Mitochondrial DNA polymorphisms reveal additional genetic heterogeneity within the Leptasterias hexactis (Echinodermata: Asteroidea) species complex

Sea stars of the Leptasterias hexactis complex are important members of the rocky intertidal community of the temperate and subarctic North American Pacific coasts. Recent attempts to determine the genetic structure and taxonomic relationships of these sea stars have not been completely successful, due to the combination of extreme morphological variability and relative scarcity of taxonomically-informative genetic (allozyme) polymorphisms. The present study used mitochondrial DNA restriction-site polymorphisms for PCR (polymerase chain-reaction)-amplified products to provide additional information about this complex. Five restriction-site haplotypes were identified in southern Alaska and in the Puget Sound region that were allozymically distinct. Because the allozymic differences (Nei's unbiased genetic differences >0.08) are maintained in sympatry, the haplotypes behave evolutionarily as conventional species. Although most haplotypes show deficiencies of heterozygotes for polymorphic allozyme loci, there is no evidence for the existence of cryptic species within any of the haplotypes. Rather than consisting of a few polytypic and cosmopolitan species, the L. hexactis complex is apparently composed of numerous species with more restricted ranges. Determining the exact ranges and taxonomic relationships of the various species will require additional study.     

Responses of Leptasterias hexactis (Echinodermata: Asteroidea) to low salinity

The six-rayed starfish Leptasterias hexactis (Stimpson, 1862) is seasonally exposed to low salinities in southeastern Alaska. Individuals that were gradually exposed to reduced salinities in the laboratory had a 28-d TLm of 12.9 S. The activity of L. hexactis, as measured by its activity coefficient, varied directly with salinity. Individual feeding rates of the starfish on similarly exposed Mytilus edulis, measured daily for 21 d at salinities of 30, 20 and 15 S, also varied directly with salinity. The dry weight of mussel tissue consumed was 8.84, 8.49 and 0.58 mg ·starfish^-1·d^-1 at 20, 20 and 15 S. Expressed as percent of dry starfish weight, the daily feeding rate was 1.35, 0.76 and 0.10% at 30, 20 and 15 S. Absorption efficiency decreased from 64% at 30 S to 49% at 20 S, further reducing the energy available for metabolism. Growth, measured in terms of changes in total dry weight or dry weight of soft tissues, also varied directly with salinity. Although exposure to hyposmotic conditions did induce stress responses, as indicated by reductions in activity, feeding, absorption efficiency and growth rates, L. hexactis maintained positive growth for at least a 3-wk period in the laboratory at 20 S and 13°CC. The population of L. hexactis investigated must be considered euryhaline and brief periods of exposure to hyposmotic conditions should not limit its distribution.     

Burrow morphology,biometry, age and growth of piddocks (Mollusca: Bivalvia: Pholadidae) on the south coast of England

Biometry and growth of three piddock species Pholas dactylus, Barnea candida and B. parva, from chalk and clay substrata were investigated between 1999 and 2000 at five low shore locations along the south coast of England. Piddock burrow shape was significantly different (P<0.01) amongst the locations. Burrows at Lyme Regis showed the largest (height/maximum diameter) ratio (4.86±2.00) whilst those at Compton the lowest (3.73±1.62). Using the method of Bhattacharya, the population structure of P. dactylus, B. candida and B. parva was separated into eight, three and five modal size classes, respectively. Age and shell growth were determined from the number and spacing, respectively, of annual growth lines present in acetate peel replicas of shell sections. The von Bertalanffy growth (VBG) equation fitted the size at age data obtained for P. dactylus and B. candida (L and K were 79.3±13.8 mm and 0.0011±0.22 and 29.6±1.5 mm and 1.17±0.47, respectively) whilst the size at age data for B. parva were linearly related and did not fit the assumptions of the VBG equation. Male and female gonads were mature and piddocks competent to spawn between June and September 1999, with settlement of juveniles observed between November 1999 and February 2000. A significant relationship between burrow aperture diameter and age of the occupant piddock was established for the three species. Burrow morphology and spatial distribution of burrows were influenced by substratum hardness and population density. Based on estimates at Lyme Regis, piddocks are capable of removing up to 41% of the shore substratum to a depth of 85 mm over their lifespan (12 years), significantly compromising the structural stability of the soft rock shores they inhabit and contributing to bioerosion.Communicated by O. Kinne, Oldendorf/Luhe     

