The occurrence and effects of Mytilicola intestinalis in Mytilus edulis

Mytilicola intestinalis was observed in the mussel Mytilus edulis in increasing numbers for the first time at Brighton (England), in October 1966; the populations here and at Whitstable were examined. Mussels exposed high in the littoral zone were less heavily infected than those lower down, the degree of infection being directly related to the duration of exposure in each tidal cycle. Silt in the intestine of the mussel is considered to act as a controlling factor in numbers of parasites present at Whitstable. Egg-bearing copepods were present in samples throughout the year, suggesting that breeding is not interrupted by the winter. Evidence indicates that juvenile stages of the parasite cause most damage to the host, due in part to their presence in the ramifications of the hepatopancreas. Recovery of the mussel from the effects of parasitation is rapid, following a reduction in parasite population density and number of juveniles. In the laboratory, M. edulis is more rapidly affected by lack of food at 10 °C than M. intestinalis. No dead parasites were seen during 4 months of laboratory storage. Juvenile parasites continned to mature, indicating that this period of time may be required for Mytilicola intestinalis to reach maturity at 10°C.     

Reproductive habits and life cycle of the small clam Kingiella chilenica (Bivalvia: Cyamiidae) in an estuarine sand flat from the south of Chile

A population of the small clam Kingiella chilenica Soot Ryen 1959 was studied from March 1986 to December 1988 in an intertidal flat at Queule River, in the south of Chile. The life cycle and life history pattern of the bivalve were established contrasting population structure and dynamics to its reproductive habits. Individuals are gonochoristic and semelparous, presenting a typical annual life cycle. The species is a sequential brooder whose embryos undergo direct development. After the brooding season (summer through autumn), the adults disappear gradually (autumn through winter). Recruited juveniles overwinter during a relatively long period, undergoing rapid growth during the spring to attain the adult stage during the summer. The number of brooded embryos increases in proportion to adult length cubed. Life history traits of this bivalve are compared to those reported for other small brooder clams. Some basic tendencies become apparent when traits for semelparous versus iteroparous species are contrasted. As in other semelparous sequential brooders (Transennella tantilla, Gaimardia bahamondei), the relationship between brood size and shell length obseved in K. chilenica does not fit the allometry hypothesis for marine brooding invertebrtes that allometric constraints on the brooding space limit the fecundity of larger individuals. Also contrary to theoretical predictions, small body size does not limit the diversity of a clam's developmental patterns. Possible explanations for this finding are discussed.     

Seasonal cycles in the carotenoid content in Mytilus edulis

The carotenoid content of Mytilus edulis was studied over a period of 3 years from 1965 to 1967; a seasonal cycle was observed which appeared to be controlled by the spring bloom of phytoplankton, and the sexual cycle of the mussel. Infestation by the copepod Mytilicola intestinalis and hours of exposure in the tidal cycle were found to contribute to variations in carotenoid content. Maturation of the gonads and spawning occurred in the late spring and early summer of the period studied. In addition, a second sexual phase was recorded in October 1967. The possible effect of exposure on the time of spawning is discussed. Lack of food affects both carotenoid content and the maturation of the gonads. M. edulis kept at 10 °C with little food showed no seasonal variation in the carotenoid content when compared with animals fresh from the shore. Lack of food resulted in regression in gonadal tissue after a period of time, and an absence of mature and spawning individuals.     

Seasonal variations in distribution and population structure of Microcalanus pygmaeus and Ctenocalanus citer (Copepoda: Calanoida) in the eastern Weddell Sea,Antarctica

