The turtle and tortoise fauna of the central Chaco of Paraguay

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HABITAT OBSERVATIONS

RADIATARADIATARADIATARADIATARADIATA 10 (3), 2001

Thomas & Sabine Vinke

The turtle and tortoise faunaof the central Chaco of Paraguay

Card 1. Road map of theParaguayan Chaco: There areno asphalted roads except forthe communication roadbetween the capital Asunciónand Mcal. Estigarribia.

The Gran Chaco undoubtedly belongs tothe most impressive areas of South America.It reaches from the Rio Paraguay in North-Argentina and Paraguay over Bolivia to theAndes. The area has an expanse of 800,000km2. The precipitation steadily decreases fromthe east to the west. This is also indicated bythe vegetation. Palm savannas with marshlanddominate the landscape in the east whereascactus and dry forest with thorn bushes, whichshed their foliage, characterize the centraland western Chaco. Large partsof this area are still unexploitedand offer wild animals safepossibilities of withdrawal.Unfortunately man is beginningto destroy this paradise (VINKE

& VINKE 2000a). But it is stillpossible to find a wildlife thatleaves an unforgettable impres-sion.

IntroductionFrom January 1999 until

February 2001 we visited theMennonite colony Fernheimfour times. It is situated inNorthwest-Paraguay. One of the

journeys’ goals was the observation of thecentral Chacos chelonians in their naturalhabitat. We succeeded in finding the fiveknown species.

Filadelfia was the starting point of allexcursions. From that point we went northalong the national park ”Defensores delChaco” to ”Agua Dulce”. In this area westopped several times for a couple of daysand intensively observed the wildlife. Wemainly did so in the national park near Cerro



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Ill. 1. The Chaco appearsin luxuriant green duringthe rainy season.

Ill. 2. The dry forest is shed-ding its leaves at the end of

the dry winter.

Ill. 3. A frequent finding in the Chaco:Kinosternon scorpioides scorpioides, theScorpion Mud Turtle.

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Ill. 4. The distinct jaw is used bothfor eating apple snails and fordefence.

Ill. 5. Typical habitat of Kinoster-non scorpioides scorpioides nearMariscal José Felix Estigarribia.

Ill. 6. The colouring of the plastron variesfrom almost yellow to completely black.

Ill. 7. Kinosternon scorpioides scorpioides witha very dark plastron.



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Ill. 8. The throat barbles at the lower jaw arecharacteristic for this species.

Ill. 9. Kinosternon scorpioides scorpioides canfrequently be found in ditches along dirtroads.

León, the highest elevation (365 m) of theGran Chaco. Another rewarding area ofinvestigation is situated near a deserted placecalled ”KM 145” which is named after aformer railway line. From there we headed 30km east for Porto Casado near Rio Paraguayto the edge of the palm savannah. The pointof our journey situated furthest in the westwas near ”La Patria” approximately 200 kmnorthwest of Filadelfia. In the south we onlyjust reached the Mennonite colony Neuland.

We will refer to the various places wherewe found turtles and tortoises in the chapterswhere we have a closer look at the species.All turtles were measured and photographed.

There are many speculations especially aboutthe turtle fauna of the Paraguayan Chaco andthere are hardly any written records (MÜLLER

& HELLMICH 1936, NORMAN 1994, FRITZ & PAULER

1992a, FRITZ & PAULER 1992b, FRITZ & PAULER

1999, VINKE & VINKE 2000b).In order to investigate the turtle fauna of

Paraguay more intensively we provided theStaatliches Museum für Tierkunde in Dres-den with several specimens that we collectedduring a journey we made with JAKOB &MARIA UNGER. These animals had been killedeither in road accidents or by predators.

