CSIRO PUBLISHING Australian Journal of Zoology, 2011, 59, 236–241 http://dx.doi.org/10.1071/ZO11094

Distribution, ecological attributes and trade of the New Guinea carpet python () in Indonesia

Daniel J. D. Natusch A,B and Jessica A. Lyons A

ASchool of Biological, Earth and Environmental Sciences, University of , Sydney, NSW 2052, Australia. BCorresponding author. Email: [email protected]

Abstract. Carpet pythons (Morelia spilota) are medium-sized non-venomous snakes inhabiting most of continental Australia and a small area of New Guinea. They have been relatively well studied in Australia, but little is known about the New Guinea population, even though it is harvested and exported from Indonesia for the international pet trade. In total, 281 locality records were compiled for two distinct populations south of latitude 7S in Papua New Guinea and the Indonesian province of Papua. Traders in Papua collected 274 carpet pythons (most of which were recently hatched neonates) for the pet trade. Data from a sample of 174 individuals revealed little sexual dimorphism in any traits, although males appear to grow larger than females despite females maturing at greater lengths. Fecundity was high (average 17 eggs) and reproduction was highly seasonal, with hatching in December and January. Harvest quotas for the province of Papua were exceeded in all years between 2000 and 2009 due to 50% of the national quota being allocated to West Papua province where this species is yet to be recorded. The present study provides distribution, trade and ecological information to inform conservation management of this species in Indonesia.

Additional keywords: harrisoni, quota, sustainable harvest, wildlife management, wildlife trade.

Received 21 November 2011, accepted 27 January 2012, published online 6 March 2012

Introduction The New Guinea carpet python is, however, popular in the The carpet python (Morelia spilota) is a widespread Australasian international pet trade and has been collected from the wild in species whose ecological traits show remarkable variation Indonesia and traded for this purpose since at least 1992 across their range (Shine and Fitzgerald 1995; Pearson et al. (CITES 2011). Indonesia is currently the only country that allows 2002). They are large (to 4 m) snakes and are found in a variety the legal harvest and export of wild carpet pythons, which are of habitats in temperate and tropical Australia (Wilson and listed in Appendix II of the Convention on International Trade in Swan 2008) and in the Trans-Fly and Port Moresby areas on the Endangered Species of Wild Fauna and Flora (CITES). The island of New Guinea (O’Shea 1996). Studies in Australia have nominated CITES Management Authority in Indonesia, the shown that body sizes and reproductive traits vary considerably Directorate General of Forest Protection and , allocates between areas, and one population from south-western Australia annual provincial harvest quotas for all species listed in exhibits the highest degree of sexual size dimorphism in any Appendix II of CITES. Quotas are reviewed annually under the species (Pearson et al. 2002). guidance of the Indonesian Institute of Science to ensure that In contrast to the considerable number of ecological studies harvest levels are not detrimental to the survival of the species of carpet pythons in Australia (e.g. Slip and Shine 1988a, (Siswomartono 1998). In both 2010 and 2011 the harvest 1988b, 1988c; Shine and Fitzgerald 1996; Fearn et al. 2001; quota for carpet pythons was 400 individuals, divided evenly Heard et al. 2004; Pearson et al. 2005; Corey 2007), the New between the provinces of Papua and West Papua (Anon. 2010, Guinea population has received relatively little attention. Hoser 2011). (2000) assigned specific status to New Guinea carpet pythons The current study provides the first data on distribution, and named them Morelia harrisoni. However, because disjunct ecological attributes (i.e. body sizes, reproduction and diet) and distribution was the only information provided by Hoser (2000) trade of carpet pythons in Indonesian New Guinea and compares to justify their differentiation from Australian conspecifics, the this information with that from previous studies of Australian scientific community has been reluctant to adopt the name carpet pythons. The data enable preliminary assessment of the (Wüster et al. 2001). To date, there have been no ecological sustainability of the carpet python trade in Indonesia and can be studies of carpet pythons in New Guinea and even basic used to make more informed conservation management decisions information such as geographic distribution is still not known. regarding this species.

