Fission-fusion dynamics in the social networks of a North American pitviper.

Many animal species exist in fission-fusion societies, where the size and composition of conspecific groups change spatially and temporally. To help investigate such phenomena, social network analysis (SNA) has emerged as a powerful conceptual and analytical framework for assessing patterns of interconnectedness and quantifying group-level interactions. We leveraged behavioral observations via radiotelemetry and genotypic data from a long-term (>10 years) study on the pitviper Crotalus atrox (western diamondback rattlesnake) and used SNA to quantify the first robust demonstration of social network structures for any free-living snake. Group-level interactions among adults in this population resulted in structurally modular networks (i.e., distinct clusters of interacting individuals) for fidelis use of communal winter dens (denning network), mating behaviors (pairing network), and offspring production (parentage network). Although the structure of each network was similar, the size and composition of groups varied among them. Specifically, adults associated with moderately sized social groups at winter dens but often engaged in reproductive behaviors-both at and away from dens-with different and fewer partners. Additionally, modules formed by individuals in the pairing network were frequently different from those in the parentage network, likely due to multiple mating, long-term sperm storage by females, and resultant multiple paternity. Further evidence for fission-fusion dynamics exhibited by this population-interactions were rare when snakes were dispersing to and traversing their spring-summer home ranges (to which individuals show high fidelity), despite ample opportunities to associate with numerous conspecifics that had highly overlapping ranges. Taken together, we show that long-term datasets incorporating SNA with spatial and genetic information provide robust and unique insights to understanding the social structure of cryptic taxa that are understudied.

Mechanisms underlying interaction frequencies and robustness in a novel seed dispersal network: lessons for restoration.

As human-caused extinctions and invasions accumulate across the planet, understanding the processes governing ecological functions mediated by species interactions, and anticipating the effect of species loss on such functions become increasingly urgent. In seed dispersal networks, the mechanisms that influence interaction frequencies may also influence the capacity of a species to switch to alternative partners (rewiring), influencing network robustness. Studying seed dispersal interactions in novel ecosystems on O'ahu island, Hawai'i, we test whether the same mechanisms defining interaction frequencies can regulate rewiring and increase network robustness to simulated species extinctions. We found that spatial and temporal overlaps were the primary mechanisms underlying interaction frequencies, and the loss of the more connected species affected networks to a greater extent. Further, rewiring increased network robustness, and morphological matching and spatial and temporal overlaps between partners were more influential on network robustness than species abundances. We argue that to achieve self-sustaining ecosystems, restoration initiatives can consider optimal morphological matching and spatial and temporal overlaps between consumers and resources to maximize chances of native plant dispersal. Specifically, restoration initiatives may benefit from replacing invasive species with native species possessing characteristics that promote frequent interactions and increase the probability of rewiring (such as long fruiting periods, small seeds and broad distributions).

Camera traps are an effective tool for monitoring insect-plant interactions.

Insect and pollinator populations are vitally important to the health of ecosystems, food production, and economic stability, but are declining worldwide. New, cheap, and simple monitoring methods are necessary to inform management actions and should be available to researchers around the world. Here, we evaluate the efficacy of a commercially available, close-focus automated camera trap to monitor insect-plant interactions and insect behavior. We compared two video settings-scheduled and motion-activated-to a traditional human observation method. Our results show that camera traps with scheduled video settings detected more insects overall than humans, but relative performance varied by insect order. Scheduled cameras significantly outperformed motion-activated cameras, detecting more insects of all orders and size classes. We conclude that scheduled camera traps are an effective and relatively inexpensive tool for monitoring interactions between plants and insects of all size classes, and their ease of accessibility and set-up allows for the potential of widespread use. The digital format of video also offers the benefits of recording, sharing, and verifying observations.

Ecological correlates of species' roles in highly invaded seed dispersal networks.

Ecosystems with a mix of native and introduced species are increasing globally as extinction and introduction rates rise, resulting in novel species interactions. While species interactions are highly vulnerable to disturbance, little is known about the roles that introduced species play in novel interaction networks and what processes underlie such roles. Studying one of the most extreme cases of human-modified ecosystems, the island of O'ahu, Hawaii, we show that introduced species there shape the structure of seed dispersal networks to a greater extent than native species. Although both neutral and niche-based processes influenced network structure, niche-based processes played a larger role, despite theory predicting neutral processes to be predominantly important for islands. In fact, ecological correlates of species' roles (morphology, behavior, abundance) were largely similar to those in native-dominated networks. However, the most important ecological correlates varied with spatial scale and trophic level, highlighting the importance of examining these factors separately to unravel processes determining species contributions to network structure. Although introduced species integrate into interaction networks more deeply than previously thought, by examining the mechanistic basis of species' roles we can use traits to identify species that can be removed from (or added to) a system to improve crucial ecosystem functions, such as seed dispersal.

