Seasonal variation in terrestrial resource subsidies influences trophic niche width and overlap in two aquatic snake species: a stable isotope approach

Quantifying diet is essential for understanding the functional role of species with regard to energy processing, transfer, and storage within ecosystems. Recently, variance structure in the stable isotope composition of consumer tissues has been touted as a robust tool for quantifying trophic niche width, a task that has previously proven difficult due to bias in direct dietary analyses and difficulties in integrating diet composition over time. We used carbon and nitrogen stable isotope analyses to examine trophic niche width of two sympatric aquatic snakes, banded watersnakes Nerodia fasciata and black swamp snakes Seminatrix pygaea inhabiting an isolated wetland where seasonal migrations of amphibian prey cause dramatic shifts in resource availability. Specifically, we characterized snake and prey isotope compositions through time, space, and ontogeny and examined isotope values in relation to prey availability and snake diets assessed by gut content analysis. We determined that prey cluster into functional groups based on similarity of isotopic composition and seasonal availability. Isotope variance structure indicated that the trophic niche width of the banded watersnake was broader (more generalist) than that of the black swamp snake. Banded watersnakes also exhibited seasonal variation in isotope composition, suggesting seasonal diet shifts that track amphibian prey availability. Conversely, black swamp snakes exhibited little seasonal variation but displayed strong ontogenetic shifts in carbon and nitrogen isotope composition that closely paralleled ontogenetic shifts in their primary prey, paedomorphic mole salamanders Ambystoma talpoideum. Although niche dimensions are often treated as static, our results demonstrate that seasonal shifts in niche dimensions can lead to changes in niche overlap between sympatric species. Such short‐term fluctuations in niche overlap can influence competitive interactions and consequently the composition and dynamics of communities and ecosystems.     

High resource overlap and small dietary differences are widespread in food-limited warbler (Parulidae) communities

Although both interspecific competition and coexistence mechanisms are central to ecological and evolutionary theory, past empirical studies have generally focused on simple (two-species) communities over short time periods. Experimental tests of these species interactions are challenging in complex study systems. Moreover, several studies of ‘imperfect generalists’, consistent with Liem's Paradox, raise questions about the ability of evolved species differences to partition niche space effectively when resources vary considerably across the annual cycle. Here we used a recently developed theoretical framework to combine past research on population-level processes with observational data on resource use to test for ongoing interspecific competition and understand the nature of resource overlap. We compared species diet overlaps and differences in several distinctive communities centred on a focal species, the American Redstart Setophaga ruticilla replicated both spatially and seasonally, in combination with documentation of population regulation to assess the ability of similar species to partition dietary niche space and limit interspecific competition. Our results document high dietary overlap in most of the communities studied, with only subtle differentiation consistent with known species differences in foraging behaviour and morphology. These findings are largely consistent with species foraging as imperfect generalists. However, in contrast to past studies, the high diet overlaps observed here during times of inferred resource scarcity were driven by low-value prey taxa (e.g. small ants) and did not involve truly ‘private’ resources. All of these factors increase the potential negative impacts of interspecific competition, and limit the ability of these birds to avoid competition if food availability deteriorates further than observed in our study, either seasonally or at longer intervals.     

Resource partitioning in sympatric langurs and macaques in tropical rainforests of the central Western Ghats, south India

In a competitive sympatric association, coexisting species may try to reduce interspecific interactions as well as competition for similar resources by several ecological and behavioral practices. We studied resource utilization of three sympatric primate species namely, lion-tailed macaques (Macaca silenus), bonnet macaques (M. radiata) and Hanuman langurs (Semnopithecus entellus) in a tropical rainforest of the central Western Ghats, south India. We studied resource use, tree-height use, foraging height, substrate use when consuming animal prey and interspecific interactions. The results revealed that across the year, there was very limited niche overlap in diet between each species-pair. Each primate species largely depended on different plant species or different plant parts and phenophases from shared plant species. Primate species used different heights for foraging, and the two macaque species searched different substrates when foraging on animal prey. We also recorded season-wise resource abundance for the resources shared by these three primate species. While there was low dietary overlap during the dry season (a period of relatively low resource abundance), there was high dietary overlap between the two macaque species during the wet season (a period of high resource abundance for the shared resources). We observed only a few interspecific interactions. None of these were agonistic, even during the period of high niche overlap. This suggests that the sympatric primate species in this region are characterized by little or no contest competition. Unlike in some other regions of the Western Ghats, the lack of interspecific feeding competition appears to allow these primates, especially the macaques, to remain sympatric year-round.     

