I

Asiatic Herpetological Research

June 1995

Vol. 6, pp. 30-35

The Systematic Relationships of Dravidogecko anamallensis(Gunther 1875)

AARON M. BAUER AND ANTHONY P. RUSSELL 21

1

2

Biology Department, Villanova University, Villanova, Pennsylvania 19085, USA

Vertebrate Morphology Research Group, Department of Biological Sciences, University of Calgary, 2500
University Drive N.W., Calgary, Alberta, Canada T2N 1N4

Abstract. -The relationships of the monotypic gekkonine genus Dravidogecko are assessed by comparativeevaluation of its external and internal morphology. A suite of shared-derived features is possessed by

Hemidactylus and a variety of satellite genera, including Dravidogecko. These similarities are advocated as
being so compelling, and the ostensible defining features of Dravidogecko to be so weak that the latter issubsumed as a junior synonym of Hemidactylus. The biogeographic consequences of this taxonomic shift areconsidered.

Key words: Dravidogecko, Hemidactylus, Teratolepis, digits, scansors, phalanges, paraphalangeal elements,
muscles, biogeography, India.

Russell (1976: 238;

ceratobranchial arch.

Introduction

Fig. 14) suggested that Dravidogecko had adigitalisa monotypic genus ofDravidogecko
gekkonid lizards endemic to south India.
The single species, D. anamallensis, wasoriginally described as a member of thegenus Hoplodactylus (Gunther, 1875;
Strauch, 1887), but following the work of
Smith (1933), it was assigned to a new
genus, primarily on the basis of differencesin the distal scansors and in preanal porearrangement. Subsequently it has been

demonstrated that Dravidogecko is agekkonine gecko, whereas Hoplodactylus
isastrictosensu
diplodactyline
1954;(Underwood,
Kluge, 1967). The

have

of

Dravidogecko
relationships
remained obscure, and the systematic statusof the species has never been investigated
It is known from only a few
the Anaimalais, Palnis andfromspecimens

adequately.

was most closely

structure thatapproached by that of Hemidactylus and its
close allies.
digitsis

While external form of theparticularly sensitive to functional
to exhibiting

demands and thus prone

convergence and parallelism (Russell,1979), details of the internal anatomy aremuch more

helpful at indicating true
and,therefore, affinity (Russell,
homology
Russelland Bauer, 1990).
1976, 1979;
herein present the results of a comparative
survey of both the external and internalWe

anatomyofHemidactylus (andand digits inclose relatives) and

feet

the

its

use these to demonstrate both the wide rangeof variation present and the shared derived
features that circumscribe this cluster andhelp clarify the relationships of the enigmatic

Dravidogecko.

We further relegate thegeneric name Dravidogecko

into thethroughout forested areas of southernsynonymy of Hemidactylus as there are no
derived features of Dravidogecko that are
not also shared by at least somepeninsular India (Daniel, 1983).

Hemidactylus.

Tirunelveli Hills (Satyamurti, 1962; Murthy,
1985) but is reportedly widely distributedItisprobable thatH.

anamallensis is a primitive hemidactyl.
Russell (1972) considered Dravidogecko
to belong, on morpho-functional grounds,
in the Hemidactylus group, along with

Materials and Methods

Hemidactylus, Briba, Teratolepis andSpecimens of Dravidogecko were
examined or borrowed from the collectionsof The Natural History Museum, London(BMNH) and the Institute Royal desCosymbotus. Kluge (1983) placedalongwith the other gekkonine genera previously
mentioned, in the tribe Gekkonini on thebasis of the absence of the secondit,© 1995 by Asiatic Herpetological Research

Sciences Naturelles de Belgique, Brussels

Vol.6, p. 31

Asiatic Herpetological Research

June 1995

Comparative material of othergekkonines, especially Hemidactylus, were
and theborrowed from the
California Academy of Sciences, San
Observations on toe
Francisco (CAS).
structure were made using a Nikon SMZ-10
microscope. The specimens examined arelisted below. All numbers refer to BMNH

(IRSNB).