A review of conspecific attraction for habitat selection across taxa.

Many species across taxa select habitat based on conspecific presence, known as conspecific attraction. Studies that document conspecific attraction typically provide social information (i.e., cues that indicate the presence of a given species) and then determine if a given species is more likely to settle at locations where the social information is provided compared to those locations that do not. Although the number of studies examining conspecific attraction has grown in recent years, a comprehensive review has not yet been undertaken. Here, we conducted a review of the literature and found 151 studies investigating conspecific attraction across eight taxa. We found that conspecific attraction is widespread with between 80% and 100% of studies, depending on taxa, documenting positive associations between habitat selection and the presence of conspecific cues. Conspecific attraction has been documented more frequently in bird and fish species with less attention given to invertebrate and mammal species. We use the patterns we found to (a) provide an overview of the current state of research on conspecific attraction and (b) discuss how important factors, such as cue characteristics and life history traits, may play a role in shaping conspecific attraction patterns within and across taxa.

Ecomorphological relationships and invasion history of non-native terrestrial bird species on O'ahu, Hawai'i, suggest ecological fitting during novel community assembly.

The widespread introduction of species has created novel communities in many areas of the world. Since introduced species tend to have generalized ecologies and often lack shared evolutionary history with other species in their communities, it would be expected that the relationship between form and function (i.e., ecomorphology) may change in novel communities. We tested this expectation in a subset of the novel bird community on O'ahu, Hawai'i. By relating foraging behavior observations to morphology obtained from live birds at four sites across the island, we found many relationships between species' morphology and foraging ecology that mirrored relationships found in the literature for native-dominated bird communities. Both movement and certain foraging behaviors were related to a species' tarsus-to-wing ratio. Further, bill morphology was related to gleaning, frugivory, and flycatching behaviors. The commonness of significant ecomorphological relationships suggests that, within O'ahu's novel bird community, form is strongly related to function. We hypothesize that ecological fitting likely played a major role in the assembly of this novel community conserving the relationships between form and function found in many other bird communities. To further support this hypothesis, we used niche data from EltonTraits 1.0 to determine whether the establishment of bird species introduced to O'ahu was related to the distinctiveness of their ecological niche from the incumbent community. Introduced species were more likely to establish on O'ahu if their diets were less similar to the bird species already present on the island. Our results support the idea that ecological fitting is an important mechanism in shaping ecological communities, especially in the Anthropocene, thereby influencing novel community assembly and functioning.

Identification of Factors Affecting Predation Risk for Juvenile Turtles using 3D Printed Models.

Although it is widely accepted that juvenile turtles experience high levels of predation, such events are rarely observed, providing limited evidence regarding predator identities and how juvenile habitat selection and availability of sensory cues to predators affects predation risk. We placed three-dimensional printed models resembling juvenile box turtles (Terrapene carolina) across habitats commonly utilized by the species at three sites within their geographical range and monitored models with motion-triggered cameras. To explore how the presence or absence of visual and olfactory cues affected predator interactions with models, we employed a factorial design where models were either exposed or concealed and either did or did not have juvenile box turtle scent applied on them. Predators interacted with 18% of models during field trials. Nearly all interactions were by mesopredators (57%) and rodents (37%). Mesopredators were more likely to attack models than rodents; most (76%) attacks occurred by raccoons (Procyon lotor). Interactions by mesopredators were more likely to occur in wetlands than edges, and greater in edges than grasslands. Mesopredators were less likely to interact with models as surrounding vegetation height increased. Rodents were more likely to interact with models that were closer to woody structure and interacted with exposed models more than concealed ones, but model exposure had no effect on interactions by mesopredators. Scent treatment appeared to have no influence on interactions by either predator group. Our results suggest raccoons can pose high predation risk for juvenile turtles (although rodents could also be important predators) and habitat features at multiple spatial scales affect predator-specific predation risk. Factors affecting predation risk for juveniles are important to consider in management actions such as habitat alteration, translocation, or predator control.

Structure, spatial dynamics, and stability of novel seed dispersal mutualistic networks in Hawai'i.

