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2019 the Association for Tropical Biology Wileyonlinelibrary.Com/Journal/Btp Biotropica Received: 26 September 2018 | Revised: 20 December 2018 | Accepted: 27 December 2018 DOI: 10.1111/btp.12632 ORIGINAL ARTICLE The spatial configuration of taxonomic biodiversity along a tropical elevational gradient: α- , β- , and γ- partitions Michael R. Willig | Steven J. Presley Center for Environmental Sciences and Engineering & Department of Ecology Abstract and Evolutionary Biology, University of Biodiversity at larger spatial scales (γ) can be driven by within- site partitions (α), with Connecticut, Storrs, Connecticut little variation in composition among locations, or can be driven by among- site parti- Correspondence tions (β) that signal the importance of spatial heterogeneity. For tropical elevational Michael R. Willig, Center for Environmental Sciences and Engineering & Department of gradients, we determined the (a) extent to which variation in γ is driven by α- or β- Ecology and Evolutionary Biology, University partitions; (b) elevational form of the relationship for each partition; and (c) extent to of Connecticut, Storrs, CT. Email: [email protected] which elevational gradients are molded by zonation in vegetation or by gradual vari- ation in climatic or abiotic characteristics. We sampled terrestrial gastropods along Funding information National Science Foundation, Grant/Award two transects in the Luquillo Mountains. One passed through multiple vegetation Number: DEB-0218039, DEB-0620910, zones (tabonuco, palo colorado, and elfin forests), and one passed through only palm DEB-1239764 and DEB-1546686; to the Institute of Tropical Ecosystem Studies, forest. We quantified variation in hierarchical partitions (α, β, and γ) of species rich- the University of Puerto Rico, and the ness, evenness, diversity, and dominance, as well as in the content and quality of lit- International Institute of Tropical Forestry as part of the Long-Term Ecological Research ter. Total gastropod abundance linearly decreased with increasing elevation along Program in the Luquillo Experimental both transects, but was consistently higher in palm than in other forest types. The Forest; USDA Forest Service; Center for Environmental Sciences and Engineering at gradual linear decline in γ- richness was a consequence of opposing patterns with re- the University of Connecticut gard to α- richness (monotonic decrease) and β- richness (monotonic increase). For evenness, diversity, and dominance, α- partitions and γ- partitions evinced mid- elevational peaks. The spatial organization of gastropod biodiversity did not mirror the zonation of vegetation. Rather, it was molded by: (a) elevational variation in pro- ductivity or nutrient characteristics, (b) the interspersion of palm forest within other forest types, and (c) the cloud condensation point acting as a transition between low and high elevation faunas. Abstract in Spanish is available with online material. KEYWORDS diversity, dominance, evenness, Gastropoda, hierarchical partitioning of diversity, Puerto Rico, richness, tropical montane forest 1 | INTRODUCTION varies with respect to environmental or geographic gradients have attracted much attention (e.g., Gering, Crist, & Veech, 2003; Higgins, Understanding the spatial organization of biodiversity is necessary 2010; Summerville, Boulware, Veech, & Crist, 2003; Wagner, Wildi, for determining the scales at which mechanisms operate to generate & Ewald, 2000; Wilsey, 2010), with a recent surge in attention to variation in the identity and abundances of species. Consequently, β- diversity (Mori, Isbell, & Seidel, 2018) because of its potential the manners in which the hierarchical configuration of biodiversity relationship with multifunctionality of ecosystems. Importantly, 104 | © 2019 The Association for Tropical Biology wileyonlinelibrary.com/journal/btp Biotropica. 2019;51:104–116. and Conservation WILLIG AND PRESLEY | 105 patterns of biodiversity are often scale- dependent (e.g., Jackson & Elevational gradients are useful for assessing environmental Fahrig, 2014; Scheiner et al., 2000), highlighting the role of spatial characteristics to which species respond because changes in eleva- compartmentalization in heterogeneous landscapes. Biodiversity tion result in predictable and appreciable variation in plant species at larger spatial scales (γ) can be driven by within- site diversity (α- composition and structure, as well as in abiotic factors (e.g., tem- partitions), if little compositional variation characterizes communi- perature, precipitation, N- content, or N to C ratio). Although eleva- ties (Cornell, Karlson, & Hughes, 2007; Gering & Crist, 2002; Pegg tional changes in abiotic characteristics and associated vegetation & Taylor, 2007), or biodiversity can be driven by among- site