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Asian Herpetological Research 2015, 6(2): 73-131 
DOI: 10.16373/j.cnki.ahr.140066 

Herpetological Diversity of Timor-Leste: Updates and a Review of 
Species Distributions 

Mark O'SHEA!, Caitlin SANCHEZ’, Andrew KATHRINER’, Sven MECKE', Venancio 

! Faculty of Science and Engineering, University of Wolverhampton, Wulfruna Street, Wolverhampton WVl 1LY, United 
Kingdom; and West Midland Safari Park, Bewdley, Worcestershire DY12 LF, United Kingdom 

* Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, California 92395, USA 

* Department of Biology, Villanova University, 800 East Lancaster Avenue, Villanova, Pennsylvania 19085, USA; 
present address: Department of Herpetology, Bronx Zoo, 2300 Southern Boulevard, Bronx, New York 10460, USA 

* Department of Animal Evolution and Systematics, Faculty of Biology, Philipps Universitat Marburg, Karl-von-Frisch- 
Strafe 8, 35032 Marburg, Germany 

` Universidade National Timor-Lorosa e, Faculdade de Ciencias da Educaçao, Departamentu da Biologia, Avenida 
Cidade de Lisboa, Liceu Dr. Francisco Machado, Dili, Timor-Leste 

* Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 
20013, USA 

Abstract We report the results of five herpetological surveys during 2011—2013 that included visits to all districts 
of Timor-Leste (Aileu, Ainaro, Baucau, Bobonaro, Dili, Covalima, Ermera, Lautém, Liquicga, Manatuto, Manufahi, 
Viqueque) except the Oecusse exclave. Our fieldwork culminated in the discovery of one putative new frog species 
(genus Kaloula), at least five putative new lizard species (genera Cyrtodactylus, Cryptoblepharus, and Sphenomorphus), 
and two putative new snake species (genera Stegonotus and Indotyphlops). In addition, we present new distribution 
records of amphibians and reptiles for 11 of the country's 12 contiguous districts, along with additional natural history 
data. Results from our surveys increase the number of amphibian and reptiles known to occur in Timor-Leste from 22 
species before our surveys began to over 60, including over 20 as yet undescribed species. 

Keywords Timor-Leste, Kaloula, Cyrtodactylus, Eremiascincus, Stegonotus 

1. Introduction Island, a volcanogenic island geographically positioned 

in the Inner Banda Arc and situated ~25 km north of 

Timor-Leste (Figure 1) comprises four separate land 
areas, (1) the eastern half of Timor Island in the Outer 
Banda Arc of the Lesser Sunda Archipelago, with an 
area of approximately 14,480 km’; (2) the small (12 km’) 
uninhabited Jaco Island, a landmass of corallogenic origin 
lying ca. 750 m off the coast of Timor's easternmost 
point; (3) the much larger (105 km’) inhabited Ataüro 

" Corresponding author: Dr. Hinrich KAISER, from Victor Valley 
College, USA, with his research focusing on the diversity, morphology, 
and conservation of Southeast Asian amphibians and reptiles. 

E-mail: hinrich.kaiser(g) 

Received: 24 September 2014 Accepted: 11 February 2015 

mainland Timor-Leste's northern coast at the capital city 
Dili; and (4) the Oecusse District, an exclave (810 km’) 
on the northern coast of Timor, 53 km air-line distance 
west of contiguous Timor-Leste and surrounded on all 
landward sides by Indonesian West Timor. Timor-Leste's 
position at the southeastern edge of Wallacea makes 
the country an interesting area from a biogeographical 
perspective, as it is inhabited by a mosaic of faunal 
elements with either Southeast Asian or Australo-Papuan 
origin (see Monk et al., 1997). Due to its turbulent history 
as the Portuguese colony farthest from the mother country 
(nominally since 1515; West, 2009) and because of a 

74 Asian Herpetological Research 

Vol. 6 

violent annexation by Indonesia between 1975 and 1999, 
the area could not be properly surveyed until after Timor- 
Leste regained independence in 2002. A summary of the 
geography, geology, and habitats of Timor-Leste and a 
history of herpetological collecting in the country since 
the early 19" Century, was presented by Trainor (2010) 
and Kaiser et al. (2011). 

Timor-Leste is a country with very diverse habitats 
(Figure 2), ranging from beaches and rocky shores 
(Figure 2A) to montane meadows and mountains (e.g., 
Figure 2E). Much of the habitat has been altered by the 
presence of humans to a greater or lesser degree, ranging 
from active agricultural sites (e.g., Figure 2C) to patches 
of old-growth forest used by livestock (e.g., Figure 2D). 
The most pristine habitats include those demarcated 
by precipitous slopes (e.g., Figure 2E) or periodically 
inundated areas (e.g., Figure 2F), as well as those of 
particular cultural or religious significance where human 
alterations are prohibited (e.g., Figure 2G, H; pers. obs.). 
As we reported previously (Kaiser et al., 2011), it appears 
that the herpetofauna of Timor-Leste has shown some 






resilience to disturbance, and species diversity may be 
high locally despite low-level human disturbance, and 
even after the dramatic shift from primary tropical forest 
to coffee forest. 

Beginning with an initial survey in 2009, we have 
been conducting fieldwork in all 13 districts of Timor- 
Leste under the banner of the Victor Valley College 
Tropical Research Initiative. The present report on the 
field seasons of 2011—2013, with the addition of some 
more limited surveys conducted by AVR, LLA, and ZAS, 
supplements our reports for 2009 (Kaiser et al., 2011) and 
2010 (O’Shea et al., 2012). Reports for the politically and 
geographically isolated Oecusse District (Sanchez et al., 
2012) and Ataüro Island (Kaiser et al., 2013b) have been 
published elsewhere. 

2. Methods 

Surveys were conducted during both the wet season 
(Phase IV: 18 January-6 February 2011; Phase VI: 24 
January-7 February 2012) and the dry season (Phase V: 


(East Timor) 


Figure 1 Map of Timor-Leste and its position in the Lesser Sunda Islands. Numbered localities are listed in Table 1. 

No.2 Mark O'SHEA et al. 
19 June-5 July 2011; Phase VII: 21 June-10 July 2012; 
Phase VIII: 18 June-2 July 2013). Shorter wet season 
surveys were also conducted by ZAS, LLA, and AVR 
(11-14 October 2010, 10-12 November 2010, and 7 
January 2011, respectively). During 2011—2013, fieldwork 
was carried out at 35 main localities (Table 1) with 
smaller sub-localities clustered around some of these. The 

Timor-Leste Herpetofauna Updates 75 

general methods applied during fieldwork, the preparation 
of voucher specimens, and any associated scientific tasks, 
follow the protocols detailed by Kaiser et al. (2011). Most 
roadkills, depending on their state of decomposition, 
were skin- or scale-snipped to obtain tissue samples 
for molecular studies. All vouchered specimens 
have been deposited in the United States National 

Table 1 List of localities surveyed by the Victor Valley College Herpetofaunal Survey of Timor-Leste during Phases IV-VIII (2011—2013). 
Each locality includes a superscripted Roman numeral to indicate during which phase they were surveyed (locations only visited during 
Phases I-III, on Ataüro Island, or in the Oecusse exclave are omitted (for these locations see Kaiser et al., 2011, 2012; O’Shea et al., 2012; 

Sanchez et al., 2012). 

nd District Locality E cu GPS coordinates! 
[=H Dili W Dili (Timor Lodge Hotel; Comoro; Tasi Tolu) 2-25 S 08°33' E 125°31' 
ae Dili E Dili (Becora; Cristo Rei) 20 S 08°33' E 125°35' 
go en Dili Metinaro mangrove swamp 1 S 08°31' E 125°47' 
4" Dili S Dili (Dare) 545 S 08°36' E 125°32' 
S. Dili Comoro River (Beduku) 60 S 08°35' E 125°32' 
6" vm Dili, Liquiga, Aileu Comoro River (confluence with Bemos River)” 115 S 08°37' E 125°32' 
7" Aileu Lake Be Matin 1105 S 08°42' E 125232" 
grum Ermera Eraulo (Meleotegi River; Sta. Bakhita Mission) 1100-1250 S 08°47' E 125°27' 
9" Bobonaro Balibo (Fiuren) 463 S 08°57' E 125°04' 
10 Bobonaro W Maliana (Ramaskora; Soto River) 196—230 S 08°59' E 125?12' 
117 Bobonaro E Maliana (Maganuto, Mt. Leolaco) 1040—1063 S 08°59' E 125?16' 
12" Bobonaro E Maliana (Galosapulu swamp) 712 S 09?01' E 125?16' 
13h Covalima Suai & surrounds (Castelo Fronteira Guest House) 30-53 S 09?19' E 125?15' 
14" Covalima Kasabauk rice-paddies 9 S 09*24' E 125°09' 
15 Covalima Tilomar (Tilomar Forest Reserve; Maubesi; Mt. Debululik) 260—900 S 09?20' E 125?06' 
16^ Ainaro Maubisse (Pousada Maubisse) 1495 S 08°50' E 125?36' 
17^ Manufahi Same (Ailelehun Guest House; Trilolo River; Ladiki; Mirbuti) 340-1200 S 09?00' E 125°39' 
187 * Manufahi Betano (Dry site; Wet site) 20-44 S 09°10' E 125°42' 
19 Y vr Manufahi Fatucahi (Convent of St Antony d'Lisboa; Lake Lenas) 36-38 S 09?02' E 125?59' 
20^ Manatuto Nancuro, Natarbora, S Umaboco 3 S 09?02' E 126°04' 
219. Viqueque N Ossa (Liamida; Mt. Mundo Perdido) 930-1160 S 08°44' E 126°22' 
22k Baucau Venilale caves, N Venilale 675 S 08°37' E 126°23' 
23"! Baucau Uatubala, S. Afacaimau (Carlia spot) 370 S 08°33' E 126°26' 
24! Baucau Baguia (Vila Rabilhi Guest House; Pousada de Baguia) 440 S 08?38' E 126?39' 
25 Baucau Ossohuna (Ossohuna; Afaloicai) 938 S 08°41' E 126°37' 
26" m Lautém Com (Com Beach Resort; Com wharf; Pousada de Com) 2-15 S 08?21' E 127°03' 
OTe Lautém Raga caves & surrounds 400—553 S 08°26' E 126°59' 
28" Lautém Tutuala (Pousada de Tutuala) 373 S 08?24' E 127?15' 
29" Lautém Malahara (Mainina sinkhole; Lake Ira Lalaro) 336-424 S 08?29' E 127?11' 
30" Lautém Jaco Island 10-40 S 08?25' E 127?19' 

'GPS coordinates are approximate to define the area in which the survey work was carried out. Exact localities are not provided to protect 

some of the unique and fragile habitats in Timor-Leste. 

?The confluence of the Comoro and Bemos Rivers marks the border between Dili, Liquica and Aileu Districts, with specimens collected on 

both banks in Liquica and Aileu Districts. 

76 Asian Herpetological Research Vol. 6 

Museum of Natural History, Smithsonian Institution, of the USNM and are listed here with herpetological 
Washington D.C., USA (USNM). Those specimens not image numbers (accessioned as USNM-HI). Museum 
yet accessioned have field tags of the USNM (USNM- abbreviations are taken from Sabaj Pérez (2014). 

FS). Photographs of road-killed specimens, CITES- In the species accounts, we provide information to 
protected species, and other unvouchered specimens have aid in field identification of amphibians and reptiles, 
been deposited in the herpetological image collection particularly for taxa not included in one of our earlier 

Figure 2 Sample habitat types surveyed in Timor-Leste during 2011—13. Localities are listed numerically (see Table 1). (A) Rocky shore at Cristo 
Rei Protected Area on the outskirts of Dili (Locality 2). The part of this habitat along the tidal and splash zones is a habitat of Cryptoblepharus cf. 
schlegelianus and Laticauda colubrina, whereas in the wooded area at higher elevation, sun skinks (Eutropis cf. multifasciata) and Timor monitors 
(Varanus timorensis) have been observed. (B) Montane habitat at Maganuto (Locality 11). This area has stands of tall bamboo in boulder-strewn 
areas, intermixed with a grassy meadow and a montane forest on the upper slopes. Whereas we found the forest to be unproductive in our search, the 
bamboo yielded Hemidactylus cf. garnotii, and in the grassy meadow we found a Polypedates cf. leucomystax under a flat rock some distance away 
from any vegetation. (C) The banana plantation at the confluence of the Bemos and Comoro Rivers (Locality 6; photo taken towards Aileu District) 
turned out to be an unexpectedly important site at which one of only two recent specimens of Cylindrophis cf. boulengeri was found. Other species 
recorded in this area include Duttaphrynus melanostictus, Fejervarya sp., Polypedates cf. leucomystax, sun skinks (Eutropis cf. multifasciata), and 
house geckos (e.g., Hemidactylus frenatus). (D) Disturbed forest at Fiuren (Locality 9). Overtly a nice patch of forest with an expansive growth of 
large trees, this area is beset by domestic pigs that scour the leaf litter and the root matter for food. We located P. cf. leucomystax and several gecko 
taxa (Cyrtodactylus, Gekko, Gehyra, Hemidactylus) in this area. (E) View of the mountains above our survey area near Baguia (Locality 25). A 
promising habitat with extreme topography, this is the only area in Timor-Leste where we have found individuals of Hemiphyllodactylus cf. typus. 
(F) The Mainina sinkhole (Locality 29) in Nino Konis Santana National Park. This locality is the only outflow of Lake Ira Lalaro, the largest lake 
in Timor-Leste. The area is seasonally inaccessible due to variations in the lake's water level, and it lies right at the foot of the steep-sided Paitxau 
Mountains karst formation. (G) The road passing through tropical wet forest in the Nancuro Protected Area (Locality 20). On either side of this road 
is dense, mixed coastal forest that includes some large trees. The ground is partially inundated after rains. This has been a very productive collection 
locality with a high diversity of herpetofauna, including Kaloula, Cyrtodactylus, Sphenomorphus, Dendrelaphis, Stegonotus, and Trimeresurus. (H) 
Dry coastal forest on Jaco Island (Locality 30). Even though this corallogenic island appears to be very dry, we have found species that we have more 
commonly encountered in moist habitats elsewhere in Timor-Leste, including Cyrtodactylus, Eremiascincus, and Sphenomorphus. Photos (A), (C), 
and (E)-(H) by Hinrich Kaiser, (B) and (D) by Mark O'Shea. (Continued on facing page). 

No.2 Mark O'SHEA et al. 
reports (Kaiser et al., 2011, 2013b; O’Shea et al., 2012; 
Sanchez et al., 2012), comment on new locality records 
for taxa previously recorded during Phases I-II (2009— 
2010), provide full accounts for taxa not previously 
recorded during our surveys, and discuss the natural 
history of the species and the manner in which they were 
encountered. The recording or collection of taxa during 
particular phases is indicated by the phase designation 
in bracketed superscripted Roman numerals, following 
taxon names. Thus, a species encountered in Phases IV 
and VII would carry the superscript "YY". 

Common names are provided in English (E), German 
(G) and the country's lingua franca, Tetun (T). We made 
a number of decisions with regards to the use or coining 
of common names in Tetun and the interested reader 1s 
referred to O’Shea et al. (2012) for a discussion of our 

Timor-Leste Herpetofauna Updates 77 

arguments. Proposed common names for house geckos 
incorporate the commonly used Tetun name for small 
geckos and the scientific name or a descriptive term. 

3. Results 

Family Bufonidae — True Toads 

Duttaphrynus melanostictus (Schneider, 1799 

{IV-V, VII- 

Common names. (E) Black-spined Toad, Common Asian 
Toad. (G) Schwarznarbenkróte. (T) Manduku Interfet 
(manduku = frog, INTERFET = International Force for 
East Timor). 

Known distribution. Duttaphrynus melanostictus (Figure 
3) has heretofore been reported from nine of Timor- 
Leste's 13 districts (Table 2): Aileu, Bobonaro, Covalima, 

Figure 2 Continued. 

78 Asian Herpetological Research 

Vol. 6 

Table 2 Amphibian records for the districts of Timor-Leste. Black dots indicate previously known records, red dots denote new records. 

e € : 
Taxon P g z £ 3 . r " g E " 3 References 
4i å à & BÓ à E à 4 3 3 3 6 p 

Duttaphrynus melanostictus @ e. e. e. e. e. e. e. e. e. e. 1-5 
Fejervarya spp. e. e. e. e. e. e. e. (J e. e. e. e. 2-4 
Limnonectes timorensis e e 2,5 
Litoria everetti e 2,6 
Kaloula sp. e. e. 5 
Polypedates cf. leucomystax e. e. e. e. e. e. e. e. e e. e. e. 2-5 

"References are identified numerically as follows: 1 = Trainor, 2009; 2 = Kaiser et al., 2011; 3 = O’Shea et al., 2012; 4 = Sanchez et al., 2012; 5 = this 

paper; 6 = Menzies, 2006. 

Dili, Ermera, Liquiga, Manufahi, Oecusse, and Viqueque 
(Kaiser et al., 2011; O’Shea et al., 2012; Sanchez et al., 
2012; Trainor, 2009). 

New localities. We collected additional specimens from 
the Comoro River valley (Localities 5 and 6; Table 1), 
which included a series of tadpoles from the confluence 
of the Comoro and Bemos Rivers, which occurs at the 
boundaries of Aileu, Dili, and Liquiga Districts. Tadpoles 
were captured in riverine kolks, where back eddies create 
a respite from rushing water, on the Aileu side (Locality 
6). An adult was captured at Beduku Aldeia (Dili District; 
Locality 5). We vouchered single specimens from the Soto 
River (Bobonaro District; Locality 10) and the Franciscan 
Convent of St. Antony d’Lisboa (Manufahi District; 
Locality 19), and took voucher photographs for four other 
localities where we recorded this species: Sta. Bakhita 
Mission (Eraulo, Ermera District; Locality 8); Nancuro 
coastal forest (Natarbora, Manatuto District; Locality 20), 
Ossu (Baucau District; USNM-HI 2823), and Liamida 
(Viqueque District; Locality 21). The Manatuto and 
Baucau records constitute new district records and bring 
to a total of 11 (Table 2) the number of mainland districts 
that have been colonized by D. melanostictus since its 
arrival less than a decade ago. Based on our observations, 
the species has so far (mid-2013) not expanded into 
Lautém District, the country’s easternmost and the site of 
Nino Konis Santana National Park, and it has not yet been 

documented from Ainaro District. 

Natural history. This is an introduced species that is 
believed to have arrived in Timor-Leste with INTERFET 
peacekeeping troops. The first reports appear to have 
come from Oecusse District in 1999, a date that 
coincides with the arrival of South Korean INTERFET 
peacekeepers. From there the toad appears to have 
gradually spread eastwards, arriving in Dili District in 
2007 (Trainor, 2009). We recorded it further southeast at 
Same (Manufahi District) in 2009 (Kaiser et al., 2011), 
concurring with Trainor (2009), who also recorded it in 
the area during the same year, and on the south coast at 
Uma Boot (Viqueque District) in 2010 (O'Shea et al., 

Sanchez et al. (2012) reported this species from the 
Oecusse exclave. Our surveys so far have not revealed 
the presence of D. melanostictus or any other amphibian 
species on Ataüro Island (Kaiser et al., 2013b). During 
2011 and 2012 we were able to report a much wider range 
for the black-spined toad, across the contiguous districts 
of mainland Timor-Leste, from Bobonaro (Locality 10), 
in the extreme west near the border with West Timor, to 
Ossu Subdistrict of Viqueque District (Locality 21) in 
the east. We have now recorded D. melanostictus from 
nine of the 12 contiguous districts, plus Oecusse, from 
sea level to elevations of 930 m (Liamida, Viqueque 
District; Locality 21) and 1225 m (Sta. Bakhita Mission, 

No.2 Mark O'SHEA et al. 

Ermera District; Locality 8), in habitats ranging from 
anthropogenic (roadways, convent grounds) to coastal 
forests, rocky river beds, and upland boulder-strewn 
grasslands. Based on our observations this introduced 
toad species favors anthropogenically-modified habitats, 
where it can be found in great numbers; it appears to be 
absent in pristine habitats. In drainage ditches and rice 
paddies, D. melanostictus is frequently found in sympatry 
with frogs of the genus Fejervarya. 

Our vouchers include adult toads and a series of 
tadpoles (USNM 581259—63) collected from muddy 
rivulets and pools alongside the Comoro riverbed. 
Duttaphrynus melanostictus was also found to be very 
common in the grounds of the Franciscan Convent 
of St. Antony d'Lisboa, Fatucahi (Manufahi District; 
Locality 19) but we vouchered only a single specimen 
(USNM 565895) that had predated and begun to pass a 
blindsnake (/ndotyphlops braminus; O’Shea et al., 2013). 
Another specimen was found sitting atop the 2.0 m stone 
convent wall, demonstrating the climbing ability of these 
terrestrial bufonids. 

Although we initially did not collect voucher 
specimens of this non-Timorese amphibian, in our efforts 
to monitor its effects on native taxa, we collected 87 
specimens in several districts in 2013 to be able to carry 
out a gut content analysis to study the diet of this exotic 
(Dóring et al., in prep.). Our most recent observations 
continue to confirm the absence from Timor-Leste of the 
much larger and elsewhere harmful cane toad (Rhinella 
marina), with which D. melanostictus has been confused 
by Timorese and expatriates alike. 

Family Dicroglossidae — Fork-tongued Frogs 

Genus Fejervarya ' \""! 

Common names. (E) Rice Paddy Frogs. (G) Reisfrósche. 
(T) Manduku natar (manduku = frog, natar = rice paddy). 
Known distribution. Frogs of the genus Fejervarya 
(Figure 4) have been reported from seven of Timor- 
Leste’s 13 districts (Table 2): Baucau, Dili, Ermera, 
Lautém, Manufahi, Oecusse, and Viqueque (Kaiser et al., 
2011; O’Shea et al., 2012; Sanchez et al., 2012). 

New localities. For 2011, we report additional voucher 
specimens from the localities at the confluence of 
the Comoro and Bemos Rivers (see D. melanostictus 
account), from the Aileu bank (Locality 6). We also 
added vouchers from a roadside marsh at the junction of 
the Com-Bauro road with the North Coast Road (Com, 
Lautém District; Locality 26), and from the southern 
shore of Lake Ira Lalaro (Malahara village, Lautém 
District; Locality 28). We also provide the first records 
of Fejervarya spp. from southern Timor-Leste, namely 

Timor-Leste Herpetofauna Updates 79 

for Manatuto District, from coastal forest (Locality 20); 
for Manufahi District from the grounds of the Franciscan 
Convent of St. Antony d'Lisboa and the southern shore 
of Lake Lenas (both near Fatucahi; Locality 19); and 
for Covalima District from the grounds of the Castelo 
Fronteira Guest House (Suai town; Locality 13) and the 
extensive rice-paddies at Kasabauk (Locality 14). The 
Aileu, Manatuto, and Covalima specimens represent new 
district records (Table 2). ZAS also provided our first 
records for Bobonaro District with vouchers from the 
Soto River and Ramaskora (Locality 10), and a single 
voucher from the Galosapulu swamp (Locality 12). 

In 2012 we obtained additional vouchers from west 

of Dili town (Timor Lodge Hotel grounds, Dili District; 
Locality 1) and the Meleotegi River (Ermera District; 
Locality 8), and made collections in two new localities: 
Lake Be Matin (Aileu District; Locality 7), and the 
Afaloicai and Ossohuna rice paddies (Baucau District; 
Locality 25). Fejervarya spp. have now been reported 
from 11 of the 12 contiguous mainland districts in 
addition to Oecusse (Sanchez et al., 2012), but they have 
not been recorded from Ainaro District; based on the 
limited environments suitable for Fejervarya, we do not 
anticipate their presence on Atauro Island (Kaiser et al., 
2011; 2013b). 
Natural history. Recorded widely on all previous 
phases, our additional collection confirms that rice- 
paddy frogs occupy a much broader variety of habitats 
than their common name indicates. Along the mostly dry 
Comoro riverbed (Locality 6), an adult (USNM 579397) 
was found under a rock right at the edge of the narrow 
flow, whereas a tadpole (USNM 581584) was collected 
from a nearby shallow pool shared with tadpoles of 
Duttaphrynus melanostictus. Near this locality, we 
observed a wolf spider (family Lycosidae) that appeared 
to follow the movements of Fejervarya tadpoles grazing 
near the surface (Figure 5), and we consider it likely that 
these spiders take tadpoles as prey. Lycosid spiders have 
already been documented as hunting in this way (Jara and 
Perotti, 2004). 

Specimens were also vouchered from the Soto River 
(Locality 10; USNM 579287—92) and Meleotegi River 
(Locality 8; USNM 579710, 580466) during both the 
wet and the dry seasons. A population of rice paddy frogs 
from far-eastern Timor-Leste (Locality 26) was initially 
located based on their vocalizations along the edge of the 
road, where a leaking water pipe had created puddles. 
This population (USNM 579398—401) extended into 
a marshy area next to the road. In Bobonaro District, 
a single specimen was collected in a swamp at 712 m 

80 Asian Herpetological Research 

Vol. 6 

Figure 3 Duttaphrynus melanostictus found in a streamside 
refugium along the Comoro River (Locality 1). This specimen was 
not vouchered. Photo by Mark O'Shea. 

elevation (Locality 12; USNM 279297). Near Malahara 
village (Lautém District; Locality 28) several individuals 
were seen in the marshy area along the edge of Lake Ira 
Lalaro, and a single specimen was vouchered (USNM 

Our south coast records for a Fejervarya species come 
from pristine wet coastal forest (Locality 20; USNM 
579279); grassy areas of a residential compound (Locality 
19; USNM 579276-—77); the grassy edge of a small lake 
(Locality 19; USNM 579278); and an ornamental fountain 
in a residential compound (Locality 13; USNM 579280- 
81), where they occurred in the company of a large 
number of tadpoles (USNM 581264—77). As expected, 
Fejervarya were found to be especially common in rice 
paddy habitats, from near sea level at Kasabauk (9 m, 
Locality 14; USNM 759284—86) and medium elevations 
(e.g., 229 m at Ramaskora; Locality 10; USNM 279293- 
96) to higher elevations (e.g., 775 m at Afaloicai and 
Ossohuna (Locality 25; USNM 580468—72, 581287-93), 

Figure 4 A female rice-paddy frog (genus Fejervarya) from active, 
inundated rice paddies near Baguia (Locality 25; USNM 580467). 
Photo by Mark O'Shea. 

and over 1105 m at Lake Be Matin (Locality 7; USNM 
57797706—09). Individuals were also encountered crossing 
or occupying rain puddles on the road (e.g., at Baucau 
District, between Localities 24 and 25; USNM 580467). 

As during previous phases we found rice-paddy frogs 
to be extremely abundant where they occurred, and 
although numerous specimens were initially collected, 
only a few were selected as vouchers. The physiological 
plasticity of these species and their adaptability to 
anthropogenic habitats is discussed elsewhere (Kaiser et 
al., 2011; O'Shea et al., 2012) and will not be expanded 
upon further here. 

Limnonectes timorensis (Smith, 1927) ^ Y™ 

Common names. (E) Timor River Frog. (G) Timorfrosch 
(T) Manduku mota (manduku = frog, mota = river). 
Known distribution. Limnonectes timorensis (Figure 6A) 

has heretofore been reported from only a single locality in 
Ermera District (Table 2; see Kaiser et al., 2011) 

Figure 5 Tadpole of Fejervarya sp. (arrow) with its potential predator, a wolf spider. The spider was observed in close proximity to tadpoles 
along a slow-flowing side arm of the Comoro River (Locality 1). Photo by Hinrich Kaiser. 

No.2 Mark O'SHEA et al. 

