• TABLE OF CONTENTS
HIDE
 Front Cover
 Copyright
 Table of Contents
 Acknowledgement
 Ecology of Komodo
 Systematics
 Zoogeography
 Literature cited
 Back Cover






Group Title: Bulletin of the Florida State Museum
Title: The Herpetofauna of Komodo, with notes on adjacent areas
CITATION THUMBNAILS PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00095815/00001
 Material Information
Title: The Herpetofauna of Komodo, with notes on adjacent areas
Series Title: Bulletin - Florida State Museum ; volume 25, number 2
Physical Description: p. 40-156 : ill. ; 23 cm.
Language: English
Creator: Auffenberg, Walter
Donor: unknown ( endowment )
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1980
Copyright Date: 1980
 Subjects
Subject: Reptiles -- Indonesia -- Komodo Island   ( lcsh )
Amphibians -- Indonesia -- Komodo Island   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
Spatial Coverage: Indonesia
 Notes
Bibliography: Bibliography: p. 151-156.
General Note: Cover title.
Statement of Responsibility: Walter Auffenberg.
 Record Information
Bibliographic ID: UF00095815
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 07513195
lccn - 81621399

Table of Contents
    Front Cover
        Page 37
    Copyright
        Page 38
    Table of Contents
        Page 39
    Acknowledgement
        Page 40
        Page 41
        Page 42
    Ecology of Komodo
        Page 43
        Page 44
        Page 45
        Page 46
        Page 47
        Page 48
        Page 49
        Page 50
        Page 51
        Page 52
        Page 53
    Systematics
        Page 54
        Page 55
        Page 56
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        Page 130
        Page 131
        Page 132
        Page 133
        Page 134
        Page 135
        Page 136
    Zoogeography
        Page 137
        Page 138
        Page 139
        Page 140
        Page 141
        Page 142
        Page 143
        Page 144
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        Page 147
        Page 148
        Page 149
        Page 150
    Literature cited
        Page 151
        Page 152
        Page 153
        Page 154
        Page 155
        Page 156
        Page 157
    Back Cover
        Page 158
Full Text








of the
FLORIDA STATE MUSEUM
Biological Sciences


Volume 25 1980 Number 2




THE HERPETOFAUNA OF KOMODO,
WITH NOTES ON ADJACENT AREAS


WALTER AUFFENBERG


UNIVERSITY OF FLORIDA


GAINESVILLE:







Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM, BIOLOGICAL
SCIENCES, are published at irregular intervals. Volumes contain about 800 pages and
are not necessarily completed in any one calendar year.





OLIVER L. AUSTIN, JR. Editor

RHODA J. RYBAK, Managing Editor

Consultants for this issue:

ALAN E. LEVITON

GEORGE R. ZUG





Communications concerning purchase or exchange of the publications and all
manuscripts should be addressed to: Managing Editor, Bulletin; Florida State
Museum; University of Florida; Gainesville, Florida 32611.





Copyright 1980 by the Florida State Museum of the University of Florida.


Publication date: September 22, 1980


This public document was promulgated at an annual cost of
$5,801.46, or $5.801 per copy. It makes available to libraries,
scholars, and all interested persons the results of researches in
the natural sciences, emphasizing the circum-Caribbean region.


Price: $5.80









THE HERPETOFAUNA OF KOMODO,
WITH NOTES ON ADJACENT AREAS

WALTER AUFFENBERG1

SYNOPSIS.-The reptiles and amphibians of Komodo are reviewed; their taxonomy,
distinguishing features, habitat preferences, parasites, and food habits are discussed.
Zoogeography of the Lesser Sunda Islands is reconsidered in the light of new geologic
data. A number of the included species have never before been reported from
Komodo-some constitute considerable range extensions. New taxa are Typhlops
schmutzi (n. sp. from Komodo), HemiphyUodactylus typus pallidus (n. ssp. from
Komodo), and Cyrtodactylus laevigatus uniforms (n. ssp. from Flores). Leilopisma
kadarsani Darevsky is placed in the synonomy of Emoia similis Dunn, and Sphenomor-
phus mertensi Darevsky is placed in S. emigrans emigrans (Lidth de Jeude). Cryp-
toblepharus boutonii burden (Dunn) is given specific status as C. burden Dunn.

TABLE OF CONTENTS

[NTRODUCTION .................................................... 40
MATERIALS AND METHODS ........................................ 40
ACKNOWLEDGEMENTS .............................. 40
ECOLOGY OF KOMODO .............................................. 43
Edaphic Factors ..................................... .............. 43
Climatic Factors ............ ..... ................... ........... 43
Vegetational Factors ............... ............................. 45
SYSTEMATICS ............ ......................................... 54
Family Microhylidae ............................................... 54
Family Ranidae .................................... ............ 56
Family Cheloniidae ............................................... 60
Family Crocodylidae.............................................. 62
Family Gekkonidae ................................................ 63
Family Scincidae .............. ................... .............. 75
Family Agamidae ..................................... .......... 94
Family Dibamidae ....................................... ......... 95
Family Varanidae ............. ................................... 96
Family Typhlopidae ...............................................113
Family Aniliidae ............................. ..................... 115
Family Acrochordidae ............... ............................ 117
Family Colubridae ............. ..................... ............117
Family Elapidae ............. ....................... ............129
Family Viperidae ............. ..................... .............. 131
ZOOGEOGRAPHY .............. ................... ............137
Geology and History .................................. ............137
Zoogeography of Lesser Sunda Reptiles ................................. 142
A Hypothesis of the Zoogeography Development of the Lesser
Sundanese Herpetofauna .......................................149
LITERATURE CITED ...............................................151

'The author is Curator in Herpetology at the Florida State Museum, University of Florida, and Professor of Zoology,
Department of Zoology, University of Florida, Gainesville, Florida 82611.

AUFFENBERG, WALTER. 1980. The Herpetofauna of Komodo, with Notes on
Adjacent Areas. Bull. Florida State Mus., BioL Sci. Vol 25(2):39-156.









BULLETIN FLORIDA STATE MUSEUM


INTRODUCTION
During 1969-70, 1971, and 1973 I collected reptiles and amphibians
on Komodo and nearby islands in connection with a 17-month study of
the Komodo monitor (Varanus komodoensis, Auffenberg, in press).
This report deals with the material collected on Komodo incidental to
that study, though comparative material from other Sunda Islands is
often referred to. Earlier reports dealing with the herpetology of
Komodo were results of two previous major field studies in the area:
the Douglas Burden expedition (Dunn 1927a, 1927 b, Burden 1927)
and the joint Indonesian-Soviet Expedition (Darevsky 1964a, 1964b,
1964c, Darevsky and Kadarsan 1964). The present contribution shows
that the fauna is considerably more diverse than previously believed.
MATERIALS AND METHODS
Three collections form the basis of this study: those of the Florida State Museum,
University of Florida (UF), the American Museum of Natural History (AMNH), and the
Museum Zoologicum Bogoriense (Bogor, Java, MZB). Fortunately I was able to see in
the wild all but two of the species reported from the island and to study the types of all
the endemic forms.
Synonymies in the species accounts include only the original reference, the earliest
use of the present name, and all synonymies used in papers dealing with Komodo and
nearby islands. The spelling of many place names in Indonesia is remarkably varied,
and no gazetteers include all the local names. I have tried to be consistent in using the
most approved local spellings.
ACKNOWLEDGEMENTS
Primary support for the investigation of the behavior of Varanus komodoensis came
from the New York Zoological Society and the Florida State Museum. Critical aid in the
field came from various agencies of the Indonesian government. In particular, I owe
thanks to the official assistance and cooperation of Hasan Basjarudin Nasution, Direc-
torate of Forestry, and Walman Sinaga, Department of Conservation, Bogor. Without
their help it would have been impossible to study the monitors on any of the islands.
Otto Soemarwoto, National Academy of Biology, S. Somadikarta, Museum Zoologicum
Bogoriense, and especially Sjamsiah Achmad, Indonesian Institute of Sciences, were
instrumental in facilitating our research program. I am very grateful for the varied
assistance rendered the project by the late Hilmi Oesman, Surabaja Zoological
Gardens, and Benjamin Galstaun, Jakarta Zoological Garden.
The field work would have been considerably less successful and pleasant without
the able assistance of Putra Sastrawan (Denpasar, Bali), William Holzmark (Sarasota,
Florida), Robert Armistead (Gainesville, Florida), Fred G. Thompson and Howard H.
Converse (Florida State Museum), and particularly my sons Kurt and Garth. Of utmost
importance was the cooperation and interest in the project shown by the Komodo
villagers. In the laboratory, I owe many thanks to Richard Franz and Sylvia Scudder
(Florida State Museum) without whose help the more tedious parts of the study might
have remained incomplete. The suggestions and (in some cases) loans of specimens in
their care by Illya Darevsky, S. Kadarsan, G. Zug, E. Williams, H. Oesman, and R.
Zweifel are hereby gratefully acknowledged. Finally, this work would not have been
possible without the continued interest shown by F. Wayne King, formerly of the New
Zoological Society, currently Director of the Florida State Museum.
Those drawings not done by myself (uninitialed) are the work of Nancy R. Halliday
and Glen Rogers. Rhoda J. Rybak typed and copystyled the manuscript. I am deeply
indebted to Alan Leviton and George Zug, whose critical reviews greatly improved the
manuscript.


Vol. 25, No. 2











KALIMANTAN
(BORNEO)


BANDA
SEA


Babar 7


ARAFURA
SEA


INDIAN OCEAN


150 km.
I ====I


AUSTRALIA


FIGURE 1.-Map showing location of Komodo (in marked rectangle, enlarged as Fig. 2).


od a
Kai 1'. If Aru Is.



















FLORES SEA



S BANTA =





.... .....



u "m"aw .. .:"....-." *




OEWADA SAM GILl MOTA *





SUMBA SEA o



FIGURE 2.-Komodo Island and vicinity. Dotted line represents 100 m contour below sea level; black area indicates land surface above 500 m.
P0








AUFFENBERG: KOMODO HERPETOLOGY


ECOLOGY OF KOMODO
Komodo is a small island (ca 160 km2) of the Lesser Sunda group
between Java and New Guinea (Figs. 1 and 2, 8S, 119E). Much of
the surface is hilly to mountainous (maximum elevation to 827 m).
Komodo localities mentioned in the text are shown on Figure 3.
The general aspect of the island is typical of all arid, uplifted
regions, because the mountains have steep angular slopes and alluvial
fans. Stream beds and valley floors tend to be rocky with shallow
residual or alluvial soils. Rapid runoff leads to deep, nearly vertical-
sided gullies. At the same time, vegetation is relatively simple with
the low seasonal rainfall producing open, xerophilous communities
dominated by the savanna. Long drought and fires deplete the already
sparse vegetation and lead to considerable erosion during the short
wet season.

EDAPHIC FACTORS
GEOLOGIC.-Although climate is obviously responsible for the
altitudinal zonation of Komodo plants, regional zonation is determined
largely by geologic factors. The driest habitats are in lowlands where
the ground is impermeable and the slopes steep, e.g. the dense coralline
limestones near Ntodo Klea and the many igneous hilltops. These
areas are often sparsely covered with grass. Under these conditions
detritus collects only at the foot of most hills and along intermittent
stream courses. The permeable ground of these areas receives and
holds runoff from the hills and can support an open forest.
The western and southern parts of Komodo are built on an igneous
core comprised largely of fine-grained, slightly basic rocks. Flanking
this volcanic mass are lenses of tuffs, sandstones, and conglomerates
with intercalated limestones, sandy shales, and clays. The eastern part
is composed mainly of steeply tilted beds of coralline limestone; Padar,
Rintja, and Flores are very similar. Detailed geological accounts of the
area were given by Ehrat (1928a, b), Brouwer (1942a, b), Bemmelen
(1949), and Umbgrove (1949).
Soils of the Lesser Sunda Islands have not been studied. They have
little or no visible duff during most of the year. Leaf litter collects at
the base of some perennial shrubs but generally disappears quickly.

CLIMATIC FACTORS
Because the climate in this part of Indonesia is so poorly known, I
collected weather data comprising an almost complete annual cycle at
Loho Liang, Komodo. Techniques and additional details are given in
Auffenberg (in press).


1980











BULLETIN FLORIDA STATE MUSEUM


Vol. 25, No. 2


Telok Gililodo


C,

o0

C,


Slowi NUSA MIAru

TO O






0o
KM


,.tN-0,


.0


FIGURE 3.-Komodo place names used in text. Symbols used are: houses=villages,
tent=base camp.







AUFFENBERG: KOMODO HERPETOLOGY


PRECIPITATION.-Komodo lies in the driest part of Indonesia, with
an annual total rainfall of about 50 cm in the lowlands and a minimum
of 125 cm in the uplands. The heaviest precipitation occurs during the
monsoon (December to March). The low annual average is largely due
to the fact that no rain falls on the island during the June monsoon. As
a result only two amphibian species occur on the island. One
(Oreophryne jeffersoniana) is found in moist upland forests; the other
(Kaloula baleata) is adapted to lowland xeric conditions and spends
most of the year in aestivation. Aestivation is also common among
most snakes, including the fossorial ones.
GROUND WATER.-Free water is available in pools at all elevations
during the rainy season. The largest and longest-lasting are those at
high elevations, particularly along stream beds. Free water is rare near
the coast, except during the monsoon when small, wild boar wallows
often form temporary breeding ponds for Kaloula baleata and water
holes for large reptiles such as Varanus komodoensis.
TEMPERATURE.-Maximum air temperature was 430C, minimum
17C. Black bulb temperatures varied from 23C at night and on
cloudy days to 540C after the short rainy season and before the begin-
ning of the dry period. Average annual temperatures were 26.7C at
sea level and 22.0C at 400 m.
VEGETATIONAL FACTORS
Three major lowland plant formations can be recognized on Komodo
and adjacent islands: tropical deciduous (monsoon) forest, savanna,
and steppe. Iih addition, most islands in this area have small fringing
mangrove forests. Quasi-cloud forest is generally found in the moun-
tains.
QUASI-CLOUD FOREST.-On Komodo, shade producers are found only
in the pinnacle and ridge forests above 500 m. Moist and cool (the local
rural folk call the community tanah dingan, meaning cool ground),
they harbor a relictual flora of many endemic species. Floristically it is
characterized by moss-covered rocks, rattan, bamboo groves, and
many tree species generally absent at lower elevations (Terminalia zoll-
ingeri, Podocarpus nerifolia, Uvaria rufa, Ficus orupacea, Calophyllum
spectabile, Mischocarpus sundaicus, Colona kostermansiana, and
Glycosmis pentaphylla). Though this forest's distribution is only 6
km2 on Komodo, it is widespread on larger nearby islands. The typical
composition of this forest type at its lower limits is illustrated in
Figure 4 (G. Wasedoo, Flores).
The reptiles and amphibians most commonly encountered within
this forest are listed in Table 1. Species restricted to it are Sphenomor-
phus schlegeli Sphenomorphus striolatus, and Oreophryne jefferson-
iana


1980






















Borassus Colona Terminalia Tamarindus


Cladogynos Zizyphus Zizyphus
horsfieldi timorensis


Zizyphus
jujube


Philostigma


Lumnitzea Rhizophora


Jatropha Microcos


Abutilon Hypoestes


Tabernae-
montana


Annonia


Harrisonia


Murraya Callicarpa Azyma


Sterculia
foetida


Bredelia


Shouteria


Bambusa


Avicennia


ts










AUFFENBERG: KOMODO HERPETOLOGY


Base camp Zizyphus savanna, Komodo


Nggoer mangrove area


Kali Besar gallery forest, Komodo


- downslope G. Wasedoo, Flores, el. 400 m.


Monsoon forest, Loho Liang, Komodo


FIGURE 4.-Graphic transects through major forest communities in the study area,
showing major larger plants only. The Gunung Wasedoo transect represents the transi-
tional zone to quasi-cloud forest. A 2 m vertical bar is shown on the right side for scale.
Identification of symbols is given on the facing page.


1980













TABLE 1.-Ecological distribution of Komodo reptiles and amphibians (A=arboreal, T=terrestrial, F=fossorial, W=aquatic).

Quasi-cloud Monsoon Mangrove
Species forest forest Savanna Steppe forest Littoral

FROGS
Oreophryne jeffersoniana T
Kaloula baleata T T T
REPTILES
Cyrtodactylus darmandvillei A,T
Cyrtodactylus laevigatus A
Hemidactylus frenatus A A A
Lepidodactylus lugubris A A
Gekko gecko A A A
Cosymbotus platyurus A A
Hemiphyllodactylus typus A
Gehyra mutila A A A
Varanus komodoensis A(young,) AT T T A,T
Draco volans A A
Mabuya multifasciata T T
Sphenomorphus florensis A,T A,T
Sphenomorphus striolatus A,T
Sphenomorphus emigrans A,T A,T
Sphenomorphus schlegeli T
Emoia similis A,T T






O0





Cryptoblepharus boutonii T
Cryptoblepharus burden T
Dibamus novaeguineae F F
Typhlina polygrammica F
Typhlina bramina F F
Typhlops schmutzi F
Elaphe subradiata A,T A,T
Dendrelaphis pictus A,T A,T
Psammodynastes puluerulentus T
Chersydrus granulatus W 0
Cylindrophis opisthorhodus F
Cerberus rynchops W W
Lycodon aulicus T T 0
Boiga cynodon A A
Trimeresurus albolabris A A
Vipera russelli T T
Naja naja A,T A,T A,T
Crocodylus porosus W W
Chelonia mydas W(nesting)
Eretmochelys imbricata W(nesting)








BULLETIN FLORIDA STATE MUSEUM


SAVANNA FOREST.-Most of Komodo is covered with this associa-
tion (80 km2). Insolation is usually high, especially in the dry season
when the leaves have fallen. The understory is comprised mainly of
grasses (0.5-4.0 m high); the dominant grasses are Eulalia leschenaul-
tiana, Heteropogon contortus, Themeda frondosa, and T. triandra.
Common trees are Borassus filiformis, Tamarindus indicus, and
Schleichera oleosa.
Two facies of the savanna community are recognizable. One,
dominated by the lontar palm Borassus filiformis, develops on well
drained soils (Fig. 5). The dominant trees of the other are Zizyphus ju-
jube and scattered Tamarindus indicus (Fig. 6). The soil, sometimes
waterlogged for a few weeks in rainy season and often desiccated the
rest of year, is primarily a slightly halomorphic, azonal, alluvial type.
Savannas of both facies are distributed from sea level to about 500 m.
In general the herpetofauna of the savanna is much more varied
than that of the quasi-cloud forest. Kaloula baleata is the only frog
regularly found there. Typical reptiles include Emoia similis, Varanus
komodoensis, and Vipera russelli. A complete listing is given in Ta-
ble 1.


FIGURE 5.-Savana, Borassus facies in foreground, Loho Kalo, Komodo; steppe on the
background hills.


FIGURE 6.-Savanna, Zizyphus facies, Loho Kalo, Komodo.


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


TROPICAL DECIDUOUS (MONSOON) FOREST.-Shown in Figure 7, this
association is composed mainly of phreatophytes and some
xerophytes. Like the savanna forest, it is largely fire resistant, and
many of the trees are almost leafless in the dry season. Some dominant
trees, like Tamarindus indicus, occur in both associations, but the
monsoon forest canopy is higher and more continuous. The crowns
commonly touch and the shrub layer may be dense. Dominant trees in-
clude Tamarindus indicus, Sterculia foetida, Zizyphus horsfeldi, Mur-
raya paniculata, Schoutenia ovata, Schleichera oleosa, Cladogynos
orientalis, and Tabernaemontana floribunda. Tamarindus and Ster-
culia attain large size and have breast high diameters to 80 cm.
Tropical deciduous forest covers approximately 34 km2-less than
half of the total area covered by savanna (80 km2). It is found at all
elevations below approximately 500 m, i.e. the most continually dry,
hot climate region, locally known as tanah panas (hot ground). It may
form a gallery along stream courses and often extends in a band-like
strip along the base of hills. Typical reptiles include Varanus komo-
doensis, Sphenomorphus florensis, Trimeresurus albolabris, Den-
drelaphis pictus, Lycodon aulicus, all typhlopid species, and most of
the geckos.
DISTURBED AREAS.-Disturbance is usually by slash and burn
methods. Within 8 to 10 years, these plots are usually dominated by
Hypoestes malaccensis or Abutilon indicum. Other common shrubs in-
clude Solanum junghulnii, Callicarpa sappan, Azyma sarmentosa, and
Microcos paniculata. The most common trees are Annonia squamosa
and Tamarindus indicus; the common grasses are mainly Setaria ver-
ticillata, Brachiaria ramosa, and Digitaria adscendens. Common rep-
tile species include Dendrelaphis pictus and Naja naja.














FIGURE 7.-Monsoon forest clothing hill on right and along dry stream beds, Loho
Kalo; mixed savanna on drier hillsides.


1980








BULLETIN FLORIDA STATE MUSEUM


MANGROVE FORESTS.-Only two mangrove communities are found
on Komodo; both are flooded daily by tides. Saline tolerance sorts
them out, leading to horizontal zonation along the shoreline.
Rhizophora mucronata usually grows well within the tidal zone,
closest to the open water. The tangled roots form impenetrable
thickets. On many islands this association forms an almost unbroken
fringe along the entire coastline, but on Komodo it is restricted to the
east coast near Sabita.
Lumnitzera racemosa is dominant in a few areas behind the
Rhizophora zone. Though not tall, this species grows very dense, form-
ing large thickets that are occasionally broken by small Cyperus
javanicus marshes. The mangroves Ceriops tagal and Avicennia alba
are not widespread on Komodo, and the communities of which they are
the dominant species are not important habitats. However, they are
important on Flores. Varanus komodoensis commonly occurs in Lum-
nitzera swamps; less so in Rhizophora. Mangrove communities have
relatively few reptiles and no amphibians (Table 1). Only the highly
aquatic Cerberus rynchops and Chersydrus granulatus are found there
in any numbers. Both are most common in tidal creeks flowing
through the regularly flooded forests.
LITTORAL COMMUNITIES.-These communities comprise a hot, dry,
sandy or rocky, saline, and windy zone at and immediately behind the
beach. Dominant shrubs include Spinifex littoreus, Crotalaria retusa,
and Barleria prionitis; common trees are Pandanus sp., Jatropha cur-
cas, and Thespesia populnea. Relatively few reptiles and no amphi-
bians live here (Table 1), though Cryptoblepharus boutoni renschi is
commonly encountered. Cryptoblepharus burden is restricted to
rocky coastal areas.
STEPPE.-Tropical grassland or steppe is another important floristic
formation (40 km2), often grading into savanna or forming patches in
forested areas. The great variation in occurrence and composition of
grassland types has generated a vast literature, especially in respect to
their formation and continuance.
Part of the confusion and uncertainty regarding grassland forma-
tion in the Lesser Sundas is undoubtedly based on the often repeated
error that the steppe communities of these islands are composed large-
ly of alang-alang, Imperata cylindrica (Horst 1926, Lallemont 1929,
Pfeffer 1959, Kern 1968, et al). Actually, the extensive grass-covered
areas within the range of Varanus komodoensis are dominated by
Themeda and Chrysopogon. Imperata cylindrica normally occurs on
heavy soils, often seasonally waterlogged. On Komodo it is found only
in the moist gardens of the main village and in a few small stands in


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


the mountains (Hoogerwerf 1954); it is much more extensive in the
larger disturbed parts of western Flores.
The major grassland type in this area, usually dominated by
Themeda and Chrysopogon, covers extensive parts of the islands being
considered. Unlike Imperata stands, the Themeda-Chrysopogon com-
munity usually occurs on well drained soils and is particularly exten-
sive in hilly areas (Fig. 5), but it is also common on the valley floor.
None of the villagers remembered seeing any change in the character
or distribution of this grassland type, whether burned or not. Though
this community has relictual steppe and savanna species, no reptiles or
amphibians are restricted to it on Komodo.
ECOLOGICAL CORRELATES OF THE KOMODO HERPETOFAUNA.-Of the
38 species of reptiles and amphibians now known from Komodo and
small offshore islets, 4 are restricted to quasi-cloud forest, 5 to mon-
soon forest, none to steppe and savanna (unless they are combined, in
which case there are 2), 1 to mangrove, and 1 to littoral situations. The
largest number of species is found in monsoon forest (24), then savan-
na (14), quasi-cloud forest (12), littoral (7, plus 2 nesting species), and
finally steppe (4) and mangrove communities (4). A large proportion
(51%) of the Komodo fauna is comprised of species at least partly ar-
boreal in habits. The purely terrestrial species constitute only 19% of
the total herpetofauna; even the fossorial fauna is nearly as large with
12% total (Table 1). Comparable figures for other areas of high relief,
such as the habitats in the State of Michoacan, Mexico (after
Duellman 1965), show that, based on the distribution of the 166
Michoacan species, the terrestrial species (39-72% of fauna in each
area sampled) are nearly twice as common as the arboreal ones (5-25%).
Essentially the same pattern obtains in forested areas of low relief, as
in southern Florida where the arboreal herpetofauna makes up only
25% of the total (80 species). The preponderance of arboreality in the
Komodo fauna remains unknown, but may be related to the selection
processes concerned with waif dispersal.
Inger (1966) listed Kaloula baleata, Rana limnocharis, Rana can-
crivora, Polypedates leucomystax, Hemidactylus frenatus, Cosym-
botus platyurus, Mabuya multifasciata, Typhlina bramina, and
Gehyra mutilata as commensal with man, being found mainly in large
man-made clearings, towns, and cultivated land. Of these, only
Kaloula baleata is widespread on Komodo and is here considered part
of the native xeric-adapted community of the island. On Komodo,
Mabuya multifasciata is apparently restricted to higher elevations and
is best considered part of the mesic fauna. Of those listed above, the re-
maining ones are found only at the coast and could have been
transported to Komodo and neighboring islands by man.


