Physical and biological factors influencing the spatial distribution of age-0 walleye pollock (Theragra chalcogramma) around the Pribilof Islands, Bering Sea

The waters around the Pribilof Islands, in the southeast Bering Sea, are a main nursery area for age-0 pollock. Each summer, the islands are surrounded by a well-mixed inshore region, separated from a stratified offshore region by a frontal zone. To study the spatial distribution of age-0 pollock around this frontal structure in relation to physical and biological factors that are likely to influence it, such as advection, age-0 pollock feeding, and predation, samples were collected during September of four consecutive years, 1994–97, along two transects. Samples collected included water column hydrography and currents, acoustic backscatter, and groundfish predator density.Our analysis suggested that different mechanisms may be involved in controlling age-0 pollock distribution north and south of the islands. On the shelf area north of the islands, high age-0 pollock density was significantly associated with areas of high potential for growth only in years or portions of the frontal transect in which predator numbers were relatively low, indicating the importance of predation in controlling fish distribution in this area. In contrast, south of the islands, age-0 pollock distribution was associated more with prey availability, which appeared to be determined by vertical spatial overlap between predators and prey. Moreover, south of the islands, the stronger geostrophic currents, typical of the slope region, were more likely to affect the overall standing biomass of juvenile pollock, by constantly advecting fish away from the area.     

Size-dependent, Spatial, and Temporal Variability of Juvenile Walleye Pollock (Theragra chalcogramma) Feeding at a Structural Front in the Southeast Bering Sea

Abstract. The waters surrounding the Pribilof Islands are an important nursery ground for juvenile walleye pollock (Theragra chalcogramma), an important forage fish in the pelagic food web of the productive Bering Sea shelf region. The diet of juvenile pollock was studied in two consecutive years along a transect line crossing from a well‐mixed coastal domain, through a frontal region to stratified water farther offshore. Variability in stomach fullness was high and evidence for increased feeding intensity in the front was weak. Prey diversity and prey size generally increased with increasing fish size, shifting from predominantly small copepods to larger, more evasive prey items such as euphausiids, crab megalopae and fish. The diet of the fish reflected changes in the relative abundance of copepods and euphausiids in the prey fields between years. Juvenile pollock showed increased feeding rates at dusk, and stomach fullness as well as prey condition were generally lowest just before sunrise; however, the proportion of euphausiids increased in the diet of pollock caught at night, suggesting that some food was also ingested during darkness. Juvenile pollock and their euphausiid prey both vertically migrated above the thermocline at night, although each had a different daytime depth.     

The abundance and distribution of euphausiids and zero-age pollock on the inner shelf of the southeast Bering Sea near the Inner Front in 1997–1999

Acoustic data and net samples were collected during late spring and early fall 1997–1999 to assess zooplankton and micronekton abundance and distribution relative to the Inner Front at three sampling grids (Port Moller, Cape Newenham and Nunivak Island) on the inner shelf of the southeast Bering Sea. Epibenthic scattering layers were observed during May–June and August–September in all three years. Acoustic data were scaled to euphausiid biomass using target strength models. Mean euphausiid biomass determined acoustically for each transect line was 0.7–21 g m⁻², with most values below 5 g m⁻². There was no consistent relationship between the distribution and biomass of euphausiids and the location of the Inner Front. Zero age pollock were observed on the inner shelf in August–September during all years, but were confined primarily to the stratified side of the Inner Front and to the frontal regime. The acoustic data for pollock were scaled to biomass using laboratory measurements of gas bladder dimensions and target strength models. Acoustic determinations of mean transect biomass for euphausiids did not differ from literature values for the inner shelf of the southeast Bering Sea, and pollock biomass on the inner shelf did not differ from that around the Pribilof Islands. Despite recent anomalies in climate and oceanographic conditions on the inner shelf, and high mortality of shorttail shearwaters during 1997, we found no evidence of significant interannual differences in the biomass of euphausiids or zero-age pollock on the inner shelf of the southeast Bering Sea.     

