed: 

(1) That the molar crowns in Palzomastodon are mainly tetrabunodont, i.e., each protoloph (superior) and 
each protolophid (inferior) is composed of a main external bunoid cone and a main internal bunoid cone; in the 
superior molars (Fig. 94D) where the conules persist the main crown is hexabunodont. The presence of proto- 
and metaconules blocking the median sulcus forbids the ancestry of Palzomastodon to Mastodon. 

(2) The vestigial intermediate protoconules and metaconules are observed in the hexabunodont superior 
molars of Palxomastodon intermedius, thus the crested upper grinders are hexabunodont or six coned, whereas 
the lower grinders are subtrilophodont (Fig. 93, M*-Ms;, Fig. 94). This primitive condition of the cones connects 
Palzomastodon with its undiscovered sexitubercular-quadritubercular ungulate ancestors; the conules observed in 
the third superior molar, M°, of P. intermedius (Figs. 93 and 89) are not seen in M? of the same species (Fig. 92) ; 
the conules are vestigial or disappearing structures. 

(3) The protoconules and metaconules of Palzomastodon (Fig. 93) obstruct the suleus of Mastodon. In ad- 
vanced stages of Palzomastodon and of Miomastodon the superior grinders, like the inferior, become trilophodont. 

(4) To the protoloph and -lophid also metaloph and -lophid the superior and inferior grinders, M'*-M,., 
add a rudimentary tritoloph and -lophid or third superior and inferior transverse ridge-crest (Figs. 92 and 94). 
The genesis of the tritoloph and -lophid is very clearly shown in Palzomastodon intermedius (Fig. 92); this tri- 
lophodont structure in all the superior and inferior true molars becomes a distinctive character of the grinding teeth 
in all the Proboscidea, except Deinotherium in which Dp 4 and M, are trilophodont (Figs. 54, 60, 71); conse- 
quently it is an ordinal character. 

(5) In Palzomastodon the fourth superior and inferior premolars exhibit the protoloph and -lophid 
and metaloph and -lophid only, i.e., no tritoloph and -lophid, like the fourth superior premolar of Phiomia 
(Fig. 179). 


The fourth deciduous premolars of Phiomia, Dp‘-Dp,, also acquire a tritoloph and -lophid. The fourth 
deciduous premolars of Miomastodon and Pliomastodon are not known. The fourth deciduous premolars of Mastodon 
americanus have a fully developed tritoloph and -lophid (see Warren, 1852, p. 213, Pl. v). Consequently a 
trilophodont fourth deciduous premolar appears to be a progressive ordinal character of the Proboscidea. 

(6) Thus the third lophs, tritoloph and tritolophid, are successively established in M*-Ms, in M?-Mg, in M?-M,, 
and in P*-P, by the addition of a third crest. 

Amer. Mus, 18237 Type P. matthewi Amer. Mus. 19248 a Ref. 

fr. mg (rey.) 

outer views 

1/4 nat. size 

Amer. Mus. 13449 Paratype Palaeomastodon intermedius Amer. Mus. 14547 Type 

Fig. 88 

Fig. 87. Crown view of inferior dentition and jaw of Mastodon americanus, 
containing Dps, Mi, Mo, and Mz; anterior lophs. After Warren, 1852, Pl. v, 
one-sixth natural size. Compare figures 150 and 151 of Zygolophodon borsoni. 

‘ REN Fig. 88. Comparison of third inferior and superior grinding teeth of Plio- 
Fig. 87 an mastodon matthewi (upper) and Palzomastodon intermedius (lower), one-fourth 

No natural size. 

(7) The main internal and external bunoid cones in the Mastodontine phylum, namely, in Miomastodon, 
Pliomastodon, and Mastodon, remain separate, they never unite transversely into a completed ridge-crest; thus all 
the Mastodontine remain bunolophodont, they never become fully zygolophodont. 

(8) The passage from extreme brachyodonty to subhypsodonty is observed as follows: (a) In Palzomastodon 
the paired cones or lophs are extremely brachyodont; (b) the cones or lophs in the three successive stages of 
elevation are seen in the ascending species of Miomastodon merriami, Pliomastodon matthewi, Miomastodon 
tapiroides americanus, and Pliomastodon americanus praetypica, as described in the Miomastodon section; (c) 
the summits of the cones are converted into sharply acute ridge-crests, as observed in the unworn grinders of 
Mastodon acutidens (Fig. 135 and PI. 1, L). 

(9) Observe wide differences between Palzomastodon and Phiomia in conules and trefoils: (a) In Palzo- 
mastodon the central conule (Fig. 90) is arrested, proto- and metaconules close the sulcus (Fig. 93); (b) 
in Phiomia the central conules progress strongly (Fig. 90); in the inferior molars of (c) Miomastodon and 
Pliomastodon the ectoconelets expand into trefoil spurs (Fig. 89); in (d) Mastodon americanus rudimentary 
trefoil spurs are observed on ectoconelets below and entoconelets above, the mesoconelets being reduced. 

Summing up, the progressive ‘Mastodontine’ characters are as follows: (1) Early loss of the conules; (2) 
very gradual development of the tritoloph; (3) relatively rapid development of the tritolophid; (4) persistent 
separation by a sulcus of the internal cones from the external cones, hence bilobed; (5) early development of the 
internal cingulum on the superior molars, retarded development of the external cingulum on the inferior molars. 

M. americanus 
Amer. Mus. 14293 Ref. 

M. americanus 
Amer. Mus. 14294 Ref. 

| M. pretypica (Schlesinger) 

1 1 
t H f 
i i : \ P. matthew! 
4 \ \ - matthew! 
P, matthewi 
' i j \ \ Amer. Mus. 18238, 18239 
A fi Amer. nibs. Lar Type \ \ i Paratypes 
‘ : i 
i i Lt a eee 
‘ vi ‘ xa 
fs H ‘ 
ea i : 
ae 1 ’ 
x } ; ' 
“A H 
H ! 1 
nN | ' of 
p! / / 1 
7 oSe a} 
x M 1 r yee <7 ~---j 
\ ~ , 1 H 1 ' 
\ i q ‘ ' : 1 
: \ ‘ H ' M. merriami ' 
\ r 1 ' : \ Colo. Mus, 92 Type 
‘ \ it 4 {Drawn from cast of Type Amer. Mus. 14471 
‘ H ' — 
ag \ i H : ' 
Se BS x All Z natural sige be 
~~ & Q 4 
£ 8 8 Ss 
rae ghee S 
8s 8 g *. 
ace, sok Pk 
\ 7 
\ \ \ 
n \ ' 
1 ‘ 
\ \ 

Palaeomastodon intermedius 

Amer. Mus. 13449 Paratype 
‘ 1 H 



i Img ™2 (rev.) 
‘\ Palaeomastodon intermedius 

‘\ Amer. Mus. 14547 Type 
. ‘ 

Compare PLs. I-rv, pp. 134-135 
Superior Series 
Mastodon americanus, r.M2; r.M® with 4} ridge-crests. 

Fig. 89. Evonurion or THE Morar RivGe-crests IN THE MAsTODONTS, OLIGOCENE TO PLEISTOCENE 
Pliomastodon americanus praetypica, r.M?; r.M*with 4) 

Inferior Series 
Mastodon americanus, r.M2; r.M3 with 4) ridge-crests. 

Pliomastodon americanus praetypica, r.M2; r.Ms with 5 
Pliomastodon matthewi type, r.M° with 3% ridge-crests. 

Pliomastodon matthewi paratypes, M3 with 4}; ridge-crests. 
Palxomastodon intermedius, r.M® with 2} ridge-crests. 

Miomastodon merriami type, r-Ms with 4) ridge-crests. 

Palxomastodon intermedius, 1.Mg3 with 3 ridge-crests. 
Observe: (1) Inferior molars more progressive than superior molars; (2) a) anterior ridge-crests (protoloph, -lophid) rela- 

tively constant, b) second ridge-crests (metaloph, -lophid) progressively broadening, c) third ridge-crests (tritoloph, -lophid) 
progressively broadening, d) fourth ridge-crests (tetartoloph, -lophid) progressively broadening, e) fifth ridge-crests (pentaloph, 
-lophid) more progressive in M3 than in M®; (3) third inferior molar elongate, gently narrowing posteriorly, third superior molar 
less elongate, more rapidly narrowing posteriorly; (4) absence of median trefoil spurs in Palzomastodon, presence of external 
trefoil spurs in Miomastodon and Pliomastodon: (5) rudimentary ectotrefoil spurs in Mastodon inferior molars. Observe (6) 
that in this figure the grinding teeth of all the true Mastodontine exhibit a median sulcus and four primary conelets on each 
ridge-crest; also a rudimentary trefoil spur extending from the entoconelets cf the superior molars and from the ecto- 
conelets of the inferior molars. (7) In Pal#omastodon intermedius the proto- aud metaconules close the median sulcus. 



Fina Ripce Formut™.—The ridge formula of the deciduous and permanent teeth of Mastodon americanus 
is as follows (see Warren, 1852, pp. 64-73): 

Dp: small, two ridge-crests, bilophodont. Dp? (?) two ridge-crests, bilophodont. 
Dp; larger, two ridge-crests and talon, bilophodont. Dp? two ridge-crests, bilophodont. 
Dp, three ridge-crests, cingulum, trilophodont, six fangs. Dp? three ridge-crests, trilophodont. 
M, larger, three ridge-crests. M! three ridge-crests, trilophodont. 
Mz three ridge-crests. M? three ridge-crests, trilophodont. 
M; four ridge-crests, with complex talon, tetralophodont. M? four ridge-crests, and small talon, tetralophodont. 

The ridge formula of Mastodon americanus appears to be as follows: 
Dp2% Dp3?# Dp4%? M1% M2? M3 

4 Sia 

The dental formula of Mastodon americanus accords with the observations of Hays (1834) and of Warren 
(1852, p.63). It would appear from Warren’s description that the fourth permanent premolar, P*-P,, does not erupt 
in Mastodon americanus. It is known in Serridentinus productus to be a bilobate tooth. Thus the final true 
Mastodon stage in the grinding teeth of the Mastodontine appears to give us the following dental formula: 

Deciduous premolars = Dp**-Dp.4. 

Permanent premolars= P4 in germ, P2, P3 suppressed. 

Permanent molars=M'?-M,3. 

Total number of grinding teeth which come into use in both jaws = 24. 

A ee 
Phiomia osborni 
Amer. Mus. 13468 Type m3 

Palzomastodon beadnelli 
Drawn from cast of type Amer. Mus. 9984 

1/4 nat, size 

Fig. 90. Superior view of largest known jaws of (A) Phiomia and (B) Palzomastodon superposed for comparison. One-fourth natural size. 
A, Type jaw of Phiomia osborni Matsumoto (Amer. Mus. 13468). Observe: (1) The relatively long, narrow grinding teeth; (2) the ex- 
tremely elongate symphysis extending back between right Ps and left P3; (3) an ancestral shovel-tusker (cf. Amebelodon); incisive alveoli elongate. 
B, Type jaw of Palzomastodon beadnelli Andrews (Cairo Mus. C. 10014), drawn from cast of type (Amer. Mus. 9984). Observe: (1) The 

relatively broad inferior molars; (2) the abbreviated inferior grinding series; (3) the extreme anterior position of mandibular symphysis; (4) 
incisive alveoli rounded, abbreviate. 



Famity: MASTODONTID Girard, 1852 
Subfamily: PAL#moMASTODONTIN subfam. nov. 

Osborn, 1934: The genus Palxomastodon was originally placed by Osborn, following Matsumoto, in the 
family Mastodontid#, subfamily Mastodontine, on the basis of the broad similarity of its grinding teeth to those 
of Mastodon. Recent and more intensive observation compels Osborn to remove Palzomastodon' from its supposed 
ancestral relationship to Mastodon or to any other genus of the Mastodontinee. We must await further knowledge 
of this relatively rare animal before we can determine its phyletic relationships. 

Genus: PALZ OMASTODON Andrews, 1901 

Original reference: Zoologist, 1901, (4), V, Aug. 15, pp. 318, 319, (Andrews, 1901.1); Tageblatt des V Internat. Zool.-Cong., 
Berlin, No. 6, Aug. 16, p. 4. (Published volume, Verhandlungen, 1902, p. 528). (Andrews, 1901.2). 
Genotypic species: Palzomastodon Beadnelli (Andrews, 1901.1, pp. 318, 319). 

Generic CHaracters.—(Matsumoto, 1924.1, p. 3): “A genus of Proboscidea. Skull imperfectly 
known. Palate rather short and wide, as compared with that of the next genus [Phiomia]; judging from 
the form of the palate this genus might be less long-skulled than the next one. Mandible elongated 
anteroposteriorly; mandibular symphysis rather short as compared with that of Phiomia; posterior end 
of symphysis lying a considerable distance anterior to the first cheek tooth (P;); largest and most con- 
spicuous one of mental foramina lying just below the first cheek tooth, and far behind the posterior end 
of the symphysis. Dental formula: It. Ct. P. M$. Ridge formula: Dm+}7;. P#}*. [M*%%"].” Cheek teeth 
markedly brachyodont; last premolars and all molars short and wide, bunolophodont, wearing like 
typically lophodont teeth, attaining rather sharp ridges and very widely open valleys when moderately 
worn; a rudimentary intermediate cusp is present in the anterior valley of each lower molar; no trefoil 
pattern of cusps; surface of enamel rather smooth; basal cingula neither very strong nor very rough.”’ 

As pointed out clearly in Chapter IT, Sections I-IV, the genus Palxomastodon, based on the genotype P. beadnelli 
and now amplified by our present knowledge of the species P. parvus Andrews and P. intermedius Matsumoto, 
represents a group of species entirely distinct from the Phiomia group. Inasmuch as Andrews’ final definitions 
(1906, p. 130) both of the genus Palzomastodon and of the family Paleomastodontide based on this genus are 
entirely founded on the generic characters of Phiomia, we are compelled with regret to abandon Andrews’ defini- 
tion both of the genus Palxomastodon and of the family Paleeomastodontide and to substitute, with a few omis- 
sions and slight modifications, the definition of Matsumoto which is founded on a correct diagnosis of the characters 
of Palxomastodon so far as known. In Palxomastodon the lower incisors are apparently short and rounded; in 
Phiomia they are long and flattened. 

The above generic characters are confirmed and amplified in the three ascending species included within this 
genus and distinguished by measurements as follows: 

Palzomastodon parvus Palxomastodon intermedius Palzomastodon beadnelli 
Andrews, 1905 Matsumoto, 1922 Andrews, 1901 
Length of lower molar series, M.-s, Length of lower molar series, M)-, Length of lower molar series, My-s, 
130-131 mm.; of lower premolar-molar 159-161 mm.; of lower premolar-molar 191-194 mm.; of lower premolar-molar 
series, Ps-M3, 169 mm. series, P3-Ms, 190e mm. series, P3-M;, 279 mm. 

Rarity or PaLaomMAstopon Remarns.—The relative rarity of specimens referable to the three species of 
Palzomastodon is very significant indeed; in the collections of the British, Cairo, and American museums com- 
bined the total number of specimens of teeth and jaws referable to these three species may be approximately 
enumerated as follows: 

*(1935) Now placed in the new subfamily PALAaoMASTODONTIN® (see Appendix). 
*The Matsumoto ridge formula (1924.1, p. 3) of Palwomastodon is apparently a lapsus calami or a misprint. [The ridge formula in Palwomastodon 
beadnelli and P. intermedius is: M 1 24 M 2244 M3 *4) 



Palzomastodon parvus: 4 specimens Brit. Mus., 1 specimen Amer. Mus., total... ..............-. 5 
Palzomastodon intermedius: 4 specimens Amer. Mus............ 260-0000 eee eee rt eee 4 
Palzomastodon beadnelli: 2 specimens Cairo Mus., 2 specimens Amer. Mus., total................. 4 

Grand total skull, tooth, and jaw specimens certainly referable..................+++++++--- 13 

In the American Museum collection there are seven specimens of Palzomastodon and seventy-nine specimens of 
Phiomia, the relative frequency of Palzomastodon to Phiomia being as one to eleven. 

Osborn, 1934: We observe (Figs. 91-93) that the progressive third superior and inferior molars of Palzomasto- 
don intermedius exhibit the presence of proto- and metaconules, hence hexabunodont; these conules block the median 
sulcus which distinguishes the genera Mastodon, Zygolophodon, Turicius, and Stegolophodon (as shown in Pls. 
L-tv, pp. 134-135). These characters remove Palzomastodon from the ancestry of the Mastodontine. 

| m, Paratyoe /3449 

Fig. 91. Type and paratype specimens of Palzomastodon intermedius Matsumoto, 1922, one-fourth natural 
size. Drawings prepared under the direction of Dr. H. Matsumoto. 

B, Type of Palzomastodon intermedius Matsumoto, 1922, Amer. Mus. 14547 (reversed in drawing). B1, 
Crown view of the same type specimen (reversed in drawing). 

A, Paratype palate of Palzomastodon intermedius (Amer. Mus. 13449); P%, P* drawn in from Amer. Mus. 
14548 (reversed). A1, The same paratype specimen, palatal view. 

outer side 

Fig. 92. Type and paratype 
lower and upper grinding teeth 
of Palzomastodon intermedius 
Matsumoto, 1922. Both figures ee ‘ m2 
two-thirds natural size. as 

B, Crown view, left lower mo- 
lars of type (Amer. Mus. 14547). 

A, Crown view, left upper den- 
tition of paratype (Amer. Mus. 
13449); P*®, P* drawn in from 
paratype specimen (Amer. Mus. B 

Palaeomastodon intermedius 
Amer. Mus. 13449 Paratype 

Palaeomastodon intermedius 

Amer. Mus. 14547 Type outer side 
2/3 nat. size 

a « 

Palaeomastodon intermedius Palaeomastodon intermedius 

Amer. Mus. 13449 Paratype Amer. Mus. 14547 Type 
Di upper m. JF f7 rid 
external ieracl 

metacone _ paracone 

OE oie BIRT 
& metacond 

By Qi 


Grd crest” 

/ypocone ' yrotocone Pay 
Internal cinguluin Natural size hypocomd central conule 

Fig. 93. Key to the hexabunodont upper and lower molar crowns of Palzomastodon intermedius; M3 drawn from the type (Amer. 
Mus. 14547), M® from the paratype (Amer. Mus. 13449). See figure 92A, B, for same type and paratype. Natural size. 

Observe: (1) Relatively broad proportions; (2) hexabunodont crowns of M’, namely, protocone, protoconule, paracone, hypo- 
cone, metacone, metaconule; (3) fundamental arrangement in two transverse crests, i.e., protoloph, metaloph; (4) two intermediate 
cusps, i.e., protoconule, metaconule; (5) lower molar M3 has rudimentary trilophodont crown; (6) both upper and lower molars wholly 
distinct in proportions from those of any species of Phiomia; (7) central conule rudiment in My; (8) absence of median suleus. 


AM. 13431 

AM.13437 AM.14547 

Tig. 94. Detailed studies of: True Palwomastodon intermedius third superior and inferior hezabunodont (D, E) teeth, M*-Ms, 
compared with the essentially tetrabunodont (A, B, C) Meritherium teeth, superior and inferior. From originals and casts in the 
American Museum. All figures natural size. 

A, Meritherium lyonsi ref., right M'* drawn from cast Amer. Mus. 15898. 

B, Meritherium trigodon ref., right superior grinders drawn from Amer. Mus. 13431. 

C, Maritheriwm andrewsi ref., left M-3 drawn from Amer. Mus. 13437. 

D, Palzxomastodon intermedius paratype, third right superior molar, r.M*, Amer. Mus. 13449. 

E, Palxomastodon intermedius type, third left inferior molar, 1.Ms, Amer. Mus. 14547 (reversed in drawing). 




portion of Andrews’ descriptions and all his text figures of Palzomastodon beadnelli, except those of the type 
jaw and teeth, belong not to this species but to Phiomia wintoni. All the principal specimens of skulls and upper 
jaws referred by Andrews to Palzxomastodon beadnelli also appear to belong to Phiomia wintoni, except the 
specimen (Cairo Mus. C.10014a) illustrated in his Pl. xv, fig. 2, representing a fragment of a maxilla with the 
second and third molars, M?-M®, in situ which appears to truly belong to P. beadnelli; these teeth are clearly 
shown to be bilophodont, a generic character of the Palzomastodon superior molars. 

Palwomastodon parvus Andrews, 1905 

For original description and type figure, see pp. 59, 60, of the present 

Fluvio-marine formation of the Fayim, Egypt=Upper Eocene of 
Andrews = Lower Oligocene of the present Memoir. 

This is the smallest and most primitive Palzomastodon known; 
it is probably of much greater geologic age than P. beadnelli, the 
first species discovered. The full characterization of this species 
in the systematic revision of Chapter II above and the compara- 
tive measurements as given by Matsumoto and corrected by Osborn 
leave little to be added. It is a relatively rare animal because it is 
known at present from only five specimens, namely, the type 
(Brit. Mus. 8479a) and three other specimens in the British Mu- 
seum, also referred Amer. Mus. 13497. 

Sprciric Cuaracters.—The specific characters cited in 
Chapter II from Andrews (1905, p. 562) and Matsumoto (1922, 
p. 2, 1924.1, p. 4), together with corrected measurements by Osborn, 
may be summed up in the following sentence: P. parvus is distin- 
guished by (1) Mis 131 mm. as compared with 161 mm. in P. 
intermedius and 194 mm. in P. beadnelli, (2) Ps-M; 169 mm. as 
compared with 190e mm. in P. intermedius and 279 mm. in P. 
beadnelli, (3) molar proportions of M;-s substantially similar to 
those in P. intermedius and P. beadnelli. 

Paleomastodon intermedius Matsumoto, 1922 

Yor original description and type figure, see pp. 63, 64. 

Fluvio-marine formation of the Fayim, Egypt=Upper Eocene of 
Andrews = Lower Oligocene of the present Memoir. 

As the name intermedius indicates this animal is intermediate 
in size between Palzomastodon parvus and P. beadnelli; it is more 
progressive than the former species in its molar teeth, the molars 
being relatively narrower. It is known from four specimens only. 

Grotocic Levet or Four Specrmens.—The quarry records 

Larger, more progressive 
Intermediate in character 
Smaller, more primitive 
Smaller, more primitive 

show that the level of the type (Amer. Mus. 14547) is not recorded; 
one paratype (Amer. Mus. 13480) is from Quarry B, another 
paratype (Amer. Mus. 13449) is from Quarry A, while a third, a 
maxilla (Amer. Mus. 14548), is not recorded. As shown in Chapter 
II, systematie revision of Palzomastodon intermedius above, there 
is no means of ascertaining the exact geologic levels of the different 
species of Palzomastodon; all that we can say is that the species 
P. intermedius is well represented by the four specimens in the 
American Museum collection, clearly illustrated in figures 39, 

Palxomastodon beadnelli 
Palzomastodon intermedius 
Palzxomastodon parvus 

Spreciric CHARACTERS.—Osborn confirms Matsumoto’s separa- 
tion and definition of this species and adds the following characters 
which are clearly displayed in figures 92 and 94: (1) The hexa- 
bunodont structure of M®%, less strongly indicated in M? which is 
tetrabunodont; (2) the sublophodont arrangement of the cones and 
conules both in the superior and inferior molars; (3) the arcuate 
arrangement of the superior and inferior molars of opposite sides. 
In addition to the specific characters quoted in full from Matsu- 
moto in Chapter II (Matsumoto, 1922, p. 2, 1924.1, p. 10, and ex- 
tended by Osborn), it may also be pointed out that the grinding 
teeth of P. intermedius, besides being of smaller dimensions 
throughout, are relatively narrower than those in the type of P. 
beadnelli, namely: 

M, M, Ms; 
Palxomastodon beadnelli, type 
INGICESS... nosemene ere: 77 78 68 
Palxomastodon intermedius, 
typennadicesiaarce ser: 64 66 62 

It may be premature, however, to establish these indices as 
specific characters until we secure more material. 

PHYLOGENETIC RELATIONSHIP.—It would also be premature to 
say whether Palxomastodon parvus, P. intermedius, and P. bead- 
nelli constitute an ascending series of species increasing in size, 
losing certain primitive characters and acquiring certain progres- 
sive characters. We must await further evidence which will be 
afforded, first, by the discovery of the complete dental and cranial 
characters of these primitive mastodonts, and, second, by the 
determination of the geologic levels in which they occur. Compar- 
ing Palzomastodon with Phiomia the same question will arise in 
respect to the latter genus, namely, whether the smaller, inter- 
mediate, and larger species constitute an ascending series. Tenta- 
tively the two lines may be arranged as follows: 

Phiomia osborni 
Phiomia wintoni 
Phiomia serridens 
Phiomia minor 

CRANIAL ProportTions.—The cranium of Palxomastodon 
intermedius (Amer. Mus. 13449) constitutes the only evidence we 
have at present as to the proportions of the skull in Palzomastodon 
(see Figs. 91, 39). These proportions are more brachycephalic 
than those of Phiomia, as evidenced (1) by the relatively widely 
arching palatal series, (2) by the harmonically broader and shorter 
grinding series, and (3) by the more widely arching zygomata. 
We may expect to find Palzomastodon with a broader and a shorter 
skull than that of Phiomia, with a large pair of upper and lower 
incisive tusks, probably rounder than those of Phiomia, with a broad 

— | 


enamel band on the concave outer surface of the upper tusks, as in 
the Miocene species of Miomastodon. 


(Matsumoto, 1924.1, p. 9), as well as the following new compara- 
tive measurements by Osborn: 


Palzxomastodon beadnelli type, Cairo Mus. C.10014 
Amer, Mus. 13481 

“oe “ee 

as intermedius type, Amer. Mus. 14547 

ss & paratype, Amer. Mus. 13480 

a parvus type, Brit. Mus. 8479a (cast Amer. 
Mus. 9976) 

Amer. Mus, 13497 
SupeRIOR MoLars 

| “ “ce 

Palzomastodon intermedius paratype, Amer. Mus. 13449 1. M'M%=150 | 42 34 81/50 42 84/57 46 81 
| r. M'M?=149 | 438 35 811] 50 438 86] 57 45 

DentaL CuHaracters.—The dental characters in P. inter- 
medius are beautifully illustrated in figures 93, 94, and especially 
in figure 92, showing the detail of the upper and lower grinding 
teeth. We observe that both upper and lower grinders are progress- 
ing towards trilophodonty; at the same time the three lobes are 
showing a tendency to unite into three transverse crests, so that 
the term trilophodont exactly describes them. This dental strue- 
ture does not sustain Matsumoto’s generalization that the genus 
Palxomastodon includes the ancestral forms of Mastodon. 

Palwomastodon beadnelli Andrews, 1901 

For original description and type figure, see p. 54 
Fluvio-marine formation of the Fayfiim, Egypt=Upper Eocene of 
Andrews = Lower Oligocene of the present Memoir. 

This is important as the largest and probably the most pro- 
gressive, as well as the first discovered and genotypic species of 

Grotocic Lrevrt.—Andrews (letter, 1922) informs the writer 
that the large type jaw (see original type figure, Fig. 26) was 
found at the very base of the Fluvio-marine beds, 50 or 100 feet 
below the typical Phiomia level; this genotype jaw (Cairo Mus. 
C.10014), with a very large referred femur (Cairo Mus. C.10017), 
tibia (C.10015), scapula (C.10016), humerus (C.10013), and axis 
vertebra (C.10061), was the only true P. beadnelli material found 
at this locality. This referred limb material (Fig. 96) is very 
important because of (1) its primitive Mastodon-like characters 
and (2) its use in estimating the size and limb proportions of 
Palzomastodon, as shown in Andrews’ restorations. 

