A. Kurtto, P. Uotila & A. Sennikov Alchemilla in Mediterranean Europe as revealed by Atlas Florae Europaeae Abstract Kurtto, A., Uotila, P. & Sennikov, A.: Alchemilla in Mediterranean Europe as revealed by Atlas Florae Europaeae. — Bocconea 23: 221-235. 2009. — ISSN 1120-4060. A phytogeographical analysis of the Alchemilla (Rosaceae) flora of Mediterranean Europe is presented on the basis of maps in Atlas Florae Europaeae vol. 14. In Mediterranean Europe, Alchemilla species are confined to the mountains, and 92% of them (346 species) are endemic to Europe. Many of them have very small ranges, limited to one or to only a few 50 km × 50 km AFE grid cells. The known diversity and endemism are especially high in the Alps (149 species, of which 61% are endemics) and in the Cordillera Cantábrica – Pyrenees (79 species, 46% endemics). The Mediterranean is therefore well represented in the country-specific totals for Alchemilla species, with three countries in the European top ten: (1st) France 121, (3rd) Italy 92 and (6th) Spain 81. Many of the species concentrated in the Alps have additional stations further south along the Apennines, but only 3 species are known to be endemic to the Apennines. The Balkan Alchemilla flora (60 species) includes the southernmost outposts of many of the Alpine and/or Carpathian species. In total there are 24 Balkan endemics (40%). In Crimea 16 of the known 20 species are endemic to the territory. Introduction Atlas Florae Europaeae (AFE) is a programme for mapping native and naturalized vascular plants of Europe. Furthermore, information on taxonomy, nomenclature, chromosome numbers, biosystematics and total ranges of the taxa are added alongside the maps. For the history of the project and the mapping system, see Uotila & al. (2005) and Kurtto & al. (2004). AFE volume 14, published in late 2007 (Kurtto & al. 2007), covers the genera Alchemilla and Aphanes. Reproduction mode in Alchemilla is almost obligate apomixis; according to established tradition the smallest taxonomic entities recognised by constant differences in morphology are given specific rank there. Alchemilla has 433 species in Europe (231 in Flora Europaea; Tutin & al. 1968) and well over 1 000 in the world. Most species are confined to various types of meadow habitats; on lowlands many of them are apophytes with uncertain natural limits of distribution areas, whereas in montane regions they are less dependant on the human activity. In AFE 14 the genus is mapped for the first time in its entirety on a European scale. There are many possible approaches to the analysis of the AFE dataset, including numerical classifications of distributions and regional comparisons. The present paper is the first attempt to use the AFE 14 data in such a way. 222 Kurtto & al.: Alchemilla in Mediterranean Europe as revealed by Atlas ... Floristic elements of the Alchemilla flora of Mediterranean Europe Due to the fact that in Mediterranean Europe the genus Alchemilla is restricted to high mountains, the following chorological classification is principally based on presence or absence of species in distinct mountain ranges or their divisions. The elements have been arranged in a sequence roughly running west to east. Only records of native species and only records of native occurrences with certain identifications and localities are accepted as basis for the classification and comparison, but dubious records are discussed when they may, if substantiated, change the classification. Taxonomy and nomenclature follow Kurtto & al. (2007). Delimitation and subdivision of the mountain ranges follow Tutin & al. (1968), except that in this paper the southwestern Alps include S. Jura and the central Alps include the rest of Jura and Vosges. Iberian mountains 55 species (12.8%) of European Alchemilla are confined to the Iberian mountains (including French Pyrenees; Fig. 1). 27 of these species are endemic to the Pyrenees, some being present all along the range, others restricted to shorter or longer stretches of it (for details, see Fröhner 1998). The number of endemics of the Cordillera Cantàbrica is 11, and eight Iberian endemics are present only in the Pyrenees and Cordillera Cantàbrica. Thus, the pooled number of endemics present on the two ranges alone is 46. Alchemilla atropurpurea is concentrated in the Pyrenees but has disjunct occurrences over 400 km to the south and southwest in the mountains of Avila and León. A. spectabilior has occurrences in both the Cordillera Cantàbrica and Pyrenees, as well as outside them Fig. 1. Species richness for the Alchemilla species confined to the Iberian mountains (incl. French