Distribution of the South American Chelid Turtle Phrynops Gibbus, With Observations OnHabitat and Reproduction

8/3/2019 Distribution of the South American Chelid Turtle Phrynops Gibbus, With Observations OnHabitat and Reproduction

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Herpetologists' League

Distribution of the South American Chelid Turtle Phrynops gibbus, with Observations onHabitat and ReproductionAuthor(s): Russell A. Mittermeier, Anders G. J. Rhodin, Federico Medem, Pekka Soini,Marinus S. Hoogmoed, Nelly Carrillo de EspinozaReviewed work(s):Source: Herpetologica, Vol. 34, No. 1 (Mar., 1978), pp. 94-100Published by: Herpetologists' LeagueStable URL: http://www.jstor.org/stable/3891617 .

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DISTRIBUTIONOF THE SOUTHAMERICANCHELIDTURTLEPHRYNOPSGIBBUS, WITH OBSERVATIONSONHABITATAND REPRODUCTIONRUSSELL A. MITTERMEIER, ANDERS G. J. RHODIN, FEDERICOMEDEM,

PEKKA SOINI, MARINUS S. HOOGMOED,AND NELLY CARRILLODE ESPINOZAABSTRACT: The South American chelid turtle Phrynops gibbus has previously beenreported from Trinidad, Surinam, Guyana, French Guiana, the Iquitos region of Peru,eastern Ecuador, northeasternVenezuela, the Departamento of Meta, the Intendencias ofCaqueta and Casanare and the Comisariaof Vaupes in Colombia, and the states of Para andMato Grosso in Brazil. New locality recordsreportedhere extend the range of P. gibbus intosouthern Venezuela, the Vichada of Colombia and the upper Amazon region of Colombiaand Brazil. Examination of the Mato Grosso specimen revealed that it is not P. gibbus.Clutch size in P. gibbus varies from 2 to 4 and eggs are laid in a variety of sites. Incubationperiod ranges from 178 to 200 days and hatchlings weigh 10 to 15 g and range in size from43 to 48 mm. Phrynops gibbus lives in a variety of aquatic habitats, most of them stagnant

or slowly flowing and muddy bottomed.FEW data are available on the SouthAmerican chelid turtle Phrynops gibbus(Schweigger,1812)1. Its rangeis at presentpoorlydefined,little is knownof its habitatpreferences and the only informationonreproductions providedby Medem(1973).In this paper, we review the publishedlocalitydata for P. gibbus, cite severalnew

localities which extend the range of thisturtle, provide a distributionmap, discussthe animals habitat in Peru, Colombia,Surinamand Trinidad,and add to the in-formationon reproduction.Distribution.-Wermuth and Mertens(1961) and Pritchard (1967) give therangeof P. gibbus as Trinidad, he Guianasand northand centralBrazil,but the litera-ture includes records from several othercountries as well. Phrynops gibbus hasbeen recorded from: Trinidad (Gray,1868; Boulenger, 1889; Mole and Urich,1894; Underwood, 1962; Kearney, 1972);Guyana (Gray, 1873; Boulenger, 1889;Pritchard, 1964); Surinam (Siebenrock,1905; Medem, 1973); French Guiana(Dumeril and Bibron, 1835; Fretey, 1975,

'Most earlier authors refer to this species as Meso-clemmys gibba (e.g., Wermuth and Mertens, 1961; Pritch-ard, 1967). Zangerl and Medem (1958) suggested thatboth Mesoclemmys and Batrachemys are at best subge-nerically distinct from Phrynops, and Medem (1973) re-ferred to P. gibbus as Phrynops (Mesoclemmys) gibbus.Bour (1973) elected to drop the subgeneric distinctionand referred to the animal as simply Phrynops gibbus andwe follow his decision here.

