Depth-related shift in life history strategies of a brooding and broadcasting deep-sea asteroid

Combining field and laboratory work, this study investigated the reproductive cycle, aggregative behavior, spawning periodicity, development and early growth of the sea star Henricia lisa living at bathyal depths off eastern Canada. Marked differences were found between individuals from ~1,300 and ~600 m deep. The former had a male biased sex ratio and an aperiodic reproductive cycle, whereas the latter displayed an equal sex ratio and a biannual breeding pattern. Furthermore, the maximum size was larger and female fecundity roughly five times higher in shallower compared to deeper populations. In the tanks, aggregative behavior was recorded twice a year during the summer and winter breeding periods. The onset of aggregations and spawning coincided with a temperature of 3–4°C. Males spawned first and females typically responded inside 30–60 min. Between 12 and 20 eggs were retained to be brooded under the arched arms of the female, whereas the remainder were broadcasted and developed without parental care. The fertilized eggs underwent a first cleavage after 12 h, reached the brachiolaria stage in 1 month, became juveniles within 3–4 months and reached ~ 4 mm in diameter after 14–17 months of growth. The embryos and juveniles developed at the same rate whether brooded or not, and development of winter cohorts was typically slower due to lower prevailing temperatures. This study of H. lisa provides the first evidence of lecithotrophy in a seasonally breeding deep-sea echinoderm and of brooding in a deep-sea asteroid.     

Reproductive periodicity,spawning and development of the deep-sea scleractinian coral <Emphasis Type="Italic">Flabellum angulare</Emphasis>

Increasing threats to deep-sea corals highlight the need to expand knowledge of these taxa so that conservation measures can be developed. The present study focused on the reproductive patterns of the deep-sea solitary coral Flabellum angulare. A series of samples (n = 398) collected in 2006–2008 in the northwest Atlantic at depths of 925–1,430 m revealed that gametogenesis was synchronous among males and females and fluctuated seasonally. Initiation of gamete synthesis was estimated to be in August–September and spawning in June. Further analysis and daily monitoring of 30–60 individuals maintained in a flow-through mesocosm showed that gamete release occurred in March–June with a peak in May. Release of oocytes coincided with rising seawater temperatures and high deposition rates indicative of elevated water column productivity. Oocytes (900–1,200 μm diameter) were released through the oral cavity, generally in bundles of 3–5 surrounded by mesenterial filaments and attached to a thread (30–50 mm long). As oocytes became free in the water column, ovulation occurred followed by fertilization. Eggs/embryos initially remained on the tentacles of the spawner before either falling onto the substratum or floating to the surface. The embryos developed into planula larvae measuring 2–3 mm in length within about 24 h. Together, these findings shed new light on the strategies used by deep-sea corals to maximize their reproductive success.     

Reproductive biology of a deep-sea brittle star Amphiura carchara (Echinodermata: Ophiuroidea)

 New information on the brooding reproduction of Amphiura carchara, and previous studies of several deep-sea congeners, contradict the prevailing notion that direct development is exceptional among deep-sea echinoderms. Over 500 specimens of A. carchara from 2,850 to 4,100-m depths off the coast of California, USA, were studied. The species was found to be gonochoric, although most brooding species of ophiuroids are hermaphroditic. Females each brooded up to 72 embryos at a time, with up to 10 in a single bursa. The embryos of individual adults were often at different stages of development, but those brooded in a single bursa tended to be at the same stage. Thus, the species has characteristics that breach the distinctions between sequential and simultaneous brooding exhibited by shallow-water ophiuroids. The embryos developed from yolky eggs that are large for an ophiuroid, with the mean largest oocytes 0.45 mm, and ranging to 1.28 mm in diameter. Almost all late-stage embryos were positioned with their mouth and arms pressed against the wall of the bursa, possibly to facilitate the uptake of nutrients from the parent. Thus, A. carchara may be matrotrophic. The largest embryos examined had a rudimentary disk skeleton, and arms with four joints and a terminal plate. Emerging juveniles probably differ in disk diameter. The species appears to brood year round, although differences in gonad size, the incidence of brooding, and the relative numbers of early developmental stages in summer and winter samples indicated that there are seasonal trends in reproduction. Received: 14 February 2000 / Accepted: 3 July 2000     