Speciric CHARACTERS OF P. BEADNELLI.—The specific char- 
acters are shown in the second type figure by Andrews (Fig. 95), 
in the type jaw as refigured by Osborn in comparison with Phiomia 
osborni (Fig. 90), also in Amer. Mus. 13481 (Fig. 187, Pl. 1, p. 134), 
a right lower third molar, Ms, attached to a small fragment of a 
mandibular ramus from Quarry B, measuring 81 mm. in length 
and 45 mm. in width. See Matsumoto’s definition in the systematic 
revision above; also his detailed comparative measurements 

M,-M3=194 | 48 37 77/65 51 

M,-M3=161 | 42 27 64 | 53 35 66 | 

M 1 M 2 P 3-M 3 

ap. tr. U aps tr. -L-)|aps sures 
78 | 78 53 68 
|81 45 56 
65 40 62 | P;-Ms;=190e 
60 37 62 | 

| P3-M;=279 

M,-M3;=131 | 41 28 68 | 44 32 73 | 438 30 10. | Pale 160 

52 30 soil 

PM? =200e 
79 | P*M?=200e 

SrieniricaNt Morar Inpices.—The dental proportions of 
Palzomastodon are shown in the molar indices M'-M,;, M*Mbp, 
M*-M; as above. The molar indices (I.) in the three species of 
Palxomastodon are printed above as showing the ratio of transverse 
(tr.) to anteroposterior (ap.) measurement; although greatly 
modified by crushing of the teeth (as for example in comparison of 
Amer. Mus. 13481 with Cairo Mus. C.10014) they nevertheless 
serve to distinguish Palzomastodon very clearly from Phiomia, as 
shown in the following contrast: 

Ratios (I.) or BREADTH (TR.) TO LENGTH (AP.) IN PAL&o- 

M: M. Ms 
Palzomastodon beadnelli type indices 77 78 68 
Phiomia wintoni average............ 68 62 56 

Fig. 95. Second type figure of Palzomustodon beadnelli Andrews, 1901. 
After Andrews’ “Descriptive Catalogue of the Tertiary Vertebrata of the 
Fayim, Egypt,” Pl. xv, figs. 1, 1a, 1906. Original type in Geological Museum, 
Cairo (C. 10014). One-sixth natural size. 

From this comparison we see that whereas in species of 
Palzxomastodon the breadth of Ms; is from 62% to 70% of the 
length, in species of Phiomia the breadth of Ms is from 54% to 

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63% of the length. Thus the indices in Palzomastodon of 62% to 
70% are uniformly broader than those in Phiomia of 54% to 63%. 
These molar indices therefore are very profound and significant 
characters, because the molar indices of Palxomastodon point 
towards those of Mastodon; the molar indices of Phiomia point 
towards those of Trilophodon. For example, in Serridentinus 
productus (Amer. Mus. 10582) the third molar indices are 
47-60, the proportions of M; being about the same as in Phiomia 

In Palzomastodon the lower molars are relatively shorter and 
broader; in Phiomia they are relatively longer and narrower. 
There is a less wide contrast in the upper molars. 

CHARACTERS OF THE SKELETON.—Andrews (1906, pp. 145- 
147) observes of the only bones found in the same locality as the 
type of P. beadnelli as follows: 

“ Fore limb.—An incomplete left scapula collected in the same 
locality as the type mandible is the only evidence of that bone yet 
discovered. It is very closely similar to the scapula of Hlephas, 
differing only in the rather slighter prominence of the coracoid 
process and in widening out less rapidly posteriorly above the 
glenoid cavity, the whole blade being probably rather less expanded 
above than in Elephas. The glenoid cavity is an elongated oval, 
the prescapular fossa is extremely narrow, and the process project- 
ing backwards from the middle of the spine in Hlephas here seems 
to be situated a little nearer the glenoid end. In the case of the 
humerus (Pl. xvi. figs. 2, 24) also the only specimen that can be 
definitely determined as belonging to Palxomastodon beadnelli, 


was collected near the type mandible. In its general character 
this bone is like that of the later Proboscideans, but is somewhat 
stouter and broader in proportion to its length, and approaches 
more nearly some specimens of the humerus of Mastodon with 
which it has been compared... . / A femur (Pl. xvi. figs. 3, 3a) 
from the same locality is the only specimen of that bone in the 
collections of Cairo and London that is definitely Proboscidean in 
character and can be referred to Palzomastodon—a circumstance 
which still further emphasises the extraordinary rarity of bones of 
the skeleton of that animal. . . . A left tibia (Pl. xv. figs. 4, 4) 
from the same locality is, on the whole, like that of the Elephant. 
It is, however, rather more slender in the shaft, and the distal 
articulation differs in several important points. . . . No bones of 
the foot were found with the limb-bones just described, but a 
single calcaneum (text-fig. 51), from some distance off, so nearly 
resembles in essential characters the caleaneum of the later Pro- 
boscideans, more particularly of the Lower Miocene Tetrabelodon 
angustidens, that it may be safely referred to Palzomastodon, and 
from its large size probably belongs to P. beadnelli.”’ 

Andrews combined these materials, probably chiefly refer- 
able to Palzomastodon beudnelli, with other bones probably refer- 
able to Phiomia wintoni to produce his reconstructions of the 
animal. It is probable that the skeletal parts which he referred 
to Palxomastodon parvus belong to the species Phiomia minor. 

Osborn’s restorations of Palzomastodon as a forest browser 
(Figs. 2, 97) are entirely different both as to form and function from 
Osborn’s restorations of Phiomia (Figs. 2, 17) as an ancestral 


By Margret Flinsch, under the direction of Henry Fairfield Osborn 
Skeletal parts after Andrews, 1906, Pls. xv, xv1 (Fig. 96 of the present Memoir) 

Fig. 97. The rarity of Paleomastodon beadnelli in the Fluvio-marine beds, as compared with the great abundance of Phiomia, 

indicates that this was a forest-living animal which rarely entered the Phiomia habitat. 

Forest-living animals throughout the entire 

Tertiary are not often found fossil in association either with plains-living, palustral, or amphibious forms. For limb proportions of this 
restoration, see figure 96 opposite. Osborn, 1934: The restoration of the primitive proboscis and of the single pair of lower incisive 
tusks may prove to be incorrect; we await further knowledge of the anterior portions of the upper and lower jaws and of the tusks. 

M. americanus 
Amer. Mus. 9845 Ref. 

M. merriami 
Colo. Mus. 92 Type 
Drawn from cast of Type Amer. Mus. 14471 

w=-- M. matthewi 
Amer. Mus. 18238, 18239 


Miomastodon merriami. ref. LMx 

M. matthewi ! 
Amer. Mus. 18237 Type 


M, aPiroides. 
amencanus Schles. 

f tapiroiges Schies. 

All 1/4 nat. size 


Fig. 98. Comparison of the type molars of Miomastodon and of Pliomastodon of the Miocene and Pliocene and of the true Mastodon americanus of the 
Pleistocene of America and Europe. After Osborn, 1921.522, fig. 1, except Al, which is after Frick, 1933, fig. 28. All figures one-fourth natural size. 

A, Type, r.M%, of Miomastodon |= Pliomastodon] matthewi Osborn, Snake Creek B, Pliocene, Nebraska. Amer. Mus. 18237. 

A2, A3, Paratypes of M. [=P.] matthewi. Same locality as type. Amer. Mus. 18238, 18239. 

Al, L.M; of Miomastodon merriami ref., Pawnee Creek formation, Middle Miocene, Colorado (F:A.M. 23345). After Frick, 1933, fig. 28. 

B, Type, |.Mz and r.Ms, of Miomastodon merriami, Virgin Valley formation, Middle Miocene, Nevada, Colo. Mus. 92; cast Amer. Mus. 14471. See also 
figure 99 for superior molars and tusks of same type. 

C, Referred molar, |.M;, of Mastodon americanus, from the phosphate beds of South Carolina, in the American Museum (Amer. Mus. 9845). 

D, D1, Originals after photographs by Schlesinger. Types, 1.Mo-I.Ms, |.M*, Mastodon tapiroides americanus Schlesinger, Tasnid, Usztaté Kom., Hungary. 
Lower Pliocene. Compare Schlesinger, 1922, Taf. x1v, fig. 1 [=Miomastodon tapiroides americanus of the present Memoir]. 



SuBFAMILY: MasroponTIN® Brandt, 1869—Osborn, 1910. 

Genus: MIOMASTODON Osborn, 1922 

Original reference: Amer. Mus. Novitates, No. 49, p. 4 (Osborn, 1922.564). 
Genotypic species: Mastodon merriami Osborn, 1921. 

Mastodontine apparently directly ancestral to Pliomastodon and Mastodon. Skull and skeleton 
unknown. Dental formula incompletely known: 1:3 Dp} P} M+. Incisive tusks with broad enamel 
band, broadly suboval in section, downturned (M. merriami). Grinding teeth somewhat more elevated 
than in Palzomastodon; much less elevated (subhypsodont) than in Mastodon. Ridge formula prob- 
ably: P4% M1%M2%M3;45. Crests on the ‘intermediate molars’ probably fully formed, i. e., 
trilophodont. Internal and external lobes (i.e., cones) of each loph separated by median or longitudinal 
sulcus, as in Mastodon; summit of each lobe (i.e., cone) double or bifid (Fig. 98B) as in Mastodon, unlike 
Palzomastodon in which there is no longitudinal sulcus. Each lobe transversely oval in horizontal section 
but less compressed anteroposteriorly than in Mastodon. Lower tusks rodlike, horizontal, vertically 

The above generic characters are confirmed, amplified, and slightly modified in the four ascending species 
included within the genera Miomastodon and Pliomastodon and distinguished as follows: 

Miomastodon' Pliomastodon 
Miomastodon merriami Osborn, Miomastodon tapiroides ameri- Pliomastodon americanus prae-  Pliomastodon matthewi Osborn, 
1921. Genotypic — species canus (Schlesinger in Osborn, typica Schlesinger, 1919-1922. 1921. Type (Amer. Mus. 
(Colo. Mus. 92). 1921). Type. (Fig. 98D, Cotypes. (Fig. 103.) 18237); paratypes (Amer. 
D1.) Mus. 18238, 18239). (Fig. 


Tusks concave on outer side, Incisive tusks with broad Incisive tusks not determined. Superior(?) incisive tusks 
with broad enamel band, down- enamel band on convex external rounded, upturned, enamel 
and out-turned. Lower tusks surface. Estimated length of third band vestigial or absent. 
rodlike, horizontal, vertically superior molar, M’, 144 mm., Lobes of grinders slightly 
oval. j of third inferior molar, Ms, 190 more elevated or subhypsodont. 

M; measures 179 mm. antero- Ridge-crests brachyodont or mm. M2? measure 246 mm, . Serondinfenor molar, Mz, meas- 
posteriorly, transverse 83 mm.: subhypsodont, as in M. mer- ME, measure 304 mm. In ures85e mm., anteroposteriorly ; 
M23 measure 10}3 in. (=269  riamz; less erect than in P. go. nearly equal to molars of third superior molar, r.M?, 
mm.). matthew?; Mos measure 284 mm. Mastodon — americanus: MP3. gcasunes.. J45e- xine. aankeeb: 

anteroposteriorly. measure 311 mm., Mes measure posteriorly. 
330 mm. 
Middle Miocene of Nevada Lower Pliocene of Hungary. Middle Pliocene of Hungary. Lower Pliocene of Nebraska. 

and Colorado. 

The genotypic species of Miomastodon is Mastodon merriami Osborn (Figs. 98B, 99) from the Middle Miocene, 
Virgin Valley formation, Nevada; it is distinguished by low-crowned grinding teeth, especially by the persistence 
of a broad enamel band on the outer side of the superior incisive tusks. The geologically succeeding species 
Mastodon matthewi becomes genotypic of the new genus Pliomastodon, which entirely lacks the enamel on the 
incisive tusks, as also probable in Mastodon americanus praetypica Schles.; moreover, the ridge-crests in Plio- 
mastodon are somewhat more elevated (compare Figs. 89, 98, 102, 394). 

The genotypic specimens (Fig. 98B, 99) from the Middle Miocene of Nevada are now reinforced by the 
more complete referred specimen (Fig. 100) recently discovered by Frick in the Pawnee Creek, Colorado, which 
fortunately reveals the lower jaw and tusks. 

'The new and very primitive species Miomastodon depereti of the Lower Miocene of France is described in the Appendix at the close of the present 
Volume. See also comparative Pl. 1, pp. 134-135, also figs. 138A, and 230D. 



Discovery oF MIOMASTODON AND PLIOMASTODON IN EuROPE AND AMERICA.—We owe to Schlesinger the dis- 
covery of several very distinctive specimens related to the true Mastodon from the Lower and Middle Pliocene of 
Hungary, to which he gave the name M. tapiroides americanus; original photographs of these specimens were 
kindly forwarded to the author of the present Memoir, who commented (Osborn, 1921.522, p. 2) as follows: 
“There cannot be the least doubt, however, as to the affinity [to the true Mastodon] of the grinding teeth found 
in the Lower Pliocene of Hungary, to which Schlesinger applies the name M. tapiroides americanus. ‘These teeth 
are reproduced herewith (Fig. 1, D, D1) from unpublished photographs, kindly forwarded by the author, to the 
same scale with corresponding grinders (A, Al, A2, A3) from the Lower Phocene, Snake Creek formation, of 
western Nebraska, also with lower teeth (B) from the Middle Pliocene [Middle Miocene, Virgin Valley formation], 
near Thousand Creek, Nevada, and with (C) the posterior lower molar of M. americanus from the American 
Pleistocene.” The figure alluded to in this citation (Fig. 1) is reproduced as figure 98 of the present Memoir, 
the lettering of the plate being unchanged, but with the substitution of M. merriami ref. (A1) for the erroneous 
paratype of P. matthewt. 

Osporn, 1921.—In the same article Osborn (Osborn, 1921.522, p. 2) published under the name of Schlesinger 
the species ‘‘Mastodon tapiroides americanus Schlesinger’’ based, as described in detail below, upon “upper and 
lower grinders from the Lower Pliocene, Tasndd, Usztatoé Kom., Hungary,” embracing “a third left superior 
molar (Fig. 1, D1, see Pl. xm, fig. 5 [error, H. F. O.], Schlesinger), also two left inferior molars, m.-m; (Fig. 1, D, 
see Pl. xrv, fig. 1, Schlesinger). The linear measurement of the crowns agrees closely with that of the Pleistocene 
M. americanus, but the vertical measurement is apparently less, i.e., less hypsodont. This indicates that already 
in the Lower Pliocene the mastodonts had attained the massive proportions of their Pleistocene descendants. 
The lophs are similarly composed and show no trace of a trefoil ridge. There is nothing to debar these Lower 
Pliocene mastodonts of Hungary from the true ancestral line of our Pleistocene Mastodon.” 

The type incisive tusks of Mastodon tapiroides americanus Schlesinger from the Lower Pliocene of Hungary 
agree with the type incisive tusks of Mastodon merriami from the Middle Miocene, Virgin Valley formation, near 
Thousand Creek, Humboldt County, Nevada, in the presence of a broad enamel band; as Osborn observed (op. 
cit., p. 4), ‘“A very important character is the presence of broad enamel bands on the upper tusks (Fig. 2, C, D, E), 
which are perhaps similar to the enamel bands observed by Schlesinger in the true Miocene and _ Plio- 
cene mastodons of Hungary.”’ 

In the same communication (pp. 4, 6, Figs. 1B and 2) the species Mastodon merriami Osborn was founded 
upon two left lower molars and portions of the upper molars and of the two upper tusks (Colo. Mus. 92) discovered 
in April, 1909, in the Virgin Valley formation, Middle Miocene age, near Thousand Creek, Humboldt County, 
Nevada. These teeth, constituting the first evidence of the arrival of the true Mastodon in America, were discovered, 
as described by the geologist Mr. Richard C. Hills, in a formation consisting of more or less stratified voleanic ash 
containing much opalized wood. Mr. Hills presented his priceless specimens to the Colorado Museum of Natural 
History (Colo. Mus. 92) and they were loaned by Director Figgins to the present author and figured as the types 
of Mastodon merriami (op. cit., figs. 1, 2). 

In Science, 1921 (Osborn, 1921.523), these discoveries were summed up as follows: 

If the Mastodon merriami of Nevada proves to be of Middle Miocene age, it will demonstrate that these true 
mastodons came to this country much earlier than we have been led to suppose. 

I am greatly surprised and interested by the Middle Miocene appearance of the true mastodons in America, if the above 
report by Dr. Merriam is correct, as I have no doubt it is. Middle Miocene age is, in fact, quite consistent with the structure of 
the superior . . . tusks, which bear a broad enamel band on a concave outer side, a fact that puzzled me greatly because Dr. 
Schlesinger describes the Lower Pliocene mastodons of Hungary as bearing an enamel band on a convex outer surface. We 

should expect the earlier mastodons to show just the difference in the curvature of their tusks which these two observations 
would indicate. 


Mipp_e Miocene or AMERICA AND EurAstA.—A year later (1922) Osborn realized that the presence of a 
broad enamel band on the incisive tusks in these three species necessitated their separation from the true genus 
Mastodon, and he accordingly proposed (Osborn, 1922.564, p. 4) for the reception of these species the new genus 
Miomastodon which he defined as below. The genotypic species Mastodon merriami from the Middle Miocene, 
Virgin Valley formation, of Nevada, is the only specimen in which the incisive tusks were found in association 
with the grinding teeth, thus constituting an excellent type. 

The rarity of the true mastodonts in Eurasia and America is attributable to their forest-living habits; they 
occur rarely in Austria, as recently described by Schlesinger of Vienna. The Mastodon merriami of Nevada, as it 
proves to be of Middle Miocene age, demonstrates that these true mastodonts came to America much earlier than 
we have hitherto been led to suppose. Middle Miocene age is quite consistent with the structure of the superior 
incisive tusks, which in M. merriami bear a broad enamel band on a concave external surface, whereas Schlesinger 
describes the Lower Pliocene mastodont M. tapiroides americanus as bearing an enamel band on a convex outer 
surface. We should expect the early true mastodonts to show just the progressive differences in the curvature 
and enamel of the tusks which these two observations would indicate. The species Mastodon merriami from the 
Middle Miocene of Nevada is apparently much more primitive than the species Mastodon matthewi from 
the Pliocene of the Snake Creek formation of Nebraska. 

OsBorn, 1922.—Osborn’s formal description of the genus Miomastodon follows (Osborn, 1922.564, p. 4): 

Miomastodon, new genus 

Genoryric Srecres.—Mastodon merriami Osborn, 1921 . . . from the Virgin Valley formation, Middle Miocene of Nevada. 
Generic Cuaracters.—A member of the true Mastodontine phylum leading into the Mastodon americanus type, distin- 
guished from the true Palzomastodon beadnelli of the Lower Oligocene of the Faytim, Egypt, by rounded, greatly enlarged, up- 
curved superior tusks; form of inferior tusks not certainly known, probably rounded and more or less encased in enamel; distin- 
guished from the true Pleistocene Mastodon americanus by the presence of a broad enamel band extending from the base to the 


summit of the tusk. Ridge formula: M 2 3, M 3 #4“, as compared with the Palzomastodon beadnelli ridge formula: M 2 =| 
M 3 *44; as compared with the Mastodon americanus ridge formula: M 2 3, M 3 745. 

To this genus may at present be referred four species, namely: 

Genotypic species, Mastodon merriami, Middle Miocene, Nevada= Miomastodon merriami. 

Mastodon proavus Cope, 1873, late Middle Miocene, Pawnee Creek, Colorado=Miomastodon proavus {subsequently 
referred by the present author to Serridentinus]. 

Mastodon matthewi Osborn, 1921, Middle Pliocene, western Nebraska=Miomastodon matthewi [now Pliomastodon 
matthew]. : 

Also probably Mastodon tapiroides americanus Schlesinger, Lower Pliocene of Hungary=Miomastodon tapiroides 

The distinctive grinding tooth characters in all these Mastodontinz are: (1) that each loph (protoloph, metaloph, et seq.) is 
composed of a main internal and external bunoid cone; (2) the intermediate conule region does not develop; (3) the earliest 
grinder is tetrabunodont; (4) as the third loph is added it becomes hexabunodont; (5) as the fourth loph is added it becomes 
octabunodont; (6) whereas these four, six, and eight cones heighten (hypsodonty), they never unite transversely into a crest; 
thus none of the Mastodontinz becomes [truly] zygolophodont. 

The broad enamel band of the tusks is apparently placed on the concave surface of the tusk in Miomastodon merriamz, on 
the convex surface of the tusk in Miomastodon tapiroides americanus. 

Muwp te Puiiocene oF Huncary.—In his Memoir of 1922 Schlesinger figures (Taf. xtv-xrx) and describes (pp. 
115, 116, 227-230) a number of beautifully preserved upper and lower grinders under the name M. (Mammut) 
americanus Pennant forma praetypica from the Paleontological Collections of Budapest. Unlike the previously 




mentioned Mastodon tapiroides americanus of Lower Pliocene age, these teeth are regarded as of “Levantin”’ 
or Middle Pliocene age preceding that of Mastodon [Anancus] arvernensis. Consequently the species M. prae- 
typica is geologically the most recent of all the specific stages heretofore described. 

Through our present knowledge, therefore, there are five species known in Europe and in North America, in 

descending geologic order as follows: 


Mastodon americanus praetypica Schlesinger, Hungary, of 
“Levantin”’ or Middle Pliocene age (fide Schlesinger)...... ......=Pliomastodon americanus praetypica 
Mastodon tapiroides americanus (Schlesinger in Osborn), 

Hungary, of Lower Pliocene age (fide Schlesinger). . : 
Mastodon matthewi Osborn, Lower Pliocene, Snake Creek 

. = Miomastodon tapiroides americanus 

formation, western Nebraska (fide Matthew).. .... ..........=Pliomastodon matthewi 
Mastodon merriami Osborn, Middle Miocene, Humboldt 
County, Nevada, Virgin Valley formation (fide Merriam).......... = Mviomastodon merriami 

Miomastodon merriami ref., Middle Miocene, Pawnee Creek, 
Colorado. . Ltr enh Bie) oR Species nT eee 

Miomastodon depereti ‘sp. nov., |, Lower Miocene, Sables de 
LOrléanaiss Hrancesescmeeeeeecne een: 

. = Miomastodon merriami ref. 

Kereta eons Gas Ge asteacneee = Miomastodon depereti sp. nov. 

Miomastodon merriami Osborn, 1921 
Figures 89, 98, B, 99 

Virgin Valley formation, Middle Miocene, Thousand Creek, Humboldt 
County, Nevada; also Pawnee Creek, Colorado. 

Mastodon merriami Osborn, 1921. “First Appearance of the 
True Mastodon in America,’’ Amer. Mus. Novitates, No. 10, June 
15, 1921, pp. 4-6 (Osborn, 1921.522). Typr.—(Osborn, op. cit., 
pp. 4-6). ‘“‘The type specimens include several bone fragments, 
portions of the two upper tusks, and parts of the upper molars, in 
addition to the well-preserved two lower molars here figured as 
the type. A very important character is the presence of broad 
enamel bands on the upper tusks (Fig. 2, C, D, E), which are 
perhaps similar to the enamel bands observed by Schlesinger in the 
true Miocene and Pliocene mastodons of Hungary. . . . Type: 
Colo. Mus. 92, found in 1909 in the Thousand Creek formation 
[Virgin Valley formation], Humboldt County, Nevada, includes 
two left [now identified as left and right] inferior molars (Fig. 2, A, 
B [Fig. 1,B]; cast Amer. Mus. 14471, also portions of two upper 
tusks [and two superior molars].” TypPr Freurr.—Osborn, 
1921.522, fig. 1,B, p. 3, and fig. 2, p. 5 (Figs. 89, 98, and 99 of the 
present Memoir). 

Type DESCRIPTION (OsBORN, 1921.522, p. 6).—‘‘ The contours 
of these grinding teeth, as seen from above (Fig. 1, B), are convexo- 
(inner side) concave (outer side); the first crest is relatively 
narrow; the second, third, and fourth crests are relatively broad; 
the rudimentary fifth crest is little if any more advanced than in 
M. matthewi; crests two to four exhibit rudimentary intermediate 
cones and the spurs of a trefoil. The presence of an enamel band 
on the tusks and the somewhat more brachyodont character of the 
grinding teeth separate this stage from the M. americanus (Fig. 1, 


Fig. 99. Type superior molars and tusks of Miomastodon merriami 
Osborn, 1921 (Colo. Mus. 92), Virgin Valley formation, Middle Miocene, 
Nevada. See also figure 98,B. After Osborn, 1921.522, p. 5, fig. 2. About one- 
third natural size. Cast Amer. Mus. 14471. 

A, Two superior grinding teeth, M?, M®, of the right maxilla, internal 
view. B, Same teeth, external view, with 11-inch scale indicated. C, Superior 
tusk of the left side, lateral view. HE, External view of the same tusk. D, 
Superior tusk of the right side, external view. The presence of a broad 
enamel band on the concave outer surfaces of the right and left superior 
tusks is clearly indicated in D and E. 


C) of the Pleistocene. This species is dedicated to Professor John 
C. Merriam, in recognition of his pioneer work in describing the 
fauna of Thousand Creek.” 

Miocene Type Locauiry.—The assignment of the type 
specimen to the Middle Pliocene of the Thousand Creek formation, 
Humboldt County, Nevada, was erroneous; this error was re- 
peated by the writer in his paper of September 1, 1921 (Osborn 
1921.526, p. 332), but was corrected in the following footnote in 
the same publication: ‘Immediately after the publication of this 
paper the author learned [from Prof. John C. Merriam] that the 
type of Mastodon merriami occurred in beds of Middle Miocene 
age, which makes it geologically older than Mastodon matthewi.”’ 
(Letter, Merriam, June 24, 1921): . “TI appreciate your 
courtesy in naming the Virgin Valley species in my honor. .. . I 
note that Mastodon merriami is referred to the Thousand Creek 
formation. The [type] locality described by Mr. Hills, namely, 
that at which G. D. Mathewson secured his material, is, however, 

BS Ove 


= AM23345 



summer of 1909 I went to Thousand Creek to examine the occur- 
rence near there of precious opal. . . . I continued the journey to 
Denio, Oregon. The following morning I left for a ranch on 
Thousand Creek. At the ranch I found specimens of the opal and 
fragments of a Mastodon tooth which my guide, George D. 
Mathewson, said he had found in digging one of the excavations 
along the opal outcrop. This outcrop is situated on a precipitous 
hill about 500 feet above the level of {the stream known as] Thou- 
sand Creek and between the main forks of the creek. . . . Mr. 
Mathewson had already dug out most of the bone fragments and 
thrown them out on the side of the excavation. I dug out a few 
remaining fragments myself and went very carefully over all the 
stuff that had been excavated and thrown out. Sifting was out of 
the question at that time as the ash was so wet as to form a stiff 
mud. I placed all the fragments, including the tusks, into a sack 
which was carried down to the ranch and later packed into a box 
for transportation on the stage 130 miles to the railroad and thence 


Modified after Frick, 1933, Figs. 28, 24 

Jaw with tusk (F:A.M. 23345), one-eighth natural size. 

Molars (F:A.M. 23345, 23337), one-third natural size. 

A-A3, Mandible with second and third molars, also inferior and superior tusks and sections (F:A.M. 23345). 

This appears to be a typical Miomastodon. 

B-B2, Superior molar and tusk, also section, of another individual (F:A.M. 23337). 

This third superior 

molar, M®, differs from the typical Miomastodon molar with trefoils. 

in the Virgin Valley formation, which is of approximately Middle 
Miocene age, not far from the zone of the Mascall of the John 
Day region. . . . this evidently brings the appearance of these 
Mastodons back to near Middle Miocene.” 

Osborn corrected this geologic error specifically in Science 
(Osborn, 1921.523, p. 108). 

