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CARYOLOGIA Vol. 64, no. 4: 446-452, 2011

Chromosome numbers in sect. Cammarum (Ranuncu- laceae) from the Carpathians

Ilnicki1 Tomasz and Józef Mitka2*

1Department of Cytology and Embryology, Institute of Botany, Jagiellonian University, Grodzka 52, 31-044 Kraków, Poland. E-mail: [email protected] 2Botanical Garden, Jagiellonian University, Kopernika 27, 31-501 Kraków, Poland. E-mail: [email protected]

Abstract — Chromosome numbers for four and two nothospecies of the sect. Cammarum ser. Variegata, ser. Toxicum, and nothoser. Toxigata in the Carpathians amount to 2n = 16, and in one case to 2n = 32. For the nothospecies: A. ×gayeri Starmühl. and A. ×pawlowskii Mitka et Starmühl. chromosome numbers are given for the first time. According to the cytological evidence a new combination Aconitum degenii subsp. degenii var. intermedium (Zapał.) Mitka [=A. hebegynum non DC.] is proposed.

Key words: Aconitum sect. Cammarum, chromosome numbers, hybridization, Linnaean , Ranuncu- laceae.

INTRODUCTION subgen. Aconitum is still far from being satisfac- torily understood (Luo et al. 2005). The reasons The Aconitum L. is circumscribed of taxonomic inconsistency include the high level within three subgenera: Aconitum, Lycoctonum of morphological variation due to morphologi- (DC.) Peterm. and a monospecific Gymnandrum cal plasticity and the notorious interspecific hy- (Stapf.) Rapaics (Liangqian and Kadota 2001). bridization (Kadota 1981; Oh and Park 1998), Additionally, some authors distinguish the sub- also between species of the different ploidy levels gen. Anthora DC. (Peterm.) (= Anisanthora Na- (Zieliński 1982a, b). kai) and subgen. Tangutica (W.T. Wang) Kadota The general taxonomy of sect. Aconitum in the (2001), though their rank in the Linnean system- Carpathians and the chromosome numbers were atics of the genus is not clear. In Europe, the taxo- given in our previous paper (Ilnicki and Mitka nomic treatment of subgen. Aconitum is clear: the 2009). Now, we report the chromosome numbers section Cammarum (Aconitum variegatum-Group for A. sect. Cammarum in the mountain range. sensu Götz 1967) includes only diploid species, The Carpathians are the most important cen- and sect. Aconitum (A. napellus-group sensu tre of Aconitum evolution in Europe (Mucher Seitz 1969) possesses tetraploid species. The 1993; Starmühler and Mitka 2001; Mitka 2003). species of the sect. Aconitum grow in open, high- The geomorphology of the mountain range is mountain alpine and subalpine zones, and of the linked with its geographical and geobotanical di- sect. Cammarum mainly in forest montane zones, vision into the Western, Eastern, and Southern sometimes having the status of the montane spe- Carpathians (Kondracki 1989; Zemanek 1991). cies descending to the lowlands (Mitka 2003). Especially, the Western Carpathians are isolated Opposite to Europe, the systematics of the Asian from the rest of the mountain range through the low situated the Beskid Niski (Low Beskyd Mts.) and the lowest lying in the Carpathian arch the Dukla Pass - 1000 m above the sea level. The geo- graphical isolation is accompanied by schizoen- demism in various genera in the Western Car- *Corresponding author: e-mail: [email protected] pathians (e.g. Mráz and Szeląg 2004; Ronikier et chromosome numbers in aconitum 447

