Phylogeography of subterranean and surface populations of water lice Asellus aquaticus (Crustacea: Isopoda)

The water louse Asellus aquaticus is a widespread, euryoecious species, mostly uniform throughout its range. However, six subspecies are known from the Dinaric karst in the northwestern Balkans. They include some specialized subterranean populations. The pattern of genetic variation among subterranean and surface populations in this hydrographically highly fragmented karst region was investigated using a 653 bp fragment of the mitochondrial gene (COI). Sequencing of 168 individuals from 25 localities revealed 72 haplotypes. amova and methods of phylogenetic reconstruction all uncovered hydrographic structuring of genetic variation of the populations. Nested clade analysis pointed out several fragmentation events, along with some range expansions within hydrographical systems. By superimposing the subterranean mode of life on the phylogeographical pattern, three independent cave colonizations could be inferred within a distance of < 100 km. Caves were invaded after the ancestral surface populations became isolated through vicariant fragmentation. A possible scenario of hydrographic history of the region was constructed combining the molecular data with palaeogeographical information.     

Resource-mediated effects of stream pollution on food absorption of Asellus aquaticus (L.) populations

The role of interactions between chemical perturbations and biological constraints on detritivores occurring in polluted streams were investigated by analysing food absorption variation with stress. Absorption rate and efficiency of four Asellus aquaticus (L.) populations from differently polluted habitats were quantified with respect to the microbial guilds colonizing detritus. A twin tracer method was used. Detritus was microbially colonized in standard conditions and on each stream bottom to control for potential resource-independent variations among individuals. The relationship between length and weight was also determined on a random sample of individuals of each population. Differences of 14.6% in potential absorption efficiency and 11.3% in potential absorption rate were observed between populations from the least and the most polluted habitat. Actual (realized) variations were much stronger: from a minimum of a 60.1% reduction in absorption efficiency to a maximum of 93.8% for the rate. The realized food absorption and the individual weight per length showed the same pattern of variation among populations. This suggested that the availability of energy to isopods in nature was related to stream pollution and resource quality. Bottomup interactions appear to be the most relevant pathway through which chemical water pollution affects the Asellus populations studied. The potential resource-independent variations among individuals are also likely to be explained by temporal cascading of resource-mediated effects.     

Toxicity of heavy metals to Asellus aquaticus (L.) (Crustacea,Isopoda)

Effects of heavy metals on the isopod Asellus aquaticus (L.) are studied by static toxicity tests. Results demonstrate that the species is sensitive to Cd⁺², Cr⁺⁶, Cu⁺², Fe⁺³, Hg⁺², Ni⁺² Pb⁺² and Zn⁺², but the toxicity of each metal is different. Differences are also found between adults and between adults and juveniles. The comparative analysis of all data on the toxicity has been performed on the concentrations of metal ions and not on metal compound concentrations.Criteria for establishing water quality in order to guarantee protection of the environment are discussed.     

Geography of morphological differentiation in Asellus aquaticus (Crustacea: Isopoda: Asellidae)

We implemented a detailed morphometry and multivariate statistics to establish a general, large-scale racial differentiation in Asellus aquaticus (L.) sensu Racovitza. We ascertained that in surface populations a set of 11 morphometric characters might equivalently be represented by the pleopod respiratory area size alone. The analyses resulted in a distinct distribution pattern, with the large respiratory area populations disposed mainly along the Dinaric karst between southern Slovenia and western Macedonia and surrounded by the medium respiratory area morph, spatially irregularly substituted by the small area morph. This pattern is in contradiction with the distribution pattern of molecularly defined clades (as shown by Verovnik et al. 2005 ). We could find no ecological, hydrographical or paleogeographical explanations for such distribution pattern either. The only hypothetical explanation would be a preservation of the large respiratory area as a plesiomorphic character in the comparatively sheltered karst habitats, while throughout the easier accessible parts of the species range it was replaced by the 'modern' smaller area size. While a diminution of the respiratory area functionally means an increased sclerotization – hardening of pleopod IV–V exopodites, endopodites of pleopods III–V remain less sclerotized, probably respiratory and osmoregulatory functional.     

