Distribution of abundance across the range in eastern North American trees Helen T Murphy1, Jeremy VanDerWal and Jon Lovett-Doust 1 Author for correspondence Department of Biological Sciences University of Windsor Windsor Ontario N9B 3P4 Canada Phone: 1-519-253 3000 ext 2722 Fax: 1-519-971 3609 Email: [email protected] Running title: Macroecology of eastern North American trees. 1 ABSTRACT Aim We analyzed spatial datasets of abundance across the entire, or near entirety of the geographic ranges of 134 tree species to test macroecological hypotheses concerning the distribution of abundance across geographic ranges. Location Our abundance estimates come via the USDA Forest Service Forest Inventory and Analysis Eastwide Database, which contains data for 134 eastern North American tree species. Methods We extracted measures of range size and the spatial location of abundance relative to position in the range for each species to test four hypotheses: (a) species occur in low abundance throughout most of their geographic range; (b) there is a positive interspecific relationship between abundance and range size; (c) species are more abundant in the centre of their range; and (d) there is a bimodal distribution of spatial autocorrelation in abundance across a species range. Results Our results demonstrate that (a) most species (85%) are abundant somewhere in their geographic range; (b) species achieving relatively high abundance tend to have larger range sizes; (c) the widely held assumption that species exhibit an ‘abundant-centre distribution’ is not well supported for the majority of species; we suggest ‘abundant-core’ as a more suitable term; and (d) there is no evidence of a bimodal distribution of spatial autocorrelation in abundance. Main Conclusions For many tree species, high abundance can be achieved at any position in the range, though suitable sites are found with less frequency toward range edges. Competitive relationships may be involved in the distribution of abundance across tree ranges and species with larger ranges (and possibly broader niches) may be affected more by biotic interactions than smaller ranging species. 2 Keywords abundance, distribution, range, abundant-centre, macroecology, spatial autocorrelation, occupancy 3 INTRODUCTION Increased understanding of landscape- and regional-scale processes in the distribution and abundance of organisms has led to the relatively recent emergence of macroecology (Gaston & Blackburn, 2000), and an awareness of patterns and processes that are otherwise indistinguishable at smaller scales. Macroecology involves the ecology of “wide expanses of space, long periods of time and large numbers of taxa” (Blackburn & Gaston, 2003), and addresses patterns of distribution and abundance of species at geographical spatial scales and evolutionary time scales (Brown, 1995). Thus complex relationships between variables, such as species range size and individual abundances, and location in the range and abundance, underpin most macroecological theories. Whereas distribution and abundance have been the major emphases of investigation thus far, here we focus on a different perspective: the distribution of abundances across geographic ranges. Tree-dominated communities are among the richest in ecological structure and biotic diversity, in both above- and below-ground aspects. How large numbers of competing species manage to coexist remains a major unresolved problem in community ecology and it is particularly acute for plants, given that most plants require remarkably similar resources and acquire them in similar ways (Silvertown, 2004). Here we use available spatial datasets of abundance (in the form of phytosociological ‘importance values’) and range boundaries for each of 134 eastern North American tree species (representing 29 taxonomic families) (see Table S1 in Supplementary Material) to test four major macroecological hypotheses: (a) most species occur in low abundance throughout most of their geographic range; (b) there is a positive interspecific relationship between abundance and range size; (c) species 4 are more abundant in the centre of their range; and (d) there is a bimodal distribution of spatial autocorrelation in abundance across a species range. We discuss the results in terms of problems of plant coexistence and the role of both abiotic and biotic factors in regulating species distribution and abundance. When abundance distributions have been examined over entire geographic ranges (or nearly entire ranges) it has generally been observed that species exist in low abundance at most sites and at high abundance in only a few sites (Brown et al., 1995). Furthermore, Murray & Lepschi (2004) showed that the majority of species that are locally rare in open forest communities in Australia were abundant at other locations within their geographic range (existing as ‘somewhere abundant’ species); thus sampling at any given point in a landscape should capture most species at low abundance and a few species in high abundance. The tendency for individual species to show positive relationships between abundance and range size (or occupancy) has been widely reported for a variety of taxa and appears to be remarkably robust over multiple spatial scales and across a variety of measures (Gaston et al., 2000). At least eight major mechanisms have been described, both biological and artifactual, that may account for positive interspecific abundance-range size relationships (Gaston et al., 1997). These include sampling artifacts and phylogenetic non-independence, individual species attributes and population dynamics. Studies of interspecific abundance-occupancy relationships in plants, although generally supporting the positive relationship have tended to focus on narrowly defined habitat types, or on abundance in only a small proportion of the species’ range (Gotelli & Simberloff, 1987; Collins & Glenn, 1990). 5 The ‘abundant–centre distribution’ describes the widely held assumption in biogeography and macroecology that species abundances tend to be greater toward the centre of the geographical range and lower in the periphery (Brown, 1984; Sagarin & Gaines, 2002). In an influential paper Brown (1984) argued that local abundance reflects how well a particular site meets a species’ physiological and ecological requirements. Brown suggested that spatial autocorrelation in these axes (representing dominant dimensions of the niche) could underpin the abundant-centre distribution. Increasing the distance from the optimal site should decrease the probability of a site meeting the multidimensional needs of that species (Brown, 1984). Thus there are three main predictions of the abundant-centre distribution: (1) abundance should be higher in sites closer to the centre of the range; (2) species should occupy more sites towards the centre of the range; and (3) abundance and occupancy should decline linearly towards the edge of the range. Brown et al. (1995) reported that abundance exhibited a distinctive bimodal pattern of spatial variation within the geographic ranges of four passerine birds. Spatial autocorrelation analysis produced two peaks, one at short distances (corresponding to proximal sites within the geographic range), and one at the maximum distance (corresponding to sites at opposite ends of the geographic range). Thus sites close together were more likely to have similar abundances irrespective of where they were located in the range, and sites at the range periphery tended to have consistently low abundance, producing the second peak at maximum distances. This second peak in spatial autocorrelation at maximum distances implies that species respond similarly to all aspects of the range edge. 6 Critical examination of macroecological theories in the past has suffered from a paucity of good quality datasets (Sagarin & Gaines, 2002; Mathius et al., 2004) and analyses have often been based on inadequate range size information, restricted spatial coverage of abundance data, and/or limited taxonomic extent (Sagarin & Gaines, 2002; Kotze, 2003; Blackburn et al., 2004). Fortunately, a growing availability of large scale geographic information system (GIS) datasets and sophisticated tools for spatial analysis has enabled patterns of distribution of abundance within species’ ranges to be examined in more detail. The spatial data on both abundance and distribution which we use here has been collected and compiled in a consistent way for all species in the dataset. Furthermore, the complete distribution of species’ abundances within the geographic range is known for most species. Thus many of the problems and biases relating to limited, inadequate or unreliable data previously identified in macroecology studies are avoided (see e.g., Sagarin & Gaines, 2002; Blackburn et al., 2004). However, our use of importance values rather then absolute abundance does have some limitations which are discussed further below. METHODS Abundance estimates (phytosociological importance values) were available at the Eastwide Database from U.S.D.A. Forest Service Forest Inventory and Analysis data (Prasad & Iverson, 2003), and comprise >100,000 plots and records for c. three million trees in thirty-seven states. Importance values are calculated for each species as: 7 IV(x) = 50*BA(x) / BA(all species) + 50*NS(x) / NS(all species) where x is a particular species at a plot, BA is basal area, and NS is number of stems (summed for overstory and understory individuals). In monotypic stands, the IV would reach the