The Colorado Museum record of this specimen is (letter, 
Figgins, February 25, 1921): “No. 92: from the John Day forma- 
tion (Miocene), Thousand Isle Creek, Humboldt county, Nevada, 
found April, 1909, by Geo. D. Mathewson, Denio, Ore.’’ Mr. 
Richard C. Hills writes (letter, March 9, 1921): ‘Early in the 

by express to Denver. The parts must all belong to one individual. 
I spent considerable time in assembling the fragments. In addi- 
tion to the two lower molars, there are parts of the upper molars 
on one side embedded in a portion of the upper jaw. The tusks are 
not entire. You can, however, see the black enamel bands in the 
photographs. We can, if you so desire, furnish other photographs 
showing the upper molars.”’ 

Osborn, 1922: The writer is greatly indebted to Director 
Figgins, also to the geologist, Richard C. Hills for the privilege of 
examining and describing this most important specimen which 
demonstrates the arrival of a member of the true Mastodontine 

phylum in the Nevada region of America in Middle Miocene time. 
The presence of a broad enamel band on the upper tusks is a very 
important character; these enamel bands are perhaps similar to 
the enamel bands observed by Schlesinger in some of the true 
Lower Pliocene mastodons of Hungary (as recorded in his published 
Memoir) to which he has assigned the name Mastodon tapiroides 
americanus. A significant difference is that in Miomastodon mer- 
riami the enamel is on the outer concave surface of the tusk, where- 
as in the photographs of M. tapiroides americanus furnished by 
Doctor Schlesinger the enamel is on the outer convex surface. 

MIoMASTODON MERRIAMI OF CoLorADo (Fiac. 100).—These 
most welcome specimens (F:A.M. 23345, 23337), discovered by the 
Frick Expedition of 1932 in the classic Pawnee Creek horizon of 
Colorado, were referred by Frick (1933, p. 611, fig. 28) to ‘Mzo- 
mastodon proavus Cope.’ They constitute a most timely and im- 
portant addition to our knowledge of the Middle Miocene genus 
Miomastodon and greatly strengthen Osborn’s theory of the year 
1921 that Miomastodon merriami represents the arrival of ancestors 
of the true Mastodontine in America in Middle Miocene time. 

The specific reference (Frick, 1933) of these specimens to the 
Pawnee Creek type of ‘Mastodon’ proavus Cope was an error, be- 
cause the two fragmentary grinding teeth of M. proavus (see Chap. 
X, fig. 363) clearly exhibit the characteristic serrate spur on both 
sides of the ectoconelet, both in the premolar and the fragmentary 
molar, also the reduction of the contiguous mesoconelet; these 
two features are highly characteristic of the Serridentinz, as pointed 
out in Chapter X, pages 403, 404, where a detailed description with 
figures of Serridentinus proavus is given. In the lower grinders of 
Miomastodon merriami, on the contrary (see especially Fig. 98B, 
type lower grinders of Amer. Mus. 14471), there is no serration on 
the sides of the ectoconelet, but there is a distinct trefoil expansion 
of the contiguous mesoconelet. 

The expanded trefoils 6n the ectoconelets! of the Mastodon- 
tine become more conspicuous as the teeth are worn down, as 
shown in the much worn r.M; of F:A.M. 23345 of the Pawnee 
Creek (Fig. 100). This mesial expansion of the ectoconelet is also 
observed in Miomastodon tapiroides Schles. (Fig. 98 D1), in Plio- 
mastodon matthewi (Figs. 98 A2 and 102), as well as in certain speci- 
mens of Mastodon americanus (Fig. 89—Amer. Mus. 14293, r.M?, 
Amer. Mus. 14294, r.M3); this mesial expansion is therefore char- 
acteristic of the phylum Mastodontine. 

The comparative measurements are as follows: 

Type (Colo. Mus. 92): EY}, Uy, I. 
R.M3 179 80 45 
Referred (F:A.M. 23345): 
R.M; 164 77 47 
Incisive tusk (upper) 
Length 450 mm. 
Vertical diameter 67 
Transverse diameter 56 
Width of enamel band 49 
Incisive tusk (lower) 
Length 250+ 
Vertical diameter 40 
Transverse diameter 35 


New generic and specific characters revealed (Fig. 100) in the 
Pawnee Creek specimens are: (1) Meszal expansion of rudimentary 
trefoils in ento- and ectoconelets, upper and lower; (2) mandibular 
rostrum laterally compressed, abbreviate as compared with T'ri- 
lophodon or Serridentinus; (3) rodlike inferior incisors, vertical 
oval section; (4) downturned superior incisors of moderate length, 
broad enamel band, broadly oval vertical section. 

Miomastodon tapiroides americanus Schlesinger, 
1921, 1922 
Figure 98 
Lower Pliocene, Tasndd, Usztat6 Komitat, Hungary. 

Mastodon tapiroides americanus Schlesinger (in Osborn). 
“First Appearance of the True Mastodon in America,’ Amer. 
Mus. Novitates, No. 10, June 15, 1921, p. 2. Schlesinger’s descrip- 
tion: “Die Mastodonten der Budapester Sammlungen,”’ 1922, Geol. 
Hungarica, II, Fasc. 1, pp. 224-227, Taf. xm, figs. 6, 7, and xiv, 
figs. 1-4. Typn.—A third left superior molar, 1.M?, also two 
left inferior molars, 1.Mo-3. Horizon AND Locatity.—Lower 
Pliocene, Tasnad, Usztat6 Kom., Hungary. Type Fic- 
uRE.—Op. cit., Osborn, 1921.522, p. 3, fig. 1, D, D1 (see Fig. 98D, 
D1 of the present Memoir). 

Typr Description.—(Op. cit., Osborn, 1921.522, p. 2): 
“There cannot be the least doubt, however, as to the affinity [to 
the true Mastodon] of the grinding teeth found in the Lower 
Pliocene of Hungary, to which Schlesinger applies the name M. . 
tapiroides americanus. These [type] teeth are reproduced [redrawn] 
herewith (Fig. 1, D, D1) from unpublished photographs, kindly 
forwarded by the author, to the same scale with corresponding 
grinders (A, ...A2, A8) from the Lower Pliocene, Snake Creek 
formation, of western Nebraska, also with lower teeth (B) from 
the Middle Pliocene [Middle Miocene, Virgin Valley formation], 
[near] Thousand Creek, Nevada, and with (C) the posterior lower 
molar of M. americanus from the American Pleistocene.”’ 

“The upper and lower grinders from the Lower Pliocene, 
Tasnad, Usztaté Kom., Hungary, embrace a third left superior 
molar (Fig. 1, D1, see Pl. xm, fig. 5 [error H. F. O.], Schlesinger), 
also two left inferior molars, mo-m; (Fig. 1, D, see Pl. xrv, fig. 1, 
Schlesinger). The linear measurement of the crowns agrees 
closely with that of the Pleistocene M. americanus, but the vertical 
measurement is apparently less, i.e., less hypsodont. This indi- 
cates that already in the Lower Pliocene the mastodonts had 
attained the massive proportions of their Pleistocene descendants. 
The lophs are similarly composed and show no trace of a trefoil 
ridge. There is nothing to debar these Lower Pliocene mastodonts 
of Hungary from the true ancestral line of our Pleistocene Mastodon.” 

Osborn, 1925: Since the above type description was written 
for Doctor Schlesinger the species has been transferred to the genus 
Miomastodon. The incisive tusks discovered in the same locality 
and probably attributable to the same species appear in the photo- 
graphs furnished by Doctor Schlesinger to bear a broad enamel 
band on the convex external surface; if this determination is 
correct, it would seem that in these Pliocene mastodonts the tusks 
are outwardly convex, as in Mastodon americanus, a progressive 
character; as observed in Miomastodon merriami, the tusks are 
outwardly concave. 

‘Unworn molars of the Mastodontinae clearly demonstrate that the rudimentary trefoil spur springs from the ectoconelets in the inferior molars, from the 
entoconelets in the superior molars. The worn molar gives a false impression that the trefoil spurs arise from the mesoconelets. 


SuBFAMILY: MasroponTIN# Brandt, 1869—Osborn, 1910 

Genus: PLIOMASTODON Osborn, 1926 
Original reference: Amer. Mus. Novitates, No. 238, Noy. 30, 1926, p. 1 (Osborn, 1926.706) 

Generic CHARACTERS.—Incisive tusks with enamel band vestigial or wanting, suboval to rounded 
in section, upturned (P. vexillarius) ; ridge-crests with expanded ectotrefoils (P. matthewi); intermediate 
in hypsodonty between Miomastodon and Mastodon. 

Mastodon (Miomastodon) matthewi, the genotypic species of Pliomastodon, is of Lower Pliocene age, whereas 
Mastodon merriami, the genotypic species of Miomastodon, is of Middle Miocene age. In the superior incisive 
tusks of Pliomastodon (genotype) the lateral enamel band is absent; in the superior incisive tusks of Miomastodon 
the lateral enamel band is present. Both of these generic stages are imperfectly known at present. 

This Pliocene stage is based on the species first described by Osborn as Mastodon matthewi (Osborn, 
1921.522, pp. 2, 4), then as Miomastodon matthewi (Osborn, 1922.564, p. 4), and finally made the genotype 
of the genus Pliomastodon (Osborn, 1926.706). This generic stage (Fig. 98) is typified by a right third 
superior molar, r.M® (Amer. Mus. 18237), and by paratypes r.M; (Amer. Mus. 18238) and r.M, (Amer. 
Mus. 18239), also by a referred r.M,; (Amer. Mus. 19248a) and a referred tusk (Amer. Mus. 19248b)—all 
from the Snake Creek B formation of western Nebraska, quarries 1 and 5, as shown in the table (Chap. X, p. 427). 
Associated with these specimens of Pliomastodon were the type and referred specimens of Serridentinus anguirivalis, 
S. nebrascensis, and Rhynchotherium anguirivalis. 

AMERICAN PLIOCENE SERRIDENTINUS AND Mastopon.—During the summer of 1908 an American Museum 
party under W. D. Matthew and Harold J. Cook first collected in the ‘Lower Pliocene” deposits of western Nebraska, 
subsequently known as the Snake Creek beds (Matthew and Cook, 1909, p. 363). The first proboscidean molars 
discovered belonged to the serrated molar group and are now referred to Serridentinus anguirivalis. 

Subsequently, in the seasons of 1916 and 1918, several bunolophodont grinders (Amer. Mus. 18237, 18238, 
18239, and 17217) were also discovered in the same Snake Creek formation; three years later, under the title 
“First Appearance of the True Mastodon in America’’ (Osborn, 1921.522, pp. 2,4), Osborn selected these specimens 
(Fig. 98 A-A3) as the type and paratypes of a new species which he named Mastodon matthewt, in honor of Dr. W. D. 
Matthew the author who first described the new and interesting fauna of the Snake Creek formation. Amer. Mus. 
17217, however, was erroneously figured as a paratype of Mastodon matthewi and has since been made the type of 
Serridentinus anguirivalis Osborn, 1926. The reader is referred to the new figures of the type, paratypes, and 
referred specimens of Pliomastodon matthewi from the Snake Creek B level (Fig. 102), a description of which species 

Pliomastodon matthewi Osborn, 1921-1926 
Figures 89, 98, 101, 102 

Snake Creek formation, Lower Pliocene, level B, Sioux County, western 
Nebraska. Collected by the American Museum expeditions of 1916 and 1918. 

Mastodon matthewi Osborn, 1921. ‘First Appearance of the 
True Mastodon in America,’ Amer. Mus. Novitates, No. 10, June 
15, 1921, pp. 2-6 (Osborn, 1921.522). 

Miomastodon matthewi Osborn, 1922. ‘‘Dibelodon edensis 
(Frick) of Southern California, Miomastodon of the Middle Mio- 

cene, New Genus.” Amer. Mus. Novitates, No. 49, Oct. 23, 1922, 
p. 4 (Osborn, 1922. 564). 

Pliomastodon matthewi Osborn, 1926. “Additional New 
Genera and Species of the Mastodontoid Proboscidea.’’ Amer. 
Mus. Novitates, No. 238, Nov. 30, 1926, p. 1 (Osborn, 1926.706). 

Typr DescripTion AND Type Locauity.—(Osborn, 1921.522, 
p. 2): . . . “Subsequently, in 1918, several distinctive specimens 
were found in the same beds which may now be named as the type 
and paratypes of a new species of Mastodon (Mastodon matthew?), 



A112 18237 7: m3trev) 
ae (=== 

A. 19248 b 

Sw 4 Type 
A./7./9248a Te m3 

=e ae ee 
A / 18239 


Fig. 101. Second type and paratype 
figures of Pliomastodon matthewi Osborn, 
one-third natural size. Compare figure 
98A, A2, A3, also figures 102 and 99. 

in honor of Dr. W. D. Matthew, the author who first described this 
interesting fauna. . . [p. 4] Typx: the right third superior 
molar, Amer. Mus. 18237. ParatyPEs: the right second in- 
ferior [superior] molar (unworn), Amer. Mus. 17217! [this tooth has 
now been made the type of Serridentinus anguirivalis]; the posterior 
portion of a right third inferior molar (more worn), Amer. Mus. 
18238, also of a right second [first] inferior molar, Amer. Mus. 18239. 
The type and paratypes probably belong to four different individu- 
als. . . . The type and Nos. 18238 and 18239 are from the Snake 
Creek B level (Procamelus-Hipparion Zone) of Sioux County, 
Nebraska; the level of No. 17217 is not recorded. The type (Fig. 
1, A) is distinguished by the rapid narrowing of the posterior half 
of the crown of the third upper molar, including the third and 
fourth crests; the fourth crest is extremely narrow and bilobed; 
the rudimentary fifth crest consists of a single cusp. In these 
features M. matthewi is more primitive than the corresponding 
tooth of M. tapiroides americanus (Fig. 1, D1) of the Lower Plio- 
cene of Hungary. The association of the lower molars, Amer. 
Mus. 17217 [error H. F. O.], 18238, 18239, as paratypes is provi- 
sional, because the Snake Creek deposition extended over a long 
period of time and may represent more than two life zones. Of 
these teeth, [m*] presents three unworn pointed crests with the rudi- 
ments of a trefoil (Fig. 1, Al'); in a second [first] molar (. . . A2) 
the trefoil is less apparent: in the third lower molar (. .. A3) it is 
not apparent at all. In the latter tooth, which is probably the 
posterior half of a third lower molar of the right side, the third 
and fourth crests are partly preserved; crest five is represented by a 
broad tuberculate talon.”” Compare figure 98 of the present 

Specrric CHARACTERS (Fic. 102).—As compared with the 
type molars of Miomastodon merriami (Fig. 98, B), the type and 
paratype molars of Pliomastodon matthewi (Fig. 98, A, A2, A3, 

'See Chap. X, fig. 391, p. 425. 


All 4, ratural Size 


. 4 





Fig. 102. Type and paratypes of Pliomastodon matthewi, Snake Creek 
B horizon (upper), western Nebraska, one-half natural size. Type (Amer. 
Mus. 18237), a right third superior molar, r.M*. Paratypes (Amer. Mus. 
18238), a right third inferior molar, r.M3, partly restored; (Amer. Mus. 
18239), a right first inferior molar, r.Mi. For comparison with other 
mastodonts of Snake Creek, see figure 394. 


and Fig. 102) show a greater elevation (subhypsodonty) and a 
greater anteroposterior compression (lophodonty). The summits 
of the paired lobes, i.e., cones, seem to be somewhat more approxi- 
mated. Another distinctive feature is the posterior narrowing of 
the crown of the third superior molar (Fig. 98, A and Fig. 102). 

Since the original description and figures were prepared (Fig. 
98), another molar tooth (Amer. Mus. 19248a), collected in 1918, 
has been identified as a third inferior molar of the right side, r.Ms. 
In the same quarry (Quarry 5) was found a section of a superior 
incisive tusk (Amer. Mus. 19248b) somewhat abraded by river 
erosion, which shows no sign of the superior enamel band. These 
two specimens have been mentioned as indicating a more recent 
geologic age and with correspondingly progressive characters 
which signalize a new generic stage for which the name Pliomastodon 
(see Osborn, 1926.706, p. 1) seems appropriate (Fig. 101). 

Miomastodon Pliomastodon 
COMPARATIVE MEASUREMENTS = _merriami matthewi 
Type Type and Ref. 
Third superior molar, M®, 
anteroposterior......... 154e 160 
Height of internal cone of 
HICHAIODO 2 eee Soler te 60e 57 (type) 
Third inferior molar, Ms, 
anteroposterior......... 179 194 
tramsverse............. 80 
breadth-length index....... <a 
Height of internal cone of 
metalophid: «.... argc. « 54e 6le 
Superior incisive tusk, I? 
Vertical diameter....... | 92, 90 
Transverse diameter. ... { : 81 

The progressive development of the molar crowns in these 
respective Miocene and Pliocene species may also be carefully 
examined and compared from the following figures: 

Pliomastodon matthewi. Figure 98, A, A2, A3, original type and 
paratype figures; figure 89, type and paratypes in comparison 
with Miomastodon merriami, Pliomastodon americanus praetypica, 
Mastodon americanus, and Palzomastodon intermedius; figure 394, 
new type and paratype figures in comparison with other Snake 
Creek B mastodonts; figures 102 and 101, new type and paratype 

Miomastodon merriami. Figure 99, type superior molars and 
tusks; figure 98, type molars in comparison with Pliomastodon 
matthewi, Mastodon tapiroides, and M. americanus; figure 89, type 
molars in comparison with Pliomastodon matthewi, P. americanus 
praetypica, M. americanus, and Palzxomastodon intermedius. 

The above comparative measurements and figures reveal that 
in both Miomastodon merriami and Pliomastodon matthewi the 
crowns of the superior and inferior third molars, M* and Ms, 
narrow posteriorly, the tetartoloph and -lophid have not attained 
the width seen in Mastodon americanus, the pentaloph and -lophid 
are very rudimentary, whereas in Pliomastodon americanus 
praetypica the pentalophid is almost as strong as in M. americanus. 
The distinction between Miomastodon merriami and Pliomastodon 
matthew? is partly seen in the height of the crowns, the lophs in M. 
merriami (e.g., Fig. 98, B) being somewhat less elevated than in 



P. matthewi (e.g., Fig. 98, A). The progressive difference is not so 
well marked in the grinding teeth as it is in the tusks, as observed 
by comparison of the type of M. merriami (Fig. 99) and new type 
figure of P. matthewi (Fig. 102). 

Pliomastodon americanus praetypica 
Schlesinger, 1919, 1922 
Figures 89, 103, and Pl. 1, pp. 134-135 

Upper(?) Pontian and Levantin age, Middle Pliocene. Batta-Erd, 

Rikoskeresztur, Szentlérinez, and Ajndesk6, Hungary. 

Schlesinger originally described this true species of Mastodon 
from Hungary in his paper of 1919, as cited below, figuring his 
excellent types in Taf. v1, figs. 24. His fina] and more complete 
description appears in his Memoir of 1922. 

Mastodon (Mammut) americanus Penn. forma praetypica 
Schlesinger, 1919. ‘‘Die stratigraphische Bedeutung der euro- 
pdischen Mastodonten,”’ Mitt. Geol. Ges. Wien, XI, p. 142. 

Spy WV 




Fig. 103. Cotypes of Mastodon (Mammut) americanus Penn. forma prae- 
typica Schlesinger selected in the present Memoir, namely, second and third 
right superior molars, r.M**, and second and third left inferior molars, 1.M2-s, 
one-third natural size. Compare Schlesinger, 1922, Taf. xv, figs. 2 and 4. 
Originals in Hungarian National Museum, Budapest. 

Right superior molars, r.M**: Szabadka (=Maria-Theresiopel), Komitat 
Pest, Hungary. 

Left inferior molars, 1.Me-s: Batta-Erd, Komitat Pest, Hungary. 

ORIGINAL DerscripTion.—Schlesinger, 1919, p. 142): ‘‘a) 
Die intermediiiren Molaren (vgl. Tafel v1, Figur 2-4) dieser, friiher 
mit M. Borsoni zusammengeworfenen Spezies, schliessen sich im 
Bau engstens an die vorbesprochene Art an, nur sind die Kronen 
breiter, die Tiler enger. b) Fir die letzten Molaren (vgl. Tafel 
vi, Figur 1, 3, 4) gilt, abgesehen von der héheren Jochformel 
(4X im Ober-, 4X —5 im Unterkiefer) das gleiche. Hier tritt meist 
noch das Merkmal der geringeren und etwas anderen Ausbildung 
der Sperrleisten helfend dazu. Trotzdem ist die Bestimmung 
schwierig und ohne breitere Vergleichsbasis die Trennung von der 
mioziinen Form oft unméglich.” 


M. [Mastodon] (Mammut) americanus Pennant forma praetypica 
n.f., 1922. ‘Die Mastodonten der Budapester Sammlungen.” 
Geol. Hungarica, Ed. Sep., Tome II, Fase. 1, pp. 115, 116, 

Coryprs.—The cotypes selected for illustration in this Memoir 
are the second and third superior and inferior molars from two local- 
ities, as follows (Schlesinger, 1922, Taf. xv, figs. 2 and 4—same as 
Figs. 3a and 4 of Schlesinger, 1919, Taf. v1, on larger scale): “ Figur 
2: M2%dext. von der Kaufliche [Szabadka (= Maria-Theresiopel), 
(Kom. Pest). Levantin]. . . . Figur 4: Linke Mandibel mit Mo +5. 
Fundort: Batta-Erd, Komitat Pest. Horizont: Oberstes Pontikum 
(Unterplioziin). Wiedergabe aller vier Figuren: } natiirl. Gr.” 
Originals of figures 1 and 3 in the Hungarian National Museum 
of Budapest; of figures 2 and 4 in the Hungarian Reichsanstalt of 
Budapest. Coryrr Locauitres.—Aside from Batta-Erd, Schle- 
singer mentions three other cotype localities, namely, Rakoskeresz- 
ttir, Szentlérinez, and Ajnacsk6é, Hungary. Upper(?) Pontian or 
Levantine age, (?) Middle Pliocene. Coryrr Ficgures.—Schle- 
singer, 1922, Taf. xtv-xrx. These enlarged cotype figures include 
the original figures of Schlesinger, 1919, Taf. v1, figs. 2-4. 

1922, ‘Die Mastodonten der Budapester Sammlungen,” illustrates 
a large number of grinding teeth of the true Mastodon americanus 
in the collections of Budapest and of Vienna (op. cit., 1922, Taf. 
xy-xxi1) from different localities in the United States, and com- 
pares in great detail the grinding tooth characteristics of Mastodon 
americanus typica with the grinding tooth characteristics of I. 
americanus praetypica and of M. [Zygolophodon| borsoni. He also 
summarizes in several parts of his invaluable Memoir of 1922 the 
resemblances and differences between the Mastodon americanus 
and the M. [Z.] borsonz phyla. 

PRAETYPICA.—The cotypes selected in this Memoir (Fig. 103) are 
from Batta-Erd and Szabadka (= Maria Theresiopel), Komitat 
Pest, Hungary; Schlesinger mentions three other cotype localities, 
namely, Rékoskereszttir, Szentlérincz, and Ajnaesk6. (Schle- 
singer, 1922, p. 229): “Die Zahl der gut horizontierten Fund- 
punkte des M. americanus f. praetypica ist zwar nicht reichlich, 
doch sind es qualitativ hochwertige Stellen. An dreien von ihnen 
(Rakos, Szentlérincz und Batta-Erd) ist eine konkordante Auf- 
einanderfolge oberpontischer und levantiner Bildungen erwiesen; 
in allen Fiillen kamen die Reste von M. americanus f. praetypica 
aus dem mittelplioziinen Niveau.” (Op. cit., p. 230): ‘“Solche 
Belege kénnen bei der Beurteilung einer Frage natiirlich nicht 
iibergangen werden. Ich bin vielmehr der festen Uberzeugung, 
dass M. americanus f. praetypica mit dem Ende des Levantins aus 
unserem Gebiete endgiltig abgewandert ist.” 

Osborn, 1926: In his volume of 1922, Taf. x1v—xrx, Schle- 
singer figures numerous beautifully preserved upper and lower teeth 
from the Pliocene of Hungary which he refers to Mastodon (Mam- 
mut) americanus forma praetypica. As to the geologic age of these 
specimens, Schlesinger considers (op. cit., p. 227) that undoubtedly 
they are of “Levantin age” similar to that of M. arvernensis, 
namely, Upper Pontian [?Middle Pliocene]. As to geographic 
distribution and migration, he remarks as quoted above. 

We may consider as the cotypes the eight grinding teeth 
figured in Taf. x1v—xrx of Doctor Schlesinger’s Memoir. 


Pliomastodon sellardsi Simpson, 1930 
Figures 104, 105 

Brewster, Florida; Lower Pliocene, Bone Valley formation. 

The type of Pliomastodon sellardsi was first described (1916) 
by Dr. E. H. Sellards, at that time State Geologist of Florida, as 
referable to Mammut progenium? (see Sellards, 1916, p. 95, Pl. x). 
The author pointed out, however, its close resemblance to Mastodon 
americanus but emphasized the longer symphysis. The specimen 
was discovered in the mine of the American Cyanamid Company at 
Brewster and was sent to Doctor Sellards by the late Anton 
Schneider, then superintendent of the company. The right half 
of the mandible was returned to Mr. Schneider at his request and 
has since been mislaid, but the left half of the mandible is available 
at the Florida Geological Survey. Doctor Simpson in connection 
with his description gives three views of the specimen, the missing 
right mandible, internal and external aspects, also superior view of 
both sides, which are reproduced as figures 104 and 105 of the 
present Memoir. 

Pliomastodon  sellardsi Simpson, 1930. ‘‘Tertiary Land 
Mammals of Florida,’ Bull. Amer. Mus. Nat. Hist., LIX, Art. 
III, pp. 203-206. Typr.—‘Left lower jaw with Mp3. 





ES Vesee72 

Fig. 104. Pliomastodon sellardsi Simpson, type (Fla. Geol. Surv. V3822), 
from Brewster, Florida. Lower jaw with right and left Mo-3, crown view. 
One-fifth natural size. After Simpson, 1930, fig. 30. Compare figure 87, M. 



Right lower jaw of same individual known 
from photographs, present location un- 

known. Presented by Anton Schneider.”’ 
Fla. Geol. Surv. V3822 (old number 
- 6160). Horizon AND LocaLiry.— 

“Lower Pliocene, Bone Valley Formation, 
pit of American Cyanamid Company, 
Brewster, Florida.” Type Figure.— 
Op. cit., fig. 30, p. 204, fig. 31, p. 205. 

Tyrpr Description.—(Simpson, 1930, 
p. 203): ‘“‘Symphysis elongate, stout lower 
tusks, alveoli about 50 mm. in diameter near 
posterior end. Mefully trilophodont. Fifth 
crest of Ms; less developed than in Mastodon 
americanus, ridge-like, more distinct internal 
cusp and four closely appressed cuspules 
external to it. Cusps more elevated than in 
Miomastodon merriami, less than [in] Masto- 
don americanus, about as in Pliomastodon 
matthewi. Inner side of M; strongly convex, 
outer border straight. First loph distinctly 
narrower than second, and last two lophs 
also relatively narrower than in Mastodon.” 

My, Length 110 mm. 
Width 80 
M; Length 164 

Width (max.), second loph 95 

Height internal cone of first crest—actual 55 mm., estimated 
57 mm. 

Height internal cone of second crest—actual 56 mm., esti- 
mated 59 mm. 

Osborn, 1933: Pliomastodon sellardsi, according to the above 
descriptions of Sellards (1916) and of Simpson (1930), is a most 
welcome addition to the American ancestry of Mastodon ameri- 
canus; it is distinguished by its longer rostrum, with ridge-crests 
intermediate between Miomastodon merriami and Mastodon 

To the same species may be referred a right mandibular ramus 
(Fla. Geol. Surv., V-5376, cast Amer. Mus. 26979), with Ms, Ms; 
measurements of M; 18198 mm., index 54, found at a locality 
not very far from that of the type. 

