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Home , Allothrissops, Chondrosteidae, Ichthyodectiformes, Ophiopsis, Pholidophorus, Ptycholepis

66 AMERICAN MUSEUM NOVITATES NO. 2796

versus the traditional or evolutionary meth- example, the Sundance endemic Occithris- od (e.g., Mayr, 1982). The first leads to un- sops willsoni is the earliest ichthyodectiform certainty, expressed in liberal use of incertae known, and could be treated as ancestral to sedis. The second leads to spurious certainty, other members ofthe group, or at least to the when phenetic or "adaptive" groupings are ichthyodectoids. Ancestry is thus one way of treated as real. interpreting the meaning of a monotypic tax- As for "detailed biogeographical work" on in a trichotomous cladogram like figure (Campbell's 1975 phrase), it begins for the 30C. One could then develop a narrative of cladist with the question "what is the sister- ichthyodectiform dispersal, with the Sun- group, and where does it live?" The question dance Sea as the center of origin. But just as is asked for many taxa in a diversified fauna. evolutionary systematics may lead to spu- In our Sundance/Wanakah assemblage, the rious certainty, so evolutionary biogeography question cannot be answered for any taxon, may lead to spurious stories. Our inability to and analytical biogeography cannot be start- resolve the relationships of Occithrissops is ed. For the traditionalist, biogeography can surely no basis for an inferred center oforigin. be sketched out with a broader brush. For

APPENDIX: THE DISTRIBUTION OF A resume of Jurassic distribution has been Forcing all Jurassic fish localities into either a ma- included in this Appendix in the form of two ta- rine or a nonmarine setting involved some judg- bles, one for marine (table 3) and the other for ments that may be incorrect, but in our view that nonmarine (table 4), and by a series of paleogeo- possibility is preferable to lumping all occurrences, graphic maps (figs. 37-39) showing most of the or to introducing a third set of tables for indeter- areas in which Jurassic fishes have been found. In minable or possible brackish-water occurrences. our present state of knowledge, this compilation The basis for the classification is necessarily a must be regarded as tentative. The reasons for this combination of the "classical" and phylogenetic have been set forth in the preceding discussion, approaches. For the Chondrichthyes, the recent which emphasizes some ofthe problems that arise studies of Thies (1983), Maisey (personal com- in analyzing and comparing any fossil fish assem- mun.), and Patterson (1965) have been utilized. blages of about the same age from different areas The actinopterygian classification is mostly based or parts of the world. Our justification for includ- on Gardiner (1967) for the chondrosteans, on Pat- ing these tables is simply that no effort of this sort terson (1973) for the non- neopterygians, has been attempted before, and that the tables will and on Patterson and Rosen (1977) for the . provide a summary of current information about Taxa of unknown or uncertain status are framed both the systematics and the distribution of Ju- by quotation marks or are placed in an incertae rassic fishes. sedis category. The preparation of worldwide taxonomic, The abbreviations in the marine and non-ma- stratigraphic, and geographic records ofextinct an- rine tables represent the stages of the Jurassic, as imals, such as are represented in these tables, nec- follows: essarily involves data compression and various Portlandian Po arbitrary decisions. If such compilations are un- Upper K critical, they cease to be informative. Doubtful or Ox questionable identifications, which are often per- petuated in tables such as these, can mislead the Callovian Ca specialist as well as the non-specialist. On the other Middle Bathonian Bt hand, to omit a record because of doubt about Bajocian (incl. Aelenian) Bj provenance or identification can also lead one astray. We have tried to steer a middle course, T using specialist knowledge judiciously, checking Pliensbachian P1 specimens where possible, and omitting records Lower S only when we believe that they lack any real factual Hettangian H basis. The distinction between the marine and non- In some areas, and particularly for nonmarine marine tables has been based on the literature, on deposits, the Jurassic has been divided only into paleogeography, and on our own opinion and/or Lower (L), Middle (M) and Upper (U), and in a field experience of the associated fauna and flora. few places only a Jurassic age (J) has been recog- 1984 SCHAEFFER AND PATTERSON: JURASSIC FISHES 67

