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Download Cytotaxonomic notes on some Galium species. A - UvA-DARE
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Cytotaxonomic notes on some Galium species. A BY Prof. (Communicated by F. E. Kliphuis A. Stapleu at the of March meeting 30, 1974) Introduction Within the the frame work following species were of general cytotaxonomic studies of various Galium species included: Galium rotundifolium L., Galium arenarium Galium triflorum Mich., Galium uliginosum L., Galium pumilum Murr., Galium oelandicum Ehrend., Galium tricornutum Combined The results conditions of these Huds. morphological during several investigations Galium Hyl.) Ehrend., et verrucosum and cytological uniform under (Stern, Galium Dandy, are and studies Galium on Galium L. purpureum made were spurium L., Lois., sterneri cultivated plants years. described and discussed for each species separately. Material and methods The chromosome numbers determined The roottips In some cases plants Results a mitosis. roottips fixative, embedded in paraffin, Karpechenko’s and stained number of to according Galium plants from mentioned under are and are indicated Heidenhain’s haema- rotundifolium section by population one the an investigated. was collection same number. Such asterisk. discussions rotundifolium L. Galium This micron numbers collection 1. 15 on method. toxylin These fixed in were at sectioned made were is L. belongs characterized whorls of four, and 1). (fig. to the section by hermaphrodite plants Platygalium (DC.) Koch. three-veined leaves having in flowers with white corollas in terminal panicles. Galium East a rotundifolium has pubescent, to to 40 Asia cm Flowers loose often up to panicle, Fruits 2 mm 3 Minor into an erect the Central, South and South- Caucasus. four-angled stem, It is a perennial glabrous cm long and 0.5-0.8 turning reddish-brown, mm in diameter in greenish white or cm wide, widest and black ovate long, densely covered with with just ovate above when dried. spreading axillary white, with sometimes or high. Leaves in whorls of four, short-petioled, 1-1.5 sub-orbicular, the middle, a and Europe creeping stock and its distribution in cymes apiculate uncinate hairs. forming lobes. 346 Galium rotundifolium Fig. 1. L. 347 Flowers The from collected from middle of different several during the is uniform in its species Cytological years of the parts in the investigation until May July. morphology. This experimental this on et al. diploids do not regions Himalayas to the belong Ehrendobfer and the (1973) closely related species. area Origin of the Andorra: K-1019, Austria; wald, Heide, Ponta near K-1267, 10.3) d’Envalina. Nieder-Oesterreich; K-1083, Herschafts- Dubner Steinach, Ober-Franken, Bavaria; K-717, near do Serra Geres, Geres; K-1268, Jihlavsk6 : Wallis; K-1238, K-714, K-525, near arenarium Cytological K-713, Zagreb; Lois. investigations hexaploids with (fig. on number could not made were collected (Lesconil: Mettmen, ca. of Moravia; Eastern 1360 m, Glarus. K-569, Posazavi, part of Bohemia; K-728, K-281, Serajewo. near 2). plants of Galium arenarium Lois, showed chromosomes. In the literature Faqerlind The counts Leonte, Serra do Geres. 2n =66 chromosomes. The chromosome without additional tetraploid Rio Bohemia. Predjumavi, Galium near Northern part Vrehy, Bohemia; Yugoslavia; near K-1269, Tatri, Dolina Lejowa. K-696, Switzerland: K-566, in the on (Fig. (1937) reported be 2n =44. In material grown up from seeds K-700; is regular our study this confirmed. wild in France K-295, K-561, portrait 10.1) of the Plougernan: as Department K-554 and wel as from of Finistere Cinder: K-697), Department of Morbihan (Plouharnel: K-556) and of the Depart- ment of Loire and of the Messagnes Atlantique (near Nantes: K-312, K-350, Department K-1260; plants of Les Landes Vieux-Boucau Plage: K-1265 and The to Bad Diiben. Tsjbchoslovakia and belong to portrait is regular without satel- d’Homen, Serra do Geres; K-1047, K-1270, Poland: of the According these authors Oberwart, Burgenland. Portugal: plants these investigated: K-289, Manhartsberg, Germany: K-545, 2. by reported However, rotundifolium. plants