Influence of a deep artificial entrance channel on the biological characteristics of the blue swimmer crab Portunus pelagicus in a large microtidal estuary

The opening of a deep and permanent artificial entrance channel into the Peel-Harvey Estuary in southwestern Australia in 1994 provided an excellent opportunity to determine the biological characteristics of the blue swimmer crab Portunus pelagicus in a large microtidal estuary, after tidal exchange with the ocean had increased markedly and salinities recovered more rapidly from their winter minima. P. pelagicus was collected from the original entrance channel, which is located 15 km to the north of the artificial channel, the two large basins (Peel Inlet and Harvey Estuary) and the saline lower reaches of a major tributary at regular intervals in 1995-1998. The resultant information on densities, size and age compositions, growth rates and reproductive biology were compared with those derived from data recorded by Potter et al. [Mar. Biol. 78 (1983) 75] for P. pelagicus in the Peel-Harvey Estuary in 1980 and 1981. This led to the following conclusions regarding the ways in which this portunid has responded to the impacts of the construction of an artificial entrance channel in this estuary. (i) Late 0+ crabs enter earlier than previously the Peel Inlet and both the Harvey Estuary, which now opens directly to the sea, and a tributary river in which salinities likewise rise more rapidly in the spring. (ii) Female crabs become ovigerous earlier, in concert with the earlier restoration of high salinities in the spring. (iii) Ovigerous crabs emigrate to the ocean sooner, presumably in response to the greater stimulus provided for emigration by the marked increase that has occurred in tidal flow. (iv) The growth rate during the first few months of life has increased, possibly as a consequence of a decline in the density of crabs within the estuary during those months. (v) A cohort of small 0+ crabs, not previously found in the estuary and almost certainly derived from population(s) spawning further south, is present for a few months but then disappears from the estuary. Although large crabs do not now appear to grow towards their maximum size as rapidly as previously, this is an artefact brought about by the culling of large crabs through the marked increase that has occurred in fishing pressure during recent years.     

Variation in the sex ratio, size and age of longfinned eels within and among coastal catchments of south‐eastern Australia

Longfinned eels Anguilla reinhardtii were captured by both fishery‐dependent and independent sampling methods from three rivers in New South Wales, south‐eastern Australia. Sex ratios, catch per unit effort and population age and total length structure were examined in three zones (fresh water and upper and lower tidal) in the Hacking, Hawkesbury and Clarence Rivers. Females were found in relatively high proportions in all zones, ranging from 97% in a freshwater (non‐tidal) site down to 59% in a tidal site. Males were found primarily in tidal zones (only two of the 677 longfinned eels caught in non‐tidal fresh water were males), with the greatest proportions being found in the brackish upper tidal areas. The mean number of fish captured per trap was higher in the fresh water and upper tidal zones than in the lower tidal zones. The mean ±  s . e . age, 17·9 ± 0·3 years, and age range, 5–52 years for females were significantly higher than those of males 12·2 ± 0·4 years; range 5–22 years, which is typical of other anguillid species. Longfinned eels captured in fresh water were found be significantly larger and older than those in tidal zones due to the almost exclusive predominance of females.     

Nitrogen fixation by Nodularia spumigena Mertens (Cyanobacteriaceae). 1: Field studies and the contribution of blooms to the nitrogen budget of the Peel-Harvey Estuary,Western Australia

Variations in nitrogen fixation (acetylene reduction) by Nodularia spumigena blooms in the Peel-Harvey estuarine system were examined with respect to spatial (sampling station location, and depth) and temporal (seasonal and diurnal) distribution. The annual contributions of nitrogen fixation by the blooms to the nitrogen budget of the estuary were estimated to range from 309 to 713t. Contributions by nitrogen fixation were similar to the riverine inputs in the Harvey Estuary, but lower in the Peel Inlet.The Harvey Estuary had higher biomass and total fixation rates (to 0.4 nmol C₂H₂ · ml^–1 h^–1), but the heterocyst nitrogen fixation rates were greater in the Peel Inlet (to 9 × 10^–1 nmol C₂H₂ · heterocyst^–1 · h^–1). Nitrogen fixation decreased with depth in response to light, though other factors also appeared to be involved. The rates of fixation decreased concurrently with increasing bloom age, total soluble inorganic nitrogen and salinities. Maximum daily fixation rates occurred in the early morning.     