The abundance, vertical distribution and population structure of two important small calanoid copepod species, Microcalanus pygmaeus (G. O. Sars) and Ctenocalanus citer Heron and Bowman, were studied in the eastern Weddell Sea in summer (January/February 1985), in late winter/early spring (October/November 1986) and in autumn (April/May 1992). The population of Microcalanus pygmaeus consisted mainly of copepodite stages CII and CIII in late winter/early spring and were concentrated between 500 and 200 m depth. In summer, stage CIV was the modal stage and the bulk of the population had ascended above 300 m. In autumn the population structure was bimodal with CI and CV dominating. Most of the population was concentrated between 300 and 200 m. In all investigation periods M. pygmaeus had their maximal concentrations in the thermo-pycnocline. The developmental stages CIII to CV of Ctenocalanus citer formed the bulk of the population in late winter/early spring. In October all developmental stages had their main distribution between 500 and 200 m, except females, which were concentrated in the upper 50 m. In November most of the population occurred between 200 and 50 m. The summer population was concentrated in the upper 50 m, and numbers increased dramatically as the new cohort hatched. Copepodite stages CII and CIII dominated the population at the end of January, while CIV dominated 2 wk later. In autumn, CV was the modal stage. The majority of the population was concentrated in the upper 100 m, but there was an increase in abundance below 300 m compared to summer. Age structure changed with depth with a younger surface population and an older one in deeper water layers. The seasonal change in number of M. pygmaeus is much smaller than that of C. citer; the summer:winter:autumn ratio of the former being about one, whereas the winter:summer/autumn of the latter was about nine. Early copepodite stages and adults of M. pygmaeus occurred throughout all investigation periods. The large proportion of early copepodite stages in April and in mid-October suggests autumn and early to midwinter breeding. Apparently, M. pygmaeus may reproduce and grow year-round or perhaps has a 2-yr life-cycle. In contrast, the dramatic increase in abundance of early copepodite stages of C. citer in summer suggests springtime reproduction.     

Life history of the sandy-beach amphipod Neohaustorius schmitzi (Crustacea: Haustoriidae)

Neohaustorius schmitzi Bousfield, the dominant macroscopic invertebrate on sandy beaches in North Carolina, USA, is strictly intertidal, and distributed from mean low tide to high tide, with an average density of 790/m². Location of greatest densities within the intertidal zone is dependent upon season and sex. An aggregated distribution is shown. N. schmitzi has 2 generations/year: a winter generation which lives about 8 months, and a summer generation which lives about 4 months. The reproductive season extends from February through October, with spring and summer peaks. Each female appears to produce only 1 brood of young. The number of young ranges from 2 to 14, depending upon the length and generation of the female.     

Genetic structure of hybrid mussel populations in the west of Ireland: two hypotheses revisited

In Ireland, mussels on exposed rocky shores constitute an interbreeding mixture of two forms of mussels, the blue mussel, Mytilus edulis, and the Mediterranean mussel, M. galloprovincialis. Results from an Irish study in the 1980s, using partially diagnostic allozyme markers, indicated that mussels higher up the shore were more galloprovincialis-like than those lower down. In this study we set out to test two hypotheses: (a) recruits arriving on the shore are composed of genetically distinct cohorts that settle preferentially at different levels on the shore, and maintain genetic distinctiveness into adulthood; (b) recruits are genetically homogeneous, but once settled they diverge genetically over time, due to within-habitat site specific-selection. The diagnostic Me 15/16 DNA marker was used to analyse the genetic composition of newly-settled spat recruiting to artificial substrates, which were placed at two-week intervals from May–October 2002, on the mid- and low shore areas of two exposed sites in Galway Bay. Adult mussels were also collected on each sampling date. Results did not support the preferential settlement hypothesis, i.e., the genetic composition of primary settlers (≤ 500 μm) was similar between tidal heights and shores. Neither was there evidence of post settlement selective mortality, as adults were genetically similar to settling spat. In spat and adults the frequency of the M. galloprovincialis allele was high (0.56–0.80), due to high frequencies of M. galloprovincialis (> 37%) and hybrid (> 33%) genotypes, and correspondingly low frequencies of the M. edulis genotype (< 11%). Adult mussels from a nearby sheltered estuarine site, while significantly different to exposed shore mussels, still had low frequencies of the M. edulis genotype (< 17%), indicating no apparent advantage for the genotype in this environment. There are indications that the genetic composition of mussels may be changing on the Atlantic coasts of Ireland.     

