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ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net 1
Biogeographical Regions
biogeographical regions was essentially an attempt to tucos (Ctenomyidae), spiny rats (Echimyidae), rat chinch-
group areas of endemism into a hierarchical classification illas (Abrocomidae), hutias and coypus (Capromyidae),
according to the strengths of their relationships. chinchillas and viscachas (Chinchillidae), agouties (Dasy-
It is surprising and noteworthy that the distributions of proctidae), pacas (Cuniculidae), the pacarana (Dinomyi-
species with good dispersal abilities, including plants, dae), guinea-pigs and their relatives (Caviidae), capybaras
insects and birds, tend to fall within traditional zoogeo- (Hydrochoeridae), and the recently extinct quemi and its
graphical regional confines. The avifaunas of North allies (Heptaxodontidae). The other seven endemic Neo-
America and Europe contain several families and many tropical families are bats – bulldog bats (Noctilionidae),
genera that are not shared by the two regions, even though New World leaf-nosed bats (Phyllostomidae), moustached
dispersal across the North Atlantic and Pacific Oceans by bats, ghost-faced bats and naked-backed bats (Mormoo-
‘accidental visitors’ is noted every year. Even long-distance pidae), vampire bats (Desmondontidae, which some
migrant bird taxa tend to be confined either to the eastern authorities include with the Phyllostomidae), funnel-eared
hemisphere or to the western hemisphere, where they bats (Natalidae), smoky or thumbless bats (Furipteridae)
migrate between high and low latitudes, and appear ill- and disc-winged bats (Thyropteridae).
disposed to disperse east–west between continents. 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
Mammals Neotropical region. The families are the giraffes (Giraffi-
dae), hippopotamuses (Hippopotamidae, though those
Of the six faunal regions delineated by Sclater and Wallace, living on the Lower Nile are technically in the Palaearctic
the Palaearctic is the largest. It includes Europe, North region), the aardvark (Orycteropodidae), tenrecs (Tenre-
Africa, the Near East and much of Asia (but not the Indian cidae), the Old World sucker-footed bats (Myzopodidae),
subcontinent or Southeast Asia). Its mammal fauna is lemurs (Lemuridae), woolly lemurs (Indriidae), aye-ayes
quite rich, with some 40 families. Only two of these families (Daubentoniidae), two families of shrew, and five families
are endemic to the Palaearctic region – the blind mole rats of rodent. The shrew families are the golden moles
(Spalacidae) and the Seleviniidae, represented by one (Chrysochloridae) and otter shrews (Potamogalidae).
species, the dzhalman, which is a small insectivorous The rodent families are the scaly-tailed squirrels (Anom-
rodent. aluridae), the spring hare or Cape jumping hare (Pedeti-
The Nearctic region encompasses nearly all the New dae), cane rats (Thryonomydiae), the rock rat or dassie rat
World north of tropical Mexico. Its fauna is diverse and (Petromyidae), and African mole rats (Bathyergidae). Two
includes families with a largely tropical distribution, such other families – the elephant shrews (Macroscelididae) and
as the sac-winged or sheath-tailed bats (Emballonuridae), gundis (Ctenodactylidae) – are confined to Africa but
vampire bats (Desmodontidae), and javelinas or peccaries range into the north of the continent, which is part of the
(Tayassuidae), and largely boreal families, such as the Palaearctic region.
jumping mice (Zapodidae), beavers (Castoridae), and The Oriental region covers India, Indo-China, southern
bears (Ursidae). Only two Nearctic families are endemic China, Malaysia, the Philippines, and Indonesian islands
to the region: the Aplodontidae, which contains one as far east as Wallace’s line. It has just four endemic
species, the mountain beaver or sewellel, and the Antilo- families: spiny dormice (Platacanthomyidae), tree shrews
capridae, which also contains one species, the pronghorn (Tupaiidae), tarsiers (Tarsiidae), and flying lemurs or
antelope. Two other families are almost endemic: the colugos (Cynocephalidae). It also has one endemic bat
pocket gophers (Geomyidae) live in North America, family, the Craseonycteridae, represented by a single
Central America and northern Colombia; and the kangar- species known as Kitti’s hog-nosed bat or bumblebee
oo rats and pocket mice (Heteromyidae) live in North bat, which was discovered in Thailand in 1973.
