Histological investigation of the reproductive cycles of the limpets <Emphasis Type="Italic">Patella </Emphasis><Emphasis Type="Italic">vulgata</Emphasis> and <Emphasis Type="Italic">Patella ulyssiponensis</Emphasis>

This study devised a staging system for, and monitored, the gonad development of the limpet species Patella vulgata and Patella ulyssiponensis on the South West coast of Ireland using histological techniques. Maturation began in the males of both species in January and in the females it began in March. There was no statistical difference in gonad development between sexes and between species. Spawning in the male P. vulgata occurred from September to December 2003 and in September and October 2004. In female P. vulgata spawning occurred from October to December 2003, no spawning of females was observed in 2004. In male P. ulyssiponensis spawning occurred in November and December 2003 and from September 2004 to December 2004. Spawning was observed from November 2003 to January 2004 and in September 2004 in female P. ulyssiponensis. Sex ratios also varied between the species and between months sampled. Nevertheless more males were observed in both species.     

Compressibility and shell failure in the European Atlantic <Emphasis Type="Italic">Patella</Emphasis> limpets

The specimens of Patella intermedia, Patella rustica, Patella ulyssiponensis and Patella vulgata were analysed for morphological and morphometric characters, and for the resistance to compression and crushing to a force applied at the apex. Shell shape in these species ranged from the high symmetrical cone, with a rounded base of P. rustica, to the flat, asymmetrical, narrow-pear-shaped base of P. ulyssiponensis. P. intermedia and P. vulgata showed intermediate morphologies. Shell thickness increased linearly with age, but differed in the four limpets. P. rustica had the thickest shells, and P. ulyssiponensis and P. vulgata had the thinnest shells. P. intermedia displayed intermediate shell thickness. Considering deformability and toughness, P. intermedia shells usually needed the highest force to compress in height, the highest pressure to collapse, and were appreciably deformed at collapse. On the opposite side, P. ulyssiponensis shells usually needed the lowest force to compress in height, the lowest pressure to collapse, and were much less deformed at collapse. P. intermedia shells were therefore the most deformable and tough, and P. ulyssiponensis, the most stiff and fragile. P. rustica and P. vulgata shells displayed intermediate behaviour. However, numerical simulations based on the finite element method using the experimentally determined shells’ geometry and thickness, but considering similar shells’ material and structure in the four species, predicted that shell toughness should be decreased in the order P. rustica >> P. intermedia > P. vulgata >> P. ulyssiponensis. P. rustica shells’ geometry (a high and centred cone) and thickness (very thick) were therefore, theoretically, the most fitted for shells to resist crushing by compression. Yet, in the experimental tests, they were not the most resistant shells. It was concluded that resistance to crushing was not a direct function of shell morphology and morphometry, but appeared to be mainly determined by shell deformability. This is most probably related to differences in the internal composition and architecture of the shell in the four species. By comparison with data reported in the literature on the pressures normally exerted by ocean waves, it was concluded that these limpets have resistances to crushing in far excess to pressures normally endured in nature, being therefore unlikely that these species are crushed by the action of even very strong waves on shores. Hard objects, like logs and boulders, can be hurled onto the shore by waves, and constitute a much greater threat to limpets than the waves themselves. The high toughness of the limpet shell can be related to resistance to the impact of incidental hard objects. There was no direct relationship between the habit preferences of these limpets and the resistance to crushing. Other factors are involved in the distribution of these species in the shores.     

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     

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     

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.     

Early life histories in the bryozoan<Emphasis Type="Italic"> Schizobrachiella sanguinea</Emphasis>: a case study