Reproductive biology in two species of deep-sea squids

Deep-sea squids, Moroteuthis ingens and Gonatus antarcticus, were collected in the slope waters off the Falkland Islands and their reproductive systems preserved and investigated onshore. Changes in oocyte length-frequencies at maturation and spawning, and their fecundity were studied. These squids, as well as many other species, are characterised by a synchronous oocyte growth and ovulation. Oviducts are not used for ripe egg accumulation and consequently the universal scale of Lipinski (1979) cannot be applied to assign female maturity. M. ingens spawns near the bottom; its fecundity is 168–297 thousand eggs. Maximum egg size is 1.8–2.7 mm. G. antarcticus spawns midwater; its fecundity is 10–25 thousand eggs. Egg size is 3.2–3.3 mm. In M. ingens spawning takes place in the austral autumn and winter, in G. antarcticus—in austral winter. Our data and the literature data show that the so-called “synchronous ovulation” probably occurs in all deepwater squids. This pattern is very rare among fish, but is quite common among benthic octopods that brood their egg masses.     

Reproductive biology and ecology of a deep-demersal fish, alfonsino Beryx splendens, over the seamounts off New Caledonia

The reproductive biology of the alfonsino Beryx splendens was studied by histological examinations, gonadosomatic index and macroscopic scales of maturation of a large sample of gonads. Alfonsino is a gonochoric species. The size-frequency distribution of the sex ratio was bimodal and considered to be due to size dimorphism. In New Caledonia, the breeding period of this species occurs during the southern summer, with a peak in December to January. The spawning stage is attained at a minimum fork length of 28 cm for females and 33 cm for males. The size at which 50% of the population attain sexual maturity (FL₅₀) is 33.2 cm for females and 34.5 cm for males. Maximum potential fecundity is estimated to lie between 270 000 to 675 000 eggs for fish between 34 and 40 cm in fork length. It was possible to differentiate vegetative zones, in which juvenile alfonsino grow until they reach maturity, from reproductive zones (fishing grounds) which are inhabited by mature individuals. The larvae and juveniles could be carried from the reproductive zone to the vegetative zone by currents in an oceanic eddy system. Received: 26 April 1996 / Accepted: 20 September 1996     

Multi-year observations on the gametogenic ecology of the Antarctic seastar <Emphasis Type="Italic">Odontaster validus</Emphasis>

This study reports the first multi-year observations on the reproductive patterns for an Antarctic predator/scavenger, Odontaster validus (Koehler 1912). Seastars were collected monthly from a shallow site (15–20 m depth) near the British Antarctic Survey (BAS) Rothera Research Station (Adelaide Island, 67°34′S 68°08′W) from July 1997 to January 2001. Reproductive condition, oocyte size frequencies and spermatogenesis were examined in at least ten seastars each month using histological and image analysis techniques. Gonad indices (GI) and pyloric caeca indices (PI) were also examined in the same samples. Female and male GIs varied seasonally, in parallel with a reduction in the proportion of large oocytes and mature sperm in the gonad in August to mid-October following winter spawning. Despite there being remarkable consistency in the timing of spawning from year to year, differences in the reproductive condition of individuals were apparent. Patterns in the digestive tissues also varied with season, peaking in December and reaching a minimum in February in two of the three study years. This weaker annual pattern may partly reflect the varied diet of this predator/scavenger species, which is not directly dependant on the timing and magnitude of the annual phytoplankton bloom. Pooled oocyte size distributions and residual analysis suggested that oogenesis progressed over 18–24 months, with the largest of the two size classes (maximum diameter = 183 μm) being spawned annually. This pattern of oocyte growth and spawning was previously reported in the early 1960s for an O. validus population from McMurdo Sound, which lies south of Rothera by 10° latitude. The extremely catholic diet of this predator/scavenger suggests the reproductive patterns of the seastar will be less susceptible to changes in food supply compared to polar suspension feeders or deposit feeders. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Demography of <Emphasis Type="Italic">Nebalia</Emphasis> sp. (Crustacea: Leptostraca) determined by carrion bait trapping in Lobster Bay,Cape d‘Aguilar Marine Reserve,Hong Kong

