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Chapter 9 Biological Invasions and The CHAPTER 9 Biological Invasions and the Homogenization of Faunas and Floras Julian D. Olden 1 , Julie L. Lockwood 2 , and Catherine L. Parr 3 1 School of Aquatic and Fishery Sciences, University of Washington, Seattle, USA 2 Ecology, Evolution, and Natural Resources, Rutgers University, New Brunswick, NJ, USA 3 Environmental Change Institute, School of Geography and the Environment, University of Oxford, Oxford, UK 9.1 THE BIOGEOGRAPHY OF however, the large majority of species are not distrib- SPECIES INVASIONS uted broadly, because individuals of most species have limited dispersal capabilities. In considering the distribution of organic beings These limitations on dispersal ability have produced over the face of the globe, the fi rst great fact which the interesting phenomenon that many, perhaps even strikes us is, that neither the similarity nor the dis- most, species do not occupy all of the areas of the world similarity of the inhabitants of various regions can in which they could quite happily thrive. Instead, they be accounted for by their climatal and other physical are restricted to certain regions, where they are able to conditions … A second great fact which strikes us in interact with only those species with which they co - our general review is, that barriers of any kind, or occur. The limited geography of species is responsible, obstacles to free migration, are related in a close and in part, for the fantastic array of diversity that presently important manner to differences between the pro- carpets the Earth, as it provides opportunity for con- ductions of various regions. vergent evolution in disparate unconnected regions. (Charles Darwin, 1859 , pp. 395 – 396) With the range expansion of modern humans, ini- tially out of Africa, then across the globe, came the possibility of human - mediated dispersal of a large 9.1.1 The i nvasion p rocess variety of other species. By this, we mean that humans provided the conduit for individuals of some species to One of the fundamental elements of life on Earth is disperse much farther abroad than they could natu- change. Species appear through time via evolution and rally. Species were moved within, or on, humans as disappear by the natural actions of environmental parasites or disease organisms, in their household change (e.g. volcanic eruptions, changing sea levels, goods as hitchhikers, as their livestock or working glaciation). Species have also regularly shifted their animals, as their crop plants, as their pets, and as com- geographical ranges in response to biological and phys- modities themselves. ical forces, sometimes becoming less common and There is written evidence that intentional move- other times becoming more widespread. In general, ments of species by humans traces back to ancient Conservation Biogeography, First edition. Edited by Richard J. Ladle and Robert J. Whittaker. © 2011 by Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd. Conservation planning in a changing world 225 times, such as the introduction of the tamarind tree ( Tamarindus indica ) into China by way of commerce along the Shu - Yan trade route that linked China to India 8,000 years ago (Yan et al. , 2001 ). Some species apparently have nearly circumglobal distributions because of ancient trade activities, with many of these examples only recently coming to light thanks to the power of molecular analyses to locate the evolutionary origins of now very widespread species (e.g. Wares et al ., 2002 ). There is ample historical evidence that the number of species that were moved out of their native ranges and introduced to somewhere novel via human actions increased as the world began to become ever more interconnected (Elton, 1958 ). As this number grew, the need to understand how this process occurs, and to differentiate natural species ’ range expansions from those mediated by humans, became critical. Without making this distinction, it becomes diffi cult to untangle the mechanisms that are driving historical biodiversity changes, to understand the role of new arrivals in driving evolutionary dynamics and, more practically, to stem the fl ow of species that cause ecological or economic harm (see below). Before continuing, however, it is very important to recognize that a multitude of names have been given to species that are introduced to a novel location via human actions – such as ‘ exotic ’ , ‘ invasive ’ or ‘ alien ’ species (Lockwood et al. , 2007 ). We use the term ‘ inva- sion ’ to refer to the process whereby species expand their geographical distribution outside of their natural Figure 9.1 Generalized stages common to all species dispersal range via the actions of humans, while we invasions. A species must successfully transition through refer to populations that have become otherwise estab- each sequential stage, and the proportion of species that lished outside the bounds of their native ranges as proceed from one stage to the next is less than the previous ‘ non - native ’ . one (depicted by arrow width). A more lucid understanding of the invasion process may be achieved if it is considered as a stepwise pro- ranges and become pests. These estimates were based, gression of events, whereby individuals of some species in large part, on non - native animals and plants of are moved out of their native ranges, released into a Britain. More recently, Jeschke and Strayer (2005) novel location, establish self - sustaining populations investigated all freshwater fi sh, mammal and bird there and then spread to new locations (Figure 9.1 ; species native to Europe or North America that have Sakai et al. , 2001 ). been introduced outside their native range. They found Fundamental to this process is that not all individu- that the frequencies of transitions across all three of als successfully pass through all these stages. The tens the above stages averaged 6.1 per cent, 56.0 per cent rule of Williamson (1996) states that only ≈ 10 per and 59.7 per cent, respectively. cent of transported individuals are released into a Regardless of the specifi c percentages for each stage, foreign location, ≈ 10 per cent of these introduced it is apparent that only a fraction of the species that are species will go on to survive and successfully breed (i.e. moved by people, either on purpose or by accident, will establish a new population) and ≈ 10 per cent of these complete all stages of the invasion process. A consider- established species will expand their geographical able amount of research within invasion biology has 226 Biological invasions and the homogenization of faunas and fl oras therefore focused on attempts to understand which show a much wider array of biological traits than factors differentiate between those species that suc- those species that are likely to experience natural long - cessfully progress through all invasion stages and those distance dispersal. that do not (Lockwood et al. , 2007 ). The rate at which non - native populations are estab- lishing around the world is consistently several orders of magnitude larger than fossil - derived estimates for 9.1.2 Human - a ssisted v ersus natural dispersal events at the same locations. For p rehistoric i nvasions example, the invasion rate of terrestrial species for the Hawaiian Islands was approximately 30 species per A valid and persistent question is the extent to which million years (0.00003 per year) prior to human set- modern trends in species invasions differ from those tlement, but increased to 20,000 species per million that occur naturally. This question is especially rele- years (0.02 per year) after the arrival of the Polynesians vant to students of biogeography because range expan- and to approximately 20 per year during the past two sions are a very clear component of palaeoecological centuries (Ricciardi, 2007 ). In other words, contempo- and historical biodiversity patterns (Vermeij, 2005 ). rary rates of biological invasions are nearly one million Do modern invasions warrant the attention currently times higher than the prehistoric rate for Hawaii before given to them by scientists? How different are the human infl uence. mechanisms, spatial patterns and rates of modern The number of individuals of each species being versus prehistoric invasions? Can we use prehistoric transported is also vastly different between natural and trends to help predict the consequences of modern bio- human - assisted invasion events. Natural dispersal logical invasions? events typically involve a few individuals of a species Human - assisted dispersal of non - native species fi nding their way out of the native range and attempt- differs from natural dispersal events in several impor- ing to establish a self - sustaining population in the tant aspects (J.R.U. Wilson et al. , 2009 ). Ricciardi novel locale. Occasionally the number of individuals in (2007) detailed the differences between prehistoric these natural events can be quite high – as for instance, and human - assisted invasions, which we summarize during biotic interchanges involving episodic events of below and in Table 9.1 . mass dispersal. For example, the opening of the trans- The most obvious differences are in the number polar corridor between the Pacifi c and Atlantic oceans and frequency of ‘ dispersal ’ events. Natural dispersal and the formation of the Panamanian land bridge events are characteristically rare, both in the number between North and South America during the Great of species being transported and in the temporal fre- American Interchange permitted
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