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.     

Sexual regression,shrinkage, re-maturation and growth of spent female Euphausia superba in the laboratory

Live female Euphausia superba, Dana were collected from Prydz Bay, East Antarctica in January 1985, and observations of intermoult period, growth and maturation changes were made on individuals which spawned and were subsequently maintained in the laboratory for 12 mo. Over this experimental period an instrinsic cyclic reproductive pattern was observed. Following spawning, there was a regression of the female copulatory organ, the thelycum, coupled with a decrease in body length and an increase in the krill's mean intermoult period. This was followed by a period of sexual re-maturation, associated with positive growth and a decrease in the mean intermoult period. This study indicates multi-year spawning of E. superba and a life-span greater than 4 yr.     

Pair formation,pre-spawning waiting,and protandry in kokanee,<Emphasis Type="Italic"> Oncorhynchus nerka</Emphasis>

The timing of arrival to breeding areas can have profound effects on reproductive success. Under some conditions (restricted breeding seasons and mating systems characterized by a longer period of mating among males than females), the maximization of mating opportunities by males theoretically selects for the earlier arrival of males than females (a phenomenon called protandry). This study quantifies the relationship between the arrival timing and spawning success of male kokanee (non-anadromous Oncorhynchus nerka). The spawning behavior of kokanee was observed in a large pen and the spawning success of each male was estimated as the number of spawning events he participated in. A male's spawning success depended primarily on his success at pairing with and mate-guarding females, and less on participation in spawning events while unpaired. Males who paired earlier in the season had higher spawning success than males who paired later in the season because they experienced more opportunities to pair with new females. Among males who eventually paired (some males never did), arriving early was correlated with pairing early. However, selection for protandry was weak, largely because early arrival did not guarantee that a male would pair. Pre-spawning waiting by females also weakened the correlation between arrival day and pairing day. The random probability of pairing with respect to arrival day and pre-spawning waiting by females likely explains the weak selection for protandry in kokanee and the low amount of protandry observed in other sockeye salmon (anadromous O. nerka) populations.     

The fatty acids of antarctic phytoplankton and euphausiids. Fatty acid exchange among trophic levels of the Ross Sea

The fatty acids of 3 samples of Euphausia superba, 7 samples of E. crystallorophias, and 12 samples of phytoplankton collected in the Ross Sea, Antarctica, during Eltanin Cruise 51 were examined. The fatty acid profiles of the samples of E. superba resembled each other closely. The fatty acid profiles of the E. crystallorophias samples were also similar to each other but different quantitatively from those of E. superba. Phytoplankton fatty acid patterns varied with the geographical location and species composition of the samples. The fatty acids of euphausiids were compared to those of the phytoplankton from the corresponding locations. Rather similar fatty acid patterns in phytoplankton and E. superba corroborate the herbivorous nature of this euphausiid. On the other hand, phytoplankton and E. crystallorophias showed quite different fatty acid patterns. The differences were mostly due to the presence of waxes among the lipids of E. crystallorophias. It is not clear whether these waxes are of dietary origin or are synthesized endogenously.     

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.     

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.     

Spawning migration and returning behavior of white-streaked grouper Epinephelus ongus determined by acoustic telemetry

White-streaked grouper (Epinephelus ongus) is an important fishery target that forms fish spawning aggregations (FSAs) at specific spawning grounds. The aims of the present study were to investigate migration behavior and returning behavior in terms of FSA using acoustic telemetry. Among the 30 specimens tracked, three types of movement pattern were observed: type 1, high site fidelity at the release area before the spawning period, migration to the spawning ground during the spawning period and return to the release area just after the spawning period (17 specimens); type 2, high site fidelity at the release area, disappearance from the area during the spawning period but was not detected at the spawning ground and return to the release area (8 specimens); and type 3, high site fidelity at the release area was observed before, during and after the spawning period (5 specimens). For the 17 specimens that were categorized into type 1, the dates of departure from their release area and the dates of return to the release area of males were, respectively, significantly earlier and later than those of females. The duration between the departure date and the return date (“duration away”) of males was significantly longer than that of females. The duration away tended to be positively related to the condition factor. All 25 specimens that were categorized as type 1 and type 2 showed returning behaviors to the release area.     

