Comparison of evolutionary rates in the mitochondrial DNA cytochrome b gene and control region and their implications for phylogeny of the Cobitoidea (Teleostei: Cypriniformes)

It is widely accepted that mitochondrial DNA (mtDNA) control region evolves faster than protein encoding genes with few exceptions. In the present study, we sequenced the mitochondrial cytochrome b gene (cyt b) and control region (CR) and compared their rates in 93 specimens representing 67 species of loaches and some related taxa in the Cobitoidea (Order Cypriniformes). The results showed that sequence divergences of the CR were broadly higher than those of the cyt b (about 1.83 times). However, in considering only closely related species, CR sequence evolution was slower than that of cyt b gene (ratio of CR/cyt b is 0.78), a pattern that is found to be very common in Cypriniformes. Combined data of the cyt b and CR were used to estimate the phylogenetic relationship of the Cobitoidea by maximum parsimony, neighbor-joining, and Bayesian methods. With Cyprinus carpio and Danio rerio as outgroups, three analyses identified the same four lineages representing four subfamilies of loaches, with Botiinae on the basal-most clade. The phylogenetic relationship of the Cobitoidea was ((Catostomidae+Gyrinocheilidae)+(Botiinae+(Balitorinae+(Cobitinae+Nemacheilinae)))), which indicated that Sawada's Cobitidae (including Cobitinae and Botiinae) was not monophyletic. Our molecular phylogenetic analyses are in very close agreement with the phylogenetic results based on the morphological data proposed by Nalbant and Bianco, wherein these four subfamilies were elevated to the family level as Botiidae, Balitoridae, Cobitidae, and Nemacheilidae.     

Ontogeny of the Early Carboniferous acanthodian <Emphasis Type="Italic">Acanthodes lopatini</Emphasis> Rohon

The sequence of ossification of skeletal elements in the growth series of the acanthodian Acanthodes lopatini Rohon from the Lower Tournaisian of the Minusa Trough (Siberia) is described. Four formal ontogenetic stages are recognized. It is shown that even 20?30-mm-long juveniles have well-developed fin spines, scapulocoracoids, mandibular bones, branchiostegal rays, and incipient gill rakers. At this stage, scales cover the posterior part of the body and the orbits are marked by dark “eye stains” of uncertain nature. Further ontogenetic changes are connected with the development of sclerotic ring elements, sensory line system, squamation of the head region and fin webs, perichondral ossifications of the Meckel’s, quadrate, and palatine cartilages, and, finally, mineralization of the hyoid and gill arches. Ontogenetic features of A. lopatini are compared with those in other members of the genus and other acanthodiforms. Some developmental features shared by A. lopatini and Lodeacanthus gaujicus Upeniece suggest that acanthodids could have evolved from mesacanthids.     

Morphological variation in the Weberian apparatus of Cypriniformes

Cypriniformes (which includes the minnows, carps, loaches, algae-eaters, stone loaches, and suckers) is a morphologically diverse and incredibly speciose order of teleosts. It has been suggested that a number of evolutionary innovations, key to improved hearing and feeding, have played an important role in cypriniform fishes' success. One such innovation, the Weberian apparatus, is a novel assemblage of vertebral elements and modified ribs that relay and amplify sound pressure changes from the gas bladder to the inner ear. The Weberian apparatus unites Cypriniformes with other major orders into an extremely species-rich group of fishes, the Otophysi. Together, otophysan fishes comprise one of the largest groups of fishes in the world, as well as the majority of freshwater fishes. Here we present a detailed comparison of the Weberian apparatus in a number of cypriniform families using cleared and stained specimens. We present data regarding inter- and intrafamilial morphological variation within Cypriniformes. With few, but evolutionarily important, exceptions we find that diagnostic features of the Weberian apparatus characterize each family. Interspecific variation within each of the families Balitoridae, Gyrinocheilidae, and Catostomidae is only slight, whereas variation among subfamilies within Cyprinidae and Cobitidae is far more significant. This comparative study identifies a number of distinct morphologies, some of which appear highly correlated with ecological niche. For example, inhabiting swift-moving waters appears to be a key factor in the encapsulation of the anterior gas bladder in some cobitids, balitorids, and gobionin cyprinids.     

