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An updated review of the faunas from the Crato and Santana formations in Brazil, a close relationship to the Tethys fauna

Article · January 2011

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Paulo M. Brito Yoshitaka Yabumoto Rio de Janeiro State University Kitakyushu Museum of Natural History and …

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All in-text references underlined in blue are linked to publications on ResearchGate, Available from: Yoshitaka Yabumoto letting you access and read them immediately. Retrieved on: 27 August 2016 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 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 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 , is presented here. Contrary to the previous idea that the 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 found in the 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, , Brazil, Tethys fauna.

INTRODUCTION by Spix and Martius (1823) as Agassiz (1841). Subsequently, knowledge of the ichthyofauna was further The -bearing deposits of the Lower Cretaceous (/ enriched by the works of Agassiz (1833–1844), Woodward ) 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 , 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 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 to (comprising in the Araripe Basin, 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 ) 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 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 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 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). Extant lepisosteids braincase; the presence of a “tongue bone” or basihyal tooth are distributed in eastern North America, Central America and plate consisting of a mosaic of bony pates; the presence of the Caribbean. They are basically freshwater fishes, but may opisthocoelous vertebral centra), it also retains some characters occasionally occur in brackish water. Only one extant species, considered primitive for neopterygians, such as the presence Atractosteus tristoechus, invades marine environments in the of a free maxilla; two post-orbital bones; and paleoniscoid- vicinity of Cuba and the Isle of Pines. type scales (Wenz and Brito, 1992, 1996; Brito et al., 2000; Lepisosteiforms have a long geological history (dating as Grande, 2010). The differences between the two described far back as the Lower Cretaceous; e.g., Casier, 1961; Wenz species mainly concern the ornamentation of the dermal bones and Brito, 1992, 1996; Brito et al., 2000). They also have and scales, shape of scales, and dimensions and proportions of a wide paleogeographic distribution, mainly during the Late the body and cranial bones. Cretaceous and Tertiary in Africa (e.g., Arambourg and Two other species of obaichthyids, Obaichthys africanus Joleaud, 1943; Gayet et al., 1988; Werner, 1994; Cavin and Dentilepisosteus ?kemkemensis, were recently described and Brito, 2001), South America (e.g., Santos, 1984; Cione, by Grande (2010) from the ?Late Cretaceous Kem Kem beds 1987; Gayet and Brito, 1989; Gayet, 1991; Bertini et al., of Morocco. A detailed taxonomic and phylogenetic discussion 1993), Madagascar (Gottfried and Krause, 1998); India (e.g., regarding the validity of the family Obaichthydae is discussed Woodward, 1890, 1908; Wiley, 1976; Jain and Sahni, 1983; by Grande (2010). We agree with Grande (2010) that the Rana and Kumar, 1990), Europe (e.g., Jonet, 1981; Buffetaut Obaichthydae form a monophyletic group. et al., 1996; Cavin et al., 1996; Sigé et al., 1997), and North America (see Wiley, 1976; Russell, 1988; for a complete Amiidae listing). Currently the amiids are represented by a single extant There are two lepisosteiforms in the Araripe Basin. species, Amia calva, found in lakes and rivers of Eastern These taxa, placed by Wenz and Brito (1992) in the genus North America. However, this family has a long temporal Obaichthys, were recently reviewed by Grande (2010), who distribution, being known since the Upper Jurassic. In the proposed the new family Obaichthydae as well as a second western part of Gondwana, amiids are represented by the genus (Dentilepisosteus). Subfamily Vidalamiinae, (Grande and Bemis, 1998). Obaichthys decoratus Wenz and Brito, 1992 and Denti- There are two species of amiids in the Araripe Basin, lepisosteus laevis (Wenz and Brito, 1992) (Figs. 8, 9) are the Calamopleurus cylindricus Agassiz, 1841 and Cratoamia oldest known lepisosteiforms. Both species are very rare and gondwanica Brito, Yabumoto and Grande, 2008. Cala- are thus far known only from the Santana Formation. Recently, mopleurus cylindricus is known from both the Crato and another obaichthyid was also recorded in the Crato Formation Santana formations (Fig. 11), but Cratoamia (Fig. 12) is (Fig. 10), based on a unique specimen that seems to be more currently restricted to the Crato Formation. closely related to D. laevis than to O. decoratus. Calamopleurus cylindricus is one of the best-studied amiids Although Obaichthyidae present 14 to 16 synapomorphies (e.g., Santos, 1960; Grande and Bemis, 1998). This species shared with Lepisosteidae (e.g., nasal process of premaxilla has a maximum known size of about 1400 mm total length, forms much of the ornamented drmal roof in the snout region; and is relatively common in the Santana Formation, where it

Fig. 8. Obaichthys decoratus Wenz and Brito, 1992, holotype, DGM 1336 P (Departmento Nacional de Produção Mineral, Brazil), from the Santana Formation. Approximately 600 mm SL. 114 Paulo M. Brito and Yoshitaka Yabumoto

Fig. 9. Dentilepisosteus laevis (Wenz and Brito, 1992), MPSC 901 (Museu de Paleontologia de Santana do Cariri, Ceará, Brazil), from the Santana Formation, 365 mm SL.

