Size at maturity,fecundity and reproductive potential of a protected population of the spiny lobster<Emphasis Type="Italic"> Palinurus elephas</Emphasis> (Fabricius, 1787) from the western Mediterranean

Protected lobster populations are expected to contribute to the replenishment of fished populations through increased egg production. We studied the reproductive biology and egg production potential of a population of the spiny lobster Palinurus elephas protected from fishing since 1990 in the Columbretes Islands Marine Reserve (western Mediterranean). An index of spawning potential was derived to compare egg production potential in the Reserve and in western Mediterranean exploited populations. Females' physiological maturity (ability to reproduce) and functional maturity (ability to mate and bear eggs) occurred at a carapace length (CL) of 76–77 mm. Males' physiological maturity occurred at a slightly larger size, 82.5 mm CL. In the Reserve, P. elephas' individual fecundity increases linearly with body size up to the females' maximum size, although maximum reproductive yield (eggs per body gram) was reached at intermediate sizes. Size-specific fecundity in the protected population was similar to that of lightly fished populations off Ireland and greater than that of western Mediterranean exploited populations. The female size class of 105–110 mm CL contributed most to egg production in the protected population and is well above the minimum landing size (MLS) for western Mediterranean fisheries. Newly mature females (below MLS) generate a very small fraction (1%) of the egg production from the Reserve. Given the pattern of exploitation in western Mediterranean fisheries, egg production potential depends more on the quantity than on the mean size and fecundity of the available females. The role of the greater availability of large males for mating in unfished populations is discussed in terms of the females' individual fecundity and mating success.     

Evidence for multiple spawning in the tropical oceanic squid Stenoteuthis oualaniensis (Teuthoidea: Ommastrephidae)

Indirect evidence suggests that Stenoteuthis oualaniensis is a multiple spawner, a reproductive strategy not reported previously for squid. After spawning once, S. oualaniensis apparently continues to feed, grow, and mature additional oocytes before spawning again. Squid were caught by jigging at night in waters around the main Hawaiian Islands during four cruises from 1981 to 1987. All females from 200 mm mantle length (ML) to at least 318 mm ML were sexually mature, indicating that considerable growth takes place after reaching sexual maturity. Size-frequency analyses of ovarian oocytes revealed continuous egg production throughout adult life. Mature ova are stored in the oviducts, but oviduct fullness is unrelated to body size, indicating repeat spawning.     

Movements of lobsters (Homarus americanus) tagged in the Bay of Fundy,Canada

Of the 18 359 American lobsters (Homarus americanus) tagged and released at three locations in the Bay of Fundy during 1977–1980, 5 375 lobsters (29.3%) were recaptured within 6 yr of release. Mature lobsters (95-mm carapace length, CL) on average moved significantly greater distances than immature lobsters (<95-mm CL). Many (16.4%) of the mature lobsters were recaptured >92.6 km (50 nautical miles) from the release sites; the farthest distance moved was 798 km for a male at liberty for 3.5 yr. Examination of the direction of movements indicates some intermixing of lobsters within the Bay of Fundy, and throughout the Gulf of Maine and the adjoining Continental Shelf and slopes. Mature lobsters were recaptured in deeper waters during winter than during summer. The seasonal shallow-deep migration of mature lobsters in the Bay of Fundy is possibly associated with maximizing degree-days for molting, growth, gonadal development, and egg development. In some areas, mature females on average moved farther and seasonally earlier into deeper waters than mature males. Although the seasonal migration resulted in many lobsters returning to the original release area year after year, some lobsters made extensive long-distance movements away from the Bay of Fundy. Temperature-dependent, seasonal deep-shallow migration can explain both the local returns and the long distance migrations of mature lobsters in the Bay of Fundy and the Gulf of Maine. To achieve appropriate seasonal temperature regimes, mature lobsters have to move different distances and directions, depending on local topography.     