investigated material and Hsu (1968) respectively. of Galium (Fig. also found was (1964) in plants from different Taiwan The chromosome additional chromosomes. or exception showed without parts of Poland. Khoshoo and Bhatia (1963) from the plants on cultivated garden. material by Piotbowicz (1958) and Skalinska and area with 2n =22 chromosomes. This number 2n =22 diploids lites confirmed was distributional K-1231; les St. (Labenne Bains: Giron K-1261, Plage: K-474 and K-522) Oc6an: K-1230, K-1259 K-1262 K-1264; and K-1263; Mimizan Plage: K-1266). were cultivated during several years in the Botanical Garden 348 Galium arenarium Fig. 2. Lois 349 of the State University of Utrecht, which K-1266, constant a of years Galium arenarium is France showed remarkable a cultivation. 10-20 San Sebastian in the North of stem high, cm of the coastal typical representative a from the Channel to perennial, 1973). The plants K-1259- nrs. morphology. All characters mentioned below remained the over collected in were in their uniformity of the (with exception to prostrate regions of It is Spain. reddish at the ascending, base, glabrous, smooth, quadrangular with ribbed angles, much branched, with to leaves numerous 1.5 wide, widest just stout at the end of the cymes Petals acute black, shining. Flowering period: obtained from gardens nature both with and In and 0.3 up to 4 mm (fig. rarely in short 3 to mm, 3). seed two from the botanical received samples and (Canada) Lund is portrait these respect the triflorum is in given fig. chromosomes accessory Over K-353, (K-722, chromosomes. no cm in diameter. fruit short, and (Sweden) collected the North-American continent turned out to be morphological characters. Michaux from The chromosome pattern deep yellow, mucronate. Pedicels not Montreal on 2n=66 long cm juin-september. Galium triflorum Plants rarely at the base. Intemodes to 0.8 teeth, fleshy, darkgreen, shiny. Flowers branches, obtuse, to 10, above the middle, mucronate, linear-lanceolate to linear, bearing small 3. apically, usually defoliated Leaves in whorls of 5-8 cm. of years in hexaploid respectively). 10.6. It shows hexaploid plants cultivation a normal regular detected. could be these similar in are characters their remained constant. Galium (Northern-Norway, Switzerland, and this Within plants reported diploids The finding species of a the distribution of series level cytotypes certain as and a Northem-Europe and occurs also in Sorsa occurs. Khoshoo some parts of relic. glacial (1963) and found in Bhatia (1963) Himalayas. in separated parts of the of the area correlation between chromosome number and chorology. However, should about the considered tetraploid from it Finland), polyploid a of these indicates plant from North-America there it is species Finland a Sweden and this be the case, then still too little is known plants of different ploidy level to give a clear picture. Galium ascending prickles. triflorum or a perennial, Leaves in whorls of 6-8, ovate-lanceolate, clear is erect, quadrangular, midvein, margins rough Flowers sessile or the 30-60 angles large, 1-3 mostly high. Stem sometimes cm decumbent, rough by long and to 1.2 shortly petiolate, mucronate, usually otherwise reticulate-veined, by cm antrorse prickles or glabrous, mostly retrorse cm wide, thin with dark green, smooth. in cymes of three in theleaf axils. Corolla small, greenish. 350 trif lorum Michaux Galium Fig. Fruiting pedicel straight, to 5 mm. 3. Fruit densely covered with uncinate hairs. Flowering Galium 4. time: uliginosum From each of lands and from transported July-August. L. (fig. 4). eighteen populations of one to the in the Belgian to be experimental garden. tetraploids with uliginosum Ardennes 7-8 in the Netherlands and those from the out Galium 2n =44 The plants plants population chromosomes. in the Nether- were dug out and from two from Belgium The populations plants turned from the 351 Galium uliginosum L. Fig. other sixteen Plants from populations collected in collected in the wild, Norway in the literature. diploids with location in Denmark and one tained from seeds were 4. were also Diploids by were from 2n =22 one