Herpetology Notes, volume 7: 685-688 (2014) (published online on 17 November 2014)

New data on the naturally-occurring maximum sizes attained by Ploughshare ( yniphora)

Angelo Mandimbihasina1 and Andrea Currylow2,*

As part of larger studies in January, April, and May extreme demand in the illicit trade and incited poaching 2013, we recorded morphometrics on 127 Ploughshare from the wild, making it truly the “golden ”. Tortoises (Astrochelys yniphora) (Vaillant 1885) and As in many , the Ploughshare Tortoise found previous reports of morphometrics to be lacking exhibits sexual dimorphism, and sexes can be the breadth of size this species can achieve. The distinguished via secondary sexual characteristics. As Ploughshare Tortoise is the largest extant tortoise in compared to females, male characters include overall ; however, it has previously been described longer carapaces, elongated gular scute, concave as reaching morphometrics that fall up to 36% short of plastron, longer tail, and longer anal fork. In contrast, our findings. Using data presented herein, we propose females have shorter upturned gular scute, flat plastron, to update and add to the current species description, and wider anal notch which allows egg passage. Adult offering a new standard metric for size. Ploughshare Tortoise straight carapace length (SCL) The Ploughshare Tortoise (known locally as is described as being 250–486 mm (Juvik et al., 1997; “Angonoka”) is endemic to Baly Bay National Park Smith et al., 1999; Pedrono, and Markwell, 2001). In in northwest Madagascar. The species is known as the 2001, Pedrono and Markwell reported the maximum rarest and most endangered tortoise in the world (Curl size and weight known among both male and female et al., 1985; Curl, 1986). It is listed in Appendix 1 of wild Ploughshare Tortoises in the Baly Bay region, the Convention on International Trade in Endangered Madagascar. Species, classified as Critically Endangered on the As part of larger studies in January, April, and May IUCN’s Red List, and in the top 10 of The World’s 25+ 2013, we sampled a captive population of Ploughshare Most Endangered Tortoises and Freshwater , Tortoises and conducted line-distance surveys in the designated at “Extremely High Risk of ” wild in Baly Bay, Madagascar. Due to the sensitivity (Leuteritz, and Pedrono, 2008; Rhodin et al., 2011; of locality information, we decline to state the exact CITES, 2014; IUCN, 2014). The namesake of A. location of the wild beyond the region of yniphora, the distinctive, long gular projection (plough; Baly Bay National Park. For each , we recorded an extension of the plastron) and its rarity (fewer than morphometrics (SCL; Straight Carapace Width [SCW]; 400-600 wild adults remain; Kiester et al., 2013) make Straight Plastron Length [SPL]; Straight Plastron Width it a prized animal for collection. It is further set apart [SPW]; Curved Carapace Length [CCL]; and weight). by its unique golden colour, all of which has created We recorded data from a total of 127 Ploughshare Tortoises of all age classes (Figure 1). Of those, 106 were wild individuals found during surveys in the natural range. For the 98 animals in the adult age class (individuals over 250 mm SCL) from both captive and wild populations, we report summary morphometrics 1 Durrell Wildlife Conservation Trust, Lot IIY49 J (Table 1). In our data, SCLs differ significantly between Ampasanimalo, BP 8511 Antananarivo 101, Madagascar; each of the three classes (sub-adult, male, and female; e-mail: [email protected] F = 159.5069, p< 0.0001). 2 Department of Biological Sciences, University of Southern 2,124 California, 3616 Trousdale Parkway, AHF 107, Los Angeles, Concordant with accounts of this species, we found California 90089, USA males to be significantly larger (SCL; t = 11.08, p< * Corresponding author; e-mail: [email protected] 0.0001) and heavier (t = 5.844, p< 0.0001) than females. 686 Angelo Mandimbihasina & Andrea Currylow

