Diet and trophic structure in assemblages of montane frugivorous phyllostomid bats

Neotropical frugivorous bats display a trophic structure composed of bat species with dietary preferences of core plant taxa (Artibeus-Ficus  +  Cecropia, Carollia-Piper, Sturnira- Solanum  +  Piper). This structure is hypothesized to be an ancestral trait, suggesting that similar diets would be observed throughout a species' range. However, most evidence comes from lowlands where data from montane habitats are scarce. In high mountain environments both diversity of bats and plants decreases with altitude; such decline in plant diversity produces less plants to feed from, which should ultimately affect the trophic structure of frugivorous bats in mountain environments. Here, we present a comprehensive review of the diet of frugivorous bats in Neotropical montane environments and evaluate their trophic structure in middle and higher elevations by combining a literature database with field data. We use the concept of modularity to test whether frugivorous montane bats have dietary preferences on core plant taxa. Our database revealed 47 species of montane bats feeding on 211 plant species. We find that the networks are modular, reflecting the trophic structure previously reported. We also found that in highlands the tribe Ectophyllini are Cecropia  +  Cavendishia-specialists rather than Ficus-specialists, and we describe new interactions reflecting 14 species of plants, including three botanical families previously not reported to be consumed by bats.     

Bat frugivory in two subtropical rain forests of Northern Argentina: Testing hypotheses of fruit selection in the Neotropics

Phyllostomid bats are prominent components of mammalian assemblages in the Neotropics. With many species specialized in frugivory, phyllostomids represent major partners of fleshy-fruited plants in the mutualism of seed dispersal. Here we present dietary data from two subtropical rainforests of Argentina, where fruit diversity is low and thus offer unique opportunities to test hypotheses of diet selection originally proposed for species-rich tropical assemblages. Particularly, we tested whether frugivorous phyllostomids exhibit pronounced dietary specialization in core plant taxa where fruit offer is greatly reduced as compared to tropical rainforests. We analyzed dietary overlap and niche breadth of subtropical frugivorous bats on the basis of >1000 dietary records plus >500 samples from a previous study in the region. We show that in the subtropics, frugivores from different genera remain faithful to their respective core plant taxa with few exceptions, rather than shifting toward alternative fruit resources available in the study sites. This supports predictions of specialization, which is confirmed to have a deep historical origin. The response of phyllostomid ensembles to restricted fruit diversity is at the level of species composition: absence of species for which preferred fruits do not occur in the sites. Taken together, these data lend strong support to hypotheses that explain coexistence of frugivorous phyllostomids on the basis of dietary specialization on core plant taxa with chiropterochorous fruits.     

Trophic structure of frugivorous bats in the Neotropics: emergent patterns in evolutionary history

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Trophic diversity in the evolution and community assembly of loricariid catfishes

ABSTRACT: BACKGROUND: The Neotropical catfish family Loricariidae contains over 830 species that display extraordinary variation in jaw morphologies but nonetheless reveal little interspecific variation from a generalized diet of detritus and algae. To investigate this paradox, we collected delta13C and delta15N stable isotope signatures from 649 specimens representing 32 loricariid genera (82 species) from 19 local assemblages distributed across South America. We calculated vectors representing the distance and direction of each specimen relative to the delta15N/delta13C centroid for its local assemblage, and then examined the evolutionary diversification of loricariids across assemblage isotope niche space by regressing the mean vector for each genus in each assemblage onto a phylogeny reconstructed from osteological characters. RESULTS: Loricariids displayed a total range of delta15N assemblage centroid deviation spanning 4.9per thousand, which is within the tissue-diet discrimination range known for Loricariidae, indicating that they feed at a similar trophic level and that delta15N largely reflects differences in their dietary protein content. Total range of delta13C deviation spanned 7.4per thousand, which is less than the minimum range reported for neotropical river fish communities, suggesting that loricariids selectively assimilate a restricted subset of the full basal resource spectrum available to fishes. Phylogenetic regression of assemblage centroid-standardized vectors for delta15N and delta13C revealed that loricariid genera with allopatric distributions in disjunct river basins partition basal resources in an evolutionarily conserved manner concordant with patterns of jaw morphological specialization and with evolutionary diversification via ecological radiation. CONCLUSIONS: Trophic partitioning along elemental/nutritional gradients may provide an important mechanism of dietary segregation and evolutionary diversification among loricariids and perhaps other taxonomic groups of apparently generalist detritivores and herbivores. Evolutionary patterns among the Loricariidae show a high degree of trophic niche conservatism, indicating that evolutionary lineage affiliation can be a strong predictor of how basal consumers segregate trophic niche space.     

