Skeletal ontogeny of the Plainfin Midshipman, Porichthys notatus (Percomorphacea: Batrachoidiformes).

Batrachoidiformes are benthic fishes that utilize the undersides of rocks as spawning nests. Their larvae are attached to the nest and nourished by a large yolk sac. The evolutionary shift from feeding, free-swimming larvae to sedentary larvae that are reliant on their yolk sac for nutrition can lead to changes in skeletal development. Batrachoidiformes also have many morphological specializations, such as five pectoral-fin radials (versus four in other acanthomorphs) that are of uncertain homology, the determination of which may have phylogenetic implications. A larval series of Porichthys notatus was collected and its skeletal ontogeny is described. In P. notatus the ossification of the pharyngeal toothplates occurs relatively later than in percomorphs with free-swimming larvae. The posterior basibranchial copula cartilage (= fourth basibranchial) in Porichthys notatus has a unique development among fishes: it initially develops as a paired element at 6.8-7.1 mm NL before fusing posteriorly and forming single median cartilage at 7.4 mm SL. Cartilages of hypobranchial four are transitory, being observed in two specimens of 6.8 and 7.3 mm NL before fusing with ceratobranchial four. The previously identified dorsalmost pectoral radial is a bone formed by a hypertrophied propterygium that ossifies later in development. The earliest stages of P. notatus have three dorsal spines, but during late larval development, the growth of the third dorsal spine is interrupted. The development of P. notatus is compared and discussed in context to that of other acanthomorph.

Larval stages of the Antarctic dragonfish Akarotaxis nudiceps (Waite, 1916), with comments on the larvae of the morphologically similar species Prionodraco evansii Regan 1914 (Notothenioidei: Bathydraconidae).

The notothenioid family Bathydraconidae is a poorly understood family of fishes endemic to the Southern Ocean. There is especially little information on Akarotaxis nudiceps, one of the deepest-dwelling and least fecund bathydraconid species. Using genetic and morphological data, we document and describe the larval stages of this unique species, offer a novel characteristic to distinguish it from the morphologically similar bathydraconid Prionodraco evansii and use the sampling locations to infer a possible spawning area of A. nudiceps along the western Antarctic Peninsula. These results provide important baseline information for locating, identifying and studying the biology of A. nudiceps, an important component of the Southern Ocean ecosystem.

Climate drives long-term change in Antarctic Silverfish along the western Antarctic Peninsula.

Over the last half of the 20th century, the western Antarctic Peninsula has been one of the most rapidly warming regions on Earth, leading to substantial reductions in regional sea ice coverage. These changes are modulated by atmospheric forcing, including the Amundsen Sea Low (ASL) pressure system. We utilized a novel 25-year (1993-2017) time series to model the effects of environmental variability on larvae of a keystone species, the Antarctic Silverfish (Pleuragramma antarctica). Antarctic Silverfish use sea ice as spawning habitat and are important prey for penguins and other predators. We show that warmer sea surface temperature and decreased sea ice are associated with reduced larval abundance. Variability in the ASL modulates both sea surface temperature and sea ice; a strong ASL is associated with reduced larvae. These findings support a narrow sea ice and temperature tolerance for adult and larval fish. Further regional warming predicted to occur during the 21st century could displace populations of Antarctic Silverfish, altering this pelagic ecosystem.

A taxonomic review of the family Trachipteridae (Acanthomorpha: Lampridiformes), with an emphasis on taxa distributed in the western Pacific Ocean.