1977); Brazil (Schenkel, 1901; Siebenrock,1904); the Departamentoof Meta, the In-tendencias of Caqueta and Casanareandthe Comisaria of Vaupes in Colombia(Dunn, 1945; Niceforo Maria, 1952; Me-dem, 1958,1968, 1973); the Iquitos regionof Peru (Dixon and Soini, 1975, 1977);andeastern Ecuador (Orces, 1949). Donoso-Barros(1965) also mentionsit from north-eastern Venezuela and the savannas ofVenezuela, but does not give precise lo-calities.The Brazilianrecords for P. gibbus arefrom 2 localitieswidely separated romoneanotherand from the rest of the range ofthe species. Schenkel (1901) and Sieben-rock (1904) recorded t fromthe vicinity ofPara [now Belem], state of Para,and fromCuiaba, state of Mato Grosso. We haveexaminedthe 2 specimenson which theserecordsare based (NMW 11884:2,15762).The animal romBelemis clearlyP. gibbus.On the other hand, the one from Cuiaba(NMW 15762) is not. It representsa newspeciesof Phrynopswhichwill be describedin a laterpaper.Ruschi (1966) lists P. gibbus as a mem-ber of the fauna of the Brazilianstate ofEspiritoSanto. This is almostcertainlyanerror, probably based on another unde-scribed species resemblingP. gibbus. Un-derwood (1962) gives the range of P. gib-

HERPETOLOGICA 34:94-100. March 1978


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March 19781 HERPETOLOGICA 95bus as "Trinidad and the Guianas tosouthernBrazil." The term southernBrazilis misleadingand apparentlybased on thelocation of Cuiaba. We have no definiterecords of P. gibbus from southern Braziland do not believe that it occurs south ofthe Amazondrainage.The records from French Guiana arebased on Fretey (1975, 1977) and on aspecimen described as Platemys miliusiiby Dumeril and Bibron (1835). Wermuthand Mertens (1961) speculated, and Bour(1973) proposed that Platemys miliusii issynonymouswith Phrynopsgibbus, thoughFretey (1975, 1977) considersthem to bedistinct. Recent workby R. Bour (personalcommunication) shows that P. miliusii isin fact synonymouswith P. gibbus. Wehave also obtained 2 other specimens ofP. gibbus from French Guiana, both fromthe vicinity of Remire (AGJR 111, L-341,collectedby R. RenauFerrer,1975).The literature record of P. gibbus fromnortheastern Venezuela (Donoso-Barros,1965) was probablybased on 2 specimensin the American Museum of Natural His-tory from Caripito, state of Monagas. Inaddition, the animalhas been collected byP. Trebbau at San Carlos de Rio Negro,state of Amazonas, n the extremesouthernpart of the country(ZMA 15147,collectionof Hoogmoed, and P.C.H. Pritchard,per-sonal communication) and at La Barra,TerritorioDelta Amacura (H. Ollos, per-sonalcommunication).Phrynopsgibbus also occursin the upperAmazonregion of Braziland Colombia. Aspecimenwas collected by R. Foote in 1975in the QuebradaUrumutu, a tributaryofQuebradaTacana,3 km northeastof Leti-cia, Amazonas,Colombia(live in collectionof Medem, IRF 766). In addition, a speci-men in Harvard'sMuseum of ComparativeZoology (MCZ 57130) bears the locality"Leticia, Colombia, bought from TarponZoo"[an animal mportingcompany n Tar-pon Springs,Florida,USA] andMittermeierhas seen several P. gibbus in the compoundsof animal dealers in Leticia. Leticia issituated on the Brazilian-Colombian-