Do lizard communities partition the trophic niche? A worldwide meta‐analysis using null models

Lizards have been widely used as model organisms in community ecology over the past 30 years. I have reviewed more than 50 studies from the literature on the community ecology of lizards worldwide, with a focus on studies from 1990–2007. I determined if there is support for the hypotheses that many lizard communities 1) are non‐randomly organized along the trophic niche dimension, and 2) partition the available food resource to minimize interspecific competition. I used data on number of prey items and percentage of prey volume, to calculate dietary overlap among species. I compared true datasets to randomly generate new datasets produced by 3×10⁴ Monte Carlo permutations using two algorithms (RA2 and RA3). The great majority (more than 80%) of the communities were randomly organized along the trophic niche dimension using RA2 or RA3; the frequency of occurrence of detecting a non‐random structure in lizard community dietary studies did not differ significantly between datasets based on either number of prey items or prey volume. Thus, lizard communities usually do not partition the trophic niche axis. Concerning the number of prey items, logistic regression models showed that the presence of a structure in the dataset did not depend on number of species, method employed to obtain dietary data, matrix size, or location (continent), but instead significantly depended on whether a community was tropical. Concerning prey volume, presence of a structure in the dataset also did not depend on number of species, method employed to obtain dietary data, and tropical vs non‐tropical origin, but was marginally dependent on continent (South America was favoured for identifying a structure in the dataset) and matrix size. In general, a structure in the dataset was more often uncovered by using RA2 than RA3. Overall, I conclude that 1) lizard communities are unlikely to partition available food resources, and 2) as the method for identifying dietary items does not allow for accurate prey identification (at least at the genus level), there is a risk of obtaining false null results about community structure. This is because structure along the trophic niche axis is genuinely rare in lizards and also because inappropriate methods may erroneously lead to this conclusion. Therefore, conclusions from studies utilizing stomach dissection as a source of diet data (still common in the literature) should be cautiously considered for ethical reasons and because it is difficult to find any difference between stomach‐dissection and fecal‐pellet data when assessing the presence of a structure in datasets. My data also showed there are minor differences in the probability of detecting a structure in datasets using prey item number or prey volume data for lizards. The biases, both statistical and biological, associated with this meta‐analysis are discussed.     

Seasonally fluctuating resources and temporal variability of interspecific competition

Summary The influence of seasonal availability of two critical resources (food and substrates from which food was harvested) on interspecific competition between striped surfperch (Embiotoca lateralis) and black surfperch (Embiotoca jacksoni) was examined. There was a strong depth-related gradient in density of prey and in cover of foliose algae; both declined with increasing bottom depth. Density of prey was reduced 5–10 fold during the winter season, but cover of substrates remained constant throughout the year. Although both fishes co-occurred throughout the same depth range, striped surfperch were more common in shallow habitats and black surfperch were more abundant deeper. Local abundance and distribution patterns of both surfperch species did not change seasonally. Stepwise regression analyses suggested that availability of favored substrates was a proximate influence on local patterns of surfperch distribution and abundance, and that interspecific competition depressed abundance of the two species to the same degree. Removal experiments conducted during the cold-water season revealed that interspecific competition influenced depth distribution of black surfperch but not striped surfperch. Seasonal change in density of prey was accompanied by marked changes in overlap in use of foraging substrates by the surfperches. The pattern of change in interspecific overlap suggested that surfperch competed for food only when prey were seasonally scarce. There was no difference in the agonistic tendencies of the two fishes, and the absolute and relative frequency of interspecific chases was independent of food level. These results have important implications regarding the impact of temporal variability of interspecific competition in natural communities. In the surfperch system, competition was characterized by constant and time-varying elements that had symmetrical and asymmetrical effects and involved both interference and exploitation mechanisms.     