Laboratory observations on spawning,brood size and egg hatchability of the Antarctic krill Euphausia superba from Prydz Bay,Antarctica

Forty-three female Euphausia superba were collected from three sites in Prydz Bay, Antarctica in January 1985. Most gravid females took 3 to 14 d to spawn after capture and some less mature females released eggs after 21 d. Most females spawned only once; overall mean brood size was 2 809 eggs and mean hatchability for eggs from the premoult females was 69.7%. For seven females spawning a second time, overall mean brood size was 597 eggs. Mean hatchability for second-spawn eggs from five premoult females was 67.1%. Possible origins of the second-spawned eggs were small numbers of matured eggs left in the ovaries after the first spawning or eggs developed from those of peripheral nucleolus and vitellogenic stages. Throughout the 8 wk of observations, there was no apparent development of eggs in females which had released all their eggs at one spawning.     

Spatial and temporal variation in the size at maturity of the western rock lobster Panulirus cygnus George

The sizes at which female and male western rock lobster Panulirus cygnus become mature were examined over 32 years from records at six localities along the coast of Western Australia. The size of males at maturity was estimated from a subset of these data by a morphometric and a physiological method, with both producing statistically similar results. Males were larger at first maturity than females at the same sites. For both sexes, the sizes at first maturity at each location correlated (P<0.05) with the mean annual water temperature at that location, decreasing from south to north along the mainland coast and being smallest at the offshore Abrolhos Islands. Smaller sizes at maturity were recorded for both sexes than have been published previously. One certain explanation for these differences is that management measures protecting females with ovigerous setae have distorted size compositions and the ratio of immature to mature females, thereby increasing the likelihood of capturing small mature females. However, these fishing effects cannot fully account for the progressive decline in CL₅₀ observed over the past 20 years. Other possible hypotheses considered include increases in water temperature over this period, as well as whether this change could be consistent with a genotypic response caused by the selective removal of large lobsters combined with high exploitation rates.     

Filial cannibalism in a nest-guarding fish: females prefer to spawn in nests with few eggs over many

In fish, fecundity correlates with female body size and egg-tending males often eat small broods. Therefore, small females may prefer to spawn in nests that already contain many eggs, to ensure the brood is as large as possible. In contrast, large females may prefer nests with few eggs, if high egg number or density has a negative effect on egg survival, or if there are drawbacks of spawning last in a nest. To test the hypothesis that female body size affects nest (and male mate) choice, using the sand goby (Pomatoschistus minutus), we allowed small and large females to choose between two males that were matched in size — one guarding a small clutch and the other a large clutch, respectively. We recorded where females spawned (measure of female preference), the combined brood size, male courtship, egg care and nest building. We also quantified the effect of brood size and egg density on egg survival in a separate data set. Although the combined broods did not exceed the small brood sizes that are at risk of being eaten, both small and large females preferred to spawn in nests with smaller clutch sizes. This preference could not be explained by more courtship or male parental effort, nor by reduced survival of larger or denser broods. Instead, our result might be explained by females avoiding the danger of cannibalism of young eggs by males or the risk of reduced egg health associated with being near the nest periphery.     

Life-history traits of Plesionika martia (Decapoda: Caridea) from the eastern-central Mediterranean Sea