The feeding ecology of leptostracans is not well-known. In the present study, species of Nebalia (Crustacea: Leptostraca) were captured every month for 1 year from Lobster Bay, Hong Kong, using baited traps. Captured individuals fell into a size range of 0.36–3.64 mm and 0.95–5.52 mm in carapace and total lengths, respectively. They were divided into different reproductive stages based on the morphology of the second pair of antennae and thoracopods. Sexual dimorphism becomes apparent between 0.7–1.5 mm (CL) and 1.4–3.2 mm (TL). Most individuals collected were sexually differentiated, but immature (79.7% of the captured individuals), followed by post-ovigerous (11.4%) and ovigerous (8.9%) females. Sexually undifferentiated juveniles were rare (3.9%). The sex ratio of the immature individuals approached 1:1 with a propensity towards male dominance. Not one mature male was captured. Although females with setose thoracopod endopods, indicating that they were ovigerous, were captured frequently, only two were carrying eggs. The two brooding females had clutch sizes of 11 and 17 eggs. Alternate peaks in relative abundance of ovigerous and post-ovigerous females were identified, the latter following the former. Reproductive activities occurred mainly in April–May and October–December, indicating that two cohorts are recruited. Catches using baited traps could be explained by the reproductive season of Nebalia sp. High catches were usually associated with the months of highest reproductive activity, low catches with peaks in post-ovigerous females. It is therefore concluded that not all individuals at various reproductive stages are attracted to carrion and hence adopt a scavenging feeding mode. Catches of Nebalia sp. in Hong Kong possibly reflect their annual cycle of reproductive activity     

Sardine (Sardina pilchardus) spawning seasonality in European waters of the northeast Atlantic

Egg data from ichthyoplankton monitoring sites in the western English Channel (1988–2003) and northern Spain (1990–2000) and macroscopic maturity data from biological samples of purse seine landings in western and southern Iberia (1980–2004) are used to describe the spawning seasonality of sardine (Sardina pilchardus) in European waters of the northeast Atlantic using generalised additive models. The fitted models reveal a double peak in spawning activity during early summer and autumn in the western Channel, a wider spring peak off northern Spain and a broad winter season in the western and southern Iberian Peninsula. At all sites, a high probability of spawning activity was observed over at least 3 months of the year, with the duration of the season increasing with both decreasing latitude and increasing fish size. Off western and southern Iberia there are indications that the spawning season has been of longer duration in recent years for all size classes (reaching in some cases 8 months of the year for large fish). These patterns are in general agreement with existing literature and theoretical expectations of sardine spawning being driven locally by the seasonal cycle of water temperature, assuming preferences for spawning at 14 –15°C and avoidance for temperatures below 12°C and above 16°C. Regional quotient plots indicated that spawning tolerance to higher temperatures increases progressively with decreasing latitude. Despite the weak evidence for geographical differences in temperature tolerance that may have some genetic origin, the degree of spatio-temporal overlap in sardine-spawning activity within Atlantic European waters is unlikely to promote any reproductive isolation in that area.     