Contextual Flexibility: Reassessing the Effects of Prey Size and Status on Prey Restraint Behaviour of Macrostomate Snakes

Contextual flexibility in prey restraint behaviour has been documented in advanced snakes (Colubroidea), but the degree of flexibility for earlier snake lineages has been largely unstudied. We document the prey restraint behaviour of five snake species belonging to three early macrostomate lineages: Loxocemidae, Erycinae and Boidae. Species from these lineages were chosen for this study because they utilize similar prey resources but exhibit different ecological habits that may have important consequences on prey restraint behaviour. Snakes (n = 27) were studied in a systematic experimental design assessing the effects of mouse size (small and large) and status (live and dead) across a total of 216 feeding trials. Loxocemus and Erycine snakes were highly flexible in their prey restraint behaviour patterns and these varied across prey category. Individuals of Boa constrictor exhibited very little contextual flexibility in feeding behaviour, confirming earlier reports. Flexibility in prey restraint behaviour corresponded with loop application pattern, whether the snake bent laterally or ventrally when forming a loop around prey. Our study is the first to show that early macrostomate snakes exhibit flexible prey restraint behaviours. Thus, our results suggest that flexibility in predatory behaviour may be more widespread across snake taxa than previously thought and we offer hypotheses for the observed interspecific differences in snake feeding behaviour.     

Food Habits and Livestock Depredation of Sympatric Jaguars and Pumas in the Iguaçu National Park Area, South Brazil

Understanding coexistence between sympatric felines with similar body sizes, such as jaguars Panthera onca and pumas Puma concolor , requires knowledge of the way these predators consume and partition food resources. Yet the importance of livestock predation on jaguar and puma coexistence is poorly known. I investigated food habits and patterns of livestock depredation of jaguar and pumas in the Iguaçu National Park (INP) in southern Brazil. From 1997 to 2001, I collected scats opportunistically on trails and roads in INP and visited ranches on the border of INP. I found that jaguars relied mostly on large and medium-sized wild prey species, while pumas concentrated on medium-sized prey species. Livestock was the fifth most frequent prey found in jaguar scats but the most important one in terms of biomass consumed. Jaguar and puma diets differed significantly when all prey items were compared and also when livestock was excluded from the jaguar diet. Jaguar predation on livestock was considerably higher than predation by pumas. However, predation was not substantial relative to availability of livestock, and cattle likely constitute an alternative source of prey for jaguars. Degree of diet overlap between jaguar and puma in INP suggests that coexistence was likely driven by exploitative competition through some degree of food partitioning. My results highlight the importance of more actions toward increasing numbers of large ungulates to preserve the population of jaguars in INP.     

Variation in the diet of the viperine snake Natvix mama in relation to prey availability

The diet of the viperine snake was compared with food availability in the Ebro Delta, a wetland largely occupied by rice fields, in 1990 and 1991. Snake selection of prey type and size was studied seasonally and by snake group: males, females and immature snakes. Overall, feeding activity (percentage of individuals with prey and number of prey per stomach) increased with food availability. Diet analysis showed that viperine snakes mainly foraged on the green frog Rana perezi (adults and tadpoles) and the carp Cyprinus earpio. Conversely, viperine snakes rejected the mosquito fish Gambusia holbroki which is the most abundant species in autumn, when Natrix maura has a low feeding activity. Statistical comparisons between viperine snake diet and prey availability showed that males selected small carp, immature snakes selected tadpoles and, in spring, females selected frogs. The selection of small carp by males may reflect a sexual divergence of trophic niche related to sexual size dimorphism, as females are larger than males. As tadpoles are presumably easier to catch than fish, tadpole selection by immature individuals may reflect variance in capture abilities. In spring, the selection of frogs by females overlapped with vitellogenesis, suggesting that females compensate for the cost of reproduction by selecting green frogs, which have a greater biomass and higher energy content than fish. Carps eaten in spring were smaller than in summer. Moreover, in summer viperine snakes selected smaller carp than the available mean size. This divergent tendency between carp size selection and carp size availability reveals how seasonal diet shifts in prey size selection may be a response to an increase in prey size.     