1980








BULLETIN FLORIDA STATE MUSEUM


Kaloula baleata and Oreophryne jeffersoniana are almost entirely
terrestrial; the aquatic amphibians (Rana species) found on nearby
Sumba, Flores, and Sumbawa are absent here. Similarly the simple,
xerophilous lowland flora probably also limits the occurrence of ar-
boreal species, such as Polypedates leucomystax, which occupies the
ecologically similar parts of western Flores, but only in gallery forest
along streams. The absence of Python molurus on Komodo might be
due as much to the presence of the predacious Varanus komodoensis as
to the general absence of water.
Most of Komodo below 500 m is savanna. Thus, those species con-
fined to lower elevations are either adapted to fire and/or xeric condi-
tions or are closely associated with the activities of man. The small
fauna restricted to the area above this elevation is adapted to more
mesic conditions and, on Komodo, is comprised of only Oreophrynejef-
fersoniana and Sphenomorphus schlegeli. In general, the lowland
fauna is considerably more ecologically tolerant, being, in general,
distributed over much larger areas. Very few forms are endemic. Fur-
thermore, the ecological tolerance illustrated by this fauna undoubted-
ly increased the probability for successful interisland dispersal in the
past.



SYSTEMATICS
CLASS AMPHIBIA
ORDER SALIENTIA
FAMILY MICROHYLIDAE
Kaloula baleata baleata (MULLER)
Figure 8
Bombinator baleata Miiller 1836:96 (Type locality: Lewie Gadja, Java).
Kaloula baleata Gunther 1858:235.
Callula baleata Guinther 1864:436.
?Bufo biporcatus Elbert 1912:101.
SPECIMENS EXAMINED.-(23) UF 32580-83, 36164, Denpasar, Bali, 10 m; UF
29247-58, 29260, Loho Liang, Komodo, 5 m; UF 32584, 32586-87, Nggoer, Mangarrai
District, Flores, 12 m; UF 32585, Djarek, Mangarrai District, Flores, 120 m.
DESCRIPTION.-Small or moderate-sized frogs, Komodo adults
45-60 mm SVL; stout; head broader than long; short snout; canthus
distinct, usually rounded; tympanum usually hidden, V1-% diameter of
eye; supratympanic fold present; transverse occipital fold in most
adults; dorsal skin thick, with large mucous glands, variable in tex-
ture; two metatarsal tubercles, inner one well developed, less than
length of first toe, compressed; outer one without a sharp edge; finger


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


tips much wider than penultimate segments; no metacarpal tubercles;
usually three subarticular tubercles on fourth toe.
Color of posterior surface of thigh mottled; dorsum mottled, brown
or purplish-brown, with yellow or red blotches in axilla and on thigh.
Males with dark gular pigmentation, rest of venter cream, sometimes
mottled with brown (Fig. 8).


FIGURE 8.-Dorsal (above) and ventral (below) views of Kaloula baleata baleata, Loho
Liang, Komodo. Left, male (UF 29250); right, female (UF 29252).








BULLETIN FLORIDA STATE MUSEUM


DISTRIBUTION AND HABITAT.-A widely distributed form found
from the Malay Peninsula and the Philippines southeastward to
Flores. In the Komodo area it is known from that island (Dunn 1928),
Flores and Sumbawa (Mertens 1930), and Sumba (Kampen and
Brongersma 1931; Forcart 1949). Mertens (1930) reported that in the
Lesser Sundas it occurs up to 800 m, though all our Komodo and
Flores specimens were collected near sea level. Dunn (1928) collected
two specimens from Komodo at 650 m. Parker (1934) said the species
occurred up to 1525 m.
In savanna habitats on Komodo, the frogs remain in their burrows
for much of the last half of the dry season (July to November). During
the wet season (December through June) and early part of the dry
season the frogs tend to leave their burrows every night, often climb-
ing onto grass tussocks and trees, where they apparently feed largely
on ants. We found them on banana, tamarind, and Zizyphus trees.
Mertens (1929a, 1931) reported seeing them as high as 2.5 m on trees
on Bali and Java; Taylor (1921) reported that he found them in the leaf
axils of Pandanus as high as 7 m, as well as in burrows in the ground.
With the first monsoon rain in December, these frogs call from their
burrows at night.
REPRODUCTION.-When wild boar wallows become filled with
several inches of rain water, individuals quickly gather to reproduce.
Breeding choruses on Komodo were recorded from 16 January to 8
March from wild boar wallows near the coast and in pools along inter-
mittent streams as high as 400 m. The voice is a sonorous chuck-
chuck-chuck.
Remarks.-Dunn (1928) reported finding fragments of this frog in
the stomachs of two Trimeresurus albolabris. Mertens' (1930) state-
ment that Dunn reported K. pulchra from Komodo is incorrect, for the
latter stated that it occurs on Flores; K. baleata on Komodo.
FAMILY RANIDAE
Oreophryne jeffersoniana DUNN
Figure 9
Oreophryne monticola Mertens 1927c:236 (part).
Oreophryne jeffersoniana Dunn 1928:3 (Type locality: Komodo).
Oreophryne darewskyi Mertens 1965:189 (Type locality: Rintja).
SPECIMENS EXAMINED.-(15) AMNH 24530-31 holotypee and paratype), MCZ 14861
(paratype), Komodo; MCZ 35172, Ruteng, Flores, ca 1200 m; UF 40357-60, 1 km E.
Djarek, Flores, 60 m; UF 40361-63, 1 km E Nggoer, Flores, 30 m; UF 40364-65, Tjereng,
Flores, 800 m; UF 40366-67, Ruteng Flores, 1200 m; UF 40368, Potjo Dedeng,
Nunung, Flores, 1200 m; UF 40670, Loho Lavi, Komodo, 12 m. Also 2 Oreophryne
celebensis (MCZ 2808, 26091), 10 0. monticola (AMNH 23705-6, 24814-15; MCZ 2897,
10193-94, 14862-64) from Lombok and 0. rookmaakeri (MCZ 15816, paratype), Rana
Mesa, Mangarrai District, Flores, 1200 m.


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


Taxonomic Remarks.-The Komodo material of 0. jeffersoniana is
closely related to at least some populations from nearby islands that
currently are regarded as distinct species. These are 0. rookmaakeri
(type locality: Rana Mesa, Mangarrai District, Flores), 0. darewskyi
(Rintja), 0. variabilis (Celebes), and 0. monticola (Bali and Lombok).
Mertens (1930) recognized this strong resemblance and set all the
species (except 0. darewskyi, which was described later) apart as a
distinct group, comprised of isolated, insular populations he believed
sufficiently distinct to warrant specific status. Mertens' discovery and
description of 0. darewskyi in 1964 tended to obliterate the distinc-
tiveness of the previously described populations. He, in fact, sug-
gested that future work might show that this group represents a single
Rassenkreis. The material collected during my study suggests that
some of the characters previously believed specifically distinct (1) are
ontogenetically variable (hind leg length), (2) vary individually, on-
togenetically, and/or sexually (dorsal warts), or (3) show discordant
variation within the entire complex (dorsal color and pattern). I find no
basis for Merten's distinction of "plump" body for 0. rookmaakeri and
"not so" for 0. darewskyi.
Other characters, such as the color pattern of the belly and
posterior thigh, the relative size of finger dilations, and proportional
lengths of the fingers, seem farily constant within populations and in
my opinion serve to distinguish two groups, which at this time I prefer
to consider as being specifically distinct. The most diagnostic of these
characters is the color of the posterior part of the thigh. In 0. jefferson-
iana this surface bears a dark brown horizontal band edged with white
above and sharply demarcated from the much lighter ventral surface
(Fig. 9, not as evident in old preserved specimens). In 0. monticola and
0. rookmaakeri this area is (1) lighter in color and (2) not edged above
with white. Furthermore, the SVL for both sexes of 0. jeffersoniana is
less (X = 15.1 mm) than in 0. monticola (X = 20.8) or 0. rookmaakeri
(30.0 in the type); the dorsal area and thighs are usually quite warty in



W ." .7..
.. ...._,I._.W. ...... ..._.. .. ... --.... ..



A B
FIGURE 9.-Oreophryne jeffersoniana posterior thigh surface. (A) UF 40670, female,
Loho Lavi, Komodo; and (B) UF 40365, male, Tjereng, Flores.


1980








BULLETIN FLORIDA STATE MUSEUM


the males and females of 0. jeffersoniana, only in males (and then rare-
ly) of 0. monticola, and smooth in the type (9) of 0. rookmaakeri; the
fingers are proportionately shorter in 0. jeffersoniana with digit 1
usually barely reaching the base of digit 2 and extending up onto digit
2 in the other species; the finger pads are proportionately smaller in O.
jeffersoniana (Fig. 10) and never found on digit 1, rarely on 2 and 4,
whereas they are always on 2 and 4 in the others and often on 1; the
ventral body surface and particularly the chin are heavily mottled in
0. jeffersoniana, less in the others and sometimes uniform white; the
dorsal color and pattern (although variable in all three) is generally
darker in 0. jeffersoniana, with fewer examples of longitudinal band-
ing (a common pattern in 0. monticola); a postorbital dark spot or
short stripe is usually present in 0. jeffersoniana and absent in 0. mon-
ticola; the posterior surface of the humeral portion of the arm is heavi-
ly marbled in 0. jeffersoniana and more or less uniform in the others'.











BC D








2mm
A E F

FIGURE 10.-(A) ventral surface Oreophryne jeffersoniana, UF 40365, Tjereng, Flores;
(B) left front foot, 0, jeffersoniana, paratype, MCZ 14861, Komodo; (C) left front foot,
holotype, 0. darewski, Senckenberg Mus. ZIL-4126 Rintja; (D) left front foot, 0. jeffer-
soniana, UF 40362, Nggoer, Flores; (E) left front foot, 0. rookmaakeri, paratype, MCZ
15816, Flores; and (F) right front foot, 0. monticola, MCZ 10193, Lombok.
'Mertens (1965) stated that additional material from Bali (= monticola, fide Mertens
1930) may show that this population is to be considered distinct. He suggested that
same possibility for the single Sumbawa specimen (= jeffersoniana here and in Mertens
1930).


Vol. 25, No. 2







AUFFENBERG: KOMODO HERPETOLOGY


Thus 0. jeffersoniana (and its synonym 0. darewskyi) represent one
species group, with 0. monticola and the closely related O.
rookmaakeri in the other Sundanese species group of these frogs. My
examination of both 0. variabilis and 0. celebensis suggest that they
are not so closely related to the Lesser Sunda species as had been
previously believed.
DESCRIPTION.-A very small frog (SVL 12.0-21.0 mm; X = 15.1
mm) with a large, oval, entire tongue; short, rounded snout with round-
ed canthus; interorbital space broader than the upper eyelid; tym-
panum feebly distinct, % diameter of eye; finger and toe discs feebly
developed, with first finger much shorter than second and barely
reaching base of second when adpressed; toes free, no subarticular
tubercles, but very feeble inner metatarsal tubercle; heel reaching
posterior border of eye; skin above with few to many small to relative-
ly large warts (few and smaller in young), often with a median line of
elongate warts from tip of snout to cloaca and another from eye to
more than halfway to groin; belly finely granular; dorsal color tan to
gray-brown, with warts often outlined in black; a fine, light middorsal
line and/or a darker triangular mark, or two small dark spots on
scapular region, or nearly uniform, or spotted; laterally below glan-
dular line usually darker, often brown; chin heavily mottled (50-70%
dark) with dark brown to black on dirty white ground color (Mertens'
Sumbawa specimen dirty white below); thighs indistinctly marbled or
barred above, with brown posterior surface, sharply set off from dorsal
color by narrow white line (Fig. 10). Dunn (1928) reported a car-
tilaginous procoracoid running from base of clavicle to scapula.
DISTRIBUTION AND HABITAT.-Previously known only from
Komodo (Dunn 1928) and Sumbawa (Mertens 1930, probably
originating from the Tamboro Volcano region); specimens collected
during the NYZS expedition show this species also occurs in western
Flores. In this paper, the Rintja specimens Mertens (1965) described
as a new species, 0. darewskyi, are considered as belonging to the
species 0. jeffersoniana because of the similarly of color pattern, toe
lengths, disc sizes, and adult SVL lengths.
Dunn (1928) described the Komodo habitat as heavily forested
rocky peaks, where individuals were found hopping about on the moss-
covered rocks and surface debris of the mesic forest floor. Other
populations of the same species were described as occurring in similar
mesic montane habitats. During this study adults were collected on
Flores from beneath stones and rotting piles of vegetation. Juveniles
(recently transformed ?) were found hopping about the edges of
shallow rocky pools. Adult males were found calling (8 June) in a reedy
pool formed by a small hillside spring.


1980








BULLETIN FLORIDA STATE MUSEUM


As all previous specimens were taken above 600 m elevation, 0. jef-
fersoniana was considered a montane form (Dunn 1928; Mertens 1930,
1964), but our single Komodo specimen and some of those from Flores
were taken between 8 and 100 m above sea level in and/or near small
springs and seeps at the foot of mountains. The better known 0. mon-
ticola is not known to occur below 600 m, nor is the related O.
rookmaakeri (900-1200 m, this paper).
REMARKS.-Remains of one specimen of 0. jeffersoniana were
found in the stomach of a Psammodynastes pulverulentus taken in a
mesic montane forest on Gunung Ara, Komodo.
CLASS REPTILIA
ORDER TESTUDINATA
FAMILY CHELONIIDAE
Chelonia mydas japonica (THUNBERG)
Figure 11
Testudo japonica Thunberg 1789:178 (Type locality: Japan).
Chelone mydas Weber 1890:161.
Chelone agassizii Burden 1927:94.
SPECIMENS EXAMINED.-(7) UF 29263-69, Pulau Longo, off the east coast of
Komodo.


FIGURE 1l.-Hatchling Chelonia mydas japonica, UF 29263, Pulau Longgo, off
Komodo.


Vol. 25, No. 2



























II -- :




I' .. I I L'





I - i.; ..


5'


7-


SI ..


. ,* ..


"- ,, II '. L.l ,, ii


i

L


'io~z








BULLETIN FLORIDA STATE MUSEUM


REMARKS,-Previously reported in the Komodo area from Sumba
(Mertens 1928a, Forcart 1949) and Sumbawa and Flores (Mertens
1930). The two Komodo specimens here reported were recently
hatched, collected from shallow water off a sandy beach on 12
February. A third hatchling (not preserved) was found dead on the
same beach near Nanenepi on 29 April. Remains of an adult were found
in the fresh droppings of Varanus komodoensis on Pulau Padar on 16
January. These data suggest that in the Komodo area most nesting
probably takes place during the monsoon months of January and
February. Hatching takes place in the late monsoon months of March
and April. Varanus komodoensis regularly excavate the nests and
eat the eggs of this species on Komodo beaches. Komodo villagers do
not eat the flesh, but only the eggs of the turtles. Turtle fishermen
from Bali regularly hunt for all species of marine turtles in the bays
and backwaters around Komodo. I saw one adult they slaughtered
that contained crab remains.

ORDER CROCODYLIA
FAMILY CROCODYLIDAE
Crocodylus porosus SCHNEIDER
Crocodylus porosus Schneider 1801:159 (no type locality designated).
SPECIMENS EXAMINED.-(2) UF 34888, left dentary, Loho Lavi, Komodo; UF 36200,
tooth of very large specimen, Loho Lavi, Komodo.
DISTRIBUTION AND HABITAT.-Though widely distributed over
much of coastal southeastern Asia, the saltwater crocodile has a spot-
ty distribution in the Sunda Islands. Mertens (1930) reported it from
Timor, Lombok, and Flores, Zollinger (1850) from Sumbawa, and For-
cart (1949) from Sumba. Darevsky failed to mention it from Komodo,
although Broughton (1936) referred to a single small specimen in her
account of Varanus komodoensis there. While occasionally seen along
the northern coast of Flores, it is apparently extirpated on
Komodo-chiefly by hide hunting. A large crocodile ate a school
teacher in Reo, Flores, in 1967. Darevsky (1964a) reported specimens
from eastern Rintja and cited native accounts of them in western
Flores. My extensive search in the same part of Flores in 1972 failed to
disclose any in appropriate habitats, and local villagers claimed they
were all killed by hide hunters several years previously. In the Lesser
Sunday Islands Crocodylus porosus is usually found in estuaries and
embayments, commonly in mangrove swamps. Where not persecuted,
they are often seen basking on open sand beaches. On Komodo they
have previously been seen in Soro Masalong, the western arm of the
large bay on the southeast coast, as well as the eastern mangrove coast
from Sabita southward to Tandjung Kunning. Broughton (1936)


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


reported a 2 m specimen eating carrion at one of her baiting stations
more than 400 m from the nearest water.

ORDER SQUAMATA
SUBORDER SAURIA
FAMILY GEKKONIDAE
Cyrtodactylus darmandvillei (WEBER)
Figure 13
Gymnodactylus d'armandvillei Weber 1890:160 (Type locality: Sikka, Flores).
Gymnodactylus defossei Dunn 1927a:1 (Type locality: Komodo, el 650 m).
Cyrtodactylus darmandvillei Darevsky 1964a:564.
SPECIMENS EXAMINED.-(6) UF 28913-17, 30172, Kali Besar, Loho Liang, Komodo;
UF 32579, Bara, Flores, 200 m.
DESCRIPTION.-Head broad, large; eyes large, but smaller than the
distance between the eye and the ear opening; ear opening vertically
oval, 2/3 diameter of eye; dorsal surface of body with small scales,











A

















FIGURE 13.-Crytodactylus darmandvillei, UF 30172, Komodo. (A) lateral view of head;
(B) inferior surface of the left front foot; and (C) dorsal surface.







BULLETIN FLORIDA STATE MUSEUM


among which are large, triangular, keeled tubercles, usually
longitudinally arranged on top and sides of body and tail; ventral
scales small, imbricate; no femoral or preanal pores; tail cylindrical,
with enlarged tubercles above, wide scales ventrally.
Ground color grayish-brown with five darker, somewhat wavy
crossbands, often lighter, or even absent along middorsal line, none as
light as Weber's type from Flores; postocular dark stripe turns medial-
ly over ear to form a posteriorly directed V; tail tan to gray, with
darker crossbands; juvenile pattern similar but brighter. Adult SVL
86 mm, tail 113 mm; Komodo hatchling SVL 27 mm, tail 35 mm.
DISTRIBUTION AND HABITAT.-It occurs on Flores, Pulau Kalao
(off north coast of Flores), Sumbawa (Mertens 1930), and Komodo
(Dunn 1927c, Darevsky 1964b), and is probably closely related to Cyr-
todactylus marmoratus (from Lombok westward). It is found from sea
level to 650 m elevation.
Mertens' (1930) Sumbawa specimen was taken on a large tree, and
Darevsky (1964a) reported that his specimens were taken under the
loose bark of dead trees. Our single specimen from Flores was collected
under a large granite boulder on a grassy hillside. Our Komodo
specimens were taken at night, when they were very active and often
scurried away when the headlight beam first struck them, unlike
Gekko gecko, which usually became inactive in the bright light. We
saw many specimens on the verticle clay banks of the dry creek beds,
but they were so active, and the banks were so extensive that we
caught very few.
REPRODUCTION.-Darevsky (1964a) reported finding a single egg of
this species in a Varanua komodoensis burrow; our expedition
members found them there commonly, always on the surface and
usually near the side walls.
Four males were examined; the two less than 64 mm SVL were im-
mature. In the two mature, October individuals the testes were turgid
but still subcylindrical (length 4.5/3.8, 7.2/6.8) and the vasa deferentia
were turgid and convoluted. The remaining, immature males had flat,
smooth, translucent vasa deferentia and flaccid, laterally compressed
testes. Both females (SVL 51.5-52 mm) were immature showing very
little follicular development, with follicles less that 1 mm in diameter.
FoOD.-Spiders and various small insects (mainly termites and bee-
tle larvae) are present in half of the specimens.
PARASITES.-Unidentified, in colon of one-third of the specimens.
Cyrtodactylus laevigatus DAREVSKY
Several specimens of this species collected in western Flores are
identical to one another and clearly separable from the available


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


Komodo material. On this basis they are described below as a new
subspecies.
Cyrtodactylus laevigatus laevigatus DAREVSKY
Figure 14
Cyrtodactylus laevigatus Darevsky 1964c:171 (Type locality: Komodo, 600-700 m)
SPECIMENS EXAMINED.-(4) Paratype, Museum Zoologicum Bogoriense 979,
Komodo; UF 28910-12, Loho Liang, Komodo, 20 m.
DESCRIPTION.-Head medium, depressed; snout longer than the
diameter of the eye, which is less than its distance from the ear; ear
opening oval, oblique, about one-fourth the eye diameter. Head and
body covered with granular scales, with small flat tubercles
(sometimes keeled) beginning at occiput and scattered over back;
rostral subtriangular, about twice as broad as high, with a median cleft
above; nostril between the rostral, first labial, and three small








IC
K. c.







D



A B \ B







FIGURE 14.-Cyrtodactylus laevigatus: (A C and E) C.1. laevigatus, UF 28910, Loho
Liang, Komodo; (B and D) C. 1. uniforms, holotype, UF 32596, Nggoer, Flores.


1980








BULLETIN FLORIDA STATE MUSEUM


postnasals; 11-13 supralabials, 8-10 infralabials; triangular mental,
two pairs of chinshields of which median pair are the largest and touch
mental; ventral scales small, smooth, cycloid, imbricate, 32-34 at the
middle of the belly; subdigital lamellae strongly dilated only on the
basal half, 12-15 under fourth digit. No femoral or preanal pores. Tail
round, tapered, covered with small flat scales, larger below, and small
tubercles at the base; limbs moderately long, digits depressed at the
base, strongly compressed at the end phalanges.
Color tan dorsally with darker irregular spots, often forming short
transverse bars that are often interrupted along the vertebral line.
Head with small, oblong, irregular brown spots. A dark brown tem-
poral stripe extends from end of snout through eye, over ear, and
medially to join first transverse dorsal bar. Infralabials mottled; a
dark brown stripe from nostril to upper anterior corner of eye. Larger
tubercles of back often whitish. Lower surface whitish. SVL of females
43-44.8 mm, tail 39.3 mm; males 38 mm, 27.5 mm. My counts of 13 and
14 infralabials and 15 and 16 supralabials in this small series, rather
than 10-13 and 8-10, respectively, by Darevsky for his series, is un-
doubtedly due to the fact that both the upper and lower labials become
considerably reduced in size at the corners of the mouth, and there is
some question as to which scales should be included in the count and
which excluded. We counted all the scales to the angle of the mouth
and can only presume that Darevsky counted only the enlarged ones.
DISTRIBUTION AND HABITAT. -Unfortunately the elevation at
which Darevsky's specimens were obtained is questionable. In his
species description (1964b), he stated that the type and three
paratypes were collected from under stones on the edge of the jungle
between 600 and 700 m. However, in another paper (1964a), he
reported that the adult specimens were found on tree stumps during
the night and the juveniles under stones in dry jungle. Furthermore,
the juveniles were said to have been collected at 100 m and the adults
at 200 m. Our three specimens were taken during the daytime from
under the bark of small, dead Tamarindus trees in Loho Liang, approx-
imately 20 m above sea level.
REPRODUCTION.-Both females (UF 28910, 28192) were mature
when collected in September 1969. In one, the left ovary contained one
large, yolked follicle (6.7 X 5.2 mm) and two small, mainly translucent
follicles, of which the largest was 2.8 mm in diameter. The right ovary
was more anterior than the left and also contained one large, yolked
follicle, approximately 5.6 mm in diameter. The oviducts were large,
thick-walled, and considerably convoluted. The second specimen (UF


Vol. 25, No. 2







AUFFENBERG: KOMODO HERPETOLOGY


28912) was similar, with the left ovary posterior containing one large
yolked follicle (8.4 X 4.8 mm) and three small, translucent follicles of
approximately 2.5 mm in diameter. The right ovary was more anterior
and contained one large, yolked follicle (8.9 X 4.5 mm). In addition,
three small, transparent follicles, of which the largest was
approximately 2.4 mm in diameter, were present. The oviducts were
identical to UF 28910. The only male (UF 28911) was collected in late
July and was subadult. The testes were less than 1 mm in diarneter,
very flaccid, and almost translucent. The vasa deferentia were small,
flat, and thin-walled.
Eggs and hatchlings are found under loose bark from July through
September. There is some evidence that males are mature at the begin-
ning of their second year.
FooD.-Small beetles (mainly of the family Elateridae) and their
larvae.
PARASITES.-Eggs of unknown parasites were found in the in-
testines of one of the three specimens collected.

