The boundaries of the Palearctic region - British Birds€¦ · establish an objective southern...

602 © British Birds 99 • December 2006 • 602–618

ABSTRACT Although the western and eastern boundaries of the Palearcticregion are widely agreed, there is still much debate about the southernboundary. In an attempt to establish a definitive southern boundary, the

distribution of 1,037 breeding passerine species in the Palearctic, and adjacentparts of the Afrotropical and Oriental regions north of 5°N, were analysed.

The WorldMap computer program was used to combine these maps, to reveal species richness.The variations in the position of the southern

boundary suggested by previous authorities gives rise to three zones: north of the northernmost boundary, which is unequivocally Palearctic; south of

the southernmost boundary, which is unequivocally Afrotropical or Oriental;and a border region in between, which is analysed in detail here to

establish an objective southern boundary for the region.

The boundaries of thePalearctic region

Kees (C. S.) Roselaar

In 1973, during initial meetings of the editorial team of a planned handbook ofbirds in Europe, one of the topics of debate

was the title of the forthcoming work. Severalpeople opted for the title ‘Birds of the WesternPalearctic’ because it was short and geographi-cally clearly defined; others thought that theterm ‘Palearctic’ was too little-known in birdingcircles to attract potential buyers. After long dis-cussion, it was decided that the book should becalled Handbook of the Birds of Europe, theMiddle East and North Africa, and subtitled TheBirds of the Western Palearctic. Despite thesmaller font type given to this subtitle, thehandbook soon became widely known underthe latter name, or simply as BWP. Since then,many birders have become familiar with theterms ‘Palearctic’ and ‘Western Palearctic’,although few will claim to know exactly wherethe boundaries lie.

Although the use of the term ‘Palearctic’ isnow well established, the region is not welldefined. The name was introduced by Sclater(1858), who divided the world into a number ofzoogeographical regions, each characterised byunique faunal components with geographicranges restricted to that region. Sclater based hisregional system on the distribution of passerine

birds, but analyses of other groups of terrestrialanimals by subsequent workers, notably Wallace(1876), supported and expanded his scheme.Many books and checklists have now been pub-lished on the birds of the Palearctic. Theirauthors are more or less agreed on the westernand eastern borders of the region, which stretchfrom Iceland and Morocco in the west, across toJapan, Kamchatka and the Chukotskiy Penin-sula in northeast Russia in the east, but thenumber of bird species considered as Palearcticvaries widely. This is partly due to differences intaxonomic approach; however, even when Dick-inson (2003) is used as the authority on recog-nised species and races, there is wide variationin the total number of taxa published for theregion: the Palearctic handbook of Hartert(Hartert 1903–23; Hartert & Steinbacher1932–38) included 564 breeding passerinespecies; Vaurie (1959) included 620 species; Eck(1996) 635 species; and Beaman (1994) 693species (these counts exclude introduced oraccidental breeders). Eck (2004) presented adetailed overview of the differences among thethree first-mentioned listings. It is clear that the apparent variation in number of specieswith time is explained largely by different views of what constitutes the southern limit

of the Palearctic region.During work on the passerine volume of the

forthcoming Handbook of Geographical Varia-tion and Distribution of Palearctic Birds (Rose-laar & Shirihai in prep.), the need for awell-defined southern border of the Palearcticbecame evident. It became apparent from themany detailed distribution maps that had beenprepared for this work that an analysis of shareddistributions could provide the most accurateassessment of the position of this southernboundary.

Methods and mapsThe southern border of the Palearctic is estab-lished here by comparing separate breeding dis-tribution maps of 1,037 passerine species. TheseA3-sized maps were compiled when working onspecies texts for BWP from 1987 onwards, butalso cover all songbird species which occuroutside the Western Palearctic in Eurasia andAfrica north of 5°N. The maps are not the sameas those used in BWP (which were prepared byEuan Dunn, Dorothy Vincent and MikeWilson) but are, in part, based on the same

sources. For the region outside the WesternPalearctic, the maps are more detailed thanthose in BWP, and more than 4,000 papers andbooks on breeding-bird distribution were usedto compile them. This literature, too extensiveto be listed here, includes the most recentlypublished breeding-bird atlas data for variouscountries in Europe. Europe is, however, only asmall part of the Palearctic, and comparabledata for North Africa, Siberia, China, andvarious central and southern Asian countries isnot available; an extensive literature review,covering the past 150 years, was thus necessaryto establish breeding ranges for these areas. Inaddition, the locations named on thousands ofspecimen labels were checked in several refer-ence collections. The individual species rangemaps based on these sources will be publishedin Roselaar & Shirihai (in prep.).

For analysis, the original maps were con-verted into digital format by the computerprogram WorldMap version 4.1 (Williams2000). In terms of longitude, each grid cell was1° longitude wide, with a total of 220 cells from30°W east to 170°W covering the entire

603British Birds 99 • December 2006 • 602–618

The boundaries of the Palearctic Region

337. Temminck’s Lark Eremophila bilopha,Tagdilt Track, Boumalne du Dades, Morocco, February 2006. OnlyPalearctic passerine species extend throughout the Sahara of North Africa, and no Afrotropical species occur.

Consequently, it is considered that the entire Sahara belongs to the Palearctic region.Throughout this region, thenumber of breeding species is particularly low, but larks (Alaudidae) and wheatears Oenanthe are well

represented. Many species, including Temminck’s Lark, have adapted to survive in this harsh environment byexploiting a particular, specialised niche, which enables them to exist alongside otherwise similar species.

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Palearctic from the Azores to the ChukotskiyPeninsula. In terms of latitude, 120 grid cellscovered an area from 18°N to 86°N. One degreeof longitude in the south of this area is widerthan one degree of longitude in the north(being c. 104 km wide at 18°N, c. 64 km at55°N, and c. 10 km at 85°N). Since grid cells ofequal area were required for this analysis, thelatitudinal grid-cell dimension had to beadapted: cells were c. 39 km ‘high’ at 18°N, c. 64km high at 55°N, and c. 400 km high at 85°N. Inother words, one degree of latitude at 18°Ncovers just over three cells, but one cell coversmore than three degrees of latitude at 85°N.This has the disadvantage that distributionmaps become somewhat distorted, being verti-cally compressed in the north and stretched inthe south. However, the important point is thateach grid cell covers an area of 4,062 km2, whichenables the comparison of actual range size of anorthern breeder with that of a southernbreeder; thus a northern breeding speciesoccurring in 300 grid cells will have a similarbreeding range size to that of a southernbreeder also occupying 300 grid cells.

