Varanus Salvator Ziegleri From Moluccas Indonesian Species

The Southeast Asian water monitor lizard Varanus salvator ziegleri (Laurenti, 1768) is among the largest extant squamate reptiles of the world. It is also the most widespread member of the genus Varanus (Böhme, 2003), ranging from Sri Lanka in the west through continental Southeast Asia, the Greater and Lesser Sunda Islands to Sulawesi and the Moluccas (Koch et al., 2007a). However, taxonomy and intraspecific diversity of this giant Asian monitor lizard species complex have long been neglected. Anew era started in 2007, when the three traditionally recognized subspecies from the Philippine Archipelago were elevated to full species status due to significant morphological differences (Koch et al., 2007a). In another recent study, the Philippine species were reinvestigated morphologically and it could be shown that this vast archipelago harbors no less than five different species of the closely related V. salvator complex (Kochet al., 2010).

Varanus salvator ziegleri

Another unresolved issue is the exact limit of the distribution range of V. salvator and its occurrence on the islands east of Sulawesi (see Koch et al., 2007a). De Rooij (1915), for instance, examined historical voucher
specimens from Halmahera but also listed Bacan (Batjan) within the species range. Mertens (1930, 1942,
1959, 1963) reported V. s. salvator from the islands of Obi, Halmahera, and Bacan at the eastern boundary, although no material from either of the latter two islands was available to him. Instead, he referred to earlier authors like Bleeker (1856, 1857, 1860a, 1860b), who had reported Varanus salvator ziegleri from the Moluccas. Brandenburg (1983) adopted these earlier records from Halmahera and Obi. More recent authors, however, largely ignored the possible occurrence of water monitor lizards in the Moluccas (see, e.g., Eidenmüller, 2009).

Recently, however, populations of Varanus salvator ziegleri were encountered on several Moluccan islands such as those of the Sula Archipelago and Obi (Weijola and Sweet, 2010). Subsequent investigations of the varanid collection at the Senckenberg Museum, Frankfurt (SMF), revealed one juvenile specimen of Varanus salvator ziegleri from the island of Obi, which was donated by the Museum Zoologicum Bogoriense on Java, Indonesia, in 1957. This voucher specimen formed the base for Mertens’ (1959) novel island record for Obi. The specimen exactly matches in coloration and pattern with a live specimen from Obi as depicted by Weijola (2010). Due to unique characteristics in the color pattern of the Obi population, we here describe a new subspecies of the Southeast Asian water monitor lizard Varanus salvator ziegleri.

Field work on Sulawesi and adjacent islands was conducted between 2005 and 2007. In addition, voucher
specimens of major European and Indonesian natural history museums were studied. Museum collection acronyms used follow Leviton et al. (1985). Examination methods of standard morphometric and meristic characters used in this study are based on the works of Mertens (1942), which were later further developed and expanded by Brandenburg (1983) and Böhme et al. (1994). For each voucher specimen examined
21 external morphometric and meristic characters were recorded. In order to minimize the errors based on
observer bias, all measurements and scale counts weremade by the senior author. In addition, four proportion
indices were calculated.

Head measurements were taken with a slide-caliper to the nearest 1 mm. Body and tail lengths of subadult and adult specimens were taken with a measuring tape. Scale counts were recorded using pins and with check marks. A binocular microscope or a magnifying-glass was used for juvenile specimens with very small scales. For the measurement of the scalation character Q, the first continuously tail-spanning row of scales near the tail base was counted by excluding the first rows immediately after the cloaca, which form a non-continuous bow.

The transverse scale rows on the ventral side to the insertion of the hind limbs (character T) were counted up to the last continuous row when the scales eventually decrease and are of irregular shape. In contrast to Brandenburg (1983) and Böhme et al. (1994), “N” instead of “n” abbreviates the number of gular scales from the tip of the snout to the gular fold, to avoid confusion with the standardized variable “n” for the number of voucher specimens examined.

