N. Jb. Geol. Paläont. Mh.

2002 (11)

641 – 648

Stuttgart, November 2002

Procolophon brasiliensis n. sp., a new procolophonid reptile from the Lower Triassic of southern Brazil Juan Carlos Cisneros and Cesar Leandro Schultz, Porto Alegre With 2 figures CISNEROS, J. C. & SCHULTZ, C. L. (2002): Procolophon brasiliensis n. sp., a new procolophonid reptile from the lower Triassic of southern Brazil. - N. Jb. Geol. Paläont., 2002: 641 – 648; Stuttgart. Abstract: A new taxon within the genus Procolophon is reported from the Sanga do Cabral Formation of southern Brazil. The type-material is represented by a single partial skull and a lower jaw. It is distinguished from other species of the genus on the basis of vomerian dentition characters. The importance of the palatal region, especially the teeth, for the taxonomy of the genus and also of the Procolophonidae is stressed . Resumo: Apresenta-se um novo taxon dentro do gênero Procolophon na Formação Sanga do Cabral do Sul do Brasil. O material-tipo constitui-se de um crânio parcial e mandíbula. Distingue-se de outros representantes do gênero através de caracteres da dentição vomeriana. Salienta-se a importância da região palatal, especialmente a dentição, como ferramenta para a diagnose dentro do gênero e mesmo no clado Procolophonidae. Zusammenfassung: Eine neue Art der Gattung Procolophon wird aus der brasilianischen Sanga do Cabral Formation beschrieben. Der Holotypus besteht aus dem Teil eines Schädels nebst Kieferknochen. Die Art unterscheidet sich von anderen Arten der Gattung durch Merkmale der Zahnreihe des Vomers. Die Wichtigkeit der GaumenRegion und besonders der Zahnreihe für die taxonomische Diagnose Gattung und der Procolophonidae allgemein wird hervorgehoben.

0028-3630/02/2002-0641 $2.00  2002 E. Schweizerbart’sche Verlagsbuchhandlung, D-70176 Stuttgart

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Introduction Procolophon is one of the best known fossil tetrapods. P. trigoniceps is the second most common taxon in the Early Triassic Lystrosaurus Assemblage Zone of South Africa (GROENEWALD & KITCHING, 1995), and it is also known from Antarctica (COLBERT & KITCHING, 1975). This genus was first reported for South America (BARBERENA et al. 1981) in sediments from Sanga do Cabral Formation of Southern Brazil, and assigned to a new species, P. pricei (LAVINA, 1983). Procolophonid remains are the most common vertebrate fossils in this formation, followed by temnospondyle and archosaur fragments. The present specimen is a partial skull and lower jaw which can surely be assigned to the genus Procolophon, but it shows autapomorphies of its own in the vomerian dentition.

Systematic Paleontology Parareptilia OLSON, 1947 sensu LAURIN & REISZ, 1995 Procolophonidae SEELEY, 1888 Procolophon OWEN, 1876 D i a g n o s i s o f t h e g e n u s (sensu COLBERT, 1975): Small, advanced cotylosaurs, with deep skull, triangular in shape. Orbits very large and elongated, and pineal opening large; nares almost terminal. Quadrate forward of occiput, with articulation well below level of teeth. Otic notch large; tabular and quadratojugal expanded and spike like. Palate restricted and with elongated choana. Lower jaw deep, with strong coronoid and elongated retroarticular process. Four premaxillary teeth and seven maxillary teeth, of which five or six transversely broad and chisel-shaped. Nine dentary teeth, the last six chisel-like. Post-cranial skeleton compact. Neural arches of vertebrae expanded, in cotylosaurian fashion. Ribs essentially single-headed. Girdles strong and limbs comparatively short. Feet broad, with full complement of carpal and tarsal element. Tail short.

Type species:

Procolophon trigoniceps OWEN, 1876

G e o l o g i c a l h o r i z o n of the type species: Lystrosaurus Assemblage Zone, Katberg Formation of South Africa and Fremow Formation of Antarctica, Early Triassic. Procolophon brasiliensis n. sp.

(Fig. 1, 2)

E t y m o l o g y : After Brazil, where the new material comes from. D i a g n o s i s : Vomer possessing a single tooth row extending over the entire bone, reaching the pterigoid-vomer contact, with a small diastema in the posterior third part of the row. Three vomerine fangs in the anterior part of the bone, arranged in a “V” shape, pointing forwards.

Procolophon brasiliensis n. sp. from the Lower Triassic of southern Brazil

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Fig. 1. a: palatal view of Procolophon brasiliensis n. sp.; b-d: palatal restorations (b: P. brasiliensis; c: P. pricei, based on the holotype UFRGS PV231T; d: P. trigoniceps, redrawn after BROILI & SCHRÖDER (1936)). Abbreviations: ec, ectopterygoid; pl, palatine; pm, premaxilla; ppf, prepalatal foramen; pt, pterygoid; sof, suborbital foramen; v, vomer.

