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(2000) New Data On The Distribution, Status, and Biology of The New Caledonian Giant Geckos (Squamata, Diplodactylidae, Rhacodactylus SPP.)

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(2000) New Data On The Distribution, Status, and Biology of The New Caledonian Giant Geckos (Squamata, Diplodactylidae, Rhacodactylus SPP.)

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Copyright © 2000 Amphibian and Reptile Conservation. All rights reserved.

ARC authorizes
photocopying for internal or personal use provided the appropriate fee is paid directly to the Amphibian and Reptile Conservation 2(1):24-29.
Copyright Clearance Center. Inc., Ill Rosewood Dr.. Danvers. MA 01923-4599. USA. Tel:
(978) 750-8400; fax: (978) 750-4470: email: [email protected]: website: www.copyright.com

New data on the distribution, status, and biology of the


New Caledonian geckos (Squamata: giant
Diplodactylidae: Rhacodactylus spp.)
AARON M. BAUER AND ROSS A. SADLIER 2
1

department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085-1669, USA 'Department
'

of
Herpetology, Australian Museum, 6-8 College Street, Sydney, New South Wales 2000, AUSTRALIA

Abstract. — Recent collections and observations of the New Caledonian giant geckos (Rhacodactylus) result in range
extensions and new information regarding the biology of these lizards. Significant range extensions are reported for the
rough-snouted giant gecko (R. trachyrhynchus) and for the recently rediscovered Guichenot's giant gecko (R. ciliatus). Field
observations confirm the association of the knob-headed giant gecko (/?. auriculatus) with plants of the family Cunoniaceae
and that Leach's giant gecko (R. leachianus) feeds on fruit. Range extension of some species, and data on local abundance
allows a reassessment of their conservation status. Despite implied increases in giant gecko density and range, significant
threats from habitat loss, introduced predators, and illegal trade leave all species at risk.

Key words. Rhacodactylus, geckos. New Caledonia, distribution, diet, conservation status

Introduction New Caledonia have yielded new distributional and/or dietary


Rhacodactylus is one of three carphodactyline' gecko genera information for all six of the recognized species of Rhacodactylus.
occurring in New Caledonia. The genus includes the largest liv- In addition, preliminary assessments of genetic variation within
ing species of gecko, R. leachianus (> 250 mm snout- vent length; certain species (reported more fully in Good et al. 1997) were
Russell and Bauer 1986) and has attracted scientific attention made on the basis of tissue samples accumulated over a series of
because of unusual characteristics such as viviparity' (in R. trips since the mid-1980's. We present these data here as they
trachyrhynchus; Bartmann and Minuth 1979), the possession help to provide a more accurate picture of the geographic ranges
of and specialized dentition (in R. auriculatus; Bauer and Russell and biological requirements of the geckos and may be useful in

1990; Bauer and Sadlier 1994b). and prehensile tails (all species; establishing the conservation status of Rhacodactylus species.
Bauer 1990; Bauer and Russell 1994). Species of this genus have
also attracted much popular attention, especially among ter- Materials and methods
rarium keepers, because most species thrive and reproduce well Herpetological collections and observations were made on main-
in captivity (Henkel 1987, 1991, 1993; Henkel and Schmidt land New Caledonia during trips in 1994 and 1995. Specimens
1991; Tytle 1992). were collected under a series of permits issued by the conserva-
The systematics and morphology of Rhacodactylus have tion authorities of the Province Sud (Marcel Boulet) and Prov-
recently been the focus of significant investigation (Bauer 1990; ince Nord (Christian Papineau) of New Caledonia to the au-
Bauer and Russell 1990; Bauer et al. 1993; Seipp and Klemmer thors. Preliminary estimates of genetic divergence between popu-
1994; Good et al. 1997). However, knowledge of the distribu- lations were based on results derived from allozymic* data. De-
tion and biology of these geckos remains largely incomplete tails of the electrophoretic methodology employed are presented
(Bauer and Sadlier 1993). Aside from brief reports of various in Good et al. 1997. Specimens cited are housed in the collec-