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Varanus Macraei Breeding and Distribution

Varanus macraei was only recently described by Böhme and Jacobs (2001) and is known only from Batanta Island, northwest of the Vogelkop Peninsula of Irian Jaya, Indonesian New Guinea. The species belongs to the emerald tree monitor (V. prasinus) group, which is currently comprised of nine species: V. prasinus, V. beccarii, V. kordensis, V. bogerti, V. keithhornei, V. telenesetes, V. macraei, V. boehmei and V. reisingeri (Ziegler et al., 2007). Varanus macraei is a large, slender tree monitor with smooth, unkeeled nuchal scales and a unique dorsal color pattern consisting of numerous oblique, irregular rows of blue ocelli on a black ground color.

Varanus Macraei

Maximum size is 110 cm in total length, with a maximum snout vent length (SVL) of 36 cm in males and 31.3 cm in females (Böhme & Jacobs, 2004). Varanus macraei is a diurnal, highly specialized tree monitor, as are the other members of the V. prasinus group. Because the size of Batanta Island is only 455 km², with a maximum length of 61 km and a maximum width of 13 km, V. macraei has one of the smallest distributions among Varanus, and virtually nothing is known about its habitat and natural history (Böhme & Jacobs, 2001; 2004).

Varanus Macraei

The first breeding of V. macraei in captivity was published by Jacobs (2002). Further breeding successes in private  facilities were subsequently published by Dedlmar (2007) and Moldovan (2008). Moreover, Mendyk (2007) reported on dizygotic twinning in V. macraei and Dedlmar (2008) on a two-headed hatchling. However, to our knowledge, there have been no documented zoo breedings of this rare and certainly endangered species. Thus, the first successful zoo breedings of V. macraei, at the Plzen and Cologne Zoos are herein reported.

Breeding

On 21 January 2004, the Plzen Zoo received a subadult pair of V. macraei, which was sent from Jakarta and subsequently confiscated at Prague airport. In April 2009, the female specimen measured 300 mm SVL and
620 mm tail length (TL) (920 mm in total length), the male measured 330 mm SVL and 660 mm TL (990
mm in total length). Three years later, in January 2007, Cologne Zoo received a confiscated pair of adult V.
macraei from the Wildtier- und Artenschutzstation e.V. Sachsenhagen in Germany. On 28 April 2009, the female specimen  measured 275 mm SVL and 575 mm TL (850 mm in total length), the male measured 290 mm SVL and 640 mm TL (930 mm in total length).

Both wild-caught pairs made up the unrelated breeding stocks of the Plzen and Cologne Zoos. Both  breeding pairs from Plzen and Cologne are undoubtedly members of the V. prasinus group and assignable to the species macraei due to their prehensile tails which are roundish in cross section, in combination with unkeeled, smooth neck scales, dorsal patterns consisting of transversal rows of blue ocelli, and tails bearing alternating blue and black bands (Ziegler et al., 2007).

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Varanus Prasinus Species Waigeo Papua Indonesian

On the basis of seven specimen I describe a new monitor taxon from the Indonesian isle of Waigeo (West Irian). This taxon tends to the melanistic forms of the Varanus prasinus or V. Prasinus group and shows golden yellow spots on black ground. The prehensile circular tail as well as the outer genital structures prove it to be a member of the subgenus Euprepiosaurus and the V. prasinus group. It can be differentiated from the other melanistic taxa by the lowest number of scales around midbody and the lowest number of ventrals and dorsals. The phylogenetic relationship to the other taxa of the group has to be reserved for a taxonomic review on molecular basis. With regard to morphological results the new species is closely related to V. kordensis and V. beccarii.

