Experimental studies on physiological and behavioural response mechanisms of Nitocra spinipes (Crustacea: Harpacticoidea) from brackish-water rockpools

Volume regulation and salinity-preference tests have been made on Nitocra spinipes Boeck (Crustacea, Harpacticoidea). This species is often dominant in Baltic brackish-water rockpools. The investigation attempts to evaluate the relative importance of some of the different response mechanisms of this species to salinity changes in relation to the unstable environmental conditions of the rockpools. Volume-regulation experiments have shown that N. spinipes is capable of hypoosmotic and probably hyperosmotic regulation in the tested salinity range of 1 to 20 S. In laboratory cultures, reproduction, hatching and moulting oecurred in salinities ranging from 0.5 to 30 S. Preference experiments showed that N. spinipes has a very weak behavioural response even to very large variations in salinity concentrations between the alternative; a significant choice could only be found under conditions which never occur in the natural biotope. It is therefore concluded that, in a biotope such as the rockpool, where salinity changes will affect the whole biotope rather than produce microclimatic variation, regulation and adaptation must have a higher ecological importance than escape responses.     

Molecular phylogeny of Ostracoda (Crustacea) inferred from 18S ribosomal DNA sequences: implication for its origin and diversification

Nucleotide sequences for the 18S rDNA of 17 ostracod species, representing 4 extant orders and 12 superfamilies, were determined and compared with known sequences of other crustaceans. Resulting molecular phylogenetic trees based on maximum-likelihood, maximum-parsimony, and neighbor-joining methods consistently indicated that the Podocopida plus Platycopida forms a monophyletic group, but did not indicate monophyly of the Ostracoda consisting of the four orders, Podocopida, Platycopida, Kirkbyocopida, and Myodocopida. The tree topology and the crustacean fossil record suggest that the lineage of podocopidans and platycopidans, that of kirkbyocopidans, and that of myodocopidans should have separately been established already in the Cambrian, although there is no ostracod fossil record in that time period. Close relationships between the Bythocytheroidea and Cytheroidea and among the Macrocypridoidea, Pontocypridoidea, and Cypridoidea can safely be postulated, but other relationships among podocopidan and platycopidan superfamilies could not be resolved from the 18S rDNA data. The polychotomous relationship among five lineages of the podocopidan and platycopidan superfamilies, i.e. Cytherelloidea, Bairdioidea, Darwinuloidea, Bythocytheroidea plus Cytheroidea, and Macrocypridoidea plus Pontocypridoidea plus Cypridoidea, suggests that common ancestors of those lineages diverged from each other rapidly in short intervals in the early Paleozoic.     

Chemical composition of Leptocythere psammophila (Crustacea: Ostracoda) as influenced by winter metabolism and summer supplies

Leptocythere psammophila is well known for the ecophenotypic variations of the ornamentation of its carapace. To test the respective influences of the environmental parameters on its ornamentation, 41 living specimens were collected in the Baltic Sea, the North Sea and in the English Channel during spring, summer and winter (April 1987, September and December 1988). The carbonate carapaces of these specimens were analyzed by means of electron microprobe. Thirteen elements were detected. Statistical comparisons of means and variances were performed using classical F and t-tests. Data were submitted to Normalized Principal Component Analysis and Discriminant Analysis. There are no significant differences between chemical composition of the carapaces from the different stations. However, the variability of Baltic Sea specimens is lower than that of other stations. Nonetheless, strong and significant differences appear when samples are gathered by season. The chemical composition of summer individuals is controlled by variations in water salinity and in fine grain terrigenous sediment supply. The composition of winter samples is related to the incorporation of Mg. In the case of spring specimens, both these factors have to be considered. Discriminant analysis between winter and non-winter samples sets correctly 84% of the individuals. The wrongly classified specimens are interpreted as ostracods that did not molt during the season of sampling. By attributing them to their possible season of molting, the result of the Normalized Principal Component Analysis improves. Differences in ornamentation are slight. They occur between the samples from the Baltic Sea and those from the other stations. The former have larger and less numerous punctations and lack the smooth surface in the anteroventral part of the carapace. The punctation diameter is more variable during summer than during spring. In the range of environments investigated, ornamentation of the carapace of L. psammophila seems unaffected by the seasonal environmental variations whereas its composition exhibits strong differences.     