Cyrtodactylus laevigatus uniforms new subspecies
Figure 14
DIAGNOSIS.-Scutellation as in nominate subspecies, but dorsal col-
or darker, uniform, and not brightly marked with dark brown.
HOLOTYPE.-UF 32596, adult female, Nggoer, Mangarrai District,
Flores, 5 m, taken from beneath loose bark of standing tree.
DESCRIPTION OF TYPE.-Head medium, moderately depressed;
snout longer than eye diameter, which is less than its distance from the
ear; ear opening oval, obligue. Head and body covered with small
granular scales, among which are interspersed larger tubercles, keeled
above tail, nostril bordered by rostral, first supralabial, and two small
postnasals. Supralabials 10 and infralabials 8, mental trangular, one
pair of large chin shields, in contact behind mental scale; ventral scales
small, smooth, cycloid, imbricate, in 32 rows across belly. no femoral
or preanal pores; tail round, tapering with flat scales that are larger
ventrally. Limbs moderate. Digits depressed distally, strongly com-
pressed distally. Subdigital lamellae 10.
Generally brownish-gray above with darker brown only on labials,
and in faint head markings, as in nominate subspecies. Venter
yellowish-white, SVL 47 mm, tail broken.
PARATYPE.-UF 32597, Nggoer, Mangarrai District, Flores; as in
holotype, except 14 subdigital lamellae, 11' supralabials, 9 infralabials.
The subspecies is named uniforms because it lacks any body pattern.








BULLETIN FLORIDA STATE MUSEUM


Gekko gecko gecko (LINNe)
Lacerta gecko Linn6 1758:205 (Type locality: "Indes").
Platydactylus guttatus von Martens 1876:374.
Gecko verticillatus Weber 1890:160.
Gekko gecko Barbour 1912:31.
SPECIMENS EXAMINED.-(29) UF 3307-8, Suwang, Bali; UF 33022, Labuanbadjo,
Mangarrai District, Flores; UF 33021, 33023, 30091, 39699, Nggoer, Mangarrai
District, Flores; UF 33015-17, Nangelele, Flpres; UF 33013-14, 33017, Orang (Baneng),
Flores, 610 m; UF 33018, 33019, Pota, Flores, ca 10 m; UF'33020, Bara, Flores, 200 m;
UF 28890-97, 28686, 33009-12, Loho Liang, Komodo, 0-30 m.
DISTRIBUTION AND HABITAT.-A very widely distributed form, ap-
parently common throughout the Sunda Islands; in the Komodo area
previously reported from Lombok, Sumbawa, Pulau Endah (off Flores
south coast), Flores, Komodo, Padar, Celebes, and Sumba (see Mertens
1930, Forcart 1949, Darevsky and Kadarsan 1964, and others).
Mertens (1930) stated that it does not extend higher than 800 m, which
fits with our obversations on Komodo.
It is usually found in isolated trees with large knot-holes and other
cavities. On Flores, expedition members found as many as five in one
rotten tree. In most mesic places where various ferns and orchids cover
the surface, specimens may find refuge in the epiphytic growth;
sometimes under loose bark, particularly on dead tamarind trees.
Prime habitat on Komodo and western Flores was the savanna
biotope, especially that dominated by Zizyphus jujube trees. Density
of adult specimens in this habitat type at the Base Camp in Loho
Liang, Komodo, was calculated as about 72/km2 (on the basis of both
calling individuals and the distribution and number of appropriate
dead and dying trees). Mertens (1930) reported it was sometimes found
on mangrove trees (species?). Expedition members found it on large
hollow specimens of the mangrove A vicennia alba on Flores, but never
on Rhizophora mucronata.
Individual geckos usually began calling near dusk and continued
intermittently until at least 0300 hr. Although they eat a great variety
of food, our observations at the Base Camp suggest that they eat
many Hemidactylus and smaller individuals of their own species,
sometimes almost the same size as the attacker, so that they may take
as much as 30 min to swallow their prey completely.
Hatchlings with a total length of 85-90 mm were observed to attack
and swallow small Hemidactylus frenatus successfully.
REPRODUCTION.-Examination of nine females, representing
several size classes, showed that sexual maturity was attained at a
minimum SVL of 115 mm, but most were not mature until about 120
mm. Shelled ova were found in the oviducts during January, June, and
July. Freshly laid eggs were found in March, June, August, and


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AUFFENBERG: KOMODO HERPETOLOGY


September, suggesting egg laying throughout the year, with the possi-
ble exception of the driest season (October to early December).
Hatchlings were seen in May and July. A maximum of only two eggs
are laid, apparently each separated by several days. Sometimes only
one egg is laid, the other oviduct being empty. The number of follicles
varied from five to seven (diameter from 1 to 11 mm). As many as two
stages of corpora lutea disintegration are usually discernible at any
one time (approximately 3.5 and 8.0 mm in diameter). Egg clusters
(1-5, X = 2.3) are often found under loose bark. Single females ap-
parently return to the same spot to lay their eggs. The new eggs are
glued to the wood, next to the old hatched or spoiled eggs.
Of nine males dissected, the smallest mature individual was 134
mm SVL; specimens 112-118 mm are all immature; all over 148 mm are
mature with turgid, cylindrical testes. The vasa deferentia were highly
convoluted, flat, and thin-walled. Average SVL of six mature females
is 132 mm, for six males 145.3 mm.
FooD.-Hemidactylus frenatus, Cosymbotes platyurus, spiders,
and miscellaneous insects (mainly beetle larvae and termites).
PARASITES.-The nematodes (after Pinnell and Schmidt 1977)
Pharyngodon kuntzi and Rhabdias spp. were found in the colon,
stomach, and small intestines of seven of the eight specimens
examined.
Gehyra mutilata (WIEGMANN)
Figure 15
Hemidactylus (Peropus) mutilatus Wiegmann 1834:235 (Type locality: Manila,
Philippines).
Peropus mutilata Gray 1845:159.
Gehyra mutilata Boulenger 1885:148.
SPECIMENS EXAMINED.-(9) UF 33051-57, Ruteng, Flores, 1200 m; UF 28888, 28896,
Loho Liang, Komodo, 10 m.
REMARKS.-A common gecko, often found in houses and very wide-
ly distributed. In the Komodo area, it is known from Flores (Mertens
1931), Sumba (Forcart 1949), Komodo (Dunn 1927c, this paper), and
Sumbawa (Mertens 1931). A juvenile (UF 33053) from Ruteng, Flores,
has distinct dorsal white spots that are faint in larger individuals from
the same locality. The Komodo specimens (UF 28888, 28896) collected
from the side of the Base Camp building, bear indistinct, wavy,
transverse bands.
Hemidactylus frenatus DUMERIL AND BIBRON
Figure 15
Hemidactylus frenatus Dumeril and Bibron 1836:366 (Type locality: restricted by
Loveridge 1947:127 to Java).







BULLETIN FLORIDA STATE MUSEUM


SPECIMENS EXAMINED.-(54) UF 33027-29, 33033-35, Bara, Flores, 200 m; UF 32595,
33036-46, Nggoer, Mangarrai District, Flores, 10 m; UF 33030-31, Labuanbadjo,
Flores, 10 m; UF 33032, Nunang, Flores, 600 m; UF 28879-87, 28889-95, 28897-99,
28900, 28902-5, 28986-7, all from Loho Liang, Komodo, 0-30 m; UF 33026, Padar
Island; UF 33024, Jakarta, Java.
REPRODUCTION.-I examined 18 females in varying stages of
maturity. These suggested that females become mature at an SVL of
about 44 mm (average 46.3). Follicular development clearly shows only
one egg is laid at a time. Females with eggs ready to be laid were cap-
tured in July and October. Newly laid eggs (N = 6) have a diameter of
6.4 7.9 mm, and usually are found in rotten knot holes or under loose
bark on the shaded side of Tamarindus trees. Of 19 males examined, 10
were considered mature on the basis of turgid, highly convoluted, and
thick-walled testes. The average SVL of mature males is 46.7 mm, and
maturity is reached at an SVL length of about 42 mm.
FooD.-Gecko skin found in several stomachs suggest they regular-
ly eat parts of their shed skins. Food remains in the stomachs were
comprised of ants, beetle larvae, spiders, and mainly termites.
PARASITES.-Pharyngodonid nematodes were found in the colon of
8 out of 23 specimens examined. UF 28881 has an unidentified skin
parasite on several parts of its neck and body.
REMARKS.-A common lizard over a large part of the tropics, par-
ticularly near the coast, commonly transported accidentally on native
boats. Darevsky (1964a) reported finding them on Komodo outrigger
praus. In the Komodo region the species is found on all the islands.
Vertical distribution is from sea level to 500 m (Mertens 1931). In near-
by areas it is known from Sumba (Forcart 1949). Mertens (1930)
stressed the wide range in coloration of individuals from the night to
daylight conditions. Maximum SVL on Komodo is 54.3 mm.
Prey of Gekko gecko.

Hemidactylus garnoti DUMERIL AND BIBRON
Figure 15
Hemidactylus garnoti Dumeril and Bibron 1836:368.
SPECIMENS EXAMINED.-(1) UF 32571, Djarek, Mangarrai District, Flores, 200 m.
REMARKS.-This species is often a commensal of man, and the
single specimen reported here was collected from the wall of a village
house. It has previously been reported from Flores and Sumbawa by
Mertens (1930). We found none on Komodo.
Cosymbotes platyurus (SCHNEIDER)
Figure 15
Stellio platyurus Schneider 1792:20 (Type locality: not designated).
Hemidactylus (Stellio) platyurus Wiegmann (1834:238.
Cosymbotus platyurus Stejneger 1907:178.


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


Platyurus platyurus Smith 1935:102.
SPECIMENS EXAMINED.-(16) UF 28901, 28906-09, Loho Liang, Komodo, 0-32 m; UF
33048-50, 33086-87, Denpasar, Bali; UF 33047, Jakarta, Java; UF 32588, 32590-93, Ng-
goer, Flores, 12 m; UF 32589, Labuanbadjo, Flores.
DESCRIPTION.-Snout longer than the distance between the eye and
the ear opening, 1'/2 times the diameter of the eye; ear opening small,
oval, oblique. Rostral trapezoidal, with median cleft above, nostril
borders rostral, first supralabial, and 3 postnasals; 9-11 supralabials
and 7-8 infralabials; mental large; two pair of chin shields, of which the
median pair largest, in contact with each other. Body somewhat
depressed, covered aboe with uniform small granules. A small flap of
skin extends from axilla to groin, and another along posterior edge of
hindlimb (particularly in adults). Ventral scales cycloid, imbricate,
smooth. Males with an uninterrupted series of 34-36 femoral pores.
Tail depressed, with a sharply denticulated lateral fringe of scales;
uniform dorsally granular scales and ventrally with a median series of
transversely dilated plates. Limbs moderate, depressed; digits strong-
ly dilated, partially webbed, 7-9 lamellae under the fourth toe. Gray to
grayish-brown above, marked with irregular spots or stripes dorsally,
darker gray, brown, or black; generally a dark stripe from eye to
shoulder. White ventrally. Young much more brightly colored. SVL
about 60 mm in adults, tail 65 mm.










A B







D E

FIGURE 15.-Lower surfaces of right front feet of: (A) Cosymbotes platyurus (UF
28907); (B) Gehyra mutilata (UF 33055); (C) Hemidactylus frenatus (UF 33027); (D)
Hemiphyllodactylus typus pallidus, holotype (UF 28985); and (E) Hemidactylus garnoti
(UF 32571).








BULLETIN FLORIDA STATE MUSEUM


DISTRIBUTION AND HABITAT.-Widely distributed in Southeast
Asia and extending throughout the archipelago. It is often the most
common lizard in both urban and agrarian habitats. Komodo and
Flores specimens were taken from large trees near sea level. Mertens
(1931) reported that it is not found above 700 m. Although he failed to
find it in Bali, it is common near sea level in several places along the
southeast coast. It is one of several similar-sounding geckos that
regularly call in the early evening; the sound produced is "tchack-
tchack-tchack."
REPRODUCTION.-Two eggs (7.2-822 mm diameter) are usually laid
in moist, rotten wood; my records are for January and March. Average
SVL of mature females (N = 8) is 52 mm (49-55); average SVL mature
males (N = 5) is 51 mm (48-54).
FooD.-Spiders and winged and wingless ants.
PARASITES.-Pharyngodonid nematodes were found in the colon of
2 out of 10 specimens examined.
REMARKS.-Sometimes eaten by Gekko gecko.
Hemiphyllodactylus typus BLEEKER
Figures 15, 16
Hemiphyllodactylus typus Bleeker 1860:327 (Type locality: Gunung Parang, Java).
Spathodactylus mutilatus Gunther 1872:594.
Spathoscalabotes mutilatus Boulenger 1885:157.
Previously known only from Sumbawa westward, this rare little
gecko is reported from Komodo for the first time. The distinctive color
and lack of pattern suggests this population be recognized as a
separate subspecies (Fig. 16).
Hemiphyllodactylus typus pallidus new subspecies
DIAGNOSIS.-Adults differ from those of the nominate form in lack-
ing the dark brown markings on the side of the head and dorsum;
juveniles faintly marked.


FIGURE 16.-(A) Hemiphylodactylus typus typus (fide deRooij 1915); and (B); holotype,
H.t. pallidus (UF 28985).


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


HOLOTYPE.-UF 28985, adult female, collected October 1969 from
beneath dry, fallen leaves in gallery forest along Vai Nggulung, Loho
Liang, Komodo, 30 m.
DESCRIPTION OF TYPE.-Digits slender, cylindrical at base, the
penultimate joint ending in a strong dilation with two series of
oblique lamellae below, separated by a median groove; the inner toe of
both back and front feet rudimentary, clawless; the other digits with a
free, clawed terminal phalanx projecting considerably beyond the
dilated part. Four pairs of lamellae under all normal toes. Pupil
vertical. Head longer than broad; snout as long as the distance be-
tween eye and ear opening, 1 V2 times the diameter of the eye; ear open-
ing very small, oval, oblique. Distinct, thickened glandular patch on
either side of neck above and in front of insertion of forelimbs. Rostral
broad, nearly pentagonal; nostril bordered by the rostral, the first
labial, a supranasal, and two small posterior nasals; 11 supra- and in-
fralabials; mental small, triangular; no chin shields. Limbs slender;
mental tail cyclindrical. Body long and slender, covered with small
granular scales, those on the snout and limbs somewhat enlarged,
those on the tail tubercular. Ventral scales larger, smooth, imbricate.
No preanal pores.
Light tan above, almost uniformly and finely stippled with dark
brown over entire body, tail, and head, very slightly darker from
nostril through and behind eye; uniform white below. SVL 32 mm, tail
26 mm.
PARATYPE.-UF 28878, juvenile male, SVL 21 mm, tail 15 mm,
from gallery forest along Kali Besar, Loho Liang, Komodo, 20 m.
Scalation as in type, except 10 preanal pores on escutcheon of slightly
enlarged scales arranged in a shallow V. Ground color light tan, with
few faint brown mottlings dorsally, forming a V-shaped stripe on top
of snout, very faint stripe from nostril through eye to above ear open-
ing; nine narrow, broken transverse bars, darkest over both hind and
front legs; tail faintly ringed; infralabials mottled; chin and venter
white anteriorly, becoming finely stippled posteriorly onto base of tail.
TAXONOMIC REMARKS.-Apparently closely related to the nominate
form on the basis of identical scalation and similar, but greatly sub-
dued coloration.
DISTRIBUTION AND HABITAT.-The nominate subspecies is known
from Sumatra, Simalur, Nias, Borneo, Java, Bali, and Sumbawa, at
elevations of 500 to 1300 m. Hemiphyllodactylus typus pallidus is
known only from Komodo, where it was taken near sea level from
beneath leaves in gallery forest. Both it and the nominate form are
evidently not common wherever they occur (Mertens 1930).
FooD.-Presumably small miscellaneous insects.


1980








BULLETIN FLORIDA STATE MUSEUM


Lepidodactylus lugubris intermedius (DAREVSKY)
Figure 17
Lepidodactylus intermedius Darevsky 1964c:13 (Type locality: Komodo).
Lepidodactylus lugubris Kluge 1968:338.
SPECIMENS EXAMINED.-(1) Paratype, Museum Zoologicum Bogoriense 978,
Komodo; none found during the 1969-70 NYZS expedition.
DESCRIPTION.-Based on type series (after Darevsky 1964c). Snout
longer than distance between eye and ear opening, 112 to 2 times
diameter of eye. Forehead concave. Ear opening small, round, straight
back from angle of mouth. Head scales granular, largest on snout.
Rostral four-sided, broad. Nostril between rostral, first labial, and
three nasals; 10-11 supralabials, 8-10 infralabials. Mental small, nar-
row, about as large as adjacent infralabial; chin shields small, in 3-4
transverse rows. Body and throat with minute granules; ventral scales
larger, flat, juxtaposed. Femoral and preanal pores 24, in a long
angular series. Tail cylindrical, without sharp lateral edge, covered
with small equal scales in feebly expressed groups of 9-10 transverse
rows. Limbs moderate; digits with rudimentary webs, inner digit well
developed, without claw, middle finger with 9 and middle toe with
11-12 subdigital lamellae.
Light gray above with 7 transverse, undulating, narrow, dark
brown transverse bands, darker bands on tail; no contrasting dark
streak through eye (though faint in Darevsky's [1964c] Figure 2); lower
surfaces greenish-white. SVL 39 mm, tail 35 mm.


FIGURE 17.-Lepidodactylus lugubris intermedius, (fide Darevsky 1964b); Komodo.


Vol. 25, No. 2







AUFFENBERG: KOMODO HERPETOLOGY


REMARKS.-I cannot explain how I failed to find this lizard on
Komodo, except that it is apparently rare and thus was simply missed
by chance alone. Darevsky placed it close to and intermediate between
Lepidodactylus lugubris and L. lombocensis, but it seems part of a
geographic cline in certain scale characters (number of femoral pores,
labials, and chin shields generally reduced to eastward, and number of
subdigital lamellae increased in the same direction). Kluge (1968) cor-
rectly suggested relegation of several named species, including this
one, to the synonomy of L. lugubris. The characters of L. intermedius
suggest it should be recognized as a geographic race of lugubris.
Apparently occasionally eaten by Dendrelaphis pictus (Darevsky
1964c).
DISTRIBUTION AND HABITAT.-Known only from Komodo and Rint-
ja, where specimens were collected from under the loose bark of dead
tamarind trees (Darevsky 1964c).

FAMILY SCINCIDAE
Cryptoblepharus
In his monographic treatment Mertens (1931) concluded that the
36 populations of Cryptoblepharus boutonii he recognized should all be
considered geographical races of one another. Of these, 31 occurred on
islands. Cryptoblepharus boutonii thus became one of the most
widespread of all lizard species in the world, having a discontinuous
distribution from the east coast of Africa to Hawaii. On the basis of in-
formation now available from both Komodo and Padar (and suggested
for other islands, such as Timor) it is highly likely that Mertens' inter-
pretation of a single species is perhaps oversimplified. In fact, Mertens
himself suggested that the various populations of Cryptoblepharus
boutonii represent incipient species, and that some may be much more
differentiated than others. Data provided here suggest that the two
populations in the Komodo area, C. burden and C. renschi, are distinct
at the species level.
Dunn (1927c) described C. burden as a new and distinctive
nonstriped subspecies of Cryptoblepharus boutonii based on three
specimens collected from rocks at the tideline on the east coast of
Padar. No Cryptoblepharus were found on Komodo. Until this time, no
additional specimens of this interesting form were found. Darevsky
(1946b) reported the presence of a vividly striped, tree-dwelling Cryp-
tobelpharus on Padar, referred to C. b. renschi (type locality Sumba).
He also reported the same form from Komodo for the first time, but
failed to find C. burden on Padar. During the New York Zoological







BULLETIN FLORIDA STATE MUSEUM


Society study, both forms were found on both islands, and each was
restricted to a specific habitat type.

Cryptoblepharus boutonii renschi (MERTENS)
Figure 18
Cryptoblepharus boutonii renschi Mertens 1928a:20 (Type locality: northeastern
Sumba, Kambaniru, near Waingapu).
Ablepharus boutonii renschi Mertens 1931:154.
SPECIMENS EXAMINED.-(19) UF 29921-25, east coast of Padar, 10 m; UF 29919-20,
Loho Boro, near Loho Lavi, Komodo, 3 m; UF 29918, Vai Nggulung, Loho Liang,
Komodo, 30 m; UF 29960-61, 29963-64; 28856-59, 28867, 28872, all from near Base
Camp, Loho Liang, Komodo, 3-25 m; UF 28871, near Kampung Komodo, Komodo,
20 m.
DESCRIPTION.-A somewhat small, flat- and pointed-headed
subspecies of Cryptoblepharus boutonii with a markedly striped black
and white pattern, composed of five bluish-white to white or cream
(rarely light brown) bands and six black to brownish-black bands. Me-
dian dorsal stripe, as well as the dark parietal bands, begins on point of
snout; maxillary band begins in subtympanal region, somewhat as in
C. b. keiensis, extending to hindlimb. Sometimes light band begins
from white or cream supralabials running posteriorly to ear opening.
Belly bluish-white; palmar surfaces white. (Newly hatched individuals
have the same color and pattern as the adults [Mertens 1931; UF
29919, 28858, 28860, 28864].) Midbody scales in 24-28 rows in
Komodo-Padar series (X = 26.0), 22-25 (usually 24) rows on Sumba, ac-
cording to Mertens (1930); 20-28 in other races. Both middle scale rows
larger than those on either side. Lower eyelid immovable, transparent,
covering the eye. Nostril in a nasal scale; postnasal well developed in
only one of specimens examined. Usually 4 supraciliaries, 5 in three
specimens, 3 in four specimens, 2 in four, and 3 on one side and 4 on the
other in remaining ones. All have 4 supralabials in front of subocular;
anterior loreal as long or longer than high. Lamellar scales of hind digit
IV varies from 18 to 24. Size small, SVL about 50 mm, tail about 60
mm. Hind leg reaches to ankle of appressed front leg in males and to
axilla in females.
TAXONOMIC REMARKS.-C. b. renschi is most similar to the Kei
Islands subspecies (keiensis). In the latter (as well as certain pattern
phases of C. b. intermedius) the parietal bands on the head extend for-
ward only to the supraocular region; in C. b. renschi they always ex-
tend to the very tip of the snout. C. b. keiensis also differs from C. b.
renschi in having (1) stripes more greenish than silver-white; (2)
somewhat less contrasting dark longitudinal stripes; and (3) up to 22
middorsal scale rows, whereas in C. b. renschi the largest number of
scale rows is 28. However, Mertens (1931) believed that these two


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


subspecies are very closely related to one another. The differences are
apparently bridged by the subspecies C. b. intermedius (Mertens
1931).
Darevsky (1964a) reported that their 3 specimens from Komodo
and 10 from Padar were typical of the subspecies, but with a tendency
for increased scale rows when compared to specimens of the same race
from Sumba. Our material suggests the same (see description above).
No significant difference is evident in the middorsal scale rows of
specimens from Padar (X = 25.8) and Komodo (X = 26.1). Two pro-
bably closely related subspecies are found near Komodo-C. b. sum-
bawanus on the eastern half of Sumbawa (C. b. balinensis is apparently
found in the western half of the island [Mertens 1936]) and C. b.
leschenault on Flores and other islands eastward.
DISTRIBUTION AND HABITAT.-Various localities on Sumba (For-
cart 1949), Padar, Komodo, and Pulau Longo (off Komodo). It pro-
bably also occurs in western Flores, for in 1971 I failed to capture a
specimen seen in a coastal savanna habitat near Nggoer, Mangarrai
District. It was certainly a member of this subspecies.
All the specimens reported, including those taken by Darevsky and
Kadarsan, were taken in savanna habitats, often on trees or under
their bark. With the exception of Forcart's remark (1949) that he found
specimens in dry leaves, all other references imply that this subspecies


^......i..A. .w.. .
I ~L / A n) rCC ^~~~~^ ^-4K;*
Ji A 1'i


FIGURE 18.-(A) Cryptoblepahrus boutoni renschi, Loho Liang, Komodo (UF) 29918);
and (B and C) C. burden, Gunung Klinta, Komodo (UF 30043).