Metabolic responses of the sympagic amphipodsGammarus wilkitzkii andOnisimus glacialis to acute temperature variations

Acutely elevated seawater temperatures had pronounced metabolic effect on the Arctic under-ice amphipodsGammarus wilkitzkii andOnisimus glacialis, collected in May 1986 and 1988 in the Barents Sea. An increased rate of oxygen consumption vs temperature was observed for both species. In the range from 0° to 10 °CG. wilkitzkii andO. glacialis exhibit Q₁₀ values of 3.4 and 3.6, respectively. The results also indicate increased ammonia excretion rates forG. wilkitzkii andO. glacialis by an elevation of temperature from 0° to 10°C, with an overall Q₁₀ of 1.9 and 2.3, respectively. The present study demonstrates an increased O:N ratio with ambient temperature elevation from 0° to 10°C forG. wilkitzkii andO. glacialis, with overall Q₁₀ values of 2.0 and 1.6, respectively. This indicates a temperature-induced change in the metabolic substrate towards lipids.     

Differences in morphology and habitat use among the native mussel <Emphasis Type="Italic">Mytilus trossulus</Emphasis>, the non-native <Emphasis Type="Italic">M. galloprovincialis,</Emphasis> and their hybrids in Puget Sound,Washington

Mytilus galloprovincialis (Mg), the Mediterranean blue mussel, is sympatric with the native M. trossulus (Mt) throughout much of the north Pacific, likely as the result of human introduction. We investigated the distribution of the two species and their hybrids (Mgt) in Puget Sound, Washington, to determine whether differences occur in habitat preference between the two species and hybrids. In addition, we investigated whether there were consistent size and shape differences between the native and introduced mussels and hybrids. Measurements of over 6,000 mussels from 30 sites, of which 1,460 were genotyped for a species-specific genetic marker, revealed that Mg and Mgt can be found throughout Puget Sound. Mg and Mgt were larger and exhibited a greater height:length ratio than Mt. Frequencies of Mg and Mgt were higher in subtidal habitats, such as docks, than on intertidal rocks, walls or pilings. Within intertidal habitats, Mg and Mgt were more frequent than Mt in the lower reaches of the intertidal. At slightly more than half the sites the frequency of the three genotypes accorded with random mating expectations suggesting no consistent barriers to gene flow between species. The standardized random sampling methods and simple morphometric identification techniques described here can be used to test whether the frequency of invasive mussels changes over time and space in Puget Sound.     

Population interconnectivity and implications for recovery of a species of concern, the Pacific hake of Georgia Basin

To aid the recovery of a species, understanding the extent to which populations are connected is useful for targeting conservation efforts. Pacific hake within waters of Puget Sound, Washington State, USA, and Georgia Strait, British Columbia, Canada are listed as a species of concern under the U.S. Endangered Species Act due to dramatic declines in the Puget Sound population. To assess the role of dispersal in the recovery of Pacific hake, we sought to quantify patterns of connectivity between populations in Puget Sound and Georgia Strait. Using natural chemical markers from otoliths of fish sampled from these two populations, we linked natal signatures of fish to signatures of individuals from known spawning grounds. Results indicated that 82 % of individuals collected from Puget Sound (n = 78) were estimated to have originated there, while 40 and 92 % of the individuals collected from two cohorts within Georgia Strait (n = 9 and 24, respectively) had originated from Puget Sound. A trend of “population abandonment” of fish from Puget Sound suggests that recovery of this Pacific hake population will depend on local management practices.     