Species coveredThe breeding distributions of all passerinespecies breeding north of 18°N in Africa,Europe, and Asia were plotted in the WorldMapprogram. The Cape Verde Islands were includedby moving the plots three degrees to the north.No attempt was made to plot distributions inGreenland, the Philippines or Alaska, althoughthese regions are partly visible at the fringe ofthe map. The taxonomy used in this paper isbased on Dickinson (2003); scientific andEnglish names follow the BB ‘List of Birds of theWestern Palearctic’ (www.britishbirds.co.uk/bblist.htm), and Dickinson (scientific names)and Beaman (1994) for other regions. UsingDickinson’s taxonomy, some 1,037 species camewithin the area defined above. However, somespecies have been split since Dickinson waspublished, and the following were acknowl-edged as separate species: Richard’s Pipit Anthusrichardi (northern Asia south to northernChina) and Paddyfield Pipit A. rufulus(southern China southwards); AfricanStonechat Saxicola torquatus (Afrotropics),Common Stonechat S. rubicola (western Europe

and northern Africa eastto the west Caucasusregion) and SiberianStonechat S. maurus(Ural Mountains andCaucasus eastward);Eastern OlivaceousWarbler Hippolais pallida(Sahara and the Balkanseastward) and WesternOlivaceous Warbler H.opaca (Iberian Peninsulaand northwest Africa);African Desert WarblerSylvia deserti (westernSahara) and Asian DesertWarbler S. nana (centralAsian deserts); BrownFlycatcher Muscicapadauurica (northern Asia)and Williamson’s Fly-catcher M. williamsoni(mainland southeastAsia); and Golden OrioleOriolus oriolus (Europeeast to central Siberia)and Indian GoldenOriole O. kundoo (IndianPeninsula north to

604 British Birds 99 • December 2006 • 602–618


338. Thick-billed Lark Rhamphocoris clotbey,Tagdilt Track, Boumalne du Dades,Morocco, February 2006.A number of Saharan specialities, including Thick-billedLark,Temminck’s Lark Eremophila bilopha and Red-rumped Wheatear Oenanthemoesta, are restricted to the northern fringe of the Sahara and the northern

Arabian Peninsula, and show a closer affinity to the Palearctic than to theAfrotropics.Thick-billed Lark is a desert specialist with a nomadic lifestyle, and

numbers in any particular region can vary greatly from year to year as it exploitsthe available food sources.

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central Asia, the TienShan Mountains andnorthwest China) (seeWittmann et al. 1995;Helbig & Seibold 1999;Sangster et al. 1999; Ras-mussen & Anderton 2005and Ottoson et al. 2005for the reasoning behindsome of these splits). Inaddition, seven specieswere omitted becausethey have extensive orlocalised (relict?) rangesin both the Palearctic andthe Afrotropical and/orOriental regions: Ori-ental Lark Alaudagulgula, Red-rumpedSwallow Cecropis daurica,Long-billed Pipit Anthussimilis, Zitting CisticolaCisticola juncidis, Clam-orous Reed WarblerAcrocephalus stentoreus,Large-billed Crow Corvusmacrorhynchos andHouse Sparrow Passerdomesticus. Each of theseseven may potentially besplit (e.g. Rasmussen &Anderton 2005), but thetaxonomic limits in each case are uncertain forthe moment.

Variation in the southern boundary of thePalearcticMany authors are uncertain where the southernlimits of the Palearctic lie; they accept a fairlysouthern boundary, but include only ‘trulyPalearctic’ species, while excluding ‘species ofpredominantly Oriental or Afrotropical genera’(Vaurie 1959), even if the range falls largely orentirely north of their perceived southern limit.In this analysis, a species is assigned to a certainfaunal region if more than half of its breedingrange (established from the number of gridcells occupied) falls within the boundary of thatregion. Beaman (1994) and others also includedas Palearctic those species which have aninsignificant breeding population in the region,but which otherwise occur extensively withinanother faunal region; this premise is notadopted here.

For the Western Palearctic, Hartert(1903–23) accepted a southern limit ‘throughthe Sahara, a little bit more to the south in theNile Valley, but excepting southern Arabiawhich is purely [Afro]tropical’. Other authorsare more precise for the Sahara; Eck (1996)adopted 19°N as the southern border, whileHall & Moreau (1970) and Snow (1978) put thenorthern limit of the Afrotropical region at20°N. Vaurie also accepted a southern limit at c.19°N, but included southward extensions to c.16°N in the Aïr (northern Niger) and Ennedi(northeast Chad) because some Palearctic taxaoccur there (although the vast majority ofbreeding species are Afrotropical). Voous(1973–77) established the southern border inthe western and central Sahara at 21°N, and tothe southern Egyptian border in the east,though with southern extensions to 20°N alongthe Atlantic coastline and at 18°E to include theBanc d’Arguin (Mauritania) and the TibestiMountains in northern Chad. The editorial



339. Long-billed Pipit Anthus similis, Fujairah, United Arab Emirates, December2004. Long-billed Pipit provides a typical example of the difficulties involved inassigning species to a particular zoogeographical region.Although much of the

breeding range lies in both the Afrotropical and Oriental regions, several disjunctpopulations exist along the southern boundary of the Palearctic. Of these, A. s.

captus breeds in Lebanon, Jordan, Syria and Israel, A. s. decaptus breeds in Iraq, Iran,Pakistan and Afghanistan, while A. s. arabicus breeds in the Afrotropical mountains

of the southwest Arabian Peninsula.Whether arabicus or decaptus breeds innorthern Oman and northernmost UAE has not definitely been established, but

the avifauna here is predominantly Palearctic.

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team of BWP adopted the Voous list for tax-onomy and species sequence, and also adoptedVoous’s southern boundary. In contrast,botanists, including Takhtadzhyan (1978) andCox (2001), draw the southern limit of thePalearctic much farther north, along thesouthern fringe of the Mediterranean sub-region, which approximates to the northernfringe of the Sahara. Although all ornithologicalauthorities include the Cape Verde Islandswithin the Palearctic, botanists exclude thisarchipelago (fig. 1).