New localities. Additional specimens were collected 
at the Meleotegi River location (Locality 8) during the 
dry seasons of 2011 and 2012, and a single voucher was 
obtained from the Afaloicai rice paddies near Baguia 
(Baucau District; Locality 24). This single voucher is a 
juvenile and its location at Afaloicai is over 130 km E of 
the Meleotegi River. 

Natural history. Previously we had encountered this frog 
only along the Meleotegi River (near the Sta. Bakhita 
Mission (Eraulo, Ermera District; Locality 8), with only 
two vouchers collected during the dry season of 2009. 
During the dry season of 2011 we again encountered L. 
timorensis along the Meleotegi River, where several males 
(e.g., USNM 579403-07) were discovered sheltering 
under rocks at the water's edge. Two of these specimens 
(USNM 579404, 579407) contained parasitic cestodes 
(Platyhelminthes: Cestoda) in their leg muscles (Figure 
6C, D), which likely constitute another host record for 
spargana (Goldberg et al., 2010). A return to the same 
location, at the same time of year, in 2012 produced a 
series of nine tadpoles (Figure 6B) collected from a rock 
pool on a large mid-stream rock (USNM 581278-86). The 
tadpoles were euthanized and photographed at sequential 
stages of development over the following ten days. 

All specimens of L. timorensis collected at the 

Meleotegi River (elev. 1175-1185 m) were found in close 
association with the river, albeit in the dry season, on 
rocks along its periphery. The single juvenile collected 
at Afaloicai, near the Baucau-Viqueque boundary at an 
elevation of 775 m (USNM 580371) was taken at night in 
the grass surrounding a rice-paddy. 
Taxonomic comment. The generic status of some of the 
frogs in the genus Limnonectes 1s being re-evaluated, 
and it appears that both molecular evidence and some 
morphological characteristics align the Timor population 
with ranids in the genus Hylarana (Che et al., 2007; 
Kaiser et al., 2014). If this generic concept is confirmed, 
this species should be assigned to the genus Hylarana and 
transferred to the Ranidae. 

Family Hylidae — Treefrogs 

Litoria everetti (Boulenger, 1897) '"! 

Common names. (E) Everett’s Timor Treefrog. (G) 
Everett-Laubfrosch. (T) Manduku ai Timor (manduku = 
frog, ai = tree). 

Known distribution. Litoria everetti (Figure 7) is 
reported from a single locality in Ermera District (Table 
2; see Kaiser et al., 2011). 

New localities. None, but an additional specimen was 
collected at the known locality. 

Natural history. As with Limnonectes timorensis (see 

Timor-Leste Herpetofauna Updates 81 

above), our previous encounters with this frog were in 
2009, only on the Meleotegi River (Ermera District; 

Figure 6 (A) Juvenile Limnonectes timorensis from a grassy 
patch at Afaloicai (USNM 580473, Locality 25). (B) Tadpole of L. 
timorensis from the Meleotegi River (USNM 581286; Locality 8). 
(C) Upper leg of an adult L. timorensis from the Meleotegi River 
(USNM 579404, Locality 8), showing an embedded parasite (box). 
(D) Tapeworm extracted from the animal in (C), presumably a 
sparganum that is part of the host-parasite interaction described by 
Goldberg et al. (2010). Photos (A) and (B) by Mark O'Shea, (C) 
and (D) by Hinrich Kaiser. 

82 Asian Herpetological Research 

Vol. 6 

Locality 8), when we collected two specimens. During 
2011 we collected a third specimen (USNM 579408) 
at the same location. This specimen was discovered 
underneath a rock on a small rocky island in midstream, 
and it attempted to escape by jumping into the flowing 
water. After this initial escape attempt, it remained 
motionless on the bottom of a slow-flowing portion of the 
river, where it was easily captured. 
Family Microhylidae — Narrow-mouthed Toads 
Kaloula sp. "* ""! 
Common names. (E) Timor Pumpkin Bullfrog. (G) 
Timor-Ochsenfrosch. (T) Manduku lakeru (manduku = 
frog, lakeru = pumpkin). 
Identification. Kaloula sp. is a small rotund frog with a 
blunt head and highly tuberculate dorsum (Figure 8A). 
The limbs are short, the toes unwebbed. Coloration 
consists of a mixture of olive green and light brown 
blotches. The only Timorese frogs with which this 
species could be confused are Asian black-spined toads 
(Duttaphrynus melanostictus), from which it can be 
separated by its smaller size, longer fingers and toes, 
discrete tympanum, the lack of cranial crests, parotoid 
glands, and black tipped tubercles. 
Known distribution. There are no previous reports of 
this species from Timor Island or Timor-Leste. 
New localities. Specimens collected in 2011, in southern 
Manufahi District (within the grounds of the Franciscan 
Convent of St. Antony d'Lisboa, Fatucahi; Locality 
19) constitute the first records of this species, genus, 
and family for Timor Island. Two relatively juvenile 
specimens collected in 2012, in the wet coastal forest at 
Nancuro (Locality 20) represent the first records of the 
taxon from Manatuto District. These two localities are 
only 10 km apart. 
Natural history. Within the grounds of the Franciscan 
Convent of St. Antony d'Lisboa we encountered four 
species of anurans, three of which (Duttaphrynus 
melanostictus, Fejervarya sp., Polypedates cf. 
leucomystax) are widespread in Timor-Leste. However, 
we also collected numerous specimens of Kaloula sp. 
at night in the vegetable gardens, on the rubbish dump, 
and around the convent wall. One specimen was found 
on a low tree axil approximately 45 cm from the ground, 
whereas all others were encountered at ground level, 
including under rocks together with D. melanostictus. A 
series of ten specimens was vouchered (USNM 579246- 

The juvenile specimens collected at Nancuro (USNM 
580464—65) were found on the forest floor in deep leaf 
litter. They demonstrated much more vivid markings than 

the adults from Fatucahi, in the form of a series of black- 
edged, bright orange flashes across the flanks anterior to 
the hind limbs, on the inner surfaces of the thighs, and 

on the proximal portions of the hind limbs (Figure 8B). 

Figure 7 Female Litoria everetti found underneath a flat rock 
alongside the Meleotegi River, Ermera District (Locality 8; USNM 
5779394). Photo by Mark O'Shea. 

Figure 8 (A) Adult female Kaloula sp. (USNM 579254) from 
the grounds of the Convent of St. Antony d'Lisboa near Fatucahi, 
Manufahi District (Locality 19). (B) Juvenile Kaloula sp. from wet 
forest in the Nancuro Protected Area (Locality 20; USNM 580464) 
showing the characteristic flash colors on the posterior part of the 
body in juveniles of this form. Photos by Mark O'Shea. 

No.2 Mark O'SHEA et al. 

These markings were exposed when the frogs made short 
hops and presumably constitute aposematic eyespots to 
deter potential predators, as has been well-documented 
for frogs of the genus Physalaemus (Wells, 2010). 
Taxonomic comments. There are no previous records 
of Kaloula, or any microhylid frog, from the island of 
Timor, but three species of Kaloula are reported to occur 
on other islands in Indonesia. Kaloula pulchra Gray, 
1831 has been reported from Sumatra, Borneo, Sulawesi, 
and Flores (Barbour, 1912; Dunn, 1928; Mertens, 1930), 
and it 1s unclear whether these populations are native 
or introduced. The same problem exists for K. baleata 
(Müller, 1833), which occurs on Bali, Borneo, Java, 
Komodo, Sulawesi, Sumatra, and Sumba (Dunn, 1928; 
Iskandar, 1998; McKay, 2006); the Western Australian 
Museum holds specimens from Flores and Lembata 
(Paul Doughty, pers. comm.). A population listed as “K. 
albotuberculata" by Inger and Voris (2001) may represent 
an endemic taxon found exclusively in central Sulawesi; 
the listing of this name, based on a manuscript by Djoko 
Iskandar, has created a nomen nudum, which was referred 
to as “Kaloula sp. n.” by Koch (2012). Kaloula baleata, 
as currently defined, is certainly a polymorphic taxon that 
contains at least three species in addition to the recently 
described K. indochinensis Chan et al., 2013 and K. 
latidisca Chan et al., 2014 (Rafe Brown, pers. comm.), 
and it seems likely that K. pulchra is a species complex as 
well. Specimens from the Lesser Sundas may well have 
been allocated to these two species without comparison 
to specimens from the type localities (Java for K. baleata 
and China for K. pulchra) and may therefore constitute 
undescribed species. Detailed examinations by one of us 
(HK) of the Timor specimens now housed in the USNM 
leads us to believe that they represent an undescribed 
species of Kaloula. True K. baleata and K. pulchra may 
be separated by the degree of toe webbing (webbing 
reaching the middle subarticular tubercle on the inner 
edge of the fourth toe in K. baleata, but not extending 
beyond the basal subarticular tubercle in K. pulchra 
(Inger, 1966), but the Timor material does not comply 
with either arrangement. The detailed morphological and 
morphometric information provided by Chan et al. (2013, 
2014) to assist with delineating species boundaries in the 
K. baleata complex provides an excellent opportunity for 
determining the taxonomic status of the Timor population. 
On the basis of our discovery, and pending comparative 
examination of the Timor specimens with other Lesser 
Sunda and extralimital material of K. baleata sensu lato, 
we assign the specimens from Fatucahi and Nancuro to 
Kaloula sp. 

Timor-Leste Herpetofauna Updates 83 

Family Rhacophoridae — Afro-Asian Foam-nest 

Polypedates cf. leucomystax ! 
Common names. (E) Striped Treefrog, Four-lined 
Treefrog. (G) Weifbart-Ruderfrosch. (T) Manduku ai- 
riskadu (manduku = frog, ai = tree, riskadu = striped) or 


manduku loron (manduku = frog, loron = sunlight). 
Known distribution. Polypedates cf. leucomystax 
(Figure 9) is so far known from eight of Timor-Leste’s 13 
districts (Table 2): Ainaro, Baucau, Dili, Ermera, Lautém, 
Manufahi, Oecusse, and Viqueque (Kaiser et al., 2011; 
O'Shea et al., 2012; Sanchez et al., 2012). 

New localities. In 2011 we collected specimens at the 
confluence of the Comoro and Bemos Rivers, along the 
Liquiça bank (see Duttaphrynus melanostictus account; 
Locality 6), on the south coast (grounds of the Franciscan 
Convent of St. Antony d'Lisboa, Fatucahi, Manufahi 
District; Locality 19), and in the Mt. Debululik area 
(north of Tilomar, Covalima District; Locality 15). The 
Liquiga and Covalima specimens represent new district 
records (Table 2). 

Bobonaro specimens were collected by ZAS from all 
three of his study sites: the Soto River and Ramaskora 
rice paddies (Locality 10) and the Galosapulu swamp 
(Locality 12). These are our first records of Polypedates 
from Bobonaro District. 

In 2012 we collected two specimens in the grounds of 
the Timor Lodge Hotel (Comoro, Dili District; Locality 
1) and vouchered a series of specimens from near Baguia 
(Baucau District; Localities 24 and 25), near Balibo 
(Bobonaro District; Locality 9), and from the south coast 
in the wet coastal forest at Nancuro (Locality 20), this last 
record constituting a first record for Manatuto District. 
The only district without records of Polypedates cf. 
leucomystax is Aileu, and despite of our efforts on three 
collecting trips to Ataüro Island no frog vouchers were 
collected there (Kaiser et al., 2013b). 

Natural history. These treefrogs were found exclusively 
in microhabitats with some form of water storage 
capacity, sometimes in unexpected circumstances. For 
example, specimens found in village gardens along the 
Comoro River (Dili District; Locality 6; USNM 579395— 
96) were discovered by peeling back the stem leaves at 
the bases of banana plants (Musa sp.), where runoff water 
collects. These banana plots had recently undergone an 
agricultural slash-and-burn, yet several of the banana 
plant stems were found to harbor treefrogs. Striped 
treefrogs were also seen in the freshly mown grassy 
vegetation of the grounds of the Franciscan Convent 
(USNM 579256—62), where moisture is retained well and 

84 Asian Herpetological Research 

Vol. 6 

where disturbed insects provide abundant food. Treefrogs 
were also fairly abundant near upland rice paddies at 
Ossohuna and Afaloicai (Baucau District; Locality 25; 
USNM 580453) at an elevation of 712 m. Another of 
our highest-elevation specimens (elev. 706 m; USNM 
579263) came from a rock pile in the Mt. Debululik area 
(Covalima District; Locality 15), while specimen found 
on the ground in bamboo litter at Ossohuna (Baucau 
District; Locality 25; USNM 580452) occurred at 938 m. 
The highest elevation recorded for P. cf. leucomystax was 
1074 m for a specimen curiously found under a rock on an 
exposed step-cultivated grassy hillside above Maganuto 
village, almost midway between, but still a considerable 
distance from, a large stand of bamboo and a stunted elfin 
forest growing in the shadow of Mt. Leolaco, Bobonaro 
District (Locality 11; USNM 580457). 

Striped treefrogs found in anthropogenic habitats 

included a small series (USNM 580458—63) collected in a 
significantly degraded forest at Fiuren village, Bobonaro 
District. Curiously, these frogs were found in leaf litter 
or buttress roots close to the forest floor, despite the 
entire area being drastically altered by a large population 
of domestic pigs. The presence of the pigs resulted in a 
low number of terrestrial reptiles encountered, yet the 
treefrogs endured. Another treefrog was found at night, 
perched on the branch of a tree growing within the ruins 
of an old school (Escola do Reino de Haudere) near 
Baguia (Baucau District; Locality 24; USNM 580451). 
In a more natural environment, our single Manatuto 
specimen (Locality 20; USNM 580456) was found inside 
a hollow log. 
Taxonomic comments. The taxonomic status of P. cf. 
leucomystax is discussed in previous reports (Kaiser et 
al., 2011; O’Shea et al., 2012) and will not be elaborated 
upon here. The taxonomy of the P. leucomystax species 
complex is currently under investigation (Rafe Brown, 
pers. comm.; Hidetoshi Ota, pers. comm.). Polypedates 
leucomystax has generally been considered a widespread 
Asian species that also occurs on many islands across the 
Sunda Shelf. However, P. leucomystax sensu stricto may 
not extend further east than Bali, into the Lesser Sunda 
Archipelago, although molecular data for the Lesser 
Sundas is still lacking (Brown et al., 2010; Kuraishi et 
al., 2013; Kuraishi et al., 2011). Specimens from Nusa 
Tenggara Province, including those from the island of 
Timor, could represent introduced populations originating 
in the Greater Sunda area, or they could be a regionally 
endemic, hitherto unrecognized Lesser Sunda species. 

Family Agamidae—Agamas and Dragon Lizards 

Draco timoriensis Kuhl, 1820 "YY" V! 

Common names. (E) Timor Flying Dragon, Timor 
Flying Lizard. (G) Timor-Flugdrache. (T) Teki liras (teki 
= gecko, liras = winged). In direct translation, the Tetun 
name more accurately describes the gekkonid genus 
Ptychozoon, which is not found east of Wallace’s Line. 
We believe that the common name of D. timoriensis is 
not an indication that local residents are unable to tell a 
gecko from an agamid lizard. Instead, it may reflect the 
assumption that lizards of comparable size are likely 
geckos, an error culturally perpetuated by the lack of 
opportunities to catch more than a fleeting glance at an 
individual because of the Draco lifestyle. 

Known distribution. Draco timoriensis (Figure 10) 
is currently reported from five of Timor-Leste’s 13 
districts (Table 3): Baucau, Lautém, Liquiga, Oecusse, 
and Viqueque (Kaiser et al., 2011; O’Shea et al., 2012; 
Sanchez et al., 2012). There are no records of any Draco 
species from Ataüro Island (Kaiser et al., 2013b), a 
location surrounded by islands where Draco have been 
recorded, but islanders are adamant that they do not occur 
there (O'Shea and Kaiser, 2013). 

New localities. During 2011 and 2012 we obtained 
additional specimens from Lautém District, from Com 
(Locality 26) and Raga (Locality 27). New district records 
were established for Manufahi District through the 
collection of specimens in the Betano area on the south 
coast (Locality 18), and from several localities around 
Same (Locality 17): in the mountains, in the grounds of 
the Ailelehun Guest House, and at Ladiki village, 5 km 
NE of Same. Southern coastal records came from the 
Nancuro coastal forest (Manatuto District; Locality 20), 
the grounds of Castelo Fronteira Guest House in Suai 
(Covalima District; Locality 13), and two sites outside 
of Tilomar (Covalima District; Locality 15), namely 
the Tilomar Forest Reserve and just N of Maubesi. On 
the north side of Timor we obtained a single specimen 
from the Meleotegi River (Ermera District; Locality 8), 
as well as a specimen from Dare, in the hills above Dili 
(Dili District; locality 4). The vouchers from Manufahi, 
Manatuto, Covalima, Ermera, and Dili are first district 
records and this doubles the number of districts from 
which Draco timoriensis has been recorded to ten (Table 

Natural history. Although a relatively small and slender 
species, Draco timoriensis is a fairly conspicuous lizard. 
It is usually seen running up the trunks of coconut 
palms or smooth-barked eucalypts and 1f pursued will 
easily leap and glide gracefully to another tree. It clearly 
exhibits a wide distribution, both on the southern and 

Mark O'SHEA et al. 

Figure 9 Adult Polypedates cf. leucomystax from a creek-side 
tree near Ossohuna (USNM 580454, Locality 25). Photo by Mark 

northern coasts, including the Oecusse exclave (Sanchez 
et al., 2012), and it is relatively common at elevations 
only marginally above sea level (e.g., at 3 m in the 
Nancuro wet coastal forest, Manatuto District; Locality 
20; USNM 579298; at 3 m on a tree opposite the wharf 
at Com, Lautém District; Locality 26; USNM 579491). 
We also obtained specimens at significantly higher 
elevations, on the upland limestone plateaus and central 
mountains of Timor-Leste (e.g., at 412 m elevation on 
trees around Raga village, Lautém District; Locality 27; 
USNM 579310—12, 579490; on forest trees at 442 m 
elevation near Tilomar, Covalima District; Locality 15; 
USNM 579302—04; and at 600 m elevation on a large 
tree, opposite the war memorial at Dare, Dili District; 
Locality 14; USNM 579711). The Meleotegi River 
specimen (Ermera District; Locality 8; USNM 579492) 
was collected at 1177 m, and constitutes the highest 
elevation record for D. timoriensis we have observed on 
Timor Island. 

Family Gekkonidae — True Geckos 

Cyrtodactylus spp. "Y ‘™" 

Common names. (E) Bent-toed Geckos, Bow-fingered 
Geckos. (G) Bogenfinger-Geckos. (T) Teki ain-fuan kleuk 
(Teki = small gecko, kluek = bent, ain-fuan = toe). 
Known distribution. During our initial survey in 2009 
we collected two geckos of the genus Cyrtodactylus in 
Timor-Leste (see Taxonomic comment below). This 
population, currently referred to as Cyrtodactylus sp. 
‘Trilolo River,’ was collected 4 km north of Same 
(Manufahi District; Locality 17; Kaiser et al., 2011). In 
2010 we collected a single specimen of Cyrtodactylus sp. 
*Manucoco' on the northwestern slopes of Mt. Manucoco 
on Ataüro Island (Kaiser et al., 2013b), and a series of ten 
vouchers of a third population, Cyrtodactylus sp. ‘Abanat 

Timor-Leste Herpetofauna Updates 85 

Figure 10 Adult female Draco timoriensis from a tree at Dare 
(USNM 579711, Locality 4). Photo by Mark O'Shea. 

River,’ in the Oecusse exclave (Sanchez et al., 2012). 
Populations of Cyrtodactylus are therefore known from 
two mainland districts (Manufahi, Oecusse) and from 
Ataüro Island (Dili District) so far (Table 3). 

New localities. During 2011 we discovered further 
populations of Cyrtodactylus (Figure 11) over a wide 
area of Timor-Leste. Specimens collected at sea level on 
Ataüro Island (USNM 579712-25) are being treated as 
Cyrtodactylus sp. *Ataüro coast’ (Kaiser et al., 2013b). 
Additionally, we located two more populations in Lautém 
District, along the north coast at Com (Figure 11B; 
Locality 26; USNM 579411—-23) and on the adjacent 
limestone plateau, at Raga (Figure 11A; Locality 27; 
USNM 579313, 579408-09), and near the Mainina 
sinkhole (Locality 29; USNM 579410, 579424) (see 
Taxonomic comment below). 

During 2012 four further populations of Cyrtodactylus 
were discovered and sampled: from a network of man- 
made tunnels at Venilale, Baucau District (Figure 11C; 
Locality 22; USNM 580474—84); in the coastal forest 
at Nancuro, Manatuto District (Figure 11D; Locality 
20; USNM 580485-86); near Maganuto village, in the 
mountains surrounding Maliana, Bobonaro District 
(Figure 11E; Locality 11; USNM 580457), and in Fiuren 
village, near Balibo, Bobonaro District (Figure 11F; 
Locality 9; USNM 580488). At an altitude of 1036 m, the 
Maganuto locality is the highest record for these geckos 
in Timor-Leste. 

In the summer of 2013, we discovered yet another 
population of bent-toed geckos in the vicinity of 
Com (Locality 26; USNM 581153—54), one clearly 
distinct from the small-bodied form we found in 2011. 
Cyrtodactylus spp. have now been recorded from six 
mainland districts and Ataüro Island (Table 3). 

Natural history. The only general habitat requirement 

86 Asian Herpetological Research Vol. 6 

Table 3 Records of lizard species for the districts of Timor-Leste. Black circles indicate previously known records, red circles denote new 
records. The black open circle refers to a literature record only. Records listed in grey denote literature records from West Timor, with open 
circles representing known museum specimens. 


Taxon o E S E mi © S g S $ E g - S References" 

2 5 § 3 'c S 85 8 $ s 5 43 8 |E 

2 B8 8 8 6 zBuilb 8 e 3 8 $3 P E 

«x 4 à m o ACASA A A Ss z 6 P JE 
Draco timoriensis e e e e (3 e e e e e 1-4 
Cyrtodactylus sp. ‘Trilolo River’ e. 1 
Cyrtodactylus sp. ‘Manucoco’ e. 5 
Cyrtodactylus sp. ' Ataüro coast? e. 5 
Cyrtodactylus sp. ‘Abanat River’ e. 3 
Cyrtodactylus sp. ‘Plateau’ e. 4 
Cyrtodactylus sp. 'Com small e. 4 
Cyrtodactylus sp. ‘Com large’ e. 4 
Cyrtodactylus celatus O 6 
Cyrtodactylus sp. incertae sedis e. e. e. 4 
Gehyra mutilata e. e. e. e. e. e. 1,4 
Gekko gecko e. e. e. e. e. e. e. e. e. e. 1-5 
Hemidactylus frenatus e. e. e. e. e. e. e. e. e. 1-5 
Hemidactylus cf. garnotii e. 4 
Hemidactylus platyurus e. e. e. e. e. e. e. 1-4 
Hemidactylus tenkatei [7 e. e. e. 1-4 
Hemiphyllodactylus cf. typus e. 4 
Carlia peronii O 7 
Carlia spinauris O 7,8 
Carlia sp. ‘Maubisse’ e. 1 
Carlia sp. ‘Meleotegi River’ e. 1,2 
Carlia sp. ‘South Coast e. e. ° e 1,2,4 
Carlia sp. ‘Baucau’ e. 1,2 
Carlia sp. ‘Abanat River’ e. 3 
Carlia sp. incertae sedis e. 4 
Cryptoblepharus leschenault e. e. e. 9 1,2,4,5 
Cryptoblepharus sp. ‘Bakhita’ e. 2 
Cryptoblepharus cf. schlegelianus e. e. O 4,9 
Eremiascincus antoniorum O 10 
Eremiascincus cf. timorensis e. O 4,10 
Eremiascincus sp. ‘Ermera’ e. 4 
Eremiascincus sp. ‘Montane’ e. e. 1,2 
Eremiascincus sp. *Lautém" e. 1,2 
Eremiascincus sp. ‘Jaco’ e. 4 
Eremiascincus sp. * Ataüro* e. 5 
Eutropis cf. multifasciata e. e. e. e. e. e. 1-5 
Lamprolepis smaragdina cf. elberti e. e. e. e. e. e. e. e. e. 1-5 
Sphenomorphus cf. melanopogon e. e. 1,2,4 
Sphenomorphus sp. ‘Highland large’ e. 1,2 
Sphenomorphus sp. incertae sedis e. e. e. e e 1,2,4 
Varanus timorensis e e e e O e 1,2,4,11 
Varanus cf. salvator e. 5 

"References are identified numerically as follows: 1 = Kaiser et al., 2011; 2 = O'Shea et al., 2012; 3 = Sanchez et al., 2012; 4 = this paper; 
5 = Kaiser et al., 2013b; 6 = Kathriner et al., 2014; 7 = Zug, 2010; 8 = Smith, 1927; 9 = Brongersma, 1942; 10 = Aplin et al., 1993; 11 = 
Bethencourt Ferreira, 1898. 

No.2 Mark O'SHEA et al. 

for representatives of this versatile gecko genus on Timor 
appears to be the availability of hiding places. Beyond 
this, we have encountered representatives of putative, 
undescribed species in habitats ranging from the wall of a 
cave in limestone karst (Raga, Locality 27) to the vertical 
walls of man-made tunnels (Venilale, Locality 22), and 
from wet lowland forest (Nancuro, Locality 20) to dry 
montane forest (Maganuto, Locality 11). Having had all 
of these encounters, it appears obvious to us that members 
of Cyrtodactylus on Timor display a considerable 
ecological plasticity when it comes to colonizing new 

Timor-Leste Herpetofauna Updates 87 

habitats and adapting. 

On Ataüro Island, the lowland population appears to 
occur in most sampled habitats from near-coastal cliffs 
to disturbed localities, such as plantations or residential 
areas. Whereas the majority of our specimens came 
from areas near a source of water (e.g., in proximity to 
a riverbed, a shallow ravine with water run-off), some 
were found under rocks in Barry's Place Ecoresort, or by 
rolling palm logs and pulling apart palm leaf piles in a 

Some of the microhabitats where we discovered 

Figure 11 Species of Cyrtodactylus in Timor-Leste. These six individuals represent populations of bent-toed geckos we consider distinct at 
the species level (Kathriner et al., in prep.). We refer to them here by their localities. (A) Adult specimen (sex not determined, SVL = 60 mm) 
of the ‘Plateau’ population from the wall of a limestone karst cave near Raça (USNM 579408, Locality 27). (B) Adult male (SVL = 42 mm) 
of the small north coast bent-toed gecko from the ruin of the Portuguese pousada at Com (USNM 579412, Locality 26). (C) Adult male (SVL 
= 55 mm) from a wall in the man-made tunnels at Venilale (USNM 580474, Locality 22). (D) Adult female (SVL = 41 mm) from inside a 
rotten log in the coastal wet forest at Nancuro (USNM 580486, Locality 20). (E) Adult individual (sex not determined, SVL — 44 mm) from a 
dry rotting tree in the alpine habitat at Maganuto (USNM 580487, Locality 11). (F) Adult male (SVL = 40 mm) from a fallen log in disturbed 
dry forest at Fiuren (USNM 580488, Locality 9). 