Two populations of the marine fish Porichthys notatus,one lacking in luciferin essential for bioluminescence

Porichthys notatus is a common benthic fish found along the Pacific coast of North America. It possesses more than 700 small dermal photophores on its head and trunk. When P. notatus collected in Monterey Bay, California, is injected subcutaneously with norepinephrine, the photophores emit a long-lasting luminescence that is readily visible to the dark-adapted eye. The light emission is due to the oxidation of luciferin (substrate) by molecular oxygen, catalyzed by luciferase. In contrast, P. notatus collected in Puget Sound, Washington, is nonluminous, even though the photophores do not differ ultrastructurally from those of the California fish. The inability of the Puget Sound fish to luminesce is due to lack of luciferin in its photophores. Luminescence capability, however, may be induced in the Puget Sound fish by the oral or intraperitoneal administration of a small amount of luciferin from the tiny luminescent marine ostracod crustacean, Vargula hilgendorfii, suggesting that the origin of luciferin in P. notatus may be from the diet. In this study, specimens of P. notatus were collected over the entire known range of the species, between November 1981 and September 1987, and the presence of luciferin in the photophores determined. The results indicate that there are two populations of P. notatus: a luciferin-deficient, nonluminescent population located north of northern California and a luciferin-containing, luminescent population extending south of Cape Mendocino, California, to Baja California, Mexico. At the northern end of the southern population, a mixture of luminescent and nonluminescent P. notatus was found.     

Toxicity and attraction effects produced byMorinda citrifolia fruits on theDrosophila melanogaster complex of species

Summary The ripe fruit of the Indian mulberry,Morinda citrifolia, is the host plant forDrosophila sechellia but is highly toxic for three closely related species (D. melanogaster, D. simulans, D. mauritiana). A simple bioassay is described with which a clear dose response to the fruit was found for these species. Significant differences in reactivity to the ripe fruit were found among species. Tested strains ofD. simulans andD. mauritiana adults were more sensitive to the toxic properties of the fruit thanD. melanogaster. A marked intraspecific variability was shown inD. melanogaster. Reciprocal interspecific hybridizations betweenD. sechellia andD. mauritiana suggested an autosomal dominant control of resistance. MoreoverD. melanogaster intraspecific crossings suggested the influence of an additional X-linked factor. Responses of flies toMorinda fruit in different states were tested in a T olfactometer. The less resistant strains ofDrosophila generally showed less preference for the ripe fruit.     

Association of allozymes and temperature in the crested blenny Anoplarchus purpurescens

The crested blenny Anoplarchus purpurescens occurs intertidally in the Puget Sound area (USA), where it is subjected to substantial variation in temperature. A study of a polymorphic esterase in A. purpurescens has revealed a latitudinal cline and a correlation of allelic frequencies with summer water temperature. This finding agrees with previous results for a lactate dehydrogenase polymorphism in A. purpurescens, suggesting a pervasive genetic adaptation to temperature in this species. For each of these enzymes, comparisons within and between two species of Anoplarchus andicate an association between heterozygosity and temporal variability of the thermal environment.     

Phosphate uptake by intertidal algae in relation to zonation and season

The removal of phosphate from ambient seawater by whole plants of five species of fucoid algae, collected from the east coast of N. Ireland in 1988 and 1989, was followed over 6-h periods. A transient uptake pattern was observed forPelvetia canaliculata (L.) Dcne. et Thuret,Fucus spiralis L.,F. vesiculosus L. andF. serratus L., consisting of an initial period of high uptake, followed by a phase of zero uptake and then a period at an intermediate rate.Ascophyllum nodosum (L.) Le Jolis had a constant slow rate of uptake over 6 h. The initial uptake rate ofF. spiralis was significantly greater than that of any other species. Phosphate uptake over a 2-h period was measured at concentrations ranging from that of ambient seawater to 25µg-at. l^–1 for whole plants ofF. spiralis andF. serratus, using a large scale batch method. A small scale batch method was used for whole plants ofP. canaliculata and sections of the other four species investigated. Uptake abilities of the algae at low concentrations of phosphate were compared using the parameterV ₁ (the uptake rate at 1µg-at. l^–1) and at high concentrations usingV _max, the maximum uptake rate. These kinetic parameters of uptake were calculated using a method that avoids bias and permits statistical evaluation of the results. The fucoid algae studied could be divided into two distinct groups on the basis of their abilities to take up phosphate from seawater.P. canaliculata andA. nodosum had low values ofV ₁ in winter, which were also correlated with their positions on the shore and did not vary between winter and summer. TheFucus species had higher values ofV ₁ in winter, which were also correlated with their positions on the shore. In summer, however,V ₁-values for these species decreased and no longer correlated with their shore heights. TheV _max-value forF. spiralis was higher in winter than in summer but was signifcantly greater than that of any other species at all times of year. The ecological significance ofV _max is discussed in relation to nutrient limitation and the possible occurrence of patches of high nutrient concentration in the intertidal environment.     

