L.) Klinge Two New Genera: Orchis (L.) Vermln
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Cytological observations in relation to the taxonomy of the Orchids of the Netherlands E. Kliphuis (.Botanical Museum and Herbarium, Utrecht) (;received December 10 th, 1962) Abstract 1. The Orchids in the Netherlands have been subjected to a cytological in- vestigation. 2. The division of the Orchis into genera (L.) Klinge two new genera: Orchis (L.) Vermln. and Dactylorchis (Kl.) Vermln. (Vermeulen, 1947), could be confirmed. 3. In Listera ovata (L.) R. Br. the diploid chromosome number is 34. Deviating numbers 2n 35 = and 2n = 36 were counted. Because aberations in chromosome number do not cause differences these aberations be morphological seem to un- important. 4. Out of the material it investigated might be concluded that for the moment it does not to be correct to consider Vermln, seem Dactylorchis fuchsii (Druce) as a besides maculata separate species Dactylorchis (L.) Vermln. It seems more likely that D. and fuchsii D. maculata represent two types within a complex-species. Introduction For time the a long Orchidaceae of the temperate regions have had the attention of The Dutch botanist Treub cytologists. was among the first to mention the chromosomes of the Orchidaceae in his study on the mitosis in Orchis and Some laterGuignard Epipactis (1879). years published his investigations on the meiosis of Listera ovata (1884). Already Strasburger (1882) suggested that the chromosome number is generally the same within a species, though closely related number. species may have the same Since the end of the nineteenth have century many investigators determined the chromosome number of numerous species of plants. For the Orchidaceae of the regions of the northern hemisphere the following investigators are to be mentioned: Afzelius (1922, 1943), Barber (1942), Fuchs and Ziegenspeck (1924), Guignard (1884), Hagerup (1938, 1941, 1944a, b, 1945, 1947), Heusser (1938), Hoffmann (1929, 1930), Love and Love (1942, 1944, 1956), Mauer (1938), Modelewski (1918, 1938), Muller (1912), MacMahon (1936), Richardson (1933, 1935), Rosenberg (1905), Sokolovskaja and Strelkova (1940, 1960), Staner (1929), Tuschnjakova (1929), Vermeulen (1938, 1947, 1949), Weijer (1952). Apart from the number, however, the size and the shape of the chromosomes may be a character of importance to taxonomy. The morphology of the chromosomes inListera ovata in particular has been amply investigated, partly on account of the striking differences in size and and also because be sub- shape this species was the first to 172 CYTOLOGICAL OBSERVATIONS - ORCHIDS 173 jected to thorough cytological investigations (Guignard (1884), Rosenberg (1905), Muller (1912), Tuschnjakova (1929), Hoff- mann (1929, 1930), Richardson (1935), MacMahon (1936). A and more general extensive work on the morphology of Orchid chromosomes has been published by Heusser (1938). His investiga- tions comprised the basitonic Orchids of Switzerland. With the exception of those belonging to the Dactylorchis- group*)) and in a lesser those of the Orchis- the Orchids of the degree group Nether- lands have been extensive not subjected to any cytological research yet. So it is of importance to have a chromosome study on the Orchids still be found in the to