Aquatic food‐web structure along a salinized dryland river

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Molecular characterization of trophic ecology within an island radiation of insect herbivores (Curculionidae: Entiminae: Cratopus)

The phytophagous beetle family Curculionidae is the most species‐rich insect family known, with much of this diversity having been attributed to both co‐evolution with food plants and host shifts at key points within the early evolutionary history of the group. Less well understood is the extent to which patterns of host use vary within or among related species, largely because of the technical difficulties associated with quantifying this. Here we develop a recently characterized molecular approach to quantify diet within and between two closely related species of weevil occurring primarily within dry forests on the island of Mauritius. Our aim is to quantify dietary variation across populations and assess adaptive and nonadaptive explanations for this and to characterize the nature of a trophic shift within an ecologically distinct population within one of the species. We find that our study species are polyphagous, consuming a much wider range of plants than would be suggested by the literature. Our data suggest that local diet variation is largely explained by food availability, and locally specialist populations consume food plants that are not phylogenetically novel, but do appear to represent a novel preference. Our results demonstrate the power of molecular methods to unambiguously quantify dietary variation across populations of insect herbivores, providing a valuable approach to understanding trophic interactions within and among local plant and insect herbivore communities.     

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.     

Functional differences in echolocation call design in an adaptive radiation of bats

All organisms have specialized systems to sense their environment. Most bat species use echolocation for navigation and foraging, but which and how ecological factors shaped echolocation call diversity remains unclear for the most diverse clades, including the adaptive radiation of neotropical leaf‐nosed bats (Phyllostomidae). This is because phyllostomids emit low‐intensity echolocation calls and many inhabit dense forests, leading to low representation in acoustic surveys. We present a field‐collected, echolocation call dataset spanning 35 species and all phyllostomid dietary guilds. We analyze these data under a phylogenetic framework to test the hypothesis that echolocation call design and parameters are specialized for the acoustic demands of different diets, and investigate the contributions of phylogeny and body size to echolocation call diversity. We further link call parameters to dietary ecology by contrasting minimum detectable prey size estimates (MDPSE) across species. We find phylogeny and body size explain a substantial proportion of echolocation call parameter diversity, but most species can be correctly assigned to taxonomic (61%) or functional (77%) dietary guilds based on call parameters. This suggests a degree of acoustic ecological specialization, albeit with interspecific similarities in call structure. Theoretical MDPSE are greatest for omnivores and smallest for insectivores. Omnivores significantly differ from other dietary guilds in MDPSE when phylogeny is not considered, but there are no differences among taxonomic dietary guilds within a phylogenetic context. Similarly, predators of non‐mobile/non‐evasive prey and predators of mobile/evasive prey differ in estimated MDPSE when phylogeny is not considered. Phyllostomid echolocation call structure may be primarily specialized for overcoming acoustic challenges of foraging in dense habitats, and then secondarily specialized for the detection of food items according to functional dietary guilds. Our results give insight into the possible ecological mechanisms shaping the diversity of sensory systems, and their reciprocal influence on resource use.     

Cladocerans respond to differences in trophic state in deeper nutrient poor lakes from Southern Norway

We investigated the role of trophic state in structuring the cladoceran assemblages in 35 relatively deep lakes of low nutrient levels in Southern Norway. The lakes cover gradients of altitude, latitude, longitude, area and total phosphorus concentration. The environmental control of the cladoceran assemblages was analyzed using canonical correspondence analysis. Total phosphorus was the most important variable in the minimal adequate model. Hence, the cladoceran assemblage changed with trophic state. Typical clear water species were displaced by indicators of eutrofication with increasing trophic levels. Littoral species constituted the majority of the species recorded. Littoral and pelagic species richness showed a unimodal relationship with trophic state. Along with the change in trophic state and shift in cladoceran assemblage, we observed an increase in the ratio of pelagic to littoral species as well as in the ratio of pelagic efficient bacterial feeders to total pelagic filter feeders. The study indicates that zooplankton could provide a valuable indicator of ecosystem’s structure and function in deeper nutrient poor lakes if included in water quality assessments according to the EU Water Framework Directive. Including sampling of the littoral zooplankton would improve diversity estimates and incorporate the response of the littoral zone to eutrophication.     

