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.     

Analysis of the genetic structure of European eel (Anguilla anguilla) using microsatellite DNA and mtDNA markers

The spawning population of European eel (Anguilla anguilla L.) has been considered panmictic on the basis of genetic markers and morphometric studies. This hypothesis was tested by screening glass eel from five locations (Ireland, Italy, Morocco, Sweden and U.K.), belonging to two cohorts at the cytochrome b (cyt b) locus (392 bp) of the mitochondrion and at five nuclear microsatellite loci. Seventeen cyt b haplotypes were detected, of which ten were singletons; the most common haplotype occurred in 47% of all fish. Haplotype number increased significantly with latitude. Phylogeographical structure based on the cytoplasmic marker was weak (F_ST=0.014) and non-significant. Close similarity was revealed between British and Irish glass eel populations, and weak differentiation among the British/Irish, Atlantic Moroccan, Italian and Swedish Baltic populations, respectively. No hierarchical genetic structure was obvious. Levels of genetic variation detected with five microsatellites were much higher levels than found with allozymes in previous studies (mean number of alleles per locus=11.1; mean expected heterozygosity=0.68). Overall among-population microsatellite variance was low but significant (F_ST=0.004), and caused by the linked microsatellite loci Aan03 and Aan04. The Hardy-Weinberg-Castle equilibrium and the absence of gametic disequilibria at these loci in the Moroccan population might point to its genetic isolation, although the impact of just two out of five loci is puzzling. Given the weak differentiation typical for marine species and the limitations of our data, the results should be interpreted with caution. However, combined with recent evidence from a related study, the paradigm that the European eel constitutes a panmictic population becomes difficult to maintain.     

Evidence of a spawning area of <Emphasis Type="Italic">Anguilla</Emphasis><Emphasis Type="Italic">marmorata</Emphasis> in the western North Pacific

Leptocephali of the tropical eel Anguilla marmorata have been consistently collected in the same area of the North Equatorial Current (NEC) in the western North Pacific during three consecutive cruises in June and July of 1991 (N=28) and 1994 (N=20), and July and September of 1995 (N=27), indicating that this is a spawning area of this species. These leptocephali were collected from 130°E to 142°E and 12°N to 20°N, to the west of the Mariana Islands, in 20 tows in 1991, in 13 tows in 1994 and in 17 tows in 1995, indicating a widespread presence, but a relatively low abundance. Six of these specimens (16.3-36.0 mm total length) from the 1995 cruise, which were of the typical size range of these leptocephali, were genetically confirmed to be A. marmorata in a previous study. The consistent presence of recently spawned A. marmorata leptocephali (9-20 mm) in all 3 years, suggests that the western region of the NEC is the spawning area of the northern population of A. marmorata that was identified in a recent population genetics study. These leptocephali would thus be transported westward by the NEC and then transported north into the Kuroshio Current and toward Taiwan and Japan, or south toward the southern Philippines and into the Celebes Sea by the Mindanao Current. Available evidence indicates that A. marmorata may have potentially year-round spawning, and the presence of a spawning area of this species in the same region as that of Anguilla japonica suggests that the northern population of A. marmorata has evolved a spawning migration from East Asia, the Philippines and the Celebes Sea region to the NEC area, but differs from A. japonica by having some individuals that recruit to more southern areas.     

Allozyme and mitochondrial DNA evidence of population subdivision in the purple sea urchin Strongylocentrotus purpuratus

Despite high potential for dispersal, the purple sea urchin Strongylocentrotus purpuratus was found to have significant genetic subdivision among locations. Ten geographic locations along the coast of California and Baja California were sampled between 1994 and 1995. Samples from some locations included both adult and recruit urchins. Allozyme analyses revealed a genetic mosaic, where differentiation over short geographic distances could exceed differentiation over much larger distances. Significant allozyme differentiation was found among subpopulations of adults (standardized variance, F _ST =0.033), among subpopulations of recruits (F _ST =0.037), and between adults and recruits from the same location. DNA-sequence data for the mitochondrial cytochrome oxidase I gene also showed significant heterogeneity among locations, with a mild break in haplotype frequencies observed 300 km south of Point Conception. California. Repeated sampling over time is necessary to determine whether these patterns of differentiation are stable and to begin to understand what forces produce them.     

