Feeding guild structure of a rocky intertidal fish assemblage in central Chile

The identification and analysis of the guild structure of vertebrate assemblages has played a fundamental role in the understanding of the underlying mechanisms responsible for their community organization and structuring. This approach generally has not been undertaken for temperate water intertidal fish assemblages. In central Chile, fishes are important components of the intertidal community, but no studies attempting to understand their organization and structuring have been done. In the present study, the diets of 13 of the most abundant species which inhabit tidepools in the rocky intertidal zone of central Chile were determined. A total of 660 fishes was collected at 5 sites: Los Molles, Con-Cón, Quintay, El Tabo, and Las Cruces. Dietary overlap between all species pairs was calculated and a phenogram of dietary similarity was constructed and analyzed using a bootstrapping technique to objectively determine guild membership. The results showed that the intertidal fish assemblage of central Chile can be divided into three feeding guilds: two guilds consisting of carnivorous species and one guild of omnivorous and herbivorous species. The possible causes and implications of the resulting guild structure and the potential effects of predation by these fishes on other components of the intertidal community are discussed.     

Feeding guilds and food resource partitioning in a lake fish assemblage: an ontogenetic approach

Ontogenetic diet patterns and trophic guild structure of a 15 species temperate lake fish assemblage were analysed over wide size intervals (up to seven orders of magnitude in body mass), representing practically the whole life span in most species. A two-step objective clustering technique supplemented with other multivariate statistical tools proved that size-related diet changes clearly played an important role in structuring trophic organization of fishes inhabiting Lake Balaton. As many as 13 out of the 15 fish species showed marked size-related dietary changes with two to four ontogenetic feeding stages. At the assemblage level, 11 trophic guilds were separated. Guild membership was size-dependent in 11 fish species that participated in two to four trophic guilds during their life span. The most complex trophic ontogeny was observed in roach Rutilus rutilus and asp Aspius aspius with four guild memberships. This study showed that trophic status of fishes may be very size-sensitive and thus a universal classification of fish species to general trophic guilds, such as 'planktivore', 'benthivore', 'piscivore' or 'herbivore', should be applied very carefully even in environmental monitoring and fisheries management applications, unless it is supported by relevant results of life span diet analyses.     

Food partitioning among carnivores within feeding guild structure of fishes inhabiting a mudflat ecosystem of Indian Sundarbans

The present study was aimed to determine dietary composition and feeding guild structure of the fishes inhabiting mudflat habitat of Indian Sundarbans. In addition, partitioning of food resources by the fish species belonging to the carnivorous feeding guild was also performed to understand the survival strategies of fish in a mudflat estuarine habitat. Seventeen prey categories were isolated from the stomachs of 31 studied fish species. Overall, five feeding guilds (viz. plankti-benthivore: 12 species, herbivore: one species, detritivore: three species, omnivore: one species and carnivore: 14 species) were identified on the basis of the prey abundance within their stomachs, considering 64 % Bray–Curtis similarity. Among the carnivorous, maximum trophic richness was obtained for Uroconger lepturus followed by Ophichthus apicalis. Teleost and decapods were the main animal prey items preferred by majority of the carnivorous fishes. However, O. apicalis and Terapon jarbua showed their preference toward maximum number of prey categories among carnivores, which was also ratified by the high values for standardized niche breadth presented by them. The maximum degree of interspecific dietary overlap was found between Uropterygius marmoratus and Pseudapocryptes elongatus as both of them were recognized as cranci-piscivore. The lowest was observed between Hyporhamphus limbatus and Coilia neglecta. As food resources are not limiting in the mudflats of Indian Sundarbans, the general patterns of resource partitioning and niche differentiation in resident ecological communities will improve the understanding of the mechanisms underlying species coexistence and community structure.     

Seasonal and spatial variations of fish trophic guilds in a shallow,semi-enclosed tropical estuarine bay

A study on seasonal and spatial variations of feeding habits and trophic guilds of dominant fish species in Pattani Bay during March 2003 to February 2004 was aimed at classifying diet composition, identifying dominant food components of each species, categorizing trophic guilds of the community and evaluating effects of habitat characteristics and seasonality on guild organization. Most fishes showed high food intake, fed on a diverse range of food items but relied heavily on calanoid copepods and shrimps. All species, with the exception of Epinephelus coioides, were classified as specialist feeders. Four main dietary guilds were classified. Three of these were classified as the guilds dominated by at least two major food items. Significant variations in trophic guilds of 28 fish species based on habitat types and seasons were also identified. They could be divided into three seasonal groups and three site-groups and a single site. Trophic organization for each season and habitat ranged from two to four groups. This information identifies groups of fishes that seasonally and spatially utilize different food resources within a semi-enclosed estuarine bay ecosystem.     