Life-history traits of Plesionika martia (Milne Edwards, 1883) were studied through data collected during six seasonal trawl surveys carried out in the Ionian Sea (eastern-central Mediterranean) between July 1997 and September 1998. P. martia was found at between 304 and 676 m depth, with the highest density in the 400-600 m range. Intraspecific, size-related depth segregation was shown. Recruitment occurred in summer at the shallowest depths. Juveniles moved to the deepest grounds as they grew. The largest female and male were 26 and 25 mm carapace length, respectively. The sex ratio was slightly in favour of females at depths >400 m. Although a seasonal spawning peak was shown, the reproduction appears to be rather prolonged throughout the year. Females with ripe gonads were found from spring to autumn. Ovigerous females with eggs in late maturity stage were found year round. Large females could spawn more than one time within their annual reproductive cycle. The size at first maturity (50% of the ovigerous females) was 15.5 mm CL. Average brood size of eggs with a well-developed embryo was 2,966ǃ,521. Iteroparity, low fecundity and large egg size patterns were observed. Brood size increased according to the carapace length. Two main annual groups were found in the field population of the Ionian Sea. Estimates of the Von Bertalanffy growth parameters are: L_X=30.5 mm, k=0.44 year^-1 in females; L_X=28.0 mm, k=0.50 year^-1 in males. A negative allometry was detected mostly in the ovigerous females. The life cycle of P. martia is discussed in the light of life-history adaptations shown in other deep-water shrimp species.     

Importance of rock lobster size–structure for trophic interactions: choice of soft-sediment bivalve prey

Ecologists are becoming increasingly interested in how variation in predator demographics influences prey communities. In northeastern New Zealand, the contrasting populations of previously exploited predators in highly protected marine reserves and fished areas have been used to investigate the effects of predation in soft-sediment habitats. However, these experiments have been unable to separate the role of predator size from that of density. This study provides evidence to support the model that foraging by different sizes of the rock lobster Jasus edwardsii affects soft-sediment bivalve populations in different ways. Feeding trials were conducted to investigate whether rock lobsters of different sizes vary in their choice of taxa and size of their bivalve prey. Trials with two morphologically similar species, Dosinia subrosea and Dosinia anus, indicated that lobsters of all sizes choose D. subrosea more frequently than the heavier shelled D. anus. Further results indicated that both large (>130 mm carapace length (CL)) and small (<100 mm CL) lobsters are capable of preying on a wide size range of D. subrosea (20–60 mm). However, small lobsters more frequently chose smaller shells (<30 mm) and large lobsters more frequently chose larger shells (>40 mm). Patterns in the abundance and size class distributions of these two bivalve species at protected and fished sites supported the feeding choices observed in the laboratory. These results suggest that populations of rock lobsters with large individuals inside reserves are capable of controlling the demography of bivalve populations in adjacent soft-sediment systems.     

Size-related reproductive biology of the infaunal bivalve<Emphasis Type="Italic"> Abra alba</Emphasis> (Bivalvia) in Kinsale Harbour (south coast of Ireland)

The reproductive behaviour of 248 specimens comprising four size classes of the infaunal bivalve Abra alba (Wood) was investigated between September 1994 and November 1995 in Kinsale Harbour. Small males returned a very small proportion of sexually mature and spawning animals. By contrast, no spawning females were found in this size class, suggesting that males develop and mature at a smaller size than females. Mean gonad maturity indices indicated that, except for the smallest size class, A. alba exhibits two spawning events in 1 year, i.e. a lesser spring and a major autumn spawning event. The spring event was well synchronised between males and females, while the autumn spawning was more broadly synchronised between the sexes. Only one spawning event was recorded for very large males and females, also occurring in spring. Spawning stages were recorded for both sexes over all size classes between February and November, indicating only a short period of sexual repose, if any. Oocyte diameters did not show a discrete size range from developing to spawning stages. All stages were seen to contain varying proportions of smaller, less mature oocytes, suggesting that large A. alba females do not develop and spawn en bloc, but release their gametes periodically. Maturity indices showed only a broad similarity with off-bottom temperatures.Communicated by O. Kinne, Oldendorf/Luhe     

Circadian shelter occupancy patterns and predator–prey interactions of juvenile Caribbean spiny lobsters in a reef lagoon