Increasing rates of human-caused species invasions and extinctions may reshape communities and modify the structure, dynamics, and stability of species interactions. To investigate how such changes affect communities, we performed multiscale analyses of seed dispersal networks on O'ahu, Hawai'i. Networks consisted exclusively of novel interactions, were largely dominated by introduced species, and exhibited specialized and modular structure at local and regional scales, despite high interaction dissimilarity across communities. Furthermore, the structure and stability of the novel networks were similar to native-dominated communities worldwide. Our findings suggest that shared evolutionary history is not a necessary process for the emergence of complex network structure, and interaction patterns may be highly conserved, regardless of species identity and environment. Introduced species can quickly become well integrated into novel networks, making restoration of native ecosystems more challenging than previously thought.

Rapid morphological change of nonnative frugivores on the Hawaiian island of O'ahu.

Novel ecosystems have become widespread created, in part, by the global spread of species. The nonnative species in these environments can be under intense evolutionary pressures that cause rapid morphological change, which can then influence species interactions. In Hawaii, much of the native frugivore community is extinct, replaced by nonnative bird species. Here, we determined if the passerine species of the nonnative frugivore community on O'ahu have morphologically diverged from their native ranges. We compared a variety of traits, all important for frugivory, between museum specimens from the species' native ranges to wild individuals from O'ahu. All four species tested exhibited significant divergence ranging in magnitude from 2.3% to 13.0% difference in at least two traits. Using a method developed from quantitative genetics, we found evidence that a mixture of nonadaptive and adaptive processes worked in concert to create the observed patterns of divergence. Our results suggest that rapid morphological change is occurring and, based on the traits measured, that these changes may influence seed dispersal effectiveness. As these species are largely responsible for seed dispersal on the island, the rapid morphological change of these species can influence the stability and maintenance of plant communities on O'ahu.

Variation in individual temperature preferences, not behavioural fever, affects susceptibility to chytridiomycosis in amphibians.

The ability of wildlife populations to mount rapid responses to novel pathogens will be critical for mitigating the impacts of disease outbreaks in a changing climate. Field studies have documented that amphibians preferring warmer temperatures are less likely to be infected with the fungal pathogen Batrachochytrium dendrobatidis (Bd). However, it is unclear whether this phenomenon is driven by behavioural fever or natural variation in thermal preference. Here, we placed frogs in thermal gradients, tested for temperature preferences and measured Bd growth, prevalence, and the survival of infected animals. Although there was significant individual- and species-level variation in temperature preferences, we found no consistent evidence of behavioural fever across five frog species. Interestingly, for species that preferred warmer temperatures, the preferred temperatures of individuals were negatively correlated with Bd growth on hosts, while the opposite correlation was true for species preferring cooler temperatures. Our results suggest that variation in thermal preference, but not behavioural fever, might shape the outcomes of Bd infections for individuals and populations, potentially resulting in selection for individual hosts and host species whose temperature preferences minimize Bd growth and enhance host survival during epidemics.

An efficient and inexpensive method for measuring long-term thermoregulatory behavior.

Thermoregulatory ability and behavior influence organismal responses to their environment. By measuring thermal preferences, researchers can better understand the effects that temperature tolerances have on ecological and physiological responses to both biotic and abiotic stressors. However, because of funding limitations and confounders, measuring thermoregulation can often be difficult. Here, we provide an effective, affordable (~$50 USD per unit), easy to construct, and validated apparatus for measuring the long-term thermal preferences of animals. In tests, the apparatus spanned temperatures from 9.29 to 33.94°C, and we provide methods to further increase this range. Additionally, we provide simple methods to non-invasively measure animal and substrate temperatures and to prevent temperature preferences of the focal organisms from being confounded with temperature preferences of its prey and its humidity preferences. To validate the apparatus, we show that it was capable of detecting individual-level consistency and among individual-level variation in the preferred body temperatures of Southern toads (Anaxyrus terrestris) and Cuban tree frogs (Osteopilus septentrionalis) over three-weeks. Nearly every aspect of our design is adaptable to meet the needs of a multitude of study systems, including various terrestrial amphibious, and aquatic organisms. The apparatus and methods described here can be used to quantify behavioral thermal preferences, which can be critical for determining temperature tolerances across species and thus the resiliency of species to current and impending climate change.

Do seasonal patterns of rat snake (Pantherophis obsoletus) and black racer (Coluber constrictor) activity predict avian nest predation?

