Evidence for seasonal mate limitation in populations of an intertidal amphipod, Corophium volutator (Pallas)

Potential rates of reproduction (PRR) differ between the sexes of many animal species. Adult sex ratios together with PRR are expected to determine the operational sex ratio (OSR) defined as the ratio of fertilizable females to sexually active males at any given time. OSR is expected to determine the degree to which one sex competes for another—the limiting sex. We explored the potential for mate limitation in an intertidal amphipod, Corophium volutator (Pallas). Males have higher PRR than females, but males may be limiting because of extreme female-biased sex ratios observed in this species. Consistent with this idea, late season females were less likely to be ovigerous and had smaller size-specific clutches, both of which were associated with seasonal declines in availability of males of reproductive size. Seasonal changes in ovigery could not be explained by seasonal changes across sites in other factors (e.g., female body size or phenology of breeding). Smaller females were less likely to become ovigerous later in the season at three of four sites. Seasonal reductions in clutch size also occurred among small females expected to be reproducing for their first time. In complimentary laboratory experiments, reduced likelihood of ovigery and reduced fecundity occurred when the number of receptive females was increased relative to availability of a reproductively active male. Our results suggest male mate limitation can occur seasonally in this species and that male limitation is regionally widespread and may affect recruitment.     

Bioturbation effects of the amphipod Corophium volutator on microbial nitrogen transformations in marine sediments

Microcosms containing different densities of Corophium volutator, ranging from 0 to 6000 ind m^-2, were incubated in a flow-through system. Benthic fluxes of CO₂, O₂, NO₃ ^- and NH₄ ⁺ were measured regularly. Thirteen days after setup the microcosms were sacrificed and sediment characteristics, pore water NO₃ ^-, NH₄ ⁺ and exchangeable NH₄ ⁺ concentrations, and potential nitrification activity were measured. The presence of C. volutator increased overall mineralization processes due to burrow construction and irrigation. The amphipods increased the ratio CO₂/O₂ fluxes from 0.73 to 0.86 in microcosms inhabited by 0 and 6000 ind m^-2, respectively. Burrow ventilation removed NH₄ ⁺ from the sediment, which was nitrified in the oxic layer and transported NO₃ ^- to the burrow sediment, where denitrification potential was enhanced. Nitrification and total denitrification rates (denitrification of NO₃ ^- coming from the overlying water and of NO₃ ^- generated within the sediment) were calculated and discussed. Bioturbation by C. volutator increased both nitrification and denitrification, but denitrification was stimulated more than nitrification. Denitrification of NO₃ ^- coming from the overlying water was stimulated 1.2- and 1.7-fold in microcosms containing 3000 and 6000 ind m^-2 relative to control microcosms. The presence of C. volutator (6000 ind m^-2) stimulated nitrogen removal from the system, as dinitrogen, 1.5-fold relative to non-bioturbated microcosms. C. volutator individuals used in our study were collected from Norsminde Fjord, Denmark, in 1990.     

Particle size-selection of two deposit feeders: the amphipod Corophium volutator and the prosobranch Hydrobia ulvae

The feeding biology of the deposit-feeding amphipod Corophium volutator is compared to that of the coexisting, deposit-feeding prosobranch Hydrobia ulvae. Regarding ingestion of particles, both forms show size selection which alone can explain their coexistence. Particle size-selection also explains some qualitative differences in the composition of the food of the two forms; thus, diatoms play a relatively larger role in the diet of H. ulvae than in the diet of c. volutator, whereas bacteria are probably relatively more important for the latter. Results of experiments with feeding of C-14 labelled microorganisms are in accordance with the findings on particle size-distribution of the gut contents, and show that (1) C. volutator can only utilize bacteria adsorbed to particles within the size range 4 to 63 (this is why the presence of clay and silt particles in the sediment are necessary for efficient feeding of this amphipod); (2) C. volutator can utilize bacteria suspended in the water pumped through its burrow for respiration if silt and clay particles are present in the sediment. (3) H. ulvae can utilize large particles, and also browses on surfaces, and some evidence is brought forward that it also utilizes mucus for trapping microorganisms. The coexistence of deposit-feeding animals is discussed. It is concluded that the number of coexisting, closely related species is usually small, and that their resource partitioning is probably mainly based on particle-size selectivity. In the case of unrelated forms (e.g. H. ulvae and C. volutator, a number of behavioural, physiological and morphological differences, and also the widespread ability of deposit feeders to utilize alternative feeding mechanisms may also lead to resource partitioning. Thus, there are often several niche dimensions related to feeding allowing a certain diversity of coexisting deposit feeders.     

