BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, researchlibraries, and research funders in the common goal of maximizing access to critical research.

A New Species of Araripichthys (Teleostei, Elopocephala) from the TlayúaFormation (Cretaceous, Albian), MexicoAuthor(s): Jesús Alvarado-Ortega and Paulo M. BritoSource: Journal of Vertebrate Paleontology, 31(6):1376-1381. 2011.Published By: The Society of Vertebrate PaleontologyURL: http://www.bioone.org/doi/full/10.1080/039.031.0603

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, andenvironmental sciences. BioOne provides a sustainable online platform for over 170 journals and books publishedby nonprofit societies, associations, museums, institutions, and presses.

Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance ofBioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use.

Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiriesor rights and permissions requests should be directed to the individual publisher as copyright holder.

Journal of Vertebrate Paleontology 31(6):1376–1381, November 2011© 2011 by the Society of Vertebrate Paleontology

SHORT COMMUNICATION

A NEW SPECIES OF ARARIPICHTHYS (TELEOSTEI, ELOPOCEPHALA) FROM THE TLAYUAFORMATION (CRETACEOUS, ALBIAN), MEXICO

JESUS ALVARADO-ORTEGA*,1 and PAULO M. BRITO2; 1Departamento de Paleontologıa, Instituto de Geologıa, UniversidadNacional Autonoma de Mexico, Ciudad Universitaria, Coyoacan, D. F., C.P. 04510, Mexico, alvarado@geologia.unam.mx;2Departamento de Zoologia, Instituto de Biologia, Universidade do Estado do Rio de Janeiro, Sao Francisco Xavier 524, Maracana,Rio de Janeiro 20559-900, Brasil, pbritopaleo@yahoo.com.br

Among the fish faunal components related to the Creta-ceous Tethys Sea western domain, Araripichthys castilhoi wasdescribed by Santos (1985; also see Santos, 1983) based onseveral specimens from the Santana Formation, Araripe Basin,Brazil. This author placed his deep-bodied fish within the OrderBeryciformes as a single member of his suborder and family,Araripichthyoidei and Araripichthyidae, respectively. Laterother species of Araripichthys were described. An incompletespecimen discovered in the Turonian marine sediments at Goul-mima, Morocco, was described as Araripichthys corythophorusCavin, 1997a. Description of Araripichthys axelrodi Maisey andMoody, 2001, is based on another isolated specimen from Aptianmarine sediments of the Apon Formation at Rosarito Quarry,near Villa del Rosario, Venezuela.

The taxonomical position given to Araripichtys also has beenchanged. Patterson (1993:627) vaguely suggested its possible in-clusion among the pachyrhizodontids. Later Maisey and Blum(1991) considered Araripichthys as incertae sedis taxon withinElopocephala, mainly because its caudal endoskeleton resem-bles those of the generalized elopocephalan fishes and it lacksthe diagnostic characters of higher teleost groups. Other au-thors have followed this last proposal (Cavin, 1997; Arratia andChorn, 1998; Maisey and Moody, 2001; Blanco-Pinon, 2003).More recently, Cavin (2001) considered this genus as a mem-ber of Protobramoidei (also see Cavin and Blanco, 2003; andTaverne and Gayet, 2004), which is supported in the absenceof pelvic fins, the occurrence of stout neural spines, and scalesat least covering the dorsal fin. Currently, none of these hy-potheses have been reassessed with new arguments or fos-sil findings; therefore the authors assume that the position ofthe genus Araripichthys is still uncertain among Elopocephalafishes.

Although fossil record of Araripichthys is comparativelyscarce and poor, the morphological diversity of this genus, aswell as its temporal and spatial distribution, is consistent withthe phylogenetic and biogeographical hypotheses suggestedby Maisey and Moody (2001). According to these authors, theVenezuelan species A. axelrodi is the earliest and the most basalrepresentative of this genus and Albian is the minimum agefor divergence of the Brazilian and Moroccan lineages. Suchdistribution of species belonging to Araripichthys suggests adispersal process within the Tethys Sea as a consequence of amid-Cretaceous transgression that allowed a wide distributioninto the Caribbean area, along the shallow epicontinental marinecarbonate platforms of Western Gondwana, and the Brazilianinterior basins (the Araripe Basin among others). Therefore,any additional data on Araripichthys fossil record is desirable, inorder to test these hypotheses.

