Trophic Structure of One Deep-sea Benthic Fish Community in the Eastern Canadian Arctic: Application of Food, Parasites and Multivariate Analysis

Synopsis As new arctic marine fisheries develop there is need for a comprehensive ecosystem approach to long-term management. This approach recognizes the importance of community interactions such as food web structure and trophic patterns. We determined whether hierarchical clustering (guild formation) is an effective method of trophic evaluation in deep-sea Artic fish communities using stomach content and parasite data with size class, and evaluated the application of endohelminth communities (parasite species transmitted in the food) as indicators of trophic status. Cluster analysis using food group abundance with size class of fish revealed the presence of 11 guilds within the community, however the same analysis using parasite data showed little correlation between food and parasites. Redundancy analysis (RDA) within the 11 guilds also revealed no significant correlations between food group and parasite abundance suggesting that this type of ordination is not suited for environments containing mainly generalist feeders. RDA of individual taxa without a priori guild designation found that taxa in benthic deep-sea communities are defined by their ability to exploit prey species in more than one habitat zone. Benthic fish species were significantly correlated with benthic food groups and parasites that utilize benthic intermediate hosts whereas benthopelagic–pelagic species fed on a higher diversity of prey species and were infected by a larger number of non-host specific parasites. Eigenanalysis and Monte Carlo results showed that parasites and food groups are highly correlated, indicating that parasite community analysis is an effective tool for predicting feeding strategies in Arctic marine environments. It also suggests that in most cases endoparasite infections alone could be used for trophic evaluation in the absence of stomach content data.     

Guild structure of carnivorous intertidal fishes of the Chilean coast: implications of ontogenetic dietary shifts

Although ontogenetic changes in resource use within species are common in animals, these changes have not been widely considered in studies of guild structure within communities. The occurrence of one or more shifts in resource use in an individual of a given species during its life should mean that it would also belong to different guilds at different life stages. We specifically addressed this issue by describing the feeding habits of ten species of carnivorous fishes occurring in tidepools in rocky intertidal areas along the coast of central Chile. Most of these species undergo clear ontogenetic dietary shifts and a feeding guild structure of this group of fishes was established that takes these dietary shifts into account. Each species was divided into a number of size classes. Dietary overlap values between both intraspecific and interspecific size-class pairs in the entire group of ten species were used to construct a phenogram of dietary similarity through an UPGMA cluster analysis. Numbers of guilds and their memberships were established objectively by applying a bootstrapping procedure. Four “ontogenetic” feeding guilds (OFGs), each consisting of size-classes of species, were recognized. The majority of species belonged to more that one guild. Interestingly, when the bootstrapping procedure was applied to a phenogram based on the diets of “taxonomic” or complete species, only one significant guild was found. The implications of these ontogenetic dietary shifts for interspecific interactions are substantial because the identity of the species with which each fish species shares resources change through their lives. The usefulness of taxonomic species for investigating potential competitive interactions in this assemblage is greatly undermined. Received: 26 September 1997 / Accepted: 15 December 1997     

Spatial and temporal variations among fish with similar strategies: patterns of reproductive guilds in a floodplain

Seasonal flooding regimes are closely related to the life history of neotropical fish, especially with regard to their reproduction. The classification of fish into reproductive guilds serves to identify broad patterns in reproductive types, which are important in developing management and conservation measures. We tested the hypothesis that the fish reproductive guilds in the Upper Paraná River floodplain exhibit spatial and temporal distributions. Samples were taken each quarter in 2010 from several environments (i.e., biotopes) and rivers of the plain. Fish were categorized into four reproductive guilds based on migration, external fertilization, and parental care. Moreover, the abundance and species richness within each guild were used to evaluate their patterns. Guilds were spatially structured, possibly due to species dispersion as well as the influence of the hydrological and limnological characteristics of the biotopes and rivers of the plain. The Paraná River presented lower analyzed metric values, which demonstrates the negative effects of the dam built upstream of the study area. The guilds also presented temporal structure. During the flood season, the guilds presented similar spatial structures, and the local environmental characteristics led to spatial differentiation in the structure of guilds (i.e., flooding promoted the homogenization of the reproductive guilds).     

