Latitudinal variation in the reproductive biology of the commensal crab Pinnaxodes chilensis (Decapoda: Pinnotheridae) along the Chilean coast

The pinnotherid crab Pinnaxodes chilensis is a common commensal of the edible sea urchin Loxechinus albus along the Chilean coast. Several aspects of the reproductive biology of P. chilensis were examined between April and June 1999, along temperature and salinity gradients, at three sampling sites along the Chilean coast (23°45'S-39°24'S). Results demonstrated significant differences in egg number, egg volume, dry egg weight, and reproductive output of ovigerous females, between the studied populations of northern and central-southern Chile. Egg volume, egg dry weight, and reproductive output of females decreased from high to low latitudes, while egg number increased from high to low latitudes, exhibiting a clear trade-off with egg volume. It has been shown that changes in environmental conditions (e.g. temperature, salinity) along a latitudinal gradient, can generate clines in reproductive characteristics in both free-living and parasitic species.     

Population genetics of the fissiparous holothurians Stichopus chloronotus and Holothuria atra (Aspidochirotida): a comparison between the Torres Strait and La Réunion

Collections of about 50 individuals from each of five populations of the fissiparous holothurian species Stichopus chloronotus and four populations of Holothuria atra were made in 1999. These populations were located in the Torres Strait (western Pacific) and La Réunion (western Indian Ocean). Allozyme electrophoretic surveys of five (S. chloronotus) and six (H. atra) loci were conducted to compare patterns of asexual reproduction and to investigate connectivity between regions separated by large geographic distances. Deviations from genotype frequencies expected under Hardy-Weinberg equilibrium, mostly heterozygote excesses, were observed in all populations of both species. The maximum contribution of sexual reproduction (calculated as the maximum number of sexually produced individuals: sample size=N*/N_i) was similar for all S. chloronotus (58-64%) and H. atra (76-92%) populations, and on the same level as previously reported for midshelf reefs of the Great Barrier Reef. The higher values in the latter species indicated greater contributions of asexual reproduction to S. chloronotus populations. Variability was strongly reduced in S. chloronotus populations at La Réunion, with only one locus being variable in that population. When the dataset was reduced to one representative per multi-locus genotype per population to reduce the effect of asexual reproduction on calculations on gene flow, F_ST values were not significantly different from zero, suggesting high gene flow between these regions. However UPGMA cluster analyses using Rogers' genetic distance, roughly clustered populations by region. In the case of H. atra, pooled populations within each region were significantly different from those of the other region. Thus, although some restrictions in gene flow and greater genetic distances between the regions may exist, those differences are distinctly less than those reported in previous studies on echinoderms over similar geographic scales. Despite the importance of asexual reproduction for the maintenance of local population size, this study also confirmed that the potential for widespread dispersal mediated by sexually produced larvae is large.     

Geographical subdivision,demographic history and gene flow in two sympatric species of intertidal snails, <Emphasis Type="Italic">Nerita scabricosta</Emphasis> and <Emphasis Type="Italic">Nerita funiculata</Emphasis>, from the tropical eastern Pacific

The patchy distribution of rocky intertidal communities in the tropical eastern Pacific (TEP) may impose severe constraints on the genetic connectivity among populations of marine invertebrates associated with this habitat. In this study, we analyzed a portion of the mitochondrial cytochrome c oxidase subunit I (COI) gene in two sympatric species of marine snails, Nerita scabricosta and Nerita funiculata, common inhabitants of the rocky intertidal from the Gulf of California (Sea of Cortez) and outer Pacific coast of the southern Baja California (Baja) peninsula to northern South America, to assess genetic connectivity among populations of each species. One of our aims was to determine whether the morphological, behavioral, and ecological differences observed among populations of both species throughout their range in the TEP corresponded to population genetic differences. In addition, we were interested in elucidating the demographic history of both species. We found no evidence of genetic structure throughout the Gulf of California and outer coast of the Baja peninsula region for either species. Comparisons between Gulf of California/Baja and Panama populations, however, showed significant genetic differentiation for N. scabricosta, but not for N. funiculata. The genetic differences between Mexican and Panamanian populations of N. scabricosta were consistent with previously reported ecological and behavioral differences for this species between these two distant regions. However, previously reported size differences between northern and central/southern Gulf of California individuals of N. scabricosta do not correspond with our findings of genetic connectivity among these populations. Results from neutrality tests (Tajima’s D and Fu’s F _S), the mismatch distribution, and Bayesian skyline analyses suggested that both species have experienced dramatic population expansions dating to the Pleistocene.     

