Isozyme genetics and the phytogeny of Italian freshwater gobies (Teleostei: Gobioidei)

Although predominantly a marine group in the Atlantic/Mediterranean fish fauna, three species of exclusively freshwater gobies (Teleostei: Gobiidae) occur in Italy. These are Padogobius marltmsii of north Italy and parts of Croatia, P. nigricans from the Tyrrhenian catchment area of Tuscany and Lazio, and Knipowitschia punctatissima from north-eastern Italy. Using horizontal starch-gel electrophoresis, isozyme polymorphism in 12 enzymes encoded at 18 loci was investigated in these species and related euryhaline forms. Genetic distance, and morphological cladistic analysis, indicates that the Italian freshwater gobies do not form a monophyletic group, with the Padogobius species well separated from other taxa, and K. punctalissima closest to its morphological congener, the lagoonal K. panizzae . No close correlation was found between genetic variation and environmental heterogeneity.     

Encephalization in Gobioidei (Teleostei)

We have measured the brain and body weight and determined the encephalization index for 180 species of fishes belonging to six families of the suborder Gobioidei. Within the Teleostei, these fishes exhibit a remarkably broad range in the values of their encephalization indices, but most values are in the low to middle range. Within the Gobioidei there is relatively little difference in the degree of encephalization among the different families and subfamilies except the Kraemeriidae and Amblyopinae which have low encephalization indices and the Oxudercinae (including Periophthalmus) and Rhyacichthyidae which are highly encephalized. We have shown that the form of the body has an effect on the degree of encephalization. Elongate fishes have low values, probably because of the excessive mass of their body skeleton which raises the body weight relative to the brain size. The environment in which the fishes live is correlated, in general, with their relative brain size. The values of the encephalization index arranged from low to high by habitat are as follows: muddwelling fishes, freshwater fishes, brackishwater fishes, burrowing marine fishes, freeliving marine fishes, torrent fishes and amphibious fishes. The low values of the Amblyopinae and Kraemeriidae can be explained in terms of their being both mud-dwelling and elongate.     

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.     

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.     

Patterns of gonad structure in hermaphroditic gobies (Teleostei Gobiidae)

Synopsis Hermaphroditism has been reported for a small number of gobiid fishes, but the extent of this sexual pattern within the family is not known. Gonad structure was examined in one or more species from twenty-one gobiid genera. No evidence of hermaphroditism was found in the species selected from 14 genera. Laboratory studies supported the conclusion of gonochorism for the examined species in four of them:Asterropteryx, Bathygobius, Gnatholepis, andPsilogobius. Currently, the absence of precursive testicular tissues associated with the ovary in females, in conjunction with no retained ovarian features in the testes of males, appear to be reliable indicators of a gonochoristic sexual pattern in gobiid fishes. Evidence for hermaphroditism was observed in seven genera:Eviota, Trimma, Fusigobius, Lophogobius, Priolepis, Gobiodon, andParagobiodon. Protogyny was experimentally confirmed inE. epiphanes, and the gonad structure in another nine of ten species ofEviota suggested either protogyny or protogynous tendencies. With the exception ofGobiodon andParagobiodon, which exhibited similar gonadal structure, ovarian and testicular structure varied considerably among the hermaphroditic genera examined, both with regard to the configuration and to the degree of development of ovarian and testicular tissues, or testicular tissue precursors. Findings of this study indicate that hermaphroditic gonad structure will prove to be a useful trait in determining evolutionary relationships within the Gobiidae.     

The species of Pseudaphya (Teleostei: Gobiidae) and the evolution of aphyiine gobies

The Mediterranean gobiid genus Pseudaphya Iljin, 1930, formerly containing two nominal species, the type Ps. ferreri (O. De Buen & Page, 1908) and Ps. pelagica De Buen, 1931, is now believed to be monotypic after examination of Adriatic material originally identified as Gobius pusillus Canestrini, 1861. The last is here provisionally regarded as Pomato-schistus knerii (Steindachner, 1861). The genus Pseudaphya is redefined, with additional osteological features, and its single species, Ps. ferreri , redescribed. In comparison with other small transparent gobiids, Pseudaphya is shown to possess certain primitive features and a possible mode of evolution for this group of tiny gobies by neoteny from postlarvae of hill-stream sicydiines is discussed. An origin from less specialized gobiines seems more acceptable.     

A unique teleost dermal organ in a new European genus of fishes (Teleostei: Gobioidei)

The freshwater fish fauna of Greece is a diverse assemblage of mediterranean, Ponto-Caspian, and endemic western Greek elements. In the last category, a gobiid from Epirus has been found to possess a dermal perianal organ unique among teleosts, as well as systematic features warranting recognition of a new genus, Economidichthys gen. nov. ( type-species Gobius pygmaeus Holly, 1929). The perianal organ is a sharply-demarcated, pitted area on the rear abdomen. Light-and electron-microscopy has revealed numerous specialized microvillous cells lining the pits and evidently secretory in function. Presence of the organ in both sexes, its ventral position and the secretory activity originally suggested an attachment role rather than involvement in reproduction, but obvious correlation with habitat is lacking and alternative functions are discussed.     

