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A Morphometric Study on Anthyllis Vulneraria (Fabaceae) from Poland and Its Taxonomic Implications

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A Morphometric Study on Anthyllis Vulneraria (Fabaceae) from Poland and Its Taxonomic Implications Biologia 67/2: 296^309, 2012 Section Botany DOI: 10.2478/sl 1756-012-0013-1 A morphometric study on Anthyllis vulneraria (Fabaceae) from Poland and its taxonomic implications Kaja R o l a Department of Plant Taxonomy, Phytogeography & Herbarium, Institute of Botany, Jagiellonian University, Kopernika 27, PL-31-501 Krakow, Poland; e-mail: kajaskubalaQi.nteria.pl A bstract: The paper presents results of morphometric analysis of Anthyllis vulneraria L. in Poland based on 828 herbarium specimens. This study investigates live taxa which have been recognized in Poland: Anthyllis vulneraria L. ssp. vulneraria, Anthyllis vulneraria ssp. polyph.ylla (DC.) Nyman, A. vulneraria ssp. m aritim a (Schweigg.) Corb., A. vulneraria ssp. alpestris Asch. et Graebn. and A. vulneraria ssp. carpatica (Pant.) Nyman as well as three intermediate taxa of presumably hybrid origin. Multivariate statistical analyses of 12 quantitative and 10 qualitative characteristics revealed conspicuous variation within A. vulneraria. Analysis of variance (ANOVA), principal coordinate analysis (PCoA), correspondence analysis (CA) and discriminant analysis (DA) proved the taxonomic usefulness of quantitative characteristics such as number of rosette leaves, calyx length, inflorescence length, the ratio of leaf length/width, bract length as well as qualitative characteristics such as stem hairiness, calyx colour, calyx indumentum, distances between stem leaves and form of rosette leaves. The issues concerning intermediate morphotypes occurring in Poland and their status are discussed. Finally, a key for determination of tax a within A. vulneraria in Poland and distribution maps based on the material examined are provided. Key words: Anthyllis vulneraria', biometry; Fabaceae; morphological variability; multivariate statistics; Poland; systematic key; taxonomy Introduction ten morphologically indistinct (Juzepczuk 1945; Puidet et al. 2005). Most taxa are morphologically similar, Anthyllis L. belongs to the tribe Loteae (Polhill 1981; and some of them have presumably hybrid origin. The Allan & Porter 2000) within the family Fabaceae. An­ wide variation in morphology observed nowadays sug­ thyllis L. is taxonomically very complex. It consists of gests that there are a few taxa within A. vulneraria numerous taxa wherein the status of most of them re­ and intermediates, which could be commonly found on mains questionable. The number of species within the the border ranges. The species was probably divided genus has been previously reported to range from 25 into several taxa with distinct geographical ranges in (Cullen 1986) up to 60 (Minjaev & Akulova 1987). the past (Jalas 1950). This differentiation has not, Anthyllis vulneraria L. is a very polymorphic so far as is known, led to complete genetic isolation species, consisting of 30 intraspecific taxa (many of (Marsden-Jones & Turrill 1933). As a result of many them frequently recognized as species), between which factors such as climate change and the ability to oc­ intermediates occur, often over a large area (Cullen cupy disturbed habitats, there was a partial blurring 1968). Some researchers distinguish only one poly­ of the primary border-lines of particular morphotypes typic species - A. vulneraria (Stankov & Taliev 1949; as well as diagnostic morphological differences (Jalas Jalas 1950; Jalas 1952). On the other hand, Rothmaler 1950). Therefore, in spite of the fact that it is not so (1941) and Juzepczuk (1945) claim that there are sev­ widespread to recognize A. vulneraria as species com­ eral separate and more or less independent species dis­ plex, it should be considered in such a way, because the tinguishable in Europe. These inter-grading variants complexity of this taxon is evident (Krall 1983; Köster might be, to some extent, ecologically and geograph­ 2010). ically separated (Cullen 1986); however, their classifi­ Anthyllis vulneraria has wide distribution through­ cation is complicated. It should be noted here that in out most of Europe (Becker 1911) from the Atlantic spite of wide variability within A. vulneraria, chromo­ coast to the Baltic region; south to the Mediter­ some numbers of all the taxa examined in this study ranean Basin as well as in North Africa (Algeria, Mo­ are the same (2n = 12) (Bolkhovskikh et al. 1969; rocco, Ethiopia) (Bennett et al. 2001); east to SW Van Loon & Kieft 1980; Yeh et al. 1986; Dempsey Asia - mainly Turkey (Cullen 1976) and the Caucasus et al. 1994; Lövkvist & Hultgprd 1999), which means and Iran (Marsden-Jones & Turrill 1933). It has also they are diploid. Furthermore, A. vulneraria consists been introduced into North America and New