aspects of natural history (e.g., Mertens 1964; Meier 1979; tions of the Australian Museum Sydney )-AMS,
( the Natural
Sameit 1985; Bauer 1990; Bauer and Vindum 1990; Henkel 1991 ), History Museum (London)-BMNH, and the California Acad-
field-based data are limited to a few investigations of diet (Bauer emy of Sciences (San Francisco)-CAS.
and DeVaney 1987; Bauer and Sadlier 1994b) and general ac-
counts of behavior and ecology in nontechnical works (de Vosjoli Results and discussion
1995; de Vosjoli and Fast 1995). However, increasing awareness Knob-headed giant gecko (Rhacodactylus auriculatus)
of the uniqueness of the flora and terrestrial fauna of New [Plate 1]. Bohme and Henkel (1985) reported a striped color
Caledonia (Myers 1988; Mittermeieret al. 1996) has given new phase of R. auriculatus, now known to be common. Although
impetus to the collection of basic distributional and biological polymorphisms were noted within a single population of this

data for members of the New Caledonian herpetofauna, and


all species, there were no fixed differences among population samples
previously understudied areas are being surveyed more system- from four different localities and no suggestion of significant
atically (e.g., Isle of Pines, Bauer and Sadlier 1994a). Observa- intraspecific genetic variation. This tends to corroborate mor-
tions made by the authors during several recent expeditions to phological observations that this species is generally
polymorphic'' throughout its range but that there are no geo-
Correspondence. Tel: (610) 519-4857; fax: (610) 519-7863:
graphically related trends in character variation (Bauer 1990).
email: [email protected]

24
Plate 3A Plate 3B

wHBjefr"'
K&
fflKf?^^]

BJJjgA ^ jH

SIsIm£•" ?

Plate 5A

Plate 5B Plate

Plate captions: 1. Knob-headed giant gecko, Rhacodactylus auriculatus, from Riviere Bleue. 2 Bavay's giant gecko Rhacodactylus
chahoua. Photo courtesy of R. D. Bartlett. 3A Guichenot's giant gecko, Rhacodactylus ciliatus, from Riviere Bleue (adult with autoto-
mized tail). 3B. Rhacodactylus ciliatus, from Riviere Bleue (subadult with complete original tail and complex body patterning). 4.
Leach's giant gecko, Rhacodactylus leachianus, from Mt. Koghis. 5A. Roux's giant gecko, Rhacodactylus sarasinorum, from Kwa
Neie (adult retaining bold white dorsal markings). 5B. Rhacodactylus sarasinorum, from Riviere Bleue (adult with mottled dorsal
pattern). 6. Rough-snouted giant gecko Rhacodactylus trachyrhynchus, from Mt. Aoupinie. Photos 1, 3A and B, 4, 5A and B, and 6: Ross
A. Sadlier.

25
AARON M. BAUER AND ROSS A. SADLIER

Klemmer 1994; Kullmann 1995) and several smaller offshore


islands (de Vosjoli 1995). De Vosjoli (1995) and de Vosjoli and
Fast ( 1 995) recorded this species as common on the Isle of Pines
but stated that it was not present on the mainland of New
Caledonia. Despite their claims, Bavay ( 1 869), whose data have
proved to be very accurate (see Bauer and Sadlier 1994b), re-
ported collecting seven specimens of this species at several (un-
specified) localities on the mainland, and the type locality given
by Guichenot (1866) was at Canala (21°35 S, 165°56' E), also T

on the mainland. The persistence of this species on the mainland


was verified by (AMS 146594-5) collected by
two specimens
R. A. Sadlier near Pont German, Riviere Bleue(22°06' S, 166°38'
E) in the extreme south of New Caledonia (Fig. 1 ). It has subse-
quently been taken at other localities on the mainland (Girard
and Heuclin 1998; Bauer and Sadlier 2000), suggesting that it

may be relatively widespread.

