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.     

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.     

What do caprellids (Crustacea: Amphipoda) feed on?

The present study represents the first comprehensive work dealing with the dietary analysis of Caprellidea. We studied 743 specimens of 31 genera and 62 species from all around the world. Analysis of digestive contents revealed that caprellids are mainly detritivores (detritus represented the 86% of the caprellid diet), but a small percentage of species, those lacking molars belonging to the Phtisicinae, can be considered as obligate predators and feed mainly on small crustaceans (copepods and other amphipods) and polychaetes. The contribution of sponges, hydroids, macroalgae, diatoms and dinoflagellates to the caprellid diet was very low (<2%). Apart from the clear relationship between the absence of molar process and predators, no other correlations between digestive contents and mouthpart structure were found. Cluster analysis based on the dietary data indicated a patent segregation between the Phtisicinae (obligate predators) and Caprellinae (detritivores, detritivores/predators or opportunistics) and could support the hypothesis of basal divergence in two major lines of evolution: Phtisicinae and Caprogammaridae–Caprellinae. Consequently, the debate questioning if the taxon Caprellidea is a monophyletic group or not is still open. Additional morphological, molecular and behavioral studies are necessary to define the evolutionary relationships in this group of crustaceans.     

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.     

Comparative fecundity and reproductive strategies in seventeen species of the Cumacea (Crustacea: Peracarida)

The fecundity of 17 species of cumaceans belonging to 5 families from the eastern and western temperate regions of the Atlantic and the Canadian Arctic and Subarctic Oceans are discussed separately and then compared. The species range from 0.78 to 10.11 mm (mean carapace length) with fecundities ranging from 15.1 to 168.9 (mean number of eggs brood^-1 female^-1). Interspecific comparisons show that carapace volume versus volume of brood yields a very strong relationship (r=0.99). Therefore, it is a combination of carapace volume plus egg volume which determines the fecundity in the Cumacea. A discussion of the various life-history and reproductive strategies employed by the Cumacea is included.     

Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998     

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.     

Population dynamics and reproductive biology of the commensal isopod Colidotea rostrata (Crustacea: Isopoda: Idoteidae)

Colidotea rostrata (Benedict, 1898) is the only known commensal idoteid isopod, living on and mimicking the color of two northeastern Pacific sea urchins of the genus Strongylocentrotus. The population dynamics and reproductive biology of C. rostrata on its host urchins were studied at a low rocky intertidal area in southern California (33°40N; 118°30W) from December 1984 to December 1986. Isopod populations remained relatively stable throughout the 2 yr study, with isopods inhabiting an average of 56.1% of the urchins at 6.1±0.6 (x±2 SE) isopods per urchin. Female isopods reach sexual maturity at a length of 6.8 mm. Fecundity in C. rostrata averaged 11.8±0.9 (x±2 SE) embryos per brood, and increased with female body size. Breeding occurs all year long in C. rostrata, with a main reproductive period between the warmer spring and summer months. Newly released mancas and juvenile isopods were present during all months of the study. C. rostrata differs from the free-living Idoteidae in its smaller maximum size, reduced fecundity, 1:1 sex ratio, and low juvenile mortality. These features may represent adaptations to a commensal life style that reflect a reduced mortality pressure on these isopods.     

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     

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).     

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.     

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.     

Population and reproductive biology of two sibling amphipod species from ascidians and sponges