88 Asian Herpetological Research 

Vol. 6 

Cyrtodactylus include oddities. For example, our highest 
elevation specimen (1036 m, Maganuto; Locality 11) was 
recovered from the inside of a decaying tree that stood 
isolated in an alpine meadow surrounded by very little 
vegetation. Our search there was focused on H. garnotii 
(see below), individuals of which we had found nearby 
in decaying bamboo microhabitat, and when breaking 
apart the decaying wood, a single Cyrtodactylus was 
discovered. A second unusual locality was the rather 
disturbed forest habitat in Fiuren (Locality 9) that 
appeared to endure regular disturbance from the foraging 
activity of a group of free-roaming domestic pigs. The 
most unusual locality, however, were the roadside tunnels 
at Venilale (Locality 22). These tunnels were dug by 
forced labor during the Japanese occupation of Timor in 
the 1940s, and upon our first visit to the locality in 2009 
we did not even consider conducting a careful search 
for reptiles there. While showing this locality to some 
of our team members in 2012, however, we chanced 
upon a gecko at head height on the surface of a vertical 
tunnel wall. Our subsequent, careful search revealed 
several additional specimens distributed throughout 
the tunnel system, including all tunnel surfaces (sides, 
floor, and ceiling), and in both exposed positions as well 
as underneath rocky debris. There was no evidence of 
standing or running water in the tunnels, although the air 
was cool and the humidity high. 
Taxonomic comments. Cyrtodactylus is the largest genus 
in the Gekkonidae, indeed the most speciose in the entire 
Gekkota, a highly diverse group that comprises seven 
families, over 100 genera, and around 1400 species. 
Cyrtodactylus currently comprises one eighth of that 
diversity (199 species; Uetz and Hošek, 2014; Wood 
et al., 2012), distributed from Tibet, China and India 
to northern Australia (Western Australia, Queensland) 
and east across the Indonesian island chain and New 
Guinea into the Solomon Islands, with new species 
being described at considerable frequency. Despite the 
geographic position of Timor near the center of this range, 
the only previous record of the genus from the territory 
of what is now Timor-Leste was a single specimen of 
*Gymnodactylus timorensis’ listed by Duméril and Bibron 
(1836). However, this specimen is neither a member 
of the genus Cyrtodactylus, nor did it originate from 
anywhere close to the island of Timor (L. L. Grismer, in 
prep.; HK, unpubl. data). Therefore, prior to the initiation 
of this project, Timor-Leste was considered devoid of any 
representatives of the genus Cyrtodactylus. 

Our fieldwork soon proved this not to be the case 
as the first six populations sampled during the project 

were found to represent six different species, from at 
least two different larger clades (AK, unpubl. data). 
The Ataüro coastal population has similarities to the 
regionally endemic C. darmandvillei (Weber, 1890) but 
some morphological characteristics lead us to consider 
this population as a potentially new species, here called 
* Ataüro coast’ (Cyrtodactylus sp. 2 of Kaiser et al., 
2013b). We only possess a single specimen of the montane 
Cyrtodactylus sp. ‘Manucoco’ but it clearly represents a 
different taxon from its lowland neighbor based on both 
morphological and molecular data (AK, unpubl. data). 
The five mainland populations also represent distinct 
and separate species, which currently lack names and are 
therefore documented here as Cyrtodactylus sp. ‘Trilolo 
River,’ Cyrtodactylus sp. ‘Abanat River,’ Cyrtodactylus 
sp. ‘Plateau’ (Figure 11A), Cyrtodactylus sp. ‘Com 
small’ (Figure 11B), and Cyrtodactylus sp. ‘Com large.’ 
The taxonomic status of the four more recently sampled 
populations, from Venilale (Baucau District; Figure 11C), 
Nancuro (Manatuto District; Figure 11D), and Maganuto 
and Fiuren (both Bobobaro District; Figs. 11E and 11F, 
respectively) has yet to be determined, and we list them 
here as populations incertae sedis. In addition Kathriner 
et al. (2014b) described C. celatus from near Kupang, 
West Timor, from a single specimen collected in 1924 by 
M. A. Smith, and deposited, then essentially forgotten, in 
the Natural History Museum, London, United Kingdom. 
Three of the populations we discovered stand out 
by their body size (up to 75 mm SVL), including the 
highland karst dwellers at Raça and Mainina (Localities 
27 and 29, respectively), the lowland coastal limestone 
form at Com (Locality 26), and the lowland form 
on Ataüro. A preliminary analysis of molecular data 
(Kathriner et al., unpubl. data) indicates that the larger 
mainland populations likely constitute a separate radiation 
from the small-bodied forms (up to 46 mm SVL). While 
it 1s too early to determine their exact phylogenetic 
affinities or the vector by which they arrived on Timor, 
there appears to have been sufficient time elapsed for the 
two major radiations to adapt to diverse niches and to 
diversify into an unexpectedly rich bent-toed gecko fauna. 

) [IV-vir] 

Gehyra mutilata (Wiegmann, 1834 
Common names. (E) Mutilated Gecko. (G) Vierklauen- 
Gecko. (T) Teki kulit kanek (Teki = small gecko, kulit = 
skin, kanek = injured). 

Known distribution. Gehyra mutilata (Figure 12) has so 
far been reported from only two districts (Table 3), from 
Dili and Lautém, as well as on Mt. Manucoco, Ataüro 
Island, Dili District (Kaiser et al., 2011, 2013b). 

New localities. During the last four surveys additional 

No.2 Mark O'SHEA et al. 

specimens were obtained from sea level to an elevation 
of 572 m on Ataüro Island (Kaiser et al., 2013b), and in 
Lautém District from sea level habitats at Com (Locality 
26) to the elevated central limestone plateau at Raca 
(elevation > 400 m; Locality 27). Additional lowland 
records from Phases IV-VII on the mainland came from 
the confluence of the Comoro and Bemos Rivers (8 km 
S of the Comoro River bridge, Liquiga District; Locality 
6; USNM 579425), and the wet coastal forest at Nancuro 
(Natarbora, 8 km S Umaboco, Manatuto District; Locality 
20; USNM 581759), while upland localities include the 
ruins of Escola do Reino de Haudere, Baguia (Baucau 
District; Locality 24; USNM 580489), and the grazed 
forest at Fiuren, near Balibo (Bobonaro District; Locality 
9; USNM 580490—91). On the mainland Gehyra mutilata 
has now been recorded from five districts, on both the 
northern and southern coastal lowlands, at altitudes ^ 400 
m in the central massif of Timor and > 570 m on Ataüro 
Island (Table 3). 

Natural history. Specimens of G. mutilata have been 
recovered from the standard set of microhabitats typically 
frequented by house geckos (see natural history comments 
on the species of the genus Hemidactylus below). Most 
frequently, these geckos were found associated with dry 
wooden structures, such as the loose bark on decaying 
trees, in dry leaf litter, or in the building materials used to 
make traditional huts. They were also collected from the 
walls of houses. Occasionally, a specimen was retrieved 
from underneath dry rocks (such as in a rock pile) or by 
rolling rocks in dry habitats. 

Gekko gecko (Linnaeus, 1758) "YY"! 

Common names. (E) Tokay Gecko. (G) Tokeh, Tokee, 
Panthergecko. (T) Toke. 

Known distribution. Gekko gecko (Figure 13) has so 
far been reported from five of Timor-Leste’s 13 districts 
(Lautém, Liquiga, Manufahi, Oecusse, and Viqueque; 
Kaiser et al., 2011; O’Shea et al., 2012; Sanchez et al., 
2012) and from Atatro Island, Dili District (Kaiser et al., 
2013b) at elevations from near sea level to over 500 m 
(Table 3). 

New localities. During 2011 and 2012 four more 
mainland districts were documented as part of the range 
for Gekko gecko in Timor-Leste (Baucau, Bobonaro, Dili, 
Manatuto). Since this is an introduced species and there 
are no arguments regarding its identity or taxonomy, we 
collected only few voucher specimens whenever it was 
encountered; some of our records therefore comprise a 
voucher photograph rather than a specimen. In addition, 
this is the most vocal member of the Gekkonidae on 
the island of Timor, and it possesses a characteristic, 

Timor-Leste Herpetofauna Updates 89 

eponymous vocalization. Individuals issuing the 
onomatopoeic “toh-kay” call are frequently heard in 
forests, on rocky outcrops or buildings, both by night and 

Voucher specimens were collected along the Comoro 

River (Dili District; Locality 1; USNM 579314), at 
Betano ‘dry site’ (Manufahi District; Locality 18; 
USNM 579315), and near Raga (Lautém District; 
Locality 27; USNM 579316-17). Voucher photographs 
were contributed for the wet coastal forest at Nancuro 
(Natarbora, 8 km S Umaboco, Manatuto District; Locality 
20; USNM-HI 2824), Com village (Lautém District; 
Locality 26; USNM-HI 2764), the ruins of Escola do 
Reino de Haudere (Baguia, Baucau District; Locality 
24; USNM-HI 2759-60), and from the grazed forest at 
Fiuren (Bobobaro District; Locality 9; USNM-HI 2762). 
Aural observations were made in the forest on Jaco 
Island (Lautém District; Locality 30) and along a rocky 
outcrop at Maganuto (Bobonaro District; Locality 11) for 
specimens that could be heard but not seen. Gekko gecko 
is now known from ten districts, including Ataüro Island 
(Table 3). 
Natural history. Gekko gecko is the largest member of 
the Gekkonidae in the Lesser Sunda Archipelago, and one 
of the most visible elements of the gecko fauna anywhere. 
As a predator of significant size (we encountered adult 
specimens with SVL in excess of 22 cm), this is not a 
species that needs to hide itself but tends to threaten 
when disturbed. Our relatively frequent encounters with 
this species have happened during both day and night 
and we have seen adults, juveniles, and eggs containing 
developing embryos (but destroyed by local children 
as sport) during both wet and dry season surveys. This 
species is familiar to the local population throughout 
the country, and we believe its range encompasses all of 

Hemidactylus frenatus Schlegel, 1836 "1 

Common names. (E) Common House Gecko. (G) 
Asiatischer Hausgecko. (T) Teki uma baibain frenatus 
(teki = small gecko, uma = house, baibain = common). 
Known distribution. Hemidactylus frenatus (Figure 14) 
has so far been reported from seven of Timor-Leste’s 
13 districts (Baucau, Dili, Lautém, Liquiga, Manatuto, 
Oecusse, and Viqueque; Kaiser et al., 2011; O’Shea et al., 
2012; Sanchez et al., 2012) and from Atatro Island, Dili 
District (Kaiser et al., 2013b) (Table 3). 

New localities. We here report additional voucher 
specimens from mainland Dili District on the shoreline 
at Tasi Tolu, the grounds of the Timor Lodge Hotel, the 
mangrove swamp at Metinaro, and from the Comoro 

90 Asian Herpetological Research 

Vol. 6 

Figure 12 Adult male Gehyra mutilata from a fallen log at Fiuren 
(USNM 580490, Locality 9). Photo by Mark O'Shea. 

Figure 13 Subadult Gekko gecko still showing the distinct juvenile 
tail pattern. This specimen was not vouchered. Photo by Mark 

Figure 14 Adult Hemidactylus frenatus (sex not determined) from 
the tidal rocks at Tasi Tolu, near Dili (USNM 580494, Locality 1). 
This individual is a good example of the habitat plasticity displayed 
by house geckos, as it was discovered in an area near the tidal 
splash zone that it shared with individuals of Cryptoblepharus cf. 
schlegelianus. Photo by Mark O'Shea. 

River (Localities 1, 3 and 5; USNM 579726, 579731-32, 
579736, 580494, 581746). Vouchers were also taken 
at the confluence of the Comoro and Bemos Rivers, on 
the Liquiga bank (Locality 6; USNM 579425, 579431). 
Further specimens were obtained from Lautém District, 
from Com at sea level, and from Raga on the central 
limestone plateau (Localities 26 and 27; USNM 579428— 
40, 581755—56). Other low-lying locations sampled 
during 2011 and 2012 produced vouchers from both the 
‘wet site’ and ‘dry site’ at Betano (Manufahi District; 
Locality 18; USNM 581753—54), and the grounds of 
the Castelo Fronteira Guest House, Suai (Covalima 
District; Locality 13; USNM 581747). Vouchers were 
also obtained from upland localities, such as the ruins 
of the Escola do Reino de Haldere, Baguia (Baucau 
District; Locality 24; USNM 580492-93). The Covalima 
and Manufahi District records constitute first records for 
these districts, elevating the number of districts where H. 
frenatus has been recorded to nine (Table 3). We believe 
that this species is likely found associated with human 
disturbances almost anywhere on Timor Island, certainly 
at elevations between sea level and 600 m (see Natural 
history for H. cf. garnotii). 

Natural history. Throughout all of our surveys, this 
species is clearly the most frequently encountered gecko. 
Due to its perianthropic lifestyle, it is encountered on the 
walls of almost any human habitations. These geckos 
are able to colonize even new construction rapidly and 
indiscriminately, and they appear to live in clean hotel 
rooms just as well as in natural vegetation, rock piles, 
or even trash. We have not encountered them 1n pristine 
habitats, with the exception of healthy-looking forest 
areas experiencing some minor form of human impact, 
such as those adjacent to coffee plantations. We believe 
that the species exists in all of Timor-Leste's districts, 
and we believe its arrival on the island and its dispersal 
throughout the country may be correlated with historic 
and current local trade patterns. 

Hemidactylus cf. garnotii "™ 

Common names. (E) Indo-Pacific House Gecko. (G) 
Indopazifischer Halbfinger-Gecko, Jungfern-Halbfinger- 
Gecko. (T) Teki uma baibain garnotii (teki = small gecko, 
uma = house, baibain = common). 

Identification. Hemidactylus cf. garnotii (Figure 15) is 
the fourth house gecko species (genus Hemidactylus) 
recorded from Timor-Leste, and especially in preservative 
it is one easily confused with the more common species 
(e.g., H. frenatus, H. tenkatei). Specimens encountered in 
Bobonaro were dark brown when collected, with several 
longitudinal rows or dark-edged light spots on the dorsum 

No.2 Mark O'SHEA et al. 

and a prominent ventrolateral series of white spines along 
the edge of the tail. The dorsal color paled in captivity 
but the light spots and white caudal spines were still in 
evidence. Hemidactylus cf. garnotii can be distinguished 
from H. platyurus by a tail that is not dorsoventrally 
flattened and by the absence of skin webbing and fringing 
associated with tail, digits, limbs, and flanks; from 
H. tenkatei by the absence of that species' distinctive 
longitudinal rows of raised dorsal tubercles, and from 
H. frenatus by a series of small scales that separate the 
2™ pair of postmental scales from the infralabials (both 
pairs of postmentals are in contact with the infralabials 
in H. frenatus). Hemidactylus frenatus also has four 
longitudinal rows of elevated spines on its original tail, 
whereas in H. cf. garnotii the character of tubercle rows 
is limited to two lateral rows. Hemidactylus cf. garnotii 
is easily distinguished from Gehyra mutilata by its longer 
and flatter snout and the pattern of chin scales. Several 
of our specimens had symmetrical calcium deposits 
in the neck area (Figure 15B), which is something we 
never observed in the other house gecko species found in 

Known distribution. There were no previous records of 
this species for Timor-Leste. 

New localities. Hemidactylus cf. garnotii was 
encountered only during the survey in 2012 (Phase VII) 
when seven vouchers were collected in the bamboo 
forest above Maganuto, near Maliana (Bobonaro District; 
Locality 11; USNM 580495—501) and a single voucher 
obtained from the degraded, grazed forest at Fiuren, near 
Balibo (Bobonaro District; Locality 9; USNM 580502). 
The Fiuren specimen came from an elevation of 463 m 
but the Maganuto specimens were collected at 1041—1063 
m on the slopes of Mt. Leolaco at an altitude far above 
that recorded for any other Hemidactylus species in 

Natural history. The seven specimens collected in the 
bamboo forest above Maganuto were mostly sheltering 
at the bases of bamboo leaf-axils or in termite-inhabited 
dead bamboo stalks, but one specimen was found 
under a rock and another behind the bark of a tree in 
close proximity to the bamboo. Several specimens had 
regenerated tails, and one (USNM 580498) had lost both 
its left fore- and hind limbs but had healed and survived 
the trauma. The Fiuren specimen, containing two eggs, 
was also found inside a clump of bamboo. 

Hemidactylus garnotii Duméril and Bibron, 1836 
is an all-female parthenogenetic species and should be 
considered a good colonizer: only a single adult female 
is needed to produce eggs to establish a new colony. It is 

Timor-Leste Herpetofauna Updates 91 

therefore somewhat surprising that its reproductive ability 
has not made this species more prevalent in Timor-Leste. 
We believe that it may be its reduced genetic variability, 
inherent in clonally reproducing organisms, that gives this 
species only few options to successfully compete with 
aggressive bisexual species, such as H. frenatus or H. 
tenkatei. If it is difficult for H. garnotii to live in sympatry 
with other house geckos, unlike Gehyra mutilata or H. 
platyurus, its presence and apparent success on the slopes 
of Mt. Leolaco at elevations above 1000 m might be 
explained by the fact that no other house geckos have yet 
been found above 563 m in mainland Timor-Leste. 
Taxonomic comments. Hemidactylus garnotii is a 
colonizing species, which we would most expect to 
encounter in coastal lowland beachheads. Whilst the 
Fiuren record came from a locality which was at an 
intermediate elevation (463 m) and heavily influenced by 
human activities, both being common factors associated 
with colonizing species, the majority of our specimens 
were collected at Maganuto, on the slopes of Mt Leolaco 
at an elevation considerably above that documented 
for any other Timor Hemidactylus (1041-1063 m), in a 
habitat that seemed to us incompatible with a colonizing 
species such as H. garnotii due to its remoteness and 
high elevation. This leads us to wonder if this taxon is an 
undescribed species of garnotii-like Hemidactylus, but in 
the absence of any males we cannot as yet differentiate it 
morphologically from true H. garnotii. We therefore refer 
to it as Hemidactylus cf. garnotii. 

Hemidactylus platyurus (Schneider, 1792) "Y 
Common names. (E) Common Flat-tailed Gecko. (G) 
Saumschwanz-Hausgecko. (T) Teki ikun belar (teki = 
small gecko, belar = flat, ikun = tail). 

Known distribution. Hemidactylus platyurus (Figure 
16) has so far been reported from six of Timor-Leste’s 13 
districts (Dili, Lautém, Liquica, Manatuto, Oecusse, and 
Viqueque; see Kaiser et al., 2011; O’Shea et al., 2012; 
Sanchez et al., 2012). It has not been recorded on Ataüro 
Island (Table 3; see Kaiser et al., 2013b). 

New localities. Additional specimens were collected 
in 2011 in Lautém District, near sea level on the north 
coast at Com (Locality 26; USNM 579445-47) and at 
520 m elevation, 5 km N of Maubesi (Tilomar, Covalima 
District, Locality 15; USNM 581757—58). This is a little 
lower than our elevation record for H. platyurus at 545 
m near Dare, Dili District (Locality 4; USNM 579112) 
during Phase III. Covalima is the seventh district from 
which we have recorded H. platyurus (Table 3). 

Natural history. This is another of the perianthropic 
house gecko species, though it is seen around human 

92 Asian Herpetological Research 

Vol. 6 

Figure 15 Adult Hemidactylus cf. garnotii from a bamboo stand 
above Maganuto (USNM 580502, Locality 11). This individual 
shows the presence of mature eggs and gular calcium deposits. 
Photo by Mark O'Shea. 

habitations in considerably lower numbers than either 
H. frenatus or H. tenkatei. In each of the new localities, 
other house geckos were present, although not all 
cohabiting gecko species were vouchered. The two 
Maubesi specimens were found on a roadside tree that 
initially caught our attention because of the presence of 
a monitor lizard (Varanus; see below). After capturing 
the monitor lizard, we managed to obtain both specimens 
from a height of ca. 5 m above ground level. Both of 
the specimens caught in Com were found along with 
individuals of H. frenatus and H. tenkatei 1n the rafters 
of the cabins at Com Beach Resort and on stone walls 
surrounding the compound. 

Hemidactylus tenkatei van Lidth de Jeude, 1895 "YY"! 
Common names. (E) Roti House Gecko. (G) Roti- 
Hausgecko. (T) Teki uma baibain Roti (teki = small 
gecko, uma = house, baibain = common). 

Identification. Hemidactylus tenkatei (Figure 17) can be 
distinguished from H. frenatus by the presence of 16-20 

longitudinal rows of large, strongly keeled tubercles, 

Figure 16 Adult Hemidactylus platyurus (sex not determined) from 
the wall of a building at the Com Beach Resort (USNM 579447, 
Locality 26). Photo by Mark O'Shea. 

as opposed to the numerous scattered, small conical 
tubercles of its more common congener. It also lacks 
the broad, flattened, filamentous-edged tail and strongly 
webbed toes of H. platyurus. Hemidactylus tenkatei may 
be distinguished from Gehyra mutilata by its chin shields, 
which are arranged to form a smoothly arched posterior 
border in the latter species, and from H. garnotii by the 
presence of enlarged keeled tubercles on its dorsum. 
Known distribution. Hemidactylus tenkatei had 
previously only been recorded from Liquiga and Oecusse 
Districts (Table 3; see O’Shea et al., 2012; Sanchez et al., 

New localities. We here report new district records for 
H. tenkatei from Dili District at Timor Lodge Hotel, Dili 
(Locality 1; USNM 579728—30) and Metinaro mangrove 
swamp (Locality 3; USNM 579733), and from Lautém 
District at Com Beach Resort (Locality 26; USNM 
579417, 579430, 579441—44), elevating the number of 
districts from which this introduced species has been 
recorded to four. All records are from elevations below 
25 m and from northern coastal locations, indicative of 

Figure 17 Adult Hemidactylus tenkatei (sex not determined) from 
a wall in the grounds of the Timor Lodge Hotel in Dili (USNM 
581158, Locality 1). Photo by Mark O'Shea. 

an invading species establishing bridgeheads. The lack 
of any specimens of H. tenkatei further inland could be a 
result of its recent arrival, its inability to compete with the 
already established H. frenatus, H. platyurus, or Gehyra 
mutilata, or its adaptation to a microhabitat that currently 
remains undiscovered. At our accommodation in Dili, the 
Timor Lodge Hotel, we have noticed an increase in the 
abundance of H. tenkatei relative to H. frenatus over the 
five-year period of our survey work, but this observation 
will require further verification. 

Natural history. We collected six specimens of what we 
initially believed to be H. frenatus from trees and rocks in 
the center of a seasonally dry riverbed, west of Maubara 
(Liquiga District: locality 3) on 6 February 2010 (Phase 
II). Upon later examination, one of these (USNM 579064) 

No.2 Mark O'SHEA et al. 

was re-identified as H. cf. tenkatei (A. M. Bauer, pers. 
comm.), the first specimen of the perianthropic H. brookii 
complex recorded from Timor-Leste. With a distribution 
of this species complex ranging from Pakistan and Indian 
Ocean islands to the Philippines and south into the Lesser 
Sunda archipelago (Bauer et al., 2010), ancestors of 
Timorese H. tenkatei may have found their way onto 
Roti Island, the type locality of H. tenkatei, and later on 
to Timor Island by stowing away with neolithic human 
migrants and their chattels. 

Taxonomic comments. Geckos called Hemidactylus 
brookii exist 1n museum collections from throughout 
South and Southeast Asia, and the broad distribution 
and the likely influence of historical human trading and 
colonization patterns has led to an inconsistent use of 
names for these forms. Recently, Bauer et al. (2010) 
completed a molecular analysis, in which they restricted 
the distribution of true H. brookii to Borneo, Peninsular 
Malaysia, Burma, and Karnataka State in India. However, 
their analysis conspicuously excluded data from islands 
of the Indonesian Archipelago, notated with a centrally 
placed question mark in their distribution map (Figure 1 
in Bauer et al., 2010). 

The species H. tenkatei was described by van Lidth de 
Jeude (1895) based on three specimens from Roti, a small 
(1200 km’) island off the extreme southwestern corner 
of Timor. Two decades later, de Rooij (1915) placed the 
species into the synonymy of H. brookii after a limited 
study of specimens from Flores and Wetar, presumably 
with literature accounts then available, but without the 
presentation of data. In two recent revisions of the H. 
brookii group, of which H. tenkatei is a member, Rósler 
and Glaw (2010) and Mahony (2011) removed H. tenkatei 
from the synonymy of H. brookii, but did not examine 
the relevant type material. Addition of these important 
specimens to the analysis, along with the Bornean type 
material of H. brookii and molecular data for specimens 
from Timor-Leste to the data set of Bauer eft al. (2010), 
shows that H. tenkatei is a species distinct from H. 
brookii and that Timorese populations are indeed identical 
to those on Roti (Kathriner et al., 2014a). Furthermore, it 
appears that the species H. tenkatei is a widespread and 
successful colonizer found not only in the Lesser Sundas 
but also in Sarawak, Borneo, and Penang Island, Malaysia 
(Kathriner et al., 2014a), and that these populations can 
therefore all be referred to H. tenkatei. 


Hemiphyllodactylus cf. typus 
Common names. (E) Dwarf Tree Gecko. (G) 
Zigeunergecko, Gewóhnlicher Halbblattfinger-Gecko. (T) 
Teki ai isin lotuk (teki = gecko, ai = tree, isin lotuk = very 

Timor-Leste Herpetofauna Updates 93 

small body). 

Identification. Hemiphyllodactylus cf. typus (Figure 18) 
is the smallest gecko in the region and easily overlooked, 
as it had been during six previous phases of our survey. 
This is an extremely slender, etiolated gecko, its body 
so elongated that the adpressed limbs do not overlap or 
even come into contact. It can be distinguished from 
Hemidactylus spp. by its clawless 1* digit, a characteristic 
it only shares with members of the genus Lepidodactylus, 
a taxon as yet unrecorded from Timor, and the complete 
lack of any enlarged postmental scales in the chin region. 
In L. lugubris the clawless 1“ digit is otherwise well 
developed, being at least two-thirds the length of the 2" 
digit, whereas in H. cf. typus the 1* digit is much reduced 
in size. 

Known distribution. There were no previous records of 
this genus from Timor-Leste. 

New localities. Two specimens of Hemiphyllodactylus cf. 
typus were collected at Ossohuna, near Baguia (Baucau 
District; Locality 25) during Phase VIII, the first record of 
the taxon from Timor Island. 

Natural history. The only two specimens of H. cf. typus 
(USNM 580503—04) found in Timor-Leste so far were 
collected in a clump of bamboo in a dry river gorge, 
sheltering behind the leaf-like culm sheaths that protect 
the base of the bamboo shoots. Their movements, when 
uncovered, were slow, meaning they did not ‘scamper’ as 
do many species of Hemidactylus. 

Taxonomic comments. The Indo-Pacific genus 
Hemiphyllodactylus contains as many as 20 species 
although most exhibit fairly or extremely localized 
distributions (Zug, 2010b; Grismer et al., 2013, 2014). 
The one widespread species is the parthenogenetic H. 

typus Bleeker, 1860, which is found from southern 
Myanmar and Taiwan of China to New Guinea and across 
the South Pacific to Fiji and Tonga, with established but 
isolated populations in Sri Lanka, the Mascarene Islands, 

Figure 18 Adult Hemiphyllodactylus cf. typus from a bamboo 
stand near Ossohuna (USNM 580503, Locality 25). Photo by Mark 

94 Asian Herpetological Research 

Vol. 6 

and the Hawaiian Islands (Zug, 2010b, 2013). This is a 
colonizing species that often goes undetected due to its 
small size and secretive nature, so its true distribution 
is incompletely documented (Zug, 2010b). Small 
beachhead populations of parthenogenetic geckos are 
most commonly found in lowland coastal localities where 
they have become established, either through the actions 
of man or by some other means, such as rafting. 