Feeding ecology of North American gopher snakes (Pituophis catenifer, Colubridae)

Studies of food relations are important to our understanding of ecology at the individual, population and community levels. Detailed documentation of the diet of large‐bodied, widespread snakes allows us to assess size‐dependent and geographical variation in feeding preferences of gape‐limited predators. Furthermore, with knowledge of the food habits of sympatric taxa we can explore possible causes of interspecific differences in trophic niches. The feeding ecology of the North American gopher snake, Pituophis catenifer, was studied based on the stomach contents of more than 2600 preserved and free‐ranging specimens, and published and unpublished dietary records. Of 1066 items, mammals (797, 74.8%), birds (86, 8.1%), bird eggs (127, 11.9%), and lizards (35, 3.3%) were the most frequently eaten prey. Gopher snakes fed upon subterranean, nocturnal and diurnal prey. The serpents are primarily diurnal, but can also be active at night. Therefore, gopher snakes captured their victims by actively searching underground tunnel systems, retreat places and perching sites during the day, or by pursuing them or seizing them while they rested at night. Gopher snakes of all sizes preyed on mammals, but only individuals larger than 40 and 42 cm in snout–vent length took bird eggs and birds, respectively, possibly due to gape constraints in smaller serpents. Specimens that ate lizards were smaller than those that consumed mammals or birds. Gopher snakes raided nests regularly, as evidenced by the high frequency of nestling mammals and birds and avian eggs eaten. Most (332) P. catenifer contained single prey, but 95 animals contained 2–35 items. Of the 321 items for which direction of ingestion was determined, 284 (88.5%) were swallowed head‐first, 35 (10.9%) were ingested tail‐first, and two (0.6%) were taken sideways. Heavier gopher snakes took heavier prey, but heavier serpents ingested prey with smaller mass relative to snake mass, evidence that the lower limit of prey mass did not increase with snake mass. Specimens from the California Province and Arid Deserts (i.e. Mojave, Sonoran and Chihuahuan Deserts) took the largest proportion of lizards, whereas individuals from the Great Basin Desert consumed a higher percentage of mammals than serpents from other areas, and P. catenifer from the Great Plains ate a greater proportion of bird eggs. Differences in prey availability among biogeographical regions and unusual circumstances of particular gopher snake populations may account for these patterns. Gopher snakes have proportionally longer heads than broadly sympatric Rhinocheilus lecontei (long‐nosed snake), Charina bottae (rubber boa) and Lampropeltis zonata (California mountain kingsnake), which perhaps explains why, contrary to the case in P. catenifer, the smaller size classes of those three species do not eat mammals. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77 , 165–183.     

Does interspecific competition drive patterns of habitat use in desert bat communities?

Bodies of water are a key foraging habitat for insectivorous bats. Since water is a scarce and limiting resource in arid environments, bodies of open water may have a structuring effect on desert bat communities, resulting in temporal or spatial partitioning of bat activity. Using acoustic monitoring, we studied the spatial and temporal activity patterns of insectivorous bats over desert ponds, and hypothesised that sympatric bat species partition the foraging space above ponds based on interspecific competitive interactions. We used indirect measures of competition (niche overlap and competition coefficients from the regression method) and tested for differences in pond habitat selection and peak activity time over ponds. We examined the effect of changes in the activity of bat species on their potential competitors. We found that interspecific competition affects bat community structure and activity patterns. Competing species partitioned their use of ponds spatially, whereby each species was associated with different pond size and hydroperiod (the number of months a pond holds water) categories, as well as temporally, whereby their activity peaked at different hours of the night. The drying out of temporary ponds increased temporal partitioning over permanent ponds. Differences in the activity of species over ponds in response to the presence or absence of their competitors lend further support to the role of interspecific competition in structuring desert bat communities. We suggest that habitat use and night activity pattern of insectivorous bats in arid environments reflect the trade-offs between selection of preferred pond type or activity time and constraints posed by competitive interactions.     

Interspecific competition and torpor in golden spiny mice: two sides of the energy-acquisition coin

We studied the occurrence of torpor in golden spiny mice in a hot rocky desert near the Dead Sea. In this rodent assemblage, a congener, the nocturnal common spiny mouse, competitively excluded the golden spiny mouse from the nocturnal part of the diel cycle and forced it into diurnal activity; this temporal partitioning allows the two species to partition their prey populations, particularly in summer when the diet of the two species is comprised mainly of arthropods, and largely overlap. We studied the effect of the presence of the common spiny mice at two resource levels (natural food availability and food added ad libitum) on populations of golden spiny mice in four large outdoor enclosures: two with common spiny mice removed and two enclosures with populations of both species. We hypothesized that with interspecific competition and/or reduced resources, golden spiny mice will increase their use of torpor. As we expected, supplemented food reduced the total time spent torpid. In summer, when the different activity periods of the two species results in prey species partitioning, removal of the congener did not affect torpor in the golden spiny mouse. However, in winter, when insect populations are low and the two species of mice overlap in a largely vegetarian diet, removal of the common spiny mouse reduced torpor in golden spiny mice, whether food was supplemented or not. This result suggests that torpor, a mechanism that allows small mammals to sustain periods of low availability of resources or high energetic requirements, may also help them to tolerate periods of enhanced interspecific competition. This may be a significant short-term mechanism that reduces competition and hence increases fitness, in particular of individuals of the subordinate species whose accessibility to resources may be limited.     

