Molecular population structure of the marine benthic copepod Microarthridion littorale along the southeastern and Gulf coasts of the USA

Relationships among populations (southeast Atlantic and northern Gulf coast, USA) of a ubiquitous, estuarine, harpacticoid copepod (Microarthridion littorale Poppe) were estimated using sequence data from two loci: one mitochondrial [cytochrome b (Cyt b)] and one nuclear [first internal transcribed spacer of ribosomal DNA (ITS-1)]. Copepods were collected from seven estuaries in 1997/98. Allelic phylogenies based on both genes were generally concordant, and suggested that M. littorale populations are structured over large geographic scales (hundreds of kilometers). Three well-supported groups were found in both gene trees comprising clades of alleles sampled from South Carolina, Florida, and Louisiana. Alleles from the Savannah, Georgia sample formed a monophyletic group using the Cyt b data, but this clade was not distinguishable with comparable ITS-1 data. A single specimen from Louisiana was classified in different clades depending on the locus assayed. Received: 11 June 1998 / Accepted: 29 April 1999     

Impact of epibenthic predation on estuarine intertidal harpacticoid copepod populations

The extent to which energy is transferred directly from benthic meiofauna to epibenthic predators was investigated on an intertidal sand-flat in the Exe estuary, southwest England, during 1981–1982 and compared with data obtained from an intertidal mud-flat in the Lyhner estuary, also in south-west England, between 1978 and 1981. Two species of flatfish (Pleuronectes platessa L. and Platichthys flesus L.), two species of goby [Pomatoschistus microps (Krøyer) and P. minutus (Pallas)], brown shrimp (Crangon crangon L.) and shore crabs (Carcinus maenas L.) are the most common epibenthic predators feeding on the benthic invertebrates in these locatites. Harpacticoid copepods are the only component of the meiofauna to form a significant part of the diet of early juvenile stages of these predators, particularly the invertebrates. Harpacticoids are a more important source of food for predators feeding over the sand-flat than for those feeding on the mud-flat because in the sand-flat alternative prey of suitable size, such as small annelids, are absent. Moreover, the impact of predation on the mud-flat is spread over the whole harpacticoid species spectrum whereas on the sandflat it is confined almost entirely to a single species, Asellopsis intermedia (T. Scott). Flatfish, gobies and shrimp consume daily an estimated 0.01 to 0.1% of the standing stock of A. intermedia and account for between 12 and 22% of the observed reduction in the population of this species between July and October. Therefore, only a very small proportion of total meiofauna biomass is transferred directly to higher trophic levels.     

Photosynthetic responses and daily carbon balance ofColpomenia peregrina: Seasonal variations and differences between intertidal and subtidal populations

The seasonal photosynthetic responses and daily carbon gain of upper intertidal, low intertidal and subtidal (3 to 4 m depth) populations ofColpomenia peregrina were examined over a 2 yr period (1986–1988) in Santa Catalina Island, California, USA. The populations showed significant differences in their photosynthetic responses, daily carbon balance and carbon-specific growth rates when normalized to tissue area or to chlorophyll content. The substantial plasticity with respect to photosynthetic responses shown byC. peregrina is considered to be an important factor in facilitating the colonization of both intertidal and subtidal habitats. This species appears to have a cellular carbon metabolism influenced by responses to season and tidal elevation. Highest net daily carbon balance, predicted carbonspecific growth rates and net growth efficiency were achieved in upper intertidal habitats during summer. These parameters decreased in winter and progressively declined with increasing depth as plants become increasingly exposed to low-light regimes. The diminishing net daily carbon balance and predicted carbon-specific field growth rates found during winter suggest that standing stock and lower subtidal limits of distribution ofC. peregrina are at least partly controlled by these two factors.     

