Genetic relationships of Brachygobius and related brackish Indo-west Pacific and Australasian genera (Teleostei: Gobiidae)

Horizontal starch gel electrophoresis was employed to analyse genetic variation, at 19 isozyme loci, in six putatively related species of gobies ( Brachygobius aggregatus, B. doriae, Pandaka lidwilli, Pseudogobius olorum, Pseudogobius sp. and Redigobius sp.), using seven other gobiine species as an outgroup. Allozyme data was analysed both cladistically and phenetically, all treatments confirming the monophyly of the group. Pandaka is confirmed as the sister genus to Brachygobius , these genera forming a crown group, with Redigobius and Pseudogobius forming successive sister groups. This scenario is in agreement with morphological hypotheses of phylogeny. However, this lineage appears to be relatively unrelated to other groups within the Gobiinae     

Isozyme criteria in the testing of phyletic relationships between species of Gobius and related eastern Atlantic-Mediterranean genera (Teleostei: Gobüdae)

Horizontal starch gel electrophoresis was employed to investigate allozyme variation at 19 isozyme loci, in nine closely-related Atlantic-Mediterranean gobiid fish species, to assess genetic relationships, and to compare this with a phyletic hypothesis based on morphological apo-morphies. The species examined were Gobius niger, G. auratus, G. cruentatus, G. paganellus, Mauligobius maderensis, Zosterisessor ophiocephalus, Thorogobius ephippiatus, Padogobius martensii and P. nigricans . Various phenetic and cladistic analyses were performed on isozyme and morphological data. The phenetic and cladistic results from morphological data, and the cladistic results from isozyme data, were largely comparable, with P. martensü and P. nigricans forming a sister group to all the other taxa, species of Cobius forming a crown group within the latter, and stem lines formed by Mauligobius, Zosterisessor and Thorogobius , respectively. The isozymic estimates of genetic distance differed from these chiefly in the relative positions of Zosterisessor and G. auratus , which appear less similar to other species of Gohius. G. auratus shows many alleles which are unique within the genus Gobius ; possible explanations for this are put forward. The position of nigricans as congeneric with Padogobius martensü is confirmed by both phenetic and cladistic analyses of isozyme data.     

Molecular phylogeny of the gobioid fishes (Teleostei: Perciformes: Gobioidei)

The phylogeny of groups within Gobioidei is examined with molecular sequence data. Gobioidei is a speciose, morphologically diverse group of teleost fishes, most of which are small, benthic, and marine. Efforts to hypothesize relationships among the gobioid groups have been hampered by the prevalence of reductive evolution among goby species; such reduction can make identification of informative morphological characters particularly difficult. Gobies have been variously grouped into two to nine families, several with included subfamilies, but most existing taxonomies are not phylogenetic and few cladistic hypotheses of relationships among goby groups have been advanced. In this study, representatives of eight of the nine gobioid familes (Eleotridae, Odontobutidae, Xenisthmidae, Gobiidae, Kraemeriidae, Schindleriidae, Microdesmidae, and Ptereleotridae), selected to sample broadly from the range of goby diversity, were examined. Complete sequence from the mitochondrial ND1, ND2, and COI genes (3573 bp) was used in a cladistic parsimony analysis to hypothesize relationships among the gobioid groups. A single most parsimonious topology was obtained, with decay indices indicating strong support for most nodes. Major phylogenetic conclusions include that Xenisthmidae is part of Eleotridae, and Eleotridae is paraphyletic with respect to a clade composed of Gobiidae, Microdesmidae, Ptereleotridae, Kraemeriidae, and Schindleriidae. Within this five-family clade, two clades are recovered. One includes Gobionellinae, which is paraphyletic with respect to Kraemeriidae, Sicydiinae, Oxudercinae, and Amblyopinae. The other contains Gobiinae, also paraphyletic, and including Microdesmidae, Ptereleotridae, and Schindleriidae. Previous morphological evidence for goby groupings is discussed; the phylogenetic hypothesis indicates that the morphological reduction observed in many goby species has been derived several times independently.     

