Estimation of age in the sex-changing, coral-inhabiting snail Coralliophila violacea from the growth striae on opercula and a mark–recapture experiment

Change of sex in the coral-inhabiting snail Coralliophila violacea (Lamarck) may occur in a wide range of sizes in the field. One proposed explanation for this is that the snails change their sex at a certain age and that individuals have different growth rates caused by microhabitat differences. In this study, we attempt to establish a method to determine the age of this snail and age at sex change. The growth striae on the operculum were studied and compared to the age estimated by the Gompertz growth function based on growth data obtained from mark-recapture experiments in southern Taiwan. There is a significant correlation between the number of striae on the operculum and the age estimated from the Gompertz growth function, and the relationship is 1:1. These results suggest that the number of striae on the operculum can be used as an age index, with each stria representing 1 year of age. The age of sex change of this snail, according to our estimates by both stria number and aperture-length inferences, occurs between 4 and 6 years old. Growth rates of the snails are negatively correlated to size. Furthermore, individuals undergoing sex-change grow faster than males and females.     

Intraspecific density mediates sex-change in the territorial patellacean limpet Lottia gigantea

The effects of intraspecific density and agonistic interactions on sex-change were studied in the territorial limpet Lottia gigantea. In a one-year field experiment (1982–1983) on San Nicolas Island off the southern California coast, USA, male limpets transplanted to large enclosures changed sex more frequently than those transplanted to small enclosures (9 of 13 vs 1 of 10; p=0.013), indicating that intraspecific density can profoundly influence the probability of sex change. Large limpets were more likely to change sex than small ones. Observations of gender-age distributions as well as field behavior suggested that each limpet's territorial status prior to the experiment may have been an important component of this size effect, although other interpretations including an effect of age are possible. Pooling the results with those of two previous studies confirmed that sex-change is enhanced by low density. This enhancement was observed among the largest members of a local population in the first year of each experiment, while among the smaller members the enhancement was delayed until the second or third year. Low density may be a correlate of high mortality, and therefore an adaptive cue for an earlier age of sex change. Dominant territorial status correlates with an individual's size, and therefore egg-producing capacity, relative to its neighbors, and thus may also be a good cue for the initiation of sex-change.     

Genetic structure of a scleractinian coral, Mycedium elephantotus, in Taiwan

The genetic structure of Mycedium elephantotus (Pallas, 1766) populations from three regions around Taiwan was examined using allozyme electrophoresis. Eight loci were scored from seven enzyme systems. Seven loci were polymorphic under the 95% criterion. The high ratio of observed to expected genotypic diversities at the collection sites (G _O:G _E=0.8 to 1.0) indicate that M. elephantotus propagates predominantly by sexual reproduction. Allele frequencies of M. elephantotus differed significantly among regions (D=0.024 to 0.256, F _ST=0.032 to 0.218, p < 0.001), while populations among collection sites within each region were homogeneous (D=0.000 to 0.015, F _ST=0.010 to 0.022, p > 0.05). Genetic differentiation between populations from southern Taiwan and the Penghu Islands is greater than that between southern and northern Taiwan populations, although the former two regions are much closer geographically. The UPGMA dendrogram based on Nei's unbiased genetic distance showed a clear subdivision of populations into two groupings, northern Taiwan/Penghu Islands and southern Taiwan. A higher level of gene flow was found between M. elephantotus populations in northern Taiwan and the Penghu Islands (N _m=7.56) than that between populations in southern Taiwan and other regions (N _m=0.90 to 1.72). The pattern of genetic subdivision among regions is consistent with the pattern of ocean currents, indicating that genetic differentiation is likely driven by surface circulation vicariance. Received: 2 December 1997 / Accepted: 15 September 1998     

Population structure,northern range limit,and recruitment variation in the intertidal limpet Lottia scabra

