Genetic variation in seven goby species (Perciformes: Gobiidae) assessed by electrophoresis and taxonomic inference

Data on the electrophoretic patterns of 20 enzyme loci were used to investigate relationships among seven Mediterranean goby species. The study evidenced different levels of genetic variability among the species. On the whole, the degrees of genetic differentiation were consistent with the current taxonomic recognition of species and genera. Phenetic and phylogenetic analyses agreed in clustering the samples in three groups, i.e. the Gobius (including Zosterisessor ophiocephalus), the Lesueurigobius species and Pomatoschiarua minutus. The taxa analysed probably evolved following the complex geological and climatic events that have been affecting the Mediterranean basin ever since the Miocene. Received: 20 June 1998 / Accepted: 5 February 1999     

Comparison of gut morphology and gut microflora of seven species of mud shrimp (Crustacea: Decapoda: Thalassinidea)

The gut morphology is described for seven species of Thalassinidea; Callianassasubterranea (Montagu), Jaxea nocturna Nardo, Axius stirhynchus Leach, Calocaris macandreae Bell, Upogebia pusilla (Petagna), U. deltaura (Leach) and U. stellata (Montagu). Differences were observed in the structure of the dorsal and lateral teeth of the gastric mill. Callianassa subterranea and A. stirhynchus had simple robust dorsal teeth and proportionally larger lateral teeth with flexible spines. The three upogebiids had complex dorsal teeth that were highly ridged and proportionally smaller lateral teeth with inflexible spines. The dorsal and lateral teeth of J. nocturna were intermediate in form. The dorsal tooth of Calocaris macandreae, although relatively simple in design, possessed accessory projections, or “wings”, on either side of the main structure. The lateral teeth were proportionally quite large, but lacked ridges and spines. The epithelia surfaces of the digestive gland, midgut and posterior diverticulum were similar for all seven species. Differences were observed in the morphology of the hindgut. The hindgut lining was completely smooth for the three upogebiid species. The other thalassinidean species had four distinct rows of cuticular projections situated on top of the ridges formed by the longitudinal convolutions of the hindgut. The arrangement of these projections varied between species. A gut flora was observed for six of the seven species, the exception being C. macandreae. The differences observed in the structure of the gut are discussed in relation to feeding and thalassinidean phylogeny. Received: 3 July 1998 / Accepted: 8 October 1998     

Cryptic species of Clavelina (Ascidiacea) in two different habitats: harbours and rocky littoral zones in the northwestern Mediterranean

Marinas and harbours provide ideal sites for the study of population genetics of marine invertebrates with restricted dispersal capabilities. They combine a confinement effect, particular ecological conditions (pollution, turbidity), and the possibility of high gene flow through ship-borne propagules, which greatly increases the natural dispersal capability of sexual and asexual propagules in many species with short-lived larvae. We studied the genetic structure of populations of the ascidian Clavelina lepadiformis living inside and outside harbours in the north-western Mediterranean. A 500-bp segment of the cytochrome c oxidase subunit I (COI) mitochondrial gene was sequenced in three populations from inside harbours (interior form) and in three populations from the rocky littoral (exterior form). Two congeneric Mediterranean species, Clavelina sp. and C. dellavallei, were used for comparison. We found that the interior and exterior forms of C. lepadiformis belong to two distinct clades, with a genetic divergence of 5%. Gene-flow values among these forms were insignificant. The lack of gene flow and the genetic divergence suggest that the interior and exterior forms of C. lepadiformis are in fact cryptic species rather than differentiated populations of the same species. Levels of gene flow were higher among interior habitats than among exterior habitats, a pattern likely maintained by genetic exchange through ships. We discuss the possible origins of the present-day distribution of these cryptic species. We contend that the study of species living both inside and outside these particular habitats will reveal more instances of genetic discontinuities allowing local adaptations.     

