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The Purple Saxifrage, Saxifraga Oppositifolia, in Svalbard: Two Taxa Or One?

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The Purple Saxifrage, Saxifraga Oppositifolia, in Svalbard: Two Taxa Or One? The Purple Saxifrage, Saxifraga oppositifolia, in Svalbard: two taxa or one? ANNE K. BRYSTING, TOVE M. GABRIELSEN, OVE SBRLIBdmN, ODDMUND YTREHORN and CHRISTIAN BROCHMA" Brysting, A. K., Gabrielsen, T. M., S~rlibrlten,O., Ytrehorn, 0. & Brochmann, C. 1996: The Purple Saxifrage, Saxifraga oppositifolia, in Svalbard: two taxa or one? Polar Research 15(2), 93-105. Several studies have demonstrated high levels of genetic (DNA), ecophysiological, ecological, and morphological variation within the species Purple Saxifrage, Saxifraga oppositifolia, in Svalbard. It has recently been proposed that S. oppositifolia is represented by two conspicuously different subspecies in this archipelago: ssp. reptans, a late-flowering, prostrate ecotype of snow-protected, damp habitats, and ssp. pulvinata, an early-flowering, cushion-like ecotype of dry, wind-exposed heaths and ridges. It has also been suggested that the subspecies may be differentiated at the tetraploid and diploid levels, respectively, which would promote reproductive isolation. These hypotheses are tested by examining variation in morphology, ecology, and pollen size and stainability in 150 plants of S. oppositifolia growing in 50 vegetation samples at four sites in the Kongsfjorden area. Although analyses of the various data sets demonstrated the large variation within the species, the material could not be separated into distinct groups. The morphological variation was continuous along local ecological gradients. The pollen grains were fully stainable and the pollen diameter data showed a unimodal distribution, suggesting that the plants analysed represent only one ploidal level. These results reject a hypothesis that the morphologically intermediate plants are hybrids between two taxa at different ploidal levels. Thus, the conspicuous variation in S. oppositifolia in Svalbard probably results from local, in situ ecoclinal differentiation. Although this variation clearly is wifhout taxonomic significance, it is important in the broader context of arctic conservation biology and the potential impact of global warming on arctic vegetation. Anne K. Brysting, rove M. Gabrielsen, Oddmund Ytrehorn and Christian Brochmann, Botanical Garden and Museum, University of Oslo, Trondheimsveien 238, N-0562 Oslo, Norway; Ove S@rlibriten, Department of Biology, Division of Zoology, University of Oslo, P.O. Box 1050 Blindern, N-0316 Oslo, Norway. Introduction evolution is probably associated with the turbulent climatic history of arctic areas, with repeated It has been shown recently that, although fewer in glaciations/deglaciations, massive plant migra- number than in more southern areas, many arctic tions, and frequent hybridisation. In diploid arctic plants possess extraordinary high levels of varia- species, high levels of variation may be caused by tion. High levels of intraspecific variation in arctic differential selection along gradients in snow plants have been documented in their ecological cover and duration of the growth season. Such traits (McGraw & Antonovics 1983; McGraw factors vary considerably over short distances in 1995a, b), ecophysiology and morphology (Craw- the treeless arctic and alpine areas, and this is ford et al. 1993), and molecular genetics (Broch- probably the main reason for the ecotypic mann et al. 1992a, b, c; Haraldsen & Wesenberg differentiation observed in the arctic diploid 1993; Nordal & Iversen 1993; Gabrielsen et al. Dryas octopetala (McGraw & Antonovics 1983; 1995; Hagen et al. 1995; Tollefsrud et al. 1995; McGraw 1995b). Bauert 1996; Brysting et al. in press). For The high level of intraspecific variation in polyploid species, which constitute the majority arctic plants has important implications for arctic of arctic floras, use of a wide species concept is conservation biology. It is widely accepted that often necessary because many such species are genetic variation and ecotypic differentiation is composed of races of different hybrid origins that necessary for the long-term persistence of a bridge the morphological discontinuities origin- species, which is of particular relevance today ally found between their diploid progenitors with regard to potential global warming (Craw- (Brochmann 1992b, 1995a, b). This type of ford et al. 1993; McGraw 1995b). It is paradoxi- 94 A. K. Brysting et al. cal, however, that traditional conservation biology Recently, there has been particular focus on the focuses on taxon numbers and rare taxa rather variation within S. oppositifolia in Svalbard. This than on the genetic diversity occurring within focus has been accentuated by the publication of a widely defined, widespread taxa, which probably colo;r