The spiny lobster, Panulirus argus, is predominantly nocturnal, remaining inside shelters during the day and foraging outside at night, presumably to minimize predation risk. Predation risk generally decreases with increasing lobster size. Therefore, this study examined the hypothesis that size would influence this basic circadian pattern. Video cameras continuously recorded the shelter occupancy of juvenile lobsters (n = 72) having a carapace length (CL) of 30–62 mm that were tethered to shelters in a shallow reef lagoon. The lobsters’ shelter occupancy was 100% during the day, but declined linearly from shortly before sunset to a minimum of 50% shortly after midnight and then increased linearly, reaching 100% by 1 h after sunrise. The percent time the lobsters spent in the shelters followed a similar trend, but there was wide variability at night (0–100%) for individual lobsters. Lobsters left their shelters 2–30 times night⁻¹, with a majority of excursions lasting <10 min. These results suggest that juvenile P. argus minimize predation risk by remaining in their shelters as long as possible but offset the energetic cost of this behavior by foraging close to their shelters for several short periods at night. This emergence pattern contrasts with those of early benthic phase lobsters (<15 mm CL), which seldom leave their shelters, and adults (>80 mm CL), which have a dusk/early evening peak in activity and leave the shelter for extended periods of time during the night. Furthermore, a minimum shelter occupancy in the middle of the night appears especially well adapted to avoid exposure to daytime predators. Videotaped observations also included interactions between lobsters and two dominant lobster predators, the triggerfish, Balistes capriscus, and the octopus Octopus cf. vulgaris. Lobsters responded differently to these predators: remaining in the shelter when attacked by a triggerfish and fleeing the shelter when attacked by an octopus. Triggerfish were nearly twice as likely to attack a lobster that was outside of the shelter than inside. Once under attack, however, a lobster had nearly the same chance of surviving if it was inside or outside. Results suggest that the patterns of shelter use and emergence change as lobsters grow, probably reflecting the interplay between perception of predation risk and the need to forage. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Offshore spawning by the portunid crabScylla serrata (Crustacea: Decapoda)

The portunid crabScylla serrata (Forskål) is shown to migrate offshore to spawn. Records of 447S. serrata caught as bycatch by trawlers in the tropical waters of northern Australia were analysed with respect to area, depth, distance offshore and month of capture as well as the sex and size of the crabs and whether the females were ovigerous. The crabs were caught mainly in three areas that correspond to the tiger prawn trawl fishery, at between 10 and 60 m depth (mean 28.5 m), 3 to 95 km offshore (mean 17.9 km). Most (87%) of the crabs were captured in October and November, which suggests they move offshore in September and October. No crabs were reported from offshore by February. Over 97% of the crabs caught offshore were female, of these 61.5% were ovigerous. The size range of females (100–109 to 200–209 mm carapace-width size classes) and males (120–129 to 200–209 mm carapace-width size classes) caught was similar. The frequent occurrence in coastal waters of females that are larger than the modal size at spawning, as well as mature females with spent ovaries, suggests that many females return to the coast after spawning. Although some species of portunid crab are euryhaline, mature females of estuarine species migrate to the sea to spawn. The migration byS. serrata described here is far more extensive than would be required to reach sea water salinities; it probably provides a dispersal mechanism for larvae to enable the megalopa stage to recruit to habitats distant from those of the parents.     

Reproductive biology of the lanternfish Benthosema pterotum from the Indian Ocean

Samples of Benthosema pterotum (Alcock) were obtained from the Arabian Sea (1976, 1979, 1981, 1983, 1984), off Mozambique (1978) and from the Bay of Bengal (1979). The sexual maturity stages are described. For females, the size of the largest oocytes was found to be most useful for size of the largest oocytes was found to be most useful for assessing maturity. Mature and spawning fish were found in all seasons. Samples from November 1983 differed from all others in having very few mature females. A few apparently spent fish were observed in July to August 1979 and November 1983. Some females mature at 25 mm and males at 20 mm, whereas others can still be immature at 45 and 40 mm, respectively. Of the total number of stations, four had significantly more males than females, twelve significantly more females, and forty-seven displayed no difference in the sex ratios. The vertical migration of the mature and spawning fish did not differ significantly from that of other individuals. Batch fecundity ranged from ca. 200 to 3 000 eggs fish ranging from 27 to 52 mm, relative fecundity from ca. 2 000 to 7 000 eggs g^-1 maternal dry weight. There are indications that B. pterotum spawns only once, but no firm conclusions could be drawn.     

