bs. GEOLOGY LIBRARY no.35 KIELDIANA Geology NEW SERIES, NO. 35 Cervifurca nasuta n. gen. et sp., an Interesting Member of the Iniopterygidae (Subterbranchialia, Chondrichthyes) from the Pennsylvanian of Indiana, U.S.A. Rainer Zangerl SEP 15 189/ March 31, 1997 Publication 1483 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY Information for Contributors to Fieldiana General: Fieldiana is primarily a journal for Field Museum staff members and research associates, although manuscripts from nonaffiliated authors may be considered as space permits. The Journal carries a page charge of $65.00 per printed page or fraction thereof. Payment of at least 50% of page charges qualifies a paper for expedited processing, which reduces the publication time. Contributions from staff, research associates, and invited authors will be considered for publication regardless of ability to pay page charges, however, the full charge is mandatory for nonaffiliated authors of unsolicited manuscripts. 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Author-generated changes in page proofs can only be made if the author agrees in advance to pay for them. ” © This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). LLINOIS LIBRAK URBANA-CHAMP/ Arr mma FIELLDLA: Geology NEW SERIES, NO. 35 Cervifurca nasuta n. gen. et sp., an Interesting Member of the Iniopterygidae (Subterbranchialia, Chondrichthyes) from the Pennsylvanian of Indiana, U.S.A. Rainer Zangerl Curator Emeritus Department of Geology Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 Present address: R.R. 4 Box 252A Rockville, Indiana 47872 Accepted May 21, 1996 Published March 31, 1997 Publication 1483 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1997 Field Museum of Natural History ISSN 0096-2651 PRINTED IN THE UNITED STATES OF AMERICA } j* Table of Contents PABSTRAGH naaasicscecsssenvisontevises cope onr une nceats eos 1 INTRODUCTION: $a5430sc sauce vssaapaps- svboccssecues sa cbey 1 SYSTEMATIC PALEONTOLOGY. s.5scc3ccess SES SCCM OOOOCBDODOOIOO V9ec09 C0056 00 20mm "ESS CSBSSSS SCD 00G0D0e0000000 Fic. 21. toxic to all forms of infauna, but an ideal envi- ronment for bacteria and other decomposers. Be- cause of the unusually rapid deposition of organic debris, the mud environment was very favorable for the preservation of fish skeletons, including the calcified cartilage of chondrichthyans. For ev- idence supporting the above conditions and a de- tailed analysis of the paleoecological parameters of the black shales, see Zangerl and Richardson (1963); Coveney and Glascock (1989) corrobo- rated many of the findings of Zangerl and Rich- ardson using entirely different evidence. Given the stated aspects of the black shale de- positional environment, one must conclude that the animals entombed in the shales were not reg- ZANGERL: CERVIFURCA NASUTA, NEW INIOPTERYGID FROM INDIANA SSW 220000D DOeDBG Fr“, C= Rr aN RAPA LO SOSEDB000000000000 Reconstruction of the skeleton of Cervifurca nasuta, in ventral view. ular inhabitants of those environments, but rather were visitors from the deeper, more central parts of the basins that entered these marginal areas. There many, perhaps most, of the invaders be- came entrapped in residual ponds and succumbed to predation and injuries inflicted by predators (Zangerl & Richardson, 1963). There is, however, some evidence that not all of the fishes preserved in the black shales were inhabitants of the epicontinental waters of the central North American basin complex. The pres- ence of teeth of such sharks as Cobelodus aculea- tus (Cope), Stethacanthus altonensis (St. John and Worthen), and Phoebodus sp. in the wash residues of central United States Pennsylvanian carbonates 21 Md Lisiseanir | oeoocaco Fic. 22. Reconstruction of the skeleton of Cervifurca nasuta, in lateral aspect. and drab shales suggests that these taxa inhabited the epicontinental marine basins and were widely distributed within them (Tway & Zidek, 1982, 1983a,b; Hansen, 1986). But remains of the most conspicuous and very abundant elements of the Mecca fauna, namely the iniopterygians, are miss- ing in the mentioned wash residues. The depositional history of the Pennsylvanian rock sequence of the structural Illinois basin, for example, includes numerous transgressions of ma- rine waters over a large, shallow, subsiding epi- continental pan. These transgressions inundated and destroyed the coal forests that grew on the low and swampy lands adjacent to the deeper parts of the pan. The muds that accumulated on the former forest floor beneath mats of floating vegetation, and which have become the black car- bonaceous shales, represent the initial conse- quences of some of the periodic transgressions. It is thus entirely possible that the iniopteryg- ians and some other cartilaginous fishes in the Mecca fauna (e.g., the caseodontoid and