Climate change and control of the southeastern Bering Sea pelagic ecosystem

We propose a new hypothesis, the Oscillating Control Hypothesis (OCH), which predicts that pelagic ecosystem function in the southeastern Bering Sea will alternate between primarily bottom-up control in cold regimes and primarily top-down control in warm regimes. The timing of spring primary production is determined predominately by the timing of ice retreat. Late ice retreat (late March or later) leads to an early, ice-associated bloom in cold water (e.g., 1995, 1997, 1999), whereas no ice, or early ice retreat before mid-March, leads to an open-water bloom in May or June in warm water (e.g., 1996, 1998, 2000). Zooplankton populations are not closely coupled to the spring bloom, but are sensitive to water temperature. In years when the spring bloom occurs in cold water, low temperatures limit the production of zooplankton, the survival of larval/juvenile fish, and their recruitment into the populations of species of large piscivorous fish, such as walleye pollock (Theragra chalcogramma), Pacific cod (Gadus macrocephalus) and arrowtooth flounder (Atheresthes stomias). When continued over decadal scales, this will lead to bottom-up limitation and a decreased biomass of piscivorous fish. Alternatively, in periods when the bloom occurs in warm water, zooplankton populations should grow rapidly, providing plentiful prey for larval and juvenile fish. Abundant zooplankton will support strong recruitment of fish and will lead to abundant predatory fish that control forage fish, including, in the case of pollock, their own juveniles. Piscivorous marine birds and pinnipeds may achieve higher production of young and survival in cold regimes, when there is less competition from large piscivorous fish for cold-water forage fish such as capelin (Mallotus villosus). Piscivorous seabirds and pinnipeds also may be expected to have high productivity in periods of transition from cold regimes to warm regimes, when young of large predatory species of fish are numerous enough to provide forage. The OCH predicts that the ability of large predatory fish populations to sustain fishing pressure will vary between warm and cold regimes.The OCH points to the importance of the timing of ice retreat and water temperatures during the spring bloom for the productivity of zooplankton, and the degree and direction of coupling between zooplankton and forage fish. Forage fish (e.g., juvenile pollock, capelin, Pacific herring [Clupea pallasii]) are key prey for adult pollock and other apex predators. In the southeastern Bering Sea, important changes in the biota since the mid-1970s include a marked increase in the biomass of large piscivorous fish and a concurrent decline in the biomass of forage fish, including age-1 walleye pollock, particularly over the southern portion of the shelf. Populations of northern fur seals (Callorhinus ursinus) and seabirds such as kittiwakes (Rissa spp.) at the Pribilof Islands have declined, most probably in response to a diminished prey base. The available evidence suggests that these changes are unlikely the result of a decrease in total annual new primary production, though the possibility of reduced post-bloom production during summer remains. An ecosystem approach to management of the Bering Sea and its fisheries is of great importance if all of the ecosystem components valued by society are to thrive. Cognizance of how climate regimes may alter relationships within this ecosystem will facilitate reaching that goal.     

Variations in age-0 pollock distribution among eastern Bering Sea nursery areas: A comparative study through acoustic indices

Cohort abundance of walleye pollock (Theragra chalcogramma) is subject to strong interannual variation in the eastern Bering Sea, and this variation is known to be determined largely at the age-0 stage. We estimated the spatial distributions and densities of age-0 walleye pollock in five nursery areas around the eastern Bering shelf in three successive years (1997–1999) from acoustic survey data. Concurrently, we calculated estimates of the spatial distribution of euphausiids, a major prey of age-0 walleye pollock, and estimates of spatial overlap of groundfish predators with the age-0 walleye pollock. The analyses showed that all nursery areas had low densities of age-0 walleye pollock in 1997, which ultimately produced the weakest adult year-class. In the intermediate year of 1998, age-0 densities were low to medium, and in 1999, which produced the strongest of the three adult year-classes, all nursery areas had medium to high age-0 walleye pollock densities. Euphausiid distributions had a consistently positive spatial relationship with age-0 walleye pollock. Groundfish predator density ratios were positively related to age-0 walleye pollock density when age-0 walleye pollock were displaced relatively northward. Our results suggest that abundance of age-0 walleye pollock, and hence of adult cohorts in the eastern Bering Sea, can be predictable from a concise set of indicators: the densities of age-0 walleye pollock at nursery areas in mid- to late-summer, their spatial relationship to euphausiids and groundfish predators, and the latitudinal trend of their distributions. The 3 years 1997–1999 had significant differences of physical conditions in the eastern Bering Sea, and represent an advantageous framework for testing these hypotheses.     

