7
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. P. L. Sclater, A. R. Wallace and the Foundational Units of Biogeography Different places harbour different kinds of animals and plants. The fauna of Africa is unlike the fauna of North America; the flora of Japan is unlike the flora of South Africa. These regional differences in the distribution of species became increasingly manifest as the world was explored. George Leclerc, Compte de Buffon (1707–1788) 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 identified 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 modified this scheme in 1835, by taking account of the ‘five recorded varieties of humans’, to give five 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 Australia and New Guinea. The New World he divided into North America and South America. Sclater’s schema prompted a flurry of papers by English-speaking zoolo- gists, including Thomas Henry Huxley and Joel Asaph Allen, each of whom promulgated his own favoured geographical classification. In his The Geographical Dis- tribution of Animals (1876), Wallace reviewed the compet- ing systems, arguing persuasively in favour of adopting Sclater’s six regions, or realms as Wallace dubbed them. Sclater’s system and Wallace’s 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 identified 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). Wallace’s contribution was to identify subregions, four per region, which correspond largely to de Candolle’s botanical regions. Indeed, the nineteenth-century classification of Article Contents Introductory article . 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 . Summary Nearctic Neotropical Palaearctic Ethiopian Oriental Australian Figure 1 The six faunal regions delimited by Sclater and Wallace. 1 ENCYCLOPEDIA OF LIFE SCIENCES © 2002, John Wiley & Sons, Ltd. www.els.net

eLS || Biogeographical Regions

  • Upload
    richard

  • View
    227

  • Download
    0

Embed Size (px)

Citation preview

Page 1: eLS || Biogeographical Regions

Biogeographical RegionsRichard 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.

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

Different places harbour different kinds of animals andplants. The fauna of Africa is unlike the fauna of NorthAmerica; the flora of Japan is unlike the flora of SouthAfrica. These regional differences in the distribution ofspecies became increasingly manifest as the world wasexplored. George Leclerc, Compte de Buffon (1707–1788)studied the then known tropical mammals from the OldWorld (Africa) and the New World (Central and SouthAmerica). He found that they had not a single species incommon. Later comparisons of African and SouthAmerican plants, insects and reptiles evinced the samepattern.

By the nineteenth century, it was clear that the landsurface couldbedivided intobiogeographical regions, eachofwhich carries a distinct set of animals and adistinct set ofplants. Augustin-Pyramus de Candolle considered plantsand identified areas of endemism, that is botanical regions,each possessing a certain number of plants peculiar tothem. He listed 20 such botanical regions or areas ofendemism in 1820, and by 1838 had added another score,bringing the total to 40. In 1826, James Cowles Prichard, azoologist, distinguished seven regions of mammals: theArctic region, the temperate zone, the equatorial regions,the Indian isles, the Papuan region, the Australian region,and the extremities of America and Africa. WilliamSwainson modified this scheme in 1835, by taking accountof the ‘five recorded varieties of humans’, to give fiveregions: the European (or Caucasian) region, the Asiatic(orMongolian) region, theAmerican region, theEthiopian(or African) region, and the Australian (or Malay) region.

The early ideas of Prichard and Swainson on animaldistributions were eclipsed by the seminal work of anEnglish ornithologist, Philip Lutley Sclater, and theeminent English biogeographer and naturalist, AlfredRussel Wallace. Using bird distributions, Sclater (1858)recognized two basic divisions (or ‘creations’, as he termedthem) – the Old World (Creatio Paleogeana) and the NewWorld (Creatio Neogeana) – and six regions. The OldWorld he divided into Europe and northern Asia, Africasouth of the Sahara, India and southern Asia, and

Australia and New Guinea. The New World he dividedinto North America and South America. Sclater’s schemaprompted a flurry of papers by English-speaking zoolo-gists, including Thomas Henry Huxley and Joel AsaphAllen, each of whom promulgated his own favouredgeographical classification. In his The Geographical Dis-tribution of Animals (1876), Wallace reviewed the compet-ing systems, arguing persuasively in favour of adoptingSclater’s six regions, or realms as Wallace dubbed them.Sclater’s system and Wallace’s minor amendments to itprovided a nomenclature that survives today (Figure 1).Later suggestions were minor variations on the Sclater–Wallace theme. Sclater andWallace identified six regions –Nearctic, Neotropical, Palaearctic, Ethiopian, Orientaland Australian. Together, the Nearctic and Palaearcticregions form Neogaea (the New World), while otherregions form Palaeogaea (the Old World). Wallace’scontribution was to identify subregions, four per region,which correspond largely to de Candolle’s botanicalregions. Indeed, the nineteenth-century classification of