Peruvian border and dealers obtain ani-malsfrom all 3 countries.Finally, we have specimens that extendthe range of P. gibbus into the Comisariaof Vichada in Colombia. This record isbased on 2 animals obtained by Medemin 1968 (IRF 225-6). They were collectedby a Guahibo Indian and lack preciselocality data, but come from the Rio Mucoarea.All known ocalitiesfromwhichP. gibbushas been recorded are indicated in Fig. 1.Included in this figure are literaturerec-ords and previouslyunpublisheddata frommuseum specimens or specimens that wehave collected.Reproduction.-Data on reproduction nP. gibbus arebased on 5 egg clutches fromColombia, Peru and Surinam (Table 1).Eggs are white, elongate, hard-shelled,fairlyuniform n size, and weigh from22.5to 28.0 g. Clutch size varies from 2 to 4eggs. Eggs are apparently aid in a varietyof sites. Two nests have been observedinthe wild. A nest from the Cano Negro,Colombia,was found in the vegetable gar-den of a local villager. It was situated8 mfrom the bank of a small stream and re-ceived some sunlight, but was in the shademost of the day. The soil was dark andmoist and the nest hole 10 cm deep.The last egg of the 4 in this clutch wasonly about 1 cm below the surface. A"nest" n Surinamwas found between theroots of a tree by the side of a smallstream.The 2 eggs were on the surface of theground and were not covered. Data arealso availableon 2 clutcheslaid by captiveP. gibbus. A female from the Rio Vaupeslaid 3 eggs in a 2-cm excavationunderrot-ten bananaleaves in an outdoorenclosure.Another female from the Rio Momon inPeru laid 3 eggs in sand in her enclosure.One of the 3 eggs from the latter clutchwas opened after 153 days and found tocontain a well-formedhatchlingthat livedfor 3 days (Dixon and Soini, 1977). Thesecond hatched spontaneously after 200days. The incubationperiodsof the 4 ani-mals which hatched spontaneouslyranged


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96 HERPETOLOGICA [Vol. 34, No. 1

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University;MPM =Milwaukee Public Museum; TCWC = Texas Cooperative Wildlife Col-lection, Texas A&M University;UK = Universityof Kansas; UMMZ = University of Mich-igan Museum of Zoology; USNM - United States National Museum, Washington, D.C.;ACJR =personal collection of Rhodin; BMNH = British Museum of Natural History,London; ENPQ =Escuela Nacional Politecnica, Quito, Ecuador; FT = personal collection ofFretey; ILS = Instituto La Salle, Bogota, Colombia; IRF = Instituto Roberto Franco, Villa-vicencio, Colombia; MHNP = Museum d'Histoire Naturelle, Paris; MHNJP = Museo deHistoria Natural "Javier Prado," Lima, Peru; MZUSP = Museu de Zoologia, Universidadede Sao Paulo, Brazil; NMW = NaturhistorischesMuseum Wien, Austria; RMNH = Rijks-museum van Natuurlijke Historie, Leiden, Netherlands;SMF - Senckenberg Museum, Frank-furt, West Germany; ZMA = Zoologisch Museum Amsterdam, Netherlands.ECUADOR 1. Santa Cecilia, Napo [0?05'N, 76050'W] (UK 105438, 109857, 148409-10)2. Loreto, Lo~ver Rio Suno, Rio Napo, Napo [0?040'S, 77022'WV]( ENPQ 4612, 4614,MCZ 68859)3. Rio Conamba,Pastaza[2?05'S, 76?25'W] ( MCZ68858)PERU 4. a. Iquitos, Loreto [3?42'S, 73?15'W] (Dixon and Soini, 1977, MHNJP 17, MPM8580, MCZ uncatalogued 9 collectedy Soini,1972)b. Rio Itaya [3?45'S, 73?17'W] (Dixon and Soini, 1977, TCWC 41806)c. Rio Momon, Rio Nanay [3?037'S, 730 17'WV] (live specimen obtained by Soini, 1973)5. Maipuco, Rio Maranon, Loreto [4?47'S, 75?10'W] (Dixon and Soini, 1977, AGJRL-296--live specimen obtained by Soini, 1972)6. Yarinacocha,Rio Ucayali, Loreto [8?15'S, 74035'W]1 FMNH 45668-71)COLOMBIA 7. a. Villavicencio, Meta [40?10'N, 73?37'W] (ILS 79, USNM 108581)b. Acacias [4?00'N, 73?45'W] (Niceforo Maria, 1952, ILS 76)c. Rio Ocoa and Aguas Claras [4?08'N, 73?35'W] ( Medem, 1958, FMNH 73422-30,SMF 51477)


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March 1978] HERPETOLOGICA 97d. Cano Catatumbo[4008'N, 73040'W] (MCZ 127406)e. Cano Negro [4008'N, 73037'W] (Medem, 1973)8. Sabanade San Juande Arama,Meta [3033'N, 73050'W] (Medem, 1958)9. Quebrada Honda, upper Rio Guejar,Meta [3015'N, 73?45'W] (Medem, 1958)10. PuertoLopez, Meta [4006'N, 72?56'W] (Medem, 1958)