Phylogeny and biogeography of Sicydiinae (Teleostei: Gobiidae) inferred from mitochondrial and nuclear genes

In the Indo-Pacific area, the Caribbean region and West Africa, insular systems are colonised by particular Gobiids of the Sicydiinae subfamily. These species spawn in freshwater, the free embryos drift downstream to the sea where they undergo a planktonic phase, before returning to rivers to grow and reproduce; an amphidromous lifestyle. These gobies are the biggest contributors to the diversity of fish communities in insular systems and have the highest levels of endemism, yet their phylogeny has not been explored before with molecular data. To understand the phylogeny and the biogeography of this subfamily, sequences from the mitochondrial 16S rDNA and cytochrome oxidase I and from the nuclear rhodopsin gene were obtained for 50 Sicydiinae specimens of seven genera. Our results support the monophyly of the subfamily and of all the genera except Sicyopus, which is polyphyletic. Five major clades were identified within this subfamily. One clade clusters Sicyopterus and Sicydium as sister genera, one contains the genus Stiphodon split into two different groups, two other clades include only Sicyopus (Smilosicyopus) and Cotylopus, respectively, and the last clade groups Akihito, Lentipes and Sicyopus (Sicyopus). As a result, the subgenus Smilosicyopus is elevated herein as a genus. A molecular dating approach helps the interpretation of these phylogenetic results in terms of amphidromy and biogeographical events that have allowed the Sicydiinae to colonise the Indo-Pacific, West African and Caribbean islands.     

Mating system and alternative male mating tactics in the grass goby Zosterisessor ophiocephalus (Teleostei: Gobiidae)

 Peculiar fertilization dynamics, with males releasing sperm in mucous trails lasting several hours, characterize some demersal spawning fish. The mating system was investigated in a natural population of one of these species: the grass goby Zosterisessor ophiocephalus (Pallas, 1814), a large coastal goby inhabiting seagrass meadows in shallow brackish water. Adult males ranged in size from 7.4 to 23 cm total length, but only larger ones were observed to dig and defend a burrow, where they performed parental care on eggs laid by one to several females. Field observations together with analyses of age, sperm production, trail sperm content and sperm competition tests indicated the occurrence of alternative male mating tactics, likely the expression of an ontogenetic gradient. Larger males are older than smaller ones, and while the former are territorial, the latter “sneak” territorial male spawns. The ejaculate characteristics indicate that grass goby males have functionally polymorphic spawns: in fact sperm trails of larger males last longer and release fewer sperm than those of smaller males. Sperm production over several days is more constant in larger than in smaller males, but the total number of sperm released is higher in the latter. The influence of seminal fluid in the functional intraspecific variability in sperm release in this species is discussed. Received: 30 December 1999 / Accepted: 31 July 2000     

Use of space by Gobius cruentatus and Thorogobius ephippiatus (Teleostei: Gobiidae) in south-west Ireland

The mechanisms of spatial organisation exhibited by two co-existing species of Gobiidae, Gobius cruentatus Gmelin, 1789 and Thorogobius ephippiatus (Lowe, 1839) were examined to determine the nature of, and the factors contributing to, these mechanisms. This study was carried out in Lought Hyne, south-west Ireland, during 1990. Both species were found to occupy territories exclusive to conspecifics, whilst G. cruentatus was also found to maintain a territory exclusive to T. ephippiatus. This and other interspecific differences in the spatial patterns exhibited by these species are suggested to be related to size-based dominance abilities and their differential requirements of space-related resources.     

Associative behavior of the fish Cryptocentrus cryptocentrus (Gobiidae) and the pistol shrimp Alpheus djiboutensis (Alpheidae) in artificial burrows

Observations and experiments on the associative behavior of the fish Cryptocentrus cryptocentrus (Valenoiennes) and the shrimp Alpheus djiboutensis De Man from the Red Sea were carried out in artificial burrows. A signal system, bringing about correlated behavior in fish and shrimp, consisted of antennal contact of the shrimp with the fish, tail undulations by the fish and, finally, emergence of the shrimp. The shrimp emerged from the burrow only in the presence of the fish. A constant antennal contact was maintained by the partners while the shrimp was outside the burrow. Emerging from the burrow, the shrimp pushed the fish towards the entrance. Following visual stimulation, the shrimp always retreated back into the burrow with the fish. However, the further back the shrimp was located inside the burrow, the less intense was its retreat with the fish. In the absence of the fish, visual stimulation of the shrimp had no effect. Cleaning of fish by shrimp was observed within the burrow.     

