Life cycle of the jellyfish <Emphasis Type="Italic">Lychnorhiza lucerna</Emphasis> (Scyphozoa: Rhizostomeae)

The life cycle of Lychnorhiza lucerna (Scyphozoa: Rhizostomeae) and the settlement preferences of its larvae were studied using laboratory-based rearing experiments. Mature medusae of L. lucerna were collected from the beach of the Río de la Plata estuary, Argentina. This species displayed the typical metagenetic, (i.e. medusoid and polypoid), life cycle reported for other rhizostomes. The fertilized eggs developed into motile and short lived planulae. The majority of planulae settled on the air-water interface (p < 0.001). Of those that settled on the settlement plates provided, no significant differences were observed between styrene slides, glass slides and shells of the bivalve Mactra isabelleana (p > 0.05). No planulae settled on stones. Several hours after planulae settled, they metamorphosed into sessile four-tentacled scyphistomae. Most scyphistomae attached onto the air-water interface. At 19–22°C, the scyphistomae grew up to 22 tentacles and reached 1,500 μm height. The scyphistomae increased their numbers by means of formation of podocysts from which new polyps emerged and strobilated. Strobilation occurred 46 days after settlement. Only polydisk strobilation was observed and each strobila always produced three ephyrae. After releasing ephyrae, strobilae returned to normal scyphistomae and were capable of repeating strobilation. A single founder polyp was estimated to produce up to 60 ephyrae over 4 months. Ephyrae developed into metephyrae 15 days after release at 19–22°C. In this paper we describe the morphological and some behavioural features of L. lucerna in the polypoid and early medusoid stages.     

Developmental and genetic evidence for the existence of three morphologically cryptic species of<Emphasis Type="Italic"> Crepidula</Emphasis> in northern Chile

The taxonomy of Crepidula has been based principally on shell morphology, but shell characteristics in this genus may be strongly influenced by the substrates they inhabit and are thus of limited use in classification. Four species in this genus are currently recognized from the Chilean coast, of which two, Crepidula dilatata and C. fecunda, are sympatric. These species are morphologically cryptic and differ only in their larval development: C. dilatata has indirect development with a planktonic phase, and C. fecunda direct development within egg capsules. The other two species, C. coquimbensis and C. philippiana, are species that occur within gastropod shells inhabited by hermit crabs and differ from the other species in internal morphology of the shell, developmental features, and geographic distribution. Analyses of nine morphological characters, eight developmental characters, and 26 enzymatic loci were carried out on four Crepidula populations in the Coquimbo region of northern Chile. Four populations of Crepidula were identified based on morphological, developmental, and protein electrophoresis features. Our results suggest the presence of C. coquimbensis plus three morphologically cryptic species including C. dilatata, C. fecunda, and a new previously unrecognized cryptic species. All four populations showed differences in larval development pattern. C. fecunda have indirect development, with free-swimming larvae. The remaining three species have intracapsular metamorphosis, but different modes of larval nutrition. C. dilatata have nurse eggs that are consumed by mechanical destruction; C. coquimbensis engulf nutritional embryos; and the new species eat nutritive embryos by rotation. Protein electrophoresis data support the specific condition of each group with different larval development and endorse the use of developmental differences in taxonomy of Crepidula. Genetic distance shows that C. dilatata and C. fecunda are closely related, and that C. coquimbensis is closer to them than to the new cryptic species.     

Disinfection efficacy on cyst viability of<Emphasis Type="Italic"> Artemia franciscana</Emphasis> (Crustacea),<Emphasis Type="Italic"> Hexarthra fennica</Emphasis> (Rotifera) and<Emphasis Type="Italic"> Fabrea salina</Emphasis> (Ciliophora)

Resting stages (cysts) of Artemia franciscana, Hexarthra fennica and Fabrea salina were exposed for two periods (acute and chronic exposures) to different concentrations of five aquaculture disinfectants (formalin, sodium hypochlorite, potassium permanganate, organic iodine, copper sulphate). The effects of chemical treatments were ranked according to their action on cyst hatching: total inhibition (no cyst hatching), heavy inhibition (significant decrease of cyst hatching), alteration of hatching synchrony, and occurrence of anomalous hatching (death of emerging individuals or birth of malformed organisms). Resting-stage viability was not completely impaired by disinfectant exposures at the ordinary doses used in aquaculture, so that usual treatments are possibly ineffective against the protist and metazoan cysts potentially present in breeding systems. On the other hand, the high resistance shown by cysts suggests the possibility of using these chemicals in live-feed disinfection.     

