Osteology of Paedocypris,a miniature and highly developmentally truncated fish (Teleostei: Ostariophysi: Cyprinidae)

Species of the cyprinid genus Paedocypris are among the smallest and most developmentally truncated fishes and vertebrates. Our analysis of their skeletal structure reveals a puzzling combination of extreme developmental truncation and an increased morphological complexity in sexually dimorphic characters. The skeleton of Paedocypris is characterized by reduction and loss and resembles in many aspects that of a larval/early juvenile stage of its close relatives. We found 61 characters that have been affected by developmental truncation. A comparison with the skeletal development of a close relative, the zebrafish Danio rerio, demonstrates that the majority of the absent bones or skeletal structures in Paedocypris are those that appear late in the ossification trajectory of the zebrafish. Thus, their absence in Paedocypris seems to be due to the simple developmental truncation of terminal stages in the ossification sequence. Our study of the sexually dimorphic structures in Paedocypris demonstrates that predominantly the male exhibits the more complex state. In relation to the female, male Paedocypris uniquely possess a cleithrum with a pointed posterior process that covers the scapula laterally, and a more medially situated posterior flange that contacts the dorsal area of the coracoid; a massive and heavily ossified uppermost pectoral radial tightly bound to the scapula; thickened and enlarged three uppermost pectoral‐fin rays; a large triangular, dorsolaterally directed process on the outer arm of the massive os suspensorium; and a enlarged and shovel‐like anterodorsally directed basipterygium; and a hypertrophied first pelvic‐fin ray with additional anterior flanges that support keratinized pads of skin. Female Paedocypris show only one structure that is better developed than in males: the first proximal‐middle radial and the anteriormost fin ray of the dorsal fin are more massive and more heavily ossified. Although the function and biological role of these dimorphisms is still unknown, we hypothesize that they are related to a special reproductive behavior. Paedocypris is a prime example for the recent claim that miniaturization among cyprinids is associated with evolutionary novelty only in developmentally truncated miniatures and not in proportioned dwarfs. Paedocypris offers a strong challenge to Schindleria as the most extreme example of developmental truncation known among fishes. We highlight the difficulties that developmentally truncated taxa frequently pose to the resolution of their phylogenetic position and propose an approach to overcome this problem. Our phylogenetic comparison to determine the systematic position of Paedocypris among cyprinids reveals that it shares not only a number of unique absences, but also highly unusual progressive characters with Sundadanio and Danionella, two other Asian miniature cyprinids. We hypothesize that the three genera form a monophyletic group. We further found that Paedocypris and Danionella share a number of uniquely derived characters pointing to a sister group relationship of the two. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Osteology of Priocharax and remarkable developmental truncation in a miniature Amazonian fish (Teleostei: Characiformes: Characidae)

Establishing phylogenetic relationships of miniature fishes is challenging in taxa with developmental truncation. Within the Characiformes, developmental truncation appears to be relatively rare, with the Neotropical genus Priocharax being an example. Priocharax includes three miniature species among the smallest of the order and has been hypothesized to belong to the Heterocharacinae. The pronounced reduction in its skeleton, however, prevented a clearer evaluation of its relationships. The present detailed osteological study was designed to address this question and revealed that 21 bones are absent and nine other skeletal structures are simplified in Priocharax when compared to other characids. Comparison of the skeleton of adult Priocharax with early developmental stages of other characids demonstrated that most of the absences and simplifications can be interpreted as developmental truncations. The most striking developmental truncations are in the pectoral girdle, in which the endoskeleton remains entirely cartilaginous. Other interesting truncations are in the ethmoid region of the skull, infraorbital series, and Weberian apparatus, in which the claustrum is absent. Our study also revealed some unusual sexual dimorphisms in the pelvic girdle. Two cladistic analyses were performed to assess the relationships of Priocharax within the Heterocharacinae. The first consisted of a traditional analysis in which all absences and reductions of Priocharax were coded in the same way as in the remaining taxa. This resulted in three equally most parsimonious topologies, all of which have Priocharax as the most basal taxon of the Heterocharacinae. The second analysis incorporated ontogenetic information, and most absences and reductions of Priocharax were reinterpreted as apomorphic conditions and thus, coded differently from similar conditions in outgroups. This resulted in a single phylogenetic hypothesis with Priocharax and Gnathocharax as sister groups based on seven synapomorphies. Our approach demonstrates the importance of developmental studies to better understand morphological evolution of miniaturized, truncated taxa, and to generate hypotheses of their relationships. J. Morphol. 277:65–85, 2016. © 2015 Wiley Periodicals, Inc.     

