An updated review of the fish faunas from the Crato and Santana … · 2017-06-13 · An updated...

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] of Natural History, Kitakyushu Museum of Natural History and Human History, 2-4-1 Higashida,

Yahatahigashi-ku, Kitakyushu, Fukuoka, 805-0071, JapanE-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.

Bull. Kitakyushu Mus. Nat. Hist. Hum. Hist., Ser. A, 9: 107–136, March 31, 2011

INTRODUCTION

The fossil-bearing deposits of the Lower Cretaceous (Aptian/Albian) Crato and Santana formations of the Araripe Basin (Northeastern Brazil) have yielded numerous vertebrates, including fishes, frogs, turtles, lizards, pterosaurs, dinosaurs, and bird feathers (Figs. 1–3). The fishes, which form the dominant element of this biota, are exceptionally abundant and are famous worldwide for their quantity and excellent state of preservation, being frequently articulated and sometimes three-dimensionally preserved. In some cases, there is even the preservation of phosphatic soft tissues, including muscle fibers, gill filaments, and stomach contents (martill, 1988; WilBY and martill, 1992).

The first account of fossil fishes from Araripe appears in the atlas of the “Reisen Brasilien” expedition between 1817 and 1820 (Spix and martiuS, 1823). This publication was soon followed by those of Gardiner (1841) and aGaSSiz (1841), who drew up a list of fishes and identified one of the forms shown

by Spix and martiuS (1823) as Rhacolepis aGaSSiz (1841). Subsequently, knowledge of the ichthyofauna was further enriched by the works of aGaSSiz (1833–1844), WoodWard (1887, 1890, 1895, 1901, 1908), Jordan and Branner (1908), Jordan (1919, 1921), D’eraSmo (1938), dunkle (1940), and SantoS (1945, 1947, 1950, 1958, 1960). However it was not until the publication of SantoS and Valença (1968) that the first review of the Araripe ichthyofauna became available.

Since then, numerous authors have described a quantity of new taxa (CampoS and Wenz, 1982; SantoS, 1985a; maiSeY, 1986; Wenz and kellner, 1986; Wenz, 1989; Brito and Ferreira, 1989), developed descriptions of known fossils, and/or discussed the phylogenetic affinities of the various components of the fauna (e.g., taVerne, 1974, 1976; ForeY, 1977, 1998; Wenz, 1977; patterSon and roSen, 1977; oliVeira, 1978; patterSon, 1984; SantoS, 1985b, 1990a, b, 1994a, b, 1995; Brito, 1988, 1992b, 1999; Brito and Wenz, 1990; maiSeY and Blum, 1990; Brito and martill, 1999; Brito and meunier, 2000; Brito et al., 1998, 2000, 2010;

Paulo M. Brito and Yoshitaka YaBumoto108

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 Brito, 2004; ForeY and maiSeY, 2010).

maiSeY (1991) presented the most comprehensive account of the Araripe fish fauna published until then. However, as knowledge of this assemblage has continued to grow (e.g., Wenz and Brito, 1992; maiSeY, 1993; Brito, 2000; YaBumoto, 2002; Brito and Gallo, 2003; FiGueiredo and Gallo, 2004; Brito et al., 2008) we present in this paper an updated review of the fish faunas from the Crato and Santana formations.

The Crato and Santana formations have provided what are clearly the richest Lower Cretaceous fish faunas in South America (SantoS and Valença, 1968; Wenz and Brito, 1990; maiSeY, 1991; martill, 1993). This ichthyofauna comprises at least 28 nominal species.

The valid species of the Crato and Santana formations are all briefly reviewed below, and the relative phylogenetic relationships among these species, as currently hypothesized, 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.


Fig. 4. Tribodus limae Brito and Ferreira, 1989, MNRJ 105 (Museu Nacional do Rio de Janeiro), from the Santana Formation.

THE ICHTHYOFAUNA

HybodontidaeThere is one species representing the family Hybodontidae

in the Araripe Basin, Tribodus limae Brito and Ferreira, 1989 (Fig. 4). This species occurs in the Santana Formation and is unknown in the Crato Formation. It is a relatively frequent component of the fauna.

This species is known to reach over 600 mm in total length. It differs from most other hybodontids (except Acrodus and Asteracanthus) in having a crushing dentition (Brito, 1992a) and a hyostylic jaw suspension (maiSeY and CarValho, 1997). Recently, an anatomical revision of the pectoral girdle of Tribodus as well as a revision of the braincase, based on CT scan had been published (lane and maiSeY, 2009; lane, 2010).

Rajiformes family incertae sedisThere is one guitarfish species from the Santana Formation

of the Araripe Basin, Iansan beurleni (SantoS, 1968) (Fig. 5), which reaches close to 900 mm in total length (for a revision of the anatomical patterns of this taxon see Brito and Sérret, 1996).

