Taxonomy and ecology of the genus Aconitum L. in the Ukrainian Carpathians

Summary: In the Ukrainian Carpathians 12 species of the genus Aconitum occur, circumscribed by three subgenera. General descriptions, ecological characteristics, chorology and threat of 20 Aconitum taxa are given based on herbarium materials and fi eld investigations. It was found out that all studied taxa belong to three ecological groups, which are distributed in diff erent altitudinal belts, to four biomorphological groups and to two main life forms.

In spite of the long history of investigation, taxonomy of the genus Aconitum is still much discussed and perhaps will remain unresolved.Many taxonomical problems arise from the notorious hybridity within the genus and subsequent origin of putative genetic hybrids, which form a morphological continuum between the parental species (Kadota 1987;Starmühler 2001).In the Ukrainian keys for the species identifi cation often labile and indistinct features as shape of leaves, size of fl owers and height of plants are used.For example, in one of the most popular Ukrainian keys, the 'Plants identifi cation guide for Ukrainian Carpathians' (Chopik 1977), ten species of the genus Aconitum are cited.In another work 'Identifi cation guide for higher plants of Ukraine' (Prokudin 1987) eleven monkshood species are recognized.In both cases, some species were not taxonomically clearly sound.They were clarifi ed in recent taxonomical revisions of the genus Aconitum by Mitka (2000Mitka ( , 2003Mitka ( , 2008) ) and Starmühler & Mitka (2001).Here, the morpho-geographical species concept (Davis & Heywood 1963) was used.Mitka (2003Mitka ( , 2008) ) points to 13 taxa in the genus Aconitum in the Eastern Carpathians.All species belong to the three subgenera Aconitum, Lycoctonum and Anthora.They are described on the basis of root and fl ower morphology.Among them the subgenus Aconitum is the richest in the Ukrainian Carpathians and consists of two sections (Aconitum, Cammarum DC.), one nothosection Acomarum Starm.(sect.Aconitum × sect.Cammarum) and ten species.The second subgenus Lycoctonum (DC.) Peterm. is represented by only one species, A. moldavicum Hacq.Although A. lycoctonum is considered possible for this territory (Mitka 2003), no native localities have been found up to now.The third, controversial subgenus Anthora (DC.) Peterm. is represented by just one species, A. anthora L. From the total number, six species are endemics to the South-Eastern Carpathians, one is endemic to the Carpathians and three are Carpathian subendemics.In the Ukrainian part of the Eastern Carpathians theoretically should occur altogether 13 species, but a complete revision of the genus Aconitum in this territory has not yet been carried out.The high level of endemism and a lack of relevant information on the genus make the territory a very interesting object of the taxonomic studies.We propose here the revised chorology of the genus Aconitum in the Ukrainian part of the Eastern Carpathians alongside with a new determination key.