Pyrenees; N=55). Fig. 2. Verified distribution of Alchemilla coriacea according to Kurtto et al. (2007). Bocconea 23 — 2009 223 in Sierra de la Demanda some 100 km south of the eastern end of the Cordillera Cantàbrica. A. hypercycla and A. perspicua are present in Sierra de Satrústegui in addition to sites about 100 km further north in the Pyrenees and Cordillera Cantàbrica, respectively. The mainly Pyrenean A. oscensis has a disjunct additional outpost in the mountains of Castellón near the east coast, c. 250 km south of the Pyrenees. Four Iberian endemics are present only in mountains south of the Cordillera Cantàbrica and Pyrenees: A. atriuscula in Sierra de Andía less than 40 km south of the western end of the Pyrenees, A. crenulata in the mountains of León just south of the western end of the Cordillera Cantàbrica, A. serratisaxatilis in Sierra de Béjar and Sierra de Guadarrama in the central inland region, and A. font-queri in the Sierra Nevada. Mountains from N.W. Spain to the Alps, Apennines or Balkan Peninsula The range of Alchemilla hybrida (syn. A. lapeyrousii) extends from the Cordillera Cantàbrica and the mountains of Castellón to the western central Alps, with core areas in the Pyrenees and French Massif Central, whereas A. inconcinna is mainly a species of the Cordillera Cantàbrica – Pyrenees range and western central Alps but is also present in the Massif Central. The range of A. lucida has its core in the southwestern Alps, and only widely scattered occurrences are known from other mountains from the Cordillera Cantàbrica to the central Italian Alps. The ranges of five species extend from N.W. Spain to the Alps and further to the Apennines. Alchemilla coriacea has a wide and coherent distribution in and around the Alps (north to the Vosges and Schwarzwald; Fig. 2) and in the Massif Central, complemented by numerous occurrences in N. Spain in the Pyrenees and further westwards to the mountains of León and southwestwards to the mountains of Avila. Furthermore, the Fig. 3. Verified native distribution of Alchemilla Fig. 4. Species richness for the Alchemilla species nitida according to Kurtto et al. (2007). endemic to the Alps (N=91). 224 Kurtto & al.: Alchemilla in Mediterranean Europe as revealed by Atlas ... species is known from the highest peaks of central Apennines, some 500 km from the nearest stations in the Alps. The range of A. saxatilis is quite similar in the Iberian Peninsula and Massif Central, but the species is also present far to the south in the Sierra Nevada. In the east, the species reaches its northern limit in southern Switzerland and its southern limit in the Ligurian and Tuscan-Emilian Apennines. A. saxatilis was once present also in Corsica. A. transiens has a more scattered distribution than the two previous species, but it extends from Sierra de Guadarrama and the Cordillera Cantàbrica to the eastern Alps, Corsica and the Apennines (Ligurian and Tuscan-Emilian Apennines, as well as Abbruzzi). A. alpigena is widely present in the Cordillera Cantàbrica, Pyrenees and Alps, but absent from the Massif Central, and has additional stations in the Ligurian and Tuscan-Emilian Apennines. Alchemilla lunaria has a peculiar distribution comprising central parts of the Cordillera Cantàbrica, the Pyrenees and central parts of the Alps, as well as one very disjunct (by more than 1000 km) station in western Rodopi Planina of Bulgaria. Mountains from the Pyrenees to the Alps or Apennines Five of the eight species of this element have intermediary stations in the Massif Central of France, viz. A. demissa, A. rubristipula, A. tenuis, A. trunciloba and A. vetteri. A. tenuis is also present in Sierra Cebollera southwest of the western Pyrenees and A. vetteri in mountains of Castellón in E. Spain (for details of the distribution of the latter species, see Kalheber 1982). The ranges of A. demissa, A. tenuis and A. trunciloba extend even to the Apennines as scattered occurrences. A. conjuncta, A. pentaphyllea and A. tenerrima are absent from the Massif Central and Apennines and thus present only in the Pyrenees and Alps. A. conjuncta is principally a species of the western central Alps and A. pentaphyllea a species of the southwestern and central Alps. Both have only one verified station in the Pyrenees (Pyrénées Atlantiques of France and Valle de Boí of Spain, respectively). A. tenerrima is a rare species of the eastern Pyrenees and southwestern Alps (one station in Turin, Italy). The elements of the three first groups seem to belong to the Erysimum duriaei element of the scheme of Finnie & al. (2007) based on a wide selection of species mapped in AFE. Massif Central of France and