For the Arabian Peninsula, opinions on thesouthern border differ widely. BWP placed thisfarther north than earlier authorities, at 28°N.In contrast, Vaurie (1959) drew it, rather impre-cisely, ‘south to the region of Mecca in the westand to Oman in the east’, which is approxi-mately 21°N. Many others include the entireArabian Peninsula within the Palearctic,including Voous (1973–77), Dowsett & Forbes-Watson (1993), Beaman (1994) and Eck (1996).Martins & Hirschfeld (1994) included the entirepeninsula, with the exception of the mountainsof southwest Saudi Arabia, Yemen, andsouthern Oman. Botanists have equally variedopinions; Takhtadzhyan (1978) placed theboundary even farther to the north, andexcluded the Syrian Desert from the Palearctic,while Cox (2001) included the entire ArabianPeninsula in the Palearctic.

For Pakistan, ornithologists agree that themountains of the west and north constitute theboundary between the Palearctic and Oriental

regions, and that the Indus Plain lies entirelywithin the Oriental. However, most are unclearabout the lower altitudinal limit for Palearcticbirds, apart from Beaman (1994), who definedthe boundary in Pakistan at 2,000 m. The coastof Baluchistan, in southwest Pakistan, and theneighbouring region of extreme southern orsoutheastern Iran are sometimes included in theOriental region and sometimes in the Palearctic.

To the east, the Himalaya forms an obviousdivision between these two faunal regions butdefining the boundary has again proved contro-versial. Takhtadzhyan (1978) considered thesouthern border of the Palearctic to lie justabove the foothill zone where the subtropicalzone and the lower montane zones meet, at c.2,000 m in Nepal. Martens & Eck (1995) alsoidentified the zone below 2,000 m in Nepal as(sub)tropical, with a temperate zone at2,000–2,900 m (1,700–3,000 m), the latter dom-inated by evergreen oak forests including Sym-plocus, Castanopsis and Quercus spp. at lowerelevations, and mixed evergreen and coniferousforest including Himalayan Hemlock Tsugadumosa and Globe Magnolia Magnolia globosaover 2,600 m. Both Vaurie (1959) and Voous(1973–77) considered those species which occurupwards from the temperate mixed deciduousforest in the Himalaya and central China to bePalearctic. According to Beaman (1994), thistype of forest occurs down to c. 2,000 m in Pak-istan and Kashmir; to c. 2,500 m in HimachalPradesh and Uttar Pradesh, northwest India; toc. 2,800 m from Nepal to Arunachal Pradesh,

northeast India, andnorthern Myanmar; to c. 2,500 m in Yunnanand southern Sichuanprovinces, southwestChina; and to 2,000 min central and northernSichuan Province. Thelower altitudinal limitfor the Palearctic inNepal is thus unclear.The same is true in themountains of centralChina; Schäfer (1938)stated that the Palearcticzone in central SichuanProvince starts abovethe coniferous forest(upper montane zone)at 3,500 m, rather than



Fig. 1. The southern border of the Palearctic, as suggested by various authors,drawn on a biodiversity map of all passerine species breeding in the area of the

map. Line 1: Hartert (1903–23); Line 2: Schäfer (1938); Line 3:Vaurie (1959);Line 4: Hall & Moreau (1970); Line 5: Cramp (1977); Line 6:Takhtadzhyan (1978);

Line 7: Beaman (1994); and Line 8: Eck (1996).The key on the right-hand side of this figure (and subsequent figures) shows number of breeding passerine

species from high (red) to low (blue).

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318 species

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the 2,000 m adhered to by Beaman.Opinions on the southern boundary in

eastern China differ widely too. Takhtadzhyan(1978) considered the flora of eastern China tobe Palearctic, with the exception of the southernlowlands of Guangxi and Guangdongprovinces. Voous (1973–77, 1985) took theYangtze River as the boundary between thePalearctic and Oriental regions, while othershave considered eastern China to be predomi-nantly Oriental, with the Palearctic border at c.33°N (Eck 1996); at 34°N (Beaman 1994);‘south of northern Hubei and Shandong’, i.e. c.32–34°N (Vaurie 1959); or even ‘in the region ofBeijing’, i.e. c. 40°N (Hartert 1903–23).

Hartert (1903–23) considered that Taiwanand the islands to the south of Japan are reintropisch [purely tropical], and thus fall withinthe Oriental region. However, all other authorsincorporate the southern Japanese islands,including the Nansei-shoto (Ryukyu Islands),Kazan-retto (Volcano Islands) and Ogasawara-shoto (Bonin Islands), in the Palearctic, andTaiwan in the Oriental.

The western and eastern borders of thePalearctic are not discussed here: most authorsagree that the Palearctic reaches west to JanMayen, Iceland and the Azores, and east to theChukotskiy Peninsula, the Commander Islands(Komandorskiye Ostrova), and Ogasawara-shoto. Only Vaurie (1959) and Voous (1985)included eastern Greenland, while Vaurie(1965) included the whole of Greenland. In ananalysis based on breeding ranges of passerines,it makes little sense for Greenland to be

included, as the few passerines occurring therehave circumpolar ranges, and are thus sharedbetween the Palearctic and Nearctic.

Unequivocally Palearctic birds and apreliminary southern boundaryThe first stage of the analysis was to establishtwo lines: (Line 1) the northernmost southernborder of the Palearctic ever proposed (thenorthern limit of Takhtadzhyan (1978) in theSahara and Syria, Beaman (1994) in Iran, Pak-istan, and the Himalaya, Schäfer (1938) incentral China, and Hartert (1903–23) in easternChina and southern Japan); and (Line 2) thesouthernmost of all boundaries proposed, fol-lowing Vaurie (1959) for the Sahara, Voous(1973–77), Beaman (1994) and Eck (1996) forthe Arabian Peninsula, Takhtadzhyan (1978)and Martens & Eck (1995) along the foothills ofthe Himalaya at c. 1,700–2,000 m, and thecapricious line of Takhtadzhyan through centralMyanmar and southern China. All authorsagree that birds occurring largely or completelynorth of Line 1 are unequivocally Palearctic,while those occurring south of Line 2 are trulyAfrotropical or Oriental. Fig. 2 shows the com-bined distributions of all Palearctic passerinesbreeding predominantly north of Line 1, whilefig. 3 shows the distributions of Afrotropicaland Oriental species to the south of Line 2.

A parallel can be drawn between these twolines, Line 1 and Line 2, and the lines definingthe boundary between the Oriental and Aus-tralasian regions in Indonesia. Here, theunequivocal eastern boundary of the Oriental

region is formed byWallace’s Line (just eastof Borneo and Bali), theunequivocal westernborder of the Aus-tralasian region isLydekker’s Line, justwest of New Guinea andKepulauan Kai (KaiIslands), while the tran-sition zone betweenthese lines (‘Wallacea’) isdivided halfway byWeber’s Line (just westof the Moluccas) wherethe fauna of bothregions reaches equilib-rium.