A redescription of Pomatoschistm norvegicus (Collett, 1903) (Teleostei: Gobioidei) based on syntype material

A deeper water species of the Mediterranean-Atlantic gobiid genus Pomatoschistus, P. norvegicus (Collett, 1903), is redescribed from syntype material, including details of the lateral-line system and a summary of geographical distribution. The systematic status of P. norvegicus is briefly discussed with reference to the other nominal species, P. minutus (Pallas) and P. lozanoi (De Buen), of the 'P. minutus' complex.     

Gobies are deeply divided: phylogenetic evidence from nuclear DNA (Teleostei: Gobioidei: Gobiidae)

Gobies (Gobiidae sensu Gill & Mooi, 2012 Gill, A. and Mooi, R. 2012. Thalasseleotrididae, new family of marine gobioid fishes from New Zealand and temperate Australia, with a revised definition of its sister taxon, the Gobiidae (Teleostei: Acanthomorpha). Zootaxa, 3266: 41–52.  [Google Scholar]) are one of the most diverse families of vertebrates, and comprise over 1700 species of marine, brackish and freshwater fishes. Phylogenetic studies based on morphological characters and mtDNA have suggested that goby diversity is asymmetrically split between a speciose clade of predominantly marine species, and a less rich, but ecologically diverse, clade comprising predominantly freshwater and brackish species. This study is the first to explore this deep divide in gobies and their relationships at the family level using phylogenetic data from nuclear genes (RAG1, rhodopsin). Our results confirm the split within the Gobiidae, and agree with prior molecular studies on the inclusion of the following taxa within the two goby clades: (i) the more diverse of the two clades of gobies (the ‘Gobiidae’ sensu stricto of Thacker 2009 Thacker, C. E. 2009. Phylogeny of Gobioidei and placement within Acanthomorpha, with a new classification and investigation of diversification and character evolution. Copeia, 1: 93–104. doi:10.1643/CI-08-004[Crossref], [Web of Science ®] , [Google Scholar]) comprises the gobiines, microdesmines, ptereleotrines and kraemeriines; (ii) the less diverse of the two gobiid clades (‘Gobionellidae’ sensu Thacker 2009 Thacker, C. E. 2009. Phylogeny of Gobioidei and placement within Acanthomorpha, with a new classification and investigation of diversification and character evolution. Copeia, 1: 93–104. doi:10.1643/CI-08-004[Crossref], [Web of Science ®] , [Google Scholar]) includes the gobionellines, oxudercines, amblyopines, sicydiines, as well as the European sand gobies. Some relationships within the two major gobiid clades remain unclear. Specifically, there remains confusion regarding the monophyly and interrelationships between the northern Pacific gobionellines, the Mugilogobius group gobionellines, and the European sand gobies. Additionally, within Thacker's (2009 Thacker, C. E. 2009. Phylogeny of Gobioidei and placement within Acanthomorpha, with a new classification and investigation of diversification and character evolution. Copeia, 1: 93–104. doi:10.1643/CI-08-004[Crossref], [Web of Science ®] , [Google Scholar]) Gobiidae sensu stricto, there are several well-supported groups (e.g. the wormfishes and dartfishes, the Coral Gobies, the Gobiosomatini), yet relationships among these groups are still poorly resolved despite the use of data from two conserved nuclear genes. Future phylogenetic analyses of gobies will benefit greatly from taxon sampling that includes groups that have been historically under-represented in molecular studies (e.g. European sand gobies, northern Pacific gobionellines, African species), as well as deeper genetic sampling including large numbers of independent loci from throughout the genome (i.e. a phylogenomic approach).     

Spatial and temporal patterns of abundance of coral reef gobies (Teleostei: Gobiidae)

This study investigated the spatial and temporal patterns of abundance of four substratum-associated species of Gobiidae on a heterogeneous reef flat comprised of four distinct habitat zones, and examined microhabitat use within each zone. Asterropteryx semipunctatus had the widest distribution and was the most abundant species in each habitat zone, followed by Amblygobius bynoensis , Valenciennea muralis and Amblygobius phalaena . Significant temporal and spatial differences in mean density were evident. The highest density of A. semipunctatus (312 individuals 10 m^–2) was recorded in a habitat zone dominated by algal-covered rubble, whereas A. bynoensis and V. muralis were most abundant (mean summer density 5·5–5·8 individuals 10 m^–2) in habitats containing both sand and hard substrata. In contrast, A. phalaena was uncommon (mean density ≤ 0·4 individuals 10 m^–2) in all four habitat zones. Significant seasonal differences in abundance were due to the large influx of recruits in summer. Ontogenetic shifts in habitat use were not evident at either the macrohabitat ( i.e. among habitat zones) or microhabitat scale ( i.e. substratum use within zones). At the microhabitat scale, V. muralis consistently exhibited a strong positive association with sand and was rarely associated with hard substrata. In contrast, the two Amblygobius species were commonly associated with both sand and hard substrata, but patterns of microhabitat use differed among habitat zones. Substratum composition at the microhabitat scale may influence spatial patterns of abundance at larger spatial scales by providing essential resources and, or influencing carrying capacity and predation risk.     