Leach's giant gecko (Rhacodactylus leachianus) [Plate

4]. This species has a broad distribution in the wetter areas of


the New Caledonian mainland, especially along the east coast
(Bauer 1990). Boulenger(1885) first recorded the species from
the Isle of Pines (BMNH 53. 8. 16. 13). Bauer and Sadlier (1994a)
confirmed the presence of this species on the island with a 194
mm female (CAS 182197). Subsequently, the species has been
recorded as fairly common on the Isle of Pines and nearby off-
shore islands (de Vosjoli 1995; de Vosjoli and Fast 1995). Al-
though most known mainland New Caledonian localities are in
low- to middle-elevation forests (Bauer 1990), specimens have
been recorded from up to 1 100 m (Mertens 1964). Sight obser-
Plate 7. Geissois sp. from Mt. Do, southern New Caledonia.
This plant is apparently utilized as a food source by Rhaco-
vations in January 1995 at 540 m on Mt. Mandjelia (20°24'15"
dactylus auriculatus. S, 164°3ri8" E) extend the confirmed distribution of the spe-
cies to the northwest, almost to the limit of the humid forest on
Bavay (1869) first reported on the diet of this species, the main island of New Caledonia.
indicating that it eats the flowers of Geissois (Cunoniaceae). The Isle of Pines population has recently been described
This was verified by the recovery of anthers, stamens, and (pos- by Seipp and Obst (1994) as a distinctive subspecies,
sibly) pollen belonging to either a member of this family (or the Rhacodactylus leachianus henkeli. The validity of this form is
Myrtaceae) from the stomach of one specimen (Bauer and Sadlier challenged on the basis of morphological and allozyme charac-
1994b). On 9 January 1995 further evidence of the specific ters by Good et al. 1997. They found the henkeli color pattern
association between R. auriculatus and Geissois was obtained to occur among geckos in at least two regions of the New

when geckos were found active on flowering specimens of Caledonian mainland and regarded behavioral differences as at-

Geissois spp. (Plate 7) 1 .3-2.6 km from the summit of Mount tributable to reduced predation pressure on the insular form.
(Mt.) Do (21°45' S, 166°00' E) in south central New Caledonia. Because genetic distance data' indicated no long separation of
Isle of Pines R. leachianus from mainland populations Good et

Bavay's giant gecko (Rhacodactylus chahoua) [Plate al. 1997 regarded the split of the insular population to be very
2]. The known distribution of this species in central and south- recent. Indeed sea level minima of 100 in or more would have
ern mainland New Caledonia has been expanded by the capture connected New Caledonia to the Isle of Pines as recently as
of a specimen from Sarramea (AMS R 44 171) and by speci- 1 16,000-20,000 years ago (Stevens 1973; Holloway 1979).
mens from unstated localities on the Isle of Pines (de Vosjoli Although the diet in captivity of Rhacodactylus leachianus
1995; de Vosjoli and Fast 1995) [Fig. 1]. This species is rather has been well documented (Mertens 1964; Bauer and DeVaney
polymorphic with respect (Bavay 1869; Bohme
to coloration 1987; Henkel and Schmidt 1991 ), and a few stomach contents
and Henkel 1985; Bauer 1985), but the comparison of allozymes have been reported (Roux 1913), the natural diet remains poorly
from individuals separated by more than 100 km suggests rela- documented. At Mt. Aoupinie, in January 1995, we observed
tive genetic uniformity. individuals feeding on fruit in humid forest trees. Examination of
feces of freshly captured individuals revealed only fig seeds and
Guichenot's giant gecko (Rhacodactylus ciliatus) [Plate partially digested fig fruit. It appears likely that this, and per-
3A and 3B]. This species was numerous for the first 20 years haps other Rhacodactylus species, take advantage of seasonal
after its description (e.g., Bavay 1869) and then was not seen and local availability of figs and may play a role in seed dis-
again for over 1 00 years, despite extensive searches by several persal.

