Analysis of individual year-classes of a marine fish reveals little evidence of first-generation hybrids between cryptic species in sympatric regions

As settled juveniles and adults, blue rockfish (Sebastes mystinus) are nonmigratory inhabitants of kelp and rocky reef habitats along the California coast, USA, and prior to settlement, they possess a pelagic larval and juvenile stage lasting 3–5 months. A previous study of adults revealed two cryptic species within S. mystinus and evidence of reproductive isolation in a region where both cryptic adults co-occur. Given this pattern of reproductive isolation, we investigated the degree of hybridization or introgression in individual year-classes shortly after juvenile settlement in two different years (2001 and 2002). Using microsatellite markers, we found little indication of hybridization in new juvenile year-classes despite an adult population that comprised both cryptic species. However, we found an average of two percent of hybrid or introgressed individuals in regions with a low frequency of one of the two species. Therefore, while the lack of hybrids or introgression supports the hypothesis of reproductive isolation between the cryptic species within S. mystinus, the age-structured analysis also revealed a spatial pattern of low-frequency differences in the number of introgressed individuals. These results suggest that reproductive barriers may breakdown when one of the two species predominates the regional adult gene pool.     

Population dynamics and reproduction of Mysella bidentata (Bivalvia: Galeommatacea) in Galway Bay,Irish west coast

Mysella bidentata (Montagu) is found in high densities in an Amphiura filiformis (O. F. Müller) community in Inner Galway Bay, west coast of Ireland. As part of an ongoing investigation at one station within this community, the population dynamics and reproductive pattern of this bivalve were investigated during the period November 1978-December 1981. Monthly sampling over the 3 yr period revealed a mean population density of 745±672 individuals m^-2. Recruitment occurred during August-October and three year-classes were identified. Growth rates were assessed by analysis of length-frequency histograms and winter growth checks. Individuals from the Y1 and Y2 year-classes functioned as hermaphrodites during the 1980 and 1981 reproductive seasons. The Y0 year-class contained both males and hermaphrodites. Results are compared with available information on the same species from other locations.Contribution No. 225 from the School of Marine Sciences, University College, Galway Ireland     

Reproductive ecology of the brooding,hermaphroditic clam Lasaea subviridis

The reproductive ecology of a small intertidal clam was studied for two years (September 1980-November 1982) at two mid-intertidal sites on the northern shore of Monterey Bay, California. Lasaea subviridis is hermaphroditic and incubatory; the young are released as juveniles. Brooding adults are found throughout the year. All embryos in a clam are at the same stage of development, but the number and stage of embryos varies among clams at any one time. Individuals are iteroparous and produce small numbers of oocytes and sperms simultaneously. All stages of gonad development are found in different individuals at any one time. There was no evidence of seasonality. Young reared in the laboratory after being dissected from the parent reached brooding size (shell length of 1.7 mm) within 6 mo. Laboratory experiments indicated that juveniles crawl away soon after release from the parent. These reproductive characteristics may contribute to this taxon's ability to colonize new environments rapidly.     

Genetic population structure of sardine (<Emphasis Type="Italic">Sardina pilchardus</Emphasis>) off Morocco detected with intron polymorphism (EPIC-PCR)

The upwelling systems along the coast of Morocco support some of the largest populations of sardine (Sardina pilchardus) in the world. Although these populations provide a base for a substantial fishing industry, virtually nothing is known about the genetic stock structure of this fish. Samples (n = 346), collected from seven sites along the Atlantic coast and in the Alboran Sea, were examined for exon-primed intron-crossing PCR (EPIC-PCR) polymorphism. Two markers, CaM-4 and Ops-1, had 6 and 9 alleles, respectively, after the pooling of gel fragments into 5 bp length classes, Correspondence analysis and the distribution of F _st among samples indicated that Moroccan populations were divided into two groups with F _st = 0.034 (P < 0.05) across the Gibraltar Strait. Populations along the Atlantic coast of Morocco comprise one genetic unit, except for a weak genetic boundary south of Cape Ghir and the peculiar behavior of the Safi sample would indicate a genetic drift. Complex ocean hydrodynamics around Gibraltar Strait and across Cape Ghir, likely, contributes to these genetic isolations. These results point out the usefulness of population genetic studies in stock management for sardine populations that may be particularly vulnerable to overexploitation especially during upwelling intensity shifts.     

