Variable within-brood maternal provisioning in newly extruded embryos of Homarus gammarus

Offspring quality of decapod crustaceans has been widely studied, with special emphasis on the sources of variability determining embryonic and larval quality. Nevertheless, maternal provisioning has commonly been overlooked as a potential source of offspring within-brood variability. In the present study, the existence of variable maternal provisioning was assessed through the analysis of the fatty acid (FA) profile of newly extruded embryos from different regions of the brooding chamber of Homarus gammarus. Significant differences in the FA profile of embryos sampled from different pleopods and sides of the brooding chamber were recorded. Significant deviations of the overall mean of each surveyed female were also observed for essential FA, particularly 20:4n-6, 20:5n-3 and 22:6n-3. Lipid energy available to fuel embryogenesis also varied among embryos sampled from different regions of the brooding chamber. Results suggest variable female investment at oocyte production, which may be amplified during the incubation period of developing embryos by differential lipid catabolism. For the first time, maternal provisioning is evidenced as an additional source for within-brood variability in the FA profile of embryos.     

Role of temperature on lipid/fatty acid composition in Pacific cod (Gadus macrocephalus) eggs and unfed larvae

During early development, oviparous fish species must use finite lipid and fatty acid (FA) reserves for both catabolism and structural components. In cold environments, developing fish have the additional constraint of maintaining membrane fluidity for metabolic efficiency (homeoviscous adaptation), resulting in further demand on lower melting point FAs like n-3 polyunsaturated fatty acids (PUFAs). To examine whether marine fish embryos physiologically adapt to changing temperature environments, we incubated Pacific cod (Gadus macrocephalus) eggs at 5 temperatures (0, 2, 4, 6, and 8 °C) in the laboratory and sampled them repeatedly during development to measure changes in lipid/FA composition. Pacific cod embryos increased n-3 PUFA content during the egg stage in all temperature treatments, with the possible exception of 0 °C, where poor survival and hatch success limited our ability for continued sampling. At the beginning of the hatch cycle, free-swimming embryos shifted from lipogenesis to lipid catabolism. The rates of lipogenesis and catabolism were temperature dependent, and the distinct increase in unsaturated fatty acids at temperatures <8 °C was consistent with homeoviscous adaptation theory. However, with the possible exception of embryos at 0 °C, the relative amounts of essential fatty acids (e.g., EPA, DHA, AA) were conserved in a similar manner across incubation temperatures. Collectively, these data suggest Pacific cod are capable of homeoviscous adaptation but cannot tolerate temperatures approaching 0 °C despite their possible ability to biosynthesize PUFAs from other energetic sources.     

Factors determining the hatching success of Antarctic krill <Emphasis Type="Italic">Euphausia superba</Emphasis> embryo: lipid and fatty acid composition

The present study addresses the effect of maternal diet on hatching success and condition of embryos and larvae of Antarctic krill Euphausia superba. Lipid and fatty acid content and composition were determined in field and laboratory samples. Developmental stages analyzed in embryos included: multiple-cell, gastrula, and limb-bud stages. Larval stages analyzed included: nauplius I, nauplius II, and metanauplius. Laboratory-reared embryos were spawned by gravid females incubated under three feeding groups: (1) phytoplankton mixture, (2) phytoplankton mixture and minced clam, and (3) phytoplankton mixture, minced clam, and commercial larval food. Hatching success was highest in group 3 (100%), lowest in group 1 (0%), and highly variable in field samples (0–48%). Lipid decreased slightly in embryos during embryonic development, while large decreases in lipid were found during nauplius development. High levels of 18:2(n-6), 20:4(n-6), and 22:6(n-3) observed with group 3 samples coincided with high hatching success in krill embryos. The ratio of 22:6(n-3)/20:5(n-3) also correlated to hatching success of embryos. The fatty acid profile of embryos in group 3 was similar to that of the field-collected embryos, reflecting the contribution of the commercial larval food in the maternal diet. In our study, the maternal diet was found to influence the fatty acid composition of embryos and in turn affects the hatching success of krill. Specific polyunsaturated fatty acids appeared to play important roles in embryogenesis in krill.     