Fig. 10. Obaichthid fossil, UERJ-PMB 233 (Universidade do Estado do Rio de Janeiro), from the Crato Formation, 170 mm SL. Fish faunas from the Crato and Santana formations 115 is represented by the type species C. cylindricus (= Enneles dermosphenotic not tightly sutured to the other bones of the audax). The genus Calamopleurus was diagnosed by Grande braincase; and posterior margin of the gular deeply scalloped and Bemis (1998) on the basis of the following characters: with a series of sharp points and concavities. posterior margin of caudal fin with an almost vertical outline; Of the three nominal species of Calamopleurus, C. opercular process of the hyomandibular extremely elongate; cylindricus is the best known due to its relative abundance in teeth of the coronoid bones distributed in a single marginal the Santana Formation. Two other species (C. africanus Forey row; teeth of the vomer arranged in a single anterior row, with and Grande, 1998, from the Late Cretaceous Kem Kem beds of one or more teeth in a longitudinal row perpendicular to the Morocco; and C. mawsoni (Woodward, 1902) from the Early marginal row; dermopterotic with a bundle of elongate splint- Cretaceous Recôncavo Basin) were described based on unique like bones (dermopterotic ribs) fused to its posterior end; disarticulated specimens. Calamopleurus africanus is known

Fig. 11. Calamopleurus cylindricus Agassiz, 1841, KMNH VP 100,236 (Kitakyushu Museum of Natural History and Human History), from the Santana Formation, 1240 mm SL.

Fig. 12. Cratoamia gondwanica Brito, Yabumoto and Grande, 2008, KMNH VP 100,260 (Kitakyushu Museum of Natural History and Human History), from the Crato Formation. 116 Paulo M. Brito and Yoshitaka Yabumoto only from a skull and differs from the type species primarily in fin. The pectoral fin is very well developed, with a convex the shape of its gular plate, which carries a median ridge on its margin. The caudal fin is deeply forked, with the upper lobe ventral surface and bears 10 small projections on the posterior larger than the lower lobe. margin (see Forey and Grande, 1998). Calamopleurus Another species of Placidichthys, P. tucanensis Brito and mawsoni is known only from the type material, consisting of Alvarado-Ortega, 2008, was recently described from the an articulated skeleton, lacking the cranial and pectoral girdle Aptian Marizal Formation of the Tucano Bahia. Placidichthys regions. This species differs from C. cylindricus in the number tucanensis is clearly distinguishable from the species in the of dorsal radials and higher number of preural vertebrae (see Araripe Basin by the absence of an anal fin; the lower number Grande and Bemis, 1998). of flank scales in the caudal region; and the much more slender Cratoamia gondwanica is a medium-sized amiid of body shape. approximately 700 mm total length. This species is easily characterized by the presence of a long with between Oshuniidae 27 and 30 rays, as well as by the weak ornamentation of its The Family Oshuniidae was recently proposed by Grande dermal bones. and Bemis (1998), and is represented by only one genus and one nominal species (Oshunia brevis Wenz and Kellner, 1986; Ophiopsidae Fig. 14; from the Santana Formation), although Maisey (1991) Ophiopsids are a typical Jurassic/Cretaceous group found proposed that the genus may contain additional species. in Europe, North America, Africa and South America. In Oshunia brevis is easily characterized by its high, trian- Brazil, ophiopsids were previously recorded by Santos and gularly shaped dorsal fin and body covering of typical amioid Valença (1968), for a specimen that unfortunately could not scales. Recently, a taxon previously considered a member of be diagnosed even into a familial level, identified as Ophiopsis Ionoscopidae, Quetzalichthys perrilliatae Alvarado-Ortega cretaceous. For this reason, this species is considered a Nomen and Espinosa-Arrubarrena, 2008, was described from the nudum. Albian Tlayúa Quarry of Mexico and considered to represent There is one species representing the ophiopsids in the the sister-group of Oshunia on the basis of two morphological Araripe Basin, Placidichthys bidorsalis Brito, 2000 (Fig. 13). characters (presence of paired of longitudinal fossae on the Placidichthys is a small ophiopsid known to reach 80 mm total sides of all vertebral centra; and the presence of only three length, and several young juveniles are also known, especially urodermals). in the Crato Formation. Placidichthys is easily diagnosed by its very elongate body. It differs from other members of the family Pycnodontidae in having a long dorsal fin subdivided into two parts. The There are two pycnodontids in the Araripe Basin, known deepest part of the body is between the head and the anterior exclusively from the Santana Formation: Neoproscinetes part of the dorsal fin, with body depth decreasing strongly penalva (Santos, 1970), and Iemanja palma Wenz, 1989. toward the posterior part of the dorsal fin. The body is covered Neoproscinetes penalva (Fig. 15) is a relatively common by small, thin, diamond-shaped ganoid scales distributed from fish, reaching about 350 mm total length and having a nearly the pectoral girdle to the base of the axial lobe of the caudal circular shape in lateral outline. This fish presents a dentition