Feeding and digestion in Corophium volutator (Crustacea: Amphipoda)

New observations on the feeding behavior of Corophium volutator, collected from the Menai Strait, Anglesey, UK in 1981, show that detritus drawn into one entrance of the U-shaped burrow is pushed out at the opposing entrance either uningested or as faeces. Periodically, the amphipod turns around in the burrow and recommences feeding, so both the uningested material and the faeces may be reworked by the mouthparts. A feeding individual produces, on average, 16 faecal pellets every 10 min, at a seawater temperature of 18°C. When fed on detritus dyed with azo-carmine, approximately 12 pellets are produced before the foregut, intestine and hindgut are completely clear of previously ingested material and the dye appears in the faeces. Dye which enters the ventral caeca takes between 12 and 48 h to clear this tissue and, furthermore, it is only cleared when the amphipod is feeding. It is concluded that ingested coarse material is subjected only to a primary phase of digestion in the stomach, and that it takes 4 to 24 min to clear the gut. Fine material which enters the ventral caeca is subjected to a secondary phase of digestion and absorption which takes 24 to 48 h. Particles of thorium dioxide and ferritin incorporated in the diet are absorbed by the B- and ageing RF-cells in the ventral caeca. Non-specific esterases occur in the apical regions of the same cells, the mature cells of the anterior dorsal caeca and the cells at the anterior of the intestine. Protease activity is greatest in the lumen of the ventral caeca and the large vacuole of the B-cells. Apart from the lumen of the posterior caeca, there is some protease activity throughout the lumen of the gut. Carbohydrates were localised primarily in the R/F-cells towards the proximal region of the ventral caeca. Lipid was confined to the R/F-cells in the same region.     

Biological and environmental influences on the trophic ecology of leatherback turtles in the northwest Atlantic Ocean

Understanding the causes and consequences of variability in trophic status is important for interpreting population dynamics and for identifying important habitats for protected species like marine turtles. In the northwest Atlantic Ocean, many leatherback turtles (Dermochelys coriacea) from distinct breeding stocks throughout the Wider Caribbean region migrate to Canadian waters seasonally to feed, but their trophic status during the migratory and breeding cycle and its implications have not yet been described. In this study, we used stable carbon and nitrogen isotope analyses of bulk skin to characterize the trophic status of leatherbacks in Atlantic Canadian waters by identifying trophic patterns among turtles and the factors influencing those patterns. δ¹⁵N values of adult males and females were significantly higher than those of turtles of unknown gender (i.e., presumed to be subadults), and δ¹⁵N increased significantly with body size. We found no significant differences among average stable isotope values of turtles according to breeding stock origin. Significant inter-annual variation in δ¹⁵N among cohorts probably reflects broad-scale oceanographic variability that drives fluctuations in stable isotope values of nutrient sources transferred through several trophic positions to leatherbacks, variation in baseline isotope values among different overwintering habitats used by leatherbacks, or a combination of both. Our results demonstrate that understanding effects of demographic and physiological factors, as well as oceanographic conditions, on trophic status is key to explaining observed patterns in population dynamics and for identifying important habitats for widely distributed, long-lived species like leatherbacks.     

