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Home , Amiidae, Axelrodichthys, Calamopleurus, Cladocyclus, Crato Formation, Dastilbe

ORYCTOS.Vol. 3 : 3 - 8. Décembre2000

FIRSTRECORD OT CALAMOPLEU RUS (:: ) FROMTHE (LOWER ) OF NORTH-EAST BRAZTL

David M. MARTILL' and Paulo M. BRITO'z

'School of Earth, Environmentaland PhysicalSciences, University of Portsmouth,Portsmouth, POl 3QL UK. 2Departmentode Biologia e Vegetal,Universidade do Estadode Rio de Janeiro, rua SâoFrancisco Xavier 524. Rio de Janeiro.Brazll.

Abstract : A partial skeleton representsthe first occurrenceof the amiid (Actinopterygii: Halecomorphi: Amiidae) from the Nova Olinda Member of the Crato Formation () of north east . The new spe- cimen is further evidencethat the Crato Formation ichthyofauna is similar to that of the slightly younger Romualdo Member of the Santana Formation of the same sedimentary basin. The extended temporal range, ?Aptian to ?,for this rules out its usefulnessas a biostratigraphic indicator for the .

Key words: Amiidae, Calamopleurus,,Brazil

Première mention de Calamopleurus (Actinopterygii: Halecomorphi: Amiidae) dans la Formation Crato (Crétacé inférieur), nord est du Brésil

Résumé : la première mention dans le Membre Nova Olinda de la Formation Crato (Aptien ; nord-est du Brésil) de I'amiidé (Actinopterygii: Halecomorphi: Amiidae) Calamopleurus est basée sur la découverted'un squelettepar- tiel. Le nouveau spécimen est un élément supplémentaireindiquant que I'ichtyofaune de la Formation Crato est similaire à celle du Membre Romualdo de la Formation Santana, située dans le même bassin sédimentaire. L'extension temporelle de ce genre (?Aptien à ?Cénomanien)ne permet pas de le considérer comme un indicateur biostratigraphiquepour le bassin de l'Araripe.

Mots clés : Amiidae, Calamopleurus, Crétacé inférieu4 Brésil

INTRODUCTION Araripina and at Mina Pedra Branca, near Nova Olinda where cf. sp. and Vnctifur sp. have are very common in the Nova been collected from laminated black shalesbeneath Olinda Member of the Crato Formation where the massive evaporite deposits. However, although ichthyofaunais dominated(>95Vo) by the gonorhyn- Dastilbe crandalli is exceptionally abundantin the chiform Dastilbe crandalli Jordan, 1910 (Davis & Crato Formation, it has not been certainly reported Martill, 1999). Other speciesinclude the ich- from the Romualdo Member of the Santana thyodectid sp. Maisey (1996) which Formation, and as a result the two fish faunashave occurs rarely and single occuffences only of the been considereddistinct (Maisey, 1991; Martill, semionotid cf. Araripelepidotes (Brito et aI., 1998) 1993; Davis & Martill, 1999).Here we report on the and the sp. (Brito & occurrenceof an incomplete specimenof an amiid Martill, 1999). These rarer taxa are well known and resembling Calamopleurus from the Crato relatively corrmon in the slightly youngerRomualdo Formation: a that is common in the Romualdo Member of the SantanaFormation in the samebasin. Member of the SantanaFormation. Fishesare alsoknown from the Ipubi Formationnear ORYCTOS,Vol.3,2000

LOCALITY AND STRATIGRAPITY

The new specimenwas obtainedfrom a quarry ,t l-,it.i ! worker who reported that it was collected from the Exu Formation laminated limestonesworked in the region of Nova i.;.14.; Olinda and Santanado Cariri, southernCearâ,BrazlI * ?Cenomanian SantanaFormation (fig 1). More precise locality details were unavai- (nodulebeds) ? lable. The matrix is entirely consistentwith a deriva- a< tion from this region, where numerousquarries are lpubiFormatron worked for building, paving and ornamental stone (Martill, 1993). The limestonesworked in this region were ter- CratoFormation , Lower med the Nova Olinda Member by Martill (1993),and 'Aptian they form a distinctive lithological unit of between NovaOlinda Member -1 and 11.5 metres lying at the base of the Crato Formation. These limestonesare generally conside- BataleirasFmn. red to be of Aptian age on palynological grounds (Ponset a1.,1990)and lie within a sequenceof sand- stones, clays, shales and siltstones of the Araripe Group (Fig 2). They have been consideredto repre- sent freshwater lacustrine environmentsby Maisey Figure2. Simplifiedstratigraphic column for theAraripe Group of (1996) whereasMartill (1993) and Davis & Marrill the Araripe Basin.The new specimensof cf. Calamopleurusare (1999) suggesta stratified hypersaline/anoxiclagoo- from the Nova OlindaMember of the CratoFormation. Fish svm- nal systemfor their accumulation. bols indicatehorizons known to vield fishes.