FIG. 37. A. Paleogeographic map showing approximate distribution of land and sea during the Hettangian and Sinemurian. B. Same for the Pliensbachian and Toarcian. Modified after Hallam (1975). Geographic areas in tables 3 and 4 are shown in figure 39.

nized. In the Lower and Middle Jurassic, stages Bajocian. There are problems toward the end of are non-controversial, except that we have not dis- the Jurassic, where, as Rawson et al. (1978, p. 7) tinguished the Aalenian, including it as (Lower) put it, "Ammonite provincialism had reached such 68 AMERICAN MUSEUM NOVITATES NO. 2796

FIG. 38. A. Paleogeographic map showing approximate distribution ofland and sea from the Bajocian to the early Callovian. B. Same for the late Callovian to Kimmeridgian. Modified after Hallam (1975). Geographic areas in tables 3 and 4 are shown in figure 39.

a peak ... that it is extremely difficult to correlate the Jurassic/ boundary. Hence a three- the Tethyan and boreal ammonite sequences near fold stage terminology has evolved for the top of 1984 SCHAEFFER AND PATTERSON: JURASSIC FISHES 69

FIG. 39. Paleographic map showing approximate distribution ofland and sea during the Portlandian. Modified after Hallam (1975). Numbers refer to geographic areas in tables 3 and 4. the Jurassic: Tithonian for the 'standard' Tethyan Solnhofen, Cerin) appear as Kimmeridgian, and sequence and Volgian or Portlandian for the boreal the Lulworth Beds (formerly Purbeckian) appear regions." as Portlandian. There is also a problem with the Purbeckian, The earliest Cretaceous fish assemblages from traditionally regarded as the terminal stage of the Great Britain have also been included in the tables, Jurassic in Britain. We follow Casey (1973) and to emphasize their close identity with the Late Rawson et al. (1978) in taking the mid-Purbeck Jurassic ones, as follows: Cinder Bed as the base of the Cretaceous. The lower (Jurassic) half of the old Purbeckian is now Wealden and Purbeckian above the named the Lulworth Beds, and the upper (Creta- Cinder Bed (freshwater) We ceous) half the Durlston Beds. In attempting a Marine equivalents of Wealden Ne world survey of Jurassic fishes, our selection of The geographic subdivisions in tables 3 and 4 Upper Jurassic stage names is influenced by the are based on regional or paleogeographical con- fact that we have used the British Museum (Nat- siderations. The numbered paragraphs in the dis- ural History) collections, and their catalogues and cussion below refer to these subdivisions, as do records, as an up-to-date and verified list of Ju- the numbers in figure 39. The paleogeographic rassic taxa that will serve as a standard for the maps (figs. 37-39) should be examined in con- identification and composition ofassemblages from the remarks on occurrence. other areas. We have therefore followed the Upper nection with Jurassic correlation chart recently published by the Marine Occurrences Geological Society of London (Cope et al., 1980), (Table and used Kimmeridgian and Portlandian as the 3) last two Jurassic stages. This means that the well- 1. Great Britain is treated separately because, as known European Tithonian assemblages (e.g., noted above, the extensive collections of Jurassic uedrf 0 ; ; P.,..... IHssnl ol | Cl.. vpeurD VSf 6 ...

