Investigating Carnivore Guild Structure: Spatial and Temporal Relationships amongst Threatened Felids in Myanmar

The co-occurrence of felid species in Southeast Asia provides an unusual opportunity to investigate guild structure and the factors controlling it. Using camera-trap data, we quantified the space use, temporal activity, and multi-dimensional niche overlap of the tiger, clouded leopard, Asiatic golden cat, marbled cat, and leopard cat in the Htamanthi Wildlife Sanctuary, Myanmar. We hypothesised that the spatio-temporal behaviour of smaller cats would reflect the avoidance of the larger cats, and similar-sized guild members would partition their niches in space or time to reduce resource competition. Our approach involved modelling single-species occupancy, pairwise spatial overlap using Bayesian inference, activity overlap with kernel density estimation, and multivariate analyses. The felid assembly appeared to be partitioned mainly on a spatial rather than temporal dimension, and no significant evidence of mesopredator release was observed. Nonetheless, the temporal association between the three mesopredators was inversely related to the similarity in their body sizes. The largest niche differences in the use of space and time occurred between the three smallest species. This study offers new insight into carnivore guild assembly and adds substantially to knowledge of five of the least known felids of conservation concern.

Terrestrial mesopredators did not increase after top-predator removal in a large-scale experimental test of mesopredator release theory

Removal or loss of top-predators has been predicted to cause cascading negative effects for ecosystems, including mesopredator release. However, reliable evidence for these processes in terrestrial systems has been mixed and equivocal due, in large part, to the systemic and continued use of low-inference study designs to investigate this issue. Even previous large-scale manipulative experiments of strong inferential value have been limited by experimental design features (i.e. failure to prevent migration between treatments) that constrain possible inferences about the presence or absence of mesopredator release effects. Here, we build on these previous strong-inference experiments and report the outcomes of additional large-scale manipulative experiments to eradicate Australian dingoes from two fenced areas where dingo migration was restricted and where theory would predict an increase in extant European red foxes, feral cats and goannas. We demonstrate the removal and suppression of dingoes to undetectable levels over 4–5 years with no corresponding increases in mesopredator relative abundances, which remained low and stable throughout the experiment at both sites. We further demonstrate widespread absence of negative relationships between predators, indicating that the mechanism underpinning predicted mesopredator releases was not present. Our results are consistent with all previous large-scale manipulative experiments and long-term mensurative studies which collectively demonstrate that (1) dingoes do not suppress red foxes, feral cats or goannas at the population level, (2) repeated, temporary suppression of dingoes in open systems does not create mesopredator release effects, and (3) removal and sustained suppression of dingoes to undetectable levels in closed systems does not create mesopredator release effects either. Our experiments add to similar reports from North America, Asia, Europe and southern Africa which indicate that not only is there a widespread absence of reliable evidence for these processes, but there is also a large and continually growing body of experimental evidence of absence for these processes in many terrestrial systems. We conclude that although sympatric predators may interact negatively with each other on smaller spatiotemporal scales, that these negative interactions do not always scale-up to the population level, nor are they always strong enough to create mesopredator suppression or release effects.

Variable strategies to solve risk–reward tradeoffs in carnivore communities

Mesopredator release theory suggests that dominant predators suppress subordinate carnivores and ultimately shape community dynamics, but the assumption that subordinate species are only negatively affected ignores the possibility of facilitation through scavenging. We examined the interplay within a carnivore community consisting of cougars, coyotes, black bears, and bobcats using contemporaneous Global Positioning System telemetry data from 51 individuals; diet analysis from 972 DNA-metabarcoded scats; and data from 128 physical investigations of cougar kill sites, 28 of which were monitored with remote cameras. Resource provisioning from competitively dominant cougars to coyotes through scavenging was so prolific as to be an overwhelming determinant of coyote behavior, space use, and resource acquisition. This was evident via the strong attraction of coyotes to cougar kill sites, frequent scavenging of cougar-killed prey, and coyote diets that nearly matched cougars in the magnitude of ungulate consumption. Yet coyotes were often killed by cougars and used space to minimize encounters, complicating the fitness benefits gained from scavenging. We estimated that 23% (95% CI: 8 to 55%) of the coyote population in our study area was killed by cougars annually, suggesting that coyote interactions with cougars are a complex behavioral game of risk and reward. In contrast, we found no indication that bobcat space use or diet was influenced by cougars. Black bears avoided cougars, but there was no evidence of attraction to cougar kill sites and much lower levels of ungulate consumption and carcass visitation than for coyotes. Interspecific interactions among carnivores are multifaceted, encompassing both suppression and facilitation.