Pliomastodon vexillarius Matthew, 1930 
Figures 106-108 
From southeast of Coalinga, Fresno County, California. 
goin [=late Pliocene]. Plesippus proversus life zone. 

] Faunal Level. (Stirton, August 8, 1932): ‘The type of Pliomastodon 
vevillarius was found in the upper portion of the Etchegoin section in the 
North Coalinga region. This section has been worked by Ralph Stewart 
(U. 8. Geol. Surv.) who collected isolated teeth of Plesippus proversus and 
Castor californicus as his index fossils. Late Pliocene.” 

The Standard Oil Company of California, in excavating on its 
properties at the north end of the Kettleman Hills anticline, south- 
east of Coalinga, California, discovered the specimen described 

Upper Etche- 




Fig. 105. Pliomastodon sellardsi Simpson. Missing part of type. Right lower jaw with Mo2-3, internal 
and external views. One-fifth natural size. After Simpson, 1930, fig. 31. 

by Dr. W. D. Matthew as Pliomastodon vexillarius, the specific name 
signifying “‘standard bearer,” chosen ‘‘in recognition of the very 
important contributions to paleontology made by the Standard 
Oil Company of California in the course of its operations and re- 
searches, and the care which the officers of the company have 
taken to preserve and turn over to scientific museums the collec- 
tions made by their parties and to facilitate in many ways field 
studies and collecting work of this [University of California] and 
other institutions.” Doctor Matthew states (p. 338) that ‘‘ What- 
ever be the final determination as to nomenclature and taxonomic 
status, the skull represents a type of American mastodon hitherto 
known only from teeth and jaw fragments. It is distinguished 
from the long-jawed species by the short jaws and upward curving 
enamelless tusks, from the mirificus group by simple molars with 
little trefoil development, from the ‘American Mastodon’ by the 
shortened skull base with overhanging occiput and the marked 
approach to anteroposterior succession of teeth.” 

Pliomastodon vexillarius Matthew, 1930. ‘‘A Pliocene Masto- 
don Skull from California Pliomastodon vexillarius, N. Sp.,” 
Bull. Dept. Geol. Univ. Calif., XIX, No. 16, pp. 335-348, Pls. 
XLI-XLIv, text. figs. 1, 2. Typr.—Back of skull and 
palatal region approximately complete, zygomata broken but 
parts of them preserved, frontonasal and orbital regions partly 
preserved. Dentigerous portions of both sides of lower jaws com- 
plete, but of the symphysis, angles, and coronoid, little is left. 
Atlas complete, one femur, an astragalus, patella, and parts of 
other limb bones. Crowns of teeth largely missing; most of M?, 


right and left, and small parts of M’, posterior halves of Ms, 
right and left, preserved, but only the bases of the crowns of the 
remainder of the second and third molars. First molars had been 
lost during life and their alveoli partly closed; second molars well 
worn, third molars unworn in the posterior half, probably lightly 
worn in front. Original in the Museum of Paleontology, Uni- 
versity of California (No. 28301). Horizon AND Locauitry.— 
North end of the Kettleman Hills anticline, southeast of Coalinga, 
Fresno County, California. Upper Etchegoin [=late Pliocene]. 
Tyrer Figure.—Op. cit., Pls., text figs. 1 and 2. 
Grotocic Acre.—(Matthew, op. cit., p. 338): ‘The type and 
only known specimen of the genus is a last upper molar from the 
Snake Creek beds (referred by Matthew to Zygolophodon in 1918 
{made by Osborn in 1921 the type of Mastodon (Pliomastodon) 


of Vacek’s genus or as a distinct genus, according to the relative 
weight given to one or another of the progressive characters.” 
p. 336). Upper tusks large, cylindrical, close together and sub- 
parallel at base, curving gently upward and outward, a narrow 
strip of thin enamel showing at the base on the outer side but no 
clear evidence of enamel beyond the alveolus. Lower tusks prob- 
ably absent. Molar teeth wide, moderately crested, less so than in 
Mastodon, trefoils single, not prominent, valleys open, M 2 tri- 
lophodont, M 3 with 4% crests. The basicranial region of the skull 
is little elevated above palatal plane, much shortened as compared 
with Mastodon, occiput very low and wide, heavily overhanging 
backward instead of nearly vertical as in Mastodon and other 
genera. Posterior nares crowded forward so as to be opposite the 
back of M*; in Mastodon they are considerably behind it. The 
entire basicranial region is thus crowded forward along with the 
condyles, which are scarcely over half as far behind the teeth as in 
Mastodon. This forward crowding is paralleled in some of the 
other mastodontines and in the elephants, but mostly with a much 

Fig. 106. Type skull and jaws of Pliomastodon vexillarius Matthew (Mus. Univ. Calif. 28301). After drawings and photographs kindly furnished the 

present author by the University of California. Compare Matthew, 1930. 

(Left) Palatal and side view of type skull, one-fifteenth natural size. Compare Matthew, 1930, text figs. 1 and 2. 

(Right) Same type skull. Compare Matthew, 1930, Pl. xxi. 

matthewi|) and owing to the crowns of the upper teeth being broken 
off in our skull, exact comparison is impossible. It agrees, however, 
in geological age, and the fauna of the Lower Etchegoin is largely 
composed of the same genera as the Snake Creek fauna, so that 
provisionally I refer the skull to Pliomastodon. As at present 
understood, it represents an advanced stage of the Zygolophodon 
phylum and might be regarded as a progressive species or subgenus 

One-fifteenth natural size. 

higher elevation, especially in the latter group, of the basicranial 
above the palatal plane. The author states that the skull ap- 
parently cannot be placed in any of the better known Mastodon- 
tine genera, and gives its distinguishing characters from Gompho- 
therium, ‘ Serridentinus,’’ Mastodon, Cuvieronius, Anancus, Cordil- 
lerion, Tetralophodon, Pentalophodon, Rhynchotherium, and Choe- 


Speciric CHaractrers.—(Op. cit., Matthew, p. 339): ‘‘ Last 
upper molar with four full crests and a considerable heel behind 
the fourth; in P. matthewi type there are three full crests with the 
fourth half-formed and a small rudiment of the fifth. Comparative 
measurements of this tooth are: 

[Pliomastodon| [Cordillerion'| 
verillarius matthewi — bensonensis 
Antero-posterior length 158 144 191 
Width at anterior end 80 80 99 
Width at fourth crest 71 48 

Fig. 107. Pliomastodon vezillarius, type. Lower jaws, outside and 
occlusal views. One-fifth natural size. After Matthew, 1930, Pl. xxi. 


Other measurements of teeth and skull— 
P. vexillarius |C.) bensonensis 

M?, dimensions 107 X71 125 88.5 
Diameter of tusk (about a foot be- 

yond alveolus) 165 130 
Width of palate between M' alveoli= 155 120 
Width of palate between M® alveoli = 88 76” 

“The skull compares in bulk with the largest specimens of the 
American Mastodon. ... From ‘Tetralophodon’ edensis this 
species is separated by upcurved tusks lacking enamel band. . . . 
The skeletal parts preserved do not show any remarkable features.”’ 

Osborn, 1933: This interesting communication by Matthew, 
published posthumously, positively establishes Pliomastodon as an 
ancestral stage of Mastodon clearly distinguished by a far more 
primitive cranium, by a “narrow strip of thin enamel” within the 
alveolar base of the tusk, by the probable absence of inferior tusks, 
by the approximation of the condyles to the superior molars (M°), 
by the extreme backward recession of the anterior nares and the 
shortening of the lower jaw, probably also with an unusual width 
of the occipital crest, suggesting an exceptionally large develop- 
ment of the proboscis. 

As to bodily proportions, the type femur (op. cit., Pl. xtrv— 
Fig. 108) is much more massive than that of the Alaskan mammoth 
(Mammonteus primigenius) but indicating a massive body about 
2800 mm., or 9 ft. 24 in., in height at the shoulder. The cranium 
apparently debars this species from direct ancestry of Mastodon 
americanus, for it embraces a number of peculiar characters. It is 
nevertheless most welcome as exhibiting a very distinctive, low- 
browed, flat-crested, abbreviated postcranial region, a very broad- 
crested occipital region, and broadly divergent premaxillary region 
with sharply upturned tusks. These features are clearly displayed 
in the outline and photographs (Fig. 106) after Matthew, 1930. 
An outline restoration of this animal, to the same scale as the 
Warren specimen of M. americanus, is shown in figure 110. 

Fig. 108. Type femur of Pliomastodon vezxillarius Matthew, front and 
side views. After photograph kindly forwarded to the present author by the 
University of California (cf. Matthew, 1930, Pl. xuiv). About one-sixteenth 
natural size. 

'Gidley’s type of ‘Anancus’ bensonensis is now referred to Cordillerion bensonensis (see p. 565 below). 

One-fiftieth natural size 

This restoration is based largely upon a record specimen (Fig. 114) of the American Mastodon in the Geological Museum of the Ohio State University, 
namely, with an estimated shoulder height in the flesh of 10 ft. 2 in.; it also embodies certain characters of the Warren Mastodon. The background is 
taken directly from a snow scene in the Hudson Highlands, near Newburgh, New York, where the Warren Mastodon was found. 


Cu ef 






One-hundredth natural size 
Mastodon americanus, Mus. Ohio State Univ., adult bull of maximum size, Turicius turicensis, restored after type specimen of Schinz, in the Zurich 
restored after skeleton (Fig. 114). Collection (Fig. 162). 
Zygolophodon borsoni, restored after grinding teeth from Asti (Fig. 154) 
Pliomastodon vezillarius, Mus. Univ. Calif. 28301, restored aftertype 0d superior and inferior tusks after Schlesinger (p. 211 below). 
cranium and tusks, also femur, as described and measured by Matthew (pp. Palzomastodon beadnelli, as restored by Osborn (Fig. 97) from jaw, palate, 
161-163 above). and skeleton described and figured by Andrews, 1906 (Fig. 96). 
(Osborn, 1934) Palzomastodon is now removed from its supposed ancestral 
Miomastodon merriami, restored after type specimen from Nevada and _ relationship to Mastodon. As here restored, also in fig. 97, p. 149, the mouth 
referred specimen from Colorado (pp. 154-156 above). parts and proboscis may be incorrectly drawn. 



NoMENCLATURE.—The full history of this classic name is related in Chapter I, ‘Discovery of the Living and 
Extinct Proboscideans,” in Chapter V, ‘‘History of the Classification of the Mastodontoidea, Families and 
Subfamilies (1705-1927),”’ and again in the introduction of the present chapter (Chapter VI), ‘‘History of the 
Subfamily Mastodontine, the True Mastodonts.” 

The first full summary of the vast generic and specific synonymy was that of Joseph Leidy in his great Memoir 
of 1869, p. 392. In our more recent and searching revision of the nomenclature of the Proboscidea we have found 
only one additional quaint and abortive name, Mastotherium megalodon Fischer de Waldheim, but doubtless 
further bibliographic research will in the future reveal still others. 

Elephas americanus Kerr, 1792, Big-Bone Lick, Kentucky, Tetracaulodon Collinsii Hays, 1834. 

near the Ohio River. Mastodon Ohioticum 1832, Neues Jahrb. f. Min., p. 355, 
Ohio-Incognitum Blumenbach, 1797, same (?) locality. Gervais, 1848-1852. 
Mammut ohioticum Blumenbach, 1799, same locality. Tetracaulodon Godmani Hays, 1834. 
Elephas macrocephalus A. Camper, 1802, same (?) locality. Missourium kochii Koch, 1840, Jefferson Co., Missouri. 
Le Grand Mastodonte Cuvier, 1806, same locality. Leviathan Missourii Koch, 1841, Missouri. 
Mastodonte de V Ohio, Cuvier, 1806, same locality. Tetracaulodon Tapyroides Koch, 1841, Missouri. 

Harpagmotherium canadense Fischer de Waldheim, 1808, Tetracaulodon Osagii Koch, 1841, Missouri. 
Ohio (?) River (ef. Sherborn, 1924, p. 1022, ‘““Harpagono- Tetracaulodon kochii, Koch, 1842, Missouri. 
therium canadense .. . Anim. foss. Siberie.’’). Tetracaulodon Haysii Grant,1842. 

Mastotherium megalodon Fischer de Waldheim, 1814, Ohio. Tetracaulodon Bucklandi Grant, 1842. 

Mastodon giganteum Cuvier, 1817, Big-Bone Lick, Kentucky, Missowrium Theristocaulodon Koch, 1843. 

Ohio River. Mastodon rugatum Koch, 1845. 

Mastodon maximus Cuvier, 1824, Big-Bone Lick, Kentucky, Elephas Ohioticus de Blainville, 1839-1864. 
Ohio River. Mastodon Ohioticus Fale. and Caut., 1845. 

Tetracaulodon Mastodontoideum Godman, 1830, near New- Elephas Rupertianus Richardson, 1854, Swan River, Lake 
burgh, Orange Co., New York. Winnipeg basin, Canada. 

Mastodon Cuviert Hays, 1834. Trilophodon ohioticus Faleoner, 1868. 

Mastodon Jeffersoni Hays, 1834. Mastodon americanus Leidy, 1868. 

To our present knowledge the above names are chiefly synonyms of Elephas [Mastodon] americanus Kerr, but 
it is probable that some of these names were applied to specific or subspecific stages distinct from the type speci- 
men Kerr had in mind, which came from the Big-Bone Lick of Kentucky. We may at present consider only the 
names of Kerr and of Blumenbach, and of the species defined subsequent to Leidy’s Memoir of 1869, which are as 

Elephas americanus Kerr, 1792. Mastodon moodiei Barbour, 1931. 
Ohio-Incognitum Blumenbach, 1797. Mastodon raki Frick, 1933. 

Mammut ohioticum Blumenhach, 1799. Mastodon americanus alaskensis Frick, 1933. 
Mastodon rugosidens Leidy, 1890. Mastodon grangeri Barbour, 1934. 

Mammut progenium Hay, 1914. Mastodon pavlowi sp. nov. 

Mastodon americanus plicatus Osborn, 1926. Mastodon acutidens sp. nov. 

1778, AND To CuviER, 1806-1836! 

Cuvier (1821, p. 250) states that the first engraving of “une grande molaire de |’Ohiv”’ is that of Guettard 
(Hist. Acad. Roy. Sci. avee Mémoires, Paris, 1752 [1756], p. 360, Pl. m1, fig. 1 [Pls. 11, 12 of Memoirs]), but that 
the teeth of this animal did not attain real celebrity in Europe until between 1760 and 1770 through the 
Memoirs of Collinson (Phil. Trans., London, LVII, Pt. II, 1768, p. 468) and of Hunter (Phil. Trans., London 
LVIII, 1769, p. 34). 

‘Continued from Chapter V, pp. 118-129 of the present Memoir. 


The name Ohio, moreover, is historic, since as early as 1765, as we learn from George Croghan’s Journal and 
from Buffon (Buffon, 1778, p. 505): 

dans les contrées voisines de la riviére d’Ohio, environ a 4 milles sud-est de cette riviére, éloignée de 640 milles du fort 
de Quesne, (que nous appelons maintenant Pitsburgh) . . . a vu, aux environs d’un grand marais salé, ot les animaux sauvages 
s’assemblent en certains temps de l’année, de grands os & de grosses dents, & qu’ayant examiné cette place avec soin, il a décou- 
vert, sur un bane élevé du cété du marais, un nombre prodigieux d’os de trés-grands animaux, & que par la longueur & la forme 
de ces os & de ces défenses, on doit conclure que ce sont des os d’éléphans. 

Extracts from Croghan’s Journal, communicated to Benjamin Franklin, appear in Buffon’s Supplement (op. 
cit., p. 507) accompanied by six admirable life-size steel engravings of mastodont grinding teeth, also citations 
from two little Memoirs by Collinson read before the Royal Society of London, in which Collinson remarks 
(Buffon, op. cit., p. 509): 

Le marais salé ot !’on a trouvé les os d’éléphans, n’est qu’é quatre milles de distance des bords de la riviére d’Ohio, . . Les 
ossemens d’éléphans se trouvent sous une espéce de levée ou plutdét sous la rive qui entoure & surmonte le marais 4 cing ou six 
pieds de hauteur; on y voit un trés-grand nombre d’os & de dents qui ont appartenu A quelques animaux d’une grosseur 
prodigieuse ; 

Burron, 1778. Superior GrinDING TreetH (A, Al) oF ZYGOLOPHODON AND (B) oF Mastopon. ORIGINALS SAID TO BE IN THE Parts Museum 
(FIDE Paviow, 1894) 

Fig. 111. Grinding teeth of Zygolophodon borsoni referred (A, Al) and of Mastodon americanus referred (B), after Buffon, 1778, Pls. 1, u, and ry, one- 
fourth natural size. [Inverted by H. F. O.] 

A, Al, Buffon’s figures of the molar of an animal which we now know to be related to Zygolophodon borsoni, found in Russia (“‘la petite Tartarie”), pre- 
sented to Buffon in 1770 by M. le Comte de Vergennes. Observe three to four lobes, i.e., cones or conelets, in each transverse crest. (1778, Pls. rand 11.) 

B, Buffon’s figure of the molar of an animal now known as Mastodon americanus, from Big-Bone Lick, Kentucky, near the Ohio River, sent to Buffon 
by M. Collinson. Observe two lobes, i.e., cones, in each transverse crest, the true Mastodon type. (1778, Pl. rv.) 

Both of these beautifully engraved molar teeth appear to be third superior grinders, M*, although they lack the rudimentary fifth crest or pentaloph 
characteristic of the third superior molars of Mastodon americanus (Fig. 133). 

To Collinson’s characterization, Buffon (op. cit., p. 510) adds: 

Ce que dit ici M. Collinson, est trés-vrai; ces grosses dents molaires différent absolument des dents macheliéres de 
l’éléphant, & en les comparant a celles de l’hippopotame, auxquelles ces grosses dents ressemblent par leur forme quarrée, on 
verra qu’elles en différent aussi par leur grosseur, étant deux, trois & quatre fois plus volumineuses que les plus grosses dents 
des anciens hippopotames trouvées de méme en Sibérie & au Canada, quoique ces dents soient elles-mémes trois ou quatre 
fois plus grosses que celles des hippopotames actuellement existans. 

Buffon (op. cit., p. 511) rightly compared the mastodont and hippopotamus teeth and rightly concluded: 

. comme on peut le voir en comparant les figures des planches 1, 111 & Iv, avec cellesde la planche vy. Il paroit done certain 
que ces grosses dents n’ont jamais appartenu 4 |’éléphant nia l’hippopotame; . . . je crois done pouvoir prononcer avec fonde- 
ment que cette trés-grande espéce d’animal est perdue. 

In brief, Buffon fully characterized this animal as a mastodont and designated it as an extinct species belong- 
ing to the epoch of the elephants but did not name it. This is sufficient evidence that the name Ohio was very 
prominent in the minds of the naturalists of the period, because the writings of Buffon dominated Europe at the 


Fig. 112. Cuvier’s types of Le Grand Mastodonte, or Mastodonte de l’Ohio, 1806. After Cuvier, 1806.2, Pl. 49 [1], one-half 
natural size. 

(Cuvier, 1836, Atlas, Pl. 19): Fig. 2. Molaire a dix pointes encore intactes, donnée par le duc de Plaisance. Fig. 1. Dent 
i dix pointes en partie usée, du cabinet de Joubert, vue de profil. Fig. 3. Le méme, vue par sa couronne. Fig. 4. Dent a dix 
pointes, rétrécie en arriére, communiquée par feu M. Tonnellier. Fig. 5. Molaire a six pointes A demi usée, d’aprés un dessin de 
M. Blumenbach. 

Cuvier, 1806-1836.—Cuvier practically defined the genus ‘Mastodonte’ in his admirably engraved Planches 
and in the descriptive legends of these plates. In the descriptions he acknowledges his indebtedness for one of the 
drawings to Blumenbach and for one or two of the cotype specimens to Fabri, to Daubenton, to Joubert, and to 
Tonnellier. Referring to Buffon’s idea that the mastodont grinders were those of the “hippopotames gigan- 
tesques,”’ he pointed out that they were distinguished by the worn ‘losanges’ which differed greatly from the worn 
“tréfles de l’hippopotame.”’ 

(Cuvier, 1806.2, p. 293): ‘Notre pl. 1 représente quatre de ces dents de mastodonte A moitié grandeur, Fig. 5 en est une a 
six pointes 4 demi-usées: elle est copiée d’aprés un dessin qu’a bien voulu m’envoyer M. Blumenbach. Nous en avons au Muséum 
trois pareilles, anciennement rapportées par Fabri. Ce sont elles que Daubenton (Hist. nat. XII, n.° 1106, 1107, 1108), et Buffon 
(Epoques de la nature, pl. v) ont prises pour des dents d’hippopotames gigantesques. Elles sont aisées A distinguer par ces losanges, 
dont notre figure donne une idée fort juste, et qui différent beaucoup des tréfles de ’hippopotame. D’ailleurs ’hippopotame n’a 
jamais que quatre tréfles et non pas six.” Compare Daubenton, 1764, p. 77, No. MCXI “ Dent pétrifiée qui a quelque rapport a 
celles de U'hippopotame.”’ 


Succeeding Buffon’s description (1778) of the American Mastodon but anticipating Cuvier’s description of 
Le Grand Mastodonte, 1806, the Scottish naturalist Robert Kerr proposed the specific name Elephas americanus 
for tusks and grinders found in the ‘Big-bone-swamp’ on the banks of the Ohio. He pointed out that these teeth 
were specifically different from those of the elephant and furnished with a double row of high conic processes 
resembling those of carnivorous animals. This contribution appears in a translation of portions of at that time 
a recent edition of the “Systema Nature” of Linneus, with improvements by Professor Gmelin, entitled, “The 
Animal Kingdom or Zoological System, of the Celebrated Sir Charles Linnzeus,” published in 1792, as follows: 

(Kerr, 1792, p. 116): 2. American Elephant.—Hlephas americanus. In America, on the banks of the Ohio, are found, 
several feet below the surface, in a marshy place called Big-bone-swamp, great numbers of tusks and grinders, supposed by many 
to belong to the Elephant: But the grinders are totally different, being covered uniformly with enamel, and furnished with a 
double row of high conic processes, like those of carnivorous animals; whereas those of the Elephant are composed of alternate 
perpendicular layers of bone and enamel, and are ribbed transversely on their upper surfaces, like those of graminivorous quad- 
rupeds: Hence the species must be entirely different; and Mr. Pennant has chosen to suppose that they have belonged to an 
unknown species of this genus, which he names the American Elephant. Hist. of Quad. n. 71. 


The next name applied to the animal was that of Johann Friedrich Blumenbach, pioneer vertebrate pale- 
ontologist of Germany, who received from the British Museum his type specimen (Fig. 113), a third superior 
molar tooth of the left side. Blumenbach subsequently corresponded and exchanged drawings and notes with 
Cuvier, as mentioned above. 

(Blumenbach, 1797.2, No. 19): Bey jenem fossilen Ungeheuer der priadamitischen 
Vorwelt sind die zackichten Kronen ganz mit einer starken Lage vom so genannten 
Schmelz (substantia dentium vitrea) iiberzogen: da hingegen die Backenzihne beider 
Gattungen des Elephantengeschlechts aus vertical liegenden abwechselnden Schichten 
von Schmelz und Knochensubstanz bestehen. . . . Der vom fossilen Incognitum 
hingegen nach einem ungeheuern am Ohio ausgegrabenen Exemplar, das mir aus 
dem Britischen Museum fiir meine Sammlung iiberlassen worden. 

Osborn, 1922: Ohio-Incognitum is the name and figure cited by de 
Blainville (1839-1864, p. 245). We observe that Blumenbach’s figure of 
Ohio-Incognitum (1797. 2, No. 19) is of a third superior molar of the left side, 

Fig. 113. A. Type left third superior molar with very pronounced internal cingulum. There are four and a third lophs; 
of Ohio-Incognitum Blumenbach, 1797. Repro- ‘ i 
duced in facsimile after Blumenbach, 1797.2, there are two mamillate lobes on each crest in the figure. The above 
No. 19. Same figure in edition of 1810 desig- qeseription by Blumenbach (op. cit., No. 19) shows that he secured this speci- 

nated on the Plate as Ohzo-Incognitum. Techni- = P ‘ A , 
cally a type figure. [=Mastodon americanus.] men from the British Museum as part of its Ohio River collection. 


Speciric Name.—Leidy concludes his full revision of the American species of Mastodon, in his great Memoir 
of 1869, with the sentence (Leidy, 1869, p. 240): ‘One of the species, the well known Mastodon ohioticus, or M. 
americanus as I shall hereafter call it, appears to have roamed throughout the continent during the quaternary 
period.” The dates of these species are Hlephas americanus Kerr, 1792, and Mammut ohioticum Blumen- 
bach, 1799; to the best of our knowledge Blumenbach (1799) was the first to use the name Mammut 


ohioticum; in the Neues Jahrbuch, 1832, p. 355 (see Eichwald), the term Mastodon ohioticum is used, and Leidy 
states in his Memoir of 1869, p. 394, that Gervais used this term ‘ohioticum’ in his ‘‘Zoologie et Paléontologie 
Francaises,” 1848-1852, I, p. 187. The name Mastodon americanus was first used by Leidy in 1868, in Volume 
XX of the Proceedings of the Academy of Natural Sciences of Philadelphia, p. 175, to embrace the Mastodon 
ohioticus or M. giganteus of authors. 

GENERIC NAME.—The generic name Mastodon was universally accepted by paleontologists in all parts of 
the world until 1902, when the vernacular generic term Mammut of Blumenbach (1799) was revived by Hay, 
although subsequent to the vernacular term Ohio-Incognitum of Blumenbach. Unfortunately this vernacular 
synonym ‘Mammut’ entered the American literature between 1902 and 1925 and occurs in the writings of Palmer, 
Lull, Barbour, Stock, Daggett, and others. In Chapter I of the present Memoir, ‘‘Discovery of the Living and 
Extinet Proboscideans,”’ 
Subfamily Mastodontine, the True Mastodonts,’”’ ample reasons are given for the banishment from all future 
proboscidean literature of this barbaric term ‘Mammut.’ 

and in the foregoing pages of the present chapter (Chap. VI), “History of the 

Subsequent to Leidy’s Memoir of 1869 nine species have been proposed, namely, Mastodon rugosidens Leidy, 
1890, ‘Mammut’ progenium Hay, 1914, Mastodon americanus plicatus Osborn, 1926, Mastodon moodiei Barbour, 
1931, Mastodon raki Frick, 1933, and Mastodon americanus alaskensis Frick, 1933. The validity of Leidy’s M. 
rugosidens, 1890, and of Osborn’s M. americanus plicatus, 1926, is a question of further research. Mastodon 
pavlowi, also Mastodon acutidens, a very progressive stage, are proposed by Osborn in the present Memoir (see 
Appendix). In 1934 Barbour described the straight-tusked species Mastodon grangeri from Nebraska (see 
Appendix at close of Volume I for description and figure). 


— "Opie State Univ. 

Tre Larcest Furry Aputr Mate SKELETON oF REcorD; Estimatep SHouLtper Hereut 10 Ferr 2 INcuEs IN THE FLEsH. (SEE Fre. 109) 
Fig. 114. Mounted skeleton of Mastodon americanus from Catawba, Ohio, in the Ohio State University, Geological Museum, after photograph kindly 
sent the present author by Prof. J. Ernest Carman, Chairman of the Department of Geology. Reproduced about one twenty-fourth natural size. 


