Besprechung

COPEIA,

2002, No. 2, May 21, p. 532-535

MESOZOIC FISHES 2. SYSTEMATICS AND FOSSIL RECORD. Proceedings of the International Meeting, Buckow, 1997. Gloria Arratia and Hans-Peter Schultze (eds.). 1999. Dr. Friedrich Pfeil. Wolfratshauser Straße 27, D-81379 München. Tel.: +49-89-7428270; Fax: +49-89-7242772; E-mail: 100417.1722@compuserve.com. ISBN 3-931516-48-2. 604 p. US$ 182.50 (hardcover). – This collection of 31 papers represents much of the research originally presented at the second Mesozoic Fishes meeting, held just east of Berlin in Buckow from 6-10. July 1997, and organized by G. Arratia and H. P. Schultze. Mesozoic Fishes 2 maintains the high standards and elegant style of the first Mesozoic Fishes volume (Arratia and Viohl, 1996), including its austere price, which, however, is partly justified by the book's exceptional quality, with glossy pages and sharp reproduction of illustrations and halftones. "Mesozoic fishes" obviously does not circumscribe a monophyletic group, and many groups of fishes that appear only later in the fossil record (Tertiary) have their origins in the Mesozoic as well. But even if the time span may seem somewhat artificial, there is something in this volume for everyone interested in the systematics and evolution of fishes.

Most chapters address the systematics, morphology, and/or historical biogeography of a particular group of fishes, and almost all are based on more than fragmentary isolated remains (such as teeth or scales). There is much morphology presented in detail here, but not all of it is restricted to fossils, including two of the most informative contributions by W. E. Bemis and L. Grande (on median fin development in the paddlefish Polyodon spathula) and by E. J. Hilton and W E. Bemis (on skeletal variation in specimens of the sturgeon Acipenser brevirostrum). Bemis and L. Grande also review the literature concerned with the development of fins, exploring the classical lateral fin-fold theory (which attempts to explain the development of paired fins from lateral continuous fin folds in a model similar to the unpaired fins, but for which there is no empirical evidence; Coates, 1994). They perceive this hypothesis as "antiphylogenetic," entrenched in deveIopmental biology and paleontology, and rightly conclude that detailed empirical studies of fin development in numerous taxa should take precedence over idealistic (i.e., model-based) theories. The chapter by Hilton and Bemis addresses the par amount issue of intraspecific morphological variation, a subject that is at best difficult to properly address in fossils, and at worst simply ignored by paleontologists. These authors document the extensive variation in certain skeletal structures within a single population of the shortnose sturgeon (including dermal bones of the skull), in a style similar to the tour de force by L. Grande and Bemis (1998) on the bowfin, Amia calva. Even though the particular case-study of Hilton and Bemis may represent an extreme example of morphological variation, paleoichthyologists struggling with dermal bone terminology (that is to say, homology), small sample sizes, and taxonomic variation based solely on meager dermal bone variation, should find their paper insightful and relevant to their own work.

Three chapters are devoted to chondrichthyans: B. J. Stahl provides an overview of Mesozoic holocephalans (mostly concerning SquaIoraja, myriacanthoids, and fossil chimaeroids), L. A. Cione describes the first non-Tethyan Jurassic batoid (possessing a "rhinobatoid" Gestalt), and A. Leidner and D. Thies describe isolated euselachian and neoselachian remains (placoid scales and teeth) from the Late Jurassic of southern Germany and France. Stahl's contribution briefly reviews Mesozoic holocephalan morphology, reinforcing the fact that our understanding of these fishes is mostly ·based on their commonly fossilized tooth plates, which are difficult to interpret and may display much homoplasy (a situation not alleviated by fossil taxa known from more complete remains but which are highly modified, such as the Jurassic Squaloraja and Metopacanthus). Stahl devotes much attention to tooth plates, noting that "[i]t is to the tooth plates of Mesozoic chimaeriforms that one must look for material that allows a comprehensive comparative study of these fishes" (p. 14). However; on the very next line, Stahl affirms that "[r]ather than clarifying the relationships among the three [Mesozoic chimaeriform] suborders ... tooth plate structure onIy makes the puzzle more difficult to solve" (p. 14). Clearly, what is needed here is a comprehensive phylogenetic study of all evidence, and Stahl does not cite the most recent attempt by Lund and Grogan (1997; possibly because it was not available to her). Stahl leaves open the possibility that Mesozoic holocephalans are closely related to certain Devonian forms, which is of course true despite their "differences in age" (age is not an indicator of relationships anyway). Stahl cites for support here, among other papers, the study of Janvier and Dingerkus (1991) on the "synarcual" of Pucapampella, a taxon described as a holocephalan from the Devonian of Bolivia. This synarcual is now known to represent the parachordal (posterior) moiety of the neurocranium of an early chondrichthyan, the only one in which the ventral otic fissure remains open and possibly the sister group of elasmobranchs plus holocephalans (Maisey, 2001).

