Comparative analysis of feeding habits and dietary niche breadth in skates: the importance of body size,snout length,and depth

Skates (Elasmobranchii, Rajiformes) are a morphologically conservative group of bentophagous chondrichthyan fishes with a high degree of endemism, that occur on marine soft bottoms. Subtle morphological aspects and bathymetric distribution are traits that vary among skate species that could have implications for their feeding ecology. We test how body size, snout length and bathymetric distribution influence the feeding habits and dietary niche breadth in skates using data on 71 species taken from the literature. We hypothesized that snout length has an effect on diet composition. We also hypothesized that dietary niche breadth increases with increasing depth range and decreases with increasing body size of skate species. Generalized additive models for location scale and shape were fitted with taxonomic level (genera nested within family) included as a random effect term in each model. A model selection approach to test the level of support for alternative models was applied. We found that skate species that forage on large prey have the largest body size and skate species with the smallest body size prey on small and medium-sized invertebrates. The results indicated that body size has an effect on feeding habits of skates, whereas an effect of snout length was not supported. Bathymetric variables have an effect on the diet of skates. Our prediction that dietary niche breadth increases with increasing depth range and decreases with increasing body size of skate species was supported in part: in a first phase the relationship between dietary niche breadth and body size is positive, then in a second phase, including species larger than 1000 mm total length, the relationship become negative.     

Feeding habits and trophic ecology of Dasyatis longa (Elasmobranchii: Myliobatiformes): sexual, temporal and ontogenetic effects

Sexual, ontogenic and temporal effects in the diet of Dasyatis longa were evaluated to determine feeding habits and trophic ecology. Numeric indices and the index of relative importance were applied to establish the feeding strategy of the species. Independence of the diet with respect to sex, dry or rainy season and size was evaluated with contingency tables, correspondence analysis and multivariate analysis (MANOVA). The trophic relationships of D. longa (by sex and size intervals) were determined using Levin's niche breadth index and the Pianka's diet overlap index and their significance was determined by null models. The trophic level for each size interval and the species was also calculated. Dasyatis longa showed a narrow niche breadth feeding mainly on shrimps and fishes and its diet was dependent on size, but not on sex or season. Juvenile individuals (class I) fed on shrimps, sub-adults (class II) on fishes and adults (class III) on stomatopods. Significant overlaps between size classes I and II and classes II and III were found. The trophic level shows D. longa to be a secondary or tertiary consumer. Due to considerable fishing pressures on shrimps, the principal prey of D. longa, it will be important to determine their ability to adapt to changes in prey populations.     

Modulation and variability of prey capture kinematics in clariid catfishes

Species with narrow or limited diets (trophic specialists) are expected to be less flexible in their feeding repertoire compared to species feeding on a wide range of different prey (trophic generalists). The ability to modulate prey capture kinematics in response to different prey types and prey position, as well as the overall variability in prey capture kinematics, is evaluated in four clariid species ranging from trophic generalist (Clarias gariepinus) to species with morphological specializations and a narrow diet (Channallabes apus and Gymnallabes typus). High-speed video recordings were made of prey captures on two prey that differ in shape, attachment strength and hardness. While the observed amount of strike-to-strike variability in prey capture kinematics is similar for all species and not influenced by prey type, only the two less specialized species showed the ability to modulate their prey capture kinematics in function of the presented prey types. All species did, however, show positional modulation during the strike by adjusting the magnitude of neurocranial elevation. These results indicate that the narrow dietary breadth of trophic specialists is indeed indicative of functional stereotypy in this group of fishes. Although most studies focussing on prey processing found a similar result, the present study is one of the few that was able to demonstrate this relationship when focussing on prey capture mechanics. Possibly, this relationship is less frequently observed for prey capture compared to prey processing because, regardless of prey type, the initial capture of prey requires a higher amount of variability.     

