distribution patterns in the genus peltigera...

Distribution patterns in the genus Peltigera Willd.

Isabel MARTIuNEZ Ana Rosa BURGAZ Orvo VITIKAINEN andAdrian ESCUDERO

Abstract: The distribution of sixty-six Peltigera species in 230 biogeographic provinces or 40 regionsare presented. A hierarchical clustering approach, used to identify clusters of species with similardistribution patterns (floristic elements), resolved four groups made up of Neotropical, SouthernHemisphere, Antarctic and mainly Holarctic species. The Holarctic Kingdom is species rich with thehighest number of Peltigera species and also the highest number of endemic species; the Australianand Cape Kingdoms have the lowest number of species and endemics. The species rich provinces arebriefly discussed.

� 2003 The British Lichen Society. Published by Elsevier Ltd. All rights reserved.

Key words: biogeographic provinces, lichens, Peltigera species, world distribution patterns.

Introduction

The world distribution of organisms hasfascinated scientists curious about naturesince early times (Brown & Lomolino 1998).However, studies analysing the geographicaldistribution of lichens are very scarce, es-pecially compared with those that considerthe distribution of angiosperms or pterido-phytes. However, lichens have severalcharacteristics that could provide valuablebiogeographical information: they are con-sidered to be ancient organisms (Lutzoniet al. 2001), their distribution can be easilyand directly related to climate (Mies &Losch 1995), and they display a great dis-persal capacity, mostly due to the lightnessof their spores and vegetative propagules(Kappen & Straka 1988; Mies & Losch

1995). The scarcity of lichen biogeographi-cal investigations is probably a consequenceof the poor chorological knowledge of mostlichen species.

Earlier papers dealing with lichen bio-geography largely presented data in a sub-jective way and most of them presentexplanations based on inductive reason-ing (Yoshimura 1968; Kurokawa 1972;Almborn 1985; Galloway 1988, 1991a,b,1994; Jørgensen 1994; Moberg 1994;Goward & Ahti 1997; Caceres et al. 2000;Otte et al. 2002). More recently Printzen& Lumbsch (2000) explored molecularvariability to determine when and whereBiatora and Phyllospora have diversified.Lucking (2003) undertook a comparativestudy of Takhtajan’s floristic regionsand foliicolous lichen biogeography andproposed six lichenogeographical regions.

Currently, phytogeographic studies inlichenology, as in other disciplines, aredependent on accurate taxonomic studies.The lichen genus Peltigera comprises 66 rec-ognized species and includes terricolous andmuscicolous foliose macrolichens, which arecommon and widespread on most conti-nents. Although numerous taxonomic and

I. Martınez and A. Escudero: Area de Biodiversidad yConservacion, ESCET, Universidad Rey Juan Carlos,E-28933, Mostoles, Madrid, Spain.A. R Burgaz: Departamento de Biologıa Vegetal I,Facultad de CC. Biologicas, Universidad Complutensede Madrid, E-28040-Madrid, Spain.O. Vitikainen: Botanical Museum (Mycology),P.O. Box 47, FIN-00014 University of Helsinki,Finland.

Lichenologist 35(4): 301–323 (2003)doi:10.1016/S0024-2829(03)00041-0

0024-2829/03/040301+23 $30.00/0 � 2003 The British Lichen Society. Published by Elsevier Ltd. All rights reserved.

T 1. List of Peltigera species considered in the study, the presence of each species in the different Kingdoms andcontinents and main characteristics

Kingdom* Continent‡ Summary§

Species H N P AU HL C NA SA E AS AF AN Prv Reg Suk

P. andensis Vitik. + + 2 1 1P. aphthosa (L.) Willd. + + + + 41 8 3P. aubertii Dodge + + 2 1 1P. austroamericana Zahlbr. + + + 10 6 5P. britannica (Gyeln.) Holt.-Hartw. &