Life cycle of <Emphasis Type="Italic">Oithona</Emphasis> <Emphasis Type="Italic">similis</Emphasis> (Copepoda: Cyclopoida) in Kola Bay (Barents Sea)

The annual population dynamics (nauplii, old copepodites CIV–CV and adults) and seasonal variations in reproductive parameters of the cyclopoid copepod Oithona similis were investigated on the basis of the data 1999–2006 in Kola Bay, a large subarctic fjord in the Barents Sea. Population density of O. similis ranged from 110 to 9,630 ind m⁻³ and averaged 1,020 ± 336 ind m⁻³. The relative abundance of adults was high during winter (~60%). At the end of winter (mid-March), the population included a large percentage of later-stage copepodites (stage CIV 23% and stage CV 57%). There were two periods of mass spawning, in late June and September. Autumn and summer generations strongly differed in abundance, average prosome length (PL), clutch size (CS), egg diameter (D), egg production rates (EPR and SEPR) and female secondary production. Average PL decreased with increasing water temperature, while D and CS were strongly correlated with PL but unaffected by temperature. Annual average EPR and SEPR were 0.55 ± 0.18 eggs female⁻¹ day⁻¹ and 0.0011 ± 0.003 day⁻¹, respectively. Female secondary production averaged 0.8 ± 0.3 μg C m⁻³ day⁻¹ (range 0.001–3.58). There were positive relationships between abundance, EPR, SEPR, production and water temperatures. Reproductive parameters appeared to be controlled by hydrological factors and food conditions.     

An adaptive annual rhythm in the sex of first pigeon eggs

When the reproductive value of male and female offspring varies differentially, parents are predicted to adjust the sex ratio of their offspring to maximize their fitness (Trivers and Willard, Science 179:90–92, 1973). Two factors have been repeatedly linked to skews in avian offspring sex ratio. First, laying date can affect offspring sex ratio when the sexes differ in age of first reproduction, such that the more slowly maturing sex is overproduced early in the season. Second, position of the egg in the laying sequence of a clutch may affect sex ratio bias since manipulating the sex of the first eggs may be least costly to the mother. We studied both factors in two non-domesticated pigeon species. Both the Wood pigeon (Columba palumbus) and the Rock pigeon (Columba livia) have long breeding seasons and lay two-egg clutches. In the field, we determined the sex of Wood pigeon nestlings. In Rock pigeons, housed in captivity outdoors, we determined embryo sex after 3 days of incubation. On the basis of their sex-specific age of first reproduction, we predicted that males, maturing at older age than females, should be produced in majority early and females later in the year. This was confirmed for both species. The bias was restricted to first eggs. Rock pigeons produced clutches throughout the year and show that the sex of the first egg followed an annual cycle. To our knowledge, this study presents the first evidence of a full annual rhythm in adaptive sex allocation in birds. We suggest that this reflects an endogenous seasonal program in primary sex ratio controlled by a preovulatory mechanism.     

Influence of environmental factors and female size on reproductive output in an invasive temperate marine gastropod <Emphasis Type="Italic">Rapana</Emphasis><Emphasis Type="Italic">venosa</Emphasis> (Muricidae)