Histology and structural integration of the major morphologies of the Cypriniform Weberian apparatus

The Weberian apparatus, a diagnostic feature of otophysan fishes, is a novel hearing adaptation integrating several developmental and morphological systems (ear-vertebral column-swim bladder). Otophysan fishes are one of the largest and most successful freshwater clades, with over 10,000 species across most continents. The largest otophysan order, Cypriniformes, dominates the freshwaters of Asia, Europe, North America, and Africa. Spanning such a wide variety of environments, the Weberian apparatus undergoes morphological modifications to maintain functionality. Within Cypriniformes, we propose three distinct morphological classes of the Weberian apparatus based on the level of skeletal expansion around the swim bladder: simple (typical of most Cyprinidae), anterior plate (found in families such as Gyrinocheilidae, Catostomidae, and Botiidae), and encapsulated (either single-capsule as found, e.g., in Gobionidae and Cobitidae, or double-capsule as found, e.g., in Nemacheilidae and Balitoridae). Little ontological or comparative data exists regarding the construction or integration of these different morphologies, and less is known about the tissue level integration and variation within these morphologies. We used paraffin histology to document the hard and soft tissue anatomy of the Weberian apparatus in six species representing all morphological classes. We found sites of similarity across the morphologies including size and structure of the saccule, aspects of ossicle ossification, and swim bladder tunica composition, indicating potential sites of developmental and functional constraint. In contrast, we found differences across both auditory and nonauditory features in otic chamber size, ossification within ossicles and other vertebral elements, and composition of ligaments, indicating likely sites of adaptability. Some of these changes are likely evolutionary (taxonomic), but may be influenced by the environmental niche occupied by the clade. These results show a clear need for increased ontological and comparative study of the complete cypriniform Weberian apparatus, particularly histologically, as well as increased auditory studies across morphological types.     

Host specificity among Unionicola spp. (Acari: Unionicolidae) parasitizing freshwater mussels

Water mites of Unionicola spp. are common parasites of freshwater mussels as adults, living on the gills, or mantle and foot of their hosts and using these tissues as sites of oviposition. The present study addresses specialization among North American Unionicola mussel-mites using 2 measures of host specificity: (1) the number of host species used by a species of mite; and (2) a measure that considers the taxonomic distinctness of the hosts utilized by mites, weighted for their prevalence in the different hosts. Results of this study indicate the Unionicola spp. mussel-mites are highly host specific, with most species occurring in association with 1 or 2 species of hosts. If 2 or more host species are utilized, they are typically members of the same genus. These data are consistent with studies examining the dispersal abilities and host recognition behavior for members of the group. When the average values of host specificity for Unionicola subgenera were mapped on a phylogenetic tree for these taxa, a clade comprised of gill mites appeared to be more host specific than a clade consisting of mantle mites. There were, however, no apparent patterns of host specificity within each of the clades. Differences in specificity between the 2 lineages may reflect either a long evolutionary history that gill mites have had with host mussels or the intense competition among gill mites for oviposition sites within unionid mussels, leading to increased host specialization.     