Ontogenetic feeding shifts in the meiobenthic harpacticoid copepod Nitocra lacustris

¹⁴C-radiolabelling experiments indicate that adult stages of the salt-marsh harpacticoid copepod Nitocra lacustris (Schmankevitsch) receive a large part of their nutrition through the ingestion and assimilation of certain diatoms. An abundance of empty diatom frustules occurs in the gutpellet contents of field-collected individuals. Naupliar stages do not ingest diatoms in the laboratory, and nauplii from the field do not contain frustules in their gut pellets. Ingestion of diatoms in the laboratory first occurs during the second or third copepodite stage. ³H-radiolabel expeiments and grazing experiments using bacterium-sized beads adhering to the diatoms indicated that both adults and nauplii ingest bacteria adhering to the outer mucus coating of the diatoms (and probably ingest the diatom mucus itself). Adults ingest bacteria (and probably mucus exopolymer) coincidently while ingesting diatoms. The nauplii ingest these components by scraping the outer surface of the diatoms. SEM observations indicate that diatoms are not punctured by the nauplii during feeding. While diatom mucus and associated bacteria play an (as yet unquantified) role in the nutrition of the adults, these components may comprise the bulk of food resources for naupliar stages.     

Mating system of the intertidal copepod Tigriopus californicus

Male Tigriopus californicus clasp immature females (copepodid stages II–V) for a period of up to a week prior to the female's terminal molt; upon maturation (stage VI) the female is inseminated and released. While females can mate anytime after their terminal molt, experiments using electrophoretically-detected genetic markers indicate that each mates only once in her lifetime. No evidence of sperm displacement was observed. Hence, male mating behavior can be interpreted as pre-copulatory mate guarding, a strategy employed to assure that a potential mate has not been previously inseminated. Males minimize the time investment required to insemiate a single female successfully by preferentially choosing to clasp more developmentally-advanced females; males clasped to stage III females will release them in order to clasp stage V females if the latter are present. Since males are capable of multiple mating, under most conditions of population sex ratio, this mating system results in low availability of unclasped, developmentally-advanced females; consequently, males must clasp successively younger (i.e. developmentally less-advanced) females in order to obtain a successful insemination.     

Variation in life-history patterns of the grass shrimp Palaemonetes pugio in two South Carolina estuarine systems

A population of Palaemonetes pugio Holthuis 1949, inhabiting a fairly constant high salinity estuarine environment (North Inlet), exhibited more rapid growth, earlier first reproduction, a smaller clutch size, more fluctuating sex ratio, and shorter life span. A population in a less saline environment (Minim Creek) showed relatively slower growth, delayed first reproduction, larger clutch size, female-dominated sex ratio, and longer life span. Growth in both areas was rapid in summer and slower in winter, with the females growing much larger than the males. Summer generation females first reproduced at the age of 3.5 months in North Inlet and at 4.8 months in Minim Creek. Minim Creek females larger than 30-mm TL carried more eggs than North Inlet females of similar sizes. Life span in North Inlet was calculated to be 6–7 months for the summer generation and 9–10 months for the winter generation; in Minim Creek, the corresponding longevity estimates were 9–10 months and 12–13 months, respectively. Variations in life history patterns are hypothesized to be the results of numerous environmental factors acting differentially on the various life-stages of the organism. The results suggest that the reproductive flexibility of P. pugio enhances its ability to persist in a variety of environments.Contribution no. 426 of the Belle W. Baruch Institute for Marine Biology and Coastal Research     

Variation in egg numbers between populations and between years in the holothurian Cucumaria curata

The relationship between egg number and latitude was studied in a common intertidal holothurian from the West Coast of North America. Nine populations of Cucumaria curata Cowles, 1907, ranging in location from Monterey, California to Vancouver Island, British Columbia, were sampled from 1972 to 1974. Egg number versus body weight regressions were determined for each population and examined within and between years for any patterns of variation. Very little variability was found in the slopes of the regressions, indicating a constant allometric relationship between egg number and body weight from year to year and population to population. However, the variability in the intercepts of the regressions indicated variable reproductive output between populations and from year to year.Contribution No. 3 of the Bodega Marine Sciences Association and PACTREX Report No. 3.     