Sexual differences in morphology and niche utilization in an aquatic snake,Acrochordus arafurae

Filesnakes (Acrochordus arafurae) are large (to 2 m), heavy-bodied snakes of tropical Australia. Sexual dimorphism is evident in adult body sizes, weight/length ratios, and body proportions (relative head and tail lengths). Dimorphism is present even in neonates. Two hypotheses for the evolution of such dimorphism are (1) sexual selection or (2) adaptation of the sexes to different ecological niches. The hypothesis of sexual selection is consistent with general trends of sexually dimorphic body sizes in snakes, and accurately predicts, for A. arafurae, that the larger sex (female) is the one in which reproductive success increases most strongly with increasing body size. However, the sexual dimorphism in relative head sizes is not explicable by sexual selection.The hypothesis of adaptation to sex-specific niches predicts differences in habitats and/or prey. I observed major differences between male and female A. arafurae in prey types, prey sizes and habitat utilization (shallow versus deep water). Hence, the sexual dimorphism in relative head sizes is attributed to ecological causes rather than sexual selection. Nonetheless, competition between the sexes need not be invoked as the selective advantage of this character divergence. It is more parsimonious to interpret these differences as independent adaptations of each sex to increase foraging success, given pre-existing sexually-selected differences in size, habitat or behavior. Data for three other aquatic snake species, from phylogenetically distant taxa, suggest that sexual dimorphism in food habits, foraging sites and feeding morphology, is widespread in snakes.     

Community ecology of African reptiles: historical perspective and a meta-analysis using null models

Community ecology and resource-partitioning patterns of African reptiles have been increasingly studied over the past 30 years. In this paper, I review these studies and provide a meta-analysis based on null models and Monte Carlo simulations (RA2 and RA3 algorithms) to compare patterns of resource partitioning among reptiles in tropical versus nontropical Africa. In general, studies on community ecology of African reptiles have had a strong impact on the global study of resource-partitioning patterns in ectotherms, with such authors as Robert Barbault and Eric Pianka being among the most famous having ever worked in Africa. Despite previous claims, in this paper I suggest that there is no true evidence that reptile biomass is depressed in tropical Africa in comparison with tropical regions of other continents. My null-model reanalysis of 32 independent study systems (twenty from tropical Africa and twelve from nontropical Africa) shows that resource-partitioning patterns are nearly always found in tropical assemblages of reptiles, although this cannot be said for nontropical assemblages of species. Microhabitat (especially for lizards) and food (especially for snakes) are the resource axes typically partitioned by African reptiles.     

Foraging in corallivorous butterflyfish varies with wave exposure

Understanding the foraging patterns of reef fishes is crucial for determining patterns of resource use and the sensitivity of species to environmental change. While changes in prey availability and interspecific competition have been linked to patterns of prey selection, body condition, and survival in coral reef fishes, rarely has the influence of abiotic environmental conditions on foraging been considered. We used underwater digital video to explore how prey availability and wave exposure influence the behavioural time budgets and prey selectivity of four species of obligate coral-feeding butterflyfishes. All four species displayed high selectivity towards live hard corals, both in terms of time invested and frequency of searching and feeding events. However, our novel analysis revealed that such selectivity was sensitive to wave exposure in some species, despite there being no significant differences in the availability of each prey category across exposures. In most cases, these obligate corallivores increased their selectivity towards their most favoured prey types at sites of high wave exposure. This suggests there are costs to foraging under different wave environments that can shape the foraging patterns of butterflyfishes in concert with other conditions such as prey availability, interspecific competition, and territoriality. Given that energy acquisition is crucial to the survival and fitness of fishes, we highlight how such environmental forcing of foraging behaviour may influence the ecological response of species to the ubiquitous and highly variable wave climates of shallow coral reefs.     