Does the spawning of Calanus finmarchicus in high latitudes follow a reproducible pattern?

In the spring of 1989, an experimental study of the spawning behaviour of Calanus finmarchicus was carried out in Malangen, northern Norway. Here, a single cohort of females reproduce from mid-March to May, approximately coinciding with the wax and wane of the spring phytoplankton bloom. An evaluation of population characteristics such as the proportion of adults, sex ratio, as well as gonad maturation and daily productivity of the females clearly reveals three phases within the population's reproductive period. In between incline and decline, the highest spawning rates (on average >20 eggs female^-1 d^-1, equivalent to 5.7% body C d^-1) occur after the males have disappeared from the population and almost all females have mature gonads. During this period, the ratio of adults to copepodid Stage Vs changes from dominance of adults to that of CVs. Although first egg production was observed prior to the phytoplankton increase, it is suggested that the onset of the phytoplankton spring bloom in the first few days of April enhances the final maturation of ovaries in the females and therefore triggers the onset of the main spawning period. The clutch sizes (max. 95 eggs clutch^-1) vary with the age of the females, while the spawning frequencies depend on the available food quantities. The overlap of an estimated minimal 4 wk spawning period for the individuals leads to a main reproductive phase for the population of ca. 3 wk, during which time mean clutch sizes and spawning frequencies are maximal (highest average clutch size: 70 eggs female^-1 clutch^-1, 100 to 60% of the females spawning). This period ends before the end of the phytoplankton bloom. Calculated by stepwise interpolation and summation of the mean daily egg production in the population, an average female produced ca. 600 eggs during the spring bloom in Malangen 1989. We suggest that reproduction and population development of C. finmarchicus in spring follows a reproducible pattern for a given temperature regime and non-limiting food conditions. In the case of clearly identifiable cohorts, it seems possible to trace the state of reproduction by evaluating population parameters.     

Do cod form spawning leks? Evidence from a Newfoundland spawning ground

Catch and acoustic telemetry data were used to test three predictions of lekking behaviour in Atlantic cod (Gadus morhua L.) at a small-scale spawning ground (∼25 km²) in Placentia Bay, Newfoundland: (1) that cod form male-skewed aggregations during spawning; (2) spawning arenas occur at consistent sites; and (3) on the arenas, there will be higher proportions of spawning fish. Forty-two fishing sets from 1998 to 2003 indicated male-skewed sets early in the spawning season (April) in 5 of 6 survey years. Male-skewed sets were consistently distributed at depths < 50 m and mostly in one part of the ground. The proportions of spawning females and spent males were significantly higher in male-skewed sets, whereas sets with equal sex ratios had significantly higher proportions of immature males and spent females. In addition, cod of both sexes were significantly larger in male-skewed sets. Telemetric tracking of 25 cod in 2002 and 2003 (12 males, 13 females) indicated that both males and females were highly mobile while present on the spawning ground. Overall, our results add support to the hypothesis that cod employ a lekking mating system.     

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     

Length-weight relationships and coefficient of condition of Euphausia superba and Thysanoessa macrura (Crustacea: Euphausiacea) in southwest Indian Ocean during summer

Samples of antarctic euphausiids, Euphausia superba and Thysanoessa macrura were obtained during the MD 25 FIBEX expedition of the R. V. Marion Dufresne, in February 1981, to the southwest Indian Ocean. Individual and mean coefficients of condition (K _m) were calculated from variations in weight as a function of total length and of carapace length. In E. superba, no significant differences were found as a function of sex or development stage in either total length and weight or carapace length and weight. In T. macrura, a significant allometric difference emerged in the wet weight of juveniles, males and females as a function of total length, and in wet weight as a function of carapace length between adults and subadults. In both E. superba and T. macrura, wet weight as a function of total length differed significantly among stations, and also between species, with a greater weight increase for T. macrura in summer. The coefficients of condition calculated for all wet weight-total length relationships showed that the morphologically different Group II males were heavier than Group I males and mature females. K _m in T. macrura was higher for females than for juveniles or males, indicating a greater weight gain by the females. These differences probably reflect real differences in physiology and may affect the distribution of these two species.     