Trophic relationships among concentrated populations of small fishes in seagrass meadows

Changes in available food and utilized foods for a densely populated guild of animals can uncover periods of niche displacement among particular consumer species or their separate size classes. Dense populations of small fishes and their foods were çensussed for 13 months and studied experimentally in shoalgrass and turtlegrass meadows of Redfish Bay, Texas. Feeding habits were determined with respect to prey availability, and illustrated the extent to which seasonal partitioning of food corresponded to food depletion among these abundant consumers in seagrass meadow food webs. The darter goby, code goby, pinfish, and Gulf pipefish were the most common species throughout the year. Although the darter goby did not show distinct ontogenetic changes in food habits, the code goby, Gulf pipefish, and pinfish demonstrated major ontogenetic progressions of foods selected.Food availability in the seagrass meadows changed seasonally. When major prey such as amphipods were abundant, during spring, many fish species showed high overlap in food use. Regardless of food availability, the code goby and Gulf pipefish fed mainly on amphipods or copepods. The more common darter goby and pinfish seasonally changed their diets with changes in food availability; the darter goby and pinfish were more carnivorous during spring, but they largely consumed epiphytic algae during summer. Cage experiments were used to monitor foods confined with elevated densities of pinfish and darter gobies, relative to control cages at natural overall densities. Prey items in the former cages decreased sharply, with corresponding dietary shifts by these common fishes. During resource depletion, these changes in resource use by these naturally concentrated consumers appeared as temporary partitioning of available resources. This shift occurred during both natural (seasonal) and experimental depletions of food, and appears to result from increased interspecific and intraspecific competition during periods of depleted preferred foods.     

Feeding habits and seasonal trophic guilds structuring fish community in the bay mouth region of a tropical estuarine habitat

Understanding trophic relationships of fish in estuarine ecosystem is an important element for sustainable resource management. This study examined the feeding habits of 29 dominant fish species, characterized the trophic guilds, assessed the impact of season and clarified the role of diets in structuring the fish community in the mouth region of Pattani Bay, Thailand. Samples of 5792 fishes collected monthly by gillnets from March 2019 to February 2020 were used for stomach content analyses. It was found that the number of food types and fullness index differed between fish taxa (P < 0.001). Most fishes were specialist feeders feeding on specific food components and were categorized into five trophic guilds: piscivore, shrimp-fish feeder, polychaete feeder, zooplanktivore and planktivore. Six species were piscivorous, considered as apex predators, that fed almost entirely on fishes. High diet overlaps among some species (>0.6) were recorded. Not much variation in seasonal guilds was observed: four guilds in the dry season, three in the moderate rainy season and four in the rainy season. Some species remained in the same guild the whole year round, but some fishes changed seasonally. Two fish communities from different regions of the bay were segregated based on feeding habits. The inner bay community comprised mainly copepod and plankton feeders, but there were more piscivores in the deeper bay mouth area. Results from this study help us to understand the feeding habits and trophic guilds of dominant fish species at the mouth of this tropical estuarine bay.     

A partial food web involving predatory gastropods on a Pacific fringing reef

An analysis of the feeding habits of a diverse assemblage of predatory gastropods on a Guam fringing reef, shows that seven major prey categories are eaten, namely polychaetes, sipunculans, gastropods, bivalves, crustaceans, ophiuroids, and fish. The three largest feeding groups are the polychaete feeding guild which mainly consists of species of Conus, the gastropod guild dominated by species of the Muricidae and the sipunculan guild consisting mainly of the Mitridae. There is considerable dietary overlap between members of feeding guilds, but interaction between guilds is low and accounted for by the relatively few generalist species in the community.     

