Resource partitioning in a novel herbivore assemblage in South America.

Human-induced species declines and extinctions have led to the downsizing of large-herbivore assemblages, with implications for many ecosystem processes. Active reintroduction of extirpated large herbivores or their functional equivalents may help to reverse this trend and restore diverse ecosystems and their processes. However, it is unclear whether resource competition between native and non-native herbivores could threaten restoration initiatives, or to what extent (re)introduced species may influence local vegetation dynamics. To answer these questions, we investigated the diets of a novel South American herbivore assemblage that includes resident native species, reintroduced native species and introduced non-native species. We examined plant composition, diet breadth and the overlap between species to describe the local herbivory profile and the potential for resource competition. Using DNA metabarcoding on faecal samples (n = 465), we analysed the diets of the herbivore assemblage in the Rincón del Socorro rewilding area of Iberá National Park, Argentina. We compared the species richness of faecal samples, the occurrence of plant families/growth forms and the compositional similarity of samples (inter- and intraspecifically). Our results indicate species-level taxonomic partitioning of plant resources by herbivores in this system. Differences in sample richness, composition and diet breadth reflected a diverse range of herbivory strategies, from grazers (capybara) to mixed feeders/browsers (brocket deer, lowland tapir). Differences in diet compositional similarity (Jaccard) revealed strong taxonomic resource partitioning. The two herbivores with the most similar diets (Pampas deer and brocket deer) still differed by more than 80%. Furthermore, all but one species (axis deer) had more similar diet composition intraspecifically than compared to the others. Overall, we found little evidence for resource competition between herbivore species. Instead, recently reintroduced native species and historically introduced non-natives are likely expanding the range of herbivory dynamics in the ecosystem. Further research will be needed to determine the full ecological impacts of these (re)introduced herbivores. In conclusion, we show clear differences in diet breadth and composition among native, reintroduced and non-native herbivore species that may be key to promoting resource partitioning, species coexistence and the restoration of ecological function.

La disminución y extinción de especies ocasionada por el hombre ha llevado a la reducción de tamaño de las comunidades de grandes herbívoros, con implicaciones para muchos procesos ecosistémicos. La reintroducción activa de grandes herbívoros extirpados, o sus equivalentes funcionales, puede ayudar a revertir esta tendencia y restaurar diversos ecosistemas y sus procesos. Sin embargo, no está claro si la competencia por recursos entre herbívoros nativos y no nativos podría amenazar las iniciativas de restauración, o en qué medida las especies (re)introducidas pueden influir la dinámica de la vegetación local. Para responder a estas preguntas, investigamos las dietas de una comunidad de herbívoros sudamericanos que incluye especies nativas, especies nativas reintroducidas y especies no nativas introducidas. Examinamos la composición de plantas, la amplitud de la dieta y la superposición entre especies para describir el perfil herbívoro local y el potencial de competencia por los recursos. Utilizando metabarcoding de ADN en muestras fecales (n = 465), analizamos las dietas de la comunidad de herbívoros en el sitio de rewilding Rincón del Socorro dentro del Parque Nacional Iberá, Argentina. Comparamos la riqueza de especies en las muestras fecales, la ocurrencia de familias de plantas/formas de crecimiento y la similitud en la composición de las muestras (interespecíficamente e intraespecíficamente). Nuestros resultados indican la partición taxonómica a nivel de especie de los recursos vegetales por parte de los herbívoros en este sistema. Las diferencias en la riqueza de las muestras, la composición y la amplitud de las dietas reflejaron una amplia gama de estrategias de herbivoría, desde pastoreadores (capibara) hasta herbívoros mixtos/ramoneadores (corzuela, tapir amazónico). Las diferencias en la similitud de la composición de la dieta (Jaccard) revelaron una fuerte partición taxonómica de los recursos. Los dos herbívoros con las dietas más similares (venado de las pampas y corzuela), aún así diferían en más del 80%. Además, todas las especies menos una (ciervo axis) tenían una composición dietética más similar intraespecíficamente que en comparación con las demás. En general, encontramos poca evidencia de competencia por recursos entre las especies de herbívoros. En cambio, las especies nativas reintroducidas recientemente y las no nativas introducidas históricamente probablemente estén ampliando el rango de dinámica de herbivoría en el ecosistema. Se necesitarán más investigaciones para determinar todos los impactos ecológicos de estos herbívoros (re)introducidos. En conclusión, mostramos diferencias claras en la amplitud y composición de la dieta entre especies de herbívoros nativas, reintroducidas y no nativas que pueden ser clave para promover la partición de recursos, la coexistencia de especies y la restauración de las funciones ecológicas.