Competition and facilitation between germlings of<Emphasis Type="Italic"> Ascophyllum nodosum</Emphasis> and<Emphasis Type="Italic"> Fucus vesiculosus</Emphasis>

Competitive interactions between germlings of Ascophyllum nodosum (L) Le Jolis and Fucus vesiculosus L. were studied both in the laboratory and on a shore of the Isle of Man, in the Irish Sea. Both intra- and interspecific competition were investigated by comparing the performance of algal germlings both in monocultures and mixed populations of the two species. The growth of germlings of both species reduced with increasing density. F. vesiculosus always grew faster than Ascophyllum and did best in mixed cultures, whereas Ascophyllum did least well when mixed with Fucus germlings. Clearly the adverse effects of F. vesiculosus on A. nodosum were greater than those of Ascophyllum cohorts. At the same total density, the survival and growth of Ascophyllum declined with an increasing proportion of Fucus germlings, implying that poor recruitment of A. nodosum results from strong competition with F. vesiculosus. However, under desiccation stress on the shore, F. vesiculosus enhanced the survival of A. nodosum at the early germling stage even though competition may occur again at the late stage. Thus, whether interactions between germlings take the form of competition or facilitation depends on the environmental conditions.Communicated by O. Kinne, Oldendorf/Luhe     

Adaptations of subtropical Venus clams to predation and desiccation: endurance of<Emphasis Type="Italic"> Gafrarium tumidum</Emphasis> and avoidance of<Emphasis Type="Italic"> Ruditapes variegatus</Emphasis>

Intertidal endobenthic bivalves are often dislodged from sediments by hydrodynamic forces. As a result, they encounter the dangers of predation and desiccation, which are generally harsh near the sediment surface. To cope with such dangers, the bivalves possibly possess: (1) a strong body to endure predation and desiccation stress, (2) quick mobility to avoid the stresses, or (3) a high growth rate for attaining a size refuge. The present study examined which of these modes are adopted by the subtropical cobbled-shore Venus clams Gafrarium tumidum (Röding, 1798) and Ruditapes variegatus (Sowerby, 1852), revealing the following interspecific differences. (1) G. tumidum survived better than R. variegatus did in harsh experimental conditions, namely: the experimental cages exposed to predation and desiccation on a cobbled shore; a laboratory aquarium with a predatory crab Scylla serrata; and ovens with high temperatures (27°C and 34°C). (2) R. variegatus was more mobile than G. tumidum was, digging into the sediment on a cobbled shore more rapidly at both high and low tides. (3) The two species with shell lengths 20–30 mm showed similar growth rates (median: –0.2 to 44.5 m day^–1) in seasonal mark–recapture surveys over 2 years. Overall, to cope with the dangers of predation and desiccation G. tumidum appears to have a strong body, while R. variegatus displays rapid mobility, and neither species seems to attain a size refuge through rapid growth. Such species-specific modes are discussed in relation to the interspecific differences found in shell morphology.Communicated by T. Ikeda, Hakodate     

Effect of sexual maturation on the tissue biochemical composition of<Emphasis Type="Italic"> Octopus vulgaris</Emphasis> and<Emphasis Type="Italic"> O</Emphasis>.<Emphasis Type="Italic"> defilippi</Emphasis> (Mollusca: Cephalopoda)