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H o l o t y p e : MCN PV 1905, partial skull and jaw originally in occlusion, separated during preparation to permit further examination of the material. From Rincão dos Weiss Village, Mata County, Rio Grande do Sul State, Southern Brazil (S29033'27.35”/W53026'56.43”). In the collections of Museu de Ciências Naturais, Fundação Zoobotânica do Rio Grande do Sul. Collected by Fundação Zoobotânica do Rio Grande do Sul on 14.03.1997. G e o l o g i c a l H o r i z o n : Sanga do Cabral Formation, Rio Grande do Sul State, southern Brazil, Lower Triassic.

Description The material (Fig. 2) is imbedded in a carbonatic concretion and was heavily eroded externally. The skull roof, as well as the posterior part of the cranium and jaw are not preserved, and there is no information about the shape or the size of the quadrate-jugal process. Most of the information of the specimen is obtained from the palatal view. The skull and lower jaw, as a whole, were laterally compressed. The premaxillae are damaged. The right one is more complete, it has three partially preserved teeth, with space for one more tooth. These teeth, although broken, seem to be conical in shape, as expected in Procolophon. The premaxilla has a small area of contact with the maxilla laterally. There are no teeth preserved in the left premaxilla. The contact with the vomers has a “V” shape. The external surfaces of the maxillae are worn, and all the teeth have their pulp cavities exposed. The first tooth is preserved only in the right maxilla and is conical. The following teeth are bulbous and transversely expanded, characteristic of Procolophon. They are best seen in the left maxilla, which has seven of them. There seem to exist pulp remains of a small replacement tooth, behind the last bulbous tooth. Altogether, the maxilla of MCN PV1905 seems to bear nine teeth. Teeth implantation is protothecodont, as in most procolophonids. On the palatal surface, the vomers are typical of Procolophon, but the tooth arrangement is different from the previously known species. The vomer possesses a single longitudinal row of teeth, starting in the suture with the premaxilla and ending in the vomerpterygoid contact. There seems to be a prepalatal foramen in front of the tooth row, best seen in the right vomer. For a better understanding of the vomerian dentition, each

Fig. 2. Procolophon brasiliensis n. sp.; a: skull, palatal view (stereophotograph); b: occlusal view of the mandible; c: skull and mandible, right lateral view; d: skull and mandible, left lateral view.

Procolophon brasiliensis n. sp. from the Lower Triassic of southern Brazil

Fig. 2. (Legend see p. 144)

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vomerian tooth is here referred to with a number (vomerian tooth number = Vn). There are thirteen teeth on the left vomer and twelve on the right one. V1 to V3 in each vomer are fangs, bigger than the other ones, and follow a diagonal line in each vomer. Both lines contact antero-medially and separate postero-laterally, following an inverted “V” pattern. V4 in each vomer is small and positioned more medially in the row of teeth. V5 to V12 in the left vomer are small and there is a small diastema between V8 and V9. In the right vomer, V5 to V13 are also small, with the exception of V9 which is a fang. There is no fang present on the left vomer in the respective position. Between V9 and V10 there is also a small diastema. V10 and V13 were broken before collecting the material. The alveoli of these teeth are preserved. The vomerian tooth rows extend through the palatine. The left palatine bears four teeth. In the right palatine, there are two alveoli preserved, but there is space for at least two more, so it also could have borne a fourth teeth. Lateral to the left palatine, the suborbital foramen can be seen. Only the most anterior part of the pterygoids are preserved, although part of them have been slightly moved below the vomers due to compression. There are two alveolus visible in the left pterygoid and three teeth in the right one. Only the anterior part of the left ramus of the lower jaw is well preserved. A part of the right ramus has been preserved as a natural mold, with some teeth remains. In lateral view of the left ramus the dentary can be seen, the posterior part of which is broken, leaving the splenial exposed. These are the only bones preserved in the lower jaw of MCN PV1905. In medial view, the wide splenial and the anterior part of the dentary can be seen, showing the meckelian groove. The first incisor is not preserved in our material, the next tooth should also be an incisor but it is not complete. It is followed by two conical teeth. Backwards in the jaw there are seven bulbous, transversely expanded teeth, which increase in size posteriorly. The sixth one is the biggest, and is followed by a smaller replacement tooth. Altogether, this specimen should have possessed at least eleven lower jaw teeth.

Discussion The palatal dentition seems to be an important tool in the comparison between the species of the genus Procolophon, and it could be valid for other procolophonids as well. Nevertheless, when Procolophon trigoniceps was described by OWEN in 1876, the palate of the holotype was not prepared (and has remained unprepared), so that no palatal tooth characters were taken in account in the original diagnosis of the form. The same occurred with other Procolophon species proposed by OWEN (P. minor), SEELEY (P. griersoni, P. cuneiceps, P. laticeps, P. platyrhinus, and P. sphenorinus) and BROOM (P. baini). These species were proposed mostly on the basis of differences in size and/or proportions, most of them being better explained by individual and ontogenetic variations, and/or taphonomic factors and were synonymized by BROOM (1936) and others. None of these species are accepted at present with the exception of P. laticeps, “rescued” by HAMLEY & THULBORN (1993). During the examination of Procolophon specimens in different collections, we have noted that there are certain variations in palatal tooth arrangement in the several specimens attributed to P. trigoniceps in different parts of the world, so that it is a little difficult to define which is the pattern of

