1 Pacific Science (1995), vol. 49, no. 1: by University of Hawai'i Press. All rights reserved Why Do Introduced Species Appear to Devastate Islands More Than Mainland Areas?! DANIEL SIMBERLOFF 2 ABSTRACT: Island biotas are viewed popularly as much more fragile than those of mainland areas and much more prone to damage from invaders. There are far too few data to assess this view thorougwy; for example, failed invasions are often unrecorded, and claims that an introduced species has displaced a native one are often based on correlated population changes rather than experiment and/or detailed field observations. If there is a tendency for invasions to affect island communities more than mainland ones, it is far from universal; virtually every kind of damage wrought by invaders on islands has also been wrought in mainland areas. It is unlikely that, by virtue of their reduced species richness alone, island communities pose less "biotic resistance" to invaders than mainland communities do. Rather, certain entire groups of species, like terrestrial mammals, are often missing from islands, and these absences can predispose certain invaders to be especially likely to survive and to produce particular impacts. CONVENTIONAL WISDOM (e.g., Elton 1958, (1968) saw islands as evolutionary "backcarlquist 1965, Wilson 1965) is that islands waters and dead ends," populated by species are more easily invaded than mainland areas, whose relatives had long ago been exand ecological effects of invasions are greater tinguished in mainland areas. Greuter (1979) on islands than in mainland situations, be- depicted islands as repositories for old relicts, cause island biotas are somehow fragile and relatively unimportant in plant evolution. the species weaker than those of the main- Williamson (1981) believed that slower analand. Yoon (1992: 88) eloquently expressed genetic evolution on islands leaves island enthis view with respect to the Hawaiian Is- dernics likely to be extinguished by competilands: "But the isolation of these gentle is- tion from introduced mainland species. In lands from mainland predators and diseases addition to the susceptibility of island biotas created a flora and fauna ill equipped to engendered in the long term by evolutionary handle the rigors of competition with the processes, the small number of species and outside world. Hawaii is home to thistles supposedly simplified food web structures without prickles, blackberries without thorns were believed to make island communities and many flightless insects and birds." particularly invasible (e.g., Wilson and BosThe idea is that island species have been sert 1971). This belief was part of the dogma subjected to much less intense selective pres- that diverse ecosystems are more stable by sures. In particular, the smaller number of virtue of their diversity (MacArthur 1955, species on islands suggests that competition is Elton 1958). The notion that introduced speless intense, so island species evolve to be less cies would be much more likely to survive on "competitive." Mayr (1965) found island islands than in mainland areas even has avifaunas to offer "far less resistance" to im- management implications. For example,.-. migfants-than-mainland-faunas- do~-preston.. Immst193-1-)-felt-that-intro-ducin-glfaturaI-enemies for biological control would be an effective method only in the simplified com1 Manuscript accepted 27 April munities of islands. 2 Department of Biological Science, Florida State University, Tallahassee, Florida Nowadays, key aspects of the arguments 87
2 88 PACIFIC SCIENCE, Volume 49, January 1995 underpinning this conventional wisdom are questionable on numerous empirical grounds. First, is competitive pressure really stronger on the mainland? Though Crowell (1962) and MacArthur et al. (1972) have seen competition among species in bird communities as lower on islands, Schoener (1965), Grant (1968), Keast (1970), and others have argued that competition on islands may be stronger than in mainland areas because the resource base is reduced and that this more intense interspecific competition is reflected in greater morphological divergence of close relatives on islands than in mainland situations. Perhaps generic statements about the relative intensity of competition on islands and the mainland are not very useful, and one must always speak of a specific island and mainland and also determine the relevant resource base empirically. Second, the relationship between stability and diversity or complexity is now seen as complicated and not monotonic (Begon et al. 1990, and references therein); some diverse, complex systems are very fragile and some simple ones seem stable. That islands tend to have fewer species than mainland areas is likely to be far less