Geochelone Platynota) in Myanmar

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Herpetological Review, 2017, 48(3), 570–575. © 2017 by Society for the Study of Amphibians and Reptiles Back From the Brink: Ex-situ Conservation and Recovery of the Critically Endangered Burmese Star Tortoise (Geochelone platynota) in Myanmar

The Burmese Star Tortoise (Geochelone platynota) is a medi- (Behler 1997; Altherr and Freyer 2000) where G. platynota was um-sized tortoise (carapace length [CL] to ca. 300 mm) endemic sold as food and incorporated into traditional medicines (Platt to the dry zone of central Myanmar (Fig. 1; Platt et al. 2011b). et al. 2000). Demand from Chinese markets was soon eclipsed by Chronic over-harvesting of G. platynota, mainly for domestic illegal collecting for the high-end international pet trade, which consumption by rural Burmese (Blyth 1863; Theobald 1868), resulted in the near extinction of wild populations (Platt et al. coupled with habitat loss has resulted in widespread, albeit grad- 2011b). By the early 2000s viable populations could no longer be ual population declines throughout its range (Platt et al. 2011b). found and G. platynota was considered ecologically and func- Harvesting greatly intensified during the mid-1990s, driven by tionally extinct in the wild, even for those populations within the the burgeoning demand from wildlife markets in southern China national protected area network (Platt et al. 2011a, 2011b). Con- sequently, G. platynota was designated a member of “extinction row” along with other high-risk chelonian taxa (Buhlmann et al. STEVEN G. PLATT 2002), listed among the 25 most endangered chelonians in the Wildlife Conservation Society–Myanmar Program, Aye Yeik Mon world (Rhodin et al. 2011), and assessed as Critically Endangered 1st Street, Yadanamon Housing Ave., Yangon, Myanmar by the IUCN (IUCN 2014). Captive-breeding was recognized as KALYAR PLATT Turtle Survival Alliance–Myanmar Program, Aye Yeik Mon 1st Street, the only remaining option for preventing biological extinction of Yadanamon Housing Ave., Yangon, Myanmar G. platynota and ultimately restoring this species as a functional LAY LAY KHAING member of the dry zone ecosystem (Horne et al. 2012). Nature and Wildlife Conservation Division of Ministry of Environment, In 2004 the Nature and Wildlife Conservation Division Conservation and Forestry, Minzontaung Wildlife Sanctuary, (NWCD) of the Myanmar Ministry of Environment, Conserva- Natowgyi Township, Myanmar tion and Forestry, in collaboration with the Wildlife Conservation THIN THIN YU Society (WCS)/Turtle Survival Alliance (TSA) Myanmar Program Nature and Wildlife Conservation Division of Ministry of Environment, established three ex-situ assurance colonies (defined as demo- Conservation and Forestry, Shwe Settaw Wildlife Sanctuary, graphically and genetically viable captive breeding groups of im- Minbu, Myanmar periled taxa maintained as a hedge against the extinction of wild SHWE HTAY AUNG Nature and Wildlife Conservation Division of Ministry of Environment, Conservation and Forestry, Lawkanandar Wildlife Sanctuary, Bagan, Myanmar SAN SAN NEW Nature and Wildlife Conservation Division of Ministry of Environment, BY ELEANOR BRIGGS ELEANOR BY Conservation and Forestry, Minzontaung Wildlife Sanctuary,

Natowgyi Township, Myanmar PHOTO ME ME SOE KHIN MYO MYO TINT LWIN WIN KO KO SWANN HTET NAING AUNG Wildlife Conservation Society–Myanmar Program, Aye Yeik Mon 1st Street, Yadanamon Housing Ave., Yangon, Myanmar THOMAS R. RAINWATER* Tom Yawkey Wildlife Center & Belle W. Baruch Institute of Coastal Ecology and Forest Science, P.O. Box 596, Clemson University, Georgetown, South Carolina 29440, USA Fig. 1. Captive-bred Burmese Star Tortoise (Geochelone platynota) re- *Corresponding author; e-mail: [email protected] leased in scrub forest of Minzontaung Wildlife Sanctuary.