Nowadays, two Lauransian records of Araripichthys discov-ered in marine Cretaceous Mexican localities achieve our knowl-

*Corresponding author.

edge on the biogeography of these fishes. On one side, Blanco-Pinon (2003) and Blanco and Cavin (2003) reported the discoveryof Araripichthys from the Turonian marine sediments of AguaNueva Formation at Vallecillo Quarry, Nuevo Leon State,Mexico. Despite the importance of this finding, these fossils fromVallecillo are not described in detail. On the other side, afteralmost 30 years of careful and systematical fossil collecting in theAlbian limestones of Tlayua Quarry, near Tepexi de Rodrıguez,Puebla, Mexico, a single and fragmented fossil belonging toAraripichthys has been finally identified. Because this specimenfrom Tlayua has features that do not match the diagnosticscharacters of the nominal species within the genus Araripichthys,the aim of the present paper is to describe this fossil as a newspecies.

Although the fossil fish assemblage of Tlayua Quarry is mostlyrepresented by Actinopterygii, by far this is the most importantCretaceous fossil fish locality in Mexico due the noticeablediversity, high richness, and extraordinary conservation of itsfossils (Applegate et al., 2006). The important efforts recentlygiven to fossil prospecting and collecting in new fish localitiesinto the country are providing unexpected results, which allowcomplementing our knowledge on the history and biology ofthis group of vertebrates along the western Tethys Sea domainduring the early Cretaceous. Largely, these Mexican fish assem-blages require to be studied more closely; however, these areimportant because they contain a peculiar association involvingthe following: (1) The youngest and first American records offishes belonging to families or groups well represented in Jurassicstrata deposited under the eastern Tethys Sea domain, todaywidely exploited in the other side of the world along Europe(i.e., Macrosemiidae, Ophipsidae, Ionoscopiformes, Semiono-tiformes, Aspidorynchiformes, among others). (2) Typical taxaalready known among the Early-”middle” Cretaceous localitiesdeposited under the domain in both sides of the Tethys Sea,the eastern and western, which today we are still discoveringin South America, Africa, Europe, Middle East, and Australia(i.e., Ellimichthyiformes, Pycnodontiformes, Ionoscopiformes,Aulopiformes, Ichthyodectiformes, among others). (3) Fishesknown from North American strata deposited into the InteriorSea Way (i.e., Enchodontoidei, Ichthyodectiformes). In thissense, the Mexican fossil fish localities are some of the missingpieces within the paleontological puzzle that requires to beanalyzed for a comprehensive understanding on the evolutionaryhistory of Cretaceous fishes.

The rich fossiliferous limestones exploited at Tlayua Quarrywere described as the Middle Member of the Tlayua Formationby Pantoja-Alor (1992). This lithological unit is interpreted asan Albian (Benammi et al., 2006) epicontinental marine deposit,within a basin under restricted and shallow conditions or into anopen marine basin, with weak continental influence (see Apple-gate et al., 2006, and Suarez, et al., 2009, to complement the geo-logical information).

1376

SHORT COMMUNICATIONS 1377

FIGURE 1. IGM 6232, Araripichthys weberi holotype; A, complete specimen; B, close-up of jaws showing the diagnostic species characters; C,granular ornamentation on the opercle and preopercle surfaces; D, scales showing their posterior fringed borders. Abbreviations: ang, angular; apn,anterior premaxilla notch; den, dentary; le, lateral ethmoid; mx, maxilla; op, opercle; pmx, premaxilla; pop, preopercle; psp, parasphenoid; smx,supramaxilla; rar, retroarticular; left or right bone positions are noted as (l) and (r), respectively.

