Preventing extinction in an age of species migration and planetary change.

International and national conservation policies almost exclusively focus on conserving species in their historic native ranges, thus excluding species that have been introduced by people and some of those that have extended their ranges on their own accord. Given that many of such migrants are threatened in their native ranges, conservation goals that explicitly exclude these populations may overlook opportunities to prevent extinctions and respond dynamically to rapidly changing environmental and climatic conditions. Focusing on terrestrial mammals, we quantified the number of threatened mammals that have established new populations through assisted migration (i.e., introduction). We devised 4 alternative scenarios for the inclusion of assisted-migrant populations in mainstream conservation policy with the aim of preventing global species extinctions. We then used spatial prioritization algorithms to simulate how these scenarios could change global spatial conservation priorities. We found that 22% (70 species out of 265) of all identified assisted-migrant mammals were threatened in their native ranges, mirroring the 25% of all mammals that are threatened. Reassessing global threat statuses by combining native and migrant ranges reduced the threat status of 23 species (∼33% of threatened assisted migrants). Thus, including migrant populations in threat assessments provides a more accurate assessment of actual global extinction risk among species. Spatial prioritization simulations showed that reimagining the role of assisted-migrant populations in preventing species extinction could increase the importance of overlooked landscapes, particularly in central Australia, Europe, and the southwestern United States. Our results indicated that these various and nonexhaustive ways to consider assisted-migrant populations, with due consideration of potential conservation conflicts with resident taxa, may provide unprecedented opportunities to prevent species extinctions.

Prevención de la extinción en una época de migración de especies y cambios planetarios Resumen Las políticas de conservación nacionales e internacionales casi siempre se enfocan en la conservación de las especies dentro de su distribución histórica y nativa, por lo que se excluyen especies que han sido introducidas por el humano y algunas que se han extendido por cuenta propia más allá de su distribución. Ya que muchas de estas especies migrantes están amenazadas dentro de su distribución nativa, los objetivos de conservación que excluyen explícitamente a estas poblaciones pueden ignorar las oportunidades para prevenir extinciones y responder de forma dinámica a las condiciones ambientales y climáticas que cambian con rapidez. Nos enfocamos en los mamíferos terrestres para cuantificar el número de especies amenazadas que han establecido poblaciones nuevas mediante la migración asistida (introducción). Diseñamos cuatro escenarios alternativos para la inclusión de las poblaciones con migración asistida dentro de las políticas de conservación generales con el objetivo de prevenir extinciones globales de especies. Después usamos algoritmos de priorización espacial para simular cómo estos escenarios podrían cambiar las prioridades de conservación espacial en todo el mundo. Descubrimos que el 22% (70 de 765 especies) de todos los mamíferos con migración asistida están amenazados dentro de su distribución nativa, lo que es similar al 25% de especies amenazadas de todas las especies de mamíferos. La reevaluación de los estados mundiales de amenaza mediante la combinación de la distribución nativa y migrante redujo el estado de amenaza de 23 especies (∼33% de los migrantes asistidos amenazados). Por esto, incluir a las poblaciones migrantes en la evaluación de amenazas proporciona una evaluación más certera del riesgo de extinción que existe entre las especies a nivel mundial. Las simulaciones de priorización espacial mostraron que reinventar el papel que tienen las poblaciones con migración asistida en la prevención de la extinción de especies podría incrementar la importancia de los paisajes ignorados, particularmente en Australia central, Europa y el suroeste de los Estados Unidos. Nuestros resultados indican que estas maneras diversas y no exhaustivas de considerar a las poblaciones con migración asistida, con la debida consideración de los potenciales conflictos de conservación con los taxones residentes, puede proporcionar oportunidades sin precedentes para prevenir la extinción de las especies.

Introduced herbivores restore Late Pleistocene ecological functions.