researchers. It was regarded as extinct (see Bauer and Sadlier


1 993 ). In 1 994, the species was rediscovered and has since been Roux's giant gecko (Rhacodactylus sarasinorum)
found at a variety of localities on the Isle of Pines (Seipp and [Plate 5 A and 5B]. Bauer ( 1 990) figured the type of R. sarasinorum

26
NEW CALEDONIA
j
New Caledonia is a French overseas territory, consisting of the large island ofNew Caledonia and the Loyalty Islands. Its location is approximately 1 ,200
km east of Australia (Geographic Coordinates 2 °30' S, 65°30' E) in the South Pacific Ocean. These islands have an extraordinary diversity of fauna and
1 1

flora with anextreme level of endemism in many taxa including birds and reptiles. Naturally occurring plant species number 3,380 (vascular plants), birds
116, mammals 9, and reptiles 87 (71 terrestrial and 16 marine). No naturally occurring amphibian species exist on New Caledonia though a nonnative
species has been introduced from Australia (green and golden bell frog Litoria aured). Total area is 1 9,060 square (sq) km (land 1 8,575 sq km and water
485 sq km) comparatively, slightly smaller than New Jersey. The terrain is wesfcoastal plains with interior mountains (highest point Mont Panie 1 ,628
m) making up two-thirds of the island. The climate is subtropical (warm and humid) modified by southeast trade winds. There is little temperature change
throughout the year, averaging between 7 1°F and 75°F (22°C and 24°C). The natural vegetation comprises tropical evergreen rain forest up to 1 ,000 and m
tropical montane rain forest above 1 ,000 m. Mangroves occur along western coasts. The major vegetation types are dense evergreen forest (22.8% of total
land area), Niaouli savanna woodland (1 3.8%), maquis vegetation in mining areas (25. 1%), savanna grassland (21 .7%), and scrub "(8.3%). New Caledonia's
human population numbers 191,003 (July 1997 estimate) with a 1.68% (1997 estimate) annual growth rate. New Caledonia's moderately developed
economy is based on mining and has more than 20% of the world's known nickel resources as well as other natural resources as chrome, iron, cobalt,
manganese, silver, gold, lead, and copper (thus mining is an important environmental issue). Only a negligible amount of the land is suitable for cultivation
and food accounts for about 25% of imports. In addition to nickel, financial support from France and tourism are key to the health of the economy. The
principal threats to the natural flora and fauna are mining, logging, and bushfires, reducing the forest cover from an estimated 90% cover to just 20%.'

and noted variation in color pattern and body pro-


portions in this species but did not elaborate. At
two color morphs have been illustrated and
least

described by Henkel (Bohme and Henkel 1985; U I i i T I i i i I

Henkel 1987, 1988), but there has been no sug-


gestion of subspecific or specific distinction be-
tween these forms. Bauer's ( 1 990) and Bauer and
Vindum's (1990) concept of R. sarasinorum was
based in part on typical specimens and in part on
an individual from Touaourou that is larger, darker,

and differs from other specimens in a number of


scale counts. Allozyme analysis (Good etal. 1997)
revealed that this specimen differed from a typi-
cal R. sarasinorum from Riviere Bleue (AMS R
146596) by four fixed differences. This is a greater
genetic difference than that between R. ciliatus
and R. chahoua. Both allozyme and morphologi-
cal data thus suggest significant variation and are
Fig. 1. Distribution of Rhacodactylus ciliatus
being analyzed separately, which may result in (triangles). R. chahoua (squares), and R.
trachyrhynchus (circles) in the New Caledonia
the recognition of a new Rhacodactyhts sara- region. Open symbols represent older records
sinorum-\ike species. as summarized by Bauer and Henle (1994), new
records are indicated by solid symbols. Note
especially the range extension of R trachyrhny-
chus to the west coast at Pindai and the docu-
Rough-snouted giant gecko (Rhacodacty- mentation of the occurence of the other species
lus trachyrhynchus) [Plate 6]. Bauer (1990) on the Isle of Pines.