Seasonal fluctuations in the photosynthetic capacity of photosystem II in symbiotic dinoflagellates in the Caribbean reef-building coral <Emphasis Type="Italic">Montastraea</Emphasis>

The photosynthetic capacity of photosystem II (PS II) in symbiotic dinoflagellates (Symbiodinium sp.), as measured by analysis of chlorophyll fluorescence, was investigated in the primary Caribbean reef-building corals, Montastraea annularis and Montastraea faveolata, for 5 years and Montastraea franksi over 2 years in the Bahamas. Significant seasonal fluctuations in the quantum yield of charge separation (F_v/F_m) of PS II were found in all species at all depths, with the highest photosynthetic capacity consistently recorded between mid-winter and early spring and the lowest photosynthetic capacity occurring in the mid to late summer. Corals residing in shallow depths of 1-2 m showed the greatest fluctuations in F_v/F_m, whereas deeper corals (3-4 and 14 m depths) had consistently higher values of F_v/F_m. Densities of symbiotic dinoflagellates and photosynthetic pigments followed a similar pattern. Fluctuations of photosynthetic capacity showed a strong correlation with seasonal patterns of water temperature and light. Such seasonal shifts in photosynthetic capacity are most likely due to several biochemical processes in the algae that lead to alterations of both photoprotection and photodamage. While symbiont density changed significantly on a seasonal basis, visual signs of coral bleaching were noted only in the fall of 1995 and the spring and summer of 1998. Comparisons of photosynthetic capacity and the decrease in the number of symbionts and their subsequent recovery indicated that symbiont populations in this study had the ability to recover quickly following bleaching events, as long as continued physical perturbation (e.g. thermal stress) did not shorten the recovery phase. Large-scale bleaching events are best viewed as the end points of seasonal physiological variation in which photosynthetic capacity and density of symbiotic dinoflagellates are reduced to a lower level than during "non-bleaching" years.     

Population genetic structure of the brown tiger prawn, Penaeus esculentus, in tropical northern Australia

Eight polymorphic microsatellite loci were analysed in six population samples from four locations of the Australian endemic brown tiger prawn, Penaeus esculentus. Tests of Hardy–Weinberg equilibrium were generally in accord with expectations, with only one locus, in two samples, showing significant deviations. Three samples were taken in different years from the Exmouth Gulf. These showed no significant heterogeneity, and it was concluded that they were from a single panmictic population. A sample from Shark Bay, also on the west coast of Australia, showed barely detectable differentiation from Exmouth Gulf (F _ST = 0 to 0.0014). A northeast sample from the Gulf of Carpentaria showed low (F _ST = 0.008) but significant differentiation from Moreton Bay, on the east coast. However, Exmouth Gulf/Shark Bay samples were well differentiated from the Gulf of Carpentaria/Moreton Bay (F _ST = 0.047–0.063). The data do not fit a simple isolation by distance model. It is postulated that the east–west differentiation largely reflects the isolation of east and west coast populations that occurred at the last glacial maximum when there was a land bridge between north-eastern Australia and New Guinea.     

Molecular genetic variation in tarpon (<Emphasis Type="Italic">Megalops atlanticus</Emphasis> Valenciennes) in the northern Atlantic Ocean

