Genetic divergence and phylogenetic inferences in five species of Mugilidae (Pisces: Perciformes)

Allozyme electrophoresis was used to compare the genetic divergence of global populations of Mugil cephalus Linnaeus, 1758 and two congeneric [M.␣curema Cuvier & Valenciennes, 1836; M. gyrans (Jordan & Gilbert, 1885)] and two more distantly related [Liza ramada (Risso, 1826); Xenomugil thoburni (Jordan & Starks, 1896)] species on the basis of 18 enzyme loci. The amount of genetic divergence among the species examined is in agreement with their present systematic status, the differences being larger among non-congeneric species than among species of the same genus. Intraspecific genetic distances in M. cephalus (average Nei's D= 0.154), although higher than those reported for conspecific populations of fish, appear to be small when compared to the interspecific values among mugilid species (0.821 ≤ Nei's D ≤ 1.744). Phylogenetic trees obtained by genetic distance methods and discrete character parsimony analysis were of similar topology, except for the relationships within the genus Mugil and for the arrangement of M. cephalus populations. Received: 7 April 1997 / Accepted: 4 February 1998     

Genetic differentiation in two sibling species of the brackish-water polychaete Hediste japonica complex (Nereididae)

Genetic divergence among ten populations of small- and large-egg forms of the brackish-water polychaete Hediste japonica complex was investigated on 14 isozyme loci by electrophoretic analysis. The two forms were distinguishable by complete allele substitutions at five loci, resulting in high genetic differentiation (Nei's D: 0.533 to 0.662). No genetic evidence of hybridization between the two forms was detected in sympatric populations in three rivers. These results indicate that the two forms are reproductively isolated, clearly showing that the two forms are distinct species. The genetic differentiation among populations was higher in the large-egg form (D: 0.005 to 0.111, G _ST: 0.435) than that in the small-egg form (D: 0.000 to 0.001, G _ST: 0.020). This genetic difference between the two forms seems to be attributable to a difference in their life histories. The average expected heterozygosity was low in populations of both the large-egg form (0.005 to 0.068) and the small-egg form (0.014 to 0.038) in comparison with other marine invertebrates. Received: 11 April 1997 / Accepted: 8 September 1997     

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     

Feeding macroecology of territorial damselfishes (Perciformes: Pomacentridae)

The present study provides the first analysis of the feeding macroecology of territorial damselfishes (Perciformes: Pomacentridae), a circumtropical family whose feeding and behavioral activities are important in structuring tropical and subtropical reef benthic communities. The analyses were conducted from data collected by the authors and from the literature. A strong positive correlation was observed between bite rates and sea surface temperature (SST) for the genus Stegastes. A negative correlation was found between bite rates and mean body size for the genera Stegastes and Pomacentrus, but this relationship was not significant when all territorial pomacentrids were analyzed together. A negative correlation between body size and SST was observed for the whole group and for the genera Stegastes, and Pomacentrus. No relationship was found between territory size and feeding rates. Principal Components Analysis showed that differences in feeding rates accounted for most of the variability in the data. It also suggested that body size may be important in characterizing the different genera. In general, tropical species are smaller and have higher bite rates than subtropical ones. This study extended the validity of Bergmann’s rule, which states that larger species or larger individuals within species occur towards higher latitudes and/or lower temperatures, for an important group of reef fishes. The identification of large-scale, robust ecological patterns in the feeding ecology of pomacentrid fishes may establish a foundation for predicting large-scale changes in reef fish assemblages with expected future changes in global SST.     

Temperature tolerances of two southern African Ecklonia species (Alariaceae: Laminariales) and of hybrids between them

The temperature tolerances in culture of Ecklonia maxima (Osbeck) Papenf., the dominant kelp in the Benguela upwelling region on the west coast of southern Africa, and the smaller E. biruncinata (Bory) Papenf., growing on the warmer south coast, have been investigated. Vegetative growth and reproduction of gametophytes show very similar patterns in the two species, although temperature optima in E. biruncinata are consistently 2 ° to 3 °C higher than those of E. maxima. Reciprocal crosses between clonal female and male gametophytes of the two species produced healthy juvenile sporophytes, indistinguishable from the parent crosses. All parents and hybrids grew well over a wide range of temperatures (8 ° to 22°C), with E. biruncinata sporophytes again having slightly higher optima than E. maxima. Both reciprocal crosses exhibited optima intermediate between the parent crosses. The results are discussed with respect to the origins and evolution of Ecklonia species in the region.     