Does the zooplankton prey availability limit the larval habitats of pike in the Baltic Sea?

The objectives of this study were to (1) investigate whether the availability of suitable zooplankton prey limits the distribution of the coastal larval areas of pike (Esox lucius) in two archipelago areas of the northern Baltic Sea and (2) compare the availability of zooplankton prey in spring between different types of coastal littoral habitat. According to the results, reed belt habitats formed by Phragmites australis constitute hot spots for zooplankton prey in the coastal ecosystem. During the spring, reed-covered shores of the inner archipelago maintained more than 10 times higher densities of copepods and cladocerans, the preferred prey for larval pike, compared to the other studied shores. Temperature conditions were also most favourable in the reed belt habitat. Thus, the reed belts of the inner and middle archipelago were shown to form the best habitat for larval pike in the coastal area of the northern Baltic Sea, and this was also the only habitat where pike larvae were found. Our results suggest that the poor survival and recruitment of pike in the outer archipelago, however, cannot exclusively be explained by sub-optimal feeding conditions of the larvae. There are also other important factors, presumably connected to the exposure to the open sea, that affect the distribution of the pike larvae. Our results, however, highlight the importance of sheltered coastal reed belt shores as reproduction habitat for spring-spawning fish in the northern Baltic Sea. Further, this study disproves the assumption that the seaweed bladder wrack (Fucus vesiculosus) forms a reproduction habitat for pike in the coastal area.     

Variations in crab claw morphology and diet across contrasting inter‐tidal habitats

Crabs are important predators of inter‐tidal ecosystems, controlling the abundance and distribution of their prey populations. Often the same crab species occupies several habitats and, although their effects on prey have been quantified across habitats, crabs’ dietary and morphological responses to differing environmental influences have been overlooked. Here, we used the crabs Eriphia verrucosa and Pachygrapsus marmoratus as model species to examine differences in claw morphometry – size and wear – and diet between rocky shore and heterogeneous sand flat habitats. We predicted that, intra‐specifically, crabs from rocky shores would consume more hard‐shelled prey owing to their high availability and consequently, would display chelipeds with the following claw characteristics: a higher degree of claw damage, stronger musculature (higher propel height) and increased mechanical advantage (defined as the ratio of input lever length to output lever length) than crabs in the heterogeneous sand flat habitats. Sampling was performed in heterogeneous sand flat habitats and rocky shores of the Central Portuguese coast. For each crab species, carapace width, diet composition and several claw morphometric measures were recorded, revealing significant intra‐specific differences (using multivariate analysis) between shore types. We found that E. verrucosa and P. marmoratus consumed more hard prey on rocky shore than on sand flat habitats, which resulted in rocky shore crabs having more accentuated dentition wear and larger musculature than their sand flat habitat counterparts. We suggest that the strong response of crab claw morphometry to environmentally induced diet variations is an important mechanism in the successful adaptation of crab species to inhabit differing habitats. A major implication is that the impact of the same species on prey may vary largely with habitat type as a result of predation efficiency varying with claw condition.     

Modeled connectivity between northern rock sole (Lepidopsetta polyxystra) spawning and nursery areas in the eastern Bering Sea

Connectivity between spawning and potential nursery areas of northern rock sole, Lepidopsetta polyxystra, in the eastern Bering Sea was examined using an individual-based biophysical-coupled model. Presumed spawning areas were identified using historical field-collected ichthyoplankton data, and nursery habitats were characterized based on previously described settlement areas. Simulated larvae were released from spawning areas near the Pribilof Islands, south of the Pribilof Islands along the outer continental shelf, on the north side of the Alaska Peninsula, and in the Gulf of Alaska south of Unimak Island. Simulated larvae were transported along two general pathways: 1) northwards along the outer continental shelf from Unimak Island towards the Pribilof Islands and further north offshore of mainland Alaska, and 2) eastward along the Alaska Peninsula. At the end of the 2-month simulation, drift pathways placed pre-settlement stage larvae offshore of known nursery areas of older juveniles near mainland Alaska, consistent with a hypothesis that initial settlement may be followed by substantial post-settlement redistribution.     

