Norntates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y
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AMERICAN MUSEUM Norntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2866, pp. 1-30, figs. 1-26, 1 table December 18, 1986 Coelacanths from the Lower Cretaceous of Brazil JOHN G. MAISEY1 ABSTRACT Specimens of Mawsonia sp. (possibly repre- dermal bones, the dermosphenotic morphology, senting the type species, M. gigas), plus a new and the number of paired posterior elements in genus and species of fossil coelacanth, are de- the skull roof suggest that the new coelacanth is scribed from the Romualdo Member of the San- related to Mawsonia. A phylogenetic hypothesis tana Formation, Chapada do Araripe, Ceara (Bra- is offered in which Mawsonia plus the new taxon zil). The new form, Axelrodichthys araripensis, is represent the sister group ofMacropoma and Lati- distinguished from other coelacanths on the basis meria. The new coelacanth is represented by sev- ofits cranial anatomy. The posterior moiety ofthe eral complete, articulated skeletons, among the first skull roofhas a median dermal element plus three to be discovered in South America. paired ossifications. Heavy rugose ornament ofthe INTRODUCTION Fossils of coelacanth fishes from South 40 km north-south across the Ceara-Pernam- America are extremely rare and until now buco border and into Piaui (Lima, 1978). The have consisted mostly of incomplete and principal collecting areas in Ceara are around fragmentary specimens. The discovery of Santana, Jardim, and Missao Velhao. Access complete examples in the Lower Cretaceous is generally difficult and hazardous. The Cha- Santana Formation of Ceara (northeastern pada do Araripe is capped by the Exu For- Brazil) is therefore ofconsiderable interest to mation, some 200 m of current-bedded red paleontologists. sandstones and siltstones above the Santana The Santana Formation is exposed along Formation. The latter has much greater lith- the flanks of the Chapada do Araripe, a pla- ological variation, with interbedded lime- teau extending some 180 km east-west and stones, evaporites, marls, and siltstones 1 Associate Curator, Department of Vertebrate Paleontology, American Museum of Natural History. Copyright © American Museum of Natural History 1986 ISSN 0003-0082 / Price $3.60 2 AMERICAN MUSEUM NOVITATES NO. 2866 (Mabesoone and Tinoco, 1973). Fossil fishes tion. One of these is referable to Mawsonia, occur in concretionary limestones at various but according to Campos and Wenz (1982, levels in the Santana Formation, but partic- p. 1152), "Les proportions relatives de ularly toward the base of the upper (Romu- l'avant et de l'arriere-crane, notamment la aldo) Member. Slight differences of lithology presence d'un museu etroit et allonge, indi- allow recognition of concretions from differ- quent que le Mawsonia de Ceara est speci- ent localities and horizons. Santana Forma- fiquement distinct de l'espece de Bahia et des tion coelacanths in the AMNH collection formes Africaines, toutes datees du Neocom- come mostly from Santana, some 50 km west ien a l'Albien." Unfortunately in the other of Juazeiro do Norte, but also from Jardim, coelacanth taxon from Ceara (here regarded slightly under 50 km to the southeast. Some as a new genus), the anterior skull roofis even specimens (e.g., AMNH 11759, from San- more elongate and narrow than in Santana tana) are associated with ostracods resem- Formation specimens referred to Mawsonia, bling Pattersoncypris; the low diversity ofos- and so it is impossible to determine which of tracods within the concretionary limestones the taxa Campos and Wenz (1982) were re- is said to indicate a fresh or brackish envi- ferring to Mawsonia. As discussed below, the ronment (Bate, 1972), although this may only two coelacanth taxa from the Santana For- be true of the lower part of the Romualdo mation share several peculiar features, in- Member (Mabesoone and Tinoco, 1973). It cluding postparietals (also found in other is uncertain whether the coelacanths from Mawsonia spp.), suggesting that they are Araripe were fully marine or else tolerant of closely related. They are certainly separate brackish to fresh water. species, but it is my view that the two are Despite the great abundance and diversity sufficiently distinct to merit generic separa- of the fossil fishes from the Santana For- tion. mation (Silva Santos and Valenca, 1968), the There is some disagreement concerning fauna has still not been adequately surveyed. coelacanth bone terminology. In the present This is surprising when one considers the ex- work I have mostly followed Forey's (1981) cellent state ofpreservation and the ease with terminology, except for features not dis- which these fossils may be prepared in acid cussed by him. (e.g., Toombs