A Semionotid Fish from the Crato Formation (Aptian, Lower Cretaceous) of Brazil: Palaeoecological Implications
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ORYCTOS,Vol. 1 :37 - 42 Octobre1998 A SEMIONOTIDFISH FROMTHE CRATOFORMATION (APTIAN, LOWER CRETACEOUS)OF BRAZIL: PALAEOECOLOGICALIMPLICATIONS Paulo M. BRITO1 , David M. MARTILL, and SvtvieWENZ3 tDepartmento Biologia Animal e Vegetal, Universidade do Estado do Rio de Janeiro, R. J. Brazil & Laboratoire Ichthyologie, Museum national d'Histoire naturelle, Paris, France. '?Departmentof Geology, University of Portsmouth, Portsmouth,pOl 3eL, UK. 3 Laboratoire de Paléontologie- URA 12 CNRS, 8 Rue Buffon, F-75005 Paris, France. Abstract : A partial example of an articulated fish cf. Araripelepidotes sp. from the Nova Olinda Member of the Crato Formation (Aptian, Lower Cretaceous,Araripe Basin, Bra\zil) is the first record of this taxon from this for- mation. Fishes other than Dastilbe spp. are extremely rare in the erato Formation and we consider the implications in terms of the overall ichthyofauna diversity and palaeoecology. Key words: Pisces,Lower Cretaceous,Brazil Un poissonsemionotidé de la Formation Crato (Aptien, Crêtacéinférieur) du N. E. du Brésil: implications paléoécologiques Résumé : Un exemplaire incomplet, articulé, d'un poisson semionotidé,provenant du Membre Nova Olinda de la Formation Crato (Aptien du Bassin d'Araripe, Brésil) est identifié comme étant un cf . Araripelepidotes sp. C'est la première fois que ce type de poisson est signalé dans ce niveau. A l'exception de Dastilbe spp. les poissons sont extrêmementrares dansla Formation Crato. Le problème de la diversité de I'ichthyofaune et les implications paléo- écologiquesde cette découvertesont abordées.(traduit par la rédaction). Mots clés: Pisces, Crétacé inférieu4 Brésil INTRODUCTION thoughts on the palaeoenvironment, especially with regard to the palaeosalinity. Although the Nova The Crato Formation of the Araripe Basin, north Olinda Member of the Crato Formation (Aptian, east Brazll is one of the most important Meso zoic Lower Cretaceous,Araripe Basin, north east Brazll) fossil conservation lagerstâtte, and is particularly is well known for its exceptional preservation and well-known for yielding abundant, diverse and well abundanceof fossils, the ichthyofauna is of very low preservedinsects and plants (Martill, 1993).As of yet, diversity (only two genera so far being recorded) and no detailed palaeoecological analysis of the forma- only the gonorhynchiform teleost Dastilbe elongatus tion has been published and questions remain as to (Silva Santos, 1947) occurs abundantly.On the other the exact nature of the palaeoenvironment. The pur- hand, winged insects are extremely abundant and pose of this paper is to document the first discovery include a wide range of insect orders. Other verte- of a semionotid fish; cf. Araripelepidotes sp., from brates such as birds, pterosaurs, crocodiles, lizards the Crato Formation and consider the palaeoecologi- and frogs have all been reported (Martill & Filgueira, cal implications of this fossil in the light of current 1994; Maisey, I99I). Perhaps remarkably, if one 37 ORYCTOS,VOL.1,1998 excludesDastilbe spp. pterosaursappear to be more THE NEW SPECIMEN abundantthan fish (seeMartill & Frey, this volume). Very rare examples of the ichthyodectiform fish The presentstudy is basedon a single specimen Cladocyclussp. from the Crato Formation were first (PMB.VP-I44) housedin the Departmentof Biology reportedby Wenzand Campos (1985) from Barbalha, of the State University of Rio de Janeiro (UERJ). Cearâ.More recently Maisey (1996) claimed that Although the new specimenis incomplete,it is an specimensof Cladocyclusfrom the Crato limestones articulatedexample of a semionotidfish on a slab of representeda freshwater speciesof this taxon. We creamcoloured limestone typical of the Nova Olinda discussthis considerationbelow. Memberof the CratoFormation (Figs . l, 2).It lacks the skull and anterior trunk region, thus preventing accurategeneric and specificidentification. SYSTEMATICS ClassOsteichthyes Subclass Actinopterygii Figure 1. cf . ' Araripelepidotes sp. from the Nova Olinda InfraclassNeopterygii Member of the Crato Formation (Aptian, Lower Cretaceous) Order Semionotiformes of the Araripe Basin, n.e. Brazll. Scale bar in centimetres. entire Family Semionotidae Specimennumber PMB.VP-I44. Photographshowing specimen on slab of typical Crato limestone. Note that the Genuscf . AraripelepidotesSilva Santos,1985 skull and anterodorsal part of the trunk appears to have (seealso Silva Santos, 1990) decayedaway prior to preservation. 