diploids. diploids. Both cytotypes Homeyer (1932) chromosomes. in France all Two other are also ob- plants reported in material from unknown origin; by Hancock (1942) for Britain; for Central-Europe by Garajova (1959) and Ehrendorfer, lands by by Kliphtjis Love and Love (1962) (in Love and Love, 1961) and for the and Gadella and Kliphuis (1956) for Iceland, by Rohweder Nether- (1963), tetraploids (1937) for Germany 352 and from htjis the Netherlands and (1932) Fagerlind (1934, satellites or Galium could be observed uliginosum is rather height of the plants, leaves, petals and fruits, values plants calculated SE =0.22 and X =30.42 distribution decumbent in 1 to be and diploid plant respectively) a of a 10 It is experimental factor. discriminating The X =30.15 are: observation: fi N =200; ± and respectively. (Number European species with 10-60 perennial a glabrous, weak, little variable. are plants). Central-West Scandinavia. as: of the size level. This could be ploidy tetraploid number a B-chromo- cm quadrangular a northern high. with Stem four ribs prickles. whorls of long, to up 2 veined, one to 7 small cm. the glabrous, Anthers and long cm Pedicels axillary acute. period: the 5-8, up in ovate, mm of no 10.8b several years in the during prove with ascending, Flowers Flowering Origin L. is mucronate, lobes Fruit not the Homeyer morphology. Characters correlated with stomata per Internodes flowering. corolla 20 into or with retrorse prickles. did for uliginosum lanceolate, its ± SE =0.23 and SD =3.23 with N =200 n Leaves not cultivated SD =3.10 of observations: Galium (figs. constant in regular, are 10.8a and and the indument of the are Stomata size garden. portraits length of the internodes and pedicels, Differences if present, demonstrated on Gadella and Klip- reported by were 1937). The 2n =22 and 2n =44 chromosome somes and by Kliphuis (1962) (1963). Tetraploids from unknown origin 2.5 margins 0.5-1.5 mm, white, panicles, mm 3 wide, linear- with recurved deflexed after mm in diameter yellow. glabrous, rugulose. July-August. material investigated: Diploids: Denmark: K-206, France: K-786, Jutland, Igleso La The Netherlands : Roche en K-247*, near Brenil. near Bold Skov. Dept. Duurswoude, K-630*, K-631*, K-633*, K-634*, Speuld, K-660*, K-666*, Holland; K-152*, K-960*, Oost near prov. “de Muy”, Isle between Maarn and Eempolder, near Eemnes, of prov. of Gelderland; Putten, prov. of Gelder- Tessel, prov. prov. of Utrecht; of Noord- of Utrecht; K-37I*, K-372*, K-373*, K-1007*, former airstrip, K-374*, “de Beer”, Goeree, Zuid-Holland. K-1148, roadside near Hovringen, near South of Dovre. Tetraploids: Belgium: of Friesland; K-189*, near Oost Voorne, prov. of Zuid-Holland; Norway: prov. Maasbergen, prov. Voorne, prov. of Zuid-Holland; of d’Or. Arboretum: “De Schovenhorst”, K-190*, Putten, land; Cote K-407*, Robbertsville, Ardennes. Dombas; K-1156, 2 km 353 The Netherlands; between Ermelo 5a. Galium (Stem, et (Stern, Weerdinge, Ehrend. Hyl.) Hyl.) Ehrend. of Drenthe; prov. prov. of are (figs. (Jordan, 5a, b, has tried to closely with clarety more regard distinguished sponding often related species overlapping 1915). Sterner (1944) bring a with Galium to its which forwardly all were in in the taxonomy of European Galium oelandicum Galium pumilum (Sterner et Hyl.) Ehrend Murr. 5. characters. complex species, Europe. morphologically and et morphological study this in North-Western group pumilum Murray s.str., Fig. complex have morphological well founded a pumilum has 1909; Briquet, 1900; Briquet representatives central Galium c). pumilum Murr.. Confusion around Galium unclear and 5c. sterneri Ehrend. and Galium oelandicum 1846; Braun, 1885; Schuster, Cavxllier, especially on K-188*, Gelderland. been great because taxa of various rank within this been based He et pumilum Murr., Galium included in Galium always near Nieuw-Milligen, pumilum Murr.; 5b. Galium sterneri Ehrend. and oelandicum Galium K-32*, and a corre- Northern-European Galium sterneri Ehrend 354 taxa the on subspecific level, (0. Dahl) Nordh.; slesvicence asp. with Stern, Of the variety tetraploids, diploids is distinction in Britain in occur the largest The always