Mesozoic Fishes 1 – Systematics and Paleoecology

Preface 7
Acknowledgments 7
Marcelo R. de CARVALHO & John G. MAISEY:
The phylogenetic relationship of the Late Jurassic shark Protospinax Woodward 1919 (Chondrichthyes: Elasmobranchii) 9
Paulo M. BRITO & Bernard SERET:
The new genus Iansan (Chondrichthyes, Rhinobatoidea) from the Early Cretaceous of Brazil and its phylogenetic relationships 47
Alberto L. CIONE:
The extinct genus Notidanodon (Neoselachii, Hexanchiformes) 63
Masatoshi GOTO, Teruya UYENO & Yoshitaka YABUMOTO:
Summary of Mesozoic elasmobranch remains from Japan 73
Hans-Peter SCHULTZE:
The scales of Mesozoic actinopterygians 83
François J. MEUNIER & Mireille GAYET:
A new polypteriform from the Late Cretaceous and the middle Paleocene of South America 95
Detlev THIES & Alexander MUDROCH:
Actinopterygian teeth from the Late Jurassic (Kimmeridgian) of N Germany 105
J. Ralph NURSALL:
Distribution and ecology of pycnodont fishes 115
J. Ralph NURSALL:
The phylogeny of pycnodont fishes 125
Sylvie WENZ & Paulo M. BRITO:
New data about the lepisosteids and semionotids from the Early Cretaceous of Chapada do Araripe (NE Brazil): Phylogenetic implications 153
Andrea TINTORI:
Paralepidotus ornatus (AGASSIZ 1833-43): A semionotid from the Norian (Late Triassic) of Europe 167
Lance GRANDE:
Using the extant Amia calva to test the monophyly of Mesozoic groups of fishes 181
Gloria ARRATIA & Paul LAMBERS:
The caudal skeleton of pachycormiforms: Parallel evolution? 191
Gloria ARRATIA:
Reassessment of the phylogenetic relationships of certain Jurassic teleosts and their implications on teleostean phylogeny 219
Gloria ARRATIA:
The Jurassic and the early history of teleosts 243
Mei SHEN:
Fossil "osteoglossomorphs" from East Asia and their implications for teleostean phylogeny 261
Niels BONDE:
Osteoglossids (Teleostei: Osteoglossomorpha) of the Mesozoic. Comments on their interrelationships 273
Guo-qing LI:
A new species of Late Cretaceous osteoglossid (Teleostei) from the Oldman Formation of Alberta, Canada, and its phylogenetic relationships 285
Terry GRANDE:
The interrelationships of fossil and Recent gonorynchid fishes with comments on two Cretaceous taxa from Israel 299
Francisco José POYATO-ARIZA:
A revision of Rubiesichthys gregalis WENZ 1984 (Ostariophysi, Gonorynchiformes), from the Early Cretaceous of Spain 319
Francisco José POYATO-ARIZA:
The phylogenetic relationships of Rubiesichthys gregalis and Gordichthys conquensis (Ostariophysi, Chanidae), from the Early Cretaceous of Spain 329
Yael CHALIFA:
New species of Enchodus (Aulopiformes: Enchodontidae) from the Northern Negev, Israel, with comments on evolutionary trends in the Enchodontoidei 349
Mark V. H. WILSON & Alison M. MURRAY:
Early Cenomanian acanthomorph teleost in the Cretaceous Fish Scale Zone, Albian/Cenomanian Boundary, Alberta, Canada 369
J. D. STEWART:
Cretaceous acanthomorphs of North America 383
Paul H. LAMBERS:
A redescription of the coelacanth Macropoma willemoesii VETTER from the lithographic limestone of Solnhofen (Upper Jurassic, Bavaria) 395
Anne KEMP:
Triassic lungfish from Gondwana 409
Werner SCHWARZHANS:
Otoliths from the Maastrichtian of Bavaria and their evolutionary significance 417
Dirk NOLF & Gary L. STRINGER:
Cretaceous fish otoliths – a synthesis of the North American record 433
Mee-mann CHANG & Fan JIN:
Mesozoic fish faunas of China 461
Laurence BELTAN:
Overview of systematics, paleobiology, and paleoecology of Triassic fishes of northwestern Madagascar 479
Olivier RIEPPEL, René KINDLIMANN & Hugo BUCHER:
A new fossil fish fauna from the Middle Triassic (Anisian) of North-Western Nevada 501
Günter VIOHL:
The paleoenvironment of the Late Jurassic fishes from the southern Franconian Alb (Bavaria, Germany) 513
Shelton Pleasants APPLEGATE:
An overview of the Cretaceous fishes of the quarries near Tepexi de Rodríguez, Puebla, México 529
Luis ESPINOSA-ARRUBARRENA & Shelton Pleasants APPLEGATE:
A paleoecological model of the vertebrate bearing beds in the Tlayúa quarries, near Tepexi de Rodríguez, Puebla, México 539
David G. SENN:
Environments and functional anatomy of certain Mesozoic fishes 551
Toni BÜRGIN:
Diversity in the feeding apparatus of perleidid fishes (Actinopterygii) from the Middle Triassic of Monte San Giorgio (Switzerland) 555
Andrea TINTORI:
The field excursion in Northern Italy 567

Marcelo R. de CARVALHO & John G. MAISEY:
Phylogenetic relationships of the Late Jurassic shark Protospinax WOODWARD 1919 (Chondrichthyes: Elasmobranchii)
[pp. 9-46, 9 figs., 7 apps.]