(2001) report the largest known female (#1030) as one which is smaller than their own reported maximum SCL for females of the species (see Table 1). Accordingly, four of the adult females we recorded were larger in SCL than #1030, and seven were as heavy or heavier. The largest female we found (#3034) is in captivity in Madagascar and is part of a captive breeding program. She measured 416 mm SCL, 258 mm carapace height, and weighed 12.8 kg. During the filed surveys, we recorded two male and four female Ploughshares that exceeded any previously reported maximum SCL for the individual sexes. The largest individual, a male (#1102) we named “Rapeto” after a giant in a traditional Malagasy legend, measured 519 mm SCL, 288 mm carapace height, and easily topped out our 20-kg Pesola scale (Fig. 2A). The largest wild Figure 1. Weights and Straight Carapace Lengths of female we encountered (#1297), we named “Rasoabe” Ploughshare Tortoises surveyed during January-April 2013 in meaning “the great, good lady” which measured 411 northwest Madagascar. mm SCL, 270 mm carapace height, and weighed 15.5 kg (Fig. 2B). These two animals, along with several others, are now being monitored via radiotelemetry. Although we have reported the metric here, we found Many of our animals had their gular projections cut off weight to be an unreliable metric for larger individuals. by local people who believe that a long gular scute As can be seen in Figure 1, the fitted curve begins to prevents an animal from eating. Therefore, only our dissociate from reliably predicting the size of the larger reported ranges in SPL (not the mean SPL) are accurate adult animals. We found the largest animals were not for a proper species account. Still, two of the males we always the heaviest and adult animals often would measured were larger (SCL) than previously reported as vary by kilograms if they voided their bladders during “largest known” Ploughshare of 486 mm (Pedrono, and handling. We recorded several adult weight differences Markwell, 2001). We also did relocate the Pedrono and of more than 2.0 kg before and after voids during the Markwell largest known male (#1017) which measured same sampling visit. Similar weight variation was seen 489 mm SCL, growing only 3 mm in more than a decade in the same wild individuals across seasons in subsequent since their report. Curiously, Pedrono and Markwell surveys (Currylow unpubl. data). Therefore, we also

Table 1. Morphometrics (Straight Carapace Length [SCL]; Straight Carapace Width [SCW]; Straight Plastron Length [SPL]; Straight Plastron Width [SPW]; Curved Carapace Length [CCL]) of adult Ploughshare Tortoises in the present study compared to previous reports/species descriptions.

Previous Studies Previous Studies Males Present Study Females Present Study Morpho-metric Mean/Max % diff. Mean/Max % diff. (range; n = 41) (range; n = 57) (range; n = 32) (range; n = 40)

Weight (kg) 12.4 (8.0–20.0+) 10.3 (7.2–18.9) + 20.4 9.4 (6.0–15.5) 8.8 (5.5–12.0) + 6.8

SCL (mm) 435 (368–519) 415 (361–486) + 4.8 367 (320–416) 370 (307–426) - 0.8

SCW (mm) 291 (250–386) 317 + 21.8 267 (226–312) 280 + 11.4

Height (mm) 253 (222–288) 269 + 7.1 231 (200–319) 235 + 35.7

SPL (mm) 425 (347–508) 462 + 10.0 349 (307–389) 379 + 2.6

SPW (mm) 268 (229–311) N/A - 245 (206–300) N/A -

CCL (mm) 632 (552–742) N/A - 572 (554–604) N/A -  Naturally-occurring maximum sizes attained by Ploughshare Tortoises 687

Figure 2. Authors with (A) “Rapeto”, the largest known Ploughshare Tortoise, and (B) “Rasoabe” the largest wild female Ploughshare Tortoise on record. Photos by authors.