Fig. 13. Placidichthys bidorsalis Brito, 2000, UERJ-PMB 300 (Universidade do Estado do Rio de Janeiro), from the Crato Formation, 95 mm SL. Fish faunas from the Crato and Santana formations 117

Fig. 14. Oshunia brevis Wenz and Kellner (1986), UERJ-PMB 394 (Universidade do Estado do Rio de Janeiro), from the Santana Formation, 210 mm SL.

Fig. 15. Neoproscinetes penalva (Santos,1970), UERJ-PMB 246 (Universidade do Estado do Rio de Janeiro), from the Santana Formation. 118 Paulo M. Brito and Yoshitaka Yabumoto

Fig. 16. Iemanja palma Wenz, 1989 from the Santana Formation. A, MNHN 166b (Muséum national d'Histoire naturelle, Paris) (Holotype); B, NSM PV-20384 (National Museum of Nature and Science, Tokyo). Fish faunas from the Crato and Santana formations 119 comprising a pavement of large, low, elongate, dome-like Ariza and Wenz, 2002) suggested this taxon should be placed teeth, with the alternation of one large and two small teeth on together with other members of the pycnodontids. the main vomerine tooth row. Eight or nine teeth are present in the main vomerine tooth row. Aspidorhynchidae Iemanja palma (Fig. 16) is still a rare component within There are two aspidorhynchids in the Araripe Basin: the fauna of the Araripe Basin. It is known from specimens comptoni (Agassiz, 1841) (Fig. 17) from the Santana of approximately 600 mm total length. This species presents, Formation, and a new species of (Fig. 18) among other characters, a large supraoccipital crest; fusion of from the Crato Formation. Vinctifer and Belonostomus are the first arcocentra with the skull; an accentuated prognatism; two of four genera comprising the Aspidorhynchidae, a vomerine teeth organized in rows only in the posterior portion widely distributed Mesozoic family (the other two genera are of the vomer; vomerine teeth of the principal row with a and Richmondichthys; the latter is probably reniform shape; presence of a urodermal; scales restricted a synonym of Vinctifer) of highly elongate predatory fishes. to the antero-ventral part of the body; scales of the dorsal The aspidorhynchids are easily recognized by the presence margin and the ventral keel carrying large spines; as well as of a long rostrum formed by the premaxillae; presence of a the ventral lateral line scales passing in the first ventral scales. predentary on the lower jaw; deep elongate flank scales; and Iemanja palma was originally considered a Gyrodontid, but posteriorly located dorsal and anal fins. recently phylogenetc analysis (e.g., Nursall, 1996; Poyato- Vinctifer is one of the most common species in the Santana

Fig. 17. Vinctifer comptoni (Agassiz, 1841), FMNH 10380 (Field Museum of Natural History), from the Santana Formation.

Fig. 18. Belonostomus sp., MPSC P055 (Museu de Paleontologia de Santana do Cariri, Ceará, Brazil), from the Crato Formation. 120 Paulo M. Brito and Yoshitaka Yabumoto