1CICM 38 BRITO, MARTILL & WENZ- SEMIONOTID FROM BRAZIL As preserved the specimen is approximately 250 caudal fin is slightly forked (Fig.2). mm in length, and represents a fish with an original The body is subcylindrical with a moderate ele- estimated length of between 240 mm and 300 mm vation anterior to the dorsal fin. Dorsal and anal fins (this estimate obtained by comparing the length bet- are situated in the posterior part of the body. All the ween base of the ventral lobe of the caudal fin and fins are preceded by strongly developed fulcra. The base of pectoral fins with that reconstructed by body is coveredby rhombic, smooth, isometric scales Maisey (I99I, p. 118) for Araripelepidotes). The covered by a thick layer of ganoin. The scales differ body measuresapproximately 180 mm from the pec- depending on their position in the body. The most toral girdle to the caudal pedicle. The anomaly bet- dorsal anterior scales are larger with serrated poste- ween the actual length of the specimen and the esti- rior edges; the mid and posterior scales are diamond mated length of the complete fish is due to the tapho- shaped and lack serration. nomic forward flexure of the right pectoral fin. The It is difficult to place the Crato specimen preci- anteroventral region of the trunk shows some disrup- sely in a taxonomic position, as it lacks the skull tion, and the missing skull is due to post mortem loss bones and the body shows only the most generalised during drifting. The paired pectoral fins are present, characters' of the Semionotidae. Semionotids are as is the anal fin and parts of the dorsal fin. The cau- common in the Mesozoic rift and pre-rift basins in dal fin is entire, although the dorsal and ventral lobes the north-east of BrazII, and are known by at least 4 have "closed" slightly to give the caudal region a nar- species of Lepidotes, and one species of row appearance.A slight taper of the distal portions Araripelepidotes in the SantanaFormation. of the caudal fin suggest that in this specimen the 39 ORYCTOS,VOL.1,1998 In the Araripe basin, Lepidotes sp. is known only the anatomy of Dastilbe from the Crato Formation by disarticulated material from the Missao Velha and considers that two species may occur; a small Formation. From the Romualdo Member of the species with a dorsal fin ray count of 10 and larger SantanaFormation is known Araripelepidotes temnu- examples with a fin ray count of 13. In the Nova rus (Agassiz, 1841), a smaller and more slender Olinda Member specimens of both large and small semionotid, as well as a possible distinct semionotid Dastilbe spp. occur in abundance;smaller individuals taxon, âS yet undescribed (Brito &. Wenz in prep.). in super abundance, whereas examples of Recently, Wenz & Brito (1992) described two primi- Cladocyclus ate extremely tate, and usually represent tive gar fishes; Obaichthys decoratus and O? laevis, small individuals (size range 150 to 300 mm) compa- also from the Romualdo Member. red with examples of Cladocyclus from the The new Crato specimen can be distinguished Romualdo Member of the Santana Formation where from all other Lepidotes known in Brazllby its body lengths of over 1 m are recorded (Maisey, I99I: p. shape (subcylindrical), the absenceof a pronounced 190). The new specimen described here with pro- hump in front of the dorsal fin and the dimensions of bable affinities to Araripelepidotes indicates that its body. other fish species were entering the Crato Formation It can be distinguished from Obaichthys by the water body, but only very rarely. presence of large scales in the front part of the body The low diversity of the Crato Formation ich- (Obaichthys spp. have similar sized scales throu- thyofauna is in marked contrast to the high diversity ghout), and the absenceof opisthocoelous vertebrae. Santana Formation ichthyofauna which has to date In addition, the vertebrae of the new specimen are not yielded some twenty genera (Maisey I99I; Martill fully ossified, whereas in gars the vertebrae are well 1993). Of those recorded from the Crato Formation, ossified throughout. Araripelepidotes and Cladocyclus are also well Although no single synapomorphycan be poin- known from the Romualdo Member of the Santana ted out, we can separatethe Crato specimen from all Formation, but the gonorhynchiform Dastilbe elon- other Brazllian semionotifoffns sensu Olson and gatus has yet to be recorded. This anom aly has McCune I99I (theseauthors combine Semionotidae, important implications for any reconstruction of the Macrosemiidae and Lepisosteidae in the palaeoenvironment of the Crato Formation water Semionotiformes), except Araripelepidotes . body. The Santana Formation ichthyofauna probably However, the absenceof the skull makes any identi- represents a mixed salinity assemblage with both fication tenuous. The new specimen is certainly more marine, freshwater and brackish forms. No certain similar to Araripelepidotes (e.g. shape of the body, marine fossils (ammonites, brachiopods, etc.) have shape and dimension of the scales) than to any other been found commonly