Redescription of Ellimma branneri and Diplomystus shengliensis : and relationships of some basal clupeomorphs PDF
Preview Redescription of Ellimma branneri and Diplomystus shengliensis : and relationships of some basal clupeomorphs
AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3404, 35 pp., 13 figures, 2 tables May 22, 2003 Redescription of ¢Ellimma branneri and +Diplomystus shengliensis, and Relationships of Some Basal Clupeomorphs MEE-MANN CHANG! AND JOHN G. MAISEY? ABSTRACT Two extinct clupeomorphs, +Ellimma branneri from the Cretaceous of Brazil and +Diplo- mystus shengliensis from the Eocene of China, are redescribed. tEllimma branneri was for- merly classified within the Clupeiformes, but it lacks derived characters of clupeiforms and clupeoids. Dorsal scute “‘wings’’ are expanded and subrectangular in +Ellimma and other members of the family +Paraclupeidae Chang and Chou (1977), approximately equal to +El- limmichthyidae Grande (1982a). Consequently, tEllimma branneri is classified here within the family +Paraclupeidae. +Paraclupeidae are known from the Lower Cretaceous to the middle Eocene. In the present work, two monophyletic groups are identified within the fParaclupeidae. One group (subfamily +Paraclupeinae of Chang and Grande, 1997), known only from the Lower Cretaceous (Hauterivian—Albian), includes }Paraclupea, +Ellimmichthys, and +Ellimma. These taxa are united by strongly sculptured, skull-roofing bones with ridges radiating from the growth center, and a dorsal scute ornament of prominent ridges. +Scutatuspinosus may also belong in this group. The other group includes }Diplomystus (Upper Cretaceous—Eocene) and +Armigatus (Upper Cretaceous), which are united by a single homoplaseous character (pres- ence of a posteriorly expanded third hypural, leaving no gap between hypurals 2 and 4): this character also occurs in pristigasteroids, +Erichalcis, osteoglossids, some elopomorphs (+Le- banichthys lewisi, and most Albula spp.), and a number of ostariophysans not included in our analysis. +Paraclupeines are customarily regarded as being more closely related to the Clu- ' Research Associate, Division of Paleontology, American Museum of Natural History; Research Professor, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, People’s Republic of China. e-mail: cmeemann@ yahoo.com ?Curator and Axelrod Research Chair, Division of Paleontology, American Museum of Natural History. e-mail: maisey @amnh.org Copyright © American Museum of Natural History 2003 ISSN 0003-0082 AMERICAN MUSEUM NOVITATES NO. 3404 peiformes than to other teleosts (1.e., as clupeomorphs), although no derived characters are uniquely shared by +Ellimma branneri and modern Clupeiformes. The relationships of +E /lim- ma and certain other extinct herring-like teleosts (including other +paraclupeines) with the Clupeiformes are unclear, and they may collectively form a paraphyletic assemblage. No biogeographical hypothesis satisfactorily explains the known distribution of nonmarine +paraclupeine fishes in the Cretaceous. A substantial portion of their nonmarine fossil record is missing (as evidenced by the recent discovery of a possible +paraclupeine, +Ezkutuberezi carmeni Poyato-Ariza et al., 2000, in Spain), and some aspects of their early distribution pattern may have involved marine dispersal. Eocene +Diplomystus occurs on both sides of the Pacific Ocean, but the ‘‘Pacifica’”’ hypothesis (which lacks empirical support) is abandoned as an explanation for such Eocene (and younger) trans-Pacific distribution patterns of nonmarine fishes. Instead, a “‘freshwater Arctic Ocean’’ hypothesis is favored. According to this hypoth- esis (for which there are several independent lines of geological evidence), temporary desa- lination of the Arctic Ocean occurred during the Paleocene and early Eocene, which may have permitted freshwater fishes to move unimpeded by salt-water barriers between Asia and North America; this temporary desalination event may eventually become recognized as a significant factor in the holarctic distribution patterns of various Tertiary-Recent freshwater fishes. INTRODUCTION pramaxillae and a