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By Pat Matyot, C/o SBC, P.O. Box 321, SEYCHELLES
Abstract.— There is evidence
that the Asian agamid Calotes
versicolor (Daudin,
1802), the crested tree lizard, is now established on Ste Anne
Island in Seychelles, and it is reported to be dispersing away from
its original point of introduction. Data collected outside
Seychelles on its habitats, reproductive biology and feeding habits
show that this species is adaptable, prolific and omnivorous, and it
is considered to be an invasive alien species that competes with or
feeds on native biota in some parts of the world, such as Singapore
and Mauritius. The Ste Anne population needs to be studied and, if
possible, eradicated, to prevent this potential ecological threat
from reaching other islands in Seychelles, especially those that
harbour significant populations of native animals.
Introduction
The crested tree lizard, Calotes
versicolor (Daudin,
1802), is a strong candidate for the status of most widespread non-Gekkonid
lizard in the world. Günther
(1864) noted: ”This is one of the most common lizards,
extending from Afghanistan over the whole continent [sic] of India
to China; it is very common in Ceylon [=Sri Lanka]…” Its present
distribution stretches from Oman to the west (Loman
1997; Seufer
et al.. 1999) (the following
in Savy (1982) is
presumably a reference to Oman: “… more recently the British Museum
was sent a specimen from southern Arabia”) right across southern and
south-east Asia to Indo-China to the east (Stuart
1999), the Maldives (Hasen
Didi 1993), Réunion (Permalnaïck
1993), Mauritius (Staub
1993), (including Rodrigues (Blanchard
2000)), Seychelles (Matyot
2003) and Florida in the United States (Enge
& Krysko 2004). With growing evidence that
C. versicolor is an invasive
species with potentially harmful ecological impacts in the areas
where it is introduced (Diong
et al.. 1994;
Mauremootoo
et al. 2003), largely
because of its omnivorous feeding habits, it is necessary to gather
together the available, albeit limited, information on the status of
its presence in Seychelles. At the same time the opportunity is
taken to review the existing literature on the ecology of the
species with a view to summarizing the available data as a basis for
further research on its status as a potential threat. (Among the
various common English names used for this species (garden lizard,
bloodsucker, etc.) “crested tree lizard” is felt to be the most
appropriate in the Seychelles context, to avoid confusion with other
species found in gardens, so as not to perpetuate superstitious
beliefs, etc.)
Methods
Video footage of a specimen caught on Mahé in October 2003 filmed by
the Seychelles Broadcasting Corporation (SBC) on November 6th
2003 (Racombo 2003)
was viewed, and interviews were carried out with environment
officers and others who have been involved in sightings of
specimens. Photographs of the Mahé specimen filmed by the SBC were
sent to two authorities on the genus
Calotes,
Hidetoshi Ota (Japan)
and Scott Moody (USA),
who both confirmed it to be C.
versicolor. (Ota,
pers. comm. 2003;
Moody,
pers. comm. 2003). A
literature search was undertaken to find all references to
C. versicolor in Seychelles,
as well as references to the ecology of the species worldwide.
Presence in Seychelles
The first observation of C.
versicolor in Seychelles dates back to the 27th of
September 1982 at Barbarons on Mahé island when “a strange lizard …
was found resting in the grass by a young man who handed it over to
the Ministry of Agriculture” (Anon. 1982). The specimen measured
38.6 cm long, including the tail, and was therefore probably a
fully-grown individual (Diong
et al.. 1994). It
was sent to the Natural History Museum in London, where it was
identified as C. versicolor
(Savy 1982). In the
mid 1980s (“about 1985-1986”) a Seychellois offered to sell two
specimens that he said he had found on Mahé to a visiting American
herpetologist, Ronald
Nussbaum, but refused to provide information on where he had
captured the lizards when the offer was turned down (Nussbaum,
pers. comm., 2004). There were no further reports of
C. versicolor in Seychelles
until October 2003, when a specimen was discovered on a
Hibiscus sp. bush near Anse
aux Pins clinic on Mahé (Jeanne
Mortimer, pers. comm.
