Ecology. Michael Begon
allowed hypotheses that were once untestable to be critically evaluated. Evolutionary ecologists are not so focused now on whether or not sympatric speciation can happen, but rather how often and under what conditions.
APPLICATION 1.3 Conservation significance of hot spots of endemism
Conservationists have to make hard decisions in their quest to preserve biological diversity. Given limited resources, how can the most species be supported at minimum cost? One way is to focus attention on 'biodiversity hot spots' of species that are found nowhere else. Myers et al. (2000) took this approach when mapping the entire globe in terms of exceptional concentrations of endemic species coupled with exceptional loss of habitat (and therefore subject to a greater degree of threat to biodiversity than areas without such habitat loss). Hot‐spot boundaries were set according to the characteristic biotas they contain: examples include island groups such as the Galápagos (Section 1.3.2) and Hawaii (Section 1.4.2), and 'ecological' islands such as the East African Great Lakes (Section 1.3.3) or clearly defined continental units such as the Cape Floristic Province in South Africa. The taxa included in the analysis consisted of vascular plants, mammals, birds, reptiles and amphibians. Figure 1.13 shows 25 identified hot spots that between them contain 133 149 plant species (that is, 44% of the world's plants) and 9645 vertebrate species (35% of the world's total). Or to put it in another way that emphasises their importance, we can say that this set of hot spots provide the sole remaining habitats of 44% of the world's plant species (and 35% of animals).
Figure 1.13 Biodiversity hot spots. Twenty‐five biodiversity hot spots identified because of their exceptional concentrations of endemic species that are undergoing exceptional levels of human induced habitat loss.
Source: From Myers et al. (2000).
The five most prominent hot spots, the tropical Andes, Sundaland, Madagascar, Brazil's Atlantic Forest and the Caribbean, contain 20% of the world's vascular plants and 16% of vertebrate species but together they comprise only 0.4% of the world's surface. Moreover, they are subject to some of the heaviest levels of habitat loss: the Caribbean retains only 11.3% of its primary vegetation, Madagascar 9.9%, Sundaland 7.8% and Brazil's Atlantic Forest 7.5%. There was reasonable congruence between levels of endemism of plants and vertebrates in the hot spots, but note that no invertebrates were included in the analysis. In a geographically more restricted study in South Africa, Bazelet et al. (2016) showed that there was congruence between hot spots of the rather circumscribed diversity of katydids (bush crickets) and the biodiversity hot spots already recognised for much wider groupings, indicating that the conservation of biodiversity hot spots may often also protect non‐target organisms.
Myers et al. (2000) called for a more than 10‐fold increase in annual funding from governmental and international agencies to safeguard these hot spots.
1.4 The role of historical factors in the determination of species distributions
Our world has not been constructed by someone taking each species in turn, testing it against each environment, and moulding it so that every species finds its perfect place. It is a world in which species live where they do for reasons that are often, at least in part, accidents of history. We illustrate this first by considering continental drift, a process that operates over a timescale of tens of millions of years.
1.4.1 Movements of landmasses
Long ago, the curious distributions of species between continents, seemingly inexplicable in terms of dispersal over vast distances, led biologists, especially Wegener (1915), to suggest that the continents themselves must have moved. This was vigorously denied by geologists, until geomagnetic measurements required the same, apparently wildly improbable explanation. The discovery that the tectonic plates of the earth’s crust move and carry with them the migrating continents, reconciles geologist and biologist (Figure 1.14). Thus, whilst major evolutionary developments were occurring in the plant and animal kingdoms, populations were being split and separated, and land areas were moving across climatic zones.
Figure 1.14 Continental drift means that continents that are now separate were once joined to one another. (a) The ancient supercontinent of Gondwanaland began to break up about 150 million years (Myr) ago. (b) About 50 Myr ago (early Middle Eocene) recognisable bands of distinctive vegetation had developed, and (c) by 32 Myr ago (early Oligocene) these had become more sharply defined. (d) By 10 Myr ago (early Miocene) much of the present geography of the continents had become established but with dramatically different climates and vegetation from today; the position of the Antarctic ice cap is highly schematic.
Source: After Norton & Sclater (1979), Janis (1993) and other sources.
placental and marsupial mammals
The drift of large landmasses over the face of the earth explains many patterns in the distribution of species that would otherwise be difficult to understand. A classic example is provided by the placental and marsupial mammals. Marsupials arrived on what would become the Australian continent about 90 million years ago (in the Cretaceous period), when the only other mammals present were the curious egg‐laying monotremes (now represented only by the spiny anteaters (Tachyglossus aculeatus) and the duckbill platypus (Ornithorynchus anatinus)). An evolutionary process of radiation then occurred that in many ways paralleled that of placental mammals on other continents (Figure 1.15). The subtlety of the parallels in both the form of the organisms and their lifestyle is so striking that it is hard to escape the view that the environments of placentals and marsupials provided similar opportunities to which the evolutionary processes of the two groups responded in similar ways. Because they started to diversify from a common ancestral line, and both inherited a common set of potentials and constraints, we refer to this as parallel evolution (as opposed to convergent evolution, where structures are analogous (similar in superficial form or function) but not homologous (i.e. not derived from an equivalent structure in a common ancestry), such as the wings of birds and bats). The important point here, though, is that the marsupials are found where they are not simply because they are the best fitted to those particular environments but also because of an accident of history – in this case, geological history.