Seasonal patterns of dietary partitioning in vertebrates.

Dietary partitioning plays a central role in biological communities, yet the extent of partitioning often varies dramatically over time. Food availability may drive temporal variation in dietary partitioning, but alternative paradigms offer contrasting predictions about its effect. We compiled estimates of dietary overlap between co-occurring vertebrates to test whether partitioning is greater during periods of high or low food abundance. We found that dietary partitioning was generally greatest when food abundance was low, suggesting that competition for limited food drives partitioning. The extent of dietary partitioning in birds and mammals was also related to seasonality in primary productivity. As seasonality increased, partitioning increased during the nonbreeding season for birds and the breeding season for mammals. Although some hypotheses invoke changes in dietary breadth to explain temporal variation in dietary partitioning, we found no association between dietary breadth and partitioning. These results have important implications for the evolution of dietary divergence.

Feeding ecology and reproductive biology of small coastal sharks in Malaysian waters.

Small coastal demersal sharks form a major proportion of the sharks landed in Malaysia. However, little is known about their feeding ecology and reproduction. This study sought to elucidate the dietary patterns, role of ontogeny in prey consumption, and reproductive biology of four dominant small demersal shark species in Malaysian waters: the Hasselt's bamboo shark, Chiloscyllium hasseltii; brownbanded bamboo shark, C. punctatum; spadenose shark, Scoliodon laticaudus; and Pacific spadenose shark, S. macrorhynchos. Dietary analyses revealed a high overlap in prey taxa consumed; clear resource partitioning among co-occurring species based on the percentage Prey-specific Index of Relative Importance (%PSIRI), with higher fish %PSIRI for Chiloscyllium hasseltii, higher cephalopod %PSIRI for C. punctatum, and higher crustacean %PSIRI for both Scoliodon species; and an ontogenetic diet shift, seen through changes in prey size. Based on the examination of reproductive organs, the results showed larger sizes at maturity for males compared to females for all four species; no obvious reproductive cycles, based on hepatosomatic and gonadosomatic indices for all species; female bias in the sex ratio of the embryos of Scoliodon species; and increased reproductive output (number of eggs or embryos and size of eggs) with larger female size for C. hasseltii and Scoliodon species. The partitioning of food resources minimizes direct competition for food and supports coexistence within shared coastal habitats. The reproductive strategies of these small coastal sharks appear to be favorable for supporting short-term population productivity; although a reduction in fishing pressure, especially from bottom trawlers, is essential for the long-term sustainable use of these sharks.

Understanding carnivore interactions in a cold arid trans-Himalayan landscape: What drives co-existence patterns within predator guild along varying resource gradients?

Predators compete for resources aggressively, forming trophic hierarchies that shape the structure of an ecosystem. Competitive interactions between species are modified in the human-altered environment and become particularly important where an introduced predator can have negative effects on native predator and prey species. The trans-Himalayan region of northern India has seen significant development in tourism and associated infrastructure over the last two decades, resulting in many changes to the natural setting of the landscape. While tourism, combined with unmanaged garbage can facilitate red fox (Vulpes vulpes), it also allows free-ranging dogs (Canis lupus familiaris), an introduced mesopredator to thrive, possibly more than the native red fox. We look at the little-known competitive dynamics of these two meso-carnivores, as well as their intra-guild interactions with the region's top carnivores, the snow leopard (Panthera uncia) and the Himalayan wolf (Canis lupus chanco). To study interactions between these four carnivores, we performed multispecies occupancy modeling and analyzed spatiotemporal interactions between these predators using camera trap data. We also collected scat samples to calculate dietary niche overlaps and determine the extent of competition for food resources between these carnivores. The study found that, after controlling for habitat and prey covariates, red fox site use was related positively to snow leopard site use, but negatively to dog and wolf site use. In addition, site use of the dog was associated negatively with top predators, that is, snow leopard and Himalayan wolf, while top predators themselves related negatively in their site use. As anthropogenic impacts increase, we find that these predators coexist in this resource-scarce landscape through dietary or spatiotemporal segregation, implying competition for limited resources. Our research adds to the scant ecological knowledge of the predators in the region and improves our understanding of community dynamics in human-altered ecosystems.

Diet overlap among non-native trout species and native cutthroat Trout (Oncorhynchus clarkii) in two U.S. ecoregions.

