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Cytotaxonomic Studies on Galium Verum L

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Cytotaxonomic Studies on Galium Verum L Cytotaxonomic studies on Galium verum L. Autor(en): Kliphuis, E. Objekttyp: Article Zeitschrift: Botanica Helvetica Band (Jahr): 94 (1984) Heft 2 PDF erstellt am: 07.10.2021 Persistenter Link: http://doi.org/10.5169/seals-65878 Nutzungsbedingungen Die ETH-Bibliothek ist Anbieterin der digitalisierten Zeitschriften. Sie besitzt keine Urheberrechte an den Inhalten der Zeitschriften. Die Rechte liegen in der Regel bei den Herausgebern. Die auf der Plattform e-periodica veröffentlichten Dokumente stehen für nicht-kommerzielle Zwecke in Lehre und Forschung sowie für die private Nutzung frei zur Verfügung. Einzelne Dateien oder Ausdrucke aus diesem Angebot können zusammen mit diesen Nutzungsbedingungen und den korrekten Herkunftsbezeichnungen weitergegeben werden. Das Veröffentlichen von Bildern in Print- und Online-Publikationen ist nur mit vorheriger Genehmigung der Rechteinhaber erlaubt. Die systematische Speicherung von Teilen des elektronischen Angebots auf anderen Servern bedarf ebenfalls des schriftlichen Einverständnisses der Rechteinhaber. Haftungsausschluss Alle Angaben erfolgen ohne Gewähr für Vollständigkeit oder Richtigkeit. Es wird keine Haftung übernommen für Schäden durch die Verwendung von Informationen aus diesem Online-Angebot oder durch das Fehlen von Informationen. Dies gilt auch für Inhalte Dritter, die über dieses Angebot zugänglich sind. Ein Dienst der ETH-Bibliothek ETH Zürich, Rämistrasse 101, 8092 Zürich, Schweiz, www.library.ethz.ch http://www.e-periodica.ch Botanica Helvetica 94/2, 1984 0253-1453/84/020269-10 $ 1.50 + 0.20/0 © 1984 Birkhäuser Verlag, Basel Cytotaxonomie studies on Galium verum L. E. Kliphuis Department of Biosystematics, University of Utrecht, Padualaan 8, 3584 CH Utrecht, The Netherlands. Manuscript reeeived August 31, 1984 Abstract Kliphuis, E, 1984: Cytotaxonomie studies on Galium verum L. Bot. Helv. 94:269-278. A cytological investigation of Galium verum L. oecurring in Europe revealed diploids (2n 22) and tetraploids (2n 44). The tetraploids are distributed throughout Europe, the diploids have a more southerly distribution. A comparative morphological investigation showed three groups within the diploids. One of these is identical to the subspecies wirtgenii (F. Seh.) Ob, the other two fall entirely within the subspecies verum as far as their morphology is concerned. Within the tetraploids no well-defined correlation in morphology and geographical distribution could be demonstrated, except in the case of a taxon oecurring along the Atlantic coast of Europe. A clear-cut Separation, however, proved to be impossible due to the large degree of overlap in the distinguishing characters. Introduction The extremely Polymorphie Eurasian Galium verum group comprises several taxa whose delineation and distribution are still unclear. The group displays its highest degree of polymorphy in the Asiatic part of its distributional area. Many taxa have been described from the Caucasus and adjacent areas as well as from central and east Si- beria. Their systematic position, however, is often doubtful. In the Flora of the U.S.S.R, Pobedimova (1958) mentioned within the section Xanthogalium, described originally by De Candolle three subsections containing altogether 15 species. In Europe, with the possible exception of the Balkans the Situation does not seem to be so complicated. Ehrendorfer and Rrendl (1976) in Flora Europaea, distinguished two species within the verum group e.g.: G. tunetanum Lam - a more or less isolated taxon from the south of Spain, Sicily and North West Africa -, G. verum L. with two subspecies verum and wirtgenii (Seh.) Ob. and Galium Xpomeranicum Retz. a hybrid of the subspecies verum with the tetraploid G. album Mill. ssp. album, which is a repre- sentative of the G. mollugo group and which is widely distributed over Europe. Within G. verum L. two cytotypes are known: a diploid and a tetraploid, with 2n 22 and 2n 44 chromosomes respectively. In Europe these cytotypes are not equally distributed. This already has been known since the studies of Fagerlind (1937). Among a majority of tetraploids, distributed all over Europe, this author found a mi- nority of diploids in the south-eastern parts of Europe. This picture was confirmed by a 269 270 E. Kliphuis number of authors. Most of them report only tetraploids (Löve and Löve 1956, Piotrowicz 1959, Zhukova 1966, Leveque and Gorenflot 1969, Murin and Vächovä 1970, Schotsman 1973, Sokolovskaja 1972, Surova 1973, Murin 1974, Murin and Zabor- sky 1976, van den Brand, van Meel and Wieffering 1979). Tetraploids and diploids were reported by Kliphuis (1962) and Lovkvist (1962). A locality with diploid plants has been recorded for Bulgaria by Ancev (1974). Material of G. verum was collected from different parts of Europe. Plants obtained directly from the wild and plants grown from seed collected in the field were cultivated under uniform conditions in an experimental plot. During the