Изучение отдельных таксонов

В этом разделе приведены результаты разностороннего изучения отдельных арктических таксонов в России, Северной Америке и Западной Европе, опубликованные в монографии "The Species Concept in the High North - A Panarctic Flora Initiative" (на английском языке). Они отражают различия таксономических традиций и очень разные подходы к решению проблемы. Фенетическая пластичность и связанные с нею сложности в разграничении видов обсуждаются в статье S.G. Aiken на примере злаков и статье B.Eriksen & J.Nyléhn на примере рода Potentilla. Род лапчатка (Potentilla) особенно сложен и B.Eriksen и Б.А. Юрцев анализируют и систематизируют различные типы опушения and и обсуждают роль этих признаков для систематики этого рода. Первые результаты разностороннего изучения комплекса Potentilla nivea complex in в Северо-западной Европе приводятся в статье Nyléhn.

Обзоры двух больших таксонов представлены Б.А. Юрцевым в статье об арктических бобовых, особенно о роде Oxytropis и В.В. Петровским в статье о роде Papaver в Российской Арктике. Таксономия рода Papaver обсуждается также в статье H. Solstad et al., посвященной применению молекулярных методов, в которой обосновывается выделение рас в североатлантическом секторе Арктики. Еще один ведущий арктический род обсуждается в статьях B.Andersen et al. о комплексе Draba hirta и A.-C. Scheen et al. о Draba lactea и возможных родственных видах. В этих исследованиях использовалмсь как морфологические, так и молекулярные методы. Комплкксные подходы нашли также отражение в статьях H.R. Siegismund & M. Philipp о роде Dryas на Свальбарде, A.K. Brysting & A. Hagen о североатлантических представителях рода Cerastium (комплекс Cerastium alpinum-arcticum) и S. Fjellheim et al. о комплексе Festuca brachyphylla на Свальбарде.

Эти статьи в целом отражают широту комплекса так и не решенных проблем в таксономии арктических сосудистых растений и большое значение новых критических обработок с использованием комплексного подхода с применением молекулярнобиологических методов. Даже хорошо изученные таксоны оказываются проблеатичными и сложными при углубленном изучении с использованием методов биосистематики.


Aiken, S. G. 1999. Challenges of the species concept in Arctic grasses based on North American experience. - Det Norske Videnskaps - Akademi. I. Mat. Naturv. Klasse, Skrifter, Ny serie 38: 161-171.
Arctic grasses challenge the species concept in at least the following ways: They often have high, and sometimes, variable ploidy levels (e.g. Poa arctica and Poa alpigena). They contain taxa that are known or suspected to be hybrids (e.g. Pucciphippsia and Poa hartzii). They may be phenotypically plastic (e.g. Arctagrostis latifolia and Festuca auriculata/lenensis). They may look similar due to the harsh arctic environments, in spite of being genetical different entities (e.g. Festuca hyperborea/edlundiae).
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Keywords: phenotypic plasticity, hybridization, Arctagrostis, Festuca.
S. G. Aiken (e-mail: saiken@mus-nature.ca), Canadian Museum of Nature P.O. Box 3443, Station D, Ottawa ON K1P 6P4, Canada. Tel. +1 613 364-4073, Fax 613 374 4027.