Transient ontogenetic expression of hermaphroditic gonad morphology within the Gobiosoma group of the Neotropical seven-spined gobies (Teleostei: Gobiidae)

The Neotropical seven-spined gobies (tribe Gobiosomatini), including the Gobiosoma and Microgobius groups, constitute a speciose, monophyletic gobiid taxon. In particular, member species of the Gobiosoma group exhibit a combination of behavioral diversification and microhabitat specialization that may have played a major role in their collective rapid evolution and high rate of speciation. Functional hermaphroditism, which is common among gobiids, can promote exploitation of patchily distributed micro-niches by countering potential reductions in reproductive opportunities. However, the possible role of functional hermaphroditism in the exploitation of microhabitats and rapid speciation in the Gobiosoma group is unknown. One member species, Tigrigobius multifasciatus, is a functional hermaphrodite. Two other Gobiosoma group species are functionally gonochoric (i.e., constant-sexed), but exhibit transient hermaphroditic ovarian structure among immatures. In this study, ovarian morphology among immature and adult females of several Gobiosoma group species was examined to see if hermaphroditic gonadal features are present. Although no evidence of functional hermaphroditism in the form of precursive accessory gonadal structures (pAGS) associated with the adult ovary was found among newly examined species, all species exhibited the transient expression of hermaphroditic gonadal features associated with the immature ovary. In contrast, among six species of non-Gobiosomatini genera having no record of hermaphroditism, none exhibited similar transient hermaphroditic features associated with the immature ovary. These findings suggest that hermaphroditism may have been an ancestral trait which has been secondarily lost within some Gobiosoma group species. This study also shows that ontogenetic and morphogenic processes within the gobiid reproductive system may provide new insights into the evolution of life history traits and significantly further our understanding of the extraordinary evolutionary success of this group.     

Analysis of the mutual attraction in the association of the fish Cryptocentrus cryptocentrus (Gobiidae) and the shrimp Alpheus djiboutensis (Alpheidae)

The mutual attraction between the fish Cryptocentrus cryptocentrus and the shrimp Alpheus djiboutensis has been investigated experimentally. The fish is attracted to its partner visually, the shrimp is attracted chemically. The mutual attraction is reinforced by the strong negative phototactic reaction of both partners, orienting them towards the burrow.     

Nucleotide sequencing of highly repetitive DNA from seven species in the coral genusAcropora (Cnidaria: Scleractinia) implies a division contrary to morphological criteria

Nucleotide sequences have been determined for 31 homologous 118 base-pair highly repeated DNA sequences from seven species ofAcropora. A matrix was constructed from the sequence data and subjected to phylogenetic analysis using heuristic search routines in the PAUP (phylogenetic analysis using parsimony) program, Version 3.0L. These analyses confirm a close relationship between two species of one subgeneric group (A. pulchra andA. millepora), but identify a division in a group of six species which is contrary to taxonomic groupings based on morphological criteria.     

Foraging habitats of the seabird community of Europa Island (Mozambique Channel)