Comparison of fish impingement by a thermal power station with fish populations in the Ems Estuary

On an annual basis >10 million individuals of c . 35 fish species are impinged at the EC20 unit of the Eems power station located on the Ems Estuary. The most abundant are: herring Clupea harengus , gobies ( Gobiidae ), Nilsson's pipefish Sygnathus rostellatus , three-spined stickleback Gasterosteus aculeatus and sprat Sprattus sprattus . The impingement shows a seasonal pattern which reflects the presence of the fishes in the estuary. The results are compared with other impingement studies at power stations on the Belgian and Dutch estuaries.
In 1995, the cooling water entrance was displaced from the shoreline to 300 m off the coast at the edge of the tidal channel. The effect of this relocation on the level of fish impingement, determined by comparing the ratio of fish density in the cooling water and in the estuary before (1992–1993) and after (1996–1997) the relocation of the intake canal is discussed.     

Diel distribution patterns of fishes in a temperate large lowland river

During the day, juveniles and typical inshore species in the lower Oder River, Germany, inhabited the littoral (0·05–1·50 m deep) while potamal species inhabited the mid‐channel (40–100 m from the banks). During the night, two behavioural groups were distinguished: 1) potamal species and large individuals of other species, moved from offshore to the littoral, and 2) nocturnal foraging species, became active and increased in inshore and offshore habitats. Inshore, the most significant differences at night was the increasing density of silver bream Blicca bjoerkna increasing both total biomass and total length of fishes caught. Offshore, the increasing density of whitefin gudgeon Gobio albipinnatus at night was most significant. Additional night electrofishing improved the assessment of the abundance, age and size structure of typical potamal fish species. It should be incorporated in large river fish monitoring programmes.     

A new technique to gather 3‐D spatial information using a single camera

A new technique using a single camera and shadows to determine 3‐D spatial positions of fishes in the laboratory is described. The apparatus consisted of a large aquarium (2·0 × 1·5 × 0·4 m), a wide‐angle camera mounted above and two light sources to cast shadows to either side of the fish. Using image analysis and vector mathematics, aquarium objects were plotted within 1·5 cm of their actual location along the x ‐, y ‐ and z ‐axis. The technique was also successful in quantifying changes in 3‐D spatial pattern of juvenile fish, Atlantic cod Gadus morhua (7·4–8·6 cm standard length, L _S) and cohabitant piscivorous shorthorned sculpin Myoxocephalus scorpinus (12·0–25·8 cm L _S), at these same viewing fields. The new technique should have a wide application, largely because it is potentially less expensive, laborious and invasive than alternative methods for determining 3‐D positions of fishes.     

Evidence for independent origin of two spring‐spawning ciscoes (Salmoniformes: Coregonidae) in Germany

Combined analyses of mitochondrial DNA (mtDNA) and microsatellite loci were performed to assess the genetic differentiation of two spring‐spawning ciscoes from each other and from sympatric Coregonus albula in two German lakes. Polymorphism was screened at six microsatellite loci and mtDNA for a total of 247 and 94 ciscoes, respectively. Microsatellite data showed a weak differentiation between spring‐spawning Coregonus fontanae and sympatric C. albula in Lake Stechlin ( F _ST = 0–0·008), whereas a significant differentiation was observed between spring‐spawning Coregonus lucinensis and sympatric C. albula in Lake Breiter Luzin ( F _ST = 0·013–0·039). A more pronounced genetic difference was observed between both spring‐spawning species ( F _ST = 0·05–0·128). Shared mtDNA haplotypes among sympatric species within both Lake Stechlin and Lake Breiter Luzin were observed, whereas no haplotype was shared between C. fontanae and C. lucinensis . These results suggest an independent origin for spring‐spawning ciscoes in each lake. Evidence is also provided for mtDNA introgression of Coregonus sardinella into C. lucinensis and C. albula in Lake Breiter Luzin. Postglacially, this species or at least a population which showed mtDNA introgression has colonized the Baltic Sea basin up to the glacial margin that was located between Lakes Stechlin and Breiter Luzin.     