The tarpon (Megalops atlanticus) is a highly valued game fish and occasional food fish in the eastern and western Atlantic Ocean. Tarpon have a high capacity for dispersal, but some regional biological differences have been reported. In this study we used two molecular genetic techniques—protein electrophoresis of nuclear DNA loci, and restriction fragment length polymorphism analysis of the mitochondrial DNA (mtDNA)—to assess this species population genetic structure in the eastern (coastal waters off Gabon and Sierra Leone, Africa) and western (coastal waters off Florida, Caribbean Sea) Atlantic Ocean north of the equator. Genetic differentiation was observed between tarpon from Africa and tarpon from the western Atlantic Ocean. A unique allele and haplotype, significant differences in allozyme allele and mtDNA haplotype frequencies between the African and western Atlantic samples, and significant F_ST analyses suggest that levels of gene flow between tarpon from these two regions is low. Among the western Atlantic Ocean collections, genetic diversity values and allele and haplotype frequencies were similar. AMOVA analyses also showed a degree of genetic relatedness among most of the western Atlantic Ocean collections: however, some significant population structuring was detected in the allozyme data. A regional jackknifed F_ST analysis indicated the distinction of the Costa Rica population from the other western Atlantic populations and, in pairwise analyses, F_ST values tended to be higher (i.e., genetic relatedness was lower) when the Costa Rican sample was paired with any of the other western Atlantic samples. These data suggest that Costa Rican tarpon could be partially isolated from other western Atlantic tarpon populations. Ultimately, international cooperation will be essential in the management of this species in both the eastern and western Atlantic Ocean.Communicated by P.W. Sammarco, Chauvin     

Allozyme and mitochondrial DNA variation in yellowfin tuna (Thunnus albacares) from the Pacific Ocean

Samples of yellowfin tuna (Thunnus albacares) collected in 1991 and 1992 from the western, central and eastern regions of the Pacific Ocean were examined for genetic variability. Four polymorphic allozyme loci (ADA ^*, FH ^*, GPI-S ^* and GPI-F ^*) were examined in all samples and a fifth polymorphism (GDA ^*) was examined in western and central samples only. Samples were also screened for mitochondrial DNA variation following restriction analysis by two enzymes (BcII and EcoRI) detecting polymorphic cut sites. Eighteen mtDNA haplotypes were revealed, with an overall nucleon diversity of 0.678. A subset of individuals screened for eight restriction enzymes had an overall nucleon diversity of 0.724 and a mean nucleotide diversity per sample of 0.359%. No significant spatial heterogeneity was detected for alleles at the ADA ^*, FH ^*, GPI-S ^* and GDA ^* loci nor for the mtDNA haplotypes. Significant heterogeneity was detected for GPI-F ^*. At this locus, the two eastern samples (southern California and northern Mexico) were not significantly different from each other but were significantly different (P<0.001) from the five western/central samples (Philippines, Coral Sea, Kiribati, Hawaii-91 and Hawaii-92). GPI-F ^* 100 was the most common allele in western and central regions, GPI-F ^* 75 the most common in eastern samples.     

Effects of a complex archipelago on genetic subdivision of the intertidal limpet Siphonaria kurracheensis

Allozyme variation in the intertidal limpet Siphonaria kurracheensis was examined in 11 populations from the Houtman Abrolhos Islands, Western Australia, and four from the adjacent mainland, to test if the effects of these islands on genetic subdivision should include species with high dispersal. Consistent with extensive planktonic dispersal, genetic subdivision was low, with an average standardized variance in allelic frequencies (F_ST) of 0.010 over 400 km along the mainland, and 0.009 over 70 km in the Abrolhos Islands. Nevertheless, subdivision was statistically significant at the smallest scale examined (about 10 km), showing a pattern of local heterogeneity and large-scale homogeneity, which is common in planktonic dispersers. Among the island populations, heterozygosity was positively correlated with a multivariate index of exposure. The level of heterozygosity at the most exposed island sites is equivalent to that at the (also exposed) mainland sites, whereas it is lower in the most sheltered island sites. Although this indicates that some island populations are less well connected to the major pool of larvae, the overall effect of the archipelago on genetic subdivision is no greater than occurs on the mainland coast. This is contrary to findings in previously studied species with less potential for dispersal, suggesting that the local impediments to dispersal are selective in their effects, and are unlikely to be broadly important for genetic divergence in widely dispersing species.     