Diel vertical migration and feeding patterns of Mysis mixta (Crustacea,Mysidacea) in the Baltic Sea

The extent of the nocturnal vertical migration of Mysis mixta Lilljeborg varied between early July and late October (of 1985 and 1986) in a coastal area of the Baltic Sea. Migration was more restricted in early July and late October. Seasonal changes in surface light levels and transparency were sufficient to explain the observed differences. Mysids avoided light levels above 10^-4 lux throughout the study period. Smaller juveniles migrated higher up than larger juveniles and adults. A two-layered distribution with part of the population close to the bottom was observed at night. Zooplankton were more abundant in water layers above the main concentration of mysids. M. mixta fed on phytoplankton, detritus, copepods, cladocerans, rotifers and tintinnids. Diel changes in gut fluorescence indicated a higher intake of phytoplankton at night, but levels were low compared to primarily herbivorous zooplankton. Comparisons of stomach contents of mysids caught at the bottom in the evening and in the water column at night showed a higher ingestion of zooplankton at night and of detritus during the day. Mysids caught at the bottom at night had an intermediate diet. Copepods and cladocerans constituted between 90 and 100% of ingested material by weight in all mysid groups.     

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.     

Larval distribution and reproductive ecology of Thysanoessa macrura (Crustacea: Euphausiacea) in the Scotia Sea

In the Scotia Sea, the larvae of Thysanoessa macrura G.O. Sars, 1885 were scarce in spring (September 27 to October 6, 1971) and mainly present at the Calyptopis I stage, while in summer (December 17, 1974 to January 3, 1975) they were much more abundant and much further developed (up to Furcilia VI). The spawning season commences as early as September and, as indicated by the presence of numerous nauplii and metanauplii in summer, is rather long. T. macrura begins to spawn much earlier in West-Wind-Drift waters than in waters of the Weddell Sea. It starts to spawn simultaneously with Euphausia frigida and E. triacantha, but significantly earlier than E. superba. These timings of the breeding process are related to differential biogeographical characteristics of these species. Plankton collections made at depths of 500 or 1000 m have, for T. macrura, revealed a developmental ascent analogous to that of E. superba. Ontogenetical deep-water migrations are, consequently, characteristic not only for E. superba, but also for other Antarctic euphausiids (in particular E. frigida, but possibly not for inshore E. crystallorophias) and, as correctly postulated by several other investigators, for all oceanic euphausiid species.     

Composition élémentaire (C,H, N) de Sphaeroma serratum et S. hookeri (Crustacea: Isopoda)

The elementary composition (C, H, N) of Sphaeroma serratum and Sphaeroma hookeri has been studied in different natural populations and also in individuals maintained in laboratory cultures. Changes in elementary composition have been observed in the natural communities. Laboratory experiments demonstrated that food may influence the isopod's chemical composition. These results provide an explanation fom the intraspecific differences which appear to characterize natural populations.     

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.     

Polymorphism of mannose phosphate isomerase in North Sea and Baltic Sea populations of the amphipods Gammarus zaddachi and G. salinus

Genetic variation at the mannosephosphate isomerase (MPI) locus was examined in the euryhaline sibling species Gammarus zaddachi Sexton and G. salinus Spooner. Both crustacean amphipods share identical enzyme mobilities, following electrophoresis on vertical starch gels. The MPI locus turned out to be highly polymorphic; it is encoded by 6 alleles in G. zaddachi and 7 alleles in G. salinus. Geographic variation of allelic diversity was studied in samples from 9 G. zaddachi and 10 G. salinus populations, primarily obtained from Baltic Sea and North Sea sites. Patterns of inter- and intraspecific heterogeneity are described. Differences in allelic composition exist between Baltic and North Sea samples of G. salinus. In G. zaddachi, levels of polymorphism are higher in North Sea populations than in those from Baltic Sea areas. The significance of these findings is discussed in the light of previous biochemical genetic investigations on the population structure of the two amphipods considered.     

