Biochemical genetic variation at a leucine aminopeptidase (LAP) locus in blue (Mytilus galloprovincialis) and greenshell (Perna canaliculus) mussel populations along a salinity gradient

In several marine bivalve species, biochemical genetic variation at a leucine aminopeptidase (LAP) locus is associated with environmental variability, primarily salinity fluctuation. Population genetic variation at a LAP locus was investigated here in two sympatric mussel species (Mytilus galloprovincialis and Perna canaliculus) from three locations along a salinity gradient in Wellington Harbour, New Zealand. The data for M. galloprovincialis and P. canaliculus do not support the hypothesis that the LAP polymorphism in either species is associated with salinity variation among adult mussels. Due to the absence of small mussels among the samples it is not possible to discount the hypothesis that selection acts primarily against juveniles, as it does for M. edulis in Long Island Sound, USA. Wellington Harbour populations of M. galloprovincialis exhibited large and often highly significant heterozygote excesses at the LAP locus, whereas populations of P. canaliculus from the same locations exhibited large and highly significant heterozygote deficiencies. The reason for this inter-specific difference in population structure is unknown. If it is the result of selection, this suggests that selection acts differentially upon the two species, because demographic attributes and reproductive biology are very similar in the two species. For both M. galloprovincialis and P. canaliculus, significant levels of population genetic heterogeneity were recorded among three locations separated by only 8 to 12 km. Neither species exhibited shell length-dependent genetic variation at the LAP locus, suggesting that for these two species the LAP polymorphism is not associated with variation in shell length. Received: 30 December 1996 / Accepted: 6 January 1997     

Size-dependent, spatial and temporal genetic variation at a leucine aminopeptidase (LAP) locus among blue mussel (Mytilus galloprovincialis) populations along a salinity gradient

Biochemical genetic variation at a leucine aminopeptidase (LAP) locus is related to salinity variation in several marine bivalve molluscs. This paper details an investigation of the Long Island Sound model of LAP selection (LAP genotype-dependent mortality occurring among newly settled Mytilus edulis mussels) carried out in 1997 among three M. galloprovincialis mussel populations along the salinity gradient of Wellington Harbour, New Zealand. Significant LAP genotypic heterogeneity was observed at the LAP locus between small (<25 mm shell length) and large (>25 mm shell length) M. galloprovincialis from Petone and Eastbourne (the two sites experiencing the greatest salinity variation), whereas genotypic heterogeneity was not significantly different between small and large mussels from Seatoun (the site experiencing the least salinity variation). The Lap ³ allele decreased in frequency and the Lap ⁴ allele increased in frequency at Petone and Eastbourne, whereas the Lap ³ and Lap ⁴ allele frequencies remained effectively constant at Seatoun. Both these findings are consistent with the Long Island Sound model of selection. At all three locations, the Lap ^3,3 genotype decreased in frequency from small to large mussels, whereas the Lap ^3,4 genotype increased in frequency from small to large mussels. All other LAP genotypes occurred at low frequencies (<0.10) at all three locations and showed no evidence of frequency change from small to large-size mussels nor evidence of clinal change among the three locations. These genotype frequency data possibly indicate that the Lap ^3,3 genotype is at a selective disadvantage compared to the Lap ^3,4 genotype at all three locations, and that this selective disadvantage is related to the extent of salinity variation which exists at each location. Further investigation is required before it can be determined if the Long Island Sound model of selection best describes the size-dependent and location-specific changes in LAP allele and genotype frequencies along this salinity gradient. Comparison of the population genetic structure at the LAP locus in 1995 and in 1997 revealed a profound change from heterozygote excesses to heterozygote deficiencies for all three M. galloprovincialis populations. The reason for the change is unknown, but the change indicates that population genetic structure at the LAP locus is highly variable in time, but consistent in space, among these M.␣galloprovincialis populations. Received: 5 February 1998 / Accepted: 27 May 1998     

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.     