Zealand of local morphotypes in limited areas, which are of­ (Hulten & Fries 1986). A morphometric study on Anthyllis vulneraria (Fabaceae) from Poland 297 There are many studies concerning morpholog­ leaves often reduced to a single leaflet (Cullen 1976), ical variability and taxonomic problems of different Anthyllis vulneraria ssp. alpestris has a distinctly dif­ A. vulneraria taxa (e.g. Sagorski 1908; Cullen 1976; ferent calyx from the other analyzed taxa. Its specific Łukaszewska ei al. 1983; Akeroyd 1988; Cagibtti et al. villous to hirsute hairiness (Cullen 1968; Jasicovä 1988) 1990; Gonzalez 1998; Rich 2001; Puidet et al. 2005; gives it a unique, smoky-greyish colour (Kostrakiewicz Ilcim et al. 2008). Cullen (1976) presented a biosystem- 1959), The length of the calyx is relatively high, ranging atic study of A. vulneraria in Europe with particular on average from 1.4 to 1.6 cm. While A. vulneraria ssp. emphasis on Western Europe* The taxa included there carpatica has sparse sericeous and a usually yellowish were treated as one species divided into a number of calyx (Cullen 1968; Jasicovä 1988), which differs this subspecies and varieties. This treatment is also adopted taxa from the previous one* here. Furthermore, Cullen (1976) distinguished two ag­ Anthyllis vulneraria has been genetically investi­ gregates - Vulneraria and Alpestris aggregate - into gated. Several studies have been made in order to de­ which subspecies are grouped. These subspecies form termine molecular traits that would resolve the complex fairly natural groups corresponding to aggregates. But (Koster 2010). However, the results of different molecu­ these aggregates have no nomenclatural standing and lar analysis are quite contentious (Akeroyd 1988; Kropf intermediates do occur between them. Cullen (1976) et al. 2002; Allan et al* 2003; Nanni et al. 2004; 1 Ion­ discussed division of the group and tried to evaluate ian' et al. 2006; Van Glabeke et al. 2007; Degtjareva taxonomic characteristics of the species. In addition, he et al. 2008; Köster et al. 2008). An issue concerning A. provided distribution maps for each taxon in Europe. vulneraria was included in the Study of molecular phy- Puidet et al. (2005) analyzed morphological vari­ logeny of the genus Anthyllis. The internal transcribed ability of eight taxa of A. vulneraria in Estonia. Most spacers ITS1 and ITS2 of the nuclear ribosomal DNA of the characteristics examined were'statistically sig­ and additional polymorphic chloroplast SSR of 10 An­ nificant in species determination. This investigation re­ thyllis species, including 11 subspecies of A. vulneraria sulted in distinction of three species groups: Vulner­ were analyzed (Nanni et al. 2004), of which 3 are consid­ aria, Macrocephala and Coccinea. In Poland, the study ered in this study. The other investigation used the am­ of morphological variability of A. vulneraria was car­ plified fragment length polymorphism (AFLP) analysis ried out in northern Poland on populations from the to find differentiation in the species complex (Köster et. Baltic coast (Łukaszewska et al. 1983). Multivariate al. 2008). However, the study showed that, the taxa of A. statistical analyses confirm the differences between pop­ vulneraria cannot be differentiated by AFLP patterns, ulations primarily on the basis of fertile parts of the because the analyzed specimens of 7 taxa belonging to plants, but they also considered vegetative characteris­ A. vulneraria did not comprise groups correlated with tics (Łukaszewska ftf al. 1983)- the subspecies (Köster et al. 2008). Several characteristics which distinguish the taxa In Poland, some examinations concerning differen­ within A. vulneraria were! given in different publica­ tiation within A. vulneraria were also conducted. Kali­ tions. The most distinct and well-defined features which nowski et. al. (1983) analyzed the geographic impact differentiate the examined taxa are: the type of stem on the izoenzymatic variability of .1. vulneraria pop­ hairiness, the distribution of leaves on the stem, the ulations in Poland to determine whether there exists abundance of rosette leaves, calyx colour and length. any differentiation between populations of inland and Anthyllis vulneraria Ssp. m,ariiim,a (Schweigg.) Corb. coastal areas. The study confirmed significant differ­ can be distinguish by silky and dense calyx hairi­ ences among populations depending on geographic dis­ ness (Kostrakiewicz 1959), as well as by some inflo­ tance (Kalinowski et. al. 1983). rescences with a few flowers (sometimes not fully de­ In conclusion, different investigations concerning veloped) (Krall 1983). Anthyllis vulneraria ssp. poly- intraspecific taxa of A, vulneraria based on molecular phylla (DC.) Nyman is most readily distinguished by its studies have not. resolved the problem of distinguishing patent hairs on the stems and petioles (Kostrakiewicz
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