Chromosome Numbers in Aconitum Sect. Cammarum (Ranuncu- Laceae) from the Carpathians
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View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Jagiellonian Univeristy Repository CARYOLOGIA Vol. 64, no. 4: 446-452, 2011 Chromosome numbers in Aconitum sect. Cammarum (Ranuncu- laceae) from the Carpathians Ilnicki1 Tomasz and Józef Mitka2* 1Department of Plant 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 species 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 taxonomy, Ranuncu- laceae. INTRODUCTION subgen. Aconitum is still far from being satisfac- torily understood (Luo et al. 2005). The reasons The genus 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).