Timing of hatching inAmmodytes marinus from Shetland waters and its significance to early growth and survivorship

The significance of hatch date for the growth and survival of the sandeel,Ammodytes marinus, was investigated using otolith microstructure. Hatch dates of 2 to 6 mo-old juvenileA. marinus caught near Shetland were compared between 1990 and 1992, during which period year-class strength varied by more than an order of magnitude. The hatch-date distribution of juveniles in the 1992 year-class was compared with that estimated directly from the abundance of newly emerged larvae on the spawning grounds. The extent of larval hatching periods in 1990 and 1991 was also estimated from continuous plankton-recorder data. There were significant differences in hatching periods between all three years, hatching in 1990 and 1992 being markedly earlier than the long-term mean peak in hatching indicated from archival data. Most individuals from the 1991 year-class attained a larger size by July than those in other year-classes, despite hatching later. Variation in individual growth rates both within and between year-classes indicated that there was a seasonal cycle of growth opportunity in all years investigated. The study suggests that the degree of coupling between hatching and the onset of spring secondary production may be an important contributory factor to year-class variability in this species.     

Small effective number of parents (<Emphasis Type="Italic">N</Emphasis><Subscript><Emphasis Type="Italic">b</Emphasis></Subscript>) inferred for a naturally spawned cohort of juvenile European flat oysters <Emphasis Type="Italic">Ostrea edulis</Emphasis>

The great fecundity and very high larval mortality of most marine invertebrates and fish make possible substantial variance in the number of offspring contributed by adults to subsequent generations. The reproductive success of such organisms may thus resemble a sweepstakes lottery, in which a minority of progenitors succeeds in replacing an entire population, while the majority fails to procreate. One specific prediction of this hypothesis, that genetic diversity of newly settled cohorts should be less than that of the adult population, is tested in the present study. Microsatellite DNA markers were examined in naturally spawned juvenile European flat oysters Ostrea edulis (L.), collected over a 12-day period in 1993 from the western Mediterranean Sea, near Sète, France (43°32′N, 3°56′E) and grown out for a period of up to 10 months. Variation in these juveniles was compared to that in a pooled sample of adults collected in 1994 from two locations (Thau Lagoon and Port St. Louis) that had statistically homogeneous allelic frequencies. Though nearly twice as large as the pooled adult sample, the juvenile sample had only 60% of the adult allelic diversity. Analyses of linkage disequilibrium and kinship, as well as estimation of the effective number of parents, suggested that 10–20 adults produced this juvenile cohort. This observation supports the hypothesis of sweepstakes reproductive success and suggests that partial inbreeding may occur even in species with large populations and dispersing planktonic larvae. Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Larval abundance and recruitment of the sand dollar Dendraster excentricus in Monterey Bay,California, USA

Seasonal abundance of planktonic larvae of the sand dollar Dendraster excentricus was determined in Monterey Bay, California, USA. Larvae were counted from two offshore stations and also over a coastal sand dollar bed, and these data were compared with settlement in the sand dollar bed, with adult population structure and with adult reproductive condition. These measurements were made during the period July 1978 to October 1980 and in October 1981. Sand dollar larvae were most abundant in the plankton during the summer, a period when phytoplankton productivity tended to be high and currents were relatively slow and variable. In some years, small-scale current variations appeared to prevent many larvae in the open bay from reaching the nearshore sand dollar beds; however, in other years, massive shoreward transport of the larvae evidently did occur, since the adult population in the sand dollar bed exhibited a mode in size, indicating a large settlement. A comparison of settled individuals in 1980 and the adult size-frequency distribution in 1981 gives an estimated mortality rate of 88% yr^-1 for early juveniles of D. excentricus.     