Evidence for multiple recruitment-cohorts of Puget Sound Dungeness crab,Cancer magister

While sampling intertidally in Puget Sound, Washington, USA, for juvenile Dungeness crab (Cancer magister) in 1984, we found evidence of two distinct cohorts of the same year-class based on sizes of first-instar juveniles (J1) and the spatial/temporal patterns of settlement. In 1988, three distinct cohorts were observed to settle in Puget Sound and its approaches. Settlement of one cohort occurred during May in the Strait of Juan de Fuca and in those areas of Puget Sound closest to the Strait. J1 individuals of this cohort were large (x=7.4 mm carapace width, CW) and comparable in both size and timing of settlement to populations along the Washington coast (e.g. Grays Harbor and Willapa Bay). Initial settlement density of the May cohort was as high as 215 crabs/m² in intertidal eelgrass beds along the Strait of Juan de Fuca and decreased to <2 crabs/m² within Puget Sound and the Strait of Georgia. A second cohort apparently originated in Hood Canal (a deep inland fjord), its size upon settlement in June was significantly smaller (J1 x= 5.3 mm CW) than the May cohort, and it was limited to Hood Canal and areas of Puget Sound close to the mouth of Hood Canal. A third cohort, which settled in late July and August, was the smallest of the three cohorts (J1 x= 4.8 mm CW), and was widely distributed around Puget Sound from Seattle in the south to the USA/Canadian border in the north. We hypothesize that most juvenile recruitment in Hood Canal and Puget Sound originates from parental stocks endemic to their respective basins (Hood Canal and Puget Sound cohorts), but that, on occasion, oceanographic conditions allow substantial influx of Pacific Ocean Dungeness crab larvae (oceanic cohort) through the Strait of Juan de Fuca into Puget Sound. Tracking of spatial/temporal settlement patterns and comparison of J1 sizes proved useful for estimating the probable sources and dispersion of Dungeness crab larvae. Differences in size and time of settlement between various larval cohorts of C. magister may prove useful as biomarkers for tracing circulation patterns within and between inland waters of Washington and the Pacific Ocean. Causes of smaller size and later settlement of the Puget Sound cohort relative to oceanic conspecifics of the same year-class are discussed.Contribution No. 856, School of Fisheries, University of Washington, Seattle     

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     

The musselsMytilus galloprovincialis andM. trossulus on the Pacific coast of North America

Most recent authors have called the bay mussels of the Pacific coast of North AmericaMytilus edulis Linnaeus, 1758. Thirteen samples ofedulis-like mussels were collected from California, Oregon, and Alaska, USA, in 1985, 1986 and 1987. Electrophoretic evidence from wight loci indicates that these samples consist of two genetically distinct groups, neither of which is similar toM. edulis from the Atlantic Ocean. Mussels in southern California are very similar toM. galloprovincialis Lamarck, 1819 from the Mediterranean Sea; it is probable thatM. galloprovincialis was introduced accidentally to southern California. Mussels in Oregon and Alaska are similar to those from the Baltic Sea and parts of eastern Canada; the nameM. trossulus Gould, 1850 has priority for this taxon. In central and nothern California,M. galloprovincialis, M. trossulus and their hybrids co-occur. Despite the presence of hybrids betweenM. galloprovincialis andM. trossulus, the genetic integrity which they maintain across large areas of the world warrants their recognition as two distinct species.     