Eco-evolutionary trophic dynamics: loss of top predators drives trophic evolution and ecology of prey

Ecosystems are being altered on a global scale by the extirpation of top predators. The ecological effects of predator removal have been investigated widely; however, predator removal can also change natural selection acting on prey, resulting in contemporary evolution. Here we tested the role of predator removal on the contemporary evolution of trophic traits in prey. We utilized a historical introduction experiment where Trinidadian guppies (Poecilia reticulata) were relocated from a site with predatory fishes to a site lacking predators. To assess the trophic consequences of predator release, we linked individual morphology (cranial, jaw, and body) to foraging performance. Our results show that predator release caused an increase in guppy density and a "sharpening" of guppy trophic traits, which enhanced food consumption rates. Predator release appears to have shifted natural selection away from predator escape ability and towards resource acquisition ability. Related diet and mesocosm studies suggest that this shift enhances the impact of guppies on lower trophic levels in a fashion nuanced by the omnivorous feeding ecology of the species. We conclude that extirpation of top predators may commonly select for enhanced feeding performance in prey, with important cascading consequences for communities and ecosystems.     

Feeding ecology of pelagic fish species in the Gulf of Riga (Baltic Sea): the importance of changes in the zooplankton community

The feeding ecology of four pelagic fish species was studied in relation to their prey availability in the Gulf of Riga (Baltic Sea) during the summer 1999-2006. The zooplankton community was dominated by the cladoceran Bosmina longispina, rotifers Keratella cochlearis and K. quadrata and the copepod Eurytemora affinis, with the highest interannual variability in abundance recorded for B. longispina. The last influenced the diet of adult sprat Sprattus sprattus, juvenile smelt Osmerus eperlanus and three-spined stickleback Gasterosteus aculeatus as these were strongly selecting for B. longispina. The fish feeding activity did not match the abundance dynamics of their preferred prey, suggesting that fishes may switch to consume other prey in case the preferred diet was limited. A considerable dietary overlap indicated high potential competition between pelagic fish species. While herring Clupea harengus membras and G. aculeatus were relying on very different food, the diets of young O. eperlanus and G. aculeatus were very similar. Interannual variability in zooplankton composition and abundance significantly affected the diet composition of fishes, but those changes were insufficient to exert a consistent influence upon fish feeding activity and total amounts of zooplankton consumed.     

Trophic behaviour of juvenile reef fishes inhabiting interlinked mangrove–seagrass habitats in offshore mangrove islets

Stable isotope (δ¹³C and δ¹⁵N) and gut content analyses were used to investigate size‐related feeding habits of four reef fishes (the beaugregory Stegastes leucostictus, the french grunt Haemulon flavolineatum, the schoolmaster snapper Lutjanus apodus and the yellowtail snapper Ocyurus chrysurus) inhabiting an offshore (non‐estuarine) mangrove islet off Belize, Central America. Comparisons of isotopic niche space and Schoener diet similarity index suggested a low to moderate degree of niche overlap between fish size groups. The δ¹³C gradient between mangrove and seagrass prey as well as results of Bayesian mixing models revealed that sampled fishes relied mostly on seagrass prey items. Only small and large juveniles of the carnivorous species L. apodus derived a part of their diet from mangroves by targeting mangrove‐associated Grapsidae crabs and fish prey, respectively. Isotopic niche shifts were particularly obvious for carnivorous fishes that ingested larger prey items (Xanthidae crabs and fishes) during their ontogeny. The utilization of mangrove food resources is less than expected and depends on the ecology and life history of the fish species considered. This research highlights that mangrove‐derived carbon contributed relatively little to the diets of four fish taxa from an offshore mangrove islet.     

Feeding ecology and trophic segregation of two sympatric mesopredatory sharks in the heavily exploited coastal ecosystem of the Adriatic Sea

The ecological roles and trophic interactions of two commercially important mesopredatory shark species, Squalus acanthias and Mustelus punctulatus that co‐occur on the continental shelf of the north‐central Adriatic Sea were investigated. Both shark species are dietary specialists, with a significant dietary overlap recorded only during the spring season. They showed different patterns of feeding as they grew: S. acanthias extended its trophic niche with an increase in size, while M. punctulatus developed a more specialized diet. These two sharks partition food resources and reduce niche overlap by foraging at different trophic levels. Mustelus punctulatus is a crustacean feeder, specialized in foraging on scavenging malacostracans frequently found along trawl tracks or on discards in the Adriatic fishing zone. Conversely, S. acanthias prefers small pelagic fishes, which are commercially exploited and in decline. The different foraging strategies adopted by these two species suggest that they should be managed separately. Dietary specialization, direct competition with humans for prey and their higher intrinsic vulnerability make S. acanthias particularly susceptible to the effects of anthropogenic perturbations.     