Advantages and a disadvantage of large size for male gammarus pulex (Crustacea: Amphipoda)

Summary The breeding system of the freshwater amphipod Gammarus pulex includes a precopulatory guarding phase by a male. The length of this guarding phase is investigated with respect to a male's size and the number and size of his competitors. As the absolute number of competitors increases, so does the guarding time but as the absolute number of available females increases, the guarding time decreases. Takeovers of the females by unpaired males are more frequent in longer precopulas (Table 2). In contests for females, larger males have two advantages over smaller males; they are better able to make a takeover (Table 2) and better able to resist takeover attempts while paired (Table 3). Males increase the length of the guarding phase as the mean size of their competitors increases (Table 4). When not paired males are usually searching for available females, perhaps in the stream current. Females are unaffected by current speed but increasing current causes decreased male survivorship (Table 5) and increased precopula duration (Table 5). Searching in currents is more dangerous for larger males than smaller ones. It is proposed that the male size distribution observed is the result of selection pressure to increase size from male-male competition balanced by selection against large size while searching for females in the current.     

Trace-metal detoxification in the ventral caeca of Orchestia gammarellus (Crustacea: Amphipoda)

The talitrid amphipod crustacean Orchestia gammarellus (Pallas) was collected from metal-contaminated (Dulas Bay, Gironde) and control (Millport) sites in the UK and France. Irrespective of site of origin, the amphipods showed the same physiological mechanism of trace-metal detoxification, involving the ventral caeca. Copper was always present in lysosomal residual bodies in the ventral caeca. Following laboratory exposure to zinc and cadmium, the lysosomes usually contained both copper and zinc but cadmium was not detectable. The lysosomal copper concentration is positively correlated to that of sulphur, while the concentration of lysosomal zinc is related to that of phosphorus. Results are interpreted in terms of the differential rates of turnover of metallothioneins chelating copper, zinc or cadmium. Received: 9 February 1999 / Accepted: 2 December 1999     

Growth and reproduction ofCaprella danilevskii (Crustacea: Amphipoda) reared in the laboratory

Caprella danilevskii Czerniavski, an epifaunal amphipod crustacean, was successfully reared for two generations under laboratory conditions. Amphipods were maintained in pairs or fours in a Petri dish. Temperature and photoperiod were maintained at 20°C, and 14 h light: 10 h dark, respectively. Colonies of the diatomCylindrotheca closterium (Ehrenberg) Reumann and Lewin were provided as food source. The red algaGelidium amansii (Lamouroux) Lamouroux was used to provide an attachment substrate for the caprellids. Juveniles that emerged from the brood pouch were described as instar I. Subsequent instars were identified by molts. Juveniles molted successively at 2.5 to 6.5 d intervals. The body length of males at each instar increased exponentially, while that of females followed a sigmoid curve. A different pattern of flagellar segment addition in the antenna I of males and females was revealed. At the age of 20.8 d, females reached instar VII, and produced their first embryos. After 5.1 d, juveniles emerged from the brood pouch. On average, females produced 5.4 clutches successively at 5.0 d intervals. The number of offspring per spawning increased from 6.3 at instar VII to 22.0 at instar XII. The total number of offspring was 69.0 for a female throughout her life span.     

Effects of cadmium on Pontoporeia affinis (Crustacea: Amphipoda) in laboratory soft-bottom microcosms

Laboratory soft bottom microcosms, containing sediment with natural populations of meio-and microfauna and the amphipod Pontoporeia affinis Lindström, were used to investigate chronic effects of cadmium on a typical Baltic Sea benthic community. Temperature, salinity, organic content of the sediment and the life cycles of the amphipods in laboratory control microcosms were in good agreement with conditions in the Baltic Sea. In Experiment 1, P. affinis was exposed to four assayed cadmium concentrations, 0 (control), 6.3, 41 and 127 g Cd·1^-1, for 265 d. At the lowest cadmium concentration (6.3 g Cd·1^-1), the sediment contained about 700 times, and P. affinis about 1 700 times, the water concentration at the end of the experiments. In microcosms without P. affinis, the sediments accumulated less cadmium. Effects on P. affinis embryogenesis were detected after 105 d of exposure in all tested cadmium concentrations. Mortality of juveniles was significantly increased after 265 d of exposure in mean assayed concentrations of 41 and 127 g Cd·1^-1. In Experiment 2, P. affinis was exposed to cadmium in assayed concentrations, 0, 6.5 and 46 g Cd·1^-1 for 460 d. At the lowest water concentration (6.5 g Cd·1^-1), the prolonged exposure did not increase the sediment concentration, which indicates a steady state, while concentrations in P. affinis reached about 3 500 times the lowest water concentration after 460 d. Juvenile P. affinis matured sexually and reproduced normally in all microcosms, but their offspring died in all microcosms with added Cd. This occurred at water concentrations not uncommon in polluted marine areas. The effects are serious in comparison to those found in conventional short-term laboratory experiments with the same concentrations and demonstrate the importance of long-term experiments in ecotoxicology.     