ClimateBetween latitude 25 and 35 degrees south

the zone of high-pressure areas as basis fordry forests and deserts is located in general.In South America this region is not as straightas in other continents. The trade winds fromthe east support the existence of subtropicalpartly evergreen rain forests. That is whyEast-Paraguay clearly differs from the GranChaco where aridness is dominating accor-ding to the latitude (SEIBERT 1996). Theaverage annual precipitation in this area is758 mm. The rain is falling mainly betweenOctober and April. From May to August it iscoolest (average daily temperature: 19.8 °Cto 22.8 °C). These numbers rapidly increasein September and cause a period of extremeheat in summer (average daily temperature:28.4 °C to 29.7 °C). The highest dailytemperature then frequently rises above40 °C. From March onwards the temperaturefalls again (MÜLLER, 1996).

These facts seem relatively regular accor-ding to a climate atlas. But if daily recordsare available (records of the radio stationFiladelfia ZP30) one will soon find out thatthe weather is often characterized by extre-mes especially in winter. In the course of 24hours the temperature often rises from 0 °C to40 °C and can fall just as rapidly again. Inspring we experienced such a sudden drop intemperature after a hot day (38 °C). At 6 a. m.there still were 32 °C. Then the wind turnedsouth and it became cooler in the course ofthe day. In the evening there were only 9 °C.There is hardly any frost in the Gran Chaco,

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Ill. 10. Acanthochelys macrocephala, the Big-headedPantanal Swamp Turtle, lives a very hidden life.

and if so only in the early morning. Snow andpermanent frost never appear.

The precipitation can vary a lot. Not onlydoes the annual precipitation fluctuate verymuch but often there is only a localprecipitation. For this reason both extremedry and green areas can be found. In all theannual precipitation usually falls in summerin form of a few showers reaching 100 mmand more.

Kinosternon scorpioides scorpioides(LINNAEUS, 1766), the Scorpion MudTurtle

Reptilia: Testudines: Cryptodira (hidden-necked), family: Kinosternidae

The most common turtle is the ScorpionMud Turtle. The only Kinosternon living inParaguay was first described as Kinosternonscorpioides seriei FREIBERG, 1936. CABRERA &COLANTONIO (1997) revised this again.

The oval carapace is dark brown to nearlyblack. The three dorsal keels are easily to beseen. According to ROGNER (1996) theydisappear when the animals grow old. Thecolour of the plastron is very variable. Allanimals we found had a monochrome plastroncoloured light-yellow to nearly black. Thetwo hinges of the plastron are characteristicfor this species. They allow motility of theplastron in the front and in theback and the turtle is enabled toclose her shell completely indangerous situations. This is notits only possibility of defendingitself though. The turtle’s bite canbe very painful as its big head isprovided with powerful jaws. Inaddition it can secrete a stinkingsubstance from the anal bladder,which forces most attackers towithdraw. The Scorpion Mud Turt-le can be found in shallow waterand outside water.

We frequently found Kinoster-non scorpioides scorpioides in theearly morning. Some turtles wereeven active at noon despite theextreme heat. We found one ofthem on a sandy track. The animals

always ranged near the shallow drainageditches of the tracks. Only one animal wasalso moving along the track. We found thisspecimen near Fortín Tte. Montanía. Allbodies of water in which we found Kinoster-non were temporary ones. The ditches oftenstore the water of a shower only for a fewdays. In extreme dry weather we could notfind any Kinosternon. These periods alsooccur during the warm summery rainyseasons. The animals do not continuouslyhibernate. As soon as rain is falling, theanimals can be noticed again (UNGER perso-nal notification). Rain, however, rarely fallsin the winter.

In the temporary bodies of water aquaticplants do not develop. For this reason theydo not appear on the turtles’ menu. Insteadthere are in all of them – even in puddles –uncountable tadpoles, young and old frogsand toads as well as their spawn. Further morewe saw many apple snails (Ampullaria spec.).These findings correspond to the range offood given by ROGNER (1996).