BULLETIN FLORIDA STATE MUSEUM


is normally found on standing trees (Mertens 1927a, 1930, Darevsky
1964a). My observations on both Padar and Komodo show that on
these islands it is always found below 100 m, usually on the trunks of
the shrubby savanna tree Zizyphus jujube and usually close to the sea.
REPRODUCTION.-Newly hatched young have been reported for July
(this study and Mertens 1931), August (this study and Darevsky and
Kadarsan 1964), and October (this study). When hatched they have an
SVL of 17 mm and are approximately 32 mm at the same time the
following year.
FooD.-Spiders and very small insects, most of which are termites
and beetles.
PARASITES.-None found in specimens examined.
Cryptoblepharus burden (DUNN)
Figure 18
Cryptoblepharus boutonii burden Dunn 1927c:11 (east coast of Padar).
Ablepharus boutonii burden Mertens 1930:162.
SPECIMENS EXAMINED.-(55) UF 28967-73, west coast of Padar, 20-30 m; UF
29394-401, Tanjung Kuning, Komodo, 0-2 m; UF 30040-46, base of Gunung Klinta,
Komodo, 0-2 m; UF 29903-17, Loho Boro, nr Loho Lavi, Komodo, 0-3 m; UF 29867,
29877-86, Pulau Longo, off Komodo, 0-4 m; UF 29887-902, Kampung Sabita, Komodo,
0-2 m.
DESCRIPTION.-Scutellation as in Cryptoblepharus boutonii, except
anterior loreal as high as or higher than long and with more midbody
scale rows. Komodo sample (N = 48) mean 30.9, Padar sample (N = 7)
mean 30.6; overall range for total (N = 55) 29-34(X = 30.8). Subdigital
lamellae of 4th hind toe 20-24.
Color and pattern of C. burden quite different from that of C.
boutonii. No light stripes nor evidence of them. Entire upper surface
metallic dark brown to nearly black, with scattered, irregular lighter
flecks. Some individuals nearly uniform brassy brown with slightly
darker edge to each scale, sometimes forming faint interrupted
longitudinal lines. Ventral surface always an almost uniform bluish-
gray to gray; palmar surfaces very dark, soles lighter. SVL of large
specimens about 50 mm, tail about 60 mm. Males with hind legs
reaching to ankle or middle of forearm of appressed front leg, to axilla
in females.
DISTRIBUTION AND HABITAT.-We obtained specimens only from
the rocky shores of Padar and Komodo, but we saw others on nearby
islets and rocky headlands of extreme western Flores. The habitat is
very similar to that of the New Zealand black shore skink (Leiolopisma
suteri) as described by Towns (1975), although the habits of the two
lizards are very different. In the original description of Cryp-
toblepharus burden, Dunn (1927c) stated: "These lizards were


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AUFFENBERG: KOMODO HERPETOLOGY


discovered by Mr. Burden on rocks of the tide line on the east coast of
Padar. Later I observed them there in great number, playing about on
the wet rocks .... On a wave cut bench of rock, beset with small pools
and alive with Periophthalmus and crabs of various kinds and wet by
the waves of the rising tide, these tiny lizards scuttled about un-
concerned about their larger neighbors. When I tried to catch some
with my hands they ran into water of pools and two were caught there,
clinging under water to the rocks." Mertens (1964) pointed out that
this species and C. littoralis are apparently the most intertidal of all
Cryptoblepharus species. Some races of Cryptoblepharus boutonii,
particularly bitaeniatus, occur in similar habitats. Concerning this
subspecies in East Africa, Loveridge (1920) stated: "This little lizard
has adapted to a marine life; it is a remarkable sight to see it running
over the rocks, which a moment before were washed by waves; for com-
pany it has the peculiar fish Periophthalmus and crabs of many
species. It flies before the incoming wave and presumably manages to
avoid a wetting. It seeks refuge when pursued in the many crevices of
the rocks."
Our specimens were collected in similar situations on both Padar
and Komodo, scurrying over rocks standing in the water, as well as on
rocky cliffs immediately adjacent to the high tideline. None of the
specimens was found over 20 m from the water's edge, and never in
any place lacking small sea cliffs and rock jumbles at the shore.
TAXONOMIC REMARKS.-Mertens (1964) pointed out several in-
stances of the occurrence of two subspecies of Cryptoblepharus
boutonii on the same island. He reported both C. b. balinensis and C. b.
cursor on Bali and Lombok. Miller (in Brongersma 1934) reported both
C. b. schlegelianus and C. b. leschenault on the north coast of Samoa
were due to accidental introduction by man of one of the forms, as he
believed the extensive trans-Pacific distribution of C. b. poecilopleurus
may have been similarly produced; or by waif dispersal. Even more
confusing was his finding in Queensland, Australia, of C. b. virgatus
and C. b. littoralis within meters of one another but in different
habitats (the latter is now considered a separate species [Cogger
1975]). Also there was the problem of both C. b. renschi (reported by
Darevsky 1964a) and C. b. burden (reported by Dunn 1927c) on Padar,
both of which are now known from Komodo as well. That transport of
one subspecies (renschi?) by humans is possible in both insular situa-
tions is suggested by the range of the land snail Rachis punctata,
which is found in Africa, the west coast of India, and Padar-a range
very largely influenced by man (Djajasasmita 1972). Many native
boats visit Komodo and some stop at Padar for the easily obtainable
fresh water there, and C. b. renschi may have been accidentally in-


1980








BULLETIN FLORIDA STATE MUSEUM


produced into both islands from Sumba. Such an explanation will pro-
bably not, however, explain the presence of both forms in coastal
western Flores, or the proximity of two or more subspecies in coastal
Queensland (Mertens 1964), parts of New Guinea (George Zug, in litt.),
or Australia (Ronald Crombie, in litt.). The entire matter of the rela-
tionships of all these populations is far from clear. Even Mertens
(1933, 1934a, 1964) was not sure of their status but elected to consider
them as a Rassenkreis highly modified by man and/or waif dispersal.
At least one of the populations formerly considered a subspecies is
now designated a species (C. littoralis). The similarity of this taxon to
C. burden in both appearance and habitat suggested I examine
material of littoralis to try to determine what relationship, if any, they
have to one another. Two paratypes (AMNH 80071-2, Etty Bay by In-
nisfail, Queensland) and seven additional specimens (UMMZ 132581-2,
Cape Tribulation N. Mossman, Queensland; 132583, four specimens
from Magnetic Island, Alma Bay, Queensland; 132584, South Moule
Island, Whitsunday Passage, Queensland) show low intrapopulational
variation and are very unlike C. burden.
The presence of two highly differentiated populations of Cryp-
toblepharus on both Komodo and Padar, the lack of any indication of
intergradation between them, and the fact that both forms were taken
within several meters of one another in at least one locality on Komodo
(Loho Boro), each in its own environment, strongly suggest that Cryp-
toblepharus boutonii burden Dunn is distinctive from C. boutonii at
the species level and assignable to specific status. Additional material








.. rt


FIGURE 19.-Emoia similis, Loho Liang, Komodo (UF 28869).


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AUFFENBERG: KOMODO HERPETOLOGY


of Cryptoblepharus specimens from East Indian islands will un-
doubtedly change some of the current views regarding relationships of
the populations found there and the distribution of recognized taxa.
REPRODUCTION.-Females lay only one oval egg at a time (10.3-11.3
X 3.2-4.4 mm) during the wet season (recorded for December 16,
though yolk deposition in other specimens collected in the same month
suggests eggs are laid in January and February as well). I examined 13
females that showed sexual maturity occurs at an SVL of about 37 mm
(avg 42.5 mm). We examined 14 males. Mature specimens with thick-
walled, convoluted vasa deferentia and well developed, turgid testes
were at least 38 mm SVL (X = 40 mm).
FooD.-Almost all dissected specimens contained remains of small
insects, most of which were tiny beetles.
PARASITES.-None was recognized in any of the specimens exam-
ined.

Emoia similis DUNN
Emoia similis Dunn 1927c:10.
Leiolopisma kadarsani Darevsky 1964b:84.
Leiolopisma kadarsani padariensis Darevsky 1964b:86
Leiolopisma sembalunica rintjana Darevsky 1964b:86.
SPECIMENS EXAMINED.-(13) Holotype AMNH 31977, Komodo; UF 28862, 29965-66,
28868-70, 28873-74, 28877, all from Loho Liang, Komodo, 0-30 m; UF 28875-76, Ntodo
Liseng, Komodo, 450 m; UF 28805, macerated, from stomach of Lycodon aulicus, Loho
Liang, Komodo.
DESCRIPTION. Head medium length. Lower eyelid with undivided
transparent disc. Ear opening nearly round, smaller than eye, without
any enlarged lobules. Nostril in nasal, bordered above by supranasals.
Frontonasal wider than long, bordering rostral anteriorly and frontal
posteriorly. Frontal shorter than the single frontoparietal and in con-
tact with 2 supraoculars, 4 supraoculars, no interparietal, usually 2
nuchals, 7 supralabials. Midbody scale rows, all smooth, 26-28 (X =
27.3). Dorsal scales somewhat larger than ventral members. Preanals
slightly enlarged. Limbs slightly reduced, not meeting when appressed
against the body (overlapping in hatchlings). Distance between fore-
and hindlimbs 1.5 to 1.8 times the distance between tip of snout and
base of forelimbs; 4th digit of hindlimb longer than 3rd, with 18-23
subdigital lamellae.
Color brownish-black, brown, or brownish-gray above, with 5
silvery-white longitudinal stripes (sometimes paravertebral and dor-
solateral stripes are cream or yellow). Each stripe passes through the
border of two adjacent rows of scales. Vertebral stripe usually begins
near snout, extending posteriorly over head, becoming clearly defined
in most specimens at occiput, where it is often bordered by a fine dark








BULLETIN FLORIDA STATE MUSEUM


edge. Stripe extends posteriorly onto base of tail, which is largely an
almost uniform bluish-gray. The paravertebral stripes extend from
above eye posteriorly to anterior third of tail. Dorsolateral stripes
often indistinct, extending from behind ear opening to insertion of
hind limb. Abdomen, throat, and chin grayish-white to white. Max-
imum length 112 mm; SVL/tail 1/1.2 to 1/1.5 (X = 1/1.3).
TAXONOMIC REMARKS.-According to the literature no specimens
of Emoia similis have been collected since the two original individuals
obtained by Dunn, now in the AMNH (one has subsequently been
skeletonized). Darevsky collected none during his trip to Komodo, and
the matter was thus rather confusing; particularly as Darevsky
(1964b) described Leiolopisma kadarsani from Komodo and a
subspecies from nearby. Dunn's (1927c) description of E. similis was
very short and provided few characters with which to compare it to
other scincids. The assignment of the two Komodo specimens to
Emoia was made on the basis of the presence of supranasal scales. My
examination of the type showed it does possess these scales. Further-
more, during 1969-70 I collected the 12 specimens reported here, and
they agree with the type in every major detail, including the presence
of supranasals (Fig. 19). What is odd is that Darevsky's 15 specimens
of identical appearance and from the same locality are said to lack
them. Figure 5 of Darevsky's description (1964b) shows an almost
completely divided nasal above the nostril. The posterior part of the
cleft is sometimes difficult to see, and investigation of the material
deposited in the Bogor Museum shows that it is present in the type
series of L. kadarsani. Therefore, I place Leilopisma kadarsani in the
synonymy of Emoia similis Dunn.
Leiolopisma kadarsani padariensis Darevsky was described on the
basis of its having a wider dorsal stripe than the nominate subspecies
(particularly near the head) and a slightly less patterned head. Other-
wise the two forms are apparently identical. Of the 13 specimens
available from Komodo all but two have a dorsal stripe about one scale
wide behind the head. UF 28869 and 28870 have wider stripes, that of
the former being two scales wide, as in L. k. padariensis, and the latter
intermediate between the usual Komodo condition and UF 28869 and
the three types of L. k. padariensis. I am tentatively placing
Leiolopisma kadarsani padariensis in the synonymy of Emoia similis.
The occurrence of the subspecific characters in at least some Komodo
specimens and that only a few individuals are presently available from
Padar suggest that the best present course is not to recognize the
Padar population nomenclatorially until more specimens become
available. In the event that the subspecific characters are found to be
valid, the Padar population should be called Emoia similis padariensis
(Darevsky).


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AUFFENBERG: KOMODO HERPETOLOGY


Mertens (1927a) described Leiolopisma sembalunicum on the basis
of one specimen from Lombok; none has been found since. Though the
skin around the nasal area was scarred, Mertens placed it in
Leiolopisma on other grounds. Emoia was disregarded on the ground
that the specimen's legs were far too short. However, their lengths
seem equivalent to those of Emoia similis. Furthermore, all remaining
characters can be matched in these specimens as well, so it is highly
likely that Leiolopisma sembalunicum Mertens may be a synonym of
Emoia similis. By the same reasoning, the type series of Leiolopisma
sembalunica rintjana Darevsky should be reinvestigated, for the figure
provided (Fig. 6 of Darevsky 1964b) shows what might be interpreted
as supranasals.
DISTRIBUTION AND HABITAT.-Definitely known from Komodo,
Padar, and Rintja. Future work may show that it is found as far
westward as Lombok.
Darevsky (1964b) stated that this small lizard was common in the
grass on the edge of bamboo thickets at altitudes of 500-600 m. Dunn
(1927c) found them among stones on grassy slopes at a similar eleva-
tion. Our specimens were collected in Zizyphus savanna from sea level
to about 30 m, usually in the grass or clambering about on the trunks
of Tamarindus or Zizyphus trees.
REPRODUCTION.-I examined 11 females for reproductive data. Of
these seven were immature, as judged by the thin-walled, smooth
oviducts and lack of extensive follicular development. Of the mature
specimens two were gravid; UF 28873 had two large oval-shelled ova
(9.7 X 3.7) in the right oviduct and none in the left (January). UF
28868 had one ovum in both the right and left oviducts (December). In-
terestingly, the former had only one corpus luteum on the right ovary,
maintaining both ova, whereas the latter had one corpus luteum on
each ovary, corresponding to the single ovum in each oviduct.
The single nongravid female had one follicle on each ovary that
showed extensive yolk deposition, whereas the other follicles (4 on left
ovary, 2 on right) were small and translucent, exhibiting little if any
yolk deposition. The oviducts of the three adult females were all thick-
walled and highly convoluted.
In all but two of 10 males examined, the left testes were slightly
posterior to the right. In the remaining two both testes were approx-
imately equidistant from the head. Two individuals were juveniles.
The testes of adult males collected November through February were
turgid and cylindrical to subcylindrical, and the vasa deferentia were
thick-walled and extremely convoluted. The right testis was missing
on one specimen.


1980







BULLETIN FLORIDA STATE MUSEUM


FooD.-Spiders and miscellaneous insects were found in nearly all
the specimens examined, including one small moth, particularly beetle
larvae and small grasshoppers.
PARASITES.-None found in specimens examined.
REMARKS.-Occasionally eaten by Lycodon aulicus.
Mabuya multifasciata multifasciata (KUHL)
Figure 20
Scincus multifasciatus Kuhl 1820:126 (Type locality: [?] Java).
Mabuia multifasciata Weber 1890:160.
Mabuya multifasciata multifasciata Mertens 1930:257.
SPECIMENS EXAMINED.-(1) UF 30030, Poreng, Komodo, 550 m.
DESCRIPTION.-Snout short, obtuse; lower eyelid scaly; ear opening
round to oval (vertical to horizontal) with 3 to 6 sharp lobules on the
anterior edge of the opening (X = 4.3) (rarely missing in specimens
from other areas). Nostril in a nasal scale; postnasal and supranasal
scales present; frontonasal twice as broad as long; prefrontal always in
contact. Frontal as long as frontoparietals and interparietal together
or shorter, in contact with the second of the two anterior supraoculars;
4 supraoculars, second largest. Frontoparietals larger than the inter-
parietal; parietals separated; a pair of nuchals; supralabial 5 largest
and below the eye. Body with 29-33 scale rows on Komodo (29-34 in
other parts of range), each scale with 3 keels, except those over the
pelvic region, which often have 2. Limbs strong, hindlimbs do not
reach the axilla. Fourth hind toe longest, with 14-18 nonkeeled sub-
digital lamellae on Komodo (14-21 elsewhere and sometimes weakly
keeled). Largest Komodo specimen SVL 82 mm, tail 124 mm, much
larger specimens available elsewhere.
Color of adult females brown or olive-brown above, scales with
blackish lateral borders, sometimes forming longitudinal lines in large
specimens. Flanks dark brown or black, with light, black-edged spots.


FIGURE 20.-Mabuya multifasciata multifasciata, Poreng, Komodo (UF 30030).


Vol. 25, No. 2







AUFFENBERG: KOMODO HERPETOLOGY


Ventrally green to yellow, throat bluish-white; side of head gray to
olive-brown; supralabials white, with dusky sutures. Males are similar-
ly colored, except that they are provided with a broad yellowish-green
or yellowish-white band from just above the insertion of forelimbs
posteriorly to slightly over half way to hindlimbs. Venter and throat
darker, often dark gray to blue-green laterally.
TAXONOMIC REMARKS.-Mabuya multifasciata balinensis was
described by Mertens (1927a) on the basis of a contact between the
first supralabial and the frontal. Mertens (1930) found the contact
missing throughout the remaining part of the archipelago from which
he had specimens. The Komodo and Flores material now available
confirms his findings. The remaining diagnostic character of balinen-
sis concerns the coloration of males, which have a reddish-brown
streak on the top of the snout of large males, and a yellow rather than
reddish lateral strip. Unfortunately, no large males are available from
Komodo, but adult males from extreme western Flores have a bright
red lateral stripe, as in the nominate subspecies. Younger individuals
are provided with a gold-colored stripe.
DISTRIBUTION AND HABITAT.-The subspecies M. m. multifasciatus
is found on Lombok, Sumbawa, Komodo, Rintja, Sumba (Forcart
1949), and Flores. In many areas it is the commonest of lizards.
However, on Komodo, Rintja, and western Flores, it is restricted to
quasi-cloud forests in mountainous areas and in streamside gallery
forests. In more humid zones it is widely distributed. Its altitudinal
range is from sea level to at least 1200 m. On Komodo it occurs from
500 to 650 m.
REPRODUCTION.-On Komodo hatchlings were recorded in July and
August. Gravid females were collected in July. Two to four round eggs
(X = 11.5 mm diameter) are laid under stones and in rotten wood.
Minimum SVL of 13 mature females is 85 mm, X SVL of mature
females is 92.5 mm. Ten mature males have a mean SVL of 93 mm and
an overall range of 79-111 mm.
Sphenomorphus (Sphenomorphus) emigrans emigrans
(LIDTH DE JEUDE)
Figure 21
Lygosoma emigrans Lidth de Jeude 1895:125 (Type locality: Sumba).
Lygosoma everetti Boulenger 1897:504 (Type locality:Sumba).
Sphenomorphus emigrans emigrans Mertens 1928a:230.
Lygosoma (Sphenomorphus) emigrans Smith 1927:220.
Lygosoma (Sphenomorphus) emigrans emigrans Forcart 1949:372.
Sphenomorphus mertensi Darevsky 1964b:81.
SPECIMENS EXAMINED.-(23) UF 28821, 28823-35, 28837, 28839-40, 28960, 28962-66,
all from Loho Liang, Komodo, 0-30 m.







BULLETIN FLORIDA STATE MUSEUM


DESCRIPTION.-Head obtuse anteriorly; lower eyelid scaly, with 2-4
central scales markedly enlarged, forming a slightly distinct,
semitransparent palpebral disc; ear opening oval to nearly round, with
three slightly distinct scale lobules anteriorly; nostril in a nasal scale,
no supranasal; frontonasal wider than long; prefrontals separated by
fronto-frontonasal contact; 5 supraoculars, with second (sometimes
third) being the largest; parietals in contact with each other posterior
to a single frontoparietal; 7 supralabials; scales of midbody 26-30 (X =
27.7); 2 slightly enlarged preanals; tail somewhat thickened along its
entire length, long tail (body/tail: 1/1.1-1/1.4); adult SVL 50-65 mm, tail
70-95 mm; limbs short, hindlimb somewhat shorter than distance from
tip of snout to base of forelimbs; distance between limbs 1.5 times
distance between snout and base of forelimbs; fourth digit of
hindlimbs larger than third, with 20-24 subdigital lamellar scales (X =
22.4).
Color yellowish-brown above, in which most scales are provided
with a brown spot, sometimes aligned longitudinally to produce a
short stripe, especially in anterior half of body and sometimes on tail.


FIGURE 21.-Sphenomorphus emigrans emigrans, Loho Liang, Komodo (UF 28851).


Vol. 25, No. 2







AUFFENBERG: KOMODO HERPETOLOGY


A wide dark brown stripe passes from nostril over canthus and eye
through temporal area to over hindlimb, sometimes faintly edged with
cream above and below. Sides of body below temporal stripes densely
covered by small speckles and spots, gradually disappearing toward
abdomen. White to cream below. Infralabials variously speckled with
brown.
TAXONOMIC REMARKS.-Darevsky (1964b) described Sphenomor-
phus mertensi from Padar without comparing his type material with
specimens of S. emigrans, of which it clearly is a synonym. No single or
combination of characters will separate S. mertensi from S. emigrans.
The distinctly striped S. e. wetariensis Mertens has a restricted range
from islands north of the Flores coast to Wetar. Mertens (1930) be-
lieved the New Guinea material may belong to an undescribed
subspecies closely related to S. e. wetariensis.
DISTRIBUTION AND HABITAT.-Sphenomorphus emigrans was
previously unknown from Komodo, though it was reported from
several localities on Sumba (Forcart 1949), Wetar, Samao, Pulau
Sukur, Teun, Pulau Besar (Gr. Bastaard, north of Flores), and
Astrolabe Bay on New Guinea. Curiously, it has not been reported
from Flores itself, although it occurs on almost all the islands sur-
rounding it. However, I expect it occurs in at least the westernmost
part of the island because of similar habitat. All the Komodo
specimens were found under debris and among the leaves on the floor
of deciduous monsoon forest. Many were taken from can traps buried
along a drift fence. None was found above 30 m or far from the sea.
REPRODUCTION.-Average SVL of 10 mature females is 55.0 mm
(range 47-61), for 5 mature males 55.7 mm (42-62). The sex ratio of the
total sample (N = 23) is almost exactly 2:1 in favor of females. No
gravid females or eggs were found, but follicular development sug-
gests that one egg is laid each year, probably during the wet season,
for specimens collected July through October show no major growth of
ova, whereas those collected in November have proportionately much
larger yolked ova. Unfortunately, none was collected during the mon-
soon months of December through February.
FooD.-Small insects, particularly bupestid beetles and their lar-
vae, and ants. Scorpions are also frequently taken. Skink scales found
in one stomach suggest that this species occasionally eats its own shed
skin-a habit apparently not common among scincids.
PARASITES.-The nematodes Parapharyngodon maplestonei and
Pharyngodon kuntzi were common in the colon.