Molecular systematics of the Awaous banana complex (River gobies; Teleostei: Oxudercidae)

Diadromous fishes can exhibit interesting evolutionary and population-level patterns given their use of freshwater and marine environments as part of their life histories. The River goby genus Awaous are prominent members of riverine ichthyofaunas and occur throughout Atlantic and Pacific slopes of the Americas from the southern United States to Ecuador and Brazil. Here we study the widespread and polymorphic Awaous banana complex to assess phylogeographic patterns and test previous hypotheses that all populations of this species in the Americas belong to the same species. Analysis of sequence data based on the mitochondrial cytochrome oxidase I gene shows multiple clades within the Atlantic and Pacific basins, which correspond to previously described species. Additionally, haplotype analysis demonstrates unique and unconnected networks between these species. Within these clades we document biogeographic patterns that are congruent with results of other co-occurring diadromous species, as well as a novel biogeographic pattern for the region. Our results support the recognition of distinct species of Awaous in the Atlantic (A. banana and A. tajasica) and Pacific (A. transandeanus) basins. These results are concordant with previously established morphological characters permitting the separation of these species.     

Innervation of the lateral line system in Rhyacichthys aspro: the origin of superficial neuromast rows in gobioids (Perciformes: Rhyacichthyidae)

The lateral line system and its innervation were examined in the most primitive gobioid taxon, Rhyacichthys aspro (Rhyacichthyidae). The infraorbital canal was present, whereas superficial neuromast rows a and c, typically present on the cheek of gobioids, were absent. Because the infraorbital canal (absent in other gobioids) and the two rows were commonly innervated by the buccal ramus, the latter were categorized as replaced rows from canal neuromasts. On an innervation basis, rows b and d on the cheek were considered to comprise superficial neuromasts only in all gobioids. The trunk lateral line system comprised canal and superficial neuromasts, the former being included in the lateral line scales (each bearing 1–7 neuromasts arranged longitudinally along the direction of a groove). Absence of bony roofs in the lateral line system was proposed as a synapomorphy of Gobioidei, and a progressive neotenic shift in the lateral line system of the suborder discussed.     

The influence of mating system on seminal vesicle variability among gobies (Teleostei, Gobiidae)

A variety of sexual selection mechanisms have been implicated to drive the variability of the male reproductive tract in internal fertilizers, while studies on external fertilizers have been largely limited to exploring the influence of sperm competition on testis size and sperm number. Males in the Gobiidae, a speciose teleost family of demersal spawners with external fertilization, are known to be characterized by accessory structures to the sperm duct called seminal vesicles. These seminal vesicles secrete a mucus-enriched seminal fluid. Seminal vesicle size and function have been demonstrated to be influenced by sperm competition at the intraspecific level. With the aim to test the factors influencing the development of these male organs at the interspecific level, an independent contrast analysis was performed on 12 species, differing in mating system type, sperm competition risk, and duration of egg deposition. The type of mating system appears to be the main factor significantly affecting development of seminal vesicles, with males of monogamous species completely lacking or having extremely reduced organs.     

Unusual cell organelles during spermiogenesis in two species of gobies (Gobiidae,Teleostei)

Summary Testes of Lesuerigobius friesii and Gobius bucchichi were studied in adult reproductive fish. During the onset of spermatid development, a peculiar system of alternating rough (RER) and smooth (SER) endoplasmic reticular tubules form rings distally to the cell nucleus. The RER tubules are seen to possess up to 12 ribosomes in cross-section, whereas the SER are strongly electron-dense. Nanotubules connect these stacks of tubules to the developing head and tail of the sperm. With ripening of the sperm these tubules disintegrate within the excessive cytoplasm. It seems likely that these are special forms of Macro-Golgi System that possibly provide protamines for the developing sperm.     