Formation, where it is known from specimens of between 50 remains unclear (see Cavin, 1995). Pachyrhizodontoidei and 900 mm total length (Brito, 1997). Vinctifer comptoni is a typical Mesozoic suborder, considered to represent a is a typical southern Tethyan genus, also known in other monophyletic group that includes , Rhacolepis, northeastern Brazilian basins (e.g., Parnaíba, Sergipe-Alagoas), , , Elopopsis, and Michin (Cavin, as well as in the Aptian-Albian of Venezuela, Colombia, 2001; Alvarado-Ortega et al., 2008). Mexico and Antarctica (Moody and Maisey, 1994; Schultze Cope (1872) erected the family Pachyrhizodontidae for and Stöhr, 1996; Applegate, 1996; Brito, 1997) and probably Pachyrhizodus and related forms. Forey (1977) reviewed also the Neocomian of Rio Muni, Equatorial Guinea (Taverne, this group, creating the suborder Pachyrhyzodontoidei 1969). Two nominal species are recognised: Vinctifer including two families, Notolepidae (for Notelops) and comptoni (the type species), and Vinctifer longirostris, known Pachyrhizodontidae (for Pachyrhizodus, Rhacolepis, and only from the Tucano Basin (also in northeastern Brazil). Elopopsis). Later, other genera such as Platnix, Greenwoodella, Aspidorhynchids are very rare in the Crato Formation, and Goulmimichthys, and Tingitanichthys were also included in this until now were known only from a single specimen, attributed suborder. to Belonostomus sp. This is a medium-sized Aspidorhynchid, Two nominal species of pachyrhizodontoids are known incompletely preserved, which was previously cited as V. in the Araripe Basin, exclusively in the Santana Formation: longirostris (Brito, 1997). However, further preparation of the Notelops brama Agassiz, 1841 (Fig. 20), and Rhacolepis specimen revealed several diagnostic features of the genus (e.g., buccalis Agassiz, 1841 (Fig. 21). elongated predentary and presence of a posterior supramaxilla). Notelops and Rhacolepis are very similar fishes, and may Within the Aspidorhynchidae, the genus Belonostomus is occasionally be confused. Both taxa have a fusiform body with considered to have the largest temporal range, being known a pointed snout. The principal differences between these taxa from the to the Late Cretaceous. This genus has are the presence of three prominent bones forming the posterior a widespread paleobiogeographical distribution, occurring in border of the eye in Notelops (dermosphenotic plus two marine strata from Europe, the Middle East, North America, infraorbital plates), versus four in Rhacolepis (dermosphenotic South America, and Africa. plus three infraorbital plates). Other differences are related to the posterior extent of the infraorbitals (just reaching Cladocyclidae the preopercle in Notelops vs. overlying the preopercle in There is one species in the Crato and Santana formations Rhacolepis) and the shape of the pterotic (without posterior representing the ichthyodectiform family Cladocyclidae, spine in Notelops vs. reduced to a short spine in Rhacolepis). gardneri Agassiz, 1841 (Fig. 19). This species, For a detailed description and diagnosis, see Maisey and Blum which is very abundant in the Santana Formation, occurs in the (1990) and Maisey (1991). Araripe Basin but is also reported from other Brazilian north- A taxonomic revision of Notelops is necessary, as this genus eastern basins such as Parnaíba and Sergipe-Alagoas (Leal presents different morphotypes (see Maisey, 1991). Notelops and Brito, 2004). Cladocyclus gardineri can attain a large size reaches approximately 600 mm total length. Rhacolepis is one (over 1000 mm standard length). It is easily recognised by of the most abundant fish in the Santana Formation, and is its slender and laterally compressed body; large and oblique commonly found in an excellent state of preservation, reaching mouth opening upwards with a single series of conical, slender, 250 mm total length. pointed teeth; large orbital opening; posterior position of the dorsal and anal fins; prominent cycloid scales; and deeply Albuliformes forked caudal fin M( aisey, 1991; Brito, 2007). The order Albuliformes is an extant group of marine Agassiz (1841) described Cladocyclus gardneri on the elopomorphs with low species diversity. The relationships basis of specimens collected from the Santana Formation by of the elopomorph clade to other basal teleostean groups is George Gardner. A century later, Santos (1950) described still debated, and their intrarelationships are uncertain. The a second species from the type locality, which he named C. best known albuliform fishes are the modern Albula and ferus. Subsequent authors attempted to distinguish between Pterothrissus. these two nominal species, until Patterson and Rosen in their At least four albuliforms are found in the Araripe Basin, 1977 revision of the proposed that the two exclusively in the Santana Formation: Brannerion latum species should be synonymized. (Agassiz, 1841); B. vestitum (Jordan and Branner, 1908); Paraelops cearensis Santos, 1971; and Bullichthys santanensis Pachyrhizodontoidei (Pachyrhizodontidae and Note- Mayrinck, Brito and Otero, 2010 (Figs. 22–25). lopidae) Brannerion is a deep-bodied fish, with a long anal fin that Although pachyrhizodontoids are generally placed within originates beneath the posterior level of the dorsal fin. Other the elopomorphs, their phylogenetic position among Teleostei diagnostic characters are large eyes and villiform teeth on Fish faunas from the Crato and Santana formations 121

Fig. 19. Cladocyclus gardneri Agassiz, 1841. A, KMNH VP 100,270 (Kitakyushu Museum of Natural History and Human History), from the Santana Formation; B, KMNH VP 100,271 from the Crato Formation; C, MN 6396 (Museu Nacional, Universidade Federal do Rio de Janeiro), from the Santana Formation; D, 679 CPCA (Centro de Pesquisas de Chapada do Araripe, Brazil), from the Crato Formation. 122 Paulo M. Brito and Yoshitaka Yabumoto

Fig. 20. Notelops brama Agassiz, 1841, KMNH VP 100,272 (Kitakyushu Museum of Natural History and Human History), from the Santana Formation.