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76 1984 SCHAEFFER AND PATTERSON: JURASSIC FISHES 77 fishes in the British Museum (Natural History), rated North and South America by the Early Ba- along with their systematic catalogues and strati- jocian (figs. 37-39). This seaway was responsible graphic records, have provided an updated, veri- for the marine, fish-bearing deposits in Cuba and fied, or verifiable, list of Jurassic taxa that can Mexico. serve as a standard for the identification ofJurassic 9. The western margin of the North American assemblages from other areas. Assignment ofgen- plate was covered by a transgressive sea during the era to stage is frequently based on our assessment (fig. 37A, B). By the mid-Jurassic of these collections. that sea extended into the western interior as a 2. Northern Europe includes France and Ger- vast embayment (figs. 1 and 38). The Fernie Group many with their diversified faunas, as well as Aus- of western Canada (Frebold, 1957), the Pliens- tria, Poland, Denmark, and Spitzbergen, which bachian Nicely Formation ofOregon (Imlay, 1980), have relatively few reported taxa. the Sundance and possibly all or part ofthe Wana- 3. Spain and Portugal are grouped separately kah Formation were deposited in, or marginal to, mainly to emphasize their Tethyan connections. this transgressive sea. As discussed earlier, all of All of these areas in western Europe were covered these rock units contain fishes. The Greenland during most of the Jurassic by the transgressive portion of the North American plate was covered eastern extension of the Tethys Sea (figs. 37-39). by the sea along a restricted part of its eastern It should be noted here that the famous Montsech, border during much of the Jurassic (figs. 38A, B, Lerida locality in Spain, long regarded as Kim- 39). One fish specimen has been described from meridgian-Portlandian, has proved to be Lower Tithonian deposits on this island (Aldinger, 1932). Cretaceous or Valanginian (Brenner, Geldmacher, 10. A single marine fish occurrence and Schroeder, 1974). has been reported from the island of Honshu, Ja- 4. Italy and Sicily were part of, or close to, the pan (Yabe, 1902), probably on the southeast side African plate in the Mesozoic and were also cov- where most of the marine facies are located. Rus- ered by the Tethys Sea. The entries for the Sine- sian marine (or presumably marine) fish localities murian in column 4 are based on preliminary de- have been reported in Upper Jurassic, probably terminations of a collection from Osteno, Lake Portlandian (Middle Volgian) deposits that are Lugano, Lombardy (Pinna, 1967; Arduini, Pinna, mostly west ofthe Ural Mountains (figs. 38B, 39). and Teruzzi, 1981) now under study by B. G. - diner and C. Patterson. 5. North Africa is represented in table 3 to in- Non-marine Occurrences clude Morocco, Algeria, and Tunisia. These areas (Table 4) were also covered by the transgressive Tethys. 6. The so-called Trans-Erythrean Trough (Hal- 11. The entire Jurassic marine and continental lam, 1975, p. 167 and fig. 10.2) was an extensive rock sequence in Zaire has been dated only to the gulf-like transgression from the Tethys Sea that Upper Jurassic, mostly on the basis ofpresumably covered parts of Arabia, Baluchistan (India), East marine fishes (Saint-Seine and Casier, 1962). This Africa, and Madagascar prior to their separation. nearly unique nonmarine fish assemblage pro- As noted in table 3, some marine Jurassic fish vides little aid in this regard. remains have been found in each of these areas. 12. The lacustrine deposits in the USSR re- The occurrence of Jurassic marine sediments and ported to contain fishes have been approximately fishes in the Songa Limestone ofZaifre close to the located on the basis of comments in Obruchev equator and about 25°E longitude, can be most (1967): Karatau, Kazak, lat. 40-45°N, long. 70°E; readily explained in terms of a limited, transitory, Semipalatinsk, Kazak, lat. 50°N, long. 80°E; Bo- marine transgression that extended westward from goslowsk, Sverdlovsk, lat. 60°N, long. 60°E; Ir- the Trans-Erythrean Trough during the Late Ju- kutsk, Irkutsk, lat. 5 1°N, long 105°E. The fish as- rassic (Cahen, 1954; Saint-Seine and Casier, 1962). semblage for each of these localities has also been 7. The western coastal areas ofthe South Amer- compiled from Obruchev (1967) which includes ican plate were covered by transgressive seas dur- most of the primary references. ing most of the Jurassic (Weeks, 1947, figs. 7, 8). 13. In China, Jurassic fishes have been found in As listed in table 3, fishes have been found in continental, presumably lacustrine deposits in a Argentina and Chile. number of provinces. They are listed here on the 8. The separation of the North American and basis of updated information kindly supplied by African plates began in the Early Jurassic. The Dr. Hsient'ing Liu (personal commun.), plus re- North Atlantic ocean extended southward be- cent opinions regarding their affinities. tween these continents and, in effect, continued 14. In India, the Kota Formation contains a westward as an epicontinental seaway that sepa- Lower Jurassic assemblage recently reviewed by 78 AMERICAN MUSEUM NOVITATES NO. 2796