Two new specimens of Protospinax annectans WOODWARD 1919 are reported. The new material provides an opportunity to re-evaluate the phylogenetic relationships of Protospinax in the light of modern cladistic analyses. A revised data matrix, including Protospinax, largely extracted from the work of SHIRAI (1992a) is presented. This data matrix contains different interpretations for some of SHIRAI's characters, as well as a few characters not considered by him (pertaining to, e.g., the basihyal and puboischiadic bar). After four iterations of successive weighting, 8 minimum-length trees were found (L = 699 steps, c = .60, r = .75), of which the strict consensus is very similar to SHIRAI's phylogeny. Squaleans share various apomorphic characters, including a basitrabecular process and loss of suborbital shelf (which are also present in Protospinax), but the pharyngobranchial blade is best optimized as a galeomorph synapomorphy. Hexanchiformes (including Chlamydoselachus) is paraphyletic, but only if Echinorhinus is coded as derived for two characters related to the ectethmoid process (as herein re-defined). A character putatively homologous for hexanchiforms (single dorsal fin) was included to further test their monophyly, but this is still not supported. Protospinax is resolved as a very derived member of the squalean clade and is the sister-group to Recent hypnosqualeans (a group comprising squatinoids, pristiophoroids and batoids), with which it shares ten apomorphic characters (eight with homoplastic distribution). Protospinax retains a basioccipital fovea which is secondarily lost in other hypnosqualeans. An anal fin, basal angle and ectethmoid process are all absent in Protospinax, which has ventrally fused coracoids, calcified vertebral centra and an orbital articulation typical of most "orbitostylic" sharks. The phylogenetic position of Protospinax is unchanged even when SHIRAI's characters are left unmodified and missing characters in Protospinax are excluded from the analysis. Without successive weighting, 252 equally most parsimonious trees (L = 163 steps, c = .60, r = .75) are produced, all depicting Protospinax as a hypnosqualean (forming a basal trichotomy with Squatina and the node uniting pristiophoroids and batoids). Although resolution within some higher squalean components varied in our different analyses, Protospinax and other hypnosqualeans remained monophyletic. SHIRAI's "hypnosqualean group" is formally recognized as the Superorder Hypnosqualea.

Paulo M. BRITO & Bernard SERET:
The new genus Iansan (Chondrichthyes, Rhinobatoidea) from the Early Cretaceous of Brazil and its phylogenetic relationships
[pp. 47-62, 7 figs., 4 apps.]

The rhinobatoid Iansan beurleni SANTOS 1968 is redescribed on the basis of new material from the Santana formation, Lower Cretaceous (Albian) of Brazil.

A data matrix built with 39 characters from NISHIDA (1990), plus six new characters and four additional taxa were used to generate an hypothesis of phylogenetic relationships of rhinobatoids. The paraphyletic condition of rhinobatoid clades, suggested by NISHIDA is confirmed. The new genus Iansan is a basal member of the clade composed of some "rhinobatoids", the rajoids and the myliobatoids.

Alberto L. CIONE: The extinct genus Notidanodon (Neoselachii, Hexanchiformes)
[pp. 63-72, 4 figs., 1 tab.]

The systematics of the hexanchid genus Notidanodon is discussed. N. pectinatus is considered a nomen dubium. Notidanodon dentatus showed bipolar distribution in the Late Cretaceous which was probably climatically controled. Notidanodon probably represents a lineage. The evolution of hexanchid size as evidenced by teeth is analyzed.

Masatoshi GOTO, Teruya UYENO & Yoshitaka YABUMOTO: Summary of Mesozoic elasmobranch remains from Japan
[pp. 73-82, 5 figs., 4 tabs.]

In Japan, most fossil fishes are known from the Cenozoic. However, in recent years, many fish teeth and scales have been found and reported from Mesozoic and Paleozoic beds. On the basis of published and unpublished data, fossil elasmobranchs from the Mesozoic of Japan are classified into 47 species: four species of hybodontoids from the Triassic, four species of hybodontoids from the Jurassic, and 39 species of hybodontoids, ptychodontoids, hexanchoids, orthacodontoids, pristiophoroids, lamnoids, palaeospinacoids, and sclerorhynchoids from the Cretaceous. All of them were yielded from marine sediments.

Hans-Peter SCHULTZE: The scales of Mesozoic actinopterygians
[pp. 83-93, 6 figs.]

Cycloid scales (elasmoid scales with circuli) are a unique character of teleosts above the level of Pholidophorus and Pholidophoroides. Cycloid scales have two layers. A bony layer, usually acellular, is superimposed on a basal plate composed of partially mineralized layers of plywoodlike laminated collagen fibres. The tissue of the basal layer is refered to here as elasmodin. Basal teleosts (sensu PATTERSON 1973) possess rhombic scales with a bony base overlain by ganoin (lepidosteoid ganoid scale). Amioid scales (elasmoid scales with longitudinally to radially arranged ridges or rods on the overlapped field) are found within halecomorphs. This scale type evolved more than once within primitive actinopterygians and other osteichthyan fishes. It may have even developed twice within halecomorphs, in Caturidae and Amiidae, from rhombic scales of lepidosteoid type. Some basal genera of halecomorphs show remains of a dentine layer between ganoin and bone that is characteristic of actinopterygians below the halecostome level. The Semionotidae placed at the base of the Halecostomi, exhibit scale histology transitional between the palaeoniscoid and lepidosteoid scale type.

François J. MEUNIER & Mireille GAYET: A new polypteriform from the Late Cretaceous and the middle Paleocene of South America
[pp. 95-103, 16 figs., 1 tab.]

SEM and paleohistological studies of scales and fragments of isolated cranial ganoid bones allow discrimination of a new polypteriform in the late Maastrichtian and middle Paleocene of Bolivia and Brazil. The scales show the three superimposed layers (stratified ganoin, dentine, and bony basal plate) and the typical orthogonal plywood structure (between dentine and bony basal plate) of polypterid scales. These scales differ from those of other polypterids by a more developped vascular network of dentine, long parallel odontoblastic canaliculi and by a more regular organization of centrifugal oriented osteocytes around ascending vascular canals of the basal plate. A new species and genus, Pollia suarezi n. g., n. sp, is erected and placed incertae sedis within the Polypteriformes.

Detlev THIES & Alexander MUDROCH: Actinopterygian teeth from the Late Jurassic (Kimmeridgian) of N Germany
[pp. 105-114, 2 pls., 5 figs.]

Bulk sampling of the Late Jurassic (Kimmeridgian) section at the village of Oker in front of the northern slope of the Harz mountains (N Germany) has yielded approximately 7,000 isolated actinopterygian teeth. The following genera were identified in our material by comparison of the isolated teeth to the dentition of articulated skeletons of actinopterygians from the lithographic limestones of France (Cerin, Kimmeridgian) and the lithographic limestones of South Germany (Tithonian): Lepidotes, Proscinetes, Macromesodon, Coelodus, Histionotus, Macrosemius? or Notagogus?, Ionoscopus, Caturus, Sauropsis, Belonostomus, and Thrissops. Within the entire material the teeth of Lepidotes are the most frequent, followed by those of Caturus. Statistical consideration shows that the composition of the fauna strongly varies from bed to bed within the section. This variation is explained best by changes of the paleosalinity.