Changes in the protein, lipid, glycogen, cholesterol and energy contents, total amino acid and fatty acid profiles of Octopus vulgaris and O. defilippi tissues (gonad, digestive gland and muscle) during sexual maturation (spermatogenesis and oogenesis) were investigated. Both species showed an increase of amino acids and protein content in the gonad throughout sexual maturation (namely in oogenesis), but allocation of these nitrogen compounds from the digestive gland and muscle was not evident. The major essential amino acids in the three tissues were leucine, lysine and arginine. The major non-essential amino acids were glutamic acid, aspartic acid and alanine. With respect to carbon compounds, a significant increasing trend (P<0.05) in the lipid and fatty acid contents in the three tissues was observed, and, consequently, there was also little evidence of accumulated lipid storage reserves being used for egg production. It seems that for egg production both Octopus species use energy directly from food, rather than from stored products. This direct acquisition model contrasts with the previous model for Octopus vulgaris proposed by ODor and Wells (1978: J Exp Biol 77:15–31). Most of saturated fatty acid content of the three tissues was presented as 16:0 and 18:0, monounsaturated fatty acid content as 18:1 and 20:1 and polyunsaturated fatty acid content as arachidonic acid (20:4n-6), eicosapentaenoic acid (20:5n-3) and docosahexaenoic acid (22:6n-3). Though cholesterol is an important precursor of steroid hormones, this sterol content exhibited variations that do not seem to be related with the maturation process. Moreover, significant differences (P<0.05) were obtained between genders, suggesting that perhaps there is a greater physiological demand for cholesterol during spermatogenesis than oogenesis. If the component sterols of octopus are of a dietary origin, considerable variation in the cholesterol content between species might be expected on the basis of the sterol composition of their prey. The glycogen reserves increased significantly in the gonad and decreased significantly (P<0.05) in the digestive gland and muscle of O. vulgaris (these trends were not evident in O. defilippi). Glycogen may play an important role in the maturation process and embryogenesis of these organisms, because carbohydrates are precursors of metabolic intermediates in the production of energy. It was evident that sexual maturation had a significant effect upon the gonad energy content, but the non-significant energy variation (P>0.05) in the digestive gland and muscle revealed no evidence that storage reserves are transferred from tissue to tissue. The biochemical composition of digestive gland and muscle may not be influenced by sexual maturation, but rather by other biotic factors, such as feeding activity, food availability, spawning and brooding.Communicated by S.A. Poulet, Roscoff     

Depth-related variability in the photobiology of two populations of<Emphasis Type="Italic"> Halophila johnsonii</Emphasis> and<Emphasis Type="Italic"> Halophila decipiens</Emphasis>

The threatened seagrass Halophila johnsonii Eiseman coexists subtidally with H. decipiens Ostenfeld in southeastern Florida, but only H. johnsonii also occurs intertidally. Pulse amplitude modulated fluorometry and fiber-optic spectrometry were used to investigate the photobiology of two populations of H. johnsonii and H. decipiens in an attempt to explain these distribution patterns. Maximum photosynthetic quantum yields (Fv/Fm) were measured in situ as a function of depth distribution within, and between, these two species at two sites (Jupiter Sound, 26°57′N; 80°04′W, and northern Biscayne Bay, 25°55′N; 80°07′W) along the east coast of Florida, USA, during 6–10 March 2001. Reciprocal transplants at the northern site were used to evaluate the plasticity of photosynthetic patterns and pigment absorption spectra and to gain insights into the mechanisms responsible for variations in the observed depth-distribution patterns. Subtidal-population Fv/Fm values were generally higher for H. johnsonii than for H. decipiens, at both sites. At the northern site, intertidal H. johnsonii had significantly lower Fv/Fm (0.494±0.138) than both subtidal H. johnsonii (0.696±0.045) and subtidal H. decipiens (0.668±0.048). In contrast, at the southern site intertidal H. johnsonii had the highest Fv/Fm (0.663±0.047) and were the largest plants. Fv/Fm values of subtidal plants of both species decreased when they were transplanted into shallow, intertidal beds. Correspondingly, Fv/Fm increased for intertidal H. johnsonii transplanted into the subtidal, 2 m deep beds. Rapid light curves indicated that H. decipiens had lower maximum relative electron transport rates (RETRmax) than did H. johnsonii. In addition, the onset of photoinhibition occurred at lower irradiances for H. decipiens (537–820 μmol photons m⁻² s⁻¹) compared to H. johnsonii (1141–2670 μmol photons m⁻² s⁻¹). RETRmax values decreased for intertidal H. johnsonii transplanted into subtidal beds, but they increased for both species when transplanted from subtidal to intertidal beds. Absorption spectra for the acetone-soluble leaf pigments of intertidal H. johnsonii exhibited a dominant peak near 345 nm; this UV peak was 30% lower for subtidal plants. Pigment absorption spectra for H. decipiens lacked the 345 nm peak and absorbances, normalized to leaf pairs, were lower across the spectrum. Our results indicate that photosynthetic tolerance to higher irradiances and presence of UV-absorbing pigments (UVP) in H. johnsonii may allow this species to exploit the shallowest waters without competition from the closely related, but UVP-lacking H. decipiens.     