Species‐level predation network uncovers high prey specificity in a Neotropical army ant community

Army ants are among the top arthropod predators and considered keystone species in tropical ecosystems. During daily mass raids with many thousand workers, army ants hunt live prey, likely exerting strong top‐down control on prey species. Many tropical sites exhibit a high army ant species diversity (>20 species), suggesting that sympatric species partition the available prey niches. However, whether and to what extent this is achieved has not been intensively studied yet. We therefore conducted a large‐scale diet survey of a community of surface‐raiding army ants at La Selva Biological Station in Costa Rica. We systematically collected 3,262 prey items from eleven army ant species (genera Eciton, Nomamyrmex and Neivamyrmex). Prey items were classified as ant prey or non‐ant prey. The prey nearly exclusively consisted of other ants (98%), and most booty was ant brood (87%). Using morphological characters and DNA barcoding, we identified a total of 1,103 ant prey specimens to the species level. One hundred twenty‐nine ant species were detected among the army ant prey, representing about 30% of the known local ant diversity. Using weighted bipartite network analyses, we show that prey specialization in army ants is unexpectedly high and prey niche overlap very small. Besides food niche differentiation, we uncovered a spatiotemporal niche differentiation in army ant raid activity. We discuss competition‐driven multidimensional niche differentiation and predator–prey arms races as possible mechanisms underlying prey specialization in army ants. By combining systematic prey sampling with species‐level prey identification and network analyses, our integrative approach can guide future research by portraying how predator–prey interactions in complex communities can be reliably studied, even in cases where morphological prey identification is infeasible.     

An Examination of Scale-Dependent Resource Use by Eastern Hognose Snakes in Southcentral New Hampshire

ABSTRACT The decline of many snake populations is attributable to habitat loss, and knowledge of habitat use is critical to their conservation. Resource characteristics (e.g., relative availability of different habitat types, soils, and slopes) within a landscape are scale-dependent and may not be equal across multiple spatial scales. Thus, it is important to identify the relevant spatial scales at which resource selection occurs. We conducted a radiotelemetry study of eastern hognose snake (Heterodon platirhinos) home range size and resource use at different hierarchical spatial scales. We present the results for 8 snakes radiotracked during a 2-year study at New Boston Air Force Station (NBAFS) in southern New Hampshire, USA, where the species is listed by the state as endangered. Mean home range size (minimum convex polygon) at NBAFS (51.7 ± 14.7 ha) was similar to that reported in other parts of the species’ range. Radiotracked snakes exhibited different patterns of resource use at different spatial scales. At the landscape scale (selection of locations within the landscape), snakes overutilized old-field and forest edge habitats and underutilized forested habitats and wetlands relative to availability. At this scale, snakes also overutilized areas containing sandy loam soils and areas with lower slope (mean slope = 5.2% at snake locations vs. 6.7% at random locations). We failed to detect some of these patterns of resource use at the home range scale (i.e., within the home range). Our ability to detect resource selection by the snakes only at the landscape scale is likely the result of greater heterogeneity in macrohabitat features at the broader landscape scale. From a management perspective, future studies of habitat selection for rare species should include measurement of available habitat at spatial scales larger than the home range. We suggest that the maintenance of open early successional habitats as a component of forested landscapes will be critical for the persistence of eastern hognose snake populations in the northeastern United States.     

Does interspecific competition with a stronger competitor explain the rarity of an endangered snake on a Mediterranean island?