Fig. 2 shows that a



Fig. 2. Northernmost suggested border of the southern Palearctic (Line 1),and distribution of all unequivocally Palearctic passerines (breeding

predominantly north of the line).This line follows southern border of Takhtadzhyan (1978) in the west, Beaman (1994) in the centre, and

Schäfer (1938) and Hartert (1903–23) in the east.

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few widespread Palearctic species also occurwidely throughout the Sahara and ArabianPeninsula (Crested Lark Galerida cristata,Eastern Olivaceous Warbler and Southern GreyShrike Lanius meridionalis) while one, RufousBush Robin Cercotrichas galactotes, has a sepa-rate population in the Sahel zone. However,even these few species breed in more grid cellsto the north of Line 1 than to the south; thesame applies to a few species which are wide-spread in the plains of the Indian Peninsula(Crested Lark, Southern Grey Shrike, Great TitParus major) and 22 species in the plains ofeastern China (e.g. Barn Swallow Hirundorustica, White Wagtail Motacilla alba, WrenTroglodytes troglodytes, Brown Dipper Cincluspallasii, Siberian Stonechat, Blue Rock ThrushMonticola solitarius, Blackbird Turdus merula,Great Tit, Coal Tit Periparus ater, EurasianNuthatch Sitta europaea, Brown Shrike L.cristatus, Eurasian Jay Garrulus glandarius,Magpie Pica pica, Tree Sparrow Passermontanus, Chestnut-eared Bunting Emberizafucata and Meadow Bunting E. cioides). In addi-tion, Taiwan has 29 truly Palearctic species(including Barn Swallow, Asian House MartinDelichon dasypus, White Wagtail, Grey WagtailMotacilla cinerea, Wren, Alpine AccentorPrunella collaris, Brown Dipper, Varied Tit Parusvarius, Coal Tit, Eurasian Jay, and NutcrackerNucifraga caryocatactes).

Conversely, some Afrotropical species breedto the north of Line 2: Plain Martin Ripariapaludicola, Common Bulbul Pycnonotus bar-

batus and Black-crowned Tchagra Tchagra sene-galus are Afrotropical species breeding in thePalearctic only in northwest Africa. Othersextend well north to reach southern Algeria(e.g. Red-billed Firefinch Lagonosticta senegala,perhaps aided by recent introductions), or intosouthern Egypt via the Nile valley (e.g. AfricanPied Wagtail M. aguimp) (fig. 3). ManyAfrotropical passerines also extend into south-west Arabia, and some reach north into theLevant region (e.g. Black Bush Robin Cer-cotrichas podobe, Blackstart Cercomela melanuraand Fan-tailed Raven Corvus rhipidurus).

The number of Oriental species extendinginto the Palearctic is even greater, with severalranging from the Indian subcontinent west ornorthwest to Iraq and southern Turkey (e.g.White-cheeked Bulbul P. leucogenys, CommonBabbler Turdoides caudata and Yellow-throatedSparrow Petronia xanthocollis). Other Orientalspecies have skirted around the western flank ofthe Tien Shan Mountains and pushed northinto central Asia, with some reaching southernor even central Kazakhstan (e.g. Pied StonechatSaxicola caprata, Asian Paradise-flycatcher Terp-siphone paradisi, Indian Golden Oriole, Long-tailed Shrike Lanius schach and Common MynaAcridotheres tristis, although the recent rangeextension of the last species in this region isprobably due to introductions). To the east,Oriental species also extend into the Palearctic,in northeast China (e.g. Black Dicrurus macro-cercus, Ashy D. leucophaeus and Hair-crestedDrongo D. hottentottus and Red-billed Blue

Magpie Urocissa ery-throrhyncha), with someeven reaching theRussian Far East (e.g.Black-naped Oriole O.chinensis and Asian Par-a d i s e - f l y c a t c h e r ) .Despite their northerlydistributions, even thosespecies which extendfarthest north occupymore grid cells in theOriental region than inthe Palearctic. Forexample, Indian GoldenOriole occupies 788 gridcells in the IndianPeninsula (Orientalregion) and 502 in thePalearctic section of its



Fig. 3. Southernmost suggested border of the southern Palearctic (Line 2),and distribution of all unequivocally Afrotropical and Oriental passerines

(breeding predominantly south of the line).This line follows southern border ofVaurie (1959) in the Sahara; Voous (1973–77), Beaman (1994) and Eck (1996) in

Arabia; and Takhtadzhyan (1978) further east. The vertical blue line through Arabia represents the boundary between the Afrotropical and Oriental

regions based on the distribution of passerine breeding species.

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range, which extends from the Tien ShanMountains east into the Xinjiang UygurAutonomous Region in westernmost China.

By comparing the number of Palearctic andAfrotropical/Oriental species in each grid cell infigs. 2 and 3, a line of equilibrium between theseregions becomes apparent (fig. 4). To the northof this line Palearctic species predominate, tothe south of it Afrotropical and Oriental speciespredominate. This line roughly follows a lati-tude of 19°N in the Afrotropics, lies close to theboundary followed by BWP in Arabia, toBeaman’s (1994) line through western andcentral Asia, and to Voous’s (1973–77) lineacross eastern China. However, this preliminaryline was based on birds breeding largely to thenorth of Line 1 and south of Line 2, and thusexcludes the species occurring predominantlybetween these lines in the Sahara, the Arabian

Peninsula, the Himalaya between 2,000 and2,800 m, central China between 2,000 and 3,500m, and throughout the lowland plains and hillsof eastern China. The species in each of theseregions are discussed below, before an attemptis made to establish a definitive southern limitto the Palearctic.

Saharan speciesEighteen species occur primarily in the Sahara,mainly larks (Alaudidae) and wheatearsOenanthe, together with Pale Crag Martin Pty-onoprogne obsoleta, African Desert Warbler,Fulvous Babbler Turdoides fulva, Brown-neckedRaven C. ruficollis, Desert Sparrow Passersimplex, Trumpeter Finch Bucanetes githagineusand House Bunting Emberiza striolata (fig. 5).Since only truly Palearctic species extendthroughout the Sahara (fig. 2) and no Afrotrop-

ical species occur here(fig. 3), it is consideredthat the Sahara in itsentirety belongs to thePalearctic. Furthermore,approximately half of allSaharan species extendinto the Palearcticregions of Iran and/orKazakhstan, while of thefour species extendingto the Oriental region inthe deserts of Pakistanor northwest India, twoalso breed north toKazakhstan. In addition,a number of species arerestricted to thenorthern fringe of theSahara and the northernArabian Peninsula, thusnearer to the Palearcticthan to the Afrotropics(e.g. Thick-billed LarkRhamphocoris clotbey,Temminck’s Lark Ere-mophila bilopha andRed-rumped WheatearO. moesta). The onlyspecies restricted to thesouthern fringe of theSahara is Black-crownedSparrow-lark Ere-mopterix nigriceps; thisand the widespread



Fig. 5. Distribution of species with a breeding range falling predominantlybetween Line 1 and Line 2 in the Sahara and Cape Verde Islands.