The selection of anthropogenic habitat by wildlife as an ecological consequence of rural exodus: empirical examples from Spain

The increasing urbanization of the landscape is a major component of global change worldwide. However, it is puzzling that wildlife is selecting anthropogenic habitats despite the availability of apparently high–quality semi–natural (i.e. less intensively modified) habitats. Definitive explanations for this process are still lacking. We have previously suggested that colonization of the urban habitat is initially triggered by ecological processes that take place outside urban areas as a consequence of past rural exodus. Here we present a diverse array of examples of selection of several types of anthropogenic habitat by wildlife in Spain (including transportation infrastructure, human–exclusion areas, urban areas under construction, cities, reservoirs, quarries and landfills) in support of this idea. Wildlife is moving out of its historical ecological refuges and losing fear of harmless urban humans. Mesopredators are rebounding by mesopredator release, due to ceased human persecution, and shrubs and trees are claiming former agricultural habitats. Together, these factors force many species to move to urbanized areas where they find open habitats, food associated with these habitats, and protection against predation. Hence, the classical balance of costs and benefits that takes place once inside urban areas, would actually be a second step of the process of colonization of urban areas. A better understanding of the initial triggers of urban colonization could help us increase the biological value of human–made habitats for wildlife in the future.

Density trends of wild felids in northern Laos

Determining the density trends of a guild of species can help illuminate their interactions, and the impacts that humans might have on them. We estimated the density trends from 2013 to 2017 of the clouded leopard Neofelis nebulosa, leopard cat Prionailurus bengalensis and marbled cat Pardofelis marmorata in Nam Et—Phou Louey National Park (NEPL), Laos, using camera trap data and spatial capture-recapture models. Mean (± SD) density estimates (individuals/100 km2) for all years were 1.77 ± 0.30 for clouded leopard, 1.50 ± 0.30 for leopard cat, and 3.80 ± 0.70 for marbled cat. There was a declining trend in density across the study years for all three species, with a ≥ 90% probability of decline for clouded leopard and leopard cat and an 83% probability of decline for marbled cat. There was no evidence that mesopredator release occurred as a result of tiger (Panthera tigris) and leopard (P. pardus) extirpations. We believe that snaring, the factor that led to the extirpation of tiger and leopard in NEPL, is now contributing to the decline of smaller felids, to an extent that over-rides any potential effects of mesopredator release on their densities and interactions. We recommend that the NEPL managers implement a more systematic and intensified snare removal program, in concert with extensive community outreach and engagement of local people to prevent the setting of snares. These actions might be the only hope for saving the remaining members of the felid community in NEPL.

Games of risk and reward in carnivore communities

Mesopredator release theory suggests that dominant predators suppress subordinate carnivores and ultimately shape community dynamics, but the assumption that subordinate species are only negatively affected ignores the possibility of facilitation through scavenging. We examined the interplay within a carnivore community consisting of cougars, coyotes, black bears, and bobcats using contemporaneous Global Positioning System telemetry data from 51 individuals, diet analysis from 972 DNA-metabarcoded scats, and data from 128 physical investigations of cougar kill sites, 28 of which were monitored with remote cameras. Resource provisioning from competitively-dominant cougars to coyotes through scavenging was so prolific as to be an overwhelming determinant of coyote behavior, space use, and resource acquisition. This was evident via strong attraction of coyotes to cougar kill sites, frequent scavenging of cougar-killed prey, and coyote diets that nearly matched cougars in the magnitude of ungulate consumption. Yet coyotes were often killed by cougars and used space to minimize encounters, complicating the fitness benefits gained from scavenging. We estimated that 23% (95% CI: 8–55%) of the coyote population in our study area was killed by cougars annually suggesting that coyote interactions with cougars are a complex behavioral game of risk and reward. In contrast, we found no indication that bobcat space use or diet was influenced by cougars. Black bears avoided cougars, but there was no evidence of attraction to cougar kill sites, and much lower levels of ungulate consumption and carcass visitation than for coyotes. Interspecific interactions among carnivores are multifaceted encompassing both suppression and facilitation.Significance StatementAn incomplete understanding of the total influence top predators exert on subordinate species hinders our ability to anticipate the effects that changing carnivore populations will have in ecological communities. Here we show that cougars are the architects of a complex behavioral game of risk and reward, as subordinate or co-occurring carnivores are both provisioned and preyed on by the top predators. Each co-occurring carnivore species considered here employed a different strategy to approach the risk-reward tradeoff suggesting there are multiple viable solutions to the game. By not considering the multitude of effects top predators have on other carnivores, we are missing important linkages in terrestrial food webs.