The population recorded here 1s located near 
Ossohuna, 22 km from the north coast and 10 km from 
the south coast of Timor-Leste, at an elevation of 938 
m. Although the distances to either coast are not great, 
the road from the north coast to Ossohuna is rough, 
long, and winding and the road from the south coast 
is only accessible seasonally by vehicles with off-road 
capability and by no means a reliable transport connector. 
This leads us to query whether the specimens represent 
the parthenogenetic H. typus or an undescribed sexual 
species, such as occur at inland locations in India, China, 
Southeast Asia, Sumatra, and Borneo. In most characters 
examined, the Timor specimens appear to fall within the 
characteristics of H. typus as detailed by Zug (2010b), 
and given that the only specimens collected to date are 
a juvenile and an adult female we are unable to disprove 
the parthenogenetic species argument. However, in light 
of the cryptic diversity seen in mainland Southeast Asian 
Hemiphyllodactylus populations (Grismer et al., 2013, 
2014), a molecular analysis of the Timor specimens is 
now being conducted (P. Wood, in prep.). 

Family Scincidae—Skinks 

Genus Carlia "YY" 

Common names. (E) Four-fingered Skinks, Rainbow 
Skinks. (G) Regenbogen-Skinke. (T) Mamór liman-fuan 
haat (mamór = skink, haat = four, liman fuan = finger). 
Known distribution. During Phases I-III we collected 
Carlia in six of Timor’s mainland districts (Ainaro, 
Baucau, Ermera, Lautém, Oecusse, and Viqueque), but 
did not locate the genus on Atatro Island (Table 3), 
despite reports of the genus from Alor to the northwest 
and Wetar to the northeast (Zug, 2010a). Our vouchers 
comprised two apparently montane forms: Carlia sp. 
*Maubisse' (Figure 19A) from Ainaro District (Maubisse; 
Locality 16), and Carlia sp. ‘Meleotegi River’ (Figure 
19B) from Ermera District (Sta. Bakhita Mission and 
Meleotegi River; Locality 8), and three seemingly 
lowland forms: Carlia sp. ‘South Coast’ (Figure 19C) 
from Loré village, southeast Lautém District and Beacu 
on the coast of Viqueque District, Carlia sp. ‘Baucau’ 
(Figure 19D) from Afacaimau, Baucau District (Locality 
23), and Carlia sp. ‘Abanat River’ from the Oecusse 

District. For reports from these localities, see Kaiser et al. 
(2011), O’Shea et al. (2012), and Sanchez et al. (2012). 
New localities. During 2011 and 2012 (Phases IV- 
VII) we collected additional vouchers of all the above 
species, except Carlia sp. ‘Abanat River.’ Carlia spp. 
*Maubisse' and ‘Meleotegi River,’ which were only 
found at their original locations. However, vouchers 
of Carlia sp. ‘South Coast’ were collected as a series 
from Nancuro (Natarbora, 8 km S Umaboco, Manatuto 
District; Locality 20; USNM 579319—27), and as single 
specimens from the southern shore of Lake Lenas (near 
Fatucahi, Manufahi District; Locality 19; USNM 579328) 
and a roadside ditch on the road between Fatucahi and 
Betano (Manufahi District; Locality 18; USNM 579329), 
greatly extending the westerly range of this taxon from 
Beacu, Viqueque District, and providing the first Carlia 
records for Manatuto and Manufahi Districts. Another 
single specimen obtained by one of us (LLA) at the 
Betano “wet site” may also belong to this taxon and 
extends the range further west, although it is currently 
documented as Carlia incertae sedis. Carlia sp. ‘South 
Coast’ was also collected for the first time along the north 
coast, when two specimens were obtained from the ruins 
of the Pousada de Com (Lautém District; Locality 26; 
USNM 579448-49). A single additional specimen of 
Carlia sp. ‘Baucau’ was collected at Afacaimau (Baucau 
District; Locality 23; USNM 580506), a site known to the 
project as the “Carlia spot," and another single specimen, 
seemingly also of Carlia sp. ‘Baucau,’ was collected on 
the sandstone cliff above the Japanese caves at Venilale 
(Baucau District; Locality 22; USNM 580505), although 
this specimen was taken at an elevation of 675 m while 
the *Carlia spot" vouchers were collected at 290—370 m. 
Carlia populations have now been documented for eight 
mainland districts (Table 3). 

Natural history. Members of the genus Carlia in Timor- 
Leste appear to be habitat generalists, found in both 
dry and moist habitats, as well as both pristine and 
disturbed areas. When out in the open, we have observed 
individuals foraging in and around leaf litter and decaying 
plant material, or basking on exposed “perches,” such 
as small boulders, tree trunks, fallen banana plants, or 
retaining walls near human habitations. These lizards 
also interact with one another by signaling (e.g., tail 
waving: Langkilde et al., 2004; O'Shea, 1993) and 
were occasionally observed chasing each other as part 
of aggressive or mating encounters. Where they occur, 
Carlia can be very abundant lizards: at the Sta. Bakhita 
Mission, Carlia sp. ‘Meleotegi’ occurs at numbers of 
perhaps as many as one or two individuals per m° on the 

No.2 Mark O'SHEA et al. Timor-Leste Herpetofauna Updates 95 

Figure 19 Representatives of four populations of four-fingered skinks (genus Carlia) we consider to be distinct at the species level. 
Important characteristics to differentiate these forms in the field include the coloration of the throat and venter of breading males as well as 
the presence, color, and extent of lateral stripes in both sexes. (A) Adult female (SVL — 44 mm) from the grounds of the Portuguese Pousada 
at Maubisse (USNM 579334, Locality 16). (B) Adult male (SVL = 46 mm) from man-made gardens at the Sta. Bakhita Mission (USNM 
5779450, Locality 8). (C) Adult male (SVL — 42 mm) from among the leaf litter in wet coastal forest at Nancuro (USNM 579324, Locality 
20). (D) Adult male (SVL — 40 mm) from banana plant debris in an agricultural environment (USNM 580506, Locality 23). Photos by Mark 


terraced hillside. In other localities, abundance clearly 
depends on the presence of a potential source of food 
(e.g., invertebrates in a pile of decaying leaves). Only 
rarely did we encounter single individuals. 

We have been unable to observe particular adaptive 
specializations among the four or five putative taxa 
occurring in Timor-Leste (see Taxonomic comments 
below), which can be expected when dealing with a 
habitat generalist. One of us (SM) was able to observe 
that male Carlia from highland locations (Bakhita and 
Meleotegi) held in captivity become flushed with color 
during the breeding season. Females, from those locations, 
however, may also show a color change towards a more 
intense coloration (limited to a mid-lateral stripe), which 
is related to reproductive readiness. On the other hand, 
such drastic changes in coloration in specimens from the 
lowland rainforest of Nancuro were not observed (SM, 
pers. obs.). More detailed observations will be possible 

once the taxonomic status of these populations has been 
Taxonomic comments. Prior to the initiation of this 
survey in 2009, two species of Carlia were documented 
for the island of Timor: Carlia peronii (Duméril and 
Bibron, 1839) and C. spinauris (Smith, 1927). Although 
Greer (1976) treated C. spinauris as a synonym of C. 
peronii, Zug (20102) recognized them to be separate but 
related species within the C. peronii species group (sensu 
Greer, 1976), a group that also extends onto other islands 
in Indonesia's provinces of East Nusa Tenggara (e.g., 
Roti, Semau, Alor) and southern Maluku (e.g., Wetar, 
Kisar). This species group also includes the recently 
described C. sukur Zug and Kaiser, 2014 from Pulau 
Sukur, a small island north of Flores (Zug and Kaiser, 

In addition to a suite of morphological and 
morphometric characters, Zug (20102) separated C. 

96 Asian Herpetological Research 

Vol. 6 

peronii and C. spinauris spatially, stating that the former 
was a lowland species, whereas the latter was a highland 
species. The type locality for C. peronii was erroneously 
given as “ile de France" (= Mauritius), having been 
reassigned to Kupang, West Timor by Greer (1976), the 
only location on Timor visited by the collector, Frangois 
Auguste Péron. This species is also known from other 
low-lying locations to the east of Kupang (e.g., Kokabris, 
Noil Toko, Djamplong = Camplong). In contrast, the 
type locality for C. spinauris is Lelogama (elevation 750 
m), where it was personally collected by M. A. Smith 
and his wife in 1924, and it was also recorded from Soé 
(elevation 800 m) by de Jong (1927). To date neither 
of these species has been recorded in Timor-Leste. The 
material available to us has already undergone preliminary 
molecular analysis and there is strong evidence to support 
the recognition of four or five different species, distinct 
from the aforementioned West Timorese taxa. 


Cryptoblepharus leschenault (Cocteau, 1832) 
Common names. (E) Leschenault's snake-eyed skink. 
(G) Leschault-Schlangenaugenskink. (T) Mamór matan 
samea leschenault (mamór = skink, matan = eye, samea = 

Known distribution. During Phase I (2009) 
Cryptoblepharus leschenault (Figure 20) was documented 
from lowland locations in Lautém and Baucau Districts, 
with three and one vouchers collected, respectively 
(Kaiser et al., 2011), and a single voucher was collected 
from a coastal location on Ataüro Island, part of 
Dili District (Kaiser et al., 2013b). Cryptoblepharus 
leschenault 1s therefore known from three districts to 

New localities. Single vouchers of C. leschenault were 
collected on each of our visits to the coastal forest 
at Nancuro (Natarbora, 8 km S Umaboco, Manatuto 
District; Locality 20; USNM 579335, 580520), bringing 
to four the number of districts where the species has been 
collected (Table 3). 

Natural history. Cryptoblepharus leschenault is an 
infrequently encountered species, but where it occurs 
it may be relatively abundant but difficult to capture. 
Specimens would run rapidly up the trunks of large 
hardwood trees, from where they could usually only be 
captured using blowpipes. Despite intensive searches 
in many locations these small skinks appeared to be 
much more patchily distributed than the other tree- 
bole inhabiting lizards, the larger Draco timoriensis and 
Lamprolepis smaragdina cf. elberti. The two Nancuro 
specimens collected during Phase IV (2011) and Phase 
VII (2012) were also found on the boles of trees, 5.0 m 

Figure 20 Adult male of Cryptoblepharus leschenault from 3 m 
above ground on the trunk of a tree in wet coastal forest at Nancuro 
(USNM 580520, Locality 20). Photo by Mark O'Shea. 

and 3.0 m from the ground, respectively. 

Taxonomic comments. Prior to the initiation of this 
survey two species of Cryptoblepharus had been recorded 
from Timor Island, C. leschenault and C. schlegelianus. 
Whereas the former was only recently documented for 
Timor-Leste (Kaiser et al., 2011), the latter is known 
only from Semau, a small island off West Timor, where 
it apparently occurs in sympatry with C. leschenault 
(Brongersma, 1942). The dorsal pattern of C. leschenault 
consists of a dark background with a pair of narrow light 
dorsolateral stripes from snout to tail and a narrow light 
vertebral stripe from the snout to a point anterior to the 
forelimbs, where it then splits, in the shape of a tuning 
fork, to continue to the tail as a pair of even narrower 
paravertebral stripes. The pattern of C. schlegelianus 
comprises a pale background without a vertebral stripe, 
but with a pair of relatively broad, pale dorsolateral stripes 
above a narrower pair of dark narrow stripes that continue 
to the tail (Horner, 2007). Without the benefit of a detailed 
review of available material, we are unconvinced that C. 
schlegelianus is present in Timor-Leste; the species has 
only been verified for Semau Island, in the absence of 
actual specimens from the western end of Timor (Horner, 
2007; Mertens, 1931), although a very similar, perhaps 
conspecific form is present in Timor-Leste (see account of 
C. cf. schlegelianus below). 

Cryptoblepharus sp. ‘Bakhita’ ™ 

Common names. (E) Bakhita snake-eyed skink. (G) 
Bakhita-Schlangenaugenskink. (T) Mamór matan samea 
bakhita (mamór = skink, matan = eye, samea = snake). 
The common name ‘Bakhita’ is used in reference to 
the Sta. Bakhita Mission, the location from which 
our exploration of the nearby Meleotegi River habitat 

Identification. This hitherto undescribed species of 

No.2 Mark O'SHEA et al. 

Cryptoblepharus (Figure 21) has a dorsal stripe pattern 
similar to that of C. leschenault, but with a critical 
difference. The dorsal pattern of C. leschenault consists 
of a black background with a pair of narrow light yellow 
dorsolateral stripes from snout to tail and a yellow 
vertebral stripe, from the snout to a point anterior to 
the forelimbs, where it then splits into two narrower 
paravertebral stripes that continue to the tail, the overall 
impression being of a ‘tuning-fork’ pattern. In the two 
Meleotegi specimens, the vertebral stripe does not fork on 
the back and continues to the tail as a single stripe. 
Known distribution. During Phase II (2010) a single 
specimen of Cryptoblepharus, collected from a tree on the 
Meleotegi River, near the Sta. Bakhita Mission (Eraulo, 
Ermera District; Locality 8), was considered sufficiently 
distinct from known species (C. leschenault and C. 
schlegelianus—see Taxonomic comments below) to 
warrant recognition as a third Timorese species, pending 
the collection of additional material. 

New localities. During Phase V (2011) a second voucher 
(USNM 579472) was obtained from the same locality as 
in Phase II. 

Natural history. With only two specimens known, 
our knowledge of this species’ natural history is 
obviously very scant. Both specimens were discovered 
at a considerable height above ground on the trunks of 
large trees (as high as 7 m), and their somewhat jerky 
movements and body aspect remind us of other small 
tree-dwelling skinks in Southeast Asia, such as Lipinia 
vittigera (Boulenger, 1894). Both individuals appeared to 
be foraging on the bark surface when first seen, moving 
downwards along the tree trunk. When disturbed they 
reversed course and began moving back up the tree, 
though unhurriedly and once again appearing to forage. 
Specimens of both C. /eschenault and Cryptoblepharus 
sp. ‘Bakhita’ are infrequently encountered, and when seen 
appear as individual lizards without conspecifics present, 
in contrast to C. cf. schlegelianus. 

Taxonomic comments. The presence/absence and 
condition of various types of dorsal and lateral stripes 
is an important characteristic in the recognition of 
Cryptoblepharus species, with a number of species 
(Horner, 2007) exhibiting the ‘tuning-fork’ vertebral stripe 
pattern. These include C. leschenault from Timor and 
Flores and C. balinensis Barbour, 1911 from Bali. Other 
taxa exhibit a non-forking vertebral stripe, including C. 
balinensis sumbawanus Mertens, 1928 from Sumbawa, 
C. renschi Mertens, 1928 from Sumba and Komodo, 
and C. keiensis (Roux, 1910) from the Kei Islands. All 
of the aforementioned taxa occur at elevations up to 

Timor-Leste Herpetofauna Updates 97 

500 m, and in this assemblage highland forms (above 
800 m) are uncommon. The presence of a population of 
Cryptoblepharus in the highlands of Timor exhibiting 
the non-forked vertebral pattern on a dark background 
is therefore indicative of a species undescribed so far 
(Kaiser et al., in prep.). 


Cryptoblepharus cf. schlegelianus 
Common names. (E) Timor north coast snake-eyed 
skink. (G) Schlegel-Schlangenaugenskink. (T) Mamór 
matan samea tasi ibun utara (mamór = skink, matan = 
eye, samea = snake, tasi ibun utara = north coast). 
Identification. Differentiation of this coastal form (Figure 
22) from both other species of Cryptoblepharus so far 
found in Timor-Leste (C. leschenault, Cryptoblepharus 
sp. ‘Bakhita’) is quite simple, considering the absence 
of prominent yellow or cream dorsal stripes. Coastal 
specimens tend to be brown or black with broad (1.5-2.0 
scales wide), lighter brown longitudinal dorsolateral 
stripes over a broader (3.0—4.0 scales wide), darker 
irregular stripe that occupies much of the upper flanks 
of the body. The mid-dorsal region is brown with lighter 
flecking on some of the keeled margins of the scales and 
occasional scattered dark-brown spots. Specimens from 
the Tasi Tolu series (Dili District; Locality 1) had more 
extensive dark markings that obscured the ground color 
and exaggerated the light brown dorsolateral stripes; one 
specimen was virtually melanistic. 

Known distribution. Cryptoblepharus schlegelianus 
Mertens, 1928 1s known from specimens collected on 
Semau Island, off the southwestern part of Timor, near the 
port of Kupang in West Timor, but we have been unable 
to find any specimens from Timor associated with this 
species name in museum collections. We are therefore 
unable to confirm the occurrence of C. schlegelianus 
on mainland Timor. Although Mertens (1928) listed the 
species for Timor in his original description, that listing 1s 
based on material the Senckenberg Museum (Frankfurt, 
Germany) obtained in an exchange from the collections at 
Gießen, Germany, in 1854. Given that the port of Kupang 
was the main shipping center in this region in the early 
part of the 19" century, and given that at least one other 
species’ origin was in error based on shipping and not 
collection locality — Malayopython timorensis (Peters, 
1876), which does not occur on Timor Island (Barker 
and Barker, 1996; O'Shea et al., 2012) — we consider 
the provenance of the Gießen material problematic and 
wonder whether the distribution of C. schlegelianus 
actually includes Timor (see Taxonomic Comment 

New localities. Cryptoblepharus cf. schlegelianus, was 

98 Asian Herpetological Research 

Vol. 6 

Figure 21 Adult male of Cryptoblepharus sp. ‘Bakhita’ from 5 m 
above ground on the trunk of a large tree in coffee forest at the 
Meleotegi River (USNM 579181, Locality 8). Note the absence of 
a forked line pattern, unlike that seen in C. leschenault. Photo by 
Mark O'Shea. 

Sorgen Wiig. 

Figure 22 Adult individual of Cryptoblepharus cf. schlegelianus 
from the rocky shore at Tasi Tolu, Dili (USNM 580513, Locality 1). 
Photo by Mark O'Shea. 

sampled as small series from each of three northern 
coastal locations. During Phase V (2011) a voucher series 
was collected on the wharf at Com (Lautém District; 
Locality 26; USNM 579455-71), in the final days of 
Phase VII (2012) a second voucher series was collected 
along the rocky beach at Tasi Tolu, near Dili (Dili District; 
Locality 1; USNM 580512-19), and during Phase VIII 
(2013) we discovered a population on the other side 
of Dili, below the Cristo Rei monument (Dili District; 
Locality 2; USNM 581114—27). 

Natural history. These skinks were found in densely 
populated colonies, exclusively in locations right at sea 
level. At Com (Locality 26) they were found hiding in 
cracks on the sloping concrete of the wharf walls, or 
hunting in the flotsam, rocks, coral debris, and seaweed 
below the wharf walls. Much of this foraging activity 
was in the saltwater splash zone and while the animals 

seemed unperturbed by the spray, they actively avoided 
swells. Individuals were more commonly encountered on 
the landward, more protected inner side of the wharf but 
were also in evidence on the seaward, outer wall, where 
they were much more exposed to wave activity. At Tasi 
Tolu (Locality 1) skinks were found in almost an identical 
scenario as in Com, on a wharf and on the rocky shore 
right at sea level. Near Cristo Rei (Locality 2), individuals 
were encountered on large boulders, in rocky crevices, as 
well as in the pebbles of the splash zone. 
Cryptoblepharus cf. schlegelianus occurs at much 
greater densities than either C. leschenault or 
Cryptoblepharus sp. ‘Bakhita’ and obviously has a much 
different ecological niche. It displays a propensity to 
forage in the saltwater splash zone, where it will have 
access to terrestrial arthropods feeding on exposed littoral 
vegetation as well as tidal invertebrates, and where 
the food supply would permit the observed population 
Taxonomic comments. In general appearance, specimens 
of C. cf. schlegelianus resemble dark specimens of C. 
schlegelianus from Semau. However, differences in 
pattern, scalation, and ecology (HK, pers. obs.), as well 
as the geographic separation between populations in 
Timor-Leste and the southwestern end of Timor Island 
where Semau is situated, lead us to question whether the 
form found in Timor-Leste is indeed conspecific with C. 
schlegelianus. We therefore conservatively assign the 
name C. cf. schlegelianus to this form. 

Genus Eremiascincus "VV" 

Common names. (E) Night Skinks. (G) Glatte 
Nachtskinke. (T) Mamór kalan (mamór = skink, kalan = 

Known distribution. Night skinks (genus Eremiascincus; 
Figure 23) have been collected on most phases of the 
project, but their status and identity has been the source 
of some confusion (see Taxonomic comments below). 
During Phases I and II, species of Eremiascincus were 
documented from four mainland districts (Ainaro, 
Ermera, Lautém, and Manufahi; see Kaiser et al., 2011; 
O’Shea et al., 2012), followed during Phase VI by the 
first specimens collected on Ataüro Island (Dili District; 
see Kaiser et al., 2013b). Eremiascincus 1s therefore 
known from five districts of Timor-Leste to date. 

New localities. During Phases IV-VIII Eremiascincus 
was again encountered and collected and those records 
pertaining to mainland Timor-Leste and Jaco Island are 
included here. Additional vouchers were obtained from 
the Meleotegi River (Ermera District; Locality 8; USNM 
579474, 579760, 580521—24, 581128—39), Maubisse 

No.2 Mark O'SHEA et al. 

(Ainaro District; Locality 15; USNM 579339-45), and 
Mirbuti village, near Same (Manufahi District; 17; 
USNM 579336-—38). New records for Lautém District 
were supported by voucher material from Raga (Locality 
27; USNM 581762) and Jaco Island (Locality 30; USNM 
579473), the former only as an autotomized tail as the 
skink escaped into a limestone hole. 

Natural history. Individuals of Eremiascincus were 
invariably found by turning over rocks and logs, in both 
moist and dry substrate, and never out in the open, either 
during the day or by night. It appears that these animals 
require shelter by day and are fairly indiscriminate how 
they find it. We have found some individuals in man- 
made rock piles and underneath large flat rocks near 
human habitations, while elsewhere (such as in the dry 
coastal forest on Jaco Island; Locality 30) we encountered 
them under rotten logs. The daytime refuges also appear 
to require a certain level of moisture. 


Timor-Leste Herpetofauna Updates 99 

Taxonomic comments. The genus Eremiascincus 
was initially formed for a group of closely related 
Australian sand-swimming skinks nested within the 
genus Sphenomorphus (Greer, 1979). It was then 
expanded to include a number of taxa from the genus 
Glaphyromorphus (Mecke et al., 2009), including the 
Lesser Sunda taxa E. antoniorum (Smith, 1927), E. 
butlerorum (Aplin et al., 1993), E. e. emigrans (van Lidth 
de Jeude, 1895), E. e. wetariensis (Mertens, 1928), and 
E. timorensis (Greer, 1990). Three Eremiascincus species 
have been documented for Timor so far (E. antoniorum, 
E. cf. emigrans, and E. timorensis), but the taxonomy 
of Eremiascincus populations in the Lesser Sunda 
Islands, let alone Timor Island, is far from resolved. All 
previous reports of E. antoniorum and E. timorensis are 
from the central mountains of West Timor (Aplin et al., 
1993; Greer, 1990; Smith, 1927), whereas reports of E. 
cf. emigrans are from the south coast at Loré, Lautém 

Figure 23 Representatives of populations of Eremiascincus. Images (A)-(B) show the timorensis morphotype, (C) depicts Eremiascincus 
*Ermera', and (D) and (E) show specimens of the emigrans morphotype. (A) Adult male E. cf. timorensis (SVL = 96 mm) from under a man- 
made rock pile at the edge of the Meleotegi River (USNM 579760, Locality 8). (B) Adult male Eremiascincus sp. ‘Montane’ (SVL = 72 mm) 
from the grounds of the Portuguese Pousada at Maubisse (USNM 579339, Locality 16). This population has undetermined species affinities 
and may represent an undescribed species. (C) Adult male E. ‘Ermera’ (SVL = 53 mm) from a dry bamboo root mass alongside the Meleotegi 
River (USNM 580522, Locality 8). (D) Adult individual of Eremiascincus sp. ‘Lautém’ (SVL = 51 mm) from underneath palm leaf litter in a 
near-coastal habitat (USNM 579194, Loré, Lautém District; see Kaiser et al., 2011). (E) Adult individual of Eremiascincus sp. ‘Jaco’ (SVL = 
39 mm) from underneath a coralline rock in dry coastal forest (USNM 579473, Locality 30). Photos by Mark O'Shea. 

100 Asian Herpetological Research 

Vol. 6 

District (Kaiser et al., 2011). 

After collecting over sixty voucher specimens from 
five districts at elevations ranging from 10—2046 m we 
believe that as many as five species of Eremiascincus are 
present in Timor-Leste. Overall morphology ranges from 
large species with stout limbs and a relatively short trunk 
(a timorensis morphotype), to small-sized species, with 
reduced limbs and an elongated body that are superficially 
similar to E. emigrans. 

Among the forms with the timorensis morphotype 
are those exhibiting an orange venter, but with the 
ventral coloration not extending onto the chin region 
(Figure 23A). These are the largest, most strongly built 
forms in Timor-Leste, and they have been collected 
on the Meleotegi River at an elevation around 1180 m 
(Ermera District; Locality 8); they are herein listed as 
Eremiascincus cf. timorensis. A second member with 
this stout morphology 1s a slightly smaller, more slender, 
yellow-bellied form, whose ventral coloration extends 
across the gular region to the snout. This form is found at 
other highland locations (e.g., Maubisse, Ainaro District; 
Locality 16), the slopes of nearby Mt. Ramelau, and 
at various locations around Same (Manufahi District; 
Locality 17); it might be conspecific with Eremiascincus 
cf. timorensis or represent an undescribed taxon, and it 
is listed here as Eremiascincus ‘Montane’ (Figure 23B). 
A third highland taxon, similar to E. antoniorum in some 
respects (Figure 23D), has a yellow venter that does not 
extend into the gular region, and displays a more slender 
and elongated body than forms with the timorensis 
morphotype. This form is known from the Meleotegi 
River and surrounds (Ermera District; Locality 8), and we 
did not find it anywhere else in Timor-Leste. We refer to 
this species as Eremiascincus ‘Ermera.’ In each case, both 
male and female specimens show the respective ventral 
coloration, but intraspecific variation or color change 
related to reproductive readiness cannot be excluded at 
this point. 

The emigrans morphotype appears to inhabit only 
lowland habitats in Timor-Leste (below 500 m elevation, 
and most frequently near the coast), which is consistent 
with the distribution of E. emigrans complex forms on 
other islands in the region. The Lautém taxon listed by 
Kaiser et al. (2011) is referred to as Eremiascincus sp. 
*Lautém' here (Figure 23C), and this population may 
inhabit the limestone habitats that make up the eastern 
end of Timor Island, at elevations from sea level up to 
462 m. The mainland Lautém form is similar to, and 
may be conspecific with, a population found on Jaco 
Island, which we call Eremiascincus *Jaco' (Figure 23E). 

Finally, the population found at coastal localities on 
northeastern Ataüro Island (Dili District) 1s referred to 
as Eremiascincus * Ataüro' (Kaiser et al., 2013b), a taxon 
certainly different from E. emigrans wetariensis from 
nearby Wetar Island. A comprehensive study of these 
forms is currently underway (Mecke et al., in prep.). 

Eutropis cf. multifasciata U" V 

Common names. (E) Common Sun Skink, Many-lined 
Sun Skink. (G) Vielstreifen-Skink. (T) Mamór loro 
(mamór = skink, loro = sun). 

Known distribution. Eutropis cf. multifasciata (Figure 
24) has so far been documented from Ermera, Lautém, 
and Oecusse Districts, on the mainland (Kaiser et al., 
2011; O'Shea et al., 2012; Sanchez et al., 2012), and also 
from Ataüro Island (Kaiser et al., 2013b). 

New localities. During Phases IV-VII additional 
vouchers of Eutropis cf. multifasciata were collected 
from the Meleotegi River and Sta. Bakhita Mission 
(Ermera District; Locality 8; USNM 579787—88), while 
first district records were obtained for Manufahi District, 
at Betano “wet site" (Locality 18; USNM 579347-49), 
and for Manatuto District, in the Nancuro coastal forest at 
Natarbora, 8 km south of Umaboco (Locality 20; USNM 
580525—26), bringing to six the districts of Timor-Leste 
where this taxon has been documented. 