Reef coral reproduction in the Abrolhos Reef Complex, Brazil: the endemic genus Mussismilia

The current contribution deals with the reproductive biology of a genus endemic to Brazil –Mussismilia Ortmann, 1890 – including all three species of the genus: M. braziliensis (Verrill, 1868), M. hartti (Verrill, 1868), and M. hispida (Verrill, 1902), which occur sympatrically in the studied area, the Abrolhos Reef Complex, Brazil. Sexuality patterns, modes of reproduction, synchrony and spawning periods are reported, and were determined by histological examination of material. All three species started to develop female and male gametes over different periods in the same breeding season. The three species are probably broadcast spawners, since no embryos or planulae were observed in any species at any given time of the year. Each reproductive cycle lasted approximately 11 months. Oogenesis and spermatogenesis started in different periods, with spermaries appearing in approximately the eighth month of ovary development and lasting about 3 months. Reproductive cycles were annual. Spawning probably occurred in consecutive months in each species. In M. braziliensis, spawning presumably happened between March and the middle of May in 1996 and 1997. Evidence suggested that spawning events of M. hispida took place between the end of April and mid-June. M. hartti may have spawned between September and November. The data presented here suggested that all studied species have at least one exclusive spawning period, asynchronically with the others. A possible exception may be the simultaneous (or close) spawnings of M. braziliensis and M. hispida in May. It is suggested that asynchrony in spawning periods among species may reduce the chance of hybridization, gamete waste and the competition for settlement surfaces. The occurrence of extended spawning periods for each species may also reduce the risks of reproductive failure, due to temporary adverse conditions. Received: 8 December 1998 / Accepted: 15 July 1999     

Public Willingness to Pay for Recovering and Downlisting Threatened and Endangered Marine Species

Abstract: Nonmarket valuation research has produced economic value estimates for a variety of threatened, endangered, and rare species around the world. Although over 40 value estimates exist, it is often difficult to compare values from different studies due to variations in study design, implementation, and modeling specifications. We conducted a stated‐preference choice experiment to estimate the value of recovering or downlisting 8 threatened and endangered marine species in the United States: loggerhead sea turtle (Caretta caretta), leatherback sea turtle (Dermochelys coriacea), North Atlantic right whale (Eubalaena glacialis), North Pacific right whale (Eubalaena japonica), upper Willamette River Chinook salmon (Oncorhynchus tshawytscha), Puget Sound Chinook salmon (Oncorhynchus tshawytscha), Hawaiian monk seals (Monachus schauinslandi), and smalltooth sawfish (Pristis pectinata). In May 2009, we surveyed a random sample of U.S. households. We collected data from 8476 households and estimated willingness to pay for recovering and downlisting the 8 species from these data. Respondents were willing to pay for recovering and downlisting threatened and endangered marine taxa. Willingness‐to‐pay values ranged from $40/household for recovering Puget Sound Chinook salmon to $73/household for recovering the North Pacific right whale. Statistical comparisons among willingness‐to‐pay values suggest that some taxa are more economically valuable than others, which suggests that the U.S. public's willingness to pay for recovery may vary by species.     

Brood or broadcast? The adaptive significance of different reproductive strategies in the two intertidal sea stars Leptasterias hexactis and Pisaster ochraceus