al. 2008; Tĕšitel et al. 2009), including Aconitum. × A. variegatum subsp. variegatum) and A. ×gay- For example, in the Western Carpathians two eri (A. lasiocarpum × A. degenii). The present infraspecific taxa of Aconitum firmum Rchb., i.e. results could be helpful in the explaining of the A. f. subsp. moravicum Skalický and A. f. subsp. taxonomic relations among taxa of the sect. Cam- maninense (Skalický) Starmühl., and nothospe- marum. The results could also contribute to bet- cies A. ×pawlowskii (Pawł.) Mitka et Starmühl., ter understanding of evolutionary pathways in the are endemic to the region (Starmühler and Mit- genus Aconitum. ka 2001; Mitka 2003). All examined Aconitum subgen. Aconitum kary- otypes, both����������������������������������������� European and Asian,����������������� are highly asym- MATERIALS AND METHODS metric and always consist of two big and six small- er chromosomes (Shang and Li 1984; Joachimiak Aconitum specimens were collected in the et al. 1999; Yang 2001). The sect. Cammarum is field and transferred to the Botanical Garden of diploid with 2n = 16 (Seitz 1969; Joachimiak et al. the Jagiellonian University, Kraków, Poland. The 1999). The results of studies on the sect. Camma- specimens used in the study are kept in the her- rum ser. Variegata, i.e. A. variegatum subsp. vari- barium of the Institute of Botany of the Jagiellon- egatum, and two other species of the sect. Camma- ian University KRA. rum ser. Toxicum: A. lasiocarpum and A. degenii, The method of staining of the chromosomes showed high similarity of their chromosome mor- is described in the paper by Joachimiak et al. phologies. The only difference concerned the sta- (1999). bility of the satellites. In A. variegatum three pairs The taxonomic system follows Starmühler of NOR chromosomes (I, III and V) had the same (2001) and geographical names follow Kon- size of the satellites, but in the karyotype of the re- dracki (1989). maining species the chromosomes of I and III pairs showed structural heterozygosity, i.e. only in one of the homologues of the pair the satellite was clearly RESULTS AND DISCUSSION visible (Joachimiak et al. 1999). In our previous study (Joachimiak et al. 1999) Aconitum sect. Cammarum DC. subsect. Cam- we also presented an accurate organization of the marum ser. Variegata Steinb. ex Starmühl. (2001). classical and C-banding karyotype of A. lasiocar- pum as well as the estimated content and distribu- Ser. Variegata is characterized by homozygos- tion of heterochromatin in A. variegatum subsp. ity of the largest pair of the chromosomes (no. 1) variegatum and A. degenii. The karyotype of the in the karyotype concerning the occurrence of examined specimens of A. variegatum was rich in satellites - Joachimiak et al. (1999). heterochromatin, in contrast to A. lasiocarpum and A. degenii, which were poor in this fraction. Aconitum variegatum L. (1753) We also stated the same structural heterozygos- Aconitum variegatum subsp. variegatum - ity of the NOR-chromosomes of I and III pairs 2n = 16 (Fig. 1A) of A. lasiocarpum and A. degenii in terms of the Locality: [Poland] Tylmanowa, Gorce Mts., size of heterochromatic segments neighbouring Western Carpathians, c. 520 m, 27.09.1999 J. with NOR on their shorter arms. These results Mitka (KRA). didn’t confirm Götz’s (1967) hypothesis on the A subspecies endemic to Europe. It belongs hybridogenous origin of A. lasiocarpum, which to the European-temperate-montane group and could presumably arise