肠呼吸抑制胁迫对大鳞副泥鳅呼吸代谢和抗氧化能力的影响

为研究肠呼吸抑制胁迫对气呼吸鱼类大鳞副泥鳅(Paramisgurnus dabryanus)的鳃和肠道呼吸代谢及抗氧化能力的影响,初步探究其生理反馈调节机制,本文选取大鳞副泥鳅成熟个体(n=60)进行肠呼吸抑制胁迫实验。分别对实验组(肠呼吸抑制,n=30)与空白对照组(n=30)的大鳞副泥鳅饲养驯化2周,测定其整体静止代谢率、呼吸频率、鳃和肠道各段的乳酸脱氢酶(LDH)、琥珀酸脱氢酶(SDH)、Na+/K+ATP酶(NKA)、过氧化氢酶(CAT)以及超氧化物歧化酶(SOD)活性。肠呼吸抑制胁迫下,实验组与对照组大鳞副泥鳅整体静止代谢率无显著差异(P0.05),而实验组大鳞副泥鳅鳃部呼吸频率显著加快(P0.05)。与对照组相比,实验组大鳞副泥鳅鳃部的琥珀酸脱氢酶活性显著升高(P0.05),而后肠琥珀酸脱氢酶和Na+/K+ATP酶显著降低(P0.05)。同时实验组大鳞副泥鳅后肠的乳酸脱氢酶活性显著升高(P0.05)。实验组和对照组之间,大鳞副泥鳅鳃和前中肠的过氧化氢酶以及超氧化物歧化酶酶活性无显著差异(P0.05),后肠则有显著性升高(P0.05)。当大鳞副泥鳅肠呼吸受到抑制时,会加强其鳃部有氧呼吸代谢,弥补肠呼吸缺失部分,满足机体生理需求,而气呼吸的后肠会有一定氧化应激反应。     

Homology of the palpebral and origin of supraorbital ossifications in ornithischian dinosaurs

Maidment, S.C.R. & Porro, L.B. 2010: Homology of the palpebral and origin of supraorbital ossifications in ornithischian dinosaurs. Lethaia , Vol. 43, pp. 95–111.
The palpebral is a small ossification that projects across the orbit in some ornithischian dinosaurs and its presence is considered a synapomorphy of the clade. By contrast, other ornithischians lack the palpebral but possess accessory ossifications, commonly termed supraorbitals, which form the dorsal margin of the orbit. The homology of the ornithischian palpebral to one or more of the supraorbitals is widely accepted in the literature, but this homology has never been explicitly tested and no hypotheses have been proposed regarding the function of the palpebral or why it was incorporated into the orbital margin. As homology is synonymous with synapomorphy, incorrect homology statements can lead to incorrect hypotheses of relationships being obtained during cladistic analysis. The primary and secondary homologies of the ornithischian palpebral and the anterior supraorbital of more derived members of the major ornithischian clades are tested and we demonstrate that these homology hypotheses can be accepted. Osteological correlates indicate that the palpebral supported a layer of connective tissue that roofed the orbit; ossification of this connective tissue resulted in the incorporation of the palpebral into the skull roof and gave rise to additional supraorbital elements, which are neomorphic ossifications. Large-scale structural changes in the ornithischian skull, including dermal ossifications associated with display or defence and the development of complex feeding mechanisms, may have led to the incorporation of the palpebral into the skull roof. □ Functional morphology , homology , Ornithischia , palpebral , supraorbital.     

Mitogenomic Evolution and Interrelationships of the Cypriniformes (Actinopterygii: Ostariophysi): The First Evidence Toward Resolution of Higher-Level Relationships of the World’s Largest Freshwater Fish Clade Based on 59 Whole Mitogenome Sequences

Fishes of the order Cypriniformes are almost completely restricted to freshwater bodies and number > 3400 species placed in 5 families, each with poorly defined subfamilies and/or tribes. The present study represents the first attempt toward resolution of the higher-level relationships of the world’s largest freshwater-fish clade based on whole mitochondrial (mt) genome sequences from 53 cypriniforms (including 46 newly determined sequences) plus 6 outgroups. Unambiguously aligned, concatenated mt genome sequences (14,563 bp) were divided into 5 partitions (first, second, and third codon positions of the protein-coding genes, rRNA genes, and tRNA genes), and partitioned Bayesian analyses were conducted, with protein-coding genes being treated in 3 different manners (all positions included; third codon positions converted into purine [R] and pyrimidine [Y] [RY-coding]; third codon positions excluded). The resultant phylogenies strongly supported monophyly of the Cypriniformes as well as that of the families Cyprinidae, Catostomidae, and a clade comprising Balitoridae + Cobitidae, with the 2 latter loach families being reciprocally paraphyletic. Although all of the data sets yielded nearly identical tree topologies with regard to the shallower relationships, deeper relationships among the 4 major clades (the above 3 major clades plus Gyrinocheilidae, represented by a single species Gyrinocheilus aymonieri in this study), were incongruent depending on the data sets. Treatment of the rapidly saturated third codon–position transitions appeared to be a source of such incongruities, and we advocate that RY-coding, which takes only transversions into account, effectively removes this likely “noise” from the data set and avoids the apparent lack of signal by retaining all available positions in the data set. [Reviewing Editor: Rafael Zardoya]     