Because of the red loamy soil the water isusually coloured reddish and contains mud.In Paraguay the water is often more or lesssalty – a partly natural phenomenon. It is alsocaused by erosion or artificial irrigation. It isstriking that we have not found Kinosternon



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Ill. 13. In this small temporary pond lives a Big-headed PantanalSwamp Turtle.

Ill. 11. This species has a very broadhead and its eyes are adjusted to lifeunder water.

Ill. 12. Also Acanthochelys macro-cephala has throat barbles at the

lower jaw.

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Ill. 14. The females (left) carapace widthclearly differs from the males (right).

Ill. 15. The sexual dimorphism is moreeasily recognised by direct comparisonof the tail structure ($ above, ! below).

Ill. 16. The colouring of the plastron is likewise variable in Acanthochelys macrocephala.



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in salty water yet even if the amount of saltwas very low.

It seems as if Kinosternon s. scorpioideslives in every water-providing ditch. That iswhy we would like to give some places wherewe found this subspecies as examples: FortínTte. Montanía, track between Filadelfia andLoma Plata, near Rosaleda and Mariscal JoséFelix Estigarribia.

There is only little information about theside-necked turtles of the Gran Chaco. It ishard to find these animals in the wildernessbecause they usually hide. Similarly to Kino-sternon s. scorpioides they can also be foundin drainage ditches of the tracks. One canfind them far more seldom though. In additionthey can often be found in artificial ponds(Tajamares), which are used as reservoirs ofrainwater in agriculture. These ponds nor-mally have clearer water than the ditchesmentioned above. The Big-headed PantanalSwamp Turtle often shares its habitat withCaiman yacare, the Yacare Caiman. The waterof the artificial ponds usually contains salt.According to our observations Acantho-chelys macrocephala seems to have a con-siderably greater acceptance towards salt incontrast to Kinosternon s. scorpioides. Wecould never observe Acanthochelys macroce-phala outside the water or sunbathing.

There is a densely by Pantanal SwampTurtles populated Tajamar in the colonyMenno. This artificial pond is situated in thecentre of natural lagoons of salt. Duringdaytime we saw many animals drifting on thesurface of the water. During dusk we couldobserve more and more animals at the sametime. They seemed to become much moreactive now. In the beginning of February wewere lucky enough to watch a mating. Thecopulating animals were drifting on thesurface of the water in the early evening.Even in complete darkness with the help ofa torch we observed an Acanthochelysmacrocephala looking for food. Swimming,the animal systematically searched the shore,which was covered with small plants. Doingso it hit plants over and over again andchased away fish and frogs living there. Thenit tried to catch them in the open water.

On our journeys we were able to findAcanthochelys macrocephala in the follo-wing places: surrounding areas of Madrejon,Fortín Madrejoncito and the colony Menno.UNGER (personal notification) observed themon tracks between Filadelfia and Loma Plataat night and found a shell in Campo Maria,which he also put at the disposal of thezoological museum.

Acanthochelys macrocephala (RHODIN,MITTERMEIER & MCMORRIS, 1984), the Big-headed Pantanal Swamp Turtle

Reptilia: Testudines: Pleurodira (side-necked), family: Chelidae

The Big-headed Pantanal Swamp Turtlewas assigned to the genus Platemys in theoriginal description of RHODIN, MITTERMEIER &MCMORRIS 1984. In 1985 this genus wasthoroughly investigated by MCBEE et al. Asa consequence Platemys became a monotypicgenus, including only Platemys platycepha-la. All the other species were assigned to thegenus Acanthochelys. DERR et al. (1987)confirmed these results. FRITZ & PAULER

(1992a) described Phrynops chacoensis usinga few specimens originating from the surroun-ding area of Filadelfia, which, however, wasreversed in 1999 (FRITZ & PAULER 1999).