Macrotidal estuaries: a region of collision between migratory marine animals and tidal power development

Macrotidal estuaries of the inner Bay of Fundy are utilized by large numbers of migratory fishes, particularly dogfish, sturgeon, herring, shad, Atlantic salmon and striped bass as well as by other migratory marine animals, many of which have large body sizes (squid, Lamnid sharks, seals and whales). Tagging experiments indicate the fishes originate from stocks derived over the entire North American Atlantic coast from Florida to Labrador. Population estimates suggest up to 2.0 times 10⁶adult American shad (Alosa sapidissima) migrate through an individual embayment each year. These migrations are an integral part of the life history of the respective species and appear to be controlled in part by the near shore movements of ocean currents. In other regions of the world similar macrotidal estuaries exist (Cook Inlet, Alaska; Severn Estuary, U.K.) and they, like the Bay of Fundy, are linked in continuum to the local ocean currents. We propose that marine animals utilize all these regions in a manner similar to the Bay of Fundy estuaries and properly designed surveys will reveal their presence. Fish passage studies utilizing the Annapolis estuary low-head, tidal turbine on the Bay of Fundy have shown that turbine related mortality of 20–80% per passage occurs depending on fish species, fish size and the efficiency of turbine operation. We suggest that introduction of tidal turbines into open ocean current systems will cause widespread impact on marine populations resulting in significant declines in abundance.     

Comparison of fish communities between above‐ and below‐dam sections of small streams; barrier effect to diadromous fishes

To analyse the effects of small‐scale dams on fish communities, species diversity, population density, biomass, migration over dams and trophic relationships were investigated in the above‐ and below‐dam sections of three Japanese streams. The dams were 1·5–3·9 m high, constructed within 400 m of the sea, and had no impoundment or fishway. Fish communities were mainly composed of diadromous fishes. Of the 17 species sampled, only a few species of gobies, Rhinogobius spp. and Gymnogobius spp. were able to climb and migrate upstream over the dams. Consequently the number of species, and total density and biomass of fishes below the dams was high, but low above the dams. One of the most dominant fish below dams, Plecoglossus altivelis altivelis , predominantly fed on benthic algae, whereas other fishes preyed on benthic invertebrates. Trophic relationships above dams were very simple, the few species of gobies preying on chironomid larvae and other aquatic invertebrates.     

Feeding ecology and habitat preferences of the catfish genus Trichomycterus in low‐order streams of the Colombian Andes

Six lower order streams in the south‐west of Colombia were sampled on a monthly basis in order to determine the status of Trichomycterus species in relation to their food supply (macroinvertebrates), and physico‐chemical habitat. Fishes were sampled by electrofishing and samples taken for dietary analysis. Macroinvertebrates were collected using a Surber sampler. Trichomycterus spp. populations were present in all streams in habitats ranging from soft to hard substrata and from relatively stagnant waters to fast flowing streams. The average density of the fishes ranged from 0·06 to 1·14 individuals m⁻² with peaks in population corresponding to recruitment of individuals of <1 g. Trichomycterus spp. were found to be benthic carnivores with a diet mostly of aquatic insects. The index of relative importance for the dietary items was 35·0, 10·5, 1·9 and 7·8% for Chironomidae, Trichoptera, Ephemeroptera and Oligochaeta respectively. Combined with physical habitat data, ordinations of fishes and macroinvertebrate data suggested that differences in abundance among sites were driven by stream size and flow regime, indicating dominance of the physical habitat, although a high correlation between Trichomycterus spp. and Chironomidae abundances ( r  = 0·81, P  = 0·049) was also recorded.     

The ichthyoplankton of a seasonally closed estuary in temperate Australia. Does an extended period of opening influence species composition?