Andersen, B., Elven, R., Nordal, I. & Spjelkavik, S. 1999. Species in the Draba 'hirta' complex in the North Atlantic area. - Det Norske Videnskaps-Akademi. I. Mat. Naturv. Klasse, Skrifter, Ny serie 38: 173-182.
Material of Draba arctica (subsp. arctica, Svalbard), D. cinerea s. l. (North Fennoscandia), D. glabella (Svalbard) and D. magellanica (South Chile) has been analyzed for isoenzymes. The first three taxa together with D. arctica subsp. arctica, subsp. groenlandica (=D. oblongata, Greenland), subsp. ostenfeldii (Greenland) and the more distantly related D. arctogena (Greenland) and D. norvegica (Svalbard) were analyzed for morphological variation. All the taxa are polyploids: hexaploids (D. arctogena, D. cinerea, D. norvegica), octoploids (D. arctica subsp. groenlandica, D. glabella) and decaploids (D. arctica subsp. arctica, subsp. ostenfeldii). In a multivariate morphological analysis, all taxa constituted clear clusters except for D. arctica subsp. arctica and subsp. groenlandica which cluster together. Draba cinerea s. l. was split in two clusters, one corresponding to D. cinerea s. str., one to the otherwise Siberian D. parvisiliquosa. The former was found in North Finland and Greenland, the latter in North Norway and West Greenland. Draba arctica subsp. ostenfeldii was distinct from the other entities of D. arctica s. l. Draba glabella was distant from the other members of the complex. No structured morphological variation was found within Svalbard D. arctica. Enzymatically, the species fell into four groups corresponding to D. glabella, D. arctica, D. cinerea and D. magellanica. The first-mentioned was the most distant. The two morphological entities within D. cinerea s. l. were represented by separate isoenzyme multilocus phenotypes, but the level of difference between these two phenotypes were the same as between phenotypes within D. arctica and D. glabella. The geographically very distant D. magellanica clearly belongs in this otherwise very northern species group. The West European and Russian separation of D. arctica s. l. from D. cinerea s. l. seems to be well-founded; both should be considered as separate species. The Russian separation of D. parvisiliquosa from D. cinerea s. str. is less evident and should be investigated further. The variation within D. arctica s. l. in Greenland is problematic; from morphological evidence, subsp. ostenfeldii merits recognition whereas subsp. groenlandica (=D. oblongata) only differs in ploidy level.
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B. Andersen & R. Elven (e-mail: reidar.elven@toyen.uio.no), Botanical Garden and Museum, University of Oslo, Trondheimsvn. 23B, N-0562 Oslo, Norway
Inger Nordal (e-mail: inger.nordal@bio.uio.no), Biological Institute, University of Oslo, Box 1045 Blindern, N-0316 Oslo, Norway
S. Spjelkavik (e-mail: sigmund.spjelkavik@unis.no), University Studies in Svalbard (UNIS), Norway

Brysting, A.K. & Hagen, A. 1999. Species in polyploid complexes? The Cerastium alpinum-C.arcticum complex. - Det Norske Videnskaps - Akademi. I. Mat. Naturv. Klasse, Skrifter, Ny serie 38:183-190.
The polyploid Cerastium alpinum - C. arcticum complex presents several phylogenetic and taxonomic problems. Molecular and morphological data argue against the use of a wide species concept in this complex. What used to be called C. arcticum Lange should be kept separate from C. alpinum L. but subdivided into an arctic species, C. arcticum sensu stricto (occurring in Greenland, Svalbard and NE Canada), and a non-arctic, southern species, C. nigrescens (H.C. Watson) Edmondston ex H.C. Watson. Relationships with some Central European taxa are also discussed.
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Key words: reticulate evolution, polyphyletic origin, Cerastium nigrescens.
A. K. Brysting (e-mail: a.k.brysting@toyen.uio.no) & A. Hagen, Botanical Garden and Museum, University of Oslo, Trondheimsvn. 23 B, N-0562 Oslo, Norway.

Eriksen, B. & Nyléhn, J. 1999. Cases of phenotypic plasticity in leaves of Potentilla L. (Rosaceae). - Det Norske Videnskaps-Akademi. I. Mat. Naturv. Klasse, Skrifter, Ny serie 38: 191-199.
Greenhouse cultivation of four species of Potentilla from arctic and alpine North America and Europe resulted in radically changed leaf morphologies in the individuals subjected to the treatment. Individuals having ternate leaves in their natural habitat produced digitate leaves in the greenhouse, and individuals with digitate leaves developed pinnate ones. Favourable temperatures in combination with increased nutrient availability seem to induce a production of supernumerary leaflets. It was previously hypothesized that hybridization was responsible for the variation in leaf morphology. Our observations show that phenotypic plasticity should not be underestimated as an important a factor behind the variation found in natural populations.
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Key words: leaf morphology, hybridization, cultivation, Alaska, Svalbard.
B. Eriksen (e-mail: bente.eriksen@systbot.gu.se), Department of Botany, Evolutionary Botany, University of G?teborg, Box 461, SE-405 30 G?teborg, Sweden.
J. Nyléhn (e-mail: jorun.nylehn@bio.uio.no), Department of Biology, University of Oslo, P.O.Box 1045 Blindern, N-0316 Oslo, Norway.

