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Breeding System, Insect Flower Visitation, and Floral Traits of Two Alpine Cerastium Species in Norway

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Breeding System, Insect Flower Visitation, and Floral Traits of Two Alpine Cerastium Species in Norway Breeding System, Insect Flower Visitation, and Floral Traits of Two Alpine Cerastium Species in Norway Ørjan Totland; Björn Schulte-Herbrüggen Arctic, Antarctic, and Alpine Research, Vol. 35, No. 2. (May, 2003), pp. 242-247. Stable URL: http://links.jstor.org/sici?sici=1523-0430%28200305%2935%3A2%3C242%3ABSIFVA%3E2.0.CO%3B2-8 Arctic, Antarctic, and Alpine Research is currently published by The Regents of the University of Colorado, a body corporate, contracting on behalf of the University of Colorado at Boulder for the benefit of INSTAAR. Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/about/terms.html. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/journals/regentsINSTAAR.html. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. The JSTOR Archive is a trusted digital repository providing for long-term preservation and access to leading academic journals and scholarly literature from around the world. The Archive is supported by libraries, scholarly societies, publishers, and foundations. It is an initiative of JSTOR, a not-for-profit organization with a mission to help the scholarly community take advantage of advances in technology. For more information regarding JSTOR, please contact [email protected]. http://www.jstor.org Fri Jan 25 09:17:32 2008 Arctic, Antarctic, and Alpine Research, Vol. 35, No. 2, 2003, pp. 242-247 Breeding System, Insect Flower Visitation, and Floral Traits of Two Alpine Cerustium Species in Norway (arjan Totland Abstract Department of Biology and Nature Many alpinelarctic flowering plant species have presumably evolved the ability to self- Conservation, The Agricultural University pollinate as a reproductive assurance mechanism under harsh abiotic environmental of Norway, P.O. Box 5014, N-1432 As, conditions that restrict insect flower visitation. We compared self-pollination, pollen Norway. [email protected] limitation, insect flower visitation, and dichogamy of low- and high-elevation populations of Cerastium alpinum, a species in established alpine communities, and Cerasrium cerasroides, a pioneer species in disturbed habitats. Cerastium alpinum has large showy Bjorn Schulte-Herbriiggen flowers, while C. cerastoides has smaller and paler flowers. The temporal separation of Institute of Ecology and Resource pollen release and stigma receptivity within a flower (dichogamy) was smallest in C. Management, The University of cerasroides, which was also more highly self-compatible. No pollen limitation on seed set Edinburgh, King's Building, Edinburgh occurred in any species, possibly due to their high selfing ability. Despite a substantially EH9 3JU. U.K. higher pollinator visitation to C. alpinum compared to C. cerastoides, the latter had the higher seed set. Pollen limitation, autogamy, and pollinator visitation did not differ between altitudes for either species. Differences in habitat and flower size, color, and development between the species are consistent with their different selfing ability. Introduction pollination is an adaptation to insufficient pollinator service, there should be differences in the ability to self-pollinate between species The evolution of self-pollination (autogamy) and the ability to or conspecific populations experiencing different pollinator visitation self-fertilize (self-compatibility) are proposed to be adaptations that rates. However, few comparative studies at the species or population insure reproductive success under conditions of chronically low or level on the relationships between pollinator visitation rates, pollen temporarily unreliable cross-pollination (the reproductive assurance limitation, and selfing ability have been conducted to explicitly test this hypothesis) (Hagerup, 1951; Baker, 1955; Lloyd, 1992; Lloyd and hypothesis. Schoen, 1992). Insufficient cross-pollination appears to be common in Here we examine the relationships between flower visitation rates flowering plants. In a literature review, Burd (1994) found that female by insects, selfing ability, pollen limitation and dichogamy in two reproductive success was limited by low pollen availability in 62% of alpine Cerastium species that differ in flower size, flower coloration, species or populations examined, in at least some years. In another and habitat. Cerastium alpinum has large and bright white flowers and analysis, Larson and Barrett (2000) found that pollen limitation