The northern range limit of the intertidal limpet Lottia scabra is Cape Arago, Oregon (43°N), where adult survival is excellent, the population is small (<300), and recruitment is low; the range limit may be set by limited recruitment. Between June 2012 and March 2013, 25 sites from the middle of the species range (33°N) to Cape Arago were sampled and population size frequency distributions, densities, and nearest neighbor distances were compared to the amount of rocky and sandy shore and kelp bed size. North and south of 37°N, the densities of new recruits averaged 22 and 86 m^?2, respectively. This shift was associated with the range limit of Macrocystis pyrifera kelp beds; we hypothesize that slower currents in M. pyrifera beds may limit larval dispersal leading to higher recruitment. North and south of 40°N, adult density averaged <1 and 458 m^?2, respectively, with the species absent from many sites to the north. This shift was associated with a sharp drop in the amount of rocky shoreline and an increase in uninhabitable sandy shore. Near the northern range limit, >80 % of the individuals were solitary and may be unable to spawn successfully. Recruitment at Cape Arago was infrequent and likely due to self-recruitment. This study suggests that the range limit was set by the absence of M. pyrifera and too little rocky shore leading to high larval wastage, low settlement, low population densities, and, due to an Allee effect, very small effective population sizes.     

Population structure and genetic diversity in green turtles nesting at Tortuguero,Costa Rica,based on mitochondrial DNA control region sequences

The green turtle (Chelonia mydas) nesting population at Tortuguero, Costa Rica, is the largest nesting aggregation in the Atlantic, by at least an order of magnitude. Previous mitochondrial DNA (mtDNA) surveys based on limited sampling (n = 41) indicated low genetic diversity and low gene flow with other Caribbean nesting colonies. Furthermore, a survey of nuclear DNA diversity invoked the possibility of substructure within the Tortuguero rookery. To evaluate these characteristics, mtDNA control region sequences were determined for green turtles nesting at Tortuguero in 2001 (n = 157) and 2002 (n = 235). The increased sample revealed three additional haplotypes; five haplotypes are now known for Tortuguero female green turtles. Analyses of molecular variance indicated that there was no significant spatial population structure along the 30-km nesting beach. In addition, no temporal population structure was detected either between the two nesting seasons or within the nesting season. As a result of the larger sample size and additional haplotypes, estimates of genetic separation among Caribbean nesting colonies have changed and the concordance of phylogenetic and phylogeographic patterns reported in the past for green turtles in the Greater Caribbean has weakened. The five haplotypes from Tortuguero represent 36% of the haplotypes identified in green turtle nesting aggregations in the Greater Caribbean and 17% of the haplotypes known to occur in nesting or foraging aggregations in the Greater Caribbean. Haplotype diversity (0.16) and nucleotide diversity (0.0034) for the Tortuguero population are substantially lower than those for the combined rookeries in the Greater Caribbean (0.44 and 0.0078, respectively). Although comprehensive evaluation of regional genetic diversity requires nuclear DNA data, our study indicates that conserving genetic diversity in Caribbean green turtles will require careful management of the smaller rookeries in addition to the Tortuguero rookery.     

Population structure and reproductive biology of Ceratonereis limnetica (Polychaeta: Nereididae) at Lower Portland,Hawkesbury River,Australia

Ceratonereis limnetica Hutchings and Glasby was collected from Lower Portland, Hawkesbury River, New South Wales, Australia, from March 1982 to September 1983. C. limnetica is one of a relatively few polychaete species in which the adults normally tolerate reverine conditions. Reproduction in this species has been modified accordingly, differing from the common pelagic spawning of many nereidids. Adults of both sexes remain essentially unmodified and spawn on the bottom. Embryos and larvae are brooded in specially constructed tubes in the sediment by the female. Both sexes have a life span of about 1 yr, dying soon after spawning. The morphological and reproductive behavioural similarities between C. limnetica and C. aequisetis are considered to indicate a close phylogenetic affinity.     

Population genetic structure of the brown tiger prawn, Penaeus esculentus, in tropical northern Australia

Eight polymorphic microsatellite loci were analysed in six population samples from four locations of the Australian endemic brown tiger prawn, Penaeus esculentus. Tests of Hardy–Weinberg equilibrium were generally in accord with expectations, with only one locus, in two samples, showing significant deviations. Three samples were taken in different years from the Exmouth Gulf. These showed no significant heterogeneity, and it was concluded that they were from a single panmictic population. A sample from Shark Bay, also on the west coast of Australia, showed barely detectable differentiation from Exmouth Gulf (F _ST = 0 to 0.0014). A northeast sample from the Gulf of Carpentaria showed low (F _ST = 0.008) but significant differentiation from Moreton Bay, on the east coast. However, Exmouth Gulf/Shark Bay samples were well differentiated from the Gulf of Carpentaria/Moreton Bay (F _ST = 0.047–0.063). The data do not fit a simple isolation by distance model. It is postulated that the east–west differentiation largely reflects the isolation of east and west coast populations that occurred at the last glacial maximum when there was a land bridge between north-eastern Australia and New Guinea.     

