Fossil‐based comparative analyses reveal ancient marine ancestry erased by extinction in ray‐finned fishes

The marine‐freshwater boundary is a major biodiversity gradient and few groups have colonised both systems successfully. Fishes have transitioned between habitats repeatedly, diversifying in rivers, lakes and oceans over evolutionary time. However, their history of habitat colonisation and diversification is unclear based on available fossil and phylogenetic data. We estimate ancestral habitats and diversification and transition rates using a large‐scale phylogeny of extant fish taxa and one containing a massive number of extinct species. Extant‐only phylogenetic analyses indicate freshwater ancestry, but inclusion of fossils reveal strong evidence of marine ancestry in lineages now restricted to freshwaters. Diversification and colonisation dynamics vary asymmetrically between habitats, as marine lineages colonise and flourish in rivers more frequently than the reverse. Our study highlights the importance of including fossils in comparative analyses, showing that freshwaters have played a role as refuges for ancient fish lineages, a signal erased by extinction in extant‐only phylogenies.     

The early evolution of ray‐finned fishes

Ray‐finned fishes (Actinopterygii) constitute approximately half of all living vertebrate species. A stable hypothesis of relationships among major modern lineages has emerged over the past decade, supported by both anatomy and molecules. Diversity is unevenly partitioned across the actinopterygian tree, with most species concentrated within a handful of geologically young (i.e. Cretaceous) teleost clades. Extant non‐teleost groups are portrayed as ‘living fossils’, but this moniker should not be taken as evidence of especially primitive structure: each of these lineages is characterized by profound specializations. Attribution of fossils to the crowns and apical stems of Cladistia, Chondrostei and Neopterygii is uncontroversial, but placements of Palaeozoic taxa along deeper branches of actinopterygian phylogeny are less secure. Despite these limitations, some major outlines of actinopterygian diversification seem reasonably clear from the fossil record: low richness and disparity in the Devonian; elevated morphological variety, linked to increases in taxonomic dominance, in the early Carboniferous; and further gains in taxonomic dominance in the Early Triassic associated with earliest appearance of trophically diverse crown neopterygians.     

The phylogenetic position of toadfishes (order Batrachoidiformes) in the higher ray-finned fish as inferred from partitioned Bayesian analysis of 102 whole mitochondrial genome sequences

In a previous study based on 100 whole mitochondrial genome (mitogenome) sequences, we sought to provide a new perspective on the ordinal relationships of higher ray-finned fish (Actinopterygii). The study left unexplored the phylogenetic position of toadfishes (order Batrachoidiformes), as data were unavailable owing to technical difficulties. In the present study, we successfully determined mitogenomic sequences for two toadfish species ( Batrachomoeus trispinosus and Porichthys myriaster ) and found that the difficulties resulted from unusual gene arrangements and associated repetitive non-coding sequences. Unambiguously aligned, concatenated mitogenomic sequences (13 461 bp) from 102 higher actinopterygians (excluding the ND6 gene and control region) were divided into five partitions (1st, 2nd and 3rd codon positions of the protein-coding genes, tRNA genes and rRNA genes) and partitioned Bayesian analyses were conducted. The resultant phylogenies strongly suggest that the toadfishes are not members of relatively primitive higher actinopterygians (Paracanthopterygii), but belong to a crown group of actinopterygians (Percomorpha), as was demonstrated for ophidiiform eels (Ophidiiformes) and anglerfishes (Lophiiformes) in the previous study. We propose revised limits of major unranked categories for higher actinopterygians and a new name (Berycomorpha) for a clade comprising two reciprocally paraphyletic orders (Beryciformes and Stephanoberyciformes) based on the present mitogenomic phylogenies.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 289–306.     