Bottom-up and top-down controls of walleye pollock (Theragra chalcogramma) on the Eastern Bering Sea shelf

Control of walleye pollock (Theragra chalcogramma) recruitment in the Eastern Bering Sea involves complex interactions between bottom-up and top-down processes, although the mechanisms are poorly understood. We used statistical models to test the leading hypotheses linking recruitment variability to biotic and abiotic factors. Consistent with a “cold-pool hypothesis”, recruitment of pollock was significantly stronger if winters preceding the larval (age-0) and juvenile stages (age-1) were mild. However, our results did not support the proposed top-down mechanism (cannibalism) underlying this hypothesis. Several empirical relationships support an “oscillating control hypothesis”. As predicted by it, the effect of ice conditions on survival during the larval and early juvenile stages was modified by the abundance of adult pollock, implying stronger bottom-up control when adult abundance (hence cannibalism) was low. The proposed bottom-up mechanism predicts that the survival of pelagic-feeding walleye pollock (benthic-feeding yellowfin sole), should be higher during years with an early (late) ice retreat, which was confirmed by our analysis. Our results also provide additional evidence for a “larval transport hypothesis”, which states that cannibalism of larval and juvenile pollock is reduced in years when strong northward advection separates juveniles from cannibalistic adults.In addition to testing existing hypotheses, we identified new relationships between spawner-to-recruit survival rates of walleye pollock and several indicators of mixed layer dynamics during the spring and summer. Survival rates and recruitment were significantly reduced when larval or early juvenile stages experienced a delay in the (non-ice-associated) spring bloom as a result of stormy spring conditions, suggesting that the timing of the spring bloom is critical to both first-feeding larvae and age-1 juveniles. Furthermore, a dome-shaped relationship between pollock survival and summer wind mixing at the early juvenile stage is consistent with modeling and laboratory studies showing an increase in survival at low to moderate levels of wind mixing, but a decrease in feeding success at high levels of wind mixing.Top-down controls also regulate recruitment of walleye pollock. At least one-third of the variability in spawner-to-recruit survival could be accounted for by predation mortality at the early juvenile stage (age-1). Predation of juvenile pollock can be attributed largely to cannibalism, which varies with the abundance of adult pollock and with the availability of juveniles to adult predators. A simple index reflecting the spatial overlap between juvenile and adult pollock explained 30–50% of the overall variability in recruitment, similar to the variability explained by the best environmental predictors. Although environmental effects are difficult to separate from the effects of predation, we conclude that bottom-up and top-down processes are equally important in controlling the survival of pollock from spawning to recruitment at age 2. However, the magnitude of top-down control is itself modified by environmental factors that control the availability of juvenile pollock to adults (through impacts on spatial distribution) and the abundance of adult predators (through effects on productivity and carrying capacity).     

Habitat use patterns of the invasive red lionfish Pterois volitans: a comparison between mangrove and reef systems in San Salvador,Bahamas

The Indo‐Pacific red lionfish Pterois volitans is widespread both in its native and its non‐native habitats. The rapid invasion of this top predator has had a marked negative effect on fish populations in the Western Atlantic and the Caribbean. It is now well documented that lionfish are invading many tropical and sub‐tropical habitats. However, there are fewer data available on the change in lionfish abundance over time and the variation of body size and diet across habitats. A recent study in San Salvador, Bahamas, found body size differences between individuals from mangrove and reef systems. That study further suggested that ontogenetic investigation of habitat use patterns could help clarify whether lionfish are using the mangrove areas of San Salvador as nurseries. The aim of the present study is to determine temporal trends in lionfish relative abundance in mangrove and reef systems in San Salvador, and to further assess whether there is evidence suggesting an ontogenetic shift from mangroves to reef areas. Accordingly, we collected lionfish from mangrove and reef habitats and calculated catch per unit effort (a proxy for relative abundance), compared body size distributions across these two systems, and employed a combination of stable isotope, stomach content, and genetic analyses of prey, to evaluate differences in lionfish trophic interactions and habitat use patterns. Our results show that populations may have increased in San Salvador during the last 4 years, and that there is a strong similarity in body size between habitats, stark differences in prey items, and no apparent overlap in the use of habitat and/or food resources. These results suggest that there is not evidence an for ontogenetic shift from mangroves to reefs, and support other studies that propose lionfish are opportunistic forages with little movement across habitats.     