Population structure and accumulation of lipids in the ctenophore Mertensia ovum

We studied the abundance, length-frequency distribution and body composition (lipid accumulation) of the ctenophore Mertensia ovum in Kongsfjorden, Svalbard (79°N) between May 2001 and April 2002. The highest abundances were found during summer in the outer and middle areas of the fjord, possibly caused by the concentration of advected zooplankton. Polymodal length-frequencies indicated a life span of 2 years, with 0, 1 and 2-groups being present in summer. Total lipid for a standardized animal decreased from May to August, and increased to the highest level in September. Relative lipid content was always highest in the 0-group, probably reflecting reduction of lipid levels due to increased energy costs during reproduction for the older age groups. Lipids probably are used to fuel prolonged reproduction from May to August until spawning ends by September. The overwintering age groups (0 and 1) subsequently build up lipid reserves for the next years’ spawning.     

Host effect on size structure and timing of sex change in the coral-inhabiting snail<Emphasis Type="Italic"> Coralliophila violacea</Emphasis>

The distribution, size and reproductive characteristics of the snail Coralliophila violacea (Lamarck), which inhabits the surface of both the branching coral Porites nigrescens and the massive corals P. lobata and P. lutea, were surveyed to examine the host effect on: (1) population structure and (2) reproductive characteristics, including the size at sex change of symbionts. On branching hosts, most snails were solitary, whereas on massive hosts, most had formed multiple-snail patches. Significantly smaller snails as well as proportionally more females were found on branching than on massive hosts. Furthermore, the fecundity of the females on the branching hosts was significantly lower than that on the massive hosts. The size at sex change (male to female) of the snails was smaller on the branching hosts than on massive hosts. Patch composition differences can partly explain the smaller size at sex change for snails on branching hosts; however, there was also evidence that host morphology had a significant effect on the timing of sex change.Communicated by T. Ikeda, Hakodate     

Population structure in the sexually reproducing sea anemone Oulactis muscosa

The intertidal sea anemone Oulactis muscosa (Drayton) is dioecious and most individuals are sexually mature throughout the year. Biochemical genetic evidence was used to determine the genetic structure of populations and to infer the relative contributions of sexual and asexual reproduction to recruitment. Data were collected for six enzyme-encoding loci from local populations spread along 735 km of the south east coast of Australia. The genetic structure of each of the nine local populations studied was consistent with recruitment by sexually produced individuals. In almost all cases, the observed single-locus genotypic frequencies closely matched those expected for hardy-Weinberg equilibria, however, consistent deficits of heterozygotes were detected for all loci. No apparent subdivision of the population was detected within the sampling area. Low levels of genetic differentiation were found between local populations and standardised variance (F _ST ) values were similar to those for other species with widespread planktonic dispersal of larvae.Contribution No. 60 from the Ecology and Genetics Group of the University of Wollongong     

Variation in size at sex-change among natural populations of the protandrous hermaphrodite, <Emphasis Type="Italic">Crepidula fornicata</Emphasis> (Gastropoda,Calyptraeidae)

Protandrous hermaphrodites are predicted to change sex from male to female when relative reproductive fitness of females surpasses that of males. How size at sex transition varies with population, mating group and individual parameters was investigated for five populations of the protandrous hermaphrodite slipper snail, Crepidula fornicata. The populations varied for density, size distribution, average mating group size and sex ratio. Size at sex-change was correlated with the population sex ratio. Comparisons of multiple hypotheses revealed that variables predicting the sex of a snail vary among positions in the mating group. The variables included body size, the relative size of the snail sitting atop the focal snail and population density. Our data support the conclusions that size at sex-change (and by inference, the size at which one sex has relatively greater fitness) is not fixed for these hermaphrodites and that individual size, social conditions and population differences all influence variation in relative fitness.     

