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.     

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.     

Comparative life-history study on sympatric hyperiid amphipods (<Emphasis Type="Italic">Themisto pacifica</Emphasis> and <Emphasis Type="Italic">T</Emphasis>. <Emphasis Type="Italic">japonica</Emphasis>) in the Oyashio region,western North Pacific

Life-history features of the sympatric amphipods Themisto pacifica and T. japonica in the western North Pacific were analyzed based on seasonal field samples collected from July 1996 through July 1998, and data from laboratory rearing experiments. T. pacfica occurred throughout the year, with populations peaking from spring to summer. In contrast, T. japonica were rare from autumn to early winter, but became abundant in late winter to spring. Mature T. pacifica females and juveniles occurred together throughout the year, indicating year-round reproduction. Mature T. japonica females were observed only in spring, and juveniles occurred irregularly in small numbers, suggesting limited, early-spring reproduction in this study area. Size composition analysis of T. pacifica identified a total of eight cohorts over the 2 years of the study. Due to the smaller sample size and rarity of mature females (>9.6 mm) and males (>7.1 mm), cohort analyses of T. japonica were not comparable. Laboratory rearing of specimens at 2°C, 5°C, 8°C and 12°C revealed that a linear equation best expressed body length growth by T. pacifica, while a logistic equation best expressed body length growth by T. japoncia. Combining these laboratory-derived growth patterns with maturity sizes of wild specimens, the minimum and maximum generation times of females at a temperature range of 2–12°C were computed as 32 days (12°C) and 224 days (2°C), respectively, for T. pacifica, and 66 days (12°C) and 358 days (2°C), respectively, for T. japonica. The numbers of eggs or juveniles in females marsupia increased with female body length and ranged from 23 to 64 for T. pacifica and from 152 to 601 for T. japonica. Taking into account the number of mature female instars, lifetime fecundities were estimated as 342 eggs for T. pacifica and 1195 eggs for T. japonica. Possible mechanisms for the coexistence of these two amphipods in the Oyashio region are also discussed.Communicated by O. Kinne, Oldendorf/Luhe     

Spawning and development in <Emphasis Type="Italic">Osedax</Emphasis> boneworms (Siboglinidae,Annelida)

We report observations on spawning and early development in bone-eating worms of the genus Osedax. Individual females of Osedax rubiplumus were observed at 1820 m depth freely spawning hundreds of oocytes, and females of an undescribed species, Osedax “orange collar”, were observed spawning in laboratory aquaria. Cytological and molecular analysis of the spawned oocytes of two Osedax species revealed no evidence for the bacterial endosymbionts that the female worms require for their nutrition, suggesting that the bacteria must be acquired later from the environment, as they are in other siboglinids. Individual O. “orange collar” females released an average of 335 (±130) eggs per day, but the number of oocytes spawned per day varied greatly, suggesting that not all the females spawned daily. Fertilization rates of the spawned oocytes varied from 0 to 100%, though most females showed nearly 100% fertilization rates. Oocytes spawned in the laboratory at 4–6°C were negatively buoyant. If fertilized, these oocytes extruded polar bodies and then after at least four hours cleaved unequally. Subsequent cleavages occurred in a spiral pattern at roughly 2-h intervals, resulting in free-swimming trochophore larvae after 24 h. These lecithotrophic trochophores swam for 9–16 days before settling with several hooked chaetae, similar to those of dwarf Osedax males. The larval life span of the Osedax species studied in the laboratory appears to be shorter than in closely related Vestimentifera. Osedax rubiplumus, on the other hand, has much larger oocytes and so may have greater dispersal potential than these other Osedax species. The high fecundity and apparently continuous reproduction of Osedax boneworms permits the opportunistic exploitation of sunken vertebrate bones. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Factors affecting the interval between clutches in the hermit crab <Emphasis Type="Italic">Pagurus nigrivittatus</Emphasis>