The areas. On by diploids are such is 1957). for that cytotypes sterneri by the other hand 1955, (Goodway, the two ssp. Isles. This Ehrend.) degree of overlapping possible. Isles subspecies: one most extensive investigations a part of a pumilum belongs. (1949, Ehrendorfer, that Galium Galium as study reason are Tutin and War- (Clapham, 1953, 1954, forms pusillum complex. through This regular Leptogalium 1958, 1956, investigations, of part a complex expanded from perfectly 1955, the is a an diploid level in Galium (Ehrendorfer, barriers in becoming lesser has been able, with expand Galium hexaploid CentralIt is through pumilum of species West non all nearly partly autoploidy, its occupy present Galium pusillum flowering, smooth, white, to Flowering Plant to pumilum slightly and strong barriers 1973). These 1972, the effectively the means complex to (Ehrendorfer, l.c.). as the octoploid, rarely it and complex, occurs in glabrous hairy, or linear-falcate, 4 mm in leaves to in to erect whorls stems. of 6-9, oblanceolate-falcate, Stems 1-2 cm the upper mucronate. diameter. Fruit with tiny papillae, almost mm. time: remaining Galium species 1.5 to large through alloploidy, perennial with ascending long, the lower narrowly oblanceolate Corolla area. the number Europe. caespitose linear-lanceolate to such Murr. should be understood the are 1970, partly complex, a normal phenomenon a polyploids, higher ploidy levels. By time and and a the the lower in 1963). higher ploidy levels, There 1955, 1958; Kliphuis, effective extremely are larger to against intercrossing between the various cytotypes, 1962, the conclusion to aggregate of by to which Galium 1960, came much series. euploid 1946 onwards made from were of the section result of his a pumilum s.l. of taxa that have basic comprising area sterneri tetraploids. variable and the not suecicum mainly represented disjunct slender habit than very are Ehrendorfer as 5a. and asp. 1962). burg, often has specific rank (as Galium the Flora of the British united under septentrionale Stern.; Hyl. Galium sterneri Ehrend. is a more tetraploids a et Stem.) septentrionale elevated to was smaller and of In variety (sap. normanii ssp. Denmark and the Northern part of the British Ehrendorfer (1960). that vestrogoticum morphologically different one Hyl.; oelandicum Stern, ssp. these the subspecies subspecies of them with one islandicum Stern, et ssp. Stern.; South-Norway, the and comprising six geographically isolated group May-August. green acid which when dried. Murr. occurs, soil, belongs in to on dry grassland the to moderately vegetations Mesobromium dry which well drained are (Ehrendorfer, rich in 1953). 355 because of Probably is restricted species Galium faithful pumilum of species hellinggrasland”) Plants from chromosome is kind a belonging the and the regular, France: Netherlands 2n =88 idem, population 7-10 The plants were were plants transported morphological to the characters plants were of investigated: K-169, K-170, Thoisy K-785, slope Galium Ten K-171*, la near Berchere idem K-992*, near Baelartshoven. (Dyon). somes. Ehrend. taken from These plants (fig. flowering (fig. shoots three populations is portrait 10.7b). or hairy. Leaves in whorls of 7-8, linear, The Fruits Origin pyramidal to of Denmark, 1.25 in cm caespitose, relic pedicels Vorgod (Sterner mm. non erect, up to long. cm wide, oblanceolate mm retrorse 1-2 acute prickles. Inflorescence forming on near et from seeds) the coast; K-1199*, Hyl.) Ehrend. (fig. K-1195*, near Logstor. 5c). (Sweden). It is a diploid, 1960). perennial, densely caespitose, with tubercles. (obtained the isle of Oland (Ehrendorfer, ascending four or investigated: K-233*, is endemic a perennial, 1.5 chromosomes accessory July. material Jutland: The plant is shoots a long, and long, glabrous with mm Galium oelandicum postglacial with 2n =44 chromo- panicle. June and the species is Internodes up to to 1.5 Blavand, Dunes K-1196*, Klim, This regular, no and five investigated cyto- were tetraploids margins with curved, mucronate, Flowers K-995*, of Galium sternen Denmark tetraploid Flowers to 4 mm, creamy white, compact K-994*, prostrate, the flowering shoots ascending glabrous cm, K-993*, 5b). turned out to be The chromosome observed, province of Limburg, Schwei- Kunraderberg; Eyserbos. sterneri plants logically. 