Materials and methods
In 2008 -2010 the first author made several field studies in all regions of the Ukrainian Carpathians and collected herbarium material, which is kept in LW, LWS and LWKS.For the morphological study he investigated 576 specimens, excluding A. variegatum, A. lycoctonum and A. × cammarum.The first two species are possible for the Ukrainian Carpathian's native flora and the last one is ornamental.Raw data were processed in StatSoft Statistica 8.0 program.
For the chorological and morphological investigations we also used the herbarium collections from CHER, KRA, KRAM, KW, LW, LWS, LWKS, W, WU and YALT (for abbreviations see Holmgren et al. 1990).For the chorology we also used published data on Aconitum species.
Localities found in the field are marked on a map (Carpathians 2007) by rings, localities based on herbarium data are marked by squares, localities taken from the literature, but not confirmed in the field or by herbarium investigations, are marked by triangles.We also used the asterisk for marking suspicious localities cited in literature.The synonyms follow Mitka (2003) and others (Paczoski 1927;Domin 1929;Steinberg 1937;Kmeťová 1982;Skalický 1990;Voroshilov 1990;Tasenkevich 1998;Mosyakin & Fedoronchuk 1999;Tzvelev 2001).Some of them should be checked in future because their names are not typified.Taxonomic names were also checked by IPNI (2010), GRIN (2010) andESFEDS (2010) databases.Threat categories to the species follow the IUCN (2001) criteria.
Distribution: In the Ukrainian Carpathians A. anthora is a rare species.It occurs in Chornohora, Nyzki Polonyny, Marmarosh, Chyvchyny and Hrynyava (Fig. 6).It mainly grows in the mountane and subalpine belts on calcareous outcrops, on rocks and stony places.It is also distributed in the lowlands of the whole Ukraine, particularly in Podillya, Kharkiv region and Crimea.
Status: A. anthora is a relict species with disjunctive distribution in Eurasia.It forms morphologically distinct biotypes and needs further taxonomic studies.
Threat category in Ukraine: According to the 'Red Book of Ukraine' A. anthora is protected as A. jacquinii with the category R and A. pseudanthora with the category VU (Ziman et al. 2009;Kagalo 2009).A. jacquinii has also been included in the 'Carpathian list of endangered species' (Tasenkevich 2003) with category VU and according to Kricsfalusy & Budnikov (2007) it has the status CR.We conclude that this species should gain the category VU because of the limited numbers of localities (Novikoff 2010b).
Remarks: A. anthora is a polytypic species with high level of morphological variability, e.g.hairiness, hood shape and color of perianth.The species not only grows in the mountains but it stretches all through the Eurasian continent.Most botanists of the Ukraine and Russia described A. anthora within several taxa of different ranks (Steinberg 1937;Voroshilov 1990;Mosyakin & Fedoronchuk 1999;Tzvelev 2001).In the Ukrainian Carpathians, there are two morphotypes of A. anthora with yellow perianth: a) with hooked hairiness of the hood and pedicels; b) with long protruded hairiness of the hood and protruded or mixed hairiness of the pedicels (Novikoff 2009a(Novikoff , 2010a)).On the Podillyan plateau there grows a third morphotype with bluish or violet color of the perianth and hooked hairiness of hood and pedicels.The morphotype was decribed as A. pseudanthora Błocki.Some experimental works are needed to check its morphological stability on garden conditions.Description: Semi-rosette, often with long and procumbent stem, up to 170 cm of height; rhizomes divided by particulation; stem and pedicels pubescent; leaves divided into 5 -7 broad lacinias; inflorescence loose ramified, pubescent with hooked hairs; perianth yellow or yellowish, falling after blooming; hood hooked pubescent, cylindrically elongated, 2.5 -3 higher than wide; 3 carpels densely pubescent; filaments dentate or not; nectary spurs coiled, not reaching the top of the hood; pedicels hooked pubescent; seeds without wings.
Distribution: In the Ukrainian Carpathians A. lycoctonum subsp.lycoctonum is known from only one locality in the Chornohora Mts., where it probably has been introduced (Fig. 6).
Status: European species.6).They grow on wet places, along torrents and rivers, in forest ecotones, etc.; mainly in the submontane and montane belts.In some cases they are found in the subalpine belt (Chyvchyn, Gorgany Mts.) in open habitats.
Threat category in Ukraine: According to Kricsfalusy & Budnikov (2007) the typical subspecies has the status EN.We propose the status LC for A. moldavicum subsp.moldavicum and subsp.hosteanum and the status DD for subsp.simonkaianum and subsp.porcii (Novikoff 2010b).
Remarks: A. moldavicum is one of the most common species of the genus, distributed in almost the whole Carpathians (it is absent, for example, in the Tatra Mts., Western Carpathians).localities is unknown (Fig. 7).It grows along streams and rivers, in wet grassland, in wet meadows of the Calthion alliance and in the Carpathian alder-carr Alnetum incanae; mostly in the montane and submontane belts, rarely in the subalpine belt (Chyvchyn Mt.).
Threat category in Ukraine: Protected according to the 'Red Book of Ukraine' and the 'Bern Convention' by the VU category.Included in the 'Carpathian list of endangered species' with the VU category (Tasenkevich 2003) and according to Kricsfalusy & Budnikov (2007) it has the status EN.
Remarks: A. lasiocarpum is a rare species which occurs sporadically, mostly in mixed populations with both subspecies.Description: Upright standing, erosulate plant, 60 -160 cm high; tubers elongated; stem and leaves hooked pubescent; leaves 5(7), incised more than ½; inflorescence dense ramified, rarely loose, pubescent; perianth of different colour from white to violet and variegated, falling; hood glabrous, hemispherical or elongated, 1.2-2 times higher than wide; 3(4)5 glabrous carpels, sterile; filaments pubescent, dentate; nectary spurs bent or coiled, reaching the top of hood; pedicels pubescent with hooked short hairs just above the bracteoles and glabrous below.
Distribution: It is cultivated in gardens and sometimes occurs in nature in the abandoned villages.