Fish feeding guilds along a gradient of bay biotopes and coral reef depth zones

The study of fish feeding guild structure is a useful method to compare fish communities of complex marine ecosystems. Guild structure was determined in four coral reef depth zones, viz. the fringing reef at depths of 2, 5, 10, and 15 m, as well as in seven shallow-water biotopes within a single bay, viz. notches in fossil reef rock, mangroves, fossil reef boulders, seagrass beds, algal beds at a depth of 2 m, algal beds at a depth of 5 m, and the channel. The study was done in an inland bay on the Caribbean island of Curaçao, using a visual census technique. Total fish densities within the different feeding guilds varied considerably between the biotopes, and were generally higher in the reef biotopes and on the boulders than in the remaining bay biotopes. Cluster analysis revealed that the greatest dissimilarity in guild structures in terms of fish densities was that between the algal beds and all other biotopes, followed by that between the reef depth zones and other bay biotopes (notches, mangroves, seagrass beds, channel). The species composition of the guilds also differed considerably among the various biotopes. Species richness within the various guilds showed much smaller differences between the biotopes, but was generally somewhat higher in the reef biotopes. Cluster analysis of guild structures in terms of species richness revealed little dissimilarity among the various biotopes. The coral reef was dominated by omnivores and zooplanktivores, whereas the bay was dominated by zoobenthivores and herbivores. Differences in guild structure between the bay and the adjacent reef indicate differences in food availability.     

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.     

Experimental effects of habitat fragmentation on old-field canopy insects: community, guild and species responses

We examined the effects of habitat fragmentation on the species distributions, guild membership, and community structure of old-field insects using a fine-scale experimental approach. A continuous 1-ha goldenrod field was fragmented into four treatments that varied in both patch size and degree of isolation. Each treatment was replicated four times and arranged in a Latin square design. Canopy insects in fragmented patches were sampled with sweep nets during early and late summer 1995. The species richness of insects was significantly lower in fragmented than in unfragmented treatments during July, but was similar among treatments in September. Overall community abundance showed no treatment effect during either month. We also found significant row and column effects, suggesting there was spatial heterogeneity in species richness and abundance apart from treatment effects. Differences in species richness during July were primarily due to the loss of rare species in highly fragmented plots. Overall abundance was less responsive to community change because deletions of rare species in fragmented areas were not detected in abundance analyses. Four feeding guilds showed different responses to fragmentation: the species richness of sucking herbivores and the abundance of parasitoids were significantly reduced by fragmentation but predators and chewing herbivores were largely unaffected. Analyses of a subset of individual species within guilds suggest that the greater effects of fragmentation on sucking herbivores and parasitoids may be due to the degree of habitat specificity of guild members. The effects of small-scale habitat fragmentation were therefore detectable at the level of community, guild, and individual species. Changes in species richness, guild structure and species distributions were likely due to differential effects of habitat alteration on individual movements and patch selection rather than dispersal or demographic change. Nonetheless, the selective loss of rare species, differential guild effects and changes in species occupancy that we found in this small-scale experiment are also factors that are likely to operate in fragmented habitats over broader spatial scales. Received: 11 May 1998 / Accepted: 27 September 1998     

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.     

