Comparison of size-structured and species-level trophic networks reveals antagonistic effects of temperature on vertical trophic diversity at the population and species level

It is predicted that warmer conditions should lead to a loss of trophic levels, as larger bodied consumers, which occupy higher trophic levels, experience higher metabolic costs at high temperature. Yet, it is unclear whether this prediction is consistent with the effect of warming on the trophic structure of natural systems. Furthermore, effects of temperature at the species level, which arise through a change in species composition, may differ from those at the population level, which arise through a change in population structure. We investigate this by building species-level trophic networks, and size-structured trophic networks, as a proxy for population structure, for 18 648 stream fish communities, from 4 145 234 individual fish samples, across 7024 stream locations in France from 1980 to 2008. We estimated effects of temperature on total trophic diversity (total number of nodes), vertical trophic diversity (mean and maximum trophic level) and distribution of biomass across trophic level (correlation between trophic level and biomass) in these networks. We found a positive effect of temperature on total trophic diversity in both species- and size-structured trophic networks. We found that maximum trophic level and biomass distribution decreased in species-level and size-structured trophic networks, but the mean trophic level decreased only in size-structured trophic networks. These results show that warmer temperatures associate with a lower vertical trophic diversity in size-structured networks, and a higher one in species-level networks. This suggests that vertical trophic diversity is shaped by antagonistic effects of temperature on population structure and on species composition. Our results hence demonstrate that effects of temperature do not only differ across trophic levels, but also across levels of biological organisation, from population to species level, implying complex changes in network structure and functioning with warming.     

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.     

Feeding ecology of two sillaginid fishes and trophic interrelations with other co-existing species in the southern part of South China Sea

Feeding ecology of Sillago sihama and Sillago ingenuua and interrelationship with other 15 fish species, including Stolephorus commersonii, Escualosa thoracata, Lutjanus russelli, Terapon puta, Ambassis kopsii, Ambassis vachellii, Dendrophysa russelli, Gerres filamentosus, Leiognathus decorus, Leiognathus splendens, Leiognathus equulus, Thryssa hamiltonii, Carangoides praeustus, Lethrinus lentjan, and Pomadasys maculatum, utilizing coastal waters off the southern part of the South China Sea were investigated. Dietary contents from 4308 guts of those species collected during July 2003–May 2005 were examined. They were significantly grouped, based on similarity coefficient, into five feeding clusters (P < 0.01). Of these, most of the diets were dominated by calanoid copepods, shrimps, polychaetes and gammarid amphipods. Comparative study on two sillaginid species showed that both Sillago sihama and Sillago ingenuua were carnivores fed mainly on polychaetes and other benthic organisms. Dietary ontogenetic change of Sillago sihama was very distinctive where small sized classes started feeding mainly on calanoid copepods before changing almost completely to polychaetes. The two species applied both sharing and partitioning strategies for food, depending on size classes, to live with each other.     

Nymphal trophic behaviour of two Nemouridae species (Insecta,Plecoptera) in the Curone creek (northern Apennines,Italy)

Feeding habits of the Euholognathan stoneflies Nemoura hesperiae and Amphinemura sulcicollis are studied in a little stream located in northern Italy. The former fed mainly on coarse particulate organic matter (CPOM), together with detritus, while the latter fed principally on detritus and, in a lower concern, on CPOM. Thus they can be catalogued fundamentally as shredder and collector‐gatherer, respectively. Shifts in diet composition were detected in relation to size in both taxa. Probably niche overlapping does not exist despite both species having similar size and nymphal development duration, and coinciding in time. This work supports that, though functional feeding groups are very useful, they should not be inferred from related species or higher taxonomical levels. Even intra‐population differences and ontogenetic shifts should be considered in fine scale studies.     

Reciprocal trophic niche shifts in native and invasive fish: salmonids and galaxiids in Patagonian lakes