Genetic population structure of the seagrass Thalassodendron ciliatum in sandy and rocky habitats in southern Mozambique

The genetic diversity of the dioecious seagrass Thalassodendron ciliatum in six populations in southern Mozambique was studied by means of random amplified polymorphic DNA (RAPD). Samples were taken from sandy and rocky habitats over a span of 880 km. Analysis of molecular variance (AMOVA) showed that most of the genetic variability (71.6% of total genetic variation) was observed within populations. There were also significant genetic differences among populations within each habitat (sandy and rocky). We did not find any significant overall genetic difference between sandy versus rocky populations, indicating that the morphological differences between plants from these two habitats are not maintained by reproductive isolation. There was no significant correlation between geographical and genetic distance, which is discussed in the light of current patterns along the coast. All sampled populations consisted of several genetically distinct individuals, indicating that sexual reproduction is widespread.     

Genetic, morphological, and chemical divergence in the sponge genus Latrunculia (Porifera: Demospongiae) from New Zealand

We undertook a comprehensive study of Latrunculia in New Zealand to determine the relationship between taxonomic, environmental, and chemical variation within the genus. Sponges were collected from five locations around New Zealand: Three Kings Islands, Tutukaka, Wellington, Kaikoura, and Doubtful Sound. Allozyme electrophoresis at nine polymorphic loci indicated that sponges from each geographic location were genetically distinct, and that they displayed genetic differences of the magnitude usually associated with reproductively isolated species (Nei's D between locations =0.375-2.476). Additionally, the comparisons revealed that the green and brown colour morphs of Latrunculia that are sympatric at Three Kings Islands and Kaikoura are distinct from each other, and that there are two genetic groups within the green sponges in Doubtful Sound. On the basis of genetic data we conclude that there are at least eight species of Latrunculia in New Zealand waters, not one to four as had been previously thought. Morphological comparisons of the eight genetic species based on skeletal characters (i.e. skeletal organisation of the choanosome, spicule composition, size, and geometry) indicated that the eight Latrunculia species fell into only two morphological groups that could be easily diagnosed on the basis of discorhabd type. Within these two primary morphological groups, skeletal characteristics among the eight species largely overlap and are not diagnostic. These findings emphasise the limitations of traditional taxonomic methods based solely on skeletal characters for distinguishing species of Latrunculia. However, multivariate analysis (MANOVA and CDA) based on six measured skeletal variables did reveal significant morphological variation among the species (Pillai's Trace=3.28, F=6.90, P<0.0001), supporting the division of the genus into eight species. Comparisons of chemical extracts from Latrunculia also showed that the amounts of five different bioactive compounds (discorhabdins A, B, C, D, and J) varied predictably among the eight species. This finding suggests that discorhabdin variation within Latrunculia, previously thought to be associated with intra-specific environmental variability, is more likely to reflect differences among species rather than phenotypic plasticity. Our results also highlight the importance of thorough taxonomic studies associated with marine natural products research to understand fully the variation in bioactive properties among individuals. The potential processes underlying the unusually high speciation rates in New Zealand Latrunculia that are indicated in our study are discussed.     