Eriksen, B. & Yurtsev, B.A. 1999. Hair types in Potentilla sect. Niveae (Rosaceae) and related taxa – terminology and systematic distribution. - Det Norske Videnskaps - Akademi. I. Mat. Naturv. Klasse, Skrifter, Ny serie 38: 201-221.
This paper is a descriptive study concerned with hair types and their occurrence in primarily Potentilla sect. Niveae. One of our tasks has been to describe and illustrate the hair types found in the group and bring consistency into the terminology of leaf indumentum. We confirm that hair types in general carry important systematic information, and that hair characters, together with various floral characters, are fundamental for the presently accepted classification into subgenera and to some extent into sections. Our studies show, that by adding ultrastructural hair characters it is possible to distinguish between three of the major species aggregates described within the Niveae section on hair characters alone. Plants with pinnate leaves (sect. Multifidae) and plants with palmate leaves (proposed hybrids) are, when their hair type is considered, distributed throughout the aggregates formed by the ternately leaved members of section Niveae. Considering the evidence accumulating from cultivation of species belonging to these groups, where leaflet number is shown to be phenotypically plastic and probably related to nutrient status of the individual, we suggest that proposed intersectional hybrids are investigated in more detail and that the currently accepted classification into sections is reevaluated and perhaps redefined.
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Keywords: glands, tomentum, crispate, floccose, hair surface ultrastructure, verrucae.
B. Eriksen (e-mail: bente.eriksen@systbot.gu.se), Department of Botany, Evolutionary Botany, University of Göteborg, Box 461, SE-405 30 Göteborg, Sweden

Fjellheim, S., Guldahl, A.S., Elven, R. & Spjelkavik, S. 1999. The Festuca brachyphylla complex in Svalbard: nomenclatorial chaos but taxonomic simplicity? - Det Norske Videnskaps - Akademi. I. Mat. Naturv. Klasse, Skrifter, Ny serie 38: 223-237.
The Festuca brachyphylla complex is represented by four species in Svalbard: F. baffinenis Polunin (2n=28), F. brachyphylla Schultes & Schultes fil. (2n=42), F. edlundiae S. Aiken, Consaul & Lefkovitch (2n=28), F. hyperborea Holmen (2n=28). The four species are clearly supported by isoenzyme and RAPD analyses. Some deviating plants from the airport area near Longyearbyen might represent hybrid taxa or an introduced foreign strain of the elsewhere rather variable F. brachyphylla. Most of the taxa are bound to more or less open ground, the exception being F. baffinensis which often is found in closed meadows. F. brachyphylla is usually associated with relatively thermophilous species in dry sites. F. hyperborea is found in more mossy damp sites, often with a high-arctic species composition. F. edlundiae has a wide range with main concentration on exposed gravel.
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Keywords: morphometry, isoenzymes, RAPD, distribution.
S. Fjellheim, A.S. Guldahl & R. Elven (e-mail: reidar.elven@toyen.uio.no), Botanical Garden & Museum, University of Oslo, Norway.
S. Spjelkavik (e-mail: sigmund.spjelkavik@unis.no), University Studies in Svalbard (UNIS), Norway

Nyléhn, J. 1999. Preliminary notes on the Potentilla nivea complex in Northwestern Europe. - Det Norske Videnskaps-Akademi. I. Mat. Naturv. Klasse, Skrifter, Ny serie 38: 239-242.
Taxa in the Potentilla nivea complex are collected from Svalbard and Fennoscandia. The material is or will be analysed by RAPDs, isozymes and morphometry. Crossing experiments are carried out on the plants from Svalbard (within and between taxa), and the offspring will be analysed by isozymes and morphometry. Based on the investigations so far, the collected taxa P. chamissonis, P. insularis, P. nivea ssp. nivea and P. nivea ssp. subquinata are probably separate, but closely related, taxa.
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J. Nyléhn (e-mail: jorun.nylehn@bio.uio.no), Division of Botany and Plant Physiology, University of Oslo, P.O.Box 1045 Blindern, N-0316 Oslo.