1. Rainbow (Oncorhynchus mykiss) and brown trout (Salmo trutta) are widespread and invasive salmonids with important lethal effects as predators, although indirect effects are also possible. We used stable isotope analyses (δ¹⁵N, δ¹³C) to explore how the density of invasive trout in 25 Patagonian lakes alters the trophic niche (TN) of a widespread native fish, Galaxias platei (Galaxiidae). We also explored how the density of the galaxiid influences the TN of invasive trout. 2. We quantified two aspects of the TN: (i) the proportion of littoral carbon (PL) and (ii) trophic height (TH) (i.e. the ‘height’ at which the fish feeds in the food web). We related these measures of TN in a given species to the density of other species (as estimated by catch‐per‐unit‐effort). 3. As G. platei body size increased, their PL increased (increasing littoral feeding) in several lakes. However, none of the fish species investigated showed changes in PL with increasing density of the other fish species. TH increased with body size in all three species. In addition, the TH of large G. platei declined with increasing trout density and, reciprocally, the TH of large S. trutta decreased with decreasing G. platei density. 4. The reciprocal effects of native and the invasive fish on TH were as large as a shift of one trophic level. This pattern is consistent with an exhaustion of galaxiid prey for both piscivorous G. platei and S. trutta in lakes with high trout density. 5. These finding support the suggested management strategy of culling trout from overpopulated lakes, which should simultaneously protect native fish and enhance a lucrative sport fishery for large trout.     

Assessing the overlap between the diet of a coastal shark and the surrounding prey communities in a sub-tropical embayment

An elasmobranch survey of sub-tropical Hervey Bay, Australia, captured the slit-eye shark Loxodon macrorhinus at only one of three sites sampled. The dietary composition of this small shark species was compared to the prey communities within Hervey Bay to test whether prey availability was driving this observation. Dietary analysis of prey groups revealed that teleosts dominated the diet, per cent index of relative importance, % I(RI) (79·5%) and per cent geometric index of importance, % G(II) (52·7%), with shrimp-like invertebrates and cephalopods identified as the most important invertebrate prey groups. Baited remote underwater video (BRUV) used to sample prey communities at each site, demonstrated a highly diverse and significantly different community composition among the sites. There was no significant overlap between the diet of L. macrorhinus and any of the prey communities detected by BRUVs according to one-way analysis of similarities and the simplified Morisita index. Habitat electivity analysis revealed affinity of L. macrorhinus for the site with the highest water clarity (Secchi disc depth), opposing that of three other shark species. Overall, the results suggest that the distribution of L. macrorhinus is not driven by prey availability but other factors such as water clarity, predator avoidance or a reduction in interspecies competition.     

Feeding ecology and growth of age 0 year Platichthys flesus(L.) in a vegetated and a bare sand habitat in a nutrient rich fjord

A study was carried out to investigate the diet and feeding strategies of age 0 year juvenile flounder Platichthys flesus in two different micro‐tidal habitats in the nutrient enriched Mariager Fjord on the Danish east coast. Juvenile flounder and benthic macrofauna were sampled monthly from June to October 1999 in a bare sand habitat and a habitat covered by filamentous and mat forming macroalgae. The presence of the 'opportunistic' macroalgae created a shift in the dominance of surface dwelling prey such as epifaunal amphipods to more infaunal groups such as oligochaetes and polychaetes. The diet of the flounder varied considerably between the two habitats mainly reflecting prey availability relative to their abundance, prey spatial distribution, habitat structure and ontogenetic prey shifts as a function of total length. Flounder in the vegetated site fed on a diverse diet of copepods, polychaetes and oligochaetes, whereas those caught in the bare sand site fed primarily on the amphipod Corophium volutator which was numerically dominant at this site. During the growth season, two diet shifts were observed: from copepods early in the season to macrofauna organisms and, later in the season, the inclusion of more hyperbenthic prey such as Mysidea spp; Idotea spp. and the common goby Pomatoschistus microps .     

Diet of brown-nosed coatis and crab-eating raccoons from a mosaic landscape with exotic plantations in southern Brazil

We described the diets of two procyonids, the brown-nosed coati Nasua nasua and the crab-eating raccoon Procyon cancrivorus, through analysis of stomach contents of road-killed specimens in southern Brazil. We compared them with previously published dietary information for another syntopic mesopredator, the crab-eating fox Cerdocyon thous. The landscape of the study area includes native grasslands, forests, exotic tree plantations, and other crops. Food items were represented by frequency of occurrence (FO) and relative volume (RV). Stomach contents of 23 coatis were analyzed. Animal and plant items were equally frequent, although the volume of plants was greater. Exotic plant species were consumed more than native plants. Among prey items, invertebrates were more important than vertebrates, mainly because of the frequency of coleopterans and annelids and the volume of necrophagous dipteran larvae. Five specimens of raccoons were analyzed, in which animal items had the highest FO and plant items had the highest RV. Both invertebrate and vertebrate prey had the same FO, although vertebrates had a higher RV. Our data suggest that these procyonids are opportunistic hypocarnivores, utilizing anthropic sources, with diet overlap. The coatis also overlapped their diet substantially with the foxes. The stomach-contents analysis revealed the importance of fruit pulp biomass, soft-bodied animals such as larvae, and also soil, which might indicate feeding habits such as scavenging and geophagy.     