Herpetological Review 48(3), 2017 CONSERVATION 571 BY STEVEN PLATT STEVEN BY BY STEVEN PLATT STEVEN BY PHOTO PHOTO PHOTO PHOTO

Fig. 3. Tortoise eggs are incubated in-situ within the breeding enclo- Fig. 2. Food is placed on wooden trays for sanitary reasons and to sures. Temporary wooden markers denote the location of the buried disperse feeding activity within the breeding enclosure. Note shallow clutch, number of eggs, date of laying, and identification number of pool for drinking and soaking in middle of photograph. the nesting female.

populations; Buhlmann et al. 2002) of G. platynota at Lawkanan- placed in the assurance colonies when it became apparent these dar, Minzontaung and Shwe Settaw wildlife sanctuaries (LWS, populations could no longer be effectively protected in the face MWS, and SSWS, respectively) in Myanmar with the objectives of overwhelming poaching pressure. With the exception of wild of 1) ensuring the biological survival of G. platynota by establish- tortoises collected from MWS and SSWS, founder provenance ing a secure captive metapopulation and 2) producing sufficient cannot be determined with any degree of certainty. Additionally, numbers of offspring for eventual reintroduction into protected precisely enumerating the founders is challenging because in- natural habitat. We herein provide a brief history of these efforts, adequate records were maintained, tortoises were not individu- review husbandry practices, summarize progress towards attain- ally marked, stock was occasionally transferred among colonies, ing the objectives of the assurance colonies, and discuss the con- some tortoises were stolen, and others were released during a servation implications of this program. premature and ill-conceived reintroduction attempt (Platt et al. 2011a). All things considered, our best estimate is that the found- Site Description ing stock consisted of 175 tortoises of an approximately equal sex ratio. These numbers are in keeping with the recommendation The three wildlife sanctuaries (MWS, SSWS, and LWS) hosting of Horne et al. (2012) that chelonian assurance colonies should the assurance colonies are located within the central dry zone of consist of at least three groups of 50 individuals each (1:1 sex ra- Myanmar, a desert-like region formed by the rain shadow of the tio) maintained at separate locations in order to maximize ge- western mountains (Platt et al. 2011b). The dry zone is character- netic diversity and minimize the likelihood of catastrophic loss ized by a monsoonal climate (annual precipitation = 500–1000 through disease, accident, or natural disaster. mm) with a truncated wet season (mid-June through September) followed by a prolonged dry season (October to early June); most Housing rainfall occurs during August and September. High diurnal temperatures (to 43°C) are typical of the dry season with low noctur- Juvenile (> 2 years-old), subadult, and adult G. platynota are nal temperatures (to 4°C) occurring during January and February maintained in outdoor enclosures and exposed to ambient en- (FAO/UNDP 1982). Physiographic descriptions of SSWS (46,500 vironmental conditions throughout the year. Housing initially ha) and MWS (2260 ha) are provided elsewhere (Platt et al. 2001, consisted of makeshift enclosures of wood, chain-link fencing, 2003). LWS is a small (47 ha) urban protected area of planted and low concrete walls, but underwent significant redesign and forest on the outskirts of Bagan, an archaeological and cultural improvement in the wake of repeated thefts of tortoises during site of international importance. All three wildlife sanctuaries the early years of the project. Security considerations and hus- are located within the presumed historic geographic range of G. bandry requirements have driven renovation of the enclosures platynota (Platt et al. 2011b); both SSWS and MWS harbored wild and remain of paramount concern. The renovated enclosures populations of G. platynota until these were extirpated in late consist of a high (to 3 m) perimeter wall constructed of concrete 1990s (Platt et al. 2001) and early 2000s (Platt et al. 2003; Thanda blocks and heavy gauge fencing, topped with concertina wire, Swe 2004), respectively. sharpened bamboo (pungi) stakes, or electrically charged wires. Entry to each enclosure is through locked gates with round-the- Founding Stock clock security provided by sanctuary staff, who reside on-site. No tortoises have been stolen from the assurance colonies since The majority of founding stock for the three assurance colo- these security measures were implemented. nies included juvenile, subadult, and adult G. platynota, most Within each enclosure, low walls (ca. 80 cm high) divide the of which were confiscated from wildlife traffickers by NWCD facility into separate pens of variable size. Overhanging thatch enforcement agents. Lesser numbers of wild tortoises were col- shelters along the interior wall of each pen, and piles of dried lected from MWS and SSWS during the early to mid-2000s and vegetation, leaves, and palm fronds provide tortoises with shade

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and concealment. These refugia are extremely important because the availability of secure hiding places is known to reduce stress among captive tortoises and enhance breeding success (Ebenhack 2012). Moreover, piles of vegetation buffer against BY STEVEN PLATT PLATT STEVEN BY temperature extremes and create a humid microenvironment

which may be important for proper shell growth (Wyrwich et al. PHOTOS 2015). Each pen contains one or two shallow (maximum depth ca. 10 cm) concrete pools for drinking and soaking. The latter is a critical consideration during April and May when diurnal temperatures regularly exceed 40°C. Because tortoises frequently defecate while soaking, water is changed at least once daily. The ground is swept and feces are removed from pens twice daily; in the morning and again in late evening.