edestoid eugeneodontids) entered the epicontinental waters from deeper, oceanic realms during episodes of transgressive inundation, and were not denizens of the shallow basin provinces, but were members of the oceanic fish community of that time. Smith et al. (1994) compiled a most interesting set of 31 paleogeographic maps of the world from the Pliocene back to the Early Triassic showing the changing configuration of the land masses and the paleocoastlines. The Early Triassic map shows Pangaea with North America plus Greenland tightly joined to Africa and South America, which, in the aftermath of a major (Permian) gla- 22 ciation, means that the landmasses stood high rel- ative to mean sea level. In Pennsylvanian (e.g., Desmoinesian) time, by contrast, marine waters transgressed repeatedly over much of the lower North American continental area. Wanless (1969) suggested that two belts of transgression originat- ed from an area in southern California or north- western Mexico. One arm extended up to Mon- tana and from there to the Midwest; the other reached eastward south of the Ozark uplift to the Appalachian basins. However, the elevation of the Ouachita deformed belt blocked the eastern part of this seaway, so that Desmoinesian and later transgressions had to spread north on the west side of the Ozarks to reach the Illinois basin, for example. If my interpretation of the iniopterygians and some of the elasmobranchs of the Mecca fauna as denizens of oceanic waters is correct, these fishes would have had to travel some 3,000 km or more to reach the Illinois basin. More likely seems to be the existence of an oceanic geosyncline (per- haps alluded to by Wanless [1969, Fig. 1] under the name of Possible Gulf of Mexico Basin) from which by Desmoinesian time transgressions might have surged northward. Under this interpretation, the Mecca fish fauna would represent a mixture of basin elements and truly oceanic ones, a matter of considerable inter- est, because the Carboniferous record of oceanic chondrichthyans is rather meager indeed. Of the oceanic chondrichthyan fish community, the Mecca fauna appears to have sampled both open water endurance swimmers (e.g., Caseodus and relatives) with large forked tail fins at least FIELDIANA: GEOLOGY as stiff as those of Recent swordfish, as well as the iniopterygians, forms that perhaps inhabited the shelf areas of an oceanic seaway. Extensive collecting at many black shale out- crops in Parke, Vermillion, Fountain, Vigo, and Pike counties, and four quarries in Parke and Pike counties has shown that the various taxa of the Mecca fish fauna are not represented in equal pro- portions everywhere; rather, in each locality one or more taxa are common, whereas the rest are rare or absent. This phenomenon probably reflects the geographically uneven distribution of the fish- es immediately prior to and during the transgres- sions, when many of them entered the flotant en- vironments. Returning to the question of the great rarity of female iniopterygians, and also of females of the elasmobranch Denaea meccaensis (Williams, 1985) we can exclude, in the case of the iniop- terygians, the possibility of preferential predation affecting the unarmored females, because all the male specimens represent individuals that died of the effects of predator-inflicted injuries. In De- naea neither sex has any kind of armature. The uneven representation of the sexes perhaps merely indicates that the females preferred the deeper and presumably clearer waters of the more central parts of the basins, shunning whatever induce- ments the dark, shallow, and treacherous flotant environments offered to some 38 species of chon- drichthyans now preserved in the black shales that resulted from those rich organic mud accumula- tions. Conclusions The present account introduces a new genus of iniopterygian, Cervifurca nasuta, with a some- what dorsoventrally flattened body and eyes that faced dorsolaterad, very large pectoral fins, a pe- culiarly modified pelvic fin skeleton of uncertain function, and enormously enhanced pterygopodia. This new form enlarges our knowledge of the adaptive radiation of the Iniopterygia, which, along with some sharks, most likely were not part of the fish fauna of the epicontinental basin com- plex of North America but may have ventured, with repeated transgressions of oceanic waters, into the shallow basins with their marginal flotant environments. Acknowledgments I am most grateful to the Trustees of the Inter- state Commercial Corporation, who kindly per- mitted the establishment of the Bethel Quarry on corporate land, and I am especially indebted to John L. Barnett, Executive Director of the cor- poration, for his enduring interest and help with the project. The Indiana Department of Natural Resources, in particular John Wade, Manager of the Patoka State