Journal compilation Ó CSIRO 2011 www.publish.csiro.au/journals/ajz Ecology of New Guinea carpet pythons Australian Journal of Zoology 237

Materials and methods were found close to the coast in tropical woodlands and were Distribution data for New Guinea carpet pythons were obtained not present further inland where rainforest vegetation was more from visiting villagers who collected snakes, and from dominant. Villagers in Kimaam, on the island of Yos Sudarso published records and specimens in museum registers (Museum (Pulau Dolak), claimed they had never seen carpet pythons on , Australian Museum Sydney, Queensland Museum, the island despite the occurrence of seemingly suitable habitat. fi Bernice P. Bishop Museum, American Museum of Natural Traders in Merauke con rmed this, claiming they had never History, Californian Academy of Sciences and Museum received carpet pythons from Yos Sudarso despite receiving Zoologicum Bogoriense). other woodland species, such as the Papuan taipan (Oxyuranus Villagers and wildlife traders known to collect carpet scutellatus canni). One carpet python specimen, kept in the pythons in the Indonesian province of Papua were visited from Museum Zoologicum Bogoriense, supposedly collected in 1920 7 December 2010 to 1 January 2011 and from 1 March 2011 to by Koey from Pionierbivak on the Mamberamo River, may have 15 March 2011. Data were gathered by conducting interviews been incorrectly labelled due to the low probability of such a with villagers and traders to obtain information about how the disjunct distribution (Fig. 1) and lack of suitable woodland trade worked, the number of carpet pythons collected, collection habitat on the Mamberamo River (D. Natusch, pers. obs.). locations, and trade history. In total, 174 carpet pythons in the possession of traders were measured to record: (1) snout-to-vent Harvest length (SVL) and (2) tail length (TL), to the nearest 0.5 cm; During our study we recorded 247 wild-caught carpet (3) head width (HW) and (4) head length (HL) to the nearest pythons collected for the pet trade in the Indonesian province of 0.1 mm, at the widest point and from the tip of the snout to the Papua. Most were collected opportunistically near human base of the skull, respectively; (5) mass, to the nearest 1 g using habitation, or while crossing tracks at night, and taken to traders Pesola spring scales, and (6) sex, by inserting a blunt probe into located in the major port town of Merauke. However, a few the cloacal bursae and recording the depth. Although some snakes were brought from villages situated much further away authors have had difficulty determining sex of carpet pythons (such as Okaba, 100 km). There was no collection bias towards due to recording intermediate probe depths (Barker and either sex (null hypothesis of 50% male: c2 = 0.67, d.f. = 1, Barker 1994), males and females in our study were easily P = 0.41, restricted to individuals captured and not hatched). distinguishable and sex was often corroborated by voluntary Pythons of all sizes were collected; however, most were hemipenal eversion in males. Reproductive maturity was juveniles recently hatched from gravid females. Villagers determined for a sample of snakes either by measuring females collected gravid females and allowed them to lay and brood their found brooding eggs or by dissecting individuals recently killed eggs until hatching. Both the mother and neonates were then sold by local people. Males were recorded to be mature if they had separately to traders. While at traders premises, carpet pythons enlarged turgid testes or convoluted efferent ducts. Females were housed in plastic boxes or cotton bags to await shipment to were mature if they possessed large ovarian follicles (>5 mm) or Jakarta. It was claimed that all carpet pythons observed in this were gravid (sensu Slip and Shine 1988b). Prey types captured study were destined for Indonesian breeding farms and then by carpet pythons were determined by examining stomach exported for the international pet trade. Fig. 2 depicts the number contents or faecal samples. of carpet pythons exported from Indonesia between the years Several females found with clutches were allowed to hatch 2000 and 2009. their eggs and the resulting hatchlings were measured. Simple linear regression was used to test for correlations between Body sizes and morphology maternal body size, clutch size and hatchling traits, while Female carpet pythons attained larger average SVLs than males analysis of variance (ANOVA) was used to test for sex differences (Student’s t-test, t = 4.25, P = 0.001) (Table 1); however, males in the above traits between clutches. Because morphological reach sexual maturity at smaller sizes than females so more traits in adult pythons change with increasing SVL we used small males were represented in the sample. To account for this, analysis of covariance (ANCOVA) with SVL as the covariate to only the largest 20% of individuals of each sex were compared compare morphological measurements between the sexes. and no significant difference in SVL with sex was found. Variables were log10-transformed where necessary to meet Similarly, sexual dimorphism was not apparent in TL, HL, HW assumptions of normality and homogeneity of variance. All or mass relative to SVL (all P > 0.05). The sizes of newly hatched statistical tests were conducted using Minitab 16 software. neonates are listed in Table 2. Finally, export data were obtained from CITES (2011) for Indonesian exports between the years 2000 and 2009. Reproduction Results Reproduction is highly seasonal, with eggs observed to hatch in either December or early January. This timing is congruent with Distribution the claims of villagers that carpet pythons were found to be In total, we obtained 281 records of New Guinea carpet gravid in September and brooding eggs in November pythons. Most of the records from Indonesia were based on each year. The smallest sexually mature female (found brooding interviews with villagers who collected snakes, whereas those eggs) measured 147 cm SVL, while the smallest mature male from Papua New Guinea came from museum specimens and (dissected) measured 92 cm SVL. Eight female carpet pythons published records (see McDowell 1975 and references therein). were found with clutches (Table 2) and there was no significant Interviews with traders indicated that, in Papua, carpet pythons difference in hatchling SVL, HW, HL, TL or mass between the 238 Australian Journal of Zoology D. J. D. Natusch and J. A. Lyons

Fig. 1. Distribution of carpet pythons in New Guinea, showing locality records (*), the record by Koey from the Mamberamo River in 1920 ($) and the approximate boundary between the provinces of Papua and West Papua (black line).