Acanthochelys macrocephala is the largestspecies of its genus. The turtles’ big head(macro=big, cephalos=head) was eponymousand has got distinctive cheeks. The top ofhead and neck is coloured grey to black, theunderside monochromatically yellow, partlywith grey patches. The head is covered withrough scales and the skin of the neck isscraggly. The extremities are yellowish androughly scaled, but the outside is greyish.The monochrome dark carapace is greyish-black to dark olive-green. There is a clear,flat dent along the central carapace scutes.The plastron is coloured like horn. The easiestway of differentiating between the sexes is tocompare the shells’ shapes. The carapace isconsiderably more slender with males thanwith females. A differentiation with the helpof the tails is difficult and only possible indirect comparison.

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Acanthochelys pallidipectoris (FREIBERG,1945), the Chaco Side-necked Turtle

Reptilia: Testudines: Pleurodira (side-necked), family: Chelidae

The second side-necked turtle indigenousto the Chaco is Acanthochelys pallidipec-toris. First it was also described as a speciesof the genus Platemys (FREIBERG 1945) butthen assigned to the genus Acanthochelys inthe revision of MCBEE et al. (1985).

The Chaco Side-necked Turtle remainsconspicuously smaller than Pantanal SwampTurtle and seems to be much more delicate.The head of this turtle is far less striking; itis only a little wider than the neck. The headand the extremities are very scaly, on thetopside they are grey-brown and on theunderside yellowish. Some particularly strongscales of the extremities can be yellow attheir tips. The skin of the neck is very roughand slightly pigmented with yellow and redtubercles. Both femoral have a big spursurrounded by several smaller spurs. This isthe most outstanding distinctive featurebetween Acanthochelys pallidipectoris andA. macrocephala. The carapace is reddish-brown. This colour is an optimal camouflageon the loamy soil of the Chaco. The plastronis light yellow, the sutures of the individualshells have got a black seam. Acanthochelyspallidipectoris also has the noticeable medi-al furrow along the central scutes. There ishardly any information about the Chaco Side-necked Turtle. We only found one specimenthat crossed a sandy track at night. The trackdid not have any ditches worth mentioning,so that on both sides of the track wateraccumulated up to a level of approximately10 cm only. There were uncountable tadpolesin this shallow reservoir of water and only afew metres further we found Kinosternonscorpioides scorpioides also being active.

MARIA & JAKOB KLASSEN found a severelyinjured animal in a drinking trough nearNeuwestland. It had obviously been kickedby a cow. After a short while the animal diedand was handed over to JAKOB UNGER so thatit now also belongs to the collection of theTierkundemuseum Dresden. Besides thisanimal and our observation near Rosaleda we

know of Acanthochelys pallidipectoris foundat Fortín Toledo (UNGER personal notification)and the areas surrounding Filadelfia (H. MEIER

in FRITZ & PAULER 1992b).

Geochelone chilensis (GRAY, 1870), theChaco Tortoise

Reptilia: Testudines: Cryptodira (hidden-necked), family: Testudinidae

It is speculated about the existence of thenorthern species Geochelone petersi FREI-BERG, 1973 again and again. AccordingPRITCHARD (1979) AUFFENBERG denied theexistence of Geochelone petersi and put itdown to sexual dimorphism. Our observationsalso revealed that there are considerabledifferences in shape, colour and size withinthe same area of circulation. That is why wewill use the nomenclature of IVERSON (1992).

The Chaco Tortoise is a compact medium-sized species. The carapace mostly remainsrelatively flat and has the colour of clay,which is an advantage in the Chaco. Exceptfor very old animals the rings of growth areclearly separated and at their borders coloureddarker. Geochelone chilensis tends to losecolour when it grows old so that elderlyanimals appear in a uniform grey colour. Butwe also saw a very old animal in the Chacothat stood out because of its dark-browncolour. Usually the shell is almost round. Thestrong extremities are monochromatic lightand very scaly. It is quite frequent amongmale and female specimens to have anextremely enlarged scale at the upper part ofthe front leg.