Mediterranean populations of Schizobrachiella sanguinea seem to span a wide range of contrasting benthic habitats. We have studied adult, larval, and recruit populations of S. sanguinea from sublittoral (approximately 10 m depth) communities at Blanes (Spain, NW Mediterranean). At the study site, the species colonises semi-obscure caves and, to a lesser extent, communities of sciaphilic algae. Our first goal was to show the periods of brooding, larval release, and recruitment. Plankton was collected about twice a week between July 1998 and June 2000 over an artificial reef. A total of 303 larvae were collected in 23 of the 102 days of sampling. At the study site the species shows a single, annual larval release period, which occurs from late March to June–July every year. Recruits were found from late May to late August 1999 and in late May 2000. A recruitment peak was observed in July 1999 and again in late May 2000. The presence of brooding adults corresponded to the periods of larval supply and recruitment. Our second goal was to investigate the causes of the variable distribution of the species along several communities in the north-west Mediterranean Sea. We describe adult distribution and the dynamics of larval supply, recruitment of early (15 days from settlement) and late (4 months from settlement) recruits. Patterns of larval presence and early recruit distribution along the communities on the reef did not reflect the adult distributions, whereas this was closely matched by the distribution of late recruits. Post-recruitment mortality strongly affected recruits of the PA (photophilic algae) and SA (sciaphilic algae) communities, whereas it was very low in the communities of SOC (semi-obscure caves). Mortality of recruits was density independent and colony survivorship in the SA communities, where recruitment was the highest, appeared strongly affected by competition with brown algae. Like the majority of cheilostomate bryozoans, S. sanguinea releases a coronate larva capable of active swimming and positive reaction to light. Such swimming abilities may allow larvae to actively select the substrate on which to settle and to avoid unsuitable substrates. It seems that larval behaviour does not explain the preference of S. sanguinea for sciaphilic habitats. High post-recruitment mortality due to various factors, especially competition with fast-growing algae, seems particularly relevant in determining the adult distributions observed.Communicated by S.A. Poulet, Roscoff     

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     

Population genetic structure of protected allis shad (<Emphasis Type="Italic">Alosa alosa</Emphasis>) and twaite shad (<Emphasis Type="Italic">Alosa fallax</Emphasis>)

Determining the magnitude of homing behaviour within migratory fish species is essential for their conservation and management. We tested for population genetic structuring in the anadromous alosines, Alosa alosa and A. fallax fallax, to establish fidelity of stocks to spawning grounds in the United Kingdom and Ireland. Considerable genetic differences were present among populations of both species, suggesting strong fidelity to breeding grounds and compatible with homing to natal origins. Moreover, the genetic structure of A. fallax fallax showed a clear pattern of isolation-by-distance, consistent with breeding populations exchanging migrants primarily with neighbouring populations. Spatial genetic differences were on average much greater than temporal differences, indicating relatively stable genetic structure. Comparing anadromous A. fallax fallax populations to the landlocked Killarney shad subspecies, A. fallax killarnensis (Ireland), demonstrated a long history of separation. These results demonstrating regional stock structure within the British Isles will inform practical management of stocks and their spawning habitats.     

Tropical fishes in a temperate sea: evolution of the wrasse <Emphasis Type="Italic">Thalassoma pavo</Emphasis> and the parrotfish <Emphasis Type="Italic">Sparisoma cretense</Emphasis> in the Mediterranean and the adjacent Macaronesian and Cape Verde Archipelagos

The northeastern Atlantic and the Mediterranean Sea share geological histories and display great faunal affinities. The majority of the Mediterranean species have Atlantic origins, with a few species with tropical affinities. These include the parrotfish Sparisoma cretense and the wrasse Thalassoma pavo that are restricted to the subtropical northeastern Atlantic, the Macaronesian archipelagos (Azores, Madeira, and Canaries) and the southern Mediterranean. The Pleistocene glaciations have been described as having different effects on the fauna of the two regions. During glacial peaks, Mediterranean waters remained warmer than those of the adjacent Atlantic. Within the eastern Atlantic, the effects of Pleistocene glaciations were differentiated. Here, we perform a comparative analysis focusing on T. pavo and S. cretense populations from the northeastern Atlantic and the Mediterranean to assess the effects of Pleistocene glaciations in these two species. Sequences from the mitochondrial control region were obtained and analyzed combining phylogeographic and demographic approaches. Gene flow between Atlantic and Mediterranean populations was shown to be very high. The Mediterranean populations of T. pavo and S. cretense showed high levels of genetic diversity, even in the eastern basin, pointing to an ancient colonization event. This suggests that both species must have been able to persist in the Mediterranean during the cold Pleistocene periods. Historical migration estimates revealed a Mediterranean towards Atlantic trend in the case of T. pavo, which may reflect the re-colonization of areas in the Atlantic by fish that survived the cold phases in relatively warmer Mediterranean refugia. Our data also showed that within the Macaronesian Archipelagos, migrations occurred from Madeira towards the Azores, for both T. pavo and S. cretense, thus supporting a post-glacial colonization of the Azores by fish that persisted in the warmer region of Madeira. Similar geographic distributions, thermal affinities, and means of dispersion for T. pavo and S. cretense resulted in a similar response to the effects of Pleistocene glaciations, as evidenced by identical phylogeographic patterns.     