MATERIALS AND METHODS

Institutional Abbreviations—IGM Coleccion Nacional dePaleontologıa, Instituto de Geologıa, Universidad NacionalAutonoma de Mexico; UERJ PMB, Colecao de Paleontologia,Universidade do Estado de Rio de Janeiro, Brazil.

Anatomical Abbreviations—Abbreviations in Figures 1 and 2follow Maisey and Moody (2001).

Referred Materials—Materials reviewed in this work withcomparative propose. Araripichthys castilhoi: UERJ PMB 105,head and abdominal are of a fish transferred to resin and onlypartially prepared with formic acid; head length 80 mm; Cancao,near Santana do Carirı, Ceara, Brazil; Albian marine sedimentsof the Santana Formation. UERJ PMB 106, almost completespecimen without large part of the dorsal and anal fins; stan-dard and head lengths are 258 and 80 mm respectively; fromCancao, near Santana do Carirı, Ceara, Brazil. UERJ PMB108, part (A) and counterpart (A) of a incomplete specimenlacking the tip of jaws and the dorsal, anal, and caudal fins;estimated standard length about 95 mm; from Santana For-mation, Brazil. UERJ PMB 109, specimen lacking the nasalarea; standard length about 330 mm; from Santana Formation,Brazil.

Araripichthys corythophorus: UERJ PMB 107, part (A) andcounterpart (B) of the head and anterior part the abdomen; cal-careous nodule partially prepared mechanically; Lower Turonianmarine strata near Asfla, Atlas Mountains, Goulmima, Morocco.

SYSTEMATIC PALAEONTOLOGY

Family ARARIPICHTHYIDAE Santos, 1985Genus ARARIPICHTHYS Santos, 1985ARARIPICHTHYS WEBERI, sp. nov.

Holotype—IGM 6232, part of the head and abdominal regionof a specimen showing its lateral left surface; its estimated stan-dard length is about 300 mm; the specimen was transferred toresin and prepared with acetic acid baths.

Occurrence—Tlayua Quarry limestones, near Tepexi deRodrıguez, Puebla State, central Mexico; Middle Member of theTlayua Formation, Early Cretaceous, Albian.

Etymology—The species name honors our colleague Dr. Rein-hard Weber Gobel (1938–2009) for his important contributions tothe Mexican paleontology.

Diagnosis—Araripichthys fish characterized by the occurrenceof three exclusive characters involving the obtuse angle (about100◦) between the ascending process and horizontal arm of thepremaxilla; the notch in the anterior end of the premaxilla alve-olar border; and the intense granular ornamentation on the pre-opercle and opercle. This fish also has a particular combinationof characters found in the other species of the genus, these in-clude the cycloid scales with posterior border fringed; elongatedpremaxillary ascending process accounting for at least the half ofpremaxilla total length; maxilla slightly curved and almost uni-formly wide, with its posterior end slightly wider than the middle

1378 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 6, 2011

FIGURE 2. Reconstruction of IGM 6232, Araripichthys weberi, sp. nov. holotype, based on Figure 1A. Abbreviations: ang, angular; apn, anteriorpremaxilla notch; br, branchisotegal; c, vertebral centra; cha, anterior ceratohyial; cl, cleithrum; cor, coracoid; den, dentary; exo, exoccipital; fr, frontal;hhd, hypohyal dorsal; hhv, hypohyal ventral; hm, hyomandibular; io, infraorbital; iop, infraopercle; le, lateral ethmoid; mx, maxilla; op, opercle; pfr,pectoral fin ray; pmx, premaxilla; pop, preopercle; psp, parasphenoid; pto, pterotic; ptsp, pterosphenoid; q, quadrate; r, rib; rar, retroarticular; sca,scapular; smx, supramaxilla; sob, suborbital; v, vomer; left or right bone positions are noted as (l) and (r), respectively.

SHORT COMMUNICATIONS 1379

and anterior sections; and maxilla with a long and strong articularrod-like condyle.

