and Came SS ata of the Black Wood Turtle in Costa Rica Photos by Manuel Merchan The genus Rhinoclemmys comprises eight turtle species distributed exclu- sively in the Neotropical region (PRITCHARD and ‘TREBBAU, 1984, MERCHAN and FIDALGO, 1999), which are also the only New World members of the family Bataguridae. There is still much controversy over whether to consider Bataguridae a family in itself, or a subfamily (Batagurinae) of the family Emydidae. The sepa- zation into two families — Batagur- idae and Emydidae — has been accepted by the majority of authors in recent years (SALVADOR, 1998; MERCHAN and MARTINEZ, 1999), for which reason we will use the classification in this article. The species that make up the genus Rhinoclemmys are R. annulata, R. areolata, R. funerea, R.nasuta, R. melanosterna, R. pulcherrima, R, punctularia, and R. rubida. As our objective isto discuss R. funerea, following is a description of the ehar- acteristics that distinguish it from the other species of the genus according to ERNST and BARBOUR (1989) and our own observations, Distinguishing characteristics Rhinoclemmys funerea (Cope 1876), is distinguished from R. annulata, R. pulcherrima, R. rubida, and R. areolata by its fully webbed feet. The other four species have only small or no interdigital mem- branes. R. melanosterna and R. ‘xX Dstbution map ofthe back wood tut Tninoctermys ture costa Res punctularia\haveelear dorsal head ‘markings that are never present. im R. funerea. Finally, the somewhat rounded carapace of R. funeréa is distinguishable from, the quite flat- tened carapace of R. masta. “Also, GENERAL IDENTIFYING CHARACTERISTICS — Aquatic turtle measuring up to 325 mm in length. — Smooth black carapace with no markings, rings, or ridges. — Small head (relative to the carapace); dorsal coloration, black; ventral coloration, yellow with black spots R. funerea has black spots on its chin and throat, whereas R. nasuta has continuous lines. ‘As a result of enzyme study using electrophoresis, carried out on each species of the genus Rhinoclemmys (SITES et al., 1981), it was con- cluded that R. funerea is very close- ly related to R. areolata and R. punctularia, Only three of the eight species of Rhinoclemmys inhabit Costa Rica (SAVAGE and VILLA, 1986; ACUNA-MESEN, 1993a): the black wood turtle, R. funerea; the brown wood turtle, R. annulata; and the painted wood turtle, R. pulcherrima. Description Rhinoclemmys funerea, is the largest and most aquatic species of the genus (MERCHAN, 1992). Juveniles have rounded carapaces; adults have carapaces that are more elongated, The carapaces of males ate more elongated than the cara- paces of females, which tend to remain more rounded. Seen from the side, the black wood turtle is, rather flattened (clearly an adap- tation to its aquatic lifestyle), reach- ing its maximum height at the sec~ ond and third vertebral scutes. ‘The carapace margins are somewhat ser- ‘rated in neonates, but are complete- ly smooth in older juveniles and adults. Rhinoclemmys funerea has five vertebral scutes — the largest being the second and third, and sometimesalso the fourth. The first three ver- tebral scutes are nearly square in shape; the fourth and fifth vertebral scutes have such narrow posterior ‘margins that they appear almost tri- angular. R. funerea has a small nuchal scute between the first pair of costal scutes, ahead of the first vertebral scute, Of the four pairs of costal scutes, the first are the largest — adjacent to the first and second vertebrals and the first five marginals on either side. The costal scutes are rectangular, except the fourth. which tend to be square. The 11 pairs of marginal scutes are all of similar shape and size. The supra- caudal scute is divided, and is never in contact with the fourth costal. The carapaces of adult R. funerea are practically smooth, the scutes do not present rings or any other pattern of relief. The plastron of R. funerea is large in proportion to the carapace. There is no plastral hinge, and the bridge to the carapace is bony. The black wood turtle has six pairs of plastral scutes. ‘The gular scutes extend toward the front beyond the margin of the carapace. The humeral scutes are small, as are the femoral scutes (although the latter are slightly larger). The largest plastral scutes are the pectorals and abdominals. ‘The anal scutes of R. funerea are small, and do not pro ject toward the rear. ‘We have found numerous cases of black wood turtles with additional scutes —large ones in the carapace and small ones in the plastron. In practically all of these cases, the additional plastral scutes are situat- ed at the sutures between the pec- toral scutes and abdominal scutes, and between