Life history and reproductive strategies influence population dynamics at the inter- and intra-specific level. Environmental conditions suitable for gonad development and spawning, the reproductive range, may be a smaller portion of the broader species distribution. The only known breeding population of veined rapa whelks (Rapana venosa) in North America is in Chesapeake Bay, USA. There is considerable interest in the potential reproductive range of this non-indigenous species given the rapa whelk’s negative impacts on commercial shellfish species in both its native and introduced ranges. Weight-specific reproductive output is described for wild caught Chesapeake Bay rapa whelks maintained in flow-through mesocosms for 2 years. Measured reproductive output within and between egg capsule deposition seasons (years) in relation to water temperature, salinity, daylength, and female size is used to describe the rapa whelk’s reproductive range. Egg capsule production is influenced by seasonal and absolute water temperatures as well as seasonal daylength cycles. Egg capsule deposition by Chesapeake Bay rapa whelks begins at water temperatures of approximately 18°C and continues for 11–15 weeks. Forty to 70% of female whelks deposited egg capsules in most weeks during this season, producing 150–200 egg capsules female⁻¹ week⁻¹. Water temperatures >28°C caused reduced egg capsule production relative to temperatures of 20–25°C. Egg capsule production was positively related to seasonal changes in daylength, and two peaks of egg capsule deposition were observed in the 2001 and 2002 deposition seasons. The combination of declining daylength and higher water temperatures in late summer was associated with the cessation of egg capsule deposition. A lower average weight specific reproductive output in 130–145 mm SL rapa whelks (average 12 ± 1%) than in 90–106 mm SL rapa whelks (average 22 ± 1% of body weight) may reflect a life history that balances the physiological costs of maintaining a large body mass with the production of many planktonic larvae from multiple clutches of egg capsules per breeding season over a 10–15-year lifespan. Estimates of the cumulative day-degree requirements corresponding to the annual initiation of egg capsule deposition were 238 and 236 for 2001 and 2002, respectively. Reproductive output and day-degree requirements for Chesapeake Bay rapa whelks were similar to values calculated from previous studies of native muricids (Eupleura caudata and Urosalpinx cinerea). A latitudinal range of 30–41° (N and S) is predicted as the realized reproductive range for rapa whelk populations on the basis of the day-degree requirements for native whelks and reproductively active invasive rapa whelk populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Changes in metabolic substrates during early development in anchoveta Engraulis ringens (Jenyns 1842) in the Humboldt Current

We assessed the ontogenetic changes in protein content and free amino acids (FAA) in eggs and early larvae of Engraulis ringens (anchoveta) off central Chile on different dates during the spawning season. On all sampling dates, a reduction in embryonic yolk-sac volume, proteins and FAA concentrations occurred during development. Protein electrophoresis (SDS–PAGE) of eggs and larvae showed at least 22 protein bands: 11 were consumed early and not detected after hatching. The proportion of essential FAA (EFAA) was higher than the proportion of non-essential FAA (NEFAA) in early eggs and in 7 day-old larvae (82.5-73% EFAA respectively). During egg development, the FAA pool was dominated by leucine, alanine and lysine, three amino acids contributing 35–44% of the total FAA in eggs. During larval development, histidine was the most abundant FAA. In July, total FAA constituted 13–18% of the egg dry weight. A similar proportion (45–51%) occurred in July between protein plus FAA and total lipids. The differences in egg size during the spawning season along with variability in batch composition suggests that the female spawning condition is a major factor determining egg quality and early offspring success.     

Variation in growth rate and reproductive output in British populations of the dorid nudibranch Adalaria proxima: consequences of restricted larval dispersal?

 Growth and reproduction were compared among six geographically and genetically distinct intertidal populations of the annual, semelparous, dorid nudibranch Adalaria proxima (Alder & Hancock) to evaluate variation in fitness-related traits. The six populations spanned the geographic range in the northern British Isles: NE England (Cowling Scar), E Scotland (Kinkell Braes), NW Scotland (Loch Eriboll), W Scotland (Cuan Ferry), Northern Ireland (Portaferry), and N Wales (Menai Bridge). Nudibranchs from five sites were collected in July to August 1992 as post-metamorphic juveniles and were laboratory-reared under the same conditions of ambient temperature and photoperiod for up to 10 months and the completion of spawning. Individuals from the sixth site were added to the experiment in December 1992. Growth was monitored every 2 weeks, and reproductive performance was expressed as a weight-adjusted dimensionless index (ΣRI) of each individual's spawnings summed over the reproductive period. In general, larger nudibranchs produced larger first spawn masses and more total spawn than did smaller nudibranchs, but these size-related trends were observed only in some populations. The patterns of energy partitioning to spawnings varied significantly among populations, from allocations of a large number of eggs to few spawn masses (Loch Eriboll) to production of many small spawnings over a long spawning period (Portaferry). There was no relationship between maximum body size and the amount of spawn produced after the first spawning, nor to the length of the spawning period or the number of spawn produced. Both Menai Bridge and Kinkell Braes had low mean population ΣRI, reflecting a very poor reproductive performance, given their large maximum (pre-spawning) body sizes. By contrast, the Loch Eriboll, Cuan Ferry, and Portaferry populations all displayed high mean population ΣRI, albeit as a result of differing combinations of numbers and sizes of spawn masses and duration of the spawning period. This high variance of reproductive allocation among populations, and previous evidence of relatively stable among-population differences in allozyme frequencies, adult color, and embryo characteristics suggest very restricted larval transport of lecithotrophic larvae of A. proxima. Received: 10 December 1998 / Accepted: 23 March 2000     