Varanus Prasinus

Most recently, a blue-coloured tree monitor from Batanta Island, West Irian (Indonesia) has been discovered and described as a new species: Varanus macraei BÖHME & JACOBS, 2001. In this paper, the authors referred already to another insular form from the neighbouring island of Waigeo which, according to photographs available in the internet, was easily distinguishable by its strikingly different colour pattern und was termed there as “golden speckled tree monitor” (see BÖHME & JACOBS 2001: 10). In the meantime, some live specimens could be obtained, and it turned out that my first view which was based on photographs only and which had led me first to regard the Waigeo population just as a colour variant of Varanus macraei, was not at all tenable. Rather, its colour pattern proved to differ not only in the colouration of the light body markings, but also in their arrangement, apart from a general chevron-like pattern which is more or less expressed present in all taxa of the Varanus prasinus group.

Varanus Prasinus

Varanus Prasinus

This includes currently seven taxa (prasinus, beccarii, bogerti, keithhornei, kordensis, macraei and telenesetes) whereas MERTENS (1942) recognised only four: the nominal subspecies prasinus, kordensis, beccarii, and bogerti. Despite the taxonomic review of the group by SPRACKLAND (1991), the taxonomic status of some of these taxa and their interrelationships are still not sufficiently analysed. In particular, molecular analyses are still lacking. In any case, closer examination and comparisons with the described forms led me to believe that a new, undescribed taxon, the eighth in the group, is involved, which I am going to describe below.

Varanus Prasinus

In November 2002 and January 2003 I was able to obtain six specimens of the Waigeo “golden speckled tree monitor” from private sources and from the pet trade. One of them died soon, serving now as the holotype and being deposited in the Zoologisches Forschungsinstitut und Museum Koenig, Bonn, Germany. The five remaining specimens – described below as paratypes – are still in my possession. They will be deposited after their natural death in the same institution.

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Oriental garden lizard, Bloodsucer Lizard and CALOTES VERSICOLOR

Calotes versicolor, Agamidae, Bloodsucker, Oriental Garden Lizard

MANY non-native lizard species of tropical origin have become established in southern Florida, and Florida now has more nonindigenous amphibian and reptile species than any other state (Butterfield et al., 1997). In this paper, we report the first established population of Calotes versicolor (Daudin 1802), an Asian agamid lizard, in Florida. Butterfield and co-workers (1997) included Calotes versicolor in their list of established exotic herpetofaunal species in Florida based upon a personal communication from R. D. Bartlett. However, the purported C. versicolor population was misidentified and was actually C. mystaceus (Bartlett, 2003), which occurs in two small areas in the vicinity of pet dealerships in Glades and Okeechobee counties (Bartlett and Bartlett, 1999).

Of the 21 recognized species of Calotes (Vindum et al., 2003), C. versicolor is the most widespread, ranging from southeastern Iran and Afghanistan east to Indo-China and as far south as Sri Lanka, Sumatra, and northern Peninsular Malaysia (Boulenger, 1912; Smith, 1935; Zhao and Adler, 1993; Cox et al., 1998).
Calotes versicolor is distinguished from its congeners by having lateral body scales pointing backwards and upwards, no oblique fold or pit anterior to the shoulder, two separated spines above the tympanum, and 35–52 scales around the body (Smith, 1935). Calotes versicolor exhibits obvious geographic variation in coloration, scalation, and size across its wide range, but only two subspecies have been  described (Auffenberg and Rehman, 1993, 1995); our specimens were of the nominate subspecies.

Males may measure 140 mm snout-vent length (SVL) and have a slender tail 295 mm long (Radder et al., 2001). Adults have large heads, massive shoulders, expandable dewlaps, and laterally flattened bodies with a crest extending from the neck almost to the tail. The ground color is typically dull brown, gray, or olive with
irregular dark brown spots or bars, but breeding males have pale yellow bodies with a large black patch on each side of the throat. Breeding males also develop bright orange or crimson areas around the head and shoulders, but this brilliant coloration can change rapidly depending upon mood or environmental conditions (Bhatti and Bhatti, 1986; Cox et al., 1998). This red coloration accounts for these lizards commonly being called bloodsuckers. Other names for this arboreal species are the oriental or tropical garden lizard, Indian tree lizard, crested tree lizard, and changeable lizard.