Competitive interactions between two fucoid algae with different growth forms,<Emphasis Type="Italic"> Fucus serratus</Emphasis> and<Emphasis Type="Italic"> Himanthalia elongata</Emphasis>

Competitive interactions between two fucoid algae with different growth forms, Fucus serratus L. and Himanthalia elongata (L.) S.F. Gray were examined both in the laboratory and on a shore of the Isle of Man, Irish Sea. The growth of germlings of both species declined with increasing density, irrespective of whether they were with cohorts or rival species, indicating that intra- and interspecific competition occurred between germlings. H. elongata suppressed the performance of F. serratus at the germling stage by virtue of its larger initial size, and at the mushroom stage by forming a miniature canopy with the caps of the adjacent plants. In a field experiment, the mortality of H. elongata juveniles generally increased in mixtures with F. serratus and was highest when F. serratus were 50% of the plants. At the juvenile stage, the negative effect of F. serratus on H. elongata was more severe than the other way round. This was because F. serratus grows predominantly upwards, whereas H. elongata had already begun to expand laterally at the distal end. If F. serratus survives in sparse mixed stands with H. elongata juveniles, it can overgrow them and inhibit their subsequent survivorship and growth, probably by both shading and physical sweeping. H. elongata and F. serratus maintain their discrete monospecific stands because of the varying outcomes of mutual competitive exclusion resulting from their differing growth patterns. Thus it is possible for them to co-occur at a similar shore height.Communicated by O. Kinne, Oldendorf/Luhe     

Reproduction and sex reversal of the solitary vermetid gastropod<Emphasis Type="Italic"> Serpulorbis arenarius</Emphasis>

The reproductive biology of the Mediterranean vermetid gastropod Serpulorbis arenarius, a subtidal sessile solitary species that lives in shady rocky communities, was studied at two localities south-east of the Iberian Peninsula. Its population density is usually low, with specimens sometimes sparsely distributed. Its shell often appears to be covered by epibiotic organisms. The sex ratio is biased toward males (63%) in the populations of SE Spain, and the results point towards hermaphroditism with alternating sexuality. The results obtained suggest that all specimens are first males, change to females under some as yet unknown condition and, after the ovogenesis cycle has been completed, revert to the original sexual state (male). The possible adaptive forces that favour hermaphroditism in this species are discussed, and we hypothesise that the seizure of a spermatophore by males may stimulate sex change to female. Brooding females (51% of females studied) were found from March to December. In males, histological study of testes revealed that spermatogenesis seems to occur throughout the year. Females brooded between 5 and 28 (=15) egg capsules, which were attached by stalks to the inner surface of the shell. It is noteworthy that the gonads of two of the specimens brooding egg capsules were developing testes. The newly laid egg capsules were filled with an organic intracapsular matrix, in which 40–153 (=94) eggs were embedded. The mean diameter of the uncleaved ova was 352.5 µm. The organic intracapsular matrix and some nurse eggs were used as an extra-embryonic food source. Hatching occurred at an advanced veliger stage, immediately followed by metamorphosis. Therefore, the larval development of S. arenarius is of a lecithotrophic intracapsular type, with a short, free-swimming/crawling phase.Communicated by O. Kinne, Oldendorf/Luhe     

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.     

Variation in fertilisation abilities between hemiclonal hybrid and sexual parental males of sympatric water frogs (<Emphasis Type="Italic">Rana lessonae</Emphasis>,<Emphasis Type="Italic"> R. esculenta</Emphasis>,<Emphasis Type="Italic"> R. ridibunda</Emphasis>)

In many species, males and females mate with multiple partners, which gives rise to sperm competition and multiple paternity. The experiments on water frogs presented here demonstrate that such sperm competition can affect the structure and dynamics of mixed-species communities. The hybrid frog Rana esculenta (LR) mates with one of its parental species, usually R. lessonae (LL), although in some areas R. ridibunda (RR), to regain the premeiotically eliminated parental genome ("hybridogenesis"). Mixed LL/LR-populations are stable although hybrid numbers should continuously increase at the expense of parental animals, because of differences in female fecundity and other factors. This would finally lead to the extinction of the sexual host, followed by that of the sexual parasite, unless the reproductive superiority of R. esculenta is reduced by other factors, such as lower hybrid male fertility. Eggs from LL- and LR-females were fertilised in vitro by single- and multi-male sperm suspensions of LL-, LR- and RR-males. In all experiments, the proportion of offspring sired by R. esculenta sperm was significantly lower than that sired by R. lessonae or R. ridibunda sperm. Gonad mass, sperm morphology, sperm swimming velocity, and sperm survival did not explain these differences in fertilisation success; nor did gamete recognition and compatibility. Sperm density was the only trait that paralleled fertilisation success, but it offers no explanation either, because densities were equalised for the in-vitro fertilisations. In natural LL/LR populations, the significantly smaller amount, poorer competitive ability and lower long-term survival of R. esculenta compared to R. lessonae sperm will reduce the initial reproductive superiority of hybrids and contribute to the stabilisation of mixed water-frog populations. Differences in fertilisation ability are also likely to be relevant for the structure and dynamics of several other systems with encounters between eggs and sperm from different genotypes, ecotypes, ploidy levels and/or species.     