Most of the systematic and descriptive papers of Mesozoic Fishes 2 concern fossil actinopterygians: three chapters on pycnodonts (reviews of the Mesturidae and "pycnodont Bauplan" by J. R. NursalI, a description of Cretaceous pycnodonts from Uña, Spain by J. Kriwet), two on semionotiforms (a new genus from the Cretaceous of Niger by S. Wenz, new information on Dapedium by D. Thies and A. Herzog), a review of the caudal skeleton and phylogenetic position of the Triassic Prohalecites (concluded net to be a teleost; G. Arratia and A. Tintori), a new genus of saurodontid ichthyodectiform from the Cretaceous of North America (J. D. Stewart), two papers on aspidorhynchids (morphological notes on the Cuban Jurassic Aspidorhynchus arawaki and a review of the caudal skeleton of the family, both by P. M. Brito), a new genus of incertae sedis actinopterygian from the Cretaceous of North America and New Zealand (by G. Fielitz, J. D. Stewart and J. Wiffen), a review of the contribution of fossils to percopsiform and paracanthopterygian phylogeny (by A. Murray and M. V. H. Wilson), two papers on fossil osteoglossomorphs (Chinese hiodontiforms by Li G.-Q. and M. V. H. Wilson, a review of Tongxinichthys by Zhang J. and Jin F.), and new restorations of the skull and body of the Jurassic elopiform Anaethalion angustus (E. J. Poyato-Ariza). Many of these contributions discuss the relationships of the taxa involved, and some present novel or modified phylogenetic hypotheses (Arratia and Tintori, Stewart, Li and Wilson, Zhang and Jin, and Murray and Wilson). Two further papers address the historical biogeography of halecomorphs (L. Grande and W. E. Bemis) and gonorhynchiforms (T. Grande).