In its native range, Calotes versicolor is commonly observed in parks, gardens, agricultural areas, waste land, and open forests (Cox et al., 1998). They typically live among leafy undergrowth and grass in open habitats, but males often display from fences and other conspicuous perches (Cox et al., 1998). Juveniles forage and bask mostly at ground level, whereas subadults and adults spend much of their time on tree trunks, often in a head-down posture surveying for prey (Diong et al., 1994).

Calotes versicolor is a very adaptable, prolific species that is commonly found in human-altered environments and can survive even in urban areas in Asia (Erdelen, 1988). In its native range, C. versicolor preys primarily on arthropods, but adults prey occasionally on their own young as well as small birds and nestlings, frogs, geckos, and small snakes (Rao, 1975; Dhindsa and Toor, 1983; Sharma, 1991a, 1991b, 1999; Diong, 1994). Calotes versicolor was introduced into Singapore in the 1980s, where it rapidly established populations (Chou, 1994) and partially displaced populations of a native agamid, the green crested lizard (Bronchocela cristatella) (Diong et al., 1994). We do not know what effect C. versicolor will have on populations of sympatric native lizard species in Florida, such as the green anole (Anolis carolinensis), ground
skink (Scincella lateralis), and southeastern five-lined skink (Eumeces inexpectatus).

Populations of the non-native brown anole (Anolis sagrei) and knight anole (A. equestris) are already common in the area, and non-native brown basilisks (Basiliscus vittatus) and green iguanas (Iguana iguana) are occasionally observed in adjacent areas (Flowers, 2003; Ward, 2003). Calotes versicolor is yet another exotic species that may compete with or prey upon Florida’s native species.

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Varanus Zugorum Habitats In Asia

This is the only Varanus Zugorum which was not observed during fieldwork. It appears to be almost completely unknown by locals, and only two reasonably reliable local eyewitness accounts were collected, in Akesahu (lower western side of Kao Bay) and Labi Labi (northeast Halmahera). Searches and interviews around the type locality Pasir Putih were unproductive. The holotype was collected in December 1980 by a villager, so further details from the initial discovery remain unknown (Adam Messer, pers. comm. 2008).

Varanus Zugorum

Monitor communities and niche segregation by sympatric Varanus Zugorum have been studied by several workers, including Pianka (1994), Shine (1986), Sweet (1999, 2007) and Philipp (1999). Communities range from simple one species systems up to as many as 11 species in certain areas of northern Australia (Sweet, 2007).

Varanus Zugorum

 These studies have shown that interspecific competition is typically minimized by differing use of habitat,
different body sizes, and foraging mode. Additionally, phylogenetic distance may aid in reducing niche overlap
in the most diverse communities of Australia and New Guinea.

A community comparable to the one on Halmahera, though slightly less diverse, was studied by Philipp
(1999) in West Papua. He investigated the habitat use of three closely related species of the indicus-group (V.
doreanus, V. indicus and V. jobiensis), and concluded that competition and interaction is largely avoided by
the use of different habitats and/or microhabitats.

This study found that on Halmahera and Obi, V. rainerguentheri is similar in habitat use to that of V. indicus on New Guinea (Philipp, 1999), while V. caerulivirens appears to fill a similar niche to that of V. jobiensis. Varanus yuwonoi overlaps widely in habitat use with V. caerulivirens but grows significantly larger, may use a different hunting strategy, and concentrate on larger prey items. Thus there seem to be clear niche separations in communities of V. indicus group animals in Moluccan multi-species communities as well. The ecological separation where members of Euprepiosaurus and V. salvator overlap is not equally obvious.

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Varanus Cf. Salvator Habitat In Indonesian

The distribution of Varanus Cf. Salvator  in the Moluccas has been uncertain for a long time, probably starting with what appears to be an erroneous record for Halmahera in de Rooij (1915), widely cited since (for example Bennett, 1995, Harvey and Barker, 1998). Obi, Seram and Bacan were mentioned by Mertens (1930, 1942, 1959), and Buru by Bleeker (1857). Since recent fieldwork, including this study, has not found support for many of these localities, it is suggested that all except Obi be omitted from the range of Varanus Cf. Salvator. Since the animals on Obi do not completely correspond with any of the described Varanus salvator group taxa, it is here employed with a cf. (meaning “compare with”).

