The Niche, Biogeography and Species Interactions
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Downloaded from rstb.royalsocietypublishing.org on July 18, 2011 The niche, biogeography and species interactions John J. Wiens Phil. Trans. R. Soc. B 2011 366, 2336-2350 doi: 10.1098/rstb.2011.0059 References This article cites 85 articles, 14 of which can be accessed free http://rstb.royalsocietypublishing.org/content/366/1576/2336.full.html#ref-list-1 Subject collections Articles on similar topics can be found in the following collections ecology (2529 articles) evolution (2816 articles) Receive free email alerts when new articles cite this article - sign up in the box at the top Email alerting service right-hand corner of the article or click here To subscribe to Phil. Trans. R. Soc. B go to: http://rstb.royalsocietypublishing.org/subscriptions This journal is © 2011 The Royal Society Downloaded from rstb.royalsocietypublishing.org on July 18, 2011 Phil. Trans. R. Soc. B (2011) 366, 2336–2350 doi:10.1098/rstb.2011.0059 Review The niche, biogeography and species interactions John J. Wiens* Department of Ecology and Evolution, Stony Brook University, Stony Brook, NY 11794-5245, USA In this paper, I review the relevance of the niche to biogeography, and what biogeography may tell us about the niche. The niche is defined as the combination of abiotic and biotic conditions where a species can persist. I argue that most biogeographic patterns are created by niche differences over space, and that even ‘geographic barriers’ must have an ecological basis. However, we know little about specific ecological factors underlying most biogeographic patterns. Some evidence supports the importance of abiotic factors, whereas few examples exist of large-scale patterns created by biotic interactions. I also show how incorporating biogeography may offer new perspectives on resource-related niches and species interactions. Several examples demonstrate that even after a major evolutionary radiation within a region, the region can still be invaded by ecologically similar species from another clade, countering the long-standing idea that communities and regions are generally ‘saturated’ with species. I also describe the somewhat paradoxical situation where compe- tition seems to limit trait evolution in a group, but does not prevent co-occurrence of species with similar values for that trait (called here the ‘competition–divergence–co-occurrence conundrum’). In general, the interface of biogeography and ecology could be a major area for research in both fields. Keywords: biogeography; competition; ecology; evolution; niche 1. INTRODUCTION focusing on the environmental conditions determining Ecology and biogeography are two fields with many the large-scale distribution of species (i.e. a Grinnellian obvious connections, but the relationship between niche concept; [7]). I consider these to represent equally them has often been troubled [1]. On the one hand, valid conceptualizations of different aspects of the general those studying large-scale historical (i.e. phylogenetic) Hutchinsonian niche concept [6], and they clearly have biogeography have tended to ignore ecology entirely, many important intersections (e.g. local-scale biotic with little or no consideration of the idea that distri- interactions may set range limits). butions of organisms are influenced by factors such as In this paper, I will review what the niche can tell us habitat tolerances or species interactions. This is well- about biogeography and what biogeography can tell us illustrated in the excellent and integrative textbook by about the niche. I have previously written about the Lomolino et al.[2], in which ecology goes largely gulf between biogeography and ecology [1] and how unmentioned in the chapter on historical biogeography. the niche and biogeography may be related [8]. Those On the other hand, ecologists have often tended to papers focused largely on climatic niches, with relatively ignore biogeography (but with some important excep- little emphasis on the potential role of biotic factors and tions; [3,4]). For example, ecologists often do not species interactions in creating biogeographic patterns. study how large-scale biogeographic history influences Here, I will first focus on the importance of the niche the patterns of diversity and community structure that to biogeographic patterns, and the factors that set the they study, nor how ecological processes create biogeo- range limits of species and clades (i.e. the Grinnellian graphic patterns (review in [1]). niche). I will then focus on what the combination of One place where these two fields should come biogeography, phylogeny and resource-related traits together (but often fail to) is in the concept of the can tell us about niches, particularly related