Mate choice in the face of both inbreeding and outbreeding depression in the intertidal copepod Tigriopus californicus

 In species vulnerable to both inbreeding and outbreeding depression, individuals might be expected to choose mates at intermediate levels of genetic relatedness. Previous work on the intertidal copepod Tigriopus californicus has repeatedly shown that crosses between populations result in either no effect or hybrid vigor in the first generation, and hybrid breakdown in the second generation. Previous work also shows that mating between full siblings results in inbreeding depression. The present study again found inbreeding depression, with full sibling mating causing significant fitness declines in two of the three populations assayed. In the mate choice assays, a single female was combined with two males. Despite the costs of both inbreeding and outbreeding, mate choice showed clear inbreeding avoidance but no clear pattern of outbreeding avoidance. This lack of outbreeding avoidance may be attributed either to the temporary increase in fitness in the F₁ generation or to the absence of selection for premating isolation in wholly allopatric populations with infrequent migration. If this inability to avoid unwise matings is common to other taxa, it may contribute to the problem of outbreeding depression when allopatric populations are mixed together. Received: 18 May 1999 / Accepted: 25 January 2000     

The spatial structure of interzonal copepod populations in the Southern Ocean

The vertical distributions of Calanoides acutus, Calanus propinquus and Rhinicalanus gigas have been studied during Cruises I and II of the R.V. Ob in the Indian sector of the Southern Ocean. During the spawning period, all 3 species reveal the highest concentrations, and occupy the highest positions in the water column. During the period of feeding and growth, they begin slowly to disperse and to descend to greater depths. In spring, the populations ascend again to the water surface. In both seasons, later stages migrate first. Pronounced temperature gradients may prevent the ascent, and result in the breeding and subsequent growth of the new generation occurring in subsurface waters. Details of the vertical distributions of populations depend on their seasonal state and the local hydrology.     

Biogeographic patterns on shallow subtidal reefs in the western Indian Ocean

To resolve biogeographic limits and patterns on the east coast of Africa, the presence/absence and quantitative biomass data were collected from 55 shallow subtidal reefs along 4,800 km of coastline (5.2°–31.1°S). Multivariate analysis of distributions, trophic structure and biomass revealed two distinct marine provinces, the Tropical Indo-West Pacific and Subtropical Natal, with a transitional overlap (of ca. 120 km) located between Leven Point and Kosi Mouth on north-east South Africa. This region of overlap was one of three bioregions revealed by post hoc analyses. Biomass was unexpectedly highest in the tropics. ‘Auto-heterotrophs’ (species with autotrophic symbionts such as corals and clams), deposit feeders and grazers contributed significantly more biomass to the Tropical Indo-West Pacific Bioregion, whereas filter feeders dominated the Subtropical Natal Bioregion. ‘Auto-heterotrophs’ declined with latitude while filter feeders increased; soft corals made a defining contribution in the overlap bioregion. Possible underlying causes of these patterns include productivity, nutrient levels, riverine input, light penetration and temperature.     

Genetic differentiation of populations of the planktonic copepod Labidocera aestiva