Population structure and growth in captivity of the spiny lobsterPanulirus penicillatus from Dahab,Gulf of Aqaba,Red Sea

Panulirus penicillatus (Olivier, 1791) (Decapoda: Palinuridae) is the most common spiny lobster in the Red Sea and is widely distributed in the Indian and Pacific Oceans. Lobsters (n=377) were collected on four occasions during 1986 on the coral reef at Dahab, Sinai, Egypt. Average size of the collected individuals was 70.5±24.6 mm carapace length (CL) for males and 63.2±15.9 mm CL for females. The sex ratio was 1:1.64 males to females. Length increment per molt was inversely correlated with size and ranged from 2.1 mm per molt in the 40 to 50 mm CL size class to less than 1 mm in the 60 to 70 mm CL size class. Average intermolt period was ca. 136 d for all size classes. The relationship between carapace length and body weight was expressed by the equation:W _b=6.43 × 10^–4 × (CL)^2.89.P. penicillatus from Dahab differ in size, sex ratio and growth rate compared to other palinurid populations throughout their range. This might represent the effect of isolation and location at the edge of the geographical range for this species. It may also indicate an adaptation to their unique habitat in the coral reef in comparison to other palinurid species.     

Population structure and life cycle of the mesopelagic ostracod Conchoecia pseudodiscophora in Toyama Bay,southern Japan Sea

The population structure and life cycle of the mesopelagic ostracod Conchoecia pseudodiscophora Rudjakov in Toyama Bay, southern Japan Sea, were investigated using a time-series of samples collected during 0 to 500 m vertical hausls with twin-type Norpac nets (0.35 and 0.10 mm mesh) over one full year (1 February 1990 to 30 January 1991). Additional samples were also collected with a single-type closing Norpac net (0.06 mm mesh) to examine the vertical distribution patterns of eggs and all instars of this species. The proportion of gravid females present indicated that reproduction of C. pseudodiscophora continues throughout the year, but peaks in April–July. Eggs and Instars I and II were distributed below 500 m, while the more advanced instars were most abundant in the 350 to 500 m stratum by both day and night. Based on the abundance peaks of each instar in the time-series samples, development times were estimated to be 2.5, 4, 3, 7 and 11 mo for Instars III, IV, V, VI, and VII, respectively. Thus, a total of 30 mo is required for newly spawned eggs to hatch and reach adulthood. Stomach-fullness indexes revealed no seasonality in the feeding activity of any instar stage, but that feeding activity was low in older instars, particularly in adult males. The present results are compared with those for a few other ostracod species, in an attempt to characterize the life cycle of C. pseudodiscophora inhabiting waters of subzero temperature in the mesopelagic zone of the Japan Sea.     

Population structure of two Antarctic ice-associated copepods, Drescheriella glacialis and Paralabidocera antarctica, in winter sea ice

The spatial distribution and population structure of two dominant ice-associated copepods, Drescheriella glacialis and Paralabidocera antarctica, were studied during winter at nine locations in east Antarctic fast ice. These species accounted for at least 90% of the total metazoan abundance at each location. Abundances were high, reaching 175 individuals l^-1 (190,000 m^-2) for D. glacialis and 660 l^-1 (901,000 m^-2) for P. antarctica. These abundances were probably partly supported by the high biomass of ice-algae (Pearson correlation coefficient, r=0.75), as indicated by chlorophyll-a concentrations (1.7-10.1 µg l^-1). The population structures of each species suggested very different life-history strategies. All developmental stages of D. glacialis were isolated from the ice cores, including females with egg sacs, supporting the hypothesis that this species reproduces in the sea ice during winter. This strategy might assist D. glacialis in leading a continually colonising existence, whereby it responds opportunistically to the availability of favourable habitat patches. The populations of P. antarctica were composed primarily of nauplii (>99%), consistent with past observations of a synchronised life cycle for this species. The strong coupling of the developmental cycle of P. antarctica to the growth and decay of sea ice suggests that local extinctions might occur in areas where ice break-out is unpredictable.     

Population structure of the black tiger prawn, Penaeus monodon, among western Indian Ocean and western Pacific populations

We examined the population structure of the black tiger prawn, Penaeus monodon Fabricius, 1798, in the Indo-West Pacific by analyzing the geographic distribution of elongation factor 1-alpha intron sequences from specimens collected during the winter and spring of 1997. Both the molecular phylogeny of alleles and F-statistics indicated very strong differentiation between populations from the western Indian Ocean and western Pacific. This pattern is concordant with other recent studies of marine species in this region, implying that the Indo-Australian Archipelago represents a biogeographic break between populations in the Indo-West Pacific. F _ST-values among populations in the western Indian Ocean also indicate structure within this region, whereas no structure was found among western Pacific populations. Nucleotide diversity was significantly lower in the western Indian Ocean populations than in the western Pacific, implying that the populations have regional differences in demographic history. Received: 16 November 1998 / Accepted: 26 May 1999     