Fig. 21. Rhacolepis buccalis Agassiz, 1841, KMNH VP 100,273 (Kitakyushu Museum of Natural History and Human History), from the Santana Formation.

the margins of the jaws. Although two nominal species of and parhypurapophysis; basisphenoid not contacting the this genus are currently recognized, other morphotypes of the dorsomedial limb of the parashenoid (probably having been in genus are known. However, these cannot be referred to either contact through a cartilage component); and subepiotic fossa of the existing species (see Blum, 1991). Recently, Forey and not as deep as in many other albuliform fishes. Maisey (2010) presented the most complete revision of this taxon, redescribing it and analysing its systematic relationship within the Albuloidei. The family Chanidae contains a living representative, the Paraelops is a large fusiform fish with a short, upturned near globally distributed Chanos chanos, which is sometimes snout; large eyes; and microscopic teeth forming a dense tooth referred to as the “milk fish”. There are at least two chanids in plate. Two prominent infraorbitals form the posterior border of the Araripe Basin: Tharrhias araripis (Jordan and Branner, the orbit, although a third, the dermosphenotic, is present and 1908), and Dastilbe crandalli Jordan, 1910 (Figs. 26, 27). is much smaller than the infraorbitals (for a complete diagnosis Tharrhias was erected by Jordan and Branner (1908) for and description see Maisey, 1991). Paraelops is a very rare a from the Santana Formation that was considered to component within the Santana assemblage, and a taxonomic be a leptolepid. In the same article, these authors described and systematic revision of this taxon is necessary. another monospecific genus, Cearana, which they classified Bullichthys was recently described based on certain (with some restrictions) as an osteoglossid. Cearana rochae unique characters, including the presence of hypurapophysis was later included by Jordan (1921) as a second species of Fish faunas from the Crato and Santana formations 123

Fig. 22. Brannerion latum (Agassiz, 1841), BMNH P 1959 (Natural History Museum, London), from the Santana Formation, about 100 mm SL.

Fig. 23. Brannerion vestitum (Jordan and Branner, 1908), MPSC P055 (Museu de Paleontologia de Santana do Cariri, Ceará, Brazil), from the Santana Formation. 124 Paulo M. Brito and Yoshitaka Yabumoto

Fig. 24. Paraelops cearensis Santos, 1971, KMNH VP 100,274 (Kitakyushu Museum of Natural History and Human History), from the Santana Formation.

Fig. 25. Bullichthys santanensis Mayrinck, Brito and Otero, 2010, holotype, UERJ-PMB 142 (Universidade do Estado do Rio de Janeiro), from the Santana Formation.

Fig. 26. Tharrhias araripis (Jordan and Branner, 1908), AMNH 79293b (American Museum of Natural History), from the Santana Formation. Fish faunas from the Crato and Santana formations 125

Tharrhias (T. rochae). A few years later, in his revision of shown by Brito and Amaral (2008), some characters (such as the fish fauna of the Santana Formation, D'Erasmo (1938) the shape of the dentary and the relationship between the origin questioned the validity of these two species, commenting on of the pelvic and dorsal fins) cannot be used to differentiate a possible synonymy between them. In their overview of the species, as they vary according to ontogeny. Santana Formation fauna, Santos and Valença (1968) were the first authors to disagree with the taxonomic position of Clupeocephala incertae sedis the genus Tharrhias within the leptolepids, proposing it as a Beurlenichthys ouricuriensis Figueiredo and Gallo, 2004 member of the family Chanidae. Tharrhias araripis is a very (Fig. 28) is a small fish, reaching about 10 mm total length, common taxon from the nodule level of the Santana Formation generally found in the nodules of the Santana Formation that (Romualdo Member), and has recently been collected in the contain mass mortalities of fish (see Martill et al., 2008). slightly older Crato Formation. Tharrhias is easily recognised This taxon is also present in the Riachuelo Formation of the by its terminal mouth with no teeth on the jaws, as well as its Sergipe-Alagoas Basin. palate and hyobranchial apparatus. The jaws are very short (not Beurlenichthys presents a dentiginous lamina on a large extending past the anterior margin of the orbit); a supramaxilla and curved maxilla. Its premaxilla bears teeth shaped like rose is absent; and the posteroventral process of quadrate is thorns, and possesses a short and rounded ascending process. extended posteriorly. Its caudal skeleton has two ural centra, The first preural and first ural centra are separated. two epurals, three uroneurals, and six hypurals. Dastilbe crandalli is by far the most abundant fish in the Clupeomorpha incertae sedis Crato Formation. Four nominal species of Dastilbe have been Santanaclupea silvasantosi Maisey, 1993 is still a rare described: Dastilbe crandalli Jordan, 1910, D. elongatus component within the Santana ichthyofauna (Fig. 29). Santos, 1947, D. moraesi Santos, 1955, and D. batai Gayet, Santanaclupea is a small clupeomorph, reaching approximately 1989. However, a recent revision argued for the validity of only 120 mm standard length, and is considered to belong to a single nominal taxon, Dastilbe crandalli (Brito and Amaral, Clupeomorpha incertae sedis. This taxon has slender, toothed 2008). Dastilbe is a very abundant fish in Lower Cretaceous jaws, and the diagnostic series of ventral scutes extending from western Gondwanan localities, principally found in the Araripe, behind the pectoral fins to the anal fin. Although it presents a Parnaíba, Sergipe-Alagoas, and Sanfranciscana basins, as well series of plesiomorphic caudal endoskeletal characters, such as in the Gabon Basin. Dastilbe crandalli is a medium-sized as the autogenous parahypural; lack of fusion between the first fish, up to about 210 mm standard length, and easily identified uroneural and first preural centrum; and an unreduced first ural on the basis of its expanded operculum (which is about one- centrum; it also shares the obliquely inclined suspensorium and third to one-half of head length, ovoid in shape, and smooth); elongate jaws found in engrauloid clupeiforms (see Maisey, the position of the anal fin (closer to the caudal fin than to the 1993). ); and the deeply forked caudal fin (Brito, 2007). As