The gastropod statolith: a tool for determining the age of<Emphasis Type="Italic"> Nassarius reticulatus</Emphasis>

The microstructure, shape and appearance of the growth rings in statoliths of Nassarius reticulatus (L.) were investigated. This species possesses two statocysts, each containing a single spherical statolith of calcium carbonate of up to 0.22 mm in diameter in the largest animals. The relationship between statolith diameter (SD) and total shell height (TSH) is exponential [ln(TSH)=26.3SD–0.842], although the function is site specific. Statoliths of the largest whelks (>29 mm) contained three or four clearly defined rings, corresponding to TSH values of ~1.1, 4.6–5.3, 12.0–13.5 and 18.5 mm, respectively. The first ring likely represents the metamorphic ring that was deposited at the time of larval metamorphosis when the post-larval whelk adopted a benthic lifestyle. The estimated size of the whelks at formation of the second, third and fourth statolith rings closely matched the TSH inferred from the shell rings. It is concluded that the patterns of growth rings present in statoliths can provide information about the age and growth of N. reticulatus.Communicated by O. Kinne, Oldendorf/Luhe     

DNA markers indicate that distinct spawning cohorts and aggregations of Patagonian squid,<Emphasis Type="Italic"> Loligo gahi</Emphasis>, do not represent genetically discrete subpopulations

The Patagonian squid, Loligo gahi DOrbigny, has been described as having temporally and spatially identifiable spawning aggregations. Variation at six microsatellite loci was used to assess whether seasonal and geographical spawning groups around the Falkland Islands represent distinct sub-populations. Genetic variation at these loci is high in this species (mean expected heterozygosity=0.87; mean number of alleles=14.7). No evidence of significantly different allele frequencies was found, either between samples from putative spawning cohorts or geographical areas, indicating that L.gahi around the Falkland Islands comprise a single genetically homogeneous population. Age structure analysis of samples (from statolith growth increments) indicated substantial spread in hatching dates among individuals of similar size and maturity status, suggesting the potential for extensive interbreeding between seasonal cohorts. A sample of L.gahi from the SE Pacific (Peru) displayed distinctly different gene frequencies (and allele size distribution at one locus) from SW Atlantic samples, supporting the suggestion that SE Pacific and SW Atlantic populations may represent distinct subspecies.Contributed by J.P. Thorpe, Port Erin     

Diet of<Emphasis Type="Italic"> Octopus vulgaris</Emphasis> in False Bay,South Africa

The diet of Octopus vulgaris was analysed using instantaneous daytime observations, midden counts, and stomach contents and a total of 39 prey species were identified. From stomach contents, the most important prey species were Plagusia chabrus (64.6% IRI, index of relative importance) and Haliotis midae (21.6% IRI). Crustaceans were the most frequently found prey group in octopus stomachs (63.6% frequency of occurrence), followed by molluscs (37.6%), teleosts (11.2%), and polychaetes (10.8%). Prey size and diversity increased with increasing octopus size. From middens, the mean shell lengths of H. midae consumed by small, medium, and large O. vulgaris were 53.3, 72.6, and 86.0 mm, respectively. Compared with stomach contents, midden counts were 3 times higher for shelled molluscs, but 5 times lower for crustaceans and soft-bodied organisms. Similarly, instantaneous daytime observations were 3 times higher for shelled molluscs, but 5 times lower for crustaceans and 2 times lower for soft-bodied organisms.Communicated by G.F. Humphrey, Sydney     

Mitochondrial DNA lineages in the European populations of mussels (<Emphasis Type="Italic">Mytilus</Emphasis> spp.)