Biological patterns and near-bottom population characteristics of two pasiphaeid decapod crustacean species, Pasiphaea sivado and P. multidentata, in the north-western Mediterranean Sea

Biological aspects of the only two epi-/mesopelagic pasiphaeid species distributed along the continental marginof the north-western Mediterranean Sea were studied over the period 1991-1994. The annual reproductive cycle, reproductive output, sex and size distributions by depth (near-bottom distribution), and growth were analysed using 4156 specimens of Pasiphaea sivado (Risso, 1816) and 5491 specimens of P. multidentata Esmark, 1866. The reproductive cycle in the shallower-dwelling species, P. sivado, was nearly continuous, in contrast to the reproductive cycle in P. multidentata, which was markedly seasonal in nature. Both species exhibited similar size and population structure patterns with depth, with individual size increasing along the bottom depth gradient. The shallower-dwelling species had a significantly higher reproductive output level, in terms of the relative number of eggs, than the deeper-dwelling species. Biological trends for each of the species have been compared. The comparative results for these two congeneric species revealed that the biological parameters analysed were closely linked to their population structure. The slight differences in the depth range inhabited by each species have been considered the main environmental factor affecting the life histories of these two species. The data presented supported the hypothesis that depth exerted an important influence on the biological patterns of deep-water species, with the seasonality of reproductive processes increasing in deeper-dwelling species.     

Effects of isolation by distance and geographical discontinuity on genetic subdivision of Littoraria cingulata

Littoraria cingulata (Philippi, 1846) is a Western Australian, mangrove littorine snail, represented by two morphologically distinct subspecies, whose distributions are separated by >300 km. The southern subspecies, L. cingulata pristissini, is distinguished from the northern subspecies, L. cingulata cingulata, by having a thinner, keelless shell with more primary grooves, and lower and much more numerous ribs. In contrast with these striking differences, L. cingulata cingulata is morphologically very similar to another species, L. sulculosa, with which it also shares a nearly coincident geographic range. Allozyme comparisons at 22 presumptive loci confirmed a large genetic distance between L. cingulata and L. sulculosa, and the apparent conspecificity of the morphologically divergent subspecies of L. cingulata. Based on geological evidence, the geographical separation of the morphologically divergent forms of L. cingulata has developed within the past 5000 to 10 000 yr. The extensive continuous distribution of the northern subspecies, L. cingulatacingulata, and the large geographic disjunction between the northern and Shark Bay subspecies, L. cingulata pristissini, allowed a test of the genetic importance of this relatively recent disjunction. Within the continuous distribution of the two subspecies, a pattern of isolation by distance was visible up to distances of 300 km. Beyond 300 km, genetic subdivision, measured by pairwise G _ST (the proportion of genetic diversity due to differences between populations), averaged 0.028, whereas subdivision between Shark Bay and northern populations averaged 0.055 over the same range of distances. Although the relative paucity of barriers to gene flow tends to limit genetic subdivision in marine species with planktotrophic larvae, the results for L. cingulata suggest that subdivision can occur within a continuous distribution, but that special events leading to major disjunctions can substantially increase divergence, even over a relatively short period of time. Received: 16 February 1998 / Accepted: 23 April 1998     

Morphological and genetic adaptation to a lagoon environment: a case study in the bryozoan genus Alcyonidium

The Fleet (southern England) is a stable (ca. 5,000 years) coastal saline lagoon that supports a population of Alcyonidium resembling the common coastal epiphyte, Alcyonidium gelatinosum (L.). A combination of morphological, reproductive, and ecological characters was used to compare lagoonal and non-lagoonal proximate populations. Comparisons revealed a difference in the timing of spawning, considered to be related to the temporally restricted availability of viable substrata within the lagoonal basin. Allochronous spawning and spatial separation together suggest that the lagoonal taxon is reproductively isolated. The two populations were further compared with seven other coastal populations of Alcyonidium using randomly amplified polymorphic DNA (RAPD) analysis. The results confirm the individuality of the lagoonal taxon but also a close relationship with three A. gelatinosum populations. We present and consider four hypotheses that may account for the presence of this genetically distinct taxon: (1) diversification within the Fleet; (2) colonisation from another lagoon; (3) non-indigenous species status; and (4) introduction by shipping or other anthropogenically mediated dispersal mechanism. Significant diversification on the time scale involved has been demonstrated for isolated freshwater environments and, therefore, is feasible within a saline lagoon. Hypothesis 1 and, to a lesser extent, hypothesis 2 are consistent with the recognition of individual lagoons as 'biogeographic' islands of importance for their unique or characteristic biodiversity. The study also represents the first example of concordant morphological, reproductive, and genetic diversification in a marine bryozoan.     

