Habitat harshness and morphodynamics: life history traits of the mole crab Emerita brasiliensis in Uruguayan sandy beaches

The habitat harshness hypothesis (HHH) postulates that in reflective beaches the harsh environment forces organisms to divert more energy towards maintenance and they therefore have lower abundance, fecundity, growth and survival rates than in dissipative beaches. Recent investigations have tested this hypothesis through single comparisons of only two beaches, and thus the observed trends in population level variables cannot be attributed incontestably to the beach state, but only to location. Here, abundance, reproduction, recruitment, population structure and body size of the intertidal mole crab Emerita brasiliensis were compared between populations from eight microtidal exposed sandy beaches with contrasting morphodynamics, sampled bimonthly during 22 months throughout the 180 km Uruguayan Atlantic coast. Physical variables and compound indices of the beach state were used to categorize sandy beaches. The results of this bi-annual large-scale analysis were fully consistent with the predictions of the HHH: abundance (total and population components), duration of the reproduction and recruitment seasons and the individual size of megalops and females of the mole crab E. brasiliensis decreased from dissipative to reflective beaches. This was reflected by linear or, mostly, nonlinear relationships between biological and both physical variables and compound indices of beach state. In conclusion, this multi-beach sampling provides compelling evidence of a consistent response of demographic and life history traits of an intertidal beach species to morphodynamic characteristics.     

Reproductive plasticity in mole crabs, Emerita brasiliensis, in sandy beaches with contrasting morphodynamics

Ovarian macroscopical analysis, histological validation and field sampling procedures were used to evaluate the variability in reproductive traits of the mole crab Emerita brasiliensis Schmitt, 1935 (Decapoda: Hippidae) in two exposed sandy beaches of Uruguay with contrasting morphodynamics. All developmental stages involved in the complex life cycle exhibited lower abundance, individual size and temporal occurrence in a harsh reflective beach, compared with a more benign dissipative environment. In addition, this population showed more compressed events of the reproductive cycle (e.g oogenesis, encounter of potential mate and female parental care) and recruitment period. However, the beginning of the vitellogenesis, ovary maturity, male sexual differentiation and ending of spermatogenesis occurred at smaller sizes under dissipative conditions. These results disagree with the recent findings of delayed sexual maturity in dissipative beaches. We postulate that, in dissipative conditions, high food availability might allow an overlapping of reproductive and moulting processes, and thus females may reach optimum size and sexual maturity with fewer moults than in reflective beaches. Hence, reproductive responses must be considered not only in relation to environmental harshness, but also in the context of life history traits and their phylogenetic and allometric constraints. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Geographical patterns in reproductive biology of the Pan-American sandy beach isopod<Emphasis Type="Italic"> Excirolana braziliensis</Emphasis>

Eleven populations of the Pan-American sandy beach isopod Excirolana braziliensis, distributed from tropical (9°N) to temperate (39°S) sandy beaches in Atlantic and Pacific Oceans, were analyzed to determine latitudinal variations in: breeding and recruitment patterns, sex ratios, size of ovigerous females and of juveniles and length–fecundity relationships. E. braziliensis exhibited strong latitudinal patterns in all reproductive traits throughout its distribution range. Breeding and recruitment shifted from continuous to seasonal from tropical to temperate beaches, having a predominance of females at higher latitudes. In agreement with the latitudinal gradient hypothesis, ovigerous females inhabiting tropical sandy beaches (low latitudes) were smaller, became sexually mature at smaller sizes and had lower individual fecundity than on temperate beaches. Juveniles were also smallest at low latitudes. Between-ocean comparisons showed very similar reproductive characteristics for roughly the same latitude. These linked reproductive parameters suggest that the intensity of breeding effort is associated with the duration of the breeding season and geographically size-related characteristics of the species. Geographic variations in the breeding and recruitment seasons, as well as in individual fecundity, size structure of mature females and sex ratios, are proposed to have major consequences in explaining local variations in population demography. Our paper also reinforces the notion that sandy beach animals are highly plastic.Communicated by O. Kinne, Oldendorf/Luhe     

Analysis of lipid components for determining the condition of anchovy larvae,Engraulis mordax