Temperature adaptation in eurythermal cod (Gadus <Emphasis Type="Italic">morhua</Emphasis>): a comparison of mitochondrial enzyme capacities in boreal and Arctic populations

Activities of citrate synthase (CS), cytochrome c oxidase (CCO) and the electron transport system (ETS) were investigated in white muscle and liver of laboratory-maintained cod, Gadus morhua, from the North Sea, Norwegian coast and Barents Sea for an analysis of temperature acclimation and adaptation in aerobic metabolism. Cold acclimation within each population led to elevated activities of CS, CCO and ETS in white muscle. In liver, however, only North Sea cod showed cold-compensated CS activities, with CCO and ETS unchanged. In contrast, cold-acclimated Norwegian cod displayed unchanged enzyme activities, and Arctic cod showed elevated activities for CS, but decreased activities for CCO and no change in ETS. Between-population comparisons revealed clear evidence for permanent cold adaptation in white muscle of northern (Norwegian coast and Barents Sea) compared to boreal (North Sea) cod populations, reflected by higher activities of CS, CCO and ETS at the same acclimation temperature. Cold-compensated, mass-specific enzyme activities in liver were found in northern compared to boreal cod for CS and CCO, however, for ETS, after warm acclimation only. When evaluated as capacity in total liver, such activities were only found in northern populations in the case of CS at all temperatures, and for CCO and ETS at 15°C only. Hepatosomatic index (I_H) and liver lipid contents were highest in North Sea cod, with similar I_H but lower lipid contents in cold- versus warm-acclimated animals. An acclimation effect on I_H was found in Norwegian cod only, with higher I_H but unchanged lipid contents in the cold. In conclusion, permanent cold adaptation of muscle aerobic metabolism prevails in cod populations at higher latitudes, which is in line with permanently elevated rates of oxygen consumption observed in a parallel study. These differences reflect higher maintenance costs in cold-adapted versus cold-acclimated cod.     

Panmixia in Pocillopora verrucosa from South Africa

The genetic structure of six local collections of Pocillopora verrrucosa from six coral reefs in KwaZulu-Natal, South Africa, was examined using allozyme electrophoresis. The six separate reefs lie within two different reef complexes. Twenty-two enzymes were screened on five buffer systems, but only five polymorphic loci (Gpi-1, Gdh-1, Lgg-2, Lpp-1, Est-1) could be consistently resolved. No significant differences in allelic frequencies were detected among the six sites. All local collections were genotypically diverse, with evidence of only very limited clonal replication at each site. Indeed, the ratio of observed to expected genotypic diversity (mean Go:Ge=0.64ǂ.05 SD), the ratio of observed number of genotypes to the number of individuals (mean Ng:N=0.65ǂ.04 SE), and deviations from the Hardy-Weinberg equilibrium indicate that sexual reproduction plays a major role in the maintenance of the populations. No genetic differentiation was found either within (FSR=0.026ǂ.003 SE) or between (FRT=0.000ǂ.001 SE) reef complexes. The homogeneity of the gene frequencies across the six reefs strongly supports the assumption that the KwaZulu-Natal reef complexes are highly connected by gene flow (Nem=44). The reefs in the southern and central reef complexes along the northern Maputaland coastline can therefore be considered part of a single population.     

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.     

Influence of environmental factors on burrow irrigation and oxygen consumption in the mudflat invertebrate Urechis caupo

We examined burrow irrigation activity by the mudflat worm Urechis caupo in response to suspended food, ambient hypoxia (down to 3.3 kPa PO₂), hydrogen sulfide exposure (up to 100 µmol l^-1), and short-term temperature change (range 10-22°C). In normoxic, nutrient-free water at 14°C, O₂ consumption ( [(M)\dot]O₂ ) \left( {\dot M{\rm O}_2 } \right) was 45 nmol min^-1 g^-1, water flow rate ( [(V)\dot]_W ) \left( {\dot V_{\rm W} } \right) was 27 ml min^-1 (0.66 ml min^-1 g^-1), frequency of peristaltic waves (F_P) was 2.6 contractions min^-1, stroke volume (SV) was 11 ml, and O₂ extraction coefficient (EO₂) was 0.27. Adding suspended food to the burrow water occasionally elicited stereotypical feeding behavior but had no effect on any measured variables during nonfeeding periods. Hypoxia greatly decreased [(M)\dot]O₂ \dot M{\rm O}_2 (75% reduction at 3.3 kPa PO₂) but did not affect [(V)\dot]_W \dot V_{\rm W} , F_P, SV, or EO₂. Sulfide at 50 µmol l^-1 or less had no effect on burrow irrigation activity, whereas 100 µmol l^-1 sulfide decreased [(V)\dot]_W \dot V_{\rm W} by 58% and F_P by 50% but had no effect on SV. Temperature strongly affected [(V)\dot]_W \dot V_{\rm W} (Q₁₀ of 1.9 from 10°C to 22°C). We propose that U. caupo's ability to live in the hypoxic, sulfidic mud of productive mudflat environments, combined with its very efficient mucous net, allows it to process much less water for feeding than other suspension-feeding invertebrates. This, in turn, necessitates an efficient O₂ extraction mechanism, which is provided by the water lung activity of U. caupo's unique hindgut.     