Iansan is relatively common in the Santana fish fauna, but so far has not been reported from the Crato Formation or any other Lower Cretaceous western Gondwanan formations. The phylogenetic relationships of I. beurleni to other fossil and living guitarfishes are still unclear.

SemionotidaeSemionotidae is a geographically widespread family

with a chronological range from the Middle Triassic to the Late Cretaceous. Semionotids are found in both marine and continental strata from Europe, North America, Asia, Africa,

South America and Cuba. The Western Gondwana semionotids are the most characteristic Mesozoic fishes, especially in the northeastern basins of Brazil where they have a temporal range from the Late Jurassic to Late Cretaceous (comprising nine nominal species; Brito and Gallo, 2003; Gallo and Brito, 2004). Two species are known in the Araripe Basin, each occurring in both the Crato and Santana formations: Araripelepidotes temnurus (aGaSSiz, 1841), and Lepidotes wenzae Brito and Gallo, 2003 (Figs. 6 and 7).

Araripelepidotes temnurus is relatively common in the Santana Formation, although it is very rare in the Crato Formation. This taxon, considered for a long time as a species of Lepidotes, was removed to the new genus Araripelepidotes by SantoS (1990a). This species is known to reach about 400 mm in total length, and is characterized by a very weak, edentulous lower jaw composed of a single element; reduction of the coronoid process; and separation of the dermopterotic and the frontal by the dermosphenotic. Thus far, Araripelepidotes appears to be endemic to the Araripe Basin.

Lepidotes wenzae is a relatively small-sized species, reach-ing about 250 mm standard length. This species, not very common in the Santana Formation (on the basis of known specimens), has a moderate pre-dorsal elevation, scales lacking ornamentation, and a moderate crushing dentition. Lepidotes wenzae is a rarely occurring component of the fish fauna of the Crato Formation, and is represented only by juvenile specimens (Brito, 2007). Detailed comparisons of the skull morphology between the specimens from the Santana and Crato formations are needed to resolve the validity (or synonymy) of this species. For now, we consider Lepidotes from the Araripe Basin to represent a unique species.


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.


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.


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.

Lepisosteiformes (Obaichthyidae)The family Lepisosteidae is well known nowadays by seven

nominal species belonging to two genera, Lepisosteus and Atractosteus (WileY, 1976; nelSon, 1994). Extant lepisosteids are distributed in eastern North America, Central America and the Caribbean. They are basically freshwater fishes, but may occasionally occur in brackish water. Only one extant species, Atractosteus tristoechus, invades marine environments in the vicinity of Cuba and the Isle of Pines.

Lepisosteiforms have a long geological history (dating as far back as the Lower Cretaceous; e.g., CaSier, 1961; Wenz and Brito, 1992, 1996; Brito et al., 2000). They also have a wide paleogeographic distribution, mainly during the Late Cretaceous and Tertiary in Africa (e.g., aramBourG and Joleaud, 1943; GaYet et al., 1988; Werner, 1994; CaVin and Brito, 2001), South America (e.g., SantoS, 1984; Cione, 1987; GaYet and Brito, 1989; GaYet, 1991; Bertini et al., 1993), Madagascar (GottFried and krauSe, 1998); India (e.g., WoodWard, 1890, 1908; WileY, 1976; Jain and Sahni, 1983; rana and kumar, 1990), Europe (e.g., Jonet, 1981; BuFFetaut et al., 1996; CaVin et al., 1996; SiGé et al., 1997), and North America (see WileY, 1976; ruSSell, 1988; for a complete listing).

There are two lepisosteiforms in the Araripe Basin. These taxa, placed by Wenz and Brito (1992) in the genus Obaichthys, were recently reviewed by Grande (2010), who proposed the new family Obaichthydae as well as a second genus (Dentilepisosteus).

Obaichthys decoratus Wenz and Brito, 1992 and Denti-lepisosteus laevis (Wenz and Brito, 1992) (Figs. 8, 9) are the oldest known lepisosteiforms. Both species are very rare and are thus far known only from the Santana Formation. Recently, another obaichthyid was also recorded in the Crato Formation (Fig. 10), based on a unique specimen that seems to be more closely related to D. laevis than to O. decoratus.

Although Obaichthyidae present 14 to 16 synapomorphies shared with Lepisosteidae (e.g., nasal process of premaxilla forms much of the ornamented drmal roof in the snout region;

predorsal length of fish is 75% or more of standard length; the palatal complex has laterolly sliding articulation between the metapterygoid and the basipterygoid process of the braincase; the presence of a “tongue bone” or basihyal tooth plate consisting of a mosaic of bony pates; the presence of opisthocoelous vertebral centra), it also retains some characters considered primitive for neopterygians, such as the presence of a free maxilla; two post-orbital bones; and paleoniscoid-type scales (Wenz and Brito, 1992, 1996; Brito et al., 2000; Grande, 2010). The differences between the two described species mainly concern the ornamentation of the dermal bones and scales, shape of scales, and dimensions and proportions of the body and cranial bones.

