Biogeographical Regions

Richard Huggett, University of Manchester, Manchester, UK

Species are not uniformly distributed over the land surface. Fauna and flora display regional differences. The largest regions of animals and plants are biogeographical regions, each bearing a distinctive fauna and flora. Some families and even some orders of animals are endemic to particular biogeographical regions. Other families are shared by two or more regions. A few families are cosmopolitan, being found in all biogeographical regions.

Introductory article
Article Contents
. P. L. Sclater, A. R. Wallace and the Foundational Units of Biogeography . Mammals . Floral Regions . Comparisons and Contrasts between Taxa . Transitional Zones and Filters . The Applied Use of Biogeographical Regions: Their Place in Conservation

P. L. Sclater, A. R. Wallace and the Foundational Units of Biogeography

Dierent places harbour dierent kinds of animals and plants. The fauna of Africa is unlike the fauna of North America; the ora of Japan is unlike the ora of South Africa. These regional dierences in the distribution of species became increasingly manifest as the world was explored. George Leclerc, Compte de Buon (17071788) studied the then known tropical mammals from the Old World (Africa) and the New World (Central and South America). He found that they had not a single species in common. Later comparisons of African and South American plants, insects and reptiles evinced the same pattern. By the nineteenth century, it was clear that the land surface could be divided into biogeographical regions, each of which carries a distinct set of animals and a distinct set of plants. Augustin-Pyramus de Candolle considered plants and identied areas of endemism, that is botanical regions, each possessing a certain number of plants peculiar to them. He listed 20 such botanical regions or areas of endemism in 1820, and by 1838 had added another score, bringing the total to 40. In 1826, James Cowles Prichard, a zoologist, distinguished seven regions of mammals: the Arctic region, the temperate zone, the equatorial regions, the Indian isles, the Papuan region, the Australian region, and the extremities of America and Africa. William Swainson modied this scheme in 1835, by taking account of the ve recorded varieties of humans, to give ve regions: the European (or Caucasian) region, the Asiatic (or Mongolian) region, the American region, the Ethiopian (or African) region, and the Australian (or Malay) region. The early ideas of Prichard and Swainson on animal distributions were eclipsed by the seminal work of an English ornithologist, Philip Lutley Sclater, and the eminent English biogeographer and naturalist, Alfred Russel Wallace. Using bird distributions, Sclater (1858) recognized two basic divisions (or creations, as he termed them) the Old World (Creatio Paleogeana) and the New World (Creatio Neogeana) and six regions. The Old World he divided into Europe and northern Asia, Africa south of the Sahara, India and southern Asia, and

. Summary

Australia and New Guinea. The New World he divided into North America and South America. Sclaters schema prompted a urry of papers by English-speaking zoologists, including Thomas Henry Huxley and Joel Asaph Allen, each of whom promulgated his own favoured geographical classication. In his The Geographical Distribution of Animals (1876), Wallace reviewed the competing systems, arguing persuasively in favour of adopting Sclaters six regions, or realms as Wallace dubbed them. Sclaters system and Wallaces minor amendments to it provided a nomenclature that survives today (Figure 1). Later suggestions were minor variations on the Sclater Wallace theme. Sclater and Wallace identied six regions Nearctic, Neotropical, Palaearctic, Ethiopian, Oriental and Australian. Together, the Nearctic and Palaearctic regions form Neogaea (the New World), while other regions form Palaeogaea (the Old World). Wallaces contribution was to identify subregions, four per region, which correspond largely to de Candolles botanical regions. Indeed, the nineteenth-century classication of

Nearctic Ethiopian

Neotropical Oriental

Palaearctic Australian

Figure 1 The six faunal regions delimited by Sclater and Wallace.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net

Biogeographical Regions

biogeographical regions was essentially an attempt to group areas of endemism into a hierarchical classication according to the strengths of their relationships. It is surprising and noteworthy that the distributions of species with good dispersal abilities, including plants, insects and birds, tend to fall within traditional zoogeographical regional connes. The avifaunas of North America and Europe contain several families and many genera that are not shared by the two regions, even though dispersal across the North Atlantic and Pacic Oceans by accidental visitors is noted every year. Even long-distance migrant bird taxa tend to be conned either to the eastern hemisphere or to the western hemisphere, where they migrate between high and low latitudes, and appear illdisposed to disperse eastwest between continents.