Reproductive parameters of female Hawaiian damselfish<Emphasis Type="Italic"> Dascyllus albisella</Emphasis> with comparison to other tropical and subtropical damselfishes

The reproductive ecology of female Hawaiian damselfish Dascyllus albisella (Gill) was studied at Kaneohe Bay, Oahu, Hawaii (21°27'N; 157°47'W) during two peak reproductive seasons (June–September 1997, July–August 1998). In both years, spawning occurred cyclically every 5–7 days, with all spawning in the study area in each cycle concluding within 2–3 days. The study period encompassed 16 spawning cycles in 1997 and 10 cycles in 1998. Some females spawned in almost all cycles, whereas others spawned in as few as one. Mean spawning interval of individual females ranged from 6.4 to 11.7 days in 1997 and from 5.5 to 29.0 days in 1998. Batch fecundity increased exponentially with body length, and ranged from 23,100 to 52,800. Spawning duration increased linearly with body length and ranged from 20 to 286 min. On a single day, large females spawned earlier than small females. Females mated with several different males over a season, but typically mated with only one male on a single spawning day. Female spawning frequency and interval, spawning duration, batch fecundity, as well as diel timing of spawning in D. albisella were comparable to those reported for other large, planktivorous, non-territorial damselfishes.     

Social inhibition of maturation in females of the temperate wrasse Pseudolabrus celidotus and a comparison with the blennioid Tripterygion varium

In two temperate reef fish, Pseudolabrus celidotus (Bloch and Schneider) and Tripterygion varium (Bloch and Schneider) studied near Leigh, New Zealand, most spawning activity was concentrated during the first 2 months of spawning seasons which lasted about 5 months. In P. celidotus, maturation and spawning of first-year females (0+) was delayed with respect to older (1+yr) females. Within the 0+ age class, the minimum size of maturation declined as the spawning season proceeded. However, in all females of T. varium, ripening and spawning took place over the same time ranges. Observations on the seasonal patterns of female aggression suggested that larger female P. celidotus were socially inhibiting the maturation of small females. The level of interaction was high compared to that in T. varium, and reached a peak over the onset of the spawning season. This hypothesis was tested by the removal of larger (1+yr) female P. celidotus from a field population. The remaining 0+ females increased in ovary weight and matured earlier than those in an undisturbed area. We argue from this experiment that it is of advantage for P. celidotus females to spawn early in the season and of advantage to inhibit other females from doing so. Two possible reasons are suggested, the first applying to sexchanging species such as P. celidotus, the second to temperate-water fishes in general.     

Life history of the golden ring cowry Cypraea annulus (Mollusca: gastropoda) on Okinawa Island,Japan

Two mark-recapture studies, regular population censuses, field observations, and laboratory culture were used to study the life history of the tropical marine gastropod Cypraea annulus Linnaeus, 1758 from Cape Maeda, Okinawa Island, from April 1984 to March 1986. In the field, the average rate of increase in shell length of marked juvenile snails was 1.0±0.3 mm wk^-1 (N=13), with a maximum of 1.5 mm wk^-1 (initial shell length 10.6 to 17.2 mm). In the laboratory, the maximum growth rate of juveniles was 3.0 mm wk^-1 with food ad libitum. Snails with primordial teeth on the shells grew at a rate of 0.1 to 0.5 mm wk^-1 for about 2 wk. The adults continued to grow at a similar rate for an additional 3 to 6 wk, and ceased detectable growth when some females started spawning egg masses. In the littoral zone on Okinawa Island, snails reproduced throughout the year. Egg masses brooded by three females (shell length 17.4 to 21.3 mm) in the field contained 90 000 to 133 000 ova. In the laboratory, brooding periods of three females lasted 6,8, and 9 d. The estimated average frequency of spawning was 5 egg masses female^-1 yr^-1. The mean shell length of adult females (20.3 mm) was significantly larger than that of adult males (19.6 mm). The life-history strategy of C. annulus is characterized by rapid growth, high fecundity, and repeated spawning throughout the year.     