Varanus Cf. Salvator

Varanus Cf. Salvator Distribution (in the north Moluccas): Obi Island, probably also on nearby Bisa, but not confirmed.

Varanus Cf. Salvator Ecology

Five observations were made of this species during two weeks on Obi. The sympatry between members of Soterosaurus and Euprepiosauris is interesting, since some authors have presumed that they might be ecologically exclusive (Mertens, 1942, Sprackland, 2004, Ziegler et al., 2007b). This proves to be incorrect since this study confirms sympatry of V. cf. salvator with V. rainerguentheri and V. caerulivirens on Obi, as well as two other species in the Sula islands (Weijola and Sweet, in press).

Varanus Cf. Salvator

Varanus Cf. Salvator

 All observations of V. cf. salvator on Obi occurred in coastal mangrove forest and periodically flooded littoral vegetation, where it is widely syntopic with V. rainerguentheri. They most likely also occur in inland forests and around freshwater streams side by side with V. caerulivirens. On Obi, they appear less numerous than V. rainerguentheri (5 vs. 22 observations), and similarly on Sanana, many fewer Varanus cf. salvator were observed compared to a sympatric member of the V. indicus complex during the course of fieldwork.

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Varanus Yuwonoi Origanaly Indonesian Reptiles

Varanus Yuwonoi (V. yuwonoi)

The black-backed mangrove monitor was only observed around the villages of Akesahu and Kao, both in the western part of Teluk Kao (Kao Bay) on Halmahera. The type series was collected near Jailolo (Harvey and Barker, 1998). Interviews with locals and animal dealers suggest that this species is widespread throughout the island. Eyewitness accounts of Varanus yuwonoi like animals from Morotai, Bacan, and even Obi exist, but still need confirmation.

Varanus yuwonoi is a difficult species to study. Despite frequent observations by local hunters, specimens are very rarely encountered out in the open. Considering the difference in size, which is presumably also reflected in trophic level, this species probably does not reach population densities as high as Varanus caerulivirens. A less active mode of hunting (sit and wait/ambush) may also result in the fewer encounters with V. yuwonoi.

Varanus Yuwonoi-V. Yuwonoi

Traps used by locals specifically for this species are always set up around megapode bird (scrubfowl) nests in
inland forests. The unusual pigmentation of the tail, body and neck breaks up the silhouette of the animal very well as they lie on the forest floor. Considering that the smaller and more predator-vulnerable species in the Moluccas have not evolved such an elaborate camouflage, it may have evolved rather as a means to escape detection from potential prey. The head is comparatively powerfully built and the teeth proportionately longer than any of its close relatives, even in comparison to most other more distantly related varanids.

Considering these facts, the author hypothesizes that Varanus yuwonoi is an ambush predator, favoring sites in the forest such as megapode nests, that are frequented by a variety of potential prey items. It is uncertain whether large individuals can kill adult scrubfowl, but the large nest mounds attract a number of smaller birds such as pigeons, and also lizards, snakes, and invertebrates, as well as the frequently hatching megapode chicks. According to local people, this species is also occasionally seen digging into these nests in search of eggs.

Varanus Yuwonoi

Additionally, observations of intraspecific aggression within the Varanus yuwonoi and Varanus doreanus clade (Ast, 2001) in captivity would support a hypothesis that V. yuwonoi could be an ambush predator. With few exceptions (Sweet, 1999, 2007), widely foraging monitor (and other scleroglossan lizards) species are not territorial; however, the two species (Varanus scalaris and Varanus glebopalma) known to be territorial are both ambush hunters. For a sit and wait predator favoring prey “hotspots”, a certain degree of territoriality could be expected to evolve. The largest individual measured 146 cm in total length; this specimen however did not appear to be fully grown, and according to locals they occasionally grow much larger.

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