Miniatures,morphology and molecules: Paedocypris and its phylogenetic position (Teleostei,Cypriniformes)

We review the morphological and molecular evidence that Mayden & Chen recently used to infer that the developmentally truncated fish genus Paedocypris is not a member of the teleost order Cypriniformes or carp‐like fishes, but is ‘the basal sister group to all Cypriniformes’. This hypothesis contradicts several previous studies that used molecular sequence data or morphological characters. A review of the morphological characters that Mayden & Chen discussed and mapped onto their ‘simplified tree’ shows that these, analysed alone, rather support a close relationship of the cyprinids Sundadanio, Danionella, and Paedocypris. We also present four additional analyses of morphological data, which all contradict Mayden & Chen's result. Despite its highly reductive skeleton, posing a serious problem when analysing its phylogenetic position with skeletal characters, the presence in Paedocypris of the basioccipital masticatory plate is compelling evidence that it is a member of the Cyprinoidei (Cyprinidae plus Psilorhynchidae). Our reanalysis and exploration of their molecular sequence data shows that only a single gene, EGR3, of the six nuclear genes analysed by Mayden & Chen, is responsible for the position of Paedocypris as ‘the basal sister group to all Cypriniformes’. Three independent methods to visualize and analyse phylogenetic signal and conflict of data sets (phylogenetic networks, splits analysis methods or SAMS, and site‐wise likelihood analyses) reveal a high level of character conflict and noise in Mayden & Chen's data set. The ‘basal’ position of Paedocypris seems to be the outcome of the interplay of two long‐branch effects. We apply the same analytical methods to the data set from Rüber et al.'s molecular analysis of the phylogenetic position of Paedocypris and discuss our findings. We conclude that none of the molecular data sets compiled to date can establish the phylogenetic position of Paedocypris with confidence. Morphological data suggest that Paedocypris and Danionella are sister genera, and that their closest relative is Sundadanio, although the position of these three miniatures among cyprinoids is still unclear. © 2014 The Linnean Society of London     

Molecular Phylogeny and Biogeography of Percocypris (Cyprinidae,Teleostei)

Fierce predatory freshwater fishes, the species of Percocypris (Cyprinidae, Teleostei) inhabit large rivers or lakes, and have a specific distribution pattern. Only a single species or subspecies occurs in each large-scale drainage basin of the Southeastern Tibetan Plateau. In this study, the molecular phylogenetic relationships for all but one of the described subspecies/species of Percocypris were investigated based on three mitochondrial genes (16S; COI; Cyt b) and one nuclear marker (Rag2). The results of Maximum Likelihood and Bayesian Inference analyses show that Percocypris is a strongly supported monophyletic group and that it is the sister group of Schizothorax. Combined with analyses of morphological characters, our results suggest that Percocypris needs to be reclassified, and we propose that six species be recognized, with corresponding distributions in five main drainages (including one lake). In addition, based on the results of the estimation of divergence times and ancestral drainages, we hypothesize that Percocypris likely originated in the early Miocene from a paleo-connected drainage system containing the contemporary main drainages of the Southeastern Tibetan Plateau. This study suggests that vicariance (due to the uplift of the Tibetan Plateau modifying the large-scale morphologies of drainage basins in the Southeastern Tibetan Plateau) has played an important role in the speciation of the genus. Furthermore, external morphological characters (such as the length of the fins) and an internal trait (the position of pterygiophore) appear to be correlated with different habitats in rivers and the lake.     

