Life under Arctic sea ice – reproduction strategies of two sympagic (ice-associated) amphipod species, Gammarus wilkitzkii and Apherusa glacialis

The reproductive strategies of two gammaridean amphipod species, Gammarus wilkitzkii and Apherusa glacialis, that permanently inhabit the Arctic sea ice were investigated. G. wilkitzkii reaches sexual maturity at an age of 2 years and produces 128 ± 54 eggs fem.⁻¹ yr⁻¹. Mating takes place during fall and winter, and the development of the large eggs (0.60 to 0.80 mm diam.) lasts 6 to 7 months. The sex ratio of G. wilkitzkii was dominated by males in a proportion of 1.5:1. In vivo studies showed that juveniles are released in batches from the brood pouches of the females during April and May. A. glacialis reaches sexual maturity at the age of 1 year and produces 555 ± 151 eggs fem.⁻¹ yr⁻¹. The eggs are between 0.18 and 0.23 mm in diameter, and are the smallest known for gammaridean amphipods. Eggs are kept in packages of two to eight in the brood pouches of females. The sex ratio of A. glacialis was dominated by females in a proportion of 3:1. The high fecundity of both amphipod species, the release of juveniles in batches over a period of time, a high proportion of females (A. glacialis), and an elongated life-span with multiple spawnings (G. wilkitzkii) are discussed as possible adaptations to the specific and highly variable conditions under Arctic sea ice. Received: 29 December 1999 / Accepted: 8 March 2000     

Occurrence of the semi-catadromous European eel Anguilla anguilla in the Baltic Sea

Strontium (Sr) and calcium (Ca) contents in the otoliths of yellow and silver European eels [Anguilla anguilla (L.)] collected from coastal waters of the Baltic Sea and a freshwater lake in Sweden were examined by wavelength dispersive X-ray spectrometry with an electron microprobe. The mean Sr/Ca ratios from elver check to otolith edge were significantly higher for the eels from coastal waters (5.39 ± 1.09‰) than for those from the lake (0.71 ± 0.89‰). The evidence indicates that European eels in the Baltic Sea do not necessarily migrate into freshwater streams during the growth phase. Received: 30 September 1999 / Accepted: 6 April 2000     

Parthenogenetic reproduction of Diaphanosoma celebensis (Crustacea: Cladocera): influence of salinity on feeding, survival, growth and neonate production

 Effect of salinity on the feeding rate and parthenogenetic reproduction of asexual females of the cladoceran Diaphanosoma celebensis Stingelin was studied. Short-term (10 h) grazing experiments were conducted using Isochrysis galbana as feed at 5, 17, 25 and 30 psu salinity. Gut pigment concentration showed a significantly higher rate of feeding at lower salinities. Survival, growth, maturity attainment and neonate production of asexual females reared in the above four test salinities indicated preference for lower salinities (5 and 17 psu). The mean size of adult females decreased from 909 to 593 μm, mean life span from 24 to 5 d, mean neonate production from 12 to 2 and mean size of neonates from 434 to 400 μm as the salinity increased from 5 to 30 psu. Salinity variations also affected the size and age of primiparous females. Resting egg formation and sexual reproduction did not occur at the tested salinities. The results indicate that D. celebensis is adapted to low saline, estuarine environments. Received: 14 January 2000 / Accepted: 24 March 2000     

Genetic differentiation in a pelagic crustacean (Meganyctiphanes norvegica: Euphausiacea) from the North East Atlantic and the Mediterranean Sea

Meganyctiphanes norvegica (M. Sars)is a pelagic crustacean that plays a key role in marine food webs of North Atlantic Ocean and marginal seas. We studied eight population samples collected in the European Atlantic and Mediterranean Sea. By means of single strand conformation polymorphism analysis (SSCP) and direct sequencing, we investigated a segment of 158 base pairs of the mitochondrial gene coding for the subunit 1 of NADH dehydrogenase. We found 12 sequence variants among the 385 individuals studied. Analysis of molecular variance (AMOVA) showed that 14.75% of the total genetic variability was explained by differences between populations, thus indicating absence of panmixia for these populations. Pairwise comparisons revealed three distinct genetic pools: the first one represented by Cadiz Bay, the second one by the Ligurian Sea, and the third one included all the NE Atlantic samples. We also investigated one population from the Alboran Sea (within the Mediterranean basin, east of the Strait of Gibraltar). This population was found to be genetically intermediate between the NE Atlantic samples and the Ligurian sample, suggesting that the restriction to the gene flow is not associated with the Strait of Gibraltar, but possibly with the Oran–Almeria oceanographic front. The present work indicates that M. norvegica, although endowed with a high dispersal capacity because of its pelagic habit, can develop separate breeding units inside the same oceanic basin (the Atlantic). Furthermore, the Ligurian sample should be considered as a distinct evolutionary entity, separated from the Atlantic population. Received: 2 May 1999 / Accepted: 26 November 1999     