A new gyrodactylid species from Cobitis granoei (Rendahl) (Cobitidae) in central China

Gyrodactylus granoei n. sp. (Monogenea: Gyrodactylidae) is described from the fins and body surface of the spine loach, Cobitis granoei (Rendahl) (Cobitidae), in central China. It resembles a suite of species from cyprinid and cobitid fishes that have short, compact hamuli, a ventral bar with no obvious anterolateral projections, a linguiform posterior membrane, a male copulatory organ with small hooks in multiple rows, a simple cylindrical dorsal bar, and short marginal hooks, with an expanded sickle heel. Of these, the new species resembles most closely Gyrodactylus micracanthus Hukuda, 1940 from Misgurnus anguillicaudatus (Cantor) (Cobitidae), but it is identified by the length of the hamuli and the morphology of dorsal and ventral bars. When searched using BLAST, sequence data (829 bp) spanning the ITS1, 5.8s, and ITS2 region did not return an identical match; close similarity (82-88%) was found with sequenced members of the subgenus Gyrodactylus. It is suggested that the new species is part of a freshwater lineage that has radiated successfully among cyprinid fishes in North America, Europe, and Asia, and some of their predator and amphibian neighbors.     

中国鲤形目鱼类的中轴骨骼数及其生态适应性

为探讨中国鲤形目鱼类中轴骨骼数及其与系统发育和生态习性的相关性,采用X光透视照相法对157种鲤科(Cyprinidae)、鳅科(Cobitidae)、平鳍鳅科(Balitoridae)鱼类的脊椎骨数、肋骨数、尾椎数进行了比较分析。结果显示,中国鲤形目鱼类的脊椎骨数在30-52枚之间,均值39.45?4.44;肋骨数在8-28对之间,均值15.27?3.08;尾椎数在14-34枚之间,均值21.08?2.89。依据脊椎骨数、肋骨数、尾椎数对鲤科各亚科进行聚类,显示鲤科12个亚科分为2大类：雅罗鱼类,包括雅罗鱼亚科、鲌亚科、鲴亚科、裂腹鱼亚科等4亚科;鲃类,包括鲢亚科、鮈亚科、鱼丹亚科、鲃亚科、野鲮亚科、鳅鮀亚科、鲤亚科、鱊亚科等8亚科。单因素方差分析显示,鲤科肉食性鱼类的肋骨数与脊椎骨数的比值显著少于植食性鱼类(P＜0.05),而脊椎骨数、尾椎数则显著多于植食性鱼类(P＜0.05)。中上层鱼类的脊椎骨数、尾椎数显著多于下层鱼类(P＜0.05)。极小型鱼类的脊椎骨数、肋骨数,以及肋骨数与脊椎骨数之比显著少于大型鱼类(P＜0.05)。表明鲤科鱼类的中轴骨骼数与其系统发育和生态习性及体型具有明显的相关性。     

Phylogenetic relationships of Cypriniformes and plasticity of pharyngeal teeth in the adaptive radiation of cyprinids