Feeding ecology and energetics of the Antarctic chaetognaths Eukrohnia hamata, E. bathypelagica and E. bathyantarctica

Ecological and physiological studies focused on dietary preferences, lipid biochemistry and energetics within the three Antarctic chaetognaths Eukrohnia hamata, E. bathypelagica and E. bathyantarctica from meso- and bathypelagic depths. Eukrohnia hamata and E. bathypelagica respired 0.15 μL O₂ mg dry mass (DM)⁻¹ h⁻¹, which translates to an average metabolic loss of only <1.1% of body carbon per day. Lipid storage was not substantial in E. bathypelagica (mean 11.5 ± 6.5% DM) and E. bathyantarctica (mean 15.4 ± 4.1% DM) during summer and winter, suggesting year-round feeding of these predators mainly on copepods. In E. bathypelagica, total fatty acids were dominated by the fatty acids 16:0, 20:5(n-3) and 22:6(n-3) and in E. bathyantarctica also by 18:1(n-9), a fatty acid usually found in storage lipids. Only the latter species was characterized by significant amounts of wax esters, consisting largely of the common fatty alcohols 16:0, 20:1(n-9) and the unusual fatty alcohol isomer 22:1(n-9).     

The ecological significance of lipid/fatty acid synthesis in developing eggs and newly hatched larvae of Pacific cod (Gadus macrocephalus)

The lipid/fatty acid composition of marine fish eggs and larvae is linked with buoyancy regulation, but our understanding of such processes is largely restricted to species with pelagic eggs. In this study, we examined developmental changes in the lipid/fatty acids of eggs and embryos of Pacific cod (Gadus macrocephalus), a species that spawns demersal eggs along coastal shelf edges, but as larvae must make a rapid transition to the upper reaches of the water column. Adult Pacific cod were collected in the Gulf of Alaska during the spawning season and eggs of two females were artificially fertilized with sperm from three males for each female. The eggs were subsequently reared in the laboratory to determine (1) how lipids/fatty acids were catabolized during egg and larval development, and (2) whether lipid/fatty acid catabolism had measurable effects on egg/embryo density. Eggs incubated at 4°C began hatching after 3-weeks and continued to hatch over a 10-day period, during which there was a distinct shift in lipid classes (phospholipids (PL), triacyglycerols (TAG), and sterols (ST)) and essential fatty acids (EFAs: 22:6n-3 (DHA), 20:5n-3 (EPA), and 20:4n-6 (AA)). In the egg stage, total lipid content steadily decreased during the first 60% of development, but just prior to hatch we observed an unexpected 2–3-fold lipid increase (~6–9 μg individual⁻¹) and a significant drop in egg density. The increase in lipids was largely driven by PL, with evidence of long-chained fatty acid synthesis. Late-hatching larvae had progressively decreasing lipid and fatty acid reserves, suggesting a shift from lipogenesis to lipid catabolism with continued larval development. Egg density measures suggest that lipid/fatty acid composition is linked to buoyancy regulation as larvae shift from a demersal to a pelagic existence following hatch. The biochemical pathway by which Pacific cod are apparently able to synthesize EFAs is unknown, therefore representing a remarkable finding meriting further investigation.     

Costs of extended brood protection in the Antarctic sea star, Neosmilaster georgianus (Echinodermata: Asteroidea)

Changes in size, morphology, and biochemical composition in adults and embryos of a brooding sea star, Neosmilaster georgianus (Sladen), were studied in a population adjacent to Anvers Island, Antarctic Peninsula (64°46′S; 64°04′W) during the austral spring, 1991. Five morphological stages of development were designated in 24 broods, and for each the weight and biochemical composition of the brooding adults and their embryos were determined. Between Stage 1 and 2, the dry weight (dw) and organic weight (ow) of the embryo did not change. From Stage 2 to 3, the dw and ow increased significantly by 10%. Stage 2 and 3 embryos were in clusters of a few (2 to 10) to as many as 40 individuals. In the smaller clusters, individual embryos were attached by tissue cords to another, sometimes atrophied, brood member. In the larger clusters, they were attached to a central mass of tissue containing remnants of embryos. We interpret these interactions as a form of cannibalism which may account for the weight gains between Stage 2 and 3. During Stages 4 and 5, as juvenile form was approached, the dw and ash weight of the young increased significantly and the ow decreased significantly. The calculated energy content for the juvenile (Stage 5) was not significantly different from the energy content of the earliest undifferentiated stage (Stage 1), an indication that most of the organic matter in the egg is the primary contribution to the large juvenile. In brooding females, pyloric caeca indices declined by 52% from Stage 1 to Stage 5 and pyloric energy stores declined by 63% due to proportionately equivalent declines in protein and lipid. The ovary index was low and increased only slightly during brood protection, while the size of the largest oocytes remained approximately 23% that of ova. Energy stores in the pyloric caeca of brooding N. georgianus thus become depleted over a long period of incubation during which the adults apparently starve. This may delay oocyte development and ultimately limit the frequency of individual reproduction. Received: 15 September 1998 / Accepted: 9 March 1999     