TABLE 4 World Distribution of Jurassic Fishes-Non-marinea

E X 5 XeP,X,O 3 CO) . 'O.O o v..00 1- (1, en t . n e o \0 00 Chondrichthyes Elasmobranchii Hybodontidae * ~M L Hybodus M We Acrodus Lissodus * M (=Lonchidion) Bt, We

Asteracanthus U Bdellodus L Hylaeobatis We Holuridae Palaeoniscinotus M Coccolepidae Pteroniscus U Coccolepis U M We Plesiococcolepis L Ptycholepidae Ptycholepis H-S Chungkingichthys M Yuchoulepis M Redfieldiidae Redfieldius S Chondrosteidae Peipiaosteus U Stichopterus M Halecostomi Semionotidae Semionotus H-S Lepidotes U L L-U? * * Po, We Dapediidae Paradapedium * L Tetragonolepis * L Gyrodontidae ?Tibetodus J Coelodus Po, We Uarbryichthyidae Uarbryichthys M We Ophiopsidae Neorhombolepis We Sinamia U Ikechaoamia U 1984 SCHAEFFER AND PATTERSON: JURASSIC FISHES 79

TABLE 4-(Continued)

Cu

S 0 wS ]= XS M~~~~10 o 0

N e t e £ ~~~~~~~~~~~~~~~4b oo

C - -4 I- c

Teleostei incer. sed. Catervariolus U Galkinia U Ligulella U Majokia U Pholidophoridae * M-U Baleiichthys * M M Hungkiichthys Hengnania L J~ ~ Aetheolepis L~~ Aphnelepis M * ~M Madariscus * * M Archaeomaene * * M~ Oreochima Ichthyokentemidae Ichthyokentema ?M Pleuropholidae Pleuropholis U Parapleuropholis U Austropleuropholis U - incer. sed. Pachythrissops .We* "" talbragarensis Aethalionopsis ** We "Leptolepis" sp. * * We Fuchunkiangia U Mesoclupea U Huashia U Hiodontidae Lycoptera U Tongxinichthys U? Sinolycoptera U? Elopocephala incer. sed. "Anaethalion" U Clupeomorpha incer. sed. Paraclupea U Sarcopterygii Actinistia Coelacanthidae Lualabaea U Indocoelacanthus L Diplurus S Dipnoi Ceratodontidae Ceratodus L, U P1, Po Bt? a See footnote on p. 76. 80 AMERICAN MUSEUM NOVITATES NO. 2796