in the effect of crossing to the Alpic-Central European distributional type between A. variegatum and A. degenii. On the (Zając and Zając 2009). It occurs in the Alps, other hand, the structural heterozygosity shows Pyrenees, northern Apennines, Sudetes, East- evolutionary lineage of A. lasiocarpum and A. de- ern and Western Carpathians. In the Western genii (Joachimiak et al. 1999), circumscribed in Carpathains it reaches the north-eastern limit of the ser. Toxicum (Mucher 1993). the geographical distribution along the Wisłoka In the present study, we examined eight taxa river in the Niski Beskyd (Poland). The ecological of the sect. Cammarum from the Carpathians: center of the species in the Western Carpathians A. variegatum subsp. variegatum, A. v. subsp. is in the Tatra Mts., where it can be found spo- nasutum, A. lasiocarpum subsp. lasiocarpum, A. radically up to 1500 m above the sea level. l. subsp. kotulae, A. degenii and A. toxicum and Apart from the Carpathians A. variegatum was two hybrid, i.e. A. × pawlowskii (A. lasiocarpum karyologically analyzed in Bulgaria by Koeva-To- 448 ilnicki and mitka dorovska (1992) and in Austria by Schafer and orientated tubers, thus it belongs to the sect. Cat- La Cour (1934), and Seitz (1969). In each case enata Steinb. ex H. Riedl (1978). The taxonomic diploid number 2n = 16 was noted. For informa- position of A. nasutum from Asia Minor is also tion on karyology of the species in Europe see not known. According to the karyological data by Simon et al. 1999. Diploid number of the chro- Simon et al. (1999) three ploidy levels are to be mosomes (2n = 16) for typical representative of found in A. nasutum: 2n = 16, 28, 32. The diploid the species was confirmed - see Tab. 1. We found number of chromosomes was found in the Cau- homozygosity of the satellites on the two largest casus (Tumajanov and Beridze 1968) and Bosnia chromosomes of the subspecies A. variegatum and Herzegovina (Seitz 1969), aneuploid and subsp. variegatum (Fig. 1A, marked with asterisk). tetraploid number in Turkey (Beyazoglu et al. 1994) and tetraploid number in the Alps (Seitz Aconitum variegatum subsp. nasutum (Fisch. et al. 1972) and the Big Caucasus (Gagnidze et al. ex Rchb. em. Rupr.) Götz (1967) - 2n = 16 1985). The taxon needs taxonomic and nomen- Locality: [Romania] Cluj-Napoca, Fâget for- clatural revision. Diploid number of the chro- est, Transilvania, c. 540 m, 5.08.2007 J. Mitka mosomes (2n = 16) of the studied subspecies was (KRA). confirmed. In our studies on Aconitum variega- According to Götz (1967) the subspecies tum subsp. nasutum, homozygosity of the largest occurs in Europe and in the Caucasus. In Eu- pair of the chromosomes in the karyotype con- rope it can be found in the Eastern Alps, East- cerning the occurrence of the satellites was noted. ern Carpathians (the Rodna, Cǎlimǎn-Harghita Mts.), Apuseni Mts., Banat, Transilvania, Balkan Aconitum sect. Cammarum DC. subsect. Cam- Mountains (the Velika Plazenica, Stara Planina, marum ser. Toxicum (Rchb.) Mucher (1993). Rila, Vitoša, Rodopes). Outside Europe in Asian Series Toxicum is morphologically character- part of Turkey (Davis 1965) and in the Caucasus ized by glandular hairs in the inflorescence, and (Luferov 2000). According to Luferov (2000) karyologically by heterozygosity of satellites on the Caucasian A. nasutum differs from the Euro- the largest pair of the chromosomes (no. 1) - pean morphotypes by the plagiothropic, chain- Joachimiak et al. (1999).