Population structure and genetic variation of lane snapper (<Emphasis Type="Italic">Lutjanus synagris</Emphasis>) in the northern Gulf of Mexico

Lane snappers (Lutjanus synagris), sampled from eight localities in the northern Gulf of Mexico (Gulf) and one locality along the Atlantic coast of Florida, were assayed for allelic variation at 14 nuclear-encoded microsatellites and for sequence variation in a 590 base-pair fragment of the mitochondrially encoded ND-4 gene (mtDNA). Significant heterogeneity among the nine localities in both microsatellite allele and genotype distributions and mtDNA haplotype distributions was indicated by exact tests and by analysis of molecular variance (AMOVA). Exact tests between pairs of localities and spatial analysis of molecular variance (SAMOVA) for both microsatellites and mtDNA revealed two genetically distinct groups: a Western Group that included six localities from the northwestern and northcentral Gulf and an Eastern Group that included three localities, one from the west coast of Florida, one from the Florida Keys, and one from the east (Atlantic) coast of Florida. The between-groups component of molecular variance was significant for both microsatellites (Φ _CT = 0.016, P = 0.009) and mtDNA (Φ _CT = 0.208, P = 0.010). Exact tests between pairs of localities within each group and spatial autocorrelation analysis did not reveal genetic heterogeneity or an isolation-by-distance effect among localities within either group. MtDNA haplotype diversity was significantly less (P < 0.0001) in the Western Group than in the Eastern Group; microsatellite allelic richness and gene diversity also were significantly less in the Western Group (P = 0.015 and 0.013, respectively). The difference in genetic variability between the two groups may reflect reduced effective population size in the Western Group and/or asymmetric rates of genetic migration. The relative difference in variability between the two groups was substantially greater in mtDNA and may reflect one or more mtDNA selective sweeps; tests of neutrality of the mtDNA data were consistent with this possibility. Bayesian analysis of genetic demography indicated that both groups have experienced a historical decline in effective population size, with the decline being greater in the Western Group. Maximum-likelihood analysis of microsatellite data indicated significant asymmetry in average, long-term migration rates between the two groups, with roughly twofold greater migration from the Western Group to the Eastern Group. The difference in mtDNA variability and the order-of-magnitude difference in genetic divergence between mtDNA and microsatellites may reflect different demographic events affecting mtDNA disproportionately and/or a sexual and/or spatial bias in gene flow and dispersal. The spatial discontinuity among lane snappers in the region corresponds to a known zone of vicariance in other marine species. The evidence of two genetically distinct groupings (stocks) has implications for management of lane snapper resources in the northern Gulf.     

Latitudinal trends in size-dependence of bioluminescence in the midshipman fish Porichthys notatus

The bioluminescent fish Porichthys notatus (plainfin midshipman), has a discontinuous distribution along the Pacific coast of North America. The fish is present from Cape Mendocino southward to Baja California, Mexico, absent off the coast of Oregon, USA, and abundant, northward, in Puget Sound, Washington. Interestingly, the population in Puget Sound lacks the substrate (luciferin) necessary for the luminescence reaction and, despite possessing an otherwise fully functional photophore system, is nonluminescent. The California population of P. notatus is uniformly luminescent south of Monterey Bay, but 15% of the speciments tested from San Francisco Bay and the Gulf of the Farallons have been reported to be nonluminescent. Explanations for nonluminescent midshipman in both Puget Sound and the San Francisco Bay area have focussed on a dietary requirement for luciferin. To gain further insight into reasons for nonluminescence in the San Francisco Bay region, the distribution of bioluminescence in P. notatus was studied from Monterey Bay to Cape Mendocino during 1985. A complex pattern of bioluminescence was found, in which nonluminescent individuals reflected neither a local anomaly in the San Francisco Bay region nor a simple gradient of decreasing luminescence towards the northern end of the range of the California population. Instead, a distinct size-dependent component in luminescence capability of the fish was observed. Aspects of the life history of P. notatus and related factors which might influence the bioluminescence characteristics of this population are discussed.Please address all correspondence to Dr. F.I. Tsuji at the Osaka Bioscience Institute     