Diet and feeding strategy of blackmouth catshark Galeus melastomus

Diet and feeding strategy of the blackmouth catshark Galeus melastomus in the deep waters of the eastern Ionian Sea were investigated. Sampling was carried out using experimental bottom longline fishing at depths ranging from 300 to 855 m in summer and autumn 2010. Diet variability with fish size, season, area, sex and depth zone was tested and only season was found to significantly affect the diet of the species. Of the 870 stomachs examined, only 1·4% were empty. Cumulative prey curves showed that the sample sizes were adequate to describe the main prey items of the diet for both seasons. Prey identified belonged primarily to three major groups: fishes, cephalopods and dendrobranchiatan and caridean shrimps. In autumn, the above three major groups were found as principal prey, whereas in summer cephalopods followed by fishes were the principal prey and shrimps were found as secondary prey. SIMPER analysis indicated high dissimilarity between seasons and highlighted that Sepiolidae, Myctophidae, fishes, cephalopods, shrimps and other crustaceans contributed to seasonal differences in the diet of G. melastomus. The prey diversity index was higher in autumn than in summer. A high dietary overlap was observed between the two seasons. Galeus melastomus behaved as an opportunistic predator with a variety of unimportant prey in its diet. Feeding strategy indicated that G. melastomus holds a generalist niche with a considerable specialization at the individual level. This strategy seems to be an adaptation to a food‐scarce environment, as typified in deep‐water habitats.     

Food and feeding ecology of piscivorous fishes at Lake St Lucia, Zululand

The food and feeding ecology of piscivorous fish in Lake St Lucia was monitored for two years. Piscivorous fishes feed predominantly on the planktivorous Gilchristella aestuarius and Thryssa vitrirostris but a wide range of prey species was recorded. Numbers of the predominant piscivores, Argyrosomus hololepidotus and Elops machnata , in an area appear to be related to the densities of their major prey, T. vitrirostris and G. aestuarius . Large piscivorous fishes are restricted to the deeper portions of the lake, whereas small piscivores such as Johnius belengerii and Terapon jarbua feed predominantly on small fishes in the littoral zone. The highly significant correlation between the composition of prey fish species in the lake and prey fish species in the diet of piscivorous fishes, indicates that piscivores are feeding in a density dependent manner. However, factors such as habitat, fish size and swimming speed of prey species are shown to be important in prey selection. Juvenile fish of species such as Sarotherodon mossambicus, Liza macrolepis and Acanthopagrus berda remain in shallow marginal areas, thus avoiding large piscivorous fishes. However by frequenting shallow areas these species become vulnerable to bird predators, especially egrets and herons.     

Competition versus opportunism: Diet and trophic niche relationship of two sympatric apex carnivores in a tropical forest

There is a little understanding of how apex carnivores partition their diet to coexist. We studied food habit and trophic niche overlap of two apex carnivores, tiger and leopard in the tropical forest of Similipal tiger reserve, eastern India. We used line transect and scat analysis method, to estimate the prey availability and determine the diet and prey selection of two apex carnivores. Tigers consumed mostly large and medium-sized prey, whereas leopards mostly consumed medium and small-sized prey. Both carnivores were not random in their consumption of prey, sambar and wild pigs were selectively consumed by tigers, whereas leopards selectively consumed wild pigs, barking deer and mouse deer. Dietary overlap between two carnivore species was moderate (Pianka's niche overlap index: 0.55), and trophic niche breadth (Levin's standardized niche breadth) of the leopard (0.52) was greater than that of the tiger (0.37). Overall, tigers exhibited specialized feeding habits, whereas leopards showed generalist feeding habits. Our study highlights the opportunistic nature of leopards and probably a reason for the species successfully coexist with tigers.     

Feeding patterns of the dominant benthic and demersal fish community in a temperate estuary

The diet and feeding relationships of the 12 most abundant benthic and demersal fish species of the Mondego Estuary, Portugal, were studied between June 2003 and May 2006. Fishes were caught during the night using a 2 m beam trawl. The stomach contents were analysed for prey identification, counting and weighing. According to prey importance in diets, three main feeding guilds were identified: (1) invertebrate feeders, (2) invertebrate and fish feeders and (3) plankton and invertebrate feeders. Besides these main feeding guilds, some fishes also presented fractions of algae and zooplankton in their stomach contents. The most abundant prey items were macroinvertebrates, with several polychaetes ( Nephtys spp., Capitellidae, Spionidae and Eunicidae), Corophium spp. and Crangon crangon among the dominant prey. Pomatoschistus spp. were the most preyed on fishes. Several fish species showed a tendency for a specialized diet, but almost all also showed some degree of opportunistic feeding by preying on other food resources. High diet overlap was found between some fish species, yet exploitative competition could not be concluded.     