The family Trachipteridaethe Ribbonfishes, Dealfishes, and their relativeshas a circumglobal distribution, with at least 10 species in three genera (Zu Walters Fitch 1960, Desmodema Walters Fitch 1960, and Trachipterus Goan 1770) that are characterized by elongate, extremely laterally compressed bodies, large eyes, absence of ribs, spines on lateral-line scales, greatly protrusible mouths, and a lack of pelvic fins in adults. They are also known for the drastic morphological changes that occur during ontogeny. Trachipterids are poorly represented in collections due to the fragile nature of their bodies. Most studies of the Trachipteridae have been limited by the numbers, developmental stages, and the completeness of the specimens that were examined. Along with the lack of available material, incomplete and conflicting character information compounds the taxonomic confusion of Trachipteridae. Despite the body of regional revisions that have examined trachipterid taxonomy, none have synthesized a suite of morphological characters across ontogeny. The goals of this paper are to (1) revise the family Trachipteridae, (2) revise the genera Trachipterus, Zu, and Desmodema, including information regarding ontogeny and biogeography, and 3) address the alpha taxonomy of Zu, Desmodema, and Trachipterus from the western Pacific Ocean. We recognize possibly five species of Trachipterus as being present in the western Pacific, as well as two species of both Zu and Desmodema. Despite additions to the specimen base that allows refinement of taxonomy and diagnoses, there are still large knowledge gaps associated with the taxonomic review of Trachipteridae. These reflect incomplete understanding of geographic distribution of taxa which may mask unrecognized taxonomic variability. The genus Trachipterus specifically remains problematic and will require greater detailed global study. Early life history stages remain unknown for several taxa which hinders full interpretation of ontogenetic transitions. Protracted transitions, some of which are clarified here, further confuse stage-based diagnoses and must be considered in future analyses of this family.

Sturgeons (Acipenseridae) from the Late Miocene of Ukraine, with a discussion of materials associated with Widhalms (1886) nomen nudum, †Acipenser euhuso.

The fossil record of the family Acipenseridae (sturgeons) extends to the Late Cretaceous (c. 85 MY), with a ghost lineage extending to approximately 120 MY when the first members of the family Polyodontidae are known. Much of the fossil record of Acipenseridae is formed by isolated and fragmentary dermal bones, which bear characteristic surface ornamentation. In this paper, we report on a collection of fossil sturgeons from the Upper Miocene deposits of southern Ukraine. These specimens include those used by Widhalm to establish †Acipenser euhuso, which is a nomen nudum. While we do not establish a new taxon for these specimens, the morphological variation of those elements that are preserved does suggest the presence of several species represented in this fauna.

Confronting Sources of Systematic Error to Resolve Historically Contentious Relationships: A Case Study Using Gadiform Fishes (Teleostei, Paracanthopterygii, Gadiformes).

Reliable estimation of phylogeny is central to avoid inaccuracy in downstream macroevolutionary inferences. However, limitations exist in the implementation of concatenated and summary coalescent approaches, and Bayesian and full coalescent inference methods may not yet be feasible for computation of phylogeny using complicated models and large data sets. Here, we explored methodological (e.g., optimality criteria, character sampling, model selection) and biological (e.g., heterotachy, branch length heterogeneity) sources of systematic error that can result in biased or incorrect parameter estimates when reconstructing phylogeny by using the gadiform fishes as a model clade. Gadiformes include some of the most economically important fishes in the world (e.g., Cods, Hakes, and Rattails). Despite many attempts, a robust higher-level phylogenetic framework was lacking due to limited character and taxonomic sampling, particularly from several species-poor families that have been recalcitrant to phylogenetic placement. We compiled the first phylogenomic data set, including 14,208 loci ($>$2.8 M bp) from 58 species representing all recognized gadiform families, to infer a time-calibrated phylogeny for the group. Data were generated with a gene-capture approach targeting coding DNA sequences from single-copy protein-coding genes. Species-tree and concatenated maximum-likelihood (ML) analyses resolved all family-level relationships within Gadiformes. While there were a few differences between topologies produced by the DNA and the amino acid data sets, most of the historically unresolved relationships among gadiform lineages were consistently well resolved with high support in our analyses regardless of the methodological and biological approaches used. However, at deeper levels, we observed inconsistency in branch support estimates between bootstrap and gene and site coefficient factors (gCF, sCF). Despite numerous short internodes, all relationships received unequivocal bootstrap support while gCF and sCF had very little support, reflecting hidden conflict across loci. Most of the gene-tree and species-tree discordance in our study is a result of short divergence times, and consequent lack of informative characters at deep levels, rather than incomplete lineage sorting. We use this phylogeny to establish a new higher-level classification of Gadiformes as a way of clarifying the evolutionary diversification of the order. We recognize 17 families in five suborders: Bregmacerotoidei, Gadoidei, Ranicipitoidei, Merluccioidei, and Macrouroidei (including two subclades). A time-calibrated analysis using 15 fossil taxa suggests that Gadiformes evolved $\sim $79.5 Ma in the late Cretaceous, but that most extant lineages diverged after the Cretaceous-Paleogene (K-Pg) mass extinction (66 Ma). Our results reiterate the importance of examining phylogenomic analyses for evidence of systematic error that can emerge as a result of unsuitable modeling of biological factors and/or methodological issues, even when data sets are large and yield high support for phylogenetic relationships. [Branch length heterogeneity; Codfishes; commercial fish species; Cretaceous-Paleogene (K-Pg); heterotachy; systematic error; target enrichment.].