report other shell measurements, such as carapace height, for this species (Table 1). We found that using R2(SA): 0.99 the height metric (as opposed to weight) along with R2 (F): 0.85 SCL was both informative and resulted in high linear- R2 (M): 0.83 prediction values for the distinct groups (Fig. 3). The relationship is strongest for the sub-adult age classes (r2 = 0.99), and is robust for both females (r2 = 0.85) and males (r2 = 0.83). While the sizes of the largest Ploughshares recorded during the current study are impressive, they were not unpredicted. In 1989, Reid, Durrell and Rakotobearison suggested that the largest Ploughshare Tortoise could reach a half-meter in SCL and 25 kg in weight. Curl et al. (1985) reported that local villagers estimated the Figure 3. Size metrics (carapace height and straight carapace species to reach 560 mm, but noted that this extreme length) for each of the three Ploughshare groups (sub-adult was larger than any known individual. Pedrono and [SA], female [F], and male[M]) with fitted regression lines. Markwell stated that ploughshares can be bigger than Shaded areas around the Fit Lines delineate confidence of fit. the maximum sizes they reported in 2001. However, R-square values for each fit line are in the upper left. this study now shows that males and females of this species can commonly achieve even larger and more impressive sizes than those previously reported in the literature. We propose to update and add to the current species description, offering carapace height and SCL as the new standard metrics for size comparisons. 688 Angelo Mandimbihasina & Andrea Currylow

Acknowledgements. Work was conducted under the Ministère Juvik, J.O., Kiester, A.R., Reid, D., Coblentz, B., Hoffman, J. de l’Environnement et des Forêts #09/13/MEF/SG/DGF/DCB. (1997): The conservation biology of the angonoka, SAP/SCBSE. We thank , Turtle Conservation yniphora, in northwestern Madagascar: Progress Report. Fund, Durrell Wildlife Conservation Trust, and Madagascar Conservation, Restoration, and Management of Tortoises Biodiversity Partnership for funding and support of our field and Turtles−An International Conference Turtle and Tortoise research. Permission and help was supplied by Madagascar Society. 345−350 pp. National Parks, the Ministry of Environment, Ecology and Kiester, A.R., Mandimbihasina, A.R., Lewis, R.E., Goode, E.V., Forests, as well as local field guides in Baly Bay. Ernest Bekarany Juvik, J.O., Young, R., Blanck, T. (2013): Conservation of and Tsanta Rakotonanahary assisted us with the captive breeding the Angonoka (Ploughshare Tortoise), Astrochelys yniphora. population. Lance Woolaver helped with logistical support and he Chelonian Research Monographs 6: 162−170. and Ross Kiester provided critical reviews of the earlier versions Leuteritz, T., Pedrono, M. (2008): Astrochelys yniphora. IUCN of this manuscript. 2010 The IUCN Red List of Threatened Species. 9 pp. Pedrono, M., Markwell, T. (2001): Maximum size and mass of References the ploughshare tortoise, Geochelone yniphora. Chelonian Conservation and Biology 4: 190. CITES (2014): Appendices I, II and III. Convention on International Reid, D., Durrell, L., Rakotobearison, G. (1989): The captive Trade in Endangered Species of Wild Fauna and Flora. 46 pp. breeding project for the angonoka Geochelone yniphora in Curl, D.A. (1986): The rarest tortoise on earth. Oryx 20: 35−39. Madagascar. Dodo 26: 34−48. Curl, D.A., Scoones, I.C., Guy, M.K., Rakotoarisoa, G. (1985): Rhodin, A.G.J., Walde, A.D., Horne, B.D., Dijk, P.P.v., Blanck, The Madagascar tortoise Geochelone yniphora: current status T., Hudson, R. (2011): Turtles in Trouble: The world's 25+ and distribution. Biological Conservation 34: 35−54. most endagered tortoises and freshwater turtles−2011. Turtle IUCN (2014): The IUCN Red List of Threatened Species, Conservation Coalition. 54 pp. International Union for Conservation of Nature and Natural Smith, L.L., Reid, D., Robert, B., Joby, M., Clément, S. (1999): Resources. Available at: http://www.iucnredlist.org. Last Status and distribution of the (Geochelone accessed on 4 August 2014. yniphora) of western Madagascar. Biological Conservation 91: 23−33.

Accepted by Diogo Provete; Managing Editor: Wouter Beukema