Ground gleaning in horseshoe bats: comparative evidence from<Emphasis Type="Italic"> Rhinolophus blasii</Emphasis>,<Emphasis Type="Italic"> R. euryale</Emphasis> and<Emphasis Type="Italic"> R. mehelyi</Emphasis>

The 71 species of horseshoe bat (genus Rhinolophus) use echolocation calls with long constant-frequency (CF) components to detect and localize fluttering insects which they seize in aerial captures or glean from foliage. Here we describe ground-gleaning as an additional prey-capture strategy for horseshoe bats. This study presents the first record and experimental evidence for ground-gleaning in the little-studied Blasius horseshoe bat (Rhinolophus blasii). The gleaning bouts in a flight tent included landing, quadrupedal walking and take-off from the ground. The bats emitted echolocation calls continuously during all phases of prey capture. Both spontaneously and in a choice experiment, all six individuals attacked only fluttering insects and never motionless prey. These data suggest that R. blasii performs ground-gleaning largely by relying on the same prey-detection strategy and echolocation behaviour that it and other horseshoe bats use for aerial hawking.We also studied the Mediterranean horseshoe bat (R. euryale) in the flight tent. All four individuals never gleaned prey from the ground, though they appeared to be well able to detect fluttering moths on the ground. It is not known yet whether ground-gleaning plays a role in Mehelys horseshoe bat (R. mehelyi). In a performance test, we measured the ability of these three European species of middle-sized horseshoe bats (R. euryale, R. mehelyi and R. blasii) to take-off from the ground. All were able to take flight even in a confined space; i.e. the willingness to ground-glean in R. blasii is not related to a superior take-off performance. In contrast to ground-gleaning bats of other phylogenetic lineages, R. blasii appears not to be a specialist, but rather shows a remarkable behavioural flexibility in prey-capture strategies and abilities. We suggest that the key innovation of CF echolocation paired with behavioural flexibility in foraging strategies might explain the evolutionary success of Rhinolophus as the second largest genus of bat.Communicated by T. Czeschlik     

Geographic variation and behavioral evolution in marine plankton: the case of<Emphasis Type="Italic"> Mastigias</Emphasis> (Scyphozoa,Rhizostomeae)

Although complex behavior in marine zooplankton has been considered strong evidence of adaptation, ethological studies of marine zooplankton generally have not employed either the comparative approach or evolutionary perspective necessary to distinguish adaptation from any alternative. Consequently, the potential for intra-specific variation in the behavior of marine zooplankton has received insufficient attention and conclusions of adaptation remain poorly substantiated. Intra-specific comparison of patterns of migration and behavior for seven populations of golden jellyfish, Mastigias (Scyphozoa: Rhizostomeae), inhabiting isolated marine lakes and semi-enclosed lagoon coves in Palau document population specific differences in patterns of horizontal migration, vertical migration, pulse rate, swimming speed, and turning behavior. Evidence was found for symplesiomorphic behaviors, canalization, exaptation, adaptation, and probably once-deleterious traits. Behavioral evolution likely proceeded via, at least, relaxation of selection, trade-offs with morphology, and natural selection effected by predation. Behavioral patterns also may change with ontogeny. Geographic variation in the behavior of marine plankton therefore can be substantial and patterns of evolution complex. Behavioral evolution can rapidly generate coastal biodiversity. Thus, geographic variation in marine plankton is of potential interest to ethologists, evolutionary biologists, biogeographers, and conservation biologists.Electronic Supplementary Supplementary material is available in the online version of this article at Communicated by G.F. Humphrey, Sydney     

Geographical genetic structure and phylogeography of the <Emphasis Type="Italic">Sargassum horneri</Emphasis>/<Emphasis Type="Italic">filicinum</Emphasis> complex in Japan,based on the mitochondrial <Emphasis Type="Italic">cox3</Emphasis> haplotype