This paper considers the adaptive significance of two different reproductive methods in two co-occurring, competing sea stars. The smaller (3 to 8 g mean wet weight) Leptasterias hexactis broods relatively few, large young in the winter, while the large (300 to 650 g mean wet weight) Pisaster ochraceus broadcasts relatively many, small eggs each spring. L. hexactis matures at a small size (2 g wet weight) in about 2 years, and P. ochraceus matures at a larger size (70 to 90 g wet weight) in about 5 years (Menge, 1974). As in many broadcasting asteroids, gonad and storage organ indices of P. ochraceus are inversely related over time, and maximum storage-organ index correlates with the summer feeding maximum (Mauzey, 1966). In contrast, both organ indices of L. hexactis and feeding increase and are positively correlated until early autumn, when feeding activity begins to decline. At this time the male gonad index continues to rise, and the storage-organ index drops. In contrast, both organ indices of females rise. Spawning occurs from November to January. Thereafter storage-organ indices decline in females, presumably because females draw upon energy reserves while brooding; storage-organ indices rise in males, presumably because males do not brood and can feed if food is available. The primary cause for the differences between annual reproductive cycles of P. ochraceus and L. hexactis is suggested to be patterns of food availability for the released young (planktonic food for the broadcasted young of P. ochraceus and benthic prey for the brooded young of L. hexactis. Estimates of pre-maturity survival and post-maturity longevity indicate that the probability of survival per individual of young P. ochraceus is vastly lower than that of L. hexactis. However, once mature, P. ochraceus has a much longer expected lifespan. Brooding is suggested to be a coadaptive consequence of competition-induced small size. Assuming planktonic mortality rates in this environment are roughly constant across broadcasting species, I suggest that a small broadcasting species could not produce enough offspring in its expected lifespan to replace itself. This hypothesis is partly supported by some simple simulations. Broadcasting is suggested to permit rapid location and utilization of spatially and temporally unpredictable, but highly desirable, resources by allowing rapid and widespread dispersal. Brooders presumably cannot disperse rapidly and must rely on more reliable, but perhaps less desirable, resources. Factors affecting reproductive patterns in marine invertebrates include (1) food availability for both adults and offspring, (2) planktonic mortality rates, (3) interactions between species and latitudinal changes in these factors, and (4) various physical factors. This paper suggests that competition and predation can have an important effect on the evolution of reproductive methods, a possibility heretofore largely ignored. Although several similar examples of co-occurring species' pairs which differ in reproductive method and size are available, the role of adult interactions is unknown in these examples.     

Depression of feeding and growth rates of the seastar Evasterias troschelii during long-term exposure to the water-soluble fraction of crude oil

To test the effect of petroleum hydrocarbons on predation by the seastar Evasterias troschelii (Stimpson, 1862) on the mussel Mytilus edulis (L.), we exposed the predator with the prey to six concentrations of the water-soluble fraction (WSF) of Cook Inlet crude oil. Seastars and mussels were collected at Auke Bay, Alaska, in November 1980. During a 28 d exposure in a flow-through system, seastars were more sensitive to the WSF than mussels: the LC₅₀ for the seastars was 0.82 ppm at Day 19 and, although no mussels were exposed to WSF for more than 12 d, none died. Daily feeding rates (whether in terms of number of mussels seastar^-1 d^-1 or dry weight of mussels seastar^-1 d^-1) were significantly reduced at all concentrations above 0.12 ppm. At 0.20, 0.28 and 0.72 ppm WSF, daily feeding rates (in terms of dry weight of mussels) were, respectively, 53, 37, and 5% of the control rate; at the two highest concentrations (0.97 and 1.31 ppm WSF), the seastars did not feed. Seastars at concentrations greater than 0.12 ppm WSF grew slower than individuals from the control group and the 0.12 ppm-treatment group combined. These laboratory results show that E. troschelii is more sensitive to chronic low levels of the WSF of crude oil. The possibility that such oil pollution could reduce predation and permit M. edulis to monopolize the low intertidal zone of southern Alaska remains to be studied.     