Maturation and diel reproductive periodicity of round scad (Carangidae: Decapterus punctatus)

The diel reproductive periodicity of Carangidae is poorly known but appears to be highly variable between species. Some species spawn during the day, others are believed to spawn at night, and it is demonstrated here that round scad, Decapterus punctatus, spawn at dusk. We collected D. punctatus in the eastern Gulf of Mexico during three April cruises (1995, 1996, and 1997). Based on histological criteria, size at 50% maturity was 113 mm fork length (FL) for males and 128 mm FL for females. The gonad-somatic index (GSI) of mature males was significantly different between hours and appeared to show diel periodicity. Diel periodicity was also observed in changes in female GSIs, whole oocyte diameters, and ovarian histology. The average GSI of mature females fluctuated two-fold between day and night, and the size distribution of whole oocytes in some fish was bimodal (at 0.3-0.4 and 0.7-0.8 mm diameter) at dusk rather than unimodal during most of the diel cycle. Histological preparations revealed that these rapid changes in ovarian GSIs and oocyte size distributions were the result of final oocyte maturation. Germinal vesicle migration was observed from 0900 to 1400 hours eastern standard time (EST), germinal vesicle breakdown was evident as early as 1100 hours EST, and ovulation occurred as early as 1800 hours EST. Spawning frequency (approximately every 5 days) was similar whether calculated from the proportion of females with hydrated oocytes during the afternoon or from the proportion of females with postovulatory follicles during the morning. Batch fecundity correlated with fish size and ranged from 5,500 to 34,700 hydrated eggs per individual. These findings do not support published hypotheses that young-of-the-year D. punctatus reproduce before their first winter or that D. punctatus reproductive output is bimodal within a year.     

Activity budgets in two species of bee flies (Lordotus: Bombyliidae,Diptera): a comparison of species and sexes

Summary This study compares time budgets of males and females of two sympatric species of bee flies, Lordotus pulchrissimus and L. miscellus. These species occur synchronously in sand dune habitat on the north shore of Mono Lake, Mono Co., California, and feed almost exclusively on the flowers of a composite shrub, Chrysothamnus nauseosus. The two closely related species, with similar ecological requirements and in an identical environment, allocate time and energy in very different ways, even when accomplishing the same ecological goals. L. pulchrissimus males and females engage respectively in aggressive interactions and feeding primarily in the morning. In contrast, L. miscellus males and females intersperse brief periods of resting with flying when defending territories and feeding, and they keep up these activities until later in the day. Similarly within species, males are involved in aggressive interactions for a shorter period each day, and females feed over a longer period. Flies in all sex-species classes but male pulchrissimus allocate their time energetically in similar ways. Male pulchrissimus spend more time each day in energetically costly activities; they engage in continuous hovering flight and intense interactions in aerial aggregations. While male miscellus feed little, male pulchrissimus spend a large portion of their time feeding, approximately as much as females, contrary to the expectation that males should be foraging-time minimizers. This study corroborates the conclusions of previous studies on bee-fly communities by showing that nectar and pollen are important resources for adult bee flies, at least for some species: Individuals of these two species spend a large proportion of their adult lifetimes feeding.     

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.     