Netherlands, also including those that are now extinct there, but until recently, were a part of the local flora. Material and methods The plants were collected in the field, dug out with a considerable of soil quantity and reared in pots in a non-heated greenhouse or in the open. In the preliminary stage only squash slides were made. Roottips and flowerbuds were used directly as well as after Carnoy fixation. For staining orceine, aceto-carmine or leuco-basic-fuchsin were used. The were and therefore microtome sections results, however, very poor also made later This were on. technique gave excellent results and the of was, in this case, certainly preferable to use squashes. For the microtome sectioning roottips and also flowerbuds were fixed in Karpechenko and embedded in paraffine. Microtome sections of 15 micron were stained with cristal-violet or according to Heiden- hain’s haematoxylin method, or in some cases with leuco-basic- fuchsin according to Feulgen. The best slides were obtained with cristal-violet. the of this the However, degree staining depended upon season in which the material fixed. When cristal-violet was gave no satisfactory results one of the other two mentioned staining methods was used. Drawings of the metaphase plates of the first—or second division of the pollenmothercell or of the pollendivision and/or of the metaphase plates of roottip cells were made with the aid of an Abbe Camera Lucida. Results The results with respect to the chromosome numbers are given in the following table. In the first column: the species; in the second: the determined chromosome number; in the third: the data of literature. * The nameDactylorchis Vermin. (1947) is mostly adopted in modern literature, however, Dactylorchis Vermin, is a nomenclatural of Dactylorhiza Nevski synonym - (1937) (Bullock 1959 Taxon 8: 46). 174 E. KLIPHUIS 40 2n — — 36 — 36 36 — — — — 42 42 42 — — 42 42 — — 42 42 — — — 40 40 — 80 80 80 80 80 80 60 80 20, n 18 18 18 18 18 18 18 18 18 21 21 — 2121 21 21 — 21 21 21 — 21 20 20 — — — 40 — — 40 — — — — 40 1956 1951 1949 1949 1948 1947 1948, 1956 1957 1957 1944, 1944 1956 al. 1947, 1949 1949 al. 1947, 1949 1938, 1938 1947 1957 Love Kaad Love Literature 1938 et 1938 1938 1938 1938, 1943 1938 1938 1938 1938 1938 1938 et 1938 1958 1938 1942 1942 1942 1942 & Heslop-Harrisonand Heslop-Harrison 1951 and Afzelius Barber Heusser Barber Heusser Heusser Hagerup Skalinska Vermeulen Hagerup Heusser Vermeulen Heusser Vermeulen Hagerup Heusser Skalinska Vermeulen Hagerup Heusser Skalinska Vermeulen Afzelius Barber Love Vermeulen Barber Hagerup Heusser Holman Love Love Vermeulen Vermeulen 38 38 100 2n 36 42 42 42 42 40 120 36, 36, 80, n 18 18 — 21 — 21 — 20 40 . Vermin.. Vermin. Species (Druce) (L.) L Huds Huds L L L Lam fuchsii maculata insectifera apifera morio purpurea simia militaris mascula Ophrys Ophrys Orchis Orchis Orchis Orchis Orchis Dactylorchis Dactylorchis CYTOLOGICAL OBSERVATIONS - ORCHIDS 175 82 80 2n — 40 40 40 80 80 80 — 80 80 80 80 — — 40 40 40 — 40 40 40 40 40 — 42 42 42 — 42 42 — 36 42 36 80, 40, n 20 — — — 40 — — — 40 — — — 40 20 20 40 20 — — 20 — — 20 — — 21 21 21 — 2121 — 21 18 18 21 18 1960 1949 1960 1956 1956 1947, 1947 Strelkova 1957 1957 1944, Strelkova 1957 1957 & Kaad 1947 1938, 1938, al. 1938 1947 1935 al. 1935 al. 1935 al. 1935 1935 