For definitions of these Lines, see figs. 2 & 3.

Fig. 4. Preliminary southern border of the Palearctic, separating unequivocallyPalearctic and unequivocally Afrotropical and Oriental passerine species, based onnumber of species in each grid cell as counted from figs. 2 & 3 (line drawn where

number for each region is in equilibrium). Map and southern boundary ignorethose species breeding predominantly between Line 1 and Line 2.

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340. White-throated Redstart Phoenicurus schisticeps, Jiuzhaigou, SichuanProvince, China, July 2006. Redstarts reach their greatest diversity in the

mountains bordering the southern fringe of the Palearctic in the Himalaya andwestern/central China, where they occupy a wide range of habitats.White-

throated Redstart typically breeds in scrub forest on rocky slopes and alongforest edge above 3,200 m from Nepal to Bhutan, and in western China.

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341. Indian Blue Robin Luscinia brunnea, Jamboti, Karnataka, India,April 2006.Although the stunning Indian Blue Robin is a common and widespread

summer visitor to the Himalaya and mountains of western/central China, it is secretive, usually keeps close to the ground, and sings from deep within

cover. Like many species in this region, it breeds in temperate forest between 2,400 and 3,200 m, in and above the zone of overlap between

the Oriental and Palearctic regions.

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House Bunting are the onlySaharan species whichextend exclusively to north-west India. These two areperhaps the only Saharanspecies of southern origin,because other Eremopterixspecies are restricted to theAfrotropical and the Ori-ental regions, while therelationships of HouseBunting appear to be withthe Afrotropical buntingssuch as Cinnamon-breastedEmberiza tahapisi and CapeBunting E. capensis. Clearly,the Saharan passerine faunais predominantly Palearctic.Alternatively, if a faunalregion is defined as an areawhich has more endemicspecies than species sharedwith other regions, theSahara and Syrian desertsmay form a valid zoogeo-graphical unit, the‘Eremian’ region, because18 passerines are endemicto the Sahara and Syrian(semi-)deserts, sharingtheir range with just threePalearctic and no Afrotrop-ical species.

The Cape Verde Islandsbelong to the Palearcticbecause, of the ten breedingpasserines there, Palearcticspecies predominate. Ofthese ten, three are sharedwith the Palearctic (Specta-cled Warbler Sylvia conspic-illata, Blackcap S. atricapillaand Spanish Sparrow Passerhispaniolensis); four areSaharan (and thus alsoPalearctic – Bar-tailedDesert Lark Ammomanescinctura, Hoopoe LarkAlaemon alaudipes, Brown-necked Raven and Black-crowned Sparrow-lark);and three are endemic. Ofthe last group, Raso LarkAlauda razae seems likely to

have Palearctic affinities, while the other two aremore likely to be Afrotropical (Cape VerdeWarbler Acrocephalus brevipennis is morpholog-ically close to Greater Swamp Warbler A.rufescens, and Cape Verde Sparrow P. iagoensisto an African sparrow group including RufousSparrow P. motitensis).

Arabian speciesThe overwhelming majority of the breeding birdsof south and southwest Arabia are shared withthe Afrotropics; 24 of the 70 Afrotropical speciesfalling within the geographical boundaries usedin this analysis also occur in southwest Arabia(see fig. 3), while of the Palearctic species onlyMagpie extends to the same area (fig. 2). Conse-quently, all ten endemic Arabian species (fig. 6),which are found in the southwest of the penin-sula, are considered to be Afrotropical. This isreinforced by examining the relationships ofthese ten species: eight (White-spectacled BulbulPycnonotus xanthopygos, Yemen Thrush T. men-achensis, Yemen Parisoma Parisoma buryi,Arabian Babbler T. squamiceps, Tristram’s StarlingOnychognathus tristramii, Arabian WaxbillEstrilda rufibarba, Olive-rumped Serin Serinusrothschildi and Yemen Serin S. menachensis) showa close morphological resemblance to relatedAfrotropical or, in some cases, Oriental species.Only two endemics appear to be of Palearcticorigin – Yemen Accentor Prunella fagani (geo-graphically isolated from a group of accentors ofsimilar appearance, including Radde’s Accentor P.ocularis of the mountains of Turkey and northernIran) and Yemeni Linnet Carduelis yemensis(related to Linnet C. cannabina). No passerinesare endemic to the interior deserts and coastal

plains of Arabia. Since the species occurringthroughout these areas of the Arabian Peninsulaare shared with the Sahara, the lowlands are bestconsidered Palearctic, and the mountains of thesouthwest as Afrotropical.

Mid-altitude species of the Himalaya andcentral ChinaFor the zoogeographical affinities of species inthe Himalaya, an analysis was made of speciesoccurring in Nepal, based on the altitudinal dis-tribution of breeding records supplied byMartens & Eck (1995), supplemented by datafrom Grimmett et al. (1998), the latter mainlyfor species occurring at lower elevations (a zoneless well covered by Martens & Eck). A total of104 Nepalese bird species which are widespreadin the plains and hills of the Indian Peninsulaare considered as Oriental; these are augmentedby a few species endemic to the Himalayanfoothills (if occurring entirely below 2,000 m),and by 69 Oriental species from the plains andhills of mainland southeast Asia, many of whichextend westwards through the eastern Himalaya,and occur in Nepal mainly between 1,000 and2,000 m. Palearctic taxa are represented in Nepalby 65 species which are widespread in thePalearctic north of 35°N; these occur mostlyabove 2,800 m, but some are restricted to thelowlands or foothills (see fig. 2). These are aug-mented by 70 endemic species of the southernand eastern fringes of the Tibetan Plateau, whichoccur predominantly above 2,800 m in Nepal,the altitudinal lower boundary of the Palearcticof Beaman (1994). The distribution of specieslargely confined to the zone between 2,000 and2,800 m in Nepal is plotted in fig. 7 (these are

the birds not used in thepreliminary analysis forfigs. 2 and 3). Fig. 7shows clearly that mostspecies breeding at thisaltitude in Nepal alsoextend into the easternHimalaya, and manyalso occur in the hills ofsouth and east Assam,India, the mountains ofnorthern, northeasternand western Myanmar,and into western YunnanProvince, China. Asmaller number of theseNepalese species also



Fig. 6. Distribution of species with a breeding range falling predominantlybetween Line 1 and Line 2 in the Arabian Peninsula (ten Arabian endemics).