Tønsberg+ + + 15 3 2

P. canina (L.) Willd. + + + + + + + + + + + + 87 20 11P. chilensis Gyeln. + + 2 1 1P. chionophila Goward & Go$net + + 6 2 1P. cichoracea Jatta + + + 3 3 2P. cinnamomea Goward + + 5 2 1P. collina (Ach.) Schrad. + + + + + + + + + 62 15 7P. continentalis Vitik. + + 6 3 2P. degenii Gyeln. + + + + + + + 42 9 5P. didactyla (With.) J. R. Laundon + + + + + + + + + + + + 87 26 11P. dilacerata (Gyeln.) Gyeln. + + 2 2 2P. dissecta Purvis, P. James & Vitik. + + 1 1 1P. dolichorhiza (Nyl.) Nyl. + + + + + 29 21 12P. dolichospora Vitik. + + 2 1 1P. elisabethae Gyeln. + + + + 45 8 3P. erioderma Vain. + + + 2 1 1P. evansiana Gyeln. + + + 11 5 2P. fibrilloides (Gyeln.) Vitik. + + 1 1 1P. friesiorum Gyeln. + + + 2 2 2P. frigida R. Sant. + + 1 1 1P. frippii Holt.-Hartw. + + + + 4 1 1P. horizontalis (Huds.) Baumg. + + + + + 58 7 3P. hymenina (Ach.) Delise + + + + 28 4 2P. kristinssonii Vitik. + + + + 19 5 2P. laciniata (G. Merr. ex Riddle) Gyeln. + + + 9 5 3P. lairdii Dodge & E.D. Rudolph + + 1 1 1P. lambinonii Go$net + + + + 2 2 2P. lepidophora (Vain.) Bitter + + + + + + + + 43 8 4P. leucophlebia (Nyl.) Gyeln. + + + + 53 8 3P. lyngei Gyeln. + + 2 1 1P. malacea (Ach.) Funck + + + + 49 9 3P. melanorrhiza Purvis, P. James & Vitik. + + 2 2 2P. membranacea (Ach.) Nyl. + + + + + + 58 11 5P. microdactyla Nyl. + + 1 1 1P. monticola Vitik. + + + + 16 5 3P. neckeri Hepp ex Mull. Arg. + + + + + + + 59 8 3P. neopolydactyla (Gyeln.) Gyeln. + + + + 38 7 3P. nigripunctata Bitt. + + 7 4 1P. occidentalis (E. Dahl) H. Krist. + + + + 6 2 1P. oceanica Gyeln. + + 2 1 1P. pacifica Vitik. + + 5 2 1P. patagonica Rasanen + + 2 1 1P. phyllidiosa Go$net & Mia(likowska + + 3 1 1P. pindarensis D. D. Awasthi & M. Joshi + + 1 1 1P. polydactyloides Nyl. + + 3 2 1P. polydactylon (Neck.) Ho#m. + + + + + + + + + + + + 82 20 11P. ponojensis Gyeln. + + + + 46 6 3P. praetextata (Florke ex Sommerf.) Zopf + + + + + + 78 12 4P. pruinosa (Gyeln.) Inumaru + + 3 1 1P. pulverulenta (Taylor) Nyl. + + + + + 8 7 5P. retifoveata Vitik. + + + + 13 4 2

302 Vol. 35THE LICHENOLOGIST

chorological approaches have been con-ducted during recent years (Holtan-Hartwig1993; Go$net & Hastings 1994, 1995;Purvis & James 1993; Vitikainen 1985,1986, 1994a,b, 1995, 1999; Go$net et al.1994; Goward et al. 1995; Go$net &Miadlikowska 1999; Martınez 1999;Goward & Go$net 2000), most of themwere regionally based, so they lack thebroad biogeographic perspective. However,Miadlikowska & Lutzoni (2000) carried outa phylogenetic revision of the genus basedon morphological, chemical and moleculardata. Despite these studies, knowledge ofthis genus remains geographically hetero-geneous. The Northern Hemisphere isrelatively well-studied and important collec-tions are available in herbaria, exceptfor central Asia, which remains poorlyexplored. On the other hand, the SouthernHemisphere is poorly known, althoughsome studies are currently underwayin the Neotropical (Vitikainen 1994b,1995, 1999), Holantarctic and AustralianKingdoms (Vitikainen, in prep.). Never-theless, Peltigera is one of the most exten-sively studied lichen genera, and so is avaluable subject by which to explore thedistribution features of a widely distributedlichen genus.

The present study aims to identify theexistence of groups of Peltigera species withsignificantly similar distribution patterns[floristic elements (Birks 1976)] in theworld. In addition areas with the highestnumber of species are identified anddiscussed.

Materials and Methods

A. Data collection

Only well-accepted Peltigera species [e.g. see recentrevisions by Vitikainen (1994a, 1999)] were included inthe study and taxonomic categories below species weredisregarded (Table 1). Recently, Miadlikowska &Lutzoni (2000) considered Hydrothyria venosa to beincluded in the genus Peltigera, as P. hydrothyriaMiadlikowska & Lutzoni, but this species has not beenincluded in this study because the data have alreadybeen analysed. Available data from the literature (morethan 300 floristic papers) and from revised materialfrom the most relevant herbaria for the genus (seeVitikainen 1994a) were compiled. The world wasdivided into 230 biogeographic provinces that mainlyfollow Tahktajan (1986), but with certain modificationsfrom Quezel (1978) for Northern Africa, and fromRivas-Martınez (1987, 1993) and Rivas-Martınez et al.(1999) for the Mediterranean Region and the Americancontinent (Appendix 1). Classification of these unitsinto higher levels follow Tahktajan (1986). Abundanceof each species in a province was estimated with a fivestep linear scale based on the total number of recordsper species weighted by the size and the total number of

T 1 Continued.

Kingdom* Continent‡ Summary§

Species H N P AU HL C NA SA E AS AF AN Prv Reg Suk

P. rufescens (Weiss) Humb. + + + + + + + + + + + + 89 20 9P. rufescentiformis (Gyeln.) Dodge + + 3 2 1P. scabrosa Th. Fr. + + + + 31 7 3P. scabrosella Holt.-Hartw. + + + 6 2 1P. soredians Vitik. + + 3 2 2P. spuriella Vain. + + 3 2 2P. subhorizontalis Gyeln. + + + 3 2 2P. truculenta De Not. + + + 3 2 2P. ulcerata Mull. Arg. + + + + + + + + + + 13 8 7P. vainioi Gyeln. + + 1 1 1P. venosa (L.) Ho#m. + + + + 46 8 3

*H: Holarctic Kingdom; N: Neotropical Kingdom; P: Paleotropical Kingdom; AU: Australian Kingdom;HL: Holantarctic Kingdom; C: Cape Kingdom.

‡NA: North America; SA: South America; E: Europe; AF: Africa; AS: Asia; AN: Australian-New Zealand.§Prv: number of provinces in which each species appears; Reg: number of regions in which each species appears;

Suk: number of Subkingdoms in which each species appears.

