Vicuña antipredator diel movement drives spatial nutrient subsidies in a high Andean ecosystem.

Large animals could be important drivers of spatial nutrient subsidies when they ingest resources in some habitats and release them in others, even moving nutrients against elevational gradients. In high Andean deserts, vicuñas (Vicugna vicugna) move daily between nutrient-rich wet meadows, where there is abundant water and forage but high risk of predation by pumas (Puma concolor), and nutrient-poor open plains with lower risk of predation. In all habitats, vicuñas defecate and urinate in communal latrines. We investigated how these latrines impacted soil and plant nutrient concentrations across three habitats in the Andean ecosystem (meadows, plains, and canyons) and used stable isotope analysis to explore the source of fecal nutrients in latrines. Latrine soils had higher concentrations of nitrogen, carbon, and other nutrients than did nonlatrine soils across all habitats. These inputs corresponded with an increase in plant quality (lower C:N) at latrine sites in plains and canyons, but not in meadows. Stable isotope mixing models suggest that ~7% of nutrients in plains latrines originated from vegetation in meadows, which is disproportionately higher than the relative proportion of meadow habitat (2.6%) in the study area. In contrast, ~68% of nutrients in meadow latrines appear to originate from plains and canyon vegetation, though these habitats made up nearly 98% of the study area. Vicuña diel movements thus appear to concentrate nutrients in latrines within habitats and to drive cross-habitat nutrient subsidies, with disproportionate transport from low-lying, nutrient-rich meadows to more elevated, nutrient-poor plains. When these results are scaled up to the landscape scale, the amount of nitrogen and phosphorus subsidized in soil at plains latrines was of the same order of magnitude as estimates of annual atmospheric nitrogen and phosphorus deposition for this region (albeit far more localized and patchy). Thus, vicuña-mediated nutrient redistribution and deposition appears to be an important process impacting ecosystem functioning in arid Andean environments, on par with other major inputs of nutrients to the system.

Cascading effects of a disease outbreak in a remote protected area.

Disease outbreaks induced by humans increasingly threaten wildlife communities worldwide. Like predators, pathogens can be key top-down forces in ecosystems, initiating trophic cascades that may alter food webs. An outbreak of mange in a remote Andean protected area caused a dramatic population decline in a mammalian herbivore (the vicuña), creating conditions to test the cascading effects of disease on the ecological community. By comparing a suite of ecological measurements to pre-disease baseline records, we demonstrate that mange restructured tightly linked trophic interactions previously driven by a mammalian predator (the puma). Following the mange outbreak, scavenger (Andean condor) occurrence in the ecosystem declined sharply and plant biomass and cover increased dramatically in predation refuges where herbivory was historically concentrated. The evidence shows that a disease-induced trophic cascade, mediated by vicuña density, could supplant the predator-induced trophic cascade, mediated by vicuña behaviour, thereby transforming the Andean ecosystem.

A methodological roadmap to quantify animal-vectored spatial ecosystem subsidies.

Energy, nutrients and organisms move over landscapes, connecting ecosystems across space and time. Meta-ecosystem theory investigates the emerging properties of local ecosystems coupled spatially by these movements of organisms and matter, by explicitly tracking exchanges of multiple substances across ecosystem borders. To date, meta-ecosystem research has focused mostly on abiotic flows-neglecting biotic nutrient flows. However, recent work has indicated animals act as spatial nutrient vectors when they transport nutrients across landscapes in the form of excreta, egesta and their own bodies. Partly due to its high level of abstraction, there are few empirical tests of meta-ecosystem theory. Furthermore, while animals may be viewed as important mediators of ecosystem functions, better integration of tools is needed to develop predictive insights of their relative roles and impacts on diverse ecosystems. We present a methodological roadmap that explains how to do such integration by discussing how to combine insights from movement, foraging and ecosystem ecology to develop a coherent understanding of animal-vectored nutrient transport on meta-ecosystems processes. We discuss how the slate of newly developed technologies and methods-tracking devices, mechanistic movement models, diet reconstruction techniques and remote sensing-that when integrated have the potential to advance the quantification of animal-vectored nutrient flows and increase the predictive power of meta-ecosystem theory. We demonstrate that by integrating novel and established tools of animal ecology, ecosystem ecology and remote sensing, we can begin to identify and quantify animal-mediated nutrient translocation by large animals. We also provide conceptual examples that show how our proposed integration of methodologies can help investigate ecosystem impacts of large animal movement. We conclude by describing practical advancements to understanding cross-ecosystem contributions of animals on the move. Understanding the mechanisms by which animals shape ecosystem dynamics is important for ongoing conservation, rewilding and restoration initiatives around the world, and for developing more accurate models of ecosystem nutrient budgets. Our roadmap will enable ecologists to better qualify and quantify animal-mediated nutrient translocation for animals on the move.

Author Correction: An alternative hypothesis for the evolution of same-sex sexual behaviour in animals.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

An alternative hypothesis for the evolution of same-sex sexual behaviour in animals.

Same-sex sexual behaviour (SSB) has been recorded in over 1,500 animal species with a widespread distribution across most major clades. Evolutionary biologists have long sought to uncover the adaptive origins of 'homosexual behaviour' in an attempt to resolve this apparent Darwinian paradox: how has SSB repeatedly evolved and persisted despite its presumed fitness costs? This question implicitly assumes that 'heterosexual' or exclusive different-sex sexual behaviour (DSB) is the baseline condition for animals, from which SSB has evolved. We question the idea that SSB necessarily presents an evolutionary conundrum, and suggest that the literature includes unchecked assumptions regarding the costs, benefits and origins of SSB. Instead, we offer an alternative null hypothesis for the evolutionary origin of SSB that, through a subtle shift in perspective, moves away from the expectation that the origin and maintenance of SSB is a problem in need of a solution. We argue that the frequently implicit assumption of DSB as ancestral has not been rigorously examined, and instead hypothesize an ancestral condition of indiscriminate sexual behaviours directed towards all sexes. By shifting the lens through which we study animal sexual behaviour, we can more fruitfully examine the evolutionary history of diverse sexual strategies.

Eco-Evolutionary Dynamics: The Predator-Prey Adaptive Play and the Ecological Theater.

The emerging field of eco-evolutionary dynamics has demonstrated that both ecological and evolutionary processes can occur contemporaneously. Ecological interactions, such as between predator and prey, are important focal areas where an eco-evolutionary perspective can advance understanding about phenotypically plastic and adaptive evolutionary responses. In predator-prey interactions, both species reciprocally respond and adapt to each other in order to simultaneously ensure resource consumption and predation avoidance. Here we sketch out a way to help unify experimental and analytical approaches to both eco-evolutionary dynamics and predator-prey interactions, with a specific focus on terrestrial systems. We discuss the need to view predator-prey eco-evolutionary dynamics as a perpetually adaptive interplay with constantly shifting pressures and feedbacks, rather than viewing it as driving a set evolutionary trajectory. We then outline our perspective on how to understand eco-evolutionary patterns in a predator-prey context. We propose initiating insight by distinguishing phenotypic plasticity against genetic change (i.e., "molecular reductionism") and further applying a landscape-scale perspective (i.e., "landscape holism"). We believe that studying predator-prey interactions under an eco-evolutionary lens can provide insights into how general and, consequently, predictable species' evolutionary responses are to their contemporary environments.