The effect of oceanographic variability and interspecific competition on juvenile pollock (Theragra chalcogramma) and capelin (Mallotus villosus) distributions on the Gulf of Alaska shelf

Results from this study suggest that small-scale variability in the Alaska Coastal Current (ACC) and competition between juvenile pollock and capelin are potential mechanisms affecting the distribution and abundance of fishes in the Gulf of Alaska (GOA). Fish distributions in Barnabus Trough, off the east coast of Kodiak Island, were assessed using acoustic data collected with a calibrated echosounder during August–September 2002 and 2004. Trawl hauls were conducted to determine the species composition of the fish making up the acoustic backscatter. Oceanographic data were collected from moorings, conductivity–temperature–depth (CTD) probes, trawl-mounted microbathythermographs (MBT) and expendable bathythermographs (XBT). National Centers for Environmental Prediction (NCEP) reanalysis data were used to assess area winds, and information on regional transport was derived from current meters deployed on moorings north and south of Kodiak Island. The distribution of water-mass properties and fish during 2002 showed variability at the temporal scale of weeks. Juvenile pollock (age-1 and age-2) were initially most abundant in warm, low-salinity water on the inner shelf, whereas capelin were distributed primarily on the outer shelf in cool, high-salinity waters. During a 2-week period juvenile pollock distribution expanded with the offshore expansion of warm, low-salinity water, and capelin abundance in outer-shelf waters decreased. We hypothesize that wind-driven pulsing of the ACC resulted in increased transport of warm, low-salinity water through the study area. In 2004, warm, low-salinity water characterized the inner shelf and cool, high-salinity water was found on the outer shelf. However, the distribution of water-mass properties did not show the weekly scale variability observed in 2002. Area winds were consistently toward the southwest during 2004, such that we would not expect to see the wind-driven pulsing of ACC water that occurred in 2002. Age-1 and age-2 pollock were not observed in Barnabus Trough in 2004. Instead, the midwater acoustic backscatter was composed of capelin mixed with age-0 pollock, and these capelin were not restricted to the outer-shelf waters, but were found primarily in warm, low-salinity inner-shelf waters that had been previously occupied exclusively by age-1 and age-2 pollock. We suggest that this is consistent with inner-shelf waters being preferred foraging habitat for juvenile pollock and capelin. Further study of the mechanisms linking climate change with variability in the ACC is needed, as are studies of the potential for competition between juvenile pollock and capelin.     

Effects of habitat complexity on cannibalism rates in European green crabs (Carcinus maenas Linnaeus, 1758)

The habitat in which predator–prey interactions take place may have a profound influence on the outcome of those interactions. Cannibalism is an intriguing form of predation whereby foraging by predators may contribute to the regulation of their own populations.This is particularly interesting in the case of invasive species, like the widely distributed European green crab (Carcinus maenas). This study explores how habitat complexity influences cannibalism rates in green crab populations of Prince Edward Island, Atlantic Canada. Both laboratory and field experiments were conducted to measure feeding rates by individual adult green crabs on a standard number of smaller conspecifics. In the laboratory, experimental treatments mimicked unstructured to increasingly structured habitats: water, sandy bottom, oyster shells, mussel shells, oyster shells with sandy bottom and mussel shells with sandy bottom. In those trials, adult green crabs consumed several times more juveniles on unstructured habitats than on the most structured ones, with a gradual decrease in predation rates across increasingly complex habitats. Field inclusion experiments used the same approach and were conducted in sandy bottoms, sandy bottoms with a layer of oyster shells and sandy bottoms with a layer of mussel shells. These trials showed similar patterns of decreasing feeding rates across increasingly complex habitats, but differences among treatments were not significant. These results support the idea that complex habitats have the potential to mediate predator–prey interactions, including adult–juvenile cannibalism in green crabs.     

Dietary Studies on the Predatory Fishes of the Norman River Estuary, with Particular Reference to Penaeid Prawns