Large-bodied mammalian herbivores dominated Earth's terrestrial ecosystems for several million years before undergoing substantial extinctions and declines during the Late Pleistocene (LP) due to prehistoric human impacts. The decline of large herbivores led to widespread ecological changes due to the loss of their ecological functions, as driven by their unique combinations of traits. However, recently, humans have significantly increased herbivore species richness through introductions in many parts of the world, potentially counteracting LP losses. Here, we assessed the extent to which introduced herbivore species restore lost-or contribute novel-functions relative to preextinction LP assemblages. We constructed multidimensional trait spaces using a trait database for all extant and extinct mammalian herbivores ≥10 kg known from the earliest LP (∼130,000 ybp) to the present day. Extinction-driven contractions of LP trait space have been offset through introductions by ∼39% globally. Analysis of trait space overlap reveals that assemblages with introduced species are overall more similar to those of the LP than native-only assemblages. This is because 64% of introduced species are more similar to extinct rather than extant species within their respective continents. Many introduced herbivores restore trait combinations that have the capacity to influence ecosystem processes, such as wildfire and shrub expansion in drylands. Although introduced species have long been a source of contention, our findings indicate that they may, in part, restore ecological functions reflective of the past several million years before widespread human-driven extinctions.

A novel trophic cascade between cougars and feral donkeys shapes desert wetlands.

Introduced large herbivores have partly filled ecological gaps formed in the late Pleistocene, when many of the Earth's megafauna were driven extinct. However, extant predators are generally considered incapable of exerting top-down influences on introduced megafauna, leading to unusually strong disturbance and herbivory relative to native herbivores. We report on the first documented predation of juvenile feral donkeys Equus africanus asinus by cougars Puma concolor in the Mojave and Sonoran Deserts of North America. We then investigated how cougar predation corresponds with differences in feral donkey behaviour and associated effects on desert wetlands. Focusing on a feral donkey population in the Death Valley National Park, we used camera traps and vegetation surveys to compare donkey activity patterns and impacts between wetlands with and without cougar predation. Donkeys were primarily diurnal at wetlands with cougar predation, thereby avoiding cougars. However, donkeys were active throughout the day and night at sites without predation. Donkeys were ~87% less active (measured as hours of activity a day) at wetlands with predation (p < 0.0001). Sites with predation had reduced donkey disturbance and herbivory, including ~46% fewer access trails, 43% less trampled bare ground and 192% more canopy cover (PERMANOVA, R2  = 0.22, p = 0.0003). Our study is the first to reveal a trophic cascade involving cougars, feral equids and vegetation. Cougar predation appears to rewire an ancient food web, with diverse implications for modern ecosystems. Our results suggest that protecting apex predators could have important implications for the ecological effects of introduced megafauna.

Animal cognition and culture mediate predator-prey interactions.

Predator-prey ecology and the study of animal cognition and culture have emerged as independent disciplines. Research combining these disciplines suggests that both animal cognition and culture can shape the outcomes of predator-prey interactions and their influence on ecosystems. We review the growing body of work that weaves animal cognition or culture into predator-prey ecology, and argue that both cognition and culture are significant but poorly understood mechanisms mediating how predators structure ecosystems. We present a framework exploring how previous experiences with the predation process creates feedback loops that alter the predation sequence. Cognitive and cultural predator-prey ecology offers ecologists new lenses through which to understand species interactions, their ecological consequences, and novel methods to conserve wildlife in a changing world.

Meta-analysis shows that wild large herbivores shape ecosystem properties and promote spatial heterogeneity.