Diet and Feeding

Little is known concerning the diet of G. platynota in the wild, although limited field observations suggest it consists largely of grass and other foliage, with lesser amounts of fruits, inverte- brates, and on occasion, vertebrate carrion (Thanda Swe 2004; Platt et al. 2011b; Me Me Soe et al. 2016). The standard diet of G. platynota in the assurance colonies consists of water spin- ach (Ipomoea aquatica), various locally available grasses, carrot roots and tops, cucumber fruit, and Roselle (Malvaceae) foliage, augmented with banana leaves and stems, gourd foliage, prickly- pear cactus pads, and fruits of tomato, gourd, cantaloupe, pa- paya, and watermelon, depending on availability. Oxalate-rich foods are fed sparingly (e.g., figs and squash) or avoided (e.g., legumes and cabbage). Foods are washed to remove any lingering pesticide residues, and then chopped and thoroughly mixed before being offered to tortoises. Food is provided every afternoon (1200–1400 h) on wooden trays to reduce the likelihood of fecal contamination and Fig. 4. Neonates are kept in secure, rat-proof wooden boxes for parasite transmission. Several feeding trays are placed at widely three months (above) before being transferred to a nursery building scattered locations in each pen to disperse feeding activity and (below). Neonate boxes are shaded for most of the day. Note high ensure each tortoise has access to sufficient food (Fig. 2). Uneat- security fence (behind the boxes) topped with concertina to deter en food is removed from the pens and feeding trays are washed thieves (above). with dishwashing soap and sun-dried at the end of the day. Given the size of the captive population, individual prophylactic de- worming is impractical. Therefore, fenbendazole (Panacur®, hatched before 2010 and reared at the assurance colonies were Merck Animal Health, Madison, New Jersey, USA) is mixed with now laying eggs at the time of manuscript submission. food at a dosage of 50 mg/kg of body mass and administered an- Eggs are incubated in-situ under ambient environmental nually to tortoises > 2 years old. Although high doses of fenben- conditions at all three assurance colonies. Because enclosures dazole administered over longer periods can cause toxicosis in are shaded at LWS and more open at MWS and SSWS, there is tortoises (Neiffer et al. 2005), the dosage we administer follows concern over how this might affect incubation temperatures and recommended guidelines (Carpenter and Marion 2013). hence offspring sex ratio (pivotal temperature ca. 30°C; Kuchling et al. 2011). Although secondary sexual characteristics are not yet Reproduction and Hatchling Care evident in most cohorts, we noted a moderate male bias among older cohorts at LWS and a similar degree of female bias at MWS Breeding groups of approximately equal sex ratio are main- when assembling groups of subadult tortoises for release at MWS tained together in outdoor enclosures; courtship and mating and SSWS (Platt et al. 2014, 2015). Despite the apparent bias, occurs throughout the year (Thanda Swe 2004; Platt et al., un- adequate numbers of both sexes are available for reintroduction. publ. data). Egg-laying begins in late September and continues Should it be deemed necessary to manipulate sex ratios in the through mid-May. During this period, keepers monitor breeding future, eggs could be artificially incubated in solar-powered groups throughout the day, noting when females begin excavat- incubators (continuous electricity is unavailable at MWS and ing nests. Upon completion of laying, a wooden marker is insert- SSWS). ed at the nest and labeled with the identification number of the Hatching occurs from mid-May through mid-July and hatch- female, date, and clutch size (Fig. 3). Females reach sexual ma- ing rates typically range between 50–75%. The hard-packed sub- turity when 7–8 years-old and deposit one to four clutches each strate is moistened with water every morning during this period season; mean (± 1SE) clutch size and annual reproductive output to facilitate hatchling egress from nests. Hatchlings exit the nest are 4.4 ± 0.4 eggs (range = 1–11) and 9.9 ± 3.2 eggs/female/year unassisted and are collected soon after appearing aboveground. (range = 5–16 eggs), respectively (Thanda Swe 2004). Females Staff excavate each nest 24 hours after the last neonates emerge

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to low overnight temperatures. Because neonates often develop respiratory ailments at the onset of the wet season, a multivita- min containing vitamin B complex and vitamin C (Envervon C, United Pharma, Yangon, Myanmar) is administered to all individuals < 2 years-old during this period.