Food habits of small fishes in a common reed <Emphasis Type="Italic">Phragmites australis</Emphasis> belt in Lake Shinji,Shimane, Japan

To clarify the feeding habits of reed fishes, the gut contents of 13 fish species collected in a Phragmites australis belt in Lake Shinji were examined. Six species showed ontogenetic and/or seasonal changes in food use patterns. Smaller individuals generally preyed on small planktonic items (e.g., calanoid and cyclopoid copepods) or small crustaceans (gammaridean amphipods), subsequently changing to other prey items (e.g., mysids and filamentous algae) with growth. The most important dietary items for the reed fish assemblage comprised planktonic copepods, gammaridean amphipods and mysids. However, the relative importance of these changed seasonally, gammaridean amphipods being the most important in autumn and winter, and planktonic copepods and mysids the most important in spring and in summer. Cluster analysis based on dietary overlaps showed that the reed fish assemblage comprised five feeding guilds (planktonic-copepod, mysid, gammaridean-amphipod, filamentous-algae, and detritus feeders). Of these, the three former guilds were the most abundantly represented, whereas detritivores were represented by a single species.     

Comparing the diet of two sympatric urolophid elasmobranchs (Trygonoptera testacea Müller & Henle and Urolophus kapalensis Yearsley & Last): evidence of ontogenetic shifts and possible resource partitioning

The diets of two urolophids, the common stingaree Trygonoptera testacea and the kapala stingaree Urolophus kapalensis were analysed and compared to examine resource partitioning between these two morphologically similar sympatric Australian batoids. The diet of T. testacea was polychaete-based, while that of U. kapalensis was dominated by crustaceans (mostly carid shrimps and amphipods). Intraspecific dietary compositions were examined amongst size classes within each of the two species to identify ontogenetic shifts in diet. Differences in the dietary compositions of the smaller total-length classes of T. testacea suggest that their diet shifts as they increase in size, from one dominated by carids to one almost entirely comprising polychaetes. Significant ontogenetic dietary shifts were not identified in U. kapalensis . Although the two species shared eight broad dietary categories, their overall dietary compositions were found to be significantly different. The limited overlap in the dietary compositions of these two sympatric stingarees suggests the possibility of resource partitioning, with interspecific competition being implicated as a possible cue. Possible mechanisms for the partitioning of resources within and between these two species are discussed.     

Analysis of trophic structure of two carnivore assemblages by means of guild identification

We evaluated the existence of trophic guild structure, considering seasonal and annual variation, in two terrestrial carnivore assemblages: one from Santa Cruz province (Argentinean Patagonia, composed by six carnivore species), and the other from Doñana National Park (SW Spain, composed by five carnivore species). To identify trophic guilds, we first studied seasonal and annual diets of predators, calculated trophic overlap among species pairs, and then constructed overlap matrices (similarity matrices). We determined guild membership objectively by entering the similarity matrices into the clustering technique unweighted pair-group method with arithmetic averaging. Carnivores from both assemblages were grouped, respectively, into four feeding guilds. Lagomorphs and rodents promoted the formation of two feeding guilds in both study sites, although the taxonomic composition of predator species that composed them was different. The ungulates-edentates feeding guild was only present at Santa Cruz, whereas the birds and reptiles feeding guild was only present at Doñana. Invertebrates and fruits were the base for the formation of a guild composed by species of the same taxonomic origin both in Santa Cruz and Doñana. Guild structure of Santa Cruz and Doñana assemblages did not exhibit seasonal or annual variation, although the specific guild composition changed over the two studied periods for both assemblages. This structure probably responded to discontinuities in resource spectra in Santa Cruz and fluctuations in rabbit abundance in Doñana. Our results support the hypothesis that establishes that guilds are originated by opportunistic convergence of species on abundant and energetically rewarding resources.     

Patterns of morphological traits shaping the feeding guilds in the intertidal mudflat fishes of the Indian Sundarbans