Mammals
Of the six faunal regions delineated by Sclater and Wallace, the Palaearctic is the largest. It includes Europe, North Africa, the Near East and much of Asia (but not the Indian subcontinent or Southeast Asia). Its mammal fauna is quite rich, with some 40 families. Only two of these families are endemic to the Palaearctic region the blind mole rats (Spalacidae) and the Seleviniidae, represented by one species, the dzhalman, which is a small insectivorous rodent. The Nearctic region encompasses nearly all the New World north of tropical Mexico. Its fauna is diverse and includes families with a largely tropical distribution, such as the sac-winged or sheath-tailed bats (Emballonuridae), vampire bats (Desmodontidae), and javelinas or peccaries (Tayassuidae), and largely boreal families, such as the jumping mice (Zapodidae), beavers (Castoridae), and bears (Ursidae). Only two Nearctic families are endemic to the region: the Aplodontidae, which contains one species, the mountain beaver or sewellel, and the Antilocapridae, which also contains one species, the pronghorn antelope. Two other families are almost endemic: the pocket gophers (Geomyidae) live in North America, Central America and northern Colombia; and the kangaroo rats and pocket mice (Heteromyidae) live in North America, Mexico, Central America and northwestern South America. The Neotropical region covers all the New World south of tropical Mexico. It boasts some 27 endemic families of mammals: the solenodons (Solenodontidae), the recently extinct West Indian shrews (Nesophontidae), New World monkeys (Cebidae), marmosets (Callithricidae), caeonolestids or marsupial mice (Caenolestidae), the monito del monte or monkey of the mountains (Microbiotheriidae), anteaters (Myrmecophagidae), sloths (Bradypodidae), and 12 caviomorph rodent families. The rodent families are the degus, coruros, and rock rats (Octodontidae), tuco2

tucos (Ctenomyidae), spiny rats (Echimyidae), rat chinchillas (Abrocomidae), hutias and coypus (Capromyidae), chinchillas and viscachas (Chinchillidae), agouties (Dasyproctidae), pacas (Cuniculidae), the pacarana (Dinomyidae), guinea-pigs and their relatives (Caviidae), capybaras (Hydrochoeridae), and the recently extinct quemi and its allies (Heptaxodontidae). The other seven endemic Neotropical families are bats bulldog bats (Noctilionidae), New World leaf-nosed bats (Phyllostomidae), moustached bats, ghost-faced bats and naked-backed bats (Mormoopidae), vampire bats (Desmondontidae, which some authorities include with the Phyllostomidae), funnel-eared bats (Natalidae), smoky or thumbless bats (Furipteridae) and disc-winged bats (Thyropteridae). The Ethiopian region encompasses Madagascar, Africa south of a somewhat indeterminate line running across the Sahara, and a southern strip of the Arabian peninsula. It has about 15 endemic families, almost as many as the Neotropical region. The families are the giraes (Giradae), hippopotamuses (Hippopotamidae, though those living on the Lower Nile are technically in the Palaearctic region), the aardvark (Orycteropodidae), tenrecs (Tenrecidae), the Old World sucker-footed bats (Myzopodidae), lemurs (Lemuridae), woolly lemurs (Indriidae), aye-ayes (Daubentoniidae), two families of shrew, and ve families of rodent. The shrew families are the golden moles (Chrysochloridae) and otter shrews (Potamogalidae). The rodent families are the scaly-tailed squirrels (Anomaluridae), the spring hare or Cape jumping hare (Pedetidae), cane rats (Thryonomydiae), the rock rat or dassie rat (Petromyidae), and African mole rats (Bathyergidae). Two other families the elephant shrews (Macroscelididae) and gundis (Ctenodactylidae) are conned to Africa but range into the north of the continent, which is part of the Palaearctic region. The Oriental region covers India, Indo-China, southern China, Malaysia, the Philippines, and Indonesian islands as far east as Wallaces line. It has just four endemic families: spiny dormice (Platacanthomyidae), tree shrews (Tupaiidae), tarsiers (Tarsiidae), and ying lemurs or colugos (Cynocephalidae). It also has one endemic bat family, the Craseonycteridae, represented by a single species known as Kittis hog-nosed bat or bumblebee bat, which was discovered in Thailand in 1973. The Australian region includes mainland Australia, Tasmania, New Guinea, Sulawesi, and many small Indonesian islands. It possesses some 19 endemic families of mammals: the echidnas or spiny anteaters (Tachyglossidae), the platypus (Ornithorhynchidae), marsupial mice and cats (Dasyuridae), the Tasmanian wolf (Thylacinidae), the numbat or banded anteater (Myrmecobiidae), the marsupial mole (Notoryctidae), bandicoots and bilbies (Peramelidae), burrowing bandicoots (Thylacomyidae), spiny bandicoot and mouse bandicoot (Peroryctidae), striped possum, Leadbeaters possum and wrist-winged gliders (Petauridae), feathertail gliders (Acrobatidae),

ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net

Biogeographical Regions

pigmy possums (Burramyidae), brush-tailed possums, cuscuses, scaly-tailed possums (Phalangeridae), ringtail possums and great glider (Pseudocheiridae), kangaroos and wallabies (Macropodidae), rat kangaroos, potoroos, and bettongs (Potoroidae), koalas (Phascolarctidae), wombats (Vombatidae), and the noolbender or honey possum (Tarsipedidae).
23

1 2b 4 3 9 10 28 30 25 26 27 29 16 14 31 37 34 12 13 15 33 32 21 35 11 17 18 19 22

2a 7a 8 20 24 7b 6 5

Floral Regions
In The Geography of the Flowering Plants (1974), British botanist Ronald Good summarized the distribution of living angiosperms by adapting a scheme devised by Adolf Engler during the 1870s. Good delineated six major oral regions, though he styled them kingdoms: the Boreal region, the Palaeotropical region, the Neotropical region, the Australian region, South African (Cape) region and the Antarctic oral region. Each of these comprises a number of subregions (Good called them regions), of which there are 37 in total (Figure 2). The Boreal oral region spans North America and Asia, which share many families, including the birches, alders, hazels and hornbeams (Betulaceae), mustard (Cruciferae), primrose (Primulaceae) and buttercup (Ranunculaceae). Six subregions are recognized: the Arctic and Subarctic, East Asia, Western and Central Asia, the Mediterranean, Euro-Siberia and North America. The Palaeotropical region covers most of Africa, the Arabian peninsula, India, southeast Asia, and parts of the western and central Pacic. The subregions are not rmly agreed, but Malesia, Indo-Africa, and Polynesia are commonly recognized. The Malesian subregion is exceptionally rich in forms, with about 400 endemic genera. Madagascar, which is part of the Indo-African subregion but sometimes taken as a separate region, has 12 endemic families and 350 endemic genera. The Neotropical region covers most of South America, save the southern tip and a southwestern strip, Central America, Mexico (excepting the dry northern and central sections), and the West Indies and southern extremity of Florida. It is gloriously rich oristically, housing 47 endemic families and nearly 3000 endemic genera. The Cape region of South Africa is, for its small size, rich in plants, with 11 endemic families and 500 endemic genera. The Australian region is highly distinct with 19 endemic families, 500 endemic genera, and over 6000 oweringplant species. The Antarctic region has a curious geography and includes a coastal strip of Chile and the southern tip of South America, the Antarctic and subantarctic islands, and New Zealand. The subantarctic subregion (southern Chile, Patagonia and New Zealand) carries a distinctive ora involving some 50 genera, of which the southern beech (Nothofagus) is a characteristic element.