BMNH

specimens unless otherwise identified.

muriceus 1926.9.24.13; 1966.283; H.
modestus 1946.8.25.37; H. ophiolepisH.
1937.12.5.324-325;
oxyrhinus
99.12.5.170-175; 1967.491-494; H.

persicus 1970.250; 1972.716; H. prashadi
1946.8.14.66-69; H. pumilio 1946.8.20.1;
H.
reticulatus,
1946.8.25.58-61;
H.
richardsoni
1916.5.29.1;
1901.3.8.1-3;
1919.8.16.49; H. ruspolii 1937.12.5.228229; 1937.12.5.239-246; H. sinaitus

1937.2.5.293;

Dravidogecko anamallensis 82.5.22.79-84;

97.10.28.83-86;

IRSNB 1194.

1953.1.6.97-98; H. smithi 1931.7.20.8589; 1972.745; H. somalicus 1946.8.25.7778; H.
squamulatus 9 8.1.8.2-3;
1902.5.26.2; 1923.10.9.2; 1923.10.9.1415; H. subtriedrus 74.11.11.1; H. taylori
1946.8.23.48; H. triedrus xxi.l9a-b; H.

Briba brasiliana 1971.1045.

craspedotus 1926.12.7.7,
1930.10.9.2
C.
platyurus xxi.36a, 97.6.21.4,
97.12.28.10, CAS 18565, CAS 18567
Hemidactylus albopunctatus 1946.8.22.75;
H. ansorgii 1901.1.28.22; 1966.337; H.
barodanus 1905.11.7.1-6; 1937.12.5.215216; 1958.1.6.29; 1970.1437-38; H.
bouvieri 66.4.12.3; 75.4.26.10;
bowringii 1929.12.1.7-10; 1940.4.26.2-3;
1956.1.11.15-16; H. brookii 1918.1 1.12.2-

CosymbotusH.

1931.12.10.6-7;
1930.10.6.6;
1970.2196-98; 1971.242; H. citernii
and1931.7.20.114-119
128-130;
1937.12.5.202-204; H. curlei
1946.8.25.41; H. depressus 52.2.19.21;
1948.1.7.35; H. echinus
61.2.21.5;
89.7.6.1; 1903.7.28.1-2; H. fasciatus

tropidolepis

1937.12.5.322-323;

H.

turcicus 1934.11.8.10- 14; 1971.1143-45;
H. yerburii 99.12.13.43-44; 1903.6.26.3-4;

1945.12.18.12.

Teratolepis fasciata

69.8.28.32;

1933.7.8.37; 1963.1019; 1964.930-931; T.
albofasciatus 1963.613-621

10;

1919.8.16.48; 1956.1.11.37-40; 1971.253;

H. flaviviridis

1931.7.20.153-155;

1971.1378-1382; H.forbesii 1946.8.25.43-

Al;H.frenatus 1938.10.2.1; 1952.1.4.301970.1879-1895; H. garnotii
31;
95.11.7.1; 1903.2.21.1-2; 1940.6.3.24-29;

H. giganteus 1908.12.28.27; 1969.82874.4.29.1388;
829; H. gracilis
80.11.10.47; H. granti 1957.1.9.52-66; H.
93.12.7.1; 98.3.30.21-22; H.
greeffii
homeolepis 99.12.5.38; 1953.1.7.84-85;
1967.485-489; H. isolepis 1952.1.7.79-80;
H. jubensis 1946.8.23.66; H. karenorum
68.4.3.88-89; 91.11.26.13-14; H.laevis
1946.8.25.42; H. leschenaulti 70.5.18.7071; 74.4.29.233-236 (six specimens); H.
longicephalus 1936.8.1.287-305; H.
mabouia 1923.11.9.46-50; 1964.1429-35;

Results

A considerable range of variation indigital form

existsand subdigital scansor design
among members of the genus

This variation is1).
evident in such aspects as the number ofdivided scansors (lamellae), the extent of
their division, the extent of the undivided
lamellar series at the base of the digits, andthe length, form and degree of separation of
the free, distal, claw-bearing segment of thedigits. Figure 2 illustrates the general formof the ventral aspect of the right pes of

Hemidactylus (Fig.

