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Journal of Vertebrate Paleontology 23(2):464467, June 2003 2003 by the Society of Vertebrate Paleontology



EDIO-ERNST KISCHLAT1 and SPENCER G. LUCAS2, 1Instituto de Geocie ncias, Universidade Federal do Rio Grande do Sul, Porto Alegre, Brazil; 2New Mexico Museum of Natural History, 1801 Mountain Road NW, Albuquerque, New Mexico 87104 U.S.A.,

Phytosaurs were Late Triassic archosaurs whose abundant and widely distributed fossils have made them a key element in the interpretation of Late Triassic tetrapod biostratigraphy and paleobiogeography. Phytosaur fossils are known from North America, Europe, India, Thailand, Turkey, North Africa and Madagascar (e.g., Westphal, 1976; Buffetaut, 1993). The few putative phytosaurian records previously reported from South America (Rusconi, 1947; Reig, 1961; Bonaparte, 1978; Dornelles, 1990) were either inconclusive or incorrect (Charig and Reig, 1970; Baez et al., 1993), so, until now, no diagnostic phytosaur material has been described from the long known and extensive Triassic tetrapod faunas of Brazil and Argentina. Here, we document a phytosaur fossil from the Late Triassic of Brazil, briey mentioned in earlier papers (Kischlat, 1996, 2001; Faccini et al., 2000; Schultz et al., 2000), and thereby establish the presence of phytosaurs in the South American Late Triassic. Institutional Abbreviations Instituto de Geocie ncias, Universidade Federal do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul, Brazil (UFRGS); Museu de Cie ncias Naturais, Fundac a o Zoobota nica, Porto Alegre, Rio Grande do Sul, Brazil (MCN-FZB); New Mexico Museum of Natural History, Albuquerque, New Mexico, U.S.A. (NMMNH). PROVENANCE AND AGE The specimen documented here is from the Botucara locality, 6 km west of Candela ria, Rio Grande do Sul, Brazil. This locality, a highway cut at Botucara Hill, is in crossbedded sandstone and massive-to-laminar siltstone of the Caturrita Formation (Faccini et al., 2000). The Caturrita Formation is interpreted to have been deposited by a perennial braided uvial system in a humid climate (Holz and Scherer, 2000). The fossils were exposed on the ground, beside the highway. Other vertebrate fossils from this locality are the dicynodont Jachaleria candelariensis (Arau jo and Gonzaga, 1980), archosaurian and/or dinosaurian teeth (Dornelles, 1990; Azevedo, 1993, 1999; Faccini et al., 2000; Schultz et al., 2000), the saurischian dinosaur Guaibasaurus (Bonaparte et al., 1999) and a fragmentary pelvis and vertebrae of a ?herrerasaurid theropod dinosaur (Kischlat and Barberena, 1999). This vertebrate fossil assemblage is of late Carnian (Adamanian) age (Lucas, 1998; Schultz et al., 2000). DESCRIPTION MCN-FZB 1865 (Fig. 1) is a snout fragment composed of a main part (a) that measures 87 mm (long axis) by 41 mm and 45 mm (minimum and maximum width, respectively), and seven other disarticulated fragments. There is no reason to believe that the fragments represent more than one individual. The main part includes parts of four bones sutured to each other; these are paired elements that surround the midline. It is slightly crushed and distorted to one side. The sides are convergent to one plane of fracture, interpreted by us to be rostral (the shorter end). The additional scattered fragments were collected several months after the main one, and the assumed rostral plane of fracture is sharper than the caudal plane, which is slightly polished. This suggests that the fossil was preserved with the tip of the snout into the outcrop, so that the rest of the skull is no longer preserved. Efforts to nd additional material proved fruitless. On the tooth-bearing side (Fig. 1A), there is a lateral element, the row of teeth and a medial element, sutured to its counterpart. We identify the lateral element as the dentary, and the medial one as the splenial. Between both there is a continuous rostro-caudal line on both sides that