Fatty acid composition of organs and tissues of the tiger prawn Penaeus esculentus during the moulting cycle and during starvation

The fatty acids (FA) in neutral lipid (NL) and polar lipid (PL) of digestive gland, muscle and integument of Penaeus esculentus Haswell were analysed and compared during the moulting cycle and during starvation. The prawns were collected from Moreton Bay, Queensland, Australia, by trawling during 1985–1987, and were fed with a standard semi-purified diet. Compared with a natural diet, the artificial diet had much higher levels of 18:1n-9 and 18:2n-6, but only trace amounts of 20:4n-6, but there was no evidence of dietary imbalance. The fatty acid composition (percentage of total lipid) of the digestive gland changed markedly during the moulting cycle and during starvation, but the small changes observed in both muscle and integument suggested that these tissues maintained their composition under both conditions. When the fatty acids were calculated as absolute amounts, muscle composition, as well as that of the digestive gland, changed significantly. In the digestive gland, saturated FA (SFA), monounsaturated FA (MUFA), diunsaturated FA (DUFA) and polyunsaturated FA (PUFA) all increased up to the middle of the moulting cycle and then declined; with starvation all groups decreased. In muscle, SFA, MUFA and DUFA all increased during the moulting cycle; starvation caused SFA, MUFA and PUFA to decrease, whereas DUFA did not vary. Starvation caused both 18:2n-6 (linoleic) and 18:3n-3 (linolenic) in the digestive gland to reach or almost reach zero. The other essential PUFA, 20:4n-6 (arachidonic), 20:5n-3 (eicosapentaenoic) and 22:6n-3 (docosahexaenoic), decreased during the moulting cycle, but during starvation 20:4n-6 did not decrease as much. In muscle, the levels of 18:2n-6 and 18:3n-3 increased, while 20:4n-6, 20:5n-3 and 22:6n-3 remained approximately constant during the moulting cycle. Starvation reduced 20:5n-3 and 22:6n-3 to about 60%. The data suggest that levels of 18:3n-3, 20:4n-6, 20:5n-3 and 22:6n-3 are regulated, and that 20:4n-6 can be synthesised from 18:2n-6. There is no clear evidence that 20:5n-3 and 22:6n-3 are essential in P. esculentus, but tissue catabolism of cell membranes during starvation may have provided sufficient amounts for maintenance.     

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.     

Cytochrome <Emphasis Type="Italic">b</Emphasis> (<Emphasis Type="Italic">Cyt-b</Emphasis>) gene sequence analysis in six flatfish species (Teleostei,Pleuronectidae), with phylogenetic and taxonomic insights