G. Arratia presents another detailed analysis of higher-level teleostean relationships, reinforcing her cause célèbre that it is the elopomorphs (really equivalent to elopiforms in her analyses), and not the osteoglossomorphs, that are the most basal living teleostean group (cf. Patterson and Rosen, 1977). Seven different phylogenetic analyses are presented, varying mostly in relation to the outgroup employed (whether restricted to fossil or living taxa), and achieving topologies that differ in two main regards: in relation to the succession of fossil and extant taxa falling outside of, and leading to, the Teleostei, and in relation to the phylogenetic placement of the fossil genera Ascalabos and Tharsis, which alternated between two different positions within the Teleostei in her trees [note that Arratia (p. 317, 320) erroneously claims that relationships within the Teleostei do not change between the cladograms of her figs. 19, 20, and figs. 21, 22, and that the nodes of her cladogram in fig. 22 are not labeled identically to the others]. Arratia claims that when Amia is used as the sole outgroup, the extinct "pachycormiforms become the basal taxon of Teleostei" (p. 321), but her tree (fig. 22) clearly shows the aspidorhynchids as occupying that position, with pachycormiforms forming the sister group of aspidorhynchids + Teleostei. When analyzing only extant teleosts (with Amia and Lepisosteus as outgroups), her results revert to the more traditional topology of osteoglossomorphs + (elopomorphs + all other teleosts), but osteoglossomorphs (represented by Hiodon and Heterotis) are not monophyletic (Hiodon is closer to all other teleosts), a result she obviously finds "very questionable" (p. 320). Using only extant teleosts in the ingroup, but the Jurassic Pholidophorus bechei and Leptolepis coryphaenioides as outgroups (and excluding Amia and Lepisosteus altogether), her tree once again shows elopomorphs + (osteoglossomorphs + all other teleosts), but with osteoglossomorphs monophyletic. The phylogeny Arratia seems to favor (p. 322), her fig. 19, places aspidorhynchids and lepisosteids (with the pycnodont Mesturus as a basal lepisosteid) as a monophyletic group basal to the Pachycormiformes + Teleostei (the most basal teleosts again are P. bechei and L. coryphaenoides, this phylogeny uses the Triassic Watsonulus as the outgroup). Some of Arratia's differences with earlier work on teleostean phylogeny (e.g., Patterson, 1977) concern where the dividing line between teleosts and nonteleosts is to be drawn (her claim that pachycormiforms are not teleosts but the sister group to teleosts), the relationships of aspidorhynchids, and which group is the most basal living teleost (elopomorphs or osteoglossomorphs). These issues may depend on the inclusion of fossil taxa not available to, or treated by, earlier workers (e.g., Watsonulus, varasichthyids, Orthogonikleithrus), and consideration of different characters and coding. Arratia claims (p. 322; Arratia, 1998: 1109) that Greenwood et al. (1966) also regarded elopomorphs as the most basal living teleosts, but these authors postulated a closer relationship between elopomorphs and clupeomorphs (their Division I), originating in a trichotomy from "pholidophoroid holosteans" along with osteoglossomorphs (Division II) and all other teleosts (Division III). Pycnodontiforms were mostly excluded from Arratia's matrices (except Mesturus), leaving largely untested the theory that they may be the sister-group of teleosts (cf. Gardiner et al., 1996; Nursall, 1996). Minor criticisms aside, Arratia's study on the phylogeny of basal teleosts contains a wealth of information and is another benchmark treatment of the problem but one that must be carefully distilled and compared to her earlier work (Arratia, 1997), and to the exchange between Patterson (1998) and her (Arratia, 1998). As she makes quite clear: "my results reveal that the sister group of teleosts is still unknown" (p. 327).

There are also papers reviewing regional paleoichthyofaunas (mostly actinopterygians): Triassic of Siberia (E. Sytchevskaya); Cretaceous of China (Chang M.-m.); Triassic of Switzerland (T. Bürgin); Triassic of Lombardy, Italy (A. Tintori and C. Lombardo); Mesozoic of Spain (F. J. Poyato-Arriza, A. D. Buscalioni, and J. Cartanyà); Triassic of Cataluna, Spain (J. Cartanyà); Cretaceous of southeastern Morocco (D. B. Dutheil); and Mesozoic of Chile (G. Arratia and H.-P Schultze). One paper that appears completely out of place in this book applies paleosalinity indicators (Sr isotopes) to interpret whether certain Kimmeridgian and Tithonian faunas of Germany and France were completely marine or brackish/marine (by A. Murdoch, D. Thies, and A. Baumann). Altogether, nine new genera and 11 new species are named in Mesozoic Fishes 2 (six of the genera and seven of the species, not to mention one new order, are described in the paper by Sytchevskaya). Additional taxonomic rearrangements result from some of the phylogenetic contributions, and further new taxa are mentioned but not named in some papers (e.g., by Tintori and Lombardo, and Arratia and Schultze).