Interaction between solar orientation and landscape visibility in Talitrus saltator (Crustacea: Amphipoda)

A series of experiments on the littoral amphipod Talitrus saltator (Montagu) was carried out between April and September, 1978–1981, both under natural conditions and inland with artificial landscapes of different heights on the horizon, in order to assess the visual importance of the landscape in zonal orientation in populations from the Mediterranean coast, and to determine interactions between solar orientation and orientation based on the landscape. Inland, orientation in controls (permitting only vision of the sky and the sun) was compared to that of the experimental individuals, who could see a simulated landscape positioned landwards to them and seawards. In nature, sandhoppers released in a level arena with the landscape screened from view were compared with others released in an unscreened arena and on the sand in absolutely natural conditions. Situations where solar orientation contradicted local cues were produced both by releasing the sandhoppers on a shore diversely orientated to their own, and by shifting their internal clock by nine hours. Results show that T. saltator uses the landscape as a cue in its orientation towards the sea, in conjunction with solar orientation: the latter being the principal factor involved even when the sandhoppers are separated from the local optical factor in the sky (in trials at a distance from the sea). In fact, with the artificial landscape set seawards, none of the populations we studied showed any orientation based on the landscape comparable in accuracy to orientation based exclusively on the sun. From the experiments carried out in natural conditions, it is possible to deduce that conflicting conditions gave rise to two types of results: deviation from the mean direction and an increase in dispersion, up to total dispersion when the local factors were in total contrast to solar orientation (clock-shifted sandhoppers released on the sand).     

Presence of β-carotene and a number of xanthophylls in Hyale perieri (Crustacea: Amphipoda) from the Black Sea

Investigations have been made on the carotenoids in the marine amphipod Hyale perieri (Lucas) from the Black Sea. The presence of the carotenoids -carotene, canthaxanthin, lutein, neoxanthin and astaxanthin was determined by means of columnar and thin-layer chromatography. It is highly probable that violaxanthin is also present in H. perieri.     

Fatty acid composition as a dietary indicator of the invasive caprellid, Caprella mutica (Crustacea: Amphipoda)

The invasive caprellid amphipod Caprella mutica is one of the most widely dispersed marine non-native species globally. Originating in sub-boreal north-east Asia, it has now been found in both the northern and the southern hemispheres. One potential reason why this species is such a successful invader is its ability to utilise a wide variety of food sources. The contribution of different food sources to the diet of C. mutica was estimated using fatty acids as biomarkers. Caprella mutica was collected from three field sites, including sea cages stocked with Atlantic salmon Salmo salar, shellfish longlines stocked with the blue mussel Mytilus edulis and mooring lines marking the Loch Linnhe Artificial Reef (>2 km from caged finfish aquaculture), where established populations of this species are known to occur. In addition, the fatty acid compositions of C. mutica held in aquaria and either fed the microalga, Dunaliella tertiolecta, or the diatom, Phaeodactylum tricornutum, for a period of 21 days were investigated. The fatty acid composition of the diatom and the microalgal diets was also examined. The results showed that C. mutica contained high levels of polyunsaturated fatty acids, particularly 20:5(n-3); other dominant fatty acids included 18:1(n-9), 22:6(n-3) and 16:0 (in decreasing order based on abundance). Significant differences in the fatty acid profiles between caprellids fed on the microalgae and the diatom diets and between C. mutica collected from the field sites were observed. These results provide evidence that lipid biomarkers can be successfully used to provide evidence of feeding strategy for C. mutica and that the flexibility observed in this strategy may play an important role in its invasion success.     

