Foraging ecology and movement patterns of jumbo squid (Dosidicus gigas) in the California Current System

From 2002 to 2010, the jumbo squid (Dosidicus gigas) has been regularly encountered in large numbers throughout the California Current System (CCS). This species, usually found in subtropical waters, could affect coastal pelagic ecosystems and fisheries as both predator and prey. Neither the abundance of jumbo squid nor the optimal ocean conditions in which they flourish are well known. To understand better the potential impacts of this species on both commercial fisheries and on food-web structure we collected nearly 900 specimens from waters of the CCS, covering over 20° of latitude, over a range of depths and seasons. We used demographic information (size, sex, and maturity state) and analyzed stomach contents using morphological and molecular methods to best understand the foraging ecology of this species in different habitats of the CCS. Squid were found to consume a broad array of prey. Prey in offshore waters generally reflected the forage base reported in previous studies (mainly mesopelagic fishes and squids), whereas in more coastal waters (shelf, shelf break and slope habitats) squid foraged on a much broader mix that included substantial numbers of coastal pelagic fishes (Pacific herring and northern anchovy, as well as osmerids and salmonids in northern waters) and groundfish (Pacific hake, several species of rockfish and flatfish). We propose a seasonal movement pattern, based on size and maturity distributions along with qualitative patterns of presence or absence, and discuss the relevance of both the movement and distribution of jumbo squid over space and time. We find that jumbo squid are a generalist predator, which feeds primarily on small, pelagic or mesopelagic micronekton but also on larger fishes when they are available. We also conclude that interactions with and potential impacts on ecosystems likely vary over space and time, in response to both seasonal movement patterns and highly variable year-to-year abundance of the squid themselves.     

Abundance and distribution of pelagic piscivorous fishes in the Columbia River plume during spring/early summer 1998-2003: Relationship to oceanographic conditions, forage fishes, and juvenile salmonids

From 1998 to 2003, we observed large fluctuations in the abundance and distribution of four pelagic predatory (piscivorous) fishes off northern Oregon and southern Washington, USA. Fluctuations in predatory fish species composition and abundance were strongly linked to the date of the spring transition and to ocean temperatures. Predatory fishes, forage fishes, and juvenile salmonids had distinct spatial distributions, with predators distributed primarily offshore and forage fish and salmonids onshore, but this varied depending on ocean conditions. We suggest that predatory and forage fish distributions respond to ocean temperatures, predator/prey interactions, and possibly turbidity. A shift in ocean conditions in 1999 decreased overall predator fish abundance in the Columbia River plume, particularly for Pacific hake. Marine survival of juvenile salmon started to increase in 1999, and forage fish densities increased in 2000, lagging by one year.     

Feeding of hake and a first attempt at determining their trophic role in the South African west coast marine environment

During intensive biannual demersal fish-sampling cruises since January 1983 over the South African west coast shelf region (Orange River to Cape Agulhas, coast to 500-m isobath), feeding studies on both species of Cape hake (Merluccius capensis and M. paradoxus) were undertaken. These studies were of two forms, random sampling of the dietary preference and constituents of the hake caught during daylight only and intensive sampling of stomachs with food throughout a 28-hour cycle. From the index of relative importance, crustaceans were shown to be most important in the majority of length classes of hake studied, with mesopelagics (fish and cephalopods) and other hake also important. Differences by hake species, by predator size, by geographical location and by season were evident. The Cape hakes, which constitute some three-quarters of the demersal fish biomass in the area, are deduced to play a major role in the marine faunal environment, both as predators and prey, and some base-line figures of the quantities of different food items consumed by the resource are presented. Hake are largely opportunistic feeders and it is concluded that they can adapt to perturbations in the availability of prey, whether they be triggered by environmental cause or by, for example, purse-seining of particularly mesopelagic fish.     

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.     

The importance of cephalopods as prey for hake and other groundfish in South African waters

Literature on trophic relationships in the Benguela ecosystem has stressed the importance of cephalopods as prey of groundfish. The groundfish community of the shelf and upper slope of southern Africa is dominated by the Cape hakes, and the results presented (1984–1991) confirm that both species of hake are important predators of cephalopods, especially taking into consideration the abundance of hake in the ecosystem. However, geographic, seasonal and species variability are evident in the patterns observed. The main prey species are Sepia spp. (predominantly Sepia australis), Loligo vulgaris reynaudii, Todaropsis eblanae and Lycoteuthis ?diadema. The last-named is an important food organism for fish. Its systematic status needs revision, however. Qualitative results of studies of cephalopod predation are also provided for kingklip and monkfish.     