Ontogeny and homology of cranial bones associated with lateral-line canals of the Senegal Bichir, Polypterus senegalus (Actinopterygii: Cladistii: Polypteriformes), with a discussion on the formation of lateral-line canal bones in fishes.

The association between lateral-line canals and skull bones in fishes has been the subject of several studies and raised a series of controversies, particularly with regard to the hypothesized role of lateral-line organs (i.e. neuromasts) in osteogenesis and the consequences for hypotheses of homology of the bones associated with lateral-line canals. Polypteridae, a group of freshwater fishes that occupies a key phylogenetic position as the most basal extant lineage of ray-finned fishes (Actinopterygii), provides an interesting model for the study of the relationships between lateral-line canals and skull bones. We describe the development of bones associated with lateral-line canals in the Senegal Bichir, Polypterus senegalus, and use these data to re-address previous hypotheses of homology of skull bones of polypterids. We demonstrate that the lateral-line canals constitute a separate component of the dermatocranium that may interact with a membranodermal component, thereby forming compound bones in the adult. Differences in the interactions between these components determine the characteristics of the development of each independent bone in the skull of adult P. senegalus. Our results shed light on long-standing controversies about the identity of skull bones such as the rostral, preopercle, and sphenotic in Polypteridae, and suggest the presence of an ancestral two-component pattern of formation of bones associated with lateral-line canals in bony fishes. These findings reveal the need to re-address previous hypotheses of homology of bones associated with lateral-line canals in different groups of bony fishes, especially fossil taxa.

Using multiple natural tags provides evidence for extensive larval dispersal across space and through time in summer flounder.

Dispersal sets the fundamental scales of ecological and evolutionary dynamics and has important implications for population persistence. Patterns of marine dispersal remain poorly understood, partly because dispersal may vary through time and often homogenizes allele frequencies. However, combining multiple types of natural tags can provide more precise dispersal estimates, and biological collections can help to reconstruct dispersal patterns through time. We used single nucleotide polymorphism genotypes and otolith core microchemistry from archived collections of larval summer flounder (Paralichthys dentatus, n = 411) captured between 1989 and 2012 at five locations along the US East coast to reconstruct dispersal patterns through time. Neither genotypes nor otolith microchemistry alone were sufficient to identify the source of larval fish. However, microchemistry identified clusters of larvae (n = 3-33 larvae per cluster) that originated in the same location, and genetic assignment of clusters could be made with substantially more confidence. We found that most larvae probably originated near a biogeographical break (Cape Hatteras) and that larvae were transported in both directions across this break. Larval sources did not shift north through time, despite the northward shift of adult populations in recent decades. Our novel approach demonstrates that summer flounder dispersal is widespread throughout their range, on both intra- and intergenerational timescales, and may be a particularly important process for synchronizing population dynamics and maintaining genetic diversity during an era of rapid environmental change. Broadly, our results reveal the value of archived collections and of combining multiple natural tags to understand the magnitude and directionality of dispersal in species with extensive gene flow.

Review of Marine Fishes of the Arctic Region (Mecklenburg et al. 2018).

In looking through the shelves of my library, it strikes me that there are two styles of faunal guide books within ichthyology: those that attempt to be authoritative and comprehensive (both taxonomically and biologically) and those that are more utilitarian, with a rare few that are able to pull off both formats. Both approaches have their strengths, and the results of both approaches display a wide degree of variation. The commonalities of successful examples for each approach include accessibility of information for the expert on a particular group of fishes, as well as the informed novice, the presentation of key points in clear and unambiguous fashion, and its usefulness as a contribution to future studies on the fauna, as no work will be the final word.

