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An Updated Review of the Fish Faunas from the Crato and Santana Formations in Brazil, a Close Relationship to the Tethys Fauna

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An Updated Review of the Fish Faunas from the Crato and Santana Formations in Brazil, a Close Relationship to the Tethys Fauna Bull. Kitakyushu Mus. Nat. Hist. Hum. Hist., Ser. A, 9: 107–136, March 31, 2011 An updated review of the fish faunas from the Crato and Santana formations in Brazil, a close relationship to the Tethys fauna Paulo M. BRITO 1 and Yoshitaka YABUMOTO2 1Departamento de Biologia Animal e Vegetal, Universidade do Estado do Rio de Janeiro, rua São Francisco Xavier 524, Rio de Janeiro, 20559-900, Brazil E-mail: [email protected] 2Department of Natural History, Kitakyushu Museum of Natural History and Human History, 2-4-1 Higashida, Yahatahigashi-ku, Kitakyushu, Fukuoka, 805-0071, Japan E-mail: [email protected] (Received December 14, 2010; accepted March 1, 2011) ABSTRACT — The Lower Cretaceous Crato and Santana formations have provided one of the richest Mesozoic fish faunas from South America. An updated review of this ichthyofauna, comprising, 28 nominal species, is presented here. Contrary to the previous idea that the Araripe Basin had an endemic fauna related to the opening of the South Atlantic Ocean, it is now accepted that this fauna is instead related to that of the Tethys. A marine connection with the Araripe Basin is indicated by the presence of species closely related to those of other assemblages occurring in the western part of the Tethys. However, the absence of marine invertebrates suggests non-marine conditions for this basin, with only intermittent connections to the epicontinental seaway. Some of the fishes found in the Crato Formation are juveniles of the species found in the Santana Formation, suggesting several important paleoecological implications related to the reproduction of these fishes and using there as a nursery. KEY WORDS: Paleoichthyofauna, Santana Formation, Crato Formation, Lower Cretaceous, Gondwana, Brazil, Tethys fauna. INTRODUCTION by SPIX and MARTIUS (1823) as Rhacolepis AGAssIZ (1841). Subsequently, knowledge of the ichthyofauna was further The fossil-bearing deposits of the Lower Cretaceous (Aptian/ enriched by the works of AGAssIZ (1833–1844), WOODWARD Albian) Crato and Santana formations of the Araripe Basin (1887, 1890, 1895, 1901, 1908), JORDAN and BRANNER (1908), (Northeastern Brazil) have yielded numerous vertebrates, JORDAN (1919, 1921), D’ERASMO (1938), DUNKLE (1940), and including fishes, frogs, turtles, lizards, pterosaurs, dinosaurs, SANTOS (1945, 1947, 1950, 1958, 1960). However it was not and bird feathers (Figs. 1–3). The fishes, which form the until the publication of SANTOS and VALENÇA (1968) that the dominant element of this biota, are exceptionally abundant and first review of the Araripe ichthyofauna became available. are famous worldwide for their quantity and excellent state Since then, numerous authors have described a quantity of of preservation, being frequently articulated and sometimes new taxa (CAMPOS and WENZ, 1982; SANTOS, 1985a; MAISEY, three-dimensionally preserved. In some cases, there is even the 1986; WENZ and KELLNER, 1986; WENZ, 1989; BRITO and preservation of phosphatic soft tissues, including muscle fibers, FERREIRA, 1989), developed descriptions of known fossils, gill filaments, and stomach contents (MARTILL, 1988; WILby and/or discussed the phylogenetic affinities of the various and MARTILL, 1992). components of the fauna (e.g., TAVERNE, 1974, 1976; FOREY, The first account of fossil fishes from Araripe appears in 1977, 1998; WENZ, 1977; PATTERSON and ROSEN, 1977; the atlas of the “Reisen Brasilien” expedition between 1817 OLIVEIRA, 1978; PATTERSON, 1984; SANTOS, 1985b, 1990a, b, and 1820 (SPIX and MARTIUS, 1823). This publication was soon 1994a, b, 1995; BRITO, 1988, 1992b, 1999; BRITO and WENZ, followed by those of GARDINER (1841) and AGAssIZ (1841), who 1990; MAISEY and BLUM, 1990; BRITO and MARTILL, 1999; drew up a list of fishes and identified one of the forms shown BRITO and MEUNIER, 2000; BRITO et al., 1998, 2000, 2010; 108 Paulo M. BRITO and Yoshitaka YABUMOTO Fig. 1. A, Schematic stratigraphic log for the Araripe Basin (after MARILL, 2007); B, Simplified geological map of the Chapada do Araripe (after MARTILL et al., 2007); C, Paleogeographic roconstruction of the Early Cretaceous with the location of the Araripe Basin. BRITO and AMARAL, 2008; DAVIS and MARTILL, 1999; LEAL and The Crato and Santana formations have provided what BRITO, 2004; FOREY and MAISEY, 2010). are clearly the richest Lower Cretaceous fish faunas in South MAISEY (1991) presented the most comprehensive account America (SANTOS and VALENÇA, 1968; WENZ and BRITO, 1990; of the Araripe fish fauna published until then. However, as MAISEY, 1991; MARTILL, 1993). This ichthyofauna comprises at knowledge of this assemblage has continued to grow (e.g., least 28 nominal species. WENZ and BRITO, 1992; MAISEY, 1993; BRITO, 2000; YABUMOTO, The valid species of the Crato and Santana formations 2002; BRITO and GALLO, 2003; FIGUEIREDO and GALLO, 2004; are all briefly reviewed below, and the relative phylogenetic BRITO et al., 2008) we present in this paper an updated review relationships among these species, as currently hypothesized, of the fish faunas from the Crato and Santana formations. are illustrated in Appendices 1 and 2. Fish faunas from the Crato and Santana formations 109 Fig. 2. A, A typical outcrop, with limestone nodules, of the Santana Formation near the town of Nova Olinda, State of Ceará; B, Pedra Branca gypsum mine, Nova Olinda Municipality, State of Ceará. All the sediments above the gypsum are part of the Santana Formation. Fig. 3. A, Crato Formation quarries on the road between the towns of Nova Olinda and Santana do Cariri, State of Ceará; B, A stone cutting in one of the limestone quarries (Crato Formation) near the town of Nova Olinda, State of Ceará. 