Literature 1938 1947 1938 Love & 1947 1938 1938 et 1943 1938 et 1943 1938 et 1922 1938 et 1922 1938 1938 1938 & 1939 1942 1942 and HagerupHagerup Heusser Holmen Vermeulen Maude Vermeulen Vermeulen Hagerup Heusser Skalinska Vermeulen Vermeulen Afzelius Barber Heusser Richardson Skalinska Sokolovskaja Afzelius Heusser Love Richardson Sokolovska Skalinska Afzelius Heusser Richardson Skalinska Afzelius Hagerup Heusser Richardson Barber Heusser Richardson Heusser 2n 40 80 80 80 40 40 42 42 36 — n 20 40 40 40 20 — 21 21 — 18 junialis var. Vermin Vermin, Rich Spreng Vermin Vermin Br Rich (Druce) (Druce) R. Hartm (L.) Species Rich (Gust.) (L.) (L.) (Rchb.) (L.) (L.) (L.) chlorantha hircinum incarnata praetermissa praetermissa Vermin majalis conopsea viride bifolia pyramidalis Dactylorchis Dactylorchis Dactylorchis (Vermin.)Dactylorchis Gymnader.ia Coelcglossum Platanthera Plathanthera Anacamptis Himantoglossum 176 E. KLIPHUIS 36, 36 36 40 38 34 36 38 35, 2n — 42 — 32 — — 40 — 40 — 38 40 — — 40 — — — — 35, — 34 — 38 — 36 30 30 — 40 — 34,34, 37, 32, 34, 36, 34, 34, 18 20 20 18 16 19 17 16 17 19 16 17 17, 18 15 16 n 21 21 18 16 20 — — — — 20 — — — — — — — — — — 20 14 19, (10) 17, 16, 1948 1930 1957 1944 1959 1960 1947 1944 1944 1956 1957 1956 Strelkova 1957 1929 . 1936 1933 & 1935 . al. Love al. al. Love 1884 1929,Love Love 1905 al. Love al. Literature 1947 1944 1945 1947 et et 1944, 1947 et et 1941 1944 1938 et 1938 1942 1942 1942 1952 1942 1912 1929 1942 and and and and Tuschnjakova and Sokolovskaja 1954 Barber Heusser Barber Hagerup Skalinska Hagerup Love Barber Hagerup Hagerup Skalinska Skalinska Weyer Hagerup Love Barber Guignard HagerupHoffmann Love Love MacMahon Müller Richardson Rosenberg Skalinska Staner Love Barber Skalinska Love Richardson Hagerup Heusser Hagerup 35, 2n 42 — 40 38 40 36 40 — 30 26 40 — 34, 18 19 17 18 15 14 n 21 20 — 20 . .— . Druce Br Schuit R. (Mill.) Br Crantz Crantz O.K R. Species (L.) Br Br Rich (L.) (L.) (Hoffm.) Br R. (L.) R. Rich (Mill.) (L.) damasonium R. (L.) (L.) (L.) (L.) paludosa anthropophorum palustris helleborine atrorubens repens monorchis ovata cordata nidus-avis loeselii Aceras Cephalanthera Epipactis Epipactis Epipactis Listera Listera Neottia Goodyera Liparis Hermium Hammarbya - 177 CYTOLOGICAL OBSERVATIONS ORCHIDS Ophrys insectifera L. Ophrys insectifera L. 2n = 36 Fig. 1a. 2n = 38 Ophrys apifera Huds. Ophrys apifera Huds. = 1b. = n 18 Fig. 2n 36 Orchis morio L. Orchis morio L. 36 1c. c. = 38 2n = Fig. 2n 178 E. KLIPHUIS Huds Orchis Orchis purpurea purpurea Huds. n = 21 2n = 42 Fig. 2a. Orchis militaris L. Orchis militaris L. = 21 2n 42 n = Fig. 2b. Orchis simia Lamk. Orchis mascula L. 2n = 42 2n = 42 Fig. 2c. CYTOLOGICAL OBSERVATIONS - ORCHIDS 179 Dactylorchis fuchsii (Druce) Vermln. Dactylorchis fuchsii (Druce) Vermln. n = 20 2n = 40 Fig. 3 a. Dactylorchis maculata (L.) Vermln. Dactylorchis maculata (L.) Vermln. n = 40 2n = 80 Fig. 3b. Dactylorchis maculata (L.) Vermln. Dactylorchis maculata (L.) Vermln. 2n = 100 2n = 120 Fig. 3c. 180 E. KLIPHUIS Dactylorchis incarnata (L.) Vermln. Dactylorchis incarnata (L.) Vermln. n = 20 2n = 40 4a. Fig. a. Dactylorchis praetermissa Dactylorchis praetermissa (Druce) Vermln. n = 40 Fig. 4 b. (Druce) Vermln. 2n = 80 Dactylorchis praetermissa (Druce) Dactylorchis praetermissa (Druce) Vermln.var.