1980








BULLETIN FLORIDA STATE MUSEUM


Sphenomorphus (Sphenomorphus) florensis florensis (WEBER)
Lygosoma florense Weber 1890:173 (Type locality: Maumeri, Flores)
Sphenomorphus florensis Barbour 1912:29.
Sphenomorphus florensis florensis Dunn 1927c:5.
Sphenomorphus florensis nitidus Dunn 1927c:5 (Type locality: Komodo, 1200 ft).
Lygosoma (Sphenomorphus) florense Smith 1927:219.
SPECIMENS EXAMINED.-(72) UF 29926-30, 29938-51, all from Nggoer, Mangarrai
District, Flores, 0-10 m; Holotype S. f. nitidus, AMNH 32068,650 m, Paratypes AMNH
32055-67, 32070, 32101-4, 0-650 m, Komodo; UF 28819-20, 28822, 28826, 28836, 28838,
28841-54, 28861, Loho Liang, Komodo, 0-30 m; UF 33067-68, 33070-72, Nunung,
Mangarrai District, Flores, 700 m; UF 33064, Nisar, Mangarrai District Flores, 200 m;
UF 33065-66, Ruteng, Flores, 1200 m; UF 32601-3, Padar.
DESCRIPTION.-Head medium length, snout somewhat shortened;
lower eyelid scaly; ear opening oval, smaller than orbit, with 4-7
anterior lobules (usually 6) in Komodo specimens (fewer in some other
areas, see Mertens 1930); nostril in a nasal scale, no supranasals; fron-
tonasal slightly broader than long; frontal almost as long as fron-
toparietal and interparietal together, in contact with 4 anterior
supraoculars; 6 to 7 supraoculars, first larger than the second;
parietals in contact; no nuchals; supralabials 7, 6 and 7 largest, under




















A B C D



FIGURE 22.-Sphenomorphus florensis florensis, all from Komodo. (A) (UF 28846) SVL
49 mm; (B) (UF 28844) SVL 53 mm; and (C) (UF 28851) SVL 50 mm; (D) (UF 28849) SVL
53 mm.


Vol. 25, No. 2







AUFFENBERG: KOMODO HERPETOLOGY


eyes; scales smooth, largest dorsally, 44-47 in number on Komodo (X
= 45.9) (43-50 over entire range); length from tip of snout to insertion
of forelimb is 11/3 times the length between appressed limbs; two
preanals enlarged; fourth hind toe longest, with 24-29 (X = 27.7)
smooth (rarely feebly keeled) subdigital lamellae.
Ground color light brown with metallic luster, sometimes with nar-
row lighter middorsal stripe about one scale wide; dorsally with small
brownish spots, sometimes arranged in paravertebral rows, indistinct
in large adults, most distinct in young where they sometimes nearly
form crossbands; dark brown speckles on legs and dorsal surface of
tail; dark lateral stripe beginning behind eye and extending over ear
and limb insertions onto sides of tail for '/ to /2 of its length. Dark
brown stripe widest in middle of body, but often disrupted by white
spots along lower edge. Often edged with a narrow white to cream line
or white spots above or short vertical bars extending onto dark side
stripe. Subcanthal stripe from eye to nostril less evident in Komodo
specimens than those from lowland Flores. Subocular dark stripe and
supralabial sulcal vertical bars dusky, not intense in most Komodo
specimens. Venter, throat, and chin greenish-yellow to yellow-white.
TAXONOMIC REMARKS.-Komodo is the type locality of Sphenomor-
phus florensis nitidus, which Dunn (1927c) described on the basis of
several color and pattern characters. After a thorough comparison
with topotypic material of S. f. florensis, Mertens (1930) placed S. f.
nitidus in the nominate subspecies, and I agree. All the characters of S.
f. nitidus are duplicated in specimens from both Flores and Sumbawa.
The head of S. f. florensis adult males is said to be red-tinged, whereas
that of S. f. nitidus is not. Mertens stated that only the S. f. florensis
from eastern Flores were so colored. The male specimens we collected
in July are brassy brown on the top of the head, as in most adults, but
the chin and throat, as well as the side of the neck and front legs, all
have a reddish tinge. This fades into yellow on the chest just in front of
and between the forelimbs. The colors are probably sexually cor-
related, for we did not see them in the few females collected. Further-
more, they may be seasonal. Though Weber (1890) mentioned black
flecks on the chin of Flores specimens, Mertens (1930) found none on
any of his material, nor did we on our specimens from west Flores or
Komodo, but the darker individuals from Ruteng, Flores, have flecks
on the infralabials.
DISTRIBUTION AND HABITAT.-Sphenomorphus f. florensis has been
reported from Sumbawa, Sumba (Forcart 1949), Komodo, Padar, Rint-
ja, Flores, Pulau Besar, Adonare, Alor, Samao, and Timor; S. f. bar-







BULLETIN FLORIDA STATE MUSEUM


bouri from Wetar, and S. f. weberi from Damma. On Komodo it is
distributed from sea level to 650 m; on Flores we collected it as high as
1200 m. Throughout its range it is usually found in drier areas.
Mertens (1930) reported it from small rocky islets off Sumbawa and in
the arid zones of southeastern Flores (particularly Corypha and Pan-
danus forests) as well as the moist mountain top forests in both Sum-
bawa and Flores. It is commonest in open places in monsoon forest,
particularly along the dry sandy creek beds. Here it is quite abundant
and can be found scurrying about, even at night. It climbs well and on
Komodo was often seen in trees to a height of 10 m. Both the New
York Zoological Society and the Burden Expeditions found it in the
quasi-rain forests of Komodo at about 500 m elevation, but always in
open areas, quite often in ecotonal areas.
REPRODUCTION.--The sex ratio is essentially equal (a' c 1:1.3 9 9).
In both males and females maturity occurs at an SVL of about 58 mm
(average in females 62.3 mm, males 63.5 mm). Eggs are probably laid
during the wet season, for in UF 28961, collected in November, the
right ovary contained large, yellow follicles showing advanced yolk
development (3.0 mm d.); all others were white and less than 1 mm in
diameter. The left ovary had five follicles: one large (3.4 mm d.) and
yolked, two with slight yolk deposition (1.5 mm d.), and two very small
and translucent. The oviducts were thick-walled and extremely con-
voluted.
FooD.-Food remains were found in 75% of the specimens exam-
ined. Beetle larvae were most numerous; termites, ants, and grasshop-
pers were infrequent, spiders more common.
PARASITES.-Endoparasitemia is low, with a colonic oxyuroid
(Thelandros sp.) found in only 1 of 22 specimens examined.
REMARKS.-One specimen was found in the stomach of a Lycodon
aulicus. Individuals were frequently seen abroad at night.
Sphenomorphus (Sphenomorphus) striolatus (WEBER)
Figure 23
Lygosoma striolatus Weber 1890:161 (Type locality: Reo, Flores).
Sphenomorphus striolatus Barbour 1912:185.
SPECIMENS EXAMINED.-(65) AMNH 32071, 32076-85, 32090-93, Komodo, 650 m;
UF 29931, 32573-78, all from Nggoer, Mangarrai District, Flores, 10 m; UF 33075,
Ruteng, Flores, 1200 m; UF 28925-32, Gunung Satilibo, Komodo, 600 m; UF 28933-46,
Galulai, Komodo, 500 m; UF 28947-59, Mborombampang, Komodo, el 350 m; UF
28974-83, Gunung Vau, Komodo, 500 m.
DESCRIPTION.-Head moderately short, particularly the snout;
lower eyelid scaly; ear opening oval, smaller than orbit, with no
lobules; nostril in a nasal scale, no supransals; frontonasal much
broader than long; frontal about as long as frontoparietals and inter-


Vol. 25, No. 2







AUFFENBERG: KOMODO HERPETOLOGY


parietal together; 6 to 7 supraoculars, the first longer than the second;
parietals in contact; nuchals weakly developed or absent; 5th and 6th
supralabials largest and under eye; scales at midbody in Komodo
specimens 28-42 (X = 40.5), 36-45 over entire range; length from tip of
snout to insertion of front leg is about 1% to 1 times the distance be-
tween the appressed legs; fourth toe largest, with 23-26 keeled sub-
digital lamellae.
Color not noticeably sexually dimorphic. Metallic brassy brown to
greenish-yellow middorsal stripe about two scales wide. A
paravertebral row of dark brown to black long spots or dashes,
sometimes with cream or white spots in between, dark spots continu-
ing onto tail, though faded. Sides with irregular dark stripes com-
prised of a brown or black spot on each scale, sometimes broken by
narrow irregular stripes, dashes, or spots of cream, tan, or brown.
Dark stripe from nostril to eye. Supralabials and infralabials edged
ventrally and dorsally with brown or usually black, often extending
dorsally along supralabial sulci. Legs speckled with brown or black.
Postorbital stripe variable. Ventral surface uniform cream to white.
Hatchlings and juveniles colored and patterned as adults.
DISTRIBUTION AND HABITAT.-Known only from Komodo, Rintja,
Flores, and Damma. Generally widely distributed in quasi-rain forests
in the mountains to at least 1200 m elevation, but also at low eleva-
tions in moist gallery forests along rivers, such as at Reo and Nggoer,
Flores.












B1



FIGURE 23.-Sphenomorphus striolatus striolatus; (A and B) Nggoer, Flores (UF
32574); and (C) Ruteng, Flores (UF 33075).








BULLETIN FLORIDA STATE MUSEUM


REPRODUCTION.-The sex ratio (N = 35) is probably equal (c Oc
1:1.2 9 9). I examined 18 females for reproductive data; 11 had one
shelled egg in the oviduct (11.1-13.4 x 4.5-5.7 mm). The shell was com-
pletely deposited, being white and granular. The associated ovaries
each had one corpus luteum; disc-shaped, yellow, and an indented
center. In all females I examined the oviducts were broad and con-
voluted (except UF 28948) and very thick-walled and mature (except
UF 28942 and 28929). In UF 28942 and 28929 the oviducts were
thinner-walled, though each contained shelled ova. The number of
follicles varied from 2 to 7 (X = 5.3); follicular size from less than 1 to
4.0 mm. Apparently only one egg is laid at a time, 2-7 annually, most
of which are laid in the monsoon season, December to February.
Average size of 18 mature females is 55 mm, of 15 males 49 mm.
FooD.-Food remains were found in 90% of the specimens col-
lected. Beetle larvae were most common, but ants, termites, grasshop-
pers, spiders, adult beetles, and scorpions were also found. The beetle
larvae represented several types that are most commonly found deep
in rotten wood and suggest considerable digging activity by the
lizards.
PARASITES.-Endoparasitemia is low, with the nematode
Trichoskrjabinia secundus found in only 35 specimens.
Sphenomorphus (Homolepida) schlegeli (DUNN)
Figure 24
Homolepida schlegeli Dunn 1927c:8 (Type locality: Komodo; 650 m).
Omolepida schlegeli Mertens 1930:272.
Sphenomorphus schlegeli Darevsky 1964a:569.
Sphenomorphus oxycephalus Darevsky 1964b:83 (Type locality: Rintja).
SPECIMENS EXAMINED.-(4) Holotype, AMNH 31994; UF 28983-84, Gunung Ara,
Komodo, 500 m; UF 28989, gallery forest along Kali Besar, Loho Liang, Komodo,
200 m.
DESCRIPTION.-Snout longer than S. temmincki; lower eyelid scaly;
ear opening large, round to slightly oval, nearly same size as eye, no
scaly lobules; nostril in a nasal; no supranasal; frontonasal wider than
long; prefrontals in contact; frontal about as long as frontoparietals
and half of interparietal, in contact with first two supraoculars; 5
supraoculars, third largest; parietals in contact behind the inter-
parietal; no nuchals; 5 supralabials; suboculars present; midbody scale
rows 22-24, smooth paravertebral rows slightly enlarged; preanals
slightly enlarged; limbs very short, small; fourth hindlimb digit
longest, with 9 (1 specimen) to 10 subdigital lamellae.
Brownish-gray above with bronze tinge. Two narrow white to
cream dorsolateral stripes from last supraocular or over ear opening
posteriorly onto base of tail, bordered above by a narrow dark row of


Vol. 25, No. 2







AUFFENBERG: KOMODO HERPETOLOGY


spots anteriorly. Sides dark brown, darkest below light dorsolateral
stripe and fading to yellow-gray on venter. Throat and chin im-
maculate to spotted with brown. Tail variable, from almost uniform
light brown to marked with a spotted or much darker dorsolateral
stripe extending about one half tail length. Body/tail lengths: 22/24,
33/29+, 31/35.
TAXONOMIC REMARKS.-Sphenomorphus oxycephalus Darevsky is
synonymized with S. schlegeli (Dunn) because the stated characters of
the type clearly fall within the variation of the specimens of S.
schlegeli now available from Komodo. The considerably narrow head
of the type mentioned by Darevsky (1964b) is obviously due to the fact
that the specimen was allowed to dry out, as is clearly shown in the
photograph provided in a companion paper by Darevsky (1964a).
Dunn (1927c) placed this species close to the Papuan S. crassicauda
and S. forbesi. In my opinion it seems much closer to the former.
Either schlegeli or close relatives are logically expected in moun-
tainous tracts of Flores and other more eastern islands of the Lesser
Sundas. This group (including S. unilineatum, West Irian) is apparent-
ly restricted to the New Guinea-Nusa Tenggara section of the Lesser
Sundas. The S. temmincki group (including S. parvum) is much more
widely distributed, being found from Sumatra to near Timor. S. alfredi








A B


C







D E

FIGURE 24.-(A and C) Sphenomorphus schlegeli, Gunung Ara, Komodo (UF 28983) (B)
S. schlegeli, holotype (AMNH 31994); and (D and E) S. oxyrhopus, holotype (Zool. Inst.
Sci., Leningrad, 17598).


1980








BULLETIN FLORIDA STATE MUSEUM


and S. hallieri comprise a separate subgroup of the subgenus
Homolepida, restricted to Borneo.
DISTRIBUTION AND HABITAT.-Most quasi-rain forests of Komodo
and Rintja. at elevations generally above 500 m.
REPRODUCTION.-Both females examined had 3 follicles in both
their right and left ovaries (> 1-1.5 mm). Their oviducts were flat, thin-
walled, and only slightly convoluted, suggesting egg laying does not
occur during the dry season (specimens collected in August).
Food.-No food remains were found.
PARASITES.-No parasites were found in the two female specimens
dissected.
REMARKS.-Until this study only the type was known; collected in
"heavy forest, interspersed with great rock masses which forms the
elevated center of Komodo." One specimen was taken from the
stomach of an adult Psammodynastes pulverulentus.

FAMILY AGAMIDAE
Draco volans reticulatus (GUNTHER)
Figure 25
Draco reticulatus (Giinther) 1864:125 (Type locality: Philippines).
Draco volans Roux 1911:496.
Draco reticularis Elbert 1912:329.
Draco timorensis Rooij 1915:75 (part).
Draco volans reticulatus Mertens 1930:249.
SPECIMENS EXAMINED.-(7) UF 28918-24, Loho Kiang, Komodo, 10m.
















A B C
FIGURE 25.-Flight membrane of Draco reticulatus. (A) dorsal surface of adult female
(UF 28924); (B) same, of adult male (UF 28921); and (C) ventral surface of adult male
(UF 28921), all from Komodo.


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


DISTRIBUTION AND HABITAT.-Previously known from most of the
Lesser Sunda Islands: Celebes, Flores, and Sumbawa (Mertens 1930);
Sumba (Forcart 1949); and Komodo (Darevsky 1964a, who also saw it
on Rintja). Though originally taken in the Philippines, its exact locali-
ty was unknown, and it has not been taken there since. Draco v. volans
extends westward to Thailand on the mainland.
It is a much commoner lizard than the number of captured
specimens indicate; many more could have been captured had a serious
attempt been made to do so. Mertens (1930) stated that it is more com-
mon at higher altitudes, but our experiences suggest it is abundant in
any rather mesic, forested area. The gallery forests along both Vai Ng-
gulung and Kali Besar in Loho Liang, Komodo, as well as the coconut
plantation near Kampung Komodo, had good populations as dense as
those in the higher mountain forests. I agree with Mertens (1929b)
that this lizard is most often encountered on large nearly leafless trees
in open forests.
REPRODUCTION.-Of the three females examined, only one (UF
28924), collected in February, was gravid. Each oviduct contained two
partially shelled, oval ova (14.5-16.9 x 4.1-7.5 mm) ready to be laid, so
that the total egg complement would have been four. In addition, the
left ovary had two corpora lutea, two large yolked follicles (suggesting
multiple clutches), and nine very small translucent follicles. The right
ovary had one large yolked follicle, one small translucent follicle, and
no visible corpora lutea.
FOOD.-Dissected specimens contained only ants.
REMARKS.-Only one species of Draco inhabits the Lesser Sunda
Islands chain; it is extremely variable in both color and many of its
scale characters (Mertens 1930). As in most individuals of this species,
the scales near the vertebral line of the Komodo specimens are
somewhat irregular. They also lack the keeled paravertebral scales
that are sometimes found in Flores specimens but are missing in those
from Sumbawa and Lombok. The dorsal color of the gliding membrane
is reticulated with black, brown, and white; those of the males are
generally darker than those of the females (Fig. 25). The ventral mem-
brane is uniform white, but some specimens have a few black flecks or
streaks, in males particularly near the outer edge (Fig. 25). Dewlap
lemon yellow in males.

FAMILY DIBAMIDAE
Dibamus novaeguinea (DUMERIL AND BIBRON)
Dibamus novae-guineae Dumeril and Bibron 1839:834.
SPECIMENS EXAMINED.-(5) UF 33074, female, Vai Nggulung, Loho Liang, Komodo,
300 m; UF 33073, male, Kali Besar, Loho Liang, Komodo, 160 m; UF 29460, 29449, two
males, Loho Lavi, Komodo, 30 m; UF 33488, female, 1 km S Nunung, Flores, 75 m.


1980








BULLETIN FLORIDA STATE MUSEUM


DISTRIBUTION AND HABITAT.-This distinct legless lizard has a
wide distribution-from the Nicobar Islands and the Malay peninsula
through the Sundas to New Guinea. In the Komodo area it has been
reported from Flores (Smith 1927), Sumbawa (Mertens 1930), Sumba
(Forcart 1949), and Komodo (Darevsky 1964a). The usual habitat is
under stones in moist forests, generally above 400 m. Our specimens
from below this elevation were taken from beneath stones in the
gallery forests that extend downward as moist, cooler tongues of the
quasi-rain forests on the highest slopes. Darevsky apparently collected
his two specimens on the central ridge of Komodo above 500 m. UF
33074 was found drowned after a flash flood.
REMARKS.-Mertens (1930) stated that both his specimens (Flores
and Sumbawa) had 22 midbody scale rows, whereas those from Lom-
bok had 20-24 rows, suggesting significant geogaphic variation. Two
of the Komodo males have 24 rows and the third male and the female
22 rows. One male (SVL 129 mm, tail 21 mm) is a uniform brownish-
purple above and below, with only the enlarged head scales and
perianal region lighter; another (92 mm/18 mm) is similar but with
lighter spots on the belly, and the third (86 mm/13 mm) is provided
withia few random spots over the body and a light band near the mid-
dle of the body. The female (132 mm/16 mm) is similar, except that the
belly is mottled with lighter color, and two broad bands of the same
lighter shade encircle the anterior half of the body.
I can add nothing new regarding this species' reproductive and
feeding habits.

FAMILY VARANIDAE
Varanus komodoensis OUWENS
Figures 26-33
Varanus komodoensis Ouwens 1912:1 (Type locality: Komodo).
Placovaranus komodoensis Fejevary 1927:284 (substitute name).
SPECIMENS EXAMINED.-(164) UF 29000-1, skins and skeletons, coast at Nggoer,
Mangarrai District, Flores (under permit); UF 28221, right mandible found on surface,
Gunung Ara, Komodo; UF 28220, skeleton, tagged #27W in life, killed by monitor
#19W on 16 November 1969, Loho Liang, Komodo; UF 28219 skeleton, tagged #18W in
life, killed by monitor #19W on 3 April 1970, Loho Liang, Komodo; UF 28227, parts of
skeleton from monitor fecal pellets, Ntodo Klea, Komodo; UF 28228, from fecal pellet
remains, Loho Liang, Komodo; MZB 582, 6 full term embryos, eggs laid in Surabaja
Zoo; MZB (no number), 2 embryos, same date; MZB 926-28, 946, 980-81, Loho Liang,
Komodo; MZB 929, Mburo, Flores; plus 127 live specimens examined on Komodo,
Padar, and Flores during this study.
DESCRIPTION.-A very large (max. total length ca 3 m, X = 1.70 m,
X SVL = 74.5 cm, see Fig. 26), heavy monitor when adult (max. field
weight 54 kg, X = 8.6 kg, N = 50, see Fig. 27); males slightly larger


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


-34 -54 -* 74 94 -114 -134

SNOUT-VENT LENGTH
(in cm)


FIGURE 26.-Comparative lengths of Varanus komodoensis specimens from Nggoer,
Flores (shaded) and Loho Liang, Komodo (clear). Smaller histogram delineated by
darker line represents the Nanenepi, Komodo. The latter is mangrove habitat, in which
primarily the larger animals occur (see Auffenberg, in press, for details).


1980







BULLETIN FLORIDA STATE MUSEUM


than females (X total length 10 largest males 2.26 m, 10 largest
females 1.76 m); total length of 29 hatchlings 25.3-55.5 cm (X = 30.4
cm), weight 80.3 gm. Snout broad, truncated (Fig. 28). Statements by
Lederer (1942) and various animal dealers that females have propor-
tionately narrower heads are incorrect (Fig. 29). Round narial openings
located near end of snout; tail relatively short and heavy in adults, pro-
portionately larger in young (Fig. 30); cross-section of tail base round
in adults, more compressed in young. Teeth considerably compressed


641-


,0)



I-


0,
.0


..




0*
*0*
-


100


150 200 250 300


TOTAL LENGTH (in cm)


FIGURE 27.-Length-weight relationship in Varanus komodoensis, all from Loho Liang,
Komoda.





Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


laterally, with serrated posterior edges on at least maxillary and den-
tary series (Burden 1928, Auffenberg, in press); premaxillary teeth 7,
maxillary 13, dentary 12. Males distinguished by a pair of small
precloacal pores (Fig. 31), in spite of the suggestion by Mertens (1930)
that this character is probably not sexually diagnostic (see Auf-
fenberg, in press, for details).


TABLE 2.-Scale variation in Varanus komodoensis from Loho Liang, Komodo
(N=50).

Scales Range X S.D.
Infranasal 4 4
Interoculars 5-9 6.7 0.9
Internasals 6-10 8.2 1.4
Mouth-mouth 48-51 49.3 1.8
Midbody 170-176 173.8 0.4
Ventrals 80-110 92.6 7.4
4th toe lamellae 7-8 7.3 0.2


Scale data of 50 Varanus komodoensis from Komodo are given in
Table 2. Upper head scales rounded and slightly raised, largest on
snout and smallest in temporal region; supraoculars not differentiated,
generally smaller than temporals; interparietal only slightly enlarged;
nape scales relatively large, generally conical; dorsal scales smaller
than those of neck, but larger than lateral body members, being long
and strongly keeled; lower leg dorsal scales weakly keeled; ventrals
elongated, varying from weakly keeled to smooth; all caudals strongly
keeled, dorsal ones smallest; complete caudal rings only in basal third
of tail; two median dorsal tail scale rows slightly enlarged, producing a
low double keel; inner surface of proximal part of fourth toe provided
with a row of conical lamellae, similar to those of Varanus various.
Apical pits (?) on most scales, as in other species (Smith 1935),
those anterior to vent and on thighs are larger and appear different
than those in other parts of body; cranial scale organs described in
Auffenberg (in press).
Color and pattern of three different-sized Komodo individuals (in
life) are as follows: In a hatchling (SVL 19.2 cm, tail 27.1 cm, Fig. 32)
head brownish-black dorsally with a few light yellow scattered spots;
iris light brown; neck ground color black with transverse darker bands
approximately 1500 to horizontal when viewed from side; body bands


1980








BULLETIN FLORIDA STATE MUSEUM


comprised of faint yellow spots, often forming ocelli medioposteriorly
and alternating with transverse rows of yellowish-tan or tan spots,
becoming ochre with yellowish centers mediodorsally. Body ground
color black with transverse rows of dark brown ocelli (10-12 across the
back) is a series of small yellow dots, usually in a single row. The tail
has 19 yellow bands on a grayish background; the last third is gray
laterally, yellow both dorsally and ventrally (reddish-brown medially
near its base); tail base gray laterally, interrupted by the bases of the
yellow transverse bands. Underside of hindlimbs light rust with small
gray dots. Anterior to the insertion of the hindlimbs belly yellowish
with rust-brown reticulations forming alternating wide and narrow
bands. Central; depressed area of the precloacal scale rosette brown.
Soles of hind feet pinkish-white, those of the front feet clear yellow.
Behind insertion of forelimbs on the ventral surface, the dorsal ocelli
bands pass over the chest and break into three transverse bars, each


FIGURE 28.-Head of adult Varanus komodoensis (UF 29001), Nggoer, Flores.