Colonization of the deep sea by fishes

Analysis of maximum depth of occurrence of 11 952 marine fish species shows a global decrease in species number (N) with depth (x; m): log₁₀N = ?0·000422x + 3·610000 (r² = 0·948). The rate of decrease is close to global estimates for change in pelagic and benthic biomass with depth (?0·000430), indicating that species richness of fishes may be limited by food energy availability in the deep sea. The slopes for the Classes Myxini (?0·000488) and Actinopterygii (?0·000413) follow this trend but Chondrichthyes decrease more rapidly (?0·000731) implying deficiency in ability to colonize the deep sea. Maximum depths attained are 2743, 4156 and 8370 m for Myxini, Chondrichthyes and Actinopterygii, respectively. Endemic species occur in abundance at 7–7800 m depth in hadal trenches but appear to be absent from the deepest parts of the oceans, >9000 m deep. There have been six global oceanic anoxic events (OAE) since the origin of the major fish taxa in the Devonian c. 400 million years ago (mya ). Colonization of the deep sea has taken place largely since the most recent OAE in the Cretaceous 94 mya when the Atlantic Ocean opened up. Patterns of global oceanic circulation oxygenating the deep ocean basins became established coinciding with a period of teleost diversification and appearance of the Acanthopterygii. Within the Actinopterygii, there is a trend for greater invasion of the deep sea by the lower taxa in accordance with the Andriashev paradigm. Here, 31 deep‐sea families of Actinopterygii were identified with mean maximum depth >1000 m and with >10 species. Those with most of their constituent species living shallower than 1000 m are proposed as invasive, with extinctions in the deep being continuously balanced by export of species from shallow seas. Specialized families with most species deeper than 1000 m are termed deep‐sea endemics in this study; these appear to persist in the deep by virtue of global distribution enabling recovery from regional extinctions. Deep‐sea invasive families such as Ophidiidae and Liparidae make the greatest contribution to fish fauna at depths >6000 m.     

Reproduction in Middle Triassic actinopterygians; complex fin structures and evidence of viviparity in fossil fishes

New well-preserved specimens of Peltopleurus lissocephalus Brough, 1939, a small actinopterygian fish of the Alpine Middle Triassic, are described. Some of them show a highly modified anal fin, with a brush-shaped holdfast device and a gonopodium-like structure, possibly used during mating behaviour. Additional examples of modified anal fins are reported for the genera Cephaloxenus and Habroichthys. Evidence for viviparity within the genus Saurichthys , found at the same localities, is discussed and compared with earlier reports of the phenomenon. A comparison is made of the different reproductive strategies used in some Middle Triassic actinopterygian fishes.     

A NEW SEMIONOTIFORM (ACTINOPTERYGII, NEOPTERYGII) FROM UPPER JURASSIC – LOWER CRETACEOUS DEPOSITS OF NORTH-EAST THAILAND, WITH COMMENTS ON THE RELATIONSHIPS OF SEMIONOTIFORMS

Abstract:  A new semionotiform fish, Isanichthys palustris gen. et sp. nov., is described from the Late Jurassic – Early Cretaceous Phu Kradung Formation, north-east Thailand. I. palustris is known from a single, nearly complete specimen found alongside abundant Lepidotes specimens at the Phu Nam Jun locality. I. palustris shows a mixture of semionotid-like characters, such as the pattern of cheek ossifications, and lepisosteid-like characters, such as the body shape and a dorsal fin opposed by an anal fin. I. palustris possesses only some of the characters currently used to define the Semionotidae. Cladistic analyses including various semionotid and gar taxa, together with Amia calva and Leptolepis coryphaenoides , suggest that the Semionotiformes (Lepisosteidae and 'Semionotidae') form a monophyletic clade, but the 'Semionotidae' taxa form an unresolved polytomy. The relationships between Semionotiformes, Halecomorphi and Teleostei are unresolved. When restricted to the best-known taxa, however, the analysis shows the monophyly of the Semionotidae sensu stricto ( Semionotus + Lepidotes ) and a sister-group relationship between halecomorphs and teleosts. These last two results are regarded as the preferred hypothesis for further studies. I. palustris is the only known example of a predaceous, probably piscivorous, 'semionotid'. It illustrates the great diversity and ecological adaptation of the semionotiforms during the Late Jurassic – Early Cretaceous. We question the phylogenetic relationships of 'ancient fishes' founded on molecular-based trees because we suspect that the use of very few Recent taxa as representatives of previously diverse lineages is an inevitable, but important, bias in the construction of such trees.     

Osteology and phylogenetic interrelationships of sturgeons (Acipenseridae)