Sequence length and variation in the mitochondrial control region of two freshwater gobiid fishes belonging to Rhinogobius (Teleostei: Gobioidei)

The control region (D-loop) of mitochondrial DNA (mtDNA) was amplified and sequenced for eight samples of the rhinogobies Rhinogobius maculafasciatus and R. giurinus from Taiwan and southern China. The control regions of both species are of 841–842 bp; the length of these sequences being the most compact among all known sequences in teleost fishes. Three conserved sequence blocks (CSB) were observed. The full D-loop and tRNA Phe gene sequences were determined and compared with other fishes. The interspecific sequence divergence between the two species is 11.3–11.7%; and the intraspecific variation in R. guirinus 0.8–1.8%. Results suggest that the control region of Rhinogobius is informative for phylogenetic reconstruction at both intraspecific and interspecific levels in this gobiid genus.     

The innervation of head neuromast rows in eleotridine gobies (Teleostei: Gobioidei)

The innervation of free neuromast (sensory papillae) rows is described from Sihler wholemount preparations of four species of eleotridine gobies, one ( Perccottus glenii ) representing the 'longitudinal' type of neuromast arrangement, the others ( Butis buits, Bostrychus urophthalmus, B. marmoratus ) the 'transverse' arrangement. In the latter, the characteristic transverse cheek rows (1–7) are innervated from the infraorbital trunk of the anterior lateral-line nerve. Longitudinal cheek rows b and d , and the three opercular rows, ot, os and oi , common to all species, are innervated by rami of the hyomandibular trunk of the same nerve. Two neuromast groupings are shown to have a mixed nerve supply. For the median preorbital snout rows, there is innervation from the infraorbital ( s ³ and r ) as well as the supraorbital ( s ¹ and s ²) trunks of the anterior lateral line nerve. The anterior dorsal rows are supplied both by the posterior lateral-line supratemporal ramus (rows g and m ) and the anterior lateral-line supraorbital trunk (rows o and n ). The neuromast rows, under the designations of Sanzo, are tabulated according to innervation and their putative origin in the phyletic replacement of a complete head canal system seen in more generalized percomorph fishes.     

Ubiquity of Sicyopterus lagocephalus (Teleostei: Gobioidei) and phylogeography of the genus Sicyopterus in the Indo-Pacific area inferred from mitochondrial cytochrome b gene

Sicyopterus lagocephalus is a Gobiidae Sicydiinae (Teleostei) thought to inhabit Indo-Pacific island rivers from Comoros Islands in the Indian Ocean to Australs Islands (French Polynesia) in the Pacific Ocean. Its biological cycle comprises a marine planctonic larval phase of several months allowing it to migrate from island to island, but the other species of the genus, with such a larval stage, have generally a more restricted range and are often endemic. To understand the organisation of a species with such a wide distribution, mtDNA cytochrome b sequences were amplified for 55 specimens of this genus covering most of its distribution range together with six close endemic species and other gobiids used as outgroups. The main result is the confirmation of the ubiquity of S. lagocephalus that occurs over a range of 18,000 km in the Indian and Pacific Oceans. Two clades were identified within this species, one clustering most of French Polynesian haplotypes and the other clustering most of Mascarene (including Comoros) haplotypes. The overall pattern of distribution and phylogenetic relationship suggests that the lineages leading to endemic species originated earlier than S. lagocephalus. This latter seems to be a secondary migrant species, having colonised both Indian and Pacific Oceans with a few exceptions, situated at the border of the range (Madagascar, Marquesas, Rapa). According to the results, the phylogeny of the Sicyopterus group, the age of the different lineages and the past history of the colonisation of the Indo-Pacific islands are discussed.     