important, from the standpoint of invasibility, than the nature of those species that are present (Goodman 1975). Aside from assessing the underlying reasons for believing that island communities are more likely than mainland ones to be disrupted by introduced species, it is also worth examining whether empirical data really demonstrate this heightened susceptibility. Probability That an Introduced Species Survives on an Island A problem in assessing the likelihood that an introduced species survives on both mainland and islands is general lack of information on failed introductions (Simberloff 1981, 198-6a}--Apprax-imately slll"Viving---introduced insect species are known from the contiguous United States (Anonymous 1986), and about 2500 are known from the Hawaiian Islands (Funasaki et al. 1988, Nishida 1992). But these are only surviving introduced species; who knows how many propagules of other species landed but did not lead to populations or produced small populations that went extinct before being recorded? Surely the number must be greater for the contiguous United States, but how much greater? Probably a lot, so it seems as if the probability of an introduced insect's surviving in Hawai'i is greater. Because Hawai'i has about 5000 described native species (Nishida 1992) and the contiguous United States has about 90,000 (Arnett 1983), the probability of survival seems greater in the site with fewer species. Is this probability of survival greater because there are fewer species there to resist the invader? An order-by-order examination of the introduced insects of Hawai'i is not consistent with this hypothesis (Simberloff 1986a). For example, Hawai'i has proportionally more surviving introduced Coleoptera and Lepidoptera species than does the contiguous United States, but it also has proportionally more native Coleoptera and Lepidoptera. By and large, even though the native insect orders are very differently represented in Hawai'i than in the contiguous United States (many orders are even missing), the relative proportions of surviving introduced species in the different orders are very similar in the two sites. Without knowing how many species of each order were introduced in each place, we cannot even say where the probability of survival is greatest, so we cannot really assess the hypothesis as we would wish to, but the data on surviving species suggest that the question is not trivial. Probably the best published data for survival and disappearance of introduced species come from the literature on biological control, although even here there have been many unrecorded, more or less ad hoc introductions (F. G. Howarth, pers. comm., 1991). From an encyclopedic list of biocontrol introductions (Clausen 1978), I found -six- genel'a-~five-parasiticwasp genera -and-a beetle genus) with the largest number of species that had been introduced, in various directions, among islands and mainland, with Australia considered a mainland (Simberloff ::::mwa:awsuih brim' this 1+5 iff'''i' U2~a.'.4 MAt gw
3 Impact of Introduced Species on Island Biotas-SIMBERLOFF a). I tabulated successes and failures of 1983). There are probably some competitive introductions from mainland to island, interactions between introduced birds that do mainland to mainland, island to island, and invade native forest (especially Zosterops island to mainland. If there were less biotic japonicus Temminck & ScWegel) and native resistance posed by island biotas and island birds (Scott et al. 1986). But habitat restricspecies were generally less competitive, one tions surely greatly reduce potential comwould expect mainland-to-island introduc- petition between introduced species and nations to have survived far more frequently tive ones, so that it is unlikely that resistance than island-to-mainland ones, with the other or lack thereof by native birds is the reason two classes intermediate. These six genera for success or failure of species introduced to produced 281 introductions; although the Hawai'i. probabilities of survival are in the order pre- Similarly, 20 passeriforms have been indicted, the probabilities are all very close. troduced to the Mascarene Archipelago, of For example, 49% of mainland-to-island in- which nine survived on all islands to which troductions survived, and 33% of island-to- they have been introduced, nine failed whermainland ones. A multiway contingency test ever introduced, and two survive tenuously does not reject the hypothesis that survival or on La Reunion but disappeared from Maudisappearance of a propagule does not de- ritius after more than a century (Simberloff pend on whether source or target areas are 1992a). As in the Hawaiian Islands, there is island or mainland. general habitat separation between the in- The general