Petrovsky,V. V. 1999. The problem of the species-width in poppies of Russian Arctic. - Det Norske Videnskaps-Akademi. I. Mat. Naturv. Klasse, Skrifter, Ny serie 38: 243-259.
There are at least 25 species of Papaver belonging to section Meconella Spach. The extent of their distribution, the large number of populations, the great morphologic variation, and the ecologic plasticity of the species have lead to an inequality of the natural entities included in these species and, therefore, the inequality of the species themselves. The necessity to unify the interpretation of species-width within section Meconella is obvious. It is assumed that revision of the f closely related species within the aggregate taxa may clarify their affinity and the relationships. A lot of herbarium material has been accumulated since 1975 when A. Tolmachev published the last critical review of poppies for the Russian Arctic. New data l provide background for a reassessment of arctic poppies. A comparison of taxa within species-aggregates provides an opportunity to revise and review the taxonomic rank of some species earlier described as well as to identify more correctly the systematic status ofothers.
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Keywords: Species concept, morphological variation.
V.V.Petrovsky, Dept of vegetation of the Far North, Komarov Botanical Institute, ul. prof. Popova, 2, 197376, St.-Petersburg, Russia

Scheen, A.-C., Grundt, H. H., Brochmann, C., Elven, R. & Spjelkavik, S. 1999. The arctic hexaploid Draba lactea and is putative progenitors. - Det Norske Videnskaps-Akademi. I. Mat. Naturv. Klasse, Skrifter, Ny serie 38: 261-268.
The genus Draba is well known for its complex morphological and chromosomal variation, and the genus is important and widespread in the Arctic. We are currently working with the hexaploid D. lactea and its putative diploid progenitors D. fladnizensis, D. nivalis and D. subcapitata. The aims of this study are to map the evolutionary pathways in this group of taxa and to revise the group taxonomically. Only preliminary results and conclusions are presented. We have found no signs of local formation of D. lactea in Svalbard from the diploid relatives. Based on molecular markers, D. subcapitata and D. fladnizensis seem to be closely related and to be equidistant from D. lactea, whereas D. nivalis is more distantly related to the three other taxa. The taxonomic distinction between D. lactea and D. fladnizensis is maintained, thus supporting the traditional morphological species concept in this case.
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Keywords: polyploidy, isozymes, RAPDs, morphology, evolutionary pathways, Brassicaceae, Draba fladnizensis, Draba nivalis, Draba subcapitata.
A.-C. Scheen (e-mail: a.c.scheen@toyen.uio.no), H. H. Grundt (e-mail: h.h.grundt@toyen.uio.no), C. Brochmann (e-mail: christian.brochmann@toyen.uio.no), R. Elven (e-mail: reidar.elven@toyen.uio.no), Botanical Garden and Museum, University of Oslo, Trondheimsvn. 23B, N-0562 Oslo, Norway
S. Spjelkavik (e-mail: sigmund.spjelkavik@unis.no), University Courses in Svalbard (UNIS), Box 156, N-9170 Longyearbyen, Norway