Famity: MASTODONTID Girard, 1852 

SuBFAMILY: MASTODONTIN Brandt, 1869—Osborn, 1910 

Genus: MASTODON Cuvier, 1806-1817 
Original reference: Mastodonte, “Sur Le Grand Mastodonte,’’ Cuvier, 1806.2, pp. 270, 272, 293; Mastodon, “Te Régne Animal,” 

Cuvier, 1817, pp. 232, 233. 

Genotypic species: Mastodon giganteum. 
Synonyms: Mastotherium Fischer de Waldheim, 1814, pp. 337-341. 

Generic CHaracters.—Incisive tusks without enamel; superior tusks large and rounded with 
slightly indicated annular ring growths. Inferior tusks straight, cylindrical, variable in old age. 
Marked sexual disparity in female tusks. Permanent premolars suppressed,’ except the vestigial P*-P,. 

Dental formula; [s=3:3 Dp3=#+ Pt (vestigial) Mi-3% 

Intermediate molars, Dp 4-M 2, trilophodont, i.e., with three ridge-crests. Ridge-crests progressively 
elevated, lophodont to subhypsodont. Cingulum of grinding teeth slightly stronger on the inner side 
than on the outer side (in Serridentinus much stronger on the inner side). Grinders bilobate with 
strong median sulcus between the inner and outer lobes. Very rudimentary trefoil spurs on the superior 
inner lobes and on the inferior outer lobes (compare Fig. 133). Summits of lobes simple or crowned with 
small conelets. Progressive elevation, subhypsodonty, of the lobes (ef. Figs. 112, 135). Progressive pli- 
cation of the surface enamel of the grinding teeth (Fig. 120). Progressive development of the third crest 
(tritoloph and -lophid) in intermediate molars; progressive development of the fifth crest rudiment 
(pentalophid) in third inferior molars. Progressive reduction of the rostrum of the lower jaw (see 

M. progenius and M. americanus). 

The above list of generic characters enables us to distinguish clearly the Pleistocene species of the true 

American Mastodon from their Pliocene and Miocene ancestors Pliomastodon and Miomastodon. 

Whereas M?- 

M, show a rudiment of the fifth crest in Mastodon americanus (Figs. 116, 117), no such rudiment is observed in 
Miomastodon merriami of the Middle Miocene. These progressive characters of the grinding teeth in the Masto- 
dontine phylum are well displayed in figure 98, also in Pl. 1, pp. 184-135. 

Mastodon americanus Kerr, 1792 
Figures 76, 112, 114, 116, 117, 124-126, 128-130, 1382-134, 136, Pl. 1, p. 134 

Big-Bone Lick, Boone County, Kentucky, late Pleistocene, IV Glacial = 
Wisconsin time. 

The history of this classic species is fully recited above in the 
present chapter of this Memoir. Cuvier’s types of the genus 
‘Mastodonte’ (= Mastodon) are reproduced above in figure 112. 
Unfortunately Kerr’s type of the species Elephas [= Mastodon] 
americanus was not figured. 

Tyre Description.—Kerr’s type description is cited in full 
above (p. 168) and need not be repeated here. The type locality, 
however, deserves very careful description. Mastodon americanus 
from Big-Bone Lick, Kentucky, as figured by Cuvier (see Fig. 112 
of the present Memoir) probably belongs to IV Glacial = Wiscon- 
sin time; the molars are much more primitive than those of M. 
acutidens sp. nov. 


locality both of the genus ‘Mastodonte’ (= Mastodon) Cuvier and 
of the species Elephas [= Mastodon] americanus Kerr. This famous 
salt swamp or salt lick was probably visited by the ungu- 
lates for a long period of geologic time, as shown by the latest 
analysis of its contained fauna (see Hay, 1923.1). In the list of 
species by Hay (p. 403) cited below references are made in his 
Memoir on ‘‘The Pleistocene of North America and Its Verte- 
brated Animals” to pages whereon further information is given 
regarding the respective species. 

Hay believes (op. cit., 1923, p. 403) that the Big-Bone Lick 
fauna followed Glaciation III (=Illinoian) and belongs chiefly in 
3rd Interglacial time (=Sangamon interval); he accordingly 
assumes that the animals were buried during the Sangamon stage 
in the interval between the Illinoian (III) and the Wisconsin (IV) 
glaciations; the presence of Mammonteus primigenius in the Big- 
Bone Lick deposits rests (op. cit., p. 146) on the determination of a 
single superior molar tooth with 23-24 plates, which Osborn con- 
siders may be referable to Parelephas jeffersonii. 

‘See page 138 for observations of Hays and of Warren, namely, that P 4 forms in the jaw but does not erupt; it is therefore vestigial. 



Megalonyx jeffersonii (p. 44). 
Mylodon harlani (p. 44). 

Equus complicatus (p. 202). 
?Tapirus haysii (p. 209). 
Odocoileus virginianus (p. 234). 
Cervus canadensis (p. 243). 
Cervalces scotti. 

Alces americanus. 

Rangifer caribou (p. 247). 

Mastodon americanus rugosidens Leidy, 1890 
Figure 115 

Santee Beds of Beaufort County, South Carolina; Pleistocene. 

Mastodon rugosidens Leidy, 1890. ‘Mastodon and Capybara 
of South Carolina.” Proc. Acad. Nat. Sci. Phila., Vol. XLII, 1890, 
p. 184. Typr.— .. . “complete crown of a last upper [lower| 
molar, strikingly different from that tooth in the common American 
Mastodon . . . looking asif carved out of ebony, as is the case with 
many of the fossils from the same and similar localities.”’ TYPE 

Locauiry.—‘‘It was found in the Santee Beds of Beaufort Co., 8. 

M. rugosidens 
Drawn from 14445 Cast of type 

(outer view) Al 

Tyee or MAstopon 

Fig. 115. Last inferior 
molar of the left side, 1.M3. 
Type of Mastodon rugosi- 
dens Leidy (Philadelphia 
Academy). Drawn from 
east of type (Amer. Mus. 
14445), one-third natural 

A, Crown view; Al, 
external view; A2, inter- 
nal view. 

1/3 nat. size 


C.” and ‘was presented to the Academy by Mr. James R. 
McKee.” Type Figure.—Drawn from cast of type (Amer. 
Mus. 14445—Fig. 115 of the present Memoir). 

Typr Description.—(Leidy, 1890.2, p. 184): “It is worn only 
on the summits of the anterior pair of lobes, which display the 
usual exposed dentinal areas. Notwithstanding the many species 
of Mastodon which have been recorded in North and South 
America, the present tooth seems to indicate a different one. It 
more nearly resembles the corresponding tooth of the W. floridanus, 
recently described, or that of the M. angustidens of Europe, than 
of the M. americanus. In comparison with the molars of these and 
other known species the tooth is remarkable for the greater propor- 


306therium bombifrons (p. 255). 
Symbos cavifrons (p. 255). 
Bison antiquus (p. 265). 

Bison bison (p. 270). 

Mammut americanum (p. 128). 
Elephas primigenius (p. 146). 
Elephas columbi (p. 160). 
Ursus americanus. 

| = Mastodon americanus type 
= Mammonteus primigenius ref.(?) 
= Parelephas jeffersonii ref.(?)| 

tionate length of the constituent lobes of the crown and their 
conspicuously wrinkled condition. The wrinkling is longitudinal 
and regular and apparently not the result of an abnormal state. 
Similar wrinkling is observed in some specimens of the same teeth 
in M. americanus, but mainly confined to the intermediate vallies 
of the crown, while it is well produced laterally in the present 
fossil. From the comparatively more prolonged condition of the 
lobes, the summits of the inner ones appear more tapering or 
narrowly pointed than in M. floridanus; while with the fore and 
aft extensions of the same lobes the summits form acute and not 

Fig. 116. Type left third superior molar, |.M®, 
of Ohio-Incognitum Blumenbach, 1797. Repro- 
duced in facsimile after Blumenbach, 1797.2, 
No. 19. Same figure in edition of 1810 desig- 
nated on the Plate as Ohio-Incognitum. {= Mas- 
todon americanus.} 

Fig. 117. 

Inferior molar of the left side, 1.Ma, of 
Mastodon americanus. One-fourth natural size. The 
record of the specimen, from which this beautiful 
drawing was made by Mr. R. Weber, has been lost. 

obtuse angles as in the latter. Regarding the specimen as indicat- 
ing a previously unknown species, this may be distinguished as the 

Comparative measurements of the tooth are as follows: 

M. rugosidens M. floridanus 

Length of crown fore and aft 190 mm. 190 mm. 
Breadth of base at fore part 92 mm. 96 mm. 
Length of second inner lobe 90 mm. 80 mm. 
Length of second outer lobe 80 mm. 62 mm.” 


Osborn, 1925: The type cast (Amer. Mus. 14445), from the 
original type in the Academy of Natural Sciences, Philadelphia, 
agrees exactly in measurement with Leidy’s description (Leidy, 
1890.2, p. 184) cited above; thus there is no mistaking the type. 
Leidy erred, however, in describing it as “a last upper molar’’; 
it is certainly a last lower molar of the left side, 1.M3, as shown by 
careful comparison of the type with third inferior grinding teeth 
and with the specimen beautifully displayed in figure 117. 

It is doubtful also whether the wrinkling of the enamel, to which 
the specific name rugosidens applies, is actually of specific value. 


Fig. 118. Type jaw of ‘Mammut’ progenium Hay, 1914, Pl. xurv, figs. 1 
and 2. From Cox gravel pit, Missouri Valley, Harrison County, Iowa. 
Described by the present author as Mastodon progenius. 

Mastodon progenius Hay, 1914 
Figure 118 
Cox gravel pit, Missouri Valley, Harrison County, Iowa, early Pleisto- 
cene, probably 1st Interglacial, Aftonian age, fide Hay. 

This species is of great interest as representing an early Pleisto- 
cene stage in the development of the true Mastodon. Of 1st Inter- 
glacial or Aftonian age it is much more primitive than the type of 
Mastodon americanus of 3rd(?) Interglacial or Sangamon age. 

Mammut progenium Hay, 1914. “The Pleistocene Mammals of 
Iowa.” Iowa Geol. Surv., 1914, XXIII (Ann. Rept., 1912), pp. 
368-373. Typr.—Lower jaw of a very old animal, with last 
molar in each side of jaw, worn down to the roots. Univ. of Iowa 
No. 292. Horizon AND Locaurry.—Apparently found in 
1910, in the Cox gravel pit, at Missouri Valley, Harrison County, 


Iowa; of Aftonian age. Type Ficure.—Hay, op. cit., 1914, 
Pl. xuiv, figs. 1 and 2. ‘It was described and figured by Calvin 
in 1911 (Bull. Geol. Soc. Amer., Vol. XXII, p. 218, Pls. xx and xx1) 
under the name Mastodon americanus. Figures are here presented 
which are made from the same photographs as those used by Calvin 
. . . but reduced in size” (p. 368). 

Type Description.—(Hay, 1914, p. 369) . . . ‘‘symphysis 
of lower jaw longer than in M. americanum; chin less constricted 
at the symphysis, as viewed from above, and not truncated in 
front. . . . The following are the dimensions of this jaw as furnished 
by the measurements of Calvin and the writer [Hay]. 

Length from front of symphysis to hinder border just 

above theangle:<: 2 aq.-tccraoaas cece cee meee 830 mm. 
Length from front of symphysis to rear of condyles, in 

straight dine: 2) a kracsee ean eee eee eee 920 mm. 
Height of coronoid process above lower border of the jaw 425 mm. 
Height of condyles above lower border of jaw.......... 410 mm. 
eng thot, the sympiiysise: see secereett ar eee naeteenee 212 mm. 
Height of the jaw at the front of m.3.................... 192 mm. 
Thickness of the jaw at the front of m.3................ 120 mm. 

Width of the ascending ramus, from rear of the condyle. 290 mm.” 

The symphysis of M. progenium is relatively much longer 
than in M. americanum; its length equals eighty per cent. of the 
width of the ramus. The rami cease to diverge opposite M; and 
there is even some constriction. ‘Occasionally in the lower jaw of 
M. americanum there is a single tusk, rarely two of them. . . in 
the lower jaw of the extremely old animal under description here, 
both tusks were present. Their presence is indicated by the widely 
open sockets. The vertical diameter of the socket measured, the 
right, is 73 mm.; the transverse diameter, 50 mm. These measure- 
ments show that the tusks were far larger than those now and then 
found in M. americanum. The depth of the socket is 160 mm. 
The measurements show likewise that the tusks were considerably 
compressed, the horizontal diameter being about two-thirds of the 
vertical. What the form and the length of these tusks were, 
beyond the sockets, we can only surmise. . . . In the type jaw the 
lingual gutter is somewhat peculiar in being overhung on each side 
by the upper borders of the jaw. These approach until they are 
only 28 mm. apart. On the outer face of this part of the Jaw the 
surface is concave as it rises to the dental border. In this specimen 
the penultimate molar, m.2, had been pushed out on the right side 
of the jaw before the death of the animal; that of the left side 
seems to have been lost after death, for there remains a part of one 
root. As stated, the crown of m.3 is worn down to its base. The 
grinding surface of the left molar forms a concavity which is sur- 
rounded by a ring of enamel; but on the right side a part even of 
this is missing. This attrition of the tooth had so weakened it that, 
before the death of the animal, the teeth had each split into two 
parts. The inner wall of each had broken at the middle of the 
second crest and the cleft had run backward and outward to near 
the hinder end of the tooth. That this had occurred before death 
is evident, as Calvin remarked, from the fact that the edges of the 
fracture had been rounded off. The right tooth has another cleft, 
which crosses its front; but Calvin concluded that this was a post- 
mortem break. It is not improbable, however, that it happened 
under the strain of chewing just before death of the animal. It is 
evident that the hinder molar had four transverse crests and a 
heel which was essentially a crest.’ 


REFERRED SPECIMEN.—Left ramus and symphysis of lower 
jaw, figured by Calvin under the name Mammut americanum 
(Calvin, 1909, p. 352, Pl. xxv, fig. 2), referred provisionally by Hay 
to Mammut progenium because of the resemblances of the jaw to 
the type. The referred specimen is figured by Hay on PI. xvii, 
fig. 1; Pl. un, fig. 2. 

Tyepr or Mastopon 

Fig. 119. Type figures 
of Mammut [= Mastodon} 
oregonense Hay, 1926, figs. 
1, 2, about one-half natural 
size, from Rye Valley, on 
Dixie Creek, Baker County, 
Oregon (Nat. Mus. 4911). A 
second superior molar of the 
left side, 1.M?. The crenulate 
crowns resemble those of 
Mastodon americanus plicatus 
Osborn (Fig. 120), a synonym. 

Mastodon oregonensis Hay, 1926 
Figure 119 

Rye Valley, Dixie Creek, Baker County, Oregon. Probably Pleistocene. 

Compare Mastodon americanus plicatus Osborn, 1926. 

Mammut oregonense Hay, 1926. ‘‘Two New Pleistocene 
Mastodons.” Journ. Washington Acad. Sci., Vol. XVI, No. 2, 
Jan. 19, 1926, pp. 39 to 41. Typr.—A second superior 
molar of the left side, 1.M*?, Nat. Mus. 4911. HorIzON AND 
Locauiry.—This molar was ‘‘found by the Cartwright Brothers, 
placer miners, at Rye Valley, on Dixie Creek, in township 13 
south, range 43 east [Oregon].”” Probably Pleistocene. TYPE 
Figurn.—Op. cit., p. 36, figs. 1 and 2. 

Typr Description.—(Hay, op. cit., 1926, pp. 39 and 40): 
“Tn the U.S. National Museum is a mastodon tooth (Cat. no. 4911) 
which was sent there in November, 1900, by Dr. Waldemar Lind- 
gren, from Baker City, Baker County, Oregon. It had been found 
by the Cartwright Brothers, placer miners, at Rye Valley, on Dixie 
Creek, in township 13 south, range 43 east. Dr. Lindgren reported 
that the tooth had been found in a fluviatile clay bed which had 
formed a part of a bench of auriferous gravels, overlying the 
Payette beds. He regarded the fluviatile clay as of Pliocene age. 
It appears more probable that the bed belonged to the Pleistocene, 
for in it was discovered a tooth of Elephas columbi. . . . The type 
tooth here described and figured is the upper left second molar. 
It has been regarded as belonging to M. americanum, but it is so 
different that the writer ventures to give it a distinct name. The 
tooth had apparently not yet begun to suffer wear; or, if at all, 
only slightly on the first cross-erest. The length is 111 mm.; the 
width of the front end, 74 mm.; of the rear end, 80 mm. The 


crown presents 3 cross-crests and, in the rear, a talon. The crests 
are high, and the valleys narrow. The ends of the cross-crests 
slope steeply and nearly equally. The summits of the two princi- 
pal cones of each crest are well separated, as follows: First crest, 
38 mm., second, 40 mm., third, 40 mm. . . . All around the tooth 
is a heavy cingulum composed, at the pretrite ends of the valleys, 
of 5 or 6 tooth-like conules. On the posttrite side the conules are 
smaller and more numerous.” 

Mastodon americanus plicatus Osborn, 1926 
Figure 120 

Geologic age uncertain, possibly of Postglacial (IV) or post-Wisconsin age. 
From Walnut, Bureau County, Illinois. 

The type of this species is a superior true molar series of the 
right side, r.M*, from Walnut, Illinois, in the American Museum 
collection (Amer. Mus. 10666). 

Mastodon americanus plicatus Osborn, 1926. “Additional 
New Genera and Species of the Mastodontoid Proboscidea.”’ 
Amer. Mus. Novitates, No. 238, November 30, 1926, p. 1. 
Typr.—Superior true molar series of the right side, r.M'*. Amer. 
Mus. 10666. Horizon aND Locauiry.—Walnut, Illinois. 
Geologic age uncertain, possibly of Postglacial (IV) or post- 
Wisconsin age. Tyre Ficure.—Op. cit., 1926.706, p. 2, fig. 1. 

Sprciric Cuaracters.—(Op. cit., 1926.706, p. 1): “This 
progressive subspecies is distinguished by numerous minor fold- 
ings, valleys, and plications which break up all the surfaces of 
the lophs or lobes and which to a certain degree indent even 
the summits of the ridge-crests.” 

M. americanus 
Amer. Mus. 10666 

1/4 nat. size 


Fig. 120. Type superior true molar series of Mastodon americanus 
plicatus Osborn (Amer. Mus. 10666), from Walnut, Bureau County, 
lllinois. One-fourth natural size. After Osborn, 1926.706, p. 2, fig. 1. 
Compare Mastodon oregonensis Hay (Fig. 119). 

“As in the case of the type of Mastodon rugosidens Leidy, it is 
somewhat doubtful whether these foldings and plications represent 
constant specific characters. Consequently the teeth are given 
merely subspecific rank, pending a fuller study of the dental char- 
acters in the Mastodon americanus molar teeth in America.” 


Mastodon moodiei Barbour, 1931 
Figure 121 

West Blue River, about nine miles southwest of Milford, Seward County, 
Nebraska. Pleistocene, II Glacial, Kansan. See Pleistocene Correlation Chart 
in Appendix of the present Volume I. 

The type cranium of this species, discovered during the 
winter of 1931, was unearthed, together with the atlas, axis, 
two thoracic vertebre, and ribs, between March and June of the 
same year. It was named by Doctor Barbour Mastodon moodiei, 
after Dr. Roy L. Moodie of the University of Southern California. 

Mastodon moodiei Barbour, 1931. ‘‘The Milford Mastodon, 
Mastodon moodiei, Sp. Noy. A Preliminary Report.’ Neb. 
State Mus., Bull. 24, Vol. I, December, 1931, pp. 203-210 (Barbour, 
1931.3). Typre.—Skull, together with atlas, axis, two thora- 
cic vertebree, ribs, and mandible with teeth. Neb. Mus. 21-38-31. 

Neb Mus. 21-3-31 

INastodon moodiei Barbour 

Fig. 121. 

protruded 6 or 8 feet or more. . . . By the position of the frag- 
ments the tusks were judged to have been 6 to 8 feet in length. . 


Transverse diameter of condyle, 5 inches (127 mm.) 

Depth in front of the molar, 7 inches (180 mm.) 

Depth just back of the molar, 6% inches (172 mm.) 

Greatest thickness, 7 inches (182 mm.) 

Length of molar, 6% inches (175 mm.) 

Width of molar, 3% inches (96 mm.) 

Length of mandibular tusks, 934 inches (235 mm.) 

Diameter of tusk, 2 inches (51 mm.) 

The beds which yielded the relics of Mastodon moodiei are 
plainly Pleistocene in age, probably Aftonian.” 

Typr or MAstopoN MOODIEI BARBOUR, THE Mitrorp Mastopon or Nesraska, Morritt Pataw&oNnTOLOGICAL 

Cotuections (Nes. Mus. 21-3-31) 

(Left) Adult mandible with M2, Ms, crown view. About 
one-sixth natural size. 

Rostrum relatively long, laterally compressed; short, blunt 
incisive tusks worn off squarely at tips. Compare figure 87. 

Horizon AND Locatiry.—Dam No. 7, across the West Blue River, 
about nine miles southwest of Milford, Seward County, Nebraska. 
Pleistocene, II Glacial, Kansan. Type Figure.—Barbour, 
op. cit., figs. 130, 131, 132b. See also new figures of type in the 
present Memoir (Fig. 121). 

Speciric CHaracters.— (Barbour, 1931.3, pp. 206, 207, 210): 
“Both mandibular tusks were secured uninjured. They are dense 
and strong and slightly curved. They measure 914 inches (235 
mm.) in length, and the pulp cavity is shallow, measuring but % 
inch (13 mm.) in depth. Each mandibular tusk ends squarely, 
for an indefinite amount has been worn off in some manner. To 
have thus ground them off it seems as though the creature must 
have designedly rubbed his tusks on rocks. Yet the interference 
offered by the great protruding upper tusks must have rendered 
such a procedure impossible. The incisive sheaths show that the 
upper tusks had a diameter of 4 inches, and they must have 

(Right) Type skull and mandible. About one-twelfth 

natural size. 
The mandible exhibits a downcurved rostrum with two 
blunt incisive tusks sharply worn off at the extremities. 

1931).—The mandibular tusks of Mastodon americanus (ef. Barbour, 
1931.1, pp. 163, 164) were waning inheritances from the past, 
which plainly required ages for reduction and suppression, and 
complete elimination was never a reality. In the majority of 
jaws, the inferior tusks are wholly wanting; in others the tusks 
have not been erupted, but lie embedded in the bone; sometimes 
one or two small tusks are erupted; in still other cases large ones 

The Warren mastodon in The American Museum of Natural 
History (Fig. 124) had a solitary dwarfed tusk in the mandible, 
and henée was named Tetracaulodon, and the term might still be 
used conveniently to designate mastodons possessed of mandibular 
tusks. The tetracaulodont type of mastodon is not particularly 
rare, for there are two good examples in Amherst. Hay reported 
several, and Warren, studying mastodon jaws, found nine of this 


type. ‘There is one example in the Nebraska State Museum. In 
the museums of the country many others can doubtless be found 
The presence of mandibular tusks in mastodon jaws is partly a 
sexual character, those with tusks being males, those without, 

In the Amherst Museum is a jaw from Nine Mile Bottom, 
South Carolina, displaying inferior tusks 14 in. in length (Barbour, 
op. cit., fig. 103), whereas the Warren Mastodon in the American 
Museum shows only one small vestigial mandibular tusk (op. cit., 
fig. 104). In the ‘Monroe mastodon’ of Orange County, New 
York, in the State Museum, Albany, there are two dwarfed and 
irregular mandibular tusks. In the ‘Seward County Mastodon’ 
of Nebraska (Neb. Mus. 11-7—25), the two mandibular tusks 
were probably 12-18 in. in length (op. cit., fig. 106). 

From all these geologic, phylogenetic, and sexual variations 
of the mandibular tusks, we draw the induction that at the begin- 
ning of Pleistocene time not only was the rostrum of the mandible 
much longer than at the close of Pleistocene time, as illustrated in 
Mastodon progenium Hay, but the pair of lower mandibular tusks 
were much more fully developed, as illustrated in the case of the 
‘Seward County Mastodon’ (Barbour, op. cit., fig. 106), which 
was found 75 feet below the surface in interglacial gravels. 

Mastodon grangeri Barbour, 1934 
From near Pender, Thurston County, Nebraska. 
For full description, with figures, of Mastodon grangeri, see 
Appendix of the present Volume I. 

Mastodon raki Frick, 1933 
Figure 122 

From Hot Springs, New Mexico. Pleistocene. 

Mastodon raki Frick, 1933. ‘‘ New Remains of Trilophodont- 
Tetrabelodont Mastodons,” Bull. Amer. Mus. Nat. Hist., Vol. 
LIX, Art. IX, pp. 506, 510, 630. Typr.—(Op. cit., p. 630): 
“Right ramus and symphysis and part of left ramus with me-ms. 
F:A.M. 23335.” From Hot Springs, New Mexico. Found in 1927. 
Pleistocene. Typr Ficgurn.—Op. cit., figs. 25A and 29A. 

Type Description.—(Frick, 1933, p. 630): ‘A right and 
broken left ramus with m; alveolus, m2-ms and partial symphysis, 
from the Pleistocene of New Mexico, is of the general proportions 
typical of Mastodon americanus-like forms. On the character of 


the heel and the ms crown, which is narrower and taller than usual, 
the specimen is referred to a new species.” 

Osborn, 1933: Osborn observes (Fig. 122) that the type third 
right inferior molar exhibits the following: (1) Serrated spurs from 
the ectoconelets, with reduced contiguous mesoconelet; (2) three 
to four conelets on worn inner lobes; (3) traces of cement in two 
anterior median valleys; (4) third inferior molar with well- 

developed pentalophid, crown relatively elongate and narrow (ap. 

Mastodon raki Frick 

Fig. 122. (Upper) Type third right inferior molar, r.Ms, distinguished 
by the unusual prominence of the trefoil crests extending down the slopes of 
the ectoconelets (see 5 below). After Frick, 1933, fig. 25A. About one-half 
natural size. 

(Lower) “ Right ramus and symphysis and part of left ramus with me-ms 
{rev.].” After Frick, 1933, op. cit., fig. 29A. About one-sixth natural size. 

184 mm., tr. 80 mm., index 43), as compared with corresponding 
r.M; of Mastodon americanus (Amer. Mus. 21920, ap. 181 mm., tr. 
93 mm., index 51) and with the large Shawangunk cranium (Amer. 
Mus. 2595, ap. 155 mm., tr. 84 mm., index 54). (5) This molar 



| Mastodon raki Type Mastodon americanus 
F:A.M. 23335 A.M. 21920 A.M. 17771 A.M. 2595 | 
New Mexico New York Indiana ‘Shawangunk’ | 
New York | 
-———- a | = 
NEE re 2h. seats icia's Sepeniy wn ahe ate ee 88 mm. 77.5+ mm. 
E2625 A ATs tuiscieie ate ka cca ws 114 mm. 113 114 103 
PERT eg tree Soin ovals interes tee ae 2 184 181 180 155 
Mer nat AO Crest... isc. ceate ck cost ee 80 (crown tall) 93 (crown low) 96 84 
Pier Ule ENP thik... deasel cite. site vas 800e 
Mrameteriof Gusk.:...%.s 0.68. sees hoe 45 


with serrated ectoconelets and small mesoconelet appears to be 
distinct from the “losange’’ form prevailing in Mastodon americanus. 

GeroLtogic AGrE.—Frick (op. cit., p. 630) observes: ‘‘The 
mandible was collected by Joseph Rak in the fall of 1927 from beds 
bearing teeth of Pleistocene Equus.” 

Mastodon americanus alaskensis Frick, 1933 
Figure 123 

From the vicinity of Fairbanks, Alaska. Pleistocene. 

The specimen is especially interesting and important because 
of the rarity of the Mastodon in Alaska. 