recorded five mainland New Caledonian localities I I

for R. trachyrhynchus. Several of these, Coula-


Boreare, Ciu, and Mt. Gouemba are in the eastern humid forest scincus festivus), New Caledonian skink (Caledoniscincus
region of the island. The other two localities, La Foa and near austrocaledonicus), and a new species of elf skink (Nannoscincus
Noumea are imprecise but probably are also humid forest locali- sp.). All previous records of Rhacodactylus spp. from the New
ties. All localities are at middle to low elevation. As briefly Caledonian mainland have originated from the wetter eastern
noted by Bauer (1995), recent censuses have expanded the portions of the island, or from rainforest or maquis vegetation*
known range of the species, both elevationally and geographi- in the south (Bauer 1990; Henkel 1991, 1993; Bauer and Henle
cally (Fig. 1 ). Five specimens (AMS R 464 1 1 7-9, CAS 200266- 1994). The occurrence of R. trachyrhynchus at this site is thus

8) were found during rainstorms in humid forest (Fig. 2A) at intriguing and suggests a much broader habitat tolerance range
approximately 520 m on Mt. Aoupinie in Central New Caledonia than previously suspected for this species. Although normally
(21°09'19" S, 165° 19' 12" E), 27 km north and west of the associated with large, mature rainforest trees (Meier 1979), at
previously documented range of the species. A single specimen, Pindai', this gecko was collected less than five meters from the
CAS 200269, was obtained in sclerophyll foresf at Pindai ground in the branches of a small tree. A very low genetic dis-
(21°20'02" S, 164°58'21" E) at approximately 20 m elevation tance from this specimen to specimens from Mt. Gouemba and
(Fig. 2B). This locality is also somewhat further north than Mt. Aoupinie (Good et al. 1997) suggests no significant differ-
earlier records, but it is unique in that it is a west coastal locality entiation in the dry forest population, and the specimen is typi-
in an area of low rainfall. The local vegetation is dominated by cal for the species in regard to morphology. Henkel ( 1 99 1 ) sug-
largely endemic dry forest plants and is regarded as the most gested that there are two morphs in the species, one with a
threatened terrestrial habitat in New Caledonia ( Jafrre et al. 1 993; short, wide, robust snout, the other less so, but because these
Bouchet et al. 1995). The only other reptiles collected sympat- features were noted in captive born specimens of uncertain lo-

rically with R. trachyrhynchus at Pindai were Vieillard's prehen- cality he did not imply any subspecific distinction. Our data,
sile-tailed gecko (Eurydactylodes vieillardi), Giinther's New from three widely scattered localities, do not support the recog-
Caledonian gecko (Bavayia cyclura), sclerophyll forest gecko nition of any specific or subspecific subdivisions within R.
{Bavayia exsuccida), festive New Caledonian skink (Caledoni- trachyrhynchus.

27
AARON M. BAUER AND ROSS A. SADLIER

Fig. 2. Habitat of Rhacodactylus trachyrhynchus. habitat types. The data reported on here necessi-
tate a re-evaluation of that status report.
Rhacodactylus ciliatus, previously considered
possibly extinct, is now known to be common on
islands of the south coast of New Caledonia and
present, if somewhat less common, on the main-
land. Using the terminology of Bauer and Sadlier
(1993), its distribution is now regarded as re-

stricted and its status as locally common. With


the extension of its known range to the north and
to the west coast sclerophyll forest, the distribu-
tion of Rhacodactylus trachyrhynchus can now
be upgraded from restricted to moderately wide-
spread and its status to locally common. The pres-
ence of this species, as well as a regionally en-