2003 Distribution of the genus Peltigera Willd.—Martiınez et al. 303

records per geographical unit. An abundance matrix(66 species�230 provinces) was prepared andsubjected to further analyses.

B. Data analysis

The abundance matrix of species�biogeographicalprovinces was submitted to a hierarchical clustering toclassify species into homogeneous groups. Euclideandistance was used as distance coe$cient and theUPGMA algorithm was approached to build a hier-archical classification. The optimal number of clustersor cut levels in the hierarchical classification wasobtained by means of the cluster separation coe$cient(Podani 1993). This coe$cient was calculated for eachlevel (‘groups number’). The number of clusters inwhich the asymptotic value of the coe$cient wasfirst reached was considered the optimal classificationstructure. The number of clusters considered rangesbetween 2 and 10. Analyses were conducted withSYN-TAX v.4 (Podani 1993).

Results and Discussion

The presence of each of the 66 Peltigeraspecies in the di#erent Kingdoms and asummary of their distribution characteristicsare presented in Table 1 and the number ofPeltigera species found on each continent inTable 2. The number of Peltigera species oneach continent in the Northern Hemisphereis rather similar (ranging between 30 and35), whereas in the Southern Hemispherethis number decreases from 25 in SouthAmerica to only 6 in New Zealand. More-over, the three northern land masses sharearound 25 species, while the SouthernHemisphere barely reaches 10 species(Table 2).

The endemic Peltigera species in each con-tinent are listed in Table 3. Species richnessin the Holarctic reaches a maximum amongthe floristic Kingdoms (44 species with 30

endemics). The Australian (9 species with1 endemic) and Cape Kingdoms (4 speciesand no endemics) are the poorest.

The Subkingdoms or equivalent geo-graphical units with the highest number ofspecies are the Boreal Subkingdom (40),Tethyan Subkingdom (31), MadreanSubkingdom (21), Andean Superregion(17) and African Subkingdom (14), thethree former belonging to the HolarcticKingdom. The regions with the highestnumber of Peltigera species are also situatedin the Holarctic Kingdom: Circumboreal(35), Rocky Mountains and Iranian-Turanian (28), Eastern Asiatic (26), NorthAmerican Atlantic (24) and Mediterranean(23). And finally, the provinces with thehighest number of Peltigera species are alsosituated in the Holarctic Kingdom (Fig. 1).

Fig. 2 gives the optimal classification ofspecies distributions produced by the hier-archical clustering analysis. The 66 Peltigeraspecies were classified into four floristic ele-ments; the most e$cient partition beingobtained at the four cluster level. At this stepthe separation coe$cient reached theasymptotic level thus optimizing the di#er-ences among groups. The first group,namely group A, separates the Neotropicalspecies and includes 3 subgroups. Thesecond cluster (B) mostly separates veryscarce species mainly distributed inthe Southern Hemisphere and can bedivided in 4 subgroups. The third group (C)separates Holantarctic species. Finally,the fourth cluster (D) includes mostPeltigera species, mainly distributed in theHolarctic Kingdom and includes 7 sub-groups (Fig. 2).

T 2. Number of Peltigera species in each continent are indicated in the diagonal. Species shared between continentsare indicated above and below the diagonal

Europe Asia N. America S. America Africa Australia N. Zealand

Europe 30 24 26 6 11 7 3Asia 24 35 26 8 14 8 5N. America 26 26 33 9 12 8 4S. America 6 8 9 25 8 7 3Africa 11 14 12 8 20 8 4Australia 7 8 8 7 8 9 4New Zealand 3 5 4 3 4 4 6


Group A

This group separates 10 Neotropicalspecies, which fall into three subgroups.However, due to imperfect knowledge of thetaxonomy of Neotropical Peltigera species,discussion of these subgroups is fraught withambiguities. The first two subgroups arerepresented by very scarce species, reportedfrom only one to three biogeographicprovinces.

The first subgroup comprises Peltigeraandensis, P. soredians, P. spuriella, P.fibrilloides, P. microdactyla and P. vainioi,which are endemic species from the Neo-tropical Kingdom and distributed mainly inthe Andean range. Almost all these speciesgrow at high altitudes. As Schuster (1983)suggested in the case of Neotropical bryo-phytes, these high-elevation areas constitute

biotic islands that su#ered the e#ects ofPleistocene glaciation. Rapid evolutionaryprocesses coupled with such climatic changecould explain the high levels of endemism.At least some of these taxa could be ancientbut now have very restricted ranges(palaeoendemics).

The second subgroup is represented only byP. friesiorum, which is distributed in theNeotropical Kingdom and in the Tristan daCunha Islands (Holantarctic Kingdom).

The third subgroup includes three species,reported from eight to ten biogeographicprovinces and whose distribution areas aremainly spread in the Neotropical Kingdom,although they also grow in other Kingdoms.Peltigera laciniata is a Gondwana element,restricted to Central and South America(Neotropical and Holantarctic Kingdoms).