America, Mexico, Central America and northwestern The Australian region includes mainland Australia,
South America. Tasmania, New Guinea, Sulawesi, and many small
The Neotropical region covers all the New World south Indonesian islands. It possesses some 19 endemic families
of tropical Mexico. It boasts some 27 endemic families of of mammals: the echidnas or spiny anteaters (Tachyglos-
mammals: the solenodons (Solenodontidae), the recently sidae), the platypus (Ornithorhynchidae), marsupial ‘mice’
extinct West Indian shrews (Nesophontidae), New World and ‘cats’ (Dasyuridae), the Tasmanian wolf (Thylacini-
monkeys (Cebidae), marmosets (Callithricidae), caeono- dae), the numbat or banded anteater (Myrmecobiidae), the
lestids or marsupial mice (Caenolestidae), the monito del marsupial mole (Notoryctidae), bandicoots and bilbies
monte or ‘monkey of the mountains’ (Microbiotheriidae), (Peramelidae), burrowing bandicoots (Thylacomyidae),
anteaters (Myrmecophagidae), sloths (Bradypodidae), spiny bandicoot and mouse bandicoot (Peroryctidae),
and 12 caviomorph rodent families. The rodent families striped possum, Leadbeater’s possum and wrist-winged
are the degus, coruros, and rock rats (Octodontidae), tuco- gliders (Petauridae), feathertail gliders (Acrobatidae),
2 ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net
Biogeographical Regions
ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net 3
Biogeographical Regions
scheme suggests that taxa shared between two biogeogra- Indian and Madagascan subregions. Each subregion is as
phical regions are ‘characteristic’, taxa shared between unique as it can be compared with all other subregions.
three or four biogeographical regions are ‘semi-cosmopo- Several features of Smith’s system are intriguing. First, it
litan’, and taxa shared between five or more biogeogra- reveals a close similitude between the mammal families of
phical regions are ‘cosmopolitan’. Links between regions the Ethiopian and Oriental regions. Second, it includes the
are suggested by a mixing of some faunal or floral elements. Mediterranean subregion within the Ethiopian region,
A Malesian floral element is present in the tropical thus excluding it from the Palaearctic region. Third, it
rainforests of northeastern Queensland, Australia. Ant- promotes Madagascar and the West Indies to distinct
arctic and Palaeotropical flora interdigitate in South Island island subregions, removing them from the Ethiopian
of New Zealand, Tasmania and the Australian Alps. The region and the Neotropical region, respectively.
strong affinity of the Ethiopian and Oriental faunal regions The regional richness and endemicity of mammal
is reflected in a number of shared families: bamboo rats families in Smith’s regions and subregions are as follows:
(Rhizomyidae), elephants (Elephantidae), rhinoceroses the Holarctic has 36 families, of which six (17%) are
(Rhinocerotidae), chevrotains (Tragulidae), lorises and endemic; the Latin American region has 48 families, of
pottos (Lorisidae), galagos or bushbabies (Galagonidae), which 20 (42%) are endemic; the Afro-Tethyan region has
apes (Pongidae), and pangolins or scaly anteaters (Man- 65 families, of which 29 (45%) are endemic; and the Island
idae). 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
A new look at mammal regions occurs in the Palaearctic subregion, with no endemic
families, and the highest in the Neotropical subregion, with
The similarities and differences of different biogeographi- nine endemic families. Smith’s analysis also indicated that
cal regions are brought out clearly by applying modern the Nearctic, Palaearctic, Mediterranean and Oriental
methods of numerical classification to mammal distribu- subregions have high affinities with the faunas of other
tions. By applying multidimensional scaling to data on the subregions, whereas the Argentine and Australian sub-
distribution of 115 mammal families (wholly marine regions have low affinities with the faunas of other
families and the human family were omitted) in Wallace’s subregions. Furthermore, the effects of isolation or
24 subregions, Charles H. Smith delineated similar regions inaccessibility (or both) are reflected in the nature of the
to those in the Sclater–Wallace scheme, but significant Neotropical, Argentine, Ethiopian, Australian, West
differences emerged. In Smith’s 1983 system, there are four Indian and Madagascan faunas.