Sex-specific patterns of lifetime reproductive success in single and repeat breeding steelhead trout (Oncorhynchus mykiss)

All other things being equal, the lifetime reproductive success (LRS) of iteroparous and semelparous individuals should scale with the number of breeding seasons. Deviations from this relationship may occur for many reasons, including age- or size-related fecundity or life history trade-offs, which may differ between sexes. We used 19 brood years of DNA parentage analysis in a small (N = 4–143 year⁻¹) wild, unexploited population of steelhead trout (Oncorhynchus mykiss) to compare the LRS of individuals that spawned only once [“one time spawners” (OTS), N = 355 male, 371 female] to those spawning twice [“repeat spawners” (RPS), N = 13 male, 49 female]. Female RPS had nearly twice the LRS of female OTS (1.17 offspring per female vs 0.91 offspring per female), whereas male RPS had nearly three times the LRS of male OTS (1.54 offspring per male vs 0.57 offspring per male). Female RPS produced slightly more adult offspring during their second breeding season than their first (0.78 vs 0.82 offspring per female); however, male RPS produced all of their adult offspring in their second breeding season (0 vs 1.54 offspring per male). The additional growth in body size of males between breeding seasons may give them an advantage in their second breeding season, but the lack of offspring produced in their first season suggests a trade-off between survival and future reproduction that was not expressed in females.     

Multiple spawning events and sexual reproduction in the octocoral Sarcophyton elegans (Cnidaria: Alcyonacea) on Lizard Island, Great Barrier Reef

Sarcophyton elegans is a common symbiotic (zooxanthellate) octocoral species in the shallow waters of the Great Barrier Reef (GBR). Study of a population at Lizard Island (14°40′S, 145°28′E) on the GBR from October 1991 to January 1994 revealed that, as is typical of tropical alcyonarian corals, S. elegans is a gonochoric broadcast spawner with a 1:1 sex ratio. Sexual reproduction was closely correlated with colony size, with first reproduction at 13-cm basal stalk circumference for females and 12 cm for males. Oogenesis took 19–24 months, with a new cycle commencing every year, and spermatogenesis took 10–12 months. The majority of gametes were released during the annual austral mass coral spawning event after the full moon in November, but gametes were also released after the full moon in each month between August and February. All autozooid polyps participated in reproduction, but those at the outer edge of a colony released their gametes first. During subsequent months, the polyps closer to the center of the colony released their gametes. This is a novel strategy of gamete release, reported here for the first time, which accommodates the demands of feeding and reproduction in a different way than other corals where individual polyps have separate feeding or reproductive roles. Colonies upstream in the prevailing current spawned up to 1 month earlier than those downstream and ceased 1 month earlier. The mechanism controlling this spatial differentiation in spawning time, repeatedly observed over three seasons, is unknown. Sarcophyton elegans appears to have a dual strategy of providing protection for its gametes by releasing most of them concurrently with the single, annual mass spawning of a large number of cnidarians, while also hedging its bets by individual colonies spawning a fraction of their gametes over an extended period of 6 months.     