complex pattern of sculp- ture on the surface of its dorsal scutes. He The primary purpose of our study is to re- restored the generic name fEllimma and re- examine }+Ellimma branneri, an Early Cre- ferred it to the family Clupeidae, subfamily taceous (Upper Aptian) clupeomorph from Clupeinae. Riacho Doce, in the Sergipe Basin of Ala- Our further preparation and observation of goas, Brazil. The original material was col- the original specimens of +E. branneri, from lected by J.C. Branner in 1907 and was de- both the American Museum and Carnegie posited in the Carnegie Museum, Pittsburgh. Museum collections, has provided new in- It formed the basis for Jordan’s (1910) de- formation supporting the view that }Ellimma scription under the name Ellipes, where it is a distinct taxon, but also suggesting that it was referred to the family Clupeidae. Unfor- does not belong in the Clupeidae. Instead, we tunately the name Ellipes is preoccupied by propose that it is a more primitive clupeo- a genus of orthopteran (Scudder, 1902), re- morph, closely related to three other Early quiring that another name be created for the Cretaceous taxa, ¢Ellimmichthys longicosta- fish taxon (fEllimma; Jordan, 1913). tus (Cope, 1886), TE. goodi (Eastman, 1912), +E. branneri was restudied by Schaeffer and +Paraclupea chetungensis Sun (1956). (1947) using additional material collected by The collective geographical distribution of Euphrasio Borges in 1917-1918. In 1931 Eu- these taxa is of considerable interest. +E llim- zebio Paulo de Oliveira donated this material michthys longicostatus is from the Marfim to the Department of Vertebrate Paleontology at the American Museum of Natural History Formation (late Hauterivian—early Barremi- from the Geological Survey of Brazil. an, Recdncavo Basin) of Brazil (Maisey, Schaeffer (1947) transferred the species to 2000), and }Ellimmichthys goodi is from Ap- the clupeid genus }Knightia, effectively tian—Albian strata of Equatorial Guinea, West making +Ellimma a synonym of that genus. Africa (Eastman, 1912), while +Paraclupea His proposal implied a significant strati- chetungensis is from the Lower Cretaceous graphic range extension for +Knightia, as the of southeastern China (Sun, 1956). A close genus had previously been recognized only relationship between +£llimmichthys and in the Eocene deposits of western North +Paraclupea was established by Chang and America (Jordan, 1907). Schaeffer’s proposal Grande (1997), who erected the subfamily was subsequently reversed by Grande +Paraclupeinae for these two genera, within (1982a, 1982b, 1985a), who excluded fE. the family +Paraclupeidae of Chang and branneri from the genus +Knightia, noting Chou (1977). The family was defined by a that +Ellimma can be distinguished from oth- single character (dorsal scutes broader than er clupeid taxa by the presence of two su- long), which also occurs in the genus +Di- 2003 CHANG AND MAISEY: TWO EXTINCT CLUPEOMORPHS 3 plomytus. Grande (1982a) had previously re- Cretaceous, Aptian—Albian?) from the Cabo lated }Ellimmichthys and +Diplomystus with- Basin of northeastern Brazil. This form was in his family +Ellimmichthyidae on the basis identified originally as fEllimmichthys lon- of a similar character (“‘lateral wings of the gicostatus (Costa et al., 1979), but it was dorsal scute elongated and blunted at the lat- subsequently referred to fEllimma and eral edges, giving the scute a subrectangular named fE. cruzi by Silva Santos (1990). The outline’), and Chang and Grande (1997) also same horizon has produced a small gonor- included +Diplomystus within the +Paraclu- hynchiform which was referred to }Dastilbe. peidae. The Cretaceous and Eocene paraclupeids The pattern of the dorsal scutes in }Ellim- each present a puzzling biogeographic pat- ma branneri only partially agrees with this tern for freshwater fishes. The Early Creta- family-level character, because the anterior ceous “‘southern transatlantic”’ pattern (1.e., dorsal scutes are longer than broad in this spe- restricted to western Gondwana, and repre- cies, whereas the posterior ones are broader sented by fEllimmichthys longicostatus and than