was grows in dry meadows and rock outcrops, while C. cerastoides have less intense in self-compatible and autogamous species, compared to smaller and paler flowers, and grows in late-melting wet snowbeds. On obligate outcrossing species. This finding supports the reproductive the basis of these differences between the species, we predicted that assurance hypothesis. C. cerastoides should be visited less frequently by insects, possess Low temperatures and short snow-free periods, which constrain decreased dichogamy, be less pollen limited, and be more autogamous the abundance and flight activity of insect pollinators (McCall and compared to C. alpinum. Specifically we ask: (1) if pollen limitation, Primack, 1992; Totland, 1994), characterize arctic and alpine habitats. autogamy, and selfing ability differs between the species, (2) if pol- Thus, a relatively high proportion of arctic and alpine flowering plant linator visitation rates differ between the species and between low- and species may have evolved autogamy and self-compatibility in response high-altitude populations of each species, and (3) if dichogamy differs to poor cross-pollination opportunities. In an extensive review, Molau between the species. (1993) suggests that self-compatibility is very widespread among tundra plants. He also found that species growing in late-thawing habitats, such as snow beds, were more self-compatible than species in Study Species earlier emerging habitats. Self-compatibility should be accompanied by evolutionary adjustments of morphological or phenological traits of Cerastium alpinum L. is an insect-pollinated, perennial mat- flowers that enhance self-pollination (Lloyd and Schoen, 1992). For forming herb. The hermaphroditic, five-petalled, actinomorphic example, the physical distance between stigma and anthers (herkog- flowers are bright white, and are ca. 20 mm in diameter (Clapham amy) is usually lower in highly autogamous phenotypes or taxa than et al., 1987). Flowers are visited by several species of Muscidae, in predominately outcrossing ones (Lloyd and Schoen, 1992; Take- Anthomyiidae, and Empididae (Totland, 1993). Flowering is initiated bayashi and Delph, 2000). Moreover. the temporal separation between relatively late after snow melt, around mid-July (Totland, 1993). It anther pollen release and stigma receptivity (dichogamy) should also grows in a range of habitats, but it is most common in dry, south-facing decrease with increasing autogamy (Bertin and Newman, 1993). How- slopes on rock outcrops or in dry meadow areas with a stable ever, to our knowledge few studies have actually documented a rela- vegetation cover where snow melts early in the season. There is a large tionship between dichogamy and selfing rate at the among-population amount of intraspecific morphological variation within the C. alpinum or among-species levels. Such studies could show that selection has complex (Grundt et al., 2000). Based on habitat type and hairiness, the favored traits that enhance self-pollination ability. If autogamous plants in this study probably belong to C. alpinum spp. alpinunl. 0 2003 Regents of the University of Colorado 1523-0430103 $7.00 Cerastium cerastoides (L.)Britton is a perennial herb with many- entered bags at the low-altitude C. alpinum population. These were branched, creeping stems. The hermaphroditic, five-petalled actino- removed immediately after being discovered, but the flowers were morphic flowers are dull white, and are ca. 10 mm in diameter included in the statistical analyses. (Clapman et a]., 1987). Flowers are visited by the same insects as At the low altitude, we initiated the experiment on 11 and 12 July C. alpinun7 (0.Totland, personal observation). Flowering is initiated on C. alpinunl and C. cerastoides, respectively. The early-flowering relatively late in the season, around late July to early August. group of C. alpinun7 at high altitude was initiated on 12 and 13 July, Cerastium cerastoides typically grows in moist snowbeds with and the late-flowering group at 20 and 21 July. The early-flowering a discontinuous vegetation cover. The two species have very similar group of C. cemstoides at high altitude was initiated on 22 and 23 July, distribution patterns in Europe (Jalas and Suominen, 1983). The two and the late-flowering group at 10 August. To ensure counting the species have nearly identical number of ovules per flower (C. alpinum: complete seed number, we collected capsules for each population and mean =45.7 ovules/flower, SD = 5.8, N =71; C. rerastoides: mean = period on the same day (ca. three weeks after flowering), irrespectively 45.5, SD
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