Cranial morphology and osteology of the sexually dimorphic electric fish, Compsaraia samueli Albert Crampton (Apteronotidae, Gymnotiformes), with comparisons to C. compsa (Mago-Leccia).

Sexual dimorphism of the snout has evolved independently in at least four separate clades of the gymnotiform family Apteronotidae. This phenomenon may help identify sex, except in the absence of mature individuals, and has led to confused taxonomy for several species. We examined a large collection of Compsaraia samueli collected during the breeding season from a remote stream in the Rio Negro drainage. This collection contains a wide range of sizes of both sexes, but most individuals were easily identified as mature. To quantify the sexual dimorphism of these specimens, 15 measurements were taken from the head and the body. In addition, some specimens were cleared-and-stained to study cranial osteology. We found that long-snouted males of C. samueli span a wide range of body sizes. As the snout length increases the distance between the eye and the occiput does not increase at the same rate, suggesting that it is only the anterior portion of the head that has an increased allometry. Skeletal anatomy differs between the sexes in that the lower jaw is more triangular in females and more linear in males. The coronomeckelian is small and round in females in contrast to being longer and pointed in males. There is strong interlacing of the dentary and anguloarticular bones in males, whereas this contact is not as extensive in females. We also discuss the implications of sexual dimorphism for identification of this species relative to its congener (C. compsa), and for the evolution of sexual dimorphism in the family.

Tooth development and replacement in the Atlantic Cutlassfish, Trichiurus lepturus, with comparisons to other Scombroidei.

Atlantic Cutlassfish, Trichiurus lepturus, have large, barbed, premaxillary and dentary fangs, and sharp dagger-shaped teeth in their oral jaws. Functional teeth firmly ankylose to the dentigerous bones. We used dry skeletons, histology, SEM, and micro-CT scanning to study 92 specimens of T. lepturus from the western North Atlantic to describe its dentition and tooth replacement. We identified three modes of intraosseous tooth replacement in T. lepturus depending on the location of the tooth in the jaw. Mode 1 relates to replacement of premaxillary fangs, in which new tooth germs enter the lingual surface of the premaxilla, develop horizontally, and rotate into position. We suggest that growth of large fangs in the premaxilla is accommodated by this horizontal development. Mode 2 occurs for dentary fangs: new tooth germs enter the labial surface of the dentary, develop vertically, and erupt into position. Mode 3 describes replacement of lateral teeth, in which new tooth germs enter a trench along the crest of the dentigerous bone, develop vertically, and erupt into position. Such distinct modes of tooth replacement in a teleostean species are unknown. We compared modes of replacement in T. lepturus to 20 species of scombroids to explore the phylogenetic distribution of these three replacement modes. Alternate tooth replacement (in which new teeth erupt between two functional teeth), ankylosis, and intraosseous tooth development are plesiomorphic to Bluefish + other Scombroidei. Our study highlights the complexity and variability of intraosseous tooth replacement. Within tooth replacement systems, key variables include sites of formation of tooth germs, points of entry of tooth germs into dentigerous bones, coupling of tooth germ migration and bone erosion, whether teeth develop horizontally or immediately beneath the tooth to be replaced, and how tooth eruption and ankylosis occur. Developmentally different tooth replacement processes can yield remarkably similar dentitions.

Systematics and biodiversity of eels (orders Anguilliformes and Saccopharyngiformes) of Taiwan (II) (Cover Copyright page).

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Systematics and biodiversity of eels (orders Anguilliformes and Saccopharyngiformes) of Taiwan (II) (Table of contents).

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Development of the muscles associated with the mandibular and hyoid arches in the Siberian sturgeon, Acipenser baerii (Acipenseriformes: Acipenseridae).