its connections to the east Only one species, Alchemilla grenieri, is certainly endemic to the Massif Central, since records of the Alpine A. amphisericea from the Pyrenees may be referable to A. charbonneliana, which has hitherto been considered endemic to the Massif Central. A. saxetana, a species previously known only from a couple of localities in Unterwallis, Switzerland, was found in the Massif Central in 2001. A. nitida and A. pallens both show in their distributions unique combinations of mountain ranges. A. nitida is known from the Massif Central, Alps, Vosges and Schwarzwald, as well as along the Apennines as far to the south as northern Calabria (Fig. 3). A. pallens shows a rather similar distribution pattern, but has only one station in the Apennines (Tuscan-Emilian Apennines) and a remarkably disjunct (by c. 1000 km) occurrence in the Pirin Planina of Bulgaria. Bocconea 23 — 2009 225 Endemics of the Alps or/and Apennines The Alps are extremely rich in Alchemillae: a total of 149 species, of which no less than 91 are endemic (Fig. 4; for further details, see the regional comparison below and Fröhner 1990). 52 of the endemic species of the Alps either extend to Mediterranean Europe or are endemic to it (e.g. the recently described endemics of the Italian Alps A. federiciana, A. lasenii and A. nydeggeriana; see Fröhner 2005). Only three species are endemic to the Apennines: A. ceroniana of the Tuscan-Emilian Apennines, A. marsica of the Abruzzo, and A. austroitalica of Aspromonte (Fig. 5). The total ranges of eight species are confined to the Alps and Apennines. A. compta and A. hoppeana occur discontinuously almost all along the Alps and have only a single known locality in the Apennines (Abruzzo and Tuscan-Emilian Apennines, respectively). A. subserica has a more continuous distribution from the Maritime Alps to the Ötztaler Alpen of W. Austria and two widely disjunct stations in the Apennines. A. strigosula is included in this element, though its area extends in the north from the Alps proper to the lower mountains of S. Germany and it may also be present in the Massif Central and even in the Pyrenees. In the Apennines, the species is present in Liguria and in a rather wide area from Marche in the north to Abruzzo in the south. A. undulata is mainly a plant of the eastern Alps, but it has fairly numerous occurrences in the central Alps and disjunctly in the Abruzzo, as well as a few localities in the northern parts of the soutwestern Alps (Fig. 6). Records of the species from the central Balkans are dubious. Alchemilla cinerea (excl. A. lanuginosa) is confined to the southwestern Alps, TuscanEmilian Apennines and Abruzzo. A. sinuata is principally a plant of the central Alps but has very disjunct localities in the Tuscan-Emilian Apennines and far to the south in Monte del Papa of Basilicata. According to the current view, A. cataractarum seems to be restricted to the eastern Alps and Tuscan-Emilian Apennines. The species treated in this section, as well as many of the mountain species of the fol- Fig. 5. Distribution of the Apenninian endemics Fig. 6. Verified distribution of Alchemilla undulaof Alchemilla. ta according to Kurtto et al. (2007). 226 Kurtto & al.: Alchemilla in Mediterranean Europe as revealed by Atlas ... lowing sections, belong to the Salix serpillifolia element of Finnie & al. (2007). However, mountain species with easterly bias seem to be best included in their Rumex alpinus element, Dianthus moesiacus element, or Ranunculus psilostachys element. Alps and the Balkan Peninsula A large number of Alchemillae with their main ranges in the Alps have been recorded from the Balkan Peninsula. However, most of the records are erroneous or at least dubious. The floristic connection between the Alps and the Balkan Peninsula, as reflected by Alchemilla, is therefore much weaker than was previously thought. Perhaps only six species have ranges which cover exclusively parts of the Alps and parts of the Balkan mountains, but even the taxonomy and chorology of this small group is much in need of further elucidation. A. croatica is a poorly understood species with very scattered localities in the Italian Alps, northwestern Croatia and central Bosnia-Herzegovina. A. obtusa belongs to a taxonomically very intricate species group and therefore its exact distribution is not known. It may be a plant of only the Alps and the highest mountains of the central parts of the Balkan Peninsula. A. lineata has a wide range in the central and eastern Alps and perhaps only one station in the Balkan Peninsula (Prokletije). A. racemulosa is known from widely scattered stations along the Alps