The feeding habits of slope dwelling macrourid fishes in the eastern North Pacific

The diet of slope dwelling macrourid fishes in the eastern North Pacific is poorly known. We collected several hundred stomach samples to investigate the feeding habits of Coryphaenoides acrolepis and Albatrossia pectoralis, the two dominant slope dwelling macrourids off the continental United States. Coryphaenoides acrolepis exhibited a pronounced ontogenetic shift in diet. Specimens <15 cm pre-anal fin length (PAF) consumed primarily polychaetes, amphipods, cumaceans and mysids, while larger individuals consumed increasingly larger, more pelagic prey such as fish, squid, and large crustaceans. Scavenging was also very important to specimens >15 cm with scavenged food constituting approximately 20% of the weight of total prey and occurring in approximately 20% of fish 21–29 cm. Albatrossia pectoralis consumed primarily midwater fish and squid, and we believe that it feeds in the water column. There were significant differences between the diets of A. pectoralis and C. acrolepis suggesting some degree of niche separation between macrourid species on the continental slope of the eastern North Pacific. Both species are at the top of the food web on the upper continental slope and, because of their abundance, may exert significant pressures on their prey populations.     

秋季黄海中南部鱼类群落对饵料生物的摄食量

研究鱼类与饵料生物之间食物定量关系进而为多鱼种资源评估提供依据,2000~2002年秋季(10~11月)在黄海中南部海域进行了定点底拖网调查,应用Eggers模型,计算了带鱼(Trichiurus lepturus)、小黄鱼(Pseudosciaena polyactis)、黄(Lophius litulon)、细纹狮子鱼(Liparis tanakae)等23种鱼类在秋季对饵料生物的摄食量。结果表明:黄海中南部23种鱼类在秋季对饵料生物的总摄食量约为309万t,其中,鱼(Engraulis japonicus)的摄食量最高(在250万t以上),占总摄食量的80.9%。中上层和底层鱼类对饵料生物的摄食量分别为262万t和47万t左右,占总摄食量的84.7%和15.3%,鱼和细纹狮子鱼分别是中上层和底层鱼类中最主要的捕食者。磷虾类是中上层鱼类最主要的食物来源,其次是桡足类、端足类和毛颚类;虾类和鱼类是底层鱼类最主要的食物来源,其次是磷虾类。太平洋磷虾(Euphausia pacifica)、中华哲水蚤(Calanus sinicus)、细长脚虫戎(Themisto gracilipes)、脊腹褐虾(Crangon affinis)和鱼同时是黄海中南部被摄食量最高的5种饵料生物,它们被摄食的生物量之和约为233万t,占总摄食量的75.5%。     

Characterisation of the dietary relationships of two sympatric hake species,Merluccius capensis and M. paradoxus,in the northern Benguela region using fatty acid profiles

The two sympatric species of Cape hake, Merluccius capensis and M. paradoxus, have been the main targets of bottom-trawl fisheries off Namibia for several decades. The feeding ecology of these hakes has been studied mainly using stomach content analyses and thus there remain some gaps in our knowledge about food assimilated over the longer term. In this study, we used fatty acid (FA) profiles to characterise the dietary relationships of M. capensis and M. paradoxus. Muscle samples from hake (n=110) and their known prey (n=68) were collected during trawl surveys off Namibia during 2011. Significant differences between the neutral FA profiles of the hake populations were detected in December 2011 but not in January 2011, an indication of temporal variations in diet and resource partitioning. Comparisons of the neutral FAs in hake and the total FAs of potential prey showed no clear trophic connections, with the exception of flying squid Todarodes sagittatus, which had FA profiles very similar to those of M. paradoxus in December 2011. Our results highlight the complex and temporally shifting relationships that exist between hake and the large pool of prey available to them, and between the two hake species that overlap in their feeding habits and distribution within the highly productive Benguela Current region.     