Evidence for developmental linkage of pigment patterns with body size and shape in danios (Teleostei: Cyprinidae)

Abstract. Variation in pigment patterns in fishes is known to be subject to natural and sexual selection, but the mechanisms by which that variation is generated are only beginning to be understood. Theoretical models of pigment pattern formation in animals suggest that the size and shape of the organism at the time of pattern determination as well as subsequent growth time are important determinants of pattern. However, few data document the empirical relationship of pigment patterning with size, shape, and growth. Here we document patterns of growth in relation to pigment pattern formation in the zebrafish ( Danio rerio ) and six close relatives. In all species examined, establishment of adult pigment pattern within a particular region of the body is associated with a period of substantial growth and shape change in that region of the body. Furthermore, forms with more horizontal stripes on the midbody as adults ( Danio rerio and D. rerio " leo ") are larger at the time pigment cells begin to assume their adult pattern. Finally, continued deepening of the body as the pigment pattern develops is associated with vertical distortions and reticulations in the patterns of D. malabaricus and D. browni . These results are consistent with the predictions of theoretical models that size, growth, and shape change are critical determinants of pigment patterning, and suggest that variation in pigment pattern may arise in part through differential allometric growth between species.     

Vertebral Dimensions in the Genus Carasobarbus (Teleostei: Cyprinidae)

Journal of Ichthyology - The morphological study of the axial skeleton of the nine species of the genus Carasobarbus Karaman, 1971 enabled to confirm the division of this bony structure into five...     

Masculinization mechanism of hybrids in bitterlings (Teleostei: Cyprinidae)

The sex ratio of bitterling hybrids (subfamily: Acheilognathinae) is often likely to be biased toward males. Artificial hybridization was carried out in 10 species of bitterlings (three genera) in order to elucidate the masculinization mechanism of hybrids. Tanakia himantegus never produced viable F1 hybrids with other species, while hybrids of most other species were viable. In terms of sex ratio and fertility, hybrids were clearly divided into two groups: congeneric Tanakia hybrids and others. Both male and female congeneric Tanakia hybrids were fertile. The sex ratio was nearly 1:1 in all groups of Tanakia hybrids. Except for the congeneric Tanakia hybrids, sterile males appeared predominantly in groups of hybrids in which females were very rare but remained fertile. Sterile intersexes were also observed in five hybrid groups: T. lanceolata (female) x Acheilognathus cyanostigma (male), Rhodeus uyekii (female) x T. lanceolata (male), A. rhombeus (female) x T. lanceolata (male), A. rhombeus (female) x T. limbata (male), and A. tabira tabira (female) x A. cyanostigma (male). In the development of male-predominant hybrids, although hybrid and control (parental species) hatching and survival rates do not differ, no females appeared in hybrids, contrary to the controls. Taking the female heterogametic sex-determining system (ZW) and the phylogenetic relationship of bitterlings into consideration, the masculinization mechanism of hybrids in bitterlings can be explained by the interaction of two sex chromosomes, derived from each parental species. The basic genetic sex in bitterlings is male (ZZ) and the derivative is female (ZW). When parental species are related, the sex phenotype of hybrids coincides with the genetic sex. However, when the parental species differ, the sex phenotype of the ZW genotype is reversed to become male by an abnormal interaction between the Z and W chromosomes. The rare appearance of females and intersexes in male-predominant hybrids might be due to complete or partial functional expression of the W chromosome.     

