A N N A L E S HISTORICO-NATURALES MUSEI NATIONALIS H U N G A R I C I

Tomus V I I I . Series nova 1957

The Chromosome Numbers and Cytotaxonomical Relations of Some European Plant Species

By L. B A K S A Y , Budapest

As a continuation of a similar paper published a year previously (1955), author gives the chromosome numbers of species collected in various plant associations of the Carpathian Basin and the Carpathians, the paper being also a preliminary report on the critical species.

1. Cimicifuga europaea Schipcz. n = 8 (2 x ) . Bálvány 900 m, Mts. Bükk , Acereto-Fraxinetum association, on lime. This species is not varying, and the Old and New World species of the genus have identical chromosome numbers, 2n = 16.

2. ! Spiraea media Schm. 2n = 10 (2 x) . Mt . Bélkő 750 m, Mts. Bükk, on a craggy and steep slope ; the character species of Spiraeetum mediae associa­t ion. According to S a x, the chromosome basic number of the genus is x = 9. In three sections, he examined Old and New World species, 18 in all , and 9 of the hybrids. The Old World species are, wi th few exceptions, 2n = 18 diploids, w i t h Spiraea media among them ; the New World species and hybrids are t r i - and tetraploids. The material of S a x was not taken from the natural flora, but consisted of species collected and cultivated in botanical gardens. Spiraea media in its natural site is rather variable, ranging from SE Europe to NE Asia ; its taxonomical evaluation needs further research. The above chromosome number, being also a new basic number, indicates an origin differing from that of the Asiatic species, indeed, as far as the age and locality of origin is concerned^ the SE European centre is supposedly the more ancient.

3. Astrantia major L . 2n = 28 ( 4 x ) . Bánkút 850 m, Mts. Bükk, Sesle-rieto-Fagetum association, on limestone. Rich in forms, varying, wi th several subspecies. W a n s c h e r shew a 2n = 14 species; unfortunately, his material also originated from botanical gardens and so both the locality of origin and the.

range of the diploid are unknown to us. The species examined by us is exceedingly similar to the Balkanian ssp. elatior (Friv.) Murb., its further clarification needs chromosome counting from various other Iocatilies. H a k a n s s o n ' s member for ssp. biebersteini is n = 14 (4 x).

4. ! Pimpinella major (L.) Huds. 2n = 20 (2 x) . See above locality. Accord­ing to the counting of S c h u l z - G a e b e l , 2n = 18. The whole family is euploid, there are very few polyploids among the species. Chromosome number 18 is probably not right.

5. Asperula montana W. et K . ( = Asperula cynanchica L . p. p.). 2n = 44 (4 x ) . Macskagödör, Mts. Vértes, Festuca sulcata — Carex humilis association; Mt . Nagyszénás, Nagykovácsi , Festucetum glaucae hungaricum association. The plant is of a Mediterranean origin, and has also a diploid. The name Asperula montana W. et K . refers to a species described from Hungary which is, according to the above examination, a tetraploid, to be conserved as a valid specific mame in the case the A. cynanchica named by L i n n é refers to a 2n = 22 species. A. montana is regarded by some authors as a variety and the chromosome num­ber 2n = 22 was referred to i t (H o m e y e r ) .

6. ! Mercurialis ovataSternb. et Hoppe, n = 16, 2n = 32 ( 4 x ) . From the area of the Hungarian Central Mountains : Gyenesdiás, Csidervölgy, Querceto-pubescentis ; Kígyósvölgy, Festuco-Brometum erecti ; Tihany, Csúcshegy, Quer-cetum ; Budakeszi, Fekete hegyek, Qu. pubescentis associations.

7. Mercurialis longistipes (Borbás) Baksay n = 32, 2n = 64 (8 x) . Vértes­kozma, Macskagödör ; Budapest, Hármasha tá rhegy , Quercetiim pubescentis ; Budapest : Széchenyihegy ; Szár. Zuppahegy ; Solymár, the Solymár Wall . In al l three localities, from Querceto-Carpinetum associations.

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8. Mercurialis perennis L . n = 21, 2n = 42 (2 x) . Fánivölgy, Mts. Vér­tes ; Mt . Naszál, Vác ; Mélysár, Mts. Bükk. In all three localities, from Fagetum associations. Budapest, Szarvashegy, from a Querceto-Carpinetum association.