Fig. 27. Dastilbe crandalli Jordan, 1910, UERJ-PMB 41 (Universidade do Estado do Rio de Janeiro, Rio de Janeiro, Brazil), from the Crato Formation. 126 Paulo M. Brito and Yoshitaka Yabumoto

Fig. 28. Beurlenichthys ouricuriensis Figueiredo and Gallo, 2004, UERJ-PMB 117 (Universidade do Estado do Rio de Janeiro), from the Santana Formation, about 40 mm SL.

Fig. 29. Santanaclupea silvasantosi Maisey, 1993, UERJ-PMB 71 (Universidade do Estado do Rio de Janeiro), from the Santana Formation.

Otophysi family incertae sedis Santana and Crato formations. Recently, Fillol and Maisey Santanichthys diasii (Santos, 1958) (Fig. 30), from the (2004) redescribed S. diasii as a stem characiform, due to the nodules of the Santana Formation, was originally described as a presence of a complete and large, globular leptolepid ( diasii). Later, Santos (1991) considered lagenar capsules extending far lateral to the cranium. Although this species to represent a distinct genus, Santanichthys, placed we feel it is premature to place Santanichthys within the by this author within the clupeomorphs. Santanichthys is a , we have to confirm this taxon as the earliest tiny fish, mostly under 30 mm total length, found in both the known otophysan fish. Fish faunas from the Crato and Santana formations 127

Teleostei incertae sedis Axelrodichthys araripensis is relatively common in the Araripichthydae Santana Formation, although it is very rare in the Crato Araripichthys castilhoi Santos, 1985a is still a rare com- Formation, being known only from young individuals (see ponent within the Santana ichthyofauna (Fig. 31). This deep- Brito and Martill, 1999). This species is known to reach bodied, laterally compressed teleost has a covering of small about 1000 mm in total length, and is characterized by the scales on the trunk as well as on the anterior part of the base posterior moiety of the skull roof having a medial element plus of the dorsal and anal fins, but this is absent on the pelvic fins. three paired ossifications. The systematic position of Araripichthyidae within the Mawsonia brasiliensis is still a relatively rare species is still unclear. Santos (1985a) argued that this family should in comparison with Axelrodichthys. Mawsonia brasiliensis be considered as a new suborder of beryciformes, although reaches about 1280 mm in standard length, and is easily later authors considered it as an elopocephalan incertae sedis differentiated from the other species of the Araripe Basin by its (Maisey and Blum, 1990). For a discussion of characters see scales, which lack ornamentation. Maisey and Moody (2001). Mawsoniid coelacanths are relatively common in the Araripichthys is a typical component of the western Tethys western part of Gondwana, being known from the ?Upper Sea, and is also known in the Aptian Apon Formation of Jurassic to the (see Yabumoto, 2008; Cavin and Venezuela (A. axelrodi Maisey and Moody, 2001) and the Forey, 2004). Detailed comparisons between M. brasiliensis Turonian Goulmima locality of Morocco (A. corythophorus and the other species of Mawsonia show differences in the Cavin, 1997). Recently, A. castilhoi has been recognized in proportions of the skull, gular plates, and angular, as well as in the Albian of Mexico, thus increasing the the position of the otic canal fossa. A systematic revision of geographic distribution of this fish (Alvarado-Ortega and the mawsoniids is needed to resolve the validity (or synonymy) Brito, 2010). of some of these species.