Planktonic populations of the calanoid copepod Labidocera aestiva show significant biochemical genetic heterogeneity along the Atlantic coast of the USA. In summer, 1981, copepods were collected by surface tows at Beaufort Inlet, North Carolina; Fort Pierce Inlet, Florida; and Vineyard Sound, Massachusetts. Genetic variation within each population and genetic differentiation among the three populations were studied by micro-acrylamide gel electrophoresis of six loci encoding four enzymes. All six enzyme loci were polymorphic when all populations were considered together, but the North Carolina population was monomorphic at two loci. High genetic variability was indicated by the following: (1) the number of alleles per locus averaged over all loci was 2.57±0.26 SD; (2) the average proportion of loci for which the frequency of the most common allele was not greater than 0.95 was 0.78±0.10; (3) the frequency of heterozygous individuals was 0.25±0.07. Genetic differentiation among population samples in the three regions was demonstrated in several ways: allele frequencies at one aminopeptidase-I locus, Lap-1, differed significantly among samples of the three populations, and there were unique alleles of high frequency at this locus in two population samples. Values of the statistic of genetic distance (D) averaged 0.20±0.08 for pairwise comparisons between all samples. Compared to expected heterozygosity if individuals across the entire range sampled mated at random, there were highly significant heterozygote deficiencies at five of the six loci. Genetic differentiation of populations of L. aestiva may result from (1) differential selection on populations in the three regions, or (2) restricted gene flow between the populations. Gene flow may be limited by geographic separation or differences in life history, such as seasonal presence in the plankton and diapause egg production.Contribution No. 5810 of Woods Hole Oceanographic Institution     

Sex ratio and hermaphroditism in Oregon intertidal populations of the echinoid Strongylocentrotus purpuratus

Examination of large numbers of the echinoid Strongylocentrotus purpuratus (Stimpson) in the years 1969, 1970 and 1971 indicated that the sex ratio at Yaquina Head, Oregon (USA) was significantly different from 1:1, with males predominating. During 1968, 1970 and 1971 at Boiler Bay and Whale Cove, Oregon, the sex ratio did not differ significantly from 1:1 but, in 1969 at Boiler Bay, the proportion of males was 0.61. Two functionally hermaphroditic individuals were found among the 1,354 Yaquina Head echinoids examined, but none were found at Boiler Bay in an equal number of individuals. The hypothesis is proposed that S. purpuratus is a labile gonochorist, with a multiple, autosomal, sex-determining mechanism whose expression may be influenced by the environment.     

Variation in larval life-history traits among reef fishes across the Isthmus of Panama

We tested the hypothesis that regional differences in oceanic productivity have led to the evolution of predictable patterns of regional variation in life-history traits of pelagic larvae of tropical reef fishes. To do so we compared larval traits (egg and hatchling size, larval growth rate and duration, and size at settlement) among closely related reef fishes from the Atlantic and Pacific coasts of the Isthmus of Panama. This comparison provides a control for phylogenetic effects because those regions shared a common fauna prior to the rise of the Isthmus ˜3.5 million years ago, subsequent to which each fauna evolved independently under a very different productivity regime. We measured larval traits of 12 benthic-spawning damselfishes (Pomacentridae: Abudefduf, Chromis and Stegastes) and 13 pelagic-spawning wrasses (Labridae: Bodianus, Halichoeres and Thalassoma). These included members of each genus on each side of the Isthmus and four sets of transisthmian sister species of pomacentrids. Among the pomacentrids we found consistent transisthmian differences in hatchling size, but not in other larval traits. Essentially the reverse pattern occurred among the labrids – larval growth and duration differed consistently among congeners in the two regions, but without consistent differences in hatchling size or size at settlement. Neither relationship is predicted by the regional-productivity hypothesis. Most of the differences were quite small. Stronger phylogenetic effects on larval traits (inter- and intrageneric variation within regions) occur in both families and evidently overwhelm any effect of regional variation in productivity. Reassessment of data that takes into account such phylogenetic effects questions previous conclusions about the existence of regional differences in larval traits among damselfishes in the West Pacific and the Caribbean. Received: 19 January 2000 / Accepted: 26 September 2000     

Hydrodynamic patterns in lotic intertidal sands and their bioclimatological implications

Water movement is probably the most important factor influencing life in the interstices of sandy beaches. In order to better understand this ecosystem, e.g. the distribution and migration of interstitial fauna and overall energy transfer, measurements of water flow through high energy beaches were made on the Atlantic coast of the USA. Special hot thermistor-probes and associated electronic circuits were built, capable of measuring water-flow velocity down to less than 10 m/sec. These were buried in the sand at various depths and locations in the intertidal zone and the flow velocity recorded continuously for one or more tidal cycles. In addition, an orientation sensor was constructed and used together with the flow probe so that a picture of the flow patterns in the intertidal zone of a beach and their changes during a tidal pattern could be developed and defined quantitatively.     