Population genetic structure of North American <Emphasis Type="Italic">Ophiactis</Emphasis> spp. brittle stars possessing hemoglobin

With the discovery of previously unreported populations of hemoglobin-possessing Ophiactis from the Texas coast in the Gulf of Mexico, an investigation into its population structure, including populations of O. simplex from the Pacific coast of California and O. rubropoda from the Atlantic coast of Florida, was undertaken using DNA sequence data from the mitochondrial COI gene. The reconstructed haplotype network suggests that California populations contain the ancestral source of mtDNA variation, and there is no evidence of recent introductions into Texas. Population genetic analyses reveal the California, Florida, and Texas Ophiactis populations to each be significantly differentiated from one another. Sequence divergence among the three areas is shallower than would be predicted given biogeographic history. Texas and Florida populations are equally genetically diverged from California populations as they are to one another, despite the greater potential for gene flow between these areas. The genetic distinctiveness of the Texas populations and the concordance of this pattern with phylogeographic patterns in other brittle star systems indicate an isolated and independent evolutionary history and we hypothesize that the three geographic regions included in this study each serve as hypotheses of population-level lineages that remain to be tested with independent sources of data.     

Life history of Gonatus onyx (Cephalopoda: Teuthoidea): ontogenetic changes in habitat, behavior and physiology

In the present study 670 individuals of Gonatus onyx (Cephalopoda: Teuthoidea) were observed in Monterey Bay, California from a remotely operated vehicle. The vertical distribution of this species was bimodal, with peaks at 400 and 800 m depth during the day and 300 and 500 m during the night. The bimodal distribution reflects a life stage shift between younger, schooling juveniles living in shallower water and older, solitary adults which live deeper. Ontogenetic changes in behavior associated with this life stage shift are reflected in the physiology of the organisms as well. Both aerobic and anaerobic metabolism, as estimated using mitochondrial and glycolytic enzymes, decline with increasing body mass, suggesting reduced locomotory capacity in deeper-living adults. Oxygen consumption rates were also determined in relation to oxygen partial pressure. Oxygen consumption regulation was similar between juvenile and adult squids. The critical oxygen partial pressures (29 to 30 mmHg) correspond precisely to the oxygen concentrations found at the depth of maximal abundance for day and night populations of juveniles and adults, respectively. Behavioral and physiological changes with ontogeny of G. onyx are believed to result from reduced visual predator/prey interactions in the light-limited deep sea. Received: 7 February 1999 / Accepted: 26 January 2000     

The role of the amphipod <Emphasis Type="Italic">Gammarus locusta</Emphasis> as a grazer on macroalgae in Swedish seagrass meadows

Mesograzers are thought to play a critical role in seagrass beds by preventing overgrowth of ephemeral algae. On the Swedish west coast, eelgrass Zostera marina has decreased in recent decades as a result of eutrophication and increased growth of macroalgal mats (mainly filamentous Ulva spp. and Ectocarpales), with no indication of grazer control of the algae. The aim of this study was to investigate the ability of the amphipod Gammarus locusta to control algal blooms during nutrient-enriched and ambient conditions, using a combination of laboratory, field and model studies. Laboratory experiments demonstrated that juvenile and adult G. locusta could consume both Ulva spp. and Ectocarpales, but that consumption of Ulva spp. was significantly higher. Cannibalism was common in individual treatments involving multiple size-classes of G. locusta, but only large, male gammarids consumed smaller juveniles in the presence of Ulva spp. as an alternative food source. However, no negative effects of cannibalism were found on total grazing impact. A model using size-specific grazing rates and growth rates of G. locusta and of Ulva spp. suggests that approximately 62 young juvenile, or 27 adult G. locusta are needed per gram DW of Ulva spp. to control the algal growth during ambient nutrient conditions, and approximately 2.6 times as many gammarids during enhanced nutrient conditions. On the Swedish west coast, densities and mean sizes of G. locusta in eelgrass beds are below these critical values, suggesting that the gammarids will not be able to control the growth of the filamentous macroalgae. However, in the field cage experiment, immigration of juveniles and reproduction of encaged adult G. locusta resulted in unexpectedly high densities of G. locusta (>4,000 individual m⁻²), and very low biomass of Ulva spp. in both ambient and nutrient-enriched treatments. Although the high numbers of juveniles in all cages precluded any significant treatment effects, this suggests that in the absent of predators, the population of G. locusta can grow significantly and control the biomass of Ulva spp. Furthermore, low grazing of Ectocarpales in the laboratory and high biomass of these filamentous brown algae in the field indicate a preference for the more palatable green algae Ulva spp. This study indicates that the high grazing capacity of G. locusta, in combination with high reproduction and growth rates, would allow the amphipod to play a key role in Z. marina ecosystems by controlling destructive blooms of filamentous green algae. However, high predation pressure appears to prevent large populations of G. locusta in eelgrass beds on the Swedish west coast today.     