Experimental effect of cold,La Niña temperatures on the survival of reef fishes from Gorgona Island (eastern Pacific Ocean)

The eastern tropical Pacific (ETP) reefs are affected at irregular times by extremely cold temperatures that occur principally during La Niña events. The effects of these low temperatures on the survival of reef fishes were experimentally assessed by determining the critical thermal minimum (CTM) of 15 reef fish species from Gorgona Island (ETP), and comparing these CTMs with the records of temperature during past La Niña events. Among species, mean CTMs ranged from 10.8°C to 16.3°C, which were lower than the coldest temperature recorded during the last La Niña event (18°C during La Niña 1998-1999). However, the observed ranges of CTM for two species (Thalassoma lucassanum and Eucinostomus gracilis) extended above 18°C. These results suggest that most of the reef fishes we studied are physiologically tolerant to the cold temperatures encountered during La Niña, though decreases in at least two populations may be expected as a result of the mortality of less tolerant individuals. Although tolerant to cold temperatures, reef fish populations may still experience negative changes during La Niña, because other determinants in population maintenance (e.g. reproduction and recruitment) are more temperature sensitive. The effects of other cold phenomena on reef fish survival are also discussed herein.     

Phylogeography of two squid (Loligo pealei and L. plei) in the Gulf of Mexico and northwestern Atlantic Ocean

The loliginid squids Loligo pealei LeSueur and L. plei Blaineville (both recently proposed for reclassification as Doryteuthis) are commercially important, similar in appearance, and sympatric throughout much of the northwestern Atlantic Ocean, the Gulf of Mexico, and the Caribbean Sea. To investigate possible cryptic speciation and population structure, we examined samples (collected from 1995 to 1997) of both species for restriction fragment length polymorphisms (RFLPs) in PCR products of the mitochondrial gene cytochrome c oxidase (subunit I). RFLP haplotypes were further characterized by direct sequencing. In North American waters, cryptic speciation was rejected by the far greater nucleotide sequence divergence between species (~14%) versus within species (<1%). Each species displayed about a dozen RFLP haplotypes, but only three of their respective haplotypes were found among 90% of L. pealei specimens (n=356) and 97% of L. plei specimens (n=431). For L. pealei, a genetic break existed between the northern Gulf of Mexico and the Atlantic Ocean; among sample units within each population, gene flow was consistent with panmixia. The phylogeography of L. pealei is likely a consequence of the eastward currents of the Florida Straits, the elevated temperatures of those surface waters, and the restriction of this species to the continental shelf. For L. plei, a genetic break existed between longitudes 88°W and 89°W, with the northwestern Gulf of Mexico and the northeastern Gulf-Atlantic Ocean comprising separate populations; among sample units within each population, gene flow fit an isolation-by-distance model. If the genetic break found for L. plei represents resident populations separated by nearshore physical parameters (e.g. effects of the Mississippi River and the sediment boundary at longitude 88°W), the lack of structure within the Gulf for L. pealei might be due to its distribution farther from shore. However, the two populations of L. plei probably represent annual recolonization from the southwestern Gulf of Mexico and from the eastern Caribbean Sea, whereas the populations of L. pealei probably are permanent residents within their respective regions.     