Seasonal variation in ecophysiological patterns in macroalgae from an Arctic fjord. I. Sensitivity of photosynthesis to ultraviolet radiation

Marine macroalgae inhabiting Arctic coastal ecosystems are exposed to pronounced seasonal variations in the radiation regime, including harmful UVB radiation. This study presents the first data on the seasonal changes in the sensitivity of macroalgal photosynthesis towards UV exposure by comparing under-ice, clear-water and turbid-water conditions characteristic for late winter, spring and summer. Various brown (Laminaria saccharina, L. digitata, L. solidungula, Saccorhiza dermatodea, Desmarestia aculeata), red (Palmaria palmata, Devaleraea ramentacea) and one green macroalgal species (Monostroma aff. arcticum) were collected at the same water depth throughout the seasons in the Kongsfjord (Spitsbergen, Svalbard, Norway). Maximum quantum yield (F_v/F_m) and maximum photosynthetic electron transport rates (ETR_max) were determined immediately after collection, after 2 h exposure to artificial UV radiation and after 18 h recovery in dim white light. Photosynthesis of the studied species showed different responses depending on their morpho-functional and physiological characteristics, their life strategies, phenology and depth distribution. Within the genus Laminaria, maximum quantum yield of adult specimens of the deep-water species L. solidungula was most strongly UV sensitive. Adult L. saccharina exhibited a lower UV sensitivity than a 6-month-old specimen. Inhibition of photosynthesis after UV exposure remained at the same level throughout the study period, both in adult L. saccharina and S. dermatodea. However, adult specimens of L. saccharina collected in May showed partial recovery only, whereas photosynthesis of specimens from both species collected later recovered fully. D. aculeata exhibited a remarkable decrease of UV sensitivity during the study period. Photosynthesis of specimens collected under the ice was strongly inhibited by UV, but the degree of inhibition decreased during spring and summer. Concomitantly ETR_max values were low after UV exposure in specimens collected in June, but increased later in the season. P. palmata exhibited a relatively flexible response. Photosynthesis in specimens collected under the ice in June or in turbid water in July/August was relatively strongly inhibited; specimens collected during sunny periods and in clear water in spring showed a much lower degree of photoinhibition after UV exposure. The seasonal pattern of low/high ETR_max values in spring/summer is probably a characteristic of the life strategy of this species. The UV sensitivity of D. ramentacea exhibited a similar seasonal pattern. In M. aff. arcticum, UV sensitivity increased and ETR_max values decreased during the study period, reflecting the life strategy of this annual late winter/spring species. The physiological basis for the seasonal changes in UV sensitivity of photosynthesis is presented in a companion paper (this issue).     

Metabolic properties of Northern krill, Meganyctiphanes norvegica, from different climatic zones. II. Enzyme characteristics and activities