Description

General Features—The poor preservation of IGM 6232 pre-vents the recognition of a lot of its measurements and body pro-portions; it is only possible to measure its head length of 93mm. On the basis of the body proportions on the more com-plete Araripichthys castilhoi specimen illustrated by Maisey andBlum (1991:213, 217) and Maisey and Moody (2001:fig. 4), andspecimens UERJ PMB 106 and109 analyzed here, head length ofwhich is about 33% of the standard length (SL), it is possible toestimate that IGM 6232 had a SL about 300 mm.

Skull—The skull roof of the specimen is almost lost; it pre-serves only part of the left frontal that has a smooth surfacebut some small radiating ridges and pores located along orbitaledge; a deep longitudinal depression is present in the central an-terior part of this bone. Along its posterior border, the frontaljoins a fragment of the sphenotic bone that also forms the rearof the orbit. Behind the sphenotic there is another bone withrough surface, here interpreted as a possible part of the exoc-cipital. The pterosphenoid is a rounded structure projected intothe orbit from its roof. The parasphenoid and the vomer are de-tached; they are equally thick and stout. Apparently the vomer istoothless and shows two wide, rounded anterior, projections thathouse maxilla condyles. A lateral ethmoid is preserved in IGM6232; it is a thick stout bone, is Y-shaped, and borders the orbitanteriorly.

Circumorbital Series—Bones of the circumorbital series areall displaced; at least four presumably infraorbitals are present.A suborbital cover a large surface of the lower jaw articulation.There is no evidence of any supraorbital bone. Perhaps scleroticbones fragments are also preserved.

Upper and Lower Jaws—Bones of upper and lower jaws (max-illa, premaxilla, and dentary) are totally toothless. Premaxilla isboomerang-shaped with a rounded, wide obtuse angle of about100◦. The height of the premaxillar ascending process is almostequal to the length of alveolar or horizontal limb of this bone,which seems to occupy all the upper border of the mouth. Thealveolar or horizontal limb of the premaxilla joins the ventrallimb of the maxilla, which is slightly curved, forming a ventralconcavity; thus even though the premaxilla preserved in IGM6232 is covered by the maxilla and supramaxillae, it is still pos-sible to estimate its length. In this specimen two supramaxillaeare preserved, the first is almost totally covered by a huge trian-gular supramaxilla. The alveolar border of the premaxilla showsan anterior small, rounded notch.

The maxilla is a rectangular structure with regular height andcurved tips. Its stout anterior condyle has a well-developed rod-like anterior process, its middle section is almost straight butshows a ventral curvature where the premaxilla might be placed,and its posterior third section is slightly expanded, curved, androunded.

Dentary bone is V-shaped, where the rectangular alveolar sec-tion and coronoid process form a straight angle. In IGM 6232,the angular and retroarticular are exposed but the articular boneis practically obscured. The angular is rectangular but with ante-rior concave border. The wide retroarticular attaches the poste-rior corner of the angular and covers all the lower jaw postarticu-lar process. Dorsally, this process is gently curved and forms thedeep cavity that borders the articular facet for the quadrate.

Hyomandibular and Hyoid Arch—In IGM 6232 the bones ofthe hyopalatine series are not clear in the specimen except forthe hyomandibular, which has a stout opercular process. Someelements of the hyoid arches are displaced in the specimen. Bothanterior ceratohyals are exposed; these are ovoid structures witha rounded foramen slightly displaced dorsally. The ventral hypo-

hyal and a presumably dorsal hypohyal articulate with the ante-rior ceratohyal; the former is rectangular and the latter is trian-gular and smaller. There is only one C-shaped branchiostegal raypreserved near to ceratohyals, but at least there are other threebelow of the opercular series (Fig. 2).

Opercular Series—The preopercle is a flat bone with aboomerang shape that has a short horizontal anterior arm anda deep vertical arm. Its anterior border is thick, but it is widelyexpanded posteriorly. The opercle is ovoid, almost twice higherthan wide. The surface of the preopercle and opercle show in-tense granular ornamentation. Subopercle and interopercle aresmooth bones (Figs. 1C, 2).