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
Faunal and floral regions compared
Argentine subregions; the Afro-Tethyan region comprises The major floral regions and the major faunal regions are
the Mediterranean, Ethiopian and Oriental subregions; roughly congruent, but there are important differences
and the Island region comprises the Australian, the West between them. First, owing to the superior dispersal ability
of some plants compared with terrestrial mammals, the
floral regions tend to be less sharply defined than do the
faunal regions. Second, although the boreal floral region is
equivalent to the combined Palaearctic and Nearctic
Palaearctic faunal regions (the Holarctic region), the North American
Nearctic
floral subregion differs from the Nearctic faunal region in
West Mediterranean that it does not occupy all of Florida or Baja California.
Indian The Palaeotropical floral region is equivalent to the
Neotropical
Ethiopian combined Ethiopian and Oriental faunal regions or a large
part of Smith’s Afro-Tethyan region, excluding the
Australian Mediterranean, which is floristically grouped with the
Madagascan Boreal region. The Australian floral region approximately
Argentine
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 flora of New Guinea
Latin Afro- is Palaeotropical while its fauna is Australian. The
Holarctic Island
American Tethyan
Neotropical floral region broadly matches the Neotropical
Figure 3 The four faunal regions and 10 subregions recognized by Smith. faunal region, but the floral Neotropical region, unlike the
4 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 Huxley’s modification
southern end of Florida. The Cape floral region, which of Wallace’s line
20°N
occupies the southern tip of Africa, bears no equivalent
faunal region. The Antarctic floral region, which, like the South Luzon Pacific
Cape floral region, possesses no faunal counterpart, China Sea Ocean
includes southern South America and New Zealand, and
some of its members are found in Tasmania and south- 10°N
eastern Australia. Wallace’s line
Weber’s line
Wallacea
Borneo Lydekker’s
0°
Transitional Zones and Filters Sumatra Celebes
line
New
The chief faunal and floral regions are separated from one Guinea
Indian Ocean Java
another by various kinds of barriers determined mainly by
climate, mountains and water gaps. The Nearctic is 10°S
separated from the Palaearctic by two water gaps – the 100°E 120°E 140°E
Bering Strait and the Norwegian Sea, both of which
experience cold climates. A narrow land-link (the Isthmus Figure 4 Wallacea – the transition zone between the Oriental and
Australian faunal regions.
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 genetically distinct from their relatives in the Oriental
Sahara Desert divides the Palaearctic region from the region. A very few Oriental species, all of which might have
Ethiopian region. The Ethiopian region is insulated from been introduced, occur on the islands as far east as Timor,
the Oriental region by arid lands in southwest Asia and the but no Oriental species live beyond that point. Lydekker’s
Arabian peninsula. The Himalayas and their eastward line, which passes between the Australian mainland and
extensions create a formidable barrier between the Oriental Timor and between New Guinea and Seram and Halma-
region and the Palaearctic region. In the region sometimes hera, follows the edge of Australia’s continental shelf (the
called Wallacea, a series of water gaps hinders movement Sahul Shelf). It marks the westernmost limit of a wholly
between the Oriental region and the Australian region. Australian fauna. A few Australian species are found on
The borders between biogeographical regions may be some small islands a little to the west, and as far west as
crossed with varying levels of ease or difficulty. Seldom do Sulawesi and Lombok. Weber’s line (Figure 4) runs west of
the environmental conditions in the border areas allow the Moluccas and east of Timor, and marks places with an
unhampered access between regions. A fairly open border equal mix of Oriental and Australian species. It is taken by
once existed between Alaska and Siberia when, during the some authorities as the dividing line between the Oriental
Pleistocene epoch, there was a dry-land connection across and Australian faunas. However, the search for a hard-
what is now the Bering Strait. Other borders tend to act as and-fast dividing line in such a patently transitional region
filters and prevent the passage of some species from one seems pointless.