Natural history. Skinks of this species were most 
frequently seen moving around in the open during daytime 
and were found in a variety of habitats, including rain and 
dry forests, grasslands, as well as coastal environments. 
We also encountered them on paths, roadways, and patios 
near human habitations. A single juvenile specimen was 
found underneath a flat rock along the Meleotegi River 
(Locality 8). 

Taxonomic comments. We refer to the population of 
Eutropis in Timor-Leste as E. cf. multifasicata because 
other than a resemblance to other Southeast Asian 
populations of the E. multifasciata species complex, there 
Is insufficient evidence to align it more closely with any 
other island or mainland population. The taxon currently 
referred to as E. multifasciata (Kuhl, 1820) has a very 
wide distribution, from the Southeast Asian mainland 
down to Timor and east to the Philippines. It is in dire 
need of taxonomic revision and once this has been carried 
out it may be possible to be more precise about the status 
of the Timorese populations. In the Lesser Sunda region, 
the population on Bali currently has subspecific status as 
E. m. balinensis (Mertens, 1927). 

Lamprolepis smaragdina cf. elberti ™=™ 

Common names. (E) Emerald or Green Tree Skink. 

No.2 Mark O'SHEA et al. 

Figure 24 Adult male Eutropis cf. multifasciata from a sun spot in 
leaf litter in wet coastal forest at Nancuro (USNM 579346, Locality 
20). Photo by Mark O'Shea. 

(G) Elbert-Smaragdskink. (T) Mamór modok (mamór = 
skink, modok = green). 

Known distribution. Lamprolepis smaragdina cf. elberti 
(Figure 25) has been documented from Baucau, Lautém, 
Oecusse, and Viqueque Districts on the mainland (Kaiser 
et al., 2011; O’Shea et al., 2012; Sanchez et al., 2012), 
and also from Ataüro Island (Kaiser et al., 2013b). 

New localities. During Phases IV-VII we added further 
records for Lautém District, from Raga (Locality 27; 
USNM 579357—58) and the Pousada de Tutuala (Locality 
28; USNM 579475—76), as well as the first district records 
for Manatuto District, from the Nancuro coastal forest at 
Natarbora (8 km south of Umaboco, Locality 20; USNM 
579350); for Manufahi District, from Betano (“wet site,” 
Locality 18; USNM 579356); for Covalima District, in 
western Suai (Locality 13; USNM 579351—52); and for 
Bobonaro District, from a heavily grazed forest at Fiuren 
(near Balibo, Locality 9; USNM 580527-—28), bringing 
the total number of districts where L. s. cf. elberti has 
been documented to nine. 

Natural history. Specimens of L. s. cf. elberti were 
primarily collected by blow-piping or hand-slapping 
from the trunks of trees. The majority of individuals 
was encountered fairly high above ground level (3-7 
m) on tree trunks with varying diameters (> 20 cm). 
This position is used as a perch for basking, as a base 
for foraging, and as an eyrie from which to observe the 
surroundings. Our earlier observation (Kaiser et al., 2011) 
of site fidelity for this skink appears to be confirmed by 
additional observations: specific individuals seem to 
remain on the same tree during a days-long period of 
incidental observations. 

Taxonomic comments. Preliminary examinations of 
the subspecies of Lamprolepis smaragdina undertaken 

Timor-Leste Herpetofauna Updates 101 

by HK and AK revealed that the form encountered on 
Timor would most likely be L. s. elberti (Sternfeld, 1918), 
a subspecies described from Wetar Island in the Inner 
Banda Arc, across the Wetar Strait from Timor. However, 
our examination of the holotype and topotypic specimens 
of that subspecies has revealed differences in color 
pattern and pholidosis, and we therefore find the use of 
L. smaragdina cf. elberti the most appropriate approach. 
It is interesting to note that coloration of this skink 1s 
quite variable and may deviate considerably from the 
emerald-green suggested by the name. While there are no 
individuals with entirely green body coloration in Timor- 
Leste, we have seen individuals possessing a bright green 
anterior half of the body that transforms in the medial 
section of the body into a “pepper-and-salt pattern” on 
a bronze-brown background (Figure 25A). This dorsal 
“pepper-and-salt pattern" still has the remnants of green 
coloration ventrally and on to the lower lateral parts of 
the body, but turns entirely bronze-brown on the tail. The 

alternative form is one devoid of any green coloration, 

Figure 25 Individuals of Lamprolepis smaragdina cf. elberti 
showing variation in dorsal coloration. This is not an example 
of sexual dimorphism, as both male and female individuals may 
possess either color pattern. Both specimens shown here were seen 
on tree limbs in their respective habitats. (A) Adult male presenting 
the two-part color pattern with a green anterior half of the body 
and a pepper-and-salt pattern on bronze background covering 
the posterior half and the tail (USNM 579213, Viqueque town, 
Viqueque District; see Kaiser et al., 2011). (B) Adult male from 
coastal wet forest at Nancuro presenting the unicolor dorsal pattern 
that includes light green portions of the venter and the bronze dorsal 
coloration with pepper-and-salt patterning along the entire body 
(USNM 579350, Locality 20). Photos by Mark O’Shea. 

102 Asian Herpetological Research 

Vol. 6 

with perhaps only a greenish sheen on the venter (Figure 

Genus Sphenomorphus '" \" 

Common names. (E) Forest or Wedge skinks. (G) 
Waldskinke. (T) Mamór ai laran (mamór = skink, ai laran 
— forest). 

Known distribution. The genus Sphenomorphus (Figure 
26) has so far been recorded from four districts (Ainaro, 
Ermera, Lautém, and Manufahi; see Kaiser et al., 2011; 
O'Shea et al., 2012). 

New localities. During Phases IV-VIII additional 
records for the genus Sphenomorphus were obtained 
for Lautém District at Raga (Locality 27; USNM 
579371—72, 579477-81), the Mainina sink-hole (Locality 
29; USNM 579482-83), and Jaco Island (Locality 30; 
USNM 579484—86); for Ermera District (Meleotegi 
River, Locality 8; USNM 579487-89, 579765—66, 

580539); and for Manufahi District (Betano “wet site,” 
Locality 18; USNM 579369—70). First district records for 
Manatuto District are supported by voucher specimens 
from Nancuro (Natarbora, 8 km S of Umaboco, Locality 
20; USNM 579359-67, 580529—33, 580534—38), and 
for Baucau District from the Japanese caves at Venilale 
(Locality 22; USNM 580540-41), increasing the number 
of districts from which Sphenomorphus skinks have been 
documented to six. The genus has yet to be documented 
for Ataüro Island. 

Natural history. Among the forms of Sphenomorphus 
found in Timor-Leste, it may be possible to declare a 
distinction between lowland and highland forms. Some 
lowland forms (including those in the wet coastal forest 
of Nancuro and Betano as well as the dry forest on Jaco 
Island) are likely closely related to or identical with S. 
melanopogon (Duméril and Bibron, 1839). We have 

encountered these fairly robust and long-limbed animals 

Figure 26 Representative individuals of several different phenotypes of forest skinks, genus Sphenomorphus. (A) Male individual of S. 
melanopogon (SVL — 69 mm) from a root buttress in coastal wet forest at Nancuro (USNM 579364, Locality 20, near sea level). (B) High- 
altitude color variation is seen in this adult S. cf. melanopogon (SVL = 64 mm) found on the trunk of a tree in coffee forest (USNM 579368, 
Locality 17 at 1200 m elevation). (C) Male individual of Sphenomorphus sp. ‘Highland large’ (SVL = 55 mm) from the wall of a limestone 
cave near Raga (USNM 579479, Locality 27 at 550 m elevation). (D) Male specimen of the Jaco Island population of Sphenomorphus (SVL 
— 75 mm, USNM 579486, Locality 30). Individuals of this population are seen quite commonly running across the leaf litter covering the 
limestone karst on their way into refugia that run deep into the rock. (E) Male individual of Sphenomorphus sp. ‘Highland small’ (SVL = 42 
mm) from the leaf litter outside the man-made caves at Venilale (USNM 580540, Locality 22). Photos by Mark O'Shea. 

No.2 Mark O'SHEA et al. 

most frequently in a head-down posture on the trunks 
of trees or root buttresses, from where they can launch 
themselves quickly and escape into the underbrush. 
We also found juvenile Sphenomorphus skinks in all 
areas where we recorded this genus, attesting to a fairly 
high reproductive rate and a high population density. 
In contrast, there are fewer individuals of the highland 
form found, for example, in the coffee forest along the 
Meleotegi River (Locality 8) or in the drier forest of 
the karst plateau of Lautém District (e.g., in Raga and 
Mainina, Localities 27 and 29, respectively). In addition, 
we have seldom encountered juveniles of this latter form 
(or these forms), and their bodies have a more vivid 
coloration in general, and on the belly in particular. Until 
a thorough taxonomic treatment is concluded, it 1s not 
feasible to provide detailed, taxon-specific data regarding 
the natural history. 

Taxonomic comments. The taxonomy of Sphenomorphus 
in Timor-Leste appears to be even more complex 
than that of Eremiascincus. We may have collected 
specimens belonging to different taxa but are unable to 
attribute them to any known species at this time. Shea 
(2012) investigated the Lesser Sunda and New Guinea 
populations of Sphenomorphus melanopogon, and selected 
as the lectotype for this species a syntype collected 
by Péron on Timor, presumably in West Timor. This 
confirms that S. melanopogon sensu stricto 1s a Lesser 
Sunda-Moluccan endemic, and New Guinean populations 
formerly considered conspecific with S. melanopogon 
are now treated as S. meyeri (Doria, 1874). Some of our 
lowland specimens from Lautém and Manatuto Districts 
may be referable to S. melanopogon (Figure 26A), but 
there remain some differences in coloration and gestalt 
(Figure 26B). Those with a similar overall morphotype 
but different coloration collected on the Meleotegi River 
and on the Lautém karst plateau appear distinct and are 
referred to as Sphenomorphus sp. ‘Highland large’ (Figure 
26C), but once again, there is merely similarity but not 
identity with forms from Jaco Island (Figure 26D). One 
other, small and slender form from the Venilale caves, 
with a very distinct pattern of stripes and blotches, 
may be referred to as Sphenomorphus sp. ‘Highland 
small’ (Fig, 26E). All other specimens, including those 
from the Meleotegi River collected during Phases I 
and III, are currently retained as incertae sedis within 

Family Varanidae—Monitor Lizards 

Varanus timorensis Gray, 1831 "Y V V! 

Common names. (E) Timor Tree Monitor, Spotted Tree 
Monitor. (G) Timor-Waran. (T) Lafaek rai-maran (lafaek 

Timor-Leste Herpetofauna Updates 103 

— crocodile or large lizard, rai — dirt, maran — dry). 
Known distribution. Varanus timorensis (Figure 27A) is 
the only varanid currently known to occur on Timor and 
it was recorded from Lautém during Phase I and the north 
coast of Manatuto District during Phase III (Kaiser et al., 
2011; O'Shea et al., 2012). Bethencourt Ferreira (1898) 
also reported specimens collected by Rafael das Dóres 
in Liquiga District, at Lahane, Fatunaba, and Maubara, 
which were subsequently lost to a fire at the Museu 
Bocage in Lisbon. 

New localities. Phases IV, VI, and VII produced 
additional records from northern Lautém and Manatuto 
Districts, and new records from Baucau District (along the 
coastal road; USNM-HI 2831-33), Dili District (Timor 
Lodge Hotel; Locality 1; USNM-HI 2834), and Covalima 
District (northwest of Maubesi, near Tilomar; Locality 15; 
USNM 579389). Since this is a CITES protected species 
we have voluntarily limited our collecting to either tissue 
samples or road-killed specimens, where these were fresh 

enough to be sampled. Live specimens were collected, 
photographed in situ and released. The exception to this 

ne TR APPS i 

Figure 27 (A) Adult Varanus timorensis (not vouchered) displaying 
the characteristic morphology and coloration seen in individuals 
encountered all along the northern low-lying coastal habitats 
in Timor-Leste. (B) An unusual specimen we refer to as V. cf. 
timorensis due to its aberrant color pattern, habitat, and behavior. 
We found this specimen ca. 5 m high on a roadside tree at an 
elevation of 520 m (USNM 579389, Locality 15). Photo by Mark 

104 Asian Herpetological Research 

Vol. 6 

was the specimen from Tilomar (Figure 27B), given that 
its morphology, color pattern, and occurrence at higher 
altitude (520 m) gave the appearance that it might be a 
specimen of V. auffenbergi Sprackland, 1999, a species 
described from neighboring Roti Island. However, 
according to Bóhme (2003) and Moldovan (2007), 
the status of V. timorensis populations on Timor and 
neighboring Roti and Kisar is still unresolved. Varanus 
timorensis, inclusive of the unusually colored Tilomar 
specimen, is now known to occur in five mainland 
districts at an elevational range from 6 m to as high as 
520 m. 

Natural history. Most of our observations of this 
species have been fleeting glimpses of lizards dashing 
across roads, or through the examination of road-killed 
specimens. Lizards appear to be particularly abundant in 
the vegetation associated with active and unplanted rice 
paddies, but we believe them to be present in essentially 
any lowland habitat. Even in residential areas, such as the 
compound of the Timor Lodge Hotel in Dili (Locality 1), 
these lizards are able to make a living, perhaps attracted 
by the presence of small vertebrates and invertebrates 
associated with human habitations. 

Taxonomic comments. Various varanids have been 
listed as present on Timor by previous authors, including 
Varanus timorensis, V. indicus, and V. salvator. Varanus 
timorensis 1s a tree monitor species present on both the 
northern and southern coasts but the species is also found 
further inland, although it has yet to be recorded at or 
above 600 m. The specimen from Covalima is the only 
specimen found an appreciable distance (approx. 12 
km) inland and since this specimen differed slightly in 
appearance from the usual V. timorensis and 1s perhaps 
conspecific with V. auffenbergi, it was tentatively listed 
as V. cf. timorensis. Varanus indicus 1s probably recorded 
from Timor in error, as it is known to be a mangrove- 
and estuarine-dwelling species from New Guinea and the 
Moluccan islands of Aru, Kei, Seram, and Buru (Bóhme, 
2003). Varanus salvator is not known from the main 
island of Timor but a population of V salvator-like lizards 
has been documented from Ataüro Island (Kaiser et al., 


Family Acrochordidae—Filesnakes 

Acrochordus granulatus (Schneider, 1799) /"! 
Common names. (E) Little filesnake. (G) Indische 
Warzenschlange, Zwerg-Warzenschlange. (T) Samea kulit 
krukut (samea = snake, kulit = skin, krukut = rough). 
Identification. Due to their excessively baggy, highly 
tuberculate skin, the three extant members of the genus 

Acrochordus are instantly recognizable, and afforded 
common names such as “wartsnake,” “filesnake,” or 
“elephant’s trunk snake” (this latter the case for the 
larger freshwater species). Acrochordus granulatus 
(Figure 28A) is the smallest member of the genus, with a 
maximum length of 1.6 m (McDowell, 1979), although 
most specimens are less than 1.0 m long. 

Known distribution. One historic locality record for 
the occurrence of A. granulatus exists for Timor-Leste 
(Table 4), documented from a single specimen collected 
by Francisco Newton, at “Dilly” (= Dili, Dili District, 
Locality 1), and reported by Bethencourt Ferreira (1898) 
as present in the Museu de Lisboa; this specimen was lost 
in the museum fire of 1978. Acrochordus granulatus is 
also known from West Timor (from Kupang and Tuakdale 
Lagoon; de Lang, 2011). 

New localities. One individual (Figure 28B) was 
collected by AVR in the mangrove swamp at Metinaro 
(Dili District; Locality 3; USNM-FS 255498; field tag 
only, specimen to remain on exhibit in Timor-Leste; 
USNM-HI 2825). 

Natural history. The unusual tuberculate skin is an 
essential aid for the identification of these snakes in 
Timor-Leste. Acrochordus are ambush predators or 
active foragers, that grasp and coil around their slippery 
fish prey, with the tubercles maintaining a strong 
and inescapable, constriction-like grip as the fish is 
maneuvered into a head-first ingestible position. It has 
also been suggested that the tuberculate skin may prevent 
the snakes from drying out if exposed to the air (Greer, 
1997), and tubercles may also serve a sensory purpose 
in prey location (McDowell, 1979; Shine and Houston, 

Filesnakes are completely aquatic, found in coastal, 
brackish and occasionally fresh water, being ill-adapted 
to movement on land due to their extremely small ventral 
scales and flabby bodies. What makes locomotion 
laborious and impossible in a terrestrial environment 
enables filesnakes to become efficient inhabitants of 
aquatic environments as the body can be flattened laterally 
as a broad ribbon for effortless swimming. Other notable 
external aquatic features include dorsally positioned 
valvular nostrils, small eyes, and a row of small, tight- 
fitting supralabial scales along the lips, perhaps to 
reduce water ingress into the oral cavity. More subtle 
physiological aquatic adaptations include a low metabolic 
rate and almost twice the blood content of terrestrial 
snakes of similar size, which, coupled with high levels 
of oxygen-carrying red blood cells, have enabled captive 
specimens of A. granulatus to remain submerged for up 

Mark O'SHEA et al. 

Figure 28 Individual of Acrochordus granulatus from Metinaro 
Swamp (USNM-FS 255438, Locality 3). (A) Photo taken by 
Agivedo Varela Ribeiro right after capture. (B) Specimen 
after preservation shows a color shift to brown, indicating the 
characteristic banding pattern of the species. Photo by Mark O'Shea. 

to 139 minutes (Whitaker and Captain, 2004). Filesnakes 
may remain motionless for prolonged periods of time, 
whether resting or in ambush, using their prehensile tails 
to maintain an anchorage against prevailing currents. 
Acrochordus granulatus 1s probably the most adaptable 
of Acrochordus species, occurring in marine, brackish, 
and freshwater habitats, and although there is one record 
of a specimen from an elevation of 90 m (McDowell, 
1979), this 1s a low elevation species. Although it is most 
often associated with mangrove swamps and turbid river 
estuaries, this species is also encountered in shallow, 
crystal-clear coral reef environments (MOS, pers. 
obs). Acrochordus granulatus is also the most widely 
distributed member of the genus, occurring from the 
western coastline of India, east to Indochina, southeast to 
northern Australia, and eastwards to the Solomon Islands. 

Timor-Leste Herpetofauna Updates 105 

It has been collected 10—15 km from shore and at a depth 
of 20 m (Stuebing and Voris, 1990; Voris and Glodek, 
1980), but it is considered an inshore rather than an open 
water species. 

Prey of A. granulatus is composed entirely of inshore 
or estuarine fish (McDowell, 1979); the presence of crabs 
or snails in gut contents is likely attributable to secondary 
ingestion (Greer, 1997). Acrochordus granulatus has 
been observed foraging actively, swimming and probing 
the substrate for hidden prey (Gorman et al., 1981). Both 
the chemosensory forked tongue and tactile sensory 
bristles on the tubercles may be utilized in prey location 
and capture (Greer, 1997), resulting in the opportunistic 
capture of fish coming into contact with a resting filesnake 
as much as the active foraging for prey. 

In contrast to its two larger, primarily nocturnal, 

relatives, A. granulatus is equally active both by day or 
night (Greer, 1997), although in our experience (MOS, 
pers. obs.) they are more frequently encountered surfacing 
for air in estuarine habitats after dark. Acrochordus is a 
viviparous genus, with female A. granulatus producing 
1—12 neonates (McKay, 2006). 
Taxonomic comments. In the historic literature, A. 
granulatus is frequently referred to as Chersydrus 
granulatus (e.g., Schneider, 1801; Merrem, 1820; 
Boulenger, 1893; de Rooij, 1917), distinct from the 
only other known species at the time, the much larger A. 
javanicus Hornstedt, 1787, which was itself split into two 
species by McDowell (1979): the freshwater-brackish 
Southeast Asian A. javanicus and the entirely freshwater 
Australo-Papuan A. arafurae McDowell, 1979. 

Despite its huge geographical range, and the antiquity 
of the family, with species divergence times of 20-16 
Mya, a recent study (Sanders et al., 2010) found no 
evidence that A. granulatus might be a composite of 
several different species. The family and genus are 
remarkably species-poor with one extinct species, A. 
dehmi Hofstetter, 1964 described from Pakistan (Head, 
2005; Hoffstetter, 1964). 

Family Colubridae— Typical Snakes 
Coelognathus subradiatus (Schegel, 1837) 
Common names. (E) Lesser Sunda Racer, Lesser 
Sunda Trinket Snake, Timor Racer. (G) Indonesische 
Kletternatter. (T) Samea laho (samea = snake, laho = rat). 
Known distribution. During Phases I and III we collected 
two specimens of Coelognathus subradiatus (Figure 29) 
in Baucau and Viqueque Districts, both on the outskirts of 


the towns bearing the districts’ names (Kaiser et al., 2011; 
O’Shea et al., 2012), and recorded a third specimen as a 
roadkill on the Atambua-Kefamenanu road in West Timor 


Asian Herpetological Research 

Vol. 6 

Table 4 Records of snake species for the districts of Timor-Leste. Black circles indicate previously known records, red circles denote new 
records. Black open circles are literature records. Records listed in grey denote literature records from West Timor, with closed circles 
representing road-killed specimens we found and open circles representing known museum specimens. Check marks denote encounters with 

positive identifications, but without voucher specimens. 

Taxon 2 8 B E E P E S £ & 3 E 9 B E References* 
i i$ B 6 P C S$ B à 3 B8 EB EIE 
a O Z4 = 2 5 Oo $|8 
Acrochordus granulatus e. O 4,6,10 
Coelognathus subradiatus subradiatus e. e. O e. e. e = 1,2,4-6,10 
Dendrelaphis inornatus timorensis [9] y ® Y e e. O 1,5,6 
Lycodon capucinus e e. e. e. e. e. e. e. e 1,2,4-6 
Lycodon subcinctus e e 1,4,6 
Stegonotus sp. e. e. 4 
Cylindrophis cf. boulengeri O e. O 4,6,8 
Laticauda colubrina e Y e. 3,4 
Cantoria violacea O 6,7,10 
Cerberus rynchops e. O O e. O 1,2,4,6 
Fordonia leucobalia e. O 4,6 
Liasis mackloti mackloti e. * e. e. 2,4 
Malayopython reticulatus e. O e. e. e. e. O 2-4,6 
Indotyphlops braminus e. e. e. e. e. O 1,4,6 
Indotyphlops incertae sedis e. e. e. 1,4 
Sundatyphlops polygrammicus e. O e. 4,6,9 
Trimeresurus insularis e. e. [9] o e. O e. e. e. e. 1,2,4,6 

‘References are identified numerically as follows: 1 = Kaiser et al., 2011; 2 = O’Shea et al., 2012; 3 = Sanchez et al., 2012; 4 = this paper; 5 
= Kaiser et al., 2013b; 6 = de Lang, 2011; 7 = de Rooij, 1917; 8 = Forcart, 1953; 9 = Barbour, 1912; 10 = Bethencourt Ferreira, 1898. 

(O’Shea et al., 2012). A fourth specimen, from Ataüro 
Island, was documented elsewhere (Kaiser et al., 2013b). 
Coelognathus subradiatus was also reported from two 
Lautém locations, the towns of Lospalos and Muapitine, 7 
km E of Lospalos (de Lang, 2011). 

New localities. During Phase VII two further specimens 
were recorded, both as roadkills, one on the road from 
Baucau to Venilale, just south of Baucau (USNM-HI 
2827), and on the road from Manatuto to Natarbora 
(USNM 580544), on the south side of the central 

mountain range. In March of 2011, HK visited Timor- 
Leste and photographed a road-killed individual on 
the road between Dili and Railaco, in Liquiga District 
(USNM-HI 2826a-c). During Phase VIII, one specimen 
was collected near the Dili port (USNM 581171). This 
specimen, together with the Liquica and Manatuto 
records, constitute first district records, bringing to seven 
the districts for which C. subradiatus has been confirmed 
(Table 4). 

Natural history. Coelognathus subradiatus 1s a 

Mark O'SHEA et al. 

Figure 29 Adult male Coelognathus subradiatus collected at 
Palapasu, Dili (USNM 581171, Locality 1). Photo by Mark O'Shea. 

crepuscular to nocturnal species that exhibits a 
considerable degree of habitat plasticity, occurring in a 
wide variety of environments across its Lesser Sunda 
range, which includes virtually every island from Lombok 
to Wetar and Timor, with the notable exception of Savu. 
Habitats range from coastal coconut plantations and 
low-lying steppe-grasslands to monsoon and montane 
rainforests, to elevations up to 1200 m (Schultz, 1996). 
Mertens (1930) also commented that this snake is often 
encountered in close proximity to human habitations, 
and this observation has been borne out by our own 
experiences on Timor (Kaiser et al., 2011; O’Shea 
et al., 2012). Although Schultz (1996) considered C. 
subradiatus to be primarily terrestrial, we obtained one 
particularly dark specimen (USNM 579779) on Ataüro 
Island (Kaiser et al., 2013b) after it had escaped into a 
tree to a height of approximately 6 m, then leapt to the 
ground when pursued aloft. 

The prey of C. subradiatus comprises primarily 
small mammals, such as rodents, which are killed by 
constriction by this relatively powerful, muscular species; 
birds may also be taken (Schultz, 1996). Auffenberg 
(1980) reported that juvenile Lesser Sunda racers on 
Komodo prey on geckos. A more catholic diet was 
reported by de Lang (2011), who listed “small mammals, 
birds, reptiles, amphibians, fish, and even insects." 
Coelognathus subradiatus is oviparous, but clutch size is 
largely unknown; Schultz (1996) discussed six hatchlings 
that emerged from a clutch of unknown size after having 
been laid in captivity by a wild-caught female. 

When this species feels threatened it may elevate the 
anterior portion of its body into a vertical S-shape, inflate 
its neck, and make lunging strikes, biting freely if contact 
is made; being completely nonvenomous, this display is 

Timor-Leste Herpetofauna Updates 107 

largely bluff. 

Taxonomic comments. Bethencourt Ferreira (1897) 
described Coluber melanurus var. timoriensis (a synonym 
of C. subradiatus) from Timor, presumably from the 
Portuguese eastern end of the island, now Timor-Leste. 
This specimen was collected by Francisco Newton, who 
failed to provide a precise locality; it was lost in the 
Museu Bocage fire of 1978. 

In the Lesser Sunda Islands there appear to be two 
different forms, which are referred to as “Groups” by 
Schultz (1996). Group 1 comprises slender-bodied snakes 
that achieve total lengths of 1200-1600 mm whereas 
Group 2 includes the more heavily-built snakes that 
achieve total lengths of 1500—2200 mm. Differences in 
patterning were noted by both Schultz (1996) and de 
Lang (2011). Racers found on Timor and the neighboring 
islands of Roti and Semau would fall within Group 1, 
whilst all other Lesser Sunda specimens would be part of 
Group 2. However, these groups have no taxonomic status 
and are purely subjective. A population of racers from 
Enggano Island (402.6 km?), almost 1600 km west of the 
westernmost population of Lesser Sunda C. subradiatus 
on Lombok and separated by the island of Java, was 
for a time treated as a subspecies of subradiatus, C. s. 
enganensis (Vinciguerra, 1892), but it has lately been 
treated as a full species (Das, 2012; Wallach et al., 2014). 

Dendrelaphis inornatus timorensis Smith, 1927 "! 
Common names. (E) Timor Bronzeback, Lesser Sunda 
Treesnake. (G) Timor-Bronzenatter. (T) Samea kotuk kór 
kafé (samea = snake, kotuk = back, kor kafé = brown). 
Known distribution. During the early phases of 
the project (2009 and 2010) Dendrelaphis inornatus 
timorensis (Figure 30) was documented for Lautém 
and Viqueque Districts (Kaiser et al., 2011; O’Shea et 
al., 2012), whereas de Lang (2011) included Dili (Dili 
District) in its distribution. It is a species commonly 
documented in West Timor (Table 4). 