Systematics of freshwater gobies from Greece (Teleostei: Gobiidae)

Five species of gobiid fish are recognized in the fresh waters of Greece. Most distinctive is the south-western Hellenic (West Balkanian component) endemic genus Economidichthys Bianco, Bullock, Miller & Roubal, 1987, with the type-species E. pygmaeus (Holly, 1929) from Lefkas, Epirus and Lake Trichonis, and E. trichonis sp. nov., from Lake Trichonis, the latter probably the smallest freshwater European teleost at maturity (female, 18·0 mm) and maximum size (male, 30·0 mm). In tributaries of the northern Aegean basin (north-eastern Hellenic freshwater fauna), two species of the Ponto-Caspian genus Knipowitschia can be identified, the widespread Kn. caucasica (Kawrajsky in Berg, 1916) (coastal plain of Macedonia and Thrace) and the endemic Kn. ihessala (Vinciguerra, 1921) (Pinios river system, Thessaly). The larger Ponto-Caspian tubenose goby, Prolerorhinus marmoratus (Pallas, 1811), occurs in the Evros (= Maritsa) (Thrace) and Strymon (Macedonia) systems. Only the genera Economidichthys and Knipowitschia are considered in this paper, being redefined in terms of osteology and head lateral-line system, and the species described in detail, with special reference to meristic variation and squamation, and, in Economidichthys , the unique perianal organ of this genus. The relationships of Economidichthys and Kn. thessala are discussed and possible scenarios for their evolution are suggested in the light of Miocene to Recent events.     

Global diversity of halacarid mites (Halacaridae: Acari: Arachnida) in freshwater

Halacarid mites have successfully invaded the sea and approximately 56 species colonized the freshwater. Invasion from the sea into continental waters probably started in the Mesozoic or Pre-Mesozoic and went on in the following epochs. The number of genera (14) and species (34) recorded from the Palaearctic is remarkably higher than that of other geographical regions. These numbers do not imply that the Palaearctic is a centre of origin, they reflect the sampling activity rather than reliable data on diversity. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Unusual gonad structure in the paedomorphic teleost Schindleria praematura(Teleostei: Gobioidei): a comparison with other gobioid fishes

The gonads of eight gobioid species were examined histologically, including Pandaka pygmaea and Schindleria praematura , in order to investigate the manifestations of miniaturization and paedomorphosis in the gonads. Rearrangements of reproductive structures were found only in S. praematura , and included only the gonad tissues, not the gametes. Both sexes of S. praematura maintain basic germinal components: the spermatocyst composed of Sertoli cells and spermatogonia in the testis and the follicle and oocyte complex in the ovary. The stromal ovarian tissue and testicular interstitial tissue, however, is reduced compared to other gobies, and the number and location of gonial cells is restricted. This sequestered pattern of gonial cells is known as a restricted spermatogonial type in the testis, and has been reported only in atherinomorphs. The restriction of gonial cells in the ovary is extremely rare among teleosts, known only from one other species. These restricted gonial patterns in S. praematura are probably related to the overall reduction of morphological complexity in this genus, due to its extreme paedomorphosis.     

Molecular phylogeny of gobioid fishes (Perciformes: Gobioidei) based on mitochondrial 12S rRNA sequences

The molecular phylogeny of the gobioid fishes, comprising 33 genera and 43 valid species, was examined by use of complete mitochondrial 12S rRNA and tRNA(VAL)genes. Both parsimony and neighbor-joining analyses revealed comparable results and are generally congruent with those of morphological studies. The Odontobutis, which was always placed at the base of the phylogenetic trees, can be treated as a sister group of all other nonrhyacichthyid gobioids. Within eleotrid fishes, the monophyly of the Eleotrinae is strongly supported by molecular data. The Butinae is closer to fishes with five branchiostegal rays and should be treated as a sister group of the latter. The group with five branchiostegal rays, except for sicydiines, can be divided into two groups according to their epural counts. Fish with one epural, the Gobiinae of Pezold plus Microdesmidae, form a monophyletic group which is sister to those with two epurals, the Oxudercinae and Gobionellinae of Pezold. However, Sicydiinae, which have one epural, are closer to the Oxudercinae and Gobionellinae rather than to the Gobiinae. Since progressive reduction in epural number has been observed along this lineage, the sicydiines should be treated as a derived group within the groups with two epurals.     