Article Contents

Introductory article

. 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

. Summary

Nearctic Neotropical Palaearctic

Ethiopian Oriental Australian

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

1ENCYCLOPEDIA OF LIFE SCIENCES © 2002, John Wiley & Sons, Ltd. www.els.net

Page 2: eLS || Biogeographical Regions

biogeographical regions was essentially an attempt togroup areas of endemism into a hierarchical classificationaccording to the strengths of their relationships.

It is surprising and noteworthy that the distributions ofspecies with good dispersal abilities, including plants,insects and birds, tend to fall within traditional zoogeo-graphical regional confines. The avifaunas of NorthAmerica and Europe contain several families and manygenera that are not shared by the two regions, even thoughdispersal across the North Atlantic and Pacific Oceans by‘accidental visitors’ is noted every year. Even long-distancemigrant bird taxa tend to be confined either to the easternhemisphere or to the western hemisphere, where theymigrate between high and low latitudes, and appear ill-disposed to disperse east–west between continents.

Mammals

Of the six faunal regions delineated bySclater andWallace,the Palaearctic is the largest. It includes Europe, NorthAfrica, theNear East andmuch ofAsia (but not the Indiansubcontinent or Southeast Asia). Its mammal fauna isquite rich, with some 40 families. Only twoof these familiesare endemic to the Palaearctic region – the blind mole rats(Spalacidae) and the Seleviniidae, represented by onespecies, the dzhalman, which is a small insectivorousrodent.

The Nearctic region encompasses nearly all the NewWorld north of tropical Mexico. Its fauna is diverse andincludes families with a largely tropical distribution, suchas the sac-winged or sheath-tailed bats (Emballonuridae),vampire bats (Desmodontidae), and javelinas or peccaries(Tayassuidae), and largely boreal families, such as thejumping mice (Zapodidae), beavers (Castoridae), andbears (Ursidae). Only two Nearctic families are endemicto the region: the Aplodontidae, which contains onespecies, the mountain beaver or sewellel, and the Antilo-capridae, which also contains one species, the pronghornantelope. Two other families are almost endemic: thepocket gophers (Geomyidae) live in North America,Central America and northern Colombia; and the kangar-oo rats and pocket mice (Heteromyidae) live in NorthAmerica, Mexico, Central America and northwesternSouth America.

The Neotropical region covers all the NewWorld southof tropical Mexico. It boasts some 27 endemic families ofmammals: the solenodons (Solenodontidae), the recentlyextinct West Indian shrews (Nesophontidae), New Worldmonkeys (Cebidae), marmosets (Callithricidae), caeono-lestids or marsupial mice (Caenolestidae), the monito delmonte or ‘monkey of the mountains’ (Microbiotheriidae),anteaters (Myrmecophagidae), sloths (Bradypodidae),and 12 caviomorph rodent families. The rodent familiesare the degus, coruros, and rock rats (Octodontidae), tuco-