Frick observes (p. 631): 

‘ Ve) ay 3 4 _— AB 
. Wa IANS ——gee ae he 
Type i rae hit?“ 
INastodon americanus alaskensis Frick ZA 

Fig. 123. Type of Mastodon americanus alaskensis Frick (F:A.M. 27009), 
about one-sixth natural size. 

Left ramus and symphysis, also left third superior molar, 1.M*. After 
Frick, 1933, fig. 29A. 



cs ar} 
P| a 

aN Ee ES 


“‘ Mastodon itself so far has been exampled in the Fairbanks area 
alone by a few detached molars. As associated remains of any of 
the extinct forms are almost unknown in the particular area, the 
present season’s find of a mastodon mandible associated with mi, 
superior tusk and a representation of the limb elements, is re- 

Mastodon americanus alaskensis Frick, 1933. ‘‘New Remains 
of Trilophodont-Tetrabelodont Mastodons,” Bull. Amer. Mus. 
Nat. Hist., Vol. LIX, Art. IX, pp. 506, 510, 631, 632. TypPEr.— 
(Op. cit., p. 631): “Partial mandible with left m2, both mss, partial 

M.a. alaskensis|M. americanus | M. americanus 
A.C-F:A.M. A.M. 14345 A.M. 9951 
Meteor oe 109-+mm. 123 
m3 187 [ap.] 196 (left) 
{101 tr.| (176, right) 
Humerus...... 905 910 
Radius 680 670 
Winae scenes 675 
Metacarpal 1. . 107 110 
Ne 148 150 
Boe 175 160 
4.. 152 150 
ae 123 100 


ae ee 

plat | | 

Mastodontinae type O O, referred e. Zygolophodontina 




American mastodon discoveries 

Fig. 128a. TuHrorretic Migration Lines 

Famous Warren mastodon of Newburgh ®. 

a, referred a. Ancient mastodonts type ®, referred +. 
Osborn, 1934. 



“Yoke-crested”’ mastodonts (subfam. Zygolophodontine, gen. Zygolophodon, Turicius) indicated by squares: 0 types, m referred. Compare figure 137. 

True mastodonts (subfam. Mastodontine, gen. Miomastodon, Pliomastodon, Mastodon) indicated by circles: © types, @ referred. 

Ancient mastodonts (order Mastodontoidea, gen. Palzomastodon) indicated 
Miomastodon tapiroides americanus, Tasndd, Hungary. 
Miomastodon depereti sp. nov., Chevilly, France. 

Compare figure 86. 
by crosses: ® types, + referred. 

Pliomastodon americanus praetypica, Maria-Theresiopel and Batta-lrd, Hungary. 

“Mastodon ohioticus,’ Pestchana, Podolia, Russia. 
Mastodon pavlowi sp. noy., Pestchana, Podolia, Russia. 

Zygolophodon, chiefly distributed in Italy, North Africa, France, and Hungary. Zygolophodon (?) (Matsumoto), Japan. 
Mastodon tapiroides (Pallas, 1770-1777), Belaja, Russia [ =Zygolophodon borsoni). 
Mastodon abundant in former forested regions of North America. See figure 123b. 

Miomastodon and Pliomastodon rare in western United States. 

See figure 123b. 


alveolus of m, and incisive alveoli; left m*, superior tusk, humerus, 
both radii, ulne, manus, and distal end of femur. Alaska College 
—F:A.M. 27009. Figured this paper, Fig. 29A.” Horizon 
anp Locauiry.—Vicinity of Fairbanks, Alaska. TYPE 
Ficure.—Op. cil., fig. 29A. 

Tyre Description.—(Frick, 1933, pp. 631, 632): ‘The 

mandible exhibits a heavier vertical ramus and symphysis and 
narrower symphysial trough, the alveoli suggest much larger 
incisors, and the molar tooth-crowns are lower than in the American 
The limbs ap- 

Mastodon, as seen in A. M. 14345, from Illinois. 
proximate those of the Warren mount [Amer. Mus. 9951]. 

tusk measures 7 ft. 1 in. on curve, and 19.5 in. circumference at 

Osborn, 1933: The type mandible of Mastodon americanus 
alaskensis reveals (Fig. 123) relatively broad third inferior 
grinders, namely, 1.M;, ap. 187 mm., tr. 101 mm., index 54; (2) 
the mandible is correspondingly massive, as well as the rostrum. 
Another feature, also observed in Mastodon raki Frick, is the 
serration of the ectoconelets and relative reduction of the con- 
tiguous mesoconelet. Frick observes (op. cit., p. 631) that “The 
individual, while of as large size as the Warren Mammoth [Masto- 
don], was immature, the epiphyses being unconsolidated.” 


















—* SSS 7 LAA 
Mastodon americanus in North America-after OP Hay. 1923-1930 






(1) Type locality of Mastodon americanus Kerr-Cuvier. 

(7)-(24), see full list of species on page 137. 

© Geographic distribution of subspecies and mutations of Mastodon americanus in the United States and Canada, after detailed reports and re- 
searches of O. P. Hay on the mastodonts of the Eastern, Middle, and Western United States, and Eastern Canada. 


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See figures 82, 83, 124, 126, and 130 

Hudson River near Newburgh, New York, Pleistocene, of Postglacial (IV), or post-Wisconsin age 

Every bone of this superb skeleton has been separately figured and described with great accuracy in the two 
editions of Warren’s great Memoir (1852, 1855). The reader is referred to Warren’s Memoir for all details, except 
those relating to the tusks which are fully described below in the present Memoir. 

A full history of the discovery in the summer of 1845 of this famous skeleton and of the various phases in its 
mounting and exhibition is given below (pp. 181-183). In 1906 the skeleton was presented to the American 
Museum by Mr. J. Pierpont Morgan, to whom this Memoir is dedicated. The entire year (1906-1907) was 
devoted to the work of cleansing, renewing, and remounting the skeleton; all the bones were given an alcohol 
immersion to remove the dark varnish with which they had been covered, thus the beautiful original light brown 
color at the time of discovery was completely regained. The tusks had been completely shattered and were 
restored with great difficulty; twenty-three inches of each tusk are inserted in the sockets, the projecting part 
measures 6 feet 8 inches, as shown in figures 124 and 126. 


Mastodon americanus Loxodonta africana oxyotis 
Total length of skeleton, base of incisive tusks to 
perpendicular drop of tail.................. 4.546 14ft. 11 in. 4.090 13 ft. 5in. 
Height to summit of dorsal ee Pe summit 
of scapule.. >, , 4 Macrame ths: 9 1% 3.150510 4 
Summit of ani fr om iti! er re. 7a (opts) te) te! 3.045 10 
Summit of highest part of pelvis or ossa nominees 
from the ground............... 2.668 8 9 : 2.785 9 2 
Incisive tusks: total jencth of mane uelee on nani 
curve.. : a eerolon 7S (I 0S ee Sh cic 
Incisive ie Seay oe neat ae as exposed. 2033, 6538 oo — —E eer 
Thigh bones: length of right tibia.. ete t:(/i| De a4 788 2 7 
length of night ee. 2 vat ROG 3. 4% 1.258 4 1} 
length of right pes eoedieie : .53 19 (on ground) S217 8 
pelvis, width of ossa inane 1.826 6 1.340 4 7M 
Shoulder girdle and forelimb: height of canna 916 38 yy, 900 2 1 
right humerus, 
length of......... 950). Sy rt 1.091 38 7 
right ulna-radius, : 
length of......... .658 2 2 (ulna without ole- 1.037 3 4h, 
right extended 
manus, length of . 285 11% (without carpus) 
Seven cervical vertebre, length of ................ .ol ise ys! -900 «1 10 
Dorsolumbar vertebre, total length of........... 2.250 7 4 2.0385 6 8 
Five sacral vertebre, total length of.............. 48 1 6% . 268 104 (length 4 sacral 
‘ vertebre) 
Caudal vertebre, estimated total number (28), 
POUBISIERM DUN Ofer Seiapeten s aante sl aanatne es aerate: 2.085 6 10 1.455 4 9 (est. 21 caudal ver- 

‘Amer. Mus. Dept. Mam. 3283. Prepared from specimen presented by P. T. Barnum in the year 1889. 


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The contrast between the comparatively broad, long-backed, low-bodied proportions of Mastodon ameri- 
canus and the elevated, short-backed, narrow-bodied proportions of the African elephant is similar to that which 
generally obtains as between the Mastodontoidea and the higher Elephantoidea. 

MASTODONTOIDEA: Relatively broad, low-bodied, heavy-limbed, long-backed proboscideans. 
ELEPHANTOIDEA: Relatively narrow, high-bodied, long-limbed, short-backed proboscideans. 

Tue ‘Wuitrietp Masropon’ (MAsToDON AMERICANUS) SKELETON 

Hudson River, near Newburgh, New York, Pleistocene, probably of Postglacial (IV) or post-Wisconsin age 

The Whitfield Mastodon (Fig. 125), named in honor of the late Curator Robert Parr Whitfield of the Ameri- 
can Museum, is the assembled skeleton of a male mastodon in which the tusks are of a large size, but the propor- 
tions and mounting are much less accurate. This skeleton, mounted for the American Museum in 1879, was 
purchased by the Senckenberg Museum of Frankfort, Germany, where it is now on exhibition. 

History.—(1) This skeleton was discovered in 1879 near the town of Little Britain, about nine miles south 
of Newburgh, N. Y. It was imbedded in peaty material on the edge of what had been less than fifty years before 
an open pond; the pond had been drained and was under cultivation at the time of discovery. In course of further 
drainage work, following the wet season of 1879, at a depth of fourteen inches, an apparent log was reached, 
which proved to be the leg bone of an animal. This induced the farmer to search for other bones, and in a few 
days more than one-third of the skeleton had been exhumed, including the head and lower jaws. (2) Three 
weeks later Professor R. P. Whitfield, first Curator of Geology in The American Museum of Natural History, 
visited the excavation at the request of Major T. B. Brooks of Newburgh. He found that the swamp on the side 
nearest the skeleton was bordered by a low hill of hard blue “boulder clay,” mixed with shale and gravel. This 
hard layer sloped down and passed under the peat or “muck” of the swamp to form the original bottom of the 
pond. From this peat the skeleton derived its dark color. (3) The skeleton lay with the head farthest from the 
original shore of the pond and deeply imbedded. The right limbs were near the surface, the right humerus being 
buried only fourteen inches, which proves apparently that the animal had been mired and had fallen on its left 
side. At the point where the head was found, twenty feet from the place at which the ‘“‘boulder clay”’ rose to the 
surface, the peaty material had accumulated over the head to a depth of ten feet. The humerus and other parts 
nearer the surface were consequently less perfectly preserved. 


Superior and inferior grinders, incisive tusks, age, sex, growth, cranial and mandibular development 

In the assemblage and preparation of the tusks of the Warren Mastodon skeleton (Amer. Mus. 9951), Osborn 
(1910.347 and 1923.574) made the following observations which should be permanently recorded in this Memoir: 

A Means or EstImaTING THE AGE OF THE Mastopon.—(Osborn, 1910.347, pp. 1-3): In repairing the tusks, the outer 
sheathing of the dentine was found in large part absent. The inner sheathing exposed a series of concentric constrictions and 
expansions which were observed to be approximately symmetrical on the two sides, as indicated by the series of + signs in Fig. 3, 
A and B [Fig. 126, right, of present Memoir]. Secondly, it was noted that the intervals between these constrictions are broader 
in the middle and fore part of the tusks (corresponding with the youthful stage of growth of the tusk) and become narrower 


toward the base of the tusk (corresponding with the mature or adult stages of growth). Eighteen of these rings are preserved 
on one side and thirteen on the other. They are faintly indicated also in the waving surface of the dentine of the tusk (Fig. 3). 
On the hypothesis that these are actual annular increments of growth, the right tusk (Fig. 3, A) consisted of about twenty-eight 
segments, which allowing for the milk dentition and for the part worn off at the tip would assign to the Warren Mastodon an age 
of perhaps thirty years. Similar annular constrictions are observed in the tusks of the mammoth from Alaska; and are also 
indicated in the tusks of the African elephant. Since the age of the Indian elephant and the rate of tusk growth is definitely 
known, the identification of similar concentric annular growths would be the means of testing the value of this hypothesis. 

M. americanus 
“Warren Mastodon” 
Amer. Mus, 9951 Ref. 

..M. americanus 
‘Warren Mastodon” 
Amer. Mus. 9951 Ref. 

1/24 nat. size 
V/12 nat. size, 

Fig. 126. Superior, inferior, and lateral views of Warren Mastodon (Amer. 
Mus. 9951) tusks, an adult male from Newburgh-on-Hudson, New York. Both 
figures after Osborn (1910.347) and (1923.574). 

Left. Superior aspect. Observe the deep insertion of the right superior tusk 
and the annular growth rings feebly indicated. One twenty-fourth natural size. 

Right. Enlarged inferior aspect of the tusks showing the annular growth 
rings. One-twelfth natural size. 

A, Right tusk drawn from the outer inferior side showing growth rings 1 to 
19, the center of each ring being indicated by a plus sign (+). Of great interest: 
is the contact point (c) just above the 19th growth ring, which positively 
enabled us to determine the actual length (8 ft. 7 in.) of the right tusk on the 
outside curve. 

B, Left tusk, outer view, showing 13 growth rings. 

The Warren Mastodon thus represents a male animal at least thirty years of age at the time it sank into 
a bed of marl near Newburgh, New York. The story of the restoration of these tusks is as follows (cf. Osborn, 
1923.574, p. 18): 

Metuop or Restorinc tHe WARREN Mastopon Tusks.—There still remained the problem of the tusks, which are in- 
variably the most vital part of buried skeletons of the great proboscideans of the past. It appears that the original tusks could 
not be preserved entire by the methods known at the time of exhumation. The discoverers were unable to prevent the tusks 
from splitting, warping, and falling to pieces, especially at the butt. In order to preserve what could be saved intact, the 
butts of the tusks, already split and warped, were sawed off under Doctor Warren’s direction, and only the tips, about three 
feet in length, were treated and preserved. The butts, fallen into fragments, but still lying undisturbed in two of the original 
boxes used for transporting the skeleton, were found in the Warren Museum when the skeleton was repacked to be sent to 
the American Museum. The tips, treated with preservatives, were still intact in another box; but neither had been used 
apparently for measurements in making the papier-mAché restorations fitted to the skull in the Warren Museum. This documen- 
tary evidence certainly was not used by Doctor Warren, because in his three restorations he unfortunately accepted the 


erroneous original reports that the tusks as found were more than eleven feet in length; they were so deseribed and illus- 
trated by him in the entirely impossible position shown in the photograph on p. 15 [Fig. 83 of the present Memoir]. 

When the Warren collection reached the American Museum, it was very carefully looked over in a search for remnants of 
the original tusks, and finally the fragmentary fossil ivory tips and butts were found, but inasmuch as most of the original 
records had been lost and no use of these materials had been made by Doctor Warren, it remained to be proved that the 
fragmentary butts of the tusks really belonged with the skull. The piecing together of these butts required several months of 
most ingenious and patient work on the part of one of our preparators, Mr. Charles Christman. The ends of each tusk were 
perfectly preserved, but there was no connection between these tips and the reconstructed butts of either tusk. Fortunately, 
when the butts of the tusks were sawed off, a single splinter of bone broke off, and finally this splinter was found to fit exactly 
to a fragment of the butt. There was great rejoicing in the laboratory when the relationship of these two fragments was 
discovered, because it enabled us to determine positively the length of the tusks as 8 feet, 7 inches. 

The rebuilding of the tusks, which required several months of most patient work, had two very important results: in the 
first place, it enabled us to place them properly in the sockets of the skull and to prove for the first time the exact relations of the 
mastodon ivories; secondly, a very painstaking examination of these tusks led to an important and most interesting dis- 
covery, namely, that it was possible to determine very closely the age of the Warren Mastodon. The ivory exhibits a series of 
growth rings which, counted from tip to base, seems to prove that the Warren Mastodon was perhaps thirty years of age 
at the time it sank into the bed of marl near Newburgh. The right tusk included at least twenty-eight of these segments. 
The growth rings are shortest near the tip of the tusk when the animal is young, and increase in length from the tip toward 
the middle of the tusk, but not in a regular ratio. These growth rings do not correspond exactly in the opposite tusk, but 
in both tusks they are longest in the middle region. Nine smaller rings are in the lower part. The writer’s theory regarding 
these growth rings is that during the summer season, when all the conditions of life were favorable, and perhaps during the 
rutting period, when tusk growth was hastened by internal secretions from the reproductive glands, the growth of ivory was 
very rapid, the maximum growth in the 17-18 ring being 108 centimeters, or 4) inches, perhaps the maximum growth of a 
favorable season at the most vigorous reproductive period of life. The Warren Mastodon is an adult but not an aged specimen; 
the skeleton is apparently that of a younger animal than the one represented by the Shawangunk head. Some estimate the 
maximum age of the American mastodon at between thirty and forty years,—less than half the life span of the elephant, 
which attains more than one hundred years. 

MALr AND Femae Tusxs.—In all the known Proboscidea there is a marked disparity between the male 
and female incisive tusks both in length and in diameter. The adult female 

tusks never fully attain the length of the adult male tusks, but a still more 
striking difference is their slenderness of proportion and diameter. 

The female tusks are finely displayed in the female skull of Mastodon 
americanus (Amer. Mus. 14293) found near Fulton, Indiana, and represented 
one-tenth natural size in figure 128. We observe that the divergence of the 
tusks in their sockets produces the strikingly V-shaped arrangement of the 
grinding teeth and the tusks, so that a straight line drawn from the apex of 
the tusks traverses the grinders and passes through the occipital condyle 
of the opposite side; this explains the marked divergence of the grinding 
teeth anteriorly, and the convergence of the grinding teeth posteriorly. 
Undoubtedly the incisive tusks were extremely important in females for the 
defense of the young, and this strongly divergent tusk mechanism brought 
about a strongly divergent grinding tooth mechanism. This divergence, 
however, is much less marked in the male skull (Yale Mus. 12600), beauti- 
fully represented in figure 129, than in the Warren Mastodon skull (Fig. 130). 

Fig. 127. Tusks of Elephas indicus (?) suma- 
Mae aND FeMALe Crania.—The palatal aspect (Fig. 129) of the ‘vv showing 17-18 annular growth rings. 

; Ee After photograph kindly furnished by Mr. H. 
superbly preserved adult skull of the Mastodon americanus from Otisville, | Munniks de Jongh of the Hague, who purchased 

New York, in the Peabody Museum of Yale University (Yale Mus. 12600), ‘2e™ Some years ago without a record. It is 
is sh ‘chil § . i f probable that they are from the Dutch East 
is Shown one-eighth natural size. The drawing is one of the masterpieces of Indies. One-tenth natural size. (Cf. Fig. 126.) 


Mrs. L. M. Sterling, the artist of most of the original pen illustrations in this Memoir. All the constituent bones 
and the foramina are clearly indicated by abbreviations, as lettered under the direction of Dr. William K, 

Fig. 128. Female skull. Palatal view of the finely preserved female skull 
(Amer. Mus. 14293) of Mastodon americanus from near Fulton, Indiana. One- 
tenth natural size 

Observe the strikingly V-shaped arrangement of the grinding teeth and tusks, 
the apex of the V being at the posterior nares. The tusks are relatively long, very 
slender, gently convex or outcurved, and slightly incurved at the extremities. 

Gregory. Viewed both from above (Fig. 130) and 
below (Fig. 129) the facial and cranial portions of 
the M. americanus cranium are subequal in length. 
There is a very marked development of the pre- 
maxillaries (P. mz.) and of the maxillaries (Mz.), an 
abbreviation of the palatines, a compression and an 
abbreviation of all the basicranial elements, which, 
however, are much less extreme than in the basi- 
cranial aspect of the Indian elephant (Hlephas in- 
dicus), see Volume II, Chapter XX. 

The male cranium (Amer. Mus. 14535) repre- 
sented in figure 132 differs strikingly from the 
Peabody Museum cranium (Yale Mus. 12600, Fig. 
129) in the fact that the facial portion, from the 
premaxillaries to the back of the grinders inclusive, 
is decidedly longer than the cranial portion; it 
differs also in the marked divergence of the grind- 
ing teeth of the opposite sides, whereas in the 
Peabody Museum skull (Yale Mus. 12600) the 
grinding teeth are more nearly parallel. 

The female cranium (Amer. Mus. 14292) repre- 
sented in figure 182 shows a very marked sexual 
disparity in size; this is probably the normal dis- 
parity between the male and female crania of 
Mastodon americanus, because this female cranium 
does not differ markedly from the female cranium 
(Amer. Mus. 14293) shown in figure 128. The more 
feeble and slender female tusks (Fig. 128) demand 
less prominent premaxillo-maxillary “development, 
consequently the sockets for the incisive tusks are 
relatively short and small, and elevated above the 
grinding tooth border, as seen in both the female 
skull (Amer. Mus. 14298, Fig. 128) and in the fe- 
male skull (Amer. Mus. 14292, Fig. 132). Thus 
there is a marked disparity in size between the 
females and males in (1) the grinding teeth, (2) the 
incisive tusks, (3) the sockets of the incisive tusks, 
and (4) the size of the cranium as a whole. 

JUVENILE Mag Crania.—The fully adult age 
characters pointed out in the above descriptions 


and figures of the male and female skulls should be compared with the juvenile and infantile characters of the 
immature male crania shown in figure 131. The immature male skull’ (Amer. Mus. 17727, Fig. 131 Al), from 
Rochester, Indiana, represents a stage in which the three ‘intermediate molars’ only are in use, namely, Dp 4, 
M 1, M2. All these teeth show the three transverse crests characteristic of this genus and species. The tetra- 
lophodont third superior and inferior molars, M*-M;, have been extracted from the skull and jaws and are repre- 

M. americanus M. americanus 

t / 
Yale Mus. 12600 Ref. “Warren Mastodon’ an” 1/8 nat. size 
/ Amer. Mus. 9951 Ref. Xs os L_-? 
1/8 nat. size Pe . ee ALS 

. eR gm 
Fig. 129. Mastodon americanus palate of finely preserved Fig. 130. Superior view of skull (Amer. Mus. 9951) of the 
adult skull (Yale Mus. 12600), from Otisville, New York. One- “Warren Mastodon,” an adult male with the tusks removed. 
eighth natural size. Complete skeleton collected May 7, 1875. One-eighth natural size. 

sented in figure 135. In this young male cranium! (Amer. Mus. 17727) we observe the following points: (1) The 
facial and cranial portions are subequal in length; (2) the sides of the cranium are nearly parallel; (3) the juvenile 
tusks are imbedded in slightly expanded premaxillary sockets; (4) all the crests of the grinding teeth (Fig. 131 
Al, also Fig. 135) are extremely sharp, representing a high degree of subhypsodonty. 

The calf skull of Mastodon americanus (Fig. 131 B) found near Hackettstown, Warren County, New Jersey, 
about twenty miles from Newark, represents a much more immature stage (Amer. Mus. 10459). 

IMMATURE AND Mature GRINDING TEETH.—The immature male skull! from Rochester, Indiana (Amer. Mus. 
17727, Fig. 131 A1) yields a series of immature dental crests (represented in Fig. 135) of the third left superior 
molar, 1.M°, and of the third left inferior molar, 1.M;. These ridge-crests are a revelation of the very sharp, 

!This specimen is now made the type of Mastodon acutidens (see Appendix at the close of the present Volume, also Figs. 131, Al—A4, 135, and PI. 1, L, 
of this chapter). 


Fic. 131. (Al1—A4) Type or MastTopoNn ACUTIDENS SP. NOV. 
iews of young Mastodon skull (Amer. Mus. 17727),! from Rochester, Indiana, showing Dp‘, M’, 

Type of Mastodon acutidens sp. nov. 
about twenty miles from 

Al, Left lateral, A2, superior, A3, palatal, A4, occipital v 
M3-M3, extracted from the skull and jaws, shown in figure 135. 

and M2, anterior portion only in use. 
Found near Hackettstown, Warren County, New Jersey, 

B, Calf Mastodon americanus skull (Amer. Mus. 10459), superior view. 
See footnote on page 185. 

Fig. 132. Mate anp Femate Crania oF MASTODON AMERICANUS 

(Left). Male cranium of Mastodon americanus viewed from above and from the 
14535), from Ashley, Indiana. One-twelfth natural size. 

(Right). Female cranium of Mastodon americanus, of somewhat greater age th 

cranium (Amer. Mus. 14292) was discovered two and a half miles southeast of Fulton, Indiana. 

side; also palatal view. This is a young adult cranium (Amer. Mus. 

an the male cranium, exhibiting similar aspects of the skull. This 
One-twelfth natural size. 


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almost knife-like summits of the molar ridge-crests, which give a truly subhypsodont character to these crowns. 

These sharply crested summits illustrate the very marked progression of Mastodon acutidens sp. nov. upon the low, 

compressed crowns in the two species Miomastodon merriami and Pliomastodon matthewi, and upon the rounded 

lobes, i.e., cones, in Palzomastodon. 

Hyrsopont GrinpErS.—It is probable that when all the grinding 
teeth now referred to Mastodon americanus are carefully measured and 
compared, considerable progressive evolution toward hypsodonty will be 
observed through the long period of Pleistocene time. 

Much more sharply elevated and acute transverse crests than those of 
the female specimens found near Fulton, Indiana, and represented in 
figure 134, are the grinders of the young male (Amer. Mus. 17727, Fig. 135), 
the type of Mastodon acutidens sp. nov. 

TypicaL GrinpERS.—The typical female grinders (Amer. Mus. 14293 
and 14294, Fig. 134) exhibit relatively obtuse crests both in the upper and 
lower jaws. It may be observed also: (1) That the individual lobes, i.e., 
cones, are not distinctly bilobed; (2) that the worn enamel presents 
simple, compressed loops in M*-M,; (3) that the trefoil spurs rise from 
the internal lobes of M?, from the external lobes of M,; (4) that a sharp 
median sulcus divides the internal lobes from the external in M*-M;. It 
is not surprising that these rudimentary trefoil spurs should be compared 
with the homologous trefoils of Trilophodon and of Serridentinus; in 
(5) The 
third inferior molar, Ms, is distinctly longer and relatively narrower than 
the third superior molar, M’. 
very clearly illustrate the characters of the typical Mastodon americanus 
and agree closely with Cuvier’s type grinders (Figs. 76, 112) from the Big- 
Bone Lick, near the Ohio River, Kentucky. 

position they resemble Serridentinus rather than Trilophodon. 

These beautifully drawn grinding teeth 

PLICATED GRINDERS.—A somewhat more folded or plicated condition 
of the superior grinders is shown in the palate and lower jaw (Amer. 
Mus. 12464, Fig. 133) of a middle-aged male (?) individual of Mastodon 
americanus found near Buffalo, Kansas. These plicated or folded grinders 
approach those of the type of Mastodon americanus plicatus Osborn (Fig. 
120), although the plication is somewhat less extreme. We observe in M* 
(Fig. 133 A) the marked development of the cingulum, internal, anterior, 
and external, and the marked irregularity of the enamel surfaces of the 
lobes; the tetartoloph in M* is subdivided into four conelets; in M; there 
is a strong anterior cingulum, but it exhibits no external or internal cingula. 

We conclude that the male and female grinding teeth of specimens 
from different parts of the United States, mostly of Upper Pleistocene and 




Fig. 135. Sharp summits of partly developed 
molar crowns excavated from a young male skull 
(Amer. Mus. 17727), from Rochester, Indiana. 
Compare figure 131, Al-A4. 

Al, Five crested left superior molar, 1.M*, external 

A2, Crown view of the same five crests of ].M®. 
Observe the division of the transverse crests into 
inner and outer lobes, completely separate in the 
young condition. 