Reproductive investment reduces recuperation from exhaustive escape activity in the tropical scallop Euvola zizac

In scallops, mobilization of reserves from the adductor muscle to support maintenance and reproductive activity may impinge upon a major role of the adductor muscle, the movement of the valves during swimming and escape responses. The tropical scallop Euvola ziczac (Linnaeus 1758) invests energy reserves to different degrees during its two periods of reproduction each year. We evaluated the impact of reproductive investment on the escape responses (clapping capacity and recovery after exhaustive exercise) of E. ziczac sampled at different reproductive stages (immature, mature, spawned) during the two reproductive periods in 1997. We compared the escape response capacities with the levels of muscle energetic reserves (glycogen, proteins, and arginine phosphate) and muscle metabolic capacities [activity of the glycolytic enzymes: glycogen phosphorylase (GP), pyruvate kinase (PK), phosphofructokinase (PFK), octopine dehydrogenase (ODH), arginine kinase (AK), and the mitochondrial enzyme, citrate synthase (CS)]. Gonad size, gamete volume fraction, and levels of gonadal protein and lipid were greater for mature scallops during the first than during the second reproduction. Numbers of claps during escape responses (49–57) and levels of muscle arginine phosphate remained similar throughout the different reproductive stages in both reproductive periods. In contrast, recovery was slowed during gonadal maturation in both reproductive periods and during spawning in the first reproduction. Scallops generally took more time to regain their initial clapping capacity during the first (25–40 min) than during the second reproduction (20–30 min). Muscle glycogen decreased markedly during both gonadal maturation and spawning in both reproductions; whereas, muscle proteins decreased only in the first reproduction. The levels of most enzymes decreased during gonadal maturation in both reproductions, and also after spawning, particularly during the first reproduction. We concluded that gonadal maturation and spawning did not decrease clapping capacity of E. ziczac, but decreased its capacity to recover from exhaustive exercise most likely due to decreased levels of energetic reserves and a reduced metabolic capacity of the adductor muscle. Moreover, these effects were probably stronger during the first cycle because of the greater reproductive investment coincident with decreased food availability. Received: 28 April 2000 / Accepted: 21 September 2000     

Reproductive biology of the deep-sea octocoral Drifa glomerata in the Northwest Atlantic

The present study examined the mode and timing of reproduction of poorly understood deep-water octocorals and the environmental factors that may influence their reproductive patterns. Data on reproductive characteristics of the octocoral Drifa glomerata (Alcyonacea: Nephtheidae) collected between 2004 and 2007 at ca. 100–330 m depth off Newfoundland and Labrador (eastern Canada) were compared among years, months and depth ranges. No male gonad was ever observed during the study. The ratio of fertile colonies possessing large pinkish polyps with oocytes/planulae was >50% throughout the year. The number of brooded planula larvae within a single fertile polyp varied between 1 and 10 for a total of approximately 40–3,000 in a whole colony. The size of oocytes and/or planulae was consistently greater in the polyps than in the branchlets, indicating that the development pathway of oocytes to planulae is from the branchlets to the polyps. Although larval production seemed to persist year round, the onset of major planulation events occurred in December–January of two consecutive years, when large mature planulae were released in correlation with the first increase of photoperiod and maximum temperature at 150 m. A second peak in planulation between April and early June followed the phytoplankton bloom. Seasonal trends were more apparent in colonies from <200 m, and the planula index varied among sampling depths and years. Larval release in a live colony under laboratory conditions occurred between January and June 2008, closely following predictions based on field samples.     