tucos (Ctenomyidae), spiny rats (Echimyidae), rat chinch-illas (Abrocomidae), hutias and coypus (Capromyidae),chinchillas and viscachas (Chinchillidae), agouties (Dasy-proctidae), pacas (Cuniculidae), the pacarana (Dinomyi-dae), guinea-pigs and their relatives (Caviidae), capybaras(Hydrochoeridae), and the recently extinct quemi and itsallies (Heptaxodontidae). The other seven endemic Neo-tropical families are bats – bulldog bats (Noctilionidae),NewWorld leaf-nosed bats (Phyllostomidae), moustachedbats, ghost-faced bats and naked-backed bats (Mormoo-pidae), vampire bats (Desmondontidae, which someauthorities include with the Phyllostomidae), funnel-earedbats (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 theSahara, and a southern strip of the Arabian peninsula. Ithas about 15 endemic families, almost as many as theNeotropical region. The families are the giraffes (Giraffi-dae), hippopotamuses (Hippopotamidae, though thoseliving on the Lower Nile are technically in the Palaearcticregion), the aardvark (Orycteropodidae), tenrecs (Tenre-cidae), the Old World sucker-footed bats (Myzopodidae),lemurs (Lemuridae), woolly lemurs (Indriidae), aye-ayes(Daubentoniidae), two families of shrew, and five familiesof rodent. The shrew families are the golden moles(Chrysochloridae) and otter shrews (Potamogalidae).The rodent families are the scaly-tailed squirrels (Anom-aluridae), the spring hare or Cape jumping hare (Pedeti-dae), cane rats (Thryonomydiae), the rock rat or dassie rat(Petromyidae), andAfricanmole rats (Bathyergidae). Twoother families – the elephant shrews (Macroscelididae) andgundis (Ctenodactylidae) – are confined to Africa butrange into the north of the continent, which is part of thePalaearctic region.The Oriental region covers India, Indo-China, southern

China, Malaysia, the Philippines, and Indonesian islandsas far east as Wallace’s line. It has just four endemicfamilies: spiny dormice (Platacanthomyidae), tree shrews(Tupaiidae), tarsiers (Tarsiidae), and flying lemurs orcolugos (Cynocephalidae). It also has one endemic batfamily, the Craseonycteridae, represented by a singlespecies known as Kitti’s hog-nosed bat or bumblebeebat, which was discovered in Thailand in 1973.The Australian region includes mainland Australia,

Tasmania, New Guinea, Sulawesi, and many smallIndonesian islands. It possesses some 19 endemic familiesof mammals: the echidnas or spiny anteaters (Tachyglos-sidae), the platypus (Ornithorhynchidae),marsupial ‘mice’and ‘cats’ (Dasyuridae), the Tasmanian wolf (Thylacini-dae), the numbat or banded anteater (Myrmecobiidae), themarsupial mole (Notoryctidae), bandicoots and bilbies(Peramelidae), burrowing bandicoots (Thylacomyidae),spiny bandicoot and mouse bandicoot (Peroryctidae),striped possum, Leadbeater’s possum and wrist-wingedgliders (Petauridae), feathertail gliders (Acrobatidae),

Biogeographical Regions

2

Page 3: eLS || Biogeographical Regions

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

Floral Regions

In The Geography of the Flowering Plants (1974), Britishbotanist Ronald Good summarized the distribution ofliving angiosperms by adapting a scheme devised by AdolfEngler during the 1870s. Good delineated six major floralregions, though he styled them ‘kingdoms’: the Borealregion, the Palaeotropical region, the Neotropical region,theAustralian region, SouthAfrican (Cape) region and theAntarctic floral region. Each of these comprises a numberof subregions (Good called them regions), of which thereare 37 in total (Figure 2). The Boreal floral region spansNorth America and Asia, which share many families,including the birches, alders, hazels and hornbeams(Betulaceae), mustard (Cruciferae), primrose (Primula-ceae) and buttercup (Ranunculaceae). Six subregions arerecognized: the Arctic and Subarctic, East Asia, Westernand Central Asia, the Mediterranean, Euro-Siberia andNorth America. The Palaeotropical region covers most ofAfrica, the Arabian peninsula, India, southeast Asia, andparts of the western and central Pacific. The subregions arenot firmly agreed, butMalesia, Indo-Africa, and Polynesiaare commonly recognized. The Malesian subregion isexceptionally rich in forms, with about 400 endemicgenera. Madagascar, which is part of the Indo-Africansubregion but sometimes taken as a separate region, has 12endemic families and 350 endemic genera. TheNeotropicalregion coversmost of SouthAmerica, save the southern tipand a southwestern strip, Central America, Mexico(excepting the dry northern and central sections), and theWest Indies and southern extremity of Florida. It isgloriously rich floristically, housing 47 endemic familiesand nearly 3000 endemic genera. The Cape region ofSouth Africa is, for its small size, rich in plants, with11 endemic families and 500 endemic genera. TheAustralian region is highly distinct with 19 endemicfamilies, 500 endemic genera, and over 6000 flowering-plant species. The Antarctic region has a curiousgeography and includes a coastal strip of Chile and thesouthern tip of South America, the Antarctic andsubantarctic islands, and New Zealand. The subantarcticsubregion (southern Chile, Patagonia and New Zealand)carries a distinctive flora involving some 50 genera, ofwhich the southern beech (Nothofagus) is a characteristicelement.