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


with two anterior projections. A black linear patch above insertion of
forelimbs. Dorsally thighs of hindlimbs black with transverse rows of
larger yellow spots alternating with rows of minute dark yellow dots.
Lower part of hindlimb black, changing to dark brown on foot with
scattered yellow dots and darker edges. Both fore- and hindlimbs have
yellow bands at each toe joint. Dorsal surface of front limb black with
transverse bands of small yellow dots. Claws black. A gray circle with
a yellowish-white center on side of head in front of the tympanum.
Yellow patches above and below eye. Three vertical yellow bars on a
brownish-black background between eye and nostril. Following jaw
angle from immediately behind tympanum to throat slightly pinkish-
white. Most of throat yellow with a tinge of ochre sprinkled with black
dots. Chin whitish-yellow with greenish-black spots.
The following description is based on 18W, a young male, body
length 69.1 cm, tail 84.8 cm. Dorsally head largely black with scat-
tered yellow spots. Nuchal area black with many scales having yellow
bases; a few entirely yellow. Hind- and forelimbs black above with
small scattered yellow spots. Forefeet entirely black above, hindfeet
with a few yellow spots. Body dorsally black with base of some scales



E
u
c 14

S 12 *
o 0. *

o
10 *




6
0 42 *

$J 2
I- 4 6 8 10 12 14 16 18 20 22 24

HEAD LENGTH(incm)
FIGURE 29.-Head width-length relationships of Varanus komodensis, all from Loho
Liang, Komodo.







BULLETIN FLORIDA STATE MUSEUM


reddish-brown and small yellow spots aligned in bands, larger and
more numerous laterally. Posterodorsally body and tail base more
rust-colored with both black and yellow spots; changing at more com-
pressed middle of tail to yellow-green with brown and black spots; last
2 cm brilliant yellow-green. Most of tail ventral surface yellow with a
short medial black stripe. Below tail base three fairly distinct smoky-
gray partial bands extending medially, but not to midline which is
yellow with a few black scales. Color ventrally near hindlimbs and
under thighs yellow to grayish-yellow; soles of hindfeet brilliant


E
u


I
0
Z
LU

0
0


150


125.


100


75


50


25


0
00


.s.
4*

0'*


50 75


100 125 150


TAIL LENGTH (in cm)
FIGURE 30.-Body-tail length relationships in Varanus komodoensis, all from Loho
Liang, Komodo.


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


yellow. Ventrolaterally body more rust-colored, becoming yellowish
ventrally with a scattering of black scales under forelimbs. Three
grayish-black shallow V's with apices pointed anteriorly on chest.
Neck base below speckled with black and yellow, black anteriorly. On
chin and anterior part of throat the black spots form narrow
longitudinal rows, largest being outer one immediately below lower lip.
Between these, scales clear yellow. A greenish-yellow crescent with
smoky-gray spots present along chin border. Laterally head grayish-
black, intense black temporally. Below and in front of nostrils a large
yellowish-gray blotch. Eyelids bright yellow. Sides of lower jaw below
mouth black to grayish-black with a few yellow spots.
The following description is based on 26W, body length 64.0 cm,
tail 86.4 cm. Head entirely black. Above, neck black, with yellow-green
interstitial skin. Back with black to brownish-black scales, with
orange, ochre, or sometimes yellow interstitial skin. Below, neck gray
to yellow. Dark chest bars still evident. Belly medially yellow, ven-
trolaterally with light orange countershading. Both front and hind
legs black with small yellow spots. Tail dark gray, lighter toward tip,
with yellowish-gray bands still evident.


FIGURE 31.--Scalation surrounding cloaca (=C) showing the two precloacal pores found
in male V. komodoensis (=P).


















0



























FIGURE 32.- Hatchling V. komodoensis, Loho Liang, Komodo (composite from photos and field sketches).


z
N1j
















pl








AUFFENBERG: KOMODO HERPETOLOGY


When Varanus komodoensis attain 2 m, they are almost uniformly
brown or grayish-brown dorsally on body, legs, and tail. Chest bars
light gray to absent. Interstitial skin of back rusty. Tail bands com-
pletely absent. Belly gray with faint speckling. Particularly old in-
dividuals with yellowish-green, pink, or even lavender irregular spots
on snout. Tongue pale yellow in individuals of all sizes. Iris always
brown.
DISTRIBUTION AND HABITAT.-Known only from Komodo, Padar,
Rintja, the western end of Flores, and the small islets of Gili Moto and
Oewada Sami (Fig. 33).
It is most abundant in ecotonal situations between savanna and
monsoon forests but is common in both, occasionally in steppe and
mangrove forests. It does not normally occur in the quasi-rain forest
community, which usually begins at an elevation of about 500 m.
Young are largely arboreal during their first year of life and often
climb high trees in search of shelter or food. The adults are largely ter-
restrial and rarely climb. They occasionally enter brackish or sea
water. Shelter for the young is found under loose bark and in hollow
trees. Adults may construct burrows up to 3 m long (usually much
shorter) along vertical banks of stream beds or beneath boulders and
fallen trees. The burrows are sometimes distinctly clustered. In-
dividuals spend the night sleeping in high grass, dense thickets of
bush and saplings, under overhanging shrubs or in the burrows. Large
adults use specific thermoregulatory sites, usually on hillsides, that
overlook valleys or coastal areas.
REPRODUCTION.-Data from Brongersma (1932), Lederer (1942),
Hisada (1966), Galstaun (1973), Auffenberg (in press), and Prato (MS)
constitute the basis for the following conclusions. From 1-30 eggs (X
= 11.3) are laid in a nest(s) excavated in the soil. Females may lay up
to three clutches/year from April to September (usually June-August)
and interclutch time varies from 1-8 days (X = 2.5). Incubation period
varies from 21/2-8 months, probably dependent on both temperature
and soil moisture. Eggs vary in weight from 55-210g (X = 124.7), and
their size from 59-115 mm (X = 86.5) x 42-66 mm (X = 55.5). The
young normally appear in April through May, and there is some
evidence that they remain together in small groups for several months.
Adult males have been known to eat the eggs, and there is a report
that some females may return to the nest and open it about the time
that the young hatch (Pfeffer 1959). Males and females apparently
become mature in the 5th to 7th year, at a length of approximately 1.5 m.

















FLORES SEA


.ANTA




P
o


w




z


0



cj3

Tj
p

S


OEWADA SAMI


SUMBA SEA


FIGURE 33.-Known distribution of V. komodoensis (stippled area). On Flores Island the hollow dots represent native reports and solid ones
actual sightings or other proof of occurrence.


,,








AUFFENBERG: KOMODO HERPETOLOGY


FooD.-Highly opportunistic, this species feeds upon a wide vari-
ety of prey (Table 3). While arboreal, the young feed chiefly upon
smaller lizards, insects, birds and their eggs; medium-sized individuals
eat mainly rodents and larger native birds (such as Megapodius
freycineti, Lincoln 1974; Auffenberg, in press). These are all captured
by active pursuit and/or search. Medium to large adults frequently
feed on carrion, though the largest individuals also prey on wild boar
and deer, which they usually catch from ambush along game trails.



TABLE 3.-Prey represented in Varanus komodoensis fecal pellets (N=4267) collected
on Komodo.

Prey eaten Number occurrences

INVERTEBRATES
Land molluscs (Asperites trochus) 3
Marine clam (sp.?) 1
Beetles (sev. sp.) 15
Grasshoppers (sev. sp.) 56
Crabs (sev. sp.) 3
FISHES (sev. sp.)
REPTILES
Ora (Varanus komodoensis) 66
Gecko (Gekko gecko) 24
Gecko (Hemidactylus sp.) 3
Skink (Sphenomorphus sp.) 3
Bamboo viper (Trimeresurus albolabris) 8
Rat snake (Elaphe subradiata) 29
Whip snake (Dendrelaphis pictus) 43
Cobra (Naja naja) 3
Russell's viper (Vipera russelli) 15
Hawksbill sea turtle (Eretomchelys imbricata) (+ eggs) 3
BIRDS
Megapode (Megapodius freycineti) (+ eggs) 48
Jungle fowl (GaUus various) (+ eggs) 4
Misc. small bird species 71
MAMMALS
Bat (sp. ?) 2
Mouse (Mus musculus) 36
Rat (Rattus rattus) 196
Feral dog (Canis familiaris) 8
Feral goat (Capra hircus) 26
Palm civet (Paradoxurus hermaphrodytes) 20
Mongoose (Herpestes javanicus ?) 4
Wild boar (Sus scrofa) 808
Sunday deer (Rusa timorensis) 1979
Feral water buffalo (Bos bubalis) 15


1980









BULLETIN FLORIDA STATE MUSEUM


Animals as large as horses and water buffalo are successfully attacked
and some individuals, particularly on Flores, regularly attack village
livestock. Humans are occasionally attacked and sometimes killed and
eaten (see Auffenberg 1977, in press, for details).
PARASITES.-Ectoparasites include three species of ticks:
Amblyomma helvolum, A. robinsoni, and Aponomma komodense, of
which A. robinsoni is most common. No sporazoans were found in
blood samples taken from four Flores specimens. Subcutaneous larval
pseudophyllidean cestodes were very common in Flores specimens
(estimated density 150/1.5 m adult V. komodoensis). They were not
found on any Komodo specimens. The larger larvae formed lumps
under the skin and were scattered over most of the dorsal part of the
body. The monitors were probably infected by eating aquatic
vertebrates (essentially absent on Komodo). All fecal specimens ex-
amined showed a low density of non-pathogenic amoeba (Endolimax
sp.) and cestode ova. Mature intestinal cestodes included the

TABLE 4.-Parasites of V. komodoensis and western Flores populations of V. salvator.

Parasites Place V. komodoensis V. salvator


AMOEBA
Endolimax sp.
SPOROZOA
Hemagregarine
CESTODA
Pseudophyllidae
Larvae (Sphargana type)
Duthiersia expansa
Duthiersia venustal
Duthiersia crassa
Duthiersia sp. indet.
Spirometra sp. indet.
Scyphocephalus bisulcatus
Proteocephalidea
Kapsulotaenia sandgroundi
Acanthotaenia shipleyi
NEMATODA
Spiruroidea
Tanqua sp.
ACARINA
Amblyomma helvolum
Amblyomma robinsoni
Aponomma komodense


fecal

blood


subcutaneous
intestinal
intestinal
intestinal
intestinal
intestinal
intestinal

intestinal
intestinal


gastric

skin
skin
skin


L=D. sarawakensis


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


pseudophyllideans Spirometra sp., Duthiersia cf D. expansa, D.
venusta (or sarawakensis), and the proteocephalidean Kapsulotaenia
sandgroundi.
The endoparasitic data available (Table 4) suggest a rather exten-
sive fauna, of which certain elements are shared between V. komodoen-
sis and V. salvator, whereas others are not. In general, V. komodoensis
is not heavily parasitized; V. salvator shows much higher density and
diversity.
REMARKS.-Called mainly ora in the local dialect and buaja darat
(= "land crocodile") in Malay by people not well acquainted with it.
Additional data are provided by Dunn (1927a), Burden (1928), Mertens
(1929c), Lederer (1933), Pfeffer (1959), and Darevsky and Kadarsan
(1964).


Varanus salvator salvator (LAURENT)
Figures 34-35
Lacerta monitor Linne 1758 (part) (Type locality: "Indies").
Stellio salvator Laurent 1768:56 (Type locality: "America," in error).
Monitor nigricans Cuvier 1829:27 (Type locality: Java).
Hydrosaurus salvator Gray 1838:394.
Monitor bivittatus var. javanica Schlegel 1844:10 (Type locality: Java).
Monitor bivittatus var. celebensis Schlegel 1844:10 (Type locality: Celebes).
Varanus crocodilinus Owen 1845:265 (Type locality: restricted to Java).
Varanus salvator Cantor 1847:635.
Varanus togianus Weber 1890:169.
Varanus salvator salvator Mertens 1937:178.
SPECIMENS EXAMINED.-(12) UF 29002-3, 30171, 40432, Nggoer, Flores, 2-5 m; MZB
941, Udjung Kulon, Java; MZB 942, Kerto Malang, Java; MZB 943, Bogor, Java; MZB
947, Liroeng, Salibaboe; MZB 949, Atcheh, Sumatra; MZB 952, Madiun, Java; MZB
954, Liroeng, Salibaboe; and MZB 957, Udjung Kulon, Java.
DESCRIPTION.-Dorsal head scales large, flat, smooth, largest in
supraocular region (Fig. 34); supraoculars well differentiated, 4-6 in
Flores specimens (4-8 elsewhere; fide Mertens 1942); occipitals larger


FIGURE 34.-Head of adult Varanus salvator (UF 29002), Nggoer, Flores.


1980








BULLETIN FLORIDA STATE MUSEUM


than temporals; 45-48 scales across top of head from angle of mouth on
one side to same place on other (48-60 elsewhere); neck scales smaller
than occipitals, only slightly larger than the strongly keeled oval dor-
sal body scales; 148-153 scales at midbody; ventral scales rectangular,
larger than dorsal scales, keeled, except near breast, 80-85 from gular
fold to insertion of hindlimb; tail scales keeled dorsally and ventrally,
not in obvious marked transverse rings, the two dorsovertebral rows
enlarged, their keels providing a low double caudal ridge.
Ground color brownish-black with five faint (even in the young)
crossbands produced by yellowish-white-edged scales, forming very ill-
defined ocelli (rarely seen in live animals but evident when dried skins
are held to the light), thus unlike the vividly marked Sumbawa
specimens figured in Mertens (1942); feet black except (usually) for a
yellow dot on dorsal surface of each toe just behind claw; tail almost
uniformly grayish-black above, without obvious lighter colored bands;
belly light yellow, with 8-10 faint black bars extending medially from
sides, but not meeting at midline; throat with 3-4 black crossbands; a
few black dots on anterior part of throat, chin provided with 2-3 nar-
row black cross bars that become faint in adults; sides of head yellow
with 4 vertical brownish-black bars extending onto lower jaws, becom-
ing faint from general darkening with age; dorsally head brownish-
black with 3 darker lateral head bars extending over snout and down
other side, but observable only in juveniles, head almost uniformly
black in adults.
Although this species grows to nearly 3 m on the Asian mainland,
Mertens (1942) pointed out the somewhat smaller size of specimens
from the Lesser Sunda Islands, with which I agree (X SVL of 3 Flores
adults = 42.5 cm, total length 90.0-134.5, X = 115.8 cm).
DISTRIBUTION AND HABITAT.-V. salvator has been reported from
Sumba (Barbour 1912, Mertens 1930, 1942, Forcart 1949), Flores (Bar-
bour 1912, Mertens 1942), and Sumbawa (Mertens 1942). The report
by Hoogerwerf (1954) that it occurs on Padar based on footprints in
the sand is ill-founded. I doubt that it exists there for this island lacks
appropriate habitat. DeJong (1937) believed there were no differences
in the spoor of V. komodoensis and V. salvator. The latter is clearly ab-
sent on Komodo; undoubtedly for the same reason, for on surrounding
islands V. salvator is common near streams and permanent large
ponds. On Flores, we rarely saw it farther than 100 m or so from the
water's edge, even when we attempted to draw it into the surrounding
savanna with carrion bait. However, Varanus komodoensis, the local
savanna and deciduous forest species, is easily drawn into the stream-
side habitat of V. salvator with the same bait.


Vol. 25, No. 2










AUFFENBERG: KOMODO HERPETOLOGY


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FIGURE 35.-Repeated sightings (dots) of four adult Varanus salvator along Nanga
Look in western Flores. Approximate boundaries of activity ranges are shown. A, B,
and C show location of carrion baiting sites. V. salvator specimens came to site A,
whereas V. komodoensis fed at all the stations.


" v" ^ f^
j" rfIS V-'j








BULLETIN FLORIDA STATE MUSEUM


At night they sleep on bushes overhanging the water or in burrows
excavated in the stream banks.
On Flores, V. salvator occurs from brackish coastal mangrove
swamps to elevations of at least 1000 m, dependent largely on the
distribution of the surface water. In general, there is no correlation
between its distribution and river bank ecology-all types of riverine
habitats are exploited. In agricultural areas, they are often found near
rice fields. Density along Nanga Look was estimated at 15/km along
the river above high tide effects, 8/km in coastal Lumnitzera and/or
Rhizophora mangrove forests. The core area of the adult activity
ranges are usually separated by about 500 m (Fig. 35).
FooD.-Forcart (1949) described the protection of its young by a
marsh hen (Gallinula sp.) against the attack of a large monitor and also
a monitor's attack on a wild duck (Anas giberifrons). On Flores the
water monitor is an opportunistic predator, feeding on a great variety
of live prey, carrion, and even human feces. Living prey locally record-
ed are snakes (Dendrelaphis pictus, Trimeresurus albolabris, Cerberus
rynchops), frogs (Rana cf cancrivora), lizards (Sphenomorphus sp.,
Lamprolepis smaragdina), mammals (Rattus sp., Mus musculus),
wading birds (Bucoroides sp., Nothophyx sp.), and domestic ducklings
and chicks. My data from Nanga Lo6k, a small river in extreme
western Flores, suggest that they usually forage within 40 m of the
water's edge, but during the monsoon season they move at least 500 m
inland.
PARASITES.-The only ectoparasites found on Flores specimens
were Amblyomma helvolum' (also commonly parasitized local
Varanus komodoensis). A hemogregarine infection sporozoann) was
found in 25% of the Flores V. salvator examined2. Subcutaneous infec-
tion of larval cestodes (Order Pseudophyllidea; sphargana type) were
exceedingly abundant in all V. salvator from the Nggoer, Flores, area,
probably ultimately derived from frogs2. Intestinal parasites included
a heavy infestation of the pseudophyllidian cestode Duthiersia (D. ex-
pansa in 50% of the Flores material, and an undescribed species2). One
other pseudophyllidian of the genus Scyphocephalus (bisulcatus3) was
found, as well as a protocephalidian of the genus Acanthotaenia
(shipleyi3). Spiruroid nematodes (Tanqua sp.) were found in 66% of the
specimens examined. Parasite data for Varanus salvator and V. komo-
doensis are summarized in Table 4.

'Identified by H. Hoogstral.
'Identified by S. R. Telford
'Identified by J. L. Pinnell


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


REMARKS.-Hoogerwerf (1954) and de Jong (1937) stated that eggs
and young of V. salvator are eaten by V. komodoensis. On Flores, I
found that when both species were attracted to carrion at the same
time the larger V. komodoensis dominated the relationship usually
with no or few overtly aggressive displays. Local villagers stated that
when carrion is completely eaten large V. komodoensis occasionally at-
tacked and ate the smaller V. salvator that were also attracted to the
carrion.
Varanus salvator is called biawak in Indonesian, weti in the local
Mangarrai dialect, and alu in Balinese.

SUBORDER SERPENTES
FAMILY TYPHLOPIDAE
Typhlina bramina bramina (DAUDIN)
Figure 36
Eryx braminus Daudin 1803:279 (Type locality: Vizagapatan, India).
Typhlops braminus Cuvier 1829:73.
Typhlops braminus braminus Mertens 1930:278.
SPECIMENS EXAMINED.-(40) UF 29406, 29410, 29424-38, 29441-48, 29450-51,
29454-56, 29458, all from Loho Lavi, Komodo, 20 m; UF 32600, Nggoer, Mangarrai
District, Flores, 12 m; UF 33058, Bara, Flores, 200 m.
DESCRIPTION.-Snout rounded, no keel; nostril lateral; rostral ex-
tending to level of eyes or beyond (shorter in most other populations
according to de Rooij 1915); nasal cleft completely divided; preocular
present, nearly- to as large as the ocular, in contact with the second
and third supralabials; 4 supralabials; eyes distinct. Body with 20 mid-
body scale rows. Greatest body diameter in total length (Komodo
specimens): adults 34.7-56.9 midbody (X = 59.2). Tail as long as or a
little longer than broad, ending in a spine.
Color brown, brownish-gray, or blackish-brown above, lighter
below. Snout, chin, cloacal area, and tail tip often cream or whitish.
Total length 79-165 mm (X = 135 mm) in Komodo area, to 173 mm in
Sumbawa (Mertens 1930).
DISTRIBUTION AND HABITAT.-In the Indoaustralian area,
Typhlina bramina has been reported from Australia, Sumba, Sum-
bawa, Komodo, and Flores (Mertens 1930, Forcart 1949).
Mertens (1930) stated that this blind snake is never found above
500 m in this area. On Komodo it is common in gallery forest areas
near sea level. Many of our specimens were obtained by villagers while
preparing garden plots. The snakes were found beneath the surface,
usually within the upper 15 cm. Other specimens were collected from
under stones along dry stream courses and in gallery forests. On
Flores, one specimen was collected at night, crawling in the grass
across the surface of a Zizyphus savanna.







BULLETIN FLORIDA STATE MUSEUM


REPRODUCTION.-Two to four elongate eggs (17.5 x 3.3 mm) are laid
during the wet season (January dates recorded on Komodo) by females
at least 130 mm SVL.
FooD.-The guts of all specimens examined (12) contained only ter-
mites.
REMARKS.-Called ular duwel locally.
Typhlina polygrammica undecimlineata MERTENS
Typhlops florensis undecimlineatus Mertens 1927c:239 (Type locality: Semongkat,
Sumbawa, 400 m).
Typhlops polygrammicus undecimlineatus Forcart 1949:377.
SPECIMENS EXAMINED.-(1) Uncatalogued juvenile in the private collection of the
late Hilmi Oesman, from Komodo, "in mountains" (precise location of specimen
unknown).
REMARKS.-Darevsky (1964a) recorded this subspecies from
Komodo on the basis of a subadult collected beneath stones in moist
forest at 400 m. Both his specimen and that in Oesman's collection had
11 dark brown dorsal, longitudinal stripes on a light yellow ground col-
or. The specimen that I examined had a clear yellow venter, 22 scales
rows at midbody, and a midbody diameter that went into the total
length 39.6 times. The subspecies differs from its closest relative, T. p.
florensis, in lacking stripes on the venter. T. p. brongersmai from Sum-
ba is uniform dark blackish-gray above and yellow, white, or pink
below (Mertens 1928a).
Typhlina polygrammica has been taken from 400 to 1000 m
(Mertens 1927c); above the major part of the activity range of Varanus
komodoensis, and thus infrequently visited by our party. Darevsky
(1964a) reported a single Komodo specimen collected at 400 m.

Typhlops schmutzi new species
Figure 36

HOLOTYPE.-UF 29507, adult female, Loho Lavi, Komodo, 12 m, collected January 1970
by expedition members.


FIGURE 36.-Left, lateral head view of Typhlops schmutzi (new species, UF 294323)
Loho Lavi, Komodo; right, Typhlina bramina bramina (UF 29424), Loho Lavi, Komodo.