Table 1. Sample sizes and adult SVL and mass results for carpet pythons collected by wildlife traders in Papua, Indonesia Standard errors are shown in parentheses

Males Females (n = 30) (n = 17) SVL Mean 139.5 (6.4) 179.1 (4.6) Mean of largest 20% 192.1 (7.5) 202.3 (2.6) Number of individuals Range 92–222 147–210 Mass Mean 916.3 (138) 1608 (124) Mean of largest 20% 2042.5 (247) 2203 (98.2) – – Year Range 197 3189 759 2459

Fig. 2. Numbers of carpet pythons exported from Indonesia as wild- caught (white column) and farm-bred (grey column) between the years 2000 and 2009.

Table 2. Clutch sizes and hatchling traits from female carpet pythons found brooding eggs in Papua, Indonesia Standard errors are shown in parentheses

Female # Maternal Clutch No. of Mean hatchling Mean hatchling SVL size hatchlings SVL mass Female 1 192.5 18 18 45.5 (0.23) 32 (0.21) Female 2 184 16 16 45.3 (0.3) 35.3 (0.6) Female 3 147 8 8 42.5 (0.2) 29.8 (0.32) Female 4 210 17 8 37.9 (0.8) 24.3 (1.6) Female 5 201 27 27 43.4 (0.48) 25.5 (0.9) Female 6 200 20 20 42.8 (0.15) 26.9 (0.18) Female 7 184 13 –– – Female 8 164 12 –– – Ecology of New Guinea carpet pythons Australian Journal of Zoology 239 sexes in any of the clutches (all P > 0.05). Consequently, data on 2001). The lack of sexual dimorphism in other traits (such as male and female hatchlings were combined for subsequent head dimensions and tail length) contrasts the results of Pearson analysis. As illustrated in Fig. 3, clutch size was significantly et al.(2002) for a south-western Australian population, where correlated with maternal SVL (F1,7 = 10.3, P = 0.018) and despite female snakes were heavier and had larger heads (in terms of significant differences in both hatchling mean SVL (F5,96 = 27, both length and width) than similarly sized males. However, P < 0.001) and mean mass (F5,95 = 27.4, P < 0.001) among the results fit with those of Shine and Fitzgerald (1995), who did clutches, hatchling SVL, TL, HL, HW and mass showed no not record any sexual dimorphism in similar traits in an east correlation with either maternal SVL or mass (all P > 0.05). Australian population. Furthermore, these variables were not significantly correlated Carpet pythons from temperate and semiarid Australia may with maternal condition (based on residual scores of linear take advantage of high temperatures during summer months to regressions of SVL and mass (all P < 0.05)). assist egg incubation, with hatching occurring during March and April (Slip and Shine 1988b; Pearson et al. 2002; Corey Food habits 2007). New Guinea carpet pythons, however, are exclusively Stomach contents and scat samples revealed 17 prey items, tropical and may not be constrained to oviposit during months of which all were mammals. Three large snakes (measuring with high ambient temperatures (James and Shine 1985; Brown 140, 152 and 174 cm SVL) had eaten bandicoots (family and Shine 2006). Instead, hatching appears to coincide with the Peramelidae) and the smallest (57.5 cm SVL) had eaten a onset of the summer monsoon, which may confer benefits to Rattus sp. juveniles due to increased prey activity at this time (Fitch 1982), or promote water retention in eggs and thus hatchling size Discussion and vigor (Brown and Shine 2006). This reproductive timing is ’ Natural history also seen in carpet pythons from the Top End of Australia s (R. Shine, pers. comm. 2011), which The considerable divergence in body size and reproductive experiences a similar, seasonal wet/dry, climatic regime to biology between carpet python populations in Australia southern New Guinea (Prentice and Hope 2007). suggests that, given their disjunct distribution, New Guinea The clutch and hatchling sizes of New Guinea carpet carpet pythons may exhibit similarly divergent traits. Our pythons are similar to those of carpet pythons from other results show, however, that the ecology of carpet pythons from populations (Charles et al. 1985; Slip and Shine 1988b; Barker New Guinea closely approximates that found on the Australian and Barker 1994). Positive relationships between clutch size and east coast (Shine and Fitzgerald 1996; Fearn et al. 2001). The maternal size are common in snakes (Seigel and Ford 1987). maximum sizes of male and female New Guinea carpet pythons They are also known for this species (Slip and Shine 1988b) and were similar, and despite the average size of females being were observed inthe present study (Fig. 3). However, we found no greater than that of males, the largest male captured measured significant correlation between maternal size (either SVL or 2.6 m (total length) and the mass of this individual (3189 g) mass) and offspring traits, although sample sizes were small. suggests that males may grow significantly larger than females Madsen and Shine (1996) used pathway analysis to show that (Table 1). Body size dimorphism in Australian carpet pythons offspring size and shape in Australian water pythons (Liasis is related to male combat (Shine and Fitzgerald 1995; Pearson fuscus) were indirectly affected by a complex interaction et al. 2002); males are the larger sex in populations where between maternal size, clutch size and mass, but noted that much combat is present. It is currently unknown whether combat of the variance was attributable to other, unknown, factors. occurs in wild New Guinea carpet pythons, although the large Because clutch mass and female condition before oviposition size of males and the presence of combat between captive could not be recorded during the present study, the proximate specimens (Mutton and Julander 2011) suggest that it may. causes for differences in hatchling traits in New Guinea carpet Males were found to mature at SVLs 50 cm less than females, pythons remain poorly understood. but no females smaller than 147 cm SVL were available for All carpet pythons in this study preyed on mammals, as do dissection so maturation in females may occur at shorter carpet pythons in Australia (Slip and Shine 1988c; Shine and lengths, as in other populations (Shine and Slip 1990; Fearn et al. Slip 1990; Shine and Fitzgerald 1996; Fearn et al. 2001; Corey 2007), but reference material was not available to identify prey remains to species. Most other studies have recorded in 30 the diet, particularly in young individuals. Our small sample was 25 likely the reason reptiles were absent from the diet of carpet pythons in our study. 20 New Guinea carpet pythons were found not to inhabit 15 rainforests, preferring areas of tropical woodland near the coast. By contrast, carpet pythons from a population in north