36 36

1 2

3 4 5 6 7

Boreal region Arctic and Sub-arctic EuroSiberian a. Europe b. Asia SinoJapanese W. and C. Asiatic Mediterranean Macaronesian Atlantic North American a. Northern b. Southern Pacific North American Palaeotropical region AfricanIndian Desert Sudanese Park Steppe N. E. African Highland W. African Rainforest E. African Steppe South African Madagascar Ascension and St. Helena Indian Continental S. E. Asiatic Malaysian Hawaiian New Caledonia Melanesia and Micronesia Polynesia

24 25 26 27 28 29 30

Neotropical region Caribbean Venezuela and Guiana Amazon South Brazilian Andrean Pampas Juan Fernandez

31

South African region Cape

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23

32 33 34

Australian region N. and E. Australian S. W. Australian C. Australian

35 36 37

Antarctic region New Zealand Patagonian S. Temp. Oceanic Islands

Figure 2 The six floral regions and 37 subregions mapped by Good.

Comparisons and Contrasts between Taxa

The worlds regional faunas are linked with each other in complex ways, as are the worlds regional oras. Connections at the species level are weak, except between the Palaearctic and Nearctic regions, but some regions share genera and families. Each biogeographical region possesses two groups of families: those that are endemic or peculiar to the region, and those that are shared with other regions. Although no agreed system of naming shared taxa (species, genera, families, or whatever) exists, a useful
3

ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net

Biogeographical Regions

scheme suggests that taxa shared between two biogeographical regions are characteristic, taxa shared between three or four biogeographical regions are semi-cosmopolitan, and taxa shared between ve or more biogeographical regions are cosmopolitan. Links between regions are suggested by a mixing of some faunal or oral elements. A Malesian oral element is present in the tropical rainforests of northeastern Queensland, Australia. Antarctic and Palaeotropical ora interdigitate in South Island of New Zealand, Tasmania and the Australian Alps. The strong anity of the Ethiopian and Oriental faunal regions is reected in a number of shared families: bamboo rats (Rhizomyidae), elephants (Elephantidae), rhinoceroses (Rhinocerotidae), chevrotains (Tragulidae), lorises and pottos (Lorisidae), galagos or bushbabies (Galagonidae), apes (Pongidae), and pangolins or scaly anteaters (Manidae).

A new look at mammal regions

The similarities and dierences of dierent biogeographical regions are brought out clearly by applying modern methods of numerical classication to mammal distributions. By applying multidimensional scaling to data on the distribution of 115 mammal families (wholly marine families and the human family were omitted) in Wallaces 24 subregions, Charles H. Smith delineated similar regions to those in the SclaterWallace scheme, but signicant dierences emerged. In Smiths 1983 system, there are four regions Holarctic, Latin American, Afro-Tethyan and Island and 10 subregions (Figure 3). The Holarctic region comprises the Nearctic and the Palaearctic subregions; the Latin American region comprises the Neotropical and Argentine subregions; the Afro-Tethyan region comprises the Mediterranean, Ethiopian and Oriental subregions; and the Island region comprises the Australian, the West

Indian and Madagascan subregions. Each subregion is as unique as it can be compared with all other subregions. Several features of Smiths system are intriguing. First, it reveals a close similitude between the mammal families of the Ethiopian and Oriental regions. Second, it includes the Mediterranean subregion within the Ethiopian region, thus excluding it from the Palaearctic region. Third, it promotes Madagascar and the West Indies to distinct island subregions, removing them from the Ethiopian region and the Neotropical region, respectively. The regional richness and endemicity of mammal families in Smiths regions and subregions are as follows: the Holarctic has 36 families, of which six (17%) are endemic; the Latin American region has 48 families, of which 20 (42%) are endemic; the Afro-Tethyan region has 65 families, of which 29 (45%) are endemic; and the Island region has 35 families, of which 15 (43%) are endemic. Of the 115 mammal families used in the analysis, 43 (37%) are endemic to subregions. The lowest subregional endemicity occurs in the Palaearctic subregion, with no endemic families, and the highest in the Neotropical subregion, with nine endemic families. Smiths analysis also indicated that the Nearctic, Palaearctic, Mediterranean and Oriental subregions have high anities with the faunas of other subregions, whereas the Argentine and Australian subregions have low anities with the faunas of other subregions. Furthermore, the eects of isolation or inaccessibility (or both) are reected in the nature of the Neotropical, Argentine, Ethiopian, Australian, West Indian and Madagascan faunas.