photograph of a cushion-formed plant from is a better approach to conserving arctic bio- Ny Alesund on the front cover of the journal diversity. Molecular Ecology (Abbott et al. 1995), and by The Purple Saxifrage, Saxifraga oppositifolia the recognition of two ecologically and morpho- L., is a variable species with wide circumpolar logically divergent subspecies, one cushion- distribution, extending also into the higher formed and one prostrate, in the new edition of mountains southwards in Europe, Asia, and North Rbnning’s Svalbards flora (Rbnning 1996). This America (Hultbn & Fries 1986). It is a long-lived variation in growth form within S. oppositifolia in perennial with opposite, small leaves and con- Svalbard was already reported by Anderson & spicuous, pink or purple flowers appearing soon Hesselman (1900). Recent studies have demon- after snow-melt. Although this species, in likeness strated that there not only is large morphological with most other arctic species, is self-compatible, and ecological variation in S. oppositifolia in the it strongly depends on insect pollination for seed archipelago, but that the species also shows large set (Stenstrom & Molau 1992). The plants grow in variation in DNA markers (Abbott et al. 1995; a wide range of habitats in the Arctic, including Gabrielsen et al. 1995, Gabrielsen 1996) and moist snow beds, heaths, dry ridges, and barren, ecophysiological traits (Crawford et al. 1993, recently unglaciated areas. 1994; Crawford & Abbott 1994). The broad morphological variation within the The different growth forms observed in S. species has been noted by many scientists, and oppositifolia in Svalbard have also been suggested to be correlated with other characteristics (Craw- various attempts have been made to split it into ford et al. 1993; Rbnning 1996; Elven & Elvebakk several taxa (see e.g., Hayek 1905). The most 1996). The prostrate plants grow in snow- recent taxonomic treatment of the intraspecific protected, damp habitats and have high photo- variation in is that of Webb & S. oppositifolia synthetic rates, distant leaves with marginal cilia, Gornall (1989), who closely follow Engler & and open flowers with narrow, non-overlapping Irmscher (1919). Webb & Gornall (1989) recog- petals, whereas the cushion-formed plants grow nise one widely distributed and morphologically on dry, wind-exposed ridges and have low variable subspecies, ssp. oppositifolia (the only photosynthetic rates, dense, overlapping leaves subspecies they recognise in the European Arctic), entirely or mostly without cilia, and urn-shaped four subspecies endemic to the central and flowers with broad, overlapping petals. southern European mountains, and two additional In the new edition of Rbnning’s Svalbards flora subspecies occurring in the North American (R~nning1996), the prostrate plants are referred Arctic (and probably across the Beringia into to as S. oppositifolia ssp. reptans (Anderss. & eastern Asia (Hultbn & Fries 1986). Hesselm.) Rbnning, and the cushion-formed The species also varies in chromosome number. plants are referred to as S. oppositifolia ssp. Most populations investigated are diploid pulvinata (Anderss. & Hesselm.) Rbnning. (2n = 26), but several tetraploid (2n = 52) popula- Although only tetraploid plants of S. oppositifolia tions have been reported from arctic areas in have been recorded in Svalbard, it has been Europe, Asia, and North America (Hedberg 1967; suggested that both diploid and tetraploid plants Johnson & Packer 1968; Lave & Liive 1975). are present and that the difference in ploidal level Two chromosome counts, both tetraploid, have may be correlated with growth form (Crawford et been published for S. oppositifolia in Svalbard, al. 1993; cf. also Liive & Eve 1951, 1975). one from the @fjorden area (Flovik 1940) and one The aim of the present study was to determine from the Ny Alesund area (Chapman 1995). All if it is reasonable to divide S. oppositifolia in studies from the rest of northern Europe have Svalbard into two ecologically, morphologically, reported a diploid number (e.g., Skovsted 1934; and possibly also chromosomally, distinct taxa, or Eve & Lave 1956). To what extent the if the variation merely is ecoclinal, representing chromosomal variation in S. oppositifolia is continuous morphological variation along local correlated with the morphological differentiation ecological gradients. Four different data sets were of the species is unknown. produced to answer this question: (1) Ecological The Purple Saxifrage 95 parameters were measured or estimated to detect occurred, and so that at least one plant of S. possible habitat differences correlated with oppositifolia occurred within each sample. growth form differences; (2) macro-morphologi- In each sample, the percentage cover of each cal characters were measured in field-collected vascular plant species (nomenclature
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