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


PARATYPES.-(84) UF 29415-23, 29452, 29466-95, 29500-34, all from Loho Lavi,
Komodo, 12 m; UF 29411-13, 29461-65, all from Loho Liang, Komodo, 0-30 m; UF
29414, Kampung Komodo, Komodo, 5 m; UF 37018, Poto, Flores, 52 m.
DIAGNOSIS.-Differing from all other described species of the genus
in possessing, in combination, extreme slenderness, no subocular,
nasal cleft arising.from second supralabial, and a completely divided
nasal.
DESCRIPTION OF HOLOTYPE.-Snout rounded, no horizontal keel,
projecting; nostrils lateral. Rostral parallel-sided, dorsal portion about
/3 width of head, just barely or not reaching level of eyes; nostril in a
divided nasal, anterior one in contact with rostral and first two
supralabials, posterior one with second supralabial, preocular, and
rostral; preocular present, nearly as large as ocular, in contact with
supralabials 2 and 3. Eyes distinct. Scales over eye larger than body
scales, median head scales smaller than those on body. Four
supralabials. Body with 20 middorsal scale rows. Greatest body
diameter (1.8 mm) 70.6 times in total length (127 mm). Tail slightly
longer than its base, ending in a small spine.
Color pinkish-brown, slightly lighter below; snout and underside of
tail and its tip whitish.
VARIATION.-Scalation and coloration are remarkably constant in
all the paratypes. Minor variations in scale proportions occur in the
rostral, nasals, and preocular, but none is significant. Body propor-
tions (greatest diameter into total length) 62.5-81.9 (X = 72.3). Tail
length '/ to 2 times the width of the base, caudal scales 10-12. Range
in total length 58-140 mm. Color of juveniles only slightly lighter than
that of the adults.
DISTRIBUTION AND HABITAT.-Known only from localities near sea
level on Komodo and the north coast of Flores. All the type material
was collected from the soil, usually within 15 cm of the surface.
REMARKS.-Named for Rev. E. Schmutz, a talented naturalist of
Nunung Mission, Mangarrai District, Flores.

FAMILY ANILIIDAE
Cylindrophis opisthorhodus BOULENGER)
Figure 37
Cylindrophis opisthorhodus Boulenger 1897:506 (Type locality: Lombok, 500 m).
SPECIMENS EXAMINED.-(1) UF 28763, Loho Lavi, Komodo, 10 m.
DESCRIPTION.-Eye diameter into distance from mouth 2 times, in
distance to nostril 4 times; distance between eyes more than distance
from eye to nostril; frontal as wide as the single supraocular, broader
than the parietals; nasals in contact, prefrontals in contact; 6
supralabials, third and fourth in contact with the eye; scales in 23 rows
middorsally, 21 anterior to the cloaca; ventrals only slightly enlarged,
216 (184-213 elsewhere), subcaudals 5 (5-7 elsewhere).


1980








BULLETIN FLORIDA STATE MUSEUM


Color iridescent light brown above, with very narrow dark brown
vertebral stripe, few very scattered dots laterally, tending to be ar-
ranged ie a row on the third or fourth scale row on either side of
vertebral stripe, often in groups of 2 or 3 anteriorly. Head pale brown
above, with dark brown blotches, ground color darkening to rust
brown from supraoculars to a point about one head length behind
supraoculars, with cream vertical temporal bar outlined by a broken
black edge, cream flecks in front of eye. Black to dark brown ven-
trolaterally, with large paramedian white markings, tending to form ir-
regular bars across belly, or an irregular checkerboard in some places.
Dorsal color pattern separated from that of venter by a black line on
1-2 scale rows. Anal region black with a white spot on either side of
anal plate. Tail marked somewhat like head (local Malay name, ular
kepala dua, the two-head snake), but red ventrally (faded to white in
alcohol). Mertens (1930) stated that the pattern of the young is more
contrasting than that of larger individuals. Total length of Komodo
specimen 276 mm (maximum 283 mm, Sumbawa specimen, Mertens
1930).
DISTRIBUTION AND HABITAT.-This is the first record from
Komodo; previously reported localities are western Flores, Lombok,
Sumbawa, and Flores. Mertens (1930) stated that all the known
specimens have so far been near sea level. Ours was found about 10 cm
below the surface in clay soils of a gallery forest (a second specimen,
captured in the same place and under the same conditions, escaped
before preservation).


SD
FIGURE 37.-Cylindrophis opisthorhodus (UF 28763), Loho Lavi, Komodo. (A and B)
dorsal and lateral views of head; (C) mid-ventral pattern; and (D) lateral view of the tail
(note resemblance to lateral head view).


Vol. 25, No. 2


116


A








AUFFENBERG: KOMODO HERPETOLOGY


FAMILY ACROCHORDIDAE
Chersydrus granulatus (SCHNEIDER)
Hydrus granulatus Schneider 1799:243 (Type locality not designated).
Anchrochordus granulatus Cantor 1847:59.
Chersydrus granulatus Gray 1849:61.
Achrochordus granulatus granulatus Loveridge 1948:380.
SPECIMENS EXAMINED.-(7) UF 39776-77, Suwong, Bali, at sea level; UF 28775-77,
28779, Loho Lavi, Komodo, at sea level; UF.28778, Kampung Komodo, Komodo, at sea
level.
DESCRIPTION.-Head not distinct from neck; covered with small
juxtaposed granular scales; nostrils on top of snout, close together, no
rostral; a series of larger shields on lips, separated from mouth by a
row of very small scales; no chin shields; about 100 scales around body,
of which dorsals are longer, no widened ventral scales; tail short, lat-
erally compressed, prehensile.
Gray to gray-brown with dark brownish-black to black transverse
rings.
DISTRIBUTION AND HABITAT.-Widely distributed, found in large
rivers and along sea coasts from northwestern India to Solomon
Islands, not previously reported from Komodo; de Rooij (1917) listed it
for Flores.
REMARKS.-Komodo specimens were obtained at night during low
tide from mangrove forests dominated by Rhizophora mucronata, in
the west end of the bay called Telok Slawi. Most were first seen when
they were swimming actively, apparently having been frightened by
our lights and movement. When at rest, only the tip of the nose is held
above the water. The tail is coiled about twigs, stones, or roots, and
most of the weight is carried by the widest part of the body (the
posterior third), which rests on its side on the bottom. The anterior
two-thirds of the body ascends to the surface, easily moved back and
forth by the movements of the water.
One of the Bali specimens, collected in September, contained six
embryos that would probably have been born in late October.
Mature Komodo males are at least 70 cm SVL; females are 76 cm
and much heavier-bodied.
FAMILY COLUBRIDAE
Boiga cynodon (BOIE)
Figure 38
Dipsas cynodon Boie 1827:549 (Type locality: Sumatra).
Pareas waandersii Bleeker 1860:471 (Type locality: Bali)..
Dipsadomorphus cynodon Boulenger 1896:78.
Boiga cynodon Mertens 1930:315.
SPECIMENS EXAMINED.-(6) UF 28681, 28684-85, Loho Liang, Komodo, 0-30 m; UF
28682, Kampung Komodo, Komodo, 5 m; UF 28683, Kampung Sabita, Komodo, 12 m;
UF 36203, Nangahele, Mangarrai District, Flores.


1980







BULLETIN FLORIDA STATE MUSEUM


DESCRIPTION.-Snout longer than eye diameter; rostral broader
than high, just visible from above; anterior palatine and mandibular
teeth greatly enlarged; internasals broader than long, shorter than the
prefrontals; frontal as broad as long, somewhat shorter than its
distance from snout tip, shorter than parietals; loreal broader than
high; one preocular, narrowly separated from frontal; two postoculars;
temporals 2(3)+4; 9-10 supralabials, of which 4-7 enter eye; 4 in-
fralabials in contact with anterior chin shields; dorsal scales in 19
rows, median row greatly enlarged; ventrals 247-261 (X = 257)
(248-290 elsewhere), subcaudals 130-136 (X= 129.2) (114-159
elsewhere), anal entire.
Ground color grayish-brown to reddish-brown, with 35-37 irregular
dark brown crossbands, often with black edges, sometimes a faint
band or spot in ground color. Head nearly uniform light gray to light
brown, without dorsal black n-shaped mark found in specimens farther
west. Most have a narrow, partially black postocular stripe. Belly
pinkish-brown, chin and supralabials white. Total lengths of five
Komodo adults range from 102.1 cm to 171.7 cm.
DISTRIBUTION AND HABITAT.-This is the first record ofB. cynodon
from Komodo. Previous nearby localities are Flores (de Rooij 1917)
and Sumbawa (Mertens 1930). From Flores its range extends
westward through the entire archipelago north to India and the Philip-
pines.
All specimens of this arboreal, nocturnal snake were found in lontar
and coconut palm trees, where they remained as high as 15 m in the
bushy tops during the daytime. Birds commonly roosted in the palms
at night; the snakes apparently preyed on the sleeping birds, for we
noted the snakes were often active at night. One of our specimens fell
out of a lontar palm when the wind dislodged some of the dead fronds.


FIGURE 38.-Boiga cynodon (UF 36203), Nangahele, Flores.


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


REPRODUCTION.-The Komodo data are important, for reproductive
behavior was previously unknown (Leviton 1968). On the basis of the
following data, 1-23 eggs are laid during the monsoon season. One
female specimen collected in October had a total of 26 follicles, of
which 15 were small but with much evidence of yolk deposition; the re-
maining ones were small and translucent. The oviducts were smooth,
flat. Two adult females were dissected in January; one showed 10
follicles, of which only one had advanced yolk deposition; the others
were small, flat, and translucent. The ovaries in this specimen were
small and flat. This female (SVL 78.9 cm) may have been in her first
laying year. The second female (133.0 cm) had 55 follicles, of which five
showed slight evidence of yolk deposition. That this female had recent-
ly laid her eggs was indicated by 23 still red oviductal scars. The
oviducts were very thick-walled but not very convoluted.
FOOD.-Of six specimens examined for food, one contained a Draco
volans and two had bird feathers in their digestive tract. Another was
eating a Gekko gecko when captured.
Cerberus rynchops rynchops (SCHNEIDER)
Figure 39
Hydrus rhynchops Schneider 1799:246.
Homalopsis rhynchops Cantor 1847:94.
Cerberus rhynchops Giinther 1864:279.
Hurria rhychops Stejneger 1907:304.
Hurria rhynchops Barbour 1912:123.
Cerberus rhynchops rhynchops Loveridge 1948:388.
SPECIMENS EXAMINED.-(61) UF 28601, 28604-5; 28608-12, 28614-15, Loho Lavi,
Komodo; UF 28598, 28600, (with 8 embryos), 28606-7, 28613, Sabita, Komodo; UF
30161, Nangahele, Mangarrai District, Flores; UF 30152-60, Nggoer, Mangarrai
District, Flores; UF 36171-79, 361867-95, Suwong, Bali; UF 36180-82, Benoa, Bali.
DESCRIPTION.-Frontal distinct or broken into small scales, nasal
cleft in contact with supralabials 2-4; eye bordered by 5-6 scales, 1-2
postoculars, 1-2 suboculars; supralabials 10, rarely 9; infralabials
13-15, usually 14, 3-4 in contact with anterior chin shields; dorsal
scales strongly keeled, in 23, 25, or 27 rows at midbody; ventrals on
Komodo 148-153 (X = 150.5), 122-160 elsewhere; subcaudals divided,
54-66 (X = 58.2), 49-72 elsewhere.
Color olive to dark brown above, with black spots or transverse
bands, sometimes indistinct; a light yellow to gray lateral band; black
to brown postocular stripe passing through the eye onto the side of the
neck; ventrally black, with a single row of yellowish to gray spots on
each side; the darker chin stripes absent or very faint in all Komodo
specimens. Newly hatched young are similar, though generally
brighter patterned. The postocular stripe is sometimes represented by
only a row of brown dots or a broken stripe.







BULLETIN FLORIDA STATE MUSEUM


SVL of mature individuals in the Komodo region is over 550 mm,
maximum 750 mm; Hatchlings from this area (SVL 116-128 mm,
X = 121) are much smaller than those from mainland Asia (130-160,
fide Smith 1943).
TAXONOMIC REMARKS.-The Komodo material is clearly referable
to Cerberus rynchops rynchops on the basis of the combination of scale
and color characters. Other subspecies are known from the Philippine
Islands (C. r. microlepsis), Papua (C. r. novaeguineae), and Australia (C.
r. australis), and Loveridge (1948) gives a key and discussion.
DISTRIBUTION AND HABITAT.-On Komodo this species is found
chiefly in shallow mangrove swamps dominated by Rhizophorus
mucronata or Sonnertia sp. Hoogerwerf (1954) reported finding one in
shallow water off the sand beach near Nangelele, Komodo. On both
Komodo and Bali it was frequently seen at night during low tide,
where it moved about within tidal pools and shallow creeks searching
for fishes and crabs. At low tides during the day it usually spends
most of the time in crab and fish burrows.
It is widely distributed in appropriate habitats from the west coast
of India to northern Australia. In the Komodo area it has previously
been reported from Flores (de Rooij 1917), Sumba (de Rooij 1917, For-
cart 1949), and Komodo (Hoogerwerf 1954).


FIGURE 39.-Cerberus rynchops rynchops (UF 30159), Nggoer, Flores.


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AUFFENBERG: KOMODO HERPETOLOGY


REPRODUCTION.-The sex ratio of 36 specimens examined is 1 to 12
in favor of females-a ratio that may reflect differing activity patterns
more than actual ratio differences. Eight near-term embryos were
found in females collected on Komodo in November. Total atretic
follicles in 10 mature females taken November to January vary from
18-62 (X = 45.1), total corpora lutea 5-36, and enlarged follicles show-
ing yolk deposition 6-38 (X = 20.5). All available evidence suggests
that one litter a year is usual in the Komodo area. The young, which
number from 5 to 38 per litter (X = 19.5), are brought forth from late
November through February. The testes and vasa deferentia of the
males examined showed developmental stages reflecting a similar
seasonal reproductive cycle. The SVL of mature females varies from
550-750 mm (X = 590) and for males 570-600 mm (X = 590).
FooD.-In all 18 specimens examined the guts of 50% contained
food remains, and 66% of these contained the following identifiable
parts: fishes of the families Gobiidae (Amblygobius sp.) 33% and
Synodontidae (Synodus ivermanni) 33%, unidentifiable fishes 33%,
unidentified Crustacea (genus unknown) 25%, and pistol shrimp
(Alphaeus sp.) 12.5%.
PARASITES.-NOne noted.
Dendrelaphis pictus intermedius (MERTENS)
Figure 40
Dendrophis pictus Lidth de Jeude 1890:179 (part, in Weber).
Ahaetulla boiga intermedia Mertens 1927b:241 (Type locality: Sumbawa Besar,
Sumbawa).
Dendrophis pictus intermedius Meise and Hennig 1932:288.
Dendrelaphis pictus intermedius Mertens 1934b:199.
SPECIMENS EXAMINED.-(25) UF 36198, Djarek, Flores, 80 m; UF
30092, 30164-66, 30170, Nggoer, Flores, 20 m; UF 30167, 2 km WNW
Nuneng, Mangarrai District, Flores, 900 m; UF 30163, 4 km ENE
Look, Flores, 150 m; UF 28738-40, 28742, 28744-52, 30168-69, Loho
Liang, Komodo, 1-30 m, UF 28741, 28743, Gunung Ara, Komodo, 500
m.
DESCRIPTION.-Rostral much broader than high; internasals about
as long as prefrontals; frontal slightly shorter than its distance from
rostral, shorter than parietals; loreal much longer than high; one
preocular; supralabials 8-10, of which numbers 4-6 border eye; 5 in-


FIGURE 40.-Dendrelaphis pictus intermedius (UF 30170), Nggoer, Flores.


1980


121







BULLETIN FLORIDA STATE MUSEUM


fralabials in contact with anterior chin shields; scales keeled, in 15
rows middorsally, the median vertebral row very enlarged; ventrals
185-208 (X = 196.9) on Komodo, 182-200 elsewhere; subcaudals
125-163 (X = 149.3) on Komodo, 143-160 elsewhere.
Ground color dark olive-green to brownish-green (sometimes with a
faint metallic sheen); interstitial skin blue to black. A characteristic
black longitudinal stripe beginning between eye and nostril and
extending onto neck (Fig. 39), where it breaks up into flecks and disap-
pears. A yellowish-green line on outer lateral scale rows. Ventrals
olive-green; supralabials and chin yellowish-white. Juvenile color and
pattern as in adults.
DISTRIBUTION AND HABITAT.-This subspecies is restricted to Sum-
bawa, Flores, and Komodo (Mertens 1927b, 1934b, de Haas 1950,
Meise and Hennig 1932); Darevsky (1964a) listed Padar and Rintja. D.
p. inornatus is known from Sumba and Timor on the outer volcanic arc
(Forcart 1949) and D. p. pictus from Lombok and Celebes westward to
western India and north to southern China. The Komodo specimens
were all collected from ecotonal situations, between monsoon forest
and savanna, usually in low bushes or near them, through which they
could move very quickly. On Komodo they are found from near sea
level to 650 m (latter according to Dunn 1927b).
REPRODUCTION.-The sex ratio of 15 specimens examined is 2:13 in
favor of the females and, if correctly reflecting the local situation, is
very skewed. It is much more likely that females are more often
observed than males, and that the sex ratio is nearly equal. Two
shelled ova (34.8-40.0 x 7.6-9.2 mm) were found in a female taken on
Komodo in July. The total number of enlarged yolked follicles varies
from 2-18 (X = 9.3). Follicles show gradual increase of size and yolking
during the dry and early wet seasons (August through January). The
vasa deferentia of the male collected in September showed recent ac-
tivity, while those of an individual taken in January showed inactivity;
both specimens were mature and nearly the same size. Mature females
have an SVL of 530-700 mm (X = 626), males 510-580 (X = 560).
Mertens (1930) reported four females with 5-12 eggs in April and June
and suggested they may lay earlier on Sumbawa than on Komodo.
FooD.-On Komodo they appear to subsist primarily on lizards.
Darevsky (1964a) reported that a Padar specimen had eaten a
Lepidodactylus lugubris. Mertens (1930) reported some specimens of
this subspecies had eaten two species of the genus Rana. Of the 14
Komodo specimens examined 43% contained food remains in the gut,
which included Hemidactylus frenatus (50% of those containing food),
Sphenomorphus sp. (33%), and unidentified lizards (7%).
PARASITES.-None obtained.


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Elaphe subradiata subradiata (SCHLEGEL)
Figures 41, 42

Coluber subradiatus Schlegel 1837:139 (Type locality: Timor)
Elaphe subradiatus Giinther 1858:95 (part).
Elaphe subradiata Barbour 1912:31.
Elaphe subradiata subradiata Mertens 1930:298.
SPECIMENS EXAMINED.-(14) UF 39779:80 Nggoer, Flores, 12 m; UF 39778, Djarek,
Flores, 80 m; UF 36202, 4.5 km E Maumere, Flores, ca 10 m; UF 28753, Loho Lavi,
Komodo, 18 m; UF 28754, "Komodo"; UF 28756, 28759-62, Loho Liang, Komodo, 5-20 m;
UF 28755, Gunung Ara, Komodo, 300 m; UF 28757, Kampung Komodo, Komodo, 5 m.
DESCRIPTION.-Head long, somewhat distinct from neck; eye large,
pupil round; rostral broader than high; internasals shorter than
prefrontals; frontal as long as its distance from tip of snout, shorter
than the parietals; loreal deeper than long; preocular large; a
subocular; two postoculars; temporals 2-2 or 2-3; supralabials 9, 5th
and 6th entering eye, 4th very small; 5 infralabials in contact with the
anterior chin shields, latter shorter than the posterior pair. Scale rows
23-25 at midbody, smooth dorsally, keeled laterally; ventrals 246-278
(X = 256.3), 226-248 elsewhere; anal entire; divided subcaudals,
102-121 (X = 111.1), 80-102 elsewhere.
Two color phases occur on Komodo. One is yellow-brown to olive
above, with two black paravertebral stripes on each side of anterior
part of body, often interrupted, sometimes uniform olive-brown,
sometimes another stripe 3 scales below. This phase is found in
specimens of all sizes, from hatchlings to the largest adults. The other
phase is predominately barred (Fig. 41). Ground color is similar but
alternate black X-shaped bars extend from the ventral to near the
vertebral line. The widest, darkest, and most evident parts of each bar
are the two ventral termini and two places along the bar where the
stripes would occur in the striped phase. Between these bars is a dark
gray bar or rounded spot, with the darkest zones again in the stripe
areas. Intermediates between the two phases are also represented in
the material (Fig. 42). Of the 10 specimens available from Komodo, 4


FIGURE 41.-Elaphe subradiata subradiata (UF 28762), Loho Liang, Komodo.


1980








BULLETIN FLORIDA STATE MUSEUM


are barred, 4 are striped, and 2 are intermediate. Juveniles are similar
to adults, but with a wide light brown band across the back of the head
with a thin black band or band of spots at the nape. Head yellow to
dark brown, usually with a dark postocular stripe. Lower surface
yellow to white, with faint dusky edges on many of the ventral shields
and black dots at the edge of ventrals about every six scales.
Supralabials uniform yellowish-white. The largest in the series, which
is also the largest reported to date, is a male from Flores (TL 2060 mm,
SVL 502 mm); the largest Komodo specimen (N=10) has an SVL of
223 mm, but these all seem immature.
DISTRIBUTION AND HABITAT.-Previously reported from Komodo
by Dunn (1927a) and from Rintja by Darevsky (1964a), this species is
also known from the neighboring islands of Sumba (Forcart 1949),
Flores and Sumbawa (Mertens 1930, de Haas 1950), and eastward as
far as Timor and Samao. Almost all the specimens the New York
Zoological Society Expedition collected were taken in gallery forests
extending downward from the higher heavily forested slopes. Mertens
(1930) considered it eurytopic, reporting it from humid montane
forests as well as dry coastal lowlands, including areas of alang-alang
savanna.
REPRODUCTION.-All the Komodo specimens are immature.
FooD.-Large Flores specimens contained bird feathers, remains of
Rattus sp., and Gekko gecko. The smaller Komodo specimens con-
tained only remains of Hemidactylus sp.
PARASITES.-None noted.







AB C





D E F


(B) UF 28759; (C) UF 28755; (D) UF 28757; (E) anterior body patterns of UF 28762; and
(F) middorsal pattern of the same specimen. All individuals are from Komodo and of the
approximate same size.


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AUFFENBERG: KOMODO HERPETOLOGY


Lycodon aulicus capucinus BOIE
Figure 43

Lycodon capucinus Boie 1827:551 (Type locality: Java).
Lycodon aulicus Boulenger 1893:352.
Lycodon aulicus capucinus Boettger 1898:37.
Ophites aulicus Griffin 1909:596.
SPECIMENS EXAMINED.-(18) UF 36201, Bara, Flores, 200 m; UF 28792, 28796-97,
28799, 28800-4, 28806, 36170, Loho Liang, Komodo, 2-30 m; UF 28793-95, 28798,
28805, Loho Lavi, Komodo, 5-10 m; UF 39894, Padar, 3 m.
DESCRIPTION.-Head slightly distinct from neck, eye rather small
with vertical pupil; nostril large, maxillary curved medially, anterior
teeth enlarged; internasals much shorter than prefrontals; frontal
usually shorter than distance from tip of snout, shorter than parietals;
loreal large, long, not entering eye, in contact with internasal; one
preocular, in contact with frontal in 53% of Komodo specimens; 2
postoculars; 2+3 temporals; supralabials 8-9, with 3-4 entering eye;
4-5 infralabials in contact with anterior chin shields; 17 midbody
scales, smooth, with apical pits; ventrals on Komodo specimens
181-214 (without strong sexual dimorphism as reported by Mertens
[1930], because Komodo males vary from 181-214 [X = 198.0], females
196-213 [X = 204.9], 187-210 elsewhere); anal divided, 69-89 divided
subcaudals (X = 76.8), 62-67 elsewhere.
Dorsal ground color gray-brown, with yellow to gray interstitial
skin and adjacent scales forming a network that often delineates a
series of irregular spots down back; head uniformly brown above, often
with a darker edge posteriorly, an hourglass-shaped light gray to tan
blotch on occiput; brown edges on otherwise gray to tan supralabials;
chin dark on edges, especially anteriorly, otherwise white to cream;
venter uniformly white to cream. Juveniles similar, but brighter pat-
terned throughout.
The largest specimen previously reported (Mertens 1930) is 464
mm SVL, but UF 36201 (Bara, Flores) is 470 mm; average Komodo
SVL 380 mm, tail 80 mm.


FIGURE 43.-Lycodon aulicus capucinus (UF 28806), Loho Liang, Komodo.