Clutch size 10 Queensland occur in the rainforests of the Daintree and on the 5 Atherton Tableland (Freeman and Bruce 2007). However, during 0 extensive searches (>500 person-hours) of rainforest habitats 140 150 160 170 180 190 200 210 220 SVL (cm) in Cape York Peninsula, far northern Queensland (Natusch and Natusch 2011), carpet pythons were never encountered. They Fig. 3. Clutch size versus maternal SVL of New Guinea carpet pythons. were, however, observed in nearby woodlands (D. Natusch, 240 Australian Journal of Zoology D. J. D. Natusch and J. A. Lyons unpubl. data). The causal reasons behind habitat preferences in Acknowledgements this species remain unknown and require further study. We thank the Indonesian villagers and traders who allowed us access to pythons in their possession. Thank you to Ross Sadlier (Australian Museum), Wayne Longmore (Museum Victoria), Andrew Amey and Patrick Trade Couper (Queensland Museum) and Awal Riyanto (Museum Zoologicum We recorded 69% of the annual harvest quota for carpet Bogoriense) for providing us access to specimens in their care. Thank you pythons in Indonesia during the short visits made in this study. to R. Shine and two anonymous reviewers for their comments on an earlier Recently hatched juveniles comprised the majority of snakes draft of this manuscript. This work was carried out in accordance collected and hatching time was found to occur in late 2010 with University of New South Wales Animal Ethics protocol (Permit Number and early 2011. Thus, hatchlings may be claimed to be collected 10/90A). either in the current or forthcoming harvest year, making it difficult to determine whether annual quotas have been References exceeded. Nevertheless, evaluation of the CITES Trade Database Anon. (2010). Kuota eksportum buhan alam dan satwa liar yang termasuk Appendix CITES untuk periode tahun 2010. Keputusan Direktur Jenderal (CITES 2011) revealed that Indonesia exceeded the quota in Perlindungan Hutan Dan Konservasi Alam. 2004 (Fig. 2) and suggested that quotas have been exceeded in Anon. (2011). Kuota eksportum buhan alam dan satwa liar yang termasuk other years. Nevertheless, Nijman and Shepherd (2009) found Appendix CITES untuk periode tahun 2011. Keputusan Direktur Jenderal that many Indonesian breeding farms did not have the facilities Perlindungan Hutan Dan Konservasi Alam. to adequately breed reptiles and suggested that the cost of Barker, D. G., and Barker, T. M. (1994). ‘Pythons of the World: Volume 1. producing large numbers of individuals in captivity is far greater Australia.’ (Advanced Vivarium Systems Inc.: Escondido, CA.) than that of collecting similar numbers from the wild. Further, Brown, G. P., and Shine, R. (2006). 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