the abdominal scutes and the femoral scutes. The plastral formula (relationship of scutes in order from longest to shortest) of the black wood turtle, according to ACUNA-MESEN (1993a), is as follows: abdominal- pectoral-gular-anal-humeral. The Values we obtained in a study of 35 adult specimens of R. funerea are presented in Table 1. ‘erizal view of the specimen inthe previous photo. Smal addtional scutes are vible between the pars of pectoral and abdominal 3B‘Akt ahinoclemmys unereain te wate. Ths isthe mest aquatic secs ofthe serus 34 REPTILIA ‘Table 1. Comparison of plastral scute suture lengths (mm) between male and female black ‘wood turles, Rhinoclemmys funeres. N Mean Min, Max SD. 17 338 245 392 36 18 347 2S 412 47 17 141 100 22 39 1B 14189 269 45 OS 51 554 432 610 51 @8 S514 735 64 615 541 838 80 432 330 493 41 18 473 402 539 35 LSAN-M 17 334 260 400 46 LSAN-H 18 332 260 400 36 N= Number fsecimens. SD = Standard devi ation, GSL = Gular suture length. HSL = umeral suture length. PSL = Pectoral suture length. ABSL = Abdominal suture length, FSL = Femoral suture length, ANSL. = Anal suture length, M~ Male. F= Female, The head of the black wood turtle is medium-sized and can be fully retracted into the carapace. The nostrils project upward, and. the ocular orbits are situated antero- laterally. ‘The skin of the head is smooth with no distinguishable scales. The hind limbs are fairly large'with webbed feet. The dorsal coloration of this species is black, or less commonly dark brown. The plastron is black with yellow spots of variable size. The sutures between the plastral scutes are often yellow, as is the cen- tral area of the plastron (normally including parts of the pectorals, abdominals, and femorals). The dorsal color of the head is black. The chin and throat are yellow, pro- fusely mottled with round black spots; the number and size of these spots decrease from the chin tothe throat, almost disappearing near the plastron. The insides of the four legs are completely yellow. The tail is black with a yellow underside. ‘The axillary and inguinal scutes may be slightly pinkish or orangish. Theskin of the neck and the dorsal parts of the extremities generally become ‘grayish as the animal ages. Sexual dimorphism in R. funerea includes the already-mentioned dif- ference in carapace shape — rounder in females, and more elon- gated in males, Also, tail length is significantly greater in males, as is the distance between the base of the tail and the cloaca — an adaptation facilitating the alignment of male and female cloacas for copulation. In a study of 30 specimens, ACUNA-MESEN (1993b) showed that the average carapace str length is 270.6 mm (minimum 2 mm; maximum 284 mm). This study also indicated sexual dimorphism in carapace length, carapace width, curved length, and curved width. Ina study of 35 adult specimens, we found that females reach larger sizes and greater weights in propor: tion to the sizes of their carapaces (MORA and MERCHAN, in prep.), as seen in Table 2. Table 2. Comparison of weights (g) and carapace lengths (mm) between male and {female black wood turtles. N Mean 1733294 18 36639 7 214 18 2828 7 3204 18 2988 = Number of specimens W = Weight. SL = ‘Straigh length of carapace. CL = Curved lengt ofcaraptce. M=Mate = Female Geographic variation No subspecies of Rhinoclemmys funerea have been described (PRITCHARD, 1979). In our study, we found no significant mor- phological variation among speci- mens of black wood turtles other than the differences attributable to sexual dimorphism. ‘This homo- "Adu female Rneclemys fares atte Sinn Botner 700 Distribution and habitat The black wood turtle is distrib- uted in the Caribbean lowlands of Central America from the Coco River along the border between Honduras and Nicaragua to the area of the Panama Canal (ERNST and BARBOUR, 1989). According to PRITCHARD (1979), the species has also been reported on the Panamanian island of Bastimentos. ACUNA-MESEN (1993a) describes the distribution of R. funerea in Costa Rica as extending through eneity seems to support the nonex- pp Back wood tries, hinclemmysfunerea, prefer the quet waters of mers and kes — Inthe phat, the istence of subspecies. "Ho Fi, nthe National Wife Refuge et Cano Negro (nore Cosa cs) 35‘Adu female Rhinoclemms fuerea. Notice the yellow venta coloration with Black spats on the ‘rot and legs the Caribbean lowlands to as far as the eastern slopes of the Cordillera de Guanacaste (along the Pizote and Las Haciendas rivers) in the north- west. It is found as far south as the Estrella and Telire rivers near the Panamanian border. No specimens: of this species have been found on the Pacific side of Costa Rica ‘As already