For definitions of these Lines, see figs. 2 & 3.

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extend south into the moun-tains of northern Thailand,northern Laos, and north-western Vietnam, whileothers reach north toSichuan, Gansu, andsouthern Shaanxi provincesin central China, or reappearin the hills of south-centralChina east to the hills ofFujian Province in easternChina, and some even reachTaiwan. In contrast to theOriental species which pre-dominate at 1,000–2,000 min Nepal (see above), veryfew of those species at2,000–2,800 m extend to thehills of mainland southeastAsia or Hainan Island,China.

When the number ofspecies per 100 m of altitudeis plotted for the sameunequivocally Palearctic andOriental species used in figs.2 and 3, it becomes clearthat some Palearctic speciesextend down below 100 mwhile some Oriental speciesreach up to 3,200 m, eventhough their main distribu-tions are (respectively)above 2,800 m/below 2,000m (fig. 8). The elevation atwhich Palearctic and Ori-ental passerines appear toreach an equilibrium isapproximately 2,400 m.Therefore, all 68 specieswith a main breeding distri-bution between 2,000 and2,800 m are, for furtheranalysis, divided into speciesoccurring predominantly at2,400–2,800 m (36 species,classed as Palearctic), andthose predominantly at2,000–2,400 m (32 species,Oriental).

A similar analysis wasundertaken using altitudinaldata from Schäfer (1938),who carried out a transect



342. Chestnut Thrush Turdus rubrocanus, Jiuzhaigou, Sichuan Province, China,July 2006.Two races of Chestnut Thrush breed in the mountains fringing thesouthern boundary of the Palearctic.The nominate, pale-headed, form breedsat 2,300–3,300 m in the Himalaya, from E Afghanistan to Arunachal Pradesh,

India, and the dark-headed form T. r. gouldi (illustrated) breeds at 2,800–3,800 m in montane forest in western/central China.

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343. Plain-backed Thrush Zoothera mollissima, Emei Shan, Sichuan Province,China,April 2005.Within the Palearctic, Zoothera thrushes reach their greatest

diversity in the eastern Himalaya, where seven species breed, these specieshaving ranges in the Oriental as well as the Palearctic region. Plain-backed

Thrush is an altitudinal migrant, breeding entirely within the Palearctic, mostlybetween 3,000 and 4,300 m, but descending into the Oriental region in winter,

where it occurs between 1,400 and 2,800 m.

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in central China. Starting in the Red Basin, anextension of the eastern lowland plain intocentral Sichuan, he surveyed bird distributioncontinuously to the Tibetan Plateau in eastern

Qinghai Province. Hefound a strong turnoverin species compositionat 2,000–2,600 m, withspecies of the (sub)trop-ical lowland plain occur-ring up to 2,000 m(2,600 m), and Pale-arctic ones mainly from(2,000 m) 2,500 mupwards, but with 20species of the lattergroup below 1,000 m.Based on his data, anequilibrium betweenPalearctic and Orientalpasserines occurs at c.

2,300–2,400 m. Therefore, mid-altitudeendemics with a main altitudinal range of2,000–2,400 m in central China are con-sidered as Oriental, while those inhabitingthe bamboo (Bambuseae) jungle and cloudforest at 2,400–3,800 m are Palearctic.

Eastern ChinaOf the 37 species which occur predomi-nantly between Line 1 and Line 2 inChina, and below 2,400 m, remarkably fewoccur beyond eastern China (fig. 9). Toestablish whether these species are essen-tially Palearctic or Oriental, it is necessaryto review the remaining passerine fauna inthis area. Of the unequivocal Palearcticspecies (fig. 2), 37 extend into the plainsand/or hills of eastern China, as do 50unequivocally Oriental species, whichoccur predominantly south of Line 2 (fig.3). In addition, several species found atmid elevations in the Himalaya also extendinto eastern China; 17 occur mainly at2,400–2,800 m in the Himalaya and areconsidered Palearctic, and 12 breed pre-dominantly at 2,000–2,400 m and aretreated as Oriental. This gives a combinedtotal of 54 Palearctic and 62 Orientalbreeding species in the area of easternChina under discussion (see table 1).

Clearly, the avifauna of eastern China isof mixed origin, but Oriental species pre-dominate (table 1). Consequently, eastern

China is here considered to belong to the Oriental region, as are its 37 endemic birdspecies, even though the difference in numberof Oriental and Palearctic species is small. The



Fig. 8. Number of passerine species per 100 m change inelevation in the Himalaya, Nepal, based on unequivocallyPalearctic and Oriental birds (those mapped in fig. 2 and fig. 3, respectively). Nepalese species with a distribution

predominating between 2,000 m and 2,800 m are excluded(see fig. 7 for the horizontal distribution of these species).

Key: blue = 135 Palearctic species; red = 185 Oriental species.

Fig. 7. Distribution of species with a breeding range falling predominantlybetween Line 1 (at c. 2,800 m) and Line 2 (at c. 2,000 m) in Nepal.

For definitions of these lines, see figs. 2 & 3.

0 20 40 60 80 100 120 140Number of species

5500-5600

5200-5300

4900-5000

4600-4700

4300-4400

4000-4100

3700-3800

3400-3500

3100-3200

2800-2900

2500-2600

2200-2300

1900-2000

1600-1700

1300-1400

1000-1100

700-800

400-500

75-200

Alti

tudi

nal z

one

(m)

1

2

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endemics include some interesting speciesrestricted to the hills of Sichuan, southernGansu and/or southern Shaanxi provinces, suchas Martens’s Warbler Seicercus omeiensis, EmeiLeaf Warbler Phylloscopus emeiensis, Emei ShanLiocichla Liocichla omeiensis, and Slaty BuntingLatoucheornis siemsseni.