It has been presumed that intertidal spawning by Limulus polyphemus minimizes the loss of egges to subtidal predators; however, this strategy involves considerable risks. Massive beach strandings of adults accompany seasonal spawning migrations of crabs along Cape May in Delaware Bay, (USA). At least 190000 horseshoe crabs, approximating 10% of the adult population, died from beach stranding along the New Jersey shore of Delaware Bay during the 1986 (May to June) spawning season. Abnormalities of the telson (which is used in righting behavior) were significantly more common among stranded crabs than among individuals actively spawning on the intertidal beach. The number of stranded crabs per day was not correlated with tidal height or environmental variables (wind speed, wave height) which characterized the conditions at spawning. A complex suite of factors, including the size of the available spawning population, tidal and weather conditions, and beach slope, influence the number stranded during the breeding season. Horseshoe crab stranding results in a large loss of gravid females from the population, and may represent a major input of organic matter to intertidal sandy beaches in certain regions of Delaware Bay.     

Responses of fish assemblage structures to sandy beach types in Kyushu Island, southern Japan

To determine differences in fish assemblage structures between beach morphodynamic types on an exposed sandy shore, three beach types (reflective, intermediate, and dissipative) were sampled at Fukiagehama Beach, Kyushu Island, southern Japan, in May, August, and November 2006 and 2007. Distinct differences in physical conditions and the amount of major prey invertebrates, such as copepods, mysids, and amphipods, were recognized between the three beach types. The reflective beach was characterized mainly by turbulent swashes, coarse sediment, and a low abundance of major prey invertebrates. In contrast, the dissipative beach was characterized mostly by benign swashes, fine sediment, and a high abundance of prey invertebrates. The intermediate beach tended toward an intermediate condition between the reflective and dissipative beaches. The fish assemblage structures also differed clearly between the three beach types, with species and individual numbers being greatest on the dissipative beach and lowest on the reflective beach. A similarity index indicated differences in species composition of the assemblages between the beach types. Such differences arose primarily from the differential distributions of two dominant trophic groups, zooplankton (mostly copepods) and epibenthic crustacean (mainly mysids and amphipods) feeders, between the beach types, both groups tending to increase in species and individual numbers from the reflective beach to the dissipative beach. In addition, the mean standard length of all fish collected in each month tended to decrease toward the dissipative beach. These results suggested that beach types strongly influence sandy beach fish assemblage structures.     

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     

Morphodynamic responses of nourished beaches in SW Spain

Coastal erosion in SW Spain is affecting man-made structures and beaches that represent an important economic resource in the area. In the last decade the Spanish government carried out several nourishment works that have limited durability. Most of the artificial beach fills consist of a spill of natural dredged sand on the visible beach, leading to a flat, artificial berm with an important seaward slope and narrow foreshore. As a result, the initial dissipative profile was transformed into a fully reflective one. A beach monitoring program was carried out to record morphological evolution after the nourishment works. Several field assessments of disturbance depth were also made to characterize beach morphodynamics of a nourished beach (Rota) and a natural dissipative one (Tres Piedras), whose slope was similar to the pre-nourishment gradient of Rota beach. Natural dissipative beaches were characterized by spilling breakers that did not significantly affect bottom sand. The severe erosion recorded in the nourished zones was related to the new morphodynamic regime acting on these beaches, which was controlled by high erosive plunging breakers associated with high foreshore slopes. In conclusion, other nourishment practices should be used, better adapted to the natural beach morphodynamics of the zone, taking also into account the original grain size and density of the beach sands, in order to obtain more durable artificial beaches.     

Beach erosion and geochemical factors: Influence on spawning success of horseshoe crabs (Limulus polyphemus) in Delaware Bay

Horseshoe crab spawning activity is spatially patchy within the Delaware Estuary. This study investigated the importance of geochemical and erosional factors to the selection of breeding beaches. Two sandy beaches in Cape May county, New Jersey, USA, were studied; one beach had been subjected to considerable erosion, exposing underlying peat; the second beach, less than 1 km away, had only traces of peat. Reduced sediments with high levels of hydrogen sulfide were correlated with the presence of peat, and significantly fewer crabs utilized sediments in the proximity of peat beds for reproduction. The lower spawning activity on the beach in the vicinity of exposed peat, suggests that crabs may detect, at a distance, the nature of sediments and the quality of beach for spawning activity. Active salt marsh and peat-bank sediments dominate the upper bay shore; these sediments are unsuitable, or at best marginal, for horseshoe crab reproduction. Extensive bulkheading of eroding sandy beach along several New Jersey shore communities has further restricted the availability of suitable spawning habitat, making the remaining stretches of optimal sandy beach critical to the reproductive success of this species.     