We investigated the foraging habitats of the winter breeding community of tropical seabirds from Europa Island (Mozambique Channel) in September 2003. We focused our study on the dominant species of this austral community, the sooty tern (Sterna fuscata), the red-footed booby (Sula sula), and the frigatebirds, including the great (Fregata minor) and the lesser frigatebirds (F. ariel). We considered the at-sea distribution and abundance of these species in relation to chlorophyll concentration, sea-surface temperatures, sea-surface height anomalies, depth of the thermocline, distance to the colony, and presence of surface marine predators, flying fishes and other seabirds. Although the marine environment where seabirds foraged was oligotrophic, it presents the best feeding opportunities for seabirds for the area in winter. Our study demonstrates that the winter-breeding seabird species of Europa Island tend to forage in productive waters in association with other marine predators when possible. Sooty terns and frigatebirds were widely distributed in the whole study area, whereas red-footed boobies were not found farther than 160 km from their colonies and were associated with relatively productive waters. Sooty terns and red-footed boobies were aggregated where flying fishes were abundant. The presence of other marine predators was associated with larger multispecies feeding flocks than when no association occurred. Sooty terns, which are numerically dominant at Europa and adopted network foraging, seem to be catalysts of feeding events, and represented a good target for the other foraging species, especially frigatebirds. However, when possible, frigatebirds favour association with flocks of red-footed boobies.Communicated by S.A. Poulet, Roscoff     

Larval durations and recruitment patterns of two Caribbean gobies (Gobiidae): contrasting early life histories in demersal spawners

The population dynamics of small tiger prawns (Penaeus esculentus and P. semisulcatus) were studied at three sites around north-western Groote Eylandt, Gulf of Carpentaria, Australia, between August 1983 and August 1984. Seagrasses typical of open-coastline, reef-flat and river-mouth communities were found in the shallow depths (2.5 m) at these sites. The temperature and salinity of the bottom waters did not differ among the shallowest depths of the three sites and mean values at night ranged from 21.9 to 32.0 °C, and from 30.1 to 37.5% S. Data from fortnightly sampling with beam trawls showed that virtually all post-larvae (90%) were caught in the intertidal and shallow subtidal waters (2.0 m deep). At one site, where the relationship between seagrass biomass, catches and depth could be studied in detail, high catches were confined to seagrass in shallow water, within 200 m of the high-water mark. This was despite the fact that seagrass beds of high biomass (>100 g m^-2 between August and February) were found nearby, in only slightly deeper water (2.5 m). It is likely, therefore, that only the seagrass beds in shallow waters of the Gulf of Carpentaria act as important settlement and nursery areas for tiger prawns. In general, catches of tiger prawn postlarvae (both P. esculentus and P. semisulcatus) and juvenile P. esculentus on the seagrass in the shallowest waters at each site were higher in the tropical prewet (October–December) and wet (January–March) seasons than at other times of the year. Juvenile P. semisulcatus catches were highest in the pre-wet season. While seasonal differences accounted for the highest proportion of variation in catches of tiger prawn postlarvae and juvenile P. semisulcatus, site was the most important factor for juvenile P. esculentus. In each season, catches of juvenile P. esculentus were highest in the shallow, open-coastline seagrass, where the biomass of seagrass was highest. The fact that the type of seagrass community appears to be more important to juvenile P. esculentus than to postlarvae, suggests that characteristics of the seagrass community may affect the survival or emigration of postlarval tiger prawns. Few prawns (<10%) from the seagrass communities in shallow waters exceeded 10.5 mm in carapace length. Despite the intensive sampling, growth was difficult to estimate because postlarvae recruited to the seagrass beds over a long period, and the residence times of juveniles in the sampling area were relatively short (8 wk).     

Sexual reproduction in Acropora (Isopora) species (Coelenterata: Scleractinia)

Despite the wide range of morphological diversity among the Acropora (Isopora) colonies on Heron Island (Great Barrier Reef, Australia), only two reproductively isolated species were present from 1977 to 1982: A. cuneata (Dana, 1846) and A. palifera (Lamarck, 1816). Both species released planulae lacking zooxanthellae and were simultaneous hermaphrodites with testes and ovaries occurring on separate mesenteries within the same polyp. Oogenesis preceded spermatogenesis. Seasonal cycles of gametogenesis, embryogenesis and planulation occurred in the two species. Colonies of A. cuneata developed two cycles of gametes. One cycle matured near the first quarter moon in April and the other on the same lunar phase in June. Planulae release occurred from about September to December each year and was not correlated with lunar phase. Gametes of A. palifera ripened only once per annum, a few days after the last quarter moon in November, and planulation occurred from about January to March. Embryos were brooded in the coelenteron of the polyps in both species. Ova were fertilized in the mesenteries and embryos were retained within an envelope of mesoglea and gastrodermis, remaining attached to the mesentery by a stalk until the larvae matured and were released. A. palifera and A. cuneata were abundant in the unpredictable reef flat environment. However, their life-history traits, e.g. seasonal reproduction, delayed sexual maturity, large colony size and fairly long life span, were more specialised than had been predicted for this type of environment.     