Genetic heterogeneity among adult and recruit red sea urchins, Strongylocentrotus franciscanus

Allozyme electrophoresis was used to characterize genetic variation within and among natural populations of the red sea urchin Strongylocentrotus franciscanus. In 1995 to 1996, adult urchins were sampled from twelve geographically separated populations, seven from northern California and five from southern California (including Santa Rosa Island). Significant population heterogeneity in allelic frequencies was observed at five of six polymorphic loci. No geographic pattern of differentiation was evident; neighboring populations were often more genetically differentiated than distant populations. Northern and southern populations were not consistently distinguishable at any of the six loci. In order to assess within-population genetic variation and patterns of recruitment, large samples were collected from several northern California populations in 1996 and 1997, and were divided into three size classes, roughly representing large adults (>60 mm), medium-sized individuals (31 to 60 mm, “subadults”) and individuals <2 yr of age (≤30 mm test diam, referred to as “recruits”). Comparisons of allelic counts revealed significant spatial and temporal differentiation among size-stratified population samples. Recruit samples differed significantly from adult samples collected at the same locale, and showed extensive between-year variation. Genetic differentiation among recruit samples was much higher in 1997 than in 1996. Between-year differences within populations were always greater for recruits than for adults. Potential explanations for the differentiation of recruit samples include pre- and post-settlement natural selection and high interfamily variance in reproductive success or “sweepstakes” recruitment. Unless recruit differentiation can be attributed to an improbable combination of strong and spatially diverse selection, such differentiation across northern California populations indicates that the larval pool is not well mixed geographically (even on spatial scales <20 km), despite long planktonic larval duration. Received: 6 July 1999 / Accepted: 25 January 2000     

Microsatellite DNA markers and morphometrics reveal a complex population structure in a merobenthic octopus species (<Emphasis Type="Italic">Octopus maorum</Emphasis>) in south-east Australia and New Zealand

Five polymorphic microsatellite loci were developed and then used to assess the population genetic structure of a commercially harvested merobenthic octopus species (Octopus maorum) in south-east Australian and New Zealand (NZ) waters. Beak and stylet morphometrics were also used to assess population differentiation in conjunction with the genetic data. Genetic variation across all loci and all sampled populations was very high (mean number alleles = 15, mean expected heterozygosity = 0.85). Microsatellites revealed significant genetic structuring (overall F _ST = 0.024, p < 0.001), which did not fit an isolation-by-distance model of population differentiation. Divergence was observed between Australian and NZ populations, between South Australia and north-east Tasmania, and between two relatively proximate Tasmanian sites. South Australian and southern Tasmanian populations were genetically homogeneous, indicating a level of connectivity on a scale of 1,500 km. Morphometric data also indicated significant differences between Australian and NZ populations. The patterns of population structuring identified can be explained largely in relation to regional oceanographic features.     