we identify as the splenio-dentary suture. This is not serrated, but is a plane and reaches deep into a cavity lled by sediments. It runs along the medial plane of the alveoli and, on the right side of the rostral end, this suture shifts medially, crossing the dorsal face and starting the rostral splenial wedge characteristic of phytosaurs (Fig. 2). Both splenials contact each other in a non-serrated, but planar, intersplenial suture. The dorsal face of the splenial (and probably its rostral continuation on the dentary) are at a level higher than the level of alveoli, and its dorsal surface is nearly smoothonly minute, parasagitally-oriented lineations are present. Some of the alveoli are not completely surrounded medially by bone, and the interalveolar septa make contact with the splenial. Externally (laterally and ventrally), the bone surface of the main fragment is ornamented by a sparse array of linear grooves and pits, which are rostrocaudally directed. Three main grooves (Fig. 1BC) are present: one arranged ventrally, lateral to the tooth row; the second oriented laterally; and the third oriented sagittally, coincident with the dorsal part of the intersplenial suture. The paired grooves are best preserved on the right side. On the left side, they are disrupted at a place of weakness by crushing. Ventrally, the same bones (Fig. 1B) can be discerned, also forming non-serrated, articulations (spleniodentary and intersplenial sutures). The splenials also wedge rostrally. Internally, in the caudal plane of fracture (cross-section) (Fig. 1D), both splenials have a shape resembling an upside-down mushroom, but the most internal parts are not fully preserved. This region is formed by cracked spongy bone lled with sediment. Laterally, these bones (best preserved on the right side) surround medially and ventrally a cavity here intepreted as the Meckelian canal. This cavity is surrounded by the dentary laterally and dorsally, with the alveoli positioned mediodorsally. The articular faces of both dentary and splenial are oblique in the caudal part of the preserved portion of the spleniodentary suture, but rostrally, they are nearly vertical. The intersplenial suture has the continuous coincident groove cited earlier, and is not serrated, but is also planar. Only the ventral part of the second fragment of bone (b) is preserved (Fig. 1B). It shows the interdentary suture as a planar articulation, and the articular faces show some minute, rostrocaudally positioned, lamellae of bone that t into depressions on its counterpart. A groove can also be discerned, probably the continuation of the groove at the ventral part of the intersplenial suture in the main fragment, but there is no preserved connection between the two. The sides of the second fragment are less convergent, and its position evidently was more rostral than the main fragment. Rostrocaudally directed grooves and pits are also visible. A third fragment (c), roughly articulated with the main one, also shows this ornamentation and represents part of a right dentary, where the splenial articular face is slightly oblique caudally, but fully vertical rostrally. Probably, this was the place of the tip of the wedge of both splenials, inserting between both dentaries (Fig. 2). The fourth fragment of bone (d) represents the dorsal and medial part of the right dentary, in a position dorsal to c (Fig. 1A). Unfortunately, no obvious t can be recognized. It has alveoli and the same kind of striations on the dorsal surface as in a, and the medial articular face shows rostrocaudal lamellae. The dorsal surface also has a kind of rostrocaudal bump, rostral to the splenial wedge. There is a disrupted tooth row laterally, and caudomedially there is an identation that represents the articular face for the rostral tip of the splenial wedge. This interpretation leads us to conclude that, at this point, the symphysis




FIGURE 1. Phytosaur from the Caturrita Formation, Brazil, MCN-FZB 1865. A, Dorsal view of lower jaw fragments; note that a mirror image of d is included. B, Ventral view of lower jaw fragments; note that a mirror image of e is included. C, Lateral view of fragment a showing the two lateral grooves. D, Caudal plane of fracture, showing both splenials, dentaries and the Meckelian canal (Can. cart. mand.). E, Rostral plane of fracture, showing the right splenio-dentary suture (Sut. spldent.).