Trophic ecology of two Pithecopus species (Anura: Phyllomedusidae) living in syntopy in southern Bahia,Brazil

To coexist, two species should use available resources in different ways. These differences may happen in three different dimensions: temporal, spatial and trophic. In frogs, differences in trophic dimensions are mainly due to different sizes and types of ingested prey. We studied the diet of two syntopic Pithecopus species, addressing prey selectivity and sharing of food resources. Fieldwork was undertaken at the Michelin Ecological Reserve, southern Bahia, between the years 2011 and 2012. Stomach contents were retrieved using a flushing procedure and data on prey availability was obtained. Of the 16 invertebrate categories found in the environment, 11 were consumed by P. rohdei and 12 by P. nordestinus, features that characterize a generalist diet. The most abundant category in the diet of P. nordestinus was Diptera while Hemiptera was the most consumed prey category for P. rohdei. Trophic niche overlap was high considering the size of ingested prey and lower for categories. The selectivity index showed that both species used the resources available in the environment in a slightly different way. The differences found concerning trophic ecology of these two species seem to be sufficient to allow coexistence in the studied area.     

Feeding ecology of the Common Tern Sterna hirundo in a wintering area in southern Brazil

The wintering diet of Common Terns Sterna hirundo was studied by using 714 pellets collected on roosting sites at the mouth of the Lagoa dos Patos and on adjacent coastal beaches, in Rio Grande do Sul, southern Brazil, from March 1999 to February 2000. A total of 12 340 individual prey items of 35 different food types was found. Fish was the most important food type in the diet, constituting 32% by number and 93% by mass. Insects contributed 67% by number but only 3% by mass. The main food types were sciaenid fishes Paralonchurus brasiliensis, Micropogonias furnieri, Cynoscion guatucupa and Macrodon ancylodon. Several of these are important commercial species; fisheries potentially impact food availability to the terns, and terns may contribute significantly to the natural mortality of these fishes. Clupeiform fishes, the urophycid fish Urophycis brasiliensis and flying ants (Camponotus sp.) were also important. Species composition of the diet (food types), both by number and by mass, differed significantly between months. Prey sizes ranged in length from 12.7 mm to 217.4 mm. The average estimated total length of fish taken was 77.7 mm, but the mean differed significantly among prey species. The importance of demersal sciaenids to the diet of the Common Tern, a surface predator, may be explained by their association with aquatic predators, especially adult Bluefish Pomatomus saltatrix and Striped Weakfish Cynoscion guatucupa, and the Franciscana Dolphin Pontoporia blainvillei, which drive these fish to the surface. The occurrence of flying ants in the diet was related to offshore winds, which carried these insects out to sea. The occasional high availability of insects possibly changed the cost/benefit relationship of several food types, causing diet changes. The high number of prey species, the temporal variations in the composition of the diet and the wide range of prey sizes are evidence of the high dietary plasticity of the Common Tern, at wintering areas in southern Brazil.     

Reproductive biology and population variables of the Brazilian sharpnose shark Rhizoprionodon lalandii (Müller & Henle, 1839) captured in coastal waters of south-eastern Brazil

Throughout 1 year, from October 2003 to September 2004, 88 visits to the landing site of a small urban fishery (APREBAN) in Rio de Janeiro city were conducted and 816 specimens of Rhizoprionodon lalandii were analysed. The sample, mostly females, was composed of two cohorts: young-of-the-year were abundant in spring and summer and adults predominated in autumn and winter. Gravid females were most abundant from April to June, whereas post-partum females composed most of the catch in August to September. Adult males were present all year although were more abundant between February and July. No neonates were captured during the study and most embryos collected measured slightly below the reported total length ( L _T) at time of birth ( L ₀) for the species, suggesting that parturition may occur slightly outside the main fishing grounds or that neonates were not captured in commercial gillnets set at this time of the year. The mean L _T at maturity ( L _T50) for males was 578 mm and females matured between 620 and 660 mm, although a precise estimate of L _T50 for females could not be determined. The total length ( L _T) and total mass ( M _T) relationship     was calculated for both sexes and showed no significant differences. The mean condition factor increased steadily from February to July followed by a steep decrease in values relative to females in August and September, suggesting a pupping season. The present study area can be classified as a coastal juvenile habitat and a probable mating ground for R. lalandii .     