Temporal variability in predator presence around a fin fish farm in the Northwestern Mediterranean Sea

Recently, aquaculture has generated worldwide interest as a result of the overexploitation of wild stocks combined with a growing international demand for fish and seafood products. Wild fish attracted to the marine fish farms, together with the presence of the farmed fish, are powerful attractants to predators that normally feed on similar or identical fish stocks in nature. This 9‐year study describes for the first time in Mediterranean waters the temporal variability of mammalian and avian predators in a coastal fin fish farm. In all, 99 months (1062 days during 36 consecutive seasons) were spent in the field. By examining the results of this study, it is clear that species as seagulls, shags, bottlenose dolphins and grey herons (considered to cause economic loss in aquaculture owing to direct predation) interact regularly with the fish farm. Although bottlenose dolphins and grey herons were not the most important of all predator species, predatory interactions with the fish farm occurred with what seems to be increasing regularity. Another result observed is the possible bottlenose dolphins’ attraction caused by the harvesting operations in the fish farm. The fish farm offers an alternative food source for predators; hunting at fish farms usually requires less effort on the part of the predator, and becomes a more attractive option than hunting wild fish over wide ranges. During the period of this study, individually identified dolphins feeding were regularly observed feeding on discarded fish from fish farm workers during harvesting operations, supporting the possibility that some individuals are habituated to this food supply. Based on the evidence presented in this paper, it is recommended that strategies for the management of both the aquaculture industry and marine mammal populations should take the results of this study into consideration.     

Evaluating impacts of pulse fishing on the effectiveness of seasonal closure

Seasonal fishing closures are often used in fisheries management to conserve overfished stocks.As one of the unintended consequences,fishermen often contend for maximizing catches immediately after reopening fisheries.The resultant large catch landings in a short time period(i.e.,pulse fishing)may undermine the benefit of closure.We implemented an end-to-end model OSMOSE-JZB(Object-oriented Simulator of Marine ec OSystem Exploitation OSMOSE)modelling ecosystem in the Jiaozhou Bay located in China to evaluate the impact of pulse fishing on the effectiveness of seasonal closure at levels of fish community,population,and individual.Our study demonstrated that the three-month closure was successful in conserving fish stocks.There were small variations on ecological indicators(i.e.,total biomass of the community,mean trophic level of the community,mean trophic level of the catch,and Shannon-Wiener biodiversity index)when pulse fishing occurred.Pulse fishing seemed not to result in a great shift in community structure.Compared to other species,the biomass of two large predatory fishes were more susceptible to pulse fishing.Pulse fishing could change the pressure of predators to fish stocks via food webs,especially for young individuals.Our simulations indicate that we can improve the effectiveness of seasonal closure by managing pulse fishing.Although the results derived in this study may be specific to the target ecosystem,the general approach is applicable to other ecosystems when evaluating fishing impacts.     

Changes in the northern Gulf of St. Lawrence ecosystem estimated by inverse modelling: Evidence of a fishery-induced regime shift?

Mass-balance models have been constructed using inverse methodology for the northern Gulf of St. Lawrence for the mid-1980s, the mid-1990s, and the early 2000s to describe ecosystem structure, trophic group interactions, and the effects of fishing and predation on the ecosystem for each time period. Our analyses indicate that the ecosystem structure shifted dramatically from one previously dominated by demersal (cod, redfish) and small-bodied forage (e.g., capelin, mackerel, herring, shrimp) species to one now dominated by small-bodied forage species. Overfishing removed a functional group in the late 1980s, large piscivorous fish (primarily cod and redfish), which has not recovered 14 years after the cessation of heavy fishing. This has left only marine mammals as top predators during the mid-1990s, and marine mammals and small Greenland halibut during the early 2000s. Predation by marine mammals on fish increased from the mid-1980s to the early 2000s while predation by large fish on fish decreased. Capelin and shrimp, the main prey in each period, showed an increase in biomass over the three periods. A switch in the main predators of capelin from cod to marine mammals occurred, while Greenland halibut progressively replaced cod as shrimp predators. Overfishing influenced community structure directly through preferential removal of larger-bodied fishes and indirectly through predation release because larger-bodied fishes exerted top-down control upon other community species or competed with other species for the same prey. Our modelling estimates showed that a change in predation structure or flows at the top of the trophic system led to changes in predation at all lower trophic levels in the northern Gulf of St. Lawrence. These changes represent a case of fishery-induced regime shift.     