Africa's Clarias gariepinus (Teleostei: Clariidae) appears in rivers in Brazil

As a major tributary of the Nyl River, and ultimately for the Nylsvley wetland, the Olifantspruit, Limpopo province, South Africa, was investigated during the summer (high flow) and winter (low flow) of 2007 at three sites. This preliminary study used physical and biological indicators to determine the quality of the water that ultimately flows into Nylsvley. Significant seasonal differences were found. The observed differences relate to habitat structural differences and human activities in the catchment. Key findings included higher water temperatures during the high-flow period than during the low-flow period. The higher temperatures were accompanied by lower concentrations of dissolved oxygen at all sites. In total, 13 fish species were recorded, with most being found in the pool biotopes. The Nantes site, located nearest to the Nylsvley wetland, had the highest number of fish species, possibly indicating a linkage between these two systems. The most common macroinvertebrate families were the Baetidae, Aeshnidae and Coenagrionidae. It can be assumed that this river acts as an important refuge area for fish and macroinvertebrates from the Nyl River, and therefore it is important to conserve the Olifantspruit.     

Molecular phylogeny of the genus Capoeta (Teleostei: Cyprinidae) in Anatolia,Turkey

Phylogeny of Capoeta genus distributed in Anatolia were carried out by analysing mitochondrial cytochrome b gene (1140 bp) sequences from 332 samples representing 59 populations of 15 species across their geographical distribution. Haplotype network and phylogenetic analysis (neighbor-joining, maximum-likelihood, maximum parsimony, and bayesian inference) of the 103 cytochrome b haplotypes detected in Capoeta species resulted in similar tree topologies including four distinct clades, in congruent with taxonomic classification of Capoeta based on morphological characteristics such as scale size, mouth shape, and body spotting. Based on cyt b nucleotide sequences, the present study suggests that four undescribed Capoeta species may exist in Anatolia freshwater; one species in the Kizilirmak River, the second species in the Dirgine River, the third species B. Menderes River, and the fourth species in the some Yesilirmak tributaries that run into the Black Sea Basin. Capoeta taxa distributed in the rivers of Anatolian freshwater basins are isolated from each other during middle Miocene (Serravallian)-late Pleistocene (Ionian) (about 13.75–0.41 million years). This suggests that distribution and presence of Capoeta species were shaped under paleogeographic conditions such as Pleistocene climate changes in Quarternary period as well as tectonic uplift and faulting, which probably has not changed up to now.     

Breeding tubercles of Phoxinus (Teleostei: Cyprinidae): Morphology,distribution, and phylogenetic implications

Studies of the skin with scanning electron microscopy (SEM) reveal a diverse morphology in breeding tubercles among species of Phoxinus. Based mainly on the fine structure of the surface of tubercles, nine morphotypes, coded as letters A-I, occur in Phoxinus. Most of the morphotypes are common to all Phoxinus species, but type E is present only on the dorsum of the head of P. phoxinus, type H on the breast scale of female P. phoxinus, and type I on the pectoral fin in P. erythrogaster. Multicellular breeding tubercles bearing unicellular projections, identified as unculiferous tubercles are found in type H and probably types F and G. The distribution of tubercles on head, body, and fins is described and compared among Phoxinus species. Breeding tubercles in Phoxinus and other minnow genera are compared in order to interpret the phylogenetic implication of the tuberculation in Phoxinus. The deeply embedded breast scales and the breeding tubercles on their apical margins, and a series of tubercles on the apical margins of lateral scales of the caudal peduncle in breeding males of Phoxinus species, are the characters supporting the monophyly of the genus. © 1996 Wiley-Liss, Inc.     

Diversity,phylogeny and biogeography of Systomus (Teleostei,Cyprinidae) in Sri Lanka