FIGURE 2. Dorsal view of a phytosaur lower jaw (from Camp, 1930:g. 12) showing the location of the preserved fragments of the Brazilian specimen (MCN-FZB 1865).

widens, because the medial dorsal surface of the dentary is wider rostrally than caudally. The fth (e) fragment of bone is probably the most rostral of all (Fig. 1B). There is no rostral connection with fragment d, but there is a rostrocaudal trough that ts the bump described in fragment d. So, this fragment may represent a bit of the right premaxillary. It shows traces of alveoli laterally, and the ventral surface of the bone is positioned obliquely, tting the bump caudally but continuing rostrally and forming an elongate rostrocaudal depression. The same kind of striations seen on fragments a and d are present, although they are ner. There are three more bits of bone. The rst one (f) shows grooves and pits and possibly could be part of the dentary. The other two (g and h) are featureless and only show an articular plane. In the main fragment (a), there are nine alveoli on each side. They are essentially round and are lled by broken root or crown bases of teeth that are round to slightly elliptical in cross section, more rounded rostrally than caudally. The additional fragments of bone (d and e) show four (with two broken teeth preserved) and four (with no teeth preserved) more alveoli. Completing the gaps between fragments a (nine alveoli) and d (four alveoli), there is space for more ve alveoli, so there were at least 18 teeth. Most of the teeth are broken at their bases, and do not show evidence of heterodonty or any trace of carinae. Three better preserved teeth can be noted in a. The rst one (position 8, left side, the second most caudal preserved alveolus) was not erupted, and shows an acuminate tip with ne serrations rostrally and caudally. The second one (position 6, right side, third most caudal fully preserved alveolus) was in a degree of eruption later than the rst and, although its crown is broken, shows the presence of carina rostrally and caudally and is asymmetric, with the medial face less convex than the lateral one. The third (position 4, TABLE 1. Piece of bone d Length of alveoli of MCN-FZB 1865.

right side, third most rostral fully preserved alveolus) is an unerupted tooth totally covered by hard matrix (only the very tip is exposed) and is in a stage of eruption similar to the second tooth described. The alveoli of fragment e are difcult to measure accurately, but fragments d and a show progressive enlargement of the alveoli caudally (Table 1). Dornelles (1990) documented three isolated archosaur teeth (UFRGS 319-21) from the Botucara locality, comparing them to the teeth of Parasuchus hislopi, described by Chatterjee (1978), but concluding that they can only be ascribed to archosaurs. These teeth could not come from the specimen here described, because they are larger. They are asymmetrically attened transversely and bicarinate, but only have caudal serrations. Although the teeth alone may not be denitive of a phytosaur, the presence of the jaw fragment, documented here, lends support to Dornelles comparisons. IDENTIFICATION AND DISCUSSION Although these fossils are obviously part of the elongated snout of an archosaur, their identication as being from the lower jaw is not so obvious. The canal, here interpreted as that for Meckels cartilage, is lateral to the tooth roots and is continuous on both sides rostrocaudally. Some portion of an antorbital cavity would be medial to the tooth roots if the fragments were part of the upper jaw. Even if it could be a rostrum instead of a mandibular symphysis, the medial bones in external view, here interpreted as splenials, would be identied as septomaxillaries. As such, they are much too broad to be phytosaur septomaxillaries and they should not reach the palate. So, we believe that this specimen represents a mandibular symphysis, corresponding closely to the middle portion of that found in phytosaurs (Fig. 2). When put together in a continuum and after lling the gaps, the described fragments a and b result in a length of at least 18 cm. Comparison to Chinle Group phytosaur jaws in the NMMNH collection (Pseudopalatus: NMMNH P-4975, 17323, both from the Bull Canyon Formation in eastern New Mexico: Hunt, 1994) supports identication of the Brazilian fossil as a phytosaur. Points of detailed resemblance are: (1) long, parallel-sided tooth rows; (2) numerous, closely spaced, bicarinate, serrated and asymmetrical teeth with nearly round basal cross-sections; (3) thecodont implantation with interalveolar ridges; (4) long and large rostrally converging splenials that medially separate the dentaries on ventral surface; and (5) external surface of dermal bones ornamented by a diffuse pattern of parasagittally-oriented grooves and pits. Indeed, MCN-FZB 1865 displays three characters that are uniquely derived features of phytosaur lower jaws (Hungerbu hler, 1998, 2001). These are: (1) a symphyseal platform, which is the long, at, horizontal plane formed by the dorsal aspect of the mandibular symphysis; (2) an alveolar ridge and intermaxillary groove, present on fragment e; and (3) labiolingually asymmetric teeth, a feature unique to phytosaurs among archosauromorphs. The presence of these apomorphies of phytosaurs in MCN-FZB 1865 provides decisive evidence that it represents