Genetic adaptation to local habitats and speciation processes within the genus Gammarus (Amphipoda: Crustacea)

A recent event of allopatric speciation (a time span of about 4 000 years) is reported from a comparison between two guilds of amphipod gammarids from the central Baltic Sea and from the Limfjord (inner Danish waters), respectively. For at least two of the five species populations involved, a genetically fixed habitat specialization has led to the establishment of reproductive barriers. It has, however, not been possible to correlate this speciation event with electrophoretically determined isozyme changes. It is argued that the lack of correlation is due to the presence of an initial time lag before a speciation event can be detected as isozyme differences between newly formed species populations as predicted by the neutrality hypothesis for evolution at the molecular level.     

Invertebrate introductions in marine habitats: two species of hydromedusae (Cnidaria) native to the Black Sea,Maeotias inexspectata and Blackfordia virginica,invade San Francisco Bay

The hydrozoans Maeotias inexspectata Ostroumoff, 1896 and Blackfordia virginica Mayer, 1910, believed to be native to the Black Sea (i.e. Sarmatic) and resident in a variety of estuarine habitats worldwide, were found as introduced species in the Petaluma River and Napa River, California, in 1992 and 1993. These rivers are mostly-estuarine tributaries that flow into north San Francisco Bay. Both species appeared to be well-established in this brackishwater habitat. Salinities at the collection sites were about 11 during the summer, rising to nearly 20 in the early autumn and falling to near 0 in the winter. Large numbers of all sizes of both species of medusae were observed and collected, indicating that the hydroid stages of the life cycles of the two are also well-established in these rivers. In the Petaluma River, populations of both species were at maximum in late July, with numbers of individuals declining through August and into September; the Napa River was sampled only in October, and at that time only B. virginica was found. Examination of full guts of M. inexspectata and B. virginica medusae revealed that both species had fed nearly exclusively on small crustaceans, principally barnacle nauplii, copepods and their eggs and nauplii, and crab zoea larvae (M. inexspectata only). All the M. inexspectata medusae were males, indicating that the population has probably developed from the introduction of perhaps only a single male polyp or polyp bud. In spite of its inability to reproduce sexually, this population appears to be maintained by the prodigious ability of the polyp to bud and reproduce asexually, and is fully capable of invading additional low-salinity habitats from its present Petaluma River site. Male and female B. virginica medusae were collected in both the Petaluma River and the Napa River, indicating that B. virginica may have been introduced by either the polyp or medusa stage (or both), but that multiple individuals (of both sexes) must have arrived from another port in one or more invasions. As indicated for M. inexspectata, the B. virginica population will also probably seed new populations in San Francisco Bay and elsewhere. Based on its cnidome as well as the morphology of both medusa and polyp, M. inexspectata has been reclassified by moving it from the family Olindiidae, Limnomedusae, to the family Moerisiidae, Anthomedusae.     

Karyotypes in two species of the genusHippocampus (Pisces: Syngnatiformes)

Diploid modal numbers of chromosomes 2n=48 forHippocampus ramulosus and 2n=44 forH. hippocampus, collected in the Gulf of Palermo in May 1985, are established. No sex-linked heteromorphic pairs are observed in the two species analyzed. The karyotype ofH. hippocampus seems to be a derived condition.     