Recruitment and ontogenetic habitat shifts of the yellow snapper (<Emphasis Type="Italic">Lutjanus argentiventris</Emphasis>) in the Gulf of California

We examined recruitment and ontogenetic habitat shifts of the yellow snapper Lutjanus argentiventris in the Gulf of California, by conducting surveys and collections in multiple mangrove sites and major marine coastal habitats from 1998 to 2007. Over 1,167 juvenile individuals were collected and 516 otoliths were aged to describe the temporal pattern of the settlement. L. argentiventris recruits in mangroves, where juveniles remain until they are approximately 100 mm in length or 300-days-old. Back-calculated settlement dates and underwater surveys indicated a major recruitment peak during September and October, around 8 days before and after the full moon. The majority of mangrove sites in the Gulf of California had a similar L. argentiventris average size at the beginning of the settlement season for the cohort of 2003; although there were significant differences in individual sizes at the end of the nursery stage. When sub-adults leave mangroves, they live in shallow rocky reefs and later become abundant in deeper rocky reefs. The density of migratory individuals (10–20 cm SL) decreased exponentially as the distance between a reef and a nearby mangrove site increased. This finding has important implications for local fishery regulations and coastal management plans.     

Temporal stability of gene frequencies within genetically heterogeneous populations of the queen scallop Aequipecten (Chlamys) opercularis

The widely held assumption that gene frequencies within samples of multi-aged organisms are temporally stable has rarely been tested. In order to test this assumption in the commercially exploited bivalve scallop Aequipecten (Chlamys) opercularis (L.), 12 populations were sampled from around the UK over the period 1988–1990 and each was assayed at four putative polymorphic enzyme loci. Most populations were comprised of 3 to 4 year-classes, distinguishable by annual rings laid down on the shell. Three alleles from each locus were used as discriminant-function variables. Discriminant-function analysis based on allele frequencies averaged for each inividual year-class (45 demes) revealed highly significant inter-site heterogeneity. A posteriori classification of individual year-classes based on the discriminant analysis resulted in assignment of year-classes to the correct home site in 80% of cases. If allozyme neutrality is assumed, the phenomenon of predominant local allele frequency-stability through time within populations which are genetically heterogeneous from one site to another raises the possibility that queen scallop populations may be self-recruiting. Despite a planktonic larval phase lasting up to several weeks, this possibility is not excluded by the available hydrographic data.     

The importance of life stage to population connectivity in whiting (Merlangius merlangus) from the northern European shelf

Juvenile movements are rarely considered in studies of connectivity. This study investigates movements in juvenile and adult life stages of an ecologically important temperate piscivore, the whiting (Merlangius merlangus). Adult movement was assessed from tag-recapture data collected between 1958 and 1980 around the coast of Scotland. Displacement distances ranged from 53 to 123 km. Adults rarely dispersed offshore in the North Sea, and exchange with the west of Scotland was negligible. Connectivity between nursery and spawning grounds was investigated using otolith microchemistry. Five juvenile nursery areas were discriminated with an accuracy of 75–85%. Comparing isotopic signatures of adults and juveniles indicated that several nursery areas contributed to each spawning aggregation. The lack of evidence for return migrations suggested an opportunistic and non-philopatric recruitment strategy within a single population unit. Differences in the proportion contribution of juveniles to adult aggregations also indicated that the “value” of nursery areas differed and the Scottish west coast could be viewed as a net source of recruits to the North Sea. Given the extent of juvenile dispersal, this study highlights the fact that all stages of the life cycle should be considered in studies of connectivity.     

Wave exposure effects on population structure and recruitment in the mussel <Emphasis Type="Italic">Perna perna</Emphasis> suggest regulation primarily through availability of recruits and food,not space