varia- are predictable, they differ in the form of their variation. Climatic tion (β- partitions) that signal the importance of spatial heterogene- characteristics, as well as abiotic factors associated with productiv- ity at landscape or regional spatial scales (Belmaker, Ziv, Shashar, & ity (e.g., N- content or N to C ratio), generally change in a gradual, but Connolly, 2008; Erős, 2007; Stendera & Johnson, 2005). The contri- not necessarily linear fashion with elevation (Waide, Zimmerman, & butions of α- or β- partitions to γ are largely dependent on the nature Scatena, 1998; Whiteman, 2000), whereas floral associations often of environmental variation within a domain of interest (Freestone & have more or less discrete boundaries recognized as habitat types, Inouye, 2006) and the ways in which different species respond to forest types, or life zones (e.g., Hemp, 2006; Kessler, 2000). Because spatial variation in the environment. habitat specialization and responses to abiotic characteristics are important in defining the ranges of animal species, the spatial or- ganization of biodiversity is contingent on the dominant mechanism 1.1 | Elevational gradients that molds animal species distributions. The differences in the form Montane environments provide powerful test beds for the examina- of elevational variation in abiotic factors and vegetation offer an op- tion of patterns and processes that mold spatial patterns of biodi- portunity to disentangle the effects of these potential structuring versity (e.g., Cisneros et al., 2014; González, Willig, & Waide, 2013; mechanisms. If habitat preferences or specializations determine hi- McCain, 2009; Terborgh, 1971; Whittaker, 1956; Willig & Presley, erarchical structure of biodiversity along elevational gradients, then 2016). Dramatic variation in environmental drivers (e.g., solar in- communities within a vegetation type should exhibit low among- site solation, temperature, precipitation) occurs over relatively short variation (i.e., low β- diversity), with high β- diversity among vege- geographic extents, such that ecological mechanisms, rather than tation types. In contrast, if the spatial structure of biodiversity is biogeographic or historic mechanisms, likely mold biotic responses. driven by species- specific tolerances to abiotic characteristics that This contrasts greatly with latitudinal gradients of biodiversity, in change gradually with elevation, then β- diversity among elevations which considerably greater geographic distances are necessary to within a vegetation type should be no different than β- diversity produce comparable variation in environmental drivers, making it among vegetation types. more difficult to disentangle ecological from biogeographic or his- toric mechanisms (Willig, Kaufman, & Stevens, 2003; Willig & Presley, 1.3 | Objectives and hypotheses 2013). Elevational gradients of biodiversity have received renewed attention because predicted changes in climate promise to manifest We explore the extent to which environmental characteristics first and most strongly at high latitudes and elevations, especially mold patterns of taxonomic biodiversity of terrestrial gastropods in tropical environs (Colwell, Brehm, Cardelús, Gilman, & Longino, along a tropical elevational gradient by quantifying the hierarchical 2008; Dirnbock, Essi, & Rabitsch, 2011; González et al., 2013; Malhi compartmentalization (α- , β- , and γ- partitions) of species richness, et al., 2010). This may be particularly true to the extent that tropical evenness, diversity, and dominance, along two parallel transects, species have evolved in environments with less intra- annual variabil- one passing through three elevationally stratified forest types (ta- ity in climatic conditions, leading to narrow niche breadths than in bonuco, palo colorado, and elfin) and one passing only through palm their extratropical counterparts (Janzen, 1967). forest. In addition, we determine the form and parameterization of elevational gradients of each hierarchical partition along each tran- sect. Finally, we evaluate the effects of elevation and transect on 1.2 | Gastropods in the Luquillo Mountains litter nutrient content, which may represent an alternative driver Terrestrial gastropods are taxonomically diverse and numerically of spatial structure of biodiversity for terrestrial gastropods along abundant heterotrophs that fill important roles as detritivores, elevational gradients. herbivores, and carnivores, making them useful for biogeographic, If climatic factors (e.g., temperature or precipitation) or abiotic macroecological, or conservation study (e.g., Bloch, Higgins, & factors (e.g., N- content or N to C ratio) that are related to produc- Willig, 2007; Clements et al., 2008; Stanisic, Cameron, Pokryszko, & tivity primarily
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