Effect of chronic exposure to cadmium and copper on Asellus aquaticus (L.) (Crustacea,Isopoda)

The effects of chronic exposure to 5 g·1^–1 cadmium or copper on the crustacean Isopod Asellus aquaticus (L.) were studied by analyzing survival and body growth in the first stages of the life-cycle and by determining fecundity and survival of embryo-bearing females. Juveniles survival is differently affected by the two metals in that embryonic development is more sensitive to cadmium while juvenile development is more sensitive to copper. Juvenile body growth is stimulated by cadmium and depressed by copper. Embryo-bearing female survival and fecundity are significantly reduced by cadmium but are not affected by copper. The consequence of environmental contamination by a sublethal cadmium or copper concentration is discussed.     

Sexual selection, antennae length and the mating advantage of large males in Asellus aquaticus

In crustacean species with precopulatory mate-guarding, sexual size dimorphism has most often been regarded as the consequence of a large male advantage in contest competition for access to females. However, large body size in males may also be favoured indirectly through scramble competition. This might partly be the case if the actual target of selection is a morphological character, closely correlated with body size, involved in the detection of receptive females. We studied sexual selection on body size and antennae length in natural populations of Asellus aquaticus, an isopod species with precopulatory mate guarding. In this species, males are larger than females and male pairing success is positively related to body size. However, males also have longer antennae, relative to body size, than females, suggesting that this character may also be favoured by sexual selection. We used multivariate analysis of selection to assess the relative influences of body size and antennae length in five different populations in the field. Selection gradients indicated that overall body size was a better predictor of male pairing success than antennae length, although some variation was observed between sites. We then manipulated male antennae length in a series of experiments conducted in the laboratory, and compared the pairing ability of males with short or long antennae. Males with short antennae were less likely to detect, orient to and to pair with a receptive female compared to males with long antennae. We discuss the implications of our results for studies of male body size and sexual dimorphism in relation to sexual selection in crustaceans.     

Sexual selection, antennae length and the mating advantage of large males in Asellus aquaticus

In crustacean species with precopulatory mate-guarding, sexual size dimorphism has most often been regarded as the consequence of a large male advantage in contest competition for access to females. However, large body size in males may also be favoured indirectly through scramble competition. This might partly be the case if the actual target of selection is a morphological character, closely correlated with body size, involved in the detection of receptive females. We studied sexual selection on body size and antennae length in natural populations of Asellus aquaticus, an isopod species with precopulatory mate guarding. In this species, males are larger than females and male pairing success is positively related to body size. However, males also have longer antennae, relative to body size, than females, suggesting that this character may also be favoured by sexual selection. We used multivariate analysis of selection to assess the relative influences of body size and antennae length in five different populations in the field. Selection gradients indicated that, overall, body size was a better predictor of male pairing success than antennae length, although some variation was observed between sites. We then manipulated male antennae length in a series of experiments conducted in the lab, and compared the pairing ability of males with short or long antennae. Males with short antennae were less likely to detect, orient to, and to pair with a receptive female compared with males with long antennae. We discuss the implications of our results for studies of male body size and sexual dimorphism in relation to sexual selection in crustaceans.     

Sensitivity of Asellus aquaticus (L.) and Proasellus coxalis Dollf. (Crustacea,Isopoda) to Copper

The effects of Copper on the life-cycle of two crustacean isopods, Asellus aquaticus (L.) and Proasellus coxalis Dollf., are studied. ST₅₀ indicates that females and males are differently sensitive in comparison with juveniles. The two species do not show differences in sensitivity in the range of 0.01–15 mg·1^-1 between adults and juveniles. The effect of 0.005 mg·1^-1 Copper sulphate was strongest on embryonic and larval stages. Ecological implications are discussed.     

The ultrastructure of the heart of Oniscus asellus L. and Asellus aquaticus L. (Crustacea,Isopoda)

Summary The endocardium of Oniscus asellus L. and Asellus aquaticus L. consists of lipid cells. The epicardium consists of a layer of cells with a vesiculated cytoplasm covered by a thick extracellular fibrous sheet. The myocardium is a single layer of cells, the sarcolemma invaginates at Z disc level forming transverse tubules, and longitudinal tubules branch off from these. At the A-I level' longitudinal tubules form transverse systems, which form couplings with the sarcoplasmic reticulum. The sarcoplasmic reticulum appears as perforated sheets enveloping the myofibrils. Two types of nerve terminal are found: one is embedded in a myocardial cell process, the other lies in a myocardial cell depression. They contain clear and dense-cored synaptic vesicles.This work was supported by grants from the Norwegian Research Council for Science and the Humanities     