Fig. 1 — Metaphase plates of the studied taxa of Aconitum sect. Cammarum from the Carpathians (A – A. variegatum (2n = 16), B – A. lasiocarpum subsp. kotulae (2n = 16), C – A. degenii (2n = 16), D – A. degenii (2n = 32). * satellite, scale bar = 10 μm. chromosome numbers in aconitum 449

Aconitum degenii Gáyer = A. paniculatum (2n = 16) for the studied variety of A. degenii was Lam. nom. inval. confirmed - see Tab. 1. Our cytogenetic studies A. degenii subsp. degenii - 2n = 16, 32 (Fig. 1C, showed that A. intermedium has structural hetero- D) zygosity in two NOR-chromosome pairs I and III Locality: [Ukraine] Polonina Pożyżewska, of the karyotype, typical of the ser. Toxicum. Chornogora Mts., Eastern Carpathians, c. 1300 m, 1.08.1999 J. Mitka (KRA) (Fig. 1 C); [Romania] Aconitum lasiocarpum (Rchb.) Gáyer (1911) Mt. Pietrosul, Rodna Mts., 1640 m, 17.08.1998 J. Aconitum lasiocarpum subsp. lasiocarpum - Mitka, P. Bochenek (Fig. 1 D) (KRA). 2n = 16 A. degenii subsp. degenii is a Carpathian sub- Locality: [Poland] Buk on the Solinka river, species endemic to E Carpathians, S Carpathians, Western Bieszczady Mts., Eastern Carpathians, c. and the Apuseni Mts. A distinct morphotype 490 m, 4.09.1996 J. Mitka (KRA). with capitate spur of the nectaries fo. craciunel- An Eastern Carpathian subspecies endemic to ense Gáyer (1906) occurs in E Carpathians in the the Bieszczady, Vihorlat, Chornogora, Gorgans Rodna, Chyvchyn and Chornogora Mts. Aconi- and Chywchyn Mts. (Mitka 2000). The southern tum degenii in the Alp and Balkan Mountains is limit of its geographic distribution is in the Bistricei circumscribed within the subspecies A. d. subsp. Mts. on Mt. Ceahlǎu. In the paper by Joachimiak paniculatum (Arcang.) Mucher and in the Alps as et al. (1999) heterozygosity of the I, III and V pair A. d. subsp. valesiacum (Gáyer) Mucher (1993). of the chromosomes concerning the occurrence Diploid number of the chromosomes (2n = 16) of the satellites was noted. In our paper diploid for typical representative of the species was con- number (2n = 16) for the typical representative of firmed - see Tab. 1. We also noted a tetraploid the species was confirmed - see Tab. 1. form of the subspecies A. degenii subsp. degenii 2n = 32 (Fig. 1D) and the heterozygosity of the Aconitum lasiocarpum subsp. kotulae (Pawł.) satellites on the largest pair of the chromosomes Starmühl. et Mitka in Mitka and Starmühl. (Fig. 1C, marked with asterisk). (2000) - 2n = 16 (Fig. 1B). Locality: [Poland] Mt. Cergowa Góra, Niski Aconitum degenii Gáyer subsp. degenii var. in- Beskyd, Western Carpathians, c. 700 m, 8.09.1998 termedium (Zapał.) Mitka comb. nov. - 2n = 16 = A. J. Mitka (KRA). hebegynum non DC., auct. fl. Carp. Orient. (Mit- A Western/Eastern Carpathian subendemic: ka 2000, 2003). W Carpathians - the Doły Jasielsko-Sanockie Ba- Locality: [Ukraine] Zaroślak on the Prut river sin, the Niski Beskyd, Tatra Mts.; E Carpathians and Polonina Pożyżewska, Chornogora Mts., - the Bieszczady Mts., Góry Sanocko-Turczańskie, Eastern Carpathians, c. 1275-1410 m, 1.08.1997 Vihorlat, Svidovec-Negrovec, Chornohora, Ma- J. Mitka (KRA). ramures, Rodna, Gorgans Mts. (Mitka 2000); on An Eastern-Carpathian variety endemic to E Podolia in the Ukraine (Mitka and Starmühler Carpathians: the Bieszczady, Svidovec-Negrovec, 2000). Aconitum lasiocarpum subsp. kotulae is Chornogora, Bucovina, Gorgans and Cǎlimǎn- characterised by glandular pubescent helmets and Harghita Mts. (Mitka 2000; Szajna and Mitka glabrous pedicels (Podolia and E Carpathians) or 2010). Zapałowicz (1908) described a variety curved and crisped eglandular pubescent pedicels within A. paniculatum (= A. degenii) “pedicelli sub (Novikoff and Mitka 2011) (the Tatra Mts.). The flore sparse pilosi, cassis angusta”. The taxon, typi- typical subspecies has exclusively glandular pu- fied in Mitka (2003), is characterized by pubes- bescent pedicels. Maybe, a diploid form of this cent helmet, carpels glabrous and pedicels below subspecies was found by Váchova in Váchova and bracteoles glabrous (Novikoff and Mitka 2011) Májovský (1977) - see a comment below Table 1. or carpels pubescent and helmet and pedicels be- In the present paper, heterozygosity of the satel- low bracteoles glabrous. Careful phenetic study lites on the largest pair of the chromosomes was (Szajna and Mitka 2010) revealed that A. interme- noted for Aconitum lasiocarpum subsp. kotulae dium occupies intermediate position between the (Fig. 1B, marked with asterisk). A. degenii and A. variegatum along the Detrended Correspondence Axis 1, and an isolated position Aconitum ×gayeri Starmühl. (1996) - 2n = 16. along the DCA Axis 3. Thus, A. intermedium fits (Aconitum degenii × A.lasiocarpum) the morphological hybrid characteristics, but cy- Locality: [Poland] Procisne, San river, West- togenetic line of evidence places it into the ser. ern Bieszczady Mts., Eastern Carpathians, c. 515 Toxicum. Diploid number of the chromosomes m, 6.09.1995 J. Mitka (KRA). 450 ilnicki and mitka