Activities and characteristics of two metabolic key enzymes, citrate synthase (CS) and pyruvate kinase (PK), were studied in the Northern krill, Meganyctiphanes norvegica, with respect to adaptive properties under different thermal conditions. Krill were sampled during late winter/spring and summer from the constantly warm Ligurian Sea (12-13°C below the thermocline), the colder but also comparatively constant Clyde Sea (7-8°C), and the variable Kattegat (2-16°C). Both enzymes showed distinct tissue- and organ-specific activities, which were highest in the pleopods - the principal locomotive organs. The fourth and fifth abdominal segments, however, were used for routine investigation due to lowest variability. Specific activity of CS and PK did not differ between seasons in krill from the Kattegat or the Clyde Sea. In the Ligurian Sea, in contrast, specific CS activities were significantly lower during summer. Analysis of individual data illustrated a decrease of CS activity with size and an increase of PK activity with size. Taking these allometric effects into account, as emphasized by calculating the ratio between both enzymes, variation of CS and PK activities turned out to be solely dependent on body size, which differed between locations and seasons. Ligurian krill from the summer, however, were unique in that they showed a lower CS/PK ratio than would be predicted by the scaling effect. Thermal characteristics of each enzyme were similar between locations and seasons. During the winter, in Kattegat and Clyde Sea krill, K_m values (Michaelis-Menten constant) of CS towards acetyl-coenzyme A exhibited an almost constant level over the experimental temperature range of 4-16°C. During summer, however, K_m values were lower at 8°C in the Clyde Sea and at 12°C in the Kattegat. In Ligurian krill from the summer, K_m values were consistently lower than those of winter krill over the entire experimental temperature range. In conclusion, Kattegat and Clyde Sea krill show only minor adaptations to their respective thermal environments in terms of CS and PK characteristics. Ligurian krill, in contrast, exhibited decreased specific CS activity during summer, which might be compensated by elevated enzyme-substrate affinity as indicated by lower K_m values. Since temperature was constant during both seasons, this effect cannot be explained as a reaction to thermal conditions. Consequently, oligotrophic conditions in the Ligurian Sea during summer may entail a reduction in the somatic performance of krill, which is reflected by lower CS activity.     

Genetic differentiation in Sargassum polyceratium (Fucales: Phaeophyceae) around the island of Curaçao (Netherlands Antilles)

The seaweed Sargassum polyceratium Montagne inhabits a broad spectrum of subtidal and intertidal habitats. Genetic diversity and spatial genetic structure were examined within and among 12 stands using random amplified polymorphic DNA (RAPD) phenotypes. Data were analyzed using analysis of molecular variance (AMOVA) and Shannon's information measure. In both analyses, 60-75% of the variation occurred within stands and 25-40% between stands. These values are consistent with out-crossing, high-dispersal species. Significant differentiation was found among bays ca. 25 km apart (Shannon's G'_st averaged 0.37 and pairwise AMOVA K_st values averaged 0.272) and among stands 150-200 m apart within bays (AMOVA K_st values averaged 0.149). Effects of shore (windward vs. leeward), depth, and bay on population structure were tested. These analyses revealed that the factor depth is confounded with shore, and that bays show significant differentiation from each other but are not completely isolated from one another. Mantel tests for differentiation-by-distance were significant along both sides of the island but stronger along the windward side. A neighbor-joining analysis of genetic distances among stands showed that the effects of currents around both tips of the island were especially important for shallow populations. For S. polyceratium, depth and bay promote population differentiation along shores, yet dispersal around the tips of the island simultaneously connects these populations to varying degrees. This study highlights the importance of investigating the relative contribution of habitat factorsin relation to island-scale population structure.     

Effects of emersion and elevated haemolymph ammonia on haemocyanin–oxygen affinity of Cancer pagurus

The subtidal crab Cancer pagurus (L.) experiences involuntary periods of emersion associated with practices used in their marketing and distribution. During 24 h emersion, impaired gill function caused an increase of circulating total ammonia (TA=NH₃+NH₄⁺) of 0.35 mmol TA l^-1 (167%). The oxygen-binding characteristics of the haemocyanin of C. pagurus were examined at 10°C in the presence of total ammonia (0.2-1.0 mmol TA l^-1). The haemocyanin-oxygen affinity was decreased in the presence of TA ((logP₅₀/(log[TA]=0.16). Emersion induced significant acidosis and elevated circulating levels of haemolymph TA, lactate and urate, but all had returned to normal levels within 24 h of re-immersion. The accumulation of haemocyanin-modulating substances during 24 h emersion compensated partially (40%) for the effect of the acidosis, but the net effect of the emersion period was a significant decrease in oxygen affinity, corresponding to an increase of P₅₀ (10°C ) from 1.24 kPa (immersed) to 1.96 kPa (24 h emersion). The implications of the findings are considered in terms of the effects and adaptations to emersion.     