Production, hatching success and surface ornamentation of eggs of calanoid copepods during a winter at 57°N

Close to 50 species of marine Calanoid copepods have been reported to produce diapause eggs (Engel and Hirche in J Plankton Res 26:1083–1093, 2004); eggs that are viable but require a refractory phase before they hatch, sometimes after months. Diapause eggs are often described as morphologically different with respect to egg membrane ultrastructure and having a thicker egg shell with surface ornamentation as opposed to the smooth shell found in subitaneous eggs that hatch within days (Belmonte in J Mar Syst 15:35–39, 1998; Chen and Marcus in Mar Biol 127:587–597, 1997; Castro-Longoria in Crustaceana 74:225–236, 2001). Egg production rates, egg surface ornamentation, and hatching success were monitored in large aquaculture fish enclosures during winter with close to zero water temperatures (N57°). Surprisingly, all female copepods (Acartia spp.—presumably A. tonsa, and Centropages hamatus) produced eggs all through the winter with no obvious pattern with respect to light, temperature and food availability, and no diapause eggs were observed. However, individual females produced several categories of eggs with or without surface spines even within the same egg batch as evidenced by scanning electron microscopy (SEM). Four egg categories were distinguishable: ‘no spines’, smooth eggs; ‘short spines’, 5–15 μm long; ‘truncated spines’, with the spine tips cut-off <10 μm long; and ‘long spines’, up to 30 μm long. All egg categories remained unchanged with respect to surface structures from when we took them out of the incubation bottles until they hatched. In general, the frequency of ‘no spines’ was 10–40%, and most eggs were ornamented with ‘short-’ or ‘long spines’. Further, a given egg can be ornamented with all types of surface spines simultaneously, which might even be a fifth egg category. The different egg categories were all able to hatch within days when exposed to normoxic conditions suggesting that they were subitaneous.     

Growth and reproduction in the Antarctic brooding bivalve Adacnarca nitens (Philobryidae) from the Ross Sea

We present information on the reproductive biology, population structure, and growth of the brooding Antarctic bivalve Adacnarca nitens Pelseneer 1903, from the Ross Sea, Antarctica. Individuals ranging from 0.85 to 6.00 mm were found attached to a hydrozoan colony. This species shows low fecundity and large egg size, common to other brooding species. The minimum size at which oogenesis was detected was 2.3 mm and the minimum size at which brooding was evident was 3.9 mm. Embryos of a full range of developmental stages were brooded simultaneously in females. The population showed a log–normal distribution and results suggest non-periodic reproduction with continuous embryonic development. The reproductive traits of A. nitens are discussed in the context of circum-Antarctic species distribution and limitations to dispersal in brooding benthic invertebrates.     

Reef coral reproduction in the Abrolhos Reef Complex, Brazil: the endemic genus Mussismilia

The current contribution deals with the reproductive biology of a genus endemic to Brazil –Mussismilia Ortmann, 1890 – including all three species of the genus: M. braziliensis (Verrill, 1868), M. hartti (Verrill, 1868), and M. hispida (Verrill, 1902), which occur sympatrically in the studied area, the Abrolhos Reef Complex, Brazil. Sexuality patterns, modes of reproduction, synchrony and spawning periods are reported, and were determined by histological examination of material. All three species started to develop female and male gametes over different periods in the same breeding season. The three species are probably broadcast spawners, since no embryos or planulae were observed in any species at any given time of the year. Each reproductive cycle lasted approximately 11 months. Oogenesis and spermatogenesis started in different periods, with spermaries appearing in approximately the eighth month of ovary development and lasting about 3 months. Reproductive cycles were annual. Spawning probably occurred in consecutive months in each species. In M. braziliensis, spawning presumably happened between March and the middle of May in 1996 and 1997. Evidence suggested that spawning events of M. hispida took place between the end of April and mid-June. M. hartti may have spawned between September and November. The data presented here suggested that all studied species have at least one exclusive spawning period, asynchronically with the others. A possible exception may be the simultaneous (or close) spawnings of M. braziliensis and M. hispida in May. It is suggested that asynchrony in spawning periods among species may reduce the chance of hybridization, gamete waste and the competition for settlement surfaces. The occurrence of extended spawning periods for each species may also reduce the risks of reproductive failure, due to temporary adverse conditions. Received: 8 December 1998 / Accepted: 15 July 1999