Fish eggs and larvae were collected monthly between September 1987 and April 1989 from sites throughout the main basin and within the saline regions of the two main tributary rivers of Wilson Inlet, a seasonally closed estuary in south-western Australia. Of the eggs, 43.7% belonged to Engraulis australis (Shaw) and the rest to unidentified teleosts. The larval fish assemblage comprised 17 families represented by 25 species. The Gobiidae contained the highest number of species (four) and contributed approximately 57% of all larvae caught. Pseudogobius olorum (Sauvage) and E. australis were the most abundant species, contributing 43.9 and 27.9% to the total larval catch, respectively. The larvae of species which breed within Wilson Inlet dominated the assemblage, both in terms of number of species (64%) and contribution to total catch (99.9%). The numbers of the eight marine species and one freshwater species represented in the ichthyoplankton were very low. Classification and multi-dimensional scaling ordination showed that the composition of the larval fish fauna at the various sites during a period when the estuary remained open to the sea (December 1988-April 1989) was similar to that of the corresponding sites during the same period in the previous year when the estuary had become closed (December 1987-April 1988). This can be attributed to the spatial distribution, time of occurrence and abundance of estuarine-spawned larvae being similar in the two periods and to the rarity of marine-spawned larvae, even in the spring and summer of 1988/1989 when the estuary was open for the whole time when most marine teleosts spawn in south-western Australia. The low occurrence of marine-spawned larvae in Wilson Inlet reflects the fact that tidal water movement within the basin of the system is so small that it is unable to facilitate the transport and dispersion of larvae. The ichthyoplankton of Wilson Inlet resembles that of other poorly-flushed estuaries in that it is low in species richness and dominated by estuarine-spawned larvae.     

Muscle fine structure reflects ecotype in two nototheniids

The fine structure of swimming (pectoral) and myotomal (axial) skeletal muscle and myocardium of two species of Antarctic nototheniid fishes were studied by electron microscopy, comparing the cryopelagic Pagothenia borchgrevinki and the benthic Trematomus bernacchii . Mean fibre size varied by a factor of four among muscles within each species and may have reflected the locomotory power available, being larger in pectoral oxidative (red) and axial glycolytic (white) muscle of P. borchgrevinki . Both species use labriform locomotion, and the more active P. borchgrevinki had a greater capillary supply, expressed as a capillary to fibre ratio, than T. bernacchii to both red (3·48 ± 0·36 v . 1·63 ± 0·14, mean ±  s . e .; P  < 0·01) and white (2·70 ± 0·20 v . 1·53 ± 0·18, mean ±  s . e .; P  < 0·01) regions of the pectoral musculature. The greater aerobic scope of P. borchgrevinki was strikingly demonstrated in the higher mitochondrial content of all skeletal muscle types sampled, and the ventricular myocardium (0·269 ± 0·011 v . 0·255 ± 0·012 mean ±  s . e .; P  < 0·05). Minor differences were found in other elements of fibre composition, with the exception of a five‐fold greater lipid content in pectoral red fibres of P. borchgrevinki (0·074 ± 0·014 mean ±  s . e .) v . T. bernacchii (0·010 ± 0·003; P  < 0·05). Differences in muscle fine structure among species clearly reflected differences in their ecotype.     

The biology of the bigeye grenadier at South Georgia

The biology of the bigeye grenadier Macrourus holotrachys caught as by‐catch in the Patagonian toothfish Dissostichus eleginoides longline fishery conducted around South Georgia was investigated to improve data available for fisheries management. Age estimates suggest that M. holotrachys is a moderately slow growing species ( K  = 0·10), reaching ages of >30 years and attaining total lengths ( L _T) >80 cm ( L _∞ = 33). The size at which 50% of females had started to mature ( L _int50) for M. holotrachys was 21 cm pre‐anal length ( L _PA) and occurred at c . 9 years old. Estimates of natural mortality and Pauly's growth performance index were found to be low ( M  = 0·09 and Φ = 2·82 respectively). Gonad maturity stage was described from macroscopic and histological investigation. Mature ovaries had oocytes at all developmental stages with between 22 and 55% likely to be spawned each year. Absolute fecundity ranged from 22 000 to 260 000 eggs and was positively correlated with both pre‐anal length and mass. A highly skewed sex ratio of 32 : 1, females : males, was found for specimens caught by longlines but not for a small sample of shallower trawl‐caught specimens. It is suggested that females are far more susceptible to longline capture than males. Macrourus holotrachys is a bentho‐pelagic predator and scavenger that feeds on a wide range of fishes and invertebrates. The fish are long lived, slow‐growing species typical of deep‐water grenadiers; fisheries management strategies should reflect their probable susceptibility to overfishing.     