Current Captive Population

Since 2008 the number of hatchlings produced at the three assurance colonies has been increasing at an annual rate of 37% (Fig. 5; average rate of increase calculated from slope of the lin- earized relationship between hatchling production and year; Fig. 5. Combined annual production of Burmese Star Tortoise hatchlings at three assurance colonies in Myanmar (2008–2016). Bayliss et al. 1987). At the time of this writing (mid-October 2016) the total captive population of G. platynota in Myanmar consists of 7150 individuals (Table 1) with an additional > 1000 eggs cur- Table 1. Captive population of Burmese Star Tortoises held by three rently being incubated. The captive population is dominated by assurance colonies in Myanmar (October 2016). Assurance colonies: younger cohorts, reflecting the success of propagation efforts in Minzontaung Wildlife Sanctuary (MWS); Lawkanandar Wildlife the last 5–6 years. Captive facilities are expected to reach maxi- Sanctuary (LWS); Shwe Settaw Wildlife Sanctuary (SSWS). Three mum capacity within the next 2–3 years, although the reintro- hundred subadult tortoises reintroduced into MWS and SSWS are not included in this stock assessment. duction of head-started subadults into the wild (Platt et al. 2014, 2015) will no doubt extend this period. Age class Discussion Assurance ≤ 2 yrs Subadults Breeding Total colony adults Ex-situ conservation of G. platynota is proving extremely suc- MWS 1508 567 278 2353 cessful in Myanmar because husbandry is relatively straightfor- LWS 2891 1084 185 4160 ward, and the species adapts well to captivity and reproduces SSWS 450 143 44 637 readily in confinement. Our ex-situ conservation efforts have averted the near-certain biological extinction of G. platynota and Total 4849 1794 507 7150 its continued survival is no longer in doubt, one of the original objectives of the assurance colonies. The second objective—to produce offspring for eventual reintroduction—has likewise to ascertain that all hatchlings have exited the nest and deter- been met and large numbers of captive-bred tortoises are now mine the number of unhatched eggs. Neonates are separated available for release into protected habitats. Limited success to into groups of 50 and kept in wooden rearing boxes (122 cm long this end has already been achieved at MWS and reintroduction × 122 cm wide × 25 cm high with access via a hinged top) for three efforts are now underway at SSWS (Platt et al. 2014, 2015, 2016). months (Fig. 4). After three months, neonates are transferred to Most importantly, the ultimate, albeit long-term objective put a more spacious nursery building and maintained there until forth in the national conservation action plan (K. Platt et al. 2014) reaching three years of age. Nurseries are open buildings ex- of restoring viable populations of G. platynota in every protected posed to ambient conditions, containing low-walled (ca. 30 cm area within the central dry zone is now biologically attainable, high) enclosures housing groups of neonates (Fig. 4). Because although social and political challenges remain to be resolved. rats (Rattus sp.) represent a serious threat to neonates, nursery Geochelone platynota, together with the endemic and Critically buildings are sealed with small gauge wire-mesh and rat traps Endangered Eld’s Deer (Cervus eldii thamin) are considered are maintained along exterior walls. Shelters within each pen iconic focal species for preservation of dry zone landscapes (Re- provide neonates with shade and concealment, and sunken con- public of the Union of Myanmar 2011). crete pools are available for drinking and soaking. Owing to their To date there has been no genotyping of captive tortoises small body size, neonates are particularly vulnerable to dehydra- in Myanmar, largely due to the costs associated with screening tion. Therefore, during the late dry season when air temperatures such a large population. Nevertheless, genotyping of at least the exceed 33ºC neonates are manually gathered and placed in shal- reproductive adults is warranted to avoid potential inbreeding, low water-filled basins for about 1 hour every morning. maximize the effective population size, and maintain sufficient Neonate diet and feeding protocols are similar to those of levels of heterozygosity among the captive population. Although other age classes except that foods are finely chopped and laced a transition from herd-level to individual management is imprac- with supplementary calcium. Food is provided daily on wooden tical, achieving these objectives may entail restructuring breed- trays or banana leaves and uneaten material is removed in late ing groups to maximize the likelihood that genetically high-value afternoon. The pens are swept daily to remove feces and other tortoises produce offspring. To further diversify the genetic base, debris. Young tortoises have been found to be susceptible to it may be advantageous to incorporate additional tortoises into chilling when nocturnal temperatures drop below 21°C, and re- the assurance colonies from collections outside of Myanmar, peated chilling has been associated with respiratory signs such such as the Turtle Conservancy (Ojai, California) and Taipei Zoo as serous nasal discharge and dyspnea (K. Platt et al. 2014). The (Taipei, Taiwan). Of course, any importation of tortoises into use of heat lamps to maintain optimal temperatures is precluded the Myanmar assurance colonies must follow strict quarantine by the lack of a reliable electrical source so small tortoises are protocols to prevent the introduction of novel pathogens. Ex- instead covered with dry straw and leaf litter to reduce exposure situ programs in other countries are also important as bulwarks