mentioned, R, funerea is the most aquati¢ species of the pri- ‘marily terrestrial genus Rhinoclemmys (MERCHAN and FIDALGO, 1999), It is found in the quiet waters of rivers and lakes. ‘According to. ACUNASMESEN (1993a), R. funerea prefers areas of vegetation predominated by sedges (eg, Cyperus virens), mimosa (€-8., Mimosa pigra), and common water hyacinth (Eichornia crassipes). "The ‘same author also reports that in these areas R. funerea is sympatric with other aquatic turtles: the Ecuadorian snapping turtle, Chelydra serpentina acutirostristhe natrow-bridged mud turtle, Kinosternon angustipons, the \white-Hipped thud turtle, Kinostemon leucostomum postinguinales the scorpion mud turtle, Kinosternon scorpioides, and’ the Nicaraguan slider, Trachemys}scriptayemolli. The black wood turtleycan be seen basking on fogs in the canals of the Parque Nacional de Tortuguero, in the northwestern part offCosta Rica (MORA, pers. obs). Rhinoclemmys funerea has been recorded atélevations of up to 1,049 méters. in Costa Rica’s Valle Central, witit water temperatures of 17.3-19.7°@(ACUNA-MESEN et al, 1983). Alimentation Adult black wood turtles are exclusively herbivorous, and feed mainly on guavas, Psidium guajaba; papayas, Carica papaya; bananas, Musa sapientum, and oranges, 36 REPTILIA Citrus sinensis (ACUNA-MESEN, ). At the Simén Bolivar Zoo José, we saw captive black "wood turtles being fed a dict of let- tuce, Lactuca sativa; tomato, Lycopersicon esculentum; carrot, Daucus carota; pepino, Solanum ‘muricatum, and watermelon, ferullus Janatus. However, as they Wersjhoused together with carniv- S turtles (ie., Kinosternon jes and Chelydra serpentine) that were being fed chicken and beef, we cannot be sure that the herbivorous wood turtles did not also consume meat. Rhinoclemmys funefea has) not been observed feeding on carrion. Reproduction i ‘According to MERCHAN (1992) the jreproductive period of Rhinoclemmiys funerea is the months of April, May, and June. In i petiod extends to March and August, (ACUNA-MESEN, 1993a), Courtship has been described as it appears here, from the original notesitaken by the authors: “The'male approaches the female with his neck outstretched, and walksaround her a. few times. hen, the male comes up behind the female and\tries to mount her for copulation. Failing, he repeats the procedure and tries to mount again, During this entire time, the male fhoves his head up and down ina slight nodding motion. This lasts fot a total of only a few minutes. ‘Once the male has successfully mounted, he lowers his head towards the head of the female, who at the same time retracts her head. The male tries to bite the female’s neck or head. He keeps the female held dowin with all four legs. Now they are copulating, The anterior half of the male rocks'as if hhe were losing his balance, but he keeps his posterior half held tightly to the female. “A second male approaches and hhartasses the copulating male, who continues copulating and trying to reach the head of the female with his. mouth, Four minutes havepassed. The second male, also try- ing to reach the head of the female, is bitten repeatedly by the copulat- ing male. The female walks along the bottom of the pond a few centi- meters, but the male remains securely mounted on her back. “Prior to the separation of the mating pair, two other males approach. ‘The copulating male tries to bite both of them, in addi- tion to the female. Finally, the cop- ulating male releases the female. Five and a half minutes have passed. Immediately following cop- ulating, the male’s tail appears noticeably swollen at the base. “Now a third male tries to copu- late with the same female; she responds by biting and displaying her open mouth to new male advances. The male gives up the attempt. Then two males try to bite each other. One of them succeeds in biting the other firmly on the neck. Biting is not confined to the head and neck area; attacks on the extremities and carapace margins are also observed. Before the described copulation, another took place in which a milky substance (possibly semen) was observed in the water near the cloa- cas of the copulating pair. During the reproductive period, 1-4 clutches are laid, with an aver- age of 3 eggs (1-7) per clutch (ACUNA-MESEN, 1993a; ERNST and BARBOUR, 1989). Because of the relatively large egg size (characteristic of the entire genus) and the limited amount of space inside the body of the female, eluteh size is smaller in this species than in others; on the other hand, the hatching offspring are much larger, reducing their susceptibility to numerous small predators. "According to ACUNA-MESEN (1993a), black wood turtle eggs are elongated, white, and measure about 68 x 