Interhabitat and instream movements of large Atlantic salmon parr in a Newfoundland watershed in winter

Radio‐telemetry was used to investigate movement of large, mainly mature male (80%) Atlantic salmon Salmo salar parr in Stoney River, Newfoundland during early winter (November; water temperature 6·0 ± 0·1° C) and mid‐winter (January to February; 0·8 ± 0·0° C). Site fidelity of parr in early winter was low. Parr moved between fluvial and lacustrine habitats and were active throughout the diel cycle. Parr caught in fluvial habitats in mid‐winter were smaller and younger than parr caught in early winter. Site fidelity of parr in mid‐winter was greater than in early winter. Parr in mid‐winter moved between fluvial and adjacent small lacustrine habitats, but avoided a larger pond inhabited by large piscivorous fishes. Instream movement rates in mid‐winter were lower than in early winter and occurred primarily during hours of darkness (dawn, dusk and night). Fluvial habitats were relatively stable and ice‐free throughout the study periods. These results suggested that large Atlantic salmon parr utilize a variety of habitats and remain active throughout the winter, even under stable environmental conditions.     

Characteristics of the ichthyofaunas of southwestern Australian estuaries,including comparisons with holarctic estuaries and estuaries elsewhere in temperate Australia: A review

Data on the species compositions and the ages, sizes, reproductive biology, habitats and diets of the main species in the ichthyofaunas of seven estuaries in temperate southwestern Australia have been collated. Twenty-two species spawn in these estuaries, of which 21 complete their lifecycles in the estuary. The latter group, which includes several species of atherinids and gobies with short lifecycles, make far greater contributions to the total numbers of fish in the shallows of these estuaries than in those of holarctic estuaries, such as the Severn Estuary in the United Kingdom. This is presumably related in part to far less extreme tidal water movements and the maintenance of relatively high salinities during the dry summers, and thus to more favourable conditions for spawning and larval development. However, since estuaries in southwestern Australia have tended to become closed for periods, there would presumably also have been selection pressures in favour of any members of marine species that were able to spawn in an estuary when that estuary became landlocked. Furthermore, the deep saline waters, under the marked haloclines that form in certain regions during heavy freshwater discharge in winter, act as refugia for certain estuarine species. The contributions of estuarine-spawning species to total fish numbers in the shallows varied markedly from 33 or 34% in two permanently open estuaries to ≥ 95% in an intermittently open estuary, a seasonally closed estuary and a permanently open estuary on the south coast, in which recruitment of the 0 + age class of marine species was poor. The larger estuarine species can live for several years and reach total lengths of ~ 700 mm and some estuarine species move out into deeper waters as they increase in size. Several marine species use southwestern Australian estuaries as nursery areas for protracted periods. However, sudden, marked increases in freshwater discharge in winter and resultant precipitous declines in salinity in the shallows, and in other regions where haloclines are not formed, are frequently accompanied by rapid and pronounced changes in ichthyofaunal composition, partly due to the emigration of certain marine species. In contrast, the ichthyofaunal compositions of macrotidal holarctic estuaries undergo annual, cyclical changes, due largely to the sequential entry of the juveniles of different marine species for short periods. The ichthyofaunal compositions of the narrow entrance channels, wide basins and saline riverine reaches of large, permanently open southwestern Australian estuaries vary, reflecting the marked tendency for some species to be restricted mainly to one or two of these regions. Comparative data indicate that the characteristics determined for ichthyofaunas in southwestern Australian estuaries apply in general to estuaries elsewhere in temperate Australia.     

First observations of the burrows of Anguilla japonica

The burrow morphology of the Japanese eel Anguilla japonica was studied using in situ resin‐casting in a mud bottomed tidal drainage channel adjacent to the Fukui River in Tokushima, Japan. Two eels (62·5 and 56·3 cm total length) were initially fished from the burrows to verify that they were being used by A. japonica . Casts were made of 10 burrows that were found to have from one to three openings and main tunnels that were parallel to the axis of water flow in the channel. The maximum depths of the tunnels in the mud were 17·8–30·0 cm. The diameters of main tunnels ranged from 1·2 to 7·9 cm, were almost always wider than the bodies of the Japanese eels examined, and were more variable in the horizontal axis than in the vertical axis. There were no other animals capable of constructing a long and thin burrow in this channel, so these observations indicate that anguillids are able to construct their own burrows in soft mud sediments that may be used for extended periods of time.     