demic Bavayia (B. exsuccida) and a new, appar-


ently endemic, Nannoscincus in the sclerophyll

Overview of humid forest habitat at middle elevation on Mt. Aoupinie forest in west coastal New Caldeonia (Bauer et al.
1 998) adds impetus to efforts to protect the small
remaining tracts of this habitat. Rhacodactylus
leachianus was previously regarded as widespread
and uncommon. The extension of the east coast
range to the limit of humid forest underscores the
fact that this is the most widely distributed of all
Rhacodactylus species and observations of indi-

viduals under ideal weather conditions (warm and


wet) suggest that the species may best be catego-
rized as common in appropriate habitats. The sta-
tus of the remaining species is unchanged by the
new records.
Although
^HiHi^^HHHHnHMl
B. Sclerophyll forest habitat at Pindai on the west central coast of New
at least several
Rhacodactylus do appear to be locally abundant,
even the most widely ranging species
species of

is endemic
Caledonia.
to the New Caledonian mainland and adjacent sat-
The presence of this species on the Isle of Pines remains ellite islands, an area about the size of Connecticut. Further, no
problematic. Boulenger (1878) recorded the species (as species is known to be present in all native habitat types and all

Chamaeleonurus trachycephalus) from this locality. Subsequent are excluded from agricultural or urban environments (although
collecting activity on this island, however (Bauer and Sadlier they may be present at the periphery of human-modified areas).
1994a; de Vosjoli 1995) has not verified this occurrence. The Habitat destruction and the impact of introduced predators were
unexpected findings of this and other Rhacodactylus after many cited as the primary threats to the herpetofauna of New Caledonia
years of extensive and intensive searching, however, argue against by Bauer and Sadlier in 1993. These factors remain the most
dismissal of this early record. Bauer and Sadlier ( 1 994a) identi- significant conservation concerns, but the illegal pet trade in
fied appropriate habitat for the species on the island. Rhacodactylus has increased significantly in the 1990's and has
become a potential threat to wild populations.
Conclusions
The new dietary observations confirm earlier reports of her- Acknowledgments. — We thank the permit-issuing authori-
bivory by Rhacodactylus species. It is especially noteworthy ties inNew Caledonia for their continued support of our re-
that R. auriculatus was found active on flowering heads of plants search. We also thank Jean Chazeau (IRD) and Alain Renevier
of the same genus on which Bavay observed them over 130 and his family for their encouragement of our activities. Finan-
years ago. The significance of reptiles as seed dispersers and as cial support for this work came from Villanova University, the
possible pollinators was recognized by Borzi (1911) but re- California Academy of Sciences (AMB), and the Australian
mains largely unexplored. Its recent documentation for the closely Museum (RAS).
related carphodactyline geckos of New Zealand ( Whitaker 1 987)
suggests that at least some species of giant geckos are important References
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Rhacodactylus sp. and the significance of geckos as dispersers of Rhacodactylus trachyrhynchus Bocage 1873 aus Neukale-
donien. Salamandra 15:58-60.
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Bauer, A. M. 1985. Notes on the taxonomy, morphology and be-
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Bauer and Sadlier ( 1 993) reviewed the conservation status
nidae). Bonner Zoologische Beitrage 36:81-94.
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Bauer, A. M. 1990. Phylogenetic systematics and biogeography of
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28

NEW RHACODACTYLUS DATA

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Girard. F. and Heuclin, D. 1998. Premiere mention de gecko
1
The research sources used in writing the coi sidebar on page JO (Madagt
volume, and footnoted on page 14 (footnote 2 ), also used in writing the Ne
Rhacodactylus ciliatus sur la Grande Terre (Nouvelle-Caledonie)
depuis sa description en 1866. Bulletin de la Societe Herp-
etologique de France 85-86:60-61. Manuscript received: January- 1997
Good, D. A., Bauer, A. M., and Sadlier. R. A. 1997. Allozyme evi- Accepted: April- 1998

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