The influence of intrinsic and extrinsic factors on developmental parameters and their relationships in the marine isopod Idotea linearis (Crustacea)

Developmental and reproductive parameters and their relationships were studied in the marine isopod Idotea linearis. We hypothesized that (1) the temporal patterns of molting and growth undergo complex and sex-specific changes with age as well as with the onset of sexual maturation, and that (2) sexual maturation (and dependent parameters) is controlled by the photoperiod. Both males and females were singly cultured in the laboratory at two alternative photoperiods (constant long and short days, respectively) from hatching until death. Males molted and grew throughout their life, showing a steady increase in stage duration and body size with each molt. Females, in contrast, showed much more complex modifications in molt chronology due to reproductive demands. There was some variability in the stage number, when females reached maturity. Reaching maturity early in the succession of molts was associated with smaller body size at maturity, smaller size of broods, but higher average number of broods per lifetime. Post-puberty molts in females occurred without further growth, and successive broods did not differ in size. The photoperiod strongly affected sexual maturation (and thus in turn molting and growth patterns) in females, while males remained completely unaffected by the photo regime.     

Costs of intersexual conflict in the isopod Idotea baltica

In sexual reproduction one sex can increase its reproductive success at the cost of the other, a situation known as intersexual conflict. In the marine isopod Idotea baltica, males guard females before copulation. The guarding phase is preceded by struggles as females resist males’ attempts to initiate guarding. We determined whether the struggle and/or mate‐guarding result in fitness costs in the form of decreasing fecundity and lower levels of the energy storage compounds, glycogen and lipids. Females that underwent the period of struggles with males had decreased glycogen levels compared with females maintained alone. No such cost was found for males. Females guarded by a male also had smaller eggs than females that were not guarded. Thus the intersexual conflict, imposed by the fitness maximization strategy of the males, gave rise to both a fecundity cost and an energetic cost for females. The fecundity cost confirms the existence of intersexual conflict in I. baltica. This cost is shared by males, suggesting that the intersexual conflict restrains the reproductive output of both sexes.     

Differential predation on sexes affects colour polymorphism of the isopod Idotea baltica (Pallas)

Variable selection, including spatio-temporal variation, frequency-dependent selection and differential selection due to habitat choice, may maintain polymorphism in heterogeneous environments. We studied predation as a selective agent on colour polymorphism of the aquatic isopod I baltica. Variable predation on this species can arise from at least three sources. First, apostatic selection was studied by testing the formation of preferences on colour morphs in the perch, a common predator of I baltica. Such acquired preferences should induce apostatic selection. While our results indicate some acquired preferences, there was significant heterogeneity in the behaviour of predator individuals. Second, temporal variation in selection can arise due to habitat shift from the green algae juvenile habitat to the bladderwrack adult habitat, and the consequent change in the crypsis of the morphs. Different crypsis between sexes probably promoted high predation mortality among females in the juvenile habitat. The high rate of male mortality during the breeding period, on the other hand, was presumably due to their high mate-searching activity. Third, the sex-dependent habitat choice of I baltica leads to sexual differences in the susceptibility of morphs to predation. Predators preferred the white-spotted morph over the uniform one in males but not in females, supporting the 'dimorphic niche' hypothesis as an explanation of sexual differences in morph frequencies. Finally, no evidence was found that the colouration patterns were under sexual selection. We therefore conclude diat variable predation is the most promising explanation for the maintenance of polymorphism in I. baltica.     

Electrophoresis study of hemocyanin of the polytypic species Idotea chelipes and Idotea balthica basteri (Crustacea Isopoda)]

Hemocyanin has been used as a protein marker for Crustacea speciation. The specificity of the hemocyanin fractions of two Idotea species has been confirmed by mono and bidirection electrophoresis. Four protein fractions have been found in the two Idotea species. However, the molecular weight of each Idotea chelipes fraction is slightly higher. The two fractions of high molecular weight are strongly associated. We assumed a molecular weight of 240,000 for I. chelipes and 230,000 for I. balthica basteri. The two other protein fractions are well separated and have a weight of 130,000 and 77,000 in I. chelipes; 120,000 and 75,000 for I. b. basteri. In contrast, the subspeciation of I. chelipes has not been clearly demonstrated by electrophoresis of the hemocyanin.     