T 3. Endemic Peltigera species in the different floristic Kingdoms. Total numbers of Peltigera species in eachKingdom are shown at the bottom of the Table

Holarctic Paleotropical Neotropical Australian Holantarctic

P. aphthosa P. cichoracea P. andensis P. lairdii P. aubertiiP. britannica P. erioderma P. fibrilloides P. chilensisP. chionophila P. lambinonii P. microdactyla P. frigidaP. cinnamomea P. oceanica P. soredians P. patagonicaP. continentalis P. polydactyloides P. spuriella P. truculentaP. dissecta P. rufescentiformis P. vainioiP. dolichosporaP. elisabethaeP. evansianaP. frippiiP. horizontalisP. hymeninaP. kristinssoniiP. leucophlebiaP. lyngeiP. malaceaP. melanorrhizaP. monticolaP. neopolydactylaP. nigripunctataP. occidentalisP. pacificaP. phyllidiosaP. pindarensisP. ponojensisP. pruinosaP. retifoveataP. scabrosaP. scabrosellaP. venosa44 15 18 9 18


F. 1. World map showing the number of Peltigera species in the di#erent biogeographic provinces.

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Peltigera austroamericana and P. pulverulenta,which are known from South America (Neo-tropical and Holantarctic Kingdoms), andreach Central Mexico (Holarctic Kingdom),can be included in a North America-SouthAmerican phytogeographical element (Qian1999). They exemplify the significance ofthe current landbridge between North andSouth America which facilitates the bioticexchange between South and NorthAmerican landmasses (Brown & Lomolino1998).

Group BThis group, comprising 13 species, in-

cludes mainly very rare species distributed inthe Southern Hemisphere. As in group A,di#erent subgroups are apparent.

The first subgroup includes five endemicspecies from the Paleotropical Kingdom,although with very small, non-overlapping,ranges. Peltigera cichoracea is endemic toeastern Africa and Papua New Guinea,P. polydactyloides and P. rufescentiformis areendemic to central Africa, and P. eriodermaand P. oceanica are endemic to theIndomalesian Subkingdom (Philippineanand Papuan Provinces).

The second subgroup is formed bythree species distributed in the AustralianKingdom. Peltigera lairdii appears only in theSouth-east Australian Province; P. lambino-nii occurs in New South Wales of theAustralian Kingdom and East Africa; andP. dilacerata is known from the Japanese-Korean and the South-east AustralianProvinces.

The third subgroup is represented bytwo endemic species from Asia (HolarcticKingdom). Peltigera dolichospora is endemicto central Asia (Northern Chinese andEastern Himalayan Provinces) and P. pru-inosa is an endemic of eastern and centralAsia (Japanese-Korean, Taiwanian andEastern Himalayan Provinces).

The fourth subgroup includes P. doli-chorhiza, P. ulcerata and P. subhorizontalis.These species are distributed mainly in theSouthern Hemisphere, but P. dolichorhizaand P. ulcerata are also found elsewhere.Thus, P. ulcerata grows in the Paleotropical,

Neotropical, Australian, Cape and Holant-arctic Kingdoms, and is also present in theHolarctic Kingdom (only known from theWestern Himalayan Province). Peltigeradolichorhiza is known from all floristic King-doms (except in the Cape). However, itspresence in the Holarctic Kingdom is verylimited, appearing only in the south ofMexico, the Himalaya range, Japan andTaiwan. On the other hand, the distributionarea of P. subhorizontalis is restricted toAustralia and New Zealand.

Group C

This group separates five very rarespecies known only from the HolantarcticKingdom. Peltigera aubertii and P. chilensisare limited to the southernmost tip of SouthAmerica (Argentina and Chile), P. frigidais restricted to Tristan da Cunha and Tierradel Fuego, P. patagonica is endemic tothe Valdivian-Magellanic Region (Tierradel Fuego and Antarctic Provinces), andP. truculenta is restricted to the southermostpart of South America and to Marion andPrince Edward Islands.

Group D

This group separates a very large group ofPeltigera species (38), which are exclusivelyor mainly distributed in the HolarcticKingdom. Within this wide group, sevensubgroups are recognized.

The first subgroup includes some ratherabundant species, which are present in atleast three di#erent continents, and distrib-uted mainly along circumpolar or circumbo-real areas of the Holarctic Kingdom. Thesespecies are P. aphthosa, P. neopolydactyla,P. elisabethae, P. degenii, P. leucophlebia,P. malacea, P. venosa, P. lepidophora, P.hymenina and P. scabrosa. All of them aredistributed in North America, Europe andAsia, except P. hymenina, which is knownfrom North America, Europe and Africa(Macaronesian area), and P. lepidophoraappears also in the Neotropical (SouthAmerica), Paleotropical (Hawaii) and theHolantarctic (New Zealand) Kingdoms.


The second subgroup includes five species,which are almost cosmopolitan and the mostcommon species in the world: Peltigeracanina, P. didactyla, P. polydactylon, P.praetextata and P. rufescens.

The third subgroup is also composed of fivespecies: P. collina, P. horizontalis, P. mem-branacea, P. neckeri and P. ponojensis. Thefirst three have suboceanic preferences,avoiding extreme continental situations. Allof the species are distributed in the HolarcticKingdom, but four of them have reachedother Kingdoms. Thus, P. collina is found insome areas of the Neotropical (Mexicoand Peru), Paleotropical (India), and

Holantarctic Kingdoms (Patagonia andTierra del Fuego), P. neckeri is found in theHolantarctic Kingdom (Argentina andChile), and P. membranacea reachesthe south-west of India (PaleotropicalKingdom).