Seasonal differences in population‐, ensemble‐ and community‐level responses of bats to landscape structure in Amazonia

The amount (composition) and spatial arrangement (configuration) of forest patches in fragmented landscapes influence the accessibility, as well as the abundance and diversity of resources available to bats. Moreover, tropical fruit and insect abundance differ seasonally in response to changes in precipitation, and many bats in the family Phyllostomidae employ seasonal reproductive strategies. Because reproductive activities involve constraints on time and energy as well as increased nutritional demands, foraging behavior and home range size may differ between wet and dry seasons. Nonetheless, seasonal variation in response to landscape structure by bats has not been examined previously. Consequently, population‐, ensemble‐ and assemblage‐level responses of phyllostomids to landscape composition and configuration were quantified separately during the wet and dry season at three circular focal scales (1, 3 and 5 km radii) for 14 sites in fragmented lowland Amazon forest. Responses to landscape characteristics were scale‐dependent, species‐specific, and seasonal. Abundances of frugivores responded to landscape composition in the dry season and to landscape configuration in the wet season. Conversely, abundances of animalivores responded to landscape configuration in the dry season and to landscape composition in the wet season. Divergent responses to landscape structure between seasons suggest that variation in resource abundance and diversity play a significant role in structuring population‐, ensemble‐ and assemblage‐level patterns. As such, considerations of the effects of dietary flexibility and reproductive constraints on foraging strategies and habitat use may be important when designing management plans that successfully promote long‐term persistence of biodiversity in fragmented landscapes.     

Diets and trophic‐guild structure of a diverse fish assemblage in Chesapeake Bay,U.S.A.

Dietary habits and trophic‐guild structure were examined in a fish assemblage (47 species) of the Chesapeake Bay estuary, U.S.A., using 10 years of data from >25 000 fish stomachs. The assemblage was comprised of 10 statistically significant trophic guilds that were principally differentiated by the relative amounts of Mysida, Bivalvia, Polychaeta, Teleostei and other Crustacea in the diets. These guilds were broadly aggregated into five trophic categories: piscivores, zooplanktivores, benthivores, crustacivores and miscellaneous consumers. Food web structure was largely dictated by gradients in habitat (benthic to pelagic) and prey size. Size classes within piscivorous species were more likely to be classified into different guilds, reflecting stronger dietary changes through ontogeny relative to benthivores and other guilds. Relative to predator species and predator size, the month of sampling had negligible effects on dietary differences within the assemblage. A majority of sampled fishes derived most of their nutrition from non‐pelagic prey sources, suggesting a strong coupling of fish production to benthic and demersal food resources. Mysida (predominantly the opossum shrimp Neomysis americana) contributed substantially to the diets of over 25% of the sampled predator groups, indicating that this species is a critical, but underappreciated, node in the Chesapeake Bay food web.     

Seabird species‐ and assemblage‐level isotopic niche shifts associated with changing prey availability during breeding in coastal Newfoundland

Shifting prey availability can lead to altered species interactions, indicated by variation in the dietary niche breadth and position of species within an assemblage. On the Newfoundland coast, annual inshore spawning migration of the dominant forage fish, Capelin Mallotus villosus, provides an excellent opportunity to investigate the influence of varying prey availability on dietary niche breadth and position among species. During June–August 2017, we investigated species‐ and assemblage‐level dietary responses to shifting Capelin availability of three Capelin‐eating, sympatrically breeding auk species, the Atlantic Puffin Fratercula arctica, Razorbill Alca torda and Common Murre Uria aalge. The diet of Leach's Storm Petrels Oceanodroma leucorhoa, which breed alongside the three auk species but are not known to rely on Capelin, was also examined to determine dietary shifts throughout breeding that were unrelated to Capelin availability. We quantified stable isotope ratios (δ¹⁵N, δ¹³C) in seabird blood components (plasma, cellular component) collected both before and after spawning Capelin arrived in the study area and compared isotopic niche breadth within a Bayesian framework. At the species level, auk trophic position increased and isotopic niche breadth narrowed after Capelin arrived, suggesting a more Capelin‐based diet. Simultaneously, trophic diversity of the auk assemblage, reflecting the extent of spacing among niches of species, decreased after spawning Capelin arrived inshore. Contrastingly, increased trophic position but broader isotopic niche breadth during higher relative to lower Capelin availability for Leach's Storm Petrel confirm that this species is probably not affected by the inshore arrival of Capelin, but instead that isotopic changes may be more related to a shift in breeding stage to chick‐rearing. Overall, our findings reiterate the importance of Capelin as a prey resource for breeding auks in coastal Newfoundland, but that the degree of reliance on Capelin varies among species, possibly allowing coexistence of these ecologically similar species. The findings highlight potential changing species interactions, such as increased competition, under declines in Capelin biomass.     