Male Choice and Male-male Competition in Idotea baitica (Crustacea,Isopoda)

Sexual selection in mate-guarding Crustacea may involve several processes: male choice, male-male competition, and female choice. To evaluate the relative importance of the different processes in mate choice of the aquatic isopod I. baitica we studied 1) the mate-choice criteria of males, 2) effects of sex ratio on the outcome of the mating contest, and 3) the role of size in male-male interactions. When given a choice between a small and a large female, males most often chose the one that matured earlier for parturial ecdysis. Maturity was a more important choice criterion than female size, but these also correlated positively. Large males had a mating advantage in both male- and female-biased sex ratios; pairing was size-assortative only in the male-biased ratio where guarding was also longer. If an extra male was placed with a precopulatory pair, 30 % take-overs occurred, large males surpassing. Present and earlier work suggests that male size is an asset in both intra- and intersexual interactions. There is little or no direct phenotypic sexual selection on female size: sexual selection for large males presumably contributes to the evolution of sexual size dimorphism in I. baitica.     

Evolutionary biogeography of the Microparasellid isopod Microcharon (Crustacea) in the Mediterranean Basin

The interstitial subterranean isopod Microcharon (Crustacea) is highly diversified within the Mediterranean. The present distribution of 70 species is considered the result of the joint history of this genus and its environment. This stygobiont is derived from marine surface ancestors. Both Plate Tectonics and the two-step model of colonization and evolution, the second phase of which represents vicariance during the Tethys regressions, enable an understanding of the evolutionary history of this monophyletic group. The single extant marine littoral species was separated from its sister group by the closure of the Gibraltar Straits during the Messinian. The drift of the Apulian plate, the Alboran-Kabylian-Calabrian fragment as well as the opening of the western Mediterranean through the translation of the Corsican-Sardinia plate could explain the divergences in the western and the eastern Mediterranean. The primitive sister groups in Morocco and in Spain would have been left by the Turonian Tethys regression. Such regressions of the Tethys embayments could have played a major role in the evolution of the genus from Turanian up to Tortonian in Norht Africa, and up to Pliocene and even to Pleistocene in Italian, Iberian and Greek zones. Several eastern European species may be a product of the Paratethys regressions. Everywhere, species with plesiomorphic characters are related to old regressions and vice versa. Local geological events could have resulted in further vicariance divergences. There is a good congruence between the evolutionary history of Microcharon and other stygobiont isopod, amphipods or syncarids living in the same regions studied in the Mediterranean.Presented at the Symposium of Biogeography at Subterranean Crustaceans: the effects of different scales (June 1992; Charleston, South Carolina, U.S.A.). Manuscrips prepared for print by J. R. Holsinger.     

Male dimorphism in the harem-forming gnathiid isopod Elaphognathia discolor (Crustacea: Isopoda)

Previously unreported males of a gnathiid isopod were found in reproductive aggregations of the harem-forming gnathiid Elaphognathia discolor. Although the male gnathiids were small in size and morphologically different from E. discolor males, the male sexual organ, appendix masculina, was similar to that of E. discolor males, and possible conspecific larvae and females of the small male gnathiid were never found. In the laboratory, the small male gnathiids as well as male E. discolor successfully copulated with female E. discolor, and the development of embryos in female brood pouches was observed. Offspring of small male gnathiids develop to adults of E. discolor after molting three times, or small male gnathiids after molting two times. Thus, the small male gnathiid was concluded to be an alternative male form compared to the regular large male form of E. discolor. This male polymorphism was thought to have a genetic basis, since no small male specimens appeared in offspring of regular E. discolor males. Field sampling showed that a regular large male formed a harem composed of one large male and several females and never coexisted with other large males as previously reported. However, small males were often found together with large males. Therefore, small males are thought to be sneakers intruding into harems dominated by large males.     