Procolophon brasiliensis n. sp. from the Lower Triassic of southern Brazil

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palatal dentition for this species. However, these variations seem to represent different patterns, which may indicate different species and/or geographic variations of the same species. This matter will be developed in another work. Among the Karoo specimens examined, we have found one with the pattern that characterizes P. pricei from Brazil (CISNEROS & SCHULTZ in prep.) indicating that this species was present also in that region. However, none of the arrangements observed is similar to the condition present in P. brasiliensis. The vomerian dentition of P. brasiliensis (Fig. 2) is quite different from previous Procolophon species in possessing three anterior vomerian fangs. P. pricei possesses just one, while P. trigoniceps specimens may have one or two. Besides, in the specimens with two fangs, they are arranged nearly transversely, and not diagonally as in P. brasiliensis. HAMLEY & THULBORN (1993) recognized the taxon P. laticeps, on the basis of its temporal fenestration and other skull roof characters. Unfortunately, a comparison between this species and P. brasiliensis is problematic, because the skull roof of MCN PV1905 is lost, while fangs can not be seen in the P. laticeps specimens, since these need more preparation in the palate. However, the posterior portion of the vomer seen in specimen NH 1947 (Natural History Museum, London) is different from MCN PV1905, since it has no teeth (DIAS-DA-SILVA, pers. comm.). Vomerian rows of teeth in P. brasiliensis are also unique among the genus because of its large number of teeth. Although some P. trigoniceps specimens have nearly the same number of vomerian teeth as our material, these individuals have two rows of teeth rather than the single one of P. brasiliensis. Besides, these individuals have also vomerian teeth reaching the pterygoid contact, but these are separated from the main pterygoid rows by a great diastema, thus being different from our material. This new taxa increases the previously known fauna of the Sanga do Cabral Formation of Southern Brazil, where Procolophon is the most common taxon. NEVELING et al. (1999), noted that Procolophon is also the most common taxon in the lower part of the “Impoverished Zone” between the Lystrosaurus and Cynognathus Assemblage Zones, and proposed a Procolophon Zone for that part of the record. Based on the predominance of the same taxon, it is probably that the fauna from Sanga do Cabral Formation may be stratigraphically correspondent, or at least close, to the South African Procolophon Zone. This topic is being developed in another paper. Acknowledgments Thanks are due to the kindness of A. M. RIBEIRO and J. FERIGOLO at the Fundação Zoobotânica do Rio Grande do Sul for the loaning of the material and working facilities during preparation stage and to CAPES for funding this study.

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References BARBERENA, M. C., LAVINA, E. L. & BECKER, M. R. (1981): Sobre a presença de tetrápodos na Formação Sanga do Cabral (Grupo Rosário do Sul), Triássico do Rio Grande do Sul, Brasil. - An. do II Congr. Latino-Americano Paleont., 295-304. BROILI, F. & SCHRÖDER, J. (1936): Beobachtungen an Wirbeltieren der Karrooformation. XXI: Über Procolophon OWEN. - Sber. Akad. Wiss. München, 2: 239-256, pls. 3-6. BROOM, R. (1936): The South African Procolophonia - Ann. Transvaal Mus., 18: 387391. COLBERT, E. H. & KITCHING, J.W. (1975): The Triassic reptile Procolophon in Antarctica. -Amer. Mus. Novit. 2566. HAMLEY, T. & THULBORN, T. (1993): Temporal fenestration in the primitive Triassic reptile Procolophon. - New Mexico Mus. Natur. Hist. Sci., Bull., 3: 171-174. LAVINA, E. L. (1983): Procolophon pricei sp. n., um novo réptil procolofonídeo do Triássico do Rio Grande do Sul. Iheringia. (Geol.), 9: 51-78. NEVELING, J., RUBIDGE, B. S. & HANCOX, P. J. (1999). A lower Cynognathus Assemblage Zone fossil from the Katberg Formation (Beaufort Group, South Africa). - South Afr. J. Sci. 95: 555-556. GROENEWALD G. H. & KITCHING J. W. (1995): Biostratigraphy of the Lystrosaurus Assemblage Zone. In: Biostratigraphy of the Beaufort Group (Karoo Supergroup), ed. RUBIDGE, B. S. 1995. - SACS Biostratigraphic Ser., 1: 35-39. Received: March 1, 2001. Revised manuscript accepted by the Tübingen editors: XXX

Address of the authors: JUAN CARLOS CISNEROS, CESAR LEANDRO SCHULTZ, Universidade Federal do Rio Grande do Sul - Instituto de Geociências av. Bento Gonçalves 9500, CP 15.001, 91540000, Porto Alegre, RS, Brazil. e-mail: [email protected]

Procolophon brasiliensis n. sp., a new procolophonid ...

Parareptilia OLSON, 1947 sensu LAURIN & REISZ, 1995 ... D i a g n o s i s o f t h e g e n u s (sensu COLBERT, 1975): Small, advanced cotylosaurs, with deep ...

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