Pectoral Fin and Girdle—Bones of the pectoral girdle are alsopartially preserved in IGM 6232. The lateral surface of the clei-thrum is uniformly wide inverse C-shaped. Although the scapulaand coracoids are located below the horizontal section of thecleitrum, it is not possible to describe these (Figs. 1, 2).

The pectoral fin involves 16 large rays attached to ventral sur-face of the coracoids. The most anterior ray is spine-like, about45 mm length, unsegmented, and unbranched. It is followed bythe six largest rays, which reach about 80 mm in length and arestrongly branched and segmented. The posterior pectoral fin raysshow the same aspect of the largest rays, but their length progres-sively decreases in anterior-to-posterior order (Figs. 1A, 2).

Vertebral Column—Elements of the vertebral column are al-most totally missing. There are ventral fragments of three abdom-inal centra above the opercle, showing lateral cavities. Long ribsalmost enclose the anterior abdominal cavity; these are uniformlythin and slightly wider near the top (Figs. 1A, 2).

Scales—All scales of the specimen are deeply overlappedto each other. In the abdominal region they are heart-shaped,notched anteriorly, and slightly longer than high. The externalsurface of scales is ornamented with numerous concentric circulliand the posterior border is intensely fringed (Fig. 1D).

DISCUSSION AND CONCLUSIONS

The following combination of characters, based on theemended generic diagnoses provided by Maisey and Moody(2001), allows recognizing IGM 6232 as a new member of thegenus Araripichthys. In this large Araripichthys fish, where thestandard length may reach 300 mm, the jaw bones (premaxilla,maxilla, and dentary) and the vomer are toothless; the premax-illa shows a prominent ascending anterodorsal process; there aretwo supramaxillae above the maxilla; and the cycloid scales aredeeply overlapped to each other (see Description and Figs. 1, 2).

Maisey and Moody (2001:4) also provide diagnoses for thethree species already known of Araripichthys (including thoseemended for A. castilhoi and A. corythophorus, and the originalfor their A. axelrodi). Regarding those diagnoses, it is possible torecognize that IGM 6232 represents a new species here named asAraripichthys weberi, because it shows a unique combination ofcharacters in the maxilla, premaxilla, and scales.

The premaxilla ascending process is larger than half of the totallength of this bone in A. weberi and A. castilhoi, whereas it isabout one third in A. corythophorus and less than a third in A.axelrodi (pmp in Fig. 3). On the other hand, the maxilla articularcondyle is absent in A. axelrodi and well developed in the otherspecies (ac in Fig. 3).

Additionally, in A. weberi the premaxilla shows two character-istics not seen in any other of these nominal species. This Mex-ican fossil has a small notch in the alveolar border anterior end(apn in Figs. 1B, 2, 3), and its horizontal and vertical premax-illa arms form a rounded obtuse angle (about 100◦). In contrast,this angle is almost straight (a little over 90◦) in other species (A.castilhoi and A. axelrodi) with a premaxillar ascending process(see Maisey and Moody, 2001:figs. 3, 5–7).

1380 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 6, 2011

FIGURE 3. Possible relationships among Araripichthys species (modi-fied after Maisey and Blum, 2001:fig 7, and including the possible positionof A. weberi). Node 1: Ascending process of premaxilla (pmp) slightly lessthan half of premaxilla total length; articular condyle of maxilla (ac) welldeveloped. Node 2: Ascending process of premaxilla (pmp) larger thanhalf of premaxilla total length. Node 3: Maxilla rectangular in shape (pos-terior section less high). Node 4: Wide, rounded abtuse maxillar angle;anterior premaxilla notch developed.

In addition, in Araripichthy costilhoi and A. axelrodi, the max-illa is almost triangular, its anterior tip is acute, and the poste-rior section is wider and rounded (Fig. 3). On the other hand, themaxilla is more rectangular in A. weberi and A. corythophorus,because its posterior section is just slightly higher than the rest ofthe bone. Other peculiar character in Araripichthys weberi is theposteriorly fringed heart-shaped cycloid scales (Fig. 1D), whichdiffers from other nominal species that have scales with four tosix straight and parallel posterior radii (not described for A. axel-rodi) instead the fringed border.