biogeographical region to another. In many cases, the
border area is transitional as the fauna or flora of one
biogeographical region intermixes with the fauna or flora The Isthmus of Panama
of an adjacent biogeographical region. Two cases will
illustrate these points. South America is presently connected to North America,
but for most of the last 65 million years or so it was an
Wallacea island-continent. Once during that time, from about 40 to
36 million years ago, a land connection with North
The famous zoogeographical transition zone between America, probably through a chain of islands, existed.
Lydekker’s line and Wallace’s line is sometimes called Two groups of mammal – primates and ancestors of the
Wallacea (Figure 4). It is a large area in which Oriental and caviomorph rodents – took advantage of the connection
the Australian faunas grade into one another. The faunas and invaded South America. Having arrived in South
of both these regions thin out across the transition zone. America, both groups underwent an impressive adaptive
Wallace’s line, which passes between Bali and Lombok and radiation to produce the great variety of rodents and New
along the Makassar Strait between Borneo and Sulawesi, World monkeys found in South America today. From 30
marks the easternmost extension of a wholly Oriental million to 6 million years ago, South America remained a
fauna. A few Oriental species (shrews, civets, pigs, deer and colossal island and mammals had no possibility of
monkeys) have colonized Sulawesi and Bali, but they are interaction with other faunal regions. Even as recently as
ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net 5
Biogeographical Regions
6 million years ago, the Bolivar Trough connected the realization that entire communities need conserving, and
Caribbean Sea with the Pacific Ocean and deterred the not only ‘fashionable’ species like the tiger and orang-utan.
passage of animals. However, at that time, members of two
families of mammals – the ‘field mice’ (Cricetidae) and Human introductions
racoons, cacomistles, coati-mundis, kinkajous and olingos
(Procyonidae) – rafted across the seaway on clumps of soil Humans are watering down the distinctiveness of biogeo-
and vegetation. By 3 million years ago, a land connection – graphical regions by the introduction of alien species: they
the Panamanian land bridge – had developed that supplied are homogenizing the global fauna and flora. Take the case
a gateway for faunal interchange between North and South of New Zealand. Fifty-four mammal species have been
America. A flood of mammals simply walked into South introduced to the island. Twenty came directly or indirectly
America. Members of many families were involved: from Britain and Europe, 14 from Australia, 10 from the
Cervidae (deer), camels (Camelidae), peccaries (Tayassui- Americas, six from Asia, two from Polynesia and two from
dae), tapirs (Tapiridae), horses (Equidae), mastodons Africa. The package contained domestic animals for
(Gomphotheriidae), rabbits (Leporidae), squirrels (Sciur- farming and household pets and feral animals for sport
idae), shrews (Soricidae), mice (Muridae), dogs (Canidae), or fur production. Farm animals included sheep, cattle and
bears (Ursidae), weasels (Mustelidae) and cats (Felidae). horses. Domestic animals included cats and dogs. Sporting
The passage was two-way and is known as the Great animals included pheasant, deer, wallabies and rabbits.
American Interchange. 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,
The Applied Use of Biogeographical rooks, yellow hammers, chaffinches, budgerigars, hedge-
hogs, hares, weasels, stoats, ferrets, rats and mice. Of
Regions: Their Place in Conservation course, natural invasions of alien species do take place, but
not, it would seem, at the human-induced rates prevalent
Each biogeographical region contains a combination over the last couple of centuries.
of species, genera and families, many of which are Introduced species commonly have an adverse effect
endemic. Each has a distinctive character that, without upon native species. The Indian mongoose (Herpestes
conservation measures, stands to be greatly diminished or auropunctatus), introduced to various islands worldwide in
even lost. Natural biogeographical regions are threatened the hope of controlling rats and other vertebrate pests, has
by human activities, and in particular by habitat destruc- led to the extinction of several native bird and reptile
tion and fragmentation and by the introduction of alien populations. Cats and rats introduced to islands have also
species. tended to have an inimical effect on native wildlife. The
inadvertent introduction of the sac fungus, Cryphonectria
Habitat destruction and fragmentation (Endothia) parasitica, into the United States around 1900
led within 50 years to the near elimination of the American
The human species has transformed the globe to such an chestnut (Castanea dentata) from the native eastern
extent that only fragments, admittedly some large, of hardwood forests.