Mitochondrial DNA (mtDNA) at Cyt-b gene region was sequenced for six flatfish species (in total nine sequences of at least 1,117 bp) from the Far East of Russia and compared with other sequences of Pleuronectiformes comprising altogether 34 species and 3 outgroup species (Perciformes). Analysis of the protein-coding Cyt-b gene revealed a statistically substantiated bias in (T + C):(A + G) content, supporting earlier findings. Values of P-distances, as summarized for different scales of the evolutionary history at Cyt-b gene, revealed a clear pattern of increased nucleotide diversity at four different phylogenetic levels: (1) intraspecies, (2) intragenus, (3) intrafamily, and (4) intraorder. Scores of average P-distances of the four categories of comparison in flatfishes were (1) 0.46 ± 0.19%, (2) 11.74 ± 2.26%, (3) 17.51 ± 3.13%, and (4) 25.60 ± 0.42%, respectively (mean ± SE). These data support the concept that speciation in the order Pleuronectiformes, in most cases, follows a geographic mode through the accumulation of numerous small genetic changes over a long period of time. A phylogenetic tree for 64 sequences of flatfishes and 3 other fishes belonging to ray-finned fishes (Actinopterigii) was developed using Cyt-b gene and four different analytical approaches: neighbour-joining (NJ), Bayesian (BA), maximum parsimony (MP), and maximum likelihood (ML). The analysis revealed a monophyletic origin for the representatives of Pleuronectidae, which is the principal flatfish family investigated (99% repetition level in our BA analysis). A well supported property of the phylogenetic tree was the monophyletic placement of all the five flatfish families and the order Pleuronectiformes among other representatives of ray-finned fishes of the class Actinopterigii. Species identification on per individual basis (barcoding tagging) was generally high. However, there were taxonomic complications that arose during analysis, and they are discussed. Major outcomes of this discussion are the necessity of synonymy acceptance for Hippoglossoides elassodon, and H. robustus, and for Pseudopleuronectes yokohamae and P. schrenki. Priority considerations suggest H. elassodon and P. yokohamae as valid species names.     

Brooding behaviour and reproductive success in two species of free-living simultaneous hermaphrodites

Although polyclads are amongst the most structurally simple of the triploblastic metazoans, they adopt a wide range of reproductive strategies. Parental care behaviour in this group is yet to be quantified for any species. We assessed the significance of brooding behaviour to the reproductive success of two free-living marine flatworms. Echinoplana celerrima and Stylochus pygmaeus were collected from the field and placed in pairs in containers of filtered seawater where they laid batches of eggs. Both parents were then removed from half of the containers and the brooding behaviour and hatching success of eggs were quantified. There were interspecific differences in brooding behaviour. Egg masses were covered by one E. celerrima parent for 12 ± 2% of time, whereas egg masses of S. pygmaeus were covered by one or both parents simultaneously for 85 ± 8% of time. Egg batches were abandoned by both species immediately prior to the onset of hatching (10–12 days). Hatching success was generally high (~90%) and brooding did not enhance the hatching success of eggs. We assessed the significance of parental care to hatching success of E. celerrima egg masses in the presence of three potential egg predators; in the presence of other organisms. E. celerrima devoted less time to brooding; however, hatching success was not affected. The amount of time spent brooding eggs differed greatly between the two polyclad species but was not essential to their reproductive success under benign conditions. Parental care may be of adaptive value under more stressful environmental conditions commonly experienced in estuarine environments such as lowered salinity, increased hypoxia or turbidity. Covering egg batches may play an additional role of advertising sexual status and a willingness to care for eggs.     

Functional effects of the hadal sea cucumber <Emphasis Type="Italic">Elpidia atakama</Emphasis> (Echinodermata: Holothuroidea,Elasipodida) reflect small-scale patterns of resource availability

Holothuroidea represent the dominant benthic megafauna in hadal trenches (~6,000–11,000 m), but little is known about their behaviour and functional role at such depths. Using a time-lapse camera at 8,074 m in the Peru–Chile Trench (SE Pacific Ocean), we provide the first in situ observations of locomotory activity for the elasipodid holothurian Elpidia atakama Belyaev in Shirshov Inst Oceanol 92:326–367, (1971). Time-lapse sequences reveal ‘run and mill’ behaviour whereby bouts of feeding activity are interspersed by periods of locomotion. Over the total observation period (20 h 25 min), we observed a mean (±SD) locomotion speed of 7.0 ± 5.7 BL h⁻¹, but this increased to 10.9 ± 7.2 BL h⁻¹ during active relocation and reduced to 4.8 ± 2.9 BL h⁻¹ during feeding. These observations show E. atakama translocates and processes sediment at rates comparable to shallower species despite extreme hydrostatic pressure and remoteness from surface-derived food.     