Two other species of obaichthyids, Obaichthys africanus and Dentilepisosteus ?kemkemensis, were recently described by Grande (2010) from the ?Late Cretaceous Kem Kem beds of Morocco. A detailed taxonomic and phylogenetic discussion regarding the validity of the family Obaichthydae is discussed by Grande (2010). We agree with Grande (2010) that the Obaichthydae form a monophyletic group.

AmiidaeCurrently the amiids are represented by a single extant

species, Amia calva, found in lakes and rivers of Eastern North America. However, this family has a long temporal distribution, being known since the Upper Jurassic. In the western part of Gondwana, amiids are represented by the Subfamily Vidalamiinae, (Grande and BemiS, 1998).

There are two species of amiids in the Araripe Basin, Calamopleurus cylindricus aGaSSiz, 1841 and Cratoamia gondwanica Brito, YaBumoto and Grande, 2008. Cala-mopleurus cylindricus is known from both the Crato and Santana formations (Fig. 11), but Cratoamia (Fig. 12) is currently restricted to the Crato Formation.

Calamopleurus cylindricus is one of the best-studied amiids (e.g., SantoS, 1960; Grande and BemiS, 1998). This species has a maximum known size of about 1400 mm total length, and is relatively common in the Santana Formation, where it


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.


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.

is represented by the type species C. cylindricus (= Enneles audax). The genus Calamopleurus was diagnosed by Grande and BemiS (1998) on the basis of the following characters: posterior margin of caudal fin with an almost vertical outline; opercular process of the hyomandibular extremely elongate; teeth of the coronoid bones distributed in a single marginal row; teeth of the vomer arranged in a single anterior row, with one or more teeth in a longitudinal row perpendicular to the marginal row; dermopterotic with a bundle of elongate splint-like bones (dermopterotic ribs) fused to its posterior end;

dermosphenotic not tightly sutured to the other bones of the braincase; and posterior margin of the gular deeply scalloped with a series of sharp points and concavities.

Of the three nominal species of Calamopleurus, C. cylindricus is the best known due to its relative abundance in the Santana Formation. Two other species (C. africanus ForeY and Grande, 1998, from the Late Cretaceous Kem Kem beds of Morocco; and C. mawsoni (WoodWard, 1902) from the Early Cretaceous Recôncavo Basin) were described based on unique disarticulated specimens. Calamopleurus africanus is known


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

only from a skull and differs from the type species primarily in the shape of its gular plate, which carries a median ridge on its ventral surface and bears 10 small projections on the posterior margin (see ForeY and Grande, 1998). Calamopleurus mawsoni is known only from the type material, consisting of an articulated skeleton, lacking the cranial and pectoral girdle regions. This species differs from C. cylindricus in the number of dorsal radials and higher number of preural vertebrae (see Grande and BemiS, 1998).

Cratoamia gondwanica is a medium-sized amiid of approximately 700 mm total length. This species is easily characterized by the presence of a long dorsal fin with between 27 and 30 rays, as well as by the weak ornamentation of its dermal bones.

OphiopsidaeOphiopsids are a typical Jurassic/Cretaceous group found

in Europe, North America, Africa and South America. In Brazil, ophiopsids were previously recorded by SantoS and Valença (1968), for a specimen that unfortunately could not be diagnosed even into a familial level, identified as Ophiopsis cretaceous. For this reason, this species is considered a Nomen nudum.

There is one species representing the ophiopsids in the Araripe Basin, Placidichthys bidorsalis Brito, 2000 (Fig. 13). Placidichthys is a small ophiopsid known to reach 80 mm total length, and several young juveniles are also known, especially in the Crato Formation. Placidichthys is easily diagnosed by its very elongate body. It differs from other members of the family in having a long dorsal fin subdivided into two parts. The deepest part of the body is between the head and the anterior part of the dorsal fin, with body depth decreasing strongly toward the posterior part of the dorsal fin. The body is covered by small, thin, diamond-shaped ganoid scales distributed from the pectoral girdle to the base of the axial lobe of the caudal

fin. The pectoral fin is very well developed, with a convex margin. The caudal fin is deeply forked, with the upper lobe larger than the lower lobe.

Another species of Placidichthys, P. tucanensis Brito and alVarado-orteGa, 2008, was recently described from the Aptian Marizal Formation of the Tucano Bahia. Placidichthys tucanensis is clearly distinguishable from the species in the Araripe Basin by the absence of an anal fin; the lower number of flank scales in the caudal region; and the much more slender body shape.