original fauna and flora remain in most biogeographical
regions. Natural habitats are conserved in wildlife reserves,
where efforts are made to preserve the indigenous faunas
and floras. Threatened species and communities stand an Summary
even better chance of survival if the wildlife reserves are
linked by corridors. By the mid-1980s, 13 western North The world’s terrestrial animals and plants are grouped into
American wildlife parks had lost 43% of their historical faunal and floral regions. Six faunal regions are recognized
lagomorph (rabbits, hares and pikas), carnivore and traditionally, though a modern scheme, constructed using
ungulate species. But the Kootenay–Banff–Jasper–Yoho a numerical classification technique, identifies four regions
park system, which embodied significant connections and 10 subregions. Six floral regions and 37 floral regions
between wildlife reserves, maintained all its original are commonly distinguished. The floral and faunal regions
mammal fauna. bear broad agreement with one another but display
The old idea that species could be preserved in zoos is no important differences of detail. The natural faunas and
longer seen as a workable option. Zoos have their place in floras of biogeographical regions are unique. They are
conservation, for example enabling the reintroduction of under a severe threat from habitat destruction, habitat
near-extinct species to the wild, but much conservation fragmentation, and the introduction of new species by
effort now goes into protecting species in the surviving humans. Their long-term survival depends upon local and
fragments of natural habitats. There is also a growing regional conservation schemes.
6 ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net
Biogeographical Regions
Further Reading Sclater PL (1858) On the general distribution of the members of the
class Aves. The Journal of the Linnean Society of London: Zoology 2:
Böhning-Gaese K, González-Guzmán LI and Brown JH (1998) 130–145.
Constraints on dispersal and evolution of the avifauna of the Northern Smith CH (1983) A system of world mammal faunal regions. I. Logical
Hemisphere. Evolutionary Ecology 12: 767–783. and statistical derivation of the regions. Journal of Biogeography 10:
Brown JH and Lomolino MV (1998) Biogeography, 2nd edn. Sunder- 455–466.
land, MA: Sinauer Associates. Smith CH (1983) A system of world mammal faunal regions. II. The
Cox CB and Moore PD (1993) Biogeography: An Ecological and distance decay effect upon inter-regional affinities. Journal of
Evolutionary Approach, 5th edn. Oxford: Blackwell. Biogeography 10: 467–482.
Feldhamer GA, Drickamer LC, Vessey SH and Merritt JF (1999) Vane-Wright RI (1991) Transcending the Wallace Line: do the western
Mammalogy. Adaptation, Diversity, and Ecology. Boston, MA: WCB edges of the Australian region and the Australian plate coincide?
McGraw-Hill. Australian Systematic Botany 4: 183–197.
Good R (1974) The Geography of the Flowering Plants, 4th edn. London: Wallace AR (1876) The Geographical Distribution of Animals; with
Longman. a Study of the Relations of Living and Extinct Faunas as Elucidating
Huggett RJ (1998) Fundamentals of Biogeography. London: Routledge. the Past Changes of the Earth’s Surface, 2 vols. London:
Michaux B (1994) Land movements and animal distributions in east Macmillan.
Wallacea (eastern Indonesia, Papua New Guinea and Melanesia). Wilson DE and Reeder DM (1993) Mammal Species of the World: A
Palaeogeography, Palaeoclimatology, Palaeoecology 112: 323–343. Taxonomic and Geographic Reference, 2nd edn. Washington and
Newmark WD (1987) A land-bridge island perspective on mammalian London: Smithsonian Institution Press in association with the
extinctions in western North American parks. Nature 325: 430–432. American Society of Mammalogists.
ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net 7