B1, Crests of undeveloped left lower molar, |.Ms, 
somewhat more advanced than the upper. Observe 
the bilobed condition of the fourth crest; fifth crest 
not developed. 

B2, External view of same crests of 1.M 3. Observe 
the high anteroposterior compression of these crests 
adapted to the sharp cutting of the food. 

Postglacial time, exhibit marked progressive ascending mutations and subspecific characters which may be found 

to distinguish more or less constantly the descendants of certain subspecific, specific, and phyletic branches. 



In the American Museum materials referred to Mastodon americanus, it is clear from the above brief descrip- 
tions and from the close examination and comparison of the carefully prepared figures that there is a very marked 
progressive evolution in the grinding teeth, and probably in the tusks and portions of the skeleton as well, indicating 
varied ascending mutations and subspecific if not specific stages. These characters are not mere individual varia- 
tions; they certainly mark progressive evolution extending over a very long period of time, namely, over the 
entire 1,000,000-year period, the minimum figure now assigned to the American Quaternary or Pleistocene time. 

(1) The primitive species Mastodon progenius Hay is believed to be of Aftonian or Ist [interglacial age. 

(2) The Mastodon americanus of Big-Bone Lick, Kentucky, is of much more recent geologic age, probably TV 
Glacial time, indicated by Ovibos and Rangifer. These are probably the typical deposits of the species Hlephas 
[Mastodon] americanus Kerr, which came from the Big-Bone Lick. 

(3) The ‘mastodon’ deposits of Newburgh-on-Hudson, New York, including the ‘Warren Mastodon,’ are 
probably of Postglacial (IV) age, or the period of the retreat of the great Wisconsin glacier. 

If these geologic correlations are correct, the evolution of Mastodon americanus covered a period of at least 
1,000,000 years, from the more primitive stages, such as M. progenius, to the most progressive stages, such as M, 
acutidens sp. nov. These stages, however, can only be clarified by very close and prolonged monographic research. 

Pe "4 




Left portion of the great late-Pleistocene mural in the Hall of the Age of Man in the American Museum. The faunal period of Bison latifrons 
(Bison regius) and Equus complicatus (Equus scotti) 
Fig. 136. This mural, painted under the direction of Henry Fairfield Osborn, was completed by Charles R. Knight in the year 1920 and is one of the 
finest examples of his art. Inthe group there are one bull, two cows, and a calf. 




1. History of the subfamily Zygolophodontine, 1770-1927— 6. Characters of Zygolophodon and its included species. 
1935. 7. Characters of Turicius and its included species and muta- 

2. Collateral relationship to the subfamily Stegodontine. : ; = errs aay Een «(pee 
Sela: : t as 7 l d Serri- 
3. Zygolophodon and Turicius independent phyla (Osborn, aie as compared with Zygolophoden and: errs 
1926) of the Zygolophodontinz Osborn, 1923. : SS gee = '¢ y 
4. Résumé of the generic and specific distinctions. 8. Geographic distribution and extinction of species of 
5. Progressive characters of the grinding teeth. Turicius. 

(January, 1935) Since 1928, when this chapter was written, the fundamental patterns of the grinding teeth of Paleomastodon, 
Mastodon, Zygolophodon, Turicius, and Stegolophodon have been intensively examined, compared, and figured, as shown in 
Pls. I-IV, between pp. 134 and 135. These new comparative figures positively demonstrate: (1) The complete phyletic separa- 
tion of these five genera, and (2) the possible derivation of the grinding teeth of the Stegodontoidea from those of Stegolophodon.' 
The species of Stegolophodon and of Stegodon are described in detail in Chapter XV of Volume II of this Memoir. 

We may open this chapter by calling attention to the marked sexual and progressive characters which may be 
observed in the mastodonts in the American Museum collections. These serve clearly to distinguish the Mas- 
todontine or true mastodonts from their most nearly related phylum the Zygolophodontine, because it is clear 
throughout that (1) the true Mastodon molar is profoundly different in character from the true Zygolophodon 
molar, and (2) that the final stage of evolution in the Mastodon acutidens molar (Fig. 135) converges towards 
the final evolution stage in the zygolophodont molar (Fig. 168, Turicius virgatidens). 

This chapter covers the genera Zygolophodon Vacek and Turicius Osborn. The species included within 
Turicius show several resemblances to those included within the genus Serridentinus (Chap. X), as observed by 
the present author; in many other characters Turicius is clearly separable from Serridentinus and more closely 
related to Zygolophodon. The median longitudinal sulcus (Fig. 89) of Mastodon americanus disappears in Zygolo- 
phodon and transversely perfected crests evolve. Zygolophodon has four to six main conelets, Turicius has five 
to eight small conelets (7. turicensis) to eight to nine (T. virgatidens). 

As pointed out clearly in the preceding and succeeding sections, the genus Zygolophodon, based by Vacek 
(1877) on the three genotypic species Mastodon borsoni, M. turicensis, and M. tapiroides (= M. pyrenaicus), now 
proves to include two distinet generic phyla which may be traced chiefly in western Eurasia from the base of the 
Miocene to the summit of the Pliocene, namely: 

I. Zygolophodon borsoni, which may spring from Z. pyrenaicus, and from Z. pyrenaicus aurelianensis, Lower 
Miocene of France. This generic phylum parallels the true Mastodon americanus phylum in its evolution, but it is 
nevertheless quite distinct. Like the true mastodonts, it is little known until Middle and Upper Pliocene times, 
when the remains of Z. borsoni are very abundant. 

Il. Turicius, typified by Turicius turicensis of the Upper Miocene, is a very distinct generic phylum, extend- 
ing from the Lower Miocene ‘Mastodon’ tapiroides of Cuvier to the Middle Pliocene. This phylum is abundantly 
represented especially in lignitic and fluviatile deposits. It is quite distinct (see Figs. 188 C, C1; 139, 4, 5, of 
the present chapter) from the phylum Zygolophodon borsoni and presents certain parallels with the Serridentinus 
phylum, which is widely distributed in the Miocene and Pliocene of America. 

1(1935) Stegolophodon removed from the Stegodontine and placed in the new subfamily SregoLorHopONTIN ® (see Appendix of the present Volume I). 




Original reference: Zygolophodontinz Osborn, 1923, ‘‘ New Subfamily, Generic, and Specific Stages in the Evolution of the Probo- 
scidea,’’ Amer. Mus. Novitates, No. 99, p. 1 (Osborn, 1923.601). 

Burron, 1770.—The history of discovery goes back to the year 1770, when Buffon received from the Comte 

de Vergennes an upper grinding tooth found in Russia (“‘la petite Tartarie”’) and after examination and descrip- 

tion figured the tooth in 1778, as shown in our facsimile reproduction (Fig. 111). 


Pavlow (1894) re- 

fers to Buffon’s description of this tooth in the ‘‘Epoques de la Nature,’ 1775 [1778]. Almost at the same time 
(1777) Pallas described a tooth found near the Belaja, an affluent of the Kama River, Russia, a tooth which sub- 
sequently Eichwald and de Blainville referred to Mastodon tapiroides [now referable to Zygolophodon borsona). 
Pavlow’s full description of these Russian and Siberian discoveries is cited above (Chapter VI, pp. 133-135). 

be as follows: 




1857, 1859 
1859, 1879 





Near the Belaja, Russia 

Montabusard, France 
Montabusard, France 

Montabusard, France 
Elgg, Switzerland 
Piedmont, Italy 
Simorre, France 
Vialette (environs du 
Puy), France 
Velay, France 
Auvergne, France 
Pikermi, Greece 

Locality (?) 

Fulda, Germany 

Thorn, West Prussia 
Wahlheim and Essel- 
born, Germany 

Sables de l’Orléanais, 
Chevilly (?), Casse- 
grain, Avaray, 

Simorre, France 

See Figure 137 


True zygolophodont molar, de- 
scribed and figured by Buffon 
(see Fig. 111) 

Zygolophodont molar described 
by Pallas 

Petit mastodonte Cuvier 

Mastotherium microdon Fischer 
de Waldheim 

Mastodon microdon Rafinesque 

Mastodonte tapiroide Cuvier 

Mastodon turicense Schinz 

Mastodon Borsoni Hays 

M. tapiroides-minus Lartet 

Mastodon Vialetii Aymard 
Mastodon vellavus Aymard 
Mastodon Buffonis Pomel 
Mastodon atticus G. & L. 
(in Wagner) 
M. (Triloph.) Pyrenaicus 
Mastodon affinis Jourdan (MS., 
labels in Lyons Museum); in 
Lortet and Chantre, 1879 
Mastodon virgatidens von 
Mastodon Zaddachi Jentzsch 
Mastodon wahlheimensis Klahn 

Zygolophodon pyrenaicus aure- 
lianensis Osborn 
Turicius turicensis simorrensis 



= Zygolophodon borsoni 

=Zygolophodon borsoni 
= Turicius tapiroides 

= Turicius tapiroides 
=(?)Turicius tapiroides 

= Turicius tapiroides 

= Turicius turicensis 

= Zygolophodon borsoni 

= Turicius tapiroides-minus 

=Zygolophodon borsoni vialetii 
= Zygolophodon borsoni vellavus 
= Zygolophodon borsoni buffonis 

= Turicius atticus 

= Zygolophodon pyrenaicus 

=Zygolophodon borsoni affinis 

= Turicius virgatidens 
=Zygolophodon borsoni zaddachi 
= Turicius wahlheimensis 
[see Chap. VIII, p. 282, for 
description and figure] 

=Zygolophodon pyrenaicus aure- 

= Turicius turicensis simorrensis 

The chief succession in the discovery and description of the species of Zygolophodonts in Eurasia appears to 

GroLocic AGE 

Pleistocene (?) 

Pleistocene (?) 
Lower Miocene 

Lower Miocene 
Lower Miocene 

Upper Miocene 
Middle Pliocene 

Upper Pliocene 
Upper Pliocene 
Upper Pliocene 

Lower Pliocene 

Middle Miocene 

Middle Miocene 

Middle(?) Pliocene 

Upper Pliocene 

Upper Pliocene 

Lower Miocene 

Middle Miocene 


Cuvier, 1806.—In Chapter V, ‘History of the Classification of the Mastodontoidea, Families and Sub- 
families, 1705-1927,” we have recited Cuvier’s original description (1806) of ‘‘Petit mastodonte” (= Turicius 
tapiroides), a tooth from Montabusard, compared with one from Saxony; this specific description was confirmed 
by de Blainville in 1839-1864. Schinz (1824, p. 278) recognized Cuvier’s five classic species and added the 
zygolophodont species ‘Mastodon turicense’ from the Miocene lignites of Elgg, Canton Zurich, Switzerland. Sub- 
sequently the zygolophodont relationship of ‘Mastodon’ tapiroides and of ‘M.’ turicensis was recognized. 

Hays, 1834, AyMarp, 1847, Pome., 1848.—The next step was that of Hays, clearly distinguishing the Middle 
to Upper Pliocene zygolophodont species as Mastodon borsoni. Then followed the descriptions of Mastodon 
vialetii and of M. vellavus by Aymard, of Mastodon Buffonis by Pomel, and of Mastodon zaddachi by Jentasch—all 
of which species we now believe to be referable to Zygolophodon borsoni. See Hays’ type figure and description 
on page 207 below. 

Waaner, 1857.—In 1856 Gaudry and Lartet described Mastodon pentelicus. In 1857 Wagner cited (p. 140) 
the same animal as: “11. Mastodon atticus G. & L.,” thus apparently attributing the species pentelicus to Gaudry 
and Lartet, but under another name. In PI. vu, fig. 16, he referred to this species a maxillary fragment of another 
individual, on page 142 making reference to this specimen as follows: ‘In meinen friiheren Erérterungen 

Osborn | 

Fig 137. Geographic distribution (according to the numbers in the list opposite) of species of Zygolophodon and of Turicius. The white dots within the 
bluck areas represent the approximate localities where the types of these sixteen species were discovered. The white crosses represent referred specimens. 
The Japanese record is from the Tatsunokuchi, Lower Pliocene (Matsumoto, letter, 1924). 

dieser Ueberreste habe ich es nicht gewagt, sie einer bestimmten Spezies von Mastodon zuzuweisen. Gaudry und 
Lartet haben sie jetzt als Mastodon atticus bezeichnet; ihre ausfiihrliche Abhandlung, die noch nicht erschienen 
ist, wird jedenfalls die Griinde angeben, warum sie dieselbe von M. angustidens getrennt haben.” 

Osborn (1927) in the present Memoir points out that Wagner’s maxillary fragment (PI. vu, fig. 16) belongs 
to the genus T'uricius and by courtesy adopts Wagner’s specific name atticus. 

Gaupry, 1862.—Gaudry (1862, p. 153) pointed out that Cuvier established the name tapiroides on a single 
fractured milk molar (‘‘molaire de lait’’), a specimen which had been mentioned as early as 1783 by Defay; the 
third tooth cited by Defay was described by Cuvier as M. tapiroides. 


Gaudry also pointed out, referring to Schinz, von Meyer, de Blainville, Lartet, and Kaup, that the teeth 
described throughout France as ‘M.’ tapiroides chiefly belong to the ‘M.’ turicensis Schinz. Gaudry then 
described (op. cit., p. 155) and figured (Pl. xxtv, figs. 1 and 2) the jaws of a young specimen of ‘M.’ turicensis from 
Pikermi, including three superior and inferior molars figured together in Pl. xxtv, fig. 1 (Fig. 165 of present 
Memoir); the superior premolars enlarged (Pl. xxtv, fig. 3) as compared with Cuvier’s Lower Miocene type, 
the inferior premolars also enlarged (Pl. xx1v, fig. 4—cf. Fig. 165 of present Memoirfor Dp,), and superior view of the 
jaw (PI. xxrv, fig. 2—Fig. 165 of present Memoir). These Lower Pliocene grinders of Pikermi age referred to ‘M.’ 
turicensis appear to be somewhat more progressive than the Upper Miocene stage of ‘M.’ turicensis of Schinz. 
The several characters cited by Gaudry are enumerated below in our specific description of ‘M.’ turicensis in the 
present Memoir. See Pl. m1, pp. 134-135, diagrams of Turicius molar evolution. 

LoRTET AND CHANTRE, 1879.—The next great step was that of Lortet and Chantre (1879), when they brought 
together as Group II the following six species: 

Mastodon tapiroides Mastodon borsoni Mastodon vellavus 
i turicensis me buffonis * vialetiz 

This Group II corresponds in large part with Vacek’s Zygolophodon published two years earlier. Finally, 
in 1909, practically the same zygolophodont group of species was adopted by Mayet. 

M. Borsoni Hays. M. arvernensis Cr. et Job. 
M. Twricensis vy. Pikermi----------:--+-: 1 M. atticus Wagner -+---------------~----M. Pentelici Gaud. 

Form y. Baltavar. MM. longirostris Kaup. 

Formen aus dem Flinz der Isar, 
d. Sandsteine von Veltheim, 
d. Leythakalke. ? Steinheim. 

M. tapiroides Cuv. M. pyrenaicus Lart. M. angustidens Cuv. : 
: Halbjoche in gleicher Linie. Halbjoche alternirend. 
Zygolophodon Bunolophodon 
a a ee ee, ___ _ 

Definition and phylogeny of Zygolophodon (subgenus Bunolophodon), in which the characters of the grinding teeth of 
Zygolophodon and of Bunolophodon (Mastodon angustidens, M. arvernensis) are contrasted. After Vacek, 1877, p. 45. 

Vacrexk, 1877.—Meanwhile Michael Vacek, the Austrian paleontologist, in his Memoir “Uber Osterreich- 
ische Mastodonten,” Vienna, July 1, 1877, gave a complete revision of the mastodonts of Europe from the primi- 
tive period of the contributions of Baldassari (1767), of Pallas (1777), of Buffon (1784), of Ildephons Kennedy 
(1785), to the contributions of Biedermann (1876). He especially mentions the masterly memoirs of Etienne 
Borson (Turin, 1820, 1823), the palzeontologist to whom Hays dedicated his species ‘Mastodon’ borsoni in 1834. 

The most valuable parts of Vacek’s contribution are his illuminating reviews of Mastodon tapiroides Cuv. 
(including M. turicensis, pp. 4-6), of M. borsoni Hays (pp. 6-11) in comparison with M. angustidens Cuv. (pp. 
12-25), with M. longirostris Kaup (pp. 25-33), and with M. arvernensis Croizet and Jobert (pp. 33-39). In the 
case of each species valuable observations are given upon the known geographic distribution, as well as the 
resemblances and differences of the grinding teeth in relation to other species. Vacek sums up (pp. 39-45) the 
clear separation of his ‘Group’ Zygolophodon from his ‘Group’ Bunolophodon, as presented in his diagram herewith. 


We may by courtesy take Vacek’s group name as a valid definition of the genus Zygolophodon which he based 
on the species Mastodon borsoni, M. turicensis, and M. tapiroides (= M. pyrenaicus). 

Vacek’s diagnosis and diagram above embody a clear recognition of three distinct phyla corresponding 
with the three phyla of this Memoir, namely, the Zygolophodontine, the Longirostrinze (= Bunolophodon: ‘Halb- 
joche in gleicher Linie’’), and the Brevirostrine (= Bunolophodon: ‘‘Halbjoche alternirend’’). Vacek concludes 

Das Vorkommen von Uebergangstypen in einer Zeit, wo die beiden 

Formengruppen noch nicht so scharf geschieden waren, wie wir dies in Mastodon turicensis Schinz. 

den jiingsten Ablagerungen sehen, hat demnach nichts Auffallendes an 1825, Mastodon tapirotdes. Cuvier, Ossements fossiles, t. 1. p. 267, 
sich, ja dasselbe muss sogar von vornherein erwartet werden, und die Pr i iy! Cf. aussi Guettard, Mémoires, t. V1, 10° m., 
Trennung der Formen nach Massgabe ihrer phyllogenetischen [sic] Ent- 1847. Mastodon turicensis. Schinz, Naturg. und Abbild. d. Séuge- 

. eae ; 5 5 thiere, 1827, p. 243. 
wickelung oder die Trennung in Zygolophodonte und Bunolophodonte :8{a, Mastodon tuviecnete 1 von Weyne, Pabeoulalagten wal Basokt= 
ist sonach eine natiirliche. Wollten wir uns zum Schlusse nach den in : chte der Erde und ihrer Geschopfe, Francfort. 

ae eh . q i” : rier a: 1850, Mastodon tapirofdes. Blainville, Ostéographic, G. Eléphant. 
diesem Aufsatze gegebenen Gesichtspunkten einen Ueberblick tiber die 1859. Mastodon Borsonii. G. Gervais (p. part.), Zoologie et Paléonto- 
bisher bekannten europiiischen Formen der Gattung Mastodon schaffen, logie .frangaises, 2° éd., p. 68. 

2 = ; . 2 1878. Mastodon affinis. Jourdan, Archives du Muséum de Lyon, t. II. 

so kénnte man sich die Gruppirung derselben in folgender Art denken. 1878. Mastodon tapiroldes. Lortet et Chantre, Recherohes sur lés 

Mastodontes (Archives du Muséum de Lyon, t. Il, p. 285. 
3 : pl. VII et IX). 
Mayet, 1908.—Whereas Vacek (1877) and Schlesinger (1917, 1887. Mastodon turicensis. Depéret, Vertébrés miocénes de la vallée 
pets du Rhone (Archives du Muséum de Lyon, t.1V, p. 131). 
1922) adhere to the name ‘Mastodon taptroides : throughout the 1908. Mastodon turicensis. L. Mayet, Mammiféres miocénes des sables 

de l'Orléanais et des faluns de la Touraine (Annales de 

Miocene, Mayet (1908, p. 196) abandons the above name in favor [Université de Lyon, fasc. 24); == des sables de l’Orléanais, 
‘ . «yy a 4 Res p. 194, pl. VIII, fig. 1 et 2; == des faluns du Blésois 

of ‘Mastodon turicensis’ owing to the uncertainty as to Cuvier’s p. 298, pl. XI, fig. 4 et 5. 

lost type specimen. Mayet, on the other hand, sharply distin- Type and referred species related to the Mastodon 

; : : ce tapiroides, M. turicensis, M. borsoni groups. After Mayet, 
guishes the species Mastodon pyrenaicus Lartet from M. tapiroides 909, ». 45—all referable to the Zygolophodontine. 

(=M. turicensis) and points out that M. pyrenaicus occurs in the 

Lower Miocene, Burdigalian stage, as well as in the Middle Miocene horizon of the type specimen of Lartet. 
Compare also Mayet, 1909, p. 45, Section B, ““Mastodontes 4 Mamelons Disposes en Crétes Transversales 
(Type Lophodonte)”’ herewith. 

KAHN, 1922, 1931.—-Klihn, in describing (1922) the mastodonts of Baden and Rheinhessen, cites the 
species Mastodon wahlheimensis, which the present author regards as belonging to the Zygolophodontine. 
This is included, therefore, in the list of species on page 192 as Turicius wahlheimensis; it is not described, how- 
ever, in the present chapter, but is briefly treated together with other of Klihn’s species in Chapter VIII, Sec. 
11, “Miocene and Pliocene Mastodonts of Baden and Rheinhessen,” on pages 281-283. 

Compare Vou. II, CHap. XV, STEGODONTIN&H (SEE STEGOLOPHODON, PI. Iv, pp. 134-135) 

ScHLESINGER, 1917.—Schlesinger (1917) in describing his type specimen of Mastodon (Bunolophodon) longi- 
rostre Kaup forma sublatidens n.f., discovered near Teschen (Schlesien), Austria, proposes the new subgeneric 
name Stegolophodon for ‘Mastodon’ latidens Clift (see Schlesinger, p. 115, footnote) : 

Ich schlage fiir M. latidens [Clift], dassich durch seine kurze Symphyse von dem Subgenus Bunolophodon, durch seinen 

Molarenbau von Dibunodon entfernt, den Untergattungsnamen Stegolophodon vor. Der Name bringt einerseits die nahen 

Beziehungen zum Genus Stegodon, anderseits die Loslésung der Untergattung von Bunolophodon und ihre Sonderstellung 
gengentiber Dibunodon zum Ausdruck. 

Schlesinger’s type specimen of ‘Mastodon’ sublatidens from Teschen (Schlesien) certainly suggests the zy- 
golophodont form of molar, but it more strongly resembles Clift’s type of M. latidens from Burma, and stimulates 

(FIG. 138 C, Cl, FIG. 139, 4, 5, AND FIG. 140) 

Lower Miocene. Turicrus (4, 5) anp TRILOPHODON (2) 
Fig. 139. Turicius turicensis [=tapiroides] (upper 4, 5) 

compared with Trilophodon pontileviensis (lower 2), one-half 

natural size. After Mayet, 1908, PI. x1, figs. 2, 4, and 5. 


(Upper) Fig. 4.—‘‘ Mastodon turicensis [=T. tapiroides). 
Pontlevoy. Derniére molaire inférieure. Paris, Muséum. 
Grandeur naturelle.” Fig. 5.—‘‘7d. vue d’en haut.’”’ (See also 

Fig. 188. A, B, From the Sables de Fig. 138 C, C1.) 

VOrléanais, Lower Miocene; C, from Pontle- 
voy, Lower Miocene. After Mayet, 1908, PI. 
vu, figs. 3 and 5, Pl. x1, figs. 4 and 5, reduced 
to one-third natural size. 

These molar teeth exhibit the profound difference between 
the zygolophodont molar (above), resembling that of a primi- 
tive Stegodont with uninterrupted, widely open valleys, and the 
bunolophodont molar (below), in which the valleys are closed 

e . ‘ by the central conules. 
A, (Op. cit., Pl. vit, fig. 3): “3.—Mastodon 

angustidens, Chevilly (sabliére Cassegrain, (Lower) Fig. 2.—‘‘ Mastodon angustidens. Falun de Pont- 
au Glorieux, aoit 1877). Derniére molaire levoy. Derniére molaire supérieure. Communiquée par M. 
supérieure gauche. Paris, Muséum. Grandeur Jean de Bodard, Pontlevoy. Grandeur naturelle.” [=Trilopho- 


naturelle.’ = Miomastodon depereti sp. nov. don pontileviensis. 
P I 

B, “5.—Mastodon angustidens. Chevilly. 
Derniére molaire supérieure [H. F. O. inféri- 
eure]. Paris, Muséum, Grandeur naturelle.” 
{ = Trilophodon pontileviensis.| 

We observe that the T'rilophodon grinders 
are much narrower than those of T'uricius. 

C, Cl (Op. cit., Pl. x1, figs. 4 and 5): “4.— 
Mastodon turicensis |= T. tapiroides H. F. O.). REFERRED TURICIUS TURICENSIS MOLAR 
Pontlevoy. Derniére molaire  inférieure. Fig. 140. A lower molar, 1.Me, from Murinsel, Croatia, 
Paris, Muséum. Grandeur naturelle. 5.— referred by Vacek to Mastodon [=Turicius] tapiroides, but 
Id. vue d’en haut.”” (See Fig. 139 for en- which appears to be closer to the stage Turictus virgatidens. 
larged view of these teeth.) After Vacek, 1877, Taf. vu, figs. 4, 4a, one-third natural size. 

Observe that in Trilophodon pontileviensis central conules block the two anterior valleys; there are four conelets of unequal size in each loph. Observe in 
Turicius that all four of the valleys are open and free of central conules; each loph consists of five to six conelets, foreshadowing the ecto- and entoserrate 
lophs of the genotypic species Turicius turicensis (Fig. 162). In the Simorre stage, T'uricius simorrensis (Fig. 141) displays serrate spurs from the ectoconelets. 

The Lower Miocene Trilophodon angustidens of France (cf. Trilophodon pontileviensis, Chap. VIII, fig. 230) is paralleled by 7. cooperi in India. 

The Lower Miocene Miomastodon depereti sp. nov. of France is fully described in the Appendix of the present Volume I. 



our comparison of the type molar of Mastodon sublatidens Schlesinger with the type molar of Mastodon { = Stego- 
lophodon| latidens Clift. Schlesinger does not state whether he also applies his new subgeneric name Stegolophodon 
to his species sublatidens, but it is obvious that M. sublatidens and M. latidens belong to the same genus, i.e., 
Stegolophodon; consequently Osborn has referred Mastodon sublatidens to the subfamily Stegolophodontinz! 
(Chap. XV), where it is treated in detail as Stegolophodon sublatidens. See Pl. tv, pp. 134-135. 

ScHLossER, 1903.—Long prior to Schlesinger’s note, Schlosser suggested that primitive Asiatic species of 
Stegodon may have been derived from mastodonts related to the ‘Mastodon’ turicensis of western Europe. A remote 
resemblance may be observed between specimens from the upper Middle Miocene of Simorre, France (Fig. 141), 
referred to M. tapiroides by Lartet (made the type of Turicius turicensis simorrensis by Osborn in 1926), and 
specimens from the Middle Pliocene of Perim Island, India, Lydekker’s cotypes of M. cautleyi (see Fig. 142 A, B, 
Osborn’s lectotypes of Mastodon |= Stegolophodon] cautleyi). See Pls. u-1v, pp. 134-135 (Zygolophodon, Turicius, 
and Stegolophodon). 

Fig: 3 w. 

Fic. 230. — Derniére molaire inférieure du Mastodon luricensis (lapiroides), 
aux 2/5 de grandeur. — Miocene moyen de Simorre, Gers. (D’aprés Lartet. ) 

Turicrus (C) anp StegotopHopoON (A, B) Form or Grinpinc TEEeru 

Fig. 141. (C) Type third right inferior molar, r.Mg, of Turicius turicensis simorrensis, erroneously determined by Lartet (1859, Pl. xv, fig. 3) as Mastodon 
lapiroides, two-fifths natural size. Upper Middle Miocene of Simorre. After Gaudry, 1878, p. 174, fig. 230. Reversed in drawing. See also pages 207 and 220. 