Strictly for the birds?: On ecosystem services of forage fish

Small pelagic fish like sardine, anchovy and herring feed on zooplankton and are in turn prey for fish higher up in the food chain. They are therefore expected to play a vital role in transfer of energy between levels in the food chain. Some stocks of small pelagics are extremely large and subject to very substantial fluctuations caused by environmental factors. They are also very vulnerable to exploitation due to schooling behavior and highly efficient fishing technology. Several such stocks crashed in the 20th century as a result of heavy exploitation and/or adverse environmental changes. The effect of four such stock crashes on other fisheries are reviewed and found to be limited or nonexistent. This puts into perspective a recent report from the Lenfest Foundation, which examines small pelagics and their role in the ecosystem and finds that certain sea bird populations have been severely affected by exploitation of small pelagics. The report recommends more conservative management of small pelagics to limit the effects on predators, but makes no effort to weigh this against the contribution of forage fish to food production.     

水生生态系统中的捕食与生物多样性

捕食在水生生态系统中有重要的作用，是影响生物多样性的主要因素子之一。选择性的捕食中能使被喜食的猎物种类从水体中消失，使生物多样性降低。这种捕食也可能抑制优势种的发展，占免竞争排斥，提高猎生物多样性。     

Food and population variability in five regions supporting large stocks of anchovy,sardine and horse mackerel

The Benguela, California, Humboldt and Canary currents and the coastal waters of Japan support large stocks of sardine or pilchard Sardinops spp. and Sardina sp., anchovy Engraulis spp., horse mackerel Trachurus spp. and chub mackerel Scomber japonicus, with hake Merluccius spp. also abundant in each of the systems except off Japan. In these systems many of the more numerous fish, birds and mammals are opportunistic feeders having catholic diets. As a result species often have overlapping diets, and when a particular food item is plentiful it may be consumed by a wide variety of organisms at higher trophic levels. Frequently the performances of predator populations have been related to the availability of forage species, but predators have not always been influenced similarly by gross changes in the systems. Predators with the most generalized feeding strategies appear best buffered against collapses of particular prey resources. Although seabirds are generally opportunistic feeders, limitations in their foraging range may make them more susceptible than most other predators to system perturbations. Opportunistic feeding by species at lower trophic levels suggests that they could be advantaged by an increased food supply, and it may not be unreasonable to interpret species replacements in catches as reflecting shifts in the dominance of species. Overall catches from the systems have usually been more stable than the catches of individual species, which have shown wide fluctuations. Shifts in the dominance of species in the catches have often been sustained over a number of years. Generalized feeding creates the possibility that more than one species may benefit from energy made available by the collapse of an abundant resource, and there is evidence that species more abundant in former years have in some instances been partially replaced by a number of organisms. The likelihood of a particular organism becoming the main replacing species will probably be influenced by the degree to which it is utilized during and after the collapse of the originally dominant resource.     

Predatory Promesostoma species (Plathelminthes, Rhabdocoela) in the Wadden Sea

Laboratory experiments were performed on the food ecology of four congeneric species of free-living plathelminths, Promesostoma caligulatum, P. marmoratum, P. rostratum, and P. meixneri, all inhabiting an intertidal sandflat near the island of Sylt (North Sea). Their prey spectrum is within the microcrustaceans: P. caligulatum preferred ostracods, while the other three species favoured copepods, with species-specific differences for copepod species and size classes. Daily consumption of prey species varied with the size of both the predator and the prey. On average, P. marmoratum consumed 0.76 Harpacticus flexus per day while this rate decreased to 0.06 in P. meixneri, the smallest predator. When these Promesostoma species were fed with Tachidius discipes, a smaller prey species, their predation rates were about 25% higher. While the larger predators preferred the larger harpacticoids as prey, the small P. meixneri preferred small cyclopoids over larger harpacticoids. In terms of biomass, P. marmoratum's mean consumption of T. discipes per day was about half the predator's own weight. This average varied with prey density and temperature. A comparison of these consumption rates with the field densities of the predators and their prey shows that the plathelminth predators may consume as much as 10% per day of their copepod prey populations, thus strongly influencing these prey populations on these sandflats. The predation pressure of P. caligulatum on ostracods was about 1% per day of the prey population. Since ostracods usually have fewer generations per year, the total effect on the population dynamics may be similar to that on copepods. Therefore, nocturnal swimming of copepods in the water column may be interpreted as an attempt to escape plathelminth predators.     