Position #1 #2 #3 #4 #5 #1 #2 #3 #4 #5 #6 #7 #8 #9

Left 3.5 6.5 6.4 6.1 3.0 7.8 8.0 7.6 9.3 7.2

Right 6.7 6.1 6.9 7.3 7.5 8.6 9.3 9.1 4.8

a phytosaur, and not another long-snouted archosauromorph, such as a proterochampsid. This is the rst conclusive evidence of a phytosaur from the South America. Their previous absence has generally been attributed to facies differences, with the South American Triassic tetrapods supposedly coming from drier or upland habitats in which phytosaurs did not live (Buffetaut, 1993) or to paleolatitudinal (paleoclimatic) differences between Late Triassic tetrapod faunas north and south of 30 S paleolatitude (Shubin and Sues, 1991). These previous explanations of the absence of phytosaurs in Late Triassic South America can now be abandoned. Acknowledgments We thank J. Ferigolo, curator of the FZB paleontological collection, for loaning the specimen for study and information about the stratigraphic level, and L. Maciel and F. Sedor, who collected the specimen (MCN-FZB). The rst author (EEK) thanks M. C. Barberena (UFRGS) for advising his doctoral studies. Axel Hungerbu hler and Michael Parrish provided helpful reviews of the manuscript. LITERATURE CITED Arau jo, D. C., and T. D. Gonzaga. 1980. Uma nova espe cie de Jachaleria (Therapsida, Dicynodontia) do Tria ssico do Brasil. Actas del Segundo Congreso Argentino de Paleontologia y Bioestratigraa y Primer Congreso Latinoamericano de Paleontologia 1:159174. Azevedo, S. A. K. de. 1993. Os dinossauros Tria ssicos no sul do Brasil: Dados e perspectivas. Acta Geologica Leopoldensia 16:3140. 1999. Os dinossa urios Tria ssicos do sul do Brasil: Dados atualizados e novas perspectivas. Paleontologia em Destaque 14(26):57. Ba ez, A. M., C. Marsicano, and A. L. Cione. 1993. Vertebrados Mesozoicos; pp. 341348 in V. A. Ramos (ed.), Geolog a y Recursos Naturales de Mendoza. Relatorio, XII Congreso Geolo gico Argentino y II Congreso de Exploracio n de Hidrocarburos. Instituto Argentino de Petroleo, Mendoza. Bonaparte, J. F. 1978. El Mesozoico de America del Sur y sus tetrapodos. Opera Lilloana 26:1596. , J. Ferigolo, and A. M. Ribeiro. 1999. A new early Late Triassic saurischian dinosaur from Rio Grande do Sul State, Brazil. National Science Museum Monographs Tokyo 15:89109. Buffetaut, E. 1993. Phytosaurs in time and space. Paleontologia Lombarda (Nuova Serie) 2:3944. Camp, C. L. 1930. A study of the phytosaurs with description of new material from western North America. Memoirs of the University of California 10:1174. Charig, A. J., and O. A. Reig. 1970. The classication of the Proterosuchia. Biological Journal of the Linnean Society 2:125171. Chatterjee, S. 1978. A primitive parasuchid (phytosaur) reptile from the Upper Triassic Maleri Formation of India. Palaeontology 21:83 127. Dornelles, J. E. F. 1990. Registro sobre a ocorre ncia de dentes de um arcossa urio para a formac a o Caturrita, Tria ssico Superior do Rio Grande do Sul, Brasil. Cie ncia e Natura Santa Maria 12:99101.


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