Mating behaviour of Macrophthalmus hirtipes (Brachyura: Ocypodidae)

 We examined the mating behaviour of the New Zealand ocypodid crab Macrophthalmus hirtipes in the laboratory between February and June 1998. This species has a discrete breeding season. Mating and moulting were not linked and only intermoult females with mobile gonopore opercula were attractive to males. Allometry and compatibility of gonopods and gonopores of different-sized crabs was investigated. Under laboratory conditions, the opercula of intermoult females remained mobile on average for 11.4 d, but the duration of receptivity did not appear to be under female control. The operational sex ratio in the laboratory fluctuated greatly, but was always male-dominated. During the period of opercular mobility, females mated many times with several different males. Matings in the absence of burrows were relatively short (mean duration = 23 min, max. = 122 min) and the mating behaviour of M. hirtipes lacked courtship and mate-guarding. Males used a search-intercept method to acquire mates, with very low levels of intrasexual competition. There was no evidence of mate preference in M. hirtipes, and males spent just as long mating with ovigerous females as with non-ovigerous ones. Although M. hirtipes has ventral-type spermathecae, as do several other ocypodid crabs, it is unclear whether this promotes last-male sperm precedence. The role of burrows in modifying the mating behaviour of M. hirtipes in the field remains to be established. Received: 7 January 2000 / Accepted: 5 June 2000     

Phylogeny and biogeographic history of hake (genus Merluccius), inferred from mitochondrial DNA control-region sequences

 Phylogenetic analyses of the left domain of the mitochondrial DNA control-region sequence have been used to examine the relationships among species of the genus Merluccius (Rafinesque, 1810), and to compare these with hypotheses based on morphological, meristic and allozyme characters. Analysis of aligned sequences revealed that transition bias was much lower than in mammalian mtDNA, and that nucleotide composition of control-region sequences was biased toward A and T. We have roughly calibrated a molecular clock for the genus, based on the rise of the Isthmus of Panamá, which is believed to have created a barrier to dispersal between marine species of the Atlantic and Pacific Oceans. Our mtDNA-based phylogeny was highly congruent with allozyme-based phylogenies, but poorly so with a previously described phylogeny based on morphology. Specifically, our phylogeny resolved two well-supported principal clades, one of American (west Atlantic and east Pacific) species and the other of Euro–African (east Atlantic) species. This suggests an evolutionary history during which the ancestral lineage of Merluccius was divided between two geographic regions, with subsequent dispersal and vicariant events resulting in the evolution and distribution of extant taxa. However, the relationships between some taxa within the American clade could not be resolved. We suggest that this is consistent with an hypothesis of a rapid origin and radiation of these taxa. Received: 12 December 1998 / Accepted: 15 October 1999     

Grazing rates of organic matter and living fungal biomass of decaying Spartina alterniflora by three species of salt-marsh invertebrates

 The pathway for the flow of salt-marsh grass production into marsh food-webs is still not well defined. We compared the abilities of three marsh macroinvertebrates [salt marsh periwinkles, Littoraria irrorata (Say) (=Littorina irrorata), salt-marsh coffee-bean snails, Melampus bidentatus (Say); and a talitrid amphipod, Uhlorchestia spartinophila Bounsfield and Heard] to access standing-dead leaves of smooth cordgrass (Spartina alterniflora Loisel). The invertebrates were incubated with naturally-decaying leaves, and the rates of removal of organic matter and living fungal biomass (ergosterol) were measured. The impact of invertebrate activity upon fungal growth rates was measured as rates of fungal-membrane synthesis (incorporation of radioacetate into ergosterol). The removal rates of organic leaf biomass per mg individual biomass were highest for amphipods (700 μg mg⁻¹ d⁻¹) and lowest for periwinkles (90 μg mg⁻¹ d⁻¹), but the relatively large biomass of the snails made their removal rates per individual greater than those of amphipods. Net removal of ergosterol by all three invertebrates was >50% for yellow-brown (early-decay) leaf blades. For fully-brown (advanced-decay) blades, >50% removal of ergosterol was found only for periwinkles; exposure to coffee-bean snails and amphipods resulted in a net ergosterol reduction of ≤20%. The lower net reduction of living fungal biomass by coffee-bean snails and amphipods may have been due to fungal-growth stimulation (2.3-fold stimulation in coffee-bean snails and 1.5-fold stimulation in amphipods). Grazing by periwinkles did not stimulate fungal growth, possibly because of its high intensity. Grazing by these three salt-marsh shredders may affect marsh-grass shoot-decay in different ways. Periwinkles may abbreviate the period of fungal production, and incorporate the decaying material relatively quickly into snail biomass and fecal-pellet rain to the sediments. Coffee-bean snails and amphipods may enhance and prolong fungal production, along with the formation of fecal-pellet rain. All three invertebrates fed preferentially on leaf blades rather than leaf sheaths, and feeding rates of gastropods were higher during the night than during the day. Received: 25 November 1998 / Accepted: 4 November 1999     