Comparisons and Contrasts betweenTaxa

The world’s regional faunas are linked with each other incomplex ways, as are the world’s regional floras. Connec-tions at the species level are weak, except between thePalaearctic and Nearctic regions, but some regions sharegenera and families. Each biogeographical region pos-sesses two groups of families: those that are endemic orpeculiar to the region, and those that are shared with otherregions. Although no agreed system of naming shared taxa(species, genera, families, or whatever) exists, a useful

36

Boreal region1

2

3

4

5

6

7

8

Arctic and Sub-arcticEuro–Siberiana. Europeb. AsiaSino–JapaneseW. and C. AsiaticMediterraneanMacaronesianAtlantic North Americana. Northernb. SouthernPacific North American

Palaeotropical region9

10

11

12

13

14

15

16

17

18

19

20

21

22

23

African–Indian DesertSudanese Park SteppeN. E. African HighlandW. African RainforestE. African SteppeSouth AfricanMadagascarAscension and St. HelenaIndianContinental S. E. AsiaticMalaysianHawaiianNew CaledoniaMelanesia and MicronesiaPolynesia

Neotropical region24

25

26

27

28

29

30

CaribbeanVenezuela and GuianaAmazonSouth BrazilianAndreanPampasJuan Fernandez

Australian region32

33

34

N. and E. AustralianS. W. AustralianC. Australian

South African region31 Cape

Antarctic region35

36

37

New ZealandPatagonianS. Temp.Oceanic Islands

23

20

30

37

282526

27

29

36

24

8 7b

7a

1

16

31

1413 15

12

10

6

11

9

5

2a2b

43

1817

3433

32 35

21

2219

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

Biogeographical Regions

3

Page 4: eLS || Biogeographical Regions

scheme suggests that taxa shared between two biogeogra-phical regions are ‘characteristic’, taxa shared betweenthree or four biogeographical regions are ‘semi-cosmopo-litan’, and taxa shared between five or more biogeogra-phical regions are ‘cosmopolitan’. Links between regionsare suggested by amixing of some faunal or floral elements.A Malesian floral element is present in the tropicalrainforests of northeastern Queensland, Australia. Ant-arctic andPalaeotropical flora interdigitate in South Islandof New Zealand, Tasmania and the Australian Alps. Thestrong affinity of the Ethiopian andOriental faunal regionsis reflected in a number of shared families: bamboo rats(Rhizomyidae), elephants (Elephantidae), rhinoceroses(Rhinocerotidae), chevrotains (Tragulidae), lorises andpottos (Lorisidae), galagos or bushbabies (Galagonidae),apes (Pongidae), and pangolins or scaly anteaters (Man-idae).

A new look at mammal regions

The similarities and differences of different biogeographi-cal regions are brought out clearly by applying modernmethods of numerical classification to mammal distribu-tions. By applying multidimensional scaling to data on thedistribution of 115 mammal families (wholly marinefamilies and the human family were omitted) in Wallace’s24 subregions, Charles H. Smith delineated similar regionsto those in the Sclater–Wallace scheme, but significantdifferences emerged. In Smith’s 1983 system, there are fourregions – Holarctic, Latin American, Afro-Tethyan andIsland – and 10 subregions (Figure 3). The Holarctic regioncomprises the Nearctic and the Palaearctic subregions; theLatin American region comprises the Neotropical andArgentine subregions; the Afro-Tethyan region comprisesthe Mediterranean, Ethiopian and Oriental subregions;and the Island region comprises the Australian, the West