2003; Racombo 2003;
Matyot 2003). It was
identified as C. versicolor
by its dorso-nuchal crest, the two widely separated spines on either
side of the head above the tympanum, and the absence of any fold or
pit in front of the shoulder (Ota,
pers. comm. 2003). This, the
only recent Mahé record, was followed by further sightings, on a
regular basis from November 2003 onwards, in the vicinity of the
hotel on Ste Anne island, 5 km from Port Victoria on Mahé (Lena
Desaubin, Rodney Fanchette & Maurice Loustau-Lalanne,
pers. comm. 2003;
Matyot 2003),
providing evidence that C.
versicolor is now firmly established on Ste Anne: in November
2003 there were both sexes there, including at least one male seen
in breeding coloration (anterior part of the body, including the
head, orange-red with a black patch on each side of the throat) and
a gravid female that had been crushed by a vehicle, with the eggs
that it had been carrying very much in evidence. On January 16th
2004 the Ministry of Environment and Natural Resources announced a
bounty of fifty Seychelles rupees for every specimen of
C. versicolor caught (Anon.
2004). Latest reports speak of crested tree lizards having been
observed in parts of Ste Anne away from where the hotel is situated
(Rodney Fanchette,
pers. comm. 2004).
It is not certain how C. versicolor
was introduced to Seychelles. In view of the long gap of 17-18
years between the 1985-1986 and 2003 sightings, there may have been
two separate introductions, an earlier one on Mahé that did not last
and a more recent one on Ste Anne that has been successful. One
connection between Ste Anne and Anse aux Pins, the locality for the
only recent Mahé sighting, is that some of the expatriate hotel
personnel working on Ste Anne is known to have accommodation
quarters at Anse aux Pins. Unintentional transport by humans is a
very strong possibility – it is certainly believed to have played a
big role in the spread of C.
versicolor elsewhere, e.g. the crested tree lizard is said to
have been introduced to Réunion island in around 1865 as a stowaway
in a shipment of sugarcane (Saccharum
officinarum, Poaceae) cuttings from Java, Indonesia (Permalnaïck
1993; Staub
1993). In Florida, USA, however,
C. versicolor escaped from a reptile dealer in 1978 (Enge
&Krysko 2004).
Review of literature on ecology
of C. versicolor
Since the 1970s C. versicolor
has gained importance as a laboratory animal, and researchers
in Asia in particular have been using it as a subject for a broad
spectrum of biological studies. This has resulted in a considerable
output of literature on anatomy (e.g.
Paranjape 1974),
morphometrics (e.g. Tiwari &
Schiavina 1990), endocrinology (e.g.
Ganesh & Raman 1995;
Shanbhag
et al. 2000), embryology
(e.g. Muthukkaruppan
et al. 1970;
Ji
et al. 2002) and cytology
(e.g. Ota
et al. 2000). However,
there is relatively little published material on the ecology of
C. versicolor. This is
surprising in view of its recognized status as an invasive species
that is extending its range and, in some colonized areas at least,
is impacting negatively on native biodiversity (Diong
et al.. 1994;
Mauremootoo
et al. 2003). The best
general account of the species, based on original fieldwork in
Singapore, is that of Diong
et al. (1994).
On the other hand, several of the laboratory-based studies referred
to above have produced findings on the biology of
C. versicolor that shed
light on key aspects of its ecology.
Cytology.—
Ota
et al. (2000) have suggested
the presence of cryptic taxonomic diversity in
C. versicolor, i.e. that “C.
versicolor” could in fact be a complex of different species,
after the karyotype they worked out did not match those recorded by
previous studies. It would be interesting to carry out a study of
karyotypes from specimens caught in Seychelles and elsewhere in the
western Indian Ocean to possibly elucidate the provenance of the
Seychellois population.
Habitats.— Being a
semi-arboreal, sun-loving lizard that spends a lot of time on
tree-trunks and rocks, C.
versicolor has a predilection for open scrubland, wasteland,
gardens, parks and other “man-made habitats” in Asia (Diong
et al. 1994;
Erdelen 1984).
