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Diets of Five Hake Species in the Northeast United States Continental Shelf Ecosystem

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Diets of Five Hake Species in the Northeast United States Continental Shelf Ecosystem MARINE ECOLOGY PROGRESS SERIES Vol. 204: 243–255, 2000 Published October 5 Mar Ecol Prog Ser Diets of five hake species in the northeast United States continental shelf ecosystem Lance P. Garrison*, Jason S. Link Food Web Dynamics Program, Northeast Fisheries Science Center, National Marine Fisheries Service, 166 Water St., Woods Hole, Massachusetts 02543, USA ABSTRACT: We examined the diets of 5 hake species in the continental shelf ecosystem of the north- east USA: offshore hake Merluccius albidus, silver hake M. bilinearis, white hake Urophycis tenuis, red hake U. chuss, and spotted hake U. regia. We conducted a multivariate analysis of stomach contents from bottom-trawl surveys to assess the major factors that determine diet composition, to compare and contrast diets between species, and to compare the trophic role of the hakes in the northwest Atlantic to hakes in other ecosystems. These fish consume pelagic invertebrates (e.g. eu- phausiids and other shrimps) and pelagic fish prey, including conspecifics. The incidence of piscivory generally increases with increasing size. Regional and temporal (seasonal and yearly) variation in hake diets generally reflects differences in prey availability. Hakes are among the most abundant predators in continental shelf ecosystems worldwide. Fish prey consumption by hakes can be high and may be a primary source of mortality for economically important pelagic species. Hake predation plays an important role in the dynamics of both the US northeast continental shelf ecosystem and generally on a global scale. KEY WORDS: Food habits · Hakes · Northwest Atlantic fisheries · Trophic dynamics Resale or republication not permitted without written consent of the publisher INTRODUCTION al. 1991), the southeast Pacific Ocean off the Chilean coast (M. gayi, Arancibia 1987, Stobberup 1992), the Hakes are both ecologically and commercially im- southeast African coast (M. capensis, Pillar & Wilkin- portant in continental shelf ecosystems worldwide, son 1995), and the northwest Atlantic (M. bilinearis, with global landings on the order of millions of tons per Bowman 1983, Waldron 1992, Helser et al. 1995). year (Pitcher & Alheit 1995). They are demersal gadid In the northwest Atlantic, the hakes (genera Merluc- fish with streamlined bodies (Fig. 1), well developed cius and Urophycis) are among the dominant biomass gill rakers, strong teeth, and pike-like jaws. In addi- components, and are both ecologically and economi- tion, they have large, well-developed eyes and a high cally important. Silver (M. bilinearis), white (U. tenuis), proportion of white-muscle tissue. This suite of mor- and red (U. chuss) hake have sustained considerable phological traits is consistent with the general trophic exploitation both in directed fisheries and as incidental nature of hakes as ambush predators (Pitcher & Alheit catch in otter-trawl fisheries for other groundfish over 1995). The hakes are important invertebrate and fish the last 4 decades (Helser et al. 1995, Northeast Fish- predators in continental shelf systems worldwide, eries Science Center 1998). including the northeast Atlantic Ocean (Merluccius The hakes are important both as predators and prey merluccius, Guichet 1995), the northeast Pacific Ocean in the northeast shelf ecosystem of the USA. Small (M. productus, Rexstad & Pikitch 1986, Tanasichuk et hakes are preyed upon by demersal gadids, pelagic species, and squids (Bowman & Michaels 1984). Can- nibalism and predator-prey interactions within the *E-mail: [email protected] hakes is common, particularly in silver hake, where © Inter-Research 2000 244 Mar Ecol Prog Ser 204: 243–255, 2000 frequently >10% of adult diets consist of conspecific exploited species. As fishing pressure is relaxed through juveniles (Bowman 1983). Because hakes are highly management efforts to rebuild over-exploited stocks, it abundant predators, prey consumption by hakes is will be necessary to incorporate species interactions, probably high, and may influence prey population particularly predator-prey interactions, into models to dynamics. Silver hake is a biomass dominant in this effectively predict and manage stock recovery (Jen- community, and plays an important role in the dynam- nings & Kaiser 1998). This requires a detailed under- ics of the continental shelf ecosystem of the northeast standing of the trophic relationships between taxa and United States as a predator of pelagic fish, squids, and the factors that determine the strength of predator- invertebrates (Sissenwine et al. 1984, Helser et al. prey interactions. Toward this goal, we present an 1995). analysis of the trophic patterns of the 5 major hake Fisheries management and stock-assessment sci- species (Fig. 1) in the northeast shelf ecosystem of the ence has historically