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against the potential loss of assurance colonies in Myanmar due Deb Levinson and the interlibrary loan staff at Wildlife Conservation to disease outbreaks, natural disasters, or political upheaval and Society (New York) are thanked for obtaining literature, and Bonnie accompanying civil turmoil. Raphael and Lewis Medlock commented on an initial draft of this Our ex-situ conservation efforts for G. platynota in Myan- manuscript. This paper represents technical contribution number 6544 of the Clemson University Experimental Station. mar highlight several advantages of establishing captive breeding programs within the natural range of a species (Durrell and Literature Cited Mallison 1987; Derrickson and Snyder 1992). First, local venues have a microclimate and photoperiod already appropriate for Aiello, C. M., K. E. Nussear, A. D. Walde, T. C. Esque, P. G. Emblidge, native animals making investment in potentially expensive cli- P. Sah, S. Bansal, and P. J. Hudson. 2014. Disease dynamics during mate control equipment unnecessary. This is certainly the case wildlife translocations: disruptions to the host population and po- for G. platynota at the three assurance colonies in Myanmar tential consequences for transmission in desert tortoise contact where tortoises can be maintained year-round under ambient network. Anim. Conserv. 17:27–39. environmental conditions. Second, captives are more likely to be Altherr, S., and D. Freyer. 2000. Asian turtles are threatened by extinc- adapted to the disease environment within the range country. Of tion. Turtle and Tortoise Newsletter 1:7–11. course, the possibility of introducing diseases from captive into Bayliss, P. 1987. Survey methods and monitoring within crocodile wild populations must also be considered during any reintro- management programmes. In G. J. W. Webb, S. C. Manolis, and P. J. Whitehead (eds.), Wildlife Management: Crocodiles and Alligators, duction (Cunningham 1996; Aiello et al. 2014), although with G. pp. 157–175. Surrey Beatty & Sons Pty. Ltd., Sydney. platynota this risk has been considerably reduced through rigor- Behler, J. L. 1997. Troubled times for turtles. In J. V. Abbema (ed.), Pro- ous health screening of individuals selected for release (Raphael ceedings: Conservation, Restoration, and Management of Turtles et al. 2016). Third, the transfer of animals within the borders of and Tortoises—an International Conference, pp. 13–22. New York a single country can be accomplished without being hampered Turtle and Tortoise Society, New York. by cumbersome international trade restrictions. Conway (1995) Blyth, E. 1863. A collection of sundries from different parts of Burma. rightly cautions that husbandry and medical expertise might be J. Asiatic Soc. Bengal 32:78–90. lacking in less developed range states; however, our close collab- Buhlman, K. A., R. Hudson, and A. G. J. Rhodin. 2002. A global action oration with technical experts in the international zoo and vet- plan for conservation of tortoises and freshwater turtles: strategy erinary community has largely nullified this concern (Raphael et and funding prospectus 2002–2007. Conservation International and Chelonian Research Foundation, Washington, D.C. 30 pp. al. 2016). Furthermore, a continuing emphasis on training and Carpenter, J. W., and C. J. Marion. 2013. Exotic Animal Formulary. Else- capacity building by the WCS/TSA Myanmar Program (K. Platt vier Saunders, St. Louis, Missouri. et al. 2014) has produced a cadre of highly knowledgeable and Conway, W. 1995. Wild and zoo animal interactive management and experienced staff at each assurance colony in Myanmar. habitat conservation. Biodiver. Conserv. 4:573–594. In conclusion, ex-situ conservation efforts have secured the Cunningham, A. A. 1996. Disease risks of wildlife translocations. Con- biological future of G. platynota in Myanmar. Because stockpil- serv. Biol. 10:349–353. ing animals in captivity is of little conservation benefit, the ulti- Derrickson, S., and N. Snyder. 