Figure L Map showing the region from where the new fossil came (marked with bold x). There are numeroussmall quarriesfor ornamentalstone on the outcrop of the Nova Olinda Member between Nova Olinda, Santanado Cariri and Tatajuba.Working quanies mentionedby Martill (1993) at Crato have now been abandoned.ish symbols indicate localitieswhere fish have been collected from the Crato Formation. MARTILL & BRITO -AMIID FROMBRAZIL

SYSTEMATIC PALAEONTOLOGY is assignedto the tribe Calamopleurinion accountof the coronoid bones possessingonly a single row of SubclassACTINOPTERYGtr Cope, teeth.In addition the arrangementof the teeth on the 1887(sensu Rosen et a1.,1981) vomers (e.g. an anterior row of five teeth, followed SeriesNEOPTERYGtr Regan,1923 by a row of three teeth) matchesthat describedby (sensuRosen et a1.,1981) Grande& Bemis (1998) for Calamopleurini. Division Regan, 1923 Further identification is difficult as the new spe- (sensuPatterson, I97 3) cimen doesnot display featuresthat allow us to dis- SubdivisionHALECOMORPHI Cope, 1872 tinguish between the three included in (sensuPatterson, 197 3) Calamopleurzs. Two speciesof Calamopleurus hav e Hay 1929 beenreported from the Early CretaceousofBrazil. C. (sensuGrande & Bemis,1998) cylindricus from the Romualdo Member of the Family AMMIIDAE Bonaparte,1838 SantanaFormation was describedin detail by Santos Subfamily VIDALAMIINAE Grande& Bemis, (1960), Taverne (1974), Wenz (1977) and more 1998 recently by Grande & Bemis (1998), while its Tribe CALAMOPLEURINI Grande& Bemis. 1998 nomenclatural history was reviewed by Maisey Genuscf. CalamopleurusAgassiz, 1841 (1991). C. cyclindrfczs (commonly found in the lite- rature as Ennelesaudax) is known from many hun- dreds of complete or near complete specimens(see THE NE\ry SPECIMEN Grande& Bemis 1998for an extensivelist of figured and examinedmaterial). A secondBrazilian species, The new specimenis accessionedin the collec- C. mawsoni (Woodward, 1902) from the Ilhas tion of the Museo do Santanado Cariri, specimen Formation (Early Cretaceous,Bahian local stage)of numbersMPSC (Fig. 3). The fish skeletonlies on its Bahia is known from only a single specimenlacking left side (as viewed) on a slab of cream coloured, a (see Grande & Bemis 1998 for a complete laminated limestonemeasuring 380 mm x 120 mm. review of this taxon) and cannot be comparedwith The slab is fractured with the fracture extending the new specimendescribed herein. A caudal skele- through the fossil. Although the skeleton is incom- ton attributed to Calamopleurussp. was described plete, the bonesare well preserved.The bonesof the from the Aptian-Albian marine beds of Equatorial trunk are only slightly crushed, although there is Guinea (Taverne, t997). A third species, much flattening of the skeletonsuch that the speci- Calamopleurusafricanzs Forey & Grande (1998) men is somewhat2D. has beendescribed from north westAfrica. This spe- The skeletoncomprises the skull roof, left lower cies is certainly distinct from C. cylindricus, but a jaq and anteriortwenty-two vertebraeand ribs of the synonymy with C. mawsoni cannot be ruled out. trunk (Fig. 3). The roof of the skull and braincaseis Assignmentof the specimendescribed here to a spe- seenfrom internally.Teeth are present in the left den- cies is rendered slightly difficult because of its tary but arelargely obscuredby matrix. Teethare also incompletenature as well as the incompletenature of presenton the left premaxilla (5 large teeth), the left the holotype of C. mawsoni.C. cylindricus caîreadi- maxilla (5 small teeth seen) and the vomer (a few ly be distinguishedfrom C. africanus on accountof small teeth).The left preopercleand parts of the left its broader,smooth gular; that of C. africanus is nar- opercleand subopercleare present. The anterior-most row and possessesan anterior carina (Forey & part of the vertebral column comprising twenty-two Grande,1998). vertebrae with several neural arches and pleural ribs presentis preservedstill attachedto the skull. There is no trace of the pectoralgirdle or fins and there are no scalespreserved. The systematicclassification follows Grande & Bemis, 1998.The new specimenis an amiiform and ORYCTOS.VoI.3.2000

Figure 3. Partial skeleton of cf. Calamopleurus sp. from the Nova Olinda Member of the Crato Formation. comprising an incomplete skull and anterior portion of the vertebral column. Specimen LIEzu PMB VP-88. A. Entire specimen, scalebar = 80 mm. B. Detail of skull, scale bar = 40 mm.