Changes in survivorship,behavior, and morphology in native soft‐shell clams induced by invasive green crab predators

Many studies on invasive species show reduced native densities, but few studies measure trait‐mediated effects as mechanisms for changes in native growth rates and population dynamics. Where native prey face invasive predators, mechanisms for phenotypic change include selective predation, or induced behavioral or morphological plasticity. Invasive green crabs, Carcinus maenas, have contributed to declines in native soft‐shell clams, Mya arenaria, in coastal New England, USA. We tested the hypothesis that clam ability to detect chemical cues from predators or damaged conspecifics would induce greater burrowing depth as a refuge from invasive crabs, and greater burrowing would require increased siphon growth. To determine how crab predation affected clam survivorship and phenotypic traits in the field, clams in exclosure, open, and crab enclosure plots were compared. Crab predation reduced clam density, and surviving clams were deeper and larger, with longer siphons. To determine whether the mechanism for these results was selective predation or induced plasticity, phenotypes were compared between clams exposed to chemical cues from crab predation and clams exposed to seawater in laboratory and field experiments. In response to crab predation cues, clams burrowed deeper, with longer siphons and greater siphon mass. Overall, crab predation removed clams with shorter siphons at shallow depths, and crab predation cues induced greater burrowing depths and longer siphons. Longer siphons and greater siphon mass of deeper clams suggests clams may allocate energy to siphon growth in response to crabs. By determining native behavior and morphological changes in response to an invasive predator, this study adds to our understanding of mechanisms for invasive impacts and illustrates the utility of measuring trait‐mediated effects to investigate predator–prey dynamics.     

Comparing trophic flows in the southern Benguela to those in other upwelling ecosystems

A balanced trophic flow model of the southern Benguela ecosystem is presented, averaging the period 1980–1989 and emphasizing upper trophic levels. The model is based largely on studies conducted within the framework of the Benguela Ecology Programme and updates the results of an expert workshop held in Cape Town in September 1989. Small pelagic fish other than anchovy Engraulis capensis and sardine Sardinops sagax, mainly round herring Etrumeus whiteheadi and mesopelagic fish, were important components of the food web in the southern Benguela. Severe balancing difficulties were encountered with respect to the semi-pelagic resources (hake Merluccius spp.) and demersal top predators (sharks), indicating the need for further research on the interaction of these groups with their ecosystem. The model is compared to other existing trophic flow models of ecosystems in major upwelling areas, i.e. the northern Humboldt Current (4–14°S), the California Current (28–42°N) and the southern Canary Current (l2–25°N), and to two independently constructed models of the northern Benguela ecosystem. These models are compared using network analysis routines of the ECOPATH software, focusing on the interactions between the five dominant fish species (anchovy, sardine, horse mackerel Trachurus trachurus capensis, chub mackerel Scomber japonicus and hake) that support important fisheries in all systems. The upwelling systems rank by size rather than species dominance. The ratio of catches and primary production differs between systems, partly because of differences in fishing regimes. Predation on the five dominant fish groups by other fish in the system was the most important cause of fish mortality in all models. Fishery catches are generally a larger cause of mortality for these groups than predation by mammals. The ecological cost of fishing appears to be comparatively low in the southern Benguela, because catches are low compared with the primary production, but also because the fishery is relatively low in the foodweb. However, in view of the very tight foodweb demonstrated in the model. it is likely that an increase in fishing pressure would cause severe trade-offs with respect to other components of the southern Benguela ecosystem.     

The role of macrophytes as a refuge and food source for the estuarine isopod Exosphaeroma hylocoetes ( )

The role of submerged macrophytes as refugia from fish predation and as possible food sources for the estuarine isopod Exosphaeroma hylocoetes ( Barnard, K.H., 1940) was investigated. Laboratory experiments tested the effectiveness of artificial vegetation, replicating submerged vegetation, in enabling isopods to elude selected fish predators Rhabdosargus holubi, Glossogobius callidus, Monodactylus falciformis and Clinus cottoides. Isopods preferentially hid in the vegetation (>90%), even in absence of fish. The predatory fish had varying success in finding isopods within the vegetation. Isopod mortality ranged from 2% (R. holubi) to a maximum of 87% (C. cottoides) within vegetation, depending on the fish predator present. Stable isotope and fatty acid analyses ruled out the submerged macrophyte Ruppia maritima and inundated fringing grasses as direct food sources, but highlighted the epiphytic biota (mainly diatoms) found on the submerged vegetation and sediments as more likely food sources. These findings are consistent with gut content analyses. The results suggest that the close association of E. hylocoetes with R. maritima is the result of the vegetation providing the isopod with a refuge against fish predation as well as areas of increased food availability.     

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).