Broad-scale patterns of resource utilization and the corresponding morphological evolution is a result of an integral relationship among form and function. In addition, there is also an inherent role of the latter in determining species co-interaction and assemblage pattern that forms an integral aspect of ecological research. The present study aimed to evaluate the ecomorphological relationship among 37 fish species inhabiting the intertidal mudflats of the Indian Sundarbans by outlining the following objectives: (i) identifying and characterizing feeding guilds/groups and (ii) understanding the inter-relationship between morphometry with (a) the established feeding guild classifications and (b) observed prey taxa (that characterizes these feeding groups) for determining the role of morphometry in prey acquisition followed by (iii) the evaluation of their potential phylogenetic convergence among the species. For the first objective, two approaches for feeding guild classification were made (3-Guild and 8-Guild) for assessing the prediction accuracy of morphological characters in identifying the different guilds. While the former was based on troph values, the latter classification mode relied on the similarities in diet composition among the different fish species. For addressing the second objective, we employed two different models namely, linear discriminant (LDA) and redundancy analysis (RDA). While the LDA model tested the prediction accuracy of morphological traits in classifying the different feeding guilds, RDA was applied to model the correlation between the morphological traits and the prey categories. In the LDA model, morphological characters showed higher accuracy (78.4%) in classifying three feeding groups rather than eight feeding groups (73%). Following this, the RDA model (explaining 79.78% of constrained variance) showed gill raker intensity, protrusion length, head depth, caudal peduncle, eye diameter and inter-orbital distance to be highly associated with selection of specific prey types by species, thereby characterizing a particular feeding guild. However, generalized linear models testing for correlation between troph value and feeding groups showed substantial variation (90.35%) in the dietary index being explained by the 8-Guild classification. Hence, our study maintains the assumption that broad morphological differentiation acts as one of the underlying processes resulting in dietary variations that results in the varying modes of resource utilization by the coexisting species, thereby determining the structure of a trophic guild. Furthermore, it also suggests that in terms of prey abundance or selectivity, the 8-Guild model is much more conducive in representing the feeding habits of the species while the morphological traits reflected a relatively broader scheme of classification, (i.e., 3-Guild model) with certain traits being phylogenetically conserved within these groups.     

Effect of siltation on stream fish communities

Synopsis The effect of siltation on stream fish in northeast Missouri was evaluated using community structural measurements and a functional approach that emphasized feeding and reproductive guilds. As the percentage of fine substrate increased, the distinction among riffle, run, and pool communities decreased, primarily because the number of individuals of typical riffle species decreased. Within the riffle communities the abundance of fish of two feeding guilds — benthic insectivores and herbivores — was reduced as the percent of fine substrate increased. The abundance of fish in other feeding guilds was not affected. The only reproductive guild to be similarly affected was the simple and lithophilous, whose members require a clean gravel substrate for spawning. Species within each guild affected by siltation had significantly similar trends in abundance. The guild analysis indicated that species with similar ecological requirements had a common response to habitat degradation by siltation.     

Resource partitioning in rhinolophoid bats revisited

We assessed the ecomorphological structure of a guild of rhinolophoid bats in a Malaysian rainforest first described by Heller and von Helversen (1989). These authors found that the distribution of echolocation call frequencies used by 12 syntopic species was more even than expected from allometric relationships or in randomly generated communities, and that the observed minimal ratio was greater than expected by chance alone. In this study we were able to expand their guild to 15 species, but in doing so it became apparent that call frequencies might be less evenly distributed across the total frequency range than previously proposed. We replicated Heller and von Helversen’s (1989) analyses with the full 15-species complement but were unable to support their suggestion that rhinolophoid bats exhibit resource partitioning through differences in frequency bands. We adopted a multivariate approach and incorporated measures of body size and wing morphology into the analysis. We used phylogenetic autocorrelation to ensure that the species were statistically independentand principal component analysis to describe the morphological space occupied by the 15 species in the community and four additional species representing the extremes of phenotypic variation. We derived interspecific Euclidean distances and tested the mean values and SDs of these distances against those of 100 guilds of ”synthetic” species created randomly within the principal component space. The guild of Rhinolophoidea was not distributed randomly in multivariate space. Instead we found evidence of morphological overdispersion of the most similar species, which suggests niche differentiation in response to competition. Less similar species were nearer in morphological space than expected, and we suggest this is a consequence of ecological constraints on parameter combinations. Despite this underdispersion, many of the more distant neighbours were evenly rather than randomly spaced or clumped in morphospace, suggesting that, given the environmental constraints on morphology, species in this guild do experience limits to their similarity. Finally, we tested the influence of the relative abundance of species on morphological displacement, and found no evidence that abundant, spatially correlated species reduce interspecific overlap in morphological space. Received: 10 April 1999 / Accepted: 28 February 2000     

Species packing and the competition function with illustrations from coral reef fish