We had to face special problems when welooked for the Chaco Tortoise in its naturalenvironment, because the animals obviouslyoccur exclusively in dry forests and savannas.Because of their density it is hard to searchthem effectively – especially in summer wheneverything is green. Both our observationsand questionings of natives showed that theanimals never are found near agriculturalland or meadows. That is why Geochelonechilensis is often just found coincidentally.It is most probable to find animals shortlybefore, during or after showers when they



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Ill. 17. Another Pleurodira of the Chaco isAcanthochelys pallidipectoris, the ChacoSide-necked Turtle.

Ill. 18. Its dark brown colour camouflagesAcanthochelys pallidipectoris in the loamysoil under water.

Ill. 19. Acanthochelys pallidipectorisshows the characteristic rough neck, whichis typical for the genus. The head isclearly less broad than in Acanthochelysmacrocephala.

Ill. 20. The extremely enlarged spurs arecharacteristic for the Chaco Side-neckedTurtle.

Ill. 21. Plastron colouration of Acanthochelyspallidipectoris.

Ill. 22. Acanthochelys macrocephala andAcanthochelys pallidipectoris clearly differ insize. The picture shows adult males.

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Ill. 23. Further morphological differencesbetween the two species of Acanthochelys.

Ill. 24. Geochelone chilensis, the ChacoTortoise, is widespread in the Chaco. Itspowerful front legs are highly scaled.

Ill. 25. The colour is perfectly adapted tothe habitat.

Ill. 28. Geochelone chilensis might varyclearly in colour and shape.

Ill. 26. The dense thorn bush dry forest is ahabitat for Geochelone chilensis.

Ill. 27. A semi-adult specimen of the ChacoTortoise was found on this former pasture.



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Ill. 29. A Chaco Tortoise iscrossing a road in the north ofParaguay.

cross tracks in order to change to anotherforest.

In dry forests it is possible for the ChacoTortoises to protect themselves from intenseheat and to survive cooler and dryer seasons.They use self-dug caves and also abandonedrodent burrows. Succulent plants and foliagefallen on the ground are almost the only foodavailable for tortoises in dry forests. VINKE &VINKE (2000b) have already comprehensivelydescribed that Geochelone chilensis does notcontinuously hibernate in the Chaco. As soonas it rains it can be seen again.

Meanwhile we know about the followingareas of occupancy: Fortín Toledo, surroun-ding areas of Mariscal José Felix Estigarribia(skeleton findings), Fortín Tte. Montanía,track between Fortín Tte. Montanía andMadrejon, near Fortín Defensores del Chaco.

MÜLLER 1971, MÜLLER & HELLMICH 1936). It isthe same with the morphology of ParaguayanRed-footed Tortoises. PRITCHARD & TREBBAU

(1984) tried to divide the Red-footed Tor-toises into seven variants according to theirextents of occurrence. They say that Paragua-yan animals are characterized by their re-duced size of 20 to 22 cm as a maximum andby their uniform red scales and patterns ontheir heads. Such Red-footed Tortoises arenow known as ”Cherryheads”.

We disagree on the opinion that theseanimals originate from Paraguay. We did notsee any Red-footed Tortoises there that matchthis description, neither during our obser-vations in the biotopes nor by our enquiriesin the Paraguayan Natural History Museum(MNHNP) or during our visits of the biologicalinstitutions near the reservoir Itaipú. In

Presumably Geochelone chilensis is widelyspread in the central Chaco.