Gonadal morphology and gametogenesis in the sea pen Pennatula aculeata (Anthozoa: Pennatulacea) from the Gulf of Maine

The ultrastructural features of gametogenesis are described in male and female colonies of the sea pen Pennatula aculeata. Specimens were collected for observation and fixation at 113 to 231 m depth in the Gulf of Maine, USA, in August 1993. The species is gonochoric, and all stages of gametogenesis are observed in both male and female colonies >45 mm in height. Gametogenesis shows several features that differ from sea anemones. The developing oocytes and sperm cysts are completely encompassed by gastrodermally derived follicle cells, and they are released from the mesenteries into the coelenteron before they are fully differentiated. Following maturation in the coelenteron, the eggs and intact sperm cysts are expelled through the mouths of the autozoids during spawning. The expulsion of sperm cysts suggests that they function as primitive spermatophores, perhaps as a way of reducing sperm dilution. Vitellogenesis results in the biosynthesis of lipid droplets which are the sole nutrient reserves in the egg. Heterosynthetic vitellogenesis is characterized by the importation of lipid precursors into the oocyte, and there is some indirect evidence that hypertrophic follicle cells play a role in production, transport, and/or mediation of these precursors. Spermatogenesis is similar to that of other anthozoans. The spermatozoon has a cone-shaped head, a posterior nuclear fossa, a ring of lipid-like bodies in the midpiece, a prominent cytoplasmic collar surrounding the proximal flagellum, and a single mitochondrion, but the posterior region of the sperm also contains previously undescribed concentric rings of cisternae resembling smooth endoplasmic reticulum. Received: 23 March 1998 / Accepted: 31 July 1998     

Tisbe (Copepoda: Harpacticoida) species from the Lagoon of Venice. I. Seasonal fluctuations and ecology

Distribution and abundance of Tisbe species were studied throughout a period of 3 years at 3 stations in the Lagoon of Venice (Italy). At 2 other stations samples were taken occasionally, as also at the Lido station (open sea). Twelve species may be considered as common inhabitants of the lagoon, and 3 species appear to be occasional transients, compared to the 9 species found at the Lido. A comparison of the physical data shows that the stations were similar in temperature and salinity, but differed primarily in pH values, sediment characteristics, algal substratum and faunal benthic community. The fact that remarkable differences in species composition and distribution of Tisbe could be observed between the various parts of the lagoon and the lagoon and the open sea, indicates that temperature and salinity are not so important for species diversity as are biotic factors. A particularly strong difference existed between samples taken at the bottom and from pilings, which constitute a very special biotope for the benthic communities in the lagoon and seem to be a preferred habitat for T. lagunaris and T. cucumariae. The data suggest that there is a certain interaction between T. holothuriae and the other species. Whenever T. holothuriae was abundant, the other species were rare, but T. clodiensis, T. dobzhanskii (Stations 1 and 2) and Tisbe sp. (Station 3) attained high relative abundance whenever T. holothuriae decreased considerably in number. T. holothuriae represents certainly the hardiest species of the lagoon, displaying a great ability to cope with stress conditions, even those present in the interior lagoon where strong fluctuations in salinity, temperature, pH and oxygen prevail.     

Feeding mechanisms and possible resource partitioning of the caprellidae (Crustacea: Amphipoda) from Puget Sound,USA

Caprellid amphipods feed by browsing, filter-feeding, predation, scavenging, and scraping. Food acquisition is related to the presence or absence of plumose setae on the second antenna; those species with such setae obtain a significant amount of their diet from filtering and scraping periphyton while those species without such setae usually rely on predation. Two primary predators and 8 primary filter-feeders and scrapers were investigated. Substrates inhabited by the predators did not overlap, but some filter-feeders were found together. Feeding overlap has been decreased among filter-feeders that occur together, as they either feed on different sized particles or they filter at different heights from the substrate. Species filtering the same sized particles at the same heights that were found together utilized alternate feeding mechanisms, e.g. browsing on algal filaments.     