The fourth subgroup includes six species,these being the rarest Holarctic speciesand distributed mainly in Europe, althoughthey are also present on other continents.Peltigera britannica is known from north-western and western Europe and fromnorth-western North America. This disjunctdistribution might not be surprising,because the climate conditions of western

F. 2. Dendrogram of the floristic elements. A, B, C, D represent the four floristic elements.


F. 2—continued


Europe are equivalent to western NorthAmerica, both are oceanic areas and werenot glaciated in the Quaternary (Brown &Lomolino 1998). Peltigera scabrosella is anincompletely circumpolar, boreal to arcticspecies restricted to north-western NorthAmerica, south-western Greenland andEurope (mainly in the west of theScandinavian Peninsula). Peltigera frippii, P.retifoveata and P. kristinssonii are circum-polar, boreal to arctic species, growing onthe three main landmasses of the HolarcticKingdom. Finally, P. monticola probablybelongs to the ancient Madrean-Tethyanterritory, spreading today along the calcare-ous mountains of Europe, North Americaand Asia.

The fifth subgroup includes five Holarcticspecies distributed in North America andoccasionally also in other continents. Thissubgroup is probably made up ofthree phytogeographical elements: NorthAmerican, North American-eastern Asianand tentatively North American-Europeanelements (Qian 1999). Peltigera chionophila,P. cinnamomea, P. pacifica and P. phyllidiosaare North American endemics. Peltigera cin-namomea is endemic to central and westernNorth America, being most abundant inthe Rocky Mountains. Peltigera pacifica andP. chionophila are north-western Americanendemics, while P. phyllidiosa is an easternAmerican endemic. Peltigera occidentalis is atentatively circumpolar, boreal to arcticspecies and it is known from northernNorth America and from Europe in theOroscandinavian province and probablyranges from north-western Europe over thewhole Siberian Taiga. On the other hand,P. evansiana shows an amphi-beringian pat-tern, growing in central Asia (Mongolia andNorth of China), and between 40–60(Nextending from south-east and to north-westof North America.

The sixth subgroup is made up of threeendemic Asiatic species: P. continentalisis endemic to central Asia; P. pindarensis isendemic to the Western Himalayas; andP. nigripunctata grows only in China, Korea,Japan and the Himalayas. Furthermore,some of these endemic species spread out

through China and Mongolia to Koreaand Japan, such as P. continentalis andP. nigripunctata.

Finally, the seventh subgroup includesthree very rare and outlier species from theHolarctic. Peltigera dissecta is known fromthe Azorean Province, and P. melanorrhizafrom the Azores as well as from westernIberian Peninsula. Peltigera lyngei is anarctic species known from northern Europe(Iceland and Svalbard Islands).

Provinces with the highest number ofspecies

The biogeographic provinces with thehighest number of Peltigera species arelocated in north-western North America,and the number of species decreasestowards the south (Fig. 1). This pattern isvery similar to that obtained by Goward &Ahti (1997), who in their study on theCladoniaceae in temperate and boreal west-ern North America observed that in NorthAmerica the number of species decreasestowards the south. This trend is not surpris-ing because of the geographical orientationof the principal mountain ranges, whichallowed the return to north-western NorthAmerica of many species after the glacialpulses (Brown & Lomolino 1998). On theother hand, the absence of such a latitudinaltrend in Europe may be related to the west-east orientation of main mountain ranges.

In Europe, the richest biogeographicprovinces are located in the north, with adecreasing richness towards the south,although there is a high number of species(20 or more) in some mountainous regionsof the southern half (Alps, Pyrenees,Carpathians, Cantabric mountains, etc.)(Fig. 1). In Asia, the biogeographic prov-inces with the highest number of species arelocated in the centre and east of the conti-nent. The lowest number of speciesin the northern regions of Asia probablyreflects the existence of many unexploredareas. The mountain habitats of SouthAmerica (Andes range) and Africa (easternmountains) harbour the highest number of


Peltigera species in their respective conti-nents, and in Australia it is the south-east ofthe continent where the largest number ofspecies are found (Fig. 1).

Finally, there are many areas in the worldwithout any Peltigera records. In some casesthis is to be expected, for example in thedeserts of Africa, North America and Chile(Fig. 1) but, in other cases, it may be aconsequence of the absence of floristicstudies (e.g. many areas of Brazil). It is alsoworth noting that some 25% of the speciestreated here have been recognized anddescribed during the last 20–25 years, andalso from fairly well-known areas suchas Europe. Further studies in other areasmay well reveal range extensions for manyspecies, as well as new undescribed taxa.

We are very grateful to Bruce McCune (Oregon,EEUU), Jose Miguel Olano (Soria, Spain), and twoanonymous referees for their valuable comments to themanuscript, to Gregorio Aragon (Madrid, Spain) forhis help with the elaboration of the figures and toRosario Gavilan and Daniel Sanchez-Mata (Madrid,Spain) who generously provided some biogeographicinformation. This study was supported by the SpanishProject DGES, BOS2001-0869-C04-04.

Appendix 1. List of biogeographicprovinces of the world

HOLARCTIC KINGDOM

BOREAL SUBKINGDOM

Circumboreal Region

Arctic Subregion

1. Oroescandinavian Province2. Islando-Groenlandic Province3. Asian Arctic Province.4. North American Arctic Province

Boreo-Continental Subregion

5. North European Province6. Central Russian Province7. Sarmatian Province8. Western Siberian Province9. Altai-Sayan Province

10. Middle Siberian Province11. Transbaikalian Province12. North-eastern Siberian Province13. Okhotsk-Kamchatka Province14. Canadian Boreal Province15. Yukonian Alaska Province