Geochelone carbonaria (SPIX, 1824), theRed-footed Tortoise

Reptilia: Testudines: Cryptodira (hidden-necked), family: Testudinidae

Red-footed Tortoise is another topic thatwas rarely dealt with. There are manypublications on observations in naturalhabitats in the northern equatorial extent ofoccurrence (FRETEY 1977, KORNACKER & MEYER

ZUR HEYDE 1998, MEDEM 1958, 1960, 1962,1969, MEDEM et al. 1979, MÉTRAILLER & LE

GRATIET 1996, MOREIRA 1989, MOSKOVITS &BJORNDAL 1990, PRITCHARD & TREBBAU 1984,VINKE & VINKE 2000c), but there is hardly anyinformation about the southern part (CEI 1993,

contrary the animals there are characterizedby a huge size. The size of 21 wild adultanimals from Northwest-Paraguay, which wehave measured, ranged between 26.4 and36.5 cm. Only five of these animals weresmaller than 30 cm. In the biologicalinstitution at the reservoir Itaipú the animalswere even bigger. They mainly come fromNortheast-Paraguay (Concepción, SerraníaSan Luís). Ten out of 14 adult animals wereas big as 41 to 45 cm, the other four specimensmeasured 35 to 38 cm. The biggest sample wefound in human care in Filadelfia. It had asize of 59.3 cm (VINKE & VINKE 1999). Thisanimal originates from the surrounding areasof the place called KM 145.

The Paraguayan Red-footed Tortoiseshows an extreme variety in the colour of the

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Ill. 30. In the national park ”Defensores Del Chaco”we found Red-footed Tortoises (Geochelone carbo-naria) of enormous size.

head and the extremities. We found animalswith a yellow head and yellow extremities,but also animals with orange scales. Even thered-yellow colour know from northern animalsis quite frequent. The colour of the carapacereaches from dark black to light brown, bothwith and almost without light round patches.An almost complete black plastron is uni-form. The animals seem rather ungainly, evenadult males hardly show a waist in contrast tothe northern kind. The proportions of thelength of the carapaces’ shells are extremelyvariable. There often is a distinctive bumpyscale at the upper forelimb as known from theCherryheads. Shape and size of the tail areclearly different from those of their relationsin the north.

The Red-footed Tortoise mainly inhabitsdry forests in Paraguay. It also lives in smallerembedded savannas. In all the preferredbiotopes appear to be slightly damper andtherefore a little greener than those ofGeochelone chilensis, the forests are higherand less thorny. These tortoises like to benear temporary riverbeds or at places with alower level where rainwater accumulates overa long period of time.

The dry forest provides the tortoises withan opulent menu in summer. Wewere able to observe the animalseating non-flowering plants,cactus, ferns and bromelias,which grow there in large num-bers. In the cool dry months foodis limited to dry foliage, cactusand bromelias. We frequentlysaw Geochelone carbonaria nextto Quiabentia spec. These arecactus trees with thick leaveswhich the tortoises like eating.

We found Geochelone carbo-naria in the following places:national park ”Defensores delChaco” and near KM 145, more-over UNGER found several animalson the track Loma Plata towardsFiladelfia, near Toledo, in La Patriaand the Rio Verde. Occurrencesin East-Paraguay are known fromthe biological station Serranía San

Luís (MNHNP) and from Ybabopo (FMNH). Inall the Red-footed Tortoise lives in a verylarge area of circulation in Paraguay.

Note on thanksSpecial thanks to our friends JAKOB & MARIA

UNGER who made some unforgettable excur-sions possible and unquestioningly sharedtheir comprehensive knowledge about theflora und fauna with us. Everybody who wasallowed to experience their hospitality knowswhat we are talking about.

LiteratureCABRERA, M. R. & S. E. COLANTONIO (1997):

Taxonomic Revision of the South Ame-rican Subspecies of the Kinosternonscorpioides. – Journal of Herpetology,Athens, Ohio 31(4): 507-513.

CEI, J. M. (1993): Reptiles del noroeste,nordeste y este de la Argentina : herpe-tofauna de las selvas subtropicales, Puna yPampas. Mus. Reg. di Sc. Nat. Torino.Monografía 14, 949 pp.

DERR, J. N. & J. W. BICKHAM & I. F. GREENBAUM

& A. G. J. RHODIN & R. A. MITTERMEIER

(1987): Biochemical systematics andevolution in the South American turtle



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Ill. 34. Also black-shelled Red-footed Tortoises with an orange to

red head can be found in theChaco.