long. Inclusion of this form within the tEllimma branneri from northeastern Brazil +Paraclupeidae on the basis of scute shape is and }Ellimmichthys goodi from West Africa) therefore not straightforward and necessitates is now extended into southeastern China a reevaluation of characters supporting para- (}Paraclupea chetungensis) and will perhaps clupeid monophyly. In searching for charac- be expanded farther by the new Spanish dis- ters other than scute shape, we addressed wid- covery (Poyato-Ariza et al., 2000). The Eo- er issues such as the relationships between cene forms have a “‘transpacific”’ distribu- these taxa and other basal clupeomorphs, in- tion, with the two sister species (fDiplomys- cluding +Santanaclupea, +Armigatus, +D iplo- tus shengliensis and +D. dentatus) on oppo- mystus, and +Knightia. site sides of the northern Pacific. Possible A redescription of the Chinese species of origins of this pattern are reexamined below +Diplomystus, that is, +D. shengliensis in the light of recent discoveries concerning Zhang et al. (1985), is also incorporated in the history of the Arctic region in the early this paper. According to its original descrip- Cenozoic. tion (in Chinese), }Diplomystus shengliensis is an Eocene teleost first collected from dril- ABBREVIATIONS ling cores of the Sheng-li Oil Field situated Anatomical on the southern coast of Bohai Gulf along the western coast of the Pacific Ocean. The AA angulo-articular species is very similar to {Diplomystus den- br.r branchiostegal rays tatus from the Eocene Green River Forma- BT? basibranchial toothplate? tion of North America, and hardly any dif- Ch ceratohyal ferences exist between them except for a few Cs caudal scute D dentary minor meristic characters (Zhang et al., Enpt entopterygoid 1985), a remarkable situation considering the Ep epural widely separated occurrences of these spe- Fr frontal cies on opposite sides of the Pacific. H, hypural 3 Besides the taxa considered here, there are Hm hyomandibular a few other records of Cretaceous “‘double- Io,, Io; infraorbital 2, infraorbital 3 armoured”’ clupeomorphs, such as +Scuta- Iop interopercle tuspinosus itapagipensis from the Neocomi- msc mandibular sensory canal an of Brazil (Silva Santos and Correa, 1985), Mx maxilla Op opercle an undescribed species of +Ellimmichthys P parasphenoid from the Tlayua Formation of southern Mex- Pa parietal ico (Chang and Grande, 1997: fig. 7e, f), and Pd predorsal bones +Ezkutuberezi carmeni from Spain, said to be Ph parhypural related to paraclupeids (Poyato-Ariza et al., Pmx premaxilla 2000). There are also records of a supposed Pop preopercle paraclupeid in the Cabo Formation (Lower Psp parasphenoid + AMERICAN MUSEUM NOVITATES NO. 3404 Pt posttemporal observe without some cleaning of the mate- Pu, preural centrum | rial. Fortunately the bituminous matrix has a Q quadrate very low carbonate content, and so for the S symplectic present study we were able to use dilute HCl Sc sclerotic bone to dissolve and clean broken bone from the Smxa anterior supramaxilla matrix in order to obtain clean impressions Smxp posterior supramaxilla of the skeleton. Latex peels were made from Soc supraoccipital Gy ural centrum 1 these cleaned impressions, and both reveal Ur urohyal much more detail then unprepared speci- Un, uroneural 1 mens. Institutional SYSTEMATICS AMNH American Museum of Natural History, SUBCOHORT CLUPEOMORPHA GREENWOOD New York ET AL., 1966 CM Carnegie Museum of Natural History, Pittsburgh, PA ORDER *ELLIMMICHTHYIFORMES GRANDE, FMNH _ Field Museum of Natural History, Chi- 1982 cago FAMILY +PARACLUPEIDAE CHANG AND CHOU, IVPP Institute of Vertebrate Paleontology LOFT and Paleoanthropology, Beijing SOF Shengli Oil Field, Dongying, Shandong = tELLIMMICHTHYIDAE GRANDE, 1982 Province, China SUBFAMILY *PARACLUPEINAE CHANG AND GRANDE, 1997 MATERIALS AND METHODS +ELLIMMA JORDAN, 1913 This study primarily involves specimens of }Ellimma branneri from the collections of +Diplomystus Cope, 1877: 811 (Gn part). the AMNH and CM, and of }+Diplomystus +Knightia Jordan, 1907: 136. +Ellipes Jordan, 1910: 24 (Brazilian species only). shengliensis from the