Population Diversity and Ecosystem Services

Opinion TRENDS in Ecology and Evolution Vol.18 No.7 July 2003 331 Population diversity and ecosystem services Gary W. Luck1, Gretchen C. Daily2 and Paul R. Ehrlich2 1The Johnstone Centre, Charles Sturt University, PO Box 789, Albury, NSW 2640, Australia 2Center for Conservation Biology, Stanford University, 371 Serra Mall, Stanford, CA 94305-5020, USA The current rate of biodiversity loss threatens to disrupt Population change can occur through variation in the greatly the functioning of ecosystems, with potentially number of populations for a given species, the number of significant consequences for humanity. The magnitude individuals per population, the spatial distribution of of the loss is generally measured with the use of species populations, and the genetic differentiation within and extinction rates, an approach that understates the among populations. Historically, the term ‘population severity of the problem and masks some of its most diversity’ has largely been used by geneticists when important consequences. Here, we propose a major discussing genetic differentiation (e.g., [14–16]), or as a expansion of this focus to include population diversity: measure of the number of Mendelian populations in a considering changes in the size, number, distribution given area [6]. These approaches are limited because and genetic composition of populations and the impli- they ignore important demographic characteristics of cations of those changes for the functioning of ecosys- populations that can influence the provision of ecosystem tems and the provision of ecosystem services. We also services. We argue that assessments of population outline the key components of population diversity and diversity should consider both the demographic describe a new approach to delineating a population (e.g. size, number and distribution) and genetic nature unit that explicitly links it to the services that it provides of populations. Here, we have two aims: (1) to develop a new population Species loss dramatically represents the mass extinction unit categorization that explicitly links a population with currently underway [1–4]. Yet, species extinctions are an the ecosystem service(s) that it provides; and (2) to provide inadequate measure of biodiversity loss and do not provide an extended definition of population diversity that information about changes in the capacity of particular formalizes its most important dimensions in an ecosystem species to contribute to the functioning of ecosystems. context. We present a conceptual framework for exploring Balmford et al. (this issue) provide persuasive arguments the relationships between population diversity and the for focusing on changes in population size and habitat functioning of ecosystems. Investigating these relation- extent when measuring the state of nature. We also ships empirically in even a few systems presents major emphasize that the relationship between biodiversity and challenges, but we hope our discussion stimulates debate human well being is primarily a function of populations of and innovative ideas about empirical investigations of species. Whether the benefits of biodiversity are conveyed what we believe is an area of crucial importance. The directly (e.g. food) or indirectly (e.g. pollination), their terms ‘ecosystem service’ and ‘ecosystem function’ are supply is generally determined by the diversity of largely interchangeable, as used here, although ecosystem populations producing them [5]. Hence, population change services can be defined as ecological processes that benefit can have a substantial impact on an ecosystem that is people [9], whereas ecosystem functions can be considered independent of changes in species diversity. as all ecological processes regardless of whether they are The consequences of population loss for species con- beneficial to humanity. servation are well recognized, but have been little addressed from the viewpoint of the functioning of ecosystems and the Defining population units and the importance of provision of ecosystem services [6–9]. Recent research population change has begun to explore the link between species popu- There are many approaches to defining species populations and the services that they provide [10–12], even to lations [17–22]. Consistently problematic is defining the point where species-specific services have been population boundaries so that the number of populations identified [13]. Here, we argue that this approach should can be clearly determined [6]. Geneticists use measures of be greatly expanded for two reasons: (1) to assess the gene flow and genetic differentiation to distinguish one theoretical and practical implications of population population from another [23]. In a demographic sense, this change for the functioning of humanity’s life-support can be achieved by careful measures of individual move- systems; and (2) to reflect more accurately the state of ment (e.g. [24]), which enables the delineation of popu- global biodiversity. lations that are sufficiently isolated from each other to have independent dynamics [25] (Box 1). Populations can Corresponding author: Gary W. Luck ([email protected]). also be distinguished with the use of some arbitrarily http://tree.trends.com 0169-5347/03/$ - see front matter q 2003 Elsevier Science Ltd. All rights reserved. doi:10.1016/S0169-5347(03)00100-9 332 Opinion TRENDS in Ecology and Evolution Vol.18 No.7 July 2003 defined spatial and/or temporal context (e.g. linear Imagine a single species occupying a heterogeneous distance between groups, or the presence of geographical environment such that groups of individuals can be barriers or other spatial disjunctions) or differences in distinguished on the basis of habitat suitability or the phenology, morphology or physiology. spatial characteristics of the landscape (e.g. a rainforest- The above distinctions are important from the perspec- dependent species occupying forest fragments in an tive of species conservation, but they might be inadequate agricultural landscape). The groups might be linked by when assessing the contribution that a group of individ- sufficient dispersal to classify them as belonging to a single uals from a given species makes to an ecosystem service. To evolutionary unit (EU) or demographic unit (DU; Box 1). address this issue, we suggest that it is essential to However, each group provides nonoverlapping, localized recognize the link between a group of individuals and the services to areas of the surrounding landscape, suggesting ecosystem service(s) that it provides, and use this as a that they could be categorized as different SPUs. For further criterion to delineate populations. We refer to this example, Kremen et al. [12] demonstrate that pollination new population categorization as a ‘service-providing unit’ services provided by native bee populations to watermelon (SPU; Boxes 1,2). crops are reduced on farms that are distant from native Box 1. Taxonomy of populations Evolutionary unit Evolutionary units (EUs) are populations with independent evolution- (a) EU ary dynamics. Thus, a classic Mendelian population (a reproductive community of sexual and cross-fertilizing individuals which share a common gene pool) [40] is an EU. Using neutral loci will generally circumscribe larger EUs than would using loci under strong, geo- graphically varying selection. An example of EUs is the island populations of various plant species (from the Family Asteraceae) that evolved different diaspore morphology and reduced dispersal ability, compared with the mainland populations from which they originated [41]. SPU DU Demographic unit Demographic units (DUs) are populations with independent demographic dynamics. In general, populations that fluctuate in size asynchronously, or are shown to have only a few (or no) migrants pass between them, should be considered DUs. A classic example of CU a set of DUs is the now-extinct three populations of the checkerspot butterfly Euphydryas editha in the Jasper Ridge Biological Reserve of Stanford University. They had asynchronous dynamics [42,43] and delineating the DUs was crucial to understanding those dynamics. Local populations in metapopulations are ordinarily DUs, but DUs are not necessarily elements of metapopulations. (b) SPU Conservation unit The designation of conservation units (CUs) will depend on the associated conservation goals, which vary tremendously. Goals oriented around evolution typically involve maintaining the genetic diversity of a species, or the potential for future genetic divergence or speciation. Pursuit of these goals has been formalized using concepts such as minimum viable populations [44–46], evolutionarily significant units [47–49], and so-called DU management units [48]. Here, we focus on the goal of maintaining ecological functions and their associated ecosystem services. Service-providing unit (SPU) CU The new population category that we propose is a service-providing unit (SPU). SPUs provide, or might provide in the future, a recognized ecosystem service at some temporal or spatial scale. All populations are potentially SPUs and the population categories described here can overlap (e.g. a single population might be an TRENDS in Ecology & Evolution EU, DU, CU and SPU). This will not always be the case, making it crucial to delineate SPUs when assessing the consequences of Fig. I. population change for the provision of ecosystem services. For example, population categories might occur in a nested hierarchy comprise a single SPU (Fig. Ib). For example,

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