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Mammalia: Carnivora) in the Americas: Past to Present

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Mammalia: Carnivora) in the Americas: Past to Present Journal of Mammalian Evolution https://doi.org/10.1007/s10914-020-09496-8 ORIGINAL PAPER Environmental Drivers and Distribution Patterns of Carnivoran Assemblages (Mammalia: Carnivora) in the Americas: Past to Present Andrés Arias-Alzate1,2 & José F. González-Maya3 & Joaquín Arroyo-Cabrales4 & Rodrigo A. Medellín5 & Enrique Martínez-Meyer2 # Springer Science+Business Media, LLC, part of Springer Nature 2020 Abstract Understanding species distributions and the variation of assemblage structure in time and space are fundamental goals of biogeography and ecology. Here, we use an ecological niche modeling and macroecological approach in order to assess whether constraints patterns in carnivoran richness and composition structures in replicated assemblages through time and space should reflect environmental filtering through ecological niche constraints from the Last Inter-glacial (LIG), Last Glacial Maximum (LGM) to the present (C) time. Our results suggest a diverse distribution of carnivoran co-occurrence patterns at the continental scale as a result of spatial climatic variation as an important driver constrained by the ecological niches of the species. This influence was an important factor restructuring assemblages (more directly on richness than composition patterns) not only at the continental level, but also from regional and local scales and this influence was geographically different throughout the space in the continent. These climatic restrictions and disruption of the niche during the environmental changes at the LIG-LGM-C transition show a considerable shift in assemblage richness and composition across the Americas, which suggests an environ- mental filtering mainly during the LGM, explaining between 30 and 75% of these variations through space and time, with more accentuated changes in North than South America. LGM was likely to be critical in species functional adaptation and distribution and therefore on assemblage structuring and rearranging from continental to local scales through time in the continent. Still, extinction processes are the result of many interacting factors, where climate is just one part of the picture. Keywords Ecological niche . Carnivoran . Extinction . Climate change . Late Pleistocene . Paleodistributions Electronic supplementary material The online version of this article Introduction (https://doi.org/10.1007/s10914-020-09496-8) contains supplementary material, which is available to authorized users. Understanding how species are distributed, their determinants * Andrés Arias-Alzate and constraints, and how they are spatially structured in as- [email protected] semblages through time and space is a fundamental goal of macroecology and ecology (Brown et al. 1995;Ferrazetal. 2012; Agosta and Bernardo 2013). Species can vary in size (in 1 Facultad de Ciencias y Biotecnología, Universidad CES, Calle 10A # 22-04, Medellín, Colombia mammals, which span 12 orders of magnitude), location, shape, and occupancy (Gaston 2003;Daviesetal.2009)and 2 Instituto de Biología, Universidad Nacional Autónoma de México, Circuito exterior s/n, Ciudad Universitaria, Coyoacán, are the result of responses to ecological rules such as climatic CP04510 México City, Mexico conditions, species dispersal abilities, historical events, phylo- 3 Proyecto de Conservación de Aguas y Tierras, ProCAT Colombia/ genetic inertia, and/or biotic interactions (Dynesius and Internacional, Carrera 11 # 96-43, Of. 303, Bogotá, Colombia Jansson 2000; Blomberg and Garland 2002; Steinitz et al. 4 Laboratorio de Arqueozoología “Ticul Álvarez Solórzano”, 2006; Davies et al. 2009; Blois et al. 2013, 2014). Subdirección de Laboratorios y Apoyo Académico, Instituto Understanding the main drivers of changes in such parameters Nacional de Antropología e Historia, Moneda # 16, Col. Centro, can also shed light in terms of local and global extinctions. An 06006 México City, Mexico important issue is whether these processes such as climatic 5 Instituto de Ecología, Universidad Nacional Autónoma de México, conditions have the same effect to structure assemblage pat- Circuito exterior s/n, Ciudad Universitaria, Coyoacán, terns and species distributions at different spatial and temporal CP04510 México City, Mexico J Mammal Evol scales (i.e., from continental to regional and/or to local scales) implications in the evolutionary patterns that took place con- (Arias-Alzate et al. 2017). However, accurate information on tinentally, thanks to its high taxonomic diversity and well- past and present distributional patterns for many species is resolved phylogeny (Goswami and Friscia 2010;Nyakatura often scarce at broad scales, and the mechanisms, determi- and