Positive effects of the introduced green alga, Codium fragile ssp. tomentosoides, on recruitment and survival of mussels

The green macroalga, Codium fragile ssp. tomentosoides, is an important component of sheltered low-shore assemblages on breakwaters along sandy shores in the northern Adriatic Sea. Macroscopic thalli of C. fragile are not perennial, but develop from propagules and/or undifferentiated forms in early spring, when the dominant native space-occupier, the mussel Mytilus galloprovincialis, recruits. By mid-summer, rapid growth of C. fragile leads to the formation of a dense canopy. We investigated the effects of juvenile and adult thalli of C. fragile on recruitment, survival and growth of mussels. Two experiments tested the hypotheses: (1) that recruitment of mussels is greater within patches of juvenile thalli (primordia) of C. fragile than on adjacent bare surfaces; (2) that the presence of a canopy of C. fragile affects the survival and growth of mussel recruits. The number of recruits of mussels was significantly larger within clumps of primordia of C. fragile than on bare surfaces. The removal of the canopy of C. fragile affected negatively the density of mussels after 2 months from the start of the experiment, but there were no effects on the mean size of individuals, nor on the size–frequency distribution. The same trend persisted after 4 months from the start of the experiment. These results show that re-colonisation of space by mussels is enhanced by C. fragile. Given that mussels, in turn, have the potential to reduce recruitment rates of C. fragile, quick recovery of mussel beds after disturbances could be crucial for controlling the abundance of this alga on breakwaters. Results also suggest that the effects of introduced species on native assemblages can be explained only through studies encompassing different life-stages of interacting organisms.     

Seasonal changes in abundance and development of<Emphasis Type="Italic"> Calanus pacificus</Emphasis> (Crustacea: Copepoda) in the Oyashio–Kuroshio Mixed Region

Seasonal changes in abundance and development of Calanus pacificus Brodsky were investigated by analyzing samples of different depth strata (0–150 m and 0–1000 m) collected monthly in the Oyashio–Kuroshio Mixed Region. Copepodite stage 5 (C5) emerged from dormancy and matured in early summer. A new generation appeared in July and developed into C5 during summer to autumn. Some of the summer generation arrested development at C5 and persisted below 150 m depth until the following early summer. Although the remainder matured and reproduced in October, a new generation was not observed at the surface during winter. These results suggest that C. pacificus shows two different life-cycle patterns, i.e. one generation annually, with overwintering C5 in deep waters, and two generations annually, with surface development during autumn to winter. The complex life-cycle patterns may be an adaptation to the highly fluctuating surface environment in the Oyashio–Kuroshio Mixed Region.Communicated by T. Ikeda, Hakodate     

Population dynamics of the tanaid Hargeria rapax (Crustacea: Peracarida) in a tidal marsh

The tanaidacean Hargeria rapax (Harger, 1879) was sampled along intertidal transects semi-monthly at one site and quarterly at two other sites in salt marshes on Sapelo Island, Georgia, USA, from July 1985 to July 1986. Tanaids were most abundant near the mean highwater line and became progressively less abundant at lower intertidal elevations. Population density was greatest in the winter (December to February) when there were >29 000 individuals/m² at one high intertidal station. Although reproductive individuals were present most of the year, peaks in reproductive activity occurred in autumn (late August to early November) and spring (early March to mid June). An increase in population density coincided with increased reproductive activity only in autumn. Tanaid cohorts produced in the spring and summer rarely survived beyond 6 to 8 wk, but those produced in the autumn overwintered and lived 22 to 26 wk. The sex ratio among mature individuals was 2.8:1 (females: males). Mature females ranged in size from 2.2 to 3.9 mm total length (TL) and mature males were 2.3 to 4.1 mm TL; there was no significant sexual difference (Student's t-test, P>0.05) in the mean TL of mature individuals. The mean (±SD) size of brooding females was 2.9±0.32 mm TL and the mean (±SD) nunber of offspring/brood was 8.3±4.99 young/female. The timing of tanaid reproduction together with the effects of predation by juvenile fish and crustaceans may account for most of the spatial and temporal patterns of tanaid abundance observed in this study. There was a significant linear relationship (P<0.001, r ²=0.54) between the growth rate (GR, mm/d) of individuals and average daily air temperature (°C) described by the equation: GR=0.00178 (°C)-0.00971. The potential annual contribution of tanaid production to higher trophic levels, estimated from knowledge of standing stocks, growth rates and fecundity, was 5.71, 0.91 and 0.46 g dry wt/m² for high, mid and low intertidal areas, respectively. The high intertidal marsh, which supports the largest and most persistent standing stock of H. rapax, provides a rich foraging area for aquatic predators at high tide and an important source of recruits from which tanaid populations at lower intertidal elevations are recolonized after periods of intense predation pressure.     