Faunal and floral regions compared

The major oral regions and the major faunal regions are roughly congruent, but there are important dierences between them. First, owing to the superior dispersal ability of some plants compared with terrestrial mammals, the oral regions tend to be less sharply dened than do the faunal regions. Second, although the boreal oral region is equivalent to the combined Palaearctic and Nearctic faunal regions (the Holarctic region), the North American oral subregion diers from the Nearctic faunal region in that it does not occupy all of Florida or Baja California. The Palaeotropical oral region is equivalent to the combined Ethiopian and Oriental faunal regions or a large part of Smiths Afro-Tethyan region, excluding the Mediterranean, which is oristically grouped with the Boreal region. The Australian oral region approximately corresponds with the Australian faunal region, though the dividing line with the Asian region lies between Australia and New Guinea, rather than farther west as in the case of animals. Indeed, it is puzzling that the ora of New Guinea is Palaeotropical while its fauna is Australian. The Neotropical oral region broadly matches the Neotropical faunal region, but the oral Neotropical region, unlike the

Palaearctic Nearctic West Indian Neotropical Mediterranean

Ethiopian Australian Madagascan

Argentine

Holarctic

Latin American

AfroTethyan

Island

Figure 3 The four faunal regions and 10 subregions recognized by Smith.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net

Biogeographical Regions

faunal Neotropical region, takes in Baja California and the southern end of Florida. The Cape oral region, which occupies the southern tip of Africa, bears no equivalent faunal region. The Antarctic oral region, which, like the Cape oral region, possesses no faunal counterpart, includes southern South America and New Zealand, and some of its members are found in Tasmania and southeastern Australia.

Huxleys modification of Wallaces line

20N

South China Sea 10N

Luzon

Pacific Ocean

Borneo

Transitional Zones and Filters

The chief faunal and oral regions are separated from one another by various kinds of barriers determined mainly by climate, mountains and water gaps. The Nearctic is separated from the Palaearctic by two water gaps the Bering Strait and the Norwegian Sea, both of which experience cold climates. A narrow land-link (the Isthmus of Panama), which replaced an earlier water gap, separates the Nearctic region from the Neotropical region, with arid conditions lying north of the land link in Mexico. The Sahara Desert divides the Palaearctic region from the Ethiopian region. The Ethiopian region is insulated from the Oriental region by arid lands in southwest Asia and the Arabian peninsula. The Himalayas and their eastward extensions create a formidable barrier between the Oriental region and the Palaearctic region. In the region sometimes called Wallacea, a series of water gaps hinders movement between the Oriental region and the Australian region. The borders between biogeographical regions may be crossed with varying levels of ease or diculty. Seldom do the environmental conditions in the border areas allow unhampered access between regions. A fairly open border once existed between Alaska and Siberia when, during the Pleistocene epoch, there was a dry-land connection across what is now the Bering Strait. Other borders tend to act as lters and prevent the passage of some species from one biogeographical region to another. In many cases, the border area is transitional as the fauna or ora of one biogeographical region intermixes with the fauna or ora of an adjacent biogeographical region. Two cases will illustrate these points.

Sumatra Indian Ocean 10S 100E

Wallaces line Webers line Wallacea Lydekkers 0 line Celebes New Guinea

Java 120E

140E

Figure 4 Wallacea the transition zone between the Oriental and Australian faunal regions.

genetically distinct from their relatives in the Oriental region. A very few Oriental species, all of which might have been introduced, occur on the islands as far east as Timor, but no Oriental species live beyond that point. Lydekkers line, which passes between the Australian mainland and Timor and between New Guinea and Seram and Halmahera, follows the edge of Australias continental shelf (the Sahul Shelf). It marks the westernmost limit of a wholly Australian fauna. A few Australian species are found on some small islands a little to the west, and as far west as Sulawesi and Lombok. Webers line (Figure 4) runs west of the Moluccas and east of Timor, and marks places with an equal mix of Oriental and Australian species. It is taken by some authorities as the dividing line between the Oriental and Australian faunas. However, the search for a hardand-fast dividing line in such a patently transitional region seems pointless.