Megafauna (animals ≥45 kg) have probably shaped the Earth's terrestrial ecosystems for millions of years with pronounced impacts on biogeochemistry, vegetation, ecological communities and evolutionary processes. However, a quantitative global synthesis on the generality of megafauna effects on ecosystems is lacking. Here we conducted a meta-analysis of 297 studies and 5,990 individual observations across six continents to determine how wild herbivorous megafauna influence ecosystem structure, ecological processes and spatial heterogeneity, and whether these impacts depend on body size and environmental factors. Despite large variability in megafauna effects, we show that megafauna significantly alter soil nutrient availability, promote open vegetation structure and reduce the abundance of smaller animals. Other responses (14 out of 26), including, for example, soil carbon, were not significantly affected. Further, megafauna significantly increase ecosystem heterogeneity by affecting spatial heterogeneity in vegetation structure and the abundance and diversity of smaller animals. Given that spatial heterogeneity is considered an important driver of biodiversity across taxonomic groups and scales, these results support the hypothesis that megafauna may promote biodiversity at large scales. Megafauna declined precipitously in diversity and abundance since the late Pleistocene, and our results indicate that their restoration would substantially influence Earth's terrestrial ecosystems.

Functional traits-not nativeness-shape the effects of large mammalian herbivores on plant communities.

Large mammalian herbivores (megafauna) have experienced extinctions and declines since prehistory. Introduced megafauna have partly counteracted these losses yet are thought to have unusually negative effects on plants compared with native megafauna. Using a meta-analysis of 3995 plot-scale plant abundance and diversity responses from 221 studies, we found no evidence that megafauna impacts were shaped by nativeness, "invasiveness," "feralness," coevolutionary history, or functional and phylogenetic novelty. Nor was there evidence that introduced megafauna facilitate introduced plants more than native megafauna. Instead, we found strong evidence that functional traits shaped megafauna impacts, with larger-bodied and bulk-feeding megafauna promoting plant diversity. Our work suggests that trait-based ecology provides better insight into interactions between megafauna and plants than do concepts of nativeness.

Long-term biotic homogenization in the East African Rift System over the last 6 million years of hominin evolution.

Eastern Africa preserves the most complete record of human evolution anywhere in the world but we have little knowledge of how long-term biogeographic dynamics in the region influenced hominin diversity and distributions. Here, we use spatial beta diversity analyses of mammal fossil records from the East African Rift System to reveal long-term biotic homogenization (increasing compositional similarity of faunas) over the last 6 Myr. Late Miocene and Pliocene faunas (~6-3 million years ago (Ma)) were largely composed of endemic species, with the shift towards biotic homogenization after ~3 Ma being driven by the loss of endemic species across functional groups and a growing number of shared grazing species. This major biogeographic transition closely tracks the regional expansion of grass-dominated ecosystems. Although grazers exhibit low beta diversity in open environments of the Early Pleistocene, the high beta diversity of Mio-Pliocene browsers and frugivores occurred in the context of extensive woody vegetation. We identify other key aspects of the Late Cenozoic biogeographic development of eastern Africa, their likely drivers and place the hominin fossil record in this context. Because hominins were undoubtedly influenced by many of the same factors as other eastern African mammals, this provides a new perspective on the links between environmental and human evolutionary histories.

Australia's recently established predators restore complexity to food webs simplified by extinction.

Since prehistory, humans have altered the composition of ecosystems by causing extinctions and introducing species. However, our understanding of how waves of species extinctions and introductions influence the structure and function of ecological networks through time remains piecemeal. Here, focusing on Australia, which has experienced many extinctions and introductions since the Late Pleistocene, we compared the functional trait composition of Late Pleistocene (130,00-115,000 years before present [ybp]), Holocene (11,700-3,000 ybp), and current Australian mammalian predator assemblages (≥70% vertebrate meat consumption; ≥1 kg adult body mass). We then constructed food webs for each period based on estimated prey body mass preferences. We found that introduced predators are functionally distinct from extinct Australian predators, but they rewire food webs toward a state that closely resembles the Late Pleistocene, prior to the megafauna extinctions. Both Late Pleistocene and current-day food webs consist of an apex predator and three smaller predators. This leads to food web networks with a similar total number of links, link densities, and compartmentalizations. However, this similarity depends on the presence of dingoes: in their absence, food webs become simplified and reminiscent of those following the Late Pleistocene extinctions. Our results suggest that recently established predators, even those implicated in species extinctions and declines, can restore complexity to food webs simplified by extinction.