Mitochondrial DNA variation in the European lobster (<Emphasis Type="Italic">Homarus gammarus</Emphasis>) throughout the range

The genetic differentiation of the European lobster (Homarus gammarus) was investigated in 3,283 individuals from 44 population samples throughout its geographical distribution (Norway to Greece) by means of polymerase chain reaction restriction fragment length polymorphism analysis of a 3-kb mitochondrial DNA segment. Ninety composite haplotypes were revealed with the number of haplotypes in each population sample ranging from 4 to 31. Private haplotypes were found at very low frequencies. The global exact test of sample differentiation based on composite haplotype frequencies was statistically significant. F_ST analyses also showed significant heterogeneity among the European lobster population samples (F_ST=0.078, P<0.001). This differentiation was mainly due to the population samples from northern Norway, the Netherlands, and the Mediterranean Sea. These samples differentiated from the rest due to reduced gene diversity rather than to unique haplotypes. The relationships among these samples were illustrated with cluster analyses; four major distinct groups were evident: Mediterranean, northern Norway, Netherlands, and the remaining Atlantic samples. Based on the low degree of differentiation revealed in the European lobster and its limited capacity for dispersal, the most probable hypothesis is that all populations have been established from a common refuge after the end of the last Ice Age, that is, within the past 15,000 years. The results of this study show that mitochondrial DNA is a powerful tool for the determination of the genetic structure among lobster samples, which is important for a proper management policy designed to protect and to cultivate this species.Communicated by O. Kinne, Oldendorf/Luhe     

Population genetics of the northern Pacific seastar Asterias amurensis (Echinodermata: Asteriidae): allozyme differentiation among Japanese,Russian, and recently introduced Tasmanian populations

The northern Pacific seastar Asterias amurensis Lütken was recently introduced to Tasmanian waters, possibly through ballast water discharged from ocean-going vessels. Three Tasmanian populations and seven native populations from Japan and eastern Russia were examined in 1994 for variation at 22 allozyme loci. There was significant spatial heterogeneity in gene frequencies among the native populations, especially for the ocus APK ^*, where G _ST was a very high 0.47. The Tasmanian populations could not be separated genetically from one another, suggesting they have a common origin from a single introduction. The average heterozygosity per locus of the Tasmanian populations (0.116 to 0.127, mean 0.123) was 30 to 40% less than that of the native populations (0.177 to 0.216, mean 0.192), suggesting that colonisation of Tasmanian waters was accompanied by a population-size bottle-neck. The Tasmanian seastars were genetically more closely related to the two populations from central Japan (Suruga and Tokyo Bays) than to populations from Vladivostok, northern Japan (Yoichi, Nemuro and Mutsu Bays) or southern Japan (Ariake Sea). However, there were significant differences between the populations from Tasmania and central Japan, which may be attributable to founder effects or may mean that the true source of the Tasmanian introduction has yet to be identified.     

Allozyme variation in European populations of the oyster Ostrea edulis

Variations at 22 enzyme coding loci were surveyed in 11 populations of the oyster Ostrea edulis L., which were sampled between 1988 and 1990 along the Atlantic and Mediterranean coasts of Europe. Atlantic oyster beds suffered a steady decline during the last century, and restocking of beds with oysters of foreign origin has probably resulted in a high degree of interbreeding of natural oyster stocks from all Atlantic Europe. Our study confirms the low levels of genetic variability previously reported for the oyster populations from the Atlantic coasts, and extends it to the Mediterranean coasts. The locus arginine-kinase (ARK ^*) exhibited a high degree of interpopulation differentiation (F _ST=0.289), resulting from extensive variation in gene frequencies along a geographical cline. However, the overall genetic differentiation between populations was slight, and similar to that reported for other local populations of bivalves (mean genetic distance between populations is 0.010, mean F _ST=0.062). A general pattern of increasing differentiation along the coastline in an Atlantic-mediterranean direction emerged; but genetic differentiation among the Atlantic populations was not significantly lower than that observed among the Mediterranean populations. This and other results suggest that the effects of extensive transplantation of oysters among various areas in Europe are detectable only in some particular localities. The geographical distribution of low-frequency alleles suggests a restriction to gene flow outwards from the Mediterranean Sea, across the Straits of Gibraltar.