Brooding and development of <Emphasis Type="Italic">Anasterias minuta</Emphasis> (Asteroidea: Forcipulata) in Patagonia,Argentina

Brooding, embryonic and larval development, and the influence of environmental and biological factors in tidepool habitats were studied in the sea star, Anasterias minuta, at various sites along ~220 km of the Patagonian coast. This species has a benthic, lecithotrophic development that includes eight distinct developmental stages. A larval organ, the connection cord, is developed from a small preoral lobe at early stages of development and becomes larger and thinner at advanced stages. Fecundity and average egg size increased with female body size. The regression of log egg number to log sea-star size and weight at different sites had a slope significantly less than 3.0, resulting in negative allometry and indicating that brood capacity was limited in large females. Development was generally synchronous among sites, but varied within each brood at advanced stages, with more developed brooded larvae located at the periphery of the brood mass. Brooding was synchronous among various populations at different years and spatial scales, and extended over a period of 8 months. The highest proportion of brooding females occurred during May and June (austral winter). Juveniles were released mainly during September. The likelihood of finding brooding sea stars decreased with increasing sea water temperature, tidal height, and wave exposure, and increased with increasing body size. Both body size of brooding females and brooding rate were higher in the infralittoral fringe than at midlittoral levels. A revision of the current model of brooding behavior and development among forcipulate sea stars is given.     

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.     

Reproductive biology of the spiny lobster, <Emphasis Type="Italic">Panulirus Penicillatus</Emphasis>, in the southeastern coastal waters off Taiwan

The reproductive biology of spiny lobster, Panulirus penicillatus, was studied based on 2,068 lobsters, ranging from 34.28 to 131.60 mm carapace length (CL), sampled in Taitung coastal waters from September 2003 to December 2004. The overall sex ratio approximated 1:1 (χ² = 0.02, P > 0.05), but the monthly sex ratios in 2004 showed significant differences and males were predominant in sizes larger than 80 mm CL. Reproductive activity, assessed using histology, a gonadosomatic index and percentage of ovigerous females, indicated that the mature females could be found in every month and that the major spawning occurred from May to September. The presence of re-developing/re-ripe ovaries by month and size-specific spawning time suggest that larger mature females (>60 mm CL) spawn at least three times a year while smaller new mature females spawn at least once a year. For females, the estimated sizes at 50% physiological and functional maturity were (mean ± SE) 56.46 ± 0.56 mm CL and 66.63 ± 1.07 mm CL. The estimated sizes at functional maturity were between 72 and 74 mm CL for males. The number of eggs per spawning event (brood size, BS) was related to CL by the equation Y _BS = 2.4 × 10^-3CL^4.18 (r ² = 0.902, n = 12). Female lobsters with CL ranging from 60 to 80 mm made the greatest contributions to egg production because of their high brood size and active reproductive activity. A minimum legal size should be established for the fishery to protect egg production potential of lobster population in the southeastern coastal waters off Taiwan.     

Developmental physiology of Antarctic asteroids with different life-history modes

Rates of respiration and protein synthesis were measured during embryonic and larval development of Antarctic asteroids with different life-history modes (non-feeding and feeding larvae: Acodontaster hodgsoni, Porania antarctica, Odontaster meridionalis). Patterns of respiration for these species all show an increase during embryogenesis, with subsequent maintenance of routine respiration (“starvation resistance”), even in the absence of food for ~4 months (O. meridionalis). Fractional rates of protein synthesis (i.e., rate per unit mass of whole-body protein content) in the Antarctic larvae are essentially identical to those of temperate species. Larvae of O. meridionalis had an average fractional synthesis rate of 0.52% ± 0.05 h⁻¹ at −1.0°C, which is comparable to the temperate asteroid Asterina miniata at 0.53% ± 0.14 h⁻¹ at 15°C. For embryos of the asteroids A. hodgsoni and P. antarctica, fractional rates of protein synthesis (~0.2% h⁻¹) also are comparable to those reported for embryos of temperate echinoderm species. While rates of synthesis are high, rates of protein deposition are relatively low (percent of protein synthesized that is retained for growth). During a ~4 month growth period for larvae of O. meridionalis, the average protein depositional efficiency was 5.2%. This contrasts with higher rates of depositional efficiency reported for similar developmental stages of temperate echinoderm species. The biological significance of maintaining high rates of macromolecular synthesis for species with low rates of cell division and low protein depositional efficiencies is intriguing in the context of understanding the mechanistic bases of extended life spans and dispersal potential in response to changing Antarctic environments.     