Shengli Oil Field. +Ellimma Jordan, 1913: 79 (new name for Ellipes Specimens of }+Paraclupea chetungensis Jordan, 1910, preoccupied). from the IVPP, as well as those of +Armi- +Ellimma, Jordan and Gilbert, 1919: 26. gatus brevissimus, +Diplomystus dentatus, a +Knightia, Schaeffer, 1947: 17. few species of }Knightia, plus dried skele- +Ellimma, Grande, 1982a: 22. tons and cleared-and-stained specimens of +Ellimma, Grande, 1982b: 13. Clupea harengus and Onchorhynchus mykiss +Ellimma, Grande, 1985a: 250. from the AMNH, were compared, as were EMENDED DIAGNOIS: Paraclupeine with some specimens of *Diplomystus dentatus from the FMNH. comparatively low body depth. Surface of opercle with striations. Teeth on jaws and The specimens of fE. branneri examined are all preserved in bituminous shales of the palate very weakly developed. Anterior dor- Muribeca Formation and are in a laterally sal scutes slightly longer than broad and or- compressed and flattened state. In all these namented with ridges; posterior ones broader specimens the braincase is badly crushed, than long and ornamented with tubercles or and their study was further impaired by bro- tubercles plus ridges. Sharp spine extending ken surfaces and weathering of exposed sur- from median keel of posterior few dorsal faces, which frequently makes it difficult to scutes pointing posteriorly rather than pos- examine features such as sutures between terodorsally. Origin of pelvic fin opposite to bones and other structures. The poor state of middle point of base of dorsal fin. Procurrent preservation also helps explain why features rays 5 and 6 on each side of base of caudal such as dorsal scute ornamentation, the struc- fin. ture of the caudal skeleton, and various other TYPE SPECIES: fEllipes branneri Jordan, anatomical details have not been described 1910, the only species (based on page pri- previously, for they are almost impossible to ority). 2003 CHANG AND MAISEY: TWO EXTINCT CLUPEOMORPHS 3 tEllimma branneri (Jordan), 1910 namentation, dorsal scute morphology, and Figures 1-8 entopterygoid teeth, that are not discernible in the holotype of +E. branneri. There is in- +Ellimma branneri Jordan, 1910: 25, pl. 8, fig. 3. evitably some circularity in our inclusion of +Ellimma riacensis Jordan, 1910: 28, pl. 10. these features in our description of +E. bran- +Ellimma branneri Jordan, 1913: 79. neri, but we found no evidence to retain +E/- +Knightia branneri, Schaeffer, 1947: 17. limma riacensis as a distinct species, and this tEllimma branneri, Grande, 1982a: 22; 1982b: 13; 1985a: 250. taxon is regarded here as a subjective syno- nym of TE. branneri. Thus, we conclude HOLOTYPE: CM 5249/1 and CM 5249/2. there is only one species of +Ellimma rep- Note that only CM 5249/1 was designated resented in the Riaco Doce material, rather the type by Schaeffer (1947), and this was than two sympatric species. also the type specimen chosen by Jordan for +Ellimma branneri. In fact, CM 5249/2 is the DESCRIPTION counterpart to CM 5249/1, although this was not noticed in previous descriptions. The ho- GENERAL SHAPE: The overall outline of +E. lotype is a small fish, with a standard length branneri resembles that of +Paraclupea che- of 38.5 mm (measured from the counterpart tungensis (Chang and Chou, 1977; Chang CM 5249/2, which is more complete than and Grande, 1997), with a markedly convex CM 5249/1). ventral outline and a gently curved dorsal ADDITIONAL MATERIAL: AMNH_ 10046— outline rising to an apex at the origin of the 10062; CM 5248/10, 11, 21, 36, 39; 5249/2 dorsal fin (figs. 1A, B, 2A—C). The maxi- (counterpart of 5249/1), 3, 25, 27, 33, 49, 52, mum body depth occurs at the origin of the 54, 55, 56 (counterpart of 5248/11), 125, 130 dorsal fin. The standard length of individuals (counterpart of 5249/125), 175 (counterpart in our samples varies from 20.8 (CM 5249/ of 5249/52). 