The predatory fish community and their prey in the Norman Rivere estuary, Gulf of Carpentaria Australia, are compared with the communities of other tropical inshore areas, to investigate patterns of predation in tropical inshore areas particularly in relation to penaeid prawns. Abiotic factors (turbidity, freshwater input) and diversity of habitat types affect the composition of both prey and predator communities, resulting in large differences in the communities of tropical inshore waters. The stomach contents of 2059 predatory fish from the Norman River estuary were analysed over four sampling trips in the wet and dry seasons. The stomachs of 61% (1255 fish) of 54 species contained a total of 676·2 g (dry weight) of food, while 39% (804 fish) were empty. Teleosts were the main component of the diet (by dry weight) of 13 of the 22 species analysed, followed by annelids for two species. Five species had only teleosts in their stomachs. Most predator species ate benthic or bentho-pelagic prey, while three species—Rhizoprionodon taylori, Scomberoides commersonianusandLeptobrama mulleri—ate mainly pelagic prey. Although 19 species ate some penaeids, onlyPolydactylus sheridaniate little else. This species,Lates calcariferandEleutheronema tetradactylumate 94·5% of all the penaeid prey and 97·9% of all the commercially important penaeid prey recorded in the study. Penaeid predation indices (calculated from gillnet catch rates, proportion of penaeids in the diet and a consumption rate of 3% body weight per day) were 0·23 g of penaeid per net-metre per day forP. sheridani, 0·15 forL. calcariferand 0·03 forE. tetradactylum. Commercially important penaeid predation indices were 0·11, 0·13 and 0·01, respectively. These values are intermediate between those previously recorded for the main penaeid predators in other inshore areas of the Gulf of Carpentaria (Embley River estuary and Groote Eylandt).     

南沙群岛西南大陆斜坡海域浮游动物的垂直分布

浮游动物是海洋生态系统物质和能量转移的关键环节,也是渔业资源的重要饵料,其种类组成和数量分布均有明显的垂直分层现象。利用2011年4月在南沙群岛西南大陆斜坡海域开展的18个站点90份样品的调查数据,对该海域浮游动物的垂直分布进行了研究,表明:(1)该海域浮游动物种类组成丰富,共出现18个类群580种(类);(2)浮游动物种类组成垂直变化明显,特定水层出现的种类数占总种数的43.6%,各水层均出现的种类仅占总种数的15.8%;(3)优势种组成复杂,垂直变化明显,单一种类的优势度不高;(4)浮游动物平均密度和湿重生物量分别为206.27ind/m3、94.03mg/m3,密度和生物量均以0~2m层和30~75m层较高,沿水深梯度的变化呈明显的双峰型;(5)浮游动物数量的垂直变化主要受温跃层影响,温跃层内浮游动物数量最高,温跃层上方和下方的水层内数量较低;(6)南沙西南大陆斜坡区浮游动物生产力水平较高,表明该海域渔业资源有一定的开发潜力。     

Spatial variability of the surf zone fish and macroinvertebrate community within dissipative sandy beaches in Oregon,USA

Habitat heterogeneity can influence biological communities by providing a diversity of areas that can be occupied by different species. Sandy beach surf zones are often considered homogenous environments; however, sand bars moved by currents and waves can produce trench‐like shapes or troughs that provide heterogeneity. The influence of habitat heterogeneity produced by sand movement is unclear despite the fact that surf zones are an important habitat for larval and juvenile fish and macrocrustaceans. To determine if, and how, the fish and macroinvertebrate communities present in trough and non‐trough or flat areas of Oregon surf zones differ, we compared species assemblages in both areas at three beaches adjacent to estuary mouths over 2 years. Troughs had different communities compared with flat areas, with higher total catch (mean ± SD = 123.2 ± 122.1 versus 43.6 ± 44.5 individuals × 100 m^?2) and taxon richness (6.7 ± 2.7 versus 4.0 ± 2.3 taxa); these differences were potentially due to water movement, prey availability and sediment size. The fish and macroinvertebrate communities did not vary between years but there were significant differences among beaches, with the most distinct community present at the only beach adjacent to an estuary without a jetty at its mouth, which was possibly due to higher species movement between the surf zones and estuary. Fish and macrocrustacean surf zone communities varied spatially within and among beaches in relation to habitat heterogeneity provided by sand movement and, potentially, the influence of adjacent habitats.     