and a single locality in BosniaHerzegovina. A. venosula was earlier regarded as endemic to the eastern Alps but the plant of the central Balkans called A. gracillima Rothm. is now considered conspecific with it. Alchemilla exigua is principally a plant of the central and eastern Alps but has scattered localities in the central Balkans. The species is also recorded from the Romanian Carpathians, but the identification of this material should be checked. Balkan Peninsula Currently, 24 Alchemilla species endemic to the Balkan Peninsula are known (Fig. 7). However, the actual number of endemics is evidently much higher, since Fröhner (1999) estimated that about 50 new species remain to be discovered in the area. Alchemilla velebitica and A. amphiargyrea are restricted to the western mountains of the Balkan Peninsula, the former species extending from southernmost Slovenia to southern Pindhos of Greece, the latter from southern Bosnia-Herzegovina to Sterea Ellas of Greece. A. malyi and A. vranicensis are endemic to the mountains of Bosnia-Herzegovina. The range of A. lanuginosa covers mountains from southern Bosnia-Herzegovina and Albania through southern Serbia, F.Y.R.O. Macedonia and the northernmost mountains of Greece to southwestern Bulgaria. The distribution of A. heterophylla is fairly similar but reaching central Croatia in the northwest and southern Pindhos in the southeast. The species is present on Anatolian mountains, too, and thus does not belong to the Balkan endemics proper. The high Prokletije mountains on the borders of Albania, Montenegro, Serbia and F.Y.R.O. Makedonia host several local endemic species of Alchemilla: A. albanica, A. bertiscea, A. montenegrina Plocek [nomen illeg.], A. rubidula and A. vincekii (see Plocek 1998). In central parts of the Balkan Peninsula, A. bulgarica, A. catachnoa, A. heterotricha, A. Bocconea 23 — 2009 227 indivisa and A. viridiflora have fairly wide distributions, but only the range of A. bulgarica appears continuous. A. indivisa extends farthest to the south, reaching northern Peloponnisos, and all the five species attain their northern limit in the southern half of Serbia. A. bandericensis (Jakupica, Pirin Planina), A. damianicensis (Sar Planina, Pirin Planina), A. peristerica (Perister) and A. pirinica (Sar Planina and surroundings, Pirin Planina) are much rarer endemics of the central Balkans. A. pawlowskii is restricted to the Rila Planina, and A. achtarowii, A. jumrukczalika and A. sirjaevii are known only from the Stara Planina. Alchemilla aroanica, a local endemic of the Chelmós mountains of northern Peloponnesos, is, with A. font-queri of Sierra Nevada and A. austroitalica of Aspromonte, the southernmost of the European endemics in the genus. Carpathians and the Balkan Peninsula The floristic connection between the Carpathians and the Balkan Peninsula, as reflected by Alchemilla species restricted to the two mountain chains, is roughly as weak as that between the Alps and the Balkan Peninsula, since only seven species seem to have a Carpathian – Balkan total range. Alchemilla gorcensis is centred in the central Balkans and the western Carpathians (Tatra Mts. and surroundings) and has only two known localities between the centres in the highest mountains of the Romanian and Ukrainian Carpathians (Fig. 8). A. obsoleta completely lacks such intermediary stations by being present only in the western Carpathians and highest mountains of Bulgaria. A parallel disjunction is shown by A. czywczynensis, restricted to the high mountains of the eastern Carpathians on both sides of the Ukrainian – Romanian border and to the Rodopi Planina of southern Bulgaria, and by A. zapalowiczii, which is possibly extinct in the Tatra Mts. but certainly present in the same part of Fig. 7. Species richness for the Alchemilla species Fig. 8. Distribution of Alchemilla gorcensis endemic to the Balkan Peninsula (N=24). according to Kurtto et al. (2007). 228 Kurtto & al.: Alchemilla in mediterranean Europe as revealed by Atlas ... the eastern Carpathians as the previous species and additionally in Jakupica of F.Y.R.O. Macedonia. A. serbica is widespread in the central Balkans and also seems to have one locality in the southern Carpathians. A. asteroantha, which is still in need of taxonomic clarification, is mainly a species of Stara Planina of Bulgaria but has also been recorded from the same Romanian mountain area as A. serbica. Alchemilla mollis, which is widely cultivated for ornament and widely naturalized in western and central Europe, also belongs to the species restricted to the Romanian Carpathians and the Balkan Peninsula (Stara Planina and northern