The Isthmus of Panama

South America is presently connected to North America, but for most of the last 65 million years or so it was an island-continent. Once during that time, from about 40 to 36 million years ago, a land connection with North America, probably through a chain of islands, existed. Two groups of mammal primates and ancestors of the caviomorph rodents took advantage of the connection and invaded South America. Having arrived in South America, both groups underwent an impressive adaptive radiation to produce the great variety of rodents and New World monkeys found in South America today. From 30 million to 6 million years ago, South America remained a colossal island and mammals had no possibility of interaction with other faunal regions. Even as recently as
5

Wallacea
The famous zoogeographical transition zone between Lydekkers line and Wallaces line is sometimes called Wallacea (Figure 4). It is a large area in which Oriental and the Australian faunas grade into one another. The faunas of both these regions thin out across the transition zone. Wallaces line, which passes between Bali and Lombok and along the Makassar Strait between Borneo and Sulawesi, marks the easternmost extension of a wholly Oriental fauna. A few Oriental species (shrews, civets, pigs, deer and monkeys) have colonized Sulawesi and Bali, but they are

ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net

Biogeographical Regions

6 million years ago, the Bolivar Trough connected the Caribbean Sea with the Pacic Ocean and deterred the passage of animals. However, at that time, members of two families of mammals the eld mice (Cricetidae) and racoons, cacomistles, coati-mundis, kinkajous and olingos (Procyonidae) rafted across the seaway on clumps of soil and vegetation. By 3 million years ago, a land connection the Panamanian land bridge had developed that supplied a gateway for faunal interchange between North and South America. A ood of mammals simply walked into South America. Members of many families were involved: Cervidae (deer), camels (Camelidae), peccaries (Tayassuidae), tapirs (Tapiridae), horses (Equidae), mastodons (Gomphotheriidae), rabbits (Leporidae), squirrels (Sciuridae), shrews (Soricidae), mice (Muridae), dogs (Canidae), bears (Ursidae), weasels (Mustelidae) and cats (Felidae). The passage was two-way and is known as the Great American Interchange.

realization that entire communities need conserving, and not only fashionable species like the tiger and orang-utan.

Human introductions
Humans are watering down the distinctiveness of biogeographical regions by the introduction of alien species: they are homogenizing the global fauna and ora. Take the case of New Zealand. Fifty-four mammal species have been introduced to the island. Twenty came directly or indirectly from Britain and Europe, 14 from Australia, 10 from the Americas, six from Asia, two from Polynesia and two from Africa. The package contained domestic animals for farming and household pets and feral animals for sport or fur production. Farm animals included sheep, cattle and horses. Domestic animals included cats and dogs. Sporting animals included pheasant, deer, wallabies and rabbits. The Australian possum was introduced to start a fur industry. Wild boars and goats were liberated on New Zealand by Captain James Cook. Many other species were introduced European blackbirds, thrushes, sparrows, rooks, yellow hammers, chanches, budgerigars, hedgehogs, hares, weasels, stoats, ferrets, rats and mice. Of course, natural invasions of alien species do take place, but not, it would seem, at the human-induced rates prevalent over the last couple of centuries. Introduced species commonly have an adverse eect upon native species. The Indian mongoose (Herpestes auropunctatus), introduced to various islands worldwide in the hope of controlling rats and other vertebrate pests, has led to the extinction of several native bird and reptile populations. Cats and rats introduced to islands have also tended to have an inimical eect on native wildlife. The inadvertent introduction of the sac fungus, Cryphonectria (Endothia) parasitica, into the United States around 1900 led within 50 years to the near elimination of the American chestnut (Castanea dentata) from the native eastern hardwood forests.