Interval between reproductive events is an important factor for iteroparous animals because it determines the number of clutches throughout life. This study examined whether female size, clutch size, shell size and prenuptial molting affected the clutch interval in the hermit crab Pagurus nigrivittatus. Precopulatory guarding pairs of P. nigrivittatus were sampled in the field and kept in the laboratory until the female extruded eggs. The clutch interval of each female was assessed as one of two types of relatively “short” and “long” intervals by checking whether the guarded female had eggs and/or egg cases from the preceding brood or not when the guarding pair was collected. The clutch interval was longer in females with prenuptial molting than those without molting and these females usually grew larger at the prenuptial molt. This suggests that female P. nigrivittatus with a long interval might allocate energy into growth at the expense of the number of clutches during the current reproductive season. The allocation to growth is theoretically predicted to decrease with female size. Gastropod shell size is also known to affect the reproductive activity in hermit crabs. However, female size did not significantly affect the clutch interval in P. nigrivittatus, and the effect of gastropod shell size on clutch interval was not consistent with previous empirical studies. These results may be caused by differences in the gastropod species of shell occupied by the females of P. nigrivittatus.     

Fecundity,brood loss and egg development through embryogenesis of <Emphasis Type="Italic">Armases cinereum</Emphasis> (Decapoda: Grapsidae)

The present work is a comprehensive study of reproduction and embryonic development of Armases cinereum. Ovigerous A. cinereum (Bosc, 1802) females from Sebastian Inlet, Florida (9.88–19.4 mm CW) lay 2,000–12,000 eggs per brood, depending on their CW (mm): fecundity = 24.662 CW^1.9432. A. cinereum displayed significant brood loss through development (ca. 500 eggs per brood) independently from their CW (no senescence). However, since smaller females lay fewer eggs than larger ones, the percentage of eggs lost during embryonic development is greater in smaller females. The number of eggs carried on a later stage of development (potential fertility = 5.5593 CW^2.4417) is a more accurate estimate of the reproductive output and subsequent recruitment. Egg volume increased during development (64%, 0.025–0.041 mm³ or 0.36–0.43 mm of diameter, N = 270) and was strongly correlated with egg water content increase (19.21%, r = 0.89). Lipids, particularly fatty acids, seem to be the major energy source for embryonic development, decreasing 56.31 and 37.08% (respectively) during embryonic development; both are negatively correlated with egg volume (r = −0.90). The utilization of fatty acids through the different developmental stages of A. cinereum is presented. The most consumed fatty acids are the monounsatured (43.33 μg mg⁻¹ dw), followed by the saturated (29.91 μg mg⁻¹ dw) and polyunsaturated (24.03 μg mg⁻¹). Palmitic (16:0) and linoleic (18:2n-6) acids are preferentially consumed (19.5 and 17.9 μg mg⁻¹ dw, respectively). The high proportion of essential polyunsaturated fatty acids of C₁₈ and C₂₀ reflects the consumption of primary producers such as mangrove leaves. EPA/DHA ratio (2.85–3.84) and low DHA content indicated that this species appears in a medium-low level of the trophic chain. The low ratio of 18:1n-7/18:1n-9 and high percentage of 18:1n-9 (marker of carnivory) may be a sign of the consumption of juvenile invertebrates. The high percentage of odd-numbered FA indicated the occurrence of detritivores/scavenger behaviours. The fatty acid composition of the eggs reflects adult feeding ecology (omnivorous) and habitat.     

Brooding of the sub-Antarctic heart urchin, <Emphasis Type="Italic">Abatus cavernosus</Emphasis> (Spatangoida: Schizasteridae), in southern Patagonia

The reproductive cycle of the sub-Antarctic spatangoid sea urchin, Abatus cavernosus, was examined during a 2-year period in southern Patagonia, Argentina. The population studied is the northernmost known coastal population in the austral oceans, and is influenced by a greater annual range of seawater temperature than other Abatus species. The sex ratio of the overall A. cavernosus population did not differ significantly from 1:1, but was not homogeneous across size classes. A clearly defined annual reproductive cycle was found. Spawning occurred from May to July and was synchronous between sexes. Females were observed to be brooding within a 9-month period, from May to February. Juveniles were released during the austral summer, from January to February. The length at which 50% of the females were brooding occurred at a test length of 25.9 mm. A. cavernosus had a large egg size (mean diameter = 1.4 mm) and low fecundity (maximum = 57 eggs per female) compared to closely related species. The number of eggs within each brood pouch was highest in larger anterior brood pouch, which is close to two gonopores, as opposed to the other anterior and two smaller posterior pouches. Significant interannual variation was observed in gonad cycles, fecundity, and embryo development such as: brood size decreased during 2001; adjusted gonad dry weight and fecundity were higher during 2003. Hypotheses concerning the gonadal and brooding cycles and fecundity of sub-Antarctic and Antarctic Schizasteridae are discussed. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Predation on egg capsules of <Emphasis Type="Italic">Zidona</Emphasis><Emphasis Type="Italic">dufresnei</Emphasis> (Volutidae): ecological implications