11. Orocue,upper Rio Meta,Casanare[4?48'N, 71020'W] (Medem, 1958)12. Rio Muco, Vichada [4030'N, 70?50'W] (IRF 225-6)13. a. Florencia, Caqueta [1?35'N, 75?37'W] (Niceforo Maria, 1952, ILS 74, 77, 125, 129-31, 144)b. Rio Hacha [1?40'N, 75?37'W3 (ILS 78, UK 124941)c. Rio Ortequaza [1?35'N, 75032'W] (ILS 75)d. QuebradaLa Perdiz [1?33'N, 7537'W] (ILS 145)e. Ceilan [1035'N, 75037'W] (ILS 146)f. Pueblonuevo [1035'N, 75037'W] (ILS 147)14. Rio Vaupes,Vaupes [1?30'N, 72?10'W] (Medem, 1973)15. Puerto Uva, upper Rio Apaporis,Vaupes [0030'N, 71?30'W] (IRF 254)16. a. Leticia, Amazonas[4?11'S, 69057'WI (MCZ 57130, IRF 765, FT 206)b. QuebradaUrumutu,QuebradaTacana W40WS,9056'W] (IRF 766)VENEZUELA 17. San Carlos de Rio Negro, Amazonas [1?55'N, 67004'W] (ZMA 15147-live specimen in collection of Hoogmoed, P. Pritchard, personalcommunication)18. Caripito,Monagas [10?08'N, 63?05'W] (AMNH 65540, 65555)19. La Barra,Delta Amacura [8045'N, 61?30'W] (H. Ollos, personal communication)TRINIDAD 20. a. SangreGrandeC10?35'N,61?07'W: (MCZ 66028)b. CumacaRiver, Arima [10?36'N, 61015'W] (MCZ 100901)c. Near Mt. Tamana [10t28'N, 61?12iW] (BMNH 1947.3.4.18-type of Hydraspisgordoni Gray, 1868)d. Trinidad (BMNH 1917.9.27.3)GUYANA 21. a. Dunoon, DemeraraRiver, Demerara [6?25'N, 58?20'W] (UMMZ S46672-4,53979)b. Kartabo [6023'N, 58042'W] (AMNH 64721)22. Demerara Falls [5020'N, 58032'W] (BMNH 1946.1.22.86-type of Hydraspis bicolor

Gray, 1873)23. Parabam, near Marudi River, Kuyuwini River, Essequibo [2008'N, 59?06'W] (AMNH61521)SURINAM 24. New River (Coeroeni River), 750 feet [= 228.6 ml = Kutari headwaters,Nickerie [2?15'N, 56050'WI (BMNH 1939.1.1.107)25. a. Paramaribo,Suriname[5050'N, 55010'W] (NMW 18791)b. CarolinaCreek,Para [5?25'N, 55010'W] (Medem, 1973)c. Between Powakka and Carolina, Suriname [5025 N, 55005'W] (RMNH field numberMSH 1975-301 live specimen in collectionof Hoogmoed)d. Berlijn,Para [5024'N, 55013'W] (RMNH field number MSH 1968-456 live specimenin collection of Hoogmoed)26. a. Kaboeri Creek, Nickerie [5015'N, 57015'WI (RMNH field number DGR 1971-H160)b. AvanaveroRoad, Kilometre246 west of Saronbridge [4050'N, 57?10'W] (DOR speci-men, not collected, identifiedby Hoogmoed)27. Poite Ede, upper Commewijne River, Commewijne [50 19N, 54037'WI (P. Pritchard,personalcommunication)28. 83 km from SaramaccaBridge in direction of Bitagron on Zanderij-BitagronRoad [5015'N, 55050'W] (AGJR L-544-live specimen collected by Mittermeier,1977)FRENCH GUIANA 29. a. Remire, Guyane [4052'N, 52018'W] (AGJR 111, L-341-speci-mens collected by Ramon Renau Ferrer, 1975, MHNP 1974-1005, FT 210)b. Cayenne [4?55'N, 52020'W] (MHNP 8755-type of Platemys miliusii Dumeril andBibron, 1835, FT 146, 152)BRAZIL 30. a. Corcovado,near Breves, Para [1040'S, 50030'W] (MZUSP 2684-5)b. MarajoIsland (MHNP 5339)31. Belem, Para [130'S, 48?30'W] (Schlegel, 1901, Siebenrock, 1904, NMW 11884:2, SMF42011)


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98 HERPETOLOGICA [Vol. 34, No. 1TABLE 1.-Data on reproductionin Phrynops gibbus.