How wave exposure, group size and habitat complexity influence foraging and population densities in fishes of the genus Halichoeres (Perciformes: Labridae) on tropical rocky shores

Wave exposure and habitat complexity have been used to explain variations in the distribution patterns and behavior of many reef fishes. This study analyzed the influence of both factors on density and foraging activity, and the influence of group size on foraging in three species of the genus Halichoeres (Labridae) on tropical rocky shores. It was shown that initial phases (IP) and terminal phases (TP) Halichoeres, were influenced by wave exposure, although foraging in Halichoeres brasiliensis TP was not influenced by wave exposure. The IP in all three species were positively associated with rugosity and algal cover while the TP was positively associated with depth. Habitat complexity also influences foraging in these species. Group size influenced foraging activity, except in H. brasiliensis TP. We also found variations in microhabitat patches used for foraging between species and differences in the stomach contents between species and phases, showing that functional roles change in response to ontogenetic shifts.     

Ultrastructural aspects of the symbiosis between two species of the genus Aphanocapsa (Cyanophyceae) and Ircinia variabilis (Demospongiae)

The association between two species of the genus Aphanocapsa (Cyanophyceae) and the sponge Ircinia variabilis has been studied by electron microscopy. A. feldmanni is localized in the mesohyl or inside the cells of the sponge, while the larger A. raspaigellae is located only in an extracellular position inside cavities of the mesohyl. Both algae differ from other symbiotic Cyanophyceae in having a normal cell wall. They are able to reproduce in symbiotic condition, but also undergo, in their various extracellular and intracellular positions, a massive process of disintegration. A large amount of algal material is dispersed in the sponge tissues, which is a confirmation, at the ultrastructural level, of trophic relationships in the symbiosis Aphanocapsa-Ircinia.     

Genetic differentiation in three Sepia species (Mollusca: Cephalopoda) from Galician waters (north-west Iberian Peninsula)

The systematics of the genus Sepia is not yet clear. Morphological evidence has led to S. officinalis Linnaeus, 1758 being considered as belonging to the subgenus Sepia sensu stricto, and S. orbignyana Férussac, 1826 and S. elegans Blainville, 1827 as belonging to the subgenus Rhombosepion. Samples of 30 individuals of S. officinalis and S. orbignyana from both sides of an oceanographic boundary off the north-west Iberian Peninsula, and a sample of S. elegans from the northern side, were collected in 1993–1994. Allozyme electrophoresis for 32 presumptive loci revealed low levels of genetic variability for the three Sepia species (mean expected heterozygosity estimates were <0.052). No significant differences in allozyme frequencies were detected among populations of either S. officinalis or S. orbignyana. The genetic identities (I) of S. officinalis and S. orbignyana (I=0.12) and of S. elegans (I=0.13) were significantly different from that of S. orbignyana and S. elegans (I=0.49). The former are typical of values for confamilial genera, and a new generic status is proposed for the latter two species, which become Rhombosepion orbignyana (Férussac, 1826) and R. elegans (Blainville, 1827).     

Ecology of two littoral species of caprellid amphipods (Crustacea) from Washington,USA

Ecology of 2 littoral species of caprellid amphipods is compared. Populations of Caprella laeviuscula, a periphyton scraper/filter-feeder, are most dense on eelgrass Zostera marina L. In the absence of C. laeviuscula, periphyton biomass increases 411% in protected Z. marina beds. The light absorption spectra of periphyton and Z. marina are similar, and C. laeviuscula, by its periphyton removal, may allow Z. marina to grow in areas where it would otherwise be excluded. C. laeviuscula is aggressively dominant over sympatric caprellids, and seasonal predators of C. laeviuscula are absent during winter, the period when periphyton would be most limiting to Z. marina. Populations of Deutella californica, a predator, are most dense on the hydroid Obelia dichotoma (L.). In the O. dichotoma epibiotic community, D. californica is the primary predator, but removal did not change the composition of the community structure. In the absence of other macropredators, the structure of the O. dichotoma epibiotic community depends more on the seasonality of O. dichotoma occurrence than on organismal interactions.     