Among the diverse patterns of energy allocation to the offspring of gastropods, the presence of egg capsules to protect embryos is common. Females of the edible snail Zidona dufresnei attach egg capsules to hard substrates in shallow Argentine Patagonian waters (40°45′S, 64°56′W) during spring-summer. Embryonic development takes about 30 days at 22°C. In this study, three likely capsule predator species and the marks left by each on egg capsule walls were identified in laboratory experiments in February 2010. Abundances of predators and egg capsules with evidence of predation were assessed in the field in the summers of 2010 and 2011. Under laboratory conditions (N = 10 replicates per treatment and control), the predation rate by the chiton Chaetopleura isabellei was the highest (up to 90%), followed by the gastropod Tegula patagonica and the crab Neohelice granulata (~20% each). Nearly 60% of 41 capsules found in the field showed signs of predation. According to the marks identified in the laboratory, C. isabellei was responsible for 79% of this predation, and T. patagonica for the rest. Predation appears to be important during the encapsulated early life and could be an agent for selecting for resistant capsule walls and a relatively shorter development time.     

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.     

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     

Conflict-involvement of male guinea pigs (<Emphasis Type="Italic">Cavia aperea</Emphasis> f. <Emphasis Type="Italic">porcellus</Emphasis>) as a criterion for partner preference

Polygynous mating systems can inflict considerable costs on males, often causing the emergence of alternative mating strategies to ensure reproductive success. These strategies can lead to different morphs of color, size, or behavior. The present work was done on guinea pigs, a polygynous rodent species. Some males can show a reduced readiness for conflict, perhaps employing an alternative mating strategy. To test this hypothesis, a two-stage experiment was set up, with males (N = 16) and females (N = 16) initially living in isosexual groups. Visual and olfactory contact was possible through a wire mesh. Male agonistic behavior was observed for 15 days, confirming the existence of less-conflict-involved (LCI, N = 7) and more-conflict-involved males (MCI, N = 9). Significant differences were found for a conditional parameter, body mass, and a morphometric one, testis width: LCI consistently surpassed MCI. Hormonally, cortisol was comparable, while testosterone was distinctly higher in MCI. Next, males and females were joined and observed for further 24 days. Males initially lost weight but reached original mass towards the end of the experiment. Testis width patterns were similar. Again, LCI had higher body mass. Cortisol was comparable, but testosterone release 3 days after merging was significantly higher in LCI. Behaviorally, LCI exhibited significantly less male–male aggression and more socio-sexual behavior than MCI. The former were more successful, with 57% capable of accessing females, in contrast to 11% of MCI. The existence of two distinct behavioral phenotypes in guinea pig males suggests that different reproductive strategies may exist in this species.     

Tolerance of Mediterranean seagrasses (<Emphasis Type="Italic">Posidonia oceanica</Emphasis> and <Emphasis Type="Italic">Cymodocea nodosa</Emphasis>) to hypersaline stress: water relations and osmolyte concentrations