Egg size ( mm )Eggs in - Date Date IncubationEgg source clutch (n) Length Width Egg wt (g) laid hatched time196 mm 9 from 4 44.5 30.0 25.0 ... ... ...Florencia, Caqueta, 44.0 30.0 24.0Colombia;dissected 42.0 30.0 22.522 July 1952 42.0 29.5 23.0(Niceforo Maria,1952)183mm, 700-g 9 3 44.5 29.5 24.5 23 Nov 1971 19 May 1972 178from Rio Vaupes,Colombia; 44.0 30.5 25.0 23 Nov 1971 6 Jun 1972 184eggs laid in captivity 44.0 30.0 24.2 23 Nov 1971 16 Jun 1972 194(Medem, 1973)9 from Rio Momon, Loreto, 3 44.0 32.0 ... 3 Aug 1973 openedafter153 daysPeru; eggs laid in captivity 43.5 31.5 ... 3 Aug 1973 18 Feb 1974 200(Dixon and Soini, 1977) not measured .. unknown ...Nest found in tributary 4 44.5 31.5 27.5of Cano Negro, 20 km SE 44.0 32.0 28.0Villavicencio,Colombia, 43.0 32.0 26.723 Nov. 1972 (Medem, 1973) 42.5 31.5 26.4Nest found near Carolina 2 41.0 30.0Creek,Para,Surinam, not measured12 Dec 1971 (Medem, 1973)

from 178 to 200 days. Hatchling size andweight are given in Table 2. Very small,triangular, sharply-pointed egg caruncleswere present on the 3 hatchlings from theVaupes female's clutch. They detached at22, 24 and 29 days after hatching. Althoughthe 5 hatchlings observed had access towater, they all preferred to hide in leaflitter immediately after hatching.Color in the 5 hatchlings observed showed

little variation: carapace uniformly black,with ventral surfaces of marginals orange inthe Colombian specimens, yellow in theSurinam and Peruvian specimens; suturesbetween ventral marginals black; bridgeprimarily black with yellow or orange onposterior and anterior edges; plastron uni-formly black in the Colombian and Surinamspecimens; Peruvian specimens with 2 pairsof yellow spots on lateral edges of anteriorTABLE 2.-Size and weight of Phrynops gibbus hatchlings. Measurements in millimeters; weights ingrams.

Midline lengthcarapace plastron carapace shell heightwidth (max) (max) weight

Clutch of Rio Vaupes 9Hatchling of 19 May 1972 48.0 38.0 36.0 17.5 15.6Hatchling of 6 June 1972 48.0 38.0 35.0 17.0 12.2Hatchling of 16 June 1972 48.0 37.5 35.5 17.5 13.5

Clutch of Rio Momon 9Hatchlingof 18 Feb 1974 48.0 37.0 35.0 ... .Surinamhatchling 43.0 34.0 31.0 15.5 10.0