Two lowland stream riffles – linkages between physical habitats and macroinvertebrates across multiple spatial scales

The physical structure of two riffles in a lowland Danish stream was studied and its importance for the composition and density of the macroinvertebrate communities was evaluated. The two riffles were visually assessed to be very similar, but measurements revealed that they differed in overall hydraulic conditions, stability, substratum composition and consolidation. Differences affected abundance of both burrowing and surface dwelling macroinvertebrates. The unstable unconsolidated riffle had higher total macroinvertebrate abundance (4137 m⁻² vs. 1698 m⁻²), diptera abundance (2329 m⁻² vs. 386 m⁻²) and total estimated species richness (31.7 vs. 28.8) as well as lower evenness (0.77 vs. 0.83) than the compact riffle. Among samples within the unconsolidated riffle, variations in macroinvertebrate communities were related to differences in mean substratum particle size. Here a linear log–log relationship existed between macroinvertebrate abundance, the abundance of EPT taxa and the median particle size (r ² _total = 0.46, p = 0.002; r ² _EPT = 0.73, p < 0.001). No similar relationships were evident on the consolidated riffle. Moreover, macroinvertebrate communities on the unconsolidated riffle were dominated by species with a high colonising potential. Despite being assessed to the same morphological unit, physical variation between riffles was surprisingly high as the riffles differed substantially with respect to consolidation, substratum heterogeneity and overall hydraulic structure. Macroinvertebrate community structure and composition also differed between riffles despite being drawn from the same species pool. The findings address the question if we use the correct methods and parameters when assessing the macroinvertebrate communities at the scale of the morphological unit.     

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.     

How selection structures species abundance distributions

How do species divide resources to produce the characteristic species abundance distributions seen in nature? One way to resolve this problem is to examine how the biomass (or capacity) of the spatial guilds that combine to produce an abundance distribution is allocated among species. Here we argue that selection on body size varies across guilds occupying spatially distinct habitats. Using an exceptionally well-characterized estuarine fish community, we show that biomass is concentrated in large bodied species in guilds where habitat structure provides protection from predators, but not in those guilds associated with open habitats and where safety in numbers is a mechanism for reducing predation risk. We further demonstrate that while there is temporal turnover in the abundances and identities of species that comprise these guilds, guild rank order is conserved across our 30-year time series. These results demonstrate that ecological communities are not randomly assembled but can be decomposed into guilds where capacity is predictably allocated among species.     

Fish zonations and guilds as the basis for assessment of ecological integrity of large rivers

Longitudinal zonation concepts describe the downstream changes in chemico-physical and biological properties of rivers. Including information on ecological fish guilds can enhance the usefulness of fish zonation concepts, in a way that they can be used as tools for assessment and management of the ecological integrity of large rivers. We present an ecological characterization of fish zones and fish communities in near-natural and in regulated large rivers in Europe (the River Doubs in France and the Rivers Rhine and Meuse in the Netherlands), using guild classifications of several life-history traits of fish and national Red Lists of threatened species. The Doubs data set was also analyzed using indices of the sensitivity of fish species to environmental degradation and indices for eurytopy. In these rivers, the number of ecological guilds per zone increases downstream, and there are clear shifts in the structure of the guilds. Flow preference and reproduction ecology of river fish are closely linked. The proportion of rheophilic species in the fish community decreases downstream, and the proportions of limnophilic and eurytopic species increase. Lithophilic and psammophilic spawners are dominant in the upper zones, whereas the lower zones are dominated by phytophilic and phytolithophilic spawners. The proportion of zoobenthivorous and periphytivorous species decreases downstream, and the proportion of zooplanktivorous and phytivorous species increases. However, because the European fish fauna mainly consists of feeding generalists, the discriminative abilities of simplistic feeding guild classifications are not very high. Guilds of sensitive, stenoecious species that share life history strategies that are highly adapted to specific riverine conditions (rheophils and limnophils) have declined far more than generalist species that can survive in a wide range of habitats that are not characteristic of natural river ecosystems. Because of the subsequent over-abundance of the eurytopic species the original longitudinal fish zonations are hardly recognizable anymore in heavily impacted large rivers such as the River Rhine. Hence these rivers do not meet the criteria for ecological integrity. Within a specific fish region, a suitable way of analyzing and monitoring the impact of human disturbance on the structure of the fish community is by comparing the guild structure of the present state of a fish zone with that of the reference situation.     