Non-random mating and population genetic subdivision of two broadcasting corals at Ningaloo Reef,Western Australia

Allozyme electrophoresis of two corals was used to assess whether populations at Ningaloo Reef, Western Australia are primarily self-seeding or whether recruitment is from a broader geographic pool. Significant genetic subdivision across a range of spatial scales (between 6.5 km and 155 km) was found for both Acropora digitifera and A. aspera, with mean F _ST values of 0.010 and 0.067 respectively. Large departures from Hardy-Weinberg expectations were found for both species. Without exception these were due to deficits of heterozygotes; mean D values were –0.341 for A. digitifera and –0.455 for A. aspera. The magnitude of the deficits was consistent both across loci for all sites and across all sites for each locus. Some loci were found to be in linkage disequilibrium but no consistent pattern was observed. Also, multi-locus genotypic diversity values were generally high (between 0.83 and 1.00) and so departures from equilibria cannot be attributed to asexual reproduction. The most plausible explanation for the patterns observed is restricted gene flow at both the planktonic and gametic stages, with mating between close relatives.Communicated by G.F. Humphrey, Sydney     

Dynamics of benthic vegetation standing-stock,irradiance, and water properties in central Puget Sound

The standing stock of benthic macroalgae, sediment-associated microalgae and eelgrass (Zostera marina L.) was sampled in conjunction with irradiance and water properties from June 1982 through March 1984 to examine the relationship between the dynamics of benthic primary producers and environmental factors in central Puget Sound, Washington, USA. Sediment-associated microalgal standing stock (measured by chlorophylla) peaked in April and August. The seaweed assemblage, dominated by bladed green algae (e.g.Ulva fenestrata) and eelgrass exhibited maximum standing stocks in August. Although water temperature correlated best with changes in standing stock of all vegetation types, solar irradiance appeared to trigger the onset of biomass buildup and autumn die-back by the plants. Seasonal variations in dissolved oxygen reflected the buildup and loss of plant standing-stock. Nutrient concentrations, with the exception of ammonia, exhibited seasonal trends. Most nutrients were in greatest concentration in winter and reached minimum concentration in late spring-summer. Regeneration of nutrients in autumn followed shortly after the autumn dieback of the benthic vegetation. We concluded that irradiance was the primary controlling factor in the system. Nutrient limitation (primarily nitrate) may control standing-stock accumulations from the period May–October when light is not limiting. In contrast to phytoplankton systems in deep portions of Puget Sound, shallow nearshore systems may be more susceptible to the effects of increased inorganic nutrient-loadings from anthropogenic sources.Contribution No. 808, School of Fisheries, University of Washington     

Feeding and swimming of lysianassid amphipods in a shallow cold-water bay

The potential for dispersal by lysianassid amphipods and their localization to carrion in a shallow cold-water bay in the Middle Saint Lawrence Estuary were assessed by means of endobenthic sampling, SCUBA observations, measures of swimming speeds, and by exposure of bait (50–100 g of fish) in traps. Seventy-five to 99.9% of animals attracted to traps were lysianassid amphipods belonging to five species. Lysianassid species were spatially segregated in the Bay at low tide but all were more or less dispersed at high tide. Second cohortAnonyx sarsi Steele and Brunel,Boeckosimus edwardsi andOnisimus littoralis (Krøyer) were more dispersed than the small first cohort individuals. Second cohortA. sarsi were crawlers or low (0–0.5 m off the bottom) suprabenthic swimmers in the day, but upper (0.5–2 m) suprabenthic swimmers at night. In contrast, first cohortA. sarsi were crawlers or low suprabenthic swimmers day-and-night, whileOrchomenella pinguis (Boeck) followed this swimming pattern at night but were generally akinetic in the day. Mean swimming speeds ofA. sarsi (13.6 cm s^-1) andOn. littoralis (12.1 cm s^-1) were 2 to 3 times greater than those ofOr. pinguis (7.4 cm s^-1) andPsammonyx nobilis (Stimpson) (4.4 cm s^-1). Catchability coefficients (i.e. ratio number of individuals per trap:endobenthic abundance) were 74 (A. sarsi), 8 (On. littoralis), 7 (Or. pinguis), and 0.7 (P. nobolis) m² of bottom. Gut content analysis indicated thatA. sarsi fed mostly on large carrion, whileOn. littoralis were markedly opportunistic, andOr. pinguis andP. nobilis relied on detritus, algae, and small crustaceans.