Marine mussels (Mytilus spp.) belong to a group of benthic species crucial to coastal ecosystems in Europe and are important for the cultivation industry. In the present study, the nuclear adhesive protein marker (Me15/16) was used for identification of Mytilus species in coastal areas, on a large geographic scale in Europe. Pure M. edulis populations were found in the White Sea and Iceland. M. edulis, M. trossulus and their hybrids were found in the Baltic Sea and the North Sea (Oosterschelde, The Netherlands). M. galloprovincialis, M. edulis and their hybrids occurred in Ireland. M. galloprovincialis populations were observed in the Sea of Azov (Black Sea), the Mediterranean and Portugal. The mitochondrial (mt) DNA coding-region ND2-COIII was studied by PCR (polymerase chain reaction) and RFLP (restriction fragment length polymorphism) assay methods. The mtDNA control region was studied by PCR. Substantial differentiation in the frequency of female haplotypes among the studied populations in Europe was observed. Despite isolation between the Mytilus taxa on a macro-geographic scale, considerable mitochondrial gene flow occurred between populations, with introgression in hybrid zones on a more local geographic scale in Europe. MtDNA of the Atlantic Iberian (Portugal) population of M. galloprovincialis was more similar to mtDNA in populations of M. galloprovincialis and M. edulis from the Atlantic coasts of the Ireland and M. edulis from the North Sea, than to M. galloprovincialis from the Mediterranean. Lower polymorphism of mtDNA in populations of the Baltic and Azov Sea mussels in comparison with other European populations was observed and can be explained by the recent history of both seas after the Pleistocene glaciation. In the M. galloprovincialis population from the Azov Sea, the presence of the male-inherited (M) genome was demonstrated for the first time by sequencing the control region and was observed at high frequency. Possible influence of mussel culture on geographic distribution of the Mytilus taxa in Europe is discussed.     

Phylogeographic differentiation of storm petrels (<Emphasis Type="Italic">Hydrobates pelagicus</Emphasis>) based on cytochrome<Emphasis Type="Italic"> b</Emphasis> mitochondrial DNA variation

We evaluated mitochondrial DNA (mtDNA) sequence variation in a 910 bp region of the cytochrome b gene of the storm petrel, Hydrobates pelagicus. Samples from birds collected from five populations in the North Atlantic Ocean and the Mediterranean Sea were investigated. Genetic differentiation within the Mediterranean basin was low but high in the Atlantic. Strong differences were noted between the Atlantic and the Mediterranean populations, confirming the distinction of the subspecies H. p. pelagicus and H. p. melitensis for the Atlantic and Mediterranean seabirds, respectively. Divergence between the two subspecies probably resulted from paleogeographic changes in the Strait of Gibraltar, which was likely the route used by H. pelagicus to invade the Mediterranean Sea. Current and past demography and ecology of the storm petrel is regarded as an explanation for the level of differentiation observed within each oceanic basin. We compare the phylogeographic pattern of the storm petrel to other seabirds that breed in the same regions.Communicated by S. A. Poulet, RoscoffC.C. and L.B. contributed equally to this paper.     

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     

In vitro reproduction of the turbellarian<Emphasis Type="Italic"> Urastoma cyprinae</Emphasis> isolated from<Emphasis Type="Italic"> Mytilus galloprovincialis</Emphasis>

Urastoma cyprinae is a species infecting the gills of several marine bivalves. Although there is some literature on this turbellarian, its life cycle remains unknown. In our work we have demonstrated that reproduction of U. cyprinae can be completed out of its host. More than 50% of turbellarians isolated from mussels (Mytilus galloprovincialis) secreted and cemented a cocoon to the well bottom during the first 72 h of incubation in seawater (34 salinity) at 14°C. Oviposition started at days 1–18 (average 4.8 days) and occurred in most cases inside the protective cocoon. Each Urastoma laid an average of 2.9 egg capsules (range 1–10) and 3.9 embryos were developed inside each egg capsule (range 1–11). Hatching started at days 20–43 (average 24 days). An average of 12.8 juvenile forms (range 1–64) escaped from the cocoon after hatching. The free-swimming juveniles showed a positive phototactic response and survived about a month after hatching. On the basis of our results, we propose a life cycle for U. cyprinae involving a sexual maturation parasitic period in the bivalve gills and a reproduction period including cocoon secretion, egg laying, and hatching that is entirely completed in the external environment.Communicated by S.A. Poulet, Roscoff