TaiwanA total of 91 passerine species breed regularly inTaiwan. This includes 29 species with widespreaddistributions within the Palearctic and 29 simi-larly widespread Oriental species (figs. 2 & 3). Afurther 21 species are shared with the mid-alti-tude range of the Himalaya in Nepal, or witheastern China – a range spanning a region inwhich the avifauna of the Palearctic and Orientalregions reaches equilibrium. Following analysisof these regions (see above), 17 species are con-sidered to be Oriental and four Palearctic. TheTaiwan avifauna is thus mixed, but Orientalspecies predominate (table 1). This resembles theregional affinities of species in eastern Chinaand, similarly, there is no clear altitudinal diver-gence among species, with some Palearcticspecies restricted to the lowlands, and some Ori-ental species breeding higher up in the moun-

tains. Since Oriental passerines predominate, the12 endemics are all attributed to the Orientalregion. These include species from predomi-nantly Palearctic genera (including CollaredBush Robin Luscinia johnstoniae and FlamecrestRegulus goodfellowi), although these are not par-ticularly close to any Palearctic species. Otherendemics show a morphological resemblance tomore widespread Oriental counterparts (e.g.Styan’s Bulbul Pycnonotus taivanus with Light-vented Bulbul P. sinensis, Taiwan WhistlingThrush Myophonus insularis with Blue WhistlingThrush M. caeruleus and Steere’s Liocichla Lioci-chla steerii with Emei Shan Liocichla).

The Lanyu Islands, close to southeastTaiwan, are poor in breeding species, butamong these are some shared with the Philip-pines (Lowland White-eye Zosterops meyeni andPacific Swallow H. tahitica). More detailedresearch is necessary before the islands’ zoogeo-graphic position can be established.

Southern Japanese islandsOf the passerines found on the Nansei-shoto(Ryukyu Islands), most are shared with the mainislands of Japan, and thus belong to thePalearctic; only Light-vented Bulbul and Pacific

Swallow are unequivo-cally Oriental. The rela-tionships of the threeendemic passerinesappear to lie withPalearctic species also:Ryukyu Robin Lusciniakomadori appears closelyrelated to JapaneseRobin L. akahige orRufous-headed Robin L.ruficeps of central China;Amami Thrush Zootheramajor is a close relativeof White’s Thrush Z.aurea of the northernPalearctic and Scaly



Fig. 9. Distribution of species with a breeding range falling predominantlybetween Line 1 and Line 2 in eastern China (37 species, virtually endemic to

eastern China). For definitions of these lines, see figs. 2 & 3.

Table 1. Number of Palearctic, Oriental, and endemic passerine species of some regions and islands in East Asia.

Palearctic Oriental endemic

Eastern China 54 62 37 Oriental species predominate

Taiwan 33 46 12 Oriental species predominate

Nansei-shoto (Ryukyu Islands) 11 2 3 Palearctic species predominate

Daito (Borodino) Islands 8 0 0 Palearctic species predominate

Ogasawara-shoto (Bonin Islands) 6 0 3 Palearctic species predominate

Kazan-retto (Volcano Islands) 4 0 0 Palearctic species predominate

1

2

18

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Thrush Z. dauma of theHimalaya; and Lidth’s JayG. lidthi is morphologi-cally close to Black-headed Jay G. lanceolatusof the western Himalaya.

Eight passerines breedon the Daito (Borodino)Islands. All of these(Brown-eared BulbulMicroscelis amaurotis ,Wren, Blue Rock Thrush,Japanese Bush WarblerCettia diphone, VariedTit, Japanese White-eyeZosterops japonicus, Bull-headed Shrike Laniusbucephalus, and TreeSparrow) are shared withthe main islands of Japanand are Palearctic. Six of the nine speciesbreeding, or formerlybreeding, on the Ogasawara-shoto (BoninIslands) are also shared with the main islands ofJapan, but the other three are endemic (BoninThrush Zoothera terrestris, Bonin HoneyeaterApalopteron familiare, and Bonin GrosbeakChaunoproctus ferreorostris) and the nearest rel-atives of these species are not yet satisfactorilyestablished. The four passerines breeding regu-larly on the Kazan-retto (Volcano Islands) arealso shared with the main islands of Japan.Together, the islands in southern Japan havePalearctic taxa predominating and thus areincluded in the Palearctic (table 1).

A definitive southern boundary of thePalearcticAfter establishing the affinities of species thatoccur predominantly between Line 1 and Line 2(figs. 2–9), a revised line can be determinedaccording to whether Palearctic or Afrotropicaland/or Oriental species predominate in eachgrid cell (see fig. 10). Owing to the inclusion ofthe (semi-)desert region of the Sahara in thePalearctic, this revised line takes a slightly moresoutherly track than the preliminary line (fig. 4)in Africa (at c. 19°N, but north to Gebel Elba atc. 22°N in the extreme east). Within the Arabian

Peninsula, the revisedsouthern boundaryincludes the entireregion except the moun-tains of the south andsouthwest (Afrotropical)and the coastal plain ofnorthern Oman (Orien-tal). The preliminaryand revised lines aresimilar across the Indiansubcontinent. Bothinclude the coastal plainof southern Iran andsouthern Baluchistanwithin the Orientalregion, the mountains ofwestern Pakistan in the



Fig. 10. Definitive southern border of the Palearctic region, based on distributionof all passerine species breeding within the area of the map (species richness

shown).The line is drawn where number of breeding Palearctic andAfrotropical/Oriental species reaches an equilibrium.

344. Elliot’s Laughing-thrush Garrulax elliotii, Huanglong, Sichuan Province, China,April 2005.Although most laughing-thrushes are confined to the Oriental region,

several species breed in the Himalaya and western China between 2,000 and2,800 m, where Palearctic and Oriental faunas overlap. Elliot’s Laughing-thrush isone of few species of laughing-thrush that is entirely restricted to the Palearctic,where it breeds mainly at 2,000–4,000 m in the mountains of central China from

Qinghai east to Gansu Province, and south to northern Yunnan Province.

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Palearctic (including theSafed Koh in northwest Pak-istan), although the Jalal-abad valley, just north of theSafed Koh in Afghanistan, isOriental. In China, Xizang(Tibet Autonomous Region)is Palearctic, but the altitu-dinal upper limit of theOriental region, at c. 2,400m, locally reaches intosouthernmost Xizang(Gyirong, Nyalam, part ofupper Arun Valley, Yadongarea in Chumbi Valley,Dihang/ Yarlung Valley nearBomi, Zayü Valley, parts ofNu Jiang/Salween andLancang Jiang/Mekongrivers). In Yunnan Province,the Palearctic extends southto the Dali area, and east toc. 103°E between northernYunnan and central Sichuanprovinces. Farther east theborder becomes highlycapricious, reaching northin the lowland plains to32°N near the Yangtze Riverestuary, and still farthernorth into the upper HanShui River basin in northernHubei Province. In SichuanProvince, the low-lying RedBasin is undoubtedly Orien-tal (as in fig. 4), but the sur-rounding mountains have apredominantly Palearcticpasserine fauna (above c.2,300–2,400 m).