The genetic structure and phylogeography of the brown seaweed Sargassum horneri/filicinum complex in Japan were studied based on the mitochondrial cox3 haplotype. The cox3 haplotypes found were divided into three clades in a statistical parsimony network, among which there were large numbers of steps. Contrary to the reported large amount of drifting S. horneri along the Japanese coast, the three clades were dividedly distributed on the Japanese coast: the northern Pacific, the central Pacific, and western Japan. The western Japan S. horneri had haplotypes that were phylogenetically closer to those of S. filicinum than to the northern and central Pacific S. horneri populations. The S. filicinum populations were included within the western Japan clade and grouped together with the S. horneri samples from western Japan. Taken together with the unstable morphological diagnosis, this result suggests that S. filicinum should be reduced into a synonymy of S. horneri. The TMRCA analysis suggested that the divergence time of each clade may go back to the last interglacial period and a skyline plot suggested that the last glacial maximum had only a small effect on the population size of S. horneri. The geographic subdivision of the three groups, in spite of a large amount of drifting mats, suggests a limited contribution of drifting mats to gene flow on a large geographic scale. On a small geographic scale, a small number of haplotypes were shared between S. horneri-type and S. filicinum-type populations. This result suggests that populations of these two types are partially, though not completely, isolated from each other, possibly by selfing in S. filicinum-type populations or by a difference in peak reproduction.     

Genetic evidence for the existence of cryptic species in an endangered clam <Emphasis Type="Italic">Coelomactra antiquata</Emphasis>

Coelomactra antiquata is a commercially important bivalve species, but has been suffering from severe population decline due to over-exploitation and the deterioration of environmental conditions. Previous genetic survey of C. antiquata conducted with allozymes combined with morphology revealed high levels of genetic differentiation between northern and southern populations which suggests a cryptic species might exist in C. antiquata. To test this hypothesis, amplified fragment length polymorphisms (AFLPs) and 16S rRNA gene sequence were used to re-evaluate the spatial genetic structure of six populations of C. antiquata along the coast of China. Both genetic markers display a sharp genetic break between the four northern populations (northern lineage) and two southern population (southern lineage). Large numbers of private alleles (AFLP) were found within the northern or southern populations and a deep divergence of about 6.5% in 16S rRNA gene sequence between the northern and southern lineages suggests the occurrence of potential cryptic or sibling species of C. antiquata. Applying previously published rates of mutation, divergence between the two lineages is estimated to have occurred approximately 3 million years ago and may be due to allopatric isolation during the middle Pliocene times. While no genetic differentiation was found within the northern or southern populations in both AFLP and 16S mtDNA markers, the results indicate that the northern and southern lineage should be managed separately and any translocation between the two areas should be avoided.     

House formation and feeding behaviour of<Emphasis Type="Italic"> Fritillaria borealis</Emphasis> (Appendicularia: Tunicata)

Numerous marine animals use elaborate filtration mechanisms to feed on particles suspended in the sea. Video-microscopy reveals that the common appendicularian Fritillaria borealis do so in a unique way. They live inside acellular, mucous houses, where their tail undulations act like a peristaltic pump in a close-fitting tail chamber to force water through a complex set of filters. Suspended particles, between ~0.2 and 30 µm in diameter, are retained in this process and propagated towards the mouth of the organism to be ingested. Coarser particles are back-washed out of the house during brief intervals of tail arrest, when the elastic recoil of the house walls reduces its volume to one-eighth of its inflated state. At the same time, the bifurcated tip of the animal's tail slides into a sleeve-like extension of the house. This sleeve acts like a flipper to propel the animal and collapsed house several millimetres to a few centimetres through the sea as soon as muscular activity of the tail is resumed. After a second tail arrest, a special muscular twist brings the tail out of the sleeve, whereupon undulatory pumping movements are resumed to reinflate and process particle-laden water through the house. The complete behavioural cycle lasts ~30 s. The repetitive inflation and deflation cycles of the house are probably of importance for unclogging of the coarse "inlet" filters downstream to the tail pump, as well as for agglutination of the smallest food particles trapped in the house to ease their recapture in the endostylar pharyngeal filter during the ingestion process. The unique tail sleeve and flipper at the same time enable the animal to move away from the previously filtered water and rejected coarse particles, before the house is reinflated for resumed food filtration.     

