# Reconciliation ecology and the future of species diversity

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2 Reconciliation ecology 195 fits data from the fossil record. Ecologists are not sure why a power equation fits islands or continents, but we do now have a successful mathematical theory for areas within a province. McGill & Collins (2003) deduced the species-area curve within provinces from four assumptions: Each of these processes produces SPARs with z-values in a restricted range, and the ranges abut, covering virtually the entire unit interval. Three of the puzzles fit together like a triptych. I will review all four SPAR puzzles and the processes that produce them. \$ The geographical range of each species is independently located with respect to all others. Sample-size SPARs \$ Species vary in abundance with respect to each other. Determining the number of species in an area requires \$ Species have a minimum abundance. sampling. Sampling comes with a bias, i.e. the larger the \$ Each species abundance varies significantly across number of individuals identified, the greater the number its own range, being relatively scarce more often than of species in the sample. Usually, more individuals will relatively common ( relatively means with respect to be identified from a larger area than a smaller one. A its own average abundance). SPAR generated by statistical sampling artifacts holds Data support all four assumptions. From them, McGill no biological interest. Nonetheless, one must be able to shows that there is a species-area curve and that it recognize such SPARs and eliminate them from further approximates a power equation whose z-value ranges consideration. Fisher et al. (1943) showed us one powerful between 0.05 and 0.25 with a mean of about way to do this, devising a statistic called Fisher s a that McGill s theory improves that of Leitner, which required is almost insensitive to sample size but does vary with knowledge of the relationship between abundance and S. Burnham & Overton (1979) and Lee & Chao (1994) range size (Leitner & Rosenzweig, 1997). McGill s theory have introduced other successful bias reducing statistics. also predicts the abundance-range-size relationship. These can be calculated using free software packages Leitner s theory in turn supplanted that of Preston (1962) (WS2M at and (elaborated and extended by May, 1975). Leitner found EstimateS at two cryptic errors in the Preston-May proofs, errors that Sample-size SPARs tend to have the smallest z-values. All vitiated them. Both McGill s and Leitner s theory show those I have encountered have z 0.12 (Rosenzweig, that the power equation is merely a good fit rather than 1995). an exact description of the species-area curve. That it is a good fit should not be too surprising because power equations are very plastic curves and fit a host of monotonic Sample-area SPAR relationships. All species have habitat requirements that restrict them Other ingenious attempts at a SPAR theory exist, but in space. A larger area will tend to include more habitat all have their problems, problems mostly outside the scope types than a smaller one (Williams, 1943). Thus SPARs of this paper (Wissel & Maier, 1992; Durrett & Levin, emerge from dicerent-size samples within the same 1996; Harte et al., 1999; Hubbell, 2001). Yet I must mention biological region. They tend to have z-values of one frequent problem: the assumption of a single habitat (Rosenzweig, 1995). type for all places. No model can possibly account for a pattern known to be caused by a variable that the model does not use (this is true no matter how well such a Archipelagic SPARs model may fit the data). Yet we certainly know that the The islands of an archipelago combine the habitat-sampling species-area relationship within a province depends on process with another process, which I review below. the inclusion of more habitats in larger areas (Williams, 1964). McGill s theory incorporates habitat variation in its assumption that each species abundance varies significantly across its own range. Their SPARs have z-values of (Rosenzweig, 1995). Interprovincial SPARs Scales of the species-area curve SPAR is actually four puzzles and their pieces had been mixed up together as if they belonged to the same puzzle. Thus, the SPAR puzzle pieces first had to be separated (Rosenzweig, 1995). The key was realizing that a variety of dicerent processes determine S as a function of area. Biogeographical provinces of similar environment, such as wet tropical forests, have diversities in proportion to their areas. A biogeographical province is a region whose species have evolved within it, rather than immigrating from somewhere else. Although the concept is merely an ideal every place has at least a few species that arrived as immigrants it is close to true in many places, such as dicerent continents or well-separated periods in