Greece) in Europe. However, this species is also native to Anatolia, the Caucasus and Iran. Crimea The Crimean mountain flora includes a total of 20 species of Alchemilla. At least 16 of them are Crimean endemics. A. aemula and A. taurica are present also in Caucasia, and A. lithophila and A. stevenii have been reported also from northern Anatolia. The Crimean species belong to the Dianthus capitatus element of Finnie & al. (2007) Widespread mountain species Alchemilla colorata, A. connivens, A. fissa, A. flabellata and A. straminea belong to the floras of all or almost all the principal mountain chains of South and Central Europe. Of the five species, A. flabellata has the most coherent distribution, though with notable gaps in the western Cordillera Cantàbrica and western Pyrenees (Fig. 9). The range of A. connivens is of the same kind but more scattered especially in the Carpathians and the Balkans. A. colorata is not known from the Cordillera Cantàbrica and has only a few stations in the Carpathians, whereas A. fissa is commoner in the Carpathians but absent from the Massif Central and almost absent from the Apennines. A. straminea has an extensive distribution centre in the Alps, but the species is also present in all other principal mountain areas, as well as in the Sierra Nevada, several sierras of the northern inland region of Spain, Bóhmerwald and Krkonose. Alchemilla efusa and A. fallax differ notably from all the five above-mentioned species in being totally absent from the Carpathians and from all the five except A. fissa in being absent also from the Massif Central. A. crinita, A. incisa (s. lato) and A. reniformis in turn are absent from the Iberian mountains (incl. French Pyrenees). A. crinita is known also from Morvan, a northerly extension of the Massif Central, but the other two are absent from the Massif Central too. The distribution of A. crinita probably also includes northeastern Anatolia, although the Anatolian populations show slight differentiation from the European ones. Widespread species with their southernmost localities in Mediterranean Europe Quite a number of the Alchemilla species of the temperate lowland areas and medium altitudes of more northern European countries reach Mediterranean Europe as occurrences in the mountains. However, we do not know where each species originally evolved, and Bocconea 23 — 2009 229 Fig. 9. Verified distribution of Alchemilla flabellata according to Kurtto et al. (2007). therefore it is possible, and due to the Pleistocene history of Europe in many cases even probable, that the modern distributions of these species are the result of northward migrations, with or without man’s help. Therefore, the following account (and the above accounts) must be considered merely descriptive and do not imply any particular distributional history. The distribution of A. xanthochlora (Fig. 10) is unique in its suboceanic and nemoral to boreonemoral nature and in at least approaching the Ranunculus bulbosus element of Finnie & al. (2007). In the south, the species is present in all the principal mountain ranges and extends as far south as Somosierra of the Sistema Central in Spain, Monte Pollino of Basilicata in Italy and Sterea Ellas in Greece. The distribution of A. subglobosa is also the only one of its kind. The species has a strong centre in southern Scandinavia, numerous localities in the Baltic Countries and a smaller centre in Bóhmerwald, Erzgebirge and surroundings. Towards the southwest its distribution ends at disjunct localities in the Savoie Alps in southeastern France and towards the south at a locality in the Dolomites of Italy. Of the numerous eastern boreonemoral species, only A. cymatophylla and A. propinqua reach Mediterranean Europe, the former with one locality in the Dolomites in Italy and one in the mountains of Primorska in southwestern Slovenia, the latter with a very disjunct population in the same area of the Dolomites where the former is present. Fröhner (1965) deliberated upon the history of the very similar overall distributions of the two species. The seven most widespread species of the genus in Europe – A. monticola, A. micans, A. subcrenata, A. acutiloba, A. glabra, A. filicaulis and A. glaucescens – are also members of the mountain flora of Mediterranean Europe. A. filicaulis and A. glabra have a prominently oceanic type of distribution in northwestern Europe, and at least the first-mentioned species belongs to the amphiatlantic element (A. glabra is better tending to elevated areas in Northern Europe and is probably introduced in northeastern North America). Both species extend as far south as Sierra Nevada