Genetic analysis of tuna populations,<Emphasis Type="Italic"> Thunnus thynnus thynnus</Emphasis> and<Emphasis Type="Italic"> T</Emphasis>.<Emphasis Type="Italic"> alalunga</Emphasis>

The genetic population structures of Atlantic northern bluefin tuna ( Thunnus thynnus thynnus) and albacore ( T. alalunga) were examined using allozyme analysis. A total of 822 Atlantic northern bluefin tuna from 18 different samples (16 Mediterranean, 1 East Atlantic, 1 West Atlantic) and 188 albacore from 5 samples (4 Mediterranean, 1 East Atlantic) were surveyed for genetic variation in 37 loci. Polymorphism and heterozygosity reveal a moderate level of genetic variability, with only two highly polymorphic loci in both Atlantic northern bluefin tuna ( FH* and SOD- 1*) and albacore ( GPI- 3* and XDH*). The level of population differentiation found for Atlantic northern bluefin tuna and albacore fits the pattern that has generally been observed in tunas, with genetic differences on a broad rather than a more local scale. For Atlantic northern bluefin tuna, no spatial or temporal genetic heterogeneity was observed within the Mediterranean Sea or between the East Atlantic and Mediterranean, indicating the existence of a single genetic grouping on the eastern side of the Atlantic Ocean. Very limited genetic differentiation was found between West Atlantic and East Atlantic/Mediterranean northern bluefin tuna, mainly due to an inversion of SOD- 1* allele frequencies. Regarding albacore, no genetic heterogeneity was observed within the Mediterranean Sea or between Mediterranean and Azores samples, suggesting the existence of a single gene pool in this area.     

Feeding selectivities of the marine cladocerans<Emphasis Type="Italic"> Penilia avirostris</Emphasis>,<Emphasis Type="Italic"> Podon intermedius</Emphasis> and<Emphasis Type="Italic"> Evadne nordmanni</Emphasis>

We conducted grazing experiments with the three marine cladoceran genera Penilia, Podon and Evadne, with Penilia avirostris feeding on plankton communities from Blanes Bay (NW Mediterranean, Spain), covering a wide range of food concentrations (0.02–8.8 mm³ l^–1, plankton assemblages grown in mesocosms at different nutrient levels), and with Podon intermedius and Evadne nordmanni feeding on the plankton community found in summer in Hopavågen Fjord (NE Atlantic, Norway, 0.4 mm³ l^–1). P. avirostris and P. intermedius showed bell-shaped grazing spectra. Both species reached highest grazing coefficients at similar food sizes, i.e. when the food organisms ranged between 15 and 70 µm and between 7.5 and 70 µm at their longest linear extensions, respectively. E. nordmanni preferred organisms of around 125 µm, but also showed high grazing coefficients for particles of around 10 µm, while grazing coefficients for intermediate food sizes were low. Lower size limits were >2.5 µm, for all cladocerans. P. avirostris showed upper food size limits of 100 µm length (longest linear extension) and of 37.5 µm particle width. Upper size limits for P. intermedius were 135 µm long and 60 µm wide; those for E. nordmanni were 210 µm long and 60 µm wide. Effective food concentration (EFC) followed a domed curve with increasing nutrient enrichment for P. avirostris; maximum values were at intermediate enrichment levels. The EFC was significantly higher for P. intermedius than for E. nordmanni. With increasing food concentrations, the clearance rates of P. avirostris showed a curvilinear response, with a narrow modal range; ingestion rates indicated a rectilinear functional response. Mean clearance rates of P. avirostris, P. intermedius and E. nordmanni were 25.5, 18.0 and 19.3 ml ind.^–1 day^–1, respectively. Ingestion rates at similar food concentrations (0.4 mm³ l^–1) were 0.6, 0.8 and 0.9 g C ind.^–1 day^–1.Communicated by O. Kinne, Oldendorf/Luhe     

Patterns of gamete incompatibility between the blue mussels<Emphasis Type="Italic"> Mytilus edulis</Emphasis> and<Emphasis Type="Italic"> M</Emphasis>.<Emphasis Type="Italic"> trossulus</Emphasis>