The foraging ecology of larval and juvenile fishes

Knowledge of the foraging ecology of fishes is fundamental both to understanding the processes that function at the individual, population and community levels, and for the management and conservation of their populations and habitats. Furthermore, the factors that influence the acquisition and assimilation of food can have significant consequences for the condition, growth, survival and recruitment of fishes. The majority of marine and freshwater fish species are planktivorous at the onset of exogenous nutrition and have a limited ability to detect, capture, ingest and digest prey. Improvements in vision, development of fins and associated improvements in swimming performance, increases in gape size and development of the alimentary tract during ontogeny often lead to shifts in diet composition. Prey size, morphology, behaviour and abundance can all influence the prey selection of larval and juvenile fishes. Differences in feeding behaviour between fish species, individuals or during ontogeny can also be important, as can inter- and intraspecific interactions (competition, predation risk). Temporal (diel, seasonal, annual) and spatial (microhabitat, mesohabitat, macrohabitat, regional) variations in prey availability can have important implications for the prey selection, diet composition, growth, survival, condition and, ultimately, recruitment success of fishes. For fish populations to persist, habitat must be available in sufficient quality and quantity for the range of activities undertaken during all periods of development. Habitats that enhance the diversity, size ranges and abundance of zooplankton should ensure that sufficient food resources are available to larval and juvenile fishes.     

Niche overlap and diet composition of three sympatric coastal dolphin species in the southwest Atlantic Ocean

Sympatric species are expected to differ in ecological requirements to minimize niche overlap and avoid competition. Here we assess the trophic interactions among three coexisting dolphin species from southern Brazil: the franciscana dolphin (Pontoporia blainvillei), the Guiana dolphin (Sotalia guianensis), and the Lahille's bottlenose dolphin (Tursiops truncatus gephyreus). We evaluated temporal variation in carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope values of bone collagen to examine potential dietary shifts resulting from increased fishing activity over the past three decades. We estimated the degree of niche overlap among these species and the contribution of potential prey sources to their diet. δ¹⁵N values were consistent among species and across years, while δ¹³C values increased for Guiana dolphins and decreased for bottlenose dolphins, suggesting changes in diet and/or foraging habitats through time. The similar δ¹³C and δ¹⁵N values and the high niche overlap between Guiana and bottlenose dolphins indicate that these species are primarily feeding on demersal prey. The franciscana diet is primarily composed of pelagic prey, resulting in a lower niche overlap in comparison with the other dolphin species. Our study provides further information about the foraging ecology of this unique dolphin community in southern Brazil with implications for its management and conservation.     

Trophic resource partitioning among five flatfish species (Actinopterygii, Pleuronectiformes) in a tropical bay in south-eastern Brazil

Five species of flatfishes in a tropical bay in south-eastern Brazil were studied to test the hypothesis that resource partitioning along the spatial and size dimensions has been used as a mechanism to enable coexistence. Three zones in the study area were defined according to environmental characteristics (inner, middle and outer). Sampling was conducted by otter trawl tows during daylight hours, between October 1998 and September 1999. Achiridae species ( Achirus lineatus and Trinectes paulistanus ) showed narrow niche width, indicating a specialized feeding strategy, preying on Polychaeta, and occurring mainly in the inner bay zone. Paralichthyidae ( Citharichthys spilopterus and Etropus crossotus ) and the Cynoglossidae ( Symphurus tessellatus ) showed broad niche width and a generalized feeding strategy preying on a large number of Crustacea. Symphurus tessellatus did not change diet with size-dimension feeding on Amphipoda and Polychaeta, whereas C. spilopterus and E. crossotus shifted diet with growth. Citharichthys spilopterus fed mainly on Mysida and secondarily on shrimps, with juvenile preying on large amounts of Calanoida, whereas adults consumed large amounts of fishes. Isaeidae amphipods were a significant prey for both small and large E. crossotus , whereas Polychaeta Errantia were used mainly by large fishes. Etropus crossotus and S. tessellatus share similar feeding resources in outer bay zone preying on Isaeidae and Polychaeta Errantia. Differences in the Pleuronectiformes diet composition along with spatial and size changes in the use of the available resources contributed to allow the organisms' coexistence in Sepetiba Bay. The high item diversity used by flatfishes indicates that the system plays an important role as a feeding ground, and that interspecific competition for food was unlikely.     