New localities. The only voucher specimen of this 
species obtained during the later phases of the project 
was from the Betano “wet site” (Manufahi District; 
Locality 18; USNM 579378) on the southern coast. There 
were unconfirmed sightings of “treesnakes” tentatively 
identified as D. i. timorensis in the Nancuro coastal forest 
(Manatuto District; Locality 20) and on Ataüro Island 
(Dili District) during Phases IV and VI respectively, but 
no vouchers were obtained. 

Natural history. This is a fast-moving and highly 
elusive diurnal species that often evades capture, either 
through speed or its ability to blend in with the vegetation 
when motionless. It is the only member of the genus 

108 Asian Herpetological Research 

Vol. 6 

Figure 30 Individual of Dendrelaphis inornatus timorensis (sex not 
determined) collected from low shrubs by night (USNM 573686, 
Wailakurini, Viqueque District; see Kaiser et al., 2011). Photo by 
Mark O'Shea. 

Dendrelaphis to occur on Timor and one of only two 
found in the Lesser Sunda Islands. Elsewhere in the 
Indo-Malayan region this is a commonly encountered 
and fairly well represented genus with numerous species 
in Southeast Asia and the Philippines, where they are 
known as “bronzebacks,” and nine species occurring in 
New Guinea, the Kei Islands, Palau, the Solomon Islands, 
and Australia (van Rooijen et al., 2015), where they are 
known as “treesnakes.” 

Timorese D. i. timorensis inhabit wooded hill country 
with an understory of grass and often a geology of 
black limestone outcrops (de Lang, 2011), which 1s 
precisely the habitat in which one of us (AVR) observed 
a specimen that evaded capture on Ataüro Island (Kaiser 
et al., 2013b). Smith (1927) stated that D. i. timorensis 
occured at elevations from 100-800 m, but we have 
found this species to be most abundant right at sea level 
in Loré, Lautém District (USNM 573687-88), where two 
additional specimens evaded capture (Kaiser et al., 2011); 
another evaded capture at sea level in the coastal forest at 
Nancuro (Manatuto District; Locality 20). 

Virtually no information exists regarding the natural 
history and biology of the Timorese subspecies. The 
Komodo population of the nominate form (Auffenberg, 
1980) preys on geckos (Hemidactylus) and skinks 
(Sphenomorphus), whilst frogs (Fejervarya) are known 
from the diet of Sumbawa and Flores specimens 
(Mertens, 1930). All these potential prey genera occur on 
Timor. Dendrelaphis inornatus is an oviparous species 
with clutch sizes reported from 2-18 (de Lang, 2011), but 
no data exist specifically for D. i. timorensis. 

Taxonomic comments. The subspecies D. i. timorensis 
occurs on Timor and the neighboring eastern Nusa 
Tenggara and southern Maluku islands of Roti, Semau, 

Pantar, Alor, and Wetar. The nominate subspecies is found 
on the western islands of Nusa Tenggara, west of and 
including Lomblen and Savu. 

Lycodon capucinus (Boie, 1827) "YY"! 

Common names. (E) Common (island) Wolfsnake. (G) 
Kapuzen-Wolfszahnnatter. (T) Samea lobo (samea = 
snake, lobo = wolf). 

Known distribution. During Phase I Lycodon capucinus 
(Figure 31) was recorded from Same (Manufahi District; 
Locality 17; Kaiser et al., 2011), and as a roadkill on the 
Sakato-Atambua road in West Timor in Phase III (O'Shea 
et al., 2012). During Phase VI it was also recorded as 
common on Ataüro Island (Kaiser et al., 2013b). 

New localities. During Phases IV-VIII this species was 
encountered with increasing frequency, primarily as 
roadkills. We collected live specimens in Dili District 
(grounds of the Timor Lodge Hotel, Locality 1; USNM 
579781); Lautém District (Com and Raga, Localities 
26 and 27; USNM 579381-82, 579494—95); Manufahi 
District (Ladiki coffee forest near Same, Locality 17; 
USNM 579380); and Bobonaro District (degraded forest 
at Fiuren, near Balibo, Locality 9; USNM 580547). It 
was also reported to occur at Malahara (Lautém District; 
Locality 29; de Lang, 2011). Roadkills were documented, 
and sampled when possible, from Covalima District (north 
of Suai, Locality 13; USNM 579379); Baucau District 
(near Baucau; USNM 580546); Aileu District (near Lahae 
town); and Bobonaro District (on the Maliana-Balibo 
road; USNM 580548). Lycodon capucinus has now been 
recorded from seven mainland districts, and Ataüro Island 
(Table 4). 

Natural history. Lycodon capucinus is a very common 
and widespread, but nocturnal and secretive snake that is 
easily overlooked in cursory searches, although it may be 
encountered abroad at night, especially after heavy rain. 

Figure 31 Adult male Lycodon capucinus from the leaf litter at the 
ruins of the Portuguese pousada at Com (USNM 579494, Locality 
26). Photo by Mark O'Shea. 

No.2 Mark O'SHEA et al. 

We have found it in almost every habitat investigated, 
from townships to coffee forest, and from the ruins of a 
coastal pousada close to sea level, to elevations of over 
1150 m (Ainaro District; Locality 16), greatly exceeding 
the 600 m documented for Komodo Island specimens 
(Auffenberg, 1980; Darevsky, 1964; Dunn, 1927). It 
appears to have adapted well to living alongside humans 
and is even found in major cities, such as the Indonesian 
capital at Jakarta (van Hoesel, 1959). This was easily the 
most frequently encountered snake species during our 
surveys to date, with 21 specimens documented, from 
juveniles to adults; two of these records were based on 
sloughed skins, which could be unequivocally identified 
to belong to individuals of this species based on scale 
counts and head scale morphology. 

Lycodon capucinus is a small species that rarely 
achieves a length in excess of 600 mm, although our 
highest elevation specimen (see above), a roadkill that 
was sampled for tissue only, had an SVL of 580 mm and a 
TTL of 720 mm. Although primarily a terrestrial species, 
L. capucinus is agile and may be encountered climbing in 
vegetation or on buildings. Lycodon capucinus will bite 
readily when handled. 

Prey of L. capucinus comprises primarily geckos, 

especially perianthropic species of the genera 
Hemidactylus and Gehyra, but across its extensive range 
L. capucinus is reported to have taken the skink Eutropis 
multifasciatus (Kopstein, 1936) and even mice (Mertens, 
1930). According to McKay (2006) it also eats frogs and 
reptile eggs. It is an oviparous species, and clutches of 
up to eleven eggs have been reported (David and Vogel, 
1996). On Timor, it has been reported as being parasitized 
by tapeworms (Goldberg et al., 2010). 
Taxonomic comments. Lycodon capucinus was long 
treated as either a synonym or a subspecies of the 
widespread South and Southeast Asian L. aulicus 
(Linnaeus, 1758), to which it bears a striking resemblance, 
and only relatively recently has it been consistently 
treated as a distinct and separate species based on the 
work of Taylor (1965) and David and Vogel (1996). This 
nomenclatural history has caused considerable confusion 
when the geographical range of this species needed to be 
determined (Kaiser et al., 2011). 

Lycodon subcinctus Boie, 1827 ""! 

Common names. (E) Malayan banded Wolfsnake. (G) 
(Wei-) Gebánderte Wolfsnatter. (T) Samea kadali (samea 
= snake, kadali = ring). 

Known distribution. In the early phases of the project a 
single specimen of Lycodon subcinctus (Figure 32) was 
obtained, from Mirbuti village near Same (Manufahi 

Timor-Leste Herpetofauna Updates 109 

District; Locality 17), and close to the project's first 
collection locality for L. capucinus (Kaiser et al., 2011). 
New localities. A second specimen was obtained in Raça 
village (Lautém District, Locality 27; USNM 579382). 
Natural history. Much less frequently encountered by us 
on Timor than its congener, L. capucinus, L. subcinctus is 
a secretive, nocturnal inhabitant of humid forests and dry 
woodlands, both in low-lying and montane locations up 
to elevations of 1660 m in Peninsular Malaysia (Smith, 
1930) and 1800 m in Bali (McKay, 2006). It 1s also 
reported to occur in plantations, rice paddies and other 
agricultural habitats, and around human habitations (de 
Lang, 2011). The latter location agrees with the first of 
our two specimens, which we obtained when we were 
handed a badly damaged specimen that had been killed in 
a schoolyard near Same, Manufahi District (Locality 17; 
see Kaiser et al., 2011). 

Lycodon subcinctus is a larger species than L. 
capucinus, achieving total lengths of 800-1200 mm (de 
Lang, 2011). The larger size and semi-fossorial nature 
of this infrequently (on Timor) encountered species may 
be the basis for the “Timor krait" stories circulated by 
individuals who observed this species but who were 
perhaps familiar with banded kraits from other parts 
of Indonesia (including Bali in the Lesser Sundas). 
Indeed, its pattern of white bands on a black background, 
combined with the lack of a loreal scale, afford L. 
subcinctus a startling similarity to the highly venomous 
species Bungarus candidus (Linnaeus, 1758) and B. 
fasciatus (Schneider, 1801), with which L. subcinctus 
occurs in sympatry in other parts of its range. 

Although species in the genus Lycodon are primarily 

terrestrial, L. subcinctus is also arboreal (McKay, 2006), 
with prey consisting of geckos and skinks (de Lang, 
2011). Females are oviparous, laying from 5-11 eggs (de 
Lang, 2011). 
Taxonomic comments. Three subspecies of the widely 
distributed L. subcinctus have been described. The 
nominate race is found through most of Southeast Asia 
and it is to this taxon that Lesser Sunda populations 

Stegonotus sp." Y"! 

Common names. (E) Timor Groundsnake. (G) Timor- 
Schiefernatter. (T) Samea rai kór-kafé (samea = snake, rai 
— ground, kór-kafé — brown). 

Known distribution. There were no previous records for 
the genus Stegonotus (Figure 33) from Timor, the nearest 
known populations being those of S. florensis on Flores 
and Sumba (Daan and Hillenius, 1966; de Rooij, 1917; 
Forcart, 1954). 

110 Asian Herpetological Research 

Vol. 6 

New localities. The first specimen of the genus 
Stegonotus from Timor was obtained during Phase IV, 
from the coastal forest at Nancuro, near Natarbora, 8 
km south of Umaboco (Manatuto District; Locality 20; 
USNM 579383). A second specimen was collected by one 
of us (LLA) during a personal survey, part of a research 
project from Timor-Leste's national university, at Betano 
“wet site" (Manufahi District; Locality 18; USNM 
579384). Two further specimens, one adult and one 
juvenile, were collected, in close proximity to each other 
and close to the original collection point in the Nancuro 
coastal forest, during Phase VII (USNM 580549—50). 
Stegonotus sp. is now known to occur in southern low- 
lying coastal forests in two districts (Table 4). 

Natural history. Individuals of this species were found 
exclusively in moist coastal forests within a short 
distance of the southern coast of Timor-Leste (^ 2 km). 
At Nancuro, one adult specimen was spotted moving 
through the leaf litter, while another was found in the 
hollow portion of a decaying log. The juvenile was found 
unexpectedly, in a vertical position, under loose bark of 
a standing tree. Whereas the adult in the log attempted 
to escape by retreating further into the rotting wood, the 
juvenile remained motionless when the bark was removed 
and was easily captured. 

Stegonotus is a common and well-represented genus 
in Papua New Guinea (PNG) and one well familiar to 
MOS, who identified it immediately upon capture of 
the first (Nancuro) specimen. In PNG members of this 
non-venomous genus are rarely encountered abroad 
during the day, most being found on roads or on the 
ground in the bush during the evenings and at night, 
or discovered hiding under logs or other debris during 
daylight hours (MOS, pers. obs.). Small specimens have 
also been encountered inside ant plants (Myrmecophyta), 
presumably hunting the skinks that also inhabit ant plant 
chambers. Such microhabitats should be investigated, 
should these tropical Southeast Asian-Melanesian trees 
occur in Timor-Leste. Papuan Stegonotus, particularly the 
large S. cucullatus (Duméril et al., 1854), will bite with 
vigor and little provocation (O' Shea, 1996), and some 
Timorese specimens exhibit similar behavior. Members of 
the genus Stegonotus are oviparous. 

Taxonomic comments. The genus Stegonotus currently 
comprises ten species (Uetz and Ho&ek, 2014), 
distributed throughout New Guinea (four species, at 
least one also occurring in northern Australia), the 
Bismarck Archipelago (one species), the d'Entrecasteaux 
Archipelago (one species), the Maluku Islands (one 
species), Borneo (one species), the Philippines (one 

Figure 32 Adult male Lycodon subcinctus from the leaf litter at the 
limestone caves near Raga (USNM 579382, Locality 27). Photo by 
Mark O'Shea. 



L3 T A ta ape 
Ea 7t ul 

Figure 33 Adult female Stegonotus sp., collected from the inside 
of a rotting log in coastal wet forest at Nancuro (USNM 579383, 
Locality 20). Photo by Mark O'Shea. 

species), and the Lesser Sundas (one species reported 
from Flores and Sumba). This latter taxon, S. florensis (de 
Rooij, 1917), is the only member of the genus occurring 
close to Timor. 

Comparison of Timor specimens with the type material 
of S. florensis and a variety of museum specimens 
representing the other known species of Stegonotus, has 
allowed us to determine that the Timor specimens belong 
to an undescribed species based on scale counts and head 
scale morphology. We have also been able to recognize 
that the S. florensis material represents more than one 
species, with those from Sumba most likely warranting 
the resurrection of S. sutteri from synonymy (see Forcart, 
1954). Beyond these comparisons, we have uncovered 
many inconsistencies in how names have been applied to 
Stegonotus populations throughout the range of the genus, 
and this topic is currently the subject of a comprehensive 
investigation (Christine Kaiser, unpubl. data.). 

No.2 Mark O'SHEA et al. 

Family Cylindrophiidae—Asian Pipesnakes 
Cylindrophis cf. boulengeri “™ 
Common names. (E) Boulenger's Pipesnake, Timor 
Pipesnake. (G) Boulenger-Walzenschlange. (T) Samea 
ulun rua (samea = snake, ulun = head, rua = two). This 
snake is locally known as the “two-headed snake,” given 
that the body morphology and defensive behavior of 
pipesnakes do not allow for a ready identification of the 
head and make it appear as if both ends of the snake 
might pose a threat. 
Known distribution. Cylindrophis cf. boulengeri 
(Figure 34) is recorded from Timor-Leste based on eight 
specimens collected by Prof. A. Bühler in 1935 at Baguia 
(Baucau District, no further data) and now deposited 
in the Naturhistorisches Museum Basel, Switzerland 
(NHMB 12908-15). Our visit to Baguia during Phase VII 
(2012) failed to produce any specimens or any recognition 
from the local population and villagers; when questioned 
and shown photographs, locals were unfamiliar with the 
snake (O'Shea and Kaiser, 2013). 
New localities. In late 2012 one of us (AVR) obtained 
a specimen of C. cf. boulengeri in Lospalos, Lautém 
District (USNM-FS 255499; field tag only, specimen 
remaining on exhibit in Timor-Leste; photo vouchers 
USNM-HI 2835a-c), the first specimen of the taxon 
discovered in 77 years. We subsequently captured another 
specimens in a banana plantation at the confluence of 
the Comoro and Bemos Rivers on the Aileu District side 
(Locality 6; USNM 581170). This secretive snake 1s now 
known from three districts in Timor-Leste (Table 4). 
Natural history. Pipesnakes of the genus Cylindrophis 
are nocturnal, semi-fossorial, and secretive. This lifestyle 
is the reason for our poor knowledge about the biology 
of the species currently recognized within the genus. One 
of the specimens we collected, at the confluence of the 
Comoro and Bemos Rivers (Aileu District: Locality 6) 
was found on the ground under a banana leaf. We had 
previously considered this type of habitat unproductive, 
with only a few striped treefrogs (Polypedates cf. 
leucomystax) being collected, and therefore had ignored 
such habitats during surveys. This 1s an excellent example 
for how collector's bias can influence collecting results. 
Almost nothing is known of the natural history of 
Timorese Cylindrophis, although it may be presumed 
that they prey on blindsnakes (/ndotyphlops and 
Sundatyphlops), and possibly cylindrical skinks 
(Eremiascincus) or invertebrates such as earthworms. 
All species for which reproductive biology is known 
are described as being ovoviviparous (Greene, 1997), 
a condition we consider to be a form of livebearing 

Timor-Leste Herpetofauna Updates 111 

(Blackburn, 1994). However, examination of museum 
specimens by one of us (SM) revealed that some 
populations of Cylindrophis, which likely represent 
distinct species based on morphology, may be egglaying. 
Cylindrophis exhibit an unusual defensive behavior, 
during which they hide the head in the coils of their body 
and elevate their tails, flashing the bright or contrasting 
ventral pattern in the process, a behavior reminiscent of 
Asian coral snakes (e.g., Calliophis intestinalis [Laurenti, 
Taxonomic comments. The populations historically 
associated with C. boulengeri Roux, 1911 are known 
from 12 specimens collected in the early 20* Century, 
eight from Baguia, Baucau District (Forcart, 1953; see 
above), one from an unspecified location in West Timor 
(de Lang, 2011), and three from Wetar (Brongersma, 
1933b; Roux, 1911), an island in the Indonesian province 
of Maluku to the northeast of Timor and the type locality 
of the species. We have been unable to locate additional 
museum specimens that may belong to this species. 
A specimen collected on Babar Island to the east and 
originally referred to C. boulengeri (Brongersma, 1933a) 
may represent a distinct and undescribed species of 
Cylindrophis. Two other species occur in the vicinity of 
Timor: C. opisthorhodus Boulenger, 1879 on Sumbawa, 
Flores, and Lombok to the west, and C. yamdena Smith 
and Sidik, 1998 on Yamdena Island in the Tanimbar 
Island group, to the east (Smith and Sidik, 1998). Until 
we have completed a study now underway (Kieckbusch et 
al., in prep.), we conservatively consider Timor material 
as C. cf. boulengeri. 

Family Elapidae—Cobras and their allies 

Laticauda colubrina (Schneider, 1799) "™ 

Common names. (E) Yellow-lipped sea krait, Colubrine 
sea krait. (G) Nattern-Plattschwanz, Gelblippen- 
Seeschlange. (T) Samea-tasi kor kadeli (samea-tasi = sea 
snake, kor = color, kadeli = ring). 

Known distribution. The sea krait Laticauda colubrina 
(Figure 35) was recorded from only one location and one 
specimen during the survey (Table 4), the old military 
wharf at Pante Macassar, Oecusse District (Sanchez et al., 
2012). One of us (SM) observed an individual in the 
water near the shore on Atatro Island (Dili District), but 
was unable to capture it. 

New localities. Our second specimen was obtained by 
AVR on the rocky headland at Cristo Rei, near Dili (Dili 
District; Locality 1; USNM-HI 2837) and subsequently 

Natural history. Laticauda colubrina is an amphibious 
snake, equally at home on land as in the ocean. Being 

112 Asian Herpetological Research 

Vol. 6 

oviparous, unlike true seasnakes, it is essential that L. 
colubrina be able to move onto land in order to lay its 
clutch of 6—20 eggs (Greene, 1997). This species is so 
capable on land that it may be encountered at the top 
of cliffs, aloft in low bushes, or in the center of small 
islands (O'Shea, 2005). At first glance, a sea krait even 
resembles a terrestrial elapid with its regular, imbricate, 
smooth scales arranged in transverse rows, and its large 
ventral plates for locomotion on land, but it 1s also highly 
adapted for life in the ocean with the laterally flattened, 
paddle-shaped tail typical of marine snakes, laterally 
positioned valvular nostrils (Wilson, 2005), and tight- 
fitting supralabial scales around the mouth. 

Prey of L. colubrina comprises entirely fish, including 

those species that seek protection by mimicking 
Laticauda, such as the colubrine snake eel (Myrichthys 
colubrinus), which is taken frequently (O'Shea, 1996; 
Wilson, 2005). Although a front-fanged venomous elapid, 
L. colubrina is placid and does not attempt to bite even 
when handled. 
Taxonomic comments. Two species of the genus 
Laticauda are reported from the seas around Timor, but 
only L. colubrina has been positively recorded. The 
other species, L. laticaudata (Linnaeus, 1758), could be 
mistaken for L. colubrina by a person unfamiliar with the 
characteristics that define the two species, and it is also 
possible this species does not occur this far west. 

Family Homalopsidae—Oriental and Australasian 

Cerberus schneiderii (Schlegel, 1837) "YY"! 
Common names. (E) Schneider's dog-faced watersnake, 
Schneider's bockadam, (G) Hundskopf-Wassertrugnatter, 
(T) Samea natar (samea = snake, natar = rice paddy). 
Known distribution. During the early phases of the 
survey Cerberus schneiderii (Figure 36) was found to be 
relatively common in the low-lying paddy field east of 
Baucau town (Baucau District). This species was reported 
by de Lang (2011), from Bidau, Dili (Dili District), and 
Lake Be Malae, Batugade (Bobonaro District), also on the 
north coast. We were also informed of a large specimen 
reportedly killed in the grounds of the Chinese Embassy 
in Dili, which is located on the seafront, but were unable 
to confirm this report. 

New localities. During the phases covered by this report 
we collected an extra voucher specimen, as a relatively 
fresh roadkill, from close to the original Baucau paddy- 
field location, and one of us (LLA) obtained a specimen 
from the Betano “wet site" (Manufahi District; Locality 
18; USNM 579392), the first southern coastal record of 
the species from Timor-Leste, although de Lang (2011) 

listed records from the south coast of West Timor. In 
addition we collected four specimens in the mangrove 
swamp at Metinaro (Dili District; Locality 3) where 
they were found to occur in sympatry with Fordonia 

leucobalia (see below). Including the records of de Lang 

(2011), this species is now reported from four districts of 

from a banana plantation near the confluence of the Comoro and 
Bemos Rivers (USNM 581170, Locality 6). Photo by Mark O'Shea. 

Figure 35 Adult sea krait (Laticauda colubrina) from a ruined 
wharf (USNM 579241, near Pante Makassar, Oecusse District; 
Sanchez et al., 2012). Photo by Mark O'Shea. 

Figure 36 Adult Cerberus schneiderii from the mangrove swamp 
at Metinaro (USNM 581173, Locality 3). Photo by Mark O'Shea. 

No.2 Mark O'SHEA et al. 

Timor-Leste, and confirmed with voucher specimens from 
three (Table 4). 

Natural history. Populations of snakes in the genus 
Cerberus are usually associated with inshore marine 
or brackish habitats, such as mangrove swamps and 
estuarine mud-flats, but all species are able to survive 
in freshwater and may be found in freshwater creeks or 
rivers flowing into these brackish environments (Murphy, 
2007; Murphy et al., 2012); the Philippine C. microlepis 
Boulenger, 1896 is the only land-locked freshwater lake 
dweller (Murphy, 2007). Cerberus schneiderii is also able 
to move from saltwater to freshwater habitats, but while 
we have collected it in brackish mangrove swamps on 
the north coast at Metinaro (Dili District; Locality 3) and 
on the south coast at Betano (Manufahi District; Locality 
18) we have found it in larger numbers in freshwater rice- 
paddy habitats, on the north coast at Baucau (Kaiser et 
al., 2011; O’Shea et al., 2012). Several specimens were 
found at Metinaro (USNM 580525-26, 581173—76), of 
which one was found during the late afternoon sheltering 
in a mud lobster (Thalassina anomala) burrow, while 
others were found in shallow muddy rivulets. 

It has been suggested that Cerberus feed almost entirely 
on small fish (McKay, 2006; Murphy, 2007), including 
lizardfish (Synodontidae: Synodus) and gobies (Gobiidae: 
Amblygobius), although other authors (e.g., Auffenberg, 
1980; Voris and Murphy, 2002) reported crustacean 
remains from the guts of some specimens. Whether these 
were the intended prey or secondarily ingested prey-of- 
prey is impossible to determine. Cerberus is a rear-fanged 
venomous genus possessing Duvernoy's glands, which 
contain toxic secretions to dispatch struggling prey. Since 
Fordonia is carcinophagous this would enable the two 
species to partition resources and survive in sympatry. 
However, we suspect that the C. schneiderii living in 
the rice-paddy habitat at Baucau may also be feeding on 
tadpoles and juveniles of the abundant population of rice- 
paddy frogs (Fejervarya). 

No reproductive data currently exist for C. schneiderii, 
but the genus is known to be livebearing, as are most 
obligatorily aquatic snakes, and litter sizes for Australian 
C. australis have been quoted as 6-8 (Shine, 1991a) or 
even as high as 26 (Gow, 1989), while de Lang (2011) 
provides a maximum litter size for Cerberus of 47 

Taxonomic comments. The taxonomy of homalopsid 
snakes formerly known as Cerberus rynchops (Schneider, 
1837) was recently revised (Murphy et al., 2012). The 
taxon had previously been divided into an Australo- 
Papuan population, recognized as C. australis (Gray, 

Timor-Leste Herpetofauna Updates 113 

1842), and a localized Philippine population, now 
known as C. microlepis (Murphy, 2007). The taxon C. 
rynchops was then used for all other populations until 
the latest revision restricted C. rynchops to populations 
on mainland Asia and the Andaman Islands. A new name 
was proposed for Palau populations (C. dunsoni Murphy 
et al., 2012) and the name C. schneiderii was resurrected 
for all other island and Southeast Asian populations. 

In most respects, Timor specimens fall within 
the characters given by Murphy et al. (2012) for C. 
schneiderii, although there are some differences, notably 
in the dorsal head scalation. Since few specimens have 
been collected from this southeastern corner of the C. 
schneiderii range, the precise taxonomic status of the 
intervening Wallacean populations may require additional 
research, especially as C. schneiderii is believed to be a 
species complex (John Murphy, pers. comm.). 

Fordonia leucobalia (Schlegel, 1837) "™Y™ 

Common names. (E) White-bellied mangrove snake, 
Crab-eating mangrove snake. (G) Krebs-Wassertrugnatter. 
(T) Samea parapa kabun-mutin (samea = snake, parapa = 
mangrove, kabun-mutin = white belly). 

Known distribution. Fordonia leucobalia (Figure 37) 
was not previously recorded for Timor-Leste and only a 
single record exists for its presence in West Timor (Peters, 
1876), where it was collected in the mangrove swamp at 
Atapupu, located on the northern coast between Timor- 
Leste and the Oecusse exclave (Table 4). 

New localities. During Phase VI a single specimen was 
collected from the mangrove swamp at Metinaro (Dili 
District; Locality 3; USNM 579780), the first specimen of 
the taxon from Timor-Leste, only the second from Timor, 
and the first from Timor in 135 years. We were able to 
obtain another specimen in the same locality during Phase 
VIII (USNM 581177). 