Previous research on gametic incompatibility in marine invertebrates suggests that for highly dispersive marine invertebrate species, barriers to fertilization among closely related taxa are often incomplete and sometimes asymmetric. The nature of these barriers can dramatically affect the patterns of gene flow and genetic differentiation between species, and thus speciation. Blue mussels, in the genus Mytilus, are genetically distinct in allopatry yet hybrids are present wherever any two species within the group co-occur. The present study sampled M. edulis (L.) and M. trossulus (Gould) in May and June 2001 from the East Bay section of Cobscook Bay, Maine, USA (latitude 44°56′30″N; longitude 67°07′50″W), where the two species are sympatric. Gamete incompatibility was investigated in a series of laboratory fertilizations carried out in July 2001. The proportion of fertilized eggs typically exceeded 80% at sperm concentrations of 10³–10⁴ ml^?1 among intraspecific matings (n=18), but was <30% even at sperm concentrations in excess of 10⁵–10⁶ ml^?1 for interspecific matings (n=13). Further analysis indicated that approximately 100- to 700-fold higher sperm concentrations were required to achieve 20% fertilization in interspecific matings relative to intraspecific matings, indicating strong barriers to interspecific fertilization. The proportion of fertilized eggs did not follow this general pattern in all matings, however. The eggs from two (out of five) M. edulis females were almost as receptive to M. trossulus sperm as they were to M. edulis sperm. In contrast, the eggs from all M. trossulus females (n=3) were unreceptive to M. edulis sperm, suggesting that fertilization barriers between these species may be asymmetric. Given the experimental design employed in this study, the results are also consistent with a strong maternal or egg effect on the level of interspecific gamete compatibility in M. edulis.     

Temporary cyst formation of<Emphasis Type="Italic"> Heterocapsa triquetra</Emphasis> (Dinophyceae) in natural populations

Heterocapsa triquetra (Ehrenberg) Stein is a phototrophic marine dinoflagellate with wide coastal distribution. It is known to be capable of mixotrophy and diel vertical migration. The species was particularly abundant in the Gulf of Finland (the Baltic Sea) during the summers of 1996 and 1998, leading to discolouration of water on the south-west coast of Finland. Large-scale (50 m³) coastal mesocosm experiments in the north-west Gulf of Finland (the Baltic Sea) in the summers of 1996 and 1998 with daily mineral nutrient additions provoked a biomass increase of phytoplankton dominated by H. triquetra. From the first days of the experiment temporary cysts of H. triquetra were found in the bottom sediment water of the mesocosms. Maximum temporary cyst production rates reached values up to 20×10⁶ cysts m^–2 day^–1, accounting for <1% of the depth-integrated motile population size. The environmental features favouring temporary cyst production remain uncertain; zooplankton grazing and nutrient stress are potential factors. Temporary cysts of H. triquetra were observed in a unialgal culture (f/2 medium) isolated in summer 1999 from Eel Pond (Woods Hole, Mass., USA).Communicated by M. Kühl, Helsingør     

Genetic structure of<Emphasis Type="Italic"> Linckia laevigata</Emphasis> and<Emphasis Type="Italic"> Tridacna crocea</Emphasis> populations in the Palawan shelf and shoal reefs

Allozyme variation of 10 populations of Linckia laevigata at 8 polymorphic loci and 13 populations of Tridacna crocea at 6 polymorphic loci were analyzed to compare genetic variability and genetic affinities among reefs in Palawan, Philippines. Two to five populations were sampled from each of four regions: the shelf reefs in (1) northern Palawan and (2) southern Palawan and the offshore reefs in (3) the Kalayaan island group (KIG) in the South China Sea and (4) the Tubbataha shoals in the Sulu Sea. Heterozygosity was highest in populations of L. laevigata from the south shelf of Palawan and populations of T. crocea from the Tubbataha shoals of the Sulu Sea. The lowest heterozygosity estimates were from the reefs of the KIG in the South China Sea, for both species. Overall F _ST values for both species were significant, with an estimated average number of effective migrants per generation (N _EM) of 4.85 (~5 individuals) for L. laevigata and 3.54 (~4 individuals) for T. crocea. Within-region comparisons showed N_EM ranging from 6.29 to 92.34 for L. laevigata and from 3.40 to 6.30 for T. crocea. The higher gene flow among L. laevigata populations relative to T. crocea is consistent with the greater dispersal potential of the former species. Finer scale genetic structuring was evident in T. crocea populations. For both species, the Tubbataha reefs in the Sulu Sea have higher genetic affinity with the populations from the southern shelf of Palawan, while the reefs in the KIG had higher affinity with the northern Palawan shelf reefs. The north and south shelf populations have the least genetic affinity. Genetic patchiness among reefs within regions suggests the importance of small-scale physical factors that affect recruitment success in structuring populations in small island and shoal reef systems in Palawan.     