J. Ralph NURSALL: Distribution and ecology of pycnodont fishes
[pp. 115-124, 3 figs.]

Pycnodont fishes persisted from Late Triassic (Norian) to Eocene (Ypresian-Lutetian) times, about 175 million years. They inhabited shallow, marginal, often reefal seas, located subtropically or tropically around Tethys and into the developing Atlantic. They were deep-bodied, manoeuvrable fish, restricted to a durophagous habit. Pioneers in their mode of life, they were supplanted by the teleosts, which were evolving rapidly into much more diversified forms.

J. Ralph NURSALL: The phylogeny of pycnodont fishes
[pp. 125-152, 23 figs.]

†Pycnodontiformes is a monophyletic group with two suborders, Gyrodontoidei and Pycnodontoidei, each with two families. Another family is proposed to represent a stem group to the order. Five families remain incertae sedis. The order is the sister group to teleosts, within Halecostomi. Amiiformes is the sister group to pycnodonts + teleosts. The outgroup to Halecostomi is represented by Dapedium spp.

Sylvie WENZ & Paulo M. BRITO: New data about the lepisosteids and semionotids from the Early Cretaceous of Chapada do Araripe (NE Brazil): Phylogenetic implications
[pp. 153-165, 5 figs.]

Complete or nearly complete specimens of the semionotid Araripelepidotes and the lepisosteid Obaichthys are found together in the Albian fossiliferous nodules from the Chapada do Araripe.

The Araripelepidotes dermal skull characters are very close to those of Lepidotes, although some of the braincase characters are proper to the Araripe's taxa. Obaichthys decoratus and/or Obaichthys? laevis present a typical lepisosteid braincase, although the dermal bones present a more primitive arrangement than typical lepisosteids.

The data brought about in this work leads to new insights into following two questions: firstly the affinities of Araripelepidotes within the semionotids and secondly the relationships between lepisosteids and semionotids. The braincase characters that have been used until now to bring together lepisosteids and semionotids, and the more recent use of dermal characters, are either lepisosteid synapomorphies or characters that are not unique to lepisosteids and semionotids and require precaution.

Andrea TINTORI: Paralepidotus ornatus (AGASSIZ 1833-43): A semionotid from the Norian (Late Triassic) of Europe
[pp. 167-179, 6 figs.]

Paralepidotus ornatus is a rather common semionotid in most Norian marine localities of Europe. New findings of well preserved material in northern Italy allow preliminary restoration and confirm the inclusion of Paralepidotus within the Semionotidae s.s. Since growth stages are well represented, ontogenetic variation is recorded. Juveniles are more slender than adults and their dentition is not like the powerful crushing one of large specimens. Bone and scale ornamentation appears at 11-15 cm standard length (SL) and reaches its adult aspect at about 30 cm SL. These modifications imply a change in environment during the life of an individual Paralepidotus, from superficial open waters to shallow mollusc banks when teeth reach the strictly grinding shape at 25-30 cm SL. A change in diet is also implied by the change in dentition.

Lance GRANDE: Using the extant Amia calva to test the monophyly of Mesozoic groups of fishes
[pp. 181-189, 6 figs.]

Since 1982, four phylogenies for Amiidae have been proposed, based mostly on examination of Mesozoic taxa, and apparently without detailed examination of the living amiid, Amia calva. One proposed phylogeny was based on a phenogram and provided little resolution to family interrelationships. The other three proposed phylogenies were based on cladograms. When data for Amia is corrected for the three cladograms based on examination of A. calva, the cladograms all change substantially. It is clear that the interrelationships within Amiidae are still very poorly understood.

Gloria ARRATIA & Paul LAMBERS: The caudal skeleton of pachycormiforms: Parallel evolution?
[pp. 191-218, 15 figs.]

Hypurals alone or the parhypural plus an indetermined number of hypurals form the hypural plate of pachycormiforms. An unusual series follows posterior to the series of unpaired and a few paired neural spines bearing membranous outgrowths. This series is formed by unpaired, median, thick "uroneurals" that extend posteriorly dorsal to the hypural plate, in between the hemitrichia. The "uroneurals" are interpreted here as expanded neural spines, some of them still bearing part of their arches, of the last preural vertebrae and ural vertebrae, whereas they were identified as uroneurals and the pachycormiforms placed within Teleostei sensu PATTERSON (1973). The new morphological evidence questions the teleostean synapomorphies proposed by PATTERSON (1977) as well as the inclusion of pachycormiforms and other basal "teleosts" within the Teleostei.

The broadening of the neural spines, "uroneurals", and "epurals", the thickness of haemal arch and spines, the fusion of hypurals, the increase of number of caudal rays, the peculiar structure of rays and fulcra, and the almost symmetrical position of dorsal and ventral elements of the caudal skeleton with respect to the body axis are interpreted as features increasing the stiffness and propulsion abilities of the tail.

Gloria ARRATIA: Reassessment of the phylogenetic relationships of certain Jurassic teleosts and their implications on teleostean phylogeny
[pp. 219-242, 8 figs., 3 apps.]