Natural history. Fordonia leucobalia 1s an inhabitant 
of mangrove and estuarine mud flats but it may be found 
considerable distances upstream in tidal rivers and up to 
850 km upstream in freshwater watercourses. Nocturnal 
in habit, it shelters by day in the burrows of fiddler crabs 
(Uca spp.) or mud lobsters (Thalassina anomala), only 
venturing onto the surface of the mud at night when the 
tide is returning. Fordonia is a carcinophagous species, 
preying primarily on crabs. Crustaceans recorded in the 
diet of Fordonia (Gow, 1989; Murphy, 2007; Shine, 
1991b; Voris and Murphy, 2002) include the crabs Uca sp. 
(Ocypodidae), Macrophthalmus sp. (Macrophthalmidae), 
Dotillopsis brevitarsis (Dotillidae), Sarmatium germaini, 
and an unidentified crab genus (Sesarmidae), and the 
mud lobster Thalassina anomala (Thalassinidae). Prey 

114 Asian Herpetological Research 

Vol. 6 

is grasped and dismembered with the legs broken off 
before the body is swallowed, although Voris and Murphy 
(2002) suggest that struggling crabs may autotomize their 
own legs. Fordonia is technically a rear-fanged venomous 
snake possessing Duvernoy's glands and enlarged, 
grooved rear-teeth, and its fangs are long enough to 
penetrate the carapace of decapods, and the venom is 
toxic to crabs (Kopstein, 1931; Savitzky, 1983; van 
Hoesel, 1959). Fordonia is a livebearing species, females 
producing litters of 2-17 neonates (Murphy, 2007). 
Taxonomic comments. Currently Fordonia is a 
monotypic genus occurring from mainland Asia 
(Bangladesh) to the northern Philippines (Luzon) and 
south into the Australo-Papuan realm (Murphy, 2007). 

Family Pythonidae—P ythons 
Liasis mackloti Duméril and Bibron, 1844 ""! 
Common names. (E) Macklot's water Python, White- 
lipped Python. (G) Timor-Wasserpython, (T) Fohorai-atan 
(fohorai = python, atan = slave). We have been unable to 
learn the origin of the peculiar Tetun common name. 
Known distribution. During Phase III Liasis mackloti 
(Figure 38) was documented from a single live specimen 
in Dili (Dili District) and roadkills in Baucau and northern 
Manatuto Districts (O’Shea et al., 2012). De Lang (2011) 
also reported a specimen from Dili. This species was 
also encountered as roadkills in West Timor, enroute and 
returning from the Oecusse exclave (O'Shea et al., 2012). 
Given our records and those shown on the distribution 
map in de Lang (2011), we consider the distribution of 
this species to be near-coastal and ranging throughout 
Timor in low-lying wetland habitats. 
New localities. During Phase IV Liasis mackloti 
was encountered with some frequency, unfortunately 
only as roadkills, on the roads east and west of Suai 
(Covalima District; Locality 13; USNM 579390; USNM- 
HI 2782-83; two of these specimens, Christine M. 
Dwyer field numbers 802-03, deposited in the USNM 
Biorepository, are tissue vouchers only). Five roadkills 
were documented, photographed, and locality data were 
recorded; specimens that were not in a too advanced 
state of decomposition were sampled for tissue. Liasis 
m. mackloti has now been confirmed from four mainland 
districts (Table 4). 
Natural history. Liasis mackloti is a water python that 
inhabits inundated lowland habitats such as rice paddies 
or overgrown coastal creeks. Snakes are most frequently 
encountered in the wet season, when many become road- 
killed casualties when traveling across their fragmented 

The nominate subspecies L. m. mackloti, found on 

Figure 37 Adult Fordonia leucobalia from a mud lobster burrow 
in the mangrove swamp at Metinaro (USNM 579780, Locality 3). 
Photo by Mark O'Shea. 

Figure 38 Unvouchered adult Liasis mackloti collected from 
beneath a container adjacent to the seafront in Dili (Locality 1). 
Photo by Mark O'Shea. 

Timor, is a relatively large snake that can achieve a 
maximum length of approximately 1.6 m, intermediate 
between the smaller L. m. savuensis and the larger but 
otherwise rather similar L. m. dunni Stull, 1932. It is a 
relatively powerful constrictor capable of subduing small- 
to medium-sized mammals and water birds. Various 
authors (e.g., de Lang, 2011) have included bird eggs, 
reptiles, and even frogs and fish in the diet of L. mackloti 
but there have been no studies of this species' diet in 
nature. The possibility that it may represent a climax 
predator in shallow freshwater habitats is supported by 
the feeding ecology of its close relative, L. fuscus, which 
is documented to take small crocodiles in southern New 
Guinea and northern Australia (Parker, 1982; Wilson and 
Swan, 2003). All pythons are oviparous, and females of L. 
mackloti have been reported to produce clutches of 8-14 
eggs in captivity (Ross and Marzec, 1990). 

No.2 Mark O'SHEA et al. 

Taxonomic comments. Liasis mackloti has three 
described subspecies, with only the nominate form, L. m. 
mackloti, occurring on Timor. The other subspecies are 
L. m. dunni from Wetar, northeast of Timor, and L. m. 
savuensis, from Savu, southwest of Timor. Liasis mackloti 
is very closely related to L. fuscus of northern Australia 
and southern New Guinea (Rawlings et al., 2004), 
so much so that southern Papuan water pythons were 
originally treated as L. mackloti (Parker, 1982). Water 
pythons in the Northern Territory, Australia, were found 
to be more closely related to Indonesian water pythons 
than eastern Australian/New Guinea L. fuscus (Rawlings 
et al., 2004) but the authors of this finding did not commit 
to referring to this population as L. mackloti. 

) [IV, VII] 

Malayopython reticulatus (Schneider, 1801 
Common names. (E) Reticulated Python. (G) 
Netzpython. (T) Fohorai-boot (fohorai = python, boot = 

Known distribution. During Phase III we encountered a 
number of captive adult reticulated pythons around Dili 
(Dili District) and juveniles in Viqueque District (O’Shea 
et al., 2012) (Figure 39). A captive specimen in Oecusse 
District was reported elsewhere (Sanchez et al., 2012). 
De Lang (2011) also reported specimens from Laleia 
(northern Manatuto District) and Malahara (Lautém 

New localities. During Phase IV a locally caught 
specimen was photographed and its tissue sampled at 
the Convent of St. Antony d'Lisboa (Manufahi District; 
Locality 19), and during Phase VII a dead specimen was 
encountered on the road between Natarbora and Fatucahi, 
approximately 6 km NE of the convent. This specimen 
was not a roadkill, there was evidence it had been killed 
elsewhere and dragged onto the road. The condition of the 
cadaver made tissue sampling impossible but a voucher 
photograph was taken (USNM-HI 2788). Malayopython 
reticulatus has now been confirmed for six mainland 
districts (Table 4). 

Natural history. Malayopython reticulatus is the longest 
snake species in the world, the largest potentially reliable 
account being that of a 9.98 m specimen killed and 
measured with a surveyor's tape in Sulawesi in 1912 
(Murphy and Henderson, 1997). Other large specimens 
have been reported from the Philippines, Malay 
Peninsula, Borneo, and Sumatra, but individuals on small 
islands are often considerably smaller, perhaps due to 
island miniaturization or due to the hunting pressure on 
larger individuals (McKay, 2006). The largest Timorese 
specimen encountered during the survey was a captive 
from Becora (Dili District; Locality 2), with a total length 

Timor-Leste Herpetofauna Updates 115 

just over 3.5 m. 

Malayopython reticulatus is an inhabitant of rainforests 
and monsoon forests, particularly in close proximity 
to watercourses, where young specimens sleep on 
overhanging branches and plunge into the water below 
if they detect the approach of a potential threat (O’Shea 
et al., 2004). This vegetated habitat also affords pythons 
the cover required to function as ambush predators of 
vertebrates, such as mammals. The species is also often 
found in bat caves, with these mammals providing 
a constant food source (McKay, 2006). However, 
reticulated pythons may also be found in cultivated or 
agricultural habitats, such as plantations (O’Shea, 2007), 
and individuals have been known to enter towns and even 
large cities (Cox, 1991). Several Timorese specimens 
were reportedly captured by locals on the outskirts of the 
capital, Dili. The species may occur at elevations from sea 
level to at least 1500 m (Malkmus et al., 2002; Manthey 
and Grossmann, 1997). 

The prey of M. reticulatus is composed primarily of 
mammals, with birds and large lizards occasional prey 
items (Malkmus et al., 2002). The size range of mammals 
consumed by reticulated pythons is astounding: small 
or young pythons prey on rodents, but at 3-4 m body 
length their preference changes and they are documented 
to prey upon much larger and potentially more difficult 
or dangerous mammals, such as pangolins, porcupines, 
monkeys, wild pigs, mouse deer (Shine et al., 1998), 
goats and adult deer (Taylor, 1922), sun bear (Fredriksson, 
2005), and, on rare occasions, even humans (McKay, 
2006). There exist anecdotal reports of leopards being 
killed, and one of us (MOS) encountered an injured M. 
reticulatus of approximately 3.0 m total length in Thailand 

that had obviously come off badly in an encounter with 

Figure 39 Unvouchered captive individual of Malayopython 
reticulatus from the Convent of St. Antony d'Lisboa (Locality 19). 
Photo by Mark O'Shea. 

116 Asian Herpetological Research 

Vol. 6 

a large feline. Shine et al. (1998) reported that females 
shift their attention to large prey species at a smaller size 
than males. Malayopython reticulatus is oviparous, with 
females of 5.5—6.0 m body length producing clutches of 
up to 100 eggs (McKay, 2006). 
Taxonomic comments. For most of the two centuries 
following its description by Johann Gottlob Schneider 
(1801), the reticulated python remained in the A fro-Asian 
genus Python. However, the species 1s morphologically 
and biochemically quite distinct from all other members 
of this genus, with the exception of M. timoriensis. 
Rawlings et al. (2008) determined that the taxa reticulatus 
and timoriensis were sufficiently distinct phylogenetically 
from other species in the genus Python to warrant 
separate generic recognition. In a recent paper, Reynolds 
et al. (2014), provided the genus name Malayopython 
in recognition of the type locality for the species M. 
reticulatus as the Malay Archipelago (fide Alfred Russel 
Wallace). By using the genus name Malayopython, we 
follow the recommendations of Kaiser et al. (20132). 
Malayopython reticulatus is the most widely 
distributed python in Asia. The island of Timor lies 
at its extreme southern limit but the species has been 
recorded from virtually the entire Indo-Malayan and 
Philippine Archipelagos, east of Lydekker's Line and as 
far north on mainland Southeast Asia as Myanmar. The 
northernmost limit of its range is currently Itbayat Island 
(N 20.75°, E 121.83°), in the northern Philippine Batanes 
Group, only 200 km south of Taiwan, China (O’Shea 
and Lazell, 2008). Despite this extensive geographical 
range only two subspecies are currently recognized as 
distinct from the nominate form, M. r. jampeanus (Auliya 
et al., 2002) and M. r. saputrai (Auliya et al. 2002), both 
from isolated islands south of Sulawesi (Auliya et al., 
2002); all other proposed subspecies have no scientific 
validity (Kaiser et al., 2013a). Even so, it would be 
presumptuous to assume that all other populations belong 
to the nominate subspecies M. r. reticulatus, and for that 
reason no subspecific designation is used to distinguish 
the Timorese population below the species level. 
Conservation. Malayopython reticulatus is a species 
listed on CITES Appendix II and therefore protected 
from international trade. However, unlike the smaller 
Liasis mackloti, it is being harvested for skins, meat, 
and gall bladders (e.g., Iskandar and Erdelen, 2006), and 
this highly destructive activity may exert a much greater 
pressure on wild populations than the exportation of live 
specimens to the trade. Despite its abundance in other 
parts of its range M. reticulatus does not appear to be a 
commonly encountered species on Timor. 

Family Typhlopidae—Blindsnakes 

Indotyphlops braminus (Daudin, 1803 
Common names. (E) Brahminy blindsnake. (G) 
Blumentopfschlange, (T) Samea matan delek isin lotuk 

) [IV-V1I] 

(samea — snake, matan delek — blind, isin lotuk — small 

Known distribution. During Phase I we collected 
vouchers of this widespread, parthenogenetic species 
(Figure 40), two from Ladiki, near Same Manufahi 
District, and one from Loihuna, Viqueque Districts 
(Kaiser et al., 2011). 

New localities. During the later phases of the survey, six 
specimens of 7. braminus were collected and vouchered. 
Three were taken in the gardens of the Pousada de 
Maubisse (elevation 1495 m; Ainaro District; Locality 
16; USNM 579373-75), our highest record for a snake 
in Timor-Leste. Given the means by which the ancestral 
stock of this population probably arrived at this location, 
in plant pots, we do not consider this a naturally occurring 
elevation record. At much lower altitudes individual 
specimens were collected at the Timor Lodge Hotel, Dili 
(Dili District; Locality 1; USNM 579778) and in the ruins 
of the Pousada de Com (Lautém District, Locality 26; 
USNM 579496), both north coast localities. A specimen 
was also vouchered at the Convent of St. Antony d'Lisboa, 
Fatucahi (Manufahi District; Locality 19; USNM 565896) 
after it was found protruding from the cloaca of a Black- 
spined toad (Duttaphrynus melanostictus; see O'Shea 
et al., 2013). Indotyphlops braminus has now been 
documented from six mainland districts. 

Natural history. One commonly used vernacular name 
for Indotyphlops braminus is “Flowerpot Snake,” 
(German: Blumentopfschlange) a name that these 
pencil-thin, small snakes (total length up to 180 mm) 
earned because they are often found either in the root 
balls of plants in plant pots, or in the humid darkness 
underneath plant pots. A close association with tropical 
plants exported during trade is likely the secret to how 7. 
braminus became the most widely distributed snake in the 
world. It is the only known obligatorily parthenogenetic 
snake species (no male has ever been documented; 
see Booth et al. 2014 for a review of facultative 
parthenogenesis in pythons), and as such only a single 
adult specimen is required to colonize a new habitat. 
Since these snakes often inhabit the soil of tropical plant 
root balls, they can easily be transported internationally 
within plants and establish colonies wherever they arrive. 
Snakes tend to be more resistant to the effects of plant 
quarantine than insect larvae or other invertebrates, 
and thus a bridgehead can easily be established. This is 

Mark O'SHEA et al. 

Figure 40 Specimen of /ndotyphlops braminus from the leaf litter 
at the Portuguese pousada at Com (USNM 579496, Locality 26). 
Photo by Mark O'Shea. 

undoubtedly how the population of Z. braminus became 
established at an uncharacteristically high elevation in the 
gardens of the former Portuguese Governor's pousada at 
Maubisse. /ndotyphlops braminus 1s an oviparous species, 
producing clutches of 1—8 eggs (de Lang, 2011; McKay, 

Like all blindsnakes, 7. braminus is a fossorial species 

that is more commonly found on the surface when 
flooded out of burrows by heavy rain. Its rudimentary 
eyes are simple pigmented areas under translucent scales 
that warn the snake when it has been uncovered, and this 
triggers the response to burrow rapidly. Prey comprises 
soft-bodied invertebrates, primarily termite and ant larvae 
and eggs (de Lang, 2011). This small snake may itself 
become the prey of many larger vertebrates, including the 
pipesnake Cylindrophis cf. boulengeri and the introduced 
bufonid Duttaphrynus melanostictus (O'Shea et al., 
Taxonomic comments. The genus /ndotyphlops was 
recently erected by Hedges et al. (2014) to accommodate 
the South Asian blindsnake clade. Prior to this 
revision, the species braminus was placed in the genus 
Ramphotyphlops, which is now restricted to Western 
Pacific taxa. 


Indotyphlops spp. 
Common names. (E) Blindsnakes. (G) Wurmschlangen, 
Blindschlangen. (T) Samea matan delek (samea — snake, 
matan delek — blind). 

Known distribution. A series of seemingly aberrant 
Indotyphlops were collected on Ataüro Island and 
were documented elsewhere (Kaiser et al., 2013b) as 
Ramphotyphlops sp. * Ataüro'. 

New localities. A striped /ndotyphlops (Figure 41A) 
collected at an altitude of over 905 m in a rock pile on the 

Timor-Leste Herpetofauna Updates 117 

Tilomar road in Covalima District (Locality 15; USNM 
579376) during Phase IV, could not be attributed to 
either Z. braminus or Sundatyphlops polygrammicus (see 
below) and is recorded here as /ndotyphlops sp. ‘Tilomar’. 
Similarly, an unusual /ndotyphlops with a bluish body 
coloration (Figure 41B) was collected on the trail to Mt. 
Mundo Perdido, Viqueque District (Locality 21; USNM 
580542) at an elevation of 1162 m; we recognize it here 
as Indotyphlops sp. ‘Mundo Perdido.’ We considered that 
the coloration of this individual might be due to incipient 
ecdysis, but examination of the two injured areas and 
the head, as well as of the specimen after several months 
in preservative, do not support this idea. Both of these 
mainland specimens, as well as the series from Ataüro 
Island, await closer examination. Currently all three are 
incertae sedis within Indotyphlops. 

Natural history. No natural history notes are available 

for the two aberrant /ndotyphlops specimens from 
Tilomar (USNM 579376) and Mt. Mundo Perdido 
(USNM 580542), although they were both found 
sheltering under rocks at relatively high elevations, 905 
and 1162 m respectively, the highest recorded for any 

Figure 41 (A) Aberrant specimen of /ndotyphlops from under a 
rock pile at Tilomar (USNM 579376, Locality 15, elevation 905 m). 
The fine lined pattern along the body indicates that this individual is 
not conspecific with 7. braminus, and we refer to it as Indotyphlops 
sp. ‘Tilomar.’ (B) Aberrant specimen of Indotyphlops from the 
path to Mt. Mundo Perdido (USNM 580542, Locality 21, elevation 
1162 m), showing injuries and a bluish, presumably pre-ecdysis, 
coloration. Photos by Mark O'Shea. 

118 Asian Herpetological Research 

Vol. 6 

Timorese typhlopid, excluding the artificially introduced 
I. braminus at Maubisse (see above). Both locations 
where these two snakes were found are remote, at the end 
of a road into a highland area and along a mountain trail, 
respectively, and it seems unlikely that either of these 
snakes were transported to their respective locations by 
the agencies of man. 

) [IV, VI] 

Sundatyphlops polygrammicus (Schlegel, 1839 
Common names. (E) Timor blindsnake. (G) Timor- 
Wurmschlange, Timor-Blindschlange. (T) Samea matan 
delek isin baibain (samea = snake, matan delek = blind, 
isin baibain = normal body size). 
Known distribution. No specimens of Sundatyphlops 
polygrammicus (Figure 42) were collected during Phases 
I-III, although Forcart (1953) reported eight specimens, 
now in the Basel collection (NHMB 12888-95), collected 
by Prof. A. Bühler near Baguia (Baucau District). 
New localities. During Phase IV, a large specimen of 
Sundatyphlops polygrammicus (Figure 42) was collected, 
in heavy rain as it climbed a sandstone cliff-face near 
a path, on the Trilolo River, close to Same (Manufahi 
District; Locality 17). A second specimen was collected 
during Phase VII at the “Carlia site" at Afacaimau 
(Baucau District; Locality 23; USNM 580543). 
Natural history. Sundatyphlops polygrammicus is 
an infrequently encountered species, with only two 
specimens collected in eight survey phases. As with 
most blindsnakes, S. polygrammicus 1s rarely seen on 
the surface except during or following heavy rain. Our 
Trilolo River specimen (USNM 579377) was collected as 
it climbed a sandstone cliff-face besides the path down to 
the river, the only reptile or amphibian encountered by the 
entire team during an evening search in heavy rain. 

Like other typhlopid snakes, S. polygrammicus is 
a predator of soft-bodied invertebrates, primarily the 
larvae and eggs of termites and ants, but its larger size 
(larger than species such as Z. braminus) should place 
adult ants and termites, and possibly also beetle larvae, 
within its dietary range. Large numbers of prey items 
may be consumed 1n rapid succession, from 50 to over 
500 termites (de Lang, 2011). However, the majority of 
natural history notes available for this species relate to 
the former populations from Queensland, Australia, and 
Western Province, Papua New Guinea, which are now 
treated as a separate species in a different genus, Anilios 
torresianus (see Taxonomic comments below). 
Taxonomic comments. The genus Sundatyphlops was 
recently erected by Hedges et al. (2014) to accommodate 
a clade of exclusively Lesser Sunda blindsnakes. Prior 
to this revision, the species polygrammicus was placed 

in the genus Ramphotyphlops. While this most recent 
taxonomic arrangement will still need to stand the test 
of time, Sundatyphlops 1s the most current available 
name for polygrammicus, and our use of this name here 
should not be misconstrued as a taxonomic endorsement 
but merely an acknowledgment of acceptable research. 
Sundatyphlops polygrammicus is currently believed 
to contain five subspecies distributed throughout the 
Lesser Sunda Islands, with the nominate form present on 
Timor (de Lang, 2011; Hedges et al., 2014). With three 
of the other subspecies endemic to Sumba, Lombok, 
and Flores, and a fourth reported from Sumbawa and 
neighbouring Komodo and Moyo, it is unlikely that the 
rest of this considerable range is inhabited by just the 
nominate subspecies. This is a taxon clearly in need of 
revision. As formerly recognized, S. polygrammicus was 
a polyphyletic species and caused Hedges et al. (2014) 
to resurrect torresianus (now in the genus Anilios) for 
Queensland and southern Papuan populations, and to 
confine S. polygrammicus to Lesser Sunda populations. 

Family Viperidae—True Vipers and Pitvipers 
Trimeresurus (Trimeresurus) insularis (Kramer, 1977) 

Common names. (E) Lesser Sunda Island Pitviper, Island 
Pitviper, Lesser Sunda White-lipped Pitviper. (G) Insel- 
Bambusotter, Wetar-Bambusotter. (T) Samodok (a proper 

Known distribution. During the first three research 
phases, Trimeresurus insularis was documented from three 
mainland districts (Baucau, Lautém, Viqueque; see Kaiser 
et al., 2011; O'Shea et al., 2012). It was also reported 
from Atauro Island (Kaiser et al., 2013b). Bethencourt 
Ferreira (1898) reported a juvenile specimen from Aipello 
(Liquiga District), and de Lang (2011) included Dili (Dili 
District) and additional localities in Baucau and Lautém 
Districts, bringing to five the number of districts where 
this pitviper has been recorded. Trimeresurus insularis is 
also common and widely distributed in West Timor, with 
specimens being documented enroute and returning from 
the Oecusse exclave (Sanchez et al., 2012). 

New localities. The later phases produced additional live 
specimens from Lautém District at Raça village (Locality 
27; USNM 579386-87) and Com (Locality 26; USNM 
579493); from Manufahi District at Betano (“wet site,” 
Locality 19; USNM 579385); from Manatuto District 
(Nancuro coastal forest, Natarbora, S of Umaboco, 
Locality 20; USNM 580551; Figure 43); and a road- 
killed specimen from Bobonaro District on road between 
Bobonaro and Maliana road (near Locality 12). The 
Bobonaro specimen was in too poor a condition to 

No.2 Mark O'SHEA et al. 

voucher, and we instead documented it photographically 
(USNM-HI 2791). Trimeresurus insularis is now known 
to occur in eight mainland districts and on Ataüro Island, 
but thus far not at elevations over 900 m. 

Natural history. With its lithe body shape and prehensile 
tail, the island pitviper T. insularis, is usually considered 
an arboreal species, but most specimens encountered 
during our surveys have been found on the ground, at 
night in relatively wet habitats, waiting in ambush for 
prey. Prey appears to consist largely of frogs, particularly 
rice-paddy frogs (genus Fejervarya; HK, pers. obs.) but 
it is possible that small mammals or lizards may also be 
taken on occasion, as reported by de Lang (2011) from 
other islands in the archipelago. The most common color 
phase is green but cyan coloration is known from Komodo 
(de Lang, 2011; MOS, pers. obs.), and bright yellow 
occurs on Wetar and Timor-Leste (USNM 581178). As 
with most pitvipers, T. insularis is a livebearing species. 
This species (under the generic name Cryptelytrops) 
was recorded as a paratenic host of spargana tapeworms 
(Cestoda) by Goldberg et al. (2010). 

At this point in time this is the only terrestrial snake 

known to occur on Timor or in Timor-Leste, which 
includes Ataüro Island to the north, capable of delivering 
a lethal bite to a human. Deaths following the bites 
of T. insularis are on record in Timor-Leste and at the 
very least a bite and ensuing envenomation can be an 
unpleasant experience (MOS, pers. obs.). 
Taxonomic comment. Until recently, we referred to 
this species as Cryptelytrops insularis (e.g., Goldberg 
et al., 2010; Kaiser et al., 2011). We here follow the 
nomenclature proposed by David et al. (2011), which 
we believe to be correct after a careful reading of their 
assessment. According to these authors, Trimeresurus 
viridis Lacépéde, 1804 (= T. albolabris insularis Kramer, 
1977) is the true type species of the genus Trimeresurus, 
and not Coluber gramineus Shaw, 1802, as previously 
believed. Rearrangement of the nomenclature requires 
that the species insularis bear the generic name 
Trimeresurus, with the optional use of the subgeneric 
name Trimeresurus to preserve added taxonomic 
information. The genus Cryptelytrops Malhotra and 
Thorpe, 2004 is now considered a junior synonym of 

Family Crocodylidae—Crocodiles 
Crocodylus porosus Schneider, 1801 "YY V"! 

Common names. (E) Saltwater crocodile, Estuarine 
crocodile, Naked-neck crocodile, Indo-Pacific crocodile. 
(G) Leistenkrokodil, Salzwasserkrokodil. (T) Lafa'ek tasi 

(Lafa'ek = crocodile, tasi = sea). 

Timor-Leste Herpetofauna Updates 119 

Figure 42 Individual of Sundatyphlops polygrammicus from under 
a flat rock in a disturbed area south of Baucau (USNM 580543, 
Locality 23). Photo by Mark O'Shea. 

Figure 43 Adult female Trimeresurus insularis (green phase) 
found in ground vegetation in coastal wet forest at Nancuro (USNM 
580551, Locality 20). Photo by Mark O'Shea. 

Known distribution. During earlier phases we 
documented free-living crocodiles in the Malailala River 
(Lautém District), and captives caught locally at Uma 
Boot (Viqueque District). One captive (Figure 44) from 
the south coast near Betano (Manufahi District; near 
Locality 18) has been kept in an enclosure in the town 
of Aileu (Aileu District) for nearly a decade, while a 
juvenile was kept in an old oil drum nearby (Kaiser et al., 
2009, 2013c). 

New localities. During Phase IV we documented 
another captive crocodile on the Fatucahi to Betano 
road (Manufahi District; Table 5), which local residents 
had confined to an old oil drum. During Phase V a large 
adult crocodile was seen stalking water buffalo calves 
on the southern shore of Lake Ira Lalaro at Malahara 
(Lautém District; Locality 29; USNM-HI 2798). In 2012, 
a specimen was photographed at Tibar, a popular beach 
area west of Dili (Liquiga District; USNM-HI 2836). In 

120 Asian Herpetological Research 

Vol. 6 

2013 (Phase VII), we documented crocodiles on riverine 
sand banks along the north coast road in Lautém District 
(USNM-HI 2828). 