A Three Years' Study of Life Cycle,Population Dynamics and Production of Asellus aquaticus L. in a Macrophyte Rich Stream

Populations of A. aquaticus were sampled quantitatively in 1979–82 at two localities in the river Suså which has a slope of <1 m km⁻¹ and large variations in macrophyte biomass, discharge and stream velocity. The latter two differed significantly between years. A. aquaticus had (1)−2 life cycles per year and summer and winter cohort production intervals of 85 and 290 days. Populations of A. aquaticus varied between 0 and 28,000 ind. m⁻² with an exponential increase in the spring and an exponential decrease in the autumn-winter. The A. aquaticus rate of decrease varied between 0.62 and 5.61 % d⁻¹ and increased with increasing rate of elimination of the macrophyte biomass Differences between the two localities were due to differences in physical heterogeneity. Production varied between 2.8 and 9.1 g DW m⁻² yr⁻¹ and between 12.9 and 55 g DW m⁻² yr⁻¹ at the two localities and P/B ratios were 7.4–9.9 yr⁻¹. Physical limitations are thought to be most important for the populations of A. aquaticus in the Suså, and the macrophyte biomass played an important role in modifying the physical environment. Differences between streams and lake populations are discussed.     

A comparison of the population ecology of Asellus aquaticus (L.) and Asellus meridianus rac. in the reed beds of the Tjeukemeer

The population ecology of Asellus aquaticus and Asellus meridianus was studied in a Tjeukemeer reed bed from March to October. The densities of the two species are similar at the beginning and end of the reproductive season, although A. meridianus is up to 3 times more abundant in between. In both species egg production is positively correlated to body length and summer brood sizes are smaller than in the spring. A. meridianus begins to reproduce at smaller sizes than A. aquaticus and is also more fecund for overlapping size classes. Over the season as a whole both species have the same reproductive output. The life cycles of the two species are very similar with three main periods of reproduction and the populations being replaced twice during the year. Despite the differences between the species no factors are found which gives one species an obvious competitive advantage against the other.Henry and Magniez (1970) have proposed the name Proasellus meridianus for this species.     

Light-dark cycle and mitotic index in Asellus aquaticus (L.) (Crustacea,Isopoda)

The circadian variation of the mitotic index during spermiohistogenesis was studied in Asellus aquaticus (L.). The actual number of metaphases and prometaphases was determined at the end of each hour of light or darkness over a 24 h period in animals bred under LD 12:12. The number of the metaphases and prometaphases decreases during the light period and sharply increases in the last 3 hrs of the dark period. This variation in the proliferative activity suggests that photoperiod can play a role in the synchronization of mitosis.     

Comparative ecology of Gammarus pulex (L.) and Asellus aquaticus (L.) II: fungal preferences

This paper continues to explore niche differentiation in Gammarus pulex and Asellus aquaticus by analysis of their food preferences. Individuals from both species discriminated between leaf discs colonized by different fungal species and exhibited strong preferences for Anguillospora longissima and Heliscus lugdunensis. Fungal preferences were not correlated with the relative abundance of fungi in the field and there was considerable intra-population variability in food preferences — both between individuals and for the same individual through time. Niche overlap between animals from all four study populations was high and there was no evidence of differences in the potential trophic niches of animals from sympatric and allopatric populations.     

Comparative ecology of Gammarus pulex (L.) and Asellus aquaticus (L.) I: population dynamics and microdistribution

Gammarus pulex and Asellus aquaticus generally occupy different zones in rivers; the former occurs in upper reaches but is replaced by the latter in lower reaches. Microdistribution and life-history patterns of G. pulex and A. aquaticus in sympatry and allopatry, were analyzed. Both species exhibited similar patterns of microhabitat selection, with larger individuals associated predominantly with large-sized substratum particles, and juveniles with weed. Coexisting populations of G. pulex and A. aquaticus had similar densities and population dynamics. Within each species, differences in population dynamics of allopatric and sympatric populations were observed. Although variation in population dynamics of G. pulex may be explained in terms of competition between the two species, the evidence is weak and equivocal. Differences in the dynamics of the two A. aquaticus populations were possibly a consequence of coal-mine and organic pollution, reducing the survival of offspring in the allopatric population.     