Atlantic-Central European Subregion

16. Western Alpine Province17. Eastern-Central Alpine Province18. Apennine-Padane Province19. Pyrenean Province20. Central European Province21. Subatlantic Province22. North Atlantic Province23. British Province24. Cantabrian-Atlantic Province25. Orocantabrian Province26. Azorean Province27. Carpathian Province28. Tatra Province29. Panonian Province30. Pontic Province31. Caucasian Province32. Euxine Province33. Bosnio-Illyrian Province34. Serbo-Macedonian Province

Eastern Asiatic Region

35. Manchurian Province36. Sakhalin-Hokkaido Province37. Japanese-Korean Province38. Volcano-Bonin Province39. Ryukyu or Tokara-Okinawa Province40. Taiwanian Province41. Northern Chinese Province42. Central Chinese Province43. South-eastern Chinese Province44. Sikang-Yunnan Province45. Northern Burmese Province46. Eastern Himalayan Province47. Khasi-Manipur Province

North American Atlantic Region

48. Appalachian Province49. Great Lakes and Central Lowlands

Province50. Prairies Province51. Coastal Plains Province52. Texas Prairies Province

Rocky Mountain Region

North-Western Pacific Subregion

53. Boreal Oceanic Alaskan Province54. Cascade Province

Rocky Mountain Subregion

55. Northern Rocky Mountain Province56. Central Eastern Rocky Mountain

Province


TETHYAN SUBKINGDOM

Mediterranean Region

Western Mediterranean Subregion

57. Aragonese Province58. Valencian-Catalan-Provencal Province59. Balearic Province60. Castilian-Maestracensean-Manche

Province61. Murcian-Almeriensian Province62. Carpetan-Iberian-Leonese Province63. Lusitan-Extremadurean Province64. Gaditan-Onubensean-Algarvian Province65. Betic Province66. Corse-Sardean Province67. Sıcula Province68. Ligurio-Romano-Calabrica Province69. South-western Mediterranean Province70. Southern Moroccan Province

Eastern Mediterranean Subregion

71. Puglica Province72. Etolico-Epirota Province73. Peloponnesian Province74. Crete Province75. Tracio-Tesalica Province76. Aegean Province77. Cirenico-North African Superprovince78. Steppe Eastern African Superprovince

Canarian Subregion

79. Eastern Canarian Province80. Western Canarian Province81. Madeirense Province

Saharo-Arabian Region (Saharo-Sindian)

82. Saharian Subregion83. Arabian Subregion

Irano-Turanian Region

Western Asiatic Subregion

84. Central Anatolian Province85. Armeno-Iranian Province86. Mesopotamian Province87. Turanian or Aralo-Caspian Province88. Hyrcanian Province89. Turkestanian Province90. Northern Baluchistanian Province91. Western Himalayan Province

Central Asiatic Subregion

92. Central Tien Shan Province93. Dzungano-Tien Shan Province

94. Mongolian Province95. Tibetan Province

MADREAN SUBKINGDOM

Californian Region

96. Northern Californian Province97. Southern Californian Province

Great Basin Region

98. Columbian Plateau Province99. Intermountain Province

100. Colorado Plateau Province101. Mohave Province102. Neomexican-Arizonian Middlands Province

Mexican Xerophytic Region

103. Baja Californian Province104. Sonoran Province105. Sinaloan Province106. Chihuahuan Province107. Tamaulipan Province

Madrean Region

108. Western Madrean Province109. Neovolcanic-Eastern Madrean Province

PALEOTROPICAL KINGDOM

AFRICAN SUBKINGDOM

Guinean-Congolian Region

110. Upper Guinea Province111. Nigerian-Cameroonian Province112. Congolian Province

Uzambara-Zululand Region

113. Zanzibar-Inhambane Province114. Tongoland-Pondoland Province

Sudano-Zambezian Region

Zambezian Subregion

115. Zambezian Province

Sahelo-Sudanian Subregion

116. Sahelian Province117. Sudanian Province

Eritreo-Arabian Subregion

118. Somalo-Ethiopian Province119. South Arabian Province120. Socotran Province


Omano-Sindian Subregion

121. Oman Province122. South Iranian Province123. Sindian Province

Karoo-Namib Region

124. Namib Province125. Namaland Province126. Western Cape Province127. Karoo Province

St. Helena and Ascension Region

128. Ascension Islands Province129. Province of St. Helena

MADAGASCAN SUBKINGDOM

Madagascan Region

130. Eastern Madagascan Province131. Western Madagascan Province132. Southern and Southwestern Madagascan

Province133. Comoro Province134. Mascarene Province135. Seychelles Province

INDOMALESIAN SUBKINGDOM

Indian Region

136. Sri Lanka Province137. Malabar Province138. Deccan Province139. Upper Gangetic Plain Province140. Bengal Province

Indo-Chinese Region

141. South Burmese Province142. Andamanese Province143. South Chinese Province144. Thailandian Province145. North Indochinese Province146. Annamese Province147. South Indochinese Province

Malesian Region

Malesian Subregion

148. Malay Province149. Kalimantan (Bornean) Province150. Philippinean Province151. Sumatran Province152. South Malesian Province

Papuan Subregion

153. Celebesian (Sulawesian) Province154. Moluccan Province

155. Papuan Province156. Bismarckian Province

Fijian Region

157. New Hebridean Province158. Fijian Province

POLYNESIAN SUBKINGDOM

Polynesian Region

159. Micronesian Province160. Polynesian Province

Hawaiian Region

161. Hawaiian Province

NEOCALEDONIAN SUBKINGDOM

Neocaledonian Region

162. Neocaledonian Province

NEOTROPICAL KINGDOM

CARIBBEAN-AMAZONIAN SUPERREGION

Caribbean-Mesoamerican Region

163. Floridan Province164. Balsas River and Southern Madrean Prov-

ince165. Cuba Province166. Antilles Province167. Veracruzan-Yucatanian Province168. Chiapas-Honduras Province169. Panama and Costa Rica Province170. Guajiran Province