and the central Apennines. A. filicaulis is almost absent from the Balkan Peninsula, but A. glabra has numerous localities there, the 230 Kurtto & al.: Alchemilla in Mediterranean Europe as revealed by Atlas ... Fig. 10. Verified native distribution of Alchemilla xanthochlora according to Kurtto et al. (2007). southernmost in Sterea Ellas. The other five species of the top seven have more continental distributions and clearly belong to the Eurosiberian element or, on the scheme of Finnie & al. (2007), best fit the Lychnis flos-cuculi element. The southern limits of these species vary somewhat, as follows: A. acutiloba has a very disjunct occurrence in the central Pyrenees but otherwise the southern limit runs from southernmost fringes of the Alps to Sterea Ellas; A. glaucescens, a more xerophilous and thermophilous species, reaches the Cordillera Cantàbrica and Pyrenees in the west, Monte Pollino of Basilicata in the Apennines and the northernmost mountains of Greece in the east; A. micans stops on the Alps in the west, but extends to Sterea Ellas in the east; A. monticola reaches the Pyrenees (only one verified locality), the Apennines and the northern mountains of Greece; A. subcrenata extends to the Massif Central, central Apennines and Rodopi Planina. A. plicata is endemic to the nemoral – southern boreal zones of Europe. It reaches Mediterranean Europe as only one rather disjunct station in the southwestern Alps in the Italian province of Cuneo. The amphiatlantic A. glomerulans is widespread in the boreonemoral to arctic zones of Bocconea 23 — 2009 231 northwestern Europe. It is also present in many of the principal mountains of central and southern Europe, including the Cordillera Cantàbrica, Pyrenees, French, Italian and Slovenian Alps, and highest peaks of the Balkan group of mountains from Prokletije in the north to Jakupica in the south. A. alpina is also an amphiatlantic plant but of the arcticalpine type and belongs to the Oxyria digyna element of Finnie & al. (2007). In southern Europe, the distribution of A. alpina covers mountains from the Cordillera Cantàbrica and Sistema Central of Spain in the west to western Austria and highest parts of the Apennines in the east. The species is also one of four found in the mountains of Corse. 3. Regional comparisons Out of the total of 430 native European species of Alchemilla, 374 (87%) are endemic to Europe (Table 1). The mountains of southern and central Europe host a total of 346 species, of which the proportion of endemics is as high as 92%. The Alps are richest in species, then come the Carpathians, especially the western Carpathians, and the Cordillera Cantàbrica – Pyrenees -range. The Alchemilla floras of the Pyrenees and mountains of the Balkan Peninsula have a quite similar amount of endemism. The Alchemilla floras of the Apennines and Massif Central are much poorer and less distinctive. The isolated and distinct mountain areas of Crimea have the most differentiated Alchemilla flora with 20 species, of which at least 80% are endemic. – The species richness for Alchemilla on the mountains of southern and central Europe closely parallels the taxon richness for 501 predominantly alpine species and subspecies mapped in the first 11 volumes of AFE (see Väre & al. 2003). Pair-wise similarities between the mountain ranges and their divisions (Table 2), based on all the native species of the genus, also show the relatively poor differentiation of the Alchemilla floras of the Apennines and the Massif Central and imply that the latter mountain area may have functioned as a ‘stepping stone’ in the immigration of species between the Alps and the Iberian mountains. The very low similarities between the Carpathians and the other ranges, as well as the fairly low similarities between the western Carpathians and the two other divisions of the Carpathians, are mainly due to the large number of endemics of the western Carpathians (mainly the Tatra Mts.). The high similarities between the divisions of the Pyrenees, as well as between the divisions of the Alps and between the eastern and southern Carpathians, indicate that the divisions as defined in Flora Europaea are better regarded as practical or descriptive rather than as phytogeographically justified regions. Comparisons based on smaller divisions combined with more accurate chorological data would certainly yield more meaningful results, especially for the Alps. – Using 501 alpine taxa mapped in AFE, Väre & al. (2003) calculated a Jaccard’s similarity index value of 25 between the Pyrenees and Alps. By using the Alchemilla species, the value is lower (16), which indicates that their ranges