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March 1978] HERPETOLOGICA 99plastral lobe, 1 pair on gulars, 1 on hu-merals;posterolateral dges of femoralsandedges of anals also yellow in Peruvianspecimens;mandible and dorsaland lateralsurfaces of head black with lighter grayreticulations;ympanumblack;maxilla ightgray with black spotting; feet and tail uni-formly black;iris black with a very narrowyellowish white rim aroundthe free inneredge; chin barbels white to yellowishwhite and 2 in number. It is unclearwhether differences among Peruvian, Co-lombian and Surinamspecimens representgeographicor individualvariation.Habitat.-In Peru and Surinam,P. gib-bus is found mainlyin smallforeststreams,ponds and small black water rivers, andusually staysunder closed canopy. Mostofthe streams and rivers it inhabits in Peruhave muddy bottoms. In Surinam it alsooccurs in standing bodies of water with athick layer of mud on the bottom, such asroadside ditches in more or less cultivatedregions. In Peru, P. gibbus is not knownto be common anywhere in the country(even though sympatricPhrynops nasutusis rather common in some parts of theAmazoniandrainageregion [e.g., the RioNanay near Iquitos]). In most Colombianlocalities,P. gibbuslives in streamsrunningover a muddy bed, and in muddy lagoonsand ponds in both rain forest and galleryforest in the plains. In the Vichada area,it lives mainly in the morichales (palmgroves) of Mauritia lexuosa. Within thesegroves, the habitat consists of ponds sha-dowed by the palms. The ponds have amuddy bed and may drypartiallyor totallyduring the dry season. Captive P. gibbussometimesestivate in dry weather and it isprobable that free-living animals also en-gage in such behavior. In Trinidad, thisturtle lives in clear, flowing streams withpartly muddy, partly gravel-coveredbeds(Kearney, 1972; F. Medem, personal ob-servation).Phrynopsgibbus is primarilynocturnal.In captivity, t may occasionallybaskin theearly morning or late afternoon. Captivespecimens eat fish, meat, worms, insects,

crustaceans, newborn mice, commercial dogfood, and sometimes take plant matter(e.g., leaves of Colocacia sp.).Size.-The largest recorded specimen ofP. gibbus is a female from Ecuador (ENPQ4612). It measures 232.5 mm in midlinecarapace length, 193 mm in midline plas-tron length and 67 mm in maximum shellheight.

Acknowledgments.-We thank T. Blohm, R.Bour, W. Duellman, H. Ollos, G. Orces, P. Pritch-ard, P. Trebbau and P. Vanzolini for providingunpublished locality data and other information,and J. Eiselt, A. Kluge, F. Tiedemann, E. E.Williams and R. Zweifel for allowing us to exam-ine specimens in their collections. Peter Pritchardalso read and commented on the manuscript.Mittermeier's studies in South America werepartly funded by a National Science FoundationGraduateFellowship and partly by the New YorkZoological Society. Hoogmoed's field work inSurinam in 1968 and 1974-1975 was funded bythe Netherlands Foundation for the Advancementof Tropical Research (WOTRO Grants W 956-2and W 87-78).LITERATURE CITED

BOULENGER,. A. 1889. Catalogue of the che-lonians, rhynchocephaliansand crocodiles in theBritish Museum (Natural History). BritishMuseum, London.BouR, R. 1973. Contribution a la connaissancede Phrynops nasutus (Schweigger: 1812) etPhrynops tuberculatus (Luederwaldt: 1926).Description d'une nouvelle sous-espece origin-aire du Paraguay, Phrynops tuberculatus van-derhaegei. (Testudinata-Pleurodira-Chelidae).Bull. Soc. Zool. France 98:175-190.DIXON, J. R., AND P. SOINI. 1975. The reptilesof the upper Amazon basin, Iquitos region,Peru. I. Lizards and amphisbaenians. Mlilwau-kee Public Museum, Contrib. Biol. Geol. (4):1-58. 1977. The reptiles of the upper Ama-zon basin, Iquitos region, Peru. II. Crocodil-ians, turtles and snakes. Milwaukee Public Mu-seum, Contrib. Biol. Geol. (12): 1-91.DONOSo-BARROS,R. 1965. Distribucion de lastortugas en Sudamerica. Publ. Ocas. Mus. Nac.Hist. Nat. (Santiago) (8): 1-14.DUNlERIL, A. M. C., AND G. BIBRON. 1835.Erpetologie generale. Paris. 2:431-433.DUNN, E. R. 1945. Los generos de anfibios yreptiles de Colombia. IV. Cuarta y ultimaparte: reptiles, ordenes Testudineos y Croco-dilineos. Caldasia3:307-335.FRETEY, J. 1975. Les cheloniens de Guyanefrancaise. Bull. Soc. Zoo]. France 100:674-675.


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100 HERPETOLOGICA [Vol. 34, No. 1. 1977. Les cheloniens de Guyane fran-caise. I.-Etude preliminaire. Thesis, Univ.Paris.