Fishery benefits from exploiting spawning aggregations not solely dependent on enhanced fish density

The vulnerability of spawning aggregations to exploitation varies among fisheries as a result of differences in the population-density changes associated with this behaviour. However, vulnerability to fishing is also influenced by technology, environmental factors, and fish and fisher behaviours. Focusing on a fishery for the rabbitfish Siganus sutor at Praslin Island, Seychelles, we examined how catch rate varied across spawning and non-spawning habitats in relation to in situ population-density changes and other factors known to influence catchability. Catch rates in spawning habitat were disproportionate to density changes, being only fourfold greater than catch rates in non-spawning habitat, despite the fact that spawning-aggregation formation involved nine- to thirteen-fold increases in population density. Catch rates in spawning habitat were also highly variable across the spawning season (0–23.4 fish trap-hour^?1). Current strength was of similar importance to density as a catch-rate predictor, with the highest catch rates in spawning habitats confined to months with the strongest currents. Therefore, in addition to density-dependent catchability, other factors that influence catch rates must be examined to avoid overestimation of the vulnerability of populations to aggregation fishing. The dynamics of catchability at spawning sites can limit the ability of fishers to predict and maximise returns based on increases in fish density.     

Predominance of parallel‐ and cross‐predation in anglerfish

Several studies in the last 20 years have revealed that morphological asymmetry in fish can be characterized as ‘antisymmetry’. Antisymmetry is a lateral dimorphism in which each population consists of individuals with well‐developed left sides (lefties) and well‐developed right sides (righties). This dimorphism influences predator–prey interactions. In some piscivorous fishes, it has been found that predators can catch more prey of the opposite morphological type to themselves (cross‐predation) than of the same morphological type (parallel‐predation). Our previous work clarified that the predominance of cross‐predation is caused by lateralized behaviors of predators and prey that correspond to their morphological antisymmetry. Moreover, based on the results of our behavioral observations, we hypothesized that parallel‐predation can predominate when predators encounter the potential prey frontally. To test this hypothesis, in the present study we investigated the relationship between lateral morphological types of anglerfish (Lophiomus setigerus) and those of the prey fishes found in their stomachs. Anglerfish attract potential prey using their first dorsal fin (illicium) as a lure, and their frontal encounters with potential prey fishes were photographed in situ and observed in an aquarium. The results of a stomach contents analysis indicated that parallel‐predation predominated in five benthopelagic prey fish species (perches and eels). By contrast, five benthic prey fishes (gobies and weevers) exhibited the predominance of cross‐predation. These results not only demonstrate the predominance of parallel‐predation in a natural fish community, but also suggest that the relationship between morphological types of predator and prey species can be reversed depending on the lifestyle of prey.     

Parallel structure among environmental gradients and three trophic levels in a subarctic estuary

We assessed spatial and temporal variability in the physical environment of a subarctic estuary, and examined concurrent patterns of chlorophyll α abundance (fluorescence), and zooplankton and forage fish community structure. Surveys were conducted in lower Cook Inlet, Alaska, during late July and early August from 1997 through 1999. Principle components analysis (PCA) revealed that spatial heterogeneity in the physical oceanographic environment of lower Cook Inlet could be modeled as three marine-estuarine gradients characterized by temperature, salinity, bottom depth, and turbidity. The gradients persisted from 1997 through 1999, and PCA explained 68% to 92% of the variance in physical oceanography for each gradient-year combination. Correlations between chlorophyll α abundance and distribution and the PCA axes were weak. Chlorophyll was reduced by turbidity, and low levels occurred in areas with high levels of suspended sediments. Detrended correspondence analysis (DCA) was used to order the sample sites based on species composition and to order the zooplankton and forage fish taxa based on similarities among sample sites for each gradient-year. Correlations between the structure of the physical environment (PCA axis 1) and zooplankton community structure (DCA axis 1) were strong (r = 0.43-0.86) in all years for the three marine-estuarine gradients, suggesting that zooplankton community composition was structured by the physical environment. The physical environment (PCA) and forage fish community structure (DCA) were weakly correlated in all years along Gradient 2, defined by halocline intensity and surface temperature and salinity, even though these physical variables were more important for defining zooplankton habitats. However, the physical environment (PCA) and forage fish community structure (DCA) were strongly correlated along the primary marine-estuarine gradient (#1) in 1997 (r = 0.87) and 1998 (r = 0.82). The correlation was poor (r = 0.32) in 1999, when fish community structure changed markedly in lower Cook Inlet. Capelin (Mallotus villosus), walleye pollock (Theragra chalcogramma), and arrowtooth flounder (Atheresthes stomias) were caught farther north than in previous years. Waters were significantly colder and more saline in 1999, a La Niña year, than in other years of the study. Interannual fluctuations in environmental conditions in lower Cook Inlet did not have substantial effects on zooplankton community structure, although abundance of individual taxa varied significantly. The abundance and distribution of chlorophyll α, zooplankton and forage fish were affected much more by spatial variability in physical oceanography than by interannual variability. Our examination of physical-biological linkages in lower Cook Inlet supports the concept of “bottom-up control,” i.e., that variability in the physical environment structures higher trophic-level communities by influencing their distribution and abundance across space.     