Avian nest success often varies seasonally and because predation is the primary cause of nest failure, seasonal variation in predator activity has been hypothesized to explain seasonal variation in nest success. Despite the fact that nest predator communities are often diverse, recent evidence from studies of snakes that are nest predators has lent some support to the link between snake activity and nest predation. However, the strength of the relationship has varied among studies. Explaining this variation is difficult, because none of these studies directly identified nest predators, the link between predator activity and nest survival was inferred. To address this knowledge gap, we examined seasonal variation in daily survival rates of 463 bird nests (of 17 bird species) and used cameras to document predator identity at 137 nests. We simultaneously quantified seasonal activity patterns of two local snake species (N = 30 individuals) using manual (2136 snake locations) and automated (89,165 movements detected) radiotelemetry. Rat snakes (Pantherophis obsoletus), the dominant snake predator at the site (~28% of observed nest predations), were most active in late May and early June, a pattern reported elsewhere for this species. When analyzing all monitored nests, we found no link between nest predation and seasonal activity of rat snakes. When analyzing only nests with known predator identities (filmed nests), however, we found that rat snakes were more likely to prey on nests during periods when they were moving the greatest distances. Similarly, analyses of all monitored nests indicated that nest survival was not linked to racer activity patterns, but racer-specific predation (N = 17 nests) of filmed nests was higher when racers were moving the greatest distances. Our results suggest that the activity of predators may be associated with higher predation rates by those predators, but that those effects can be difficult to detect when nest predator communities are diverse and predator identities are not known. Additionally, our results suggest that hand-tracking of snakes provides a reliable indicator of predator activity that may be more indicative of foraging behavior than movement frequency provided by automated telemetry systems.

Use of the NatureServe Climate Change Vulnerability Index as an Assessment Tool for Reptiles and Amphibians: Lessons Learned.

Climate change threatens biodiversity globally, yet it can be challenging to predict which species may be most vulnerable. Given the scope of the problem, it is imperative to rapidly assess vulnerability and identify actions to decrease risk. Although a variety of tools have been developed to assess climate change vulnerability, few have been evaluated with regard to their suitability for certain taxonomic groups. Due to their ectothermic physiology, low vagility, and strong association with temporary wetlands, reptiles and amphibians may be particularly vulnerable relative to other groups. Here, we evaluate use of the NatureServe Climate Change Vulnerability Index (CCVI) to assess a large suite of herpetofauna from the Sand Hills Ecoregion of the southeastern United States. Although data were frequently lacking for certain variables (e.g., phenological response to climate change, genetic variation), sufficient data were available to evaluate all 117 species. Sensitivity analyses indicated that results were highly dependent on size of assessment area and climate scenario selection. In addition, several ecological traits common in, but relatively unique to, herpetofauna are likely to contribute to their vulnerability and need special consideration during the scoring process. Despite some limitations, the NatureServe CCVI was a useful tool for screening large numbers of reptile and amphibian species. We provide general recommendations as to how the CCVI tool's application to herpetofauna can be improved through more specific guidance to the user regarding how to incorporate unique physiological and behavioral traits into scoring existing sensitivity factors and through modification to the assessment tool itself.

Power lines, roads, and avian nest survival: effects on predator identity and predation intensity.

1 Anthropogenic alteration of landscapes can affect avian nest success by influencing the abundance, distribution, and behavior of predators. Understanding avian nest predation risk necessitates understanding how landscapes affect predator distribution and behavior. 2 From a sample of 463 nests of 17 songbird species, we evaluated how landscape features (distance to forest edge, unpaved roads, and power lines) influenced daily nest survival. We also used video cameras to identify nest predators at 137 nest predation events and evaluated how landscape features influenced predator identity. Finally, we determined the abundance and distribution of several of the principal predators using surveys and radiotelemetry. 3 Distance to power lines was the best predictor of predator identity: predation by brown-headed cowbirds (Molothrus ater), corvids (Corvus sp. and Cyanocitta cristata), racers (Coluber constrictor), and coachwhips (Masticophis flagellum) increased with proximity to power lines, whereas predation by rat snakes (Elaphe obsoleta) and raptors decreased. In some cases, predator density may reliably indicate nest predation risk because racers, corvids, and cowbirds frequently used power line right-of-ways. 4 Of five bird species with enough nests to analyze individually, daily nest survival of only indigo buntings (Passerina cyanea) decreased with proximity to power lines, despite predation by most predators at our site being positively associated with power lines. For all nesting species combined, distance to unpaved road was the model that most influenced daily nest survival. This pattern is likely a consequence of rat snakes, the locally dominant nest predator (28% of predation events), rarely using power lines and associated areas. Instead, rat snakes were frequently associated with road edges, indicating that not all edges are functionally similar. 5 Our results suggest that interactions between predators and landscape features are likely to be specific to both the local predators and landscape. Thus, predicting how anthropogenic changes to landscapes affect nesting birds requires that we know more about how landscape changes affect the behavior of nest predators and which nest predators are locally important.