It is clear that Araripichthys weberi and A. castilhoi are closerelatives because they share an extremely large premaxilla as-cending process; nevertheless, the maxilla of the new Mexicanspecies is almost rectangular, resembling that drawn for A. cory-thophorus (Fig. 3). In this scenario, it is probable that A. weberirepresents a member into the crown group of the phylogeneticscheme previously suggested by Maisey and Moody (2001), anddirectly related to A. casthiloi or A. corythophorus.

The phylogenetic hypothesis of the species witin Araripichthys,suggested by Maisey and Moody (2001), requires to be tested,through the application of a formal cladistic algorithm. Thisalso needs to be updated with the addition of specimens be-longing to Araripichthys from the Vallecillo Quarry reported byBlanco-Pinon (2003) and the rare fish Ferrifrons rugosus Jordan,1923. The latter is a North American Turonian marine species,previously reviewed by Arratia and Chorn (1998), who placedit in the monogeneric family Ferrifronsidae within the acan-toptomorphs. Unfortunately, a direct comparison between theonly, highly incomplete single specimen of Ferrifrons and thespecies of Araripichthys is not easy, because the former lacksthe jaws, where important and specific morphological variabilityof species within Araripichthys has been recognized. AlthoughMaisey and Moody (2001) highlighted some of the shared fea-tures of Araripichthys and Ferrifrons, they did not include themin a cladistic analysis.

The discovery of Mexican fossils belonging to Araripichthys inAlbian (in Tlayua Formation) and Turonian (in Vallecillos) ma-rine sediments allows us to recognize the possibility that thesefishes are related to Ferrifrons. The new Mexican specimens willalso allow us to explore the phylogenetic and biogeographic re-lationships of these unusual fishes. The occurrence of A. weberiin Mexico clearly demonstrates that the geographic distributionof Araripihthys was wider, at least since the Albian, and involvesthe epicontinental and marginal marine areas under the westernTethys Sea domain in both Gondwana and Laurasia.

The evolutionary history and biogeography of the genusAraripichthys could be more complex than described in thispaper. According to L. Cavin (pers. comm., June 2011), thereare Cenomanian “true Araripichthys” specimens from Lebanonhoused in the Museum national d’Histoire naturelle (Paris)that still require an accurate description and taxonomicaldetermination.

ACKNOWLEDGMENTS

The fulfillment of this paper was made possible by the effortof the Aranguthy family and their Tlayua quarry workers. Weare deeply indebted to Dr. M. del C. Perrilliat, who kindly facil-itated our work within the Coleccion Nacional de Paleontologıa,UNAM, and Mr. G. Alvarez for the preparation of the speci-men described in this work. Our friend, Dr. L. P. Machado, is theauthor of the photos published here. We appreciate the techni-cal support by A. M. Rocha. The manuscript was improved onthe basis of the valuable commentaries by Ch. Underwood, J.Maisey, and L. Cavin. J.A.O. is supported by UNAM-PAPIITIN225008 and IN106011 projects. P.M.B. is supported by CNPqand Prociencia/FAPERJ fellowships.

LITERATURE CITED

Alvarado-Ortega, J., K.A. Gonzalez-Rodrıguez, A. Blanco-Pinon, L.Espinosa-Arrubarrena, and E. Ovalles-Damian. 2006. Mesozoic Os-teichthyans of Mexico; pp. 169–207 in F. J. Vega, T. G. Nyborg,M. del, C. Perrilliat, M. Montellano-Ballesteros, S. R. S. Cevallos-Ferriz, and S. A. Quiroz-Barroso (eds.), Studies on Mexican Paleon-tology. Topics in Geobiology 24. Springer, Dordrecht, The Nether-lands.