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DISCUSSION Taphonomic considerations (the specimen is incomplete) suggestthat it may not have inhabited The occurrence of cf. Calamopleurus in the the and is an allochthonousoccurrence. The Nova Olinda Member of the Crato Formation is fur- taphonomic condition hints at some post-mortem ther evidence for the similarity of its ichthyofauna drifting and decay. Thus the specimen of with that of the Romualdo Member of the Santana Calamopleurusin the Nova Olinda Member may Formation(Martill 1988,Maisey 1991).Abundance represeht an exotic transported from marine data (only one specimenof Calamopleurusreported (Calamopleurusin the RomualdoMember is thought from the Crato Formation comparedwith hundreds to be marineor brackish(Maisey, 1991)) or more fre- from the Romualdo Member of the Santana shwater environments(most post-Mesozoicamiids Formation) suggeststhat Calamopleuruswas rare in arefreshwater or from freshwaterdeposits (Grande & the Crato lagoon. Bemis,1998)) adjacent to the Cratolagoonal system. MARTILL & BRITO - AMIID FROM BRAZIL

Not withstanding the similarities between the ichthyofaunas of the Crato and SantanaFormations GENUS CRATOFORM'NIPTEIFORM'NSANTANAFORM'N NovaOlinda Mbr RomualdoMember (see Table 1), there remain notable differences.All fish generaother than Dastilbe are exceedinglyrare Iansan * in the Crato Formation, while many speciesthat are * abundant in the SantanaFormation remain to be Tribodus reported from the Crato Formation; e.g Vnctifer, Araripelepidotes *. , , B.rannerion, Neoprocinetes, Tharrias. Although these,differencesmay in part be Neoprocinetes t< attributedto age,it can be presumedthat dispa- some Iamanja {c rity is due to palaeoecologicalrather than preserva- tional factors, as a wide range of vertebratefossils Obaichthys * occurin an excellentstate of preservationin the Nova Calamopleurus Olinda Member. Such palaeoenvironmentalfactors are likely to be related to salinity and/or oxygen Oshunia *( content.We suggestthat the very high abundanceof Vinctiftr * + the single taxon Dastilbe crandalli in the Crato lagoon is evidencefor elevatedsalinities rather than Cladocyclus low salinities (see discussion in Davis & Martill, Araripichthys 1999). If this scenario is correct, then the Crato lagoon was probably a highly restricted system Brannerion T connectedto a normal marinebasin from which occa- sionally enteredmarine fishes. Much work remains Paraelops to done on the ichthyofaunas of the Crato and Rhacolepis Santana formation fishes to resolve some of the palaeoecologicalconundrums of stet fascinatingfos- Notelops sil lagerstâtte. Tharrhias +

ACKNO\ryLEDGEMENTS Dastilbe 2*

Santanaclupea + Special thanks to Dr André Herzog (URCA, Crato) and Dr Artur Andrade (D.N.P.M. Crato) for their assistancein the field. Thanks to Mr Robert Axelrodichthys x + Loveridge (Portsmouth) for photography and Drs Lance Grande (Chicago), Lionel Cavin (Esperaza) Gen Nov. x and Mike Barker (Portsmouth) for critical reading of the manuscript.Also thanks to the 1999 Portsmouth BnziI expedition team. Table 1. Generaof fish from the Araripe Basin,north-east Brazil. Datafor SantanaFormation from Maisey(1991) and Matill (1993); data for Crato Formation from Brito et al. (1998) and this paper; datafor Ipubi Formationfrom recentfieldwork by Martill. ORYCTOS.VoI.3.2000