35 mm. MERCHAN (1992) describes them as measuring 70 x 35, mm, and ERNST and BARBOUR (1989) refer to a clutch of eggs mea- suring 76 x 39 mm. The incubation period is 98-104 days, after which hatch offspring measuring 55mm. We measured 16 captive-hatched specimens at 3 months of age, and found an aver~ age straight length of 65.1 mm (min- imum 59.5 mm; maximum 74.1 mm). Wild black wood turtle hatchlings are prey to crested caracaras, Polyborus plancus, double-toothed kites, Harpagus bidentatus, buz- zards, Buteo spp.; coyotes, Canis Jatrans, spectacled caimans, Caiman erocodylus, and American croco- diles, Crocodylus acutus. These last also attack juvenile and even adult black wood turtles. Large croco- diles have also been observed prey- ing on another aquatic turtle species, the savannah sideneck tur- tle, Podocnemis vosli, in Apure, Venezuela (MERCHAN, pers. 0bs.). Current status of populations Consumption by humans repre- sents relatively little threat to Rhinoclemmys funerea compared to what is happening with other Central American aquatic turtle species. ACUNA-MESEN (1993a) indicates that indigenous peoples Iadira de San Sebestin, inthe National Wie Refuge at Cato Neg (nartnem Costa Rea). During the dy season, beck wood tutes (Rhinoctemmys fnerea) a7 nd Newaguan sides (achenysscrota emo) congegee et these water bodiesalong the northern Atlantic coast of Costa Rica consume only adult black wood turtles, captured during only one week of the year, and that no more than 10 specimens are taken per community. With current available data, we can be sure that Costa Rican popu- lations of the black wood turtle are stable, and still abundant within the natural range of the species. In Costa Rica, this range includes sev- eral protected areas known for their wetlands, such as the Tortuguero National Park, and the National Wildlife Refuge at Caio Negro. Acknowledgments ‘The authors wish to thank the fol- lowing people and institutions for their help: the Agencia Espaftola de Cooperacién Internacional (8.E.C.). for making it possible for Manuel Merchdn to visit Costa Rica; the Sim6n Bolivar Zoo and the Zoo ‘Ave, where measurements were taken and behavioral observations were made; Rafael A. Acufa- Mesén, who gave us access to his library; and Ingrid Rodriguez, Marta Coll, Ana Ferrer, and Marco A. Mora, who collaborated in the field work. Bibliography ACUNA-MESEN, R.A. 19938, Las torte Poste Rica. Universidad gas continentates de {8 Costa Rica, San José, Costa Rica. ACUNA-MESEN, R. A., 1993. Andlisis rmorfo-estadistco de Ta tortugayneara do Costa Rica Rhinoclemmys funerea Cope, 1876 (Testudines: Emydidae). Repertorio Genifco 1(3)19-28 ACUNA-MESENeRUBG A, CASTAING, and F. FLORES, 1983 Aspestos seolgios e la distribuciomde ls tortuens tenrestres semiacuaticas en el Valle Central de Costa Rica. Rev. Biol. Trop 31(2)s81-102 ERNST, C, H. and R. W. BARBOUR, 1989, Tiles of the world. Smithsonian las. Press, Washington D.C. MERCHAN, M1952. Fl mareiloso mundo de las tortugas. Fa. Antigvaria S.A. Madrid MERCHAN, Mesnd A. M. FIDALGO, 1999, “Tortugas de América del Sur! una aproximacidn & sus patrones de distribucion ‘zoogeosraticas (1) Aquamar 1926-2, MERCHAN, M. and A. MARTINEZ. SILVESTRE, 1999, Tortugas de Espana. Biolovia. patologia y conservacion de las esneciesibcricas, baleares y canarias. Ed, Antiqvaria S.A, Madrid. PRITCHARD, P. C. H., 1979. Encyclo pedia of Turtles FH. New Jersey PRITCHARD, P.C.H. and P. TREBBAU 1984, Turtles of Venezuela. Soc. Study Amph. Rept, New York SALVADOR. A. (Ed Fauna bérica, 10. Museo Nacional de Gencias Naturales CS.LC, Madrid SAVAGE, J. M. and I. VILLA, 1986, Herpetofauns de Costa Rica, Soc. Study ‘mph. Rept, Oto. SITES, I. W.,. F. GREENBAUM, and J W.BICKHAM, 1981. Biochemical sytemat ies of Neotropical turtles of the G Rhinoclemmys (Emididae: Batagurinac). Herpetologica 31 198. Reptile, Manuel Merchén end José_M. Mora Ualiersidad de Costa Rica; Escuela Biologia y Museo de Zoologi Costa Rica Permanent Ariststeles 3,2” By address: Manuel Merchén: (027 Madi; Spain nslatel from Spanish by Steven Winchell Global Gecko Association The GGA is the international organization dedicated to the needs of all people interested in. geckos. Members receive the twice yearly, full. color journal “Gekko” plus “Chit-Chat”, the GGA Members Newsletter four times a year. If you want information on geckos— JOIN THE GGA TODAY! Aainial membership is $32 United States of America) $34 Canada/Mexico, $36 overseas. Fees afe in U.S. dollars. | (VISA and MasterCard accepted—include expiry date) Send payment or inquitis to: GGA clo Matthew Henry email: membership@gekkota.com PO Box 703 ‘Website: http:inwww.gekkota.com Havikinsuile, GA 31036. USA, 38 REPTILIA