The South and South-East Asian freshwater fish genus Systomus (Cyprinidae) comprises 17 valid species. Six nominal species, including three endemics, have been reported from Sri Lanka, a continental island separated from India by a shallow-shelf sea. The species diversity of Systomus on the island has until now not been assessed; neither has an evaluation been made of their phylogenetic history. Here, based on an analysis of the nuclear recombination activating protein 1 (rag1), and mitochondrial cytochrome c oxidase subunit 1 (cox1) and cytochrome b (cytb) gene markers, and a morphological examination of 143 specimens from 49 locations in Sri Lanka, we reassess the diversity of Systomus on the island and analyse patterns of their evolution and biogeography. Divergence-time estimates, based on a substitution rate calibration, date the basal split between Systomus and its sister group, the Afrotropical small barbs, to 30.0 Ma (95% highest posterior density: 25.4–35.2 Ma). The species of Systomus belong to two distinct clades. The first includes the Sri Lankan endemics S. asoka, S. martenstyni and S. pleurotaenia, which comprise an insular diversification following the immigration of a common ancestor during the Oligocene. The second, which includes the remaining species of Indian, Sri Lankan and South-East Asian Systomus, has a crown age dating to the Late Miocene. Morphological and molecular species delimitation analyses failed to validate the two nominal species, S. spilurus and S. timbiri, previously reported from Sri Lanka: both are considered synonyms of S. sarana, as are the nomina S. chryseus, S. chrysopoma, S. laticeps, S. rufus, S. pinnauratus and S. subnasutus. Four genetically and geographically discrete lineages of S. sarana occur in the island, and three in India. Molecular species delimitation analysis suggests these all belong to a single species, S. sarana. The genetically distinct Sri Lankan populations of S. sarana result from Plio-Pleistocene dispersal or vicariance events between India and Sri Lanka—as a result of emergence and inundation of the now submerged isthmus connecting the two landmasses—as well as autochthonous insular diversification.     

Weitere elektronenmikroskopische Untersuchungen am Pinealorgan vonPhoxinus laevis (Teleostei,Cyprinidae)

Zusammenfassung Auch im Pinealorgan adulter Elritzen (Phoxinus laevis) sind die Außenglieder der Sinneszellen nicht degeneriert. Der Vergleich wurde zwischen adulten Exemplaren (8,6 cm) und 19,5 Monate jüngeren Tieren (3 cm) durchgeführt. Im Verhältnis zu den Jungtieren sind die Neuropilformationen und die synaptischen Strukturen adulter Elritzen stärker ausgeprägt. Bei den letzteren enthalten die Endfüße der pinealen Rezeptoren zahlreiche synaptische Bänder und 300–400 Å große Bläschen. Die Neuropilzonen sind außerdem reich an verdichteten Membranstellen (tight junctions). Im Pinealorgan des adulten Phoxinus finden sich eigenartige Einschlußkörper, die aus konzentrischen Lamellen bestehen; ähnliche Membran wirbel hat Takahashi (1969) bei zweijährigen Goldfischen beschrieben. Häufig lassen sich in den Epiphysen adulter Elritzen Axone beobachten, die mit 500–1000 Å großen granulierten Vesikeln gefüllt sind und im Verband bahnartig zusammengefaßter markloser Nervenfasern verlaufen. Die Herkunft und Bedeutung dieser granulierten Fasern werden diskutiert, ebenso wie die funktionelle Rolle des pinealen Sinnesorgans.

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Further electron microscopic studies of the pineal organ inPhoxinus laevis (Teleostei, Cyprinidae)

Summary The outer segments of pineal receptor cells of adult (8.6 cm) European minnows (Phoxinus laevis), like the lamellated outer segments of 19.5 months younger (3 cm) specimens, did not show signs of degeneration (see also Oksche and Kirschstein, 1967). Neuropile formations and synaptic structures were more prominent in adult than in youngPhoxinus. In the adult group, the end-feet of pineal receptors contained numerous synaptic ribbons and vesicles (300–400 Å in diameter). The neuropile zones were rich in tight junctions. A peculiar structure of the pineal organ of agingPhoxinus was the inclusion body formed by concentric arrangement of lamellae. Membrane whorls similar to these inclusion bodies were described in the pineal organ of two years old goldfish (Takahashi, 1969). In the pineal organ of adultPhoxinus considerable numbers of axons containing dense-core vesicles (500–1,000 Å in diameter) were observed within tract-like bundles of unmyelinated nerve fibers. The origin and significance of these granulated fibers and the functional role of the pineal sense organ have been discussed.

Mit Unterstützung durch die Deutsche Forschungsgemeinschaft.     