Ill. 31. At the end of the dry season herbi-vores find only little food on the forestground.

Ill. 32. The food supply is luxurious insummer – like on this clearing, where weobserved a sunbathing Red-footed Tortoise.

Ill. 33. In the Chaco Geochelonecarbonaria shows an exceptionalvariation in colour. This animal’shead is yellow and its carapace is

brown.

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Ill. 35. The plastron of Paraguayan Red-footed Tortoises is nearly uniformly almostblack.

Ill. 36. The male (middle) cannot be distin-guished from the females by its shape. Thewell-known waist of the northern type isnonexistent here.

Ill. 39. Humans are the adult Red-footedTortoises’ only enemy besides the jaguar

(Panthera onca).

Ill. 37. One often finds Geochelone carbo-naria in such caves dug by the giantarmadillo (Priodontes giganteus).

Ill. 38. A preferred feed plant: Quiabentiaspec. These cactus trees grow to an impres-sive size.



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genus Platemys (Pleurodira: Chelidae). –Copeia, Washington, 1987(2): 370-375.

FREIBERG, M. A. (1936): Una nueva tortugadel norte Argentino. – Physis, BuenosAires, 12: 169-171.

— (1945): Una nueva especia de tortuga delgénero Platemys WAGLER. – Physis, BuenosAires, 20: 19-23.

— (1973): Dos Nuevas Tortugas Terrestres deArgentina. – Bol. Soc. Biol. Concepción,46: 81-93.

FRETEY, J. (1977): Les Chéloniens de Guyanefrançaise. – 1. Etude préliminaire. Paris(Thesis, Univ. Paris), 202 pp.

FRITZ, U. & I. PAULER (1992a): Phrynopschacoensis spec. nov. (Reptilia, Chelidae),eine neue Krötenkopfschildkröte. – Mit-teilungen Zoologisches Museum, Berlin,68(2): 299-307.

— (1992b): Erstnachweis von Acanthochelyspallidipectoris (FREIBERG, 1945) für Para-guay – Herpetozoa, Wien, 5(3/4): 135-137.

— (1999): Phrynops chacoensis FRITZ &PAULER, 1992, ein Juniorsynonym vonPlatemys macrocephala RHODIN, MITTERM-EIER & MCMORRIS, 1984. – Salamandra,Rheinbach, 35(1): 53-56.

GRAY, J. E. (1870): Notice of a new Chiliantortoise (Testudo chilensis). Ann. Mag.Natur. Hist., London 4(6): 190-191.

IVERSON, J. B. (1992): A Revised Checklistwith Distribution Maps of the Turtles ofthe World. – Richmond, Indiana (Privatelyprinted): 318 pp.

KORNACKER, P. M. & C. MEYER ZUR HEYDE (1998):Herpetologische Eindrücke einer Vene-zuelareise Teil 3: El Embalse de Guri – derGuri-Stausee. – Elaphe, Rheinbach, 6(4):61-69.

MCBEE, K. & J. W. BICKHAM & A. G. J. RHODIN

& R. A. MITTERMEIER (1985): Karyotypicvariation in the genus Platemys (Testudi-nes: Pleurodira). – Copeia, Washington,1985(2): 445-449.

MEDEM, F. (1958): Informe sobre reptiliesColombrianos (II). El conocimiento actualsobre la distribución geografica de losTestudinata en Colombia. – Bol. Mus.Cienc. Nat., Caracas, 2-3: 13-45.

— (1960): Datos zoogeograficos y ecologicossobre los Rios Amazonas, Putumayo yCaquetá. – Caldasia 8: 341-351.

— (1962): La distribución geografica yecologica des los Crocodylia y Testudinataen el Departamento del Choco. – Rev.Acad. Colomb. 11: 279-303.