Reproductive effort and efficiency in the female common goby, Pomatoschistus microps (Krøyer) (Teleostei: Gobioidei)

Reproductive effort and efficiency are terms which together help to describe the reproductive investment of spawning fish. In this paper they are applied to a small and abundant inshore fish, the common goby, Pomatoschistus microps (Kroyer), based on data collected from the wild, and by modelling likely breeding patterns. Wet calorimetry was used to determine the energy content offish tissue, and this allowed reproductive effort to be expressed as both an energy and a weight ratio of gonad to somatic body. The gonadosomatic index (GSI) calculated as an energy ratio was greater than that calculated as a dry "weight ratio". For ripe females the GSI (dry weight) at the onset of breeding was 67.5%. Theoretical and observed values of the maximum number of egg batches laid by a single female during a 16-week breeding season are 9-10, and this corresponds to a total caloric investment of almost 12 kJ. Estimates of the reproductive efficiency, using the dietary energy content during the breeding season, range from 28.2% to a maximum of 44.0%. These values are among the highest calculated for a fish species, and are in keeping with the long and intensive breeding strategy observed in the wild population.     

Species and shape diversification are inversely correlated among gobies and cardinalfishes (Teleostei: Gobiiformes)

Gobies and their relatives are significant components of nearshore marine, estuarine, and freshwater fish faunas in both tropical and temperate habitats worldwide. They are remarkable for their ability to adapt to and diversify in a wide range of environments. Among gobiiform clades, species diversities vary widely, ranging from two species in Kurtidae to more than 1,000 species in Gobiidae. There is also great variation in head and body shape and in environmental preferences (fresh, brackish, or marine habitats). In this study, I used a time-calibrated molecular phylogeny, coupled with morphometric and comparative analyses, to examine evolutionary rates of both speciation and morphological diversification among gobiiform lineages. Projection of the phylogeny onto a shape-derived morphospace shows that Gobioidei is morphometrically distinct from its sister taxon Apogonoidei, but that families within Gobioidei overlap in morphospace. Analysis of species diversification rates indicates that three rate shifts have occurred over the evolutionary history of Gobiiformes. Relative to the other lineages, Kurtidae has exhibited a slowdown in speciation, whereas both Apogonidae and Gobiidae?+?Gobionellidae have experienced an increase in diversification. Comparative analyses show that in Apogonidae and Gobiidae?+?Gobionellidae, increased speciation is correlated with diminished rates of morphological diversification, differently manifested in either clade and among the various sublineages. The elevation in speciation rates and diminishment in rates of morphological change in both Apogonidae and the clade Gobiidae?+?Gobionellidae are correlated with shifts to oceanic habitats from freshwater. This pattern is the complement to that seen across the global radiation of acanthomorph fishes in which a decrease in species diversification is associated with an increase in morphological disparity.     

The importance of gobies (Gobiidae,Teleostei) as hosts and transmitters of parasites in the SW Baltic

The parasite fauna of five goby species (Gobiidae, Teleostei) was investigated in the Baltic Sea during the period 1987 to 1990. 13 parasite species were found in samples from the Lübeck Bight:Bothriocephalus scorpii, Schistocephalus sp. (Cestoda);Cryptocotyle concavum, Cryptocotyle lingua, Podocotyle atomon, Derogenes varicus (Digenea);Hysterothylacium sp. (cf.auctum),Contracaecum sp.,Anisakis simplex (Nematoda);Corynosoma sp.,Echinorhynchus gadi, Neoechinorhynchus rutili, Pomphorhynchus laevis (Acanthocephala). The number of parasite species were: 10 in the sand gobyPomatoschistus minutus, 8 in the black gobyGobius niger, 7 in the two-spotted gobyGobiusculus flavescens, 6 in the common gobyPomatoschistus microps, and 5 in the painted gobyPomatoschistus pictus. Neoechinorhynchus rutili occurred only inP. minutus, andCorynosoma sp. only inG. niger. The extent to which the gobies were parasitized clearly depended on the respective ways of life and, moreover, on the kind of prey ingested by the hosts. Additionally, the age of the hosts might be important. The highest rate of parasitism, more than 60%, was reached byHysterothylacium sp. inG. niger and byCryptocotyle concavum inP. microps. Infestation incidence lay mostly below 40% which means a satellite species status (Holmes, 1991). The number of parasite species was highest in summer; the highest intensities of single parasites occurred in spring (Podocotyle atomon) or autumn (Crytocotyle concavum).Bothriocephalus scorpii, Hysterothylacium sp. andPodocotyle infested their juvenile hosts very early, but onlyHysterothylacium was accumulated byG. niger during its whole life span, whereasBothriocephalus persisted also in older gobies in low intensities. The cercariae ofCryptocotyle spp. penetrate actively into their hosts; all the other parasites named were transmitted in larval form by prey organisms which consisted mainly of planktonic and benthic crustaceans. The gobies were final hosts for only 5 parasites; but two species may be transmitted to larger fish, and 6 species to sea birds or mammals. The parasite community of the five gobies may possibly be taken to characterize the ecological quality of the environment of the Lübeck Bight.     

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.