to species niche. Based on Hutchinson’s [5] general concept, the interactions and the Eltonian niche. Note that here niche describes the set of abiotic and biotic conditions and throughout, I refer to biogeographic patterns as where a species can persist [6]. Many ecologists favour those at the scale of species range sizes and larger a concept of the niche based on resources and species (e.g. within continents and between continents). interactions at the local scale (i.e. an Eltonian niche concept; [7]). On the other hand, more biogeo- graphically oriented ecologists often prefer a concept 2. WHY THE NICHE MATTERS FOR BIOGEOGRAPHY (a) Range limits create biogeographic patterns *[email protected] and ecological niches create range limits One contribution of 10 to a Theme Issue ‘Biogeography and Biogeographic patterns arise primarily through limits ecology: two views of one world’. on dispersal. By ‘dispersal’, I mean the process by 2336 This journal is q 2011 The Royal Society Downloaded from rstb.royalsocietypublishing.org on July 18, 2011 Review. Biogeography, ecology and the niche J. J. Wiens 2337 which species expand their ranges, including both the been addressed in some recent studies (e.g. [12,13]) movement of individuals to a new location and their using data from distributions of extant species. For successful establishment there. If there were no limits example, Paul et al.[13] quantified for each species on dispersal, every species could potentially be distrib- the ‘potential range’ (based on species-distribution uted everywhere, and spatial patterns of distribution modelling (SDM) with climatic data), actual range and diversity would be absent or due entirely to extent (how much of the span of the range is occupied) chance. But of course, we know that there are indeed and actual range occupancy (how many potentially large-scale patterns of diversity and distribution suitable pixels are occupied), and then tested how (e.g. major biogeographic provinces, more species these variables were related to species age in a genus richness in the tropics [2,9]). of Neotropical trees. They found that younger species Given this argument, the niche matters for biogeog- generally occupy less of their potential range extents raphy because the range limits of species (and clades) than older species, suggesting that species ranges (or are set primarily by ecological factors (but see below). parts of their ranges) may sometimes be determined By definition, species generally cannot spread outside by limited time for dispersal, rather than ecological of their ecological niche, the set of conditions where limits on dispersal. In contrast, Schurr et al.[12] they can persist (potentially including both abiotic and found no evidence that species range sizes increased biotic factors). Therefore, most so-called ‘geographic with their inferred age for South African Proteaceae. barriers’ and ‘physical barriers’ (e.g. oceans, rivers and Furthermore, some organisms may be able to dis- mountains) are simply areas of unsuitable habitat, and perse across barriers of unsuitable habitat to colonize are entirely organism-specific (e.g. [1]). For example, non-contiguous regions of suitable habitat (e.g. rafting oceans limit dispersal for most terrestrial and freshwater to islands). For these species, range limits may be deter- species, but not necessarily for marine organisms. mined by the combination of habitat differences over Further, many species occur in both freshwater and space and the ability of the organisms to disperse marine environments, depending on the stage in across unsuitable conditions, rather than the unsuitable their life cycle. In terrestrial environments, a river may conditions alone. be a barrier for some organisms, but not others (e.g. especially not for organisms whose niche includes freshwater habitats). Mountains may limit dispersal in (c) Which ecological factors can set range limits? some cases, but mountains are generally only barriers A variety of ecological factors may set the range limits of when they create zones of unsuitable habitat for the species to create biogeographic patterns. These factors organism in question. Many other barriers to dispersal are often categorized as being abiotic versus biotic are more subtle and more clearly related to habitat [2,14]. For terrestrial organisms, abiotic factors that differences (mesic versus arid environments, forest may set range limits include climate (temperature, versus open habitat and rivers versus streams). The precipitation), pH and unsuitable aquatic habitats (for arguments above are neither new nor surprising, but a thorough review of these and other factors see [2]). the idea that ecology influences biogeography remains For aquatic organisms, abiotic factors include tempera- surprisingly rare in both