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AMNH Digital Library AMERICAN MUSEUM Norntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2728, pp. 1-34, figs. 1-38 June 4, 1982 A Revision of the Fossil Genus tDiplomystus, With Comments on the Interrelationships of Clupeomorph Fishes LANCE GRANDE' ABSTRACT Several primitive fossil clupeomorphs are de- new genus, is proposed for tClupea brevissimus scribed, and a hypothesis of clupeomorph inter- Blainville, 1818. The relationship of tArmigatus relationships is proposed. The genus tDiplomys- brevissimus, new genus, to other clupeomorphs tus is revised and the type species for the genus, is not clear, and it forms an unresolved trichoto- tD. dentatus Cope, 1877, is redescribed. tDiplo- my with tellimmichthyids and clupeiforms. mystus dentatus is closely related to tD. birdi Many other species erroneously assigned to the Woodward, 1895, and tD. dubertreti Signeux, genus tDiplomystus are removed to make the ge- 1951, and all three species form the sister group nus monophyletic and thus useful in systematic to tEllimmichthys longicostatus (Cope, 1886). and comparative anatomical studies. Because the dorsal scutes of tE. longicostatus The comparative morphology of clupeomorph lack the pectinate posterior border diagnostic of dorsal scutes is discussed. It is found that "double tDiplomystus, Jordan's (1910 and 1919 [in Jordan armor" in clupeiforms is not restricted to a small and Gilbert, 1919]) removal of this species from specialized group, but rather is a widespread tDiplomystus is considered valid. Both tDiplo- character, occurring in clupeids, engraulids, tel- mystus and tEllimmichthys are placed in tEllim- limmichthyids, and tArmigatus. Detailed mor- michthyidae, new family (tDiplomystidae is phological study of the dorsal scute in clupeo- preoccupied by a family of South American cat- morphs shows a complex of several characters, fishes). The tEllimmichthyidae, new family, is the some of which can be used to define monophyletic sister group to the Clupeiformes. tArmigatus, groups within the Clupeomorpha. INTRODUCTION The fossil herring-like fish tDiplomystus is anatomical comparisons with other teleosts one of the most widely cited fossil teleosts by at least 40 different authors and in more in the literature. It has been cited or used for than 75 different papers. Yet, as currently 1 Department of Ichthyology, American Museum of Natural History. Copyright © American Museum of Natural History 1982 ISSN 0003-0082 / Price $3.15 2 AMERICAN MUSEUM NOVITATES NO. 2728 defined this is a non-monophyletic group and The morphology of most of the species therefore of questionable value in systematic (including the type, tD. dentatus) is poorly studies. known. "tDiplomystus" brevissimus (Blain- The genus has largely been a repository ville, 1818) is the only species described in for most any articulated clupeomorph fossil much detail, mainly by Patterson (1967); but with dorsal scutes, since its description by no evidence has ever been presented that Cope in 1877. Even Cope included a modern would indicate that "i'D." brevissimus and type of clupeoid (today called tKnightia) in tD. dentatus are closely related. Patterson the same genus with tDiplomystus. And (1967) showed that although "i'D." brevis- since Cope about 15 additional taxa have simus is a clupeomorph, it is not a clupeoid been erroneously placed into the genus. This as had been previously thought. has caused serious problems in papers using Several authors (Woodward, 1892, p. 413; tDiplomystus for anatomical and/or out- Schaeffer, 1947, p. 24; Greenwood, 1968, p. group comparisons. 265 and others) have proposed that "double- The genus tDiplomystus was described by armored" clupeiforms (those clupeiforms Cope (1877) as a clupeid genus including sev- with both dorsal and ventral scutes) consti- eral fossil species from the Green River For- tute a monophyletic group. Nelson (1970b) mation. He divided the genus into two "sec- challenged this concept by pointing out that tions." In section 1 he included tD. dentatus Clupanodon thrissa (a gizzard shad) and Cope, 1877, tD. analis Cope, 1877, and tD. Ethmidium (Chilean shad), which are both pectorosus Cope, 1877; and he included tD. double-armored, are not closely related to "humilis" (Leidy, 1856)2 and fD. altus (Lei- other double-armored clupeids. Nelson (per- dy, 1873) in section 2. He placed a sixth sonal commun.) also drew my attention to species from the Green River Formation, the fact that many engraulids are also dou- tD. theta (Cope, 1874), as "intermediate be- ble-armored (although they are peculiar in tween ... the two sections" (Cope, 1877, p. having only one or two dorsal scutes). A 811). Section 2 was removed from the genus pristigasterin (Pristigaster cayana) was also tDiplomystus by Jordan (1907) and placed in found to have dorsal scutes (see below). a new genus tKnightia Jordan, 1907. The The main objectives of the present work genus tKnightia is revised elsewhere are: to provide a detailed description of (Grande, in press). All of Cope's tDiplomys- tDiplomystus dentatus (the type species of tus species from the Green River Formation the genus); to find derived characters uniting except for tD. theta were found to be junior the type with other species, thus defining the synonyms of tD. dentatus Cope, 1877, by genus more rigorously; and to examine the Grande (1980). It is not possible to comment relationship of tDiplomystus to i'Knightia on the position of tD. theta because there is and other groups within the Clupeomorpha. no known holotype for the species, and its A comparative morphological study of the description is insufficient. dorsal scutes in clupeomorphs is also includ- More than 15 species have been assigned ed as an appendix. to the genus tDiplomystus in addition to Cope's species, but reasons for most of these ACKNOWLEDGMENTS assignments are erroneous or tenuous. All species known to the author were reviewed I thank Mr. Guido Dingerkus, Drs. Donn during this project. E. Rosen, Gareth Nelson, Colin Patterson, and C. Lavett Smith for reading and com- menting on various drafts of this manuscript. 2 Because tClupea humilis Leidy, 1856 was a hom- Dr. Gareth Nelson also assisted in translat- onym of tC. humilis von Meyer, 1848, Leidy's name is unavailable. Cope's (1870) tClupea pusilla is unavail- ing some type descriptions from French to able for a replacement because that name is preoccupied English, and by sharing his discovery that by C. pusilla Mitchill, 1814. Therefore, the valid name many anchovies have dorsal scutes. Mr. Gui- for this taxon is tKnightia eocaena Jordan, 1907 (and do Dingerkus aided with some of the pho- not tK. humilis Leidy as given in Grande, 1980). tographic work. 1982 GRANDE: tDIPLOMYSTUS 3 I also thank Dr. Daniel Goujet (Museum All other counts and measurements follow national d'Histoire naturelle) and Dr. Colin Grande (1980) and Hubbs and Lagler (1949). Patterson (British Museum, Natural History) Fossils were prepared using needles under for their help during my visit abroad. a dissecting microscope. Dorsal scutes and Part of the fieldwork and travel expenses skeletons of recent teleosts were cleared and were paid for by grants to the author, from stained following Taylor (1967). All original the Theodore Roosevelt Memorial Fund, drawings were made using a Leitz camera American Museum of Natural History. A lucida on a Bausch and Lomb stereomicro- grant from City College of New York helped scope. All illustrated specimens of tDiplo- pay for drawing equipment and photographic mystus dentatus are from locality F- 1 of supplies. Grande (1980) except for the F-2 specimen in figure lB. The names of all fossil taxa mentioned in the text are preceded by a MATERIALS AND METHODS dagger (t). Drawings of dorsal scutes are of the dorsal Fossils used here are deposited in the fol- surface unless otherwise stated. Explanation lowing institutions: Department of Verte- of morphological terminology used here for brate Paleontology, American Museum of dorsal scutes is given in figure 38. In all Natural History, New York (AMNH); De- drawings, the anterior margin faces left partment of Paleontology, Science Museum (drawings of some fossils have been re- of Minnesota, St. Paul (SMMP); British Mu- versed). seum (Natural History), London (BMNH); The taxonomic subdivisions (subfamilies) Museum national d'Histoire naturelle, Paris of clupeids and engraulids used in the Ap- (MNHN); University of Michigan, Museum pendix follow Whitehead (1968). of Paleontology (UMMP); and Department of Paleontology, California Academy of Sci, ences (CAS). Recent material examined is in ABBREVIATIONS USED IN FIGURES the Department of Ichthyology, of the Amer- Aa, angulo-articular ican Museum of Natural History, and the Bt p, basipterygoid process of parasphenoid Australian Museum, Sydney, New South C, circulus (or ridge) Wales (AM). Some uncatalogued specimens Ch, anterior ceratohyal (=ceratohyal of Mc- from the Smithsonian Oceanographic Sorting Allister, 1968) Center, Washington, D.C. (SOSC) were also De, dentary examined and accession numbers for these Dsp, dermosphenotic are given in the text. Eh, posterior ceratohyal (=epihyal of McAllister, like most 1968) tDiplomystus, clupeomorphs, Enpt t, endopterygoid teeth has two vertebrae in advance of those bear- Ep, epural ing full length ribs. These are often not vis- Fr, frontal ible in fossils because they are normally cov- HS, haemal spine ered by the superficial bones of the skull. Hy, hypural Therefore the number of anterior vertebrae lo, infraorbital in all fossils was determined by counting the lop, interopercle pairs of ribs and then adding two. Only Mx, maxilla preural centra are included in the vertebral N, nucleus (or focus) counts. Standard length was measured from NS, neural spine the anterior tip of the snout to the Op, opercle posterior Pa, parietal end of the third hypural plate. Principal ray PH, parhypural counts of the dorsal and anal fins (="major Pop, preopercle fin rays" of Grande, 1980) include those rays Pmx, premaxilla which extend all the way to the outer fin Ps, parasphenoid margin.
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