Colombian-Venezuelan Region

171. Ecuatorian-Colombian Coastal Province172. Magdalenan River Province173. Galapagos Islands Province174. Venezuelan Highland Province175. Llanos Province176. Tepuis Province

Amazonian Region

177. Loreto Province178. Negro River and High Orinoco Province179. Roraima and Lower Amazonas Province180. Guiana Province181. Amazonas Delta Province182. Acre and Madre de Dios Province183. Madeira and Tapajoz Province

CHACO-BRAZILIAN SUPERREGION

Brazilian-Paranean Region

184. Cerrado Province185. Tocantins Province


186. Beni Province187. Pantanal Province188. Atlantic Brazilian Province189. Paranean Province190. Caatinga Province

Chaco Region

191. Boreal Chaco Province192. Austral Chaco Province193. Andean Chaco Province

ANDEAN SUPERREGION

Andean Region

194. Paramo Province195. Peruvian and Bolivian Yunga Province196. Peruvian Puna Province197. Bolivian Interandean Province198. Andean Altiplano Province

Desertic Pacific Region

199. Peruvian Desert Province200. Atacama Desert Province

CAPE KINGDOM

Cape Region

201. Cape Province

AUSTRALIAN KINGDOM

North-east Australian Region

202. North Australian Province203. Queensland Province204. South-east Australian Province205. Tasmanian Province

South-west Australian Region

206. South-west Australian Province

Central Australian or Eremaean Region

207. Eremaean Province

HOLANTARCTIC KINGDOM

Pampean Region

208. Rainy Pampean Province209. Xeric Pampean Province

Middle Chile-Patagonian Region

210. Middle Chile Desert Province211. Central Chile Province212. Mediterranean Andean Province213. Monte Province214. Boreal Patagonia Province215. Austral Patagonia Province

Valdivian-Magellanic Region

216. Valdivian Province217. Austral Andean-Magellanic Province218. Tierra del Fuego Province219. Juan Fernandez Islands Province220. Antarctic Province

Region of the South Subantarctic Islands

221. Tristan-Goughian Province222. Kerguelenian Province

Neozeylandic Region

223. Lord Howean Province224. Norfolkian Province225. Kermedecian Province226. Northern Neozeylandic Province227. Central Neozeylandic Province228. Southern Neozeylandic Province229. Chathamian Province230. New Zealand Subantarctic Islands Province


Appendix 2. Matrix of 66 Peltigera species � 230 biogeographic provinces (see Appendix 1). The matrix shows the abundance on a five-step scale (1low, 5 high) of each species in each biogeographic province

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1 0 4 0 0 0 3 0 0 0 0 2 0 1 4 0 0 0 0 1 0 0 0 0 0 2 0 1 2 0 0 0 3 4 1 4 0 1 0 1 2 2 0 1 0 0 0 0 0 0 2 2 2 0 0 0 3 0 3 1 0 0 0 0 0 0 32 0 2 0 0 1 2 0 0 0 0 2 0 0 3 0 0 0 0 0 0 0 0 0 0 0 0 2 2 0 0 0 2 3 1 3 0 1 0 0 3 1 0 0 0 0 0 0 0 0 1 2 3 0 0 0 3 0 1 1 0 0 0 0 0 0 23 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 04 0 4 0 0 0 2 0 0 0 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 2 0 2 0 1 0 0 1 1 0 1 0 0 0 0 0 0 2 1 1 0 0 1 3 0 2 0 0 0 0 0 0 0 15 0 4 0 0 1 4 0 0 0 0 3 0 2 4 0 0 0 0 2 0 0 0 0 0 1 3 1 1 0 0 0 2 3 0 3 0 2 0 0 2 3 0 0 0 0 0 0 0 0 3 2 4 0 0 1 4 0 3 1 0 0 0 0 0 0 36 0 1 0 0 0 1 0 0 0 0 0 0 1 2 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 2 0 1 0 1 0 0 1 1 0 0 0 0 0 0 0 0 1 1 2 0 0 0 2 0 0 0 0 0 0 0 0 0 17 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 2 0 1 0 0 0 0 0 0 0 08 0 1 0 0 0 1 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 1 0 0 0 0 0 0 0 19 0 1 0 0 0 1 0 0 0 0 1 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1