Stuart (1999) reports
that it is found on low vegetation in open forest or disturbed areas
near human habitation in Laos. In a study of diurnal lizards in the
city of Karachi in Pakistan, it was the second most numerous species
(Khan & Mahmoud
2003). In Réunion, Mauritius and Rodrigues it favours areas of
“grassy savannah”, such as that on the west coast of Réunion (Blanchard
2000) as well as cemeteries and along roadsides (Permalnaïck
et al. 1993). In
Florida, USA specimens have been collected in an area of grasses and
weeds in the vicinity of a canal running along a dirt road (Enge
& Krysko 2004). The species appears not to favour dense
forest with closed canopies (Permalnaïck
et al. 1993;
Diong
et al. 1994;
Pawar 1999).
Erdelen (1984) found
it to be most abundant in the driest parts of Sri Lanka. It is
reported to occur from sea level to an altitude of 600m (Stuart
1999) or even 1,000 metres (Diong
et al. 1994). It has
a sit-and-wait hunting strategy, usually watching for prey from a
vantage point on a tree-trunk, but moving into the shade, including
among high grasses, when it gets too hot; adult males stay in the
open more often to exercise territoriality, while adult non-breeding
females and, especially, juveniles tend to forage on the ground in
grassy and shrubby vegetation (Diong
et al. 1994;
Sunderasan & Daniels
1994). In one study (Sundaresan
& Daniels 1994) an adult male was found on the same tree on
15 occasions. C. versicolor
is reported to be able to swim in both fresh and sea water (Permalnaïck
et al. 1993). In Florida,
USA (Enge & Krysko
2004) as well as in Mauritius (pers.
obs.) C. versicolor
roosts on vegetation at night, up to 9m from the ground, usually at
the tips of twigs, shoots and inflorescences.
Reproductive biology.—
Aspects of the reproductive biology of
C. versicolor, a
multi-clutched seasonal breeder in India (Shanbhag
2003), are summarized in Table 1. To
Bhagyashri Shanbhag
and his colleagues at the Department of Zoology of Karnatak
University in India we owe a series of studies that have revealed an
array of reproductive strategies that help to explain the
adaptability of the crested tree lizard (Shanbhag
2003), among which are the following:
a) the variation of clutch and egg size depending upon the time of
breeding: towards the end of the breeding season clutches are
smaller and eggs larger; production of heavier hatchlings at the end
of the breeding season may enhance their chance of survival in
competition for food, shelter, etc with older offspring from earlier
clutches;
b) the storage of viable sperm by the female for as long as six
months, eliminating the need for repeated mating; and
c) the retention of eggs in the oviduct for six months or even more
when conditions are not suitable for oviposition.
Feeding.— Records of prey
and other food items of C.
versicolor, most of them based on observations in India, are
summarised in Table 2. It would seem that the crested tree lizard
is predominantly insectivorous (according to
Diong
et al. 1994, “analysis of
stomach contents [in Singapore] reveals a diet comprising mainly
ants, larval and adult insects, and other small invertebrates”), but
it is clearly an opportunistic omnivore that can prey on small
vertebrates and feed on plant material as well. One particularly
interesting observation (Kalita
2000) is of a crested tree lizard charging and attempting to
bite an Oriental magpie robin (Copsychus
saularis) in a fight over a centipede (Scolopendra
sp.).
Predators.—
C. versicolor is in turn
preyed upon by a number of carnivorous reptiles, birds and mammals.
Published records of predators are summarised in Table 3.
Parasites.— The literature
on the parasites of C. versicolor
is extensive (e.g. Diong
et al. 1999;
Goldberg
et al. 2003;
Sathyanarayana & Premavathy
1994; Schmaschke
et al. 1997) The range of
identified ecto- and endoparasites is shown in Table 4, but this is
far from an exhaustive list. There may be interactions between
parasites: for example, mites have been shown to transmit a species
of Schellackia (Apicomplexa)
to other lizards (Bonorris &
Ball 1955).