The phylogenetic relationships of 15 Jurassic teleostean genera from Europe (e.g., †Allothrissops, †Anaethalion, †Ascalabos, †Leptolepis, †Orthogonikleithrus), Asia (e.g., †Lycoptera), Central America (e.g., †Luisichthys), South America (e.g., †Domeykos, †Varasichthys), and Australia (e.g., †"Leptolepis" talbragarensis) were studied using the cladistic principles. Two cladistic analyses were performed: (1) including only Jurassic teleosts, and (2) including Jurassic, certain Cretaceous and Tertiary teleosts, and extant teleosts. The two cladistic analyses show that †Ascalabos is the sister group of a clade comprising [†Domeykos + [†Varasichthys + [†Protoclupea + †Luisichthys]]]; this clade and the ichthyodectiforms are the only monophyletic assemblages that can be identified among the Jurassic teleosts. The phylogenetic position of certain advanced genera such as †Anaethalion, †Leptolepides, †Orthogonikleithrus, and †Teleost n.gen. changes when the fishes are compared only with other Jurassic teleosts or when they are compared with Jurassic plus extant teleosts. The present study confirms that †Anaethalion is the oldest elopomorph. †Teleost n.gen is the oldest known ostariophysan. The phylogenetic position of †Orthogonikleithrus is unresolved: it appears as the sister group of the clade including [†Erichalcis + [Esox + Salmonidae]] in one topology, but as the sister group of the ostariophysans, "salmoniforms" plus advanced teleosts in another topology. Basal Jurassic teleosts hold crucial evidence of the phylogenetic relationships of extant teleostean taxa. Examination of new Jurassic forms and reinterpretation of certain characters change previous hypotheses of teleostean relationships (e.g., PATTERSON & ROSEN 1977, LAUDER & LIEM 1983). The results support ARRATIA's (1990, 1991) conclusions that elopomorphs are more primitive than osteoglossomorphs, and that elopomorphs are the most primitive of the major teleostean clades. The phylogenetic relationships among the osteoglossomorphs, clupeomorphs, ostariophysans, and other teleosts are still unresolved, and therefore the higher taxa of Teleostei such as Elopocephala, Clupeocephala, and Euteleostei sensu PATTERSON & ROSEN (1977) and LAUDER & LIEM (1983) are not natural assemblages.

Gloria ARRATIA: The Jurassic and the early history of teleosts
[pp. 243-259, 6 figs., 1 tab.]

Whether the history of Teleostei begins in the Triassic or the Jurassic depends on the composition of the group. Teleostei sensu PATTERSON (1973, 1977), includes pachycormiforms, aspidorhynchiforms, and pholidophorids within the group, but is known in the Triassic only by pholidophorids. PATTERSON's (1977) cladogram indicates that pachycormiforms and aspidorhynchiforms should be older groups than the pholidophorids, but the known fossil record does not support that hypothesis because Triassic pachycormiforms and aspidorhynchiforms are unknown. New morphological evidence questions the assignment of pachycormiforms and aspidorhynchiforms to Teleostei.

Of about 450 living teleostean families, only 15 can be recognized in the Mesozoic (Cretaceous). Living teleosts are extremely diverse, both morphologically and in terms of number of species. Some major clades and morphological patterns can be traced back to the Late Jurassic, e.g., the elopomorphs with Anaethalion, the ostariophysans with an undescribed fish from southern Germany, and probably the "salmoniforms" with Orthogonikleithrus, and to the Early Cretaceous or Late Jurassic the osteoglossomorphs with numerous genera.

Three radiations of teleostean fishes can be identified in the Jurassic. The first is represented by the family Leptolepidae with two genera, which occur in the Early Jurassic. About five genera known from the Middle Jurassic represent the second teleostean radiation. About 20 genera and about five families known from the Late Jurassic represent the third radiation. Some of the Jurassic genera were described as Teleostei incertae sedis and not assigned to families. Among the Jurassic teleosts only two monophyletic assemblages are recognized; the ichthyodectiforms and the unnamed clade [Ascalabos + [Domeykos + [Varasichthys + [Protoclupea + Luisichthys]]]]. The phylogenetic relationships of the latter assemblage support the hypothesis that the European locality Solnhofen (with Ascalabos) is the sister region of the Chilean locality El Profeta (with Domeykos, Varasichthys, and Protoclupea) plus the Cuban locality Pinar del Río (with Luisichthys). This is the first evidence from the teleostean fossil record that supports a connection between European and Chilean fish faunas through the Tethys Seaway.

Mei SHEN: Fossil "osteoglossomorphs" from East Asia and their implications for teleostean phylogeny
[pp. 261-272, 4 figs., 1 tab., 1 app.]

Osteoglossomorphs are primitive teleosts which live in freshwaters of North America and almost all continents of the southern hemisphere. Many fossils, thought to be osteoglossomorphs, are also found in Mesozoic deposits of China. These fossils differ from each other in many morphological characters. Most of them possess a full neural spine on the first preural centrum and only 18 principal rays in the caudal fin. Because these characters were regarded as unique to the Osteoglossomorpha, these fishes were referred to this supercohort without exception. However, these two and most other characters of osteoglossomorphs are found in other non-osteoglossomorph teleosts, and the distribution of these characters implies that they may be primitively shared by early members of a number of teleostean groups.

Three analyses are made. The first two are phylogenetic examinations of the Osteoglossomorpha, based on the synapomorphies used by PATTERSON & ROSEN (1977) to characterize the Osteoglossomorpha, the Elopocephala, the Clupeocephala, and the Euteleostei. The results suggest that the Osteoglossomorpha is not a monophyletic group according to PATTERSON & ROSEN's definition. Analysis 3 is investigation of the relationships among Chinese fossil osteoglossomorphs. Thirty-nine characters are coded for 11 fossil osteoglossomorphs (†Lycoptera, †Paralycoptera, †Tonxinichthys, †Pulinia, †Plesiolycoptera, †Kuntulunia, †Huashia, †Changichthys, †Tanolepis, †Jiaohichthys, and †Yanbiania,) and seven representatives (Elops, Hiodon, Osteoglossum, Notopterus, Denticeps, Chanos, and Coregonus) of extant teleosts. The results illustrate that some of the so-called osteoglossomorphs in China are not related to the extant osteoglossomorphs. Only four fossil genera belong to the Osteoglossomorpha. Among this group, [†Kuntulunia + †Huashia] is the sister group to the [Osteoglossum + Notopterus], †Yanbiania and †Tanolepis are related to Hiodon. †Pulinia, on the other hand, is the sister group to Denticeps, Chanos, and Coregonus. †Plesiolycoptera and †Changichthys, together with the Osteoglossomorpha and [†Pulinia + Denticeps, Chanos, and Coregonus], constitute a clade, but their relationships within this clade is unsolved. Other fossil "osteoglossomorphs" locate between this clade and Elops. Elops is most primitive within analyzed taxa, except †Pholidophorus bechei, †Leptolepis coryphaenoides, and †Allothrissops mesogaster. They are outgroups of present analyses.

Niels BONDE: Osteoglossids (Teleostei: Osteoglossomorpha) of the Mesozoic. Comments on their interrelationships
[pp. 273-284, 4 figs.]