Krill evolution and the Antarctic ocean currents: evidence of vicariant speciation as inferred by molecular data

The phylogenetic relationships of the Antarctic krill Euphausia superba, the key species in the Antarctic food web, and other Antarctic and sub-Antarctic cuphausiids have been investigated using the 16S ribosomal mitochondrial gene. The phylogenetic reconstructions indicated that the Antarctic species form a monophyletic clade separated by the non-Antarctic species. The results revealed a large genetic divergence between the Antarctic (E. superba and E. crystallorophias) and sub-Antarctic species (E. vallentini). The time of separation between these species, estimated from the molecular data, is around 20 million years ago, which is comparable with the geological time of the formation of a circum-Antarctic water circulation and the Antarctic Polar Frontal Zone. The euphausiid molecular phylogeny therefore represents evidence for vicariant speciation.     

Larval distribution and reproductive ecology of Thysanoessa macrura (Crustacea: Euphausiacea) in the Scotia Sea

In the Scotia Sea, the larvae of Thysanoessa macrura G.O. Sars, 1885 were scarce in spring (September 27 to October 6, 1971) and mainly present at the Calyptopis I stage, while in summer (December 17, 1974 to January 3, 1975) they were much more abundant and much further developed (up to Furcilia VI). The spawning season commences as early as September and, as indicated by the presence of numerous nauplii and metanauplii in summer, is rather long. T. macrura begins to spawn much earlier in West-Wind-Drift waters than in waters of the Weddell Sea. It starts to spawn simultaneously with Euphausia frigida and E. triacantha, but significantly earlier than E. superba. These timings of the breeding process are related to differential biogeographical characteristics of these species. Plankton collections made at depths of 500 or 1000 m have, for T. macrura, revealed a developmental ascent analogous to that of E. superba. Ontogenetical deep-water migrations are, consequently, characteristic not only for E. superba, but also for other Antarctic euphausiids (in particular E. frigida, but possibly not for inshore E. crystallorophias) and, as correctly postulated by several other investigators, for all oceanic euphausiid species.     

A population study of the polychaete Capitella capitata at plymouth

A population of Capitella capitata (Fabricius) was sampled from January, 1973 to February, 1974. Size of the worms was measured and sexual maturity determined. There is an extensive breeding season, spawning occurring asynchronously throughout the year. Females containing ripe oocytes were almost always present, but most females at any one time contained immature oocytes. A high percentage of males contained active sperm. Oocytes are not released into the coelomic fluid until almost fully developed, and all are released at a single spawning. A large number of eggs are produced at each spawning and there is some evidence to suggest that larval development may be completely benthonic. Gametes first develop in 7 to 8 month old worms and take about 4 months to complete development. Analyses of the female population structure suggests that there are 3 breeding classes. The growth rate is estimated at 30 mm per year, and few worms are believed to survive longer than 2 years.     

Effect of temperature on the molting,growth and maturation of the antarctic krill Euphausia superba (Crustacea: Euphausiacea) under laboratory conditions

The effect of temperature on molting, growth, and maturation rates was studied on laboratory-maintained Euphausia superba. The length of intermolt periods (IMP's) was inversely proportional to temperature (20.10 d, SD=1.60, at 0.12°C; 16.87 d, SD=1.68, at 0.97°C; and 12.48 d, SD=0.90, at 4.48°C), and directly proportional to krill size at 0.12°C and 0.97°C. For individually maintained krill the maximum growth rate at 4.48°C (0.068 mm d^-1) was nearly twice that at 0.68°C (0.037 mm d^-1). There was no observable temperature effect on maturation rates. The maturation changes of juveniles at all temperatures indicated that more than two years are probably required to reach maturity. Mature males and females regressed to immature forms, suggesting that E. superba may reproduce in successive years. These results and previously reported field and laboratory data for E. superba and other euphausiid species suggest a 4+ year life span for this species.This work was supported by NSF grant DPP 76-23437