Pleistocene diversification of the Pomacentrus coelestis species complex (Pisces: Pomacentridae): historical biogeography and species boundaries

Pleistocene eustatic changes in sea level are often invoked to explain genetic divergence among marine organisms. However, molecular phylogenies have revealed relatively few examples of speciation events dating to the Pleistocene. We present a species-level hypothesis of the timing of evolution for the Pomacentrus coelestis species complex (Pomacentridae), based on the nuclear S7 intron and the mitochondrial Cytb gene, and reconstruct ancestral range distributions across the timetree. Ancestral range reconstruction suggests the complex originated in the Coral Triangle and East Indian Ocean, with subsequent range expansion outward from this region. We suggest that land barriers to dispersal (e.g., Indo-Pacific barrier) may be responsible for the divergence between Indian (P. alleni, P. similis, P. caeruleopunctatus, and P. caeruleus) and Pacific (P. micronesicus, P. auriventris, and P. coelestis) species groups, and subsequent isolation by Pleistocene sea-level fluctuations in certain areas of the Coral Triangle (glacial refugia) may play an important role in the diversification of this species complex. Additionally, our analyses show cryptic lineages within P. micronesicus and highlight the need for comprehensive sampling within and among species to reveal recent speciation events.     

Phylogenetic relationships of coral-associated gobies (Teleostei,Gobiidae) from the Red Sea based on mitochondrial DNA data

Bryaninops, Gobiodon, Paragobiodon and Pleurosicya are the most abundant genera of coral-associated gobies. These genera are adapted to live among coral, while other small reef gobies (e.g., the genus Eviota) show no obligate association with this living substrate. Thirteen coral-associated species and two Eviota species were sampled from different regions of the Red Sea, along with four populations/species of Gobiodon from the Indian and western Pacific Oceans. A molecular phylogenetic analysis was performed using partial sequences of 12S rRNA, 16S rRNA and cytochrome b mitochondrial genes, 1,199 base pairs in total. Several clades were consistently resolved in neighbor joining-, maximum parsimony-, maximum likelihood and Bayesian analyses. While each of the four genera Gobiodon, Paragobiodon, Bryaninops and Pleurosicya proved to be monophyletic, their relative position in the phylogeny did not support an emergence of coral-associated gobiids as a monophyletic assemblage. Instead, two separate monophyletic sub-groups were discovered, the first comprising Gobiodon and Paragobiodon, and the second Bryaninops and Pleurosicya. Our molecular phylogenetic examinations also revealed one unassigned species of Gobiodon from the Maldives as a distinct species and confirmed three putative and yet unassigned species from the Red Sea. Moreover, the uniformly black colored species of Gobiodon are not monophyletic but have evolved independently within two distinct species groups. Genetic distances were large in particular within Pleurosicya and Eviota. Estimated divergence times suggest that coral-associated gobies have diversified in parallel to their preferred host corals. In particular, divergence times of Gobiodon species closely match those estimated for their typical host coral genus Acropora.     

Validation of daily increment deposition in otoliths. Age and growth determination of Aphia minuta (Pisces: Gobiidae) from the northwest Mediterranean

The transparent goby Aphia minuta (Risso, 1810) is one of the main target species of the small-scale fishery off the Island of Majorca. Otolith microstructure and length-frequency analysis were used to study the age and growth of this species during the 1982/1983 and 1992/1993 fishing seasons. Daily periodicity of increment formation was determined by experiments with marked otoliths in individuals maintained in captivity. The length range of the catches during the 11 yr period was between 12 and 49 mm, with a main distribution (89%) between 24 and 40 mm. Otolith age-readings indicate that the population exploited in the commercial fishery consists of seven age-groups (2 to 8 mo old), with a very high proportion of individuals (95%) between 3 and 6 mo old. Population growth-curves revealed no differences between males and females. The growth parameters for the whole population are: asymptotic length, L _∞ = 53.69 mm; growth coefficient, K = 2.23 yr⁻¹; theoretical age at length zero, t ₀ = −0.005 yr. Those individuals of A. minuta caught in Majorca during the winter period reached a maximum age of 7 or 8 mo. Received: 30 December 1996 / Accepted: 16 April 1997