A nothospecies endemic to the Eastern Car- nia and Herzegovina (the Sarajevo district) and in pathians: the Bieszczady Mts., Mt. Vihorlat, Montenegro (the Durmitor Mts., see Götz 1967). Svidovec-Negrovec, Chornogora, Gorgany, Chy- Mucher (1993) recognized two other subspe- wchyn, Maramures, and Rodna Mts. (Mitka cies from S Carpathians: A. t. subsp. bucegiense 2000). The chromosome number for this taxon is (Nyár.) Mucher and A. t. subsp. crispulum (Nyár.) given for the first time. Mucher, both endemic to S Carpathians, and two nothospecies: A. ×dragulescuanum Mucher (A. Aconitum toxicum Rchb. degenii ×A. toxicum), endemic to E/S Carpathi- Aconitum toxicum subsp. toxicum - 2n = 16. ans (including the Apuseni Mts.), and A. ×bar- Locality: [Romania] Mt. Ceahlǎu, Bistricei tokianum Starmühler (2000) (A. toxicum × A. Mts., Eastern Carpathians, c. 1400 m, 8.08.2007 variegatum) reported from E and S Carpathians, J. Mitka (KRA). the Apuseni Mts., and Bosnia and Herzegovina. A subspecies endemic to the Eastern/Southern Diploids of this taxon were found by Seitz (1969) Carpathians (including the Apuseni Mts.) and the in Bosnia and Herzegovina (Trebović) and in Balkans (Götz 1967; Mucher 1993). In E Car- the Retezat Mts. (S Carpathians) - Tab.1. In this pathians in the Bistricei and Cǎlimǎn-Harghita paper we report diploid number 2n = 16 for the Mts.; in S Carpathians in the Bucegi, Piatra material collected in E Carpathians (Mt. Ceahlǎu) Cralului, Fǎgarǎş and Retezat Mts. (Grinţescu and heterozygosity of the satellites on the largest 1953; Mihok et al. 2005); in the Balkans in Bos- pair of the chromosomes.

Table 1 — Published data on the chromosome numbers of Aconitum sect. Cammarum from the Carpathians.

Taxa 2n Geographic origin Reference Ser. Toxicum A.degenii Gáyer subsp. degenii 16 Bieszczady Mts. (E Carpathians), Frey et al. (1977) Terebowiec valley A.degenii var. intermedium (Zapał.) Chornogora Mts. (E Carpathians), Joachimiak et al. (1999) 16 Mitka Zaroślak in Prut river valley and [=A. ×hebegynum non DC] Polonina Pożyżewska A. lasiocarpum (Rchb.) 16 Bieszczady Mts. (E Carpathians), Joachimiak et al. (1999) Gáyer subsp. lasiocarpum Buk on the Solinka river A.lasiocarpum subsp. kotulae 16* Niski Beskyd Mts. Váchová in Váchová and Májovský (Pawł.) Starmühl. & Mitka (W Carpathians), Čereniny (1977) 16 Niski Beskyd Mts. (W Carpathians), Joachimiak et al. (1999) Mt. Cergowa 16* Góry Sanocko-Turczańskie Mts. Kucharska in Skalińska et al. (1976) (E Carpathians), Mt. Sobień A. toxicum Rchb. subsp. toxicum 16 Retezat Mts., Gura Zlata Seitz (1969) (E Carpathians) Ser. Variegata A. variegatum L. subsp. variegatum 16 Tatra Mts. (W Carpathians) Leszczak in Skalińska (1950) 16 Tatra Mts. (W Carpathians) Leszczak (1950) 16 Tatra Mts. (W Carpathians) Szweykowski and Mendelak (1977) 16, 17 Tatra Mts. (W Carpathians) Zieliński (1982b) 16 Mt. Babia Góra (W Carpathians) Migra (1982) 16 Mt. Veľká Fatra (W Carpathians), Hindáková in Májovský et al. (1987) Ľubochnianska valley 16 Tatra Mts. (W Carpathians), Joachimiak et al. (1999) Kościeliska valley 16 Slovenský kras (W Carpathians), Uhríková in Májovský et al. (2000) Mt. Jeleni vrch

* According to authors A. degenii. chromosome numbers in aconitum 451

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