Analysis of an intertidal population of the amphipod Gammarus palustris using a modified version of the egg-ratio method

An intertidal population of Gammarus palustris Bousfield was sampled over a 23-month period and analyzed by a modified form of the egg-ratio method. Total numbers and size-frequency distributions revealed small numbers in the early spring which later increased to a peak in the summer months. This was followed by a marked reduction near the end of sumer and a subsequent rise in numbers during the early fall. Finally there was a second decrease to very low numbers in the late fall and early winter. Average brood size was calculated and used as an indication of food limitation. Attempts to correlate this with per capita rate of change (r), birth rate ( \((\hat b)\) and death rate ( \((\hat d)\) ) suggested that the population increase in the early summer was food-rather than predator-limited. The late summer decline appears to be the result of predation pressure as well as food limitation. The fall peak is attributed to juvenile release from ovigerous femals escaping previous predation and to an increase in adult males resulting from a release in predation pressure. The winter decline may be due to adverse environmental conditions.     

Accumulation of copper within the “Hepatopancreatic” caeca of Corophium volutator (Crustacea: Amphipoda)

The concentrations of various metals in specimens of Corophium volutator (Pallas) which have been taken from a site that receives freshwater drainage with a high concentration of copper, zinc and manganese, are compared with those in individuals from a site with a normal concentration of these metals for coastal waters. The amphipods at the contaminated site contain 259 g g⁺¹ dry weight of copper, which is three times that of individuals from the uncontaminated site. The concentrations of the other metals do not differ substantially between the two populations. The hepatopancreatic caeca of the midgut of amphipods with a high concentration of copper have numerous granules of homogeneous electron-dense material which can aggregate within the multivesicular bodies. Some of the amphipods with a lower concentration of copper have similar granules, but these are fewer in number. X-ray microporbe analysis shows that these granules contain copper and sulphur, and occasionally calcium. Various tests on the granules in frozen section indicate that they are relatively insoluble in a wide range of reagents and that the copper is probably complexed with organic matter.     

Space partitioning and interactions in an intertidal sand-burrowing amphipod guild

A nine-year study at a moderately exposed, fine sand intertidal beach at Long Sands, Maine, USA identified marked seasonal changes in the foci of sand-burrowing amphipod activities. These included seasonal population movements, degree of interspecific horizontal overlap, life stage segregation within and between species, and sand depth stratification. Behavioral interactions related to competition for available space are considered important in affecting the distribution and abundance of amphipod species, and community structure. Significant negative correlations were shown between the abundance and upper distributional limits of the smaller, lower shore Acanthohaustorius millsi and the abundance of the larger, higher shore species Haustorius canadensis. Tolerance limits to a variety of prevailing physical factors alone, were not considered a serious obstacle to occupation of higher intertidal level sands by A. millsi. Summer brooding females of both species occupied similar 5 to 10 cm sand depths, but were strongly segregated horizontally. Laboratory coexistence experiments during the reproductive period showed increased mortality of adults and negligible reproductive output of A. millsi when in combination with H. canadensis compared to controls where the former species was alone. A third species, Amphiporeia virginiana, occupied primarily very shallow sand (0 to 2.5 cm), and performed seasonal movements opposite in direction to A. millsi and H. canadensis. The presence, amount, and refinement of biological interactions across the wave exposure gradient requires further study. Until then, we consider as inappropriate the blanket designation of all types of intertidal sand communities as physically controlled.     

Precopulatory guarding-time of the male amphipod Eogammarus oclairi: effect of population structure

This study shows that population structure affects guarding time in the amphipod Eogammarus oclairi Bousfield. In the laboratory, mean guarding time depended on the male weight-frequency distribution, operational sex ratio, and male density. It was longer when the male population encompassed a broad range of weights than when the weight distribution was narrower; however, guarding time did not differ when the population's weight distribution was narrow and composed of either small or large males. Mean guarding time varied between a few hours when the operational sex ratio (male:female) was 1:3 to>7 d when the sex ratio was 2:1. It was also affected by male density, increasing as male density increased. The results suggest that the stimulus to guard depends on the intensity of male-male competition.     