Natural history. The saltwater crocodile is the most 
widely distributed crocodile in the Australasian region and 
the largest crocodilian in the world, achieving lengths of 
over 6.0 m (Wilson, 2005) and weights in excess of 1300 
kg (Alderton, 1991; Steel, 1989). It is the climax predator 
wherever it occurs. This species 1s the only crocodilian 
found in Timorese waters where it has achieved 
mythological status as part of the island’s creation story 
(Kaiser et al., 2009; Morris, 2011). Many local people 
respect the crocodile, but this respect is not reciprocated, 
as crocodiles are responsible for an increasing number 
of human fatalities, usually fishermen or children near 
the water's edge, every year (B. Sidelau, pers. comm.). 
Reporting of such incidences is not universal, and the 
real impact on the human population is as yet unknown 
(HK and MOS, pers. obs.). Crocodiles are most common 
in estuarine river mouths or mangrove swamps, where 
the turbid water obscures their presence. We have 
observed them lingering under bridges along the coast 
roads, basking on riverine sandbanks, and resting nearly 
submerged in water among mangrove roots, but they 
may also arrive on sandy beaches close to major towns 
or tourist locations (J. Ramos-Horta, pers. comm.). There 

exists a land-locked population, estimated to number in 
excess of 300 individuals, in Timor-Leste's largest lake, 
Lake Ira Lalaro (Lautém District: Locality 29; Middleton 
et al., 2006; M. Mendes and C. Trainor, pers. comm.). 
Taxonomic comments. It is interesting to note that the 
population in Lake Ira Lalaro, a freshwater catchment in a 
limestone polje, is isolated by distance (9 km by air to the 
nearest coastline, with the 934 m high Paitxau Mountain 
range in the way) and altitude (at an elevation of nearly 
500 m) from other saltwater crocodile populations. 
Therefore, this isolate may be considered a population of 
interest for studies of isolated crocodile populations. 
Crocodile attacks. As human activity along the coastline 
and the shores of Lake Ira Lalaro has increased, reports 
of crocodile attacks including numerous human fatalities 
have risen dramatically. While there are currently no 
formal statistics on these attacks, we have heard reports 
with greater frequency during every research phase, 
and there is increased awareness on the part of the 
government that this might need to be considered an 
important public health issue (HK, pers. obs.). It appears 
to us that in the mindset of the Timorese populace, a 
people that has experienced great violence in the recent 
past and which is fighting to emerge from extreme 
poverty, such attacks may simply be considered a fact 
of normal life, akin to motorcycle accidents or falls 

Table 5 Records of crocodilians and turtles for the districts of Timor-Leste. Black circles indicate previously known records, red circles 

denote new records. 

> © 
Taxon o 3 S 3 i References* 
o 3 E B E E S g S 8 £ 9 2. 

a 5 8 S T — — o D nd S 3 E o 

a S 3 = Ne E s g g a 

= E E: 8 $5 = = £ E SG z ES 9 S 

< < m m oO A A BA FA 3 2 2 O p 
Crocodylus porosus e. e. e. 1-4 
Chelodina mccordi timorensis e. 1 
Chelonia mydas e. e. 1,2 
Eretmochelys imbricata e. 2 
Mauremys reevesii e. 1,6 
Pelodiscus sinensis 5 

'References are identified numerically as follows: 1 = Kaiser et al., 2011; 2 = this paper; 3 = Kaiser et al., 2009; 4 = de Rooij, 1917; 5 = 

Bethencourt Ferreira, 1898; 6 — Kaiser et al., 2010. 

No.2 Mark O'SHEA et al. 

Figure 44 Captive Crocodylus porosus in an enclosure at Aileu (see 
Kaiser et al., 2009, 2013c). Photo by Mark O’Shea. 

from coconut palms. However, crocodile attacks do not 
have to be part of a valiant, post-conflict socioeconomic 
struggle. In an earlier report (Kaiser et al., 2009), we 
outlined some of the challenges resulting from the 
interactions between humans and crocodiles, as well as 
some of the misconceptions about living with crocodiles 
as neighbors. While there are no simple solutions, it 
does not appear that any systematic evaluation of the 
issue has taken place. We therefore propose that the 
Government of Timor-Leste make reports of crocodile 
attacks compulsory (including name and age of the victim 
as well as the locality, and the activity during which the 
attack occurred) and form an inter-ministerial task force, 
to include members from departments handling public 
health, internal security, environment, and tourism, to 
create and implement an educational plan so that the risk 
of death from crocodile attacks can be minimized. 

Family Cheloniidae—Sea Turtles 
Chelonia mydas (Linneaus, 1758) 
Common names. (E) Pacific Green Sea Turtle. (G) 
Suppenschildkróte, Grüne Meeresschildkróte. (T) Lenuk 
tasi kór-matak (lenuk tasi = sea turtle, kór-matak = green). 


Known distribution. There have not been any confirmed 
records of Chelonia mydas for Timor-Leste. However, 
it is listed in the IUCN Red List of Threatened Species 
(Seminoff, 2004) as native in that country. 

New localities. During Phase IV we found a dismembered 
carapace of C. mydas above the shoreline in the Nancuro 
coastal forest, Natarbora, S of Umaboco (Manatuto 
District; Locality 20; Table 5). The carapace was re- 
constructed using beach sand for support and a voucher 
photograph was obtained (Figure 45, USNM-HI 2792). 
Natural history. This turtle may achieve a carapace 
length of 1.5 m (Wilson, 2005) and weights up to 200 kg 
(Spotila, 2004). Chelonia mydas migrates long distances 
between breeding beaches, the open ocean, and shallow, 
inshore, clear water bays with sea grass where they 

Timor-Leste Herpetofauna Updates 121 

graze on algae and other marine vegetation. Adults are 
primarily, if not totally, herbivorous, but juveniles do 
include marine animals in their diets (Wilson, 2005). 
The lifespan of this turtle may exceed 50 years (Zug and 
Balazs, 1985). 

Taxonomic comments. Some authors recognize two, 
others three, subspecies of C. mydas. The population 
on the coast of Timor could be attributed to either C. m. 
agassizi (Bocourt, 1868) or C. m. japonica (Thunberg, 

Conservation. Chelonia mydas is an endangered species 
that was harvested well into the 20" century as part of the 
natural products trade, for its eggs, and for its meat for 
turtle soup (the German name Suppenschildkrote = “turtle 
used for soup" refers to the usage of the species as part 
of human diet). Although such trade is now banned, it 1s 
very difficult to prevent further exploitation of this turtle 
or its nests in economically challenged countries where 
a specimen may represent a financial windfall. We have 
on at least three occasions observed individuals offering 
sea turtle eggs, of uncertain species affinity, for sale 
along the main coastal road in Dili, with neat displays of 
four eggs per order sold alongside of the day's catch of 
fish or octopus. Whereas this type of trade is illegal in 
Timor-Leste, timing hinders enforcement, given that one 
motivated buyer may take the proof and leave no grounds 
for legal action. This same comment regarding local 
exploitation equally applies to other sea turtle species, 
including Eretmochelys imbricata, the only other species 

so far identified during our surveys. 

Figure 45 Carapace of a hunted and killed specimen of Chelonia 
mydas, of which we found and reassembled all elements, just inland 
from the shore at the Nancuro Protected Area (Locality 20). Even 
though the remaining portions of the skeleton were missing, we 
were able to determine that there was human involvement by the 
presence of recent (i.e., not healed) harpoon damage. The flashlight 
(length = 146 mm) is provided as a scale. Photo by Mark O'Shea. 

122 Asian Herpetological Research 

Vol. 6 

Eretmochelys imbricata (Linneaus, 1766) ` 

Common names. (E) (Pacific) Hawksbill (Sea) Turtle. 
(G) Echte Karettschildkróte. (T) Lenuk tasi eretmochelys 
(lenuk tasi = sea turtle, ibun = beak, makitik = hawk). 
Known distribution. No confirmed records existed for 
Eretmochelys imbricata for Timor-Leste. 

New localities. During Phase V the carapace of a 
juvenile specimen of E. imbricate was found on Jaco 
Island (Lautém District; Locality 30; Table 5). A voucher 
photograph was obtained (Figure 46, USNM-HI 2793). 
Natural history. Adults of E. imbricata may achieve 
a carapace length of up to 1.0 m (Wilson, 2005) and a 
weight approaching 80 kg (Spotila, 2004). Although 
this highly migratory species may be encountered in 
a wide variety of marine habitats, from open ocean to 
mangrove swamps and estuarine river mouths, it is most 
often associated with the clear, shallow waters of coral 
reefs, where it feeds primarily on sponges (Porifera). 
Hawksbill turtles also prey upon molluscs (Mollusca), 
jellyfish (Scyphozoa), including highly venomous 
species, sea combs (Ctenophora), sea anemones and 
soft corals (Anthozoa), and marine algae (Ernst et al., 
1994). Juveniles are solely carnivorous (Wilson, 2005). 
Hawksbill turtles may live for 30—50 years (Ernst et al., 

Taxonomic comments. Two subspecies are recognized, 
with the Indo-Pacific populations belonging to E. i. bissa 
(Rüppell, 1835). 

Conservation. Eretmochelys imbricata is a critically 
endangered species that has suffered historically from 
harvesting for the turtle shell industry, a practice now 
outlawed but continuing. Given their size, they are also 
harvested for food, and their eggs are collected from 
nesting sites or from slaughtered females. 

Family Geoemydidae—Asian Hard-shelled Turtles 
Mauremys reevesii (Gray, 1831) "Y" 

Common names. (E) Chinese pond turtle. (G) 
Chinesische Dreikielschildkröte. (T) Lenuk kakorok 
riskadu (lenuk = turtle, riskadu = striped, kakorok = 

Known distribution. During Phase I we vouchered a 
specimen of Mauremys reevesii from Baucau (Baucau 
District; Kaiser et al., 2010) and reported the presence of 
a population in Dili (Dili District; Kaiser et al., 2011). 
New localities. During Phase IV we were able to confirm 
the presence of a population of Mauremys reevesii in a 
kangkong (Ipomoea aquatica) patch in Becora, eastern 
Dili (Dili District; Locality 1; USNM 579393; see 
Kaiser et al., 2013c). We also encountered a number of 
specimens kept as garden pets in Comoro, western Dili 

(Table 5). 

Natural history. Mauremys reevesii is an introduced 
turtle that probably arrived as a deliberate introduction 
from the Asian mainland, for food or to be used in 
traditional Chinese medicine, or as a totem associated 
with Chinese culture and long life. It appears to have been 
present in the Dili area for several decades (Kaiser et al., 
2010). As shown by the presence of a male individual 
in black breeding colors (Figure 47), the population is 
reproductively active and perhaps even self-sustaining in 
their human-maintained habitat. 

Conservation. Even though the population found in 
Timor-Leste was introduced, it may be of significance in 
terms of the long-term conservation management of the 
species. In its native habitat in temperate and subtropical 
regions of mainland East Asia (China, North Korea, South 
Korea) M. reevesii has been severely exploited and has 
become very rare in the wild, having earned the IUCN 
status of Endangered A2bcd+4bcd (van Dijk, 2013). 
While populations also occur in Taiwan and Hong Kong 
of China, and Japan, these appear to have been exposed 
to hybridization with escapees from the trade (Taiwan, 
China; Fong and Chen 2010) or with M. japonica 
(Temminck and Schlegel, 1835), a close relative (Japan; 
Suzuki et al. 2011). Thus, the population in Timor- 
Leste, which by our findings appears to be a robust, self- 
sustaining population, may represent an important genetic 
reservoir of pure M. reevesii. 

Family Chelidae—South American and Australasian 
Side-necked Turtles 

Chelodina mccordi Rhodin, 1994 "! 

Common names. (E) Timor Snake-necked Turtle. (G) 
McCord-Schlangenhalsschildkróte. (T) Lenuk kakorok 
ular (lenuk = turtle, kakorok = neck, ular = snake). 
Known distribution. Snake-necked turtles on Timor 
are limited in distribution to Lake Ira Lalaro in Lautém 
District and the surrounding swampy grasslands (Table 5). 
New localities. Although we have not personally 
encountered or even pursued C. mccordi after our initial 
survey in 2009, we received several reports regarding 
their presence. These primarily came from local residents 
of Malahara village, at Lake Ira Lalaro’s southern shore, 
from forest guards working in the vicinity of the lake, and 
from expatriates working in Timor-Leste. They lead us to 
conclude that C. mccordi exists in all near-shore habitats 
around the lake (Kuchling et al., 2007). 

Natural history. Chelodina mccordi (Figure 48) is 
regarded as one of the world’s 25 most endangered turtle 
species (Rhodin et al., 2011). Amongst the reasons for 
this designation are its highly localized populations 

Mark O'SHEA et al. 

Figure 46 Carapace of a juvenile Eretmochelys imbricata from 
Jaco Island (Locality 30). The dollar bill (length = 156 mm) is 
provided as a scale. Photo by Mark O'Shea. 

Figure 47 Adult male Mauremys reevesii from the kangkong 
paddies at Balide, a part of the city of Dili (USNM 579393, Locality 
1; see Kaiser et al., 2013c). Photo by Mark O'Shea. 

Figure 48 Unvouchered adult individual of Chelodina mccordi 
from Lake Ira Lalaro, Lautém District (see Kaiser et al., 2011). 
Photo by Mark O'Shea. 

Timor-Leste Herpetofauna Updates 123 

(one in a single lake in Timor-Leste, one in two lakes on 
Roti Island), the uncertain population dynamics (e.g., 
population size, recruitment, density), and the high 
potential for extirpation due to local dietary and cultural 
customs or incipient exposure to international trade. 
Very little is known about the ecological parameters of 
C. mccordi in Timor-Leste, although a study has just 
commenced (C. Eisemberg, pers. comm.). Individuals 
of C. mccordi are most frequently encountered by local 
residents during the drier months of the year, when the 
waters of Lake Ira Lalaro recede and smaller patches of 
densely vegetated freshwater become isolated. Malahara 
villagers may have captured up to 30 specimens of C. 
mccordi from the environs of Lake Ira Lalaro during 
a single day (Kuchling et al., 2007), likely in support 
of an annual cultural event. An educational booklet 
was recently produced and is now used in schools to 
encourage conservation of the species (Eisemberg and 
Perini, 2014). 

Taxonomic comments. Even though McCord et al. 
(2007) described this population as a distinct species, 
their taxon was implicitly synonymized later that year 
by Kuchling et al. (2007), who considered the Lake Ira 
Lalaro population in Timor-Leste to be a subspecies of 
C. mccordi and gave it the name C. m. timorlestensis. 
The taxon name timorensis takes nomenclatural priority 
over timorlestensis, and thus this population should be 
referred to as C. m. timorensis (McCord et al., 2007) if a 
subspecific name were to be used. Two other subspecies 
of C. mccordi are recognized, the nominate form from 
western Roti Island and C. m. roteensis McCord et al., 
2007 from eastern Roti Island (van Dijk et al., 2014) 

4. Discussion 

Species Distribution 

The mosaic geological structure of Timor Island and 
the exploitation of organic natural resources during 
colonial times and throughout the Indonesian occupation 
(i.e., cutting of sandalwood and tropical hard woods, 
development of coffee monoculture, rice farming 
and other large-scale agricultural practices, human 
settlement) inescapably lead to certain assumptions 
about the distribution of the local herpetofauna. Habitat 
disturbances and destruction are known to effect 
significant changes in species distributions (e.g., Gardner 
et al., 2007; McKinney, 2002; Wolf et al., 2013), and this 
is perhaps most pronounced in tropical environments. We 
are therefore pleased that the reports of our herpetofaunal 
surveys to date (Kaiser et al., 2011, 2013b; O’Shea et al., 


Asian Herpetological Research 

Vol. 6 

2012; Sanchez et al., 2012; this paper) appear to 
document much higher herpetofaunal diversity than we 
had expected, which includes a series of putatively single- 
island endemics (e.g., Cyrtodactylus, Eremiascincus, 
Kaloula, Stegonotus). 

When considering species distributions, two of the 
most common ways to showcase diversity are to use 
political boundaries or habitat types. In Timor-Leste, the 
most convenient method is to use established political 
boundaries (Districts), especially since habitats are 
fragmented, disturbed, or otherwise not cohesive. It 
would be difficult to predict detailed countrywide species 
distributions for Timor-Leste based on the coverage 
of specific habitats due to the high degree of habitat 
degradation, the presence of habitat fragments of diverse 
types, sizes, and ecological qualities, and the existence of 
uncertain corridors between such habitats. Furthermore, 
it 1s clear that our sampling effort plays a significant role 
in how we can account for species distributions: while 
we have been able to sample in all 13 districts of Timor- 
Leste, some districts received a disproportionate amount 
of attention when sampling, entirely for logistical reasons. 
Whereas the political boundary method admittedly falls 
short of the most productive approach to make statements 
about species biology, it allows us to provide a geographic 
overview even while comprehensive studies of habitats 
are still very limited. The best available information 
regarding habitat distributions is still that provided by 
Trainor et al. (2007). 

As expected, there does not appear to be any general 
signal in the species diversity when looking at political 
boundaries, with the exception of Lautém District (Table 
6), which comprises the entire eastern end of Timor 
Island and includes Lake Ira Lalaro, the country's largest 

body of freshwater. Even though we visited Lautém only 
three times during Phases IV-VIII, we found 31 species, 
among them 16 lizard and nine snake species. In contrast, 
all of our surveys begin and end in Dili District, and 
our species count there 1s 21 species, with seven lizards 
and ten snakes. Aileu and Ainaro are the only districts 
with a species count below ten, and this is due to a lack 
of sampling effort. With a number of diverse mountain 
habitats, it 1s all but certain that the diversity in these 
districts should match that reported for other mountainous 
districts (e.g., Manufahi; Table 6). 

One other way to provide a general approach to species 
distribution patterns in Timor-Leste is to consider the 
north-south distribution, which largely reflects a dry-moist 
divide, respectively. The high mountains that form the 
spine of Timor act as a barrier to clouds from the south, 
effectively placing most of the habitats along the northern 
shore of Timor-Leste into a rain shadow. Portions of 
northern coastal Manatuto and Baucau Districts rank 
amongst the driest parts of Southeast Asia (Monk et al., 
1997). While some may consider lush tropical habitats 
to be those with the greater species diversity, perhaps 
due to the greater stratification of vegetation and the 
concomitant availability of niches, our data to date do 
not agree: species richness in the north is 34, in the south 
it is 35 species. As discussed above, we believe this to 
be a function of sampling effort, given that the south 
coast of Timor-Leste has limited infrastructure to support 
scientific surveys, rivers in places are unfordable even 
for 4 x 4 vehicles, and some areas are effectively isolated 
from study. This situation may improve as bridges are 
built or rebuilt, and other aspects of the infrastructure are 
improved. With increased access to the more remote areas 
we expect the list of Timorese reptiles and amphibians 

Table 6 Known species diversity of amphibians and reptiles in the districts of Timor-Leste. 

Taxon 5 * g 
E: o ‘Ss [^ 
8 3 à B g g s BP 3 3 E 
z E Bi E S = = g * | E E B 3 
= A E 8 Š = = E E RE Es S 3 S 
< < e m [9] A A a] d d z z Ó > 
Frogs 2 1 4 3 3 - 2 3 4 4 3 3 
Lizards - 3 11 5 5 10 d 16 7 12 10 9 
Snakes 1 8 3 3 10 4 - 9 1 4 9 2 6 
Turtles - - 1 - - 1 - - 3 - - - - - 
Crocodilians - - - - - - - - 1 - 1 - - 1 
TOTAL 3 4 24 11 11 21 14 11 31 11 21 23 14 16 

No.2 Mark O'SHEA et al. 

to continue to grow. The same can be expected for some 
of the more inaccessible mountainous areas. We also 
expect that the Department of National Parks of Timor- 
Leste will begin to conduct surveys for amphibians and 
reptiles independently of our own effort in the coming 
years, particularly in the Protected Areas and Nino Konis 
Santana National Park, and that this work will result in a 
more equitable sampling effort throughout the 13 districts 
of the country, as well as across the north-south divide. 


Our surveys uncovered a significant amount of single- 
island endemism. Before we began our surveys, the 
number of single-island endemics stood at eight 
(Limnonectes timorensis, Litoria everetti, Draco 
timoriensis, Carlia peronii, C. spinauris, Eremiascincus 
antoniorum, E. timorensis, Chelodina mccordi timorensis). 
For frogs, we have ascertained that at least two species of 
Fejervarya coexist in Timor-Leste, neither one of which 
is conspecific with F verruculosa (Roux, 1911), their 
geographically closest congener found on neighboring 
Wetar Island. Furthermore, the population of Kaloula 
from the southern coast of Manatuto and Manufahi is 
a species distinct from K. baleata sensu stricto as well 
as from the recently described K. indochinensis and K. 
latidisca. This more than doubles the number of endemic 
frog species, with all of the new discoveries linked by a 
Southeast Asian biogeographic ancestry. Endemism is 
even more pronounced in lizards, and in their case the 
ancestry is a mixture of cis- and trans-Wallacean elements. 
Whereas Cyrtodactylus (as many as six putative new 
species), Hemiphyllodactylus and Draco are certainly taxa 
of Southeast Asian origin, Carlia (up to five putatively 
new species) and Sphenomorphus (up to four putatively 
new species) are Indo-Papuan, and Eremiascincus (up 
to four new species) is Australian. The snake fauna also 
includes endemics, and some are still being recognized. 
In addition to the known endemics, Sundatyphlops 
polygrammicus and Dendrelaphis inornatus timorensis 
(the latter of which may deserve recognition at the species 
level; Gernot Vogel, pers. comm.), we have discovered 
at least one new species from the Australo-Papuan realm 
(genus Stegonotus), and perhaps two new species of 
Indotyphlops. Given our relatively patchy sampling effort 
in both geographic and temporal terms, we are certain that 
our estimate of endemism for Timor, the largest Lesser 
Sunda Island, is still conservative. Our findings therefore 
contradict those of Malcolm Smith, who stated, “from 
a herpetological point of view, Timor is one of the most 
disappointing places that one can visit? (Smith 1927:199). 

Timor-Leste Herpetofauna Updates 125 

Timor-Leste's Herpetofaunal Diversity in the 

Only four historical publications exist that are entirely 
devoted to the herpetofaunal diversity of the land area 
now called Timor-Leste (Bethencourt Ferreira, 1898; 
Managas, 1956, 1972; Themido, 1941), and each of these 
is very limited in scope. Several other publications focus 
on the western part of Timor Island (e.g., Smith, 1927; 
van Lidth de Jeude, 1895) or on the entire island as part of 
more general surveys (e.g., Barbour, 1912; de Rooij, 1915, 
1917; van Kampen, 1923). Where erroneous records were 
presented, either because of misidentification or due to 
errors perpetuated via faulty taxonomy, we corrected these 
in our earlier papers (Kaiser et al., 2011; O’Shea et al., 

The newly identified high degree of endemism provides 
both a challenge and an opportunity for species 
management and conservation. The challenge lies with 
the landmass itself and the economic development of a 
population whose free market economic drive has been 
regulated for centuries by external forces. The landscapes 
in Timor-Leste are made of extremes, both in terms 
of terrain (much of the habitat 1s steeply sloped) and 
climate (dry spells may be long and devastating, rains 
may be torrential and destructive). As a consequence, 
any reporting on species diversity and distribution will 
retain a certain geographic and temporal patchiness. 
While the Government of Timor-Leste has set aside a 
significant portion of Lautém District as Nino Konis 
Santana National Park and has formally protected nearly 
30 key areas (as Areas Protegidas), the establishment and 
implementation of management plans for these locales is 
only being realized very slowly due to issues with both 
capacity-building and governmental priorities. It is easy to 
enforce environmental policies in the absence of poverty, 
but it is nearly impossible or desirable when a family's 
next meal must come from the land of a protected 
area. As a consequence, the quickly developing market 
economy of Timor-Leste cannot and should not be curbed 
by copying environmental legislation from elsewhere; we 
feel that education about diversity and conservation, as 
well as the scientific use of the protected areas, is the best 
initial step to promoting broad-scale conservation in the 
country. It is fortunate that, based on our own experience 
with government leaders, the country is beginning to take 
these steps. 

The opportunity for species management and 
conservation arises through the potential scientific utility 
of the national park and the protected areas. Currently, 

126 Asian Herpetological Research 

Vol. 6 

these areas are staffed by a cadre of forest guards (Guarda 
Florestal), and several individuals may be assigned to 
serving a single protected area. The national park also 
has a special office in Lospalos, Lautém District, which 
houses the administrative base for the area. At this point, 
forest guards are under-utilized in their activities and 
represent hidden scientific potential. Once educated and 
equipped, these individuals could readily be assigned 
repeatable tasks, including: (1) twice daily recording of 
basic environmental data (e.g., temperature, precipitation, 
humidity, cloud cover, etc.); (2) conducting digital 
camera-assisted visual encounter transects (Heyer et 
al., 1994) in their area; and (3) filing monthly reports 
of photo-vouchered species encounters. In a few years, 
Timor-Leste, with its existing protected area network and 
personnel infrastructure, could perhaps become one of the 
best-researched tropical countries in Southeast Asia. 

Acknowledgements Our foremost thanks are for the 
unwavering, personal support we have received from 
Their Excellencies Xanana Gusmao, current Prime 
Minister and former President of Timor-Leste, and 
José Ramos-Horta, former President and former Prime 
Minister of Timor-Leste. Their interest in the survey 
work, in the educational opportunities this brings to 
Timorese citizens, and the welfare of wildlife in the 
country is deeply rooted in their understanding of nation- 
building and very gratifying for a team of scientists. This 
gratitude extends further, especially to Claudia Abate- 
Debat, former Senior Advisor in the Prime Minister’s 
Office, for her tireless efforts to allow us to meet the right 
people and for helping us comport ourselves with aplomb 
during important meetings, and to Manuel Mendes, 
Director of National Parks, for issuing the necessary 
permits and for his dedication to the conservation of 
Timor-Leste. Special thanks go to Damien Kingsbury, for 
his assistance with any matters requiring an historical or 
political perspective. We received able assistance in the 
field during Phases IV-VIII from Zach Brown, Kevin 
Burns, Melissa Carillo, Marissa Cox, Britta Dóring, 
Joanna Flores, Scott Heacox, Stephanie Hughes, Naveen 
Jalota, Paul Landry, Aaren Marsh, Gloria Morales, Kyle 
Olsen, Jay Paris, Julia Pozo, Justin Rader, Claudia Rivas, 
Robert Sewell, David Taylor, and Franziska Wagner. 
Without them, many specimen records would have been 
missed. For their assistance with the logistics of travel 
we thank Paulo Aniceto (Rentló Car Rental), Faridah 
Suhaimi (Air Timor), Gareth Turner (Air Timor), Ed 
Turner (Air Timor), Ian Groucott (Emirates), as well 
as the staff at Timor Lodge Hotel, particularly Kemal, 

Michelle, and Sheemon. A survey such as this requires 
an inordinate amount of specimen work covering many 
taxa in order to provide the most reliable identifications 
possible. We therefore gratefully acknowledge the 
assistance of the many individuals who were instrumental 
in facilitating loans, or permitting one or several of us 
to examine specimens under their care (for institutional 
abbreviations see Sabaj Pérez, 2014): Jeremy Jacobs, 
Steve Gotte, Robert Wilson, Kenneth Tighe, George Zug, 
and Roy McDiarmid (USNM); Annemarie Ohler and 
Ivan Ineich (MNHN); Gunther Kóhler and Linda Acker 
(SMF); Wolfgang Bóhme and André Koch (ZFMK); Pim 
Arntzen, Ronald de Ruiter, and Esther Dondorp (RMNH); 
Patrick Campbell and Colin McCarthy (BMNH); José 
Rosado, Joe Martinez, and James Hanken (MCZ); David 
Kizirian and David Dickey (AMNH); Karla Schneider 
(MLU); Rolf Beutel and Matthias Krüger (Phyletisches 
Museum, Jena, Germany); Fritz Geller-Grimm (MWNH); 
Raffael Winkler, Denis Vallan, and Urs Wüest (NHMB); 
Heinz Grillitsch, Silke Schweiger, and Georg GaBner 
(NMW); Frank Tillack and Mark-Oliver Ródel (ZMB); 
Paul Doughty (WAM); Kelvin Lim (ZRC); Stefan 
Hertwig (NMBE); Raffael Ernst and Markus Auer 
(MTKD); Andréas Schmitz (MHNG); and Alexander 
Haas and Jakob Hallermann (ZMH). Financial assistance 
for equipment and supplies was partially provided by a 
Title V Grant to Victor Valley College. Partial financing 
of student travel was provided by the Associated Student 
Body at Victor Valley College, and by donations from 
Pamela MacKay and Melinda Fisher. This paper is 
Contribution No. 15 from the Tropical Research Initiative 
at Victor Valley College. 


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