Isozyme analysis of Galaxias species (Teleostei: Galaxiidae) from the Taieri River, South Island, New Zealand: a species complex revealed

We examined genetic differentiation among 23 samples of non-migratory river galaxias from 17 streams in the Taieri River system, South Island, New Zealand. Four major genetic types were found, two of which occur in narrow sympatry in one location. These were compared with topotypical material representing Galaxias anomalus from the Clutha system (Otago) and G. vulgaris from the Waimakariri system (Canterbury) in order to establish identity. Morphological examination of these four major genetic types revealed consistent concomitant differences. The results suggest that there are at least three species of river galaxias in the Taieri system: G. anomalus, G. vulgaris and at least one previously undescribed species. We propose that the genetic structuring and subsequent speciation of this group has been promoted by the absence of the marine juvenile phase that is found in five other members of the genus native to New Zealand. This structuring may be exacerbated by population fragmentation over the last century owing to the negative influence of introduced trout. The phylogenetic diversity within the river system mirrors the diverse flora and invertebrate fauna of the region, and has conservation implications that parallel those resulting from our improved knowledge of the New Zealand herpetofauna through the application of genetic analysis.     

Phylogeny of the flyingfish family Exocoetidae (Teleostei, Beloniformes)

The phylogeny of the flyingfish family Exocoetidae is studied cladistically, using 41 morphological characters encompassing early life history, and external and internal features. The monophyly of the family is supported by 10 synapomorphies. Within the family,Oxyporhamphus is the sister group to all other genera, the monophyly of the latter being defined by 10 synapomorphies.Fodiator is the sister group of genera characterized by the presence of chin barbels in juveniles.Parexocoetus is the sister group ofExocoetus, Cypselurus, Prognichthys andHirundichthys, the latter being defined by four synapomorphies. In the latter group,Exocoetus is the sister group of the other three genera. The phylogeny of the Exocoetidae is characterized by the stepwise upgrading of gliding capability, with sequential modifications of the caudal, pectoral and pelvic fins. The subfamily Oxyporhamphinae is resurrected.     

Morphometric convergence among European sand gobies in freshwater (Gobiiformes: Gobionellidae)

The five genera of sand gobies inhabit the seas and freshwaters of Europe and western Asia and occupy habitats ranging from fully marine to exclusively freshwater. In this study, we use geometric morphometrics to quantify body shape among sand gobies, in order to investigate how shape has evolved and how it is related to habitat. We also compare body shape between preserved museum specimens and fresh specimens, to determine whether or not fixation and storage in ethanol introduce detectable bias. We confirm that the fixed specimens exhibit significant shape changes as compared to fresh specimens, and so, we perform the bulk of our analyses exclusively on fixed specimens. We find that Economidichthys, Orsinigobius, and Pomatoschistus occupy distinct regions of morphospace. Knipowitschia and Ninnigobius have intermediate forms that overlap with Pomatoschistus and Orsinigobius, but not Economidichthys. This pattern is also in rough accordance with their habitats: Pomatoschistus is fully marine, Economidichthys fully freshwater, and the others fresh with some brackish tolerance. We augment a recent phylogeny of sand gobies with data for P. quagga and interpret morphometric shape change on that tree. We then evaluate convergence in form among disparate lineages of freshwater species by constructing a phylomorphospace and applying pattern‐based (convevol) measures of convergence. We find that freshwater taxa occupy a mostly separate region of morphospace from marine taxa and exhibit significant convergence in form. Freshwater taxa are characterized by relatively larger heads and stockier bodies than their marine relatives, potentially due to a common pattern of heterochronic size reduction.     

Molecular phylogeny of basal gobioid fishes: Rhyacichthyidae, Odontobutidae, Xenisthmidae, Eleotridae (Teleostei: Perciformes: Gobioidei)