The invasion of freshwater ecosystems by non-native species can constitute a significant threat to native species and ecosystem health. Non-native trouts have long been stocked in areas where native trouts occur and have negatively impacted native trouts through predation, competition, and hybridization. This study encompassed two seasons of sampling efforts across two ecoregions of the western United States: The Great Basin in summer 2016 and the Yellowstone River Basin in summer 2017. We found significant dietary overlaps among native and non-native trouts within the Great Basin and Yellowstone River Basin ecoregions. Three orders of invertebrates (Ephemeroptera, Trichoptera, and Diptera) composed the majority of stomach contents and were responsible for driving the observed patterns. Great Basin trout had higher body conditions (k), and non-native Great Basin trout had higher gut fullness values than Yellowstone River Basin trout, indicating a possible limitation of food in the Yellowstone River Basin. Native fishes were the least abundant and had the lowest body condition in each ecoregion. These findings may indicate a negative impact on native trouts by non-native trouts. We recommend additional monitoring of native and non-native trout diets, regular invertebrate surveys to identify the availability of diet items, and reconsidering stocking efforts that can result in overlap of non-native fishes with native cutthroat trout.

Invasion Risk and Potential Impact of Alien Freshwater Fishes on Native Counterparts in Klang Valley, Malaysia.

This study explores the potential effects of alien fishes on the native fish community, well-being, and tropic preferences in selected rivers of Klang Valley, Malaysia. Following the Aquatic Species Invasiveness Screening Kit assessment, most of the alien fishes (80%) are invasive. The alien species occurrences correlated positively (p < 0.05) with poor water quality, such as rivers with high ammonia-nitrogen and nitrite, but negatively with phosphate and dissolved oxygen. Anthropogenic characteristics, such as rivers with high pollution levels and ease of accessibility to the fish habitat, are mainly associated positively (p < 0.05) with the occurrences of alien fish species. In general, the results of fish stomach contents analyses and their associated indices, together with stable carbon and nitrogen isotopes, revealed domination by alien fishes or diet overlaps between both alien and native fish species. This finding indicates that alien fishes benefited from the impacts of the anthropogenic activities in their surrounding habitats, while their plasticity in feeding habits might help them to invade, survive, and dominate in the rivers of Klang Valley, Malaysia.

What's for dinner? Diet and potential trophic impact of an invasive anuran Hoplobatrachus tigerinus on the Andaman archipelago.

Amphibian invasions have considerable detrimental impacts on recipient ecosystems. However, reliable risk analysis of invasive amphibians still requires research on more non-native amphibian species. An invasive population of the Indian bullfrog, Hoplobatrachus tigerinus, is currently spreading on the Andaman archipelago and may have significant trophic impacts on native anurans through competition and predation. We carried out diet analyses of the invasive H. tigerinus and native anurans, across four habitat types and two seasons; we hypothesized that (i) small vertebrates constitute a majority of the H. tigerinus diet, particularly by volume and (ii) the diet of H. tigerinus significantly overlaps with the diet of native anurans, thereby, leading to potential competition. We assessed the diet of the invasive H. tigerinus (n = 358), and individuals of the genera Limnonectes (n = 375) and Fejervarya (n = 65) and found a significant dietary overlap of H. tigerinus with only Limnonectes. Small vertebrates, including several endemic species, constituted the majority of H. tigerinus, diet by volume, suggesting potential impact by predation. Prey consumption and electivity of the three anurans indicated a positive relationship between predator-prey body sizes. Individuals of H. tigerinus and Fejervarya chose evasive prey, suggesting that these two taxa are mostly ambush predators; individuals of Limnonectes chose a mixture of sedentary and evasive prey indicating that the species employs a combination of 'active search' and 'sit and wait' foraging strategies. All three species of anurans mostly consumed terrestrial prey. This intensive study on a genus of newly invasive amphibian contributes to knowledge of the impact of amphibian invasions, and elucidates the feeding ecology of H. tigerinus, and species of the genera Limnonectes and Fejervarya. We also stress the necessity to evaluate prey availability and volume in future studies for meaningful insights into diet of amphibians.

Diet overlap among ruminants in Fennoscandia.

Information on overlap in resource use is central to understanding of interspecific exploitation competition and resource partitioning. Despite this, measures of diet overlap among northern ruminants in Fennoscandia is limited to one earlier study (reindeer and sheep). Diet overlap between sympatric moose and roe deer calculated with Schoener's index was 20.7% and 33.6% during summer (data from one area) and winter (data from two areas), respectively, whereas average diet overlap between moose and red deer was 32.0% during winter (data from four areas). Diet overlap between a coastal island population of red deer and sheep was 59.3% during summer and 63.9% during winter. Summer diet overlap between a sheep and a goat population and a sheep and a reindeer population calculated with data on main types of forage plants was 77.0% and 55.1%, respectively. However, overlap calculated with main plant groups was sometimes considerably higher than when calculated for individual forage species. Neither difference in feeding type nor body mass successfully predicted diet overlap between species pairs (n=9), although there tended to be negative correlation (r p =-0.586, P=0.098) between diet overlap of main plant groups (calculated across studies) and difference in feeding type.