M. perennis is a European-Mediterranean, mainly beechy species ; M. ovata is a Central European — SE Mediterranean, lanuginous oak forest species. The chromosome number of the former species was found by P ó l y a to be identical w i t h the above, whilst M e u r m a n (1925) counted n = 32 in meio-sis. The plant examined by M e u r m a n belonged therefore to M. longistipes. This latter species was considered by our specialists to be a transition between M. ovata and AL perennis, which was thought by G r a e b n e r to be a Mer­curialis Paixii (M. ovataxperennis) hybride. A i . longistipes is the polyploid of the species. M. ovata, but not an autopolyploid. The reductional division in the pollen mother cells, examined in very large quantities, was always normal,

no secundary associations, irregularities or polyvalents could be observed. We succeeded to observe a sexual chromosome in the first metaphase. The chromosome numbers of but a few Mercurialis species are known. The members of the x = 8 basic number series are : 16, 32, 48, 64. Of these, AL annua 2 x is a weed of Mediterranean origin, the 4 x and 8 x species are the above mentioned ones, M. leiocarpa 6 x is perennial, ranging in SE Asia. AL perennis, a species w i t h the basic number x = 21 (?), is genetically further away from M. ovata; the cytological examination of still more species of the genus could explain the origin of this basic number.

The range of M. longistipes is (according to the data of M e u r m a n) i n Northern and Central Europe, in the Balkans, at least in its Northern part. In M. longistipes, the margin of the leaf is finely crenate, its stalk shorter than 1 cm, never adnate as in ovata. Regarding its habitus, i t stands between M. ovata and AL perennis, and this misleading circumstance is heightened also by the sexual dimorphism. In the case of all three species, the leafes of the pistillate specimens are a broader oval, those of the staminate specimens are more narrow and longer, to which morphological changes in leaf shape are further contributed by oecological circumstances (e. g. sunshine, shadows) of the locality. M. lon­gistipes is similar to M. ovata in its oecological behaviour, i t occurs in calcareo-philous lanuginous-oak forests and hornbeam-oaklands, in Carex alba beech forests, on rocky places ; but in beech forests only M. perennis thrives. S a x e r regards this latter as a species of very high adaptability, and this view is due to the fact that, in reality, we are dealing wi th two species, genetically different and also diverging in their oecological claims. There is no M. perennis in the Pulsatilla steppe-forest mentioned by him, but AL longistipes. During mitosis examinations of M. perennis, we have once counted 2n = 84 chromosomes. Author wishes to treat the species detailedly in another paper.

9. ! Onsoma tornensis J á v . 2n = 14 (2 x ) . The seeds came, by way of the botanical garden of Kosice (Czechoslovakia), from the Várhegy of Torna. An endemic plant of the Torna-karst. 0. arenaria has 2n = 12.

m % g& 10. Hesperis matronalis ssp. Vrabélyiana (Schur.) Soó. 2n = 24 (2 x ) .

Ta rkő , Mts. Bükk, Fagetum. An endemic subspecies, the nominate species ha­ving chromosome numbers 24 and 28.

11. Arabis alpina L . 2a — 16 (2 x ) . Gerennavár, Ablakoskö valley, Mts. Bükk , Acereto-Fraxinetum association. Arabis alpina f. glabrescens Borb. 2n = 16, the Szádelő valley, from the Slovakian Karst. Also a tetraploid varia­t ion is known from Europe. Its glacial relict character in the Carpathian Basin is supported also by its chromosome number.

12. Arabis hirsuta (L.) Scop. 2n = 16 (2 x) . Szarvashegy, Budapest, Fageto-Ornetum association fragment. Rich in forms, a polyploid serial conspe-

cies w i t h chromosome numbers 32, 64 ; i t wi l l probably fall to separate species both oecologically and phytogeographically.

13. Arabis turrita L . 2n = 16 ( 2 x ) . See above locality. 14. Camelina microcarpa Andrz. 2n = 40 ( 4 x ) . Pisznice, Mts. Gerecse ;

Bélkő, Mts. Bükk. In both localities, on limestone, from Festucetum sulcatae pannonicum association, together w i t h Ferula Sadleriana, from a primary asso­ciation. Its chromosome number agrees wi th the date of M a n t o n (1932). C. microcarpa is an Eurasian steppe-element, it is especially common in secundary places, in weed-associations on a dry soil. The primary species has differentiated due to cultural effects, so much that several authors separate it into 4 species (S t e b b i n s).

15. Camelina rumelica Velen, n = 6 (2 x) . Isle Szentendre, Pócsmegyer , on the edges of locust groves on sand . Its chromosome number differs from the species examined up to now, which are all polyploids.

16. Helianthemum canum (L.) Baumg. 2n = 22 (2 x ) . Nagyszénás, Nagykovácsi , Festuceto-Brometum erecti association.

17. Telekia speciosa (Schreb.) Baumg. 2n = 20 (2 x) . in the Feketesár valley, Mts. Bükk ; in a wet, watery high-weed association. Not varying ; a montane species ranging in the Carpathian Basin, the Balkans, and the Cau­casus.

18. Solidago virga-aurea L . 2n = 18 (2 x) . Szarvashegy, Budapest, from the fragment of a Fageto-Ornetum association. An exceedingly adaptable species, a member of the most different association from the mountains to the plains.

19. ! Echinops ruthenicus Fisch. 2n = 32 ( 4 x ) . From the Botanical Gardens of Budapest. Its chromosome numbers agrees wi th that of E. sphaero-cephalus.