The Cypriniformes comprise approximately 4,200 species accounting for 25% of the diversity of all freshwater fish, which is widely distributed across the world's continents except Antarctica, South America, and Australia. The highest species diversity is found in Southeastern Asia. Despite its remarkable species diversity and broad-scale geographic patterns of distribution, the evolutionary history of this major freshwater fish group remains largely unresolved. To gain insight of the evolutionary history of Cypriniformes, we present a phylogeny of this group using 1 mitochondrial gene and 15 nuclear genes comprising a total of14,061 bp. Bayesian inference using all gene fragments yielded a well resolved phylogeny, which is mostly consistent with topologies obtained from Maximum Likelihood analyses. Our results further confirmed the monophyly of Cypriniformes and seven constituent subclades including Cyprinidae, Catostomidae, Gyrinocheilidae, Balitoridae, Cobitidae, Nemacheilidae, and Botiidae. Bayesian divergence time analysis indicated that the origin of the Cypriniformes was about 193 Mya during the early Jurassic, coinciding with the onset of the Pangaea breakup. The basal divergence of Cypriniformes is 154 Mya during the late Jurassic. Our findings from molecular divergence and biogeographical analysis indicate the most likely initial geographical range of the ancient Cypriniformes was both East and South Asia(Southeastern area of Mesozoic Laurasia). Moreover, the burst in species diversity in Cyprinidae afforded by the nearly worldwide colonization is possibly in response to the plasticity of pharyngeal dentition. The present study demonstrates that the Cypriniformes was about 193 Mya during the early Jurassic,coinciding with the onset of the Pangaea breakup. The plasticity of pharyngeal dentition of cyprinids might contribute to the burst and radiation of this lineage. The phylogenetic and biogeographic analyses in this study help to improve our understanding of the evolutionary history of this diverse and important freshwater fish group.     

Species of Proteocephalus Weinland, 1858 (Cestoda: Proteocephalidae) from cyprinid fishes in North America

A taxonomic study of Proteocephalus species parasitizing cyprinid fishes in North America, namely Proteocephalus ptychocheilus Faust, 1920 and Proteocephalus cobraeformis Haderlie, 1953, has demonstrated their conspecificity with Proteocephalus torulosus (Batsch, 1786), a common parasite of cypriniform fishes (Cyprinidae and Cobitidae) in the Palaearctic Region. Proteocephalus ptychocheilus and P. cobraeformis are considered junior synonyms of P. torulosus.     

Molecular phylogenetics of the family Cyprinidae (Actinopterygii: Cypriniformes) as evidenced by sequence variation in the first intron of S7 ribosomal protein-coding gene: further evidence from a nuclear gene of the systematic chaos in the family

The family Cyprinidae is the largest freshwater fish group in the world, including over 200 genera and 2100 species. The phylogenetic relationships of major clades within this family are simply poorly understood, largely because of the overwhelming diversity of the group; however, several investigators have advanced different hypotheses of relationships that pre- and post-date the use of shared-derived characters as advocated through phylogenetic systematics. As expected, most previous investigations used morphological characters. Recently, mitochondrial DNA (mtDNA) sequences and combined morphological and mtDNA investigations have been used to explore and advance our understanding of species relationships and test monophyletic groupings. Limitations of these studies include limited taxon sampling and a strict reliance upon maternally inherited mtDNA variation. The present study is the first endeavor to recover the phylogenetic relationships of the 12 previously recognized monophyletic subfamilies within the Cyprinidae using newly sequenced nuclear DNA (nDNA) for over 50 species representing members of the different previously hypothesized subfamily and family groupings within the Cyprinidae and from other cypriniform families as outgroup taxa. Hypothesized phylogenetic relationships are constructed using maximum parsimony and Basyesian analyses of 1042 sites, of which 971 sites were variable and 790 were phylogenetically informative. Using other appropriate cypriniform taxa of the families Catostomidae (Myxocyprinus asiaticus), Gyrinocheilidae (Gyrinocheilus aymonieri), and Balitoridae (Nemacheilus sp. and Beaufortia kweichowensis) as outgroups, the Cyprinidae is resolved as a monophyletic group. Within the family the genera Raiamas, Barilius, Danio, and Rasbora, representing many of the tropical cyprinids, represent basal members of the family. All other species can be classified into variably supported and resolved monophyletic lineages, depending upon analysis, that are consistent with or correspond to Barbini and Leuciscini. The Barbini includes taxa traditionally aligned with the subfamily Cyprininae sensu previous morphological revisionary studies by Howes (Barbinae, Labeoninae, Cyprininae and Schizothoracinae). The Leuciscini includes six other subfamilies that are mainly divided into three separate lineages. The relationships among genera and subfamilies are discussed as well as the possible origins of major lineages.     