Dravidogecko and provides comparison
with the ventral aspects of the fourth pedal
digit of Hemidactylus reticulatus andTeratolepis fasciata. While some species of
Hemidactylus, such as H. garnotii and H.
smithii (Fig. 1), have digits with a large
number of completely divided scansors, andan elongate, free distal, claw-bearing
portion, this is not so for other species, such
as Teratolepis albofasciatus (see Grandison

H.

macropholis
1931.7.20.109; 1937.12.5.250-258; H.

and Soman, 1963), Hemidactylus somalicus
and H. bouvieri (Fig. 1). In the latter threemaculatus 69.8.25.15; 1956.1.11.44; H.
megalops 1946.8.25.67; H. mercatorius

cases the number of scansors is small, only
the distal most ones are notched, and the1930.7.1.84-90; 1938.8.3.11-15; H.

distal, free, claw-bearing portion of the digit

1970.2209-15;

Vol. 6, p. 32

FIG.

Asiatic Herpetological Research

June 1995

The array of digital form in the genera
Hemidactylus and Teralolepis. All illustrations are1.

of the fourth digit of the pes; a-e, j are of the rightscale barpes, f-h, k-p are of the left pes. The 2
refers to all specimens except n, to which the 5mmmm

scale bar applies. All catalogue numbers refer to the

Natural History

Museum, London (BMNH).

a.

Teralolepis albofasciatus 1963.617; b. Hemidactylus

bowringii 1929.12.1.6; c. //. garnotii 95.11.7.1; d.
H. barodanus 1970.1438; e. H. turcicus 1971.1144;
f. H. somalicus
1946.8.25.77; g. H. ophiolepis

1937.12.5.324;h.

H. mabouia 1964.1431; j. H.

forbesii 1946.8.25.47; k. H. smithii 1931.7.20.85;
1.
H. fascial us 1919.8.16.48; m. H. ansorgii

FIG. 2.

a.

Ventral aspect of the right pes ofDravidogecko anamallensis, BMNH 82.5.22.79. b.
Ventral aspect of the fourth digit, right pes of

Hemidactylus reticulatus, BMNH 1901.3.8.1.
Ventral aspect of the fourth digit,
Teratolepis fasciata,
in millimeters.

left

c.

pes of

BMNH 1933.7.8.37. Scale bar

1901.1.28.22; n. //. richardsonii 1916.5.29.1; p. H.
bouvieri 66.4.12.3.

comparing

Dravidogecko

with

Cyrtodactylus brevipalmatus, Hemidactylus

This situation is also
seen in Hemidactylus reticulatus andTeratolepis fasciata (Fig. 2, b, c). The
almost continuous range of variation in

reticulatus and H. barodanus. While thiswas simply a depiction of change of form
assembled as a morphotypic series, it wasalso implied that there may be deeper

external digital characters, especially among

underlying anatomical clues that are
indicative of the closeness of relationship ofisrelatively short.

the west Asian and Somali species of the
Hemidactylus group of geckos has long

been recognized, and has resulted in theestablishment of several different, largely

Dravidogecko to Hemidactylus.

Thesuperficial comparisons of the digits(Figs.
1, 2; see above) provide some idea of the1942 for a discussion).

potential range of variation, but should betreated with caution when being implicateddivisionin arguments about relationship because of

there are many species that express this trait

only marginally.

the extreme plasticity of external digitalform. [Such aspects are well exemplified by
the taxonomic history of the taxon that is theRussell (1976: Fig. 14) indicated thispotential continuity in scansor form, fromundivided to completely divided, by

Detailed
subject of this contribution.]
examination of the internal anatomy of thedigits provides more convincing evidence
about the affinities of Dravidogecko.

arbitrary, generic arrangements (see Parker,

Thus, whileof the scansors is generally
characteristic of the genus Hemidactylus,June 1995

Vol. 6, p. 33

Asiatic Herpetological Research

Russell (1976) presented a mechanistic
diagram of the main features of digital
design in Hemidactylus. The chief aspectsof note here are the unusual form andrelationships of the antepenultimate phalanxof digits III-V of the pes (Russell, 1977),
the distal extent of the dorsal interossei
muscles along the digit, and the means oftendinous insertion of these muscles onto