Food resource partitioning among species of Astyanax (Characiformes: Characidae) in the Lower Iguaçu River and tributaries, Brazil

Resource partitioning allows for interspecific coexistence and is frequently reported for similar species. Here, we predicted the existence of resource partitioning among species of Astyanax that co-occur in the Low Iguaçu River and tributaries in Brazil. A total of 848 stomachs of five species of Astyanax were analyzed. Algae, terrestrial plant and fruit/seed were the most consumed resources. Astyanax bifasciatus and A. dissimilis had predominantly herbivorous diets, A. gymnodontus and A. lacustris were omnivorous, and A. minor was mainly algivorous. Permutational analysis of variance showed the species had different diets, and similarity percentage analysis indicated that fruit/seed and terrestrial plant contributed the most to this differentiation. A paired comparison indicated that the trophic breadth of A. gymnodontus differed from that of other species. The food overlap was low for 55% of Astyanax pairs. These results showed alignment with the niche theory, in which differentiation in the use of food resources facilitates the coexistence of species and minimizes competition. These adjustments to coexistence become relevant in the context of endemic species in a highly isolated basin under intense threat (dams, species introduction, deforestation, and pollution) as is the case for the Iguaçu River basin.(AU)

O particionamento de recursos permite a coexistência interespecífica e é frequentemente relatado para espécies semelhantes. Predizemos a existência de partição de recursos entre espécies de Astyanax que co-ocorrem no baixo rio Iguaçu. O total de 848 estômagos de cinco espécies de Astyanax foi analisado. Algas, plantas terrestres e frutos/sementes foram os recursos mais consumidos. Astyanax bifasciatus e A. dissimilis apresentaram dietas predominantemente herbívoras, A. gymnodontus e A. lacustris foram onívoras e A. minor foi principalmente algívora. As espécies apresentaram diferentes dietas (PERMANOVA) e a análise SIMPER indicou que frutos/sementes e plantas terrestres tiveram maior contribuição para esta diferenciação. A comparação pareada mostrou que a amplitude trófica de A. gymnodontus diferiu das outras espécies. A sobreposição alimentar foi baixa para 55% dos pares de Astyanax. Nossos resultados mostraram-se alinhados com a teoria de nicho, em que a diferenciação no uso de recursos alimentares facilita a coexistência de espécies e minimiza a competição. Estes ajustes para coexistência tornam-se relevantes no contexto de espécies endêmicas em uma bacia altamente isolada e sob intensa ameaça (barramentos, introdução de espécies, desmatamento e poluição), como é o caso da bacia do rio Iguaçu.(AU)

The partitioning of food resources between two rodents in the subtropical region of southern Brazil

ABSTRACT The capybara (Hydrochoerus hydrochaeris) and nutria (Myocastor coypus) are herbivorous semi-aquatic rodents. Although these rodents occur in sympatry in southern South America, little is known about how the two species interact in relationship to food resources. In this context, the aim of this study was to analyze the food resource overlap, the feeding strategy and the diversity of the diet of capybaras and nutria. A micro-histological analysis of feces was used to study the diets. A total of 48 plant species belonging to 10 families were identified in the diet of H. hydrochaeris, and a total of 49 species belonging to 14 families were identified in the diet of M. coypus. According to the Amundsen graphical method, both rodents adopted a specialized strategy for feeding on Poaceae and a generalized strategy for other families. The results of a multivariate analysis of the dietary data showed significant differences between the two rodent species and among the seasons. These differences between diets may be related to the different proportions of each food item eaten. However, the dietary overlap between the two rodents in the Taim wetland was high, suggesting that partitioning of other resources ensured the coexistence of the species.