The development of the osteocranium of Pleurodeles waltlii Michahelles

Under optimal conditions the development of the osteocranium covers the major part of the larval period and lasts about three and a half months. The earliest ossifications are of dermal origin and concern the tooth-bearing bones. The poorly developed enchondral ossifications found in Urodela, appear at a much later stage, just before metamorphosis.     

Comparative studies of the branchial morphology, gill area and gill ultrastructure of some thalassinidean mud-shrimps (Crustacea: Decapoda: Thalassinidea)

The morphology, gill area and branchial formulae of six thalassinidean decapods ( Calocaris macandreae, Jaxea nocturna, Callianassa subterranea, Upogebia stellata, U. deltaura and U. pusilla ) are reported. Additionally, the rarely-encountered Axius stirhynchus receives brief attention. Gill formulae are presented; the simplest arrangements are found in the Callianassidae and Upogebiidae. The deeper-burrowing, deposit-feeding species that are regularly exposed to prolonged periods of hypoxia, i.e. Callianassa subterranea and Jaxea nocturna , had significantly larger gill areas than thalassinideans that occupied more oxygenated burrows ( Upogebia spp., Calocaris macandreae ). The increase in gill area was a result of flattening of the trichobranchiate gill filaments giving rise to a phylloid gill form. It is suggested that the efficiency of gas transfer, and hence diffusing capacity, was enhanced in the phylloid gill by the larger gill area and, because of the reduced cuticle thickness, by the shorter water-haemolymph diffusion distance. The increased diffusion capacity of the phylloid gill is interpreted as a functional adaptation to the more severe physicochemical burrow water conditions experienced by Callianassa subterranea and Jaxea nocturna .     

Evolution and development of the chordates: collagen and pharyngeal cartilage

Chordates evolved a unique body plan within deuterostomes and are considered to share five morphological characters, a muscular postanal tail, a notochord, a dorsal neural tube, an endostyle, and pharyngeal gill slits. The phylum Chordata typically includes three subphyla, Cephalochordata, Vertebrata, and Tunicata, the last showing a chordate body plan only as a larva. Hemichordates, in contrast, have pharyngeal gill slits, an endostyle, and a postanal tail but appear to lack a notochord and dorsal neural tube. Because hemichordates are the sister group of echinoderms, the morphological features shared with the chordates must have been present in the deuterostome ancestor. No extant echinoderms share any of the chordate features, so presumably they have lost these structures evolutionarily. We review the development of chordate characters in hemichordates and present new data characterizing the pharyngeal gill slits and their cartilaginous gill bars. We show that hemichordate gill bars contain collagen and proteoglycans but are acellular. Hemichordates and cephalochordates, or lancelets, show strong similarities in their gill bars, suggesting that an acellular cartilage may have preceded cellular cartilage in deuterostomes. Our evidence suggests that the deuterostome ancestor was a benthic worm with gill slits and acellular gill cartilages.     

Structural adaptations for ram ventilation: Gill fusions in scombrids and billfishes

For ram‐gill ventilators such as tunas and mackerels (family Scombridae) and billfishes (families Istiophoridae, Xiphiidae), fusions binding the gill lamellae and filaments prevent gill deformation by a fast and continuous ventilatory stream. This study examines the gills from 28 scombrid and seven billfish species in order to determine how factors such as body size, swimming speed, and the degree of dependence upon ram ventilation influence the site of occurrence and type of fusions. In the family Scombridae there is a progressive increase in the reliance on ram ventilation that correlates with the elaboration of gill fusions. This ranges from mackerels (tribe Scombrini), which only utilize ram ventilation at fast cruising speeds and lack gill fusions, to tunas (tribe Thunnini) of the genus Thunnus, which are obligate ram ventilators and have two distinct fusion types (one binding the gill lamellae and a second connecting the gill filaments). The billfishes appear to have independently evolved gill fusions that rival those of tunas in terms of structural complexity. Examination of a wide range of body sizes for some scombrids and billfishes shows that gill fusions begin to develop at lengths as small as 2.0 cm fork length. In addition to securing the spatial configuration of the gill sieve, gill fusions also appear to increase branchial resistance to slow the high‐speed current produced by ram ventilation to distribute flow evenly and optimally to the respiratory exchange surfaces. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.