Postembryonic eye growth in the seashore isopod Ligia exotica (Crustacea,Isopoda)

The eye of Ligia exotica is of the apposition type and has open rhabdoms. The facets are hexagonal, and the dioptric apparatus consists of a flat cornea and a spherical crystalline cone placed in the center of two large cone cells. Each ommatidium has seven regular retinula cells and one eccentric cell; a basement membrane forms the proximal boundary of the retina. With increases in body size from 0.6 to almost 4.0 cm, facet numbers and ommatidial diameters increased from 800 to 1500 and 35 microm to 100 microm, respectively; eye length and width grew from 1.2 to 3.2 and 0.9 to 2.5 mm, respectively; and length of dioptric apparatus and width of retinal layer changed from 70 microm to 180 microm and about 70 microm to 120 microm. Visual angles and interommatidial angles of centrally located ommatidia remained constant at about 30 and 6.9 degrees, respectively. An almost perfect linear relationship was found when eye length was plotted against the product between the square root of the total number of ommatidia and the ommatidial diameter. No difference between males and females was observed in any of the relationships, but the results suggest that, compared with smaller specimens, larger ones possess increased absolute sensitivity in single ommatidia, increased sensitivity to point sources, and overall larger angular visual fields for the eye in its totality. This means that larger individuals of L. exotica (which are also faster) have an advantage over smaller individuals at night, but that smaller individuals may cope better with bright lights. Vision in L. exotica seems useful not only in detecting potential danger, but also in locating and approaching cliffs from a distance of 2-4 m when swimming in seawater.     

Burrow plasticity in the deep-sea isopod Bathynomus doederleini (Crustacea: Isopoda: Cirolanidae)

We investigated whether the deep-sea isopod Bathynomus doederleini has the capacity to change burrow length in response to changes in environmental conditions. We observed burrowing behavior in individuals that were placed on substrates with either simple (ST) or complex (CT) surface topographies. Individuals in the ST group (N = 10) constructed seven burrows. The mean ratio of the burrow length to body length was 1.8. The individuals in the CT group (N = 10) constructed eight burrows with a mean ratio of burrow length to body length of 2.5. Thus the burrows were significantly longer in the CT group. In addition, the isopods in the CT group often incorporated a chamber in the mid-section of the burrow. Our results may be used to infer the determinants of burrow morphology and speculate about the lifestyle of this species in the deep sea.     

Characterization of polymorphic microsatellite loci in the marine isopod Sphaeroma terebrans (Crustacea, Isopoda)

Nine polymorphic microsatellites were characterized in the marine isopod Sphaeroma terebrans (Isopoda, Sphaeromatidae) for phylogeographic and parentage analyses. The number of alleles ranged from 3 to 10. Observed heterozygosity ranged from 0.428 to 0.950, while expected heterozygosity ranged from 0.532 to 0.889. Heterozygote deficiency was detected for one locus, possibly the result of null alleles.     

Natural history of the Pan-American sand beach isopod Excirolana braziliensis (Crustacea: Malacostraca)

A population of the intertidal isopod Excirolana braziliensis was followed over a 10-month period at Naos Island, Panama. Average intertidal density was 2147/m² with maximum abundance occurring in the highest level. Adults were present only in the highest tidal level. Juveniles were distributed throughout all tidal levels and comprised at least 70% of the population at all times. Mean size of individuals decreased from the highest level seaward. Reproductive females were present the year round. The number of young ranged from four to 17 per brood and was dependent on the length of the female. Adults were very tolerant of changes in temperature and salinity.     