Detrimental effects of the isopod, <Emphasis Type="Italic">Edotia doellojuradoi,</Emphasis> on gill morphology and host condition of the mussel, <Emphasis Type="Italic">Mytilus edulis platensis</Emphasis>

There are many reported associations between mussels and other invertebrates, such as pea crabs, polychaetes, turbellarians and copepods, which live in their mantle cavities. The boundary between commensalism and parasitism is often indistinguishable because of insufficient knowledge or because the interaction is variable. Preliminary evidence led to a closer examination of the relationship between the mussel, Mytilus edulis platensis, and an isopod, Edotia doellojuradoi, previously described as commensalism. Monthly intertidal samples of mussels were taken from September 2004 to August 2005 at Caleta Cordova Norte (45°43′S, 67°22′W) in southern Argentina and assessed for the prevalence and abundance levels of isopods. Mussels with and without isopods were measured, examined for evidence of gill damage and their condition (soft tissue dry weight) was determined. The overall isopod prevalence in mussels was 57.9% and infestation varied with mussel length, with maximum occurence at 30.2 mm (medium-sized mussels). Experimental evidence indicated that the position of the isopod inside the mussel depended on the feeding activity of the mussel. Female isopods were observed grasping the ventral food groove of the gill demibranchs and feeding on the mucous food strand produced by the mussel. Juveniles and males were observed clustered together on the dorsal side of the single female in each occupied mussel, suggesting extended maternal care. Gill damage was observed in 58.2% of mussels at the Argentine site and was significantly associated with isopod occurrence. Categorical regression analysis showed that the most important factor associated with the degree of gill lesions was the number of male and juvenile isopods per mussel, followed by the length of female isopods and the developmental stage of juveniles. Conversely, the degree of gill damage decreased with increasing mussel length. Overall, E. doellojuradoi had a significant effect on mussel condition throughout the year, with low flesh weight in mussels with isopods, except during the austral summer and early autumn. In contrast to previous studies, which concluded that the isopod was a commensal, the present study clearly demonstrates that E. doellojuradoi is a parasite of M. edulis platensis. Other symbiotic interactions formerly classified as commensal might not be innocuous on further investigation, especially if samples are taken at multiple sites and at different times of the year.     

Reproduction in Anthelia glauca (Octocorallia: Xeniidae). I. Gametogenesis and larval brooding

Anthelia glauca Lamarck, 1816 is a gonochoric, external-brooding soft coral found in KwaZulu-Natal. It is reproductively active in the summer months. The development of gametes produced in late summer is arrested in winter. Several stages of gametes are found at the base of the polyps, and female polyps produce several cycles of larvae over an extended breeding period of 4 to 5 months. Larvae are brooded in a unique pharyngeal brooding pouch not yet described in other coral species. The brood pouch consists of an expansion of the pharynx with constrictions proximal and distal to the embryos and larvae. Our data suggest that egg transfer and fertilization occur at full moon and the mature larvae are released after new moon. Zooxanthellae are absent in A. glauca oocytes, but zooxanthella infestation commences at the immature larval stage. Received: 15 July 1997 / Accepted: 12 March 1998     

Life history strategy of a southern European population of brown shrimp (Crangon crangon L.): evidence for latitudinal changes in growth phenology and population dynamics

This study provides a comprehensive examination of the life cycle dynamics of the brown shrimp (Crangon crangon L.) in the Mondego estuary, Portugal, a habitat located near the southern edge of the range of this species in European waters. The phenology of all life stages was documented (from the occurrence of ovigerous females, developing embryos, planktonic larvae and benthic post-larvae), and migration patterns of larvae into and out of the estuary were examined. Temperature-dependent functions of egg and larval development were combined with data on field abundance to predict the timing and magnitude of occurrence of larvae and recruits. Compared to brown shrimp at higher latitudes, southern conspecifics grow slower, mature earlier and have smaller brood sizes, and larvae have a more protracted settlement period. The Mondego estuary, besides acting as nursery area, is a dynamic platform for C. crangon to use in different stages, sizes and seasons.     