Evolutionary genetics of marine Mysis spp. (Crustacea: Mysidacea)

Inter-and intraspecific allozyme differentiation in the mysid crustacean genus Mysis in the North Atlantic region was studied in order to evaluate earlier concepts of evolutionary and systematic relationships and to assess patterns of subdivision within widespread taxa. The results support a relatively ancient divergence of the marine and non-marine species of the genus, and are generally in line with the current subgeneric tridivision into Mysis s.str., Michteimysis and Auricomysis. However, the North American littoral species M. gaspensis should be returned to subgenus Mysis s.str. from its present position in Michteimysis with M. mixta. The closest observed affinities within Mysis s.str. were between M. gaspensis and the freshwater M. relicta group, and between M. oculata and M. litoralis. Intraspecific differentiation among North European coastal populations of M. oculata and M. litoralis was moderately strong (F _ST0.1), suggesting population bottlenecks and limited dispersal in the post-glacial time. On the other hand, ransoceanic differences were not essentially greater, indicating the systematic homogeneity and long-term dispersal capacity in the marine species. This contrasts with the strong genetic and systematic fragmentation earlier found within the circumboreal M. relicta species group.     

Carotenoids in the hermit crab Clibanarius misanthropus (Crustacea,Paguridae) from the Black Sea

The authors investigated the carotenoids in the hermit crab, Clibanarius misanthropus (Risso), from the Black Sea. By means of columnar and thin-layer chromatography the following carotenoids were found to be present in that species; -carotene, canthaxanthin, lutein, astaxanthin and some kind of polioxyxanthophill.     

Vertical distributions of Calanus finmarchicus and C. helgolandicus (Crustacea: Copepoda)

The vertical distributions of the spring populations of Calanus finmarchicus (Gunnerus) and C. helgolandicus Claus are described and compared. The differences we observed between the two species have probably confused the understanding of the vertical distribution and development of the populations of Calanus spp. in the shelf seas around the United Kingdom where the species occur together. The results imply that these two congeneric species have different behaviour patterns which minimise interspecific competition where the species have sympatric distributions. C. finmarchicus has its younger development stages overlying the older stages in the water column. In C. helgolandicus the converse is true; i. e., the majority of the populations of Stage I and II copepodites of the first spring generations are found below the thermocline. It is also suggested that the different behaviour patterns lead to different feeding regimes and strategies.JONSDAP Contribution No. 52     

Podotreme affinities of Raninoides sp. and Lyreidus brevifrons: evidence from spermatozoal ultrastructure (Crustacea: Brachyura: Raninoidea)

Spermatozoal synapomorphies which singly or collectively distinguish Raninoidea are: (1) the presence of single (Ranina, Raninoides) or multiple (Lyreidus) keel-like projections of the acrosomal capsule; (2) a very large, weakly electron-opaque peripheral acrosomal zone (Ranina, Raninoides) and an homologous large outer zone in Lyreidus; (3) poor differentiation of the operculum from the capsule (autapomorphy); (4) a very well developed, perforate subopercular zone, of variable form; (5) presence of unique inward longitudinal projections (septa or corrugations) in the wall of the subacrosomal chamber (autapomorphy). Shared, presumably synapomorphic characters of Ranina and Raninoides but not of Lyreidus within the Raninidae, are: (1) branching of some of the subacrosomal septa (unbranched in Lyreidus); (2) the subspheroidal form of the acrosome in Ranina with a length: width ratio (L:W) of 0.76, and, although slightly more depressed, in Raninoides (L:W 0.73), considered apomorphic relative to the more depressed form in Lyreidus (L:W 0.52); (3) single or multiple coiled perforatorial filaments (Ranina, Raninoides) contrasting with a capitate perforatorium with “amoeboid” head in Lyreidus; (4) division of the capsule wall to give one posterior (Ranina) or multiple enclaves, plesiomorphically (?) absent in Lyreidus. Similarities of Lyreidus with other podotremes include the capitate perforatorium, questionably related to the radiate spiked-wheel structure of homolids in which acrosome proportions are similar or less closely to the bilateral capitate perforatorium of dromiids and dynomenids, and basal capsular projections as in the dromiid Stimdromia (=Petamolera) lateralis and in cyclodorippoids. No spermatozoal synapomorphies support a sister-group relationship of raninoids and heterotreme-thoracotreme crabs.     