Ecological,genetic, and morphological differences among three <Emphasis Type="Italic">Pavona</Emphasis> (Cnidaria: Anthozoa) species from the Pacific coast of Panama

Ecological, genetic, and morphological differences among three Panamanian Pacific Pavona species with strongly developed collines (Pavona varians, P. frondifera, and P. chiriquiensis) were examined. Ecological factors included geographical distributions of species, habitat preferences, interspecific interactions, reproductive ecology, and tolerance to bleaching. Genetic differences were based on the electrophoretic analysis of ten allozyme loci. Morphological analyses consisted of tissue coloration, colony morphology, and measurements and counts of ten macro- and micro-skeletal characters. P. varians, present on reefs or in coral communities, is the most widely distributed and shows considerable morphological variation. P. chiriquiensis, a recently described species, encrusts basalt rock and has little morphological variation. P. frondifera is a reef dweller with a compact foliose morphology. Tissue coloration varies from light to dark brown in P. varians, from pink to brown in P. frondifera, and from brick red to brown or silvery in Pavona chiriquiensis. Also, the white to silvery polyp mouths of the latter species are a diagnostic feature that allows an easy identification in the field. Aggressive dominance during short-term interspecific interactions were as follows: Pavona chiriquiensis>P. varians>P. frondifera. P. chiriquiensis and P. varians showed contrasting responses to sea warming during the 1997-1998 El Niño Southern Oscillation. Whereas entire P. chiriquiensis bleached and died within 4 weeks of exposure to 30-31°C, colonies of P. varians did so only on their upper surfaces. The response of P. frondifera to elevated temperatures was not observed because it is mainly present in the Gulf of Panama where coral bleaching was absent in 1997-1998. The genetic data indicated that P. chiriquiensis differed strongly from both P. varians and P. frondifera, with Nei's unbiased genetic distances of 0.434 and 0.379, respectively. A fixed difference between P. varians and P. frondifera, and P. chiriquiensis exists at the triose phosphate isomerase (TPI-2) locus. A nearly fixed difference between P. chiriquiensis and P. frondifera and between P. chiriquiensis and P. varians was found at the hexokinase (HK) locus. P. varians differed slightly from P. frondifera with Nei's unbiased genetic distance of 0.068. No fixed difference was found between P. varians and P. frondifera. There were strong differences between P. chiriquiensis and P. varians in spawning times and gamete characteristics. Spawning in P. varians and P. chiriquiensis is 12 h out of phase. Also, eggs of the former species are white to beige and positively buoyant whereas those of the latter species are dark green and neutrally to negatively buoyant. No reproductive data are yet available for P. frondifera. Calicular diameters are significantly greater in P. chiriquiensis than in the other two species. In contrast, corallum thickness is greater in P. varians and P. frondifera than in P. chiriquiensis. Canonical discriminant function analysis readily separated the three species.     

Morphological and genetic evidence for vicariance and refugium in Atlantic and Gulf of Mexico populations of the hermit crab Pagurus longicarpus

The number and wide variety of southeastern United States marine taxa with significant differentiation between Gulf of Mexico and Atlantic Ocean populations suggests that these taxa may have experienced major vicariance events, whereby populations were subdivided by geological or ecological barriers. The present study compared variation in morphology, allozymes, and mtDNA in Gulf of Mexico and western Atlantic populations of the longwrist hermit crab Pagurus longicarpus Say collected during 1997 and 1998. Combined Atlantic populations had significantly fewer denticles on the second segment of the third maxilliped than did Gulf of Mexico populations, and the mean ratio of dactyl length to propodus length was significantly greater in the Atlantic crabs than in the Gulf of Mexico crabs. Allozyme allele frequencies at three loci showed genetic differentiation between a Gulf of Mexico population and two Atlantic populations. Analysis of mtDNA sequence data revealed a clear reciprocal monophyly between Gulf and Atlantic populations, with an estimated divergence age of ~0.6 million years ago. This estimated age of divergence is significantly more recent than an age previously estimated for its congener Pagurus pollicaris (~4 million years ago), suggesting that species with a similar genetic break between Gulf and Atlantic populations may not necessarily share an identical history. Surprisingly, there is evidence of geographic subdivision within Atlantic populations of P. longicarpus along the east coast of North America. This differentiation is especially strong between Nova Scotia and southern populations, suggesting that the Nova Scotia population may represent survivors from a northern refugium during the last glacial maximum.     