Developing environmental flow recommendations for freshwater mussels using the biological traits of species guilds

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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.     

Habitat Use of Fish Communities in A Virginia Stream System

Fish habitat use during summer was examined at micro- and meso-levels, to determine species associations in the upper Roanoke River watershed, Virginia. Based on multivariate-mesohabitat analysis and examination of mean microhabitat use, seven habitat-use guilds were apparent. These included four rheophilic (fast-riffle, riffle/run, fast-generalist, and shallow-rheophilic) and three limnophilic guilds (pool/run, open-pool, and pool-cover) that were reasonably robust across two river segments and two years. Although simple-hydraulic, bottom-topographic, and turbulence variables all segregated fish habitat-use guilds, turbulence variables were redundant with simple-hydraulic variables, substratum use by limnophilic fishes was related to availability, and only one guild consistently selected high cover levels. At the family level, suckers, darters, and especially minnows were notable for occupying several habitat-use guilds, because of species differences in habitat preferences. Such formulation of guilds can simplify habitat-impact analyses in biodiverse, warmwater streams, via focus on habitat needs of guilds rather than on individual species.     

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.     

The effects of fire on communities,guilds and species of breeding birds in burnt and control pinewoods in central Italy

Breeding bird communities in burnt and unburnt residual pinewoods were studied over 3 years by line-transect method, following a catastrophic fire event in Castelfusano (Rome, Central Italy; July 2000). We applied bootstrap procedures to evaluate whether the observed data were true or just produced by chance, and then examined the emerging patterns at three levels: community, guild and species levels. At the community level, fire acted on breeding bird communities by altering especially the total abundance patterns: the species abundance decreased in the burnt pinewood compared to the residual one, but other parameters were not significantly affected by fire. As a consequence of fire, the destruction and structural simplification of the canopy and shrubby component, as well as the increase of edge habitat and patchiness at landscape scale, induced a turnover in species between pinewoods. Species turnover was higher at the burnt than at the residual pinewoods, during all the 3 years of study. At the guild level, the forest species decreased strongly in terms of richness and abundance in the burnt pinewoods, contrary to the edge and open habitat species which increased in terms of richness, abundance and evenness. Edge species showed the highest turnover in burnt pinewood during the whole period of study. At species level, after an a priori subdivision (based on bibliographic search) of the various species in two ecological guilds (forest versus edge species), it was found that an a posteriori statistical analysis confirmed the expected trend, i.e. that the species which decreased significantly in burnt pinewood were essentially the forest species, whereas the species which increased were essentially the edge/open habitat ones. Overall, in order to investigate the effects of fire catastrophes on birds, the guild approach seems more exhaustive than the taxonomic community approach, where intrinsic confounding trends are present.     

Effects of siltation on stream fishes and the potential mitigating role of the buffering riparian zone

Riparian buffer zones serve several important roles in linking a stream to its watershed. A main function is controlling the dynamics of sedimentation. This paper documents how siltation impacts fish communities and how proper riparian management can mitigate the negative effects of sedimentation. Two studies examined the relation between stream siltation and fish community characteristics. Community responses to siltation were poorly described by common structural indices. Community level responses to varying siltation were most consistently described by changes in functional characteristics of the resident fish species using a guild-based analysis. Herbivores, benthic insectivores and simple lithophilous spawners were most sensitive to siltation while other guilds were not. These results were repeatable in both intraregional comparisons among sites of similar size and character, and in interregional comparisons of streams which varied in characteristics besides siltation. This suggests the index may be useful in separating the effects of siltation from other environmental variables. A discussion of bufferstrip characteristics important in mitigating against, or preventing, excess siltation is presented.     