Jain (1980). Dr. Philippe Janvier (personal com- lems in evolution. London, mun.) has provided information on the Jurassic Academic Press, pp. 271-288. fishes of Thailand. Arduini, P., G. Pinna, and G. Teruzzi 15. The age of the well-known non-marine fish 1981. Megaderaion sinemuriense n.g. n.sp., a locality at Talbragar, New South Wales, Australia new fossil enteropneust of the Sine- remains uncertain. It is regarded here as Middle murian ofOsteno in Lombardy. Atti Soc. Jurassic (White, 1981; Long, 1982). Ital. Sci. Nat., vol. 122, pp. 104-108. 16. The single actinopterygian known from Ant- Arratia, G. arctica is probably ofMiddle Jurassic age (Schaef- 1981. Varasichthys ariasi n. gen. et sp. from fer, 1972) and is apparently related to the pholi- the Upper Jurassic of Chile (Pisces, Te- dophorid Wadeichthys (Waldman, 1971) from the leostei, Varasichthyidae n. fam.). Palae- continental Lower Cretaceous ofVictoria, Austra- ontographica, vol. 175A, pp. 107-139. lia. 1982. Chongichthys dentatus, new genus and 17. In regard to Jurassic non-marine fishes from , from the Late Jurassic of Chile North America, the Pliensbachian Kayenta For- (Pisces: Teleostei: Chongichthyidae, mation (Imlay, 1980) of Utah and Colorado has new family). Jour. Vert. Paleont., vol. yielded only Ceratodus teeth. The Tithonian Mor- 2, no. 2, pp. 133-149. rison Formation also contains Ceratodus teeth plus Arratia, G., A. C. Garrido, and G. C. Diaz some scrappy remains that suggest Ophiopsis 1975a. Pholidophorus domeykanus n. sp. del (Prothero, 1981). Fishes from the Hettangian and Jurasico de Chile. Rev. Geol. Chile, no. Sinemurian zones of the Newark Supergroup are 2, pp. 1-9. discussed by Olsen, McCune, and Thomson (1982). 1975b. Sobre un pez fosil del Jurasico de Chile As interpretation ofthe depositional environment y sus posibles relaciones con clupeidos of the Todilto and Pony Express limestones re- sudamericanos vivientes. Ibid., pp. 10- mains equivocal (see section on Geologic Occur- 21. rence), the fishes from these units are included in 1975c. Leptolepis opercularis, n. sp., from the the marine list along with the taxa from the Sun- Jurassic ofChile. Ameghiniana, vol. 12, dance Formation. no. 4, pp. 350-358. 18. The British Purbeck and Wealden fishes Baltz, E. H. (Woodward, 1916, 1919) are included here either 1972. Geologic maps and cross sections ofthe under Portlandian (Purbeck below the Cinder bed) Gallinas Creek area, Sangre de Cristo or Wealden (Purbeck above the Cinder Bed, Weal- Mountains, San Miguel County, New den). Mexico. U.S. Geol. Surv., Misc. Invest., Map 1-673. Bartram, A. W. H. LITERATURE CITED 1975. The holostean fish genus Ophiopsis Agassiz. Zool. Jour. Linnean Soc., vol. Aldinger, H. 56, pp. 183-204. 1932. Uber einen Eugnathiden aus der unter- 1977. The Macrosemiidae, a Mesozoic family en Wolgastufe von Ostgronland. Med- ofholostean fishes. Bull. Brit. Mus. Nat. del. Gr0nland, vol. 86, no. 4, pp. 1-51. Hist. (Geol.), vol. 29, pp. 137-234. 1937. Permische Ganoidfische aus Ostgr6n- Baum, K., and R. Lund land. Meddel. Gr0nland, vol. 102, no. 1974. Vertebral centra in Haplolepis (Haplo- 3, pp. 1-392. lepidae, Paleonisciformes) from the Al- Allis, E. P. legheny Group, Pennsylvanian. Jour. 1889. The anatomy and development of the Paleont., vol. 48, no. 1, pp. 199-200. lateral line system in calva. Jour. Blot, J. Morph., vol. 2, pp. 463-540. 1969. Holocephales et elasmobranches. Sys- Anderson, R. Y., and D. W. Kirkland tematique. In Piveteau, J. (ed.), Traite 1960. Origin, varves and cycles ofJurassic To- de paleontologie. Paris, Masson, vol. 4, dilto Formation, New Mexico. Amer. fasc. 2, pp. 702-776. Assoc. Petrol. Geol. Bull., vol. 44, pp. Boreske, J. R. 37-52. 1974. A review of the North American fossil Andrews, S. M. amiid fishes. Bull. Mus. Comp. Zool., 1977. The axial skeleton of the coelacanth vol. 146, pp. 1-87. Latimeria. In Andrews, S. M., R. S. Bradbury, J. P., and D. W. Kirkland Miles, and A. D. Walker (eds.), Prob- 1966. Upper Jurassic aquatic Hemiptera from 1 984 SCHAEFFER AND PATTERSON: JURASSIC FISHES 81