The genetic architecture of correlations among growth‐related traits and male age at maturation in rainbow trout

Heritabilities and genetic correlations among growth‐related traits of two cultured strains (Rainbow Springs and Spring Valley) of rainbow trout Oncorhynchus mykiss were estimated using restricted maximum likelihood methods with a three‐generation pedigree. Heritability was high (>0·50 ± 0·03) for body mass and condition factor but moderate (0·35 ± 0·04) for age at sexual maturity in males. Body mass and age at sexual maturation were phenotypically correlated in the families of one experimental strain, Rainbow Springs, and had a positive genetic correlation (0·26 ± 0·03) across families from both test strains (Rainbow Springs and Spring Valley). This indicates that faster growing individuals were more likely to mature at 2 years of age than slower growing individuals in the two hatchery strains investigated. Microsatellite markers of body mass quantitative tract loci (QTL) were reconfirmed as being located on linkage groups B, G, N, 5 and new markers on Oi were detected. Some QTL effects were restricted to specific sampling dates suggesting temporal expression of QTL. QTL for condition factor were limited to linkage group G in both strains. Three suggestive QTL for precocious maturation mapped to similar regions as those for body mass in the Rainbow Springs families while no associations were evident in the Spring Valley families. The results suggest that these regions may play a role in the basis for genetic and phenotypic correlations between body mass and precocious maturation in this species.     

Allometric growth and sperm competition in fishes

The allometric relationship between body mass and gonad mass in males of 23 fish species from 11 families was examined. There was no evidence of a single allometry for all fishes. A cross species analysis suggested a scaling coefficient of 1·04, which was significantly different from a previous study that reported a scaling coefficient of 0·904. A within species analysis generated scaling coefficients from 0·68 to 3·90. Furthermore, for those species characterized by alternative mating tactics, there was no correlation between the scaling coefficients of bourgeois‐type males and parasitic‐type males. These results are discussed in the context of the gonado‐somatic index and its use in testing sperm competition theory.     

Effect of dietary cortisol administration on growth and reproductive success of channel catfish

The effect of cortisol administration on reproductive performance was investigated in channel catfish Ictalurus punctatus broodfish. Cortisol was added to a commercial catfish feed by dissolving in ethanol and spraying the feed to yield a dietary concentration of 150 mg kg⁻¹ feed. The cortisol diet and the control (no cortisol) diet were offered at a rate of 1% of biomass to three replicate ponds each containing 28 female and 14 male broodfish, respectively, three times per week for 11 weeks. Spawning began 10 days after the start of the experiment, and continued for 10 weeks. In fish fed cortisol, body mass and the hepato‐somatic index were reduced ( P  ≤ 0·02) and concentrations of plasma cortisol and glucose were significantly higher ( P  ≤ 0·0003) compared to those of controls. The relative frequency of spawning was similar between the two treatments; however, cortisol‐fed channel catfish had an average of 47·1% more spawns than the control‐fed fish. On average, there were 25·5 spawns per pond in the treated groups compared to 12·3 spawns per control pond ( P  = 0·10). No effect was observed on egg production, with individual egg mass, fecundity, and hatching success being similar ( P  ≥ 0·27) for both treatments. Despite the observed negative effects of cortisol on somatic and hepatic growth, the increased reproductive output coupled with no observable effects on the eggs or hatching success demonstrates that cortisol does not suppress channel catfish reproduction.     

Reproductive behaviour of a temperate serranid fish, Paralabrax clathratus(Girard), from Santa Catalina Island, California, U.S.A.

The reproductive behaviour of the kelp bass Paralabrax clathratus was studied on Santa Catalina Island, California, U.S.A. from April 2000 to September 2002. Adults formed aggregations of three to > 200 individuals, and spawning occurred within subgroups of three to 23 individuals that contained a single female. The gonado‐somatic index ( I _G) of collected ripe males (mean = 5·8%, range = 0·5–13·1%) indicated a large investment in sperm production that is common in group‐spawning fishes characterized by intense sperm competition. Spawning occurred 32 min before sunset to 120 min after sunset, and both males and females were capable of spawning multiple times during a single evening. Behavioural observations of adults and estimates of spawning periodicity from the collection of females with hydrated oocytes suggested that spawning occurred continuously throughout the summer months and showed no significant relationship with the lunar cycle. In general, the spawning behaviour of kelp bass was similar to other functionally gonochoric, group‐spawning serranids. The dynamics of P. clathratus spawning aggregations, however, were inconsistent with that of tropical reef fish spawning aggregations, including the transient spawning aggregations of some tropical serranids. Aggregation spawning appeared to be an important component of the annual reproduction of this species.