5c. satellites or out and dug retained their obtained from seeds all from Jutland, a found. The were 10.7c). berg; K-991*, to investigated were chromosome B-chromosomes no The Netherlands: K-990*, South of the 25 krijt- importance and consisted largely in the indument and the plants size. Belgium; were (“het as Koelerio-Gentianetum. the years of cultivation. Existing differences among the Material 5b. distinctly basic soils. limestone-grasslandvegetation to France is experimental garden. minor of or the circumstances by Westhoff and Den Held (1969) regarded Belgium, (fig. climatological in the Netherlands to neutral portrait From each during favourable Only octoploids with cytologically. observed, less up to short intemodes. Leaves in 15 cm high, flowering whorls of 6-9, linear- 356 and 1.5 panicles. Pedicels short, up oblanceolate, midal Two plants logically. 1 cm were spurium Seeds from Galium Botanical Gardens were also (K-488 diploids L. (fig. with wide. Flowers white, mm to 0.5 and Fruits K-779) 2n =22 were in lax pyra- to up 1 mm investigated chromosomes, long. cyto- 10.7a). (fig. 6). spurium collected of Udine mm. (Italy), in nature and Stockholm, received from the Erevan and Leningrad sown. Counts these long Oland from Both Galium 6. to on metaphaseplates of roottip mitosis from plants obtained from seeds, reported showed by the chromosome Fagbblind (1934, number 1937) in Galium spurium L. Fig. 6. 2n =20. material This number from is unknown 357 origin and by Podlech the Andarab in valley and Dieterle Baghlan, Afghanistan. remarkable in the genus Galium, because the number in this is genus X—11. number X =10 is known, Galium molluginoides The chromosome in the ences H. portrait is this Gadblla and Kliphuis, the Love and 1949; ques, plant same with aparine which 1963) Love, Podlech 1963; 2n =64 chromosomes, Gadblla and Kliphuis, Galium The spurium plant is quadrangular glabrous an and Central rough on flowered four Pedicels Fruits hairs cymes, 1 in mm mm in Galium are present Rahn, 1955; with decumbent 6 retrorse or prostrate Nodes prickles. cm. prickles. Flowers in greenish-white or mucronate, axillary few white with long, black, rugulose, glabrous fruits Gabriels and the there (1965) indumente, subspecies uncinate fruit-indument. e.g. is spurium and Material fruits, infestum According and to to are present, axillary flowers Udine, Val Sweden: K-455, Carlshas. U.S.S.R.: Esthonia; K-268, this the with Armenia, et Kit.) subdivision subspecies Galium Tellina. K-318, characterized subspecies (Waldst. investigated: K-301, with white uncinate subspecies spurium related to Galium closely differences two are the gated plants used in this study belong morphological or tuberculate bases. non to Italy: even of straight. differences in fruit flowers Larsen and by diameter, period: May-September. Galium 1971), long, linear-lanceolate, retrorse According with 1967; Gadella and Strid, large chromosomes high, to cm Flowering by 1967; (Pou- lobes. 1.5-3 with brous 1962, occurs metaphaseplates edges internodes up margins rough by (3-9) acute the is America. cm the Leaves in whorls of 5-10, to 3 up midrib and 1962, plates (Bocher, up to 100 annual, glabrate, or 2n =64 through the whole of Europe (except the artic North stems, the two fusion (1955) 1963). occurs West-Asia, parts), In differ- Bocher et al. the number 2n =66 and 1969). 10.4). No striking 1969; is the basic Hausskn. 1955; Kliphuis, Dieterle, 1967). species et chromosome number 1956; Kliphuis, (Kliphuis, 2n =20 chromosome present and, therefore, are but also and absent in the 2n =66 are the species basic and Dieterle, (Podlech regular (fig. very Larsen and Rahn, (Bocher, common Kliphuis, L. Krause spurium from number other two suggested for Galium aparine by as In unlikely. seems Only from The common e.g. Galium ibicinum Boiss. length of the chromosomes of chromosomes most E. in Galium (1969) aparine. spurium 2-3 with K. gla- Janchen the investi- infestum. However, having leafy bracts. clear smaller