BarredDanio species from the Irrawaddy River drainage (Teleostei, Cyprinidae)

The three barredDanio species known from the Irrawaddy River drainage are redescribed. Two species are known from southwestern Yunnan:Danio interruptus in the Husa He and Longchuan, Jian watersheds, andD. apogon abundantly in the main river and tributaries of the Daying Jiang basin. The third species,D. shanensis, occurs in the Myitnge basin of northern Shan State, Myanmar.Danio interruptus differs from other barredDanio species by having a short, incomplete lateral line, a deeper head, and shorter thicker dark vertical bars or blotches anteriorly on the sides.Danio shanensis has the lateral line either complete or incomplete, lesser head depth, and a series of deeper narrower vertical bars.Danio apogon has no lateral line, narrow or indistinct dark vertical bars, and a wide, but indistinct, horizonatal P stripe.Danio choprae, previously considered, as a barredDanio species, is more closely related to theD. dangila species group.     

Molecular systematics, phylogeny and biogeography of roaches (Rutilus, Teleostei, Cyprinidae)

In this study we used sequence data from the entire mtDNA cytochrome b gene to reconstruct patterns and times of diversification in the roach genus Rutilus. The genus is present with numerous endemic species in the Eastern peri-Mediterranean area and with a few widespread species in Central Europe. Our phylogenetic results do not support the subdivision into two subgenera proposed on morphological grounds. Within R. pigus and R. rutilus we identify highly divergent and allopatric mitochondrial lineages. The deeper splits in the genus phylogeny date back to the middle Miocene; the main diversification took place at the Miocene-Pliocene boundary.     

Brain morphology and turbidity preference in Notropis and related genera (Cyprinidae,Teleostei)

Synopsis The size of seven neural structures was compared in 51 species of Notropis, Pteronotropis, Cyprinella, Luxilus, Lythrurus, and Hybopsis, and related to the turbidity of the species& habitat. This last parameter was assessed for each species by personal communication with 42 ichthyologists. To control for size differences among species, all analyses were performed on the residuals from a regression of each character on standard length. Principal components analysis (PCA) of the residuals produced four significant PC-axes that together explained 65% of the total variation represented in the original variables. The size of brain structures concerned with vision, olfaction, and gustation was correlated with habitat turbidity. Two-way Analyses of Covariance (ANCOVAs) revealed significant differences between species in the size of all structures. Sexual dimorphism was found in the size of the olfactory bulb and the cerebellum, and significant two-way interactions (species vs. sex) were detected for the telencephalon, optic lobes, cerebellum, vagal lobe, and the eye. Cluster analysis indicated that neither similar turbidity preference nor shared phylogeny is alone sufficient to explain the observed differences in brain morphology.     

Morphology and fine structure of Barbus barbus (Teleostei: Cyprinidae) spermatozoa

Morphology and fine structure of Barbus barbus L 1758 spermatozoa were studied using scanning (SEM) and transmission (TEM) electron microscopy. The results confirm that spermatozoa exhibit morphological features typical to all teleost fishes. They are differentiated into a head, a midpiece and a flagellum with the typical '9 + 2' pairs of microtubules. Both dynein arms are present in the flagellum. The spermatozoa have spherical nuclei, 4–6 mitochondria located in the postnuclear cytoplasmic region and centriolar complex (proximal and distal centrioles). Total length, head width, length of midpiece and length of flagellum were measured to be 56.35 ± 7.42, 1.80 ± 0.06, 0.48 ± 0.14 and 54.30 ± 6.97 μm, respectively. Highly significant linear correlation was observed between posterior and anterior width of midpiece (P < 0.01). Principal component analysis (PCA) was used to explore which parameters can explain the individual variation of sperm morphology. About 44% of the total accumulated variance was absorbed by the analysis of the two first components, distinguishing different groups of parameters related to head and midpiece. The lengths of flagellum and head are more isolated; indicating that the individual variation of sperm morphology depends on these two parameters. Comparing the results of this study with information on cyprinids spermatozoa reveals that the number of mitochondria and the length of the flagellum are good characters to characterize spermatozoa of the Cyprinidae in a phylogenetic arrangement.     