Reproductive cycle of two commercial species of sea cucumber (Echinodermata: Holothuroidea) from Caribbean Panama

The reproductive status of the holothuroid species Isostichopus badionotus (Selenka, 1867) and Holothuria mexicana (Ludwig, 1875) was studied over 16 months in Bocas del Toro (Panama), from November 1999 to February 2001. Sexual reproduction was evaluated by the gonad index method, and by histology of gonad development. In addition, population structure was assessed based on sex ratio, minimum reproductive size, and length and weight distributions of males and females. The sex ratio in both species was 1:1, with a unimodal population distribution composed mainly of mature individuals. The minimum reproductive length and weight were 13-20 cm and 150 g, respectively, for both species, although reproductive individuals 10 cm in length were also found. A consistently higher gonad index was observed in H. mexicana, due to a high proportion of mature females and males and high gonad indices in most monthly samples. Gametogenesis and spawning patterns seemed to occur throughout the year, with periods of enhanced activity. Two periods of maximum reproductive activity were tentatively identified: July-November for I. badionotus and February-July for H. mexicana, but neither species had a single, sharply defined annual spawning event. Further work on these exploited holothuroids should examine the relationships between reproduction and environmental factors and between reproductive status and recruitment.     

Demographic plasticity in tropical reef fishes

We use age-based analyses to demonstrate consistent differences in growth, mortality, and longevity of coral reef fishes from similar habitats (exposed reef crests) 20 km apart. On outer-shelf reef crests of the northern Great Barrier Reef (GBR), size in four taxa of reef fishes (Chlorurus sordidus, Scarus frenatus, and S. niger and the acanthurid Acanthurus lineatus) was systematically and significantly smaller when compared with the same taxa on adjacent mid-shelf reef crests. Differences in size could be attributed to differences in growth between habitats (shelf positions). On outer reef crests the species examined had consistently lower size at age profiles and also reduced life spans compared with populations from mid-shelf reefs. To confirm this relationship, two of the most abundant species (C. sordidus and S. frenatus) were selected for more detailed spatial analysis of demographic patterns. Sampling adults of both taxa from reef crests on three mid- and three outer-shelf reefs revealed that most of the variation in growth was explained by shelf position, although C. sordidus also displayed differences in growth among mid-shelf reefs. We conclude that differences in body sizes across the continental shelf of the northern GBR are primarily determined by these trends in growth. Strong spatial patterns also existed in the mean ages of populations and longevity estimates for C. sordidus and S. frenatus between shelf positions. Both species on outer-shelf reefs displayed less variable cohort sizes, significantly reduced mean ages, and foreshortened longevity compared with populations on mid-shelf reefs. Furthermore, differences in these parameters were rare among replicate reefs within mid- and outer-continental-shelf positions. Age-based catch curves suggested that rates of S. frenatus natural mortality on the outer shelf were nearly twice as high as on the mid shelf. Visual surveys indicated that total scarid densities on outer-shelf reef crests are on average fourfold higher than for equivalent mid-shelf habitats. This fact, coupled with significantly reduced growth rates, reduced mean ages, and increased mortality rates, suggests that density-dependent processes may be responsible for observed differences among localities.     

Genetic structure of deep-water redfish, Sebastes mentella, populations across the North Atlantic

Patterns of genetic diversity of the marine deep-water redfish, Sebastes mentella Travin, from 21 sampling locations (n=46 per sample, from 1995 to 1998) were examined throughout its range in the North Atlantic. Eight microsatellite loci were used in order to: (1) define the genetic structure and consider the possible influence of dispersal, geographic distance, oceanic currents and historical factors; and (2) relate the structure to present management units and practices. Three divergent population units (mean Š=0.012) associated with distinct geographic areas were defined: Eastern (Norway and Barents Sea), Panoceanic and Western (Gulf of St. Lawrence and offshore Newfoundland). The most important observation was the lack of genetic differences (mean Š=-0.0004) and lack of genetic isolation by geographic distance (r=0.318, P=0.112) among samples within the Panoceanic zone, from the Faroe Islands to the Grand Banks (6,000 km). A combination of vicariance, historical introgression with a closely related species, S. fasciatus, and contemporary patterns of oceanic circulation are likely to have shaped the observed population structure. These results only partially support current management units, and call for more integrated practices for regulating the exploitation of S. mentella throughout its extensive range.     