Marine Gammaridea (Crustacea: Amphipoda) from the Indian region. Family Ampithoidae

This paper deals with 6 species of marine gammaridean amphipods belonging to the family Ampithoidae. The specimens described were collected from Mandapam (Gulf of Mannar), India and Thankasserry (Quilon, Kerala), India. The collection includes 3 new species, namely Ampithoe (Ampithoe) serraticauda, Ampithoe (Pleonexes) auriculata and Cymadusa imbroglio. Ampithoe (Ampithoe) kergueleni Stebbing (1888) is a new record for this region, while Ampithoe (Ampithoe) ramondi Audouin (1826) and Cymadusa filosa Savigny (1816), the two well-known tropicopolitan members, are briefly redescribed. The study supports Barnard's (1970, p. 23) contention that the genus Pleonexes Bate (1857) (since reduced to subgeneric status under the genus Ampithoe Leach, 1814) represents only a grade of evolution, but suggests that the structure of the 1st maxillary and mandibular palps and the outer lobes of the lower lip should also be taken into consideration, in combination with the telsonic processes, for distinguishing Pleonexes from Ampithoe. Based on this concept, a reappraisal of some of the species of the genera is suggested. The subgenus Ampithoe is created to hold those species of Ampithoe showing typical characteristics of the genus.     

Environmental influence on the locomotor activity rhythm of Talitrus saltator (Crustacea: Amphipoda)

The persistence and precision of the endogenous, nocturnal, circadian locomotor rhythm of the sand-beach amphipod Talitrus saltator is characterised, and the influence of substrate availability, photoperiod, temperature and relative humidity as potential environmental synchronizers of the rhythm is assessed. Individual, cyclic light-dark and temperature regimes synchronize and modify the laboratory activity pattern, but substrate availability and relative humidity have no significant long-term effect. Under simulateneous experimental regimes simulating field conditions of photoperiod, temperature and relative humidity the rhythm is entrained solely by photoperiod. The implications are discussed in relation to field behaviour.     

Microhabitat segregation of the amphipod genus Gammarus (Crustacea: Amphipoda) in the Northern Baltic Sea

Closely related species may occupy very similar niches but are often found to diverge by one or more traits when they share the same habitat. Five indigenous and sympatric Gammarus species are characteristic for the Baltic rocky littoral ecosystem. Yet, the species-specific distribution of these sympatric gammarids has not been well studied in the northern Baltic Sea. This study was undertaken to assess the spatial distribution of gammarid amphipods along wave exposure and depth gradients to study whether they show segregation in their microhabitat use. We sampled 12 rocky sublittoral shores along a wave exposure gradient over a period of 5 years. Samples differed with respect to depth and macroalgal type. Three of the five gammarid species occurred mainly in different depth zones and among different macroalgae at the exposed shores. In contrast, on protected shores, where algal zonation is weaker, a link to zonation and macroalgal type was almost absent. Moreover, the microhabitat use was strongest during the reproductive seasons of the species. The observed microhabitats of the three gammarid species fit well to their species-specific mean body sizes.     

Interspecific competition for mates and habitat selection in five species of Gammarus (Amphipoda: Crustacea)

Mating competition due to the formation of interspecific precopulas is involved in the evolution and maintenance of the habitat selection shown by the amphipod gammarids in the Limfjord, Denmark. The mating competition is discussed in relation to possible pheromones involved in initiating the formation of precopulas and in relation to the phylogeny of the species populations involved.     

Development of epiphytic communities on eelgrass (Zostera marina) along a nutrient gradient in a Danish estuary

The effect of nutrient enrichment on epiphyte development was examined by following the seasonal development of epiphyte biomass on eelgrass (Zostera marina L.) at four localities along a nutrient gradient in Roskilde Fjord, Denmark between March and December 1982. In the most nutrient-poor area, epiphyte biomass followed a distinct bimodal seasonal pattern with maxima in spring and early fall. Low nutrient availability and a high rate of eelgrass leaf renewal kept epiphyte biomass at a low level throughout the summer period. Unlike phytoplankton, the epiphytic community was not stimulated by nutrient enrichment during spring, however, from May through August, the biomass of both components increased exponentially with increasing concentrations of total N in the water. Along the nutrient gradient, phytoplankton biomass increased 5- to 10-fold, while epiphyte biomass increased 50- to 100-fold. Thus differences in nutrient conditions among study sites were more clearly reflected by epiphytes than phytoplankton.Contribution No. 419 from the Freshwater-Biological Laboratory, University of Copenhagen     