Recruitment and population structure of Perna perna in low shore mussel beds were investigated over 15 months at six sites along the south coast of South Africa. Initial, subjective classification of sites as wave exposed or wave sheltered (three of each) was confirmed using the dissolution of cement blocks to measure average water flux and dynamometers for maximum wave force. Recruitment occurred throughout the year, but recruit (1–5 mm) densities were significantly higher from January to April 1996 on both shore types. Recruit densities were positively correlated with adult (>15 mm) densities for both shore types (P < 0.05) but the correlations were extremely weak (r ² < 0.06 in each case). In areas with 100% cover, adult size (mean and maximum lengths) was greater on exposed sites, but density showed the reverse and was negatively correlated with maximum wave strength (r = −0.84). Despite differences in adult densities and sizes, biomass, which is a product of the two, showed no significant difference between the two shore types (ANOVA P > 0.05). Thus wave exposure dramatically affects density, recruitment and mussel size, but not recruitment timing or biomass where there is 100% cover, and mediates a three-way interaction among food supply, larval supply and intraspecific competition for space. In contrast to shores with saturation recruitment, mussel biomass here appears to be limited by recruit supply and constraints of food, especially on sheltered shores, while density is regulated through intraspecific competition for space primarily on exposed shores and at small spatial scales.     

Genetic structure of leopard shark (<Emphasis Type="Italic">Triakis semifasciata</Emphasis>) populations in California waters

The leopard shark (Triakis semifasciata) is an important predator in coastal marine ecosystems of California, targeted by recreational and commercial fishermen and of specific interest in fisheries management. From October 2003 to August 2006, 169 leopard sharks were collected from the coast of California (between 40.750°N and 32.678°N) and analyzed for mitochondrial and nuclear genetic structure. Analyses of mtDNA control region sequences revealed relatively low levels of genetic variation (five haplotypes, average pairwise divergence π = 0.0067). In contrast, leopard sharks were highly polymorphic for inter simple sequence repeats (ISSRs), which characterize a broad range of the nuclear genome. The null hypothesis of panmixia in California waters was rejected for both genetic markers, and ISSRs displayed a statistically significant pattern of isolation by distance (IBD) across the species range (P = 0.002). A variety of analyses showed that divergence is most pronounced in the northernmost population of Humboldt Bay. Natal philopatry in T. semifasciata was tested using Siegel-Tukey tests on data partitioned by breeding site status, and sex-specific philopatry was tested by comparing IBD plots between sexes. Although there was some evidence for natal philopatry in leopard sharks (P = 0.038), and population divergence may be related to the proximity of breeding sites (P = 0.064), we found no support for sex-specific philopatry. In addition to identifying a novel set of highly variable genetic markers for use in shark population studies, these results may be used to better inform management decisions for leopard sharks in California.     

Feeding behavior of the ctenophore <Emphasis Type="Italic">Thalassocalyce inconstans</Emphasis>: revision of anatomy of the order Thalassocalycida

Behavioral observations using a remotely operated vehicle (ROV) in the Gulf of California in March, 2003, provided insights into the vertical distribution, feeding and anatomy of the rare and delicate ctenophore Thalassocalyce inconstans. Additional archived ROV video records from the Monterey Bay Aquarium Research Institute of 288 sightings of T. inconstans and 2,437 individual observations of euphausiids in the Gulf of California and Monterey Canyon between 1989 and 2005 were examined to determine ctenophore and euphausiid prey depth distributions with respect to temperature and dissolved oxygen concentration [dO]. In the Gulf of California most ctenophores (96.9%) were above 350 m, the top of the oxygen minimum layer. In Monterey Canyon the ctenophores were more widely distributed throughout the water column, including the hypoxic zone, to depths as great as 3,500 m. Computer-aided behavioral analysis of two video records of the capture of euphausiids by T. inconstans showed that the ctenophore contracted its bell almost instantly (0.5 s), transforming its flattened, hemispherical resting shape into a closed bi-lobed globe in which seawater and prey were engulfed. Euphausiids entrapped within the globe displayed a previously undescribed escape response for krill (‘probing behavior’), in which they hovered and gently probed the inner surfaces of the globe with antennae without stimulating further contraction by the ctenophore. Such rapid bell contraction could be effected only by a peripheral sphincter muscle even though the presence of circumferential ring musculature was unknown for the Phylum Ctenophora. Thereafter, several live T. inconstans were collected by hand off Barbados and microscopic observations confirmed that assumption.