Palate Variation and Evolution in New World Leaf-Nosed and Old World Fruit Bats (Order Chiroptera)

Two bat families, the leaf-nosed (Phyllostomidae) and fruit bats (Pteropodidae), have independently evolved the ability to consume plant resources. However, despite their similar ages, species richness and the strong selective pressures placed on the evolution of skull shape by plant-based foods, phyllostomids display more craniofacial diversity than pteropodids. In this study, we used morphometrics to investigate the distribution of palate variation and the evolution of palate diversity in these groups. We focused on the palate because evolutionary alterations in palate morphology are thought to underlie much feeding specialization in bats. We hypothesize that the distribution of palate variation differs in phyllostomids and pteropodids, and that the rate of palate evolution is higher in phyllostomids than pteropodids. The results suggest that the overall level of palate integration is higher in adult populations of pteropodids than phyllostomids but that the distribution of palate variation is otherwise generally conserved among phyllostomids and pteropodids. Furthermore, the results are consistent with these differences in palate integration likely having a developmental basis. The results also suggest that palate evolution has occurred significantly more rapidly in phyllostomids than pteropodids. These findings are consistent with a scenario in which the greater integration of the pteropodid palate has limited its evolvability.     

The diversity of beetle assemblages in different habitat types in Sabah, Malaysia

The diversity of beetle assemblages in different habitat types (primary forest, logged forest, acacia plantation and oil palm plantation) in Sabah, Malaysia was investigated using three different methods based on habitat levels (Winkler sampling, flight-interception-trapping and mist-blowing). The overall diversity was extremely high, with 1711 species recorded from only 8028 individuals and 81 families (115 family and subfamily groups). Different degrees of environmental changes had varying effects on the beetle species richness and abundance, with oil palm plantation assemblage being most severely affected, followed by acacia plantation and then logged forest. A few species became numerically dominant in the oil palm plantation. In terms of beetle species composition, the acacia fauna showed much similarity with the logged forest fauna, and the oil palm fauna was very different from the rest. The effects of environmental variables (number of plant species, sapling and tree densities, amount of leaf litter, ground cover, canopy cover, soil pH and compaction) on the beetle assemblage were also investigated. Leaf litter correlated with species richness, abundance and composition of subterranean beetles. Plant species richness, tree and sapling densities correlated with species richness, abundance and composition of understorey beetles while ground cover correlated only with the species richness and abundance of these beetles. Canopy cover correlated only with arboreal beetles. In trophic structure, predators represented more than 40% of the species and individuals. Environmental changes affected the trophic structure with proportionally more herbivores (abundance) but fewer predators (species richness and abundance) in the oil palm plantation. Biodiversity, conservation and practical aspects of pest management were also highlighted in this study.     

The coexistence of fish species in streams: relationships between assemblage attributes and trophic and environmental variables

Diet overlap and niche breadth are well-known species traits from trophic ecology that can assist in explaining how species interact and coexist as well as the ecological mechanisms that influence biodiversity. In the present study, we analyzed the relationships between these trophic variables and indicators of resource availability with some attributes of fish assemblages (species richness, Shannon diversity index, evenness, density and individual body size). The physical and chemical characteristics of the biotopes (topography, water quality and conservation of slopes) were examined to identify possible patterns. Monthly sampling using electrofishing was conducted in 2003 along five streams located in the Cuiabá River watershed. The relationships between environmental variables and attributes of fish assemblages were evaluated using Spearman correlation. Species richness and abundance varied among streams, with higher values (54 and 82 species) found in low-gradient streams that drained small swampy areas discharging in Cuiabá River. Diet overlap showed significant and negative correlations with species richness, the diversity index, abundance, variation in body size and food availability and positive correlations with evenness and niche breadth. Environments that had greater food availability had a reduced variety of food items (smaller amplitude) and distinct values in terms of dietary overlap. Nevertheless, to explain resource partitioning in a fish assemblage (overlap), it is necessary to consider food availability and niche breadth (degree of trophic specialization) beyond the interaction of these variables with species richness and density. In conclusion, high diversity and abundance values were strongly associated with a high degree of trophic specialization (low amplitude of trophic niche) and a small degree of overlap in the diet.     