— (1969): Estudios adicionales sobre losCrocodylia y Testudinata del Alto CaquetáRio Caguan. – Caldasia, 10: 329-353.

MEDEM, F. & V. O. CASTANO & R.M. LUGO

(1979): Contribución al conocimientosobre la repruducción y el crecimiento delos ”morrocoyes” (Geochelone carbonariay G. denticulata) (Testudines: Testudin-idae). – Caldasia, 12: 497-511.

MÉTRAILLER, S. & G. LE GRATIET (1996) Tortuecontinentales de Guyane française -Continental Turtles of the French Guiana.– Bramois/Schweiz (Édition P.M.S.),127 pp.

MOREIRA, G. R. S. (1989): Sympatry of theturtles G. carbonaria and G. denticulatain the Rio Uatuma Basin, Central Ama-zonia. – J. of Herpetology 23(2): 183-185.

MOSKOVITS, D. K. & K. BJORNDAL (1990): Dietand food preferences of the tortoisesGeochelone carbonaria and G. denticulatain nortwestern Brazil. – Herpetologica46(2): 207-218.

MÜLLER, L. & W. HELLMICH (1936): Wissen-schaftliche Ergebnisse der Deutschen GranChaco-Expedition. Amphibien und Rep-tilien. I. Teil: Amphibia, Chelonia,Loricata. – Stuttgart (Verlag von Streckerund Schröder), 120 pp.

MÜLLER, P. (1971): Beobachtungen an brasi-lianischen Geochelone carbonaria. – AquaTerra, Biberist, 8(6/7): 69-75.

NORMAN, D. R. & L. Naylor (1994): Anfibiosy reptiles del Chaco Paraguayo –Tomo I.Amphibians and Reptiles of the ParaguayanChaco – Volume I. – Heredia, Costa Rica,281 pp.

PRITCHARD, P. C. H. (1979): Encyclopedia ofTurtles. – Neptune, NJ, (T.F.H. Pub-lications, Inc.), 895 pp.

PRITCHARD, P. C. H. & P. TREBBAU (1984): TheTurtles of Venezuela. – Cornell UniversityIthaca, New York, (Society for the Study of

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Amphibians and Reptiles), 403 pp.RHODIN, A. G. J. & R. A. MITTERMEIER & J. R.

MCMORRIS (1984): Platemys macrocepha-la, a new species of chelid turtle fromcentral Bolivia and the Pantanal region ofBrazil. – Herpetologica, Lawrence, 40(1):38-46.

ROGNER, M. (1995): Schildkröten 1. – Hürt-genwald (heiro), 192 pp.

SEIBERT, P. (1996): Farbatlas Südamerika:Landschaften und Vegetation. – Stuttgart(Ulmer), 288 pp.

SPIX, J. B. VON (1824): Animalia nova, speciesnovae Testudinum et Ranarum quas initinere per Brasiliam annis 1817-1820collegit et descipsit. München (Hübsch-mann), 53 pp.

VINKE, T. & S. VINKE (1999): Ein wahrer Gi-gant – Rekordgröße für die Köhlerschild-kröte, Geochelone carbonaria (SPIX, 1824).

– Emys, Reidling, 6(2): 27-29.— (2000a): Artenschutzdesaster in Paraguay.

– Radiata, Haan, 9(2): 29-33.— (2000b): Biotopbeobachtungen an Geo-

chelone chilensis (GRAY, 1870) im GranChaco von Paraguay. – Sauria, Berlin,22(2): 25-29.

— (2000c): Optimierung der Haltung undZucht der Köhlerschildkröte Geochelonecarbonaria (SPIX, 1824) aus der Guayana-Region. – Salamandra, Rheinbach, 36(4):233-246.

AuthorsThomas & Sabine VinkeDellerstraße 61D-42781 Haan

Translated from German by ROMI ZÄSKE, Leipzig,Germany

The turtle and tortoise fauna of the central Chaco of Paraguay

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