Arratia, G., and J. Chorn. 1998. A new primitive acanthomorph fish fromthe Greenhorn Formation (Late Cretaceous) of Nebraska. Journalof Vertebrate Paleontology 18:301–314.

Benammi, M., J. Alvarado-Ortega, and J. Urrutia-Fucugauchi. 2006.Magnetosratigraphy of the Lower Cretaceous strata in TlayuaQuarry, Tepexi de Rodriguez, State of Puebla, Mexico. Paleomag-netism and Tectonics in Latinoamerica, Earth Planets Space 58 (spe-cial issue):1295–1302.

Blanco-Pinon, A. 2003. Peces fosiles de la Formacion Agua Nueva (Tur-oniano) en el municipio de Vallecillo, Nuevo Leon, NE de Mexico.Tesis de Doctorado, Facultad de Ciencias de la Tierra, UniversidadAutonoma de Nuevo Leon, Linares, Mexico, 330 pp.

Blanco, A., and L. Cavin. 2003. New Teleostei from the Agua NuevaFormation (Turonian), Vallecillo (NE Mexico). Comptes RendusPalevol 2:299–306.

Cavin, L. 1997. Nouveaux Teleostei du gisement du Turonien inferieurde Goulmima (Maroc). Comptes Rendus, Academie des Sciences325:719–724.

Cavin, L. 2001. Osteology and phylogenetic relationships of the teleostGoulmimichthys arambourgi Cavin, 1995, from the Upper Cre-taceous of Goulmima, Morocco. Eclogae Geologicae Helvetiae94:509–535.

Jordan, D. S. 1923. A collection of fossil fishes in the University of Kansas,from the Niobrara Formation of the Cretaceous. Kansas UniversityScience Bulletin 15:219–245.

Maisey, J. G., and S. Blum. 1991. Araripichthys; pp. 208–217 in J. G.Maisey (ed.), Santana Fossils: An Illustrated Atlas. TFH Publica-tions, Neptune, New Jersey.

SHORT COMMUNICATIONS 1381

Maisey, J. G., and J. M. Moody. 2001. A review of the problematic extinctteleost fish Araripichthys, with description of a new species fromthe Lower Cretaceous of Venezuela. Americam Museum Novitates3324, 27 pp.

Pantoja-Alor, J. 1992. Geologıa y paleoambientes de la Cantera Tlayua,Tepexi de Rodrıguez, Estado de Puebla, Universidad NacionalAutonoma de Mexico, Instituto de Geologıa, Revista 9:156–176.

Patterson, C. 1993. Teleostei; pp. 621–656 in M. J. Benton (ed.), The Fos-sil Record 2. Chapman and Hall, London.

Santos, R. da S. 1983. Araripichthys castilhoi novo genero e especie deacantopterıgio da Formacao Santana, Chapada do Araripe, Brasil;p. 27 in VIII Congresso Brasileiro de Paleontologia, Rio de Janeiro,Resumos.

Santos, R. da S. 1985. Araripichthys castilhoi novo genero e especiede teleostei da Formacao Santana, Chapada do Araripe, Brasil;

pp. 133–140 in D. A. Campos, C. S. Ferreira, I. M. Brito, andC. F. Viana (eds.), Coletanea de Trabalhos Paleontologicos, SerieGeologia 27(2). Ministerio das Minas e Energia-DNPM, Rio deJaneiro.

Suarez, M. B., L. A. Gonzalez, G. A. Ludvigson, F. J. Vega, and J.Alvarado-Ortega. 2009. Isotopic composition of low-latitude pale-oprecipitation during the Early Cretaceous. Geological Society ofAmerica Bulletin 121:1584–1595.

Taverne, L., and M. Gayet. 2004. Osteologie et relations phylogenetiquesdes Protobramidae (Teleostei, Tselfatiformes) du Cenomanien(Cretace Superieur) du Liban. Cybium 28:285–314.

Submitted March 29, 2011; revisions received June 21, 2011; acceptedJune 24, 2011.Handling editor: Charlie Underwood.