As stated on the back cover, the Mesozoic Era was very important in the evolution of fishes, as many living groups became established along with the foundations for early Tertiary radiations of "higher" actinopterygians -(as revealed, e.g., in the Monte Bolca deposits of northeastern Italy). Workers on fossil fishes of this era are very grateful for the highly informative, healthy forum provided by the "Mesozoic Fishes" meetings, initiated by G. Arratia, and of which there have been three to date (Mesozoic Fishes 3 took place in Serpiano, Switzerland, from 26-31 August 2001 and was organized by A. Tintori, M. Felber, and H. Furrer). A central aspect of the meetings, and many of the papers in Mesozoic Fishes 2 (e.g., by Arratia; Murray and Wilson; Bemis and L. Grande; L. Grande and Bemis; Hilton and Bemis; T Grande), is the reciprocal illumination between the study of fossil and extant fishes, which is central to a thorough interpretation of morphological variation. Whether a character is mineralized or not is secondary to properly understanding its variation and true role as an indicator of relationships. Here again, Mesozoic Fishes 2 is definitely worth its price.

ACKNOWLEDGMENTS

I thank R. C. Schelly and W L. Smith for insightful discussion and review of the manuscript.

LITERATURE CITED

ARRATIA, G. 1997. Basal teleosts and teleostean phylogeny. Paleontol. Ichthyol. 7: 5-168.

- 1998. Basal teleosts and teleostean phylogeny: response to C. Patterson. Copeia 1998:1109-1113.

- , AND G. VIOHL. 1996. Mesozoic Fishes, Systematics and Paleoecology, Dr. Friedrich Pfeil, Munich, Germany.

COATES, M. I. 1994. The origin of vertebrate limbs, p. 169-180. In: The evolution of developmental mechanisms. M. Akam, P. Holland, and G. Wray (eds.). Development 1994 (suppl. 1).

GARDINER, B. G., J. G. MAISEY, AND T. J. LITTLEWOOD, 1996. Interrelationships of basal neopterygians, p. 117-146. In: Interrelationships of fishes. M. L. J. Stiassny, L. Parenti, and G. D. Johnson (eds.). Academic Press, San Diego, CA.

GRANDE, L., AND W. E. BEMIS. 1998. A comprehensive phylogenetic study of amiid fishes (Amiidae) based on comparative skeletal anatomy An empirical search for interconnected patterns of natural history. J. Vert. Paleontol. (suppl.) 18: 1-690. GREENWOOD, P. H., D. E. ROSEN, S. H. WEITZMAN, AND G. S. MYERS. 1986. Phyletic studies of teleostean fishes, with a provisional classification of living forms. Bull. Am. Mus. Nat. Hist. 131: 339-456.

JANVIER, P., AND G. DINGERKUS. 1991. Le synarcual de Pucapampella Janvier et Suárez-Riglos: un preuve de l'existence d'Holocéphales (Vertebrata, Chondrichthyes) dès le Dévonien. Com. Rend. Acad. Sci., Paris 312: 549-552.

LUND, R. AND E. GROGAN. 1997. Relationships of the Chimaeriformes and the basal radiation of the Chondrichthyes. Rev. Fish Biol. Fish. 7: 65-123. MAISEY, J. G. 2001. A primitive chondrichthyan braincase from the middle Devonian of Bolivia, p. 263 288. In: Major events in early vertebrate evolution, Palaeontology, phylogeny, genetics and development. P E. Ahlberg (ed.). Taylor and Francis, London.

NURSALL. J. R. 1996. The phylogeny of pycnodont fishes, p. 125-152. In: Mesozoic fishes, systematics, and paleoecology. G. Arratia and G. Viohl (eds.). Dr. Friedrich Pfeil, Munich, Germany.

PATTERSON, C. 1977. The contribution of paleontology to teleostean phylogeny, p. 579-643. In: Major patterns in vertebrate evolution. M. K. Hecht, P. G. Goody, and B. M. Hecht (eds.). Plenum Press, New York.

- 1998. Comments on basal teleosts and teleostean phylogeny, by Gloria Arratia. Copeia 1998: 1107-1109.

- , AND D. E. ROSEN. 1977. Review of ichthyodectiform and other Mesozoic teleost fishes and the theory and practice of classifying fossils. Bull. Am. Mus. Nat. Hist. 158:81-172.

MARCELO R. DE CARVALHO, Division of Paleontology, American Museum of Natural History, Present address: Department of Ichthyology, AMNH, Central Park West at 79th Street, New York, New York 10024-5192; E-mail: marcelo@amnh. org.

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