125) to 102.2 mm (CM 5248/4); in other DIAGNosIs: As for genus. words, our largest specimens are five times HORIZON AND LOCALITIES: From Riacho longer than the smallest. The proportion be- Doce, Sergipe Basin, Brazil (Muribeca For- tween the body depth and the standard length mation, Lower Cretaceous, Aptian—Albian, is therefore somewhat variable, usually being = “‘Alagoan”’ local stage). The age of this approximately 41—44% standard length, but unit is well established from palynomorphs rarely 36 or 50%. and foraminifers (Feij6, 1994). SKULL Roor: The bones of the braincase COMMENTS: The type specimen of +E/lim- in tE. branneri are usually so badly crushed ma riacensis (CM 5248/4) has a standard that it is impossible to recognize bone mar- length of 102.2 mm and is therefore a some- gins or even various recesses, fossae, and what larger individual than the type of TE. bullae within the bones; importantly, it has branneri. It is also much better preserved, not been possible to determine whether a re- and it consequently shows more morpholog- cessus lateralis was present (an important ical details than the type of +E. branneri. We clupeiform character absent in primitive clu- were however unable to identify any un- peomorphs; Grande, 1985a). Few features of equivocally unique features of }Ellimma ri- the braincase can be described. In the ante- acensis, and we consequently agree with rior part of the skull roof, the posterior por- Schaeffer (1947) that +Ellimma riacensis is tion of the dermethmoid (seen in CM 5249/ a synonym of TE. branneri. We regard +E. 49) bears a lateral process on both sides and branneri as the type species of the genus on a pair of processes extending posteriorly the basis of page priority. Our observations above the frontals. In larger specimens the led us to the conclusion that all the tE/limma posterior one-third of the frontals, as well as specimens studied by Schaeffer indeed rep- the parietals, are strongly ornamented with resent a single species, but we must also ad- ridges radiating from their growth centers. mit that the holotype of +E. riacensis (CM The dorsal part of the supraoccipital (seen in 5248/4) provides some morphological infor- CM 5249/56) is small and triangular and mation, including details of the skull roof or- shows indications of ridges radiating from 6 AMERICAN MUSEUM NOVITATES NO. 3404 Fig. 1. 4; (B) CM 5249/2, the holotype. Both scale bars = | cm. the posteromedian point of the bone antero- rietal. The supratemporal commissure passes laterally. In smaller individuals (e.g., CM through the parietals. 5248/39, standard length approximately 51 A comparatively broad and flat ridge runs mm) the skull roof bones are smooth and the length of the posttemporal along its mid- without sculpture (fig. 3). Unlike in }Para- line, and its broader posterior part is orna- clupea chetungensis there is no anterior fon- mented with ridges in larger individuals. The tanelle between the frontals. The parietals dorsal (epiotic) limb of the posttemporal is meet at the midline and are not separated by long and narrow anteriorly but becomes the supraoccipital (in clupeiforms the supra- broader posteriorly, while the ventral limb of occipital separates the parietals; Grande, this bone is thin. The lateral line sensory ca- 1985a). As far as can be determined the su- nal runs diagonally through the supracleith- praoccipital crest is small and low (CM rum. 5249/56, 5248/11, and 5248/39). The supra- ORBITAL REGION: Faint impressions of orbital sensory canal is enclosed in a bony three infraorbital bones can be seen in a few ridge and extends from the frontal to the pa- specimens, for example, AMNH 10048 (fig. 2003 CHANG AND MAISEY: TWO EXTINCT CLUPEOMORPHS 7 Fig. 2. tEllimma branneri, Muribeca Formation, Riaco Doce, Sergipe Basin, Brazil. Silicone peels prepared from AMNH specimens. (A) AMNH 10048; (B) AMNH 10057; (C) AMNH 10060. 