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.     

Reproduction of Halicarcinus planatus (crustacea, decapoda, hymenosomatidae) in the Deseado River estuary, southwestern Atlantic Ocean

The small spider crab Halicarcinus planatus, is a common resident in the rocky coasts of the Magellanic region in South America. The H. planatus population in the Deseado River estuary differs from other populations of the same species in the spatial segregation of sexes. Between March 2002 and October 2003, samples were taken monthly from the lower intertidal and upper subtidal levels. All crabs captured were females. Additional samplings were carried out at the subtidal level to search for the presence of males. A total of 3,346 females were captured and their relative age was determined. Some of these females were maintained in the laboratory to study the terminal moult and the spawning seasons. The analysis of the ovary development of adolescents (ADO) and mature (MAT) females, and the times when females of different ages appear, when spermatophores are found inside spermathecae, and when post-spawning mortality takes place allowed us to establish the whole reproductive process. Adolescents are impregnated only once before the terminal moult. There are seven or eight spawnings after the moult, with a brief resting period between two successive reproductive seasons. The life span of mature females ranges between 15 and 22 months. Results indicate that the H. planatus population of the Deseado River estuary differs greatly from that of the Kerguelen Islands.     

Annual reproductive cycles of the commercial sea urchinParacentrotus lividus from an exposed intertidal and a sheltered subtidal habitat on the west coast of Ireland

Reproduction of the commercial sea urchinParacentrotus lividus (Lamarck) from contrasting habitats on the west coast of Ireland was examined from May 1986 through August 1988. Urchins were collected intertidally from an exposed rocky shore and subtidally from a protected bay. Monthly measurements of the gonad index and histological examination of the gonads demonstrated thatP. lividus has an annual reproductive cycle. Although the two populations exhibited similar reproductive patterns over three breeding seasons, there were inter-annual and inter-population differences in the amplitude of gonadal growth and in the time when spawning started. The subtidal urchins had significantly larger gonads and exhibited a longer period of reproductive maturity compared with the intertidal urchins. This difference was also evident in the histology of the ovaries at the beginning of breeding, when most of the subtidal females contained mature ovaries, whereas most of the intertidal females contained premature ovaries. An inter-annual difference in the onset of spawning was observed, with the start of gamete release differing by as much as four weeks between years. It appears that inter-annual differences in sea temperature influence the temporal variation in spawning byP. lividus and that increasing temperature may serve as an exogenous cue for gamete release. The inter-annual variability in the onset of spawning suggests that photoperiod does not cue gamete release. Gonadal growth, on the other hand, occurs during the coldest months of the year and during the period of shortest days, suggesting that temperature and photoperiod may both influence gonadal growth during the winter. Oogenesis and spermatogenesis ofP. lividus were examined histologically. During the post-spawning recovery and growth stages, the gonads gained weight through growth of the nutritive phagocytes and the accumulation of periodic acid Schiff (PAS) + droplets by these cells. The PAS + material appears to play a nutritive role in gametogenesis. For the females, the frequencies of six ovarian maturity stages was assessed at approximately monthly intervals. Small oocytes were present throughout the year and clusters of early oocytes were most apparent during the spent and recovery stages. With the onset of vitellogenesis and subsequent accumulation of ova, the nutritive phagocytes and their PAS + droplets became depleted. The ovarian condition at the onset of breeding was variable, due to differences in the number of vitellogenic oocytes, differences in the number of ova in storage, and differences in the amount of PAS + material. In general, the nutritive phagocyte tissue is reduced by the onset of spawning and is exhausted by the termination of breeding. A similar series of events occurs during spermatogenesis. The relevance of this study for the use ofP. lividus as a brood-stock organism for mariculture is discussed.     

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.     

Gametogenesis and spawning periodicity in the fan worm <Emphasis Type="Italic">Sabellastarte spectabilis</Emphasis> (Polychaeta: Sabellidae)