Swimming performance trade-offs across a gradient in community composition in Trinidadian killifish (Rivulus hartii)

The impacts of predation and competition on life history, behavioral, and morphological traits are well established for many organisms, but effects on locomotor performance have received relatively little attention. We examined variation in sprint speed and critical swimming speed (U(crit), a measure of stamina) in the Trinidadian killifish (Rivulus hartii) across a gradient of ecological communities. R. hartii are located in (1) "high-predation" sites with large, piscine piscivores, (2) "Rivulus-guppy" sites with guppies, and (3) "Rivulus-only" sites with only R. hartii. R. hartii suffer higher mortality in high-predation sites. In Rivulus-guppy sites, population densities are reduced and growth rates increased compared with Rivulus-only sites, which likely represent indirect effects of guppy predation on young R. hartii. We show a significant negative relationship, suggesting a trade-off, between sprint speed and endurance; Rivulus from high-predation sites were faster sprinters but had reduced critical swimming speeds. This trade-off was also apparent in correlations of the nine population means. At the individual level, the correlation was weaker and only significantly negative when all nine populations (three from each site) were pooled and values were not corrected for body size. Sex had a significant effect on U(crit), with females having a lower U(crit), but sexes did not differ in sprint speed. Fish from high-predation sites also exhibited increased tail lengths and fineness ratios compared to sites without large predators. The two low-predation sites showed no statistical differences in locomotor performance or morphology.     

Macroalgae as habitat patch islands for Scutellidium lamellipes (Copepoda: Harpacticoida) and Ampithoe tea (Amphipoda: Gammaridae)

Scutellidium lamellipes and Ampithoe tea live in patches formed by the brown alga Pelvetia fastigiata. The distributions and abundance of both crustaceans were examined on P. fastigiata obtained during 1975 and 1976 from 7 collecting and 6 experimental sites in the mid-intertidal zone at La Jolla, California (USA). At each collecting site, the abundance of each crustacean generally increases with increasing plant size. Following defaunation and transplantation of algae, abundances of both crustaceans on individual plants appear to be governed by equilibria determined by in situ reproduction, immigration and emigration. Immigration rates of both animal species decrease significantly with increasing distance from areas occupied by P. fastigiata. During short-term experiments, neither crustacean was able to establish colonies on defaunated plants placed across a 30 m P. fastigiata-free zone and both species would have to reproduce to persitst on plants transplanted to that site. More frequent immigration and reproduction (and, for S. lamellipes at least, decreasing emigration rates with increasing patch size) would increase abundances and decrease probabilities of local extinction on larger and/or less distant plants. However, even within sites with isolated plants, the smallest plants often are not inhabited by either crustacean. The densitics of both species differ with plant size and distribution. Densities of S. lamellipes are greatest within a large aggregation of P. fastigiata, whereas those of A. tea are greatest on moderate-sized isolated plants at mid-tide levels.     

Does oxygen deficiency modify the functional response of Saduria entomon (Isopoda) to Bathyporeia pilosa (Amphipoda)?

The functional response of the predatory isopod Saduria entomon to the prey amphipod Bathyporeia pilosa was measured in normoxia (95% O₂ saturation), moderate hypoxia (45% O₂ saturation) and hypoxia (35% O₂ saturation) in aquarium experiments. The prey densities tested ranged from 400 to 8000 ind m⁻². Prey density influenced consumption rates of S. entomon in normoxia and 45% O₂ saturation, but there was no difference between consumption rates at these two oxygen levels. Nevertheless the form of functional response differed. In normoxia S. entomon showed a positively density-dependent functional response to B. pilosa, indicating a potentially stabilizing effect on the prey population. In moderate hypoxia the variance in consumption increased, decreasing the statistical power to distinguish between response models. The functional response of S. entomon in moderate hypoxia was best described with a density-independent response, characterized as destabilizing for the prey population. In hypoxia (35% O₂) predation by S. entomon did not respond to increasing prey density, as almost no amphipods were eaten at this oxygen level. The results are discussed in terms of the usability of theoretical models to examine predator–prey relationships in stressful environments. Received: 26 April 1997 / Accepted: 20 May 1997