Investigating carnivore guild structure: Spatial and temporal relationships amongst threatened felids in Myanmar

At least nine felid species can co-occur in Southeast Asia, thus providing an unusual opportunity to investigate poorly known guild structure and the factors controlling it. Using camera-trap data, we quantified space use, temporal activity, and multi-dimensional niche overlap of tiger, clouded leopard, marbled cat, leopard cat, and Asiatic golden cat in Htamanthi Wildlife Sanctuary of Myanmar. We hypothesized that the spatio-temporal behaviour of smaller cats can reflect avoidance of the larger cats, which are both potential competitors and predators, and similar-sized guild members would partition their niches in space or time to reduce competition for resources. Our approach involved single-species occupancy modelling to identify site covariates, pairwise spatial overlap using Bayesian inference, and activity overlap with Kernel density estimation and multivariate analyses to test hypotheses. We found tiger and marbled cats were primarily diurnal, clouded leopard and leopard cat were nocturnal and golden cat exhibited cathemeral activity. We observed a complex pattern of guild assembly and potential competition involving strong niche displacement between the golden cat and marbled cat, but high overlap between the relatively similarly-sized pairing of clouded leopard and golden cat, and the markedly differently-sized tiger – golden cat pairing. No significant evidence of mesopredator release was observed and the felid assembly in Northern Myanmar appeared to be partitioned mainly on a spatial, rather than temporal, dimension. Nonetheless, the temporal association between the three mesopredators was inversely related to the similarity in their body sizes. The insights into this felid guild revealed that the largest niche differences in the use of space and time occurred between the three smaller species, most evidently between the Asiatic golden cat-marbled cat pairing, followed by marbled cat - leopard cat pairing. This study offers new insight into carnivore guild assembly and, adds substantially to knowledge of five of the least known felids of conservation concern.

3D Geometric Morphometrics Reveals Convergent Character Displacement in the Central European Contact Zone between Two Species of Hedgehogs (Genus Erinaceus)

Hedgehogs, as medium-sized plantigrade insectivores with low basal metabolic rates and related defensive anti-predator strategies, are quite sensitive to temperature and ecosystem productivity. Their ranges therefore changed dramatically due to Pleistocene climate oscillations, resulting in allopatric speciation and the subsequent formation of secondary contact zones. Such interactions between closely related species are known to generate strong evolutionary forces responsible for niche differentiation. In this connection, here, we detail the results of research on the phenotypic evolution in the two species of hedgehog present in central Europe, as based on genetics and geometric morphometrics in samples along a longitudinal transect that includes the contact zone between the species. While in allopatry, Erinaceus europaeus is found to have a larger skull than E. roumanicus and distinct cranial and mandibular shapes; the members of the two species in sympatry are smaller and more similar to each other, with a convergent shape of the mandible. The relevant data fail to reveal any major role for either hybridisation or clinal variation. We, therefore, hypothesise that competitive pressure exerted on the studied species does not generate divergent selection sufficient for divergent character displacement to evolve, instead giving rise to convergent selection in the face of resource limitation in the direction of smaller skull size. Considering the multi-factorial constraints present in the relevant adaptive landscape, reduction in size could also be facilitated by predator pressure in ecosystems characterised by mesopredator release and other anthropogenic factors. As the function of the animals’ lower jaw is mainly connected with feeding (in contrast to the cranium whose functions are obviously more complex), we interpret the similarity in shape as reflecting local adaptations to overlapping dietary resources in the two species and hence as convergent character displacement.

Invaders induce coordinated isotopic niche shifts in native fish species

Food-web investigations inform management strategies by exposing potential interactions between native and nonnative species and anticipating likely outcomes associated with species removal efforts. We leveraged a natural gradient of compositional turnover from native-only to nonnative-only fish assemblages, combined with an intensive removal effort, to investigate underlying food-web changes in response to invasive species expansion in a Lower Colorado River tributary. Nonnative fishes caused coordinated isotopic niche displacement in native fishes by inducing resource shifts toward lower trophic positions and enriched carbon sources. By contrast, nonnative fishes did not experience reciprocal shifts when native fishes were present. Asymmetrical outcomes between native and nonnative fishes indicated species displacement may result from competitive or consumptive interactions. Native species’ isotopic niches returned to higher trophic levels after nonnative green sunfish (Lepomis cyanellus) removal, indicating removal efforts can support trophic recovery of native fishes like desert suckers (Catostomus clarkii) and roundtail chub (Gila robusta). Using stable isotope analysis in preremoval assessments provides opportunities to identify asymmetric interactions, whereas postremoval assessments could identify unintended consequences, like mesopredator release, as part of adaptive decision making to recover native fishes.