10 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 011 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 012 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 013 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 014 0 5 0 0 0 3 0 0 0 0 1 0 1 4 0 0 1 0 2 0 2 0 0 0 0 2 1 0 0 0 0 2 3 0 3 0 2 0 0 2 2 0 0 0 0 0 0 0 0 3 0 2 0 0 1 4 0 2 0 0 0 0 0 0 0 115 0 5 0 0 0 4 0 0 0 0 1 0 1 2 0 0 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 2 3 0 3 0 1 0 0 1 1 1 0 0 1 0 0 0 0 3 1 1 0 0 0 4 0 3 1 0 0 0 0 0 0 216 0 2 0 0 0 4 0 0 0 0 4 0 1 2 0 0 0 0 2 0 0 0 0 0 0 3 1 1 0 0 0 1 2 0 2 0 1 0 1 2 1 0 0 0 0 0 0 0 0 1 1 3 0 0 0 3 0 0 0 0 0 0 0 0 0 317 0 4 0 0 0 4 0 0 0 0 4 0 4 4 0 0 0 0 4 0 0 0 0 0 0 4 1 3 0 0 0 4 5 0 3 0 2 0 1 3 3 0 0 0 0 0 0 0 0 5 2 5 0 0 0 5 0 1 0 0 0 0 0 0 0 418 0 3 0 0 0 3 0 0 0 0 3 0 1 2 0 0 0 0 1 0 0 0 0 0 0 4 2 0 0 0 0 1 3 0 2 0 3 0 1 2 1 0 0 0 0 0 0 0 0 4 1 3 0 0 0 4 0 1 0 0 0 0 0 0 0 319 0 2 0 0 1 3 0 0 0 0 3 0 3 2 0 0 0 0 4 0 0 0 0 0 0 3 1 2 0 0 0 2 3 0 2 0 3 0 2 3 1 0 0 0 0 0 0 0 0 3 2 4 0 0 0 4 0 0 0 0 0 0 0 0 0 320 0 1 0 0 0 3 0 0 0 0 2 0 3 4 0 0 0 0 1 0 0 0 0 0 0 4 4 0 0 0 0 2 3 0 3 0 3 0 0 3 1 0 0 0 0 0 0 0 0 3 2 3 0 0 1 3 0 2 0 0 0 0 0 0 0 321 0 1 0 0 1 2 0 0 0 0 2 0 1 3 0 0 0 0 1 0 0 0 0 0 0 2 3 0 0 0 0 1 2 0 2 0 2 0 0 3 1 0 0 0 0 0 0 0 0 2 1 4 0 0 0 4 0 2 0 0 0 0 0 0 0 222 0 1 0 0 1 3 0 0 0 0 1 0 1 3 0 0 0 0 0 0 0 0 0 0 0 3 3 0 0 0 0 1 1 0 1 0 3 0 0 2 1 0 0 0 0 0 0 0 0 2 1 3 0 0 0 2 0 1 2 0 0 0 0 0 0 123 0 0 0 0 2 2 0 0 0 0 3 0 1 3 0 0 0 0 1 0 0 0 0 0 0 4 4 0 0 0 0 1 3 0 1 0 3 0 0 3 0 0 0 0 0 0 0 0 0 1 1 3 0 0 0 3 0 1 0 0 0 0 0 0 0 224 0 0 0 0 1 2 0 0 0 0 3 0 1 2 0 0 0 0 0 0 0 0 0 0 0 3 2 0 0 0 0 0 1 0 2 0 3 0 0 1 0 0 0 0 0 0 0 0 0 1 1 3 0 0 0 2 0 0 0 0 0 0 0 0 0 025 0 0 0 0 1 3 0 0 0 0 3 0 2 1 0 0 0 0 2 0 0 0 0 0 0 4 3 1 0 0 0 1 1 0 1 0 4 0 2 2 2 0 0 0 0 0 0 0 0 2 2 4 0 0 0 4 0 0 0 0 0 0 0 0 0 126 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 1 0 0 1 0 0 0 0 0 0 2 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 027 0 1 0 0 0 2 0 0 0 0 1 0 1 2 0 0 0 0 1 0 0 0 0 0 0 2 1 0 0 0 0 1 1 0 1 0 1 0 1 2 2 0 0 0 0 0 0 0 0 2 1 3 0 0 0 2 0 1 0 0 0 0 0 0 0 128 0 1 0 0 0 1 0 0 0 0 1 0 2 3 0 0 0 0 1 0 0 0 0 0 0 3 1 0 0 0 0 2 2 0 1 0 1 0 0 1 1 0 0 0 0 0 0 0 0 2 1 3 0 0 0 2 0 0 0 0 0 0 0 0 0 329 0 2 0 0 0 1 0 0 0 0 1 0 1 2 0 0 0 0 1 0 0 0 0 0 0 2 0 0 0 0 0 1 1 0 1 0 1 0 0 1 1 0 0 0 0 0 0 0 0 1 0 2 0 0 0 1 0 0 0 0 0 0 0 0 0 130 0 1 0 0 0 2 0 0 0 0 1 0 1 2 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 2 0 0 0 0 0 0 0 0 0 131 0 1 0 0 0 2 0 0 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 1 2 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 2 0 2 0 0 0 2 0 0 0 0 0 0 0 0 0 132 0 0 0 0 0 2 0 0 0 0 2 0 1 1 0 0 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 0 1 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 1 0 4 0 0 0 1 0 0 0 0 0 0 0 0 0 133 0 0 0 0 0 1 0 0 0 0 2 0 1 0 0 0 0 0 0 0 0 0 0 0 0 3 0 0 0 0 0 0 3 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 1 1 3 0 0 0 1 0 0 0 0 0 0 0 0 0 134 0 0 0 0 0 1 0 0 0 0 2 0 1 1 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 2 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 1 1 2 0 0 0 1 0 0 0 0 0 0 0 0 0 135 0 1 0 0 0 1 0 0 0 0 1 0 1 1 0 0 0 0 1 0 1 0 0 0 0 1 1 0 0 0 0 1 1 0 1 0 1 0 0 1 1 1 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 1 0 0 0 0 0 0 0 0

2003D

istributionof

thegenus

Peltigera

Willd.—

Martiınez

etal.

315

Appendix 2 Continued.

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Accepted for publication 7 May 2003


distribution patterns in the genus peltigera...

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