Morphological character analyses indicate that Rhyacichthyidae, Odontobutidae, Eleotridae, and Xenisthmidae are the basal families within the perciform suborder Gobioidei. This study uses DNA sequence data to infer the relationships of genera within these families, as well as determine the placement of more derived gobioids (family Gobiidae) and the identity of the outgroup to Gobioidei. Complete sequences of the mitochondrial ND1, ND2, COI, and cyt b genes (4397 base pairs) are analyzed for representatives of 27 gobioid genera and a variety of perciform and scorpaeniform outgroup candidates; the phylogeny is rooted with a beryciform as a distal outgroup. The single most parsimonious tree that results indicates that, of the outgroups sampled, the perciform family Apogonidae is most closely related to Gobioidei. Gobioidei is monophyletic, and Rhyacichthys aspro is the most basal taxon. The remainder of Gobioidei is resolved into clades corresponding to the families Odontobutidae (plus Milyeringa) and Eleotridae+Xenisthmidae+Gobiidae. Within Eleotridae, the subfamily Butinae (minus Milyeringa) is paraphyletic with respect to Gobiidae, and Eleotrinae is paraphyletic with respect to Xenisthmidae. Other than these groupings, the primary disagreement with the current morphology-based classification is that the molecular data indicate that the troglodytic Milyeringa should be placed in Odontobutidae, not Butinae, although support for this placement is weak. The most basal lineage of Gobioidei is known from the freshwaters of the Indo-Pacific, with marine-dwelling lineages arising several times independently in the group. The phylogeny also indicates that different gobioid lineages are distributed in Asia, Africa, Madagascar and the Neotropics. Five sister pairs of basal gobioid species inhabit Atlantic and Pacific drainages of Panama, with widely varying divergences.     

Length and sequence variation in the mitochondrialDNA control region of the Etruscan freshwater goby Padogobius nigricans (Teleostei, Gobiidae)

The primary structures of the mtDNA control region, the flanking tRNA^Pro and tRNA^Phe genes and the partial 12S rRNA gene of the freshwater Etruscan goby Padogobius nigricans (Teleostei, Gobiidae) were determined. A tandem array of variable 17-bp repeats, named R1–7, was found to be responsible for most of the observed length and sequence variation between individuals.     

Spermiogenesis and spermatozoal ultrastructure in Trichomycteridae (Teleostei: Ostariophysi: Siluriformes)

Spadella, M.A., Oliveira, C. and Quagio‐Grassiotto, I. 2009. Spermiogenesis and spermatozoal ultrastructure in Trichomycteridae (Teleostei: Ostariophysi: Siluriformes). —Acta Zoologica (Stockholm) 91 : 373–389. Siluriformes comprises the most diverse and widely distributed ostariophysan group, a fish assemblage that includes about three quarters of the freshwater fish of the world. In this study, the ultrastructural characterization of spermiogenesis and spermatozoa in specimens of Copionodontinae (the sister group to all other trichomycterids), Trichomycterinae (a derived trichomycterid group), and Ituglanis (a genus not assigned to any trichomycterid subfamily) is presented. The comparative analyses of the data show that trichomycterid species share six of seven analyzed spermiogenesis characters, reinforcing the monophyly of the group. Analyses of trichomycterid sperm ultrastructure showed that the species studied share the same character states for nine of seventeen characters analyzed. Copionodon orthiocarinatus and Ituglanis amazonicus each share more ultrastructural characters with species of Trichomycterus than with one another. Regarding the families of Loricarioidea, the species of Trichomycteridae share more characters of spermatogenesis, spermiogenesis, and sperm with representatives of the families Callichthyidae, Loricariidae, and Scoloplacidae than with Nematogenyidae, its hypothesized sister group. With the exception of the family Nematogenyidae, the character similarities observed reinforce the monophyly of the superfamily Loricarioidea.     

Shelter utilization by Gobius cruentatus and Thorogobius ephippiatus (Teleostei: Gobiidae)

Parameters of shelters utilized by Gobius cruentatus and Thorogobius ephippiatus were measured, in situ , using SCUBA. Both species exhibited non-random use of shelters, with T. ephippiatus , the smaller of the two species, utilizing significantly larger shelters than G. cruentatus . Interspecific size differences and competitive exclusion are not considered to be of major importance in the observed differential utilization of shelters. The size of shelters utilized by both species is suggested to be influenced by current velocities.     

Genetic variation and diadromy in some native New Zealand galaxiids (Teleostei: Galaxiidae)

We examined genetic variation at 13 isozyme loci in three diadromous and two non-diadromous species of galaxiid fish from streams in South and Stewart Islands of New Zealand. Analysis of allele frequency data revealed higher F_s7 and Nei's D values among conspecific populations of the non-diadromous species. We propose that the genetic structurings of these species are affected by their migratory behaviours. We suggest that the derived condition of non-diadromous behaviour may increase scope for speciation, but also the probability of extinction. This is exacerbated by population fragmentation over the last century.