Egg production and hatching success in North Adriatic Sea populations of the copepod acartia clausi

Egg production, fecal pellet production and hatching success are reported for Acartia clausi females sampled during three cruises in February 1997, 1998 and June 1997 at 20-24 stations along 4 transects in the North Adriatic Sea. Dramatically low hatching rates were recorded during both diatom bloom events in February as opposed to much higher rates during post-bloom conditions in June, even though A. clausi productivity during the bloom was apparently high. These results are discussed in the light of recent findings on the negative impact of diatoms on copepod reproductive potential.     

Variation in cadmium uptake by estuarine phytoplankton and transfer to the copepod Eurytemora affinis

The pathways of cadmium (Cd) uptake and transfer within an estuarine planktonic community from the Patuxent River, Maryland, USA, were investigated using an assemblage of natural phytoplankton and the copepod Eurytemora affinis Poppe. The experiment was carried out in October 1992 in replicated 500-liter, flow-through, fiberglass tanks. Growth rate, species composition, and Cd loading affected the accumulation of Cd by the phytoplankton. Uptake of Cd by phytoplankton was proportional to the amount of Cd available in the water column. Partition coefficients (K _d) for phytoplankton uptake averaged 4.4 × 10⁴. As metal loading rates and phytoplankton species composition changed during the 12-d experiment, Cd partitioning declined. Transfer of Cd to E. affinis occurred from Cd-laden phytoplankton, with levels in the copepods being approximately the same as, or somewhat less, than in the phytoplankton. Some Cd uptake occurred in copepods exposed to dissolved Cd only; however, the uptake was considerably less than that seen from food. Thus, Cd content of higher trophic levels, such as copepods, can be affected by the degree of Cd incorporation in their food source, and by ecological factors regulating phytoplankton ingestion. Received: 13 September 1995 / Accepted: 29 October 1998     

Latitudinal patterns of diversity and abundance in North Atlantic intertidal boulder-fields

In order to study taxon richness, biodiversity and abundance patterns in the North Atlantic from temperate latitudes through Arctic to high Arctic latitudes, we recorded the faunas (at ELWS level) colonizing 20 cobbles from three sites at each of seven boulder-field localities (south-west England, 50°N; Wales, 51°N; west Scotland, 56°N; Iceland, 64°N; Tromsø, 70°N; Svalbard, 77°N, 79°N). Inverse correlations were found between latitude and all measures of richness (species, orders, and phyla numbers) and biodiversity (S–W, P). However, these correlations were driven mostly by the consistently impoverished Arctic sites; an even cline of decreasing diversity from south to north did not exist. Multidimensional scaling revealed two communities, temperate–subarctic species-rich and high arctic species-poor. Evenness as measured by Pielou’s index was similar across all latitudes. Abundance data exhibited a similar trend to the species richness and diversity data with a significant negative correlation with latitude, but when Arctic data were excluded the correlation vanished.     

Reproduction and development in the pycnogonid Pycnogonum littorale

Aspects of the reproductive biology of Pycnogonum littorale (Ström) are examined. Vitellogenesis occurs between November and August, the process resembling that in Nymphon gracile, annelids, and Limulus polyphemus, in which the majority of the yolk is synthesized within the oöcyte, with only a small contribution from outside, thus contrasting with the method in insects. The mode of formation of the mature yolk platelets shows similarities with the process described in Cambarus spp. Growth occurs in the post larva by a series of moults, and sometimes in the adult by a process not involving moults. The male usually has 9 moults and the female 10 or 11.