Fig. 142. Cotypes of Mastodon [ =Stegolophodon| cautleyi Lydekker, 1884. Lectotypes (Osborn), compare Volume IT. 

A, First superior molar of the left side, 1.M’, one-third natural size. After Faleoner and Cautley, 1846 [1847, Pl. ux, figs. 3, 3a], as “ Mastodon latidens.”’ 
Length 4 inches, width 2.3 inches. Brit. Mus. M.2817. Cast Amer. Mus. 26965. Perim Island. 

B, Third superior true molar of the left side, 1.M*, one-third natural size. After Falconer and Cautley, 1846 [1847, Pl. xxx1, figs. 6, 6a] as ‘“ Mastodon 
latidens.” Length 8.5 inches, width 4.5 inches. Brit. Mus. M.2705. Cast Amer. Mus. 26966. Perim Island. See also Lydekker, 1886.1, p. 73, fig. 18. 

A further striking resemblance to the Stegodon type is seen in the molar teeth referred by von Meyer to M. 
turicensis from the Upper Miocene lignites of Elgg and Kapfnach (Fig. 162). Finally the M. virgatidens type of von 
Meyer (Fig. 168) exhibits a progressive evolution somewhat closely parallel with progressive species of Stegodon. 

It is not impossible that the forest-living Zygolophodon phylum of southern Europe may have given rise in 
Lower Miocene time to forest-living animals which spread eastward into the forests of southern Asia and developed 
into the Stegodont phylum. This whole phylogenetic problem is so important that it is treated more fully in 
Vol. II, Chap. XV, the Stegodontine, of the Stegodontoidea superfam. nov. 

OsBorN, 1935.—Osborn is inclined to favor Schlosser’s suggestion of 1903 and Schlesinger’s suggestion of 
1917 that primitive European species (Stegolophodon sublatidens) may have given rise to primitive species of 
Stegodonts of Asia to which Schlesinger has given the new generic name Stegolophodon. See Pls. mu-1v, pp. 134— 
135, also full treatment in Chapter XV (the Stegodontinz). 

'See Appendix of the present Volume I, pages 685 ef seq. 


As detailed above (pp. 192-197), beginning with Buffon in 1770 the Zygolophodontinz (Osborn, 1923.601) 
were clearly separated (Osborn, 1926.706) from the Mastodontine chiefly by (1) multiplying conelets, (2) increas- 
ing sharpness of the ridge-crests, (3) open central valleys, and (4) absence of central conules and reduction of sulcus. 

ScHLESINGER, 1917—1922.—In his detailed researches on the morphology, phylogeny, environment, and 
stratigraphy of the European mastodonts, Schlesinger gives repeated illustrations and summaries of the 
profound differences which exist between the species described in Europe as Mastodon borsoni and the species 
described as Mastodon tapiroides, the latter including the M. turicensis Schinz. We may therefore credit to 
Schlesinger (1922) the observation of clear and sharp distinctions not only between M. americanus and M. borsona, 
sufficient to place them in different phyla, but of still more importance the sharp distinctions between specimens 
referred to M. borsoni and those referred to M. tapiroides-M. turicensis. 

Osborn, 1927: Finally in studying the detailed structure and geologic stages of evolution of these Miocene and 
Pliocene Zygolophodonts, Osborn independently (1926.706, p. 3) concluded that they represent two genera, the 
included species of which may be given here as follows: 

Zygolophodon Vacek, 1877 Turicius Osborn, 1926 
Zygolophodon borsoni type, of Asti, Italy Turicius virgatidens type, of Fulda, Germany 
Turicius atticus type, of Pikermi, Greece 
Turicius turicensis type, of Elgg, Switzerland 

Zygolophodon pyrenaicus, Ile-en-Dodon, France Turicius turicensis simorrensis type, of Simorre, France 
Turicius turicensis ref., of Pontlevoy, France 
Zygolophodon pyrenaicus aurelianensis, Sables de Turicius tapiroides type, Caleaire de Montabusard, 
lOrléanais, France: France 

Whereas Vacek included Mastodon turicensis in his group Zygolophodon, Osborn points out that the type and 
referred specimens of Twuricius, extending from the Lower Miocene of Pontlevoy to the Middle Pliocene of 
Germany, are clearly separable from the type of the true Zygolophodon and constitute a distinct ascending series 
of species with certain resemblances in the grinders to the species grouped under the genus Serridentinus Osborn. 
They thus constitute the types of a distinct genus Turicius typified by the typical Mastodon turicensis of Schinz 
but embracing a whole series of stages extending from the Lower Miocene to the Middle Pliocene. 


It appears that the Zygolophodon of Vacek divides into two generic phyla. Before we can clearly summa- 
rize the generic and specific characters of Zygolophodon and Turicius and the species included within them, it is 
necessary to concentrate our attention on the type geologic locality and level and the characters of the type grinding 
teeth. Previous failure in this respect has led to great confusion which exists even to the present time. 

Geologically the oldest type is the Turicius tapiroides Cuv. from the Lower Miocene lacustrine Caleaire de 
Montabusard, France, but there is no certainty as to the precise characters of the type specimen. Geologically 
the most recent type is the Zygolophodon borsoni from the Middle to the Upper Pliocene of Asti, Italy; al- 
though there is some reason to believe that this species may have survived in eastern Europe into Lower Pleisto- 
cene time; it certainly disappeared in western Europe at the close of Pliocene time. 



Tyre Locauities.—The summary of the chief type localities, widely scattered through northern Italy 
Switzerland, Germany, and France, is as follows: 

Types Locauiries ORIGINAL NAME Speciric REFERENCE IN 

Middle Pliocene, Villanova, Asti, Italy Type of Mastodon borsoni Hays, 1834 = Zygolophodon borsoni 
Middle Pliocene, Fulda, Frankfort, Ger- Type of Mastodon virgatidens von 

many Meyer, 1867 = Turicius virgatidens 
Lower Pliocene, Pikermi, Greece Type of Mastodon atticus G. & L. (in 

Wagner, 1857) = Turicius atticus 

Upper Miocene, Elgg, Canton Zurich, Type of Mastodon turicense Schinz. 

Switzerland 1824 = Turicius turicensis 
Middle Miocene, Ile-en-Dodon (Haute- Type of Mastodon pyrenaicus Lartet, 

Garonne), France 1859 = Zygolophodon pyrenaicus 
Lower Miocene, Caleaire de Montabusard, Type of Mastodon tapiroides Cuvier- 

France Desmarest, 1806-1822 =Turicius tapiroides 

Speciric CHaracTers.—Having thus distinguished the type geologic levels and localities, let us clearly distin- 
guish the characters of the type specimens in descending geologic order; these characters are based solely on the 
type grinding teeth, since our knowledge of the incisive tusks and of the characters of the jaws is very limited. 
Doubtless the tusks exhibit other distinct characters. See Pls. m and m1, pp. 134-135. 

Type M?*, Mastodon borsoni (see Figs. 154 and 111A, after Hays and Buffon) =Zygolophodon 
Third superior molars tetralophodont (4—4% crests), each loph composed of four! distinct and subequal conelets (‘mamelons’). 
Type M;, Mastodon virgatidens (see Fig. 168, after von Meyer) = Turicius 

Intermediate molars trilophodont, posterior molars subtetralophodont, lophs acute with five to nine connate conelets (‘ Wulst- 
kanten,’ ‘mamelons’); a serrate spur on superior internal conelets. 

Type M?, Mastodon turicensis (see Fig. 162, after Schinz) = Turicius 
Second superior and inferior molars trilophodont, each loph with five to siz more or less distinct conelets, no median 
sulcus; ridge-crests slightly arched or directly transverse; strong serrate spur on superior internal conelets. 

Type M;, Mastodon pyrenaicus (see Figs. 147, 148, after Lartet) = Zygolophodon 
Third inferior molar tetralophodont plus, i.e., 44 crested, each loph composed of three to four! more or less distinct conelets; 
no median sulcus; no serrate spur on external conelets; vestigial conules in anterior valleys. 

Type r.M; of Mastodon tapiroides (see Fig. 160, after Cuvier, Fig. 161, after Mayet, also Pl. m1 A, pp. 134-135). = Turicius 
Third inferior grinders tetralophodont, each loph composed of five to siz conelets; no median longitudinal suleus; vestiges 
of conules in the middle of the valleys; trefoil spur apparently on metalophid. 

GENERIC CHARACTERS.—From the above principal characters of the type grinding teeth we observe that the 
universal generic character both of Zygolophodon and of Turicius is the absence or reduction of the longitudinal 
sulcus, fissure, or commissure, which Hays pointed out as the chief character distinguishing his type (Fig. 154) 
of M. borsoni from M. americanus. The absence or reduction of this sulcus distinguishes all the ascending species of 
Zygolophodon and of Turicius, from early Miocene to late Pliocene or early Pleistocene time, from all the ascending 
or ancestral stages of Mastodon in the corresponding period. We also observe that the fission of the cones into 
‘conelets’ is rapidly progressive and distinct in Zygolophodon and in Turicius, whereas it is decidedly retarded in 
Mastodon. In many other features (compare Schlesinger, 1922) the progressive dental and skeletal evolution of 
Zygolophodon parallels that of Mastodon, as described in the previous chapter, namely, (1) the abbreviation of 
the jaws, (2) the suppression of the premolar dental succession, (3) the reduction of the lower incisive tusks, (4) 
the progressive abbreviation of the cranium. (5) From the fact that the Zygolophodon molars are relatively 
broader and shorter than those of Mastodon, it is probable that the cranium of Zygolophodon was more brachy- 
cephalic than that of Mastodon. 

We may therefore give the following provisional generic distinctions of Zygolophodon and of Turicius from 
Mastodon, which will be amplified as we gain a more thorough knowledge of the skeleton. See also the full 
distinctions cited below (p. 211) from Schlesinger, 1922; also compare Mayet, 1908 (p. 204 of the present Memoir). 

1Four conelets in types of Zygolophodon pyrenaicus and Z. borsoni; four to six conelets in progressive stages (Pl. 1 D-G). 





Genotypic species: Elephas [= Mas- 
todon| americanus Kerr. 

1. Longitudinal sulcus persistent 
between internal and external cones. 

2. Main cones feebly subdividing 
at the summits into two conelets, pro- 
gressive to four (Fig. 89, Miomastodon). 

3. Retarded ‘trefoil spurs’ on supe- 
rior internal cones and on inferior ex- 
ternal cones. 

4. Grinders, Dp?-M®, relatively 
longer and broader, breadth-length 
index of M; not exceeding 57 (highest 
recorded index in Amer. Mus. speci- 
mens, Amer. Mus. 14345)—60. 

5. Upper incisive tusks rounded, 
enamel band persisting into Miocene 

6. Lower incisive tusks straight, 
cylindrical, variable. 


Genotypic species: Mastodon borsoni 

1. Longitudinal sulcus disappear- 
ing, vestigial. 

2. Main cones strongly subdivid- 
ing into four to five subequal conelets, 
progressive to six (Fig. 151 and PI. 1). 

3. ‘Trefoil spurs’ vestigial or absent. 

4. Grinders relatively shorter and 
broader, index of M;=43—-45. 

5. Rudimentary conules or tubercles 
present in median valleys in early 
stages, i.e., Zygolophodon pyrenaicus ref. 

6. Upper incisive tusks rounded, 
enamel band disappearing early, before 
Lower Miocene time. 

7. Lower incisive tusks reducing 
early. Symphysis more abbreviated. 


Genotypic species: Mastodon turt- 
censis Schinz. 

1. Longitudinal sulcus disappearing, 

2. Main cones subdividing into five 
to nine conelets, i.e., summits of lophs 
with quintuple, to sextuple, to octuple, 
to nonuple conelets (compare Serri- 
dentinus serridens, Figs. 388 and 389). 

3. Strong ‘trefoil spurs’ on superior 
internal and inferior external cones, 
hence ‘serridentine.’ 

4. Grinders relatively longer, in- 
creasing in breadth. 

5. Central conules vestigial in the 
median valleys in early stages (T. 

6. Upper incisive tusks oval; en- 
amel band sharp, persisting late, absent 
in T. virgatidens. 

7. Lower incisive tusks suboval, 
vestigial in Pliocene stage. Symphysis 
less abbreviated. 

GENERIC DISTINCTIONS INTO Two GRouPs.—The Mastodontine so far as we know embrace one phylum of 
species, the Zygolophodontine, however, appear to embrace two phyla or groups (I and II) of species as follows: 

I. In Zygolophodon pyrenaicus, type and referred, and in Z. borsoni, each loph of the grinding teeth sub- 
divides into not more than four, to four and a half, to five to six conelets; the lophs are directly transverse not 
arched; trefoil spurs vestigial or absent. See Plate 1. 

Il. In Turicius turicensis and T. virgatidens, each loph subdivides into from five to nine to twenty-five conelets; 
strong trefoil spurs arise on the superior internal conelets and on the inferior external conelets. See Plate 11. 

Consequently we appear to observe two ascending contemporaneous generic and specific phyla which do not 
intergrade. Geologically these phyla show two independent lines of ascent. 


I. Zygolophodon le 

Four to six persistent conelets on each 

grinding crest, ‘trefoil spurs’ vestigial 
or absent 


Five to nine conelets on each grinding 
crest, progressive to twenty-five, 
‘trefoil spurs’ more or less prominent 

Upper Pliocene 

Middle Pliocene 

Lower Pliocene 

Zygolophodon borsoni ref. 

Zygolophodon borsoni type, Asti, Italy Turicius virgatidens of Fulda, Germany 

Turicius atticus of Pikermi, Baltavar, 
Upper Miocene Stage undescribed Turicius turicensis of Elgg and Kapf- 
nach, Switzerland 
Turicius turicensis of Sansan and 
Simorre, France=T7'. simorrensis 
Turicius turicensis, Faluns de Pont- 
levoy, France 
Turicius (?) tapiroides, 
Montabusard, France 
Parallel evolution with Stego- 
lophodon and Stegodon phyla 

Middle Miocene Zygolophodon pyrenaicus, Ile-en-Dodon, 


Lower Miocene 

Lower Miocene Zygolophodon pyrenaicus aurelianensis, 

Sables de ]’Orléanais, France 

Caleaire de 

Parallel evolution with Mastodon 
americanus phylum 



Undoubtedly constituting a phylum (Group II) distinct from above 


species of Zygolophodon 

(Group I) are the numerous grinding teeth referred by all recent European authors, especially Mayet, 1908, to 
‘Mastodon’ turicensis, first observed in the Lower Miocene Faluns de Pontlevoy and Sables de |’Orléanais, again 

Fig. 143. 

Middle (=Etage Helvétien) and Upper (=Etage Tortonien) Miocene 
After Depéret (1905-1908) and Osborn (1910.346, p. 258, fig. 129). 

Horizons 1-69 arranged in ascending geologic order. 


Horizon of Sansan. FRANCE.—Calcaire 
de 1 Sansan (Gers), lacustrine. 2 Jegun (Gers). Faluns de 3 Pontlevoy, 4 Sainte-Maure, 
5 Manthelan, in the basin of the Loire, marine. Mollasse de 6 pont de l'Herbasse, 7 Bren, 
8 Clérieux, near Romans, marines GERMANY.—Meeresmolasse (in Swabia) von 9 Baltrin- 
gen, Rammingen, Heggbach, Ursendorf, Hochgeland, 10 Hausen, 11 Niederstozingen, 12 Siisscn. 
Stisswasserkalk von 13 Georgensgmiind, 14 Engelswies (Bavaria). AUSTRIA-HUNGARY. — 
Braunkohle (in Styria) von 15 Eibiswald, Wies,16 Gériach, Gamlitz, 17 Voitsberg, Parschlug, 
Neufel, Leoben, Leiding, Feisternitz. Marine sands of Grund at 17a Guntersdorf (Lower 
Austria). Horizon of Simorre. FRANCE.—2 Bonnefond, St. Cristan (Gers). 2a Villefranche 
d'Astarac, calcaire de Simorre (Gers), lacustrine. Cinérites de 18 Tournon, in south central 
France. 19 /'Isle-en-Dodon (Gers). 20 Saverdun (Ariége). Mollasse de 21 Mirabeau (Vau- 
cluse), marine. Marine deposits of 22 Sorgues (Vaucluse). Marine deposits of 28 Romans 
(Dréme). Sidérolithiques de 24 La Grive-Saint-Alban (in large part) (Istre), 25 Mont Ceindre 
(Rhone), 26 Pretty near Tournus (Saéne-et-Loire), 27 Gray (Haute-Saéne). GERMANY.— 
In Swabia, Bohnerz von 28 Willmardingen, Jungnau (in large part), Siisswasserkalk von 
Steinheim, Ries, Urlau, 29 Nérdlingen, 30 Althausen. Bohnerz von $1 Mésskirch, 32 Genkin- 
gen, 33 Heuberg, 34 Melchingen. Gips von 85 Hohenhéven (Baden) (10 meters). AUSTRIA- 
HUNGARY. — In the intra-Alpine basin of Vienna, 36 Dornbach, Vordersdorf, Loretto, Marga- 
rethen, 37 Filnfkirchen (Hungary), calcaire de 38 Bruck-a.-Leitha; 39 Breitenbrunn, Abstdorf- 
Franzensbad, 40 Mannersdorf, 41 Neudorf. 42 Trauenzinen (Silesia). 43 Krivadia (Transyl- 
vania). 44 Gyulu-Mendru (Transylvania). 45 Pesth (Hungary). 46 Ssoskut (Hungary). 
RUSSIA. —47 Wosskressensk. Horizon of Saint-Gaudens. FRANCE.—48 Valentine, Saint- 
Gaudens (Haute-Garonne). 49 Montréjau (Haute-Garonne). SWITZERLAND.—Sands of 
50 Delsberg, near Basel. 51 Le Locle, in the northwest. 52 La Chauz-de Fonds, in the north- 
west. 53 Vermes, near Basel. Mergelkalke von 54 @ningen, near-Ziirich, fresh water. Lig- 
nites of 55 Elgg, 56 Kapffnach, near Ziirich. 57, Veltheim, near Ziirich, GERMANY.—In 
Bavaria: 11 Giinsburg; Reisensburg ; Sande von 58 Hdder, Dinkelscherben, Reichenau, 59 Dies- 
sen, 60 Dasing, Fraising, Tutzing, Stdtzling, Reichertshofen, 61 Frontenhausen, Flinz of Munich. 
62 Sankt Georgen (Baden). AUSTRIA.— 36 Heiligenstadt, Tegel von Hernals, in Vienna 
basin. ITALY.— Lignites of 63 Mt. Bamboli (Tuscany). SPAIN.—64 San I/sidro, near 
Madrid. PORTUGAL.—65 Aveiras de Baixo, near Oporto. RUSSIA.—66 Kriwoi-Rog 
(Kherson). 67 Nicolaicf, at the mouth of the Dnieper. 68 Sebastopol (Crimvu). 69 Tiraspol, 
near the mouth of the Dniester, Correlation of Depéret. 


2400 mm. 7’10Vve"e 


" 2600MM., 8/64” e 



ZYGOLOPHODON Shek Vee ee ae RELIANENSIS 2725mm,Bile 

Fig. 143a. Ascending x behiciaie succession of three 
species of Turicius and one of Zygolophodon. Compare Appendix. 

4. Turicius virgatidens of Fulda, Germany. Middle Plio- 

3. Turicius turicensis of Elgg, Canton Zurich, Switzerland 
(see Fig. 143, 55). Upper Miocene. 

2. Turicius tapiroides, Calcaire de Montabusard, France. 
Lower Miocene. (See Fig. 146, 6). 

1. Zygolophodon pyrenaicus aurelianensis, Sables de |’Or- 
léanais, France. Lower Miocene. 

These outlines, restored to a one-hundredth scale, are chiefly 
after the grinding teeth and tusks, the skeleton and skull being 
relatively unknown or undescribed at the present time. 

in the Middle Miocene of Sansan and of Simorre, again in the Upper Miocene of Elgg and of Kapfnach, and 

finally in the Lower Pliocene of Pikermi and of Baltavar. 

These animals show the strong upper internal and 

lower external serrated ‘trefoil spur’ (‘Wulstkante’ of von Meyer) also characteristic of Serridentinus; in fact, 
the grinding teeth show certain resemblances to the species included within the genus Serridentinus, but the 

lower incisive tusks of Turicius are entirely different from those of Serridentinus. 

Von Meyer observes that the 


lower incisive tusks of ‘M.’ turicensis are reduced and that the upper incisive tusks are oval in section and exhibit 
an enamel band in the specimens referred to M. turicensis from Kapfnach. In brief, there are many strong 
resemblances between the grinders of Turicius turicensis and the grinders of Serridentinus; consequently we are 
disposed to think that there may be an affinity of the genus Turiczus to Serridentinus which may be demon- 
strable by fuller knowledge. 

These animals certainly constitute a distinct genus, to which the name Twuricius is applicable. Whereas 
Vacek defined Zygolophodon (1877) on three species, namely, Mastodon borsont, M. turicensis, and M. tapi- 
roides (=M. pyrenaicus), the genus Zygolophodon would now rest solely on the following species, Mastodon 
borsoni and M. pyrenaicus. 

Osborn, 1935: In the Appendix of the present Volume are described two specimens of Turicius turicensis 
in the Munich Museum, which afford a fuller knowledge of the dentition, mandibular rami, and fore- and hind- 

limbs (see also Pl. m1, pp. 184-135). 

Pontier and Anthony (Dec. 2, 1929) describe the right and left premaxillaries of a cranium referred to 
‘Tetrabelodon’ turicensis Schinz, of the following geologic age: ‘‘Villefranche d’Astarac, dans le Gers-Calcaire de 
Simorre-Helvétien-Miocéne moyen.’ This interesting specimen, reconstructed in their figure 2 (op. cit., p. 1006— 

Fig. 144. Reconstruction by Pontier and Anthony, 1929, Fig. 145. Cranium of Lozxodonta africana, from Fort 
fig. 2, of the head of Turicius turicensis Schinz, of Villefranche Sibut, Equatorial Africa, exhibiting reduplication of the 
d’Astarac, exhibiting the supplementary superior incisive tusks; superior incisive tusks. After Anthony and Prouteaux, 
inspired by Abel’s restoration of Trilophodon angustidens. 1929, Pl. 1, fig. 1. 

reproduced herewith as Fig. 144), contains on both sides a supplementary superior incisor, which emerges im- 
mediately above the normal superior incisor (I?). This was regarded by Pontier (1926, p. 162) as representing a 
vestige of, or reversion to, the first superior incisor (I) of the ancestral proboscideans. As to this question of 
homology, it is doubtful whether this anomalous tooth represents such a reversion, or a duplication of the crown 
of the normal incisor (I?). This tooth has a diameter of 36 mm. as compared with 112 mm. in a normal tooth; it 
curves upwards rather than downwards; it is certainly an anomaly and not a specific character. 

Comparison may be made with an analogous Lozxodonta africana in the duplication of the superior incisors, as 
described by Anthony and Prouteaux (1929), of a wild elephant found in the Oubangui Chari region in French 
Equatorial Africa. In this instance the paired superior tusks, beautifully illustrated by the authors (PI. 1, figs. rv), 
emerge above the normal tusks and are directly horizontal in position, while the length nearly equals the normal 
(Fig. 145). According to the authors, such anomalies are not extremely rare among African elephants but they 
are decidedly exceptional; they are regarded by the natives as princes of the elephant tribe. We accept the inter- 
pretation of the authors (op. cit., pp. 30, 31) that these anomalous teeth correspond with the second incisors (I?) 
of Meritherium. 




Genus: ZYGOLOPHODON Vacek, 1877 

Original reference: Vacek, Abh. geol. Reichsanst., VII, Heft IV, p. 45. Wien. 

Now resting on the genotypic species Mastodon borsoni and its apparently ancestral form Mastodon pyrenaicus. Compare Mastodon 
{=Stegolophodon] latidens of Burma, and Mastodon [=Stegolophodon] sublatidens Schlesinger of the Middle(?) Pliocene of Teschen (Schle- 
sien), Austria. The genus Stegolophodon Schlesinger, 1917, proves to be distinct from the genus Zygolophodon Vacek. (See Plates 
m-iv, between pp. 134-135.) 

GENERIC CHARACTERS OF ZyGOLOPHODON.—(1) Each loph divided into four to six distinct subequal 
conelets. (2) No trace of median longitudinal sulcus. (3) Conules or tubercles in median valleys disap- 
pearing in early stages. (4) Crests, i.e., lophs, directly transverse, not arched. (5) ‘Trefoil spurs’ vestigial 
or absent. (6) Fifth inferior crest, i.e., pentalophid, slowly progressive. (7) Superior incisive tusks 
rounded without enamel band (Z. borsoni). (8) Grinders permanently blunt, brachyodont (Z. borsoni), 
not becoming subhypsodont. (9) Progressive adaptations similar to those of Mastodon americanus 
rather than to those of T’uricius. Compare generic characters listed by Osborn above (p. 200) and by 
Schlesinger below (pp. 210, 211, 217). (10) No inferior enamel band (Z. pyrenaicus). 

The history of Vacek’s generic name is related above in the present chapter. Vacek founded the genus 
Zygolophodon on three species, Mastodon borsoni, M. turicensis, M. tapiroides (=M. pyrenaicus). Osborn (1926.706) 
removed the species M. turicensis and made it the type of a new genus, Turicius; he also now removes the 
species M. tapiroides' Cuvier to the genus T'uricius. 

(1) The chief genotypic species is Mastodon borsoni' Hays, Middle Pliocene of Asti, Italy. This stage is very 
distinctly characterized by the absence of a longitudinal suleus between the internal and external cones 
and by the presence of four distinct conelets in each loph, as clearly displayed in Hays’ type figure reproduced in 
facsimile (Fig. 154) below; many teeth from different parts of Europe have been referred to M. borsoni by various 
authors which lack the typical characters of the M. borsoni grinders; certain of the grinders figured by Lartet exhibit 
four and a half to five conelets in one or more of the crests. 

(2) Fortunately Lartet’s type of Mastodon pyrenaicus (Figs. 147, 148) from the Middle Miocene of Ile-en- 
Dodon clearly exhibits four distinct conelets also tubercles in the first median valley. Mayet (1908, p. 197) also 
records it in the Lower Miocene, Sables de |’Orléanais, to which stage Osborn (1926.706) assigned the new sub- 
specific name Zygolophodon pyrenaicus aurelianensis. 

(3) UNCERTAINTY AS TO M. TAPIROIDES TYPE SPECIMEN.—The type molar of Mastodon tapiroides Cuvier from 
the Lower Miocene Caleaire de Montabusard, France, has been lost and there is a difference of opinion as to 
whether it is a true molar, or a deciduous premolar (Dp*, Dp,, von Meyer, Gaudry): Cuvier’s figure reproduced in 
facsimile below (Fig. 160) certainly shows five main conelets in each transverse crest and intermediate tubercles 
or conules in the first and second valleys. The anterior portion of the crown is broken off. Mayet’s new figure 
(Fig. 161) of the type shows five conelets and a ‘trefoil spur,’ which relate this specimen to T'uricius; he regards it 
as ancestral to Twricius (see p. 205). Osborn (1935) determines this molar as an r.M; (see Pl. 1m A). 

1 Mastodon tapiroides was selected by Matthew as the type of Zygolophodon Vacek, 1877 (see Matthew, 1918.1, p. 200, footnote). 


As von Meyer and Gaudry observed, the type appears small for a true molar. It is possible that when the 
type of M. tapiroides and other molar teeth from the Calcaire de Montabusard are studied, they may be found to 
exhibit characters other than those represented in Cuvier’s type figure, on which our present description of this 
species rests. As independently observed by von Meyer and Gaudry, all the early French writers, including 
Lartet (1859), erred in referring similar specimens to Cuvier’s species M. tapiroides. Von Meyer on the other 
hand states that there is no resem