Source characterisation and mid-term spatial and temporal distribution of polycyclic aromatic hydrocarbons in molluscs along the Basque coast (northern Spain)

The variability of polycyclic aromatic hydrocarbons (PAH) measured in the soft tissues of Mediterranean mussels (Mytilus galloprovincialis) and Pacific oysters (Crassostrea gigas) are investigated. Samples were collected from estuarine waters within the Basque Country (Bay of Biscay), between 2003 and 2011. PAH bioaccumulation showed some seasonality and significant differences were observed between cold (autumn–winter) and warm (spring–summer) seasons. Sites located within the ports of Bilbao and Pasaia showed the highest PAH concentrations in molluscs, and the highest percentages of samples above the established Environmental Quality Standards and Environmental Assessment Criteria. Probably due to human activities carried out in the area, no clear trends were observed, between 2003 and 2011, for the autumn data. Since the Basque coast is an area with high population density and industrial activity, the congener profiles (which reveal the predominance of tetra-aromatics) and the diagnostic ratios identified urban/industrial combustion processes as the main PAH sources. However, natural and petrogenic sources cannot be disregarded.     

Invertebrate introductions in marine habitats: two species of hydromedusae (Cnidaria) native to the Black Sea,Maeotias inexspectata and Blackfordia virginica,invade San Francisco Bay

The hydrozoans Maeotias inexspectata Ostroumoff, 1896 and Blackfordia virginica Mayer, 1910, believed to be native to the Black Sea (i.e. Sarmatic) and resident in a variety of estuarine habitats worldwide, were found as introduced species in the Petaluma River and Napa River, California, in 1992 and 1993. These rivers are mostly-estuarine tributaries that flow into north San Francisco Bay. Both species appeared to be well-established in this brackishwater habitat. Salinities at the collection sites were about 11 during the summer, rising to nearly 20 in the early autumn and falling to near 0 in the winter. Large numbers of all sizes of both species of medusae were observed and collected, indicating that the hydroid stages of the life cycles of the two are also well-established in these rivers. In the Petaluma River, populations of both species were at maximum in late July, with numbers of individuals declining through August and into September; the Napa River was sampled only in October, and at that time only B. virginica was found. Examination of full guts of M. inexspectata and B. virginica medusae revealed that both species had fed nearly exclusively on small crustaceans, principally barnacle nauplii, copepods and their eggs and nauplii, and crab zoea larvae (M. inexspectata only). All the M. inexspectata medusae were males, indicating that the population has probably developed from the introduction of perhaps only a single male polyp or polyp bud. In spite of its inability to reproduce sexually, this population appears to be maintained by the prodigious ability of the polyp to bud and reproduce asexually, and is fully capable of invading additional low-salinity habitats from its present Petaluma River site. Male and female B. virginica medusae were collected in both the Petaluma River and the Napa River, indicating that B. virginica may have been introduced by either the polyp or medusa stage (or both), but that multiple individuals (of both sexes) must have arrived from another port in one or more invasions. As indicated for M. inexspectata, the B. virginica population will also probably seed new populations in San Francisco Bay and elsewhere. Based on its cnidome as well as the morphology of both medusa and polyp, M. inexspectata has been reclassified by moving it from the family Olindiidae, Limnomedusae, to the family Moerisiidae, Anthomedusae.     

Population dynamics of the commensal spider crab Inachus phalangium (Decapoda: Maiidae)

The ecology of the spider crab Inachus phalangium (Fabricius, 1775) (Decapoda: Maiidae) was studied in the field. I. phalangium inhabits the sublittoral on the sea anemone Anemonia sulcata Pennant. From July 1981 to April 1984 in the Mediterranean (Banyuls sur Mer, southern France) more than 3000 anemones were examined and ca 1800 I. phalangium were found on them. The population dynamics' generation cycles, reproductive activities and the dynamics of the sex ratio were investigated. The density of juveniles (crabs before the pubertal moult) on anemones changed in a yearly cycle from low in the first six months to very high in the second six months. The first occasional young crabs of a generation appeared in March/April (3rd and 4th decapodite stages) on the anemones. Their density increased enormously in the following months. The generation grew gradually on the anemones and moulted into puberty in September-January. Density of adults (crabs after the pubertal moult) on anemones changed in a yearly cycle from low to high from the summer to winter months. A new adult generation was recruited every autumn through the pubertal moult and disappeared in the following summer. Female reproductive activity continued throughout the year. Females carried several broods in succession, but the frequency of breeding females fluctuated on a yearly cycle. The highest percentage of egg-carrying females, i.e. the peak of the reproductive season, lay in the first half of the year. The maximal life span of a generation, from the hatching of the first larvae to the disappearance of the last adults, lasted 1.5 to 2 years. Males moulted into puberty ca one month later than females. The moulting distribution of adult males had roughly the same course as in females. An adult male generation diet out about one to two months before the female generation. Life expectancy was therefore 14 to 17 months for females and 12 to 15 months for males. The sex ratio of juveniles shortly before the pubertal moult was balanced. The sex ratio of adults shifted from 1:1 at the beginning of the reproductive period to ca 1:9 in favour of females at its end.     