Discussion of status as an ecological threat
The continuing extension of the range of
C. versicolor outside Asia,
to the Middle East, the western Indian Ocean and Florida in the
United States, is due to a combination of intentional and
unintentional transport and introduction by humans. At the same
time, the alteration of natural habitats through anthropogenic
factors, including deforestation, must have created suitable
habitats for the species where there were none before.
Surprisingly, although it is an adaptable, prolific and omnivorous
species, its impact on native biodiversity does not appear to have
been scrutinized very closely in the localities where it has become
established. However, Diong
et al. (1994) have
drawn attention to the fact that in Singapore it has “to some
extent” displaced the native green crested lizard,
Bronchocela cristatella (Kuhl,
1820); and Mauremootoo
et al. (2003) include
C. versicolor in their list
of introduced vertebrates thought to have a significant impact on
native biodiversity in Mauritius: they state that it competes with
native geckos and consumes native invertebrates.
Vinson (1968)
speculated that C. versicolor
may have been responsible for the disappearance or rarefaction of
phasmids (Insecta: Phasmatodea) in both Mauritius and Réunion.
Crested tree lizards have been observed in localities where phasmids
have become less common in Réunion: this is the case for
Heterophasma multispinosa in
the uplands of Saint Joseph and
Monandroptera acanthomera at Mare Longue (Nicolas
Cliquennois, pers. comm.
2003). In the United States, there is apprehension that “Calotes
versicolor is yet another exotic species that may compete
with or prey upon Florida’s native species” (Enge
& Krysko 2004).
In Seychelles, potentially suitable habitat for the crested tree
lizard is found in coastal areas and perhaps even on some of the
high-altitude “inselbergs” of the inner, granitic, islands as well
as extensively in the outer, coralline, islands. While Ste Anne,
its present focal point, is itself not a high-value island in terms
of terrestrial fauna and flora, if
C. versicolor were to disperse to other islands this could
represent a serious new threat to native biodiversity, with prey
records from India and elsewhere (Table 2) showing the wide range of
groups of organisms that could be threatened. The smaller, rat-free
islands with extensive seabird colonies and populations of endemic
terrestrial birds, invertebrates, amphibians (Caecilidae on Frégate)
and reptiles (Gerlach
1997) would seem particularly vulnerable. It is not clear to what
extent the parasites harboured by
C. versicolor are host-specific, i.e. whether some of them,
at least, can infect native lizards like skinks (Mabuya
spp.) and geckos (e.g. Phelsuma
spp.) as well. It is likely that in Seychelles
C. versicolor would be
preyed upon, especially in the juvenile stage, by the endemic snakes
Boaedon geometricus and
Lycognathopis seychellensis
and the endemic Seychelles kestrel
Falco araea as well as domestic and feral cats (Felis
domesticus); but it is doubtful that they would suppress the
population of such a prolific and fast-maturing breeder.
Research is urgently required to elucidate the status and ecology of
the established population of C.
versicolor on Ste Anne island even if eradication efforts
announced by the Ministry of Environment and Natural Resources are
ongoing (Anon. 2004). Captured specimens could be sexed, measured,
weighed and dissected for stomach contents and, possibly,
parasites. Managing the spread of invasive alien species involves
(1) prevention, preferably, to keep potential invaders from entering
a new ecosystem; (2) early detection, if prevention is not
successful, to locate the alien before it has the chance to become
established and spread; (3) eradication, mostly possible when the
alien is detected early; and (4) control as a long-term process to
limit the population size and distribution of the alien if
eradication is not possible (Reaser
2003). In the Seychelles context, to manage the spread of the
crested tree lizard would involve all of these going on at the same
time on different islands within the archipelago.
Ackowledgements
The author is grateful to the following:
Sylvain Hugel in
Strasbourg, France, who contributed to the literature search; the
staff of the Ministry of Environment and Natural Resources in
Seychelles, who provided information on sightings of
C. versicolor;
Cheong Hoong
Diong
at Nanyang Technological University in
Singapore, who shared the results of his work on
C. versicolor; and the
Seychelles Broadcasting Corporation (SBC) for logistical and moral
support.
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