taken a single-stock approach to USA from a 25 yr time-series of food habits collected in evaluating and predicting the population dynamics of seasonal resource bottom-trawl surveys. We assess the temporal, spatial, and ontogenetic trends in hake diets. Additionally, we compare and contrast the diets of these species and discuss their roles as predators in this and other ecosystems. METHODS Offshore Hake - Merluccius albidus Data collection. The data for this study were drawn from seasonal bottom-trawl surveys conducted by the Northeast Fisheries Science Center (NEFSC). The sur- veys employ a stratified random-sampling design with strata defined by depth and latitude. Sample depths ranged between 8 and 400 m. Sample stations were defined by 2.5’ latitude × 2’ longitude rectangular units Silver Hake - M. bilinearis that were randomly selected within strata. Between 350 and 400 stations on the continental shelf between Cape Hatteras, North Carolina, and southwestern Nova Scotia (Fig. 2) were sampled during each sea- sonal survey of approx. 4 to 6 wk in duration (North- east Fisheries Science Center 1988). At each station, a 36 Yankee (or similar) bottom trawl was deployed for –1 White Hake - Urophycis tenuis 30 min and towed at a speed of 6.5 km h . Sub-stan- dard tows or those with non-representative sampling resulting from gear damage were removed prior to analyses. For each tow, catch (in both mass and num- bers) at length (1 cm length classes) was recorded for all species. Details of the survey sampling design, exe- cution, and efficiency are available in Azarovitz (1981) and Northeast Fisheries Science Center (1988). Red Hake - U. chuss In addition to catch data, a subset of species were analyzed for food-habits data during each seasonal cruise. The NEFSC has conducted quantitative food- habits sampling in seasonal surveys from 1973 to the present. Prior to 1981, stomach contents were pre- served and returned to the laboratory for identification of prey items. Total stomach content mass and the mass of each prey was measured to the nearest 0.1 g. Since 1981, stomach contents have been examined at Spotted Hake - U. regia sea. The total volume of stomach contents was mea- Fig. 1. Five hake species in current study. (Drawings from sured and the proportion of stomach contents com- Flescher 1980) prised by each prey item was estimated. Garrison & Link: Diets of five hake species 245 45.0 chuss, U. regia) were included Scotian (Table 1). Shelf Prey and predator categories. 43.0 Gulf of Since stomach contents were iden- Maine tified in the laboratory prior to 1981, the taxonomic resolution of invertebrate prey is generally 41.0 ) higher in these samples. In the cur- N Georges rent study, invertebrate prey were Bank ees r Southern grouped into family or order g 39.0 New England groups to account for differences in de ( taxonomic resolution between the sampling periods. Fish and squid tude i Mid-Atlantic 37.0 at Bight prey were generally recorded at L the species level. Unidentifiable fish were an important component 35.0 of the diets of the hakes in this study. This included both well- Southern Atlantic digested remains and small fishes that were difficult to identify in 33.0 Bight both preserved and fresh samples. -80.0 -78.0 -76.0 -74.0 -72.0 -70.0 -68.0 -66.0 While it is assumed that unidentifi- Longitude (degrees E) able remains reflect the composi- tion of identifiable prey in the diet, Fig. 2. The northeast US continental shelf ecosystem and the southwestern Nova we have chosen to be conservative Scotian shelf. Heavy lines: subarea designations; light lines: sampling strata used in by including it as a separate prey NEFSC resource bottom trawl surveys type in our analyses. At points during the time- Table 1. Summary statistics and sample sizes for 5 hake species in each size class. Size series, prey amounts were mea- class ranges were same for all species (see ‘Methods: Prey and predator categories’); sured as both weights and vol- small: <20 cm; medium: 20 to 50 cm; large: >50 cm umes within the same stomachs. Using these samples, a linear Species No. of No. of Mean Mean Empty Size class tows stomachs body stomach stomachs regression was performed to con- sampled weight weight vert weights to volumes and re- (kg) (g) (%) solve the differences in prey quantification across the time- Merluccius albidus (offshore hake) series. The regression was highly Small 3 10 0.082 50.0 2 Medium 26 62 0.215 0.574 74.2 significant (r > 0.90, p < 0.0001), Large 13 20 17.600 75.0 and prey weights from the 1973 M. bilinearis (silver hake) to 1980 time period were divided Small 2324 8722 0.028 0.452 37.7 by 1.1 to convert weights to vol- Medium 4441 26070 0.144 2.589 51.9 umes based upon this analysis Large 679 1037 0.706 25.349 44.1 (Garrison & Link 2000). Further Urophycis tenuis (white hake) details of the food-habits sam- Small 154 329 0.037 0.784 25.2 pling methodology are available Medium 1634 5341 0.252 4.365 33.3 Large 1755 6049 1.344 29.203 38.1 in Link & Almeida (2000).
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