1980







BULLETIN FLORIDA STATE MUSEUM


DISTRIBUTION AND HABITAT.-The species is widely distributed in
Asia, being known from India and Ceylon east to the Philippines and
Wetar, Lesser Sundas, in the southeast. The present subspecies occurs
in the Lesser Sundas from Wetar and Serua westward to Sumatra.
Dafevsky (1964a) reported it from Komodo, Rintja, and Padar. De
Rooij (1917) and Forcart (1949) listed it from the nearby islands of
Flores, Sumbawa, and Sumba. On Komodo we found it in a variety of
habitats, usually monsoon forests, where it was under stones and the
loose bark of standing trees.
REPRODUCTION.-Mature males vary in SVL from 365-398 mm
(X = 379.5); females are much larger, from 416-470 mm (X = 436.3).
De Rooij (1917) reported that 3-7 eggs are laid in a single clutch.
Though Komodo specimens were collected in both dry and wet
seasons, it is not yet clear when eggs are laid there. Females taken in
January have 9-28 follicles, up to 16 being enlarged, but yolked in only
one specimen. Their size varies from 4.4-8.4 mm in diameter. The
oviducts are smooth, flat, wider, but generally not convoluted or over-
ly thickened. Females were collected only in the wet season. The dry
season may be the breeding season, because mature males taken from
August through November had turgid testes, with thick, convoluted,
turgid vasa deferentia. Eggs are probably laid in November and
December, just before the rainy season. Smith (1943) reported an in-
cubation time of 33 days.
FooD.-Gehyra mutilata has been reported by Mertens (1930) as
eaten by a Flores specimen. Approximately half of the 18 Komodo
specimens contained food, of which the contents of 8 could be iden-
tified. Sphenomorphus florensis and Sphenomorphus sp. were found in
50%, Mus musculus in 25%, Emoia similis in 12.5%, and a reptile egg
(Hemidactylus?) and an adult Hemidactylus frenatus in 12.5%. Smith
(1943) reported that geckos, skinks, and sometimes mice and frogs are
eaten.
PARASITES.-None obtained.
REMARKS.-Dunn (1927b) reported that his Komodo specimen was
taken at night climbing a tree. We, too, found this snake abroad at
night. Smith (1943) remarked that they are good climbers and often
nocturnal.
Psammodynastes pulverulentus (BOIE)
Figure 44
Psammadynastes pulverulenta Boie 1827:547 (Type locality: Java).
Psammadynastes pulverulentus Giinther 1858:140.
SPECIMENS ExAMINED.-(27) UF 28790, Kampung Komodo, Komodo, 5 m; UF
28769-70, 28774 (plus 5 young), 28787 (plus 3 young), Loho Liang, Komodo, 1-30 m; UF
28670, 28772-73, Kali Inaloah, Komodo, 40 m; UF 28771, 28786, 28789, Gunung Ara,


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Komodo, 700 m; UF 28767, Ntodo Puljaran, above Kampung Komodo, Komodo, el 50
m; UF 28768, Gunung.Longgo, Komodo, 150 m; UF 28781-85, 28788, 28791, Loho Lavi,
Komodo, 12 m; UF 36167, Nggoer, Flores, 30 m; UF 36196, Poku, Flores, 500 m.
DESCRIPTION.-Head somewhat triangular from above, pupil ver-
tically elliptic; anterior mandibular teeth greatly enlarged; snout
slightly turned up at tip; canthus distinct; loreal present, single; one
preocular (never 2 in Komodo material, though found elsewhere ac-
cording to Boulenger [1896]); 1-3 postoculars; temporals 2-2;
supralabials 8 (9), of which 3-5 enter eye; 8-9 infralabials of which 3
contact the anterior chin shields; 17 smooth scale rows at midbody,
without pits; ventrals 150-176 (X = 165.3), 144-175 elsewhere (fide
Mertens 1930, Boulenger 1896); anal entire; divided subcaudals 48-72
(X = 57.8), 44-70 elsewhere; greatest SVL 497 mm, tail 128 (de Rooij
1917), Komodo X = 357.6.
Brown, black, reddish-brown, or gray-brown dorsal ground color,
females generally darker than the males, particularly on the chin;
small, ill-defined darker blotches and light spots on the dorsum; head
uniform above or with a faint V-shaped darker mark finely edged in








A B







C








FIGURE 44.-Psammodynastes pulverulentus. Head, UF 28785 Loho Lavi, Komodo;
mid-ventral body patterns, above, UF 28785, a male; below, UF 28790, Kampung
Komodo, Komodo, a female.


1980







BULLETIN FLORIDA STATE MUSEUM


dark brown or black, with supralabials yellow to cream with darker
edges; ventrals with dark brown to black speckles or faint small dashes
on a light ground color in males, females darker ventrally, sometimes
with two lighter stripes (Fig. 44), chin darker, often with lighter me-
dian elongate spot or stripe. Juveniles similar, but more vividly pat-
terned.
DISTRIBUTION AND HABITAT.-Widely distributed from the eastern
Himalaya Mountains southeastward through Indochina, Philippines,
and Indonesia to Flores and Celebes. Previous records from the
Komodo area are Flores, Sumba, and Sumbawa (Mertens 1930, For-
cart 1949) and Komodo (Dunn 1927a). Barbour (1912) and Mertens
(1930) reported that it is generally found in the mountains, usually be-
tween 400 and 1200 m elevation, though the latter stated that he found
one on Flores below 200 m. Forcart (1949) reported specimens from the
coast to as high as 350 m on Sumba. That they are found at such low
elevations agrees with our observations on Komodo, for many of the
specimens were taken near sea level in gallery forests along dry stream
beds. Some were found in savanna environments, usually on rock-
strewn grassy hillsides. A few were found under loose bark on fallen
tree trunks and large branches, but most were under flat stones during
the day. One was found in a rice field on Flores.
REPRODUCTION.-Mature females were collected from August
through February. Follicles varied from 7 to 32 (X = 18.1). Enlarged
follicles 3-8 (X = 5.1); oviducts are thick walled and convoluted during
this entire period. A November female (SVL 326 mm) contained 5 well-
developed, scaled embryos (79-83 mm SVL) that would have un-
doubtedly been born in the earliest part of the wet season (December);
another set of 3 embryos (94-88 SVL) very close to being born was
found in a January female (SVL 497 mm). Females from the mainland
have more young (5-10, fide Smith 1943). Mature females have an SVL
of 320-410 mm (X = 376.3), males 288-370 (X = 336.9). Darevsky
(1964a) stated that a Padar specimen contained a near term egg, but de
Rooij (1917) and Smith (1943) stated they are ovoviviparous, fitting
with my observations on Komodo.
FOOD.-Mertens (1930) reported Rana microdisca and Sphenomor-
phus florensis as being eaten. In 20 Komodo specimens dissected the
following species were found: Sphenomorphus florensis in 10%, Cyr-
todactylus darmandvillei in 5%, Hemidactylus sp. 5%, and
Sphenomorphus schegeli 5%. One of the specimens contained the re-
mains of an Oreophryne jeffersoniana.
PARASITES.-Of the adults examined 25% were heavily parasitized
with the nematodes Hexametra quadricornis and Spinicauda komo-
doensis (J. Pinnell, pers. comm.).
REMARKS. Called ular percha locally.


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AUFFENBERG: KOMODO HERPETOLOGY


FAMILY ELAPIDAE
Naja naja sputatrix BOIE
Figure 45

Naja sputatrix Boie 1827:557 (Type locality: Java).
Naia tripudians Lidth de Jeude, in Weber 1890:179.
Naia tripudians var. sputatrix Boulenger 1896:384.
Naja naja sputatrix Stejneger 1907:395.
Naia naia Barbour 1912:30.
Naja sputatrix malayae Deraniyagala 1960:223 (Type locality: Negri Sembilan,
Malaysia).
SPECIMENS EXAMINED.-(16) UF 32572, Nggoer Flores, 15 m; UF 39833, 28674,
28679, Kali Besar, Loho Liang, Komodo, 30 m; UF 28673, 28677, 28680, 28678, Loho
Liang, Komodo, 1-30 m; UF 28675-76, Kampung Komodo, Komodo, 8 m; UF 28672,
Sabita, Komodo, 8 m; AMNH 31957-59, Komodo, "near sea level," AMNH 31974-75,
Komodo, 650 m.
DESCRIPTION.-Rostral broader than high, very evident from
above; internasals somewhat shorter than prefrontals, in contact with
a single preocular; frontal about as long as its distance from rostral, as
broad to slightly broader than supraocular; postoculars 3 (4 on one side
of one specimen); temporals consistently 2+3; supralabials 7, of which
numbers 3 and 4 enter eye; infralabials 9, first 4 in contact with first


FIGURE 45.-Naja naja sputatrix (UF 28677), Loho Liang, Komodo, head and neck.







BULLETIN FLORIDA STATE MUSEUM


pair of chin shields, which is longer than posterior pair; scale rows:
neck (through marking) 21 (23 in two specimens, 19 in the Flores
specimen); midbody consistently 19 (17 in Flores specimen); ventrals
and subcaudals remarkably consistent: males 160-173/47-55, females
175-182/51-56. Sex ratio in preserved material 1 male: 3 females.
Color light to dark brown dorsally, each scale with a yellow to
whitish anterior edge; head above uniform, usually same color as body,
sometimes darker; neck only slightly expandable, markings from
nothing to a barely discernible small light mark on either side of the
vertebral line (most common) to a faint V-shaped mark, the apex
directed posteriorly (Fig. 45); ventrally yellowish-white, sometimes a
few faint scattered spots; faint brownish-gray throat band in most
specimens, which is often interrupted. On the basis of Dunn's (1927b)
description of Komodo material, Mertens (1930) presumed the
Komodo material to be a slightly different color than specimens from
more western islands, a conclusion not justified by the present
material. Juveniles uniform brown, darker anteriorly, no lighter marks
on scales; head dark brown above, snout and sides of head yellowish-
brown; ventrally uniform yellowish-white.
In his study of the Sunda herpetofauna, Mertens (1931) stated that
if it were not for the very wide zone of intermediates, the Sumatra and
Javanese specimens (N. n. sputatrix), the Lesser Sunda material could
easily be considered a separate race.
DISTRIBUTION AND HABITAT.-Previously reported from Komodo
(Dunn 1927b) and the nearby islands of Sumbawa and Flores (Klem-
mer 1963). Dunn (1927b) reported specimens taken on Komodo from
sea level to 650 m. We found none higher than this. We found Naja na-
ja sputatrix abroad both day and night. Though Mertens (1931) found
them only on the ground, some of ours were shot from heights up to 11
m in trees overhanging dry creek beds. Specimens were also seen in
Imperata and Zizyphus savannas, as well as in gallery and deciduous
monsoon forests, particularly around rock piles and outcrops with
many fissures and holes.
REPRODUCTION.-Mature females have an SVL of 84.8 cm, males
88.3 cm. At sea level hatchlings (SVL 235-240 mm) were found in
January and February. The ovarian scars in one female suggest that as
many as 16 eggs may be laid in a clutch; yolked follicles 2.2 to 5.2 mm
(range = 1-16) were found in January-February specimens. Eggs are
probably laid at the end of the dry season (November). Males have
turgid testes and turgid, convoluted vasa deferentia in October and
flat testes and non-convoluted vasa deferentia in January-February.
FOOD.- Lycodon aulicus and Elaphe subradiata were reported as
food by Dunn (1927b), and Mertens (1931) reported mice, Rana can-


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AUFFENBERG: KOMODO HERPETOLOGY


crivora, and Mabuya multifasciata. None of my Komodo specimens
contained any identifiable food.
PARASITES.-The nematode Ophidascaris wui is common in the
digestive tract of Komodo specimens.
REMARKS.-Komodo villagers show relatively little fear of this
snake. It apparently rarely bites or spits, and then only with con-
siderable provocation or when restrained. Only one death attributed to
this species was reported to me-an old, feeble woman who had
stepped on one in the savanna and was bitten on the heel. Komodans
call this ular sendok, the "spoon snake," because of its shape when in a
reared position. Shed skins were noted from August through October.

FAMILY VIPERIDAE
Vipera russelli limits MERTENS
Figures 46, 47
Vipera russelli limits Mertens 1927b:183 (Type locality: Pulau Ende, off SE coast of
Flores).
SPECIMENS ExAMINED.-(59) UF 28616, 28619, 28621-23, 28625-26, 28629, 28631-32,
28634-35, 28637, 28639-39, 28661-64, 28666-71, Loho Liang, Komodo, 1-30 m; UF
28617-18, 28624, 28627, 28636, 28665, Kampung Komodo, Komodo, 20 m; UF 28620,
28630, 28633, 28638, 28641-42, 28660, Loho Lavi, Komodo, 12 m; UF 28628, Gunung
Puaki, above Loho Lavi, Komodo, 120 m; UF 28665, Kampung Kechil, Komodo.
DESCRIPTION.-Head small and long, moderately distinct from
neck, but habitus generally more slender in appearance than nominal
form (Fig. 46); canthus distinct; rostral broader than high, in contact
with 6 scales; head scales small and strongly keeled; supraoculars very
small, with 7 scales between them; 2 scale rows between eye and
supralabials; nostril in large nasal, in contact with nasorostral, but not
the supranasal; a very large postnasal found in 3 specimens; first
supralabial always touching lowest postnasal (usually not the case in
the nominate form); supralabials 9-11, usually 10; infralabials 11-14,
usually 12, numbers 4 and 5 in contact with anterior chin shields; dor-
sal scales keeled, except outer row, in 27 rows at midbody; ventrals
139-167 (X = 155.8) (no statistically significant sexual difference);
subcaudals divided (rarely 1-2 single members in series), 46-65
(X = 50.3) in males, 40-56 (X = 44.5) in females, the difference in
means being significant (P = > 0.05).
Ground color brownish-gray to dark gray; dorsally with a
longitudinal row of blackish-gray or dark brown blotches with white
and black edges, often round to oval on anterior third of body, and
often variously connected vertebrally to form zig-zags and other
anomalous patterns on posterior two-thirds (Fig. 47); those on tail
often fused to produce solid (in 20%) or broken (56%) stripes. Lateral


1980








BULLETIN FLORIDA STATE MUSEUM


body surface with similar smaller blotches having white and black
borders, often forming a longitudinal solid (in 45%) or broken (53%)
stripe on side of tail (newborn and 1st-year individuals with a sulfur
yellow tail line); above this series and below vertebral row a single
(10%) or double (90%) row of less evident spots; also a series of small
spots on edges of ventral scales (Fig. 47). Thus Komodo specimens
have from 5 to 9 (usually 7) rows of body spots. Head markings re-
duced when compared to nominate form. A pair of small ground-
colored oval to subtriangular spots on dorsoposterior part of head,
separated by apex of a light-colored V (which is characteristic of
nominate form) in 58% of Komodo material, tending to disappear at
end of first year of life; dorsoposterior spots in all Komodo specimens
separated from dark temporal zone by an area of ground color in all but
2 specimens; interorbital dark spot or stripe always present, heavily
pigmented and obvious in 55%, faint in 33%, and nearly absent in
12%; dark subocular spot present in all but 2 specimens; supralabials
white with black to brown flecks. Chin white, vividly marked with
numerous black to very dark brown triangles; venter white to cream
with fine gray punctations and somewhat hemispheric-shaped black,
dark brown to dark grayish-brown markings linearly arranged (in
55%), or with fewer dark markings nearly lacking gray punctations (in
40%) or without any punctations (5%).


FIGURE 46.-Vipera russelli limits (UF 28617), Loho Liang, Komodo.


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


DISTRIBUTION AND HABITAT.-This geographic race is restricted to
the Lesser Sundas and known only from Flores and Komodo. Dunn
(1927b) reported it previously from Komodo. All the Komodo
specimens were collected near the coast (1-120 m) in Zizyphus savanna
and the beachside Pandanus community. Most were found in or near
the wooded borders of more open situations, often near stone piles in
which they commonly took refuge. Many were found on the rocky
banks of dry stream beds.
REPRODUCTION.-Newborn with umbilical scars (170-275 mm SVL,
X = 189.4) were obtained on Komodo only during January. Mature
females were collected from the last third of the dry season in October
into the middle of the wet season in January. Follicles vary from 20 to
42 (X = 31.2); from October 5 to 15 they show beginnings of yolk
deposition (15-27 mm in diameter); the oviducts are smooth, thin
walled. In December the embryos (up to 13) are well formed. Males are
available from July through February. From July through September
the testes and vasa deferentia are turgid, the latter highly convoluted.
From late November through February the testes are more flaccid and
the vasa deferentia less convoluted. Thus it appears that breeding
takes place in the middle to late dry season, and the young are born in















0P.,-
:4 4:i
ii
















FIGURE 47.-Variation in mid-dorsal patterns of juvenile Vipera russelli limits from
Komodo. From left to right, UF 28669, 28639, 28644, and 28665.








BULLETIN FLORIDA STATE MUSEUM


the early part of the wet season. Mature females are 550-700 SVL
(X = 610), males 530-630 (X = 570).
FooD.-Of the 56 specimens examined, 51% contained food or food
remains, of which only those in 24% could be identified. Of these,
newborn and juvenile specimens contained elytra and other parts of
adult coleopterous insects (21.8%), of their larvae (8.3%), and of moths
(1.8%). Hemidactylus frenatus (7.2%) were found in juveniles and
adults, Cosymbotes platurus, Mus musculus (both 2.2%), skinks
(genus ?) (5.5%) in adults, Emoia similis in newborns (1%), and Kaloula
baleata (1%) in adults. Vegetable matter occurred in 7.4% and sand
and stones in 3.6%.
PARASITES.-Nematodes of the species Kalicephalis willeyi were
found in 4% of the 56 specimens examined.
REMARKs.-Indonesian populations of Vipera russelli have been ex-
tensively discussed by Dunn (1927a), Kopstein (1936), Mertens (1927c
1930, 1957), Hoesel (1953, 1958), and Brongersma (1958), all based on
relatively few specimens. The new, extensive material from the Lesser
Sunday Islands supports the validity of the trinomial, though it is not a
markedly distinct race. This is based mainly on the presence of a
subocular spot in all but 2 specimens. The light-colored V on the head
is only found in about half the specimens now available, in spite of the
importance placed on the character by Mertens (1930). The 7 rows of
dorsal spots in V. r. limits also serve to distinguish it from V. r.
siamensis (with 5) and V. r. russelli (with 3). Furthermore, the max-
imum size reported in the Lesser Sundas is 835 mm, whereas mainland
snakes become twice that length (Mertens 1930).
In general most specimens were easily angered and inflated their
bodies, hissed, and struck when annoyed, but on several occasions ex-
pedition members accidentally stepped over their well camouflaged
bodies, and the snakes made no attempt to bite. A small pet 5 kg wild
boar was struck in the forehead by a specimen 400 mm SVL, and after
much swelling and obvious discomfort recovered within a week.
Komodo villagers greatly fear this snake (which they call misa), for it
has caused more snakebite deaths on this island than any other
species. Most were seen during the late afternoon to twilight.
Trimeresurus albolabris GRAY
Figure 48
Trimeresurus albolabris Gray 1842:48 (Type locality: "China").
Lachesis gramineus Boulenger 1896:554 (part).
Lachesis fasciatus Boulenger 1896:63 (Type locality: Djampea Island, near Celebes).
Lachesis gramineus albolbris Mell 1922:126.
Trimeresurus fasciatus Dunn 1927b:5.
Trimeresurus gramineus gramineus Mertens 1930:27.


Vol. 25, No. 2








AUFFENBERG: KOMODO HERPETOLOGY


SPECIMENS EXAMINED.-(95) UF 30127-50, Nggoer, Flores, 10-20 m; UF 30151, 4.5
km ENE Look, Flores, 200 m; UF 28719-23, 28725-26, 28728-33, 28735, 28737, 28716,
28714, 28708, 28705-5, 28702, 28699, 28691, Loho Liang, Komodo, 2-30 m; UF 28734,
Nggolo Vai, Loho Liang, Komodo, 10 m; UF 28695-96, 28736, Kali Besar, Loho Liang,
Komodo, 60 m; UF 28687-88, 28706-7, 28700, 28692, 28690, 28709, 28713, 28717, Loho
Lavi, Komodo, 1-15 m; UF 28689, 28697-98, 28715, Gunung Ara, Komodo, 300 m; UF
28703, near Sabita, Komodo, 8 m; UF 28701, Kali Inaloah, near Gunung Amasobang,
Komodo, 24 m; UF 28693-94, Galong Sambe, near Poreng, Komodo, 150 m; UF
28710-12 (plus 17 young), 28718, 28724, "Komodo"; UF 36446, near Deussi, Timor,
150 m.
DESCRIPTION.-Distinct canthus, head scales small, smooth,
supraocular narrow, sometimes divided; 1-2 scales between internasals
or latter in contact; 8-13 scales between supraoculars; 2-3 postoculars;
1 subocular, sometimes in contact with supralabial 3, usually
separated from labials by 1-3 scale rows; 8-12 supralabials, second
bordering loreal pit anteriorly, third largest; temporal scales with
short keel (a few individuals with weak or no keels); dorsal scales
strongly keeled above, smooth on two lateral rows, in 19-21 rows
(usually 21); ventrals 153-168 (X = 161.2); subcaudals divided, 53-81
(X = 63.8), males 60-81, females 51-65; tail prehensile; greatest
reported SVL to 805 mm, tail 125 (Mertens 1930), largest SVL of
Komodo specimens 760 mm.
Usually bright green dorsally, often olive or even blue (in approx-
imately 1 out of 8 Komodo and western Flores specimens), often with
distinct darker transverse bands; supralabials yellow to greenish-
white (sometimes light blue) in all but largest specimens, which are
sometimes suffused with gray or olive; tail usually with rusty-colored
streak dorsally; venter greenish-yellow, greenish-white, or sometimes
light blue; iris brown to red (latter apparently more common in blue
phase individuals).
DISTRIBUTION AND HABITAT.-FrOm Sumatra and Borneo eastward
through the Sunda Islands to Timor, except for the Celebes-
Halmahera Island groups (Pope and Pope 1933). Forcart (1949), de
Haas (1950), and Mertens (1930) reported the species from the nearby


FIGURE 48.-Trimeresurus albolabris (UF 28691), Loho Liang, Komodo.


1980








BULLETIN FLORIDA STATE MUSEUM


islands of Sumbawa, Sumba, and Flores; Darevsky (1964a) reported it
from Rintja and Padar.
Certainly one of the most commonly encountered snakes on Flores
and Komodo, this species is found in a variety of habitats, though it is
most common in monsoon forest. It occurs from sea level to 700 m on
Komodo and reaches maximum abundance along stream courses,
where it is often seen abroad at night, prowling in the bordering shrubs
and small trees near the bank. Specimens were found as high as 15 m
in tamarind trees in the daytime, when they were usually asleep, coiled
in the common crotch of several branches. A few were found on the
edges of savannas, usually crawling about in the dense stands of the
tufted grasses of the genus Andropogon.
REPRODUCTION.-On Komodo up to 17 young are born alive in the
early monsoon season. Supporting data are: 3 July-August females
had small, translucent unyolked follicles, the oviducts smooth, flat and
translucent. A September female contained 11 large yolked follicles
6.2-13.2 mm in diameter, and the oviducts were large, thick-walled,
and highly convoluted. The testes of males caught at the same time
were enlarged and turgid and the vasa deferentia convoluted but flat-
tened. By October and November, the latter are nearly smooth and not
convoluted. This condition continues at least through February. Up to
17 near-term embryos (all with scalation and color pattern discern-
ble) are found in females collected in November. Newly born specimens
are found in January, and 12 females caught in December-January had
no embryos, but small follicles and flat, smooth oviducts. These data
suggest that breeding probably takes place in September, and the
young are born in late November and December. Juveniles caught in
January all show an umbilical scar and are 140-195 mm SVL
(X = 175). By the following January, they are 290-310 mm SVL. Sex-
ual maturity is probably attained in the third year.
FooD.-Of 56 specimens examined, 16 contained remains of food
that could be identified to the following species in 9: Hemidactylus
frenatus in 4 specimens, Lepidodactylus lugubris in 2, Sphenomorphus
florensis in 1, Mus musculus in 1 (also reported by Dunn 1927b), and
Kaloula baleata in 1 (also reported by Dunn 1927b). In addition, 7
others contained remains of unidentified lizards. Mertens (1930)
reported Rattus rattus, miscellaneous frogs (usually Rana cancrivora),
and lizards (usually Mabuya multifasciata) from specimens collected
on other islands.
PARASITES.-Pentastomideans (Raillietiella sp.), ascaridoidians,
oxyuroideans (Polydelphis sp.), and the nematode Hexametra
quadricornis are common in the intestines (approximately 28% are
parasitized).


Vol. 25, No. 2




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