110 Paulo M. BRITO and Yoshitaka YABUMOTO THE ICHTHYOFAUNA South America and Cuba. The Western Gondwana semionotids are the most characteristic Mesozoic fishes, especially in the Hybodontidae northeastern basins of Brazil where they have a temporal There is one species representing the family Hybodontidae range from the Late Jurassic to Late Cretaceous (comprising in the Araripe Basin, Tribodus limae BRITO and FERREIRA, 1989 nine nominal species; BRITO and GALLO, 2003; GALLO and (Fig. 4). This species occurs in the Santana Formation and is BRITO, 2004). Two species are known in the Araripe Basin, unknown in the Crato Formation. It is a relatively frequent each occurring in both the Crato and Santana formations: component of the fauna. Araripelepidotes temnurus (AGAssIZ, 1841), and Lepidotes This species is known to reach over 600 mm in total length. wenzae BRITO and GALLO, 2003 (Figs. 6 and 7). It differs from most other hybodontids (except Acrodus and Araripelepidotes temnurus is relatively common in the Asteracanthus) in having a crushing dentition (BRITO, 1992a) Santana Formation, although it is very rare in the Crato and a hyostylic jaw suspension (MAISEY and CARVALHO, 1997). Formation. This taxon, considered for a long time as a species Recently, an anatomical revision of the pectoral girdle of of Lepidotes, was removed to the new genus Araripelepidotes Tribodus as well as a revision of the braincase, based on CT by SANTOS (1990a). This species is known to reach about scan had been published (LANE and MAISEY, 2009; LANE, 2010). 400 mm in total length, and is characterized by a very weak, edentulous lower jaw composed of a single element; Rajiformes family incertae sedis reduction of the coronoid process; and separation of the There is one guitarfish species from the Santana Formation dermopterotic and the frontal by the dermosphenotic. Thus far, of the Araripe Basin, Iansan beurleni (SANTOS, 1968) (Fig. 5), Araripelepidotes appears to be endemic to the Araripe Basin. which reaches close to 900 mm in total length (for a revision Lepidotes wenzae is a relatively small-sized species, reach- of the anatomical patterns of this taxon see BRITO and SÉRRET, ing about 250 mm standard length. This species, not very 1996). common in the Santana Formation (on the basis of known Iansan is relatively common in the Santana fish fauna, but specimens), has a moderate pre-dorsal elevation, scales so far has not been reported from the Crato Formation or any lacking ornamentation, and a moderate crushing dentition. other Lower Cretaceous western Gondwanan formations. The Lepidotes wenzae is a rarely occurring component of the fish phylogenetic relationships of I. beurleni to other fossil and fauna of the Crato Formation, and is represented only by living guitarfishes are still unclear. juvenile specimens (BRITO, 2007). Detailed comparisons of the skull morphology between the specimens from the Santana Semionotidae and Crato formations are needed to resolve the validity (or Semionotidae is a geographically widespread family synonymy) of this species. For now, we consider Lepidotes with a chronological range from the Middle Triassic to the from the Araripe Basin to represent a unique species. Late Cretaceous. Semionotids are found in both marine and continental strata from Europe, North America, Asia, Africa, Fig. 4. Tribodus limae BRITO and FERREIRA, 1989, MNRJ 105 (Museu Nacional do Rio de Janeiro), from the Santana Formation. Fish faunas from the Crato and Santana formations 111 Fig. 5. Iansan beurleni (SANTOS, 1968) from the Santana Formation. A, MB. f. 12425 (Berlin Museum); B, UERJ-PMB 20 (Universidade do Estado do Rio de Janeiro); C. UERJ-PMB 18; D, UERJ-PMB 19. Scales of B to D are 50 mm. 112 Paulo M. BRITO and Yoshitaka YABUMOTO Fig. 6. Araripelepidotes temnurus (AGAssIZ, 1841), UERJ-PMB 63 (Universidade do Estado do Rio de Janeiro), from the Santana Formation. Fig. 7. Lepidotes wenzae BRITO and GALLO, 2003, holotype, MNHN-BCE 387 (Muséum national d'Histoire naturelle, Paris), from the Santana Formation. Fish faunas from the Crato and Santana formations 113 Lepisosteiformes (Obaichthyidae) predorsal length of fish is 75% or more of standard length; The family Lepisosteidae is well known nowadays by seven the palatal complex has laterolly sliding articulation between nominal species belonging to two genera, Lepisosteus and the metapterygoid and the basipterygoid process of the Atractosteus (WILEY, 1976; NELSON, 1994).
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