Aconitum × gayeri
Distribution: In all regions of the Ukrainian Carpathians from montane to subalpine belts (Fig. 7).It mainly grows in the open wet places along the streams in mixed populations with A. degenii.
Threat category in Ukraine: As one of the most frequent species in the section Cammarum DC. it is classified LC (Novikoff 2010b).
Remarks: This nothospecies resembles A. degenii in general habit.Distribution: In the whole Ukrainian Carpathians just as A. × gayeri (Fig. 7).
Threat category in Ukraine: According to Kricsfalusy & Budnikov (2007) it has the status VU.As one of the most frequent species in the section Cammarum DC. it is classified LC (Novikoff 2010b).
Distribution: In the Ukrainian Carpathians it is a rare species mostly occurring in the Chornohora and Svydovets Mts. and sporadically in the Gorgany, Nyzki Polonyny, Marmarosh and Chyvchyny Mts. (Fig. 8).It grows in Adenostylion and Calamagrostion communities in the subalpine and alpine belts, from where it descends sometimes along the streams into the higher montane belt.
Status: Species in the transitory area between the Western and Southern Carpathians.
Threat category in Ukraine: According to Kricsfalusy & Budnikov (2007) A. firmum subsp.firmum has the category LRnt, and A. firmum subsp.fissurae is CR.Because of the limited localities and the increasing tourist's impact, this species will probably gain the status of vulnerable VU in the next five years (Novikoff 2010b).
Remarks: A. firmum is a typical representative of A. napellus agg., which occurs not only in the Carpathians, but also in the Alps and other alpid mountain ranges.Many species described within A. napellus agg.form probably a morphological continuum and differ mostly by the hairiness of carpels, hood and pedicels.In the Carpathians A. firmum is circumscribed within the two subspecies: subsp.firmum and subsp.fissurae.The first has the ecological center in the Western Carpathians, the latter -in the Southern Carpathians, where it was thought to be A. romanicum.
In the Eastern Carpathians we probably meet the migratory routes of A. firmum from the Western and Southern Carpathians.The putative cline diversity of A. firmum along the mountain arch needs further molecular DNA and morphological studies.Description: In general habit similar to A. firmum; it differs by pedicel pubescence. A. × nanum has hooked hairs with admixture of rarely protruded hairs above bracteoles and with similar or without hairiness below the bracteoles.
Distribution: It mostly occurs in the Chornohora range and then in the Skolivski Beskydy region, Svydovets, Gorgany, Nyzki Polonyny and Marmarosh Mts. (Fig. 8).It often occurs in mixed populations with A. firmum.
Threat category in Ukraine: DD (Novikoff 2010b).Description: In general habit similar to A. firmum; it differs by hood and pedicels pubescence: hood densely hooked or mixed pubescent; carpels glabrous or rarely pubescent on the dorsal side (fo.bucovinense) or entirely and sparsely pubescent (fo.orthotricha Gáyer); filaments mostly pubescent; pedicels densely pubescent with mixed, protuberant and hooked hairs.The ecological groups described above are closely related to the life forms of monkshoods which have been described on the basis of the morphology of roots, shoot development and general biology.Only the members of the subgenus Lycoctonum have rhizomes, all other species have tubers (spherical in the section Cammarum and elongated napiform in the sections Aconitum and Acomarum).Moreover, the investigated species have over-ground shoots of three different morphological types: a) clinoapogeotropic (shoots growing askew or creeping), b) apogeotropic (upright standing orthotropic) with elongated internodes, c) apogeotropic with brachytic internodes (Zhmylev et al. 2005;Novikoff 2010c).The morphological shoot types are depending on the length of the plant, leaf length and length of the leaf blade, internode length and number of inflorescence branches.Together they form four biomorphological groups considered later (Tab.1).The leaves of the first morphotype are the biggest and they have the largest blades, but leaves of the second and third morphotypes differ also in length.The first morphotype has the longest internodes, while the second morphotype has middle-sized internodes.It is interesting that A. lasiocarpum has internodes which are similar to the third morphotype.We suppose that it is related to the differentiation of A. lasiocarpum (likewise A. variegatum) into the two groups: dwarf plants, growing in the subalpine belt, and typical plants, growing in the montane belt.