Mate choice,fecundity and sexual dimorphism in two pipefish species (Syngnathidae)

Summary In order to understand the causes of sexual dimorphism, mate choice and size-related fecundity were studied in two pipefish species, Syngnathus typhle and Nerophis ophidion. Sexual dimorphism is more pronounced in N. ophidion; females are larger, have sexual colourings, and are more active during courtship. In S. typhle the sexes are alike in all these respects. Males brood their offspring in both species. In N. ophidion fecundity was positively correlated with both body size and the amount of sexual colouring in females. In males no correlation between body size and fecundity, or between body size and embryo size existed. Predictably, in mate choice experiments with equal-sized females, males chose females with more extensive sexual colourings. We explain sexual dimorphism in this species as a consequence of both natural selection (fecundity increases with size in females but not in males) and sexual selection (males prefer larger females). We argue that sexual size dimorphism did not evolve by selection minimizing overlap in food niches between the sexes, because food production is high in the Zostera beds where the fishes live, and no size dimorphism was found in the sympatrically occurring S. typhle. Furthermore, in N. ophidion dimorphism is not greater in a particular mouth character than in overall body size. In S. typhle egg size and the average number of eggs transferred per spawning were positively correlated with female body size. Apparently more energy per offspring was provided by larger males than by smaller males, and larger males also carried more offspring. As predicted, large mates were preferred by both sexes in mate choice experiments. This is explicable in terms of both natural selection (fecundity increases with size in both sexes) and sexual selection (both sexes prefer large mates). As a consequence of selection acting in the same direction in both sexes, sexual dimorphism is absent in S. typhle.     

Lipid compositions,water contents,swimbladder morphologies and buoyancies of nineteen species of midwater fishes (18 myctophids and 1 neoscopelid)

Lipid compositions, water contents, swimbladder morphologies and specific gravities were studied for 19 species of oceanic midwater fishes, chiefly myctophids (lanternfishes) collected offshore from Oregon and California from 1975–1979. Three groups of species were recognizable. The first group had bodies low in both lipid and water content; they were denser than seawater, regardless of swimbladder morphology, which varied from absent through non-inflated to inflated. The second group had bodies low in lipid but high in water content; they were neutrally buoyant, evidently because of their high content of water, although their swimbladders were never inflated. The majority of the members of the aforementioned groups contained higher proportions of triglycerides than wax esters. The third group had bodies with high lipid but low water content; those with high triglyceride content had swimbladders ranging from non-inflated to (less commonly) inflated, were denser than seawater, and the lipid percentage of their body weights varied both with size and season, indicating that triglycerides function mainly as an energy store. The adults of species in the third group with a higher content of wax esters than triglycerides lacked inflated swimbladders, were neutrally buoyant, and the lipid percentage of their body weights was relatively constant with size and season, indicating that wax esters permit these fishes to attain neutral buoyancy in seawater.     

Circadian rhythms in the respiration of eight species of cardinal fishes (Pisces: Apogonidae): comparative analysis and adaptive significance

A comparative study was made of the entrained (12:12 LD) and endogenous (DD) respiratory rhythms of 8 species of cardinal fishes. Volumetric respirometers, designed specifically for long-term measurements of oxygen uptake by small fishes under controlled conditions, were used during standard 5 day experiments. Despite some individual variation, certain species-specific and genus-specific patterns of respiration emerged. Each species showed a persistant (advancing) rhythm of oxygen consumption during 3 days of DD. Light appeared to be the Zeitgeber, with the light-dark transition phasing the nocturnal peaks and the dark-light transition possibly phasing the diurnal ones. Genus-specific and species-specific trends in field behavior, habitat selection, and distribution were studied. There were correlations between the field behavior and the entrained respiratory rhythms of these nocturnally active fishes. Entrained and endogenous rhythms of each species showed characteristic similarities and were interpreted as integral components of a level of organization evolved for nocturnal life in the shallow marine environment.     