Indian and Madagascan subregions. Each subregion is asunique as it can be compared with all other subregions.Several features of Smith’s system are intriguing. First, itreveals a close similitude between the mammal families ofthe Ethiopian and Oriental regions. Second, it includes theMediterranean subregion within the Ethiopian region,thus excluding it from the Palaearctic region. Third, itpromotes Madagascar and the West Indies to distinctisland subregions, removing them from the Ethiopianregion and the Neotropical region, respectively.The regional richness and endemicity of mammal

families in Smith’s regions and subregions are as follows:the Holarctic has 36 families, of which six (17%) areendemic; the Latin American region has 48 families, ofwhich 20 (42%) are endemic; the Afro-Tethyan region has65 families, of which 29 (45%) are endemic; and the Islandregion has 35 families, of which 15 (43%) are endemic. Ofthe 115mammal families used in the analysis, 43 (37%) areendemic to subregions. The lowest subregional endemicityoccurs in the Palaearctic subregion, with no endemicfamilies, and the highest in theNeotropical subregion,withnine endemic families. Smith’s analysis also indicated thatthe Nearctic, Palaearctic, Mediterranean and Orientalsubregions have high affinities with the faunas of othersubregions, whereas the Argentine and Australian sub-regions have low affinities with the faunas of othersubregions. Furthermore, the effects of isolation orinaccessibility (or both) are reflected in the nature of theNeotropical, Argentine, Ethiopian, Australian, WestIndian and Madagascan faunas.

Faunal and floral regions compared

The major floral regions and the major faunal regions areroughly congruent, but there are important differencesbetween them. First, owing to the superior dispersal abilityof some plants compared with terrestrial mammals, thefloral regions tend to be less sharply defined than do thefaunal regions. Second, although the boreal floral region isequivalent to the combined Palaearctic and Nearcticfaunal regions (the Holarctic region), the North Americanfloral subregion differs from the Nearctic faunal region inthat it does not occupy all of Florida or Baja California.The Palaeotropical floral region is equivalent to thecombined Ethiopian andOriental faunal regions or a largepart of Smith’s Afro-Tethyan region, excluding theMediterranean, which is floristically grouped with theBoreal region. The Australian floral region approximatelycorresponds with the Australian faunal region, though thedividing line with the Asian region lies between Australiaand New Guinea, rather than farther west as in the case ofanimals. Indeed, it is puzzling that the flora ofNewGuineais Palaeotropical while its fauna is Australian. TheNeotropical floral region broadlymatches theNeotropicalfaunal region, but the floral Neotropical region, unlike the

Holarctic LatinAmerican

Afro-Tethyan Island

Nearctic

WestIndian

Neotropical

Argentine

Mediterranean

Ethiopian

Palaearctic

AustralianMadagascan

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

Biogeographical Regions

4

Page 5: eLS || Biogeographical Regions

faunalNeotropical region, takes in BajaCalifornia and thesouthern end of Florida. The Cape floral region, whichoccupies the southern tip of Africa, bears no equivalentfaunal region. The Antarctic floral region, which, like theCape floral region, possesses no faunal counterpart,includes southern South America and New Zealand, andsome of its members are found in Tasmania and south-eastern Australia.

Transitional Zones and Filters

The chief faunal and floral regions are separated from oneanother by various kinds of barriers determined mainly byclimate, mountains and water gaps. The Nearctic isseparated from the Palaearctic by two water gaps – theBering Strait and the Norwegian Sea, both of whichexperience cold climates. A narrow land-link (the Isthmusof Panama), which replaced an earlier water gap, separatesthe Nearctic region from the Neotropical region, with aridconditions lying north of the land link in Mexico. TheSahara Desert divides the Palaearctic region from theEthiopian region. The Ethiopian region is insulated fromthe Oriental region by arid lands in southwest Asia and theArabian peninsula. The Himalayas and their eastwardextensions create a formidable barrier between theOrientalregion and the Palaearctic region. In the region sometimescalled Wallacea, a series of water gaps hinders movementbetween the Oriental region and the Australian region.

The borders between biogeographical regions may becrossed with varying levels of ease or difficulty. Seldom dothe environmental conditions in the border areas allowunhampered access between regions. A fairly open borderonce existed between Alaska and Siberia when, during thePleistocene epoch, there was a dry-land connection acrosswhat is now the Bering Strait. Other borders tend to act asfilters and prevent the passage of some species from onebiogeographical region to another. In many cases, theborder area is transitional as the fauna or flora of onebiogeographical region intermixes with the fauna or floraof an adjacent biogeographical region. Two cases willillustrate these points.