Sturgeons (Acipenseridae) are an ancient and unique assemblage of fishes historically important to discussions of actinopterygian evolution. Despite their basal position within Actinopterygii, rigorous comparative morphological studies of acipenserids have never been made, and most ideas about acipenserid evolution hinge on an untested impression that shovelnose sturgeons (Scaphirhynchini) are phylogenetically primitive. This impression promoted ideas that: (1) the earliest acipenserids were highly benthic and evolved secondarily into pelagic predators, and (2) paedomorphosis has dominated mechanisms affecting their morphological change. Using cladistic methods, this study examines generic level interrelationships within Acipenseridae. Representatives of the four acipenserid genera Huso, Acipenser, Pseudoscaphirhynchus, and Scaphirhynchus, as well as their acipenseriform outgroups Polyodontidae, Peipiaosteidae, and Chondrosteidae, were surveyed for skeletal characters. Sixty-nine characters are identified and described to support the first generic level cladogram of Acipenseridae. Huso is phylogenetically primitive within Acipenseridae and the sister group to a redefined subfamily Acipenserinae. Acipenser is not supported by any characters identified in this study, but the tribe Scaphirhynchini comprising Scaphirhynchus and Pseudoscaphirhynchus is found to be monophyletic. The cladogram contradicts historical ideas about acipenserid evolution because Huso defines an outgroup morphology and life history founded on pelagic habitats and piscivory. In contrast, acipenserines, and more markedly scaphirhynchines, are benthic predators possessing character complexes for benthic feeding, respiration, locomotion, and protection. Also, the pattern of character acquisition within Acipenseridae suggests that peramorphosis played a central role in acipenserid evolution. Peramorphic addition and enlargement of the skeleton and scalation defines most characters at all nodes within Acipenseridae, and repudiates paedomorphosis as a major trend in evolution within the family Acipenseridae.     

Immunocytochemical demonstration of somatotropin-like and prolactin-like activity in the brain of Calamoichthys calabaricus (Actinopterygii)

Summary Cellular binding of anti-bSTH and anti-oPRL IgG is demonstrated in the brain and the pituitary gland of the African freshwater fish Calamoichthys calabaricus by means of the unlabeled antibody enzyme method at the light microscopic level. In the brain, somatotropin and prolactin are demonstrated in separate neurons in the preoptic area. The somatotropinergic and prolactinergic perikarya are distinct from those of the hypothalamic-hypophysial neurosecretory neurons, i.e., those stainable with aldehyde fuchsin presumed to be vasotocinergic and isotocinergic. The somatotropinergic and prolactinergic neuronal perikarya give rise to separate beaded axons which pass either ventroposteriorly into the infundibulum, terminating in the neurohypophysis, or ventro-laterally through the wall of the preoptic recess, terminating near the superficial capillary bed covering this part of the brain surface. Moreover, coarse dendrite-like processes of both kinds of immuno-reactive neurons extend towards, and end in, the third ventricle. Binding sites in the brain to antisera against hLH, hFSH, hTSH and anti-(1–24) ACTH IgG, all reactive in the pituitary, are not observed in the neurons confined to the preoptic area.Supported by the Danish Natural Sciences Research CouncilThe authors wish to thank Professor Dr K.G. Wingstrand, University of Copenhagen, Denmark, for placing two series of C. calabaricus at their disposal. They would also like to thank the National Institute of Arthritis, Metabolism and Digestive Diseases, Bethesda, USA, for the generous gift of antisera against the subunits of human LH, TSH and FSH, and likewise Dr L. Hummer, Glostrup Hospital, Copenhagen, Denmark, for the gift of the anti-(1-24)ACTH IgG     

SYSTEMATIC REVISION OF PSEUDOBEACONIA BORDAS, 1944, AND MENDOCINICHTHYS WHITLEY, 1953 (ACTINOPTERYGII: ‘PERLEIDIFORMES’) FROM THE TRIASSIC OF ARGENTINA

Abstract: The ‘perleidiform’Mendocinichthys and Pseudobeaconia from the Potrerillos and Santa Clara Abajo formations (Upper Triassic; Argentina) are reviewed. Mendocinichthys has been known from a review of this species that is not based on the type material, but on referred specimens in the American Museum of Natural History. However, those specimens are found here to represent a new species of Pseudobeaconia, P. celestae sp. nov. Consequently, Mendocinichthys is restricted here to the type material and, within it, to the only specimen that clearly represents a distinct taxon and is thus designated lectotype. We further performed a cladistic analysis and propose a new family, Pseudobeaconiidae for these two South American Triassic genera, which is mainly characterized by the presence of an incomplete dorsal ridge of spine‐like scales between the skull and the dorsal fin, and scales with straight posterior border, an elevated central region and marginal concentric ridges of ganoine. The cladistic analysis further indicates the existence of some lineages endemic to certain areas of Gondwana and Europe. Pseudobeaconia celestae sp. nov. represents the first record of Pseudobeaconia in the Cacheuta sub‐basin of the Cuyana Basin. The genus was previously known from the Santa Clara sub‐basin of the Cuyana Basin only, and the new record confirms the previous hypothesis of correlation between the sedimentary infilling of these sub‐basins.