relevance of this result is less- troduced species and the native ones, with the ened because most biological control efforts native species in upland native forest and the on both islands and mainland are in agri- introduced species in lowland open anthrocultural communities, which are both far pogenous habitat. Only Zosterops borbonicus from pristine and more similar to one an- borbonicus (Boddaert) among the natives inother, no matter whether on island or main- vades open lowland habitats, and only Fouland, than pristine communities would be. dia madagascariensis (L.) among introduced Thus the species introduced to islands for species is found in forests; for both species biological control purposes were not, by and the anomalous habitat is a very secondary large, faced at the outset with competition one (Barre 1983). The species whose invaand other interactions from the entire native sions failed were also primarily species of community. Rather, resistance was from a open habitat; it is difficult to believe that small subset of it (that lives in anthro- their failure was caused by competition from pogenous habitats), as well as from other in- native birds. troduced species. This same problem obtains The smaller number of species on islands when one considers introduced birds, for than in nearby mainland areas is the most which some data exist for failures as well as widely known characteristic of island biotas. successes. At least 50 passeriform species Invasion by a new species of a new site, ishave been introduced to the Hawaiian Is- land or mainland, is often viewed in terms of lands, of which 30 survived on all islands to the species' having to find an "empty niche" which they were introduced, 17 failed on all (Udvardy 1969, and references therein). Alislands to which they were introduced, and 3 though the static "niche" that this view survived on some islands and failed on others entails has given way to a more dynamic (Simberloff and Boecklen 1991). However, conception, the general idea still surfaces frefor the most part the introduced birds are quently and has management implications. found in agricultural lands, exotic forests, For example, many game species are inand built up-areas, and-thesewerealsotypi- --- tregueea-tefill-suppesedlyempty-nichesoncal habitats of those species that failed. The islands (Ebenhard 1988). The vacancy of native land birds are primarily found in the niches is notoriously hard to prove (Herbold native forests, almost wholly restricted to and Moyle 1986). Further, since resources uplands (Berger 1981, Moulton and Pimm are ultimately metabolized in some way or
4 90 PACIFIC SCIENCE, Volume 49, January 1995 other, even if only by decomposers, the de- specific interactions can be very difficult to piction of a niche as empty can always be elucidate without a controlled experiment or construed as simply a different way of de- very detailed observations, yet such data are fining the niches of a particular community usually lacking. Suggestive observations can (Simberloff 1991). be very misleading. For example, the rapid Nevertheless, it is clear that the reduced spread of the North American mink (Mustela number of species on islands often leads to vison Schreber) in Great Britain in the 1950s the absence of entire ways of making a living. coincided approximately with the decline of The literature on disharmony of island biotas the otter. This synchrony led many to infer a (e.g., Carlquist 1974, Begon et al. 1990) de- causal nexus; however, pesticide pollution is picts this same absence. So it is not surprising the chief reason otter populations have dethat certain types of species are especially clined (Chanin and Jeffries 1978). likely to survive on islands if introduced. The The subtlety of effects of introduced speafrican dung beetles introduced to Australia cies on natives, even some devastating effects, found a resource (300 million cowpats pro- and the absence of detailed study in the vast duced each day) that was not used in the way majority of cases make any discussion of this they use it, because the native dung beetles phenomenon highly anecdotal. For example, are not adapted to this resource. And, of it is often said that introduction of natural course, the numerous successful introduc- enemies, especially insect parasitoids and tions of predatory mammals such as rats, predators, for biological control is envirsmall mustelids, and the small Indian mon- onmentally friendly. There is good reason to goose, Herpestes auropunctatus Hodgson, to be skeptical (Simberloff 1992b). Gagne and small islands lacking such species but con- Howarth (1985) and Howarth (1985) argued taining abundant potential prey (often birds that such introduced insects might have and their eggs) can be