Reproductive isolation and reproductive output in two sympatric mussel species (Mytilus edulis, M. trossulus) and their hybrids from Newfoundland

The mussels Mytilus edulis L. and M. trossulus Gould are found sympatrically in most areas of Newfoundland, with a low frequency of hybrids. To assess the potential for reproductive isolation, we sampled mussels from three sites in an eastern Newfoundland Bay from May–October 1996 to determine if there were differences in the reproductive cycles of the two species and their natural hybrids. In mussels with sheil lengths of 38–42 mm, males and females with mature gametes were dominant in June for M. edulis and hybrids, while M. trossulus showed a lower frequency of individuals with mature gametes. M. trossulus and hybrids spawned over a prolonged period (from late spring to early autumn) compared with most M. edulis individuals that spawned over a period of 2–3 weeks in July. This asynchrony in spawning activity between the two species may partially explain the low frequency of hybrids found in previous studies of these mussel populations. Female and male hybrids between M. edulis and M. trossulus showed normal gonad development, ripening and spawning, providing an opportunity for the introgression of genes between the two species. M. trossulus had a higher reproductive output than M. edulis of similar shell length, while hybrids showed intermediate values of reproductive output. M. trossulus females produced smaller eggs than either M. edulis or hybrids. Differences in reproductive traits may partially explain the maintenance of the mussel hybrid zone in Newfoundland. Published online: 13 August 2002     

Life cycles ofCalanoides acutus,Calanus simillimus andRhincalanus gigas (Copepoda: Calanoida) within the Scotia Sea

The life cycles and distribution of three dominant copepods,Calanoides acutus, Calanus simillimus andRhincalanus gigas were studied from the Discovery collections in the Scotia Sea earlier this century.C. simillimus is a sub-Antarctic species which mates in the top 250 m mainly in spring. The rapid development of the summer generation may allow a second mating period and a smaller second generation to appear in late summer.C. simillimus remains in the surface layers for a longer period thanCalanoides acutus orR. gigas, and its depth distribution is bimodal throughout the winter.R. gigas is most abundant in sub-Antarctic waters to the north of the Polar Front. It mates within the top 750 m later in spring, and development seems less synchronised than that of the other two species, with egg laying and the growth season being more protracted. Stages CIII and CIV are reached by the first autumn and further development resumes very early the following spring. It is not clear whether the majority then spawn or whether a further year may be needed to complete the life cycle. The predominantly Antarctic species,C. acutus mates below 750 m in middle to late winter and the summer generation develops rapidly to either CIV or CV. Its lifespan seems typically 1 yr, but some of the CVs which fail to moult and spawn in winter survive into their second summer, and their subsequent fate is uncertain. The cold-water speciesCalanus propinquus is comparatively rare in the Scotia Sea and aspects of its distribution and life cycle are briefly described for comparison. Regional variations in the timing of these events were apparent forC. simillimus and possiblyCalanoides acutus, but were not seen inR. gigas. Their geographic and vertical separation, together with their asynchronous life cycles support the concept of habitat-partitioning of these dominant herbivores.     