Genetic variability in Chondrostoma from Austrian, French and Greek rivers (Teleostei, Cyprinidae)

Allozyme variation in Chondrostoma from Austrian, French and northern Greek rivers was investigated. Populations of C. nasus from the upper Danube catchment are genetically very similar and apparently connected by high level of gene flow. Chondrostoma nasus has conserved all major allozyme polymorphisms during its recent range expansion into the Rhone basin. Chondrostoma vardarense from the Axios and Aliakmonas rivers share most of their allozymes with C. nasus from the upper Danube catchment, but have less genetic variation, a possible consequence of a genetic bottleneck following river capture.     

Mitochondrial diversity of Opsariichthys bidens (Teleostei, Cyprinidae) in three Chinese drainages

We describe the phylogeographic structure of 28 Chinese populations of the cyprinid Opsariichthys bidens across three main Chinese river drainages. Our study is based on the phylogenetic analysis of the complete mitochondrial cytochrome b gene (1140 bp). We combined this analysis with population processes inferred from nested clade analysis (NCA) and mismatch distributions. Both analyses showed that Chinese O. bidens consists of five mtDNA lineages (Opsariichthys 1-5) with high genetic divergence among them. Molecular divergences (TrN+G) higher than 20% among the Opsariichthys 1-5 mtDNA lineages suggest a taxonomic underestimation at the species level. About 92% of the genetic variance among samples was explained by differences among Opsariichthys mtDNA lineages. Drainage-restricted haplotypes with high frequencies and moderate nucleotide diversity show that Opsariichthys populations have evolved independently. NCA results were congruent with the phylogeny, and unimodal mismatch distributions with negative Tajima's D values suggest population expansions in some Opsariichthys lineages. The phylogeographic structure of the Opsariichthys 1-5 mtDNA lineages appears to be related to their long-term interruption of gene flow (theta(ST)>0.97). Our results suggested that fragmentation of ancestral ranges might have caused Opsariichthys diversification in Chinese waters. However, current distribution of common haplotypes across the Yangtze and Pearl drainages suggests a recent river connection that could have favoured gene flow across drainages. Overall, the results indicated that the richness of current Asian widespread species might have been underestimated, and that the cyprinid populations of O. bidens in the Yangtze, Pearl and Hai He drainages may correspond to five species.     

Phylogeny,genetic diversity and phylogeography of the genus Codoma (Teleostei,Cyprinidae)

This study represents a thorough analysis of Codoma, a monotypic genus endemic to north‐western Mexico. A previous morphological analysis of the species concluded that there exists several morphological groups in Codoma ornata, suggesting diversity in Codoma could be underestimated. No studies have examined the genetic diversity in Codoma ornata to test this hypothesis and identify independent lineages. We present a phylogeographic analysis using one mitochondrial and two nuclear genes, and specimens from across nine major drainages in both the Chihuahuan Desert and the Sierra Madre Occidental of western Mexico. All genes and analyses recovered populations of Codoma in a well‐supported clade and sister to Tampichthys, and this clade sister to Cyprinella. Analyses of both mitochondrial and nuclear genes indicated Codoma is not monotypic and recover more diversity in the genus than currently recognized. The four (mitochondrial) and five (nuclear) genetically distinct lineages are consistent with those groups outlined in the prior morphological study of the genus. Composition and distribution of these major lineages is also consistent with prior biogeographic hypothesis for other fishes in the region, supporting an ancestral Rio Grande system extending south towards central Mexico. Fragmentation of this paleosystem was followed by allopatric speciation in the Chihuahuan Desert. These results suggest a scenario of long‐term isolation in four major regions (upper Conchos, lower Conchos, Nazas, upper Mezquital). Resolution of the diversity and biogeography of these lineages has many implications for various biological disciplines, especially for evolutionary and conservation studies.