Mawsoniidae Coelacanths were first recognized in the Araripe Basin DISCUSSIONS AND CONCLUSIONS when Campos and Wenz (1982) pointed out the presence of two taxa: Mawsonia sp. and “Forme B”. These two taxa have The last three decades have seen considerable progress in subsequently been identified as species of the monospecific our understanding of the anatomy, systematic relationships, genera Axelrodichthys M a i s e y , 1986 and Mawsonia , and biogeographic distribution of the fishes from Woodward, 1907. Axelrodichtys araripensis Maisey, 1986 is the Crato and Santana formations. Nevertheless, much work known from the Crato and Santana formations, and Mawsonia remains to be done to better understand the anatomy and brasiliensis Yabumoto, 2002 is known exclusively from the phylogenetic relationships of many of these fish taxa. In Santana Formation (Figs. 32, 33). addition to providing anatomical and phylogenetic information,

Fig. 30. Santanychthys diasii (Santos, 1958), UERJ-PMB 115 (Universidade do Estado do Rio de Janeiro), from the Crato Formation, 30 mm SL. 128 Paulo M. Brito and Yoshitaka Yabumoto

Fig. 31. Araripichthys castilhoi Santos, 1985, holotype, UERJ DZ 21P (Universidade do Estado do Rio de Janeiro), from the Santana Formation.

Fig. 32. Axelrodichthyes araripensis Maisey, 1986, MB. f.12607 (Berlin Museum), from the Santana Formation. Fish faunas from the Crato and Santana formations 129

Fig. 33. Mawsonia brasiliensis Yabumoto, 2002, holotype, KMNH VP 100,247 (Kitakyushu Museum of Natural History and Human History), from the Santana Formation, 1435 mm TL. the exquisitely preserved Crato and Santana fossils still have of the restricted links to the sea, perhaps using this lagoon as a much to reveal about faunal comparisons and relationships nursery (Brito, 2007). with other Gondwanan localities. Contrary to the idea of an endemic fauna related to the opening of South Atlantic Ocean, it is now widely accepted that ACKNOWLEDGMENTS during the Aptian and Albian a marine transgression connected the western part of Tethys with several of the Brazilian interior We thank Teruya Uyeno and Jennifer A. Lane for their basins, including the Araripe Basin (Berthou, 1990; Arai reviews and valuable comments. We are very grateful to David and Coimbra, 1990; Arai, 1999). The absence of fully marine Martill, Jésus Alvarado-Ortega, Cesar Amaral, José Artur invertebrates (such as echinoderms, cephalopods, or corals) Andrade, Gloria Arratia, Lionel Cavin, Valéria Gallo, Lance in levels where the oldest fishes occur suggests non-marine Grande, Lúcio Machado, John Maisey, Diogo de Mayrinck, conditions for this basin, with only intermittent connections and Sylvie Wenz for all their help and profitable discussions to the epicontinental seaway. A marine connection with the over all these years. We thank David Martill for allowing to Araripe Basin is indicated by species of the genera Vinctifer, use the illustrations of his papers for Fig. 1. Rhacolepis, Notelops, and Araripichthys, which appear to be PMB’s research has been partially supported by a CNPq closely related to Aptian/Albian assemblages occurring in the grant and a visiting scholar grant from the Fukuoka Cultural western part of Tethys in Colombia, Venezuela and Mexico Foundation, Japan. (Moody and Maisey, 1994; Schultze and Stöhr, 1996; Brito, 1997; Maisey, 2000; Maisey and Moody, 2001). Other Araripe genera also occur in the western Tethys of Morocco (e.g., REFERENCES Cladocyclus, Araripichthys), further hinting at marine links. The genera Tharrhias and Santanichthys are not yet known Agassiz, L. 1833–1844. Recherches sur les Poissons fossiles. outside the Araripe Basin. In the same way, some genera - 5 vols., 1420 pp., suppl.; Neuchâtel et Soleure such as Obaichthys, Dentilepisosteus, and Calamopleurus (Petitpierre). are known in the fresh water levels of the Kem Kem beds of Agassiz, L. 1841. On the fossil fishes found by Mr. Gardner Morocco (Forey and Grande 1998; Grande 2010). These in the Province of Ceará, in the north of Brazil. The taxa may represent a brackish or freshwater component of the Edinburgh New Philosophical Journal, Edinburgh, 30: fauna. 82–84. Some of the fish species of the Aptian Crato Formation are Agassiz, L. 1844. Sur quelques poissons fossiles du Brésil. juveniles of the same species found in the Santana Formation, Comptes Rendus de l’Academie des Sciences, Paris, 18: suggesting important palaeoecological implications related 1007–1015. to the reproductive biology of these fishes (Brito, 2007). Alvarado-Ortega, J. and P. M. Brito. 2010. On the occurrence Considering that the palaeoenvironmental conditions of the of Araripichthys in the Tlayúa Quarry (Albian), Central Crato Formation suggest a lagoonal system with probably Mexico. Fifth International Meeting on Mesozoic Fishes. fluctuating salinities (see Martill, 1993; Martill et al., 2007), Global Diversity and Evolution. Abstract Book and Field it is possible that marine forms entered the lagoon through one Guide. Universidad Autónoma del Estado de Hidalgo. 130 Paulo M. Brito and Yoshitaka Yabumoto