Osteoglossids comprise a family of primitive teleosts with a few living genera and species distributed in freshwater of the southern hemisphere (and SE-Asia) – apparently a typical Gondwanaland distribution. The family is most closely related to pantodontids (by some included in the family), Mormyriformes and Notopteriformes, all groups with a poor or no fossil record.

Several Early Cretaceous genera are said to be osteoglossids, some supposed to represent the extinct (mainly Early Tertiary) subfamily Phareodontinae, which would therefore indicate a very early split between the two living subgroups of the family; others are supposed to be related to the evolutionary lineage of just one Recent genus, thus indicating very early split between living sister genera of Heterotinae. These ideas of interrelationships are critically evaluated as a basis for understanding the radiation of the osteoglossids in Early Tertiary – with many members even in the seas of present day northwestern Europe.

LI Guo-qing: A new species of Late Cretaceous osteoglossid (Teleostei) from the Oldman Formation of Alberta, Canada, and its phylogenetic relationships
[pp. 285-298, 7 figs.]

A single specimen of Campanian age from the Oldman Formation of Dinosaur Provincial Park, Alberta, Canada, represents a new genus and species of osteoglossomorph fishes. It is thought to be an early member of Osteoglossinae based on the possession of five possible synapomorphies of the subfamily: 1, opercle subsemicircular in shape; 2, horizontal arm of the preopercle short with its anterior end extending only to the level of the postero-ventral corner of the third infraorbital; 3, preoperculo-mandibular canal opening in a groove on the horizontal arm of the preopercle; 4, angle of jaws posteriorly located; 5, first pectoral fin ray particularly strong and long. The combination of two synapomorphies (anterior portion of frontal nearly twice as broad as the posterior portion and a laterally-compressed deep body with a large pleuroperitoneal cavity) with one other character state (23-25 precaudal vertebrae) suggests that this new Late Cretaceous osteoglossomorph fish is the sister-species of the clade including †Phareodus and †Brychaetus.

Terry GRANDE: The interrelationships of fossil and Recent gonorynchid fishes with comments on two Cretaceous taxa from Israel
[pp. 299-318, 14 figs., 2 tabs., 3 apps.]

The interrelationships of fossil and Recent gonorynchid fishes are reviewed. The monophyly of the family is cladistically verified, in part, by caudal skeleton and dentition characters. The family can be divided into two monophyletic clades; one consisting of Gonorynchus and †Notogoneus as sister taxa and the other consisting of †Charitosomus. The systematic placement of two Cretaceous taxa, †Ramallichthys and †Judeichthys, is reviewed. Data indicate that both taxa should be included within the Gonorynchidae, and possibly synonymized with the genus †Charitosomus.

Francisco José POYATO-ARIZA: A revision of Rubiesichthys gregalis WENZ 1984 (Ostariophysi, Gonorynchiformes), from the Early Cretaceous of Spain
[pp. 319-328, 4 figs.]

The small fish Rubiesichthys gregalis, from the Early Cretaceous of Montsec and Las Hoyas (Spain), is redescribed on the basis of abundant new material; complementary anatomical information is now available, mostly concerning the anteriormost vertebrae. A modified diagnosis of the genus and the species, as well as new restorations, are presented. This revision confirms the similarities of Rubiesichthys gregalis with Gordichthys conquensis and other gonorynchiform fishes traditionally considered chanids.

Francisco José POYATO-ARIZA: The phylogenetic relationships of Rubiesichthys gregalis and Gordichthys conquensis (Ostariophysi, Chanidae), from the Early Cretaceous of Spain
[pp. 329-348, 8 figs.]

Two small teleostean fishes from the Early Cretaceous of Spain, Rubiesichthys gregalis and Gordichthys conquensis were considered related to the gonorynchiform family Chanidae when first described. Here, a cladistic analysis was carried out to establish their phylogenetic relationships. The ingroup included living and fossil forms traditionally assigned to the Chanidae, as well as other otophysans. A set of 55 characters of six genera was polarized by outgroup comparison with Elops and Diplomystus. The results confirm that the Otophysi (represented by the cyprinid Opsariichthys) is the sister group to the Gonorynchiformes, which are monophyletic. Within the order Gonorynchiformes, Chanos is the sister group to [Tharrhias + [Rubiesichthys + Gordichthys]]. This set of relationships shows that the family Chanidae is a clade; its monophyly is supported by 11 synapomorphies, eight of which are uniquely derived characters. Rubiesichthys gregalis and Gordichthys conquensis are, therefore, chanid fishes, as previously assigned; in addition, they are sister taxa. This analysis also indicates that some fossil forms are included with the Recent Chanos in the family Chanidae. This hypothesis of relationships also implies that fusion within the caudal endoskeleton arose independently in the Chanidae. In addition, the resultant interrelationships present temporal incongruences. These problems reveal the need to re-examine other fossil forms that have been traditionaly considered chanids. Such a re-examination would allow clarification of the diagnosis of the family, as well as the phylogenetic relationships within the family, and with other gonorynchiforms.

Yael CHALIFA: New species of Enchodus (Aulopiformes: Enchodontidae) from the Northern Negev, Israel, with comments on evolutionary trends in the Enchodontoidei
[pp. 349-367, 8 figs., 4 tabs.]

A new species, Enchodus zinensis (Enchodontoidei: Enchodontidae) is described from Early Maastrichtian beds in the Northern Negev, Israel. The species is characterized by its large size; an anteromedial projection on the dentary; posteroventral keel on the dentary; palatine with a smooth posterior edge directed anterodorsally; toothed maxilla; smooth and laterally compressed teeth with two cutting edges; transverse process on precaudal vertebrae; a fan-like hypural plate with a urapophysis on the terminal vertebra and the hypural plate base.

A discussion of trends in the evolution of body shape in the Enchodontoidei indicates two main lines, a trend toward shortening of the body, which occurred in Enchodontidae, and a trend toward lengthening of the body, which occurred differently in Enchodontidae and Eurypholidae.

Mark V. H. WILSON & Alison M. MURRAY: Early Cenomanian acanthomorph teleost in the Cretaceous Fish Scale Zone, Albian/Cenomanian boundary, Alberta, Canada
[pp. 369-382, 5 figs.]