Aconitum bucovinense
The mature plants of the first and the second morphological types are mainly multiflowered, they have ramified compound racemes, and the mature plants of the third group have rigid, simple or rarely dense ramified racemes with few flowers.Commonly, the semi-rosette, polycyclic clinoapogeotropic overground shoots are found only in the subgenus Lycoctonum, while the remaining species have erosulate, monocyclic apogeotropic shoots.The representatives of the section Aconitum have shoots with brachytic internodes, while the sections Cammarum and Acomarum have both types of apogeotropic shoots.Serebryakova & Polyntseva (1974) suggested that all of the representatives of the genus Aconitum belong to eleven life forms, which later (Serebryakova 1977)  main architectural models: a) perennial herbs with semi-rosette, determinate shoots (subgenus Lycoctonum) and b) perennial herbs with erosulate, determinate and indeterminate shoots (the remaining subgenera).It was found that all the species, including the subgenus Lycoctonum, have indeterminate shoots with terminal bud, which may be shift on the level below the pedicel of upper flowers (Novikoff 2009b).According to another study of the life forms in the genus (Barykina et al. 1977a), monkshoods have only seven main life forms, which include several biomorphological groups of plants and differ mainly in root morphology.
In conclusion, the monkshoods of the Ukrainian Carpathians belong to two different architectural models and two main life forms divided into four biomorphological groups (Tab.2).The fourth group includes A. anthora which is one of the most interesting species in the genus.It combines many features of the second and third group.It is worthy to note that A. anthora is in this respect an intermediate case -the tubers in early stages of the ontogenesis are spherical, but later they become elongated.Moreover, the number of tubers of A. anthora may increase and then they compound into bunch, what is also characteristic of some other Asian species (Litvinenko 1977a(Litvinenko , 1977b(Litvinenko , 1981;;Barykina et al. 1977b;Ziman 1983).Because of its wide geographical distribution, the various forms of the species may have short apogeotropic shoots, rigid inflorescences (in high-mountain localities) or very elongated apogeotropic or even clinoapogeotropic shoots and loose ramified inflorescences (in lower localities).The other significant features of A. anthora are: persistent perianth after blooming, unique morphology of the nectary (Kosuge & Tamura 1988) and seed (Cappelletti & Poldini 1984).Another species, the putative hybrid A. × cammarum, is difficult to interpret, because it has an intermediate position between the second and third biomorphological group.It has elongated tubers like the species from the third group, but all other morphological features are similar to the second group.Probably, it is conditioned by the intersectional, hybridogenous origin.

Conclusion
• The genus Aconitum in the Ukrainian Carpathians is represented by 20 taxa (12 species) which belong to 3 subgenera -Aconitum, Anthora and Lycoctonum.From the total number 11 taxa are endemics and 5 taxa have a status of threatened in the wild.
• The Aconitum species in the territory may be divided into four ecological groups according to their optimal habitats and vertical longitudinal vegetation belts.
• The species investigated belong to the two main life forms and four biomorphological groups, mainly differentiated by root and shoot morphology.

Figure 6 .
Figure 6.Distribution maps of the subgenus Anthora and subgenus Lycoctonum representatives.

Figure 7 .
Figure 7. Distribution maps of the subgenus Aconitum section Cammarum representatives.

Figure 9 .
Figure 9. Distribution of the monkshoods in different regions of the Ukrainian Carpathians.

Figure 10 .
Figure 10.Altitudinal distribution of monkshoods in the Ukrainian Carpathians.
Aconitum lycoctonum L. em.Koelle subsp.lycoctonum, The center of distribution is in Skolivski Beskydy (the Eastern Bieszczady Mts.) in the Ukrainian Carpathians, where also white-flowering plants and plants with reduced hood occur.

Table 1 .
joined into the two The results of measurements of morphological features in the Ukrainian Carpathians Aconitum species.

Table 2 .
Architectural models, life forms, and biomorphological groups in the Ukrainian Carpathians Aconitum species.