8 AMERICAN MUSEUM NOVITATES NO. 3404 mi Fig. 3. +Ellimma branneri, CM 5248/39 to show skull roof. 2A) and 10050. Two detached infraorbitals coronoid process is present on the dentary. (perhaps the second and third) are exposed The mandibular sensory canal runs above the on CM 5249/52 and 5249/175 (part and lower margin of both the dentary and the an- counterpart; fig. 4). The first two infraorbitals gulo-articular (fig. 5B). The maxilla is long seem to be narrow and elongated, but the and extends behind the jaw joint. In most third is broader and shorter. The infraorbital specimens of +Ellimma branneri the jaw sensory canal runs along the orbital margin teeth are not visible, but a few fine, conical of all three bones. Parts of the sclerotic ring teeth are recognizable on the oral margin of were seen in several specimens, although no the premaxilla and dentary in the latex peel details were evident. of AMNH 10057. Very fine serrations are PARASPHENOID AND ENTOPTERYGOID: No also present on the oral margin in one de- teeth are present on the parasphenoid. A bas- tached maxilla (CM 5248/11 and CM5249/ ipterygoid process is recognizable on the la- 56, part and counterpart; fig. 6C, D). tex cast of AMNH 10051. In most specimens Collectively, these observations suggest of +Ellimma branneri no teeth were ob- that the entire jaw dentition of }Ellimma served on the entopterygoid, although faint branneri consists only of very small teeth, traces of tiny broken tips of teeth are some- and is much reduced in comparison with times present (e.g., AMNH 10047, 10048 both +Ellimmichthys longicostatus and + Par- and CM 5248/4). In contrast, +Ellimmichthys aclupea chetungensis, where a well-devel- and +Paraclupea have an entopterygoid den- oped dentition is present both on the jaws tition that is relatively better developed, and and the palatal surface of the entopterygoid the reduced dentition in +E/limma is a distin- guishing feature of the genus within paraclu- (Chang and Chou, 1977; Chang and Grande, peids. 1997). The presence of very small entopter- JAws: In +Ellimma branneri the articula- ygoid teeth in +Ellimma branneri may be re- tion of the lower jaw is positioned more or lated to a microphagous diet, as in many Re- less below the posterior margin of the orbit cent clupeomorphs. +Ellimma branneri pos- (figs. 2A—C, 5B). The dentary, angulo-artic- sesses two supramaxillae (a primitive tele- ular, and retroarticular are seen clearly in ostean state), of which the posterior one is AMNH 10048 (fig. 5A). A well-developed larger. Both supramaxillae are ornamented by 2003 CHANG AND MAISEY: TWO EXTINCT CLUPEOMORPHS Fig. 4. +Ellimma branneri, to show detached opercle, subopercle (at bottom) and two small infraor- bitals. (A) CM 5249/52 (part) and (B) CM 5249/175 (counterpart). fine shallow grooves on their external sur- which contains three or four branches of the face. preopercular sensory canal (fig. 5B). OPERCULAR SERIES AND HyYPOBRANCHIAL The hyomandibular shaft is narrow, with APPARATUS: A detached but well-preserved a single head and no anterodorsal process. opercle with an almost complete margin is The quadrate is thick, with the symplectic present in CM 5249/52 and 175. It is deep inserted in a notch above its posteroventral and rectangular in shape, with a protruding margin, as in teleosts generally (fig. 5A, B). anteroventral corner. The ventral part of the The anterior ceratohyal is rectangular in opercle in most specimens of +£llimma shape, longer than deep, and contains an branneri is ornamented on its lateral surface elongated oval bericiform foramen (AMNH by striations which radiate ventrally from the 10048, 10050; fig. SA). This opening is tra- area of attachment (fig. 4A, B), but the sur- versed by a groove for the hyoidean artery face of the opercle in the smallest specimen on the outer face of the anterior ceratohyal. is smooth, with no striations (CM 5249/125 Absence of the bericiform foramen is con- and 130, standard length 20.8 mm). The su- sidered to be a synapomorphic character of bopercle shows the anterior ascending pro- clupeiforms by Grande (1985a). There are cess which is usually covered by the opercle approximately 10 branchiostegal rays (CM 5249/52; fig. 4A). The preopercle has (AMNH 10048; fig. 5A). An outline of the two arms forming an obtuse angle; its verti- urohyal was detected on AMNH 10048, but cal arm is longer than the horizontal one, few details are discernible because the bone 10 AMERICAN MUSEUM NOVITATES NO. 3404 Fig. 5. +£llimma branneri, silicone peels showing detail of the head in (A) AMNH 10048; (B) AMNH 10057. Both scale bars = | cm.
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