The Fish Scale Zone marks the Albian/Cenomanian boundary in the Western Interior of Canada. Exposed in northwestern Alberta within the Shaftesbury Formation, it contains scales as well as abundant disarticulated bones of teleostean fishes. The fish remains occur immediately above the presumed boundary and include †Ichthyodectidae, †Osmeroides, †Enchodontiformes, and Acanthomorpha. The acanthomorph bones are the oldest described North American acanthomorph fossils and among the oldest anywhere of that group. Numerous bones of the skull and pectoral girdle appear to belong to a single species; this is confirmed by a single associated but mostly disarticulated skull combining many of the distinctive elements. Some bones have a striking similarity to those of the paracanthopterygian genus †Sphenocephalus found previously in the Campanian of Germany. The fossils are described as †Xenyllion zonensis gen. et sp.nov. in Family †Sphenocephalidae. They extend the range of the Paracanthopterygii by about 20 My, eliminating their previously anomalous late appearance in the fossil record as compared with other acanthomorph higher taxa, but further emphasizing the late appearance of what are currently thought to be cladistically more primitive freshwater percopsiforms that do not appear in the record until the Paleocene.

J. D. STEWART: Cretaceous acanthomorphs of North America
[pp. 383-394, 2 figs.]

Prior to 1992, the oldest published acanthomorph teleost from North America was of middle Turonian age. North American acanthomorphs extend back to the late Albian or early Cenomanian stage in Utah (Mowry Shale; zone of †Neogastroplites americanus) and Alberta (Shaftesbury Formation, zone of †Neogastroplites maclearni: LECKIE et al. 1992). The same sphenocephalid genus occurs at both these localities. A single polymixiid genus occurs in late Santonian sediments of Kansas. Holocentrids are known from Turonian, Coniacian, and Santonian horizons in Kansas; the Coniacian of Colorado, and Maastrichtian rocks of New Jersey. Hoplopteryx, a trachichthyoid, occurs in Campanian rocks in Texas. Unassigned fin spines, probably from acanthomorphs, have been found in late Turonian and Campanian strata in Kansas, and in Campanian sediments of Alabama.

The relatively high energy regime of many Cretaceous formations in North America make them unsuitable for preserving relatively small remains of acanthomorph fishes. Nonetheless, at least five North American formations (Gober Chalk, Niobrara Chalk, Carlile Shale, Mowry Shale, and Hornerstown) have produced partially to fully articulated skeletons of acanthomorphs.

Only four acanthomorph families have been demonstrated in Cretaceous rocks in North America based on osteological remains. This is considerably less than in some other areas of the world. Grain size, bioturbation, evaporites, and high energy regimes have rendered many North American Cretaceous rocks as poor candidates for preserving articulated remains of these relatively small fishes.

Paul H. LAMBERS: A redescription of the coelacanth Macropoma willemoesii VETTER from the lithographic limestone of Solnhofen (Upper Jurassic, Bavaria)
[pp. 395-407, 10 figs.]

The coelacanth Macropoma willemoesii VETTER from the Upper Jurassic lithographic limestone of Bavaria is redescribed. It is shown that the species belongs to the genus Macropoma. M. willemoesii shares with other Macropoma species an entirely covered cheek, dermal bones ornamented with many large tubercles, a distinctive scale ornamentation and a reduced median lobe of the caudal fin. The sigmoid shape of the ventral border of the pterygoid is considered an autapomorphy of the genus.

Anne KEMP: Triassic lungfish from Gondwana
[pp. 409-416, 3 figs.]

Analysis of the reptilian and amphibian faunas of a number of Triassic localities in Africa, Antarctica, and Australia demonstrates that the fauna of these deposits in Australia differs from faunas found in Antarctica and Africa. A number of fish species, including lungfish, also occur in these localities. One of the lungfish species in the Lower Triassic deposits of Australia is also present in the Lower Triassic Cynognatus zone near Bergersdorp, Orange Free State, South Africa. Aquatic faunas of the Early Triassic of Australia might have been more similar to those of other Gondwanan deposits than are the terrestrial or amphibious faunas. This lungfish, previously referred to Ceratodus cf. C. philippsi, belongs in the genus Ptychoceratodus, described from extensive material from the Lower Keuper of Kupferzell in Germany. This record extends the geographic range of Ptychoceratodus to the Southern Hemisphere, and its geological range from the Middle to the Early Triassic.

Ptychoceratodus philippsi is based on well preserved skull bones, jaws and associated tooth plates from Africa and tooth plates with some skull bones from Queensland as well as poorly preserved tooth plates from Western Australia, and Tasmania. P. philippsi shares with P. serratus thick heavy skull bones, short strong jaws, and high crowned grinding tooth plates. P. philippsi differs from P. serratus in characters of the skull and in details of tooth plate shape. P. philippsi was also considerably smaller than the massive P. serratus, and reached a maximum of only 60 cm in length.

Werner SCHWARZHANS: Otoliths from the Maastrichtian of Bavaria and their evolutionary significance
[pp. 417-431, 11 figs.]

Otoliths collected from the Maastrichtian of the Gerhartsreiter Graben (Upper Bavaria) revealed the presence of 44 teleost taxa representing 13 teleost orders. Systematic allocation of otoliths, however, faces certain restrictions due to the high number of extinct early specialized taxa (particularly in Beryciformes) and some very generalized morphologies found in extinct plesiomorphic taxa. Nevertheless, some 60 % of all recorded otolith based species could be attributed to living genera or families. The interpretation of some taxa as "missing links" thought to be close to basal dichotomies of "modern" teleosts indicates the importance of Late Cretaceous fish faunas for evolutionary reconstructions. A preliminary comparison with otoliths from the Paleocene of the same area reveals a relatively sharp faunal change at the Cretaceous/Tertiary boundary. Only a fourth of the Maastrichtian taxa continue into Paleocene times. Most of the extinct early specialized taxa have disappeared in the Paleocene and the Perciformes exhibit a dramatic increase of (mostly primitive) taxa.

Dirk NOLF & Gary L. STRINGER: Cretaceous fish otoliths – a synthesis of the North American record
[pp. 433-459, 6 pls., 1 fig., 2 tabs.]