the Todilto Formation. Abstr., 1966 Fran9ois, Y. Ann. Meeting, Geol. Soc. Amer., p. 24. 1966. Structure et developpement de la ver- Brenner, P., W. Geldmacher, and R. Schroeder tebre de Salmo et des teleosteens. Arch. 1974. Ostrakoden und Alter der Plattenkalk Zool. Exp. Gen., vol. 107, pp. 287-328. von Rubies (Sierra de Monsech, Prov. 1967. Structures vertebrales des actinopteryg- Lerida, NE-Spanien). Neues Jahrb., iens. Colloques Internatl. Centre Natl. Geol. Palaont. Monatsheft, 1974, part Rech. Sci., vol. 163, pp. 155-172. 9, pp. 449-455. Frebold, H. W. L. Cahen, L. 1957. The Jurassic Fernie group in the Ca- 1954. Geologie du Congo belge. Liege, H. nadian Rocky Mountains and foothills. Vaillant-Carmanne, 577 pp. Mem. Geol. Surv. Canada, no. 287, 197 Campbell, K. S. W. pp- 1975. Cladism and phacopid trilobites. Al- Frederickson, E. A., J. M. DeLay, and W. W. Say- cheringa, vol. 1, pp. 87-96. lor Casey, R. 1956. Ralston Formation of Canon City em- 1973. The ammonite succession at the Juras- bayment, Colorado. Bull. Amer. Assoc. sic-Cretaceous boundary in eastern En- Petrol. Geol., vol. 40, pp. 2120-2148. gland. In Casey, R., and P. F. Rawson Gardiner, B. G. (eds.), The boreal Lower Cretaceous. 1960. A revision ofcertain actinopterygian and Liverpool, Seel House Press, pp. 193- coelacanth fishes, chiefly from the Low- 266. er Lias. Bull. British Mus. Nat. Hist. Cavender, T. M. (Geol.), vol. 4, pp. 239-384. 1970. A comparison ofcoregonines and other 1967. Further notes on palaeoniscoid fishes salmonids with the earliest known te- with a classification ofthe . leostean fishes. In Lindsey, C. C., and Ibid., vol. 14, pp. 143-206. C. S. Woods (eds.), Biology ofcoregonid 1984. The relationships of the palaeoniscid fishes. Winnipeg, Univ. Manitoba Press, fishes, a review based on new specimens pp. 1-32. of Mimia and from the Cope, C. W., K. L. Duff, C. F. Parsons, H. S. Upper of Western Australia. Torrens, W. A. Wimbledon, and J. K. Wright Ibid., vol. 37, pp. 173-432. 1980. A correlation of Jurassic rocks in the Gaudant, J. British Isles. Part Two: Middle and Up- 1968. Recherches sur l'anatomie et la position per Jurassic. Geol Soc. London, Special systematique du genre Lycoptera (pois- Reportno. 15,pp. 1-109. son Teleosteen). Mem. Soc. Geol. Darton, N. H. France, vol. 47, no. 109, pp. 1-41. 1899. Jurassic formations of the Black Hills Gayet, M. ofSouth Dakota. Bull. Geol. Soc. Amer., 1982. Nouvelle extension geographique et vol. 10, pp. 383-396. stratigraphique du genre Lepidotes. Dinwiddie, G. A., and A. Clebsch, Jr. Comptes Rendus Hebd. Acad. Sci. Paris, 1973. Water resources of Guadalupe County, ser. 2, vol. 294, pp. 1387-1390. New Mexico. New Mexico State Bur. Gill, E. L. Mines and Min. Res., Hydrologic Rept. 1923. The fishes of the genus Acen- no. 3, 43 pp. trophorus. Proc. Zool. Soc. London, pp. Dunkle, D. H. 19-40. 1942. A new fossil fish of the family Lepto- Gilmore, C. W. lepidae. Cleveland Mus. Nat. Hist., Sci. 1905. Osteology ofBaptanodon Marsh. Mem. Publ., vol. 8, pp. 61-64. Carnegie Mus., vol. 2, pp. 77-129. Eastman, C. R. Greenwood, P. H. 1899a. Some new American fossil fishes. Sci- 1970. On the genus Lycoptera and its rela- ence, new ser., vol. 9, pp. 642-643. tionship with the family Hiodontidae. 1 899b. Jurassic fishes from Black Hills ofSouth Bull. Brit. Mus. Nat. Hist. (Zool.), vol. Dakota. Bull. Geol. Soc. Amer., vol. 10, 19, pp. 259-285. pp. 397-408. Gregory, W. K. Edinger, T. 1923. A Jurassic fish fauna from western Cuba, 1929. Uber kn6cherne Scleralringe. Zool. with an arrangement of the families of Jahrb. (Anat.), vol. 51, pp. 163-226. holostean ganoid fishes. Bull. Amer. 82 AMERICAN MUSEUM NOVITATES NO. 2796

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