3 196 M. L. Rosenzweig the history of life. Interprovincial SPARs are much less common than those for archipelagos, but the law they follow is clear. The z-values of interprovincial SPARs begin at 0.6 and range upwards, with most about 0.9 to 1 (Rosenzweig, 1995, 2001). Fig. 1 shows a recently obtained example. Diversity in provinces appears to have been following such a law for hundreds of millions of years (Rosenzweig & Ziv, 1999), and we have no evidence to suggest that this law has been altered. On the contrary, because they appear to be so reliable and because they predict diversity far into the future, interprovincial z-values will turn out to be the most useful to conservation biologists. What makes z-values different? To understand the three SPARs of Fig. 2 (and their dicerences), we need to connect the state variable we call species diversity to its derivatives. Recognizing that, and beginning the journey to accomplish it, was one of the greatest achievements of MacArthur and Wilson (1967), who applied dynamic analysis to the problem of island diversities. They carefully defined the two derivatives that should matter most: the rate at which species not on an island arrive on it, and the rate at which species on an island become extinct there. Then, in search of a selfregulating system, they asked how these rates should vary with diversity itself. Their result was powerful and robust: An island with no species can sucer no extinctions. An island with all Fig. 1 Interprovincial species-area relationship for tropical freshwater fishes. The z-value is Data courtesy Peter Reinthal. Figure from Rosenzweig (2003) with permission. Fig. 2 Three scales of species-area curves taken from bird censuses. The steepest curve connects points from dicerent biogeographical provinces (data on frugivores of wet tropical forests from Fleming et al. (1987) and Rosenzweig (1995)). The least steep curve connects points from dicerent sized samples within the same province (Chilean matorral data from Cody (1975)). The archipelagic curve lies in between (Caribbean data from Wright (1981)). the species of its mainland source pool can experience no further immigrations. In contrast, immigrants must be arriving at some positive rate on an island with no species, and an island with as many species as possible must sucer extinctions at some positive rate. Thus there has to be at least one intermediate diversity at which the two rates neutralize each other, i.e. there has to be at least one steady-state diversity. But MacArthur and Wilson went beyond a demonstration of self-regulating diversity on islands. Their theory also predicts the existence of archipelagic SPARs: larger islands should tend to contain more habitats and, at any particular S, larger populations of species. Those influences would depress the extinction rate curve of a large island compared to a smaller one, and the steady-state should therefore increase with island size. The theory of island biogeography does not predict the shape or the z-value of archipelagic SPARs, but it does predict that their z-values should vary. At any particular diversity, islands farther from the source of colonization should receive immigrant species at a reduced rate compared to closer islands. So two islands at dicerent distances, but of the same area, will have dicerent steady states. The farther island will have the lower S. Connecting the species-area point of this farther island to that of the source area yields a steeper line than connecting the species-area point of the nearer island (with higher S) to the source area s point. But the slope is the z-value, and so an island with the same area but fewer species has a higher z-value than its closer counterpart.

8 Reconciliation ecology 201 area, these two cannot do much by themselves. No conservationist seriously believes that we can reserve much more than the 5% or 10% that now remains, and many will admit that the human population is likely to continue expanding. If we use only reservation ecology and restoration ecology, it would seem that we are doomed to lose nearly every species alive today. Reservation and restoration ecology must be supplemented. Conservation philosophy, science, and practice must be framed against the reality of human-dominated ecosystems, rather than the separation of humanity and nature underlying the modern conservation movement (Western, 2001). Fortunately, some people have begun this work. Reconciliation ecology Today, conservationists and ecologists are pioneering a new area of research that will make long-term diversity conservation possible. I call it reconciliation ecology. Reconciliation ecology discovers how to modify and diversify anthropogenic habitats so that they harbor a wide variety of wild species. In essence, it seeks techniques to give many species back their geographical ranges without taking away ours (Fig. 6). Thus it is trying to grow the earth back, to expand the area available to nature. That will establish a steady-state diversity far greater than would be available if conservation biologists restrict themselves to the areas acorded by set-asides alone. A growing number of examples demonstrate that reconciliation ecology can work (Rosenzweig, 2003). I will describe a few in the rest of the paper. Reconciliation in agricultural sites Because agricultural uses dominate most of the land areas that people have taken for themselves, perhaps the most important cases of reconciliation ecology are those associated with agriculture. Led by Gretchen Daily (Daily et al., 2001), John Vandermeer and Yvette Perfecto (Vandermeer & Perfecto, 1995), and Russell Greenberg (Greenberg et al., 1997), Countryside Biogeography is showing that some styles of land use, especially those of traditional agriculture, are already compatible with the needs of many species. Sometimes the compatibility of diversity and agriculture occurs quite accidently, sometimes it is deliberate. Such compatibility exists in pasturelands, croplands, plantations and timberlands. It comes from rich and poor countries, sponsored by private or governmental agencies. The following examples illustrate the variety. Cardamom Growers maintain many tree species in their cardamom Elettaria cardamomum plantations. They do so to provide shade for the herb and a steady supply of nectar for its pollinators, principally honey bees. Bees visit 37 tree species in the plantations, of which 10 supply both nectar and pollen, three nectar only and the rest pollen only. From May to September, flowers are not very abundant, however, and biologists are working to find more plant species to fill in this temporal gap and provide a steadier nectar supply for the bees (Kuruvilla et al., 1995). As they do so, they will be practicing deliberate reconciliation ecology. Pest control Reconciliation may have considerable potential for minimizing losses to agricultural pests. For example, California viticulturists rely on a parisitoid wasp for biological control of leafhoppers in their vineyards (Doutt & Nataka, 1973), but the wasps require prey throughout the year and the grape leafhoppers become inactive in winter. So grape growers planted patches of native blackberries in shady spots near vineyards to maintain wasp populations. Each spring, wasps quickly re-invade the vineyards from the blackberry patches, thus keeping pest populations down. Although the growers goal was not to alleviate any crisis in biodiversity, their method helps to achieve precisely that. The mosaic of habitats they designed includes patches suitable for all the native species associated with blackberries. Pasturelands Pasturelands constitute a major part of the world s agri- cultural surface, and provide some encouraging examples of reconciliation. For example, the Ocosingo Valley in Fig. 6 The capacity of land to support diversity. Reconciliation ecology treats this as a continuous variable. It seeks techniques to move land to the right along that continuum. This it accomplishes by redesigning human habitats to give some species back their geographical ranges without taking away ours.

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