The predation on Asellus aquaticus (L.) by perch,Perca fluviatilis (L.), in a small forest lake

Asellus aquaticus (L.) was the most important benthic food item for perch, Perca fluviatilis (L.), in a small, extremely humic forest lake in southern Finland. The proportion of A. aquaticus in the diet of perch varied according to the former's availability, which, in turn, depended on its life cycle. Perch 11.0–12.9 cm (total length) most frequently fed on A. aquaticus; smaller and larger perch fed more frequently on crustacean zooplankton and aquatic insect larvae, respectively. The high density of the perch population, the importance of A. aquaticus in the benthos of the study lake and the general high activity level of the prey resulted in a high predation (0.1–1.8% per day). The population of A. aquaticus was also limited by the scarcity of macrophytic vegetation and the small area of oxygenated littoral in the lake.     

Random Amplified Polymorphic DNA Diversity among Surface and Subterranean Populations of Asellus aquaticus (Crustacea: Isopoda)

The ecological and evolutionary processes leading to isolation and adaptation of cave animals compared to their surface ancestors are not yet unequivocally understood. In this study the genetic relations of four cave and three surface population of the freshwater crustacean Asellus aquaticus in the Karst region of SW Slovenia and NE Italy were assessed using RAPDs as genetic markers. The results suggest that specialized populations from two caves invaded their subterranean habitat independently, and that their morphological similarity is a result of convergent evolution. Another, less specialized cave population seems to originate from a later colonization of a cave system already inhabited by a more specialized population, but the two populations do not interbreed. This series of temporally and spatially independent invasions has generated a diversity hotspot of non-interbreeding populations of a ubiquitous freshwatercrustacean, which is uniform over most of its range. Genetic variability estimated by the percentage of polymorphic RAPD fragments was similar (86–91%) in most cave and surface populations. Substantially lower values (as low as 49%) were found in two cave populations affected by heavy pollution. Two a priori groupings of populations, traditional subspecies and hydrologically connected groups, were rejected as not significant by means of nested analysis of molecular variance (AMOVA). On the other hand, groupings revealed by UPGMA clustering displayed a significant component of among-group variance. An analysis of gene flow between populations using estimated migration rates confirmed these findings.     

Utilization of grass carp faeces by the IsopodAsellus aquaticus (L.) in the laboratory

Grass carp (Ctenopharyngodon idella Val) faecal pellets derived fromElodea nutallii consumption were efficiently utilized by the detritivoreAsellus. Immature and adult individuals of 2.1–3.2 mm and 4.9–6.0 mm respectively, exhibited a high mean specific growth (9.03%/day and 3.33%/day) when offered faecal pellets as food during two weeks experimental trials. Individuals of the same groups, when fed onElodea which had been mechanically fragmented to simulate the macerating action of the grass carp pharyngeal teeth, grew at rates (9.57%/day and 4.06%/day) not significantly different from those fed on faecal pellets. However, the growth rates of animals fed on freshElodea (6.40%/day and 2.23%/day) were significantly lower than in the above trials, indicating that the morphological destruction of the plant tissue could be responsible for the higher observed growth. The control trials, including unfed animals, showed negative growth (– 0.49%/day and – 0.48%/day).Grass carp herbivory makes a new food resource available to benthic invertebrates by converting fresh plant material to small plant fragments.     

Comparison between some epigean and hypogean populations of Asellus aquaticus (Crustacea: Isopoda: Asellidae)

Samples for benthic meiofauna were collected in the vicinity of a salmon aquaculture farm in Bliss Harbour, Bay of Fundy, Canada in early August 1990. Simultaneously, samples for water content, organic carbon, organic nitrogen were collected, and redox potential and benthic oxygen consumption were measured. Meiobenthic size-spectra of biomass and respiration (calculated using allometric equations) were examined at three locations along a gradient of sediment organic enrichment radiating from the farm. Neither biomass nor respiration size-spectra were significantly different between locations despite a decrease in taxon diversity with increasing sediment organic enrichment. Small nematodes were the single largest contributor to respiration and usually to biomass at all locations, particularly at the most organically enriched location directly under the salmon farm. Calculated meiofauna respiration accounted for a greater proportion of total benthic community respiration in organically enriched sediments than in less enriched sediments.