Population structure of the giant tiger prawn Penaeus monodon in Australian waters, determined using microsatellite markers

 We describe three highly polymorphic microsatellite loci which have been isolated from the giant tiger prawn Penaeus monodon. The number of alleles present among 312 samples at the loci Pmo9, Pmo25 and Pmo27 were 84, 34 and 35, respectively, with heterozygosities all >90%. Analyses of the distribution of length variation at three microsatellite loci among five Australian P. monodon populations revealed strong differentiation between populations from the west and those from the northern and eastern coasts. Tests for population differentiation (F st) values and an analogous measure for microsatellite loci (R st) all demonstrated that Western Australian P. monodon are a separate genetic stock which exhibits reduced genetic variation relative to the other populations. Reduced variability is consistent with a recent population bottleneck or colonization by a small founding population from the east when sea links between Indonesia, New Guinea and Australia were re-established following the last ice age. The results of this study are in agreement with previous surveys of P. monodon conducted with allozymes and mtDNA. Received: 18 December 1998 / Accepted: 27 August 1999     

Population structure and feeding deterrence in three shallow-water antarctic soft corals

Alcyonium paessleri and Clavularia frankliniana are numerically abundant soft corals in the nearshore (12 to 33 m depth) benthic communities of eastern McMurdo Sound. They are much less abundant in western McMurdo Sound where a third species, Gersemia antarctica, co-occurs in low numbers. The body tissues of these three species are comprised mainly of organic material (53 to 70% dry wt), which is primarily dervied from NaOH-soluble protein and refractory material. The energetic contents of the whole-body tissues of A. paessleri, C. frankliniana and G. antarctica are 15.9, 17.3, and 14.5 kJ g^-1 dry wt, respectively. The mean biomass per individual is 1.81, 0.008, and 45 g dry wt for each respective species. Based on population densities of 7.3, 1337.3, and 0.04 soft corals m^-2 for A. paessleri, C. frankliniana and G. antarctica, respectively, the population energetic densities are estimated to be 210.1, 185.1, and 26.1 kJ m^-2. Despite the relatively rich energetic content of the tissue and apparent vulnerability to predators, very little predation occurs on these soft corals. Two potential predators, the antarctic sea stars Perknaster fuscus and Odontaster validus, exhibited significant chemotactic defensive tube-foot retractions to hexane, chloroform, methanol, and aqueous methanol extracts of each soft coral. In addition, wholebody tissue of each soft coral was rejected by the demersal fish Pseudotrematomus bernacchii and the cryopelagic fish Pagothenia borchgrevinki. In contrast, whole soft-coral tissues sequentially extracted in four increasingly polar solvents were readily ingested by these antarctic fishes, indicating that sclerites do not play a significant role in deterring predators. Our results indicate that these antarctic soft corals contain bioactive compounds which deter common predatory seastars and fishes.     

Population distribution and structure of the free-living nematodes of Long Island Sound

The distribution and structure of nematode populations in 4 sedimentary environments (muds, muddy sands, fine sands and medium-coarse sands) in Long Island Sound were studied. Mean population densities were highest in muds and muddy sands. Cluster analysis suggested the presence of two basic faunistic units; a mud unit characterized by high species dominance, low species diversity and low species endemism, and a sand unit characterized by low species dominance, high species diversity and high species endemism. Species diversity in all habitats was a direct function of both species richness and equitability. Limited niche separation among deposit feeders, usually the dominant nematode trophic type in muddy sediments, is proposed as the cause for the high species dominance so often characteristic of shallow marine muds. The study afforded the opportunity to examine the quantitative and qualitative aspects of population structure in relation to environmental impact (as indicated by large differences in the heavy metal and organic carbon concentrations within each of the 4 sedimentary regimes). Within each sediment type no differences in population densities, species composition or species diversity of nematodes existed between heavily impacted and apparently non-impacted sediments, or between Long Island Sound and similar coastal regions. These findings cast doubt on (1) the use of heavy metal and organic carbon concentrations as indicators of environmental stress for marine nematodes; (2) the use of diversity indices alone as indicators of environmental deterioration; and (3) the usefulness of field monitoring studies alone for the assessment of pollution impact on marine nematodes.Contribution no. 100 from the Institute of Marine and Atmospheric Sciences, City College of New YorkCommunicated by M.R. Tripp, Newark