OshuniidaeThe Family Oshuniidae was recently proposed by Grande

and BemiS (1998), and is represented by only one genus and one nominal species (Oshunia brevis Wenz and kellner, 1986; Fig. 14; from the Santana Formation), although maiSeY (1991) proposed that the genus may contain additional species.

Oshunia brevis is easily characterized by its high, trian-gularly shaped dorsal fin and body covering of typical amioid scales. Recently, a taxon previously considered a member of Ionoscopidae, Quetzalichthys perrilliatae alVarado-orteGa and eSpinoSa-arruBarrena, 2008, was described from the Albian Tlayúa Quarry of Mexico and considered to represent the sister-group of Oshunia on the basis of two morphological characters (presence of paired of longitudinal fossae on the sides of all vertebral centra; and the presence of only three urodermals).

PycnodontidaeThere are two pycnodontids in the Araripe Basin, known

exclusively from the Santana Formation: Neoproscinetes penalva (SantoS, 1970), and Iemanja palma Wenz, 1989. Neoproscinetes penalva (Fig. 15) is a relatively common fish, reaching about 350 mm total length and having a nearly circular shape in lateral outline. This fish presents a dentition


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.


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).


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.

comprising a pavement of large, low, elongate, dome-like teeth, with the alternation of one large and two small teeth on the main vomerine tooth row. Eight or nine teeth are present in the main vomerine tooth row.

Iemanja palma (Fig. 16) is still a rare component within the fauna of the Araripe Basin. It is known from specimens of approximately 600 mm total length. This species presents, among other characters, a large supraoccipital crest; fusion of the first arcocentra with the skull; an accentuated prognatism; vomerine teeth organized in rows only in the posterior portion of the vomer; vomerine teeth of the principal row with a reniform shape; presence of a urodermal; scales restricted to the antero-ventral part of the body; scales of the dorsal margin and the ventral keel carrying large spines; as well as the ventral lateral line scales passing in the first ventral scales. Iemanja palma was originally considered a Gyrodontid, but recently phylogenetc analysis (e.g., nurSall, 1996; poYato-

ariza and Wenz, 2002) suggested this taxon should be placed together with other members of the pycnodontids.

AspidorhynchidaeThere are two aspidorhynchids in the Araripe Basin:

Vinctifer comptoni (aGaSSiz, 1841) (Fig. 17) from the Santana Formation, and a new species of Belonostomus (Fig. 18) from the Crato Formation. Vinctifer and Belonostomus are two of four genera comprising the Aspidorhynchidae, a widely distributed Mesozoic family (the other two genera are Aspidorhynchus and Richmondichthys; the latter is probably a synonym of Vinctifer) of highly elongate predatory fishes. The aspidorhynchids are easily recognized by the presence of a long rostrum formed by the premaxillae; presence of a predentary on the lower jaw; deep elongate flank scales; and posteriorly located dorsal and anal fins.

Vinctifer is one of the most common species in the Santana


Formation, where it is known from specimens of between 50 and 900 mm total length (Brito, 1997). Vinctifer comptoni is a typical southern Tethyan genus, also known in other northeastern Brazilian basins (e.g., Parnaíba, Sergipe-Alagoas), as well as in the Aptian-Albian of Venezuela, Colombia, Mexico and Antarctica (moodY and maiSeY, 1994; SChultze and Stöhr, 1996; appleGate, 1996; Brito, 1997) and probably also the Neocomian of Rio Muni, Equatorial Guinea (taVerne, 1969). Two nominal species are recognised: Vinctifer comptoni (the type species), and Vinctifer longirostris, known only from the Tucano Basin (also in northeastern Brazil). Aspidorhynchids are very rare in the Crato Formation, and until now were known only from a single specimen, attributed to Belonostomus sp. This is a medium-sized Aspidorhynchid, incompletely preserved, which was previously cited as V. longirostris (Brito, 1997). However, further preparation of the specimen revealed several diagnostic features of the genus (e.g., elongated predentary and presence of a posterior supramaxilla).

Within the Aspidorhynchidae, the genus Belonostomus is considered to have the largest temporal range, being known from the Late Jurassic to the Late Cretaceous. This genus has a widespread paleobiogeographical distribution, occurring in marine strata from Europe, the Middle East, North America, South America, and Africa.