Apparent resource competition among embryos in the brood pouch of a male pipefish

Embryo success was studied in the paternally brooding pipefish Syngnathus typhle. During brooding, which lasts about a month, males provide embryos in their brood pouch with nutrients and oxygen via a placenta-like structure. Egg size depends on female size. In aquaria, males were mated with differently sized females to give the following treatments: M, mixed-egg-size broods of approximately half large and half small eggs; L, single-egg-size broods of large eggs; S, single-egg-size broods of small eggs; and F, field mated males. All males were kept in aquaria for a full brooding period. For each egg-size category, the number of newborn was compared with the number of eggs the male initially fertilized in his brood pouch. Within mixed-egg-size broods, a higher proportion of large eggs survived and large eggs resulted in heavier newborn than small eggs. Indeed, small eggs from a mixed-egg-size brood had significantly lower relative success (proportion of embryos surviving to birth) than those from a brood entirely composed of small eggs. The implication is that embryos compete for resources within the brood pouch, and that competitive success depends on egg size. Given that females produce eggs corresponding in size to their body size, and that females are known to compete indirectly for access to mates (i.e., the sex-roles are reversed), this intrabrood competition could be seen as an extension of female-female competition, but alternative explanations are discussed. Received: 28 April 1995/Accepted after revision: 28 October 1995     

Geographic,population, and seasonal patterns in the reproductive parameters of the sea urchin <Emphasis Type="Italic">Paracentrotus lividus</Emphasis>

Generalized additive models were used to synthesize the data from 52 publications on the gonad growth and spawning seasonality of Paracentrotus lividus and identify spatial and temporal patterns in the reproductive processes. According to our results, Atlantic populations spawn in spring and develop large gonads (6.4 ± 0.2% of fresh body weight (mean ± SE)), which increase in size toward higher latitudes. While in the Mediterranean, the gonads are smaller (3.1 ± 0.1%), possibly as the result of successive spawnings which impede nutrients accumulating in the gonads. Differences were also observed between habitats: gonad production was higher in seagrass meadows in the Mediterranean and in subtidal rocky substrates in the Atlantic. Food availability might cause these variations as well as the negative correlation observed between depth and the gonad index for rocky substrates. Sex and body size do not seem to influence greatly gonad growth.     

Ontogenic variations in fatty acid and alcohol composition of the pelagic amphipod <Emphasis Type="Italic">Themisto libellula</Emphasis> in Kongsfjorden (Svalbard)

The fatty acid and alcohol composition of the pelagic amphipod, Themisto libellula, was monitored during the 5 first months of its life cycle (4–20 mm length) in an Arctic fjord, Kongsfjorden, Svalbard. Fatty acids of the three major lipid classes, polar lipids (PL), triacylglycerol (TAG), and wax esters (WE), were analyzed to highlight ontogenic changes in their diet and metabolism. The PL composition of T. libellula did not show any strong variations along their growth except during the first month where an important increase of 20:5(n-3) (EPA) and 22:6(n-3) (DHA) was observed. The TAG composition revealed a clear gradient corresponding to a diet shift from omnivorous juveniles toward carnivorous sub-adults and adults. Indeed, fatty acid trophic markers of diatoms were dominant in the juveniles, whereas 20:1(n-9) and 22:1(n-11), the Calanus sp. trophic markers, overwhelmed in the older stages. The WE composition highlighted the same general trend, however, differences were found with the TAG and are discussed as a result of differences in turnover rates and assimilation pathways between the two lipid classes.     

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.     

Do female roe deer in good condition produce more sons than daughters

In polygynous roe deer Capreolus capreolus, males are only slightly heavier than females and the overall sex ratio at birth is close to unity. We studied offspring sex ratio and litter size (range 1–4, n = 74) of culled females, in utero, which provided an opportunity to examine responses of sex ratio to maternal condition. Male embryos were heavier than their sisters, and male fawns (9 months old) heavier than female fawns, suggesting a higher growth rate in males. There was no evidence for differential mortality between the sexes from birth to 9 months old. Heavier adult females produced larger embryos than lighter, or primiparous females. The overall sex ratio of embryos did not differ from unity, but adult does had more male embryos (55%) than primiparous does (32%), and the proportion of male embryos in a litter increased with the mother's body mass. Litter size also tended to increase with maternal age and body mass. We argue that this pattern reflects adaptive variation in offspring sex ratio.