The sabellid polychaete Sabellastarte spectabilis (Grube 1878) was collected at approximately monthly intervals from January 2002 to December 2003 from intertidal and subtidal reefs near the Hawaii Institute of Marine Biology in Kane’ohe Bay, Hawaii, USA (21°N, 157°W). Gametogenesis and spawning periodicity were investigated using histological techniques and induction of spawning trials. Worms were characterized into four discrete reproductive stages based on histological evidence: (1) No evidence of reproductive activity in the coelom (sex cannot be determined), (2) Only coelomocytes present in the coelom (sex cannot be determined), (3) Some gametes present in the coelom (sex can be determined) and (4) Coelom densely packed with gametes (sex can be determined). The small hermaphroditic portion of the population was not used in this study. Stage 4 worms were present over an extended period of time (females, March–December and males, March–November) indicating a potentially broad reproductive season. No correlation between day length and maturation stages in S. spectabilis was detected. However, the statistical model Y = ([394.26 × X] − [7.793 × X ²]) − 4960.781 where Y the % frequency of Stage 4 worms and X the mean monthly water temperature explained 44% of the variation between water temperature and % frequency of Stage 4 worms. Maturation appeared to coincide with water temperatures of 24–25°C (March–September) after which there is a reduction in the % frequency of stage 4 individuals. Induction of spawning trials conducted between May and January showed the month of October with a significantly higher percent success than any other month investigated. According to all available information (e.g., natural spawning in water tables, histological data, induction of spawning trials, correlation of maturation stages with observed changes in average monthly water temperature.), there is an apparent peak in reproductive activity (spawning) within a broad maturational season, which may be influenced by water temperature.     

The reproductive cycle of Asterina stellifera (Möbius) (Echinodermata: Asteroidea) in the Cabo Frio region, southeastern Brazil

Reproduction of Asterina stellifera was studied in two populations from the Cabo Frio region, southeastern Brazil. One was located in an intertidal seagrass flat at Japonês Island (1994–1995); the second was on a sublittoral rocky shore at Pontal beaches (1999–2000). Reproductive cycle was studied at monthly intervals by organ indices and histological examinations of gonads. Although these two populations were studied in different habitats and at different times, the periodicity of the annual reproductive cycle was very similar for the two populations. The highest gonad index (GI) values were found in August/September (late winter), followed by a drop in GI values, indicating spawning. The GI and pyloric caeca index (PCI) tended to have an inverse relationship in both populations, but it was only statistically significant in the rocky-shore population. Gametogenesis in the two populations was described and roughly confirmed the reproductive cycles determined by GIs. Published online: 9 August 2002     

Seasonal and spatial differences in the fish assemblage of the non-estuarine Sulaibikhat Bay,Kuwait

In the period September 1986 to August 1987 fish were captured once a month, using an otter trawl, from the intertidal and subtidal regions of Sulaibikhat Bay, Kuwait. Correlation analysis revealed that numbers were not directly related to temperature at the time of sampling. The recruitment of large numbers of young of the year Liza carinata (Valenciennes) during March was related to low temperatures in November, the probable time of spawning of this species. Salinity at the time of sampling was inversely correlated with numbers. This result indicates that large numbers of 0+fish recruit to the Bay during the period of maximum fresh water outflow through the Shatt-al-Arab. Numbers of fish were significantly greater in the intertidal region, where they were present almost entirely as 0 group fish, than in the subtidal. The three dominant species of the assemblage are shown to use the two depth intervals in differnt ways. L. carinata was capture a almost exclusively in the intertidal region as 0+ fish. Pomadasys stridens (Forsskål) was captured as 0+ fish in both the intertidal and subtidal regions but in greater numbers in the intertidal region. Leiognathus brevirostris (Valenciennes) was captured as small, mostly 0+ individuals in the intertidal region and as larger fish in the subtidal region. The use made by the smaller fish of the intertidal region is related to the avoidance of sublittoral predators and reduction of intraspecific competition, whilst larger fish in the sublittoral region may be avoiding predation by piscivorous birds.     

Long-shore distribution of morphodynamic beach states along an apparently homogeneous coast in SW Spain

We present a morphodynamic study of an apparently homogeneous rectilinear coast in SW Spain. The study area covers 14 km of mesotidal sandy beaches, interrupted in some places by rocky-shore platforms. The method used consisted of a monthly monitoring of 12 beach profiles during two years. According to the results obtained from the study, which also include granulometric analyses andin situ determination of the beach disturbance depth, three main beach classes have been differentiated: low-reflective beaches, dissipative beaches and rocky-shore platform beaches. Their longitudinal distribution is not linked to their distance to the main source of sediments in the area (mouth of the river Guadalquivir). Instead, a very irregular long-shore variation of morphodynamic beach states appears. It is deduced that this long-shore variation is mainly linked to local contouring conditions (e.g. the presence of rocky shoals which affect wave-breaking processes), and not to the regional long-shore currents prevailing in the zone.     