GRAY, J. E. 1868. Notice of Hydraspis gordoni,a new species from Trinidad, living in thegardens of the Society. Proc. Zool. Soc. London1868:536-564.1873. Observationson chelonians, withdescriptions of new genera and species. Ann.Mag. Nat. Hist. London (4)11:289-308.KEARNEY, P. 1972. Nocturtles of Trinidad. Int.Turtle Tortoise Soc. J. 6(2):10-11, 32-33.MEDEM, F. 1958. Informe sobre reptiles colom-bianos (II). El conocimiento actual sobre ladistribucion geografica de las Testudinata enColombia. Bol. Mus. Cien. Nat. Caracas 2-3(1-4):13-45.. 1968. El desarrollo de la herpetologiaen Colombia. Rev. Acad. Colomb. Cien. Ex.Fis. Nat. 13(50):149-199.. 1973. Beitrage zur Kenntnis uber dieFortpflanzung der Buckel-Schildkrote,Phrynops(Mesoclemmys) gibbus. Salamandra9:91-98.MOLE, R. R., AND F. W. ULRICH. 1894. A pre-liminary list of the reptiles and batrachians ofthe island of Trinidad. J. Trinidad Field Nat.Club 2:77-90.NICEFORO MARIA, H. 1952. Testudineos del sub-order Pleurodira en el museo de La Salle. Bol.Inst. La Salle (Bogota) 39:1-8.ORCES V., G. 1949. Los Testudinata ecuatori-

anos que se conservan en las colecciones deQuito, Ecuador. Bol. Inf. Cient. Nac. 3(20-21):13-22.PRITCHARD, P. C. H. 1964. Turtles of BritishGuiana. J. Brit. Guiana Mus. 39:19-45.. 1967. Living turtles of the world.T.F.H. Publications,Jersey City, New Jersey.RuSCHI, A. 1966. Lista dos repteis do Estadodo Espirito Santo. Bol. Mus. Biol. Prof. Mello-Leitao (Zool.) 26A:1-6.SCHENKEL, E. 1901. Achter Nachtrag zumKatalog der herpetologischen Sammlung des

Basler Museums. Verhandl. Naturforsch. Ges.Basel 13:142-199.SCHWEIGGER,A. F. 1812. Prodromus Mono-graphiae Cheloniorum. Konigsberg. Arch.Naturwiss. Math. 1:271-368, 406-458.SIEBENROCK, . 1904. Schildkroten von Brasil-ien. Denkschr. Kaiserl. Akad. Wiss. (Math.-Naturwiss. KI.) 76:1-28.1905. Chelonologische Notizen. Zool.Anz. 28(12) :460-468.UNDERWOOD,G. 1962. Reptiles of the easternCaribbean.Carib. Affairs (N.S.) 1:1-192.WERMUTH, H., AND R. MERTENS. 1961. Schild-kroten, Krokodile, Bruckenechsen. GustavFischerVerlag, Jena.ZANGERL,R., AND F. MEDEM. 1958. A newspecies of chelid turtle, Phrynops (Batra-chemys) dahli, from Colombia. Bull. Mus.Comp. Zool. 119:375-390.

Received: 26 April 1976Accepted: 31 October 1977(RAM) Museum of Comparative Zoology,Harvard University, Cambridge, Massachu-setts 02138, USA and Conservation Fellow,New York Zoological Society, Bronx, NY10460, USA; (AGJR) Department of Sur-gery, Yale-New Haven Medical Center,New Haven, Connecticut 06504, USA;(FM) Instituto Roberto Franco, Apartadoaereo 22-61, Villavicencio, Meta, Colombia;(PS) Proyecto Primates, P.O. Box 621,Iquitos, Loreto, Peru; (MSH) Rijksmuseumvan Natuurlijke Historie, Leiden, Nether-lands; (NCDE) Museo de Historia Natu-ral "Javier Prado," Universidad NacionalMayor de San Marcos, Av. Arenales 1256,Apartado1109,Lima,Peru

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Distribution of the South American Chelid Turtle Phrynops Gibbus, With Observations OnHabitat and Reproduction