 The population dynamics and reproductive biology of an ascidian- and a sponge-dwelling amphipod were examined. The two undescribed amphipod species, Leucothoe“ascidicola” and L.“spongicola”, are closely related to each other, and occur in ascidians and sponges, respectively, along the Florida Atlantic coast. L. “ascidicola” was abundant in solitary ascidians during fall 1997, disappeared during spring/early summer, and became abundant again in September 1998. During the time when L. “ascidicola” were absent from their hosts, a copepod became a frequent inhabitant of the ascidians but disappeared again when L.“ascidicola” returned to the ascidians in September 1998. The numbers of L.“spongicola” in sponges increased substantially during spring, when high reproductive activity was observed. Following this reproductive peak, both adult and juvenile amphipods apparently left the sponges, and during the summer amphipod numbers in the sponges were very low. Another small amphipod species, which often co-occurred with L.“spongicola”, showed less seasonal variation and was found in sponges throughout the whole study period. The percentage of ovigerous females per host unit was usually lower in the ascidian-dwelling than in the sponge-dwelling amphipods. In solitary ascidians, L.“ascidicola” amphipods usually occurred in groups of several adults, yet there never was more than one ovigerous female per ascidian. In contrast, several ovigerous L.“spongicola” females were found to cohabit in the same spongocoel. This suggests that intrasexual aggression may be stronger among reproductive amphipod females in the ascidians than in sponges. The size distributions of juvenile cohorts indicate that juvenile L.“ascidicola” remain for relatively long time periods in the parental ascidian, where they may reach sexual maturity. In contrast, in L.“spongicola”, only cohorts of very small juveniles could be identified, indicating that juveniles disperse shortly after emerging from the female's brood pouch. It is concluded that extended parental care is of very short duration or does not occur in the sponge-dwelling amphipod L. “spongicola”, possibly because fast-growing sponges with a highly branched spongocoel system do not allow long-lasting coexistence of parent-offspring groups. In contrast, the discrete character of the solitary ascidians may enhance the potential for exclusion of other species, resource monopolization by reproductive females, and furthermore for long-lasting extended parental care in the ascidian-dwelling amphipod. Groups of single parents together with cohorts of large juveniles are reported in the literature for amphipods and isopods from brachiopods, bivalves and ascidians, suggesting that these discrete biotic microhabitats may favor the evolution of extended parental care in peracarid crustaceans. Received: 30 July 1999 / Accepted: 8 May 2000     

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.     

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.     

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.     

The effects of two acanthocephalan parasites, Pomphorhynchus laevis and Polymorphus minutus, on pairing success in male Gammarus pulex (Crustacea: Amphipoda)

Parasites can decrease male mating success in host species in various ways, in particular by affecting male competitive ability for access to females. However, male-male competition can take different forms (i.e. interference vs exploitation competition) and which type of competition is most affected by parasites is not always clear. We investigated the influence of two acanthocephalan parasites Pomphorhynchus laevis and Polymorphus minutus on the pairing success of their intermediate host, Gammarus pulex, using field-based studies and complementary laboratory-based studies. We first studied male pairing success in the field using four large samples of paired and unpaired individuals collected at different dates. In three of the samples, the effects of size and parasite infection were significant, whereas for one sample only male size had a significant effect. There was no difference in size distributions between infected and uninfected gammarids. Large males were paired more often than smaller males, and uninfected males were paired more frequently than infected males, the pairing success of P. minutus-infected males being more severely affected than that of P. laevis-infected males. We then experimentally tested the ability to enter into precopula with a receptive female in the presence or absence of competitors. In competitive situations, the pairing success of P. laevis- and P. minutus-infected males was significantly lower than that of uninfected males, with pairing success being more affected in P. laevis-infected than in P. minutus-infected males. In the absence of competition, males infected with P. laevis were significantly less likely to enter into precopula compared with uninfected males and P. minutus-infected males, whereas there was no difference between uninfected and P. minutus-infected males in their inclination to pair with a receptive female. However, for both parasites, latency time to pair formation was significantly shorter for uninfected than for infected males. In a third experiment, we tested for a potential effect of vertical segregation on the pairing success of infected and uninfected males, but found no evidence for it. We conclude that infected males may be less competitive than uninfected males in competition by exploitation between males for females.     

Genetic adaptation to local habitats and speciation processes within the genus Gammarus (Amphipoda: Crustacea)

A recent event of allopatric speciation (a time span of about 4 000 years) is reported from a comparison between two guilds of amphipod gammarids from the central Baltic Sea and from the Limfjord (inner Danish waters), respectively. For at least two of the five species populations involved, a genetically fixed habitat specialization has led to the establishment of reproductive barriers. It has, however, not been possible to correlate this speciation event with electrophoretically determined isozyme changes. It is argued that the lack of correlation is due to the presence of an initial time lag before a speciation event can be detected as isozyme differences between newly formed species populations as predicted by the neutrality hypothesis for evolution at the molecular level.