Interspecific competition has been demonstrated to be an important shaping force for snake communities worldwide, but relatively few studies have investigated its occurrence and extent with island assemblages of snakes. In Sardinia (Mediterranean Sea), two species of whip snakes (Colubridae) co-occur, one of them being abundant and widespread (Hierophis viridiflavus) and the other being localized and critically endangered (Hemorrhois hippocrepis). A previous hypothesis suggested that the latter species would be confined to suboptimal habitats by the former species, which is a stronger competitor for food. As a consequence, He. hippocrepis would not only be rare but would also be smaller in body size in Sardinia than in other regions of its range where Hi. viridiflavus does not occur. In 1999?C2010, we studied habitat selection, food habits, and body sizes of these two snakes in sympatric populations, applying a suite of statistical tools including null models and Monte Carlo simulations. We observed that dietary habits were different between species (compared to Hi. viridiflavus, He. hippocrepis more frequently preyed upon rodents and less frequently on lizards, and within rodent prey, more frequently upon Rattus and less frequently upon Mus), but not in a way compatible with competitively induced trophic niche partitioning. The two species were similar in terms of prey size and predator-size?Cprey-size relationships. They also differed in habitat selection (maquis was the preferred habitat for both, but Hi. viridiflavus was found significantly more often than He. hippocrepis in grasslands, cultivations, and artificial pinewoods) but again in a non-competitively directed way. In addition, there was no evidence for the hypothesis that He. hippocrepis was confined to suboptimal habitats. Body sizes were similar between species (with males being significantly larger than females), and Sardinian He. hippocrepis were not smaller than conspecifics inhabiting regions without Hi. viridiflavus. Overall, our study showed that the two species differed in some aspects of their ecology, but these differences could not be due to competitive interactions. The conservation implications of these results are also discussed.     

The colouration of the venomous coral snakes (family Elapidae) and their mimics (families Aniliidae and Golubridae)

The bright coloured, highly venomous coral snakes, Leptomicrurus, Micrurus and Micruroides (family Elapidae) and a series of harmless or mildly toxic mimics form an important component of the snake fauna of the Americas. Coral snake patterns are defined as any dorsal pattern found in any species of venomous coral snake and/or any dorsal pattern containing a substantial amount of red, pink or orange distributed so as to resemble that of some species of venomous coral snake. The components of coral snake colouration are described and four principal dorsal patterns are recognized: unicolour, bicolour, tricolour and quadricolour. The tricolour patterns may be further clustered based on the number of black bands or rings separating the red ones as: monads, dyads, triads, tetrads or pentads. A detailed classification of all coral snake colour patterns is presented and each pattern is illustrated. The taxonomic distribution of these patterns is surveyed for mimics and the 56 species of highly venomous coral snakes. Among the latter, the most frequent encountered patterns are tricolour monads, tricolour triads and bicolour rings, in that order. No venomous coral snakes have a tricolour dyad, tricolour tetrad or quadricolour pattern. As many as 115 species of harmless or mildly toxic species, c. 18% of all American snakes, are regarded as coral snake mimics. The colouration and behavioural traits of venomous coral snakes combine to form a significant antipredator defence of an aposematic type. The mimics in turn receive protection from predators that innately or through learning avoid coral snake colour patterns. The precise resemblances in colouration between sympatric non-coral snakes and venomous coral snakes and the concordant geographic variation between the two strongly support this view. Batesian mimicry with the highly venomous coral snakes as the models and the other forms as the mimics is the favoured explanation for this situation. It is further concluded that a number of species in the genera Elaphe, Farancia, Nerodia and Thamnophis, although having red in their colouration, should not be included in the coral snake mimic guild.     

Pitcher plant facilitates prey capture in a sympatric congener

Carnivorous plants avoid below-ground competition for nitrogen by utilizing an alternative nitrogen resource—invertebrate prey, but it remains unclear if sympatric carnivorous plants compete for prey resources. The aim of this study was to investigate if exploitative prey-resource competition occurs between the two sympatric pitcher plant species, Nepenthes rafflesiana and N. gracilis in Singapore. We first investigated if prey-resource partitioning occurs between these two species, and then investigated niche shift in N. gracilis by examining its pitcher contents along an in situ gradient of N. rafflesiana interspecific competition. Our results showed clear evidence of resource partitioning between the two species, but contrary to the expectation of competition, proximity to N. rafflesiana pitchers correlated with higher total prey numbers in N. gracilis pitchers. Our multivariate model of prey assemblages further suggested that N. rafflesiana facilitates N. gracilis prey capture, especially in several ant taxa that are trapped by both species. Concurrently, we found strong evidence for intraspecific competition between N. gracilis pitchers, suggesting that prey resources are exhaustible by pitcher-predation. Our results show that resource partitioning can be associated with facilitative interactions, instead of competition as is usually assumed. Facilitation is more typically expected between phylogenetically distant species, but divergences in resource acquisition strategies can permit facilitation between congeners.     

River habitat homogenisation enhances trophic competition and promotes individual specialisation among young of the year fish

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