Cladocyclidae There is one species in the Crato and Santana formations

representing the ichthyodectiform family Cladocyclidae, Cladocyclus gardneri aGaSSiz, 1841 (Fig. 19). This species, which is very abundant in the Santana Formation, occurs in the Araripe Basin but is also reported from other Brazilian north-eastern basins such as Parnaíba and Sergipe-Alagoas (leal and Brito, 2004). Cladocyclus gardineri can attain a large size (over 1000 mm standard length). It is easily recognised by its slender and laterally compressed body; large and oblique mouth opening upwards with a single series of conical, slender, pointed teeth; large orbital opening; posterior position of the dorsal and anal fins; prominent cycloid scales; and deeply forked caudal fin (maiSeY, 1991; Brito, 2007).

aGaSSiz (1841) described Cladocyclus gardneri on the basis of specimens collected from the Santana Formation by George Gardner. A century later, SantoS (1950) described a second species from the type locality, which he named C. ferus. Subsequent authors attempted to distinguish between these two nominal species, until patterSon and roSen in their 1977 revision of the ichthyodectiformes proposed that the two species should be synonymized.

Pachyrhizodontoidei (Pachyrhizodontidae and Note-lopidae)

Although pachyrhizodontoids are generally placed within the elopomorphs, their phylogenetic position among Teleostei

remains unclear (see CaVin, 1995). Pachyrhizodontoidei is a typical Mesozoic suborder, considered to represent a monophyletic group that includes Notelops, Rhacolepis, Goulmimichthys, Pachyrhizodus, Elopopsis, and Michin (CaVin, 2001; alVarado-orteGa et al., 2008).

Cope (1872) erected the family Pachyrhizodontidae for Pachyrhizodus and related forms. ForeY (1977) reviewed this group, creating the suborder Pachyrhyzodontoidei including two families, Notolepidae (for Notelops) and Pachyrhizodontidae (for Pachyrhizodus, Rhacolepis, and Elopopsis). Later, other genera such as Platnix, Greenwoodella, Goulmimichthys, and Tingitanichthys were also included in this suborder.

Two nominal species of pachyrhizodontoids are known in the Araripe Basin, exclusively in the Santana Formation: Notelops brama aGaSSiz, 1841 (Fig. 20), and Rhacolepis buccalis aGaSSiz, 1841 (Fig. 21).

Notelops and Rhacolepis are very similar fishes, and may occasionally be confused. Both taxa have a fusiform body with a pointed snout. The principal differences between these taxa are the presence of three prominent bones forming the posterior border of the eye in Notelops (dermosphenotic plus two infraorbital plates), versus four in Rhacolepis (dermosphenotic plus three infraorbital plates). Other differences are related to the posterior extent of the infraorbitals (just reaching the preopercle in Notelops vs. overlying the preopercle in Rhacolepis) and the shape of the pterotic (without posterior spine in Notelops vs. reduced to a short spine in Rhacolepis). For a detailed description and diagnosis, see maiSeY and Blum (1990) and maiSeY (1991).

A taxonomic revision of Notelops is necessary, as this genus presents different morphotypes (see maiSeY, 1991). Notelops reaches approximately 600 mm total length. Rhacolepis is one of the most abundant fish in the Santana Formation, and is commonly found in an excellent state of preservation, reaching 250 mm total length.

AlbuliformesThe order Albuliformes is an extant group of marine

elopomorphs with low species diversity. The relationships of the elopomorph clade to other basal teleostean groups is still debated, and their intrarelationships are uncertain. The best known albuliform fishes are the modern Albula and Pterothrissus.

At least four albuliforms are found in the Araripe Basin, exclusively in the Santana Formation: Brannerion latum (aGaSSiz, 1841); B. vestitum (Jordan and Branner, 1908); Paraelops cearensis SantoS, 1971; and Bullichthys santanensis maYrinCk, Brito and otero, 2010 (Figs. 22–25).

Brannerion is a deep-bodied fish, with a long anal fin that originates beneath the posterior level of the dorsal fin. Other diagnostic characters are large eyes and villiform teeth on


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.


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 this genus are currently recognized, other morphotypes of the genus are known. However, these cannot be referred to either of the existing species (see Blum, 1991). Recently, ForeY and maiSeY (2010) presented the most complete revision of this taxon, redescribing it and analysing its systematic relationship within the Albuloidei.

Paraelops is a large fusiform fish with a short, upturned snout; large eyes; and microscopic teeth forming a dense tooth plate. Two prominent infraorbitals form the posterior border of the orbit, although a third, the dermosphenotic, is present and is much smaller than the infraorbitals (for a complete diagnosis and description see maiSeY, 1991). Paraelops is a very rare component within the Santana assemblage, and a taxonomic and systematic revision of this taxon is necessary.

Bullichthys was recently described based on certain unique characters, including the presence of hypurapophysis

and parhypurapophysis; basisphenoid not contacting the dorsomedial limb of the parashenoid (probably having been in contact through a cartilage component); and subepiotic fossa not as deep as in many other albuliform fishes.