The pattern of digital
the scansors.
characteristics of Hemidactylus is essentiallyrepeated in Dravidogecko and is restricted toonly a few other genera (Briba, Cosymbotus
and Teratolepis). This suite of sharedderived digital features of these taxa (theshort, erect nature of the antepenultimate
phalanx of pedal digits III-V, the distal
extension of the dorsal interossei muscles as
far as the distal end of the antepenultimate
phalanx, and the tendinous insertion of thedorsal interossei muscles onto the distal
margin of each scansor in turn) unites themas a distinctive evolutionary unit.
Apartfrom Hemidactylus, all of the other generain this cluster are either monotypic (Briba
and Dravidogecko) or include only two
species (Cosymbotus and Teratolepis).

Dissection of the digits of Dravidogecko
reveals that the dorsal interossei muscles are
well-developed and robust and extend as

fleshy bellies as far distally as the digital
inflection (the point of emplacement of the
reduced, erect antepenultimate phalanx

onmanual digits III and IV and pedal digits
III-V). The dorsal interossei muscles send
individual tendons to the distal borders of
the scansors as they do in Hemidactylus (see1976) and Cosymbotus. Thissituation also pertains in Teratolepis andBriba (Russell, 1972). Dravidogecko also
shares with Hemidactylus, Briba andCosymbotus the particular morphology andplacement of paraphalangeal elements(Russell and Bauer, 1988).
Russell,

The above comparisons indicate that
Dravidogecko shares with other members ofthe Hemidactylus radiation (Hemidactylus,
Briba, Cosymbotus, Teratolepis) all of thederived digital features that distinguish thesetaxa from all other geckos.
However,
apomorphic features characteristic of many
Hemidactylus species, such as those

associated with the complete division of thescansors, are lacking in Dravidogecko. It istherefore likely that D. anamallensis is arelatively plesiomorphic member of thisradiation. As such, it is probable that therecognition ofDravidogecko renders

as presently construed
In order to maintain
paraphyletic.
monophyletic generic units we hereby place
Dravidogecko into the synonymy ofHemidactylus Gray, 1825. The correct

Hemidactylus

designation for the single known species
formerly referred to this genus thus becomes
Hemidactylus anamallensis (Giinther 1875),
new combination.

Discussion

Many lizard families include monotypic
Although in some cases thesegenera.
represent independently evolving lineages,
in most they are relatively primitive or
highly derived members of other lineages,
and their recognition renders the lattergroups paraphyletic. Hemidactylus is themost specious genus in the Gekkonidae,
with 75 species currently recognized(Kluge, 1991). Relationships within thegenus are very poorly understood (Parker,
1942;

Loveridge,

1947;

Kluge,

1969;

Bastinck, 1981) and a general uniformity
among most forms (Russell, 1976) has
rendered casual attempts at investigating its
phylogeny unsuccessful. The placement ofDravidogecko anamallensis into this morass,of course, does nothing to aid thisconfusion. It does, however, ensure that
Hemidactylus anamallensis is taken into
account if and when a generic revision of all

Hemidactylus is accomplished.

only in the interest of
maintaining monophyletic groups that therevaluation of monotypic genera isundertaken. Current nomenclatural usage
It

is

not

has implications for non-systematists. Asan endemic Indian subcontinent form,
Dravidogecko might be used to support
arguments about the uniqueness andantiquity of the Indian biota. The use bybiogeographers of classification schemesthat do not adequately reflect phylogenetic
patterns has been shown to lead to theerection of demonstrably false hypotheses

Vol. 6, p. 34(Bauer,Asiatic Herpetological Research1989).

Clearly, biogeographic
the
of
the
organisms
phylogenetic relationships
Some other Hemidactylus
considered.
interpretations must be based upongroup geckos sharing with H. anamallensis
at least partially undivided scansors are also
Indian forms (e.g., Teratolepis albofasciatusfrom the Ratnagiri District, Maharashtra,
Hemidactylus gracilis from the Madhya
Pradesh, Maharashtra and Andhra Pradesh

June 1995

E. Prickley and S. F. Milone provided
general assistance in the laboratory.

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(Smith,reticulatus

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