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Appendix 1. Fishes from the Santana Formation (Common Bullichthys santanensis Mayrinck, Brito and Otero, 2010 species of the Crato Formation in bold-faced) Clupeocephala Subdivision Ostarioclupeomorpha Class Chondrichthyes Order Gonorynchiformes Order Hybodontiformes Chanidae Hybodontidae Tharrhias araripis (Jordan and Branner, 1908) Tribodus limae Brito and Ferreira, 1989 Clupeocephala order and family incertae sedis Order Rajiformes family incertae sedis Beurlenichthys ouricuriensis Figueiredo and Gallo, 2004 Iansan beurleni (Santos, 1968) Clupeomorpha order and family incertae sedis Class Actinopterygii Santanaclupea silvasantosi Maisey, 1993 Division Holostei Otophysi family incertae sedis Subdivision Ginglymodi Santanichthys diasii (Santos, 1958) Order Semionotiformes Teleostei insertae sides Semionotidae Araripichthydae Araripelepidotes temnurus (Agassiz, 1841) Araripichthys castilhoi Santos, 1985 Lepidotes wenzae Brito and Gallo, 2003 Class Sarcopterygii Order Lepisosteiformes Order Coelacanthiformes Obaichthyidae Mawsoniidae Obaichthys decoratus Wenz and Brito, 1992 Axelrodichthys araripensis Maisey, 1986 Dentilepisosteus laevis (Wenz and Brito, 1992) Mawsonia brasiliensis Yabumoto, 2008 Subdivision Halecomorphi Order Amiiformes Amiidae Calamopleurus cylindricus Agassiz, 1841 Order Ionoscopiformes Ophiopsidae Placidichthys bidorsalis Brito, 2000 Oshuniidae Oshunia brevis Wenz and Kellner, 1986 Order Pycnodontiformes Pycnodontidae Neoproscinetes penalvai (Santos, 1968) Iemanja palma Wenz, 1989 Superdivision Teleosteomorpha Ordre Aspidorhynchiformes Aspidorhynchidae Vinctifer comptoni (Agassiz, 1841) Division Teleostei Order Ichthyodectiformes Cladocyclidae Cladocyclus gardneri Agassiz, 1841 Order Crossognathidformes Suborder Pachyrhizodontoidei Notelopidae Notelops brama Agassiz, 1841 Pachyrhizodontidae Rhacolepis buccalis Agassiz, 1841 Subdivision Elopomorpha Order Albuliformes Branneriun latum (Agassiz, 1841) Brannerion vestitum (Jordan and Branner, 1908) Paraelops cearensis Santos, 1971 136 Paulo M. Brito and Yoshitaka Yabumoto

Appendix 2. Fishes from the Crato Formation. (Common taxa of the Santana Formation in bold-faced).

Class Actinopterygii Division Holostei Subdivision Ginglymodi Order Semionotiformes Semionotidae Araripelepidotes cf. temnurus (Agassiz, 1841) Order Lepisosteiformes Obaichthyidae Obaichthyidae gen. et sp. indet. Subdivision Halecomorphi Order Amiiformes Amiidae Calamopleurus cylindricus Agassiz, 1841 Cratoamia gondwanica Brito, Yabumoto and Grande, 2008 Order Ionoscopiformes Ophiopsidae Placidichthys bidorsalis Brito, 2000 Superdivision Teleosteomorpha Ordre Aspidorhynchiformes Aspidorhynchidae Belonostomus sp. Division Teleostei Order Ichthyodectiformes Cladocyclidae Cladocyclus gardneri Agassiz, 1841 Subdivision Ostarioclupeomorpha Order Gonorynchiformes Chanidae Dastilbe crandalii Jordan, 1910 Otophysi family incertae sedis Santanichthys diasii (Santos, 1958) Class Sarcopterygii Order Coelacanthiformes Mawsoniidae Axelrodichthys araripensis Maisey, 1986