Worldwide Late Pleistocene and Early Holocene population declines in extant megafauna are associated with Homo sapiens expansion rather than climate change.

The worldwide extinction of megafauna during the Late Pleistocene and Early Holocene is evident from the fossil record, with dominant theories suggesting a climate, human or combined impact cause. Consequently, two disparate scenarios are possible for the surviving megafauna during this time period - they could have declined due to similar pressures, or increased in population size due to reductions in competition or other biotic pressures. We therefore infer population histories of 139 extant megafauna species using genomic data which reveal population declines in 91% of species throughout the Quaternary period, with larger species experiencing the strongest decreases. Declines become ubiquitous 32-76 kya across all landmasses, a pattern better explained by worldwide Homo sapiens expansion than by changes in climate. We estimate that, in consequence, total megafauna abundance, biomass, and energy turnover decreased by 92-95% over the past 50,000 years, implying major human-driven ecosystem restructuring at a global scale.

Cicada nymphs dominate American black bear diet in a desert riparian area.

American black bears are considered dependent on high-elevation forests or other montane habitats in the drylands of western North America. Black bear sign, including that of cubs, was observed throughout the summers of 2015, 2016, and 2018 along a perennial desert river in the Sonoran Desert of Arizona. We analyzed the contents of 21 black bear scats, collected from May to October of 2016 and 2018. Apache cicada nymphs (Diceroprocta apache) were the dominant food item, occurring in 90% of scats and comprising an average of 59% of scat contents. In the process of excavating these nymphs, bears created large areas of turned-over soil, a form of ecosystem engineering with potential implications for soils, vegetation, and fluvial geomorphology. Given that species distributions are shaped by physiological and ecological contexts, as well as anthropogenic legacies, it is possible that black bears once occurred more commonly in desert riparian systems prior to widespread agricultural development, hunting, and dewatering. Although more research is necessary, we suggest that desert riparian systems may be an alternative habitat for black bears. Better understanding the diet and habitat breadth of American black bears is important in the context of increasing landscape fragmentation and militarization in the U.S.-Mexican borderlands.

Equids engineer desert water availability.

Megafauna play important roles in the biosphere, yet little is known about how they shape dryland ecosystems. We report on an overlooked form of ecosystem engineering by donkeys and horses. In the deserts of North America, digging of ≤2-meter wells to groundwater by feral equids increased the density of water features, reduced distances between waters, and, at times, provided the only water present. Vertebrate richness and activity were higher at equid wells than at adjacent dry sites, and, by mimicking flood disturbance, equid wells became nurseries for riparian trees. Our results suggest that equids, even those that are introduced or feral, are able to buffer water availability, which may increase resilience to ongoing human-caused aridification.

Functional traits of the world's late Quaternary large-bodied avian and mammalian herbivores.

Prehistoric and recent extinctions of large-bodied terrestrial herbivores had significant and lasting impacts on Earth's ecosystems due to the loss of their distinct trait combinations. The world's surviving large-bodied avian and mammalian herbivores remain among the most threatened taxa. As such, a greater understanding of the ecological impacts of large herbivore losses is increasingly important. However, comprehensive and ecologically-relevant trait datasets for extinct and extant herbivores are lacking. Here, we present HerbiTraits, a comprehensive functional trait dataset for all late Quaternary terrestrial avian and mammalian herbivores ≥10 kg (545 species). HerbiTraits includes key traits that influence how herbivores interact with ecosystems, namely body mass, diet, fermentation type, habitat use, and limb morphology. Trait data were compiled from 557 sources and comprise the best available knowledge on late Quaternary large-bodied herbivores. HerbiTraits provides a tool for the analysis of herbivore functional diversity both past and present and its effects on Earth's ecosystems.