Bininda-Emonds 2012). Here, we used an ecological nants, and constraints of these ranges at these scales are still niche modeling and macroecological approach in order to as- poorly understood (Graham 2001;Prevostietal.2005; sess the influence of past climate change on carnivoran assem- Martínez-Meyer et al. 2004; Davies et al. 2009; Nogués- blage richness and composition patterns from continental to Bravo 2009). Nevertheless, the understanding of the relation- local levels over the last 130 K years in the Americas and if ship between the influence of environmental drivers and as- this influence reflects environmental filtering through ecolog- semblages measures (i.e., richness and composition) have re- ical niche constraints. It is important to note that relatively ceived more attention, especially when extreme climatic little attention has been given to understanding the underlying events (e.g., glaciations) can change the outcome of the eco- causes of non-random patterns such as geographical distribu- logical patterns and processes (Thibault and Brown 2008; tion and assemblage structuring (Collins et al. 2011;Gotelli Davies et al. 2011). and Ulrich 2012; Blois et al. 2014); therefore, revealing these These patterns are largely governed by environmental fac- patterns and some of their mechanisms would improve our tors (e.g., climatic conditions) that define part of the funda- knowledge on the macroecological history of carnivoran as- mental niche of the species, where the biotic interactions are semblages in the Americas. usually less perceptible (Martínez-Meyer et al. 2004;Soberón and Nakamura 2009; Lorenzen et al. 2011; Peterson et al. 2011;Levinskyetal.2013). However, elucidating the influ- Materials and Methods ence of these processes on species coexistence and distribu- tion is not straightforward because non-random species asso- Species Geographical Distribution (GD) Patterns Data ciations are not necessarily caused only by climatic factors nor species interactions. All these processes can operate indepen- Species GD patterns came from an ecological niche modeling dently or in synergy to determine assemblages patterns (ENM) approach done by Arias-Alzate (2016) and Arias- through time and space (Martínez-Meyer et al. 2004;Blois Alzate et al. (2017), which were estimated as described below. et al. 2014; Giarla and Jansa 2015). Therefore, the central The species studied were based on the assumption that all issue is how to differentiate or elucidate the influence pro- extant species were also present during the late Pleistocene, and duced by biotic interactions or dispersal limitations (either the extinct species disappeared at the end of the Pleistocene- by barriers or by movement capacity) from those produced early Holocene approximately 12Kyr (Webb 2006; Davies by environmental filtering (Dynesius and Jansson 2000; et al. 2009). The species criteria for that study included taxo- Svenning and Skov 2007; Blois et al. 2014; Giarla and Jansa nomic validity, proper chronological dating, and reported in- 2015). In this sense, the ecological niche played an important formation about the species’ presence during the study period. role to disentangle the combined effects of abiotic (e.g., cli- The list and the taxonomy for living species were based on matic) and biotic factors and could help us to understand in- Wilson and Reeder (2005), Wilson and Mittermeier (2009), directly about the effects of others historical factors such as and IUCN Red List of Threatened Species (version 2018.2.). barriers and accessible areas as strong forces on assemblage’s For extinct species, the taxonomy was based on the fossil re- dynamics and coexistence at different scales (Davis and Shaw cords and following Berta (1985),Barnettetal.(2005), 2001; Martínez-Meyer et al. 2004; Bofarull et al. 2008; Cisneros (2005), Cione et al. (2007), Prevosti and Rincón Peterson et al. 2011;Levinskyetal.2013; Soares 2013). (2007), Soibelzon and Prevosti (2008), Arroyo-Cabrales et al. Climate is an important determinant of species ecological (2010), and Ferrusquía-Villafranca et al. (2010). niche and its geographic expression, thus, helping to under- The study area was defined as the entire American conti- stand extinction processes through range shifts and bottleneck nent; this represents the area of accessibility M sensu Peterson events (Martínez-Meyer et al. 2004;Nogués-Bravoetal. et al. (2011), as all historical and ecological processes of colo- 2008; Arias-Alzate 2016). However, extinction processes are nization and dispersal of the species of interest occurred in the the result of many interacting factors (e.g., humans) that affect continent (during and after the Great American Biotic species differentially and could act differentially or asynchro- Interchange; GABI). Namely, during the GABI in several in- nously with additive
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