The skeleton of the jaws and neurocranium of sturgeons (Acipenseridae) are connected only through the hyoid arch. This arrangement allows considerable protrusion and retraction of the jaws and is highly specialized among ray-finned fishes (Actinopterygii). To better understand the unique morphology and the evolution of the jaw apparatus in Acipenseridae, we investigated the development of the muscles of the mandibular and hyoid arches of the Siberian sturgeon, Acipenser baerii. We used a combination of antibody staining and formalin-induced fluorescence of tissues imaged with confocal microscopy and subsequent three-dimensional reconstruction. These data were analyzed to address the identity of previously controversial and newly discovered muscle portions. Our results indicate that the anlagen of the muscles in A. baerii develop similarly to those of other actinopterygians, although they differ by not differentiating into distinct muscles. This is exemplified by the subpartitioning of the m. adductor mandibulae as well as the massive m. protractor hyomandibulae, for which we found a previously undescribed portion in each. The importance of paedomorphosis for the evolution of Acipenseriformes has been discussed before and our results indicate that the muscles of the mandibular and the hyoid may be another example for heterochronic evolution.

A review of the systematic biology of fossil and living bony-tongue fishes, Osteoglossomorpha (Actinopterygii: Teleostei)

The bony-tongue fishes, Osteoglossomorpha, have been the focus of a great deal of morphological, systematic, and evolutionary study, due in part to their basal position among extant teleostean fishes. This group includes the mooneyes (Hiodontidae), knifefishes (Notopteridae), the abu (Gymnarchidae), elephantfishes (Mormyridae), arawanas and pirarucu (Osteoglossidae), and the African butterfly fish (Pantodontidae). This morphologically heterogeneous group also has a long and diverse fossil record, including taxa from all continents and both freshwater and marine deposits. The phylogenetic relationships among most extant osteoglossomorph families are widely agreed upon. However, there is still much to discover about the systematic biology of these fishes, particularly with regard to the phylogenetic affinities of several fossil taxa, within Mormyridae, and the position of Pantodon. In this paper we review the state of knowledge for osteoglossomorph fishes. We first provide an overview of the diversity of Osteoglossomorpha, and then discuss studies of the phylogeny of Osteoglossomorpha from both morphological and molecular perspectives, as well as biogeographic analyses of the group. Finally, we offer our perspectives on future needs for research on the systematic biology of Osteoglossomorpha.(AU)

Os peixes da Superordem Osteoglossomorpha têm sido foco de inúmeros estudos sobre a morfologia, sistemática e evolução, particularmente devido à sua posição basal dentre os peixes teleósteos. Fazem parte deste grupo os "mooneyes" (Hiodontidae), "knifefishes" (Notopteridae), o "abu" (Gymnarchidae), peixes-elefante (Mormyridae), aruanãs e pirarucu (Osteoglossidae), e o peixe-borboleta africano (Pantodontidae). Esse grupo de morfologia heterogênea possui um longo e diverso registro fóssil, incluindo táxons de todos os continentes, oriundos tanto de depósitos de água doce quanto marinhos. As relações filogenéticas dentre a maioria das famílias de osteoglossomorfos é amplamente aceita. Entretanto, há muito a ser descoberto sobre a sistemática biológica desses peixes, particularmente com relação às afinidades filogenéticas de inúmeros fósseis, relações dentro de Mormyridae, e a posição filogenética de Pantodon. Neste manuscrito nós revisamos o atual estado de conhecimento dos peixes osteoglossomorfos. Nós primeiramente provemos uma abordagem geral da diversidade de Osteoglossomorpha, e então discutimos os estudos filogenéticos sobre Osteoglossomorpha sob a perspectiva morfológica e molecular, assim como uma análise biogeográfica do grupo. Finalmente, oferecemos nossas perspectivas sobre os futuros passos para pesquisa sobre a sistemática biológica de Osteoglossomorpha.(AU)

Development of the skull and pectoral girdle in Siberian sturgeon, Acipenser baerii, and Russian sturgeon, Acipenser gueldenstaedtii (Acipenseriformes: Acipenseridae).