Siegismund, H.R. & Philipp, M. 1999. The genus Dryas on Svalbard: morphological and genetic evidence for a single species, Dryas octopetala L. - Det Norske Videnskaps-Akademi. I. Mat. Naturv. Klasse, Skrifter, Ny serie 38: 269-279.
On Svalbard Dryas populations can be subdivided into individuals with or without large sessile glands on the upper surface of the leaves. The group possessing large sessile glands has been referred to Dryas punctata Juz., while those without the glands to D. octopetala L. In order to analyse whether the gland characters are correlated to other differences between these two groups of individuals, a number of morphological and isozyme characters were investigated in four populations. We found that differences among populations were larger than between the two groups of plants with or without large sessile glands. The conclusion is that there is no reason for maintaining more than one species of Dryas on Svalbard: D. octopetala.
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Keywords: morphometry, isozyme electrophoresis, population differentiation, Svalbard
H. R. Siegismund & M. Philipp (e-mail: marianp@bot.ku.dk), Department of Plant Ecology, University of Copenhagen, Øster Farimagsgade 2D, DK-1353 Copenhagen K, Denmark

Solstad, H., Elven, R. & Nordal, I. 1999. Are there too many species and subspecies in the Papaver radicatum complex? - Det Norske Videnskaps - Akademi. I. Mat. Naturv. Klasse, Skrifter, Ny serie 38: 281-294.
Analyses of isoenzymes and RAPD markers does not support subspecific delimitation of Papaver radicatum in South Norway/Iceland. The analyses of the isoenzymes revealed further that Nordic P. radicatum (decaploid), P. lapponicum (octoploid) and P. laestadianum (octoploid) constitute a group of closely related taxa, which even might be reduced to the same species, which may have originated by auto-polyploidization from P. alpinum (diploid). Within the Nordic Scapiflora poppies P. dahlianum is the most distinct species, including populations both on the northenmost, arctic, Norwegian main-land and on Svalbard. It is a wide spread circumpolar species and includes P. polare, which should be reduced to synonymy.
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Keywords: isoenzymes, RAPD, Island, Norway, Svalbard.
H. Solstad & R. Elven (reidar.elven@toyen.uio.no), Botanical Garden & Museum, University of Oslo, Trondheimsveien 23b, N-0562 Oslo, Norway
I. Nordal (inger.nordal@bio.uio.no), Department of Biology, P.O.Box 1045, Blindern, N-0316 Oslo, Norway

Yurtsev, B. A. 1999. Survey of arctic legumes with emphasis on the species concept in Oxytropis. - Det Norske Videnskaps - Akademi. I. Mat. Naturv. Klasse, Skrifter, Ny serie 38: 295-318.
The results of critical revision of the arctic legumes (in particular, the genus Oxytropis) are based on the treatment of the family for the “Arctic Flora of the USSR”. In the Arctic floristic region, here accepted in its restricted meaning, 109 indigenous species and subspecies of legumes occur, belonging to 8 genera of 5 tribes (Faboideae). Species diversity of legumes in the Arctic is uneven both geographically and taxonomically. Legumes are lacking in 6 areas of the Amphi-Atlantic sector, but reveal a high species concentration in the East-Siberian and especially Amphi-Beringian (Chukotka + Alaska–Yukon) sectors with up to 29–30 species per area. 92 species and subspecies (of the total of 109 taxa) belong to the tribus Galegeae: 27 to the genus Astragalus and 64 to Oxytropis. Obligatory outcrossing (pollination by bumblebees) has resulted in predominantly allopatric race- and species-formation which is not seldom complicated by polyploidy. In the sections Orobia and Glaeocephala of the genus Oxytropis, differentiation of polytypic species into vicarious races is widespread. Accepting wide “species” with many tens of subspecies throughout the Holarctic (like Oxytropis campestris s. ampliss.) is practically inconvenient and complicates phylogenetic analysis of the group. Acclimatization of the progenitor of a taxon at a new adaptive zone, with transformation of morphotype, favours speciation (sect. Arctobia, sect. Baicalia of the genus Oxytropis), as also does a long isolation of relict populations. Legumes provide many differential (including endemic) and co-differential species of floristic sectors and areas of the Arctic.
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Keywords: speciation, arctic legumes, Oxytropis, allopatry, species/ subspecies.
B. A. Yurtsev, Komarov Botanical Institute, Russian Academy of Sciences, ul. prof. Popova, 2, 197376 St. Petersburg, Russia.

 


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