山东近海三种底层鱼类的摄食生态与食物竞争

研究根据2016—2017年在山东近海进行的四个季节的渔业资源底拖网调查和胃含物分析数据, 应用分类树和生态位重叠指数法等, 研究了该海域高眼鲽(Cleisthenes herzensteini)、黄鮟鱇(Lophius litulon)和小眼绿鳍鱼(Chelidonichthys spinosus)3种主要底层鱼类的食物组成及其营养-空间二维生态位的重叠情况, 并分析环境因素对其摄食习性的影响, 旨在为该海域渔业资源的可持续利用和科学管理提供基础资料。研究发现, 这3种底层鱼类均主要以虾类和鱼类为食, 属于底栖动物食性。3种鱼类的摄食习性存在一定的差异, 可以通过构建分类树将其进行区分。通过CCA分析发现, 这3种鱼类的摄食习性受不同的环境因素影响, 其中高眼鲽的摄食习性主要与其体长有关, 随着体长的增加, 高眼鲽会摄食更多的鱼类和虾类; 小眼绿鳍鱼的摄食主要与季节有关, 不同季节小眼绿鳍鱼摄食鱼类和虾类的比例不同; 而黄鮟鱇的摄食主要受空间分布的影响, 高纬度的黄鮟鱇摄食更多的虾类, 而随着经度的增加, 黄鮟鱇会摄食更多的鱼类。黄鮟鱇的营养生态位和空间生态位宽度均最高, 而其余两种鱼的营养和空间生态位宽度相对较小。山东近海这3种鱼类的营养生态位重叠指数都较高, 但是空间生态位重叠指数较小, 在一定程度上可以缓和它们之间的种间竞争。     

Generalist niche, specialist strategy: the diet of an Australian percichthyid

Dietary analysis revealed that an impoundment population of Australian bass Macquaria novemaculeata holds a generalist niche, but one arising from persistent individual specialization and interindividual variation. This 'individual specialist' strategy appeared adaptive, but the strength of individual specialization was largely independent of variation in diet composition, except during blooms of Daphnia sp. Diet composition and dietary overlap showed only moderate ontogenetic variation, and niche breadth showed no relationship with ontogeny. Macquaria novemaculeata showed an asymmetric predator and prey size distribution, consistent with many aquatic predators, with positive relationships between fish size and average, maximum and minimum prey size. There was no asymmetry in the relative size-based niche breadths of individuals, however, which indicates that the niche is a fixed 'window' of relative prey sizes. The difference in the dietary niche and prey-size relationships of M. novemaculeata at the population and individual levels highlights the necessity of assessing the niche at both these levels.     

Influence of dietary specialization and resource availability on geographical variation in abundance of butterflyfish

Empirical evidence indicates that both niche breadth and resource availability are key drivers of a species' local abundance patterns. However, most studies have considered the influence of either niche breath or resource availability in isolation, while it is the interactive effects that are likely to influence local abundance. We examined geographic variation in the feeding ecology and distribution of coral-feeding butterflyfish to determine the influence of dietary specialization and dietary resource availability on their local abundance. Dietary composition and abundance of five butterflyfish and coral dietary resource availability were determined at 45 sites across five locations (Lizard Island and Heron Island, Great Barrier Reef; Kimbe Bay, Papua New Guinea; Noumea, New Caledonia; and Moorea, French Polynesia). Multiple regression models using variables representative of total dietary resource availability, availability of specific dietary resources, and interspecific competition were used to determine the best predictors of local abundance across all sites and locations for each species. Factors influencing local abundance varied between butterflyfish with specialized and generalized diets. Dietary resource availability had the strongest influence on the abundance of Chaetodon trifascialis-the most specialized species. Local abundance of C. trifascialis was best predicted by availability of the Acropora corals that it preferentially feeds on. In contrast, abundance of generalist butterflyfish was poorly described by variation in availability of specific resources. Rather, indices of total dietary resource availability best predicted their abundance. Overall, multiple regression models only explained a small proportion of the variation in local abundance for all five species. Despite their relatively specialized diets, dietary resource availability has limited influence on the local abundance of butterflyfish. Only the most specialized species appear to be consistently limited by prey availability. Local and total abundance of species are influenced by a wide range of different factors and there is definite need to conduct independent species assessments.     