Using dung beetles to evaluate the effects of urbanization on Atlantic Forest biodiversity

We used dung beetles to evaluate the impact of urbanization on insect biodiversity in three Atlantic Forest fragments in Londrina, Paraná, Brazil. This study provides the first empirical evidence of the impact of urbanization on richness, abundance, composition and guild structure of dung beetle communities from the Brazilian Atlantic Forest. We evaluated the community aspects (abundance, richness, composition and food guilds) of dung beetles in fragments with different degrees of immersion in the urban matrix using pitfall traps with four alternative baits (rotten meat, rotten fish, pig dung and decaying banana). A total of 1 719 individuals were collected, belonging to 29 species from 11 genera and six Scarabaeinae tribes. The most urban‐immersed fragment showed a higher species dominance and the beetle community captured on dung presented the greatest evenness. The beetle communities were distinct with respect to the fragments and feeding habits. Except for the dung beetle assemblage in the most urbanized forest fragment, all others exhibited contrasting differences in species composition attracted to each bait type. Our results clearly show that the degree of urbanization affects Atlantic Forest dung beetle communities and that the preservation of forest fragments inside the cities, even small ones, can provide refuges for Scarabaeinae.     

Disturbance,fish, and variation in the predatory insect guild of coastal streams

Species distributions, and thus the composition of communities, are determined by many interacting biotic and abiotic factors. We analyzed the variation in composition of the invertebrate predator guild across eight small, steep coastal streams in eastern Canada, in relation to variation in several broad categories of environmental variables: disturbance-related physical characteristics, top-down effects (fish), and bottom-up effects (prey composition and productivity-related physical/chemical variables). Similarity in composition (relative and absolute abundances of the 19 species belonging to the Rhyacophilidae, Perlodidae, and Chloroperlidae) declined significantly with distance. Streams that were most similar in predator composition were also most similar in physical factors related to disturbance, but were not more similar in prey abundance/composition or in environmental factors associated with productivity (chemistry, canopy cover, and riparian forest type). Similarity in the relative abundance of species within the Rhyacophilidae was linked to variation in the presence/abundance of brook trout. These results suggest that the invertebrate predator guilds of these small coastal streams are structured first and foremost by factors associated with the magnitude and variability of the flow regime, and second by response to top-down factors (predation by brook trout). There was no evidence that bottom-up effects altered the composition or abundance of the predator guild. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Sonja Stendera     

Caterpillar assemblages on Chusquea bamboos in southern Ecuador: abundance,guild structure,and the influence of host plant quality

1. Information on the guild structure of foliage‐associated tropical insects is scarce, especially as caterpillars are mostly considered only as herbivores feeding on living leaves. However, many caterpillar species display alternative trophic associations, feeding on dead or withered leaves or epiphylls (‘non‐herbivores’). 2. To determine the contribution of these non‐herbivores, caterpillar communities associated with Chusquea Kunth (Poaceae) in the Andes of southern Ecuador were investigated. Caterpillars were collected at two elevation levels (montane rainforest ～2000 m and elfin forest at ～3000 m a.s.l.) and assigned to three feeding guilds (strict herbivores, non‐herbivores, and switchers) based on feeding trials. Foliage quality and leaf area were recorded to test for their influence on guild composition and caterpillar density. 3. Three hundred and eighty‐nine individuals belonging to 175 Lepidoptera species associated with Chusquea bamboos were found. The species richness of caterpillars was similarly high at both elevation levels but varied between feeding guilds. Approximately half (46.5%) displayed an alternative feeding association, i.e. were non‐herbivores (31.1%) or switchers (15.4%). 4. Caterpillar density was nearly two‐fold higher in the elfin forest, but only strict herbivores and switchers increased significantly with elevation. Leaf area positively influenced the density of strict herbivores and switchers; foliage quality only affected strict herbivores. The density of non‐herbivores did not differ significantly between the forest types and was not related to leaf area or foliage quality. 5. The present study underpins that non‐herbivores make up a considerable fraction of caterpillar communities in tropical mountain ecosystems and demonstrates that elevation, foliage quality and available plant biomass further shape feeding guild composition.