ChanidaeThe family Chanidae contains a living representative, the

near globally distributed Chanos chanos, which is sometimes referred to as the “milk fish”. There are at least two chanids in the Araripe Basin: Tharrhias araripis (Jordan and Branner, 1908), and Dastilbe crandalli Jordan, 1910 (Figs. 26, 27).

Tharrhias was erected by Jordan and Branner (1908) for a teleost from the Santana Formation that was considered to be a leptolepid. In the same article, these authors described another monospecific genus, Cearana, which they classified (with some restrictions) as an osteoglossid. Cearana rochae was later included by Jordan (1921) as a second species of


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.


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.


Fig. 27. Dastilbe crandalli Jordan, 1910, UERJ-PMB 41 (Universidade do Estado do Rio de Janeiro, Rio de Janeiro, Brazil), from the Crato Formation.

Tharrhias (T. rochae). A few years later, in his revision of the fish fauna of the Santana Formation, D'eraSmo (1938) questioned the validity of these two species, commenting on a possible synonymy between them. In their overview of the Santana Formation fauna, SantoS and Valença (1968) were the first authors to disagree with the taxonomic position of the genus Tharrhias within the leptolepids, proposing it as a member of the family Chanidae. Tharrhias araripis is a very common taxon from the nodule level of the Santana Formation (Romualdo Member), and has recently been collected in the slightly older Crato Formation. Tharrhias is easily recognised by its terminal mouth with no teeth on the jaws, as well as its palate and hyobranchial apparatus. The jaws are very short (not extending past the anterior margin of the orbit); a supramaxilla is absent; and the posteroventral process of quadrate is extended posteriorly. Its caudal skeleton has two ural centra, two epurals, three uroneurals, and six hypurals.

Dastilbe crandalli is by far the most abundant fish in the Crato Formation. Four nominal species of Dastilbe have been described: Dastilbe crandalli Jordan, 1910, D. elongatus SantoS, 1947, D. moraesi SantoS, 1955, and D. batai GaYet, 1989. However, a recent revision argued for the validity of only a single nominal taxon, Dastilbe crandalli (Brito and amaral, 2008). Dastilbe is a very abundant fish in Lower Cretaceous western Gondwanan localities, principally found in the Araripe, Parnaíba, Sergipe-Alagoas, and Sanfranciscana basins, as well as in the Gabon Basin. Dastilbe crandalli is a medium-sized fish, up to about 210 mm standard length, and easily identified on the basis of its expanded operculum (which is about one-third to one-half of head length, ovoid in shape, and smooth); the position of the anal fin (closer to the caudal fin than to the pelvic fin); and the deeply forked caudal fin (Brito, 2007). As

shown by Brito and amaral (2008), some characters (such as the shape of the dentary and the relationship between the origin of the pelvic and dorsal fins) cannot be used to differentiate species, as they vary according to ontogeny.

Clupeocephala incertae sedisBeurlenichthys ouricuriensis FiGueiredo and Gallo, 2004

(Fig. 28) is a small fish, reaching about 10 mm total length, generally found in the nodules of the Santana Formation that contain mass mortalities of fish (see martill et al., 2008). This taxon is also present in the Riachuelo Formation of the Sergipe-Alagoas Basin.

Beurlenichthys presents a dentiginous lamina on a large and curved maxilla. Its premaxilla bears teeth shaped like rose thorns, and possesses a short and rounded ascending process. The first preural and first ural centra are separated.

Clupeomorpha incertae sedisSantanaclupea silvasantosi maiSeY, 1993 is still a rare

component within the Santana ichthyofauna (Fig. 29). Santanaclupea is a small clupeomorph, reaching approximately 120 mm standard length, and is considered to belong to Clupeomorpha incertae sedis. This taxon has slender, toothed jaws, and the diagnostic series of ventral scutes extending from behind the pectoral fins to the anal fin. Although it presents a series of plesiomorphic caudal endoskeletal characters, such as the autogenous parahypural; lack of fusion between the first uroneural and first preural centrum; and an unreduced first ural centrum; it also shares the obliquely inclined suspensorium and elongate jaws found in engrauloid clupeiforms (see maiSeY, 1993).


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 sedisSantanichthys diasii (SantoS, 1958) (Fig. 30), from the

nodules of the Santana Formation, was originally described as a leptolepid (Leptolepis diasii). Later, SantoS (1991) considered this species to represent a distinct genus, Santanichthys, placed by this author within the clupeomorphs. Santanichthys is a tiny fish, mostly under 30 mm total length, found in both the

Santana and Crato formations. Recently, Fillol and maiSeY (2004) redescribed S. diasii as a stem characiform, due to the presence of a complete Weberian apparatus and large, globular lagenar capsules extending far lateral to the cranium. Although we feel it is premature to place Santanichthys within the Characiformes, we have to confirm this taxon as the earliest known otophysan fish.