Multi-marker estimate of genetic connectivity of sole (Solea solea) in the North-East Atlantic Ocean

A thorough knowledge on the genetic connectivity of marine populations is important for fisheries management and conservation. Using a dense population sampling design and two types of neutral molecular markers (10 nuclear microsatellite loci and a mtDNA cytochrome b fragment), we inferred the genetic connectivity among the main known spawning grounds of sole (Solea solea L.) in the North-East Atlantic Ocean. The results revealed a clear genetic structure for sole in the North-East Atlantic Ocean with at least three different populations, namely the Kattegat/Skagerrak region, the North Sea and the Bay of Biscay, and with indications for a fourth population, namely the Irish/Celtic Sea. The lack of genetically meaningful differences between biological populations within the southern North Sea is likely due to a large effective population size and sufficient connection (gene flow) between populations. Nevertheless, an isolation-by-distance pattern was found based on microsatellite genotyping, while no such pattern was observed with the cytochrome b marker, indicating an historical pattern prevailing in the latter marker. Our results demonstrate the importance of a combined multi-marker approach to understand the connectivity among marine populations at region scales.     

Comparisons of genetic population structures in four intertidal brachyuran species of contrasting habitat characteristics

Genetic population structures along the Japanese coast, analyzed by sequence data from the mitochondrial DNA COI region, were determined for four intertidal brachyuran species in the superfamily Thoracotremata (Ocypode ceratophthalma, Gaetice depressus, Chiromantes dehaani and Deiratonotus japonicus), which were characterized by different habitat requirements. O. ceratophthalma (seashore; supratidal sand) and C. dehaani (estuarine; supratidal marsh) showed no significant genetic differentiation among Japanese populations. The Japanese populations of O. ceratophthalma, however, were found to genetically differentiated from the Philippine population. G. depressus (seashore; intertidal cobbles) exhibited significant genetic differentiation between the Amami-Ohshima population and other local populations. D. japonicus (estuarine; intertidal cobbles) showed significant genetic differentiation among many local populations separated by about 30–1,200 km. The different patterns of genetic population structure recorded for the four species, thus, do not simply correspond to habitat type.     

Comparisons of genetic population structures in four intertidal brachyuran species of contrasting habitat characteristics

Genetic population structures along the Japanese coast, analyzed by sequence data from the mitochondrial DNA COI region, were determined for four intertidal brachyuran species in the superfamily Thoracotremata (Ocypode ceratophthalma, Gaetice depressus, Chiromantes dehaani and Deiratonotus japonicus), which were characterized by different habitat requirements. O. ceratophthalma (seashore; supratidal sand) and C. dehaani (estuarine; supratidal marsh) showed no significant genetic differentiation among Japanese populations. The Japanese populations of O. ceratophthalma, however, were found to genetically differentiated from the Philippine population. G. depressus (seashore; intertidal cobbles) exhibited significant genetic differentiation between the Amami-Ohshima population and other local populations. D. japonicus (estuarine; intertidal cobbles) showed significant genetic differentiation among many local populations separated by about 30–1,200 km. The different patterns of genetic population structure recorded for the four species, thus, do not simply correspond to habitat type. An erratum to this article can be found at     

Changes in barnacle population structure along an intertidal community gradient in the Florida Keys

Population distributions, densities and percentage deaths and individual size variations of the barnacles Chthamalus stellatus and Tetraclita squamosa, and the distribution of the vermetid snail Spirogluphus annulatus are measured to clarify determinants the Florida Keys (USA). The species composition of the pilings community and barnacle population densities correspond to physical gradients dependent on piling distance from shore. Interspecific competition for space is insignificant in determining C. stellatus presence or absence on pilings. Intraspecific crowding in highdensity zones of adult T. squamosa is evidenced by a significant decrease in basal diameter as density increases. The peak percentage of dead C. stellatus coincides with peak densities. The percentage of dead T. squamosa is at a low point at peak densities due to the ability of T. squamosa cyprids to settle and survive on adult shells in high-density regions. Feasible perturbation experiments for testing the importance of interspecific competition in determining the densities and vertical distributions of the species are discussed. The results of such experiments can be used to test the assumptions and predictions of the widely applied competition coefficient measure proposed by Levins (1968). Several theoretical deficiencies of Levins' measure are discussed.     