Otolith studies revealed the presence of 43 teleost taxa in the North American Late Cretaceous. The stratigraphic and geographic distribution of the concerned taxa are summarized, and 10 new taxa are described: "genus Osteoglossidarum" tavernei, Albula campaniana, "genus Albulidarum" ripleyensis, "genus Ophidiidarum" cavatus, "genus Trachichthyidarum" oscitans, "genus Caproideorum" dockeryi, "genus Apogonidarum" maastrichtiensis, "genus Apogonidarum" zideki, "genus Pempheridarum" huddlestoni, and "genus Percoideorum" severnensis. In fact, only the Campanian and Maastrichtian records provide useful information for further analysis. All other data can be considered only as reference data, up to now isolated from their geographic and stratigraphic context. Looking to the previous fossil record of Cretaceous and Early Tertiary teleosts, two important lacunae in our data are apparent. First, there is a marked gap of at least 20 million years in the fossil record of osteichthyans between the Late Campanian and the Late Paleocene, both for the osteological and the otolith material. Secondly, no unquestionable perciform is yet recorded by Cretaceous skeletal remains. In the otolith-based fossil record of fossil fishes from North America, 40 % of the identified taxa are percoids. Comparison with unpublished data concerning a small otolith association of Campanian age from chalk facies in the Paris Basin shows that this association has no percoids, and mainly consists of strange groups that cannot be assimilated to Recent families. The Late Cretaceous greensands in North America constitute a depositional environment (near shore sandy mud bottom with abundant molluscs and a relatively slow sedimentation rate) that persisted until the present over widespread geographic areas; consequently, the fish fauna inhabiting such areas may not have been subject to rapid evolutionary change, and some aspects that we consider as "modern", e.g., the presence of percoids, can be traced far back in time. Cretaceous chalk facies, on the contrary, represents a depositional environment (continuous carbonate sedimentation with negligible clastic input, over huge epicontinental basin areas) that disappeared in the early Paleogene; consequently, many fishes associated with this environment have also disappeared.

Mee-Mann CHANG & Fan JIN: Mesozoic fish faunas of China
[pp. 461-478, 5 figs.]

The information on the geographical and geological occurrences and composition of Mesozoic fish faunas of China summarized in this paper shows successive changes of major groups of actinopterygians through Mesozoic from faunas dominated by comparatively primitive actinopterygians in Triassic and Jurassic to faunas dominated by primitive teleosts in Late Jurassic to Early Cretaceous, then to those dominated by "euteleosts" in the "Middle Cretaceous". The Triassic and Jurassic fish faunas have certain cosmopolitan character. The Late Jurassic to Early Cretaceous fish faunas from different areas of China are of a complex nature. The northern Lycoptera fauna exhibits striking endemism; the southeastern Mesoclupea fauna is similar to those of Brazil and West Africa; and the northwestern pholidophoriform Siyuichthys fauna has a not quite certain relationship with the contemporaneous one from Victoria, Australia. The "Middle Cretaceous" fish fauna from northeastern China shows again more or less cosmopolitan feature.

Laurence BELTAN: Overview of systematics, paleobiology, and paleoecology of Triassic fishes of northwestern Madagascar
[pp. 479-500, 13 figs., 1 tab., 1 app.]

Several Triassic fossil-bearing beds from northwestern Madagascar have yielded abundant ichthyofaunas which have been studied from different points of view, e.g., systematics, paleobiology, and paleoecology. An overview of these findings is presented here. Thirty-six species of actinopterygian and sarcopterygian fishes have been recovered. The fishes lived in marine water, in a semi-arid environment, with other vertebrates, invertebrates, and plants. Chemical reactions have been instigated between these Triassic sediments, derived from the nearby crystalline massifs, and biotas embedded in those. The result is that the taphonomy is uncommon, and the aspect observed in the outcrops does not reflect the paleoecology, that is to say, the biotopes of living fishes.

Olivier RIEPPEL, René KINDLIMANN & Hugo BUCHER: A new fossil fish fauna from the Middle Triassic (Anisian) of North-Western Nevada
[pp. 501-512, 5 figs.]

A new fossil fish fauna from the Middle Triassic of Nevada is presented in a preliminary overview. A bonebed of lower middle Anisian age yields a rich shark fauna which includes several species of Polyacrodus as well as teeth of Acrodus, Palaeobates and, perhaps, Palaeospinax, as well as actinopterygians (Saurichthys, Birgeria, Colobodus) and a lungfish (Ceratodus). The bonebed is a debris flow deposit in a high energy environment, and the fauna indicates coastal influence. A new taxon, Palaeospinax tregoi n. sp., is described and diagnosed.

Günter VIOHL: The paleoenvironment of the Late Jurassic fishes from the southern Franconian Alb (Bavaria, Germany)
[pp. 513-528, 15 figs.]

Upper Jurassic fishes from the Southern Franconian Alb (Bavaria, Germany) occur in plattenkalks of at least three stratigraphic units: the Torleite Formation (Late Kimmeridgian), the Solnhofen Formation, and the Mörnsheim Formation (both Early Tithonian). The plattenkalks were deposited in basins characterized by a salinity-stratification between bioherms. The fossil fish did not live in the hostile bottom waters of the basins. Their habitats were the less saline and well aerated surface waters, hardgrounds and soft bottom areas on top of ancient sponge-microbial mounds going up into the surface waters, and coral patch reefs.

Shelton Pleasants APPLEGATE: An overview of the Cretaceous fishes of the quarries near Tepexi de Rodríguez, Puebla, México
[pp. 529-538, 5 figs.]

Since 1981 workers at the Institute of Geology have been engaged in the collection and curation of an extensive fish fauna from the Tlayúa quarries. To date we have over 4,270 fossils of which 2,899 are fishes. The Halecostomi incertae sedis and Halecomorphi are known by 520 specimens which include: at least two species of Lepidotes, three genera of pycnodontids and two amiids. There are at least seven new species of Macrosemiidae, one of Belonostomus, and one of Vinctifer. The Teleostei with 2,468 specimens consist of: ichthyodectids; several Elopomorpha; at least two genera of Clupeomorpha; a large fish similar to Bananogmius; a new genus of gonorynchid; and a long-snouted teleost similar to the one described from the Cenomanian of Israel. At the present it is estimated that the ichthyofauna includes 45 new taxa. This assemblage is believed to be derived for the most part, from a nearby backreef lagoon, though there is a possibility of coral reef inhabitants, freshwater and open ocean, as well as autochthonous elements in the paleobiota. Since its discovery the number of specimens and new species has steadily increased.