Within-reach spatial variability of snails and molluscivory by brown trout

Linking habitat distributions of prey to the probability of predation is important to understanding consumptive effects of predators on prey populations. This study reports how within-reach spatial variability of two snails, the hydrobiid Potamopyrgus antipodarum and the physid Physella acuta, was linked to habitat-based predation risk by young brown trout (Salmo trutta) of different age classes. Potamopyrgus is endemic to New Zealand streams and lakes, where it commonly co-exists with the invader P. acuta, but both snails are worldwide invaders to many freshwater systems. Examination of egested snails revealed Potamopyrgus and Physella were consumed in similar numbers within age classes. However, 10-month-old trout consumed, on average, fewer snails than 20-month-old trout, and 8-month-old trout ate essentially no snails, suggesting snails were a more important prey item for larger age-1 fish than smaller age-0 fish. No Physella were egested alive by any trout age class. However, 38% and 16% of the Potamopyrgus consumed were egested alive by 10- and 20-month-old trout, respectively, with some passing live after ~70 h in digestive tracts. Physella and the spiny-shell form of Potamopyrgus were significantly denser on macrophytes than on stony sediments in midchannel, and these habitat distributions affected their odds of consumption. Risk of consumption by trout was ~10 times greater for Physella than Potamopyrgus on stones, but their risk was similar in protective macrophytes. Odds of consumption were similar for spiny and smooth shell forms of Potamopyrgus on stones, suggesting spines do not provide protection from large predators like trout. My results suggest that brown trout can potentially exert stronger population regulatory effects on Physella than on Potamopyrgus and that these effects are partly mediated by macrophyte cover.     

Characterising and comparing the spawning habitats of anchovy Engraulis encrasicolus and sardine Sardinops sagax in the southern Benguela upwelling ecosystem

The spawning habitats of anchovy Engraulis encrasicolus and sardine Sardinops sagax in the southern Benguela upwelling ecosystem were characterised by comparing their egg abundances with environmental variables measured concomitantly during two different survey programmes: the South African Sardine and Anchovy Recruitment Programme (SARP), which comprised monthly surveys conducted during the austral summers of 1993/94 and 1994/95; and annual pelagic spawner biomass surveys conducted in early summer (November/December) from 1984 to 1999. Eggs were collected using a CalVET net. Physical variables measured included sea surface temperature (SST), surface salinity, water depth, mixed-layer depth, and current and wind speeds; biological variables measured included phytoplankton biomass, and zooplankton biomass and production. Spawning habitat was identified by construction of quotient curves derived from egg abundance data and individual environmental variables, and relationships between these variables were determined using multivariate co-inertia analysis. SARP data showed that anchovy spawning was associated with cool water and moderate wind and current speeds, whereas sardine spawning was related to warmer water and more turbulent and unstable conditions (i.e. high wind speeds and strong currents) than for anchovy. SARP data also showed significant differences in selection of spawning habitat of the two species for all environmental variables. The relationship between anchovy egg abundance and salinity was strongly positive, but strongly negative with water depth, phytoplankton biomass and zooplankton production. Sardine egg abundance was strongly positively related to current speed. The spawner biomass survey data demonstrated that the spawning habitat of anchovy was characterised by warm water and high salinity, whereas sardine spawning was associated with cool water and low salinity. The survey data showed significant differences in spawning habitat selection by anchovy and sardine for SST, salinity and zooplankton biomass, but not for the other environmental variables. There was a positive relationship between anchovy egg abundance and SST, salinity and mixed-layer depth, and a negative relationship with water depth, phytoplankton biomass and zooplankton production. For sardine there was a strong positive relationship between egg abundance and current speed and wind speed. Differences in the results between the two survey programmes could be attributable to differences in their spatio-temporal coverage. Spawning habitats of anchovy and sardine appear to be substantially different, with anchovy being more specific than sardine in their preference of various environmental conditions.