Long-term experiments on lifespan, reproductive activity and timing of reproduction in the Arctic copepod Calanus hyperboreus

Several experiments were conducted with starved and fed females of the Arctic copepod Calanus hyperboreus (1) to investigate their lifespan, reproductive period, egg production and egg viability; (2) to study the effect of origin, i.e., Atlantic and Arctic Waters in the Greenland Sea, on the timing of reproduction; and (3) to study the effect of time of collection on the onset of reproductive activity as a first approach to study control mechanisms of the reproductive cycle. Females collected in October produced up to 1,000 eggs and had a maximum lifespan of 164 days without feeding, whereas fed females produced up to ca. 6,000 eggs and survived up to 806 days. These observations support earlier assumptions that females were multiannual-iteroparous, i.e., capable to spawn in successive years, which would be unique for calanoid copepods. In starved females, clutch size decreased significantly with each spawning event. Viable eggs were produced during most of the life time. There was no difference in the timing of reproductive activity between females from the West Spitsbergen Current and the Greenland Sea Gyre. Fed and starved females collected in May and June began to spawn circa 2 and 4 months after collection, respectively, whereas females collected in August and October started spawning at the same time, in the middle of October. This indicates initiation of reproductive activity in the field in August, coincident with the descent into deep waters. Potential cues for the untimely spawning of females collected in spring and ‘unnatural’ feeding in fall experiments are discussed. Their large size, robustness and combination of different types of diapause in their life cycle make C. hyperboreus a good model organism to study diapause control mechanisms.     

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.     

Developmental types of shallow-water asteroids of McMurdo Sound,Antarctica

The mode of development was ascertained for 14 of the 16 species of sea stars known to occur in shallow waters of McMurdo Sound, Antarctica (77°51S; 166°40E). The species were collected between September 1984 and December 1985. Females of three species,Odontaster validus, O. meridionalis andPorania antarctica, spawn small to moderate eggs (0.17 to 0.55 mm), have a high fecundity, and produce feeding larvae. Females of an undescribedPorania species spawn a few eggs (150 to 310) that are 0.55 mm in diameter and develop into demersal non-feeding larvae. Females ofDiplasterias brucei andNotasterias armata produce a few (<300) large eggs (2.8 to 3.5 mm) and brood their young. Females of the remaining eight species have moderate fecundity and produce pelagic non-feeding larvae, as determined from egg type (buoyant, 0.54 to 1.28 mm diam) and direct observations of spawning and development. The high incidence (11 out of 14 species; 79%) of non-feeding development is consistent with predictions that environmental conditions in high-latitude regions are unfavorable for planktotrophic development. Nonetheless, most of the species surveyed (11 out of 14) had pelagic larvae, which contradicts inferences of unusual selection for benthic development in the Antarctic.     

Life under Arctic sea ice – reproduction strategies of two sympagic (ice-associated) amphipod species, Gammarus wilkitzkii and Apherusa glacialis

The reproductive strategies of two gammaridean amphipod species, Gammarus wilkitzkii and Apherusa glacialis, that permanently inhabit the Arctic sea ice were investigated. G. wilkitzkii reaches sexual maturity at an age of 2 years and produces 128 ± 54 eggs fem.⁻¹ yr⁻¹. Mating takes place during fall and winter, and the development of the large eggs (0.60 to 0.80 mm diam.) lasts 6 to 7 months. The sex ratio of G. wilkitzkii was dominated by males in a proportion of 1.5:1. In vivo studies showed that juveniles are released in batches from the brood pouches of the females during April and May. A. glacialis reaches sexual maturity at the age of 1 year and produces 555 ± 151 eggs fem.⁻¹ yr⁻¹. The eggs are between 0.18 and 0.23 mm in diameter, and are the smallest known for gammaridean amphipods. Eggs are kept in packages of two to eight in the brood pouches of females. The sex ratio of A. glacialis was dominated by females in a proportion of 3:1. The high fecundity of both amphipod species, the release of juveniles in batches over a period of time, a high proportion of females (A. glacialis), and an elongated life-span with multiple spawnings (G. wilkitzkii) are discussed as possible adaptations to the specific and highly variable conditions under Arctic sea ice. Received: 29 December 1999 / Accepted: 8 March 2000     