Mitochondrial evolution and phylogeography in the hydrozoan<Emphasis Type="Italic"> Obelia geniculata</Emphasis> (Cnidaria)

The distribution and genetic structure of many marine invertebrates in the North Atlantic have been influenced by the Pleistocene glaciation, which caused local extinctions followed by recolonization in warmer periods. Mitochondrial DNA markers are typically used to reconstruct species histories. Here, two mitochondrial markers [16S rDNA and cytochrome c oxidase I (COI)] were used to study the evolution of the widely distributed hydrozoan Obelia geniculata (Linnaeus, 1758) from the North Atlantic and the Pacific and, more specifically, in the context of North Atlantic phylogeography. Samples were collected from six geographic localities between 1998 and 2002. Hydroids from the North Atlantic, North Pacific (Japan), and South Pacific (New Zealand) are reciprocally monophyletic and may represent cryptic species. Using portions of the 16S rDNA and COI genes and the date of the last trans-Arctic interchange (3.1–4.1 million years ago), the first calibrated rate of nucleotide substitutions in hydrozoans is presented. Whereas extremely low substitution rates have been reported in other cnidarians, mainly based on anthozoans, substitution rates in O. geniculata are comparable to other invertebrates. Despite a life history that ostensibly permits substantial dispersal, there is apparently considerable genetic differentiation in O. geniculata. Divergence estimates and the presence of unique haplotypes provide evidence for glacial refugia in Iceland and New Brunswick, Canada. A population in Massachusetts, USA, appears to represent a relatively recent colonization event.Communicated by J.P. Grassle, New Brunswick     

Reproductive strategy of<Emphasis Type="Italic"> Calanoides carinatus</Emphasis> and<Emphasis Type="Italic"> Calanus helgolandicus</Emphasis> during a summer upwelling event off NW Spain

Reproductive activity and production of the calanoid copepods Calanus helgolandicus and Calanoides carinatus were measured during a summer upwelling event off the coast of NW Spain. The upwelling pattern affected the distribution and fecundity of both species in the study area. The demographic composition of both populations and the stage of gonad maturation (e.g. the high abundance of fertilised females with mature ova) indicated active reproduction. C. carinatus, a highly fecund species associated with the African upwelling zones and considered as an upwelling specialist, showed low production rates (overall means of 15 eggs female^–1 day^–1 and 3% body C day^–1), despite the fact that the food conditions (high phytoplankton biomass dominated by diatoms) seemed to be optimal for this species. By contrast, C. helgolandicus, a temperate species that shows a strong link between spring phytoplankton blooms and reproduction time, seems to be flexible enough to take full advantage of shorter-term, enhanced feeding conditions associated with the pulsed nature of the summer coastal upwelling. Both the egg and carbon-specific production rates attained by this species (overall means of 26 eggs female^–1 day^–1 and 12% body C day^–1) were similar to values reported for a spring bloom situation. This high production would imply a long spring–summer recruitment event of C. helgolandicus in these waters. For both species the stage of gonad maturation was significantly correlated with their egg production rates and likely influenced by the food conditions; a species-specific nutritional requirement for final oogenesis is suggested. The carbon condition factor (carbon weight/prosome volume) of C. carinatus females was higher than that of C. helgolandicus, suggesting differential use of the carbon ingested; C. helgolandicus seems to use all ingested carbon to produce eggs at a high rates, whereas C. carinatus seems to store part of the ingested carbon as lipid reserves to ensure female survival and to support production during subsequent unfavourable food conditions.Communicated by S.A. Poulet, Roscoff