The head is considered the major novelty of the vertebrates and directly linked to their evolutionary success. Its form and development as well as its function, for example in feeding, is of major interest for evolutionary biologists. In this study, we describe the skeletal development of the cranium and pectoral girdle in Siberian (Acipenser baerii) and Russian sturgeon (A. gueldenstaedtii), two species that are commonly farmed in aquaculture and increasingly important in developmental studies. This study comprises the development of the neuro-, viscero- and dermatocranium and the dermal and chondral components of the pectoral girdle, from first condensation of chondrocytes in prehatchlings to the early juvenile stage and reveals a clear pattern in formation. The otic capsules, the parachordal cartilages, and the trabeculae cranii are the first centers of chondrification, at 8.4mm TL. These are followed by the mandibular, then the hyoid, and later the branchial arches. Teeth form early on the dentary, dermopalatine, and palatopterygoid, and then appear later in the buccal cavity as dorsal and ventral toothplates. With ongoing chondrification in the neurocranium a capsule around the brain and a strong rostrum are formed. Dermal ossifications start to form before closure of the dorsal neurocranial fenestrae. Perichondral ossification of cartilage bones occurs much later in ontogeny. Our results contribute data bearing on the homology of elements such as the lateral rostral canal bone that we regard homologous to the antorbital of other actinopterygians based on its sequence of formation, position and form. We further raise doubts on the homology of the posterior ceratobranchial among Actinopteri based on the formation of the hyoid arch elements. We also investigate the basibranchials and the closely associated unidentified gill-arch elements and show that they are not homologous. J. Morphol. 278:418-442, 2017. © 2017 Wiley Periodicals, Inc.

Influence of cladogenesis on feeding structures in drums (Teleostei: Sciaenidae).

Drums (family Sciaenidae) are common in tropical to temperate coastal and estuarine habitats worldwide and present a broad spectrum of morphological diversity. The anatomical variation in this family is particularly evident in their feeding apparatus, which may reflect the partitioning of adult foraging habitats. Adult and early life history stage sciaenids may display ecomorphological patterns in oral and pharyngeal jaw elements but because sciaenids are hierarchically related, the morphological variation of the feeding apparatus cannot be analyzed as independent data. Morphological patterns have been identified in three sciaenid genera from the Chesapeake Bay but it is not known if these patterns are present in other genera of the family and if such patterns are constrained by phylogenetic history. In this study, phylogenetic comparative methods were applied to two sets of oral jaw data obtained from growth series of 11 species of cleared and double-stained Chesapeake Bay sciaenids and alcohol-preserved museum specimens representing 65 of the 66 recognized genera to determine the magnitude of phylogenetic dependence present in the structure of the oral jaws using a recent molecular phylogeny of the family. Pagel's lambda, a measure of phylogenetic signal, was low for pelagic sciaenids in premaxilla, lower jaw, and ascending process lengths, indicating influence of selective forces on the condition of these traits. Conversely, for benthic sciaenids, phylogenetic signal was high for lower jaw and ascending process lengths, indicating significant phylogenetic constraint for their condition in these taxa. Pagel's lambda was intermediate for premaxilla length in benthic sciaenids, suggesting that the length of the premaxilla is influenced by a mix of selective forces and phylogenetic constraint. Although the ecomorphological patterns identified in the oral jaws of scaienids are not entirely free of phylogenetic dependence, selective forces related to foraging are likely driving the evolution of these structures.

Comparative ontogeny of the feeding apparatus of sympatric drums (Perciformes: Sciaenidae) in the Chesapeake Bay.

The anatomy of the feeding apparatus in fishes, including both oral and pharyngeal jaw elements, is closely related to the ecology of a species. During ontogeny, the oral and pharyngeal jaws undergo dramatic changes. To better understand how such ontogenetic changes occur and relate to the feeding ecology of a species, ontogenetic series of four closely related members of the family Sciaenidae (Cynoscion nebulosus, Cynoscion regalis, Micropogonias undulatus, and Leiostomus xanthurus) were examined. Sciaenids were selected because as adults they exhibit considerable specialization of the feeding apparatus correlated with differences in foraging habitats. However, it is not clear when during ontogeny the structural specializations of the feeding apparatus develop, and thereby enable early life history stage (ELHS) sciaenids to partition their foraging habitats. A regression tree was recovered from the analysis and three divergences were identified during ontogeny. There are no measurable differences in elements of the feeding apparatus until the first divergence at 8.4 mm head length (HL), which was attributed to differences in average gill filament length on the second ceratobranchial. The second divergence occurred at 14.1 mm HL and was associated with premaxilla length. The final divergence occurred at 19.8 mm HL and was associated with differences in the toothed area of the fifth certatobranchial. These morphological divergences suggest that ELHS sciaenids may be structurally able to partition their foraging habitats as early as 8.4 mm HL.