Mating behaviour and reproductive cycle of Archaster typicus (Echinodermata: Asteroidea)

This study examines the reproductive cycle, the mechanism of male-on-female pairing behavior and the spawning behavior of Archaster typicus Müller et Troschel. Field studies were conducted in the intertidal zone of the sand beaches at Penghu, Taiwan (23 32N; 119 33E) at ebb-tide in 1984 and 1985. The pairing behavior of A. typicus is a reproductive behavior which leads to simultaneous spawning, increasing the probability of fertilization. As the breeding season approaches, sea stars, especially males, display increased mobility. Because only males tend to mount another individual and because males can detect the sex of another individual by contact with the side of their arms, a male-on-female pair is formed when a male encounters a female. Eighty-five percent of the sea stars observed were paired during the pairing season. Spawning by a paired female is closely followed by spawning of its paired male; male spawning, however, does not induce spawning in its paired female. During spawning, the male turns slightly so that its arms overlap the arched arms of the female. The gonad volume of males is much less than that of females. This may result from the high efficiency of fertilization in this species, which does not require a large amount of gametes to be released, or from the higher energetic demands made on the males.     

Intertidal slope of coral sand beach as a unique habitat for fish: meiobenthic diet of the transparent sand dart, <Emphasis Type="Italic">Kraemeria cunicularia</Emphasis> (Gobiidae)

Coral sand beaches harbor gobiid sand darts (genus Kraemeria), the only fish known to live in the sand throughout their adult life. Despite the uniqueness of sand-dwelling habitat as a vertebrate, the biology of this fish remains unclear. To explore how this unique fish utilize an unusual habitat, we investigated diurnal patterns of microhabitat use and prey consumption by the transparent sand dart, Kraemeria cunicularia, at a sandy beach on Iriomote Island, the Ryukyu Archipelago, Japan. Sand darts were found in sediment in the lower intertidal zone throughout the daytime regardless of changes in tidal level, whereas at nighttime these fish were found swimming. Gut content analyses revealed that the sand dart diet was dominated by harpacticoid copepods throughout the day. Analyses of meiobenthic distribution indicated that these copepods were most abundant at lower intertidal zones where highest numbers of sand darts were found during the daytime; thus, it is possible that microhabitat use of the fish is largely determined by food availability. An extensive distributional survey throughout the Ryukyu Archipelago further indicated that sand darts prefer sandy beaches with well-sorted, coarse sand. These results provide novel insights into how sand darts respond to the tidal rhythm and highlight putative key environmental factors that determine their distribution at both regional and microhabitat scales. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Coexistence and reproductive isolation of the sympatric echinoids<Emphasis Type="Italic"> Diadema savignyi</Emphasis> Michelin and<Emphasis Type="Italic"> Diadema setosum</Emphasis> (Leske) on Kenyan coral reefs

The sympatric echinoids Diadema savignyi and D. setosum coexist in shallow reef lagoons throughout East Africa. The reproductive strategies of these echinoids were studied to investigate reproductive isolation as a possible mechanism for maintaining the coexistence of these closely related species. The annual reproductive cycle and lunar periodicity were determined by gonad index measurements, histological examination of gametogenesis, and induction of spawning with injections of KCl. The peak reproductive period of D. savignyi coincided with the north-east monsoon period (when light and temperatures are high) as gonad indices were high (>8%) beginning in February and peaked at 9.7% in May. Gonad indices subsequently rapidly decreased (by 26%) in June at the beginning of the cooler south-east monsoon period. However, the presence of sperm and ova in most months of the year indicates continuous gametogenesis with reduced reproductive effort during the cooler months. The annual cycle for D. setosum showed less of a seasonal trend as gonad indices remained above 7% throughout much of the year but tended to be highest when temperatures were lower. This is the first confirmation of continuous reproduction in these two species at the equator. The reproductive patterns of both species remained consistent over 2 years of sampling. Both species exhibited a synchronized lunar spawning periodicity during the 3 months sampled, with D. setosum spawning on lunar days 8–10 and D. savignyi spawning after the full moon (lunar days 17–18). Whereas spawning in D. savignyi was very tightly synchronized, 20% of D. setosum individuals still spawned after the peak spawning period. The coexistence of these closely related species appears to be maintained by temporal reproductive isolation during the lunar spawning period reinforced by seasonal differences in reproductive effort.Communicated by O. Kinne, Oldendorf/Luhe