Neonatal growth and survival of rodents in response to variation in maternal dietary nitrogen: life history strategy vs dietary niche

Nitrogen (N) enrichment of terrestrial ecosystems dramatically changes ecosystem diversity and structure of plant communities. Research designed to elucidate effects of nitrogen addition on mammalian assemblages is rare. We investigated nitrogen requirements of hispid cotton rats ( Sigmodon hispidus ) and fulvous harvest mice ( Reithrodontomys fulvescens ), small mammals native to the tallgrass prairie of the southern Great Plains, USA, to better understand population responses of these species to nitrogen enrichment. We studied reproductive requirements by measuring growth of offspring under varying levels of dietary nitrogen. We predicted that dietary niche would dictate nitrogen requirements, such that the larger herbivore ( S. hispidus ) would have a lower dietary need for nitrogen per unit mass than the small omnivore/granivore ( R. fulvescens ). Reproductive output (measured as mass gain of litters and offspring) was responsive to varying nitrogen in cotton rats but not in harvest mice. Nitrogen intake that supported 50% survival of juvenile harvest mice (1.34% dietary nitrogen) also was adequate for maximum growth (1.29%). Cotton rats potentially drew on maternal nutrient stores to support litter growth at low levels of dietary nitrogen (as low as 1.08%). Overall, nitrogen requirements for maximum reproduction were greater (2.31% dietary nitrogen) for cotton rats. We conclude that life history characteristics and body size constraints rather than dietary niche explain the differential species response to variation in dietary nitrogen. Our results imply that nitrogen enrichment in old-field succession in the southern Great Plains may lead to dominance by cotton rats and a reduction in diversity of the small-mammal assemblage. Consumers with similar abilities to take advantage of increased environmental nitrogen may likewise dominate other ecosystems.     

Species richness and functional structure of fish assemblages in three freshwater habitats: effects of environmental factors and management

In this study, the inverted trophic hypothesis was tested in the freshwater fish communities of a reservoir. The distribution of fish species in three freshwater habitats in the Jurumirim Reservoir, Brazil, was examined using both species richness and the relative proportions of different trophic groups. These groups were used as a proxy for functional structure in an attempt to test the ability of these measures to assess fish diversity. Assemblage structures were first described using non-metric multidimensional scaling (NMDS). The influence of environmental conditions for multiple fish assemblage response variables (richness, total abundance and abundance per trophic group) was tested using generalised linear mixed models (GLMM). The metric typically employed to describe diversity; that is, species richness, was not related to environmental conditions. However, absolute species abundance was relatively well explained with up to 54% of the variation in the observed data accounted for. Differences in the dominance of trophic groups were most apparent in response to the presence of introduced fish species: the iliophagous and piscivorous trophic groups were positively associated, while detritivores and herbivores were negatively associated, with the alien species. This suggests that monitoring functional diversity might be more valuable than species diversity for assessing effects of disturbances and managements policies on the fish community.     

Consumer vs environmental productivity control of bacterial diversity and bacteria-mediated organic matter decomposition

We experimentally investigated the determinants (consumer vs environmental productivity) and functional consequences of bacterial diversity using a simple laboratory bacterial assemblage and the bacterivorous ciliated protist Colpidium striatum . A two-way factorial design was used, with two productivity levels crossed with the presence/absence of Colpidium striatum . The effects of productivity and predation on bacterial diversity were similar to those previously reported for plant diversity in systems characterized by high productivities, with increased productivity reducing diversity and predation increasing diversity. When Colpidium was present, changes in trophic level biomass and individual bacterial species abundance in response to increased productivity were largely consistent with the keystone predation model. Importantly, Colpidium predation also resulted in an increase in particulate organic matter decomposition, largely due to an increase in the abundance of one single bacterial species that appeared resistant to predation. These results suggest that changes in community structure as a result of trophic interactions and other factors may have profound consequences for important ecosystem functions. We suggest that future research should take an integrative approach and study causes and consequences of biodiversity simultaneously.     

Responses of phyllostomid bats to forest cover in upland landscapes in Chiapas,southeast Mexico

Forests are a key habitat for bats, but tend to be lost and fragmented in some agri-environment schemes. We studied the effects of forest cover change on phyllostomid bats in agricultural landscapes with increment of open areas in an upland region in Chiapas, southeast Mexico. We tested whether with forest cover increase there is a directly proportional response on assemblage species diversity measures, on the capture success and body condition of particular ensembles. Depending on the spatial analysis window, and presumably on vagility, we found positive and significant associations with the sanguivore ensemble’s capture success, as well as with the nectarivore and shrub frugivore ensembles’ body condition. We support the idea that appropriate amounts of forest over small geographic extents may propitiate favorable environments for some phyllostomids, which can also provide important ecological services. Furthermore, the arrangement of ecologically similar species proved to be valuable for exploring adaptive traits, and adequate for conservation strategies of species-rich taxa.