Fig. 30. Santanychthys diasii (SantoS, 1958), UERJ-PMB 115 (Universidade do Estado do Rio de Janeiro), from the Crato Formation, 30 mm SL.

Teleostei incertae sedisAraripichthydae

Araripichthys castilhoi SantoS, 1985a is still a rare com-ponent within the Santana ichthyofauna (Fig. 31). This deep-bodied, laterally compressed teleost has a covering of small scales on the trunk as well as on the anterior part of the base of the dorsal and anal fins, but this is absent on the pelvic fins. The systematic position of Araripichthyidae within the teleosts is still unclear. SantoS (1985a) argued that this family should be considered as a new suborder of beryciformes, although later authors considered it as an elopocephalan incertae sedis (maiSeY and Blum, 1990). For a discussion of characters see maiSeY and moodY (2001).

Araripichthys is a typical component of the western Tethys Sea, and is also known in the Aptian Apon Formation of Venezuela (A. axelrodi maiSeY and moodY, 2001) and the Turonian Goulmima locality of Morocco (A. corythophorus CaVin, 1997). Recently, A. castilhoi has been recognized in the Albian Tlayúa Formation of Mexico, thus increasing the geographic distribution of this fish (alVarado-orteGa and Brito, 2010).

Mawsoniidae Coelacanths were first recognized in the Araripe Basin

when CampoS and Wenz (1982) pointed out the presence of two taxa: Mawsonia sp. and “Forme B”. These two taxa have subsequently been identified as species of the monospecific genera Axelrodichthys ma i S e Y , 1986 and Mawsonia WoodWard, 1907. Axelrodichtys araripensis maiSeY, 1986 is known from the Crato and Santana formations, and Mawsonia brasiliensis YaBumoto, 2002 is known exclusively from the Santana Formation (Figs. 32, 33).

Axelrodichthys araripensis is relatively common in the Santana Formation, although it is very rare in the Crato Formation, being known only from young individuals (see Brito and martill, 1999). This species is known to reach about 1000 mm in total length, and is characterized by the posterior moiety of the skull roof having a medial element plus three paired ossifications.

Mawsonia brasiliensis is still a relatively rare species in comparison with Axelrodichthys. Mawsonia brasiliensis reaches about 1280 mm in standard length, and is easily differentiated from the other species of the Araripe Basin by its scales, which lack ornamentation.

Mawsoniid coelacanths are relatively common in the western part of Gondwana, being known from the ?Upper Jurassic to the Cenomanian (see YaBumoto, 2008; CaVin and ForeY, 2004). Detailed comparisons between M. brasiliensis and the other species of Mawsonia show differences in the proportions of the skull, gular plates, and angular, as well as in the position of the otic canal fossa. A systematic revision of the mawsoniids is needed to resolve the validity (or synonymy) of some of these species.

DISCUSSIONS AND CONCLUSIONS The last three decades have seen considerable progress in

our understanding of the anatomy, systematic relationships, taxonomy, and biogeographic distribution of the fishes from the Crato and Santana formations. Nevertheless, much work remains to be done to better understand the anatomy and phylogenetic relationships of many of these fish taxa. In addition to providing anatomical and phylogenetic information,


Fig. 32. Axelrodichthyes araripensis maiSeY, 1986, MB. f.12607 (Berlin Museum), from the Santana Formation.

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


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 much to reveal about faunal comparisons and relationships 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 during the Aptian and Albian a marine transgression connected the western part of Tethys with several of the Brazilian interior basins, including the Araripe Basin (Berthou, 1990; arai and CoimBra, 1990; arai, 1999). The absence of fully marine invertebrates (such as echinoderms, cephalopods, or corals) in levels where the oldest fishes occur suggests non-marine conditions for this basin, with only intermittent connections to the epicontinental seaway. A marine connection with the Araripe Basin is indicated by species of the genera Vinctifer, Rhacolepis, Notelops, and Araripichthys, which appear to be closely related to Aptian/Albian assemblages occurring in the western part of Tethys in Colombia, Venezuela and Mexico (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., Cladocyclus, Araripichthys), further hinting at marine links. The genera Tharrhias and Santanichthys are not yet known outside the Araripe Basin. In the same way, some genera such as Obaichthys, Dentilepisosteus, and Calamopleurus are known in the fresh water levels of the Kem Kem beds of Morocco (ForeY and Grande 1998; Grande 2010). These taxa may represent a brackish or freshwater component of the fauna.