This paper concerns the contrast between guilds whose species show resource partitioning and those that show extensive overlap. Using a Lotka-Volterra model, the ease of invasion by a third species into a guild already containing two species is examined for various shapes of resource utilization curves. I show that (a) a guild is more easily invasible and allows tighter packing if its member species have leptokurtic (thick-tailed) resource utilization curves than if they have platykurtic (thin-tailed) curves; (b) the distribution of niche separation distances is bimodal in a “thin-tailed” guild and is unimodal in a “thick-tailed” guild; (c) there are three-species guilds such that removal of one particular species leaves a two-species system in which one of the remaining species excludes the other. In this context, competition pressure is a force maintaining species diversity.Groupers (Serranidae) appear to be a thin-tailed guild, and Parrotfish (Scaridae) and Surgeonfish (Acanthuridae) together appear to be a thick-tailed guild, and these guilds show many properties predicted by the model. I conjecture that thick-tailed guilds form when the constituent species are selected to be generalists and apply this idea to tropical fruit- and flower-feeding birds.     

Functional trait assembly through ecological and evolutionary time

A classic community assembly hypothesis is that all guilds must be represented before additional species from any given guild enter the community. We conceptually extend this hypothesis to continuous functional traits, refine the hypothesis with an eco-evolutionary model of interaction network community assembly, and compare the resultant continuous trait assembly rule to empirical data. Our extension of the “guild assembly rule” to continuous functional traits was rejected, in part, because the eco-evolutionary model predicted trait assembly to be characterized by the expansion of trait space and trait/species sorting within trait space. Hence, the guild rule may not be broadly applicable. A “revised” assembly rule did, however, emerge from the eco-evolutionary model: as communities assemble, the range in trait values will increase to a maximum and then remain relatively constant irrespective of further changes in species richness. This rule makes the corollary prediction that the trait range will, on average, be a saturating function of species richness. To determine if the assembly rule is at work in natural communities, we compared this corollary prediction to empirical data. Consistent with our assembly rule, trait “space” (broadly defined) commonly saturates with species richness. Our assembly rule may thus represent a general constraint placed on community assembly. In addition, taxonomic scale similarly influences the predicted and empirically observed relationship between trait “space” and richness. Empirical support for the model’s predictions suggests that studying continuous functional traits in the context of eco-evolutionary models is a powerful approach for elucidating general processes of community assembly.     

Ontogenetic niche shifts and resource partitioning in a subarctic piscivore fish guild

The feeding ecology of three piscivorous fish species (perch (Perca fluviatilis), pike (Esox lucius) and burbot (Lota lota)), was studied in the subarctic Pasvik watercourse (69 °N), northern Norway and Russia. All three species primarily occupied the benthic habitats in the watercourse. Perch and burbot exhibited distinct ontogenetic niche shifts in food resource use, perch changing from a dominance of zooplankton to zoobenthos to fish, and burbot from zoobenthos to fish. Fish prey dominated the diet of all the investigated size-classes of pike, but small-sized pike (<20 cm) were not represented in the sample. Fish prey size was positively related to predator size in all three species. Whitefish (Coregonus lavaretus) was the dominant prey of pike and large-sized burbot and perch. Nine-spined sticklebacks (Pungitius pungitius) was also an important prey and appeared to be a dietary stepping-stone enhancing the transition from invertebrate feeding to consumption of large-sized whitefish prey for all three predators. A cluster analysis separated the different size groups of the three predator species into five functional feeding groups, most of them containing two or all three species. Within these feeding groups, and especially among the piscivorous size-classes, there was a strong and significant interspecific overlap in prey selection, and the dietary similarities between the species were in general much larger than the intraspecific similarities between ontogenetic stages. All three piscivorous species are important top predators in the aquatic food web of the watercourse, and their ontogenetic diet shifts and resource partitioning patterns generate a substantial food web complexity in this subarctic ecosystem.     