Luis ESPINOSA-ARRUBARRENA & Shelton Pleasants APPLEGATE: A paleoecological model of the vertebrate bearing beds in the Tlayúa Quarries, near Tepexi de Rodríguez, Puebla, México
[pp. 539-550, 6 figs.]

The Tlayúa Formation of middle or upper Albian age is best exposed in the Tlayúa quarries near Tepexi de Rodríguez, Puebla, México. The formation is composed of three members: the upper, a gray dolomite, the middle, a honey-colored "lithographic" limestone with hematitic layers forming numerous and distinct "varves", and the lower, a gray bioturbated limestone. It is hypothesized that these beds have been formed in a backlagoon behind a barrier in front of which, a bio-rich lagoon existed and was bordered to the east by a large barrier coral reef. West of the Tlayúa backlagoon existed a low lying peneplain made up of eroded clays. This land surface was semiarid with times of seasonal rains or storms. In the rainy season, a slow-moving river or rivers emptied into the backlagoon. Water depth in the Tlayúa lagoon varied from 1.5 meters (or less) to perhaps as deep as 10 meters. The depositional features of an interesting small crocodile indicate that at one time there was a depth of 1.5 meters. Algal mats were abundant, particularly in the upper beds. The preservation could have been aided by these mats, as well as, by anaerobic and or hypersaline conditions. The excellent and numerous fossils in the Tlayúa beds offer an opportunity to study a unique paleoenvironment in conjunction with present day back bays of a similar nature.

David G. SENN: Environments and functional anatomy of certain Mesozoic fishes
[pp. 551-554.]

Body shape and arrangement of fins refer to manners of swimming in both living and fossil fishes. In addition, mobility is also reflected in the structure of the scales. The Mesozoic "holosteans" show a rich radiation; they are functionally comparable with some living marine teleosts. Before fossilization, the fishes of the Solnhofen-limestone were transported from different other environments into the area where they were conserved. Thus, fishes characteristic of various environments are represented in the fossil record; some fusiform large animals were fast swimmers and pelagic predators; fishes with a fin pattern arranged in order to perform slow and precise maneuvering movements lived in an environment of hard structures with niches such as reefs and rocky areas; some fishes related to sedimentary bottoms.

Toni BÜRGIN: Diversity in the feeding apparatus of perleidid fishes (Actinopterygii) from the Middle Triassic of Monte San Giorgio (Switzerland)
[pp. 555-565, 8 figs.]

The feeding apparatuses of the five known genera of Middle Triassic perleidid fishes from Monte San Giorgio (Switzerland) are described. The family Perleididae is a species rich taxon of marine Triassic ray-finned fishes. The genera recorded from the Middle Triassic of Monte San Giorgio include Meridensia, Aetheodontus, Colobodus, Peltoperleidus, and Ctenognathichthys.Their feeding apparatus is discussed and compared with that of the genus Platysiagum, their most probable plesiomorphic sister group. The five perleidid representatives show a broad diversity in form and function of this apparatus. The diversity ranges from taxa with a weakly developed oral dentition to such with a grasping and crushing type of dentition. Dentititon, shape of the body and size and position of the fins indicate the presumable life-styles of the taxa.

The Mesozoic era was an important time in the evolution of elasmobranch and actinopterygian fishes because it was then that most of the modern groups first entered the fossil record and began to radiate. By the end of the era, many archaic forms had disappeared and the foundation had been laid for the ichthyofauna that now exists. Despite this significant evolutionary change, there has been little concerted research done on Mesozoic fishes and no synopsis or compilation of the systematics and paleoecology of Mesozoic fishes has been published, not even for single groups. To remedy this deficiency, one of us (Gloria Arratia) initiated the symposium "Mesozoic Fishes – Systematics and Paleoecology". Its goal was to bring together paleontologists and other scientists studying Mesozoic fishes so that they might evaluate current research and form an active research group to press the investigation forward.

We organized the symposium with the help of the staff of the Jura-Museum Eichstätt (Bavaria, Federal Republic of Germany). It was held in Eichstätt from August 9 to 12, 1993. The location was ideal, because the Jura-Museum is situated in the quarry district of the Solnhofen Lithographic Limestone (Upper Jurassic). The participants were able to see the sites from which a rich and "classic" ichthyofauna has come and study specimens housed in the Jura-Museum and in other museums in the area. The material was of great interest because, although many of the forms are beautifully preserved and well known, the present systematic interpretation needs to be revised.

The symposium was attended by 53 paleontologists and biologists from 17 countries. For most of them, it was the first opportunity to meet colleagues with whom they had corresponded for many years or whom they knew only from the literature and to hear them explain their work. Lively discussions followed each presentation; questions and answers flew back and forth during the poster session. The provision of a Wild FMS microscope with camera lucida attachment gave investigators a rare chance to examine material jointly and to compare their ideas about specimens directly at hand.

The work on Mesozoic fishes reported during the meeting was set in context through activities outside the conference rooms. A one-day field trip before the opening of the symposium allowed researchers to visit the quarries near Eichstätt, test the rock with their own hammers, and study collections of fossils from the Solnhofen Lithographic Limestone in museums in two adjacent towns. In the Jura-Museum itself, a special exhibit focussed on the diversity of the fishes in the Lower Tithonian Solnhofen deposits and the manner of their preservation; this exhibit included also specimens from the Late Jurassic plattenkalks of Bavaria (Schamphaupten: Late Kimmeridgian, Malm Epsilon; Daiting: Lower Tithonian, Malm Zeta 3). After the symposium, there was a field trip of six days to survey the remarkable fish localities in northern Italy. Beginning in Friuli near the Adriatic Sea, the group traveled westward to Besano and Bergamo, inspecting Triassic, Cretaceous, and Tertiary deposits long known for the fine fish fossils they contain.

The meeting generated fruitful discussions and new information that helps to clarify the course of piscine evolution at a crucial time. Phylogenetic relationships of the different groups were the central issue; but attention was given also to questions of biostratigraphy, functional anatomy, and the evolution of histological structures. The results of the symposium presented in this volumen reflect the current state of knowledge about Mesozoic fishes. The new findings described in the 36 papers and the disagreements among authors concerning the phylogenetic relationships of the fishes they have studied is an invitation to further research.