Wallacea

The famous zoogeographical transition zone betweenLydekker’s line and Wallace’s line is sometimes calledWallacea (Figure 4). It is a large area in which Oriental andthe Australian faunas grade into one another. The faunasof both these regions thin out across the transition zone.Wallace’s line, which passes betweenBali andLombok andalong the Makassar Strait between Borneo and Sulawesi,marks the easternmost extension of a wholly Orientalfauna.A fewOriental species (shrews, civets, pigs, deer andmonkeys) have colonized Sulawesi and Bali, but they are

genetically distinct from their relatives in the Orientalregion.A very fewOriental species, all of whichmight havebeen introduced, occur on the islands as far east as Timor,but no Oriental species live beyond that point. Lydekker’sline, which passes between the Australian mainland andTimor and between New Guinea and Seram and Halma-hera, follows the edge of Australia’s continental shelf (theSahul Shelf). It marks the westernmost limit of a whollyAustralian fauna. A few Australian species are found onsome small islands a little to the west, and as far west asSulawesi and Lombok. Weber’s line (Figure 4) runs west oftheMoluccas and east of Timor, and marks places with anequal mix of Oriental and Australian species. It is taken bysome authorities as the dividing line between the Orientaland Australian faunas. However, the search for a hard-and-fast dividing line in such a patently transitional regionseems 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 anisland-continent. Once during that time, from about 40 to36 million years ago, a land connection with NorthAmerica, probably through a chain of islands, existed.Two groups of mammal – primates and ancestors of thecaviomorph rodents – took advantage of the connectionand invaded South America. Having arrived in SouthAmerica, both groups underwent an impressive adaptiveradiation to produce the great variety of rodents and NewWorld monkeys found in South America today. From 30million to 6 million years ago, South America remained acolossal island and mammals had no possibility ofinteraction with other faunal regions. Even as recently as

10°N

10°S

Sumatra

Indian Ocean

20°N

Borneo

Java

CelebesNew

Guinea

Luzon

Huxley’s modificationof Wallace’s line

Wallace’s line

Weber’s lineWallacea

Lydekker’sline

SouthChina Sea

PacificOcean

100°E 120°E 140°E

Figure 4 Wallacea – the transition zone between the Oriental andAustralian faunal regions.

Biogeographical Regions

5

Page 6: eLS || Biogeographical Regions

6 million years ago, the Bolivar Trough connected theCaribbean Sea with the Pacific Ocean and deterred thepassage of animals.However, at that time,members of twofamilies of mammals – the ‘field mice’ (Cricetidae) andracoons, cacomistles, coati-mundis, kinkajous and olingos(Procyonidae) – rafted across the seaway on clumps of soiland vegetation. By 3million years ago, a land connection –the Panamanian land bridge – had developed that supplieda gateway for faunal interchangebetweenNorth andSouthAmerica. A flood of mammals simply walked into SouthAmerica. Members of many families were involved:Cervidae (deer), camels (Camelidae), peccaries (Tayassui-dae), tapirs (Tapiridae), horses (Equidae), mastodons(Gomphotheriidae), rabbits (Leporidae), squirrels (Sciur-idae), shrews (Soricidae), mice (Muridae), dogs (Canidae),bears (Ursidae), weasels (Mustelidae) and cats (Felidae).The passage was two-way and is known as the GreatAmerican Interchange.

The Applied Use of BiogeographicalRegions: Their Place in Conservation

Each biogeographical region contains a combinationof species, genera and families, many of which areendemic. Each has a distinctive character that, withoutconservation measures, stands to be greatly diminished oreven lost. Natural biogeographical regions are threatenedby human activities, and in particular by habitat destruc-tion and fragmentation and by the introduction of alienspecies.