viewed in this light (cf. caused the unintended extinction of several King 1984, Atkinson 1989). endemic Hawaiian moth species, a charge Thus, there may be a generic tendency for contested by Funasaki et al. (1988). The specertain types of introductions to be more cifics of this case cannot be resolved with likely to survive on islands than in mainland available data (see Simberloff 1992b for deareas. About 1.6 times as many mammal tails), but it is difficult to believe that enough species and three times as many bird species attention was paid to species of pristine enhave been successfully introduced to islands vironments, especially relatively rare insects than to mainland areas (Atkinson 1989). of no obvious economic importance, to supwithout a tabulation of how many attempts port the general conclusion of Funasaki et al. were made in each setting, one cannot say (1988) that biological control introductions with certainty that the rate of survival was are not harming native nontarget species greater on islands, but I think it is likely that (Simberloff 1992b). many of these island successes were facilisimilarly, Groves (1989) claimed that no tated by absence of native species with sim- insect introduced to control an invasive plant ilar "niches." has affected a native congeneric plant, but this claim is incorrect and, in any event, there Ecological Impacts of Surviving has not been nearly enough field study of Introduced Species native plants, especially in pristine environments, to justify lack of concern. A candidate With respect to degree of ecological im- endangered species, the semaphore cactus pact, the published record on introduced (Opuntia spinosissima Miller), was found in species-is-deficient-on-tw0-g0unts.-f-irst,it-is- --the-united-states -only_on_littletorch-ke, more likely that dramatic effects will be no- Florida. All individuals there have been deticed, and the introduced species associated stroyed or are under attack by caterpillars of with them thus recorded, than if there is the moth Cactoblastis cactorum (Berg), which minimal ecological impact. Second, inter- was deliberately introduced to the island of N'f2LllJilbj s QN
5 Impact of Introduced Species on Island Biotas-SIMBERLOFF 91 Nevis in the Lesser Antilles to control another cactus and island-hopped over about 30 yr all the way to Florida. The important lesson from this introduction is that the cactus would have gone quietly extinct but for a very unlikely set of circumstances. It inhabited an area favored by plant collectors and botanists, in an intensively studied hammock that is part of a Nature Conservancy refuge, and a local botanist was tipped to the possibility of the moth's presence by a colleague who had noticed it at Guantanatno Bay, Cuba (Simberloff 1992b). How many species have been reduced or even extinguished without anyone's having thought to look? Granted that our knowledge of the true extent of ecological effects is very poor, what do the anecdotes suggest? We can begin by looking at one dramatic effect, extinction. The number of known extinctions apparently caused by introduced species is much greater on islands (Atkinson 1989). Probably the best-known reason is predation on island species that had not evolved in the presence of predatory mammals like cats, rats, and the small Indian mongoose, the key culprits aside from humans (Atkinson 1989). Absence of direct antipredatory behavior combined with habits such as nesting on the ground to predispose such species to extinction. In fact, destruction of habitat by browsing and grazing by introduced pigs, rabbits, sheep, cattle, horses, and other species has probably led to more island extinctions than predation has, either by direct elimination of an entire plant species or by reducing habitat quality and amount so much for animal and plant species that they are more prone to extinction from the various factors that place very small populations at risk (outlined by Simberloff [1986b]). Although such habitat destruction is often depicted as far greater on islands than on the mainland (e.g., Atkinson 1989), there is really no thorough study of this matter. For example, pigs devastated plant-- communities of-:a:uckland- Island; re~stricting some species to sites inaccessible to pigs (Atkinson 1989). Pigs have also greatly affected plant communities and several species of animals in the Great Smoky Mountains National Park (references in Simberloff ). Mack (1986, 1989) explained the destruction of the native plant communities of the North American intermountain West and their replacement by introduced Old World plant species as a consequence of the introduction of large congregating grazers, to which American plant species were not adapted. A similar fate befell South American and