Effects of salinity and temperature on Corophium volutator and C. arenarium (Crustacea: Amphipoda), with particular reference to distribution

The responses of the post-embryonic stages of Corophium volutator (Pallas) and C. arenarium Crawford to the combined effects of salinity and temperature show that gravid females have a wider tolerance than nongravid adult females which in turn are more tolerant than adult males. C. volutator is more tolerant of low salinity (2 to 10) than C. arenarium, but the latter is more tolerant of salinities above 45. The embryos of C. volutator develop normally and hatch at lower salinities and temperatures than those of C. arenarium, in which successful development was recorded at higher temperatures. Females undergoing a pre-copulatory moult failed to lay eggs below salinities of 3 (C. volutator) and 10 (C. arenarium), but in both species the lowest salinity at which all females moulted and laid eggs was 20. The results are discussed in relation to the distribution of both species.     

Mitochondrial DNA variation in Eritrean hamadryas baboons (Papio hamadryas hamadryas): life history influences population genetic structure

The hamadryas baboon, Papio hamadryas hamadryas, represents a rare exception from the pattern of female philopatry and male-biased dispersal predominant in mammals including primates. To elucidate the possible consequences of the dispersal pattern on the population genetic structure of hamadryas baboons, we sequenced the maternally transmitted mitochondrial hypervariable region I of 74 individuals from ten sampling locations in different ecogeographic zones of Eritrea. To this end, individual fecal samples were collected at sleeping cliffs. Upon comparing the individual sequences by means of phylogenetic tree reconstructions and AMOVA, we could not detect a population genetic structure corresponding to a geographic pattern. Tree reconstructions revealed the existence of two profoundly different lineages both present at most of the sampling locations. These findings and Mantel correlations of genetic distances and the frequency of shared haplotypes to geographic distances point to the presence of female dispersal. Female-mediated gene flow is detectable over geographic distances exceeding those between neighboring subpopulations. Our study therefore corroborates local behavioral observations on a broad geographic scale. After inclusion of geographically closely situated olive baboons, P. h. anubis, in the analyses, all anubis sequences fell within one hamadryas clade. Possible scenarios leading to this situation including long-term hybridization processes are discussed.     

Climate change and population genetic structure of marine species

Climate change influences populations by reducing or extirpating local populations, by disrupting patterns of migration and by shifting geographical distributions. These events can affect genetic population structure in several ways. Molecular markers have been used in numerous population genetic and phylogeographical studies of marine species and have detected population responses to climate change in the last few decades, such as range expansions, adaptative shifts and declines or increases in abundance. Little is known, however, about the molecular and physiological basis of adaptive responses to climate change in marine Mediterranean species. The Mediterranean Sea ecosystem is a ‘living laboratory’ with native species that are challenged by environmental change and by invasive species and a ‘gene-climate’ approach should be adopted as a way of focusing on the relationship between climate warming and genetic diversity.     

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.     

Phenotypic assortative mating and genetic population structure in the crab Trapezia digitalis

Colonies of Pocillopora spp. containing the crab Trapezia digitalis were collected from Hawaii in summer of 1979 and of 1980. Individuals of T. digitalis in Hawaii vary extensively in coloration of the carapace: crabs were separated into phenotypic classes by their degree of carapace reticulation. When presented with two females of different phenotype, males preferentially paired with the female which was similar to them in reticulation. The genetic population structure of T. digitalis was unusual by comparison with other species of Trapezia. Previously reported homozygote excesses at esterase-4, hexokinase-1, phosphoglucoseisomerase, and general protein-2 cannot be explained by the single-locus Wahlund effect, since there was not significant genetic differentiation among phenotypic classes, as measured by Wright's F _ST statistic. There was significant variation among classes in allele frequencies at a fifth locus, hexokinase-2. Two pairs of loci were in significant gametic disequilibrium. Gametic disequilibrium was not detected in seven other species of Trapezia. It is suggested that multi-locus genetic analysis and an increased geographic sampling range are needed to determine the genetic population structure of T. digitalis in relation to observed variation in coloration.