Isotopic composition, growth rates and biological behaviour of Chamelea gallina and Callista chione from the Bay of Trieste (Italy)

Shells of the bivalve molluscs Chamelea gallina L. and Callista chione L. from the Bay of Trieste (northern Adriatic) were analysed for their oxygen and carbon isotopic composition. The main purpose of this study was to obtain detailed information on the biological behaviour and, particularly, on the growth rates and the settlement time of these two species. Isotopic measurements were carried out on "spot" samples collected from the umbo to the ventral margin of each shell. The results obtained from nine specimens of C. gallina (plus some 20 specimens studied only for the isotopic composition of their initial umbo and for the last section of their ventral edge) and from two specimens of C. chione allow the following conclusions: (1) the calcium carbonate of C. gallina shells is not deposited under isotopic equilibrium conditions with the environmental variables (temperature and oxygen isotopic composition of seawater); (2) this species usually initiates precipitation of carbonate during the springtime, although some late summer-early autumn generation also occurs; (3) the rate of calcium carbonate deposition decreases with increasing age, and the overall accretion rate is a rather slow one; (4) there is a negative correlation between the carbon isotopic composition and the size of each shell; (5) the mean growth rate of the shells can be easily evaluated according to the oxygen isotopic composition of the spot measurements; (6) the specimens of C. chione do not precipitate the calcium carbonate of their shells in isotopic equilibrium with environmental conditions; (7) their accretion rate is rather fast during the first years of their life and progressively and considerably reduced after about 5 years; (8) the carbon isotopic composition of these shells does not show the marked evolution towards more negative values shown by C. gallina, suggesting that neither respiratory CO₂ nor carbon species derived from the oxidation of organic matter are systematically used for the precipitation of shell carbonate; and (9) a marked preference for the warmer half of the year is shown by the oldest specimen of C. chione (12 years).     

Genetic variability and level of differentiation in North Sea and Baltic Sea populations of the green alga<Emphasis Type="Italic"> Cladophora rupestris</Emphasis>

Cladophora rupestris is a perennial filamentous macroalga belonging to the Chlorophyta. It is widely distributed on both sides of the northern Atlantic Ocean and penetrates into the brackish Baltic Sea down to ca. 4 psu salinity. In this paper we present evidence for genetic differentiation of a Baltic form of this marine alga. We assessed genetic structure within and among 11 populations ranging along a salinity gradient from the Norwegian coast to the northern Baltic Sea proper. Samples of 328 individuals were studied using starch-gel protein electrophoresis to evaluate genetic variability and interpopulation differentiation based on allozymes. Of 11 loci examined, only one was polymorphic. For this locus, encoding superoxide dismutase (SOD-3), a total of seven alleles were distinguished. We found two genetically differentiated groups of populations of C. rupestris, one Baltic Sea group and one North Sea group, with a distinct border in the southern Kattegat near the entrance to the Baltic Sea. The genetic differentiation for SOD-3, expressed as pairwise F_ST values between the populations, was generally higher within the Baltic Sea group (0.10-0.43) than within the North Sea group (0.05-0.10); in the latter group also fewer pairs of populations differed significantly. Pairs of populations from different groups had the highest F_ST values (0.20-0.60). Hierarchical analysis of variance showed that 29.6% of the total variation in the SOD-3 locus was explained by variation between the two groups, while only 4.2% was explained by variation among the populations within the groups. The remaining variation (66.2%) was found within the populations.     