Interactions between two introduced species: Zostera japonica (dwarf eelgrass) facilitates itself and reduces condition of Ruditapes philippinarum (Manila clam) on intertidal flats

Dwarf eelgrass (duckgrass; Zostera japonica) and Manila clams (Ruditapes philippinarum) are two introduced species that co-occur on intertidal flats of the northeast Pacific. Through factorial manipulation of clam (0, 62.5, 125 clams m⁻²) and eelgrass density (present, removed by hand, harrowed), we examined intra- and interspecific effects on performance, as well as modification of the physical environment. The presence of eelgrass reduced water flow by up to 40% and was also observed to retain water at low tide, which may ameliorate desiccation and explain why eelgrass grew faster in the presence of conspecifics (positive feedback). Although shell growth of small (20–50 mm) clams was not consistently affected by either treatment in this 2-month experiment, clam condition improved when eelgrass was removed. Reciprocally, clams at aquaculture densities had no effect on eelgrass growth, clam growth and condition, or porewater nutrients. Overall, only Z. japonica demonstrated strong population-level interactions. Interspecific results support an emerging paradigm that invasive marine ecosystem engineers often negatively affect infauna. Positive feedbacks for Z. japonica may characterize its intraspecific effects particularly at the stressful intertidal elevation of this study (+1 m above mean lower low water).     

Predatory behavior and chemical communication in two Metapone species (Hymenoptera:Formicidae)

Summary. Colonies of two species of Metapone (M. madagascarica, M. new species.) were collected in Madagascar and established in laboratory nests. It could be demonstrated that both species are specialist predators of termites (Cryptotermes kirbyi). During hunting the ants sting the termites and thereby paralyze and preserve the prey alive. In this way prey can be stored in the ant nest for extended periods. During foraging and colony emigrations the ants lay chemical trails with poison gland secretions. Among the seven compounds identified in the venom only methyl pyrrole-2-carboxylate elicits trail following behavior in both Metapone species. Received 11 February 2002, accepted 23 February 2002.     

Lipid biochemistry and reproductive biology in two species of Gammaridae (Crustacea: Amphipoda)

Gammarus oceanicus Segerstråle, 1947 and Echinogammarus marinus (Leach, 1815) were sampled during the breeding season from Oslofjord in 1984, and their lipid composition examined in relation to reproductive condition. In G. oceanicus, female lipid content increased as the ovary matured. Both the amount of lipid stored and the rate of accumulation were greater in spring than in winter. Spring eggs contained 12.4 g lipid, of which 63% was triacylglycerol and 27% phospholipid. Both fractions decreased steadily during embryonic development. Winter eggs contained 19.2g lipid, of which 52% was triacylglycerol and 43% phospholipid. During the early stages of embryonic development the amount of phospholipid decreased sharply, whereas that of triacylglycerol increased, suggesting that some of the fatty acid released from phospholipid was sequestered temporarily as triacylglycerol. When newly spawned, both winter and spring eggs were richer in monoenoic fatty acids than adult amphipods and these acids were the major fuel consumed during development. 6 fatty acids were utilised more slowly than 3 acids, and egg carotenoid pigment content remained constant. Female E. marinus increased in lipid content as the ovary matured. Spring eggs contained 14.7 g lipid when newly spawned and this increased to 16.6 g during the early stages of development. This increase was entirely triacylglycerol, which declined in later stages; the source of the extra lipid was unclear. Eggs contained very little phospholipid or sterol, and both of these components remained at a steady low level during development. E. marinus eggs were not significantly rich in thonoenoic acids compared with adults, and saturated, monoenoic and polyenoic acids were utilised about equally during development. Both adults and eggs were rich in 20.46, which was utilised at a slower rate than the 3 polyunsaturated acids during embryonic development; again, egg carotenoid pigment content remained constant. In both species there was a decrease in the size of the egg (and as a result, of the newly hatched juvenile), but an increase in total reproductive output (i.e., the total weight of the egg clutch) per female as the breeding season proceeded. The reproductive output of an individual female is probably related to food availability during the period of ovarian maturation, whereas the size of an individual egg is dictated largely by feeding conditions for the juveniles once they are independent of the female. The different patterns of lipid utilisation during development found in this study emphasize the flexibility of response in the reproductive biology of gammarid amphipods. It is not yet possible, however, to relate the differing patterns in a simple way either to egg size or total female reproductive output. Two outstanding problems are the source of extra triacylglycerol during the early stages of development of E. marinus and the metabolic cost of brooding eggs.