Mating patterns and a test for female mate choice in Etheostoma spectabile (Pisces,Percidae)

Neither size nor breeding color correlated with spawning success of male orangethroat darters, Etheostoma spectabile (Pisces, Percidae), under natural field conditions. When females were presented experimentally with a simultaneous choice they spent no more time in proximity to large than small males, and were subsequently no more likely to spawn with large than with small males. Females also displayed no preference for bright versus dull males. Males and females did not differ significantly in size. Etheostoma spectabile may lack sexual size dimorphism as a result of the lack of female choice for size and the ineffectiveness of male attempts to monopolize females, or selection may be for increased size of females. Males are not dwarfs because of sperm competition. Contest competition among males appeared to be important in initiating spawnings but many males obtained spawnings by participating in ongoing spawning events. Etheostoma spectabile is an example of a sexually dimorphic species with no evident female preference for male size or color.     

Population structure and growth in captivity of the spiny lobsterPanulirus penicillatus from Dahab,Gulf of Aqaba,Red Sea

Panulirus penicillatus (Olivier, 1791) (Decapoda: Palinuridae) is the most common spiny lobster in the Red Sea and is widely distributed in the Indian and Pacific Oceans. Lobsters (n=377) were collected on four occasions during 1986 on the coral reef at Dahab, Sinai, Egypt. Average size of the collected individuals was 70.5±24.6 mm carapace length (CL) for males and 63.2±15.9 mm CL for females. The sex ratio was 1:1.64 males to females. Length increment per molt was inversely correlated with size and ranged from 2.1 mm per molt in the 40 to 50 mm CL size class to less than 1 mm in the 60 to 70 mm CL size class. Average intermolt period was ca. 136 d for all size classes. The relationship between carapace length and body weight was expressed by the equation:W _b=6.43 × 10^–4 × (CL)^2.89.P. penicillatus from Dahab differ in size, sex ratio and growth rate compared to other palinurid populations throughout their range. This might represent the effect of isolation and location at the edge of the geographical range for this species. It may also indicate an adaptation to their unique habitat in the coral reef in comparison to other palinurid species.     

Eggshell spotting in brood parasitic shiny cowbirds (<Emphasis Type="Italic">Molothrus bonariensis</Emphasis>) is not linked to the female sex chromosome

For avian brood parasites in which individual females are host-specialists, the arms race between hosts and parasites has favored egg color polymorphism in the parasite, with female lineages laying mimetic eggs that resemble those of the host species they parasitize. Female sex-linked inheritance of egg color fosters evolutionary stability of egg polymorphism if female lineages show both consistent eggshell color and host use. This co-evolutionary relationship is unlikely to occur if individual brood parasites use different hosts or if egg color is not maternally inherited. The shiny cowbird (Molothrus bonariensis) is an extreme generalist brood parasite that shows a very high degree of egg polymorphism. We tested whether egg spotting in this species has female sex-linked inheritance. If genetic factors controlling the expression of egg spotting were present on the female-specific W chromosome, we expected co-segregation between spotting patterns and mtDNA haplotypes, as both W and mtDNA are maternally inherited. In contrast to the known maternal inheritance of spotting patterns in great tits, we found no associations between eggshell spotting and mtDNA haplotypes, which suggests that eggshell spotting is not maternally inherited in this cowbird species.