Recruitment,growth, and diel vertical migration of Euphausia pacifica in a temperate fjord

The pelagic crustacean Euphausia pacifica Hansen was sampled with a multiple-sample 1.0 m² Tucker trawl and a multiple-sample 1.0 m² vertical net in Dabob Bay, Washington on 17 dates between May 1985 and October 1987. Size (stage) structure and abundance of the population were determined for each date, while vertical distribution and diel migration were determined for 13 dates. Although internannual variability in both timing and magnitude of events occurred, consistent patterns were discernable. The population produced a large pulse of larvae (2 to 5 mm) in late spring of each year, apparently in response to the vernal phytoplankton bloom. Much lower abundances of larvae occurred during summer and autumn of each year, and larvae were completely absent during winter. Recruitment to the juvenile (6 to 9 mm) and adult (10 mm) stages was strongest during the summer, with abundances of these individuals peaking in summer and autumn. Individual growth rates, determined by modal progression analysis, were calculated for E. pacifica. Rates ranged from zero for some adult cohorts during the winter to 0.12 mm d^-1 for larvae during spring. The latter are among the highest ever reported for this species in the field. The vertical distributions and diel vertical migrations (DVM) of E. pacifica varied seasonally and between size (stage) classes. At night, all size classes were distributed in the surface layer (upper 25 m) irrespective of season or year. During the day, the larger/older stages were always distributed at middepths (50 to 125 m). In contrast, the daytime distribution of the larvae was more variable, being concentrated at the surface during spring and early summer of 1985, and at increasing depths later in the summer and autumn of 1985 and again in spring of 1986. This resulted in invariant DVM in the juveniles and adults, but variable DVM in the larvae, the latter of which is hypothesized to be a response to variable abundances of zooplanktivorous fish.     

Predator-induced alarm responses in the common periwinkle, Littorina littorea: dependence on season, light conditions, and chemical labelling of predators

Chemically mediated alarm reactions of the common periwinkle, Littorina littorea (L.), were studied in laboratory experiments during two consecutive summers, and one intermediate autumn season. Responses to chemical stimuli were detected as crawl-out responses, i.e. movements of snails out of the water. Snails were exposed to extracts of injured conspecifics, extracts of the mussel Modiolus modiolus (L.), and water conditioned by the predatory crab Carcinus maenas (L.), which had been maintained on different diets. In experiments carried out during the summer, a significantly larger number of snails moved out of the water when exposed to chemical stimuli from injured conspecifics, compared to chemical stimuli from injured mussels or filtered seawater. These results suggest that chemical alarm substances are present in L. littorea. Water conditioned by crabs that had been fed L. littorea released significantly more crawl-out responses compared to water conditioned by crabs that had been kept on a fish diet. When tested in autumn, no significant differences were found in responses to the above-mentioned water samples. Crawl-out responses under different light regimes were also investigated. All series of experiments carried out in the dark evoked a higher number of responses compared to series that took place in light. These findings may indicate an adaptation of snails to night-active predators. In total, the current results suggest that a L. littorea diet may chemically “label” the predator crab with snail alarm substances, and that predator-induced responses of L. littorea are actually responses to conspecific alarm substances released from crabs that have been maintained on a L. littorea diet. The response to the alarm signal, however, appears to be dependent on season and light conditions; some ecological implications of these findings are also discussed. Received: 8 January 1999 / Accepted: 29 March 1999     

The Mytilus edulis species complex in southwest England: effects of hybridization and introgression upon interlocus associations and morphometric variation

Loglinear analysis of electrophoretic data from two hybrid Mytilus edulis x galloprovincialis populations in southwest England revealed non-significant associations between genotypes at four allozyme loci, each of which is partially diagnostic for differences between the two taxa. Significant non-random genotypic associations within the context of the non-significant model involved all four assayed loci equally, consistent with their occurrence in a relatively tight linkage group. Multivariate analyses were used to examine electrophoretic variation from the two hybrid populations, and morphometric variation in the hybrid populations and in four allopatric (two M. edulis and two M. galloprovincialis) populations from western Europe While the number of hybrid mussels is high at both sites (22% at Croyde, 53% at Whitsand) the two taxa have largely maintained the genetic differences which exist between them in allopatry. However, morphological differences between the taxa have been eroded for mussels within the hybrid zone, whereas these differences are quite pronounced for mussels from allopatric populations. It is proposed that each taxon within the genus maintains its genetic identity, despite high dispersal potential, widespread hybridization, and high levels of introgression, as a result of adaptation to different environments. The worlwide occurrence of all four Mytilus hybrid zones at ecotones between recognized biogeographical provinces which are characterized by differences in temperature and salinity is consistent with such an interpretation.