are, on average, smaller. This is also evident by visual inspection of the maps of Vàre & al. (2003) summarizing the pooled distributions of alpine plants present on each of the principal mountain ranges of Europe. The predominantly small ranges of Alchemillae are apparently largely due to the apomictic nature of the genus. Another way to compare species numbers and similarities of Alchemilla floras is to use 232 Kurtto & al.: Alchemilla in Mediterranean Europe as revealed by Atlas ... Bocconea 23 — 2009 233 countries as the geographical units. In the country-specific species numbers of Alchemilla, Mediterranean Europe is well represented in the richest end, with France as the winner and also Italy and Spain in the top 10 (Table 3; Kurtto & al. 2007). Regarding the numbers of country-specific endemics, Spain, France and Italy are among the eight richest countries. The similarity indices between countries (Table 4) are, on average, higher than those between mountain ranges, which simply reflects the facts that borders between countries often follow crests of mountain ranges and that plants do not need passports to cross the borders. The European Alchemilla flora includes a great number of local or regional endemics (e.g. Fröhner 2002). No less than 185 species are restricted to only one or two AFE grid cells with an average size of 50 km × 50 km, and 290 species are restricted to at most 10 such grid cells. The distribution of the latter species group (Fig. 11) neatly reveals the main European centres of endemism of the genus: the Cordillera Cantàbrica – Pyrenees -range, Alps (especially Unterwallis of Switzerland and Savoie of France), Jura, Tatra Mts., central parts of the eastern Carpathians, and Crimea. Regarding southern Europe, weaker centres appear especially in the Apennines and Balkan mountains. In comparison to the taxa with a narrow distribution area in the data set of 501 alpine taxa not including Alchemilla (Väre & al. 2003), the rarest European Alchemilla endemics are weakly represented in the Sierra Nevada, the southern Carpathians and some parts of the Balkan Peninsula, as well as on the Mediterranean islands, on which the genus is present only on Corsica. In the set of 501 taxa, about 10% of the species and subspecies occurred in a single AFE cell. In Alchemilla, the proportion is much higher, almost 35%. Finally, it has to be emphasized that all the above comparisons suffer from the still uneven exploration of the Alchemilla floras of various mountain areas. Numerous more or 234 Kurtto & al.: Alchemilla in Mediterranean Europe as revealed by Atlas ... Fig. 11. Number of European narrow endemics of Alchemilla in southern and central Europe. The map includes species recorded from at most 10 AFE grid cells (N=290). less local or regional endemics and also more widespread species of the genus still wait to be discovered or described from, at least, the Iberian and Balkan mountains (Fröhner 1999) and the Apennines (see Tondi 2001), not to mention the Urals (which are outside the scope of this article). Acknowledgements We are grateful to Raino Lampinen and Leena Helynranta for preparing the maps and Christopher Preston who kindly checked the language. Bocconea 23 — 2009 235 References Finnie, T. J. R., Preston, C. D., Hill, M. O., Uotila, P. & Crawley, M. J. 2007: Floristic elements in European vascular plants: an analysis based on Atlas Florae Europaeae. – J. Biogeogr. 34: 1848-1872. Fröhner, S. 1965: Die Frauenmantel-Arten Schleswig-Holsteins. – Die Heimat (Kiel) 72: 74-79. — 1990: Alchemilla. – Pp 13-242 in G. 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(Rosaceae) nell’Appennino centrale. – Inform. Bot. Ital. 33: 543-553. Tutin, T. G., Heywood, V. H., Burges, N. A., Moore, D. M., Valentine, D. H., Walters, S. M. & Webb, D. A. (ed.) 1968: Flora Europaea, 2. – Cambridge. Uotila, P., Suominen, J. & Lahti T. 2005: Erfahrung úber die langjährige Kartierung von Pflanzen: Atlas Florae Europaeae. – Hoppea, Denkschr. Regensb. Bot. Ges. 66: 119-132. Väre, H., Lampinen, R., Humphries, C. & Williams, P. 2003: Taxonomic diversity of vascular plants in the European alpine areas. – Pp. 133-148 in Nagy, L., Grabherr, G., Kórner, C. & Thompson, D.B.A. (ed.), Alpine Biodiversity in Europe – A Europe-wide Assessment of Biological Richness and Change. – Berlin, Heidelberg. Address of the authors: Arto Kurtto, Pertti Uotila, Alexander Sennikov, Botanical Museum, Finnish Museum of Natural History, P.O.Box 7, FI-00014 University of Helsinki, Finland. Email pertti.uotila@helsinki.fi, arto.kurtto@helsinki.fi, alexander.sennikov@helsinki.fi http:// www.fmnh.helsinki.fi/english/botany/afe