REFERENCES (eds),Interrelationships of fishes. Academic Press, London. AGASSZ, L. 1841.On the fossilfishes found by Mr Gardnerin the PONS,D., BERIHOU, P-Y.& CAMPOS,D. deA. 1990.Quelques Province of Cearâ, in the north of Brazil. Edinburgh New observationssur la palynologiede I'Aptien supérieuret de PhitosophicalJournal, 30 : 82-84. I'Albiendu Bassind'Araripe; pp.24l-252.1n CAMPOS, D. DE BONAPARTE,C. L. 1838.Selachorum tabula analytica. Nouvelles 4., VIANA, M. S. S., BRITO, P. M. & BEURLEN, G. (eds). Annalesdes Sciences Naturelles, 2. Atas do simposiosobre a Bacia do Araripe e BaciasInteriores BRITO, P. M. & MARTILL, D. M. 1999.Discovery of a juvenile do Nordeste,Crato, 14-16 de Junhode 1990: Crato,405pp. coelacanth in the Lower Cretaceous, Crato Formation, REGAN,C.T. 1923.The skeletonof. Lepidosteus, with remarkson NortheasternBrazTl. Cvbium.23 :209- 2lL. the origin and evolution of the lower neopterygian fishes. & WENZ. S. 1998.A Proceedingsof the hological Society,London, t923 : 445-461. semionotid fish from the Crato Formation (Aptian, Lower ROSEN,D. E., FOREY,P. L, GARDINER,B, G. & PATTERSON, Cretaceous)of Brazil: palaeoecologicalimplications. Oryctos, C. 1981., tetrapods, palaeontology and plesiomor- l:37-42. phy.Bulletin of theAmerican Museum of Natural History,167 : COPE,E. D. 1872.Observations on the systematicrelations of the 159-276. fishes. Proceedingsof the American Association for the SANTOS,R. da S. 1960.A posiçâosistemâtica de Ennelesaudax e Advancementof Science,20 : 317-343. Jordan Branner da Chapadado Araripe, Brasil. Monografia, DivisaoGeologia e Mineral, Brasil l7 t L- 25. 1887. Zittel's Manual of Palæontology[a review TAVERNE,L. 1974.Sur le premierexemplaire complet d'Enneles of this workl. American l0I4-10L9. Naturalist,2l : audaxlordan, D. S. et Branner,J. C., 1908(Pisces, , DAVIS, S. & MARTILL, D. M. 1999.The gonorynchiformfish Amiidae) du Crétacésupérieur du Brésil. Bulletin de la Société Dastilbe from the Lower Cretaceousof Brazil. Palaeontology, belgede Géologie,83: 61-71. 42:715-740. 1997.Sur la présencedu genre Calamopleurus(= (Pisces, FOREY, P. L. & GRANDE, L. 1998. An African twin to the Enneles) Actinopterygii,Amiidae) dansI'Ichthyofaune marine Éocrétacéede Guinée Équatoriale(Afrique). Mus. roy. BrazilianCalamopleurus (Actinopterygii: Amiidae). hological Afn centr, Tervuren(BeIg.), Dépt. GéoL.,Min., Rapp.Ann. 1995 Linnean Joumal of the Society,123: 179-195. &.1996:215- 220. GRANDE,L. & BEMIS,W. E. 1998.A comprehensivephylogene- V/OODWARD,A.S. 1902.On an amioidfrsh(Megalurus mawsoni, tic studyof amiid fishes(Amiidae) based on comparativeskele- sp. n.) from the Cretaceousof Bahia, Brazil. Annals and tal anatomy.An empirical searchfor interconnectedpatterns of Magazineof NaturalHistory, (ser 7),9 :87-89. natural history. Supplement to Journal of WENZ, S. 1977. Le squeletteaxial et I'endoesquelettecaudal d'Ennelesaudax, poisson Amiidé du Crétacéde Cear6(Brésil). Palaeontology,18, Memoir 4: l-690. Bulletin du MuseumNational d'Histoire naturelle.490 :341- HAY, O. P. 1929.Second bibliography and catalogueof the fossil 348. Vertebrata of . Publications of the Camegie Institute,Washington, 390 : 1-2003. JORDAN, D. S. 1910.Description of a collectionof fossil fishes from the bituminousshales of RiachoDoce, Stateof Alagôas, Brazll Annalsof the CarnegieMuseum,T :22-34. MAISEY, J. G. 1991.Santana : an illustratedatlas. Tropical Fish HobbyistPublications Inc. NeptuneCity, N.J.: l-459. 1996. Non-marine occurrenceof ichthvodecti- forms in the Lower Cretaceousof Brazil. Journal of Vertebrate ,16 : 3, Suppl., Abstractsof the 56ù Annual Meeting, New York. MARTILL, D. M. 1988.Preservation of fish in the Cretaceousof Brazil.Palaeontology, 31 : 1-l 8. 1993. Fossils of the Santanaand Crato Formations,Brazll. Palae ontolo gical Association Field Guides to Fossils,5:158 pp.; London. PATTERSON, C. 1973. Interrelationshipsof holosteans; pp. 233-305 In GREENWOOD.P. H.. MILES. R. S. & PATTERSON.C.