Feeding ecology of indigenous and non-indigenous fish species within the family Sphyraenidae

The feeding ecology of two common indigenous (Sphyraena viridensis and Sphyraena sphyraena) and one abundant non-indigenous sphyraenid species, Sphyraena chrysotaenia, of Indo-Pacific Ocean origin, was investigated in an area of the eastern Mediterranean Sea. The stomach contents of 738 individuals of varying size, collected during the period December 2008 to August 2009, were examined. The dietary analyses revealed that all three species were specialized piscivores with a diet consisting of >90% fish, both by number and mass. Concurrent sampling of the fish assemblage made it possible to calculate selectivity as well as diet breadth and overlap of these strict piscivores. Even though several prey species were found in the stomachs of the three predators examined, selectivity towards Atherina boyeri was highly significant. For all species examined, >70% of the diet by mass was made up by three indigenous species of commercial value: Spicara smaris, Boops boops and A. boyeri. Diet breadth and size of prey increased with increasing body size for all predators. With increased body size, the diet overlap between indigenous and non-indigenous species decreased. This could be attributed to increased diet breadth and the specific life-history characteristics of indigenous species developing into larger individuals. During winter, the condition factor of the non-indigenous species was significantly lower than that of the indigenous, indicating that winter conditions in the Mediterranean Sea may limit its further expansion north and westward. With this study, the gap in knowledge of the feeding preferences of the most abundant piscivorous species found in coastal areas of the study region is filled. Additionally, the results indicate that non-indigenous species familial affiliation to indigenous ones does not facilitate invasion success.     

Ontogenetic dietary shifts and feeding ecology of the rasptail skate Raja velezi and the brown smoothhound shark Mustelus henlei along the Pacific coast of Costa Rica,Central America

Stomachs from 511 Raja velezi and 340 Mustelus henlei captured as by‐catch in the commercial trawling fishery (2010–2012) were analysed to examine diet composition, ontogenetic shifts and degree of dietary overlap between species life stages in the Pacific Ocean of Costa Rica. Shrimps were the most important prey categories in the diet of R. velezi, while teleosts and cephalopods dominated the diet of M. henlei. Diet comparisons between different stages of R. velezi and M. henlei revealed clear ontogenetic dietary shifts: crustaceans (mainly shrimps, crabs and stomatopods) dominated the diet of immature individuals, and adults had a higher proportion of teleosts. The results suggest that R. velezi is an epibenthic predator that specializes in shrimps during early life stages, and to a lesser extent, teleosts as it matures, while M. henlei is an opportunistic predator with a highly diverse diet consisting of teleosts, cephalopods, shrimps and stomatopods. This study also found little evidence of dietary overlap between species or life stages and suggests that intra‐ and interspecific competition between R. velezi and M. henlei may be reduced by: (1) diet specialization in immature stages of R. velezi, (2) ontogenetic dietary shifts between immature and mature individuals, (3) prey‐size selectivity in larger individuals of R. velezi and (4) differences in depth utilization in overlapping geographical regions.     

Habitat use and trophic position determine mercury concentration in the straight fin barb Barbus paludinosus, a small fish species in Lake Awassa, Ethiopia

The diet, habitat use and mercury concentration of the small fish species, the straight fin barb Barbus paludinosus , were studied in Lake Awassa, Ethiopia, for a period of 1 year from February 2003 to January 2004. Stable isotope signatures of nitrogen and carbon in different total length ( L _T) classes were used to determine trophic positions and organic carbon sources, respectively. Barbus paludinosus mainly occupied the protected benthic habitats (littoral and profundal) of the lake. The δ¹³C values were in the range from −24 to −19‰, indicating that the carbon source for B. paludinosus was benthic, as well. Small individuals (≤ 60 mm L _T) mainly preyed upon ostracods, intermediate sizes (60–100 mm) on aquatic insects and gastropods, while a tiny cyprinodont fish Aplocheilichthys antinorii dominated the diet of large individuals (100–160 mm). The progressively increase in δ¹⁵N with increasing L _T also indicated a diet shift towards piscivory in larger individuals. The mercury concentration ranging from 0·02 to 0·74 mg kg⁻¹ wet mass (wm), was unexpectedly high in this small species, and was significantly positively related to L _T, as well as to δ¹⁵N. Some large individuals had mercury concentrations < 0·1 mg kg⁻¹ wm, and low δ¹⁵N, indicating substantial variations in diet between individuals of same size. The study suggests that other piscivorous species which include B. paludinosus in their diet may have a high mercury intake risk.     