Habitat destruction and fragmentation

The human species has transformed the globe to such anextent that only fragments, admittedly some large, oforiginal fauna and flora remain in most biogeographicalregions.Natural habitats are conserved inwildlife reserves,where efforts are made to preserve the indigenous faunasand floras. Threatened species and communities stand aneven better chance of survival if the wildlife reserves arelinked by corridors. By the mid-1980s, 13 western NorthAmerican wildlife parks had lost 43% of their historicallagomorph (rabbits, hares and pikas), carnivore andungulate species. But the Kootenay–Banff–Jasper–Yohopark system, which embodied significant connectionsbetween wildlife reserves, maintained all its originalmammal fauna.

The old idea that species could be preserved in zoos is nolonger seen as a workable option. Zoos have their place inconservation, for example enabling the reintroduction ofnear-extinct species to the wild, but much conservationeffort now goes into protecting species in the survivingfragments of natural habitats. There is also a growing

realization that entire communities need conserving, andnot only ‘fashionable’ species like the tiger and orang-utan.

Human introductions

Humans are watering down the distinctiveness of biogeo-graphical regions by the introduction of alien species: theyare homogenizing the global fauna and flora. Take the caseof New Zealand. Fifty-four mammal species have beenintroduced to the island.Twenty camedirectly or indirectlyfrom Britain and Europe, 14 from Australia, 10 from theAmericas, six fromAsia, two fromPolynesia and two fromAfrica. The package contained domestic animals forfarming and household pets and feral animals for sportor fur production. Farmanimals included sheep, cattle andhorses. Domestic animals included cats and dogs. Sportinganimals included pheasant, deer, wallabies and rabbits.The Australian possum was introduced to start a furindustry. Wild boars and goats were liberated on NewZealand by Captain James Cook.Many other species wereintroduced – European blackbirds, thrushes, sparrows,rooks, yellow hammers, chaffinches, budgerigars, hedge-hogs, hares, weasels, stoats, ferrets, rats and mice. Ofcourse, natural invasions of alien species do take place, butnot, it would seem, at the human-induced rates prevalentover the last couple of centuries.Introduced species commonly have an adverse effect

upon native species. The Indian mongoose (Herpestesauropunctatus), introduced to various islands worldwide inthe hope of controlling rats and other vertebrate pests, hasled to the extinction of several native bird and reptilepopulations. Cats and rats introduced to islands have alsotended to have an inimical effect on native wildlife. Theinadvertent introduction of the sac fungus, Cryphonectria(Endothia) parasitica, into the United States around 1900led within 50 years to the near elimination of the Americanchestnut (Castanea dentata) from the native easternhardwood forests.

Summary

Theworld’s terrestrial animals and plants are grouped intofaunal and floral regions. Six faunal regions are recognizedtraditionally, though a modern scheme, constructed usinga numerical classification technique, identifies four regionsand 10 subregions. Six floral regions and 37 floral regionsare commonly distinguished. The floral and faunal regionsbear broad agreement with one another but displayimportant differences of detail. The natural faunas andfloras of biogeographical regions are unique. They areunder a severe threat from habitat destruction, habitatfragmentation, and the introduction of new species byhumans. Their long-term survival depends upon local andregional conservation schemes.

Biogeographical Regions

6

Page 7: eLS || Biogeographical Regions

Further Reading

Bohning-Gaese K, Gonzalez-Guzman LI and Brown JH (1998)

Constraints on dispersal and evolution of the avifaunaof theNorthern

Hemisphere. Evolutionary Ecology 12: 767–783.

Brown JH and Lomolino MV (1998) Biogeography, 2nd edn. Sunder-

land, 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.

GoodR (1974)TheGeography 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: 323–343.

Newmark WD (1987) A land-bridge island perspective on mammalian

extinctions in western North American parks. Nature 325: 430–432.

Sclater PL (1858) On the general distribution of the members of the

class Aves. The Journal of the Linnean Society of London: Zoology 2:

130–145.

Smith CH (1983) A system of world mammal faunal regions. I. Logical

and statistical derivation of the regions. Journal of Biogeography 10:

455–466.

Smith CH (1983) A system of world mammal faunal regions. II. The

distance decay effect upon inter-regional affinities. Journal of

Biogeography 10: 467–482.

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: 183–197.

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 Earth’s 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.

Biogeographical Regions

7