Fish and Game Area, Winslow, Indiana, and his staff, who now constitute the en- vironmental custodians of this land, contributed most important equipment and manpower re- sources to the quarry site preparation, and helped out on many occasions with weather-related prob- lems concerning the access road. My sincere thanks are due to all of them. I also wish to ex- press my appreciation to Ralph Langenheim of the Department of Geology and the University of Illinois Museum of Natural History at Urbana for the loan of a most interesting, three-dimensionally preserved, articulated, partial skeleton of the here- in described species from a western Illinois lo- cality, and for searching the correspondence files concerning the collection history of the specimen. I also wish to thank three anonymous reviewers for their helpful suggestions. Literature Cited ABEL, O. 1914. Orimente und Rudimente. Mitteilungen des Naturwissenschaftlichen Vereins an der Univer- sitat Wien (N.S.), 12: 79-82. DEBEER, C. R. 1937. The Development of the Vertebrate Skull. Clarendon Press, Oxford, XXIV+552 pp., 143 Pls. Coveney, R. M., JR., AND M. D. GLascock. 1989. A review of the origins of metal-rich Pennsylvanian black shales, central U.S.A., with an inferred role for basinal brines. Applied Geochemistry, 4: 347-367. GEGENBAUR, C. 1872. Untersuchungen zur Vergleichen- den Anatomie der Wirbeltiere. 3. Heft: Das Kopfske- lett der Selachier, ein Beitrag zur Kenntnis der Genese des Kopfskelettes der Wirbeltiere. Engelmann, Leip- zig. HANSEN, M. C. 1986. Microscopic chondrichthyan re- mains from Pennsylvanian marine rocks of Ohio and adjacent areas. Ph.D. diss., The Ohio State University, Columbus, XXI+513 pp., 91 Figs., 11 Pls. Hoi, L. G., L. W. WELDON, AND R. D. BLACKBURN. 1969. Aquatic weeds. Science, 166(13906): 699-709. KALIN, J. A. 1945. Die Homologie als Ausdruck ganz- heitlicher Baupline von Typen. Bulletin de la Société Fribourgeoise des Sciences Naturelles, 37: 135-161. Lunpb, R., AND R. ZANGERL. 1974. Squatinactis caudi- spinatus, a new elasmobranch from the Upper Missis- ZANGERL: CERVIFURCA NASUTA, NEW INIOPTERYGID FROM INDIANA 2 sippian of Montana. Annals of the Carnegie Museum, 45(4): 43-45. PATTERSON, C. 1965. The phylogeny of the Chimae- roids. Philosophical Transactions of the Royal Society of London, Series B, 249(757): 101-219. Situ, A. G., D. G. SMITH, AND B. M. FUNNELL. 1994. Atlas of Mesozoic and Cenozoic Coastlines. Cam- bridge University Press, Cambridge, England, IX+99 pp., 31 maps. STAHL, B. 1980. Non-autostylic Pennsylvanian iniop- terygian fishes. Paleontology, 23(2): 315-324. Tway, L. E., AND J. ZIDEK. 1982. Catalog of late Penn- sylvanian ichthyoliths, part I. Journal of Vertebrate Pa- leontology, 2(3): 328-361. . 1983a. Catalog of late Pennsylvanian ichthyo- liths, part II. Journal of Vertebrate Paleontology, 2(4): 414-438. . 1983b. Catalog of late Pennsylvanian ichthyo- liths, addendum. Journal of Vertebrate Paleontology, 3(2): 67-68. WANLESS, H. R. 1969. Marine Facies of the Upper Car- boniferous of North America. Sixiéme Congrés Inter- national de Stratigraphie et de Géologie du Carboni- 24 fére. Sheffield 11—16th September 1967, Compte Ren- du, 1: 293-336. WILLIAMS, M. E. 1985. The “‘cladodont level” sharks of the Pennsylvanian black shales of Central North America. Palaeontographica A, 190: 84-158. ZANGERL, R. 1979. New Chondrichthyes from the Ma- zon Creek fauna (Pennsylvanian) of Illinois, pp. 449- 500. In Nitecki, M. N., ed., Mazon Creek Fossils. Ac- ademic Press, New York. 1981. Chondrichthyes I; Paleozoic Elasmo- branchii, pp. 1-115. Jn Schultze, H.-P., ed., Handbook of Paleoichthyology, vol. 3A. Gustav Fischer, Stutt- gart. . 1995. The problem of vast numbers of clado- dont denticles in the Pennsylvanian Excello Shale of Pike County, Indiana. Journal of Paleontology 69(3): 556-563. ZANGERL, R., AND G. R. CASE. 1973. Iniopterygia, a new order of chondrichthyan fishes from the Pennsyl- vanian of North America. Fieldiana: Geology Mem- oirs, 6: IX+67 pp., 83 Figs. ZANGERL, R., AND E. S. RICHARDSON, JR. 1963. The pa- leoecological history of two Pennsylvanian black shales. Fieldiana: Geology Memoirs, 4: IX +239 pp., 56:Pls., 51 Figs. FIELDIANA: GEOLOGY A Selected Listing of Other Fieldiana: Geology Titles Available A Preliminary Survey of Fossil Leaves and Well-Preserved Reproductive Structures from the Sentinel Butte Formation (Paleocene) near Almont, North Dakota. By Peter R. Crane, Steven R. Manchester, and David L. Dilcher. Fieldiana: Geology, n.s., no. 20, 1990. 63 pages, 36 illus. Publication 1418, $13.00 A Catalogue of Type Specimens of Fossil Vertebrates in the Field Museum of Natural History. Classes Amphibia, Reptilia, Aves, and Ichnites. By John Clay Bruner. Fieldiana: Geology, n.s., no. 22, 1991. 51 pages, 1 illus. 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