Latitudinal variation in seasonal activity and mortality in ratsnakes (Elaphe obsoleta).

The ecology of ectotherms should be particularly affected by latitude because so much of their biology is temperature dependent. Current latitudinal patterns should also be informative about how ectotherms will have to modify their behavior in response to climate change. We used data from a total of 175 adult black ratsnakes (Elaphe obsoleta) radio-tracked in Ontario, Illinois, and Texas, a latitudinal distance of >1500 km, to test predictions about how seasonal patterns of activity and mortality should vary with latitude. Despite pronounced differences in temperatures among study locations, and despite ratsnakes in Texas not hibernating and switching from diurnal to nocturnal activity in the summer, seasonal patterns of snake activity were remarkably similar during the months that snakes in all populations were active. Rather than being a function of temperature, activity may be driven by the timing of reproduction, which appears similar among populations. Contrary to the prediction that mortality should be highest in the most active population, overall mortality did not follow a clinal pattern. Winter mortality did increase with latitude, however, consistent with temperature limiting the northern distribution of ratsnakes. This result was opposite that found in the only previous study of latitudinal variation in winter mortality in reptiles, which may be a consequence of whether or not the animals exhibit true hibernation. Collectively, these results suggest that, at least in the northern part of their range, ratsnakes should be able to adjust easily to, and may benefit from, a warmer climate, although climate-based changes to the snakes' prey or habitat, for example, could alter that prediction.

Linking snake behavior to nest predation in a Midwestern bird community.

Nest predators can adversely affect the viability of songbird populations, and their impact is exacerbated in fragmented habitats. Despite substantial research on this predator-prey interaction, however, almost all of the focus has been on the birds rather than their nest predators, thereby limiting our understanding of the factors that bring predators and nests into contact. We used radiotelemetry to document the activity of two snake species (rat snakes, Elaphe obsoleta; racers, Coluber constrictor) known to prey on nests in Midwestern bird communities and simultaneously monitored 300 songbird nests and tested the hypothesis that predation risk should increase for nests when snakes were more active and in edge habitat preferred by both snake species. Predation risk increased when rat snakes were more active, for all nests combined and for two of the six bird species for which we had sufficient nests to allow separate analyses. This result is consistent with rat snakes being more important nest predators than racers. We found no evidence, however, that nests closer to forest edges were at greater risk. These results are generally consistent with the one previous study that investigated rat snakes and nest predation simultaneously. The seemingly paradoxical failure to find higher predation risk in the snakes' preferred habitat (i.e., edge) might be explained by the snakes using edges at least in part for non-foraging activities. We propose that higher nest predation in fragmented habitats (at least that attributable to snakes) results indirectly from edges promoting larger snake populations, rather than from edges directly increasing the risk of nest predation by snakes. If so, the notion of edges per se functioning as ecological "traps" merits further study.

Prey-mediated effects of drought on condition and survival of a terrestrial snake.

Drought can have severe ecological effects and global climate-change theory predicts that droughts are likely to increase in frequency and severity. Therefore, it is important that we broaden our understanding of how drought affects not only individual species, but also multitrophic interactions. Here we document vegetation and small-mammal abundance and associated patterns of Texas ratsnake (Elaphe obsoleta) body condition and survival before, during, and after a drought in central Texas, USA. Vegetation (grass and forbs) height and small-mammal capture rates were two times greater in wet years compared to the drought year. The decline of small mammals (the snakes' principal prey) during the drought was associated with a drop in ratsnake body condition, consistent with reduced food intake. During the drought, snake mortality also increased 24%. Although higher snake mortality was attributable to predation and road mortality rather than being a direct result of starvation, an increase in risk-prone behavior by foraging snakes probably increased their exposure to those other mortality factors. Drought conditions lasted only for 21 months, and vegetation, small-mammal abundance, and snake condition had returned to pre-drought levels within a year. Although estimates of snake population size were not available, it is likely that substantially more than a year was required for the population to return to its previous size.