Australian grasslands when faced with Eurasian invaders. Is the ecological impact of grazers more severe on Auckland Island than in the intermountain West? No one has addressed this issue. It is difficult to find ironclad examples on either island or mainland in which introduced species have replaced native ones by competition, rather than by occupying a changed habitat (Udvardy 1969, Ebenhard 1988). Ebenhard (1988), acknowledging that many of the cases he tabulated are not strongly supported by empirical evidence, found that, of 1559 introductions of 330 species of birds and mammals worldwide, 79 introductions may have manifested competition between the invader and one or more native species, in that at least resources were jointly used. There were relatively more such cases in mainland areas than on islands, but, for introduced mammals (not birds), documented decreases in the population of the native species (whether or not caused by competition) were more common on oceanic islands. The inferences about real impact for most of these cases are so weak that it seems impossible to say whether introductions are more likely to lead to competition on islands than in mainland areas. This is a surprising result considering the conventional wisdom described at the beginning of this paper. All one can really say for mammals and birds is that competition seems to result from an introduction very infrequently, yet both island and mainland examples are known. Disease vectored by introduced species is often viewed as a key element in the devastation- wrought by-inva-stolrs- (e.g.,erosby 1986). Some of the introduced Asian passeriform birds may have contributed to the decline of native Hawaiian species by vectoring avian malaria (Ralph and van Riper 1985,
6 92 PACIFIC SCIENCE, Volume 49, January 1995 Van Riper et al. 1986), and a similar sce- sons, relating to the geological history of nario, though with no supporting data, was Eurasia and the long association of this resuggested for native Mascarene birds by gion with human activities, that might pre Cheke (l987a,b). In neither instance, how- dispose its species to be particularly adept at ever, does the evidence suggest that these invading other regions, including islands. diseases facilitated the survival of surviving One must always consider the opportunities invaders. Rather, the more likely scenario is for dispersal to and from various sites before that habitat modification created favorable postulating a hypothesis of greater likelihood circumstances for survival of at least some of inherent superiority of species from one introduced species, simultaneously reducing biota over those of another (Simberloff native populations, and the introduction of 1989). It may well be that far more species of new diseases may have worsened the plight of both disease and free-living organisms are the natives. transported from mainlands to islands than Exactly why island species should be more vice versa and that the per-capita probability prone than mainland ones to contract patho- of surviving does not differ. The same argugenic diseases from mainland invaders is un- ment may hold for disease organisms from known. Similarly, mainland species are rela- Eurasia compared with those from elsewhere. tively impervious to island diseases. Though I Perhaps disease susceptibility on islands is know of no statistical study of these proposi- a probabilistic phenomenon related to the tions, they are surely true. In addition to the species-area relationship. One can ask if there Hawaiian bird story, other anecdotes about is a null per-species probability that a paraisland populations devastated by introduced site will be severely pathogenic, so that rediseases are numerous (e.g., Ebenhard 1988, gions with more species have, on average, Pimm 1991), but I cannot identify an exam- more pathogenic species and potential hosts pie of a disease endemic to an island that has in such regions experience more natural sedevastated a mainland species. Ofcourse, in- lection for resistance. Another possibility is troduced diseases have devastated rich con- that the smaller size of island populations tinental communities-rinderpest from Asia means that total genetic diversity is, on averinvaded large parts of Mrica in the late age, lower than in mainland situations. Carnineteenth century with enormous, lasting son (1981) argued that total genetic diversity impact on the ruminants and, through them, of continental species is far greater than that entire ecosystems (Dobson and May 1986, of island species, by virtue of the larger Barbault 1992, and references therein). The number of local populations. Thus the probphenomenon may not be only a mainland-is- ability that resistant genotypes exist is lower land effect, but may