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.     

Feeding ecology and interannual variations in diet of southern bluefin tuna, Thunnus maccoyii, in relation to coastal and oceanic waters off eastern Tasmania, Australia

The diets of 1219 southern bluefin tuna, Thunnus maccoyii, from inshore (shelf) and offshore (oceanic) waters off eastern Tasmania were examined between 1992 and 1994. Immature fish (< 155 cm fork length) made up 88% of those examined. In all, 92 prey taxa were identified. Inshore, the main prey were fish (Trachurus declivis and Emmelichthys nitidus) and juvenile squid (Nototodarus gouldi). Offshore, the diversity was greater, reflecting the diversity of micronekton in these waters. Interestingly, macrozooplankton prey (e.g. Phronima sedentaria) were prevalent in tuna > 150 cm. The offshore tuna, when in subantarctic waters, ate relatively more squid than when in the East Australia Current. In the latter, fish and crustacea were more important, although there were variations between years. No relationship was found between either prey type or size with size of tuna. Feeding was significantly higher in the morning than at other times of the day. The mean weight of prey was significantly higher in inshore-caught tuna than in those caught offshore. We estimated that the mean daily ration of southern bluefin tuna off eastern Tasmania was 0.97% of wet body weight day⁻¹. However, the daily ration of inshore-caught tuna was ∼ 3 times higher (2.7%) than for tuna caught offshore (0.8%) indicating that feeding conditions on the shelf were better than those offshore. Our results indicate that the inshore waters of eastern Tasmania are an important feeding area for, at least, immature southern bluefin tuna. This revised version was published online in July 2006 with corrections to the Cover Date.     

Trophic interactions among native and introduced fishes in a northern Patagonian oligotrophic lake

In summer the littoral zone of Lake Rivadavia contained almost all the juveniles of the native fish species present in the lake together with juveniles of the introduced salmonid Oncorhynchus mykiss . In particular, a high degree of foraging interaction was found among juveniles of the native species Aplochiton zebra and Percichthys trucha and juveniles O. mykiss. Differences in the juvenile assemblage composition, both in the presence and in the relative proportion of the species were associated with areas with different densities of vegetation. The adults of nearly all species were generalized benthic invertebrate predators. Only adults of A. zebra were absent in the littoral. Juvenile and adult O. mykiss were the only fish that preyed on planktonic organisms. Oncorhynchus mykiss was the only species that performed an ontogenetic diet shift between juveniles and adults, changing main prey categories from benthic to planktonic species.     

Varying effects of anadromous sockeye salmon on the trophic ecology of two species of resident salmonids in southwest Alaska

1. Anadromous salmon transport marine‐derived nutrients and carbon to freshwater and riparian ecosystems upon their return to natal spawning systems. The ecological implications of these subsidies on the trophic ecology of resident fish remain poorly understood despite broad recognition of their potential importance. 2. We studied the within‐year changes in the ration size, composition and stable isotope signature of the diets of two resident salmonids (rainbow trout, Oncorhynchus mykiss; Arctic grayling, Thymallus arcticus) before and after the arrival of sockeye salmon (Oncorhynchus nerka) to their spawning grounds in the Bristol Bay region of southwest Alaska. 3. Ration size and energy intake increased by 480–620% for both species after salmon arrived. However, the cause of the increases differed between species such that rainbow trout switched to consuming salmon eggs, salmon flesh and blowflies that colonized salmon carcasses, whereas grayling primarily ate more benthic invertebrates that were presumably made available because of physical disturbances by spawning salmon. 4. We also observed an increase in the δ¹⁵N of rainbow trout diets post‐salmon, but not for grayling. This presumably led to the observed increase in the δ¹⁵N of rainbow trout with increasing body mass, but not for grayling. 5. Using a bioenergetics model, we predicted that salmon‐derived resources contributed a large majority of the energy necessary for growth in this resident fish community. Furthermore, the bioenergetics model also showed how seasonal changes in diet affected the stable isotope ratios of both species. These results expand upon a growing body of literature that highlights the different pathways whereby anadromous salmon influence coastal ecosystems, particularly resident fish.     

Observations on the biology and ecology of the blue shark in the North-east Atlantic

Of 159 blue sharks Prionace glauca examined (59% female, 41% male) from oceanic waters SW of Britain and Ireland, all but two males were immature. Size-at-age and growth rate were similar to previous studies in the North Atlantic, while stomach contents included cephalopods, fish, cetaceans, and to a lesser degree, birds and crustaceans.