years of cultivation this material showed differences in its morphology and a number of characters proved to be variable. Some differences were subtle and hard to describe, whereas others were strik- ing. A number of differences were influenced by the environment. It seems worthwhile subjecting this collection to a cytological and comparative morphological investigation. The results of this investigation are given and discussed in this paper. Material and methods Plants collected in the field and plants grown from seed were cuftivated in the experimental garden of the State University of Utrecht during several years. From each seed sample 3-5 plants were used for cytological purposes. The material was collected partly by the author, partly by colleagues and partly with the aid of other botanical institutes. In this study we used only material collected in the wild and of which the place of origin was registered. The chromosome counts were made from the root tips of potted plants. The tips were fixed in Karpechenko's fixative, embedded in paraffin wax, sectioned at 15 micron and stained according to Heidenhain's method. A comparative morphological investigation was carried out on living material as well as on voucher specimens collected in the experimental plot. Results /. Cytology The chromosome number of 322 plants from 161 localities in 19 European countries was determined. The results of the counts are given in an appendix at the end of this paper. The material is arranged according to the alphabetical order of countries of provenance, and the place of origin and plant number are given. Diploids (2n 22) and tetraploids (2n 44) were observed. The tetraploids were found throughout Europe. The diploids were in the minority and were encountered in the central and southern parts of Europe only. Figure 1 shows the distribution of localities. The diploids are represented by an open dot, the tetraploids by a black dot. - The chromosome portrait of both cytotypes is regulär and no B-chromosomes or satellites were observed. 2. Morphology Comparative morphological investigations were carried out and attention was paid to the height of the plant, the length of the stem, the length of the internodes, the shape of the panicle and the length of their flowering branches, the length and width of the leaves, the length of the pedicels, the diameter of the flowers, the length of the stamens and the size of the fruits. Botanica Helvetica 94/2, 1984 271 a) Diploids Within the diploids three groups could be recognized. One group of well-developed plants, 50-80 cm high, with erect stems. The lower parts glabrous, the upper parts be Coming increasingly puberulent with short curved (not uncinate) hairs. Flowers and fruits glabrous. Flowering branches short, shorter than or exceptionally as long as their corresponding internodes, forming a narrowly cylindrical poorly flowering and in- terrupted panicle. The leaves are linear, up to 2.5 cm long and up to 2 mm wide. Com- pared to the other two groups the plants of this group have relatively long pedicels (up to 3 mm) and relatively large flowers (3-4 mm). - The plants characterized in this way were collected in the French Alps (Hautes Alpes, Alpes Maritimes and Savoie) and in ,V„w *i •^ > <*> VI) s> (7 fc 73 9 * •~l ^ »4 0 <o ö Fig. 1. Localities of the cytologically investigated plants of Galium verum L. The diploid is repre- sented by an open dot, the tetraploid by a black dot. The no. 73 represents the number of localities investigated for the Netherlands. 272 E. Kliphuis the south of Switzerland (Ticino). (Collection numbers: Kill, K224, K559, K618, and K3162). In these regions this type of G. verum is commonly distributed in meadows that are fairly moist. - The flowering time in nature as well as in the experimental plot extends from the second half of June into the first week of July. The second group comprises plants collected in Bulgaria and Greece. (Collection numbers: K 1497, K 1502, K3392, K3409, K3418, and K3427). These plants are robust, up to 125 cm high with stout erect stems. The flowering branches, which are al- ways longer than their corresponding internodes, form a loose elliptic or elliptic to nar- row-ovoid, rich-flowering panicle, which is sometimes faintly interrupted. The leaves are narrow-linear, up to 4 cm long and 2 mm wide; outside the inflorescences the leaves are often much longer than within. The pedicels are short, up to 1 mm in length and the flowers whose diameter is generally less than 3 mm, are relatively small. With exception of the flower parts the whole plants may be densely hairy. - The flowering time last from the first half of July into the first half ofAugust. The third group is formed by plants from the central parts of Europe (Poland and Hungary). These plants are up to 100 cm high with a loose faintly interrupted relatively poor-flowering panicle; the flowering branches in the lower parts are shorter than their corresponding internodes and in the upper parts they are as long as or longer to much longer than the corresponding internodes.
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