Food and habitat choice of the isopod Idotea baltica in the northeastern Baltic Sea

The isopod Idotea baltica is the most important benthic herbivore in the Baltic Sea. There exists a significant correlation between the distribution of the adult isopod and the belts of bladder wrack Fucus vesiculosus. However, following the eutrophication induced blooms of the filamentous macroalga Pilayella littoralis and the disappearance of F. vesiculosus a notable increase in idoteid abundances has been observed. The aim of this paper was (1) to evaluate experimentally whether F. vesiculosus provides either food, shelter or both to the isopods and (2) to estimate the role of associated filamentous algae in the habitat selection process. Amongst six abundant macroalgal species, about 50% of isopod population was attracted to F. vesiculosuscovered with the filamentous algae P. littoralis. The majority of the remaining part of the population was either swimming freely or attracted to non-epiphytic P. littoralis. When both live algae and artificial substrata were provided, P. littoralis growing on artificial substrata was clearly preferred by the isopods over epiphyte-free F. vesiculosus. In the grazing experiment where I. baltica was allowed to choose between F. vesiculosus and P. littoralis the latter contributed practically 100% of the diet of the isopod. The results indicate the importance of P. littoralis as a food item and F. vesiculosus as a shelter for I. baltica.     

Ultrastructure of the pleopods of the estuarine isopod Cyathura carinata (Crustacea: Isopoda: Anthuridea)

Summary The exopodal branches of the pleopods of Cyathura carinata have a thicker cuticle than the endopodites, but the hypodermis proved to be very thin and probably of respiratory function. The nuclei are found in pilaster areas together with stabilizing elements (micro-tubuli attached to cuticular pegs), which are typical for hypodermal tendon cells of other arthropods. Precipitation of Ag₂S revealed that the cuticle of the exopod is more permeable to silver ions than the cuticle of the endopods. The endopodal epithelia have the features of typical osmoregulating tissues (apical folds, mitochondrial pumps, and basal invaginations of the plasma membrane) and reduce silver nitrate. The stabilizing pilaster areas are composed of the same elements as in the exopods.     

Dynamics of intersexual conflict over precopulatory mate guarding in two populations of the isopod Idotea baltica

Aggressiveness during intersexual conflicts is predicted to depend on its costs, the value of winning and the power asymmetry of the contestants, all of which may vary between populations. In the marine isopod Idotea baltica (Pallas) a conflict occurs as females resist the attempts by males to start precopulatory mate guarding. We analysed contest dynamics with respect to female maturity stage, that is, to time left to reproductive moult, with which the payoffs of guarding for males and females change. We did this in two populations that differ in synchrony of reproduction, sex ratio and the degree of sexual dimorphism. The intensity and dynamics of contests differed between populations: in the more size-dimorphic population, females, the smaller sex, resisted less by forceful flexing but more by hooking their body than in the other population. Male aggressiveness stayed at a constant level with respect to female maturity. In the less size-dimorphic population, female resistance by flexing was intense and it decreased, while male persistence increased, with the approaching reproductive moult. Contests were more intense with small than with large males. These results fit well with the predictions from models of conflict behaviour. Assessment of the payoffs of winning versus contest costs, coadaptation of the level of aggressiveness to the other traits affecting contest outcome, and counteradaptations by the sexes to each other largely explain the dynamics and between-population differences of these contests. Copyright 2000 The Association for the Study of Animal Behaviour.     

A new genus of phreatoicidean isopod (Crustacea) from the north Kimberley region, Western Australia

A new genus and species of phreatoicidean isopod, Crenisopus acinifer, has been collected from a freshwater spring in the northern Kimberley region of Western Australia. Empirical cladistic analysis of 10 exemplars of phreatoicidean genera found a single cladogram. The new genus and species assumed a basal position in the Phreatoicidea, placing it within the family Amphisopodidae sensu lata. This family, however, was not monophyletic in the preliminary cladogram, suggesting that the taxonomic structure of the suborder must be revised. The cladogram provided evidence for the monophyly of the Phreatoicidae and its New Zealand clade. The analysis suggested that clades of modern phreatoicideans diverged from one another during the Mesozoic Era after they entered fresh water, but prior to the fragmentation of East Gondwana.