20. ! Silene densiflora d 'Urv. ssp. Sulingen Hendrych. 2n = 24 (2 x ) . In the Szádelő valley of the Slovakian Karst, in Seslerieto-Festucetum associa­tion. The primary species has a Balkanian and Near East range.

21. Allium moschatum L . 2n = 16 (2 x) . Nagyszénás, Nagykovácsi , Festucetum glaucae hungaricum. A Pontic-Mediterranean species. Our date is conform to those of B i l l e r i (1954). P ó l y a established a chromosome number 2n = 24, the oecological description of the collecting locality of his examined material is erroneous or based on some mix-up.

22. ! Carex Fritschii Waisb. 2n = 30 (2 x ) . Uzsa, Querceto-Betuletum callunetosum association (leg. Ú j h e l y i ) . Its specific value was discussed for a long time, several authors regarded i t as the subspecies of C. montana L .

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Its chromosome number differs, that of C. montana is 2n = 38. Aside of their different morphological characteristics, also their oecological claims are dissim­ilar. C. Fritschii is a species shunning limestone on the southern and eastern

submountainous areas of the Alps, and on lower places in their northern slopes. It occurs in Hungary on rocky soils and pebbles at the Bakonyalja in the Trans-•danubium. C. montana is a montane, Quercetalia-species, indifferent to soils.

23. ! Festuca pseudodalmatica Krajina. 2n = 28 ( 4 x ) . Diósjenő, Mts. Börzsöny, under the peak of the Nagymána , on andezité , in Festucetum pseudo-dalmaticae association. It cannot be synonymized either from a morphological or a cytological point of view wi th F. sulcata, having chromosome numbers 2n = 42, which, was also established by H o r á n s z k y on the basis of anatomical characters.

24. Festuca Tatrae (Czakó) Degen. 2n = 14 (2 x) . The Béla limestone Alps, in the High Tatra, Czechoslovakia (circa 1300 in), Festucetum Tatrae association, its character species. This is a subalpine-montane species on limestone and dolomite, ranging in the western part of the Northern Carpathians, the SW part of Galizia and northern Slovakia. Previous counting for F. Tatrae was made by B r a n d b e r g (1948) and S k a 1 i n s k a (1957) on the plant in the same region. The chromosome number of the related F. amethystina is 2n = 28, published first by S t ä h l i n and also corroborated by the present author in a former paper (1956). The structural pictures of the epidermis of the two spe­cies — very suitable for specific identification in the Gramineae — are identical, differring only in size characteristical of di- and tetraploids. On the basis of the concurrence of the morphological and anatomical characters, the diploid F. Tatrae has its conforming tetraploid in F. amethystina.

L i t a r d i è r e (1950) established the chromosome number of F. amethystina L . var. eu-amethystina St. Y . subvar. genuina St. Y . as 2n = 14, yet, doubtlessly, we are dealing here wi th nothing else but the diploid F. Tatrae, synonymized formerly wi th the species F. amethystina. This latter has 2 subspecies and 3 varieties, of which the description of var. genuina is highly similar to that of F. Tatrae, and i t is highly possible* that the species in question is one and the same. The plant of L i t a r d i è r e originated from the botanical gardens of Lau­sanne and so we are not cognizant of its natural locaility, yet we suppose that F. Tatrae may occur also in the area of the Alps. More extensive cytological examinations wi l l probably clarify our knowledge concerning the range of F. Tatrae which wi l l , at the same time, shed light on the place of origin and the direction of spreading of F. amethystina.

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* There is no foreign material of F. amthysiina in the herbarium of the Museum

graphic distribution in Spiraea (Journ. Arn. Arbor. 77, 1936). — 15. S a x e r, A.: Die Fagus-Abies und Picea Gürtelarten in der Kontaktzone uer Tannen u. Fichtenwälder der Schweiz (Beitr. z. Geobot. Landesaufnahme der Schweiz, H . 36, 1955). — 16. S o ó — J á v o r k a r Magyar növényvilág kézikönyve (Budapest, 1951). — 17. S t e b b i n s, L . : Variation and Evolution in Plants (New York, 1950). — 18. W a n s c h e r, J. H. : Studies on the Chromosome Numbers of the Umbelliferac (Bot. Tidsskr. 42, 1932). — 19. W o 1 f f : in Englers Pflanzen­reich : Umbelliferac (67, IV . 1913). — 20. L i t a r d i è r e, R.: Nombres chromosomiques de diverses Graminées (Bol. Soc. Brot, 24, 1950). — 21. D a r l i n g t o n — W y l i e : Chromosome Atlas of Flowering Plants (London, 1955.) — 22. L ő v e , A. et L ö v e , D. : Cytotaxonomical Conspectus of the Icelandic Flora (Acta Hort. Gotoburg. 20, 4. 1956) . — 23. S k a l i n s k a — B a n a c h - P o g a n — W c i s l o — et A l l . : Further studies in the chromosome numbers of Polish Angiosperms (Acta Soc. Bot. Polon. 26, 1. 1957) .