A Palearctic listIn this analysis, species areassigned to a certain faunalregion according to theirbreeding distribution; forexample, one with the largerpart of its range in theOrient is defined as Orien-tal, even though a smallerpart may extend well intothe Palearctic (as definedhere). Compare this with theconcept of a national list,



345. Black-browed Tit (Père Bonvalot’s Tit) Aegithalos bonvaloti, Cang Shan,Dali,Yunnan Province, China, September 2005.This attractive ‘long-tailed tit’

breeds up to 3,400 m in the mountains of northern Yunnan and centralSichuan provinces, China, which form the boundary between the Palearctic

and Oriental regions in central China.

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346. White-capped Water Redstart Chaimarrornis leucocephalus, SiguniangShan, Sichuan Province, China, June 2006.White-capped Water Redstart

inhabits fast-flowing streams of the Himalaya and hill ranges east to easternChina, where it breeds mostly between 1,800 and 4,600 m.Within such a wide

altitudinal range, it occupies suitable habitat in both the Palearctic and Oriental regions, but its main distribution lies within the Palearctic.

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where every specieswhich has occurredwithin that nation’sborders is counted. Thesame approach could beapplied to a list ofPalearctic birds, whichthen would includespecies having only a tinyfoothold in the Palearcticas a breeding bird (oreven species which occuras a migrant or vagrant),even though the speciesis mainly Afrotropical orOriental in distribution(or in some casesNearctic: Grey-cheekedThrush Catharusminimus and SavannahSparrow Passerculussandwichensis breed reg-ularly in extreme north-eastern Siberia). Thosespecies which have recog-nisable taxa in thePalearctic have a strongcase for inclusion on the Palearctic list; forexample, the Afrotropical Black-crownedTchagra, of which there is a distinct race resi-dent in northwest Africa, or the OrientalCommon Babbler, which has distinct taxa in theMiddle East. Species like these, with recognis-able taxa in different faunal regions, are prob-ably the first that should be examined in termsof potential taxonomic splits, not only thosewith distributions shared between the Palearcticand Afrotropical or Oriental regions, but alsothose with ranges which extend into bothsouthern regions, like Graceful Prinia Priniagracilis (and indeed Oriental Skylark, Red-rumped Swallow, Long-billed Pipit, Zitting Cis-ticola, Clamorous Reed Warbler, Large-billedCrow, and House Sparrow mentioned already),which are not included here because of the dif-ficulty in assigning them to a particular region.

In this zoogeographical analysis, 563passerine bird species are considered to bePalearctic. All these are also listed as Palearcticby Beaman (1994), apart from some speciesrecently discovered or split (including someSeicercus and Phylloscopus warblers, andSichuan Treecreeper Certhia tianquanensis).Beaman listed a further 43 species as Palearctic

which here are considered to have Afrotropicalaffinities (mainly in Arabia, where opinions onthe Palearctic boundary differ widely), and 92species here considered to belong to the Orien-tal region (though some do extend into thePalearctic, some quite extensively). Nonetheless,Beaman’s Palearctic list represents a valuableoverview of species occurring in the Palearctic,provided that those restricted to the south-western and southern Arabian Peninsula areexcluded.

Would the southern boundary of thePalearctic change if non-passerines wereincluded?Although the available maps for Palearctic non-passerines are more crude than those for passer-ines, many non-passerines show range sizes anddistributions comparable with those of passer-ines, especially birds of forests, shrub, and openplains, such as pheasants and quails (Phasian-idae), raptors (Accipitridae), sandgrouse (Ptero-clididae), pigeons and doves (Columbidae),cuckoos (Cuculidae), owls (Strigidae), nightjars(Caprimulgidae) and woodpeckers (Picidae).Some non-passerine landbirds are far morewidespread than any songbird (e.g. Peregrine



347. Lidth’s Jay Garrulus lidthi,Amami-Oshima, Japan, February 2006.The islands ofthe Nansei-shoto (Ryukyu Islands) form the southeastern boundary of the

Palearctic and are home to 18 passerine species, most of which are shared withthe main islands of Japan, while others are endemic to the islands. Lidth’s Jay

is found only on Amami-Oshima, but is morphologically close to Lanceolated JayG. lanceolatus of the western Himalaya.

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Falcon Falco peregrinus, Osprey Pandion hali-aetus and Barn Owl Tyto alba), but these are rel-atively few and their number would have littleeffect on the boundaries established here. Notethat the zoogeographical regions as defined bySclater (1858) are based on landbirds; seabirdssuch as petrels (Procellariidae), gannets(Sulidae) and auks (Alcidae) simply do not fitthis essentially land-based concept.

On the other hand, the passerines include veryfew waterbirds (just two dippers) or tundradwellers (mainly a few pipits, finches, andbuntings), while these are numerous among non-passerine species – e.g. waterfowl (Anatidae),divers (Gaviidae), grebes (Podicipedidae), herons(Ardeidae) and, particularly on the tundra, manywaders (Charadriidae and Scolopacidae). Thebreeding ranges of many of these are extensiveand/or extend to other continents, and includingthem in an analysis such as this may affect theoutcome, although their inclusion is more likelyto affect the northwestern and/or northeasternboundary than the southern one.

Acknowledgments

Paul Williams (The Natural History Museum, London)made the ver sion of Wor ldmap avai lable; PeterMekenkamp (Faculty of Geo Sciences, Depar tment ofCar tography, University of Utrecht) provided a finePalearctic base-map making conversion of or iginalhandmade maps into WorldMap possible. Guido Keyl(now Naturalis, Leiden) and Mansour Aliabadian (Institutefor Biodiversity and Ecosystem Dynamics IBED, Universityof Amsterdam) helped in digitising the original maps. Mapswere in par t derived from label data of specimens inzoological collections, access to which was kindlypermitted by their staff. Financial support came, in part,from Synthesys (grants DE-TAF-796 and GB-TAF-826).Comments from Ronald Sluys (IBED), Mansour Aliabadian(IBED), and Vincent Nijman (ZMA) improved the paper.

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Kees (C. S.) Roselaar, Zoological Museum ZMA, University of Amsterdam, PO Box 94766,1090 GT Amsterdam, The Netherlands; e-mail [email protected]

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