be wholly or partly a on islands, and they will evolve more slowly statistical tendency for Eurasian diseases to once an introduced species arrives. Carson be more devastating in other areas. Only (1981) saw the reduced genetic base as resyphilis among New World human diseases sponsible for a generic tendency for island has invaded the Old World, whereas no hu- species to be less "aggressive," not wellman diseases have been exported from Aus- adapted "general purpose genotypes" that tralia to elsewhere (Crosby 1986). Almost all typify invaders from the mainland. He devastating plant disease epidemics have pointed out (H. L. Carson, pers. comm., been caused by Eurasian pathogens in- 1993) that this pattern is far from universal. troduced to other continents (von Broembsen For example, among the endemic Hawaiian 1989). The pathogenicity ofeurasian human picture-winged Drosophila, most species are diseases and resistance by Eurasian humans monophagous, but five are so polyphagous -- -rnight-bea-result-ef-greater--eurasian-human. that-eagh-feeds -0n -at--ll::ast-tivl::--h0st-plant population densities, but the analogous trend families, including some introduced plant for plant pathogens cannot be so explained. species (Montgomery 1975). There has been Di Castri (1989) suggested generic rea- no precise quantification of aggressiveness
7 Impact of Introduced Species on Island Biotas-SIMBERLOFF 93 and general purpose capability, much less a tially has enormous ecosystem consequences, comprehensive comparison of all island and as do the introductions of some plants that mainland species to see if these traits are enhance fire frequency in various Hawaiian more frequent among the latter. ecosystems (Vitousek 1986). One possible generic reason why island It is not clear, however, that introduced ecosystems may be more prone to damage species are more likely to play such "keyfrom introductions is that their species' pop- stone" habitat-modification roles on islands ulations are perforce smaller and more geo- than in mainland areas. The introduction of graphically restricted, and so less likely to salt cedar (Tamarix spp.) and Russian olive have some refuge, some regions not exposed (Eleagnus angustifolia L.) along rivers in the to predators, diseases, grazers, and so forth, American Southwest has created new forests than would mainland species exposed to even more extensive than those of Hawaiian similar invaders. Thus, extinction of a native mangroves with major ecological impact species is more likely for a given introduction (references in Simberloff ). And inon an island, and, after extinction has oc- troduced nitrogen-fixers are a major threat to curred, reinvasion and succession as means dune plant communities in northern Calof repairing the ecological damage are fore- ifornia (references in Simberloff). Most closed. On the mainland, by contrast, even if of the northeastern coast of North America vast areas are devastated, some areas are has been changed from mud flats and salt more likely to be unaffected by an introduced marshes to rocks by an introduced European species, and, if the introduced species is re- periwinkle snail, Littorina littorea L. (references in Simberloff), and native forests moved or controlled, reinvasion is possible. Introduced species can have enormous of much of South Florida have been replaced impacts on target ecosystems without directly by Melaleuca quinquenervia (Cav.) from causing extinctions. Species that constitute Australia and Schinus terebinthifolius Raddi entirely new habitats can entrain major from South America (Ewel 1986). The latter changes in a biota. For example, native two species are particularly inimical to the mangroves cover soft intertidal substrates in native biota because of the enhanced fire remost tropical bays and estuaries, but there gimes they foster. were none in the Hawaiian Islands until an However, the same reduced number of introduction in Now mangroves, espe- species and disharmony noted above as cially Rhizophora, have spread, largely on characteristic of islands, especially small istheir own, to many suitable sites throughout lands, would probably make it more likely the archipelago, forming substantial forests that an introduced species would "fill an in some areas where none had previously ex- empty niche" in such a way as to become a isted. There has been little study of the ter- new habitat on an island than on the mainrestrial or marine ecology of these forests, land. but the effect of this introduction on some ecological functions must be enormous Conclusions (Simberloff 1991). For example, mangrove swamps typically drop about 10,000 kg of The data are inadequate to draw strong leaves per hectare per year and accumulate conclusions about the relative invasibility sediment that builds land (Holdridge 1940). and fragility of islands and mainland. In They are major fishery nurseries (Hutchings particular, one would have to know much and Saenger 1987, and references therein). more about frequency of different types of Thorough modification of an existing habitat introductions and number of failed introduccan also-cause-community-wide changes.--fol" - ti0ns.--f0fe~ample,- the-fact that no -island example, the invasion of young volcanic bird species has invaded continents to the areas of Hawai'i by the Atlantic plant Myr- extent that species like the house sparrow or ica faya Aiton, which fixes nitrogen, poten- the Java sparrow have invaded islands may