Some of the fish species of the Aptian Crato Formation are juveniles of the same species found in the Santana Formation, suggesting important palaeoecological implications related to the reproductive biology of these fishes (Brito, 2007). Considering that the palaeoenvironmental conditions of the Crato Formation suggest a lagoonal system with probably fluctuating salinities (see martill, 1993; martill et al., 2007), it is possible that marine forms entered the lagoon through one

of the restricted links to the sea, perhaps using this lagoon as a nursery (Brito, 2007).

ACKNOWLEDGMENTS

We thank Teruya uYeno and Jennifer A. lane for their reviews and valuable comments. We are very grateful to David martill, Jésus alVarado-orteGa, Cesar amaral, José Artur andrade, Gloria arratia, Lionel CaVin, Valéria Gallo, Lance Grande, Lúcio maChado, John maiSeY, Diogo de maYrinCk, and Sylvie Wenz for all their help and profitable discussions over all these years. We thank David martill for allowing to use the illustrations of his papers for Fig. 1.

PMB’s research has been partially supported by a CNPq grant and a visiting scholar grant from the Fukuoka Cultural Foundation, Japan.

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Appendix 1. Fishes from the Santana Formation (Common species of the Crato Formation in bold-faced)

Class ChondrichthyesOrder Hybodontiformes

HybodontidaeTribodus limae Brito and Ferreira, 1989

Order Rajiformes family incertae sedisIansan beurleni (SantoS, 1968)

Class ActinopterygiiDivision Holostei

Subdivision GinglymodiOrder Semionotiformes

SemionotidaeAraripelepidotes temnurus (AgAssiz, 1841) Lepidotes wenzae Brito and Gallo, 2003

Order LepisosteiformesObaichthyidaeObaichthys decoratus Wenz and Brito, 1992Dentilepisosteus laevis (Wenz and Brito, 1992)

Subdivision HalecomorphiOrder Amiiformes

AmiidaeCalamopleurus cylindricus AgAssiz, 1841

Order IonoscopiformesOphiopsidaePlacidichthys bidorsalis Brito, 2000

OshuniidaeOshunia brevis Wenz and kellner, 1986

Order PycnodontiformesPycnodontidaeNeoproscinetes penalvai (SantoS, 1968)Iemanja palma Wenz, 1989

Superdivision TeleosteomorphaOrdre Aspidorhynchiformes

AspidorhynchidaeVinctifer comptoni (aGaSSiz, 1841)

Division TeleosteiOrder Ichthyodectiformes

CladocyclidaeCladocyclus gardneri AgAssiz, 1841

Order CrossognathidformesSuborder PachyrhizodontoideiNotelopidaeNotelops brama aGaSSiz, 1841

PachyrhizodontidaeRhacolepis buccalis aGaSSiz, 1841

Subdivision ElopomorphaOrder Albuliformes

Branneriun latum (aGaSSiz, 1841)Brannerion vestitum (Jordan and Branner, 1908)Paraelops cearensis SantoS, 1971

Bullichthys santanensis maYrinCk, Brito and otero, 2010ClupeocephalaSubdivision Ostarioclupeomorpha

Order GonorynchiformesChanidaeTharrhias araripis (Jordan and Branner, 1908)

Clupeocephala order and family incertae sedisBeurlenichthys ouricuriensis FiGueiredo and Gallo, 2004

Clupeomorpha order and family incertae sedisSantanaclupea silvasantosi maiSeY, 1993

Otophysi family incertae sedisSantanichthys diasii (sAntos, 1958)

Teleostei insertae sidesAraripichthydae Araripichthys castilhoi SantoS, 1985

Class SarcopterygiiOrder Coelacanthiformes

MawsoniidaeAxelrodichthys araripensis MAisey, 1986Mawsonia brasiliensis YaBumoto, 2008


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

Division HolosteiSubdivision Ginglymodi

Order SemionotiformesSemionotidaeAraripelepidotes cf. temnurus (AgAssiz, 1841)

Order LepisosteiformesObaichthyidaeObaichthyidae gen. et sp. indet.

Subdivision HalecomorphiOrder Amiiformes

AmiidaeCalamopleurus cylindricus AgAssiz, 1841Cratoamia gondwanica Brito, YaBumoto and Grande, 2008

Order IonoscopiformesOphiopsidaePlacidichthys bidorsalis Brito, 2000

Superdivision TeleosteomorphaOrdre Aspidorhynchiformes

AspidorhynchidaeBelonostomus sp.

Division TeleosteiOrder Ichthyodectiformes

CladocyclidaeCladocyclus gardneri Agassiz, 1841

Subdivision OstarioclupeomorphaOrder Gonorynchiformes

ChanidaeDastilbe crandalii Jordan, 1910

Otophysi family incertae sedisSantanichthys diasii (sAntos, 1958)

Class SarcopterygiiOrder Coelacanthiformes

MawsoniidaeAxelrodichthys araripensis MAisey, 1986

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