and Rixon, 1950; Rixon, 1976, The age of the Santana fishes was set as pl. 8). Although the Santana fishes have been Aptian by Silva Santos and Valenca (1968), known since the early 19th century (Spix and on the basis of comparison with other ich- Martius, 1828; Agassiz, 1841, 1844) several thyofaunas. This determination was rein- rare taxa have only recently been discovered. forced by the discovery of supposedly iden- Among the latest additions to the Santana tical species of fossil fishes in the Riachuelo paleofauna are coelacanths, which were first Formation of Sergipe-Alagoas, below am- reported on the basis of incomplete cranial monite zones representing the Albian and and postcranial remains by Campos and Wenz Upper Aptian (Silva Santos, 1981). While this (1982), who reported two taxa, Mawsonia sp. may place an upper limit to their age, the and "Forme B." Mawsonia was recognized Santana fish fossils could nevertheless pre- on the basis of two large skulls showing sup- date the Upper Aptian by a significant amount posedly characteristic "posterior parietals" (i.e., Neocomian or Upper Jurassic), accord- (termed postparietals here). "Forme B" was ing to available palynological and ostracod founded on a fragment ofthe body, preserved data (Maisey, in prep.). in part and counterpart. Without complete specimens it was not possible for Campos and Wenz (1982) to make direct comparisons ABBREVIATIONS between these supposedly different coel- acanths. INSTITUTIONAL The AMNH collection confirms the pres- AMNH, American Museum of Natural History, ence of two distinct coelacanth taxa in the New York Romualdo Member of the Santana Forma- DGM, Divisao de Geologia e Mineralogia, De- 1986 MAISEY: COELACANTHS 3 partamento Nacional da Producao Mineral, Rio SYSTEMATICS de Janeiro CLASS OSTEICHTHYES SUBCLASS SARCOPTERYGII ANATOMICAL ORDER ACTINISTIA afs, anterior fossa of parietal FAMILY COELACANTHIDAE AGASSIZ 1844 ang, angular Genus Mawsonia Woodward antart, antotic articulation (in Mawson and antpr, antotic process of basisphenoid Woodward, 1907) apa, anterior apophysis of parietal EMENDED DIAGNOSIS: Coelacanths of large apr, ascending process of prootic size (estimated body length up to 3 + m); skull art, articular roofing bones and angular ornamented by au, autopalatine heavy rugosities; bsp, basisphenoid operculum and gular or- cc, cranial cavity namented by numerous radiating striae; an- col, supraorbital sensory canal terior moiety of skull roof 1.5 to 2 times as coron, coronoid long as posterior part and from 2 to 2.5 times den, dentary as long as broad; posterior skull roof with di, internal depression of anterior apophysis three paired bones (parietals, supratempor- dpr, dorsal process of ectethmoid als, and postparietals); dermosphenotic with dsp, dermosphenotic splintlike anterior projection, and with in- faeb1, facet for articulation of first epibranchial fraorbital sensory canal located away from foc, fossa for otic canal the anterior margin; lachrymojugal elongate, fr, frontal the posterior g, gular two-thirds almost straight, an- hm, hyomandibular facet teriorly extending to the tectal series; anterior in, intemasal margin of metapterygoid slopes obliquely j, jugular canal forward as it rises from the pterygoid. la, lachrymal process of lateral rostral TYPE SPECIES: Mawsonia gigas Woodward lj, lachrymojugal (1907); Lower Cretaceous (Neocomian), Ba- mppa, median postparietal hia Basin, Brazil. mpt, metapterygoid my, myodome Mawsonia cf. gigas Woodward na, nasal np, notochordal pit REFERRED MATERIAL op, operculum AMNH 11758 Nearly complete skull roof and pa, parietal associated elements oforobran- padl, descending lamina of parietal chial skeleton. par, parasphenoid AMNH 12216 Partial skull roof with sensory pls, "pleurosphenoid" suture canals exposed. pop, preopercular AMNH 12217 Large basisphenoid with part of postcor, "posterior coronoid" parasphenoid attached. ppa, postparietal (paired) AMNH 12218 Large badly damaged basisphe- prcon, processus connectens noid and prootic. preart, prearticular ?DGM 11 07-P, 1109-P (mentioned in Campos pro, prootic and Wenz, 1982). prof, canal for superficial ophthalmic pt, pterygoid All the above specimens are from the Ro- qu, quadrate mualdo Member, Santana Formation, Cha- so, supraorbitals pada do Araripe, Ceara, Brazil. spl, splenial sq, squamosal SYSTEMATIC NOTE st, supratemporal tec, tectal AMNH 11758 closely resembles the ho- VII, facial nerve lotype of Mawsonia gigas, BM(NH)P 10355 la I in A 5cm i pa -antpr atb' ; dpr bsp hm, j ' pro B Fig. 1. Mawsonia cf. gigas, AMNH 11758. Acid-prepared skull in (A) ventral; (B) lateral; and (C) dorsal aspects. Prior erosion has resulted in loss of parasphenoid and parts of prootics. 1 986 MAISEY: COELACANTHS 5 - foc my6: 2 cm B Fig. 2. Mawsonia cf. gigas, AMNH