8 94 be wholly or partly a consequence of the fact that such introductions of island species to mainland areas have never been performed. Virtually all attempts to introduce island birds have been from one small island to another, usually in the same region, and many have survived (Long 1981). Despite the paucity of quantitative data, it appears that introduced species are more likely to survive on islands than in mainland areas, and are more likely to produce major ecological impacts, but there are many counterexamples. If this tendency exists, it is probably not generated by an inherent, community-wide "biological resistance" posed by mainland communities to invaders but vitiated on islands. It is not simply a matter of greater species richness in mainland areas conferring greater ecological stability. Rather, it seems that the absence of entire groups of species from islands may have predisposed certain invaders from the mainland to be likely to survive on islands and certain introductions to be likely to have greater ecological effects. In other words, the proximate factor in determining the trajectory of an introduction may be the specific type of species and the specific community it is introduced to, rather than simply the size of the island community. However, over the long term, the size and composition of the island community may have led to the evolution of a community likely to be susceptible to certain types of damage from invasions. It is also possible that, if there is some inherent invasibility and fragility of island communities, at least part of it is really an aspect of an inherent invasibility and fragility of all communities when faced with Eurasian introductions. This is not a very satisfactory conclusion to a paper about the supposed vulnerability of island species and communities to invasion. Species of some native island communities seem to have resisted invasion as well as continental ones do. For example, in Ha- PACIFIC SCIENCE, Volume 49, January 1995 It would be worth exploring whether this pattern extends to other archipelagoes. In general, it is far from clear that island species do tend to be weak, unaggressive, uncompetitive evolutionary incompetents; if there is such a tendency, it is far from universal, and the reasons are still unknown. ACKNOWLEDGMENTS I thank K. Kaneshiro, H. Carson, S. Pimm, L. Hamilton, and two anonymous referees for suggestions on the manuscript and K. Kaneshiro for facilitating my participation in this symposium. LITERATURE CITED ANONYMOUS Western Hemisphere Immigrant Arthropod Database (WHIAD). USDA Agricultural Research Service, Beltsville, Maryland. ARNETT, R. H Status of the taxonomy of the insects Qf America north of Mexico: A preliminary report prepared for the subcommittee for the insect fauna of North America project. Privately printed. ATKINSON, I Introduced animals and extinctions. Pages in D. Western and M. Pearl, eds. Conservation for the twenty-first century. Oxford University Press, New York. BARBAULT, R Ecologie des peuplements. Structure, dynamique, et evolution. Masson, Paris. BARRE, N Distribution et abondance des oiseaux terrestres de l'lle de la Reunion (Ocean Indien). Rev. Eco!. Terre Vie 37: BEGON, M., J. L. HARPER, and C. R. TOWNSEND Ecology: Individuals, populations and communities, 2nd ed. Blackwell, Boston. BERGER, A. J Hawaiian birdlife, 2nd wai'i~the-plants-ofupland-nativeforests-are- ---ea-o-ynive-rsit-y 0t'-Hawaii-I>-ress,-Honolulu. not greatly affected by exotic plants (Mueller- CARLQUIST, S Island life. Natural History Press, Garden City, New York. Dombois 1981), and plant communities of extreme habitats seem not to be easily in Island biology. Columbia University Press, New York. vaded (Loope and Mueller-Dombois 1989). 5 Ed# ;am,i;
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Ecology, 77(6), 1996, pp. 1661-1666 1996 by the Ecological Society of America The varying success of invaders 1 Mark Williamson & Alastair Fitter Department of Biology, University of York, York YO1 5DD,
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