Life history of the sandy-beach amphipod Dogielinotus loquax (Crustacea: Dogielinotidae) from the outer coast of Washington,USA

Dogielinotus loquax Barnard, a common intertidal macroinvertebrate on exposed ocean beaches near Grays Harbor, Washington, USA, is distributed from mid to mean high tidal levels, with a mean density of 1 830 m^-2 in 1975–1976. Location of maximum density within the intertidal zone is related to surf intensity and varies seasonally. The species is iteroparous and has an average sex ratio of 1:1. Temperature constraints on growth and egg development rates apparently influence the timing of reproduction. Two dominant recruitments occur per year, one in early spring (the summer generation) and another in late summer (the overwintering generation). The sampling design and density estimates permitted estimation of the mortality rate for the summer generation. Shorebird predation is suspected to affect summer generation abundance. Dogielinotus loquax will soon be elevated to the type of a new genus, Proboscinotus (Bousfield and Tzvetkova, in press)Contribution no. 1271 from the Department of Oceanography, University of Washington; Seattle, Washington 98195, USA     

Ecology of two littoral species of caprellid amphipods (Crustacea) from Washington,USA

Ecology of 2 littoral species of caprellid amphipods is compared. Populations of Caprella laeviuscula, a periphyton scraper/filter-feeder, are most dense on eelgrass Zostera marina L. In the absence of C. laeviuscula, periphyton biomass increases 411% in protected Z. marina beds. The light absorption spectra of periphyton and Z. marina are similar, and C. laeviuscula, by its periphyton removal, may allow Z. marina to grow in areas where it would otherwise be excluded. C. laeviuscula is aggressively dominant over sympatric caprellids, and seasonal predators of C. laeviuscula are absent during winter, the period when periphyton would be most limiting to Z. marina. Populations of Deutella californica, a predator, are most dense on the hydroid Obelia dichotoma (L.). In the O. dichotoma epibiotic community, D. californica is the primary predator, but removal did not change the composition of the community structure. In the absence of other macropredators, the structure of the O. dichotoma epibiotic community depends more on the seasonality of O. dichotoma occurrence than on organismal interactions.     

Growth and moulting of Neomysis integer (Crustacea: Mysidacea)

The growth and moulting of Neomysis integer (Leach) was investigated in the field and the laboratory. In the Ythan estuary, Aberdeenshire, Scotland, monthly samples taken from November 1976 to October 1978 revealed that the summer generation juveniles and mature individuals grew at a rate of 4 to 5 mm and 1 to 2 mm monthly, respectively. The winter generation had a growth rate of 3 to 4 mm monthly for juveniles and about 1 mm for mature individuals; during the winter there was a period of 3 mo when growth was almost completely stopped. Mysids reared in the laboratory on Artemia sp. nauplii had an average daily growth rate of 0.06 mm at 9°C and 0.09 mm at 16°C. The growth factors of N. integer ranged from 3 to 17% for mature and immature individuals, respectively. Intermoult periods ranged from 3 to 7 d in immature mysids to 12 to 18 d in mature mysids. Average laboratory growth curves calculated from information on growth factors and intermoult periods indicate that at 9°C (winter generation) it takes N. integer 277 d to grow to be a 15 mm mature individual, whereas at 16°C (summer generation) it takes 188 d. N. integer moults 24 times as it grows from a juvenile to a mature individual.     

Interrelationships of the two families constituting the Lophogastrida (Crustacea: Mysidacea) inferred from morphological and molecular data

The Lophogastrida are primitive Mysidacea and comprise only six genera. One of these, Eucopia is considered as highly specialized and constitutes the family Eucopiidae; the other genera constitute the Lophogastridae. Among the latter family, the genus Gnathophausia is closely related to Eucopia, with two species (G. gracilis and E. sculpticauda) sharing similar morphological characteristics [i.e. ornamentation (spines) of the uropods, and the gastric mill]. This indicates that these species are phylogenetically related. To test this hypothesis, the partial 16S mitochondrial ribosomal RNA gene from various representative species of Gnathophausia and Eucopia were compared. The resulting phylogenetic tree suggests that each genus is monophyletic, and that Gnathophausia, which is the deepest-branching genus, is the most primitive, with the Eucopiidae originating from the Lophogastridae. The molecular results support the morphological hypothesis, and suggest an early separation of the two genera or a rapid divergence of Eucopia due to morphological specialization. Received: 29 June 1997 / Accepted: 16 March 1998