Seasonal dynamics of guild structure in a bird assemblage of the central Monte desert

Most studies of ecological guilds consider guild membership as a fixed attribute of species, and the guild structure as a fixed attribute of a community. Consequently, almost no study tackled the examination of temporal changes in the guild structure of a particular assemblage. We used data on foraging patterns of central Monte desert birds to assess seasonal differences in resource use by resident species, and to examine seasonal changes in the guild structure of the assemblage. From 1993 until 1999, we quantified the use of foraging substrates, manoeuvres, and height strata, and the utilisation of plants by 18 bird species. Resident species showed different kinds of seasonal dynamics, from no noticeable to drastic changes in their foraging patterns. Seasonal changes in guild structure were remarkable. Only two guilds were present all year round (ground foragers and foliage foragers), although they suffered important species turnover. Three other guilds had a seasonal occurrence: graminivores (during the non-breeding season), arboreal herbivores, and aerial feeders (during the breeding season). Two species changed from one guild to another between seasons. Both presence–absence of migrant species, and changes in the feeding ecology of resident species had an influence on the temporal dynamics of guild structure. These factors appeared to be related with changing resource availability, especially with the birds’ ability for facing lean conditions through migration and with the opportunistic use of seasonally superabundant resources. Our results undermine the trust in studies in which guild structure was assessed irrespective of temporal variations. We show also that guild membership is not a fixed attribute of a species: we should rather evaluate the individuals’ resource use in a given moment to assign them to a guild, instead of obtaining a single measure purporting to represent a species’ fixed attribute.     

Feeding biology of a guild of benthivorous fishes in a sandy shore on south-eastern Brazilian coast

The feeding biology of eight species of benthivorous fishes was studied in a sandy shore at Anchieta Island, south-eastern Brazilian coast. The fishes fed mainly on Amphipoda and Mysidacea crustaceans. The diet of the most abundant species, the drum Umbrina coroides, was analyzed in three standard length classes (20-55, 56-90 and 91-135 mm). This sciaenid showed an ontogenetic diet shift from Mysidacea to Amphipoda. The feeding behaviour of the sciaenid U. coroides and the gerreid Eucinostomus gula was recorded while snorkeling. During their foraging both species uncovered small organisms buried in the sand. Notwithstanding general similarities in diet, U. coroides and E. gula presented differences in feeding behaviour and morphology. Two carangid species of the genus Trachinotus differed in diet composition and consumed a larger array of food items than the remaining fish species. Differences in diet and feeding activity between the remaining benthivorous species were noted. These differences possibly reduce overlap in resource use and favour the coexistence of guilds of benthivorous fishes on sandy shores.     

Secondary succession and breeding bird community structure: Patterns of resource utilization

Summary Structure of breeding bird communities was compared among four habitat types representative of stages present in most old-field successions in the eastern deciduous forest formation of North America. The successional catagories, defined by vegetational structure, were designated herbaceous (type A habitats) herb, shrub and sapling (type B), young forest (type C), and older forest (type D). Density of breeding birds was lowest in A habitats, rose sharply in B habitats and reached a maximum in D habitats. Species richness and number of feeding guild showed similar patterns. Mean number of species per guilds was highest in D habitats. Generalist feeding guilds were predominant in type A and B habitats, primarily due to the importance of the graminivore-insectivore guild. Specialist guilds increased in importance with succession due to addition of several insectivorous guilds in later successional stages. Patterns of occurrence of individual feeding guilds are analyzed and discussed with respect to changes in vegetational structure. Variance in community structure was generally highest in A habitats and lowest in D habitats; this is discussed in relation to the evolution of saturated, coadapted communities.     

Structure of winter groundfish feeding guilds in Pacific herring Clupea pallasii and walleye pollock Gadus chalcogrammus nursery fjords

The contents of 1056 stomachs were included in a trophic-guild analysis to document separation amongst 16 groundfish species inhabiting Pacific herring Clupea pallasii and walleye pollock Gadus chalcogrammus nursery fjords in Prince William Sound, Alaska and to determine the relative contribution of C. pallasii and G. chalcogrammus to that separation. A total of five multi-species feeding guilds and one outlier species were determined through multivariate analyses. Major gradients of trophic separation spanned from invertebrates (mostly shrimps, crabs and unidentified decapods) to fishes (mostly unidentified fishes, C. pallasii and G. chalcogrammus) a pattern that was influenced by intra and interspecific differences in predator lengths. While C. pallasii and G. chalcogrammus were important to the overall guild structure, within-guild similarities were consistently highest due to unidentified fishes. In general, larger predators consumed the largest C. pallasii and G. chalcogrammus, with the smaller-on-average predators consuming smaller C. pallasii and fewer or smaller G. chalcogrammus. Regardless of guild inclusion, groundfishes primarily consumed pre-recruit C. pallasii and G. chalcogrammus (i.e., younger than age 3 years fishes), which has the potential to negatively influence recruitment of these forage fishes to the adult, spawning population.