The Applied Use of Biogeographical Regions: Their Place in Conservation

Each biogeographical region contains a combination of species, genera and families, many of which are endemic. Each has a distinctive character that, without conservation measures, stands to be greatly diminished or even lost. Natural biogeographical regions are threatened by human activities, and in particular by habitat destruction and fragmentation and by the introduction of alien species.

Habitat destruction and fragmentation

The human species has transformed the globe to such an extent that only fragments, admittedly some large, of original fauna and ora remain in most biogeographical regions. Natural habitats are conserved in wildlife reserves, where eorts are made to preserve the indigenous faunas and oras. Threatened species and communities stand an even better chance of survival if the wildlife reserves are linked by corridors. By the mid-1980s, 13 western North American wildlife parks had lost 43% of their historical lagomorph (rabbits, hares and pikas), carnivore and ungulate species. But the KootenayBanJasperYoho park system, which embodied signicant connections between wildlife reserves, maintained all its original mammal fauna. The old idea that species could be preserved in zoos is no longer seen as a workable option. Zoos have their place in conservation, for example enabling the reintroduction of near-extinct species to the wild, but much conservation eort now goes into protecting species in the surviving fragments of natural habitats. There is also a growing
6

Summary
The worlds terrestrial animals and plants are grouped into faunal and oral regions. Six faunal regions are recognized traditionally, though a modern scheme, constructed using a numerical classication technique, identies four regions and 10 subregions. Six oral regions and 37 oral regions are commonly distinguished. The oral and faunal regions bear broad agreement with one another but display important dierences of detail. The natural faunas and oras of biogeographical regions are unique. They are under a severe threat from habitat destruction, habitat fragmentation, and the introduction of new species by humans. Their long-term survival depends upon local and regional conservation schemes.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net

Biogeographical Regions

Further Reading
lez-Guzma n LI and Brown JH (1998) Bo hning-Gaese K, Gonza Constraints on dispersal and evolution of the avifauna of the Northern Hemisphere. Evolutionary Ecology 12: 767783. Brown JH and Lomolino MV (1998) Biogeography, 2nd edn. Sunderland, MA: Sinauer Associates. Cox CB and Moore PD (1993) Biogeography: An Ecological and Evolutionary Approach, 5th edn. Oxford: Blackwell. Feldhamer GA, Drickamer LC, Vessey SH and Merritt JF (1999) Mammalogy. Adaptation, Diversity, and Ecology. Boston, MA: WCB McGraw-Hill. Good R (1974) The Geography of the Flowering Plants, 4th edn. London: Longman. Huggett RJ (1998) Fundamentals of Biogeography. London: Routledge. Michaux B (1994) Land movements and animal distributions in east Wallacea (eastern Indonesia, Papua New Guinea and Melanesia). Palaeogeography, Palaeoclimatology, Palaeoecology 112: 323343. Newmark WD (1987) A land-bridge island perspective on mammalian extinctions in western North American parks. Nature 325: 430432.

Sclater PL (1858) On the general distribution of the members of the class Aves. The Journal of the Linnean Society of London: Zoology 2: 130145. Smith CH (1983) A system of world mammal faunal regions. I. Logical and statistical derivation of the regions. Journal of Biogeography 10: 455466. Smith CH (1983) A system of world mammal faunal regions. II. The distance decay eect upon inter-regional anities. Journal of Biogeography 10: 467482. Vane-Wright RI (1991) Transcending the Wallace Line: do the western edges of the Australian region and the Australian plate coincide? Australian Systematic Botany 4: 183197. Wallace AR (1876) The Geographical Distribution of Animals; with a Study of the Relations of Living and Extinct Faunas as Elucidating the Past Changes of the Earths Surface, 2 vols. London: Macmillan. Wilson DE and Reeder DM (1993) Mammal Species of the World: A Taxonomic and Geographic Reference, 2nd edn. Washington and London: Smithsonian Institution Press in association with the American Society of Mammalogists.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net