Blue mussels, Mytilus edulis, at the edge of the range: population structure, growth and biomass along a salinity gradient in the north-eastern Baltic Sea

The blue mussel, Mytilus edulis, is the most conspicuous animal species in the northern Baltic rocky sublittoral. In the studied area the species lives at the margin of its salinity tolerance. Although dwarfed by the low-salinity conditions, blue mussels in the northern Baltic are very abundant and have a decisive role in the benthic and pelagic ecosystems. We studied abundance, size distribution, biomass and growth rate of blue mussels along a 270 km salinity gradient in the northern Baltic Sea. Samples (n=317, 1998-1999) from moderately exposed and exposed rocky shores at seven study areas were collected in the southern Archipelago Sea in the west and into the central Gulf of Finland in the east, where the species is becoming increasingly rare. The results show a marked decline in mean mussel size and biomass from the saline west to the less saline east. The growth rate also decreased with lower ambient salinity. However, abundance of small mussels was considerably higher in the central and eastern parts of the study area. Vertically, the highest biomass was recorded at intermediate depths (5 and 8 m), being lower at both shallower (3 m) and deeper bottoms (12 m). It is concluded that salinity is the foremost factor determining size structure and growth rate among populations within the area. The results suggest that predation further influences the population structure of blue mussels living at the edge of their range in the central Gulf of Finland ultimately set by their salinity tolerance.     

Identification of Illex coindetii, I. illecebrosus and I. argentinus (Cephalopoda: Ommastrephidae) throughout the Atlantic Ocean; by body and beak characters

Shortfinned squid species of the genus Illex support commercial fisheries throughout the Atlantic Ocean and Mediterranean Sea. Previous identification of interspecific and intraspecific populations by morphological and size-at-maturity studies have not provided conclusive results. We analysed morphometric body and beak variables (24 characters) in three species of the genus (I. coindetii, I. illecebrosus and I. argentinus), using a geographic and seasonal series of 33 populations for 1,500 specimens of I. coindetii, I. illecebrosus and I. argentinus. Residuals of the regression between each morphometric body and beak variable and mantle length were used as input in a stepwise discriminant analysis. Species discrimination by body and hectocotylus characters required at least eight variables and resulted in high correct-classification percentages for I. coindetii and I. argentinus (75% and 90%, respectively), whereas the best identification resulted from beak characters (83% correctly classified). Size of the suckerless basal arm, sucker-bearing length and beak lateral wall discriminated best among I. coindetii from northern Iberia, northwest Iberia (year-1996) and Ireland in the Atlantic and western Mediterranean versus middle and eastern Mediterranean samples. Canadian shelf and American samples were discriminated from Canadian slope I. illecebrosus. Winter/shelf and winter/slope samples of I. argentinus seemed to form a single biological group separated from Falkland Island, 46°S/autumn spawners and 46°S/1996 specimens along the Patagonian Shelf. No significant sexual or maturity polymorphism was obtained. Discriminant analysis optimised population diagnosis on a morphometric basis of interest in fisheries strategies. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00227-002-0796-7.     

Cryptic species of Clavelina (Ascidiacea) in two different habitats: harbours and rocky littoral zones in the northwestern Mediterranean

Marinas and harbours provide ideal sites for the study of population genetics of marine invertebrates with restricted dispersal capabilities. They combine a confinement effect, particular ecological conditions (pollution, turbidity), and the possibility of high gene flow through ship-borne propagules, which greatly increases the natural dispersal capability of sexual and asexual propagules in many species with short-lived larvae. We studied the genetic structure of populations of the ascidian Clavelina lepadiformis living inside and outside harbours in the north-western Mediterranean. A 500-bp segment of the cytochrome c oxidase subunit I (COI) mitochondrial gene was sequenced in three populations from inside harbours (interior form) and in three populations from the rocky littoral (exterior form). Two congeneric Mediterranean species, Clavelina sp. and C. dellavallei, were used for comparison. We found that the interior and exterior forms of C. lepadiformis belong to two distinct clades, with a genetic divergence of 5%. Gene-flow values among these forms were insignificant. The lack of gene flow and the genetic divergence suggest that the interior and exterior forms of C. lepadiformis are in fact cryptic species rather than differentiated populations of the same species. Levels of gene flow were higher among interior habitats than among exterior habitats, a pattern likely maintained by genetic exchange through ships. We discuss the possible origins of the present-day distribution of these cryptic species. We contend that the study of species living both inside and outside these particular habitats will reveal more instances of genetic discontinuities allowing local adaptations.