1992. Potential limitations of captive mate goal of any conservation breeding program should be the breeding in parrot conservation. In S. Bessinger and N. Snyder restoration of the target species as a functional member of its na- (eds.), New World Parrots in Crisis: Solutions from Conservation tive landscape (Tudge 1992). Given the high value G. platynota Biology, pp. 133–163. Smithsonian Institution Press, Washington, D.C. continues to command in both the illegal and legal wildlife trade, Durrell, L., and J. Mallison. 1987. Reintroduction as a political and increasing human population density in the dry zone of Myan- educational tool for conservation. Dodo 24:6–19. mar, and limited representation of this landscape in the national Ebenhack, A. 2012. Health care and rehabilitation of turtles and tor- protected area system, restoring viable wild populations poses a toises. Living Art Publishing, New York. 392 pp. daunting, but not insurmountable, challenge to conservationists. FAO/UNDP. 1982. Shwesettaw Wildlife Sanctuary: report on a recon- Despite the difficulties involved, progress towards realizing this naissance survey and evaluation, June 1982. FAO/UNDP Nature objective is nonetheless being made using offspring produced in Conservation and National Parks Project, BUR/80/006, Field Re- the assurance colonies in Myanmar (Platt et al. 2016). Finally, our port 9/82. FAO/UNDP, Rangoon. 19 pp. experience in Myanmar suggests the ex-situ conservation model Horne, B. D., C. M. Poole, and A. D. Walde. 2012. Conservation of Asian we outline here is broadly applicable to other developing coun- tortoises and freshwater turtles: setting priorities for the next ten years. Recommendations and conclusions from the workshop in tries struggling to recover imperiled tortoises. Singapore, February 21–12, 2011. Wildlife Conservation Society, Singapore, Ltd. 32 pp. Acknowledgments.—Numerous people contributed to the suc- IUCN. 2014. IUCN Redlist of Threatened Species. [www.iucnredlist. cess of this project since its inception in the early 2000s. We are es- org; accessed 6 March 2016]. pecially indebted to U Nyi Nyi Kyaw (Director General) and U Win Kuchling, G., E. Goode, and P. Praschag. 2011. Endoscopic imaging Naing Taw (Director) for their enduring commitment to Burmese of gonads, sex ratio and temperature dependent sex determina- Star Tortoise conservation. Turtle Survival Alliance is acknowledged tion in captive bred juvenile Burmese star tortoises Geochelone for recognizing the pivotal role of ex-situ conservation in the recov- platynota. Asian Herpetol. Res. 2:240–244. ery of the Burmese Star Tortoise and supporting the improvement of Me Me Soe, San San Nwe, S. G. Platt, and T. R. Rainwater. 2016. Geoche- captive facilities at MWS and LWS. Than Myint, Rick Hudson, Colin lone platynota (Burmese star tortoise): scavenging. Herpetol. Rev. Poole, Saw Htun, Bill Holmstrom, Lonnie McCaskall, Brian Horne, 47:444–445. Andrew and Angie Walde, Cris Hagen, Bonnie Raphael, Shane Boylan, Neiffer, D. L., D. Lydick, K. Burks, and D. Doherty. 2005. Hematologic Katie Rainwater, and Paul Gibbons are thanked for their contribu- and plasma biochemical changes associated with fenbendazole tions. Financial support for tortoise conservation in Myanmar was administration in Hermann’s tortoises (Testudo hermanni). J. Zoo generously provided by Andy Sabin and the Sabin Family Founda- Wildl. Med. 36:661–672. tion, Helmsley Charitable Trust, Panaphil Foundation, Turtle Conser- Platt, K., S. G. Platt, Lay Lay Khaing, Thin Thin Yu, San San Nwe, Win Ko vation Fund, Jeff and Mara Talpin, and Joan and Howard Oestreich. Ko, Khin Myo Myo, Kyaw Moe, Me Me Soe, Tint Lwin, N. Chansue, and

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