Resource use by two electric fishes (Gymnotiformes) of the National Forest Saracá-Taquera, Oriximiná, Pará

Fishes of the order Gymnotiformes have high diversity of oral and head morphology, which suggests trophic specializations within each clade. The aim of this study was to describe resource use patterns by two fish species (Gymnorhamphichthys rondoni and Gymnotus coropinae) in the National Forest Saracá-Taquera, Oriximiná - Pará, analyzing microhabitat use, diet composition, feeding strategies, niche breadth and niche overlap. Stomach contents of 101 individuals (41 G. rondoni and 60 G. coropinae), sampled in 23 headwater streams were analyzed and volume of food items was quantified to characterize their feeding ecology. Gymnorhamphichthys rondoni was captured mainly on sandy bottoms, whereas G. coropinae in crevices. Both species had a zoobenthivorous diet and consumed predominantly Sediment/Detritus and Diptera larvae, but also included allochthonous prey in their diet. These species had high niche overlap, with small variations related to the higher consumption of Ceratopogonidae larvae by G. rondoni and of Chironomidae larvae by G. coropinae. Both species had a generalist feeding strategy, but G. coropinae had a broader niche breadth. Our results demonstrate that G. rondoni and G. coropinae occupy different microhabitats but rely on similar food resources.(AU)

Peixes da ordem Gymnotiformes apresentam alta diversidade morfológica, o que sugere a existência de especializações tróficas dentro dos clados. O objetivo deste estudo foi analisar o uso de recursos por duas espécies de peixes elétricos (Gymnorhamphichthys rondoni e Gymnotus coropinae) na Floresta Nacional Saracá-Taquera, Oriximiná - Pará, analisando o uso de microhabitats, composição da dieta, estratégias alimentares, amplitude de nicho e sobreposição de nicho. Conteúdos estomacais de 101 indivíduos (41 G. rondoni e 60 G. coropinae), capturados em 23 igarapés de cabeceira, foram analisados e quantificados volumetricamente para a caracterização da ecologia trófica. Gymnorhamphichthys rondoni foi registrada principalmente em bancos de areia, enquanto G. coropinae em fendas. Ambas as espécies apresentaram dieta zoobentívora e consumiram predominantemente sedimento/detritos e larvas de Diptera, mas também consumiram presas alóctones. Foi observada alta sobreposição de nicho trófico e a baixa variação encontrada foi relacionada à maior utilização de larvas de Chironomidae por G. rondoni e de larvas de Ceratopogonidae por G. coropinae. Ambas as espécies apresentaram estratégia alimentar generalista, porém G. coropinae apresentou maior amplitude de nicho. Nossos resultados demonstram que G. rondoni e G. coropinae ocupam microhabitats diferentes, mas dependem de recursos alimentares similares.(AU)

DISTRIBUTION AND FEEDING ECOLOGY OF THE AFRICAN TILAPIA Oreochromis mossambicus (TELEOSTEI, PERCIFORMES, CICHLIDAE) IN SURINAME (SOUTH AMERICA) WITH COMMENTS ON THE TILAPIA-KWIKWI (Hoplosternum littorale) (TELEOSTEI, SILURIFORMEs, CALLICHTHYIDAE) INTERACTION

The geographical distribution of the African Tilapia Oreochromis mossambicusin Suriname is restricted to a narrow strip of land along the Atlantic coast. Within the coastal plain, O. mossambicusoccurs in brackish lagoons, oligohaline canals, and shell-sand pit lakes. Physico-chemical characteristics and phytoplankton composition of representative Tilapia water bodies are described. Blue-green algae and fine flocculent detritus are dominant food items in the diet of the Tilapia, while Rotifera and microcrustacea are also important in the diet of larvae and juveniles. Intraspecific diet overlap among ontogenetic stages of the Tilapia did not differ significantly from 1, which means that these diets showed complete overlap. Interspecific diet overlap between the Tilapia and the indigenous armoured catfish Hoplosternum littoralewere moderate or low. The results are discussed in relation to recent developments in the Surinamese fisheries and aquaculture sector.

A distribuição geográfica da tilápia africana Oreochromis mossambicusno Suriname está limitada a uma pequena faixa de terra ao longo da costa Atlântica, Dentro desta área, O. mossambicusocorre em lagoas de baixa profundidade, canais oligohalinos e em lagos arenosos. As características fisico-quimicas e a composição em fitoplancton do meio aquático onde a tilápia ocorre são descritos. As algas azul-verde e detritos finos e floculentos são os alimentos dominantes na dieta da tilápia, enquanto que os rotíferos e os microcrustáceos são também importantes na dieta de larvas e juvenis. A sobreposição intraespecífica da dieta dentro dos estados ontogenéticos da tilápia não difere significativamente de 1, o que significa que essas dietas mostraram uma sobreposição completa. A sobreposição interespecifica da dieta entre a tilápia e o peixe-gato indigeno Hoplosternum littoralefoi moderada ou baixa. Os resultados são discutidos em relação aos recentes desenvolvimentos nos setores da pesca e aquacultura do Suriname.