Cadmium accumulation in the female shore crab Carcinus maenas during the moult cycle and ovarian maturation

 The influence of moulting and ovarian maturation on cadmium accumulation in the tissues of female shore crabs Carcinus maenas exposed to 1 mg Cd l⁻¹ in the water was investigated. Cadmium accumulation in all tissues examined was markedly increased in crabs in the postmoult stages (A and B) compared to crabs in all other moult stages. During the moult cycle, average cadmium accumulation in the midgut gland ranged from 29 μg Cd g⁻¹ dw at premoult stage (D₂) to 589 μg Cd g⁻¹ dw at postmoult stage (A). Average cadmium concentrations in the haemolymph ranged from 0.56 μg Cd ml⁻¹ at intermoult stage (C₄) to 4.6 μg Cd ml⁻¹ at postmoult stage (A), while the gills accumulated from 103 μg Cd g⁻¹ dw in intermoult stage (C₃) to 352 μg Cd g⁻¹ dw in postmoult stage (A). Cadmium concentration in gills and haemolymph was also significantly higher in crabs in late premoult stage (D₃) compared to C₄-crabs, while midgut gland cadmium concentration remained elevated in C₁- and C₃- intermoult stages relative to C₄. During ovarian maturation the cadmium accumulation in midgut gland, gills, ovaries and haemolymph decreased. Average cadmium concentration in the midgut gland decreased from 63 μg g⁻¹ dw in ovarian Stage I to 19 μg g⁻¹ dw in ovarian Stage VI. The same pattern was observed for gills, haemolymph and ovaries. The present study demonstrates that cadmium accumulation in the female shore crab strongly depends on the physiological status of the animal. A possible association between physiological calcium requirements and cadmium accumulation during moulting is discussed. Received: 20 January 2000 / Accepted: 20 July 2000     

A comparative study of feeding and growth in two coexisting species of carnivorous gastropods

The effects of variable food supply on growth rate and feeding behaviour in two coexisting intertidal gastropods were examined through field and laboratory experiments involving mark and recapture procedures, growth experiments under controlled food supply and video observations of feeding. All cohorts of both Japeuthria species grew most in summer, with the highest growth rate being achieved by the youngest cohort of both sexes. In contrast, almost all cohorts (except the youngest one) showed zero to negative growth in winter. The growth rates of Japeuthria species clearly varied depending on feeding frequencies and the initial size of individuals. Females of both species had a significantly higher growth rate than males, though the difference became less marked with decreasing food supply. The total time spent on feeding and the length of the first feeding bout generally increased as the duration of the interval between meals was increased across treatments. From both the field growth experiment and the observations of feeding behaviour it is clear that J. ferrea had a higher frequency of feeding than J. cingulata in the field; 1- to 3-d intervals of feeding frequency in the former and 3- to 6-d intervals in the latter. Most notably, the two species showed a reversal in growth rates with changing conditions of food supply. When food was relatively abundant J. ferrea achieved better growth than J. cingulata, but J. cingulata grew better than J. ferrea under low food supply. In view of the fact that J. ferrea seems to be competitively superior to J. cingulata under normal levels, the present observation that J. cingulata can perform better than J. ferrea when food availability is low gives an important advantage to the latter in terms of its coexistence with the former. Received: 9 March 1999 / Accepted: 8 October 1999     

Size-related aspects of arm damage, tissue mechanics, and autotomy in the starfish Asterias rubens

 Arm damage is a widely reported but superficially investigated aspect of the biology of the starfish Asterias rubens L. In the present study, the incidence of arm damage was surveyed in populations of A.  rubens at two sites in the Firth of Clyde, Scotland, and three sites in Gullmarsfjorden, Sweden. The mean (±SD) incidence across all sites of individuals with basal arm damage (resulting from detachment at the basal autotomy plane) was 19.69 ± 8.86%, the incidence of those with distal arm damage (resulting from amputation at more distal levels) was 7.74 ± 10.01%. The mean incidence of arms with basal damage was 5.28 ± 4.12%, of those with distal damage 1.83 ± 2.45%. There was a significant negative correlation between size and the incidence of basal damage at all but one site, but no significant correlation between size and distal damage at any site. Mechanical tests on specimens of the aboral body wall from the basal region of the arm (which included the autotomy plane) and from a more distal region revealed that with increasing body size there was a significant increase in yield stress, ultimate stress and Young's modulus (stiffness) but no significant change in yield strain and ultimate strain. There was no significant difference between the relationships for basal and distal specimens. It is hypothesised that in larger individuals increased mechanical toughness replaces autotomy as an effective antipredator strategy. Using two methods to induce autotomy, a significant positive correlation between size and the delay between the onset of stimulation and arm detachment was found; this may represent a size-related decline in the efficiency of the autotomy mechanism through the relaxation of selection pressure. Since size is an unreliable indicator of age in A.  rubens, the trends identified herein can be interpreted only tentatively as age-associated phenomena. Received: 4 September 1999 / Accepted: 17 February 2000