The present study examines for the first time the effects of increased salinity on water relations and osmolyte (carbohydrates and amino acids) concentrations in two Mediterranean seagrass species, Posidonia oceanica and Cymodocea nodosa, which are adapted to growth in environments with contrasting salinity and have a known differential sensitivity to alterations in ambient salinity. The specific aim was to obtain insights into their respective capacities to cope with natural or anthropogenically induced (e.g. desalination plants) hypersaline stress and its ecological implications. To this end, large plant fragments of both seagrass species were maintained for 47 days in a laboratory mesocosm system under ambient salinity (37 psu; control) and three chronic hypersaline conditions (39, 41 and 43 psu). Analyses of leaf-tissue osmolality indicated that both species followed a dehydration avoidance strategy, decreasing their leaf water potential (Ψ_w) as the external salinity increased, but using different physiological mechanisms: whereas P. oceanica leaves exhibited a reduction in osmotic potential (Ψ_π), C. nodosa leaves maintained osmotic stability through a decrease in turgor pressure (Ψ_p) probably mediated through cell-hardening processes. Accordingly, the concentrations of soluble sugars and some amino acids (mainly Pro and Gly) suggested the activation of osmoregulatory processes in P. oceanica leaves, but not in C. nodosa leaves. Osmotic adjustments probably interfered with leaf growth and shoot survival of P. oceanica under hypersaline stress, whereas C. nodosa showed a more efficient physiological capacity to maintain plant performance under the same experimental conditions. These results are consistent with the more euryhaline ecological behaviour of C. nodosa and contribute to understanding the high vulnerability shown by P. oceanica to even mild increments in seawater salinity.     

Egg survival,embryonic development,and larval characteristics of northern shrimp (<Emphasis Type="Italic">Pandalus borealis</Emphasis>) females subject to different temperature and feeding conditions

Laboratory experiments on ovigerous females of northern shrimp (Pandalus borealis) were used to assess the effects of temperature and food ration on female condition during incubation and examine how combined effects of temperature and female condition influenced egg survival, embryonic development, and larval characteristics. Ovigerous females were maintained at 2°C, 5°C, and 8°C and fed on a low (three times/week; 2–2.7% W/W) or high ration (five times/week at satiation). The increase in temperature accelerated the developmental time of the eggs but their survival at 8°C was reduced. Conversion efficiency of yolk reserves in developing embryos was significantly reduced at elevated temperatures and larvae hatching at 2°C and 5°C were significantly larger and heavier than those hatching at 8°C. The experimental design did not result in any effect of food ration on the energetic condition of females or on egg characteristics and their biochemical composition. However, lower energy reserves were observed for females held at 8°C.     

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     

Temperature effects on fecundity,development and survival of the benthopelagic calanoid copepod, <Emphasis Type="Italic">Pseudocyclops xiphophorus</Emphasis>

The shallow-living, benthopelagic copepod species Pseudocyclops xiphophorus Wells (R Soc Edimburg 67:1967), collected over a yearly cycle from the fouling material in the brackish water Lake Faro (North-eastern Sicily), showed marked seasonal fluctuations in population abundances, with maximum numbers recorded in autumn. Highest in situ egg production rates coincided with periods of low adult and juvenile densities and vice versa, except in autumn when peaks in egg production and adult population densities were coincident. In this period, mean daily egg production rates reached a maximum of 4–5 eggs per female, when surface water temperature was 17–18°C. Egg production rates declined drastically in winter and were completely arrested when surface temperatures dropped to 10–12°C. In March, daily egg production rates began to increase again with an increase in ambient temperatures, reaching a maximum at the end of August. In the laboratory, as in the field, mean daily egg production rates were positively correlated with temperature, with values ranging from 2.2 ± 0.3 (16°C) to 8.9 ± 2.6 (30°C) (mean ± S.D.) eggs per female per day. At 32°C, P. xiphophorus females survived but did not reproduce. At 34°C, all specimens died after a few days. In terms of total egg production for the entire female lifespan, maximum values occurred at 16°C and minimum at 24°C. Temperature also dramatically affected female life span, which was shorter at higher temperatures. Development time of eggs decreased with increasing temperature, as also development time from egg to adulthood. Remating was necessary for the continued production of fertile eggs at 16°C because female life span was longer. The unique egg-laying behaviour in this species may ensure higher survival rates of egg stages compared to free-spawning and egg-carrying calanoid species. After releasing the egg pair, the female swims over the eggs with a rotatory motion, secreting a substance which facilitates the adhesion of the eggs to the bottom; she then continues to swim over the eggs until they are attached. Although egg production rates in this species are low compared to other pelagic copepods, they are within the range of values reported for egg-carrying species. The greater fecundity at higher temperatures compared to other subtemperate species indicates that the species is well adapted to the higher temperatures of coastal lagoons and brackish water lakes where it contributes to the biofouling community.     