Food niche variation of European and American mink during the American mink invasion in north-eastern Belarus

Understanding processes allowing the co-existence of ecologically similar species is important but difficult to study in community ecology. Introductions of alien species are unplanned experiments allowing investigation of co-adaptation of both native and invasive species over a short period. We analysed food niche differentiation between native European mink and alien American mink after invasion of the latter species in Belarus. European mink feed mainly on crayfish, frogs and fish whereas American mink prefer small mammals, fish and frogs. The diet of both species varied between seasons and during the period of alien mink invasion. Concurrent with the progress of American mink invasion, the European mink food niche has narrowed to feeding mainly on frogs, with the proportion of aquatic prey (fish and crayfish) in their diet drastically reduced. In contrast, the American mink food niche became wider during invasion. The breadth was stable but included a varied proportion of different prey categories: namely an increased proportion of aquatic prey and a decreased proportion of water vole and waterfowl. The increase in abundance of American mink saw a decrease in the proportion of larger prey in their diet. When American mink preyed more often on frogs, food niche overlap of both predators increased. This result suggests that arrival of an alien competitor reduced food abundance (exploitative competition) and caused a change in native mink diet.     

Networks of prey specialization in an Arctic monomorphic seabird

Generalist predator populations are sometimes made up of individuals that specialize on particular prey items. To examine specialization in thick‐billed murres Uria lomvia during self‐feeding we obtained stomach contents and muscle stable isotope values for 213 birds feeding close to five colonies in the Canadian Arctic. Adults were less specialized during self‐feeding than during chick‐provisioning. Nonetheless, particular specialists clustered together within the foraging network. While sexes showed similar levels of specialization, individuals of the same sex clustered together within the foraging network. The significant degree of clustering regardless of sex showed that individuals specializing on one prey item tend to also specialize on another, although network topology varied from colony to colony. Adult muscle stable isotope values correlated with the stable isotope values of the prey found in stomachs, at least at the one colony with relevant prey data, suggesting that specializations are maintained over time. Degree of specialization increased with niche width across the five colonies, but similarity in gastro‐intestinal and bill morphology was independent of dietary similarity. Thus, although individual specialization is thought to play a key role in sympatric speciation through trophic specialization, we found no support for an association between morphology and foraging patterns in our species. We conclude that self‐feeding murres show clustered dietary specialization, and that specialization is highest where diet is most diverse.     

Ontogenetic diet shifts of rough scad Trachurus lathami in the North Patagonian shelf (south-west Atlantic Ocean)

Rough scad Trachurus lathami is a key pelagic fish in the Argentinean continental shelf (ACS, south-west Atlantic Ocean), with recent increases in abundance. It is a main prey of fishes and marine mammals, and shares the environment with commercially relevant pelagic species (Engraulis anchoita and Scomber colias), playing an important role linking lower and upper trophic levels in the ecosystem. This study aims to determine the ontogenetic changes in the diet composition, feeding strategy, trophic niche breadth and trophic level of T. lathami in the North Patagonian Shelf (43°–45°30′S). The stomach contents of adult fish (n = 238) were analysed. The results suggest a clear ontogenetic shift in the diet at a size of ~190 mm. Smaller individuals (160–190 mm) were specialized on misidaceans, and showed the highest trophic level, while larger T. lathami (221–230 mm) consumed decapods (Peisos petrunkevitchi) and teleosts (eggs and larvae). Trophic niche breadth was higher at the medium-sized class (191–220 mm), which mainly preyed on copepods (Calanoides carinatus) and chaetognaths (Sagitta spp.), evidencing a more diverse diet and a rather generalist strategy. Updated information on the trophic ecology of T. lathami evidences its extremely plastic feeding behaviour, being able to adapt its trophic niche to the most readily available food items from the mesopelagic community.