Limited population structure in Northern and Spotted Wolffishes (<Emphasis Type="Italic">Anarhichas denticulatus</Emphasis> and <Emphasis Type="Italic">A. minor</Emphasis>) despite low apparent dispersal potential

Northern and Spotted Wolffishes (Anarhichas denticulatus and A. minor) are demersal marine fishes listed as “threatened” in Canadian waters. Both species have unusually large benthic eggs and large size at hatch, which should reduce passive dispersal. We examined population differentiation with microsatellite and AFLP loci across the ranges of both species in the North Atlantic Ocean. Although significant population structure was documented, differentiation was less than expected based on knowledge of life history characteristics. Significant differentiation was found in Northern Wolffish between the Barents Sea and other samples based on both microsatellite and AFLP data. In contrast, population structure in the Spotted Wolffish was notably weaker, particularly with microsatellites. Both species were characterized by low genetic diversity for marine fishes and had significantly lower genetic diversity than the congeneric Atlantic Wolffish. This finding was consistent with the conservation status of these three species and suggests potential vulnerability to over-exploitation in Northern and Spotted Wolffishes.     

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     

Population structure of the planktonic copepod <Emphasis Type="Italic">Calanus pacificus</Emphasis> in the North Pacific Ocean

The pelagic copepod Calanus pacificus ranges nearly continuously across temperate-boreal regions of the North Pacific Ocean and is currently divided into three subspecies—C. pacificus oceanicus, C. pacificus californicus, C. pacificus pacificus—based on subtle morphological differences and geographic location. The relation between geography and genetic differentiation was examined for 398 C. pacificus individuals sampled from six widely distributed locations across the North Pacific, including an open ocean site and coastal sites on both sides of the North Pacific basin. For each individual copepod, the DNA sequence was determined for a 421-bp region of the mitochondrial coxI gene (mtCOI). A total of sixty-three different mtCOI sequences, or haplotypes, were detected, with a sequence divergence between haplotypes of 0.2–3.1%. The number and distribution of haplotypes varied with sampling location; 12 haplotypes were distributed across multiple sampling locations, and 51 occurred at only one location. Five genetically distinct populations were detected based on F _ST values. Haplotype minimum spanning networks, nucleotide divergence and F _ST values indicated that individuals from coastal sites in the North Pacific Ocean were more closely related to each other than to individuals from the open ocean site at Station P. These results provide genetic support for the designation of two subspecies—a coastal subspecies that consists of what is currently referred to as C. p. pacificus and C. p. californicus and an open ocean subspecies C. p. oceanicus. This work also indicates that planktonic copepods with potentially high dispersal capacity can develop genetically structured populations in the absence of obvious geographic barriers between proximate locales within an ocean basin.     

Effect of starvation on the energy budget of two Asian horseshoe crab species: <Emphasis Type="Italic">Tachypleus tridentatus</Emphasis> and <Emphasis Type="Italic">Carcinoscorpius rotundicauda</Emphasis> (Chelicerata: Xiphosura)

Energy budget is one of the most studied parameters in aquatic animals under environmental challenge. To examine how prolonged starvation would affect their energy budget, respiration rate (RR), ammonia excretion rate (ER), oxygen consumption to ammonia–nitrogen excretion (O:N) ratio and scope for growth (SfG) representing the balance between energy intake and metabolic output, two Asian horseshoe crab species, Tachypleus tridentatus and Carcinoscorpius rotundicauda, were investigated in two feeding regimes (fed and starved) over a period of 7 weeks. No significant effects of species and time course, as well as their interaction, on absorption efficiency were observed in the fed treatments. For both species, RR and ER of the starved treatments significantly decreased, while their O:N ratio significantly increased during the experiment. However, such values for the fed treatments remained relatively stable over the study period. A rapid reduction in SfG was only apparent in the first week of the starved treatments for both species; thereafter, their SfG remained relatively constant. In the fed treatments, SfG of T. tridentatus was significantly lower than that of C. rotundicauda throughout the experiment. In general, C. rotundicauda showed a greater decrease in SfG under starvation than T. tridentatus, suggesting that they may have a more competitive life-history strategy for adjusting to poor nutritional conditions.