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Between-Year Variation in Seed Weights Across Altitudes in the High-Alpine Plant Eritrichium Nanum

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Between-Year Variation in Seed Weights Across Altitudes in the High-Alpine Plant Eritrichium Nanum Plant Ecol (2010) 207:227–231 DOI 10.1007/s11258-009-9667-3 Between-year variation in seed weights across altitudes in the high-alpine plant Eritrichium nanum Lea R. Wirth Æ Rene´ Graf Æ Felix Gugerli Æ Urs Landergott Æ Rolf Holderegger Received: 6 May 2009 / Accepted: 9 September 2009 / Published online: 16 September 2009 Ó Springer Science+Business Media B.V. 2009 Abstract Seed weight is a prominent life history Introduction trait of plants affecting dispersal, establishment, and survival. In alpine environments, the few studies When comparing lowland with alpine plant species, investigating the effect of elevation on seed weight an obvious adaptation to the adverse alpine environ- within species have mainly detected a decrease in seed ment is the reduced size of alpine plants (Jenny-Lips weight with increasing elevation. This relationship is 1948;Ko¨rner 2003). In contrast, it has been observed generally attributed to the adverse climate at high that seed weight or seed size is often increased in elevations. In order to test this hypothesis, we alpine species as compared with taxonomically analyzed seed weight variation across altitudes related lowland species (Blionis and Vokou 2005; (2,435–3,055 m a.s.l.) in two consecutive years that Pluess et al. 2005; but see Baker 1972). Seed size is differed in weather conditions in the high-alpine of special interest, because it is a prominent life cushion plant Eritrichium nanum. We found a signif- history trait of plants that affects dispersal, establish- icant reduction in seed weight with increasing eleva- ment, and seedling survival (Leishman et al. 2000). tion in both years, but in the growing season with Multiple comparisons of species pairs from the Swiss more adverse weather conditions, the reduction was Alps showed heavier seeds in alpine than in lowland more substantial than in the more favorable year. We species (Landolt 1967; Pluess et al. 2005). The same conclude that alpine plants may be able to produce relationship was observed along an altitudinal gradi- well-developed seeds at low elevations in almost all ent at Mt. Olympos in Greece (Blionis and Vokou years, independent of weather conditions, whereas 2002, 2005). A possible explanation for these find- reproduction through seeds is potentially limited to ings is that embryos from heavier seeds are better years of favorable weather at high elevation. provisioned than those from lighter seeds. Under the increasingly adverse ecological conditions along Keywords Seed mass Á Seed size Á altitudinal gradients, such as shorter growing seasons Elevation Á Weather variation Á Alps and lower mean temperatures (Landolt 1967;Ko¨rner 2003), seedlings from heavy seeds might be more successful in establishing and surviving (Leishman et al. 2000; Moles and Westoby 2004) This fitness L. R. Wirth (&) Á R. Graf Á F. Gugerli Á advantage of heavy seeds could have led to the U. Landergott Á R. Holderegger evolution of heavy-seeded species at high elevations. WSL Swiss Federal Research Institute, Zu¨rcherstrasse 111, CH - 8903 Birmensdorf, Switzerland In contrast to the above mentioned species-pair e-mail: [email protected] comparisons, the few studies investigating variation 123 228 Plant Ecol (2010) 207:227–231 in seed weight within single alpine species have Fruits contain four nutlets, each with a single seed mostly reported a reduction in seed weight with (Gams 1975). increasing elevation. For instance, Eritrichium We selected 15 study sites at different elevations nanum, Saxifraga oppositifolia, Scabiosa lucida, in the Upper Engadine and adjacent valleys of and Ranunculus acris produced lighter seeds with southeastern Switzerland (Table 1). The sites were increasing elevation, while Epilobium fleischeri situated between 2,435 and 3,055 m a.s.l. and sepa- showed no directional change in seed weight (Totland rated from one another by at least 1 km. All the sites and Birks 1996; Totland 1997a; Pluess et al. 2005; consisted of large, preferably flat scree areas with Zoller et al. 2005). These results have been attributed naturally sparse vegetation on windswept crests, to low temperatures and short growing seasons at where vegetation becomes snow-free early in the high elevations, the two most important factors season. affecting seed maturation and, thus, seed weight All the 15 sites were sampled twice, once in 2005 (Totland and Birks 1996; Totland 1997a, b; Wagner and once in 2006. The growing seasons in these and Reichegger 1997; Baskin and Baskin 1998; 2 years differed in their weather, especially in July Blionis and Vokou 2005). (Table 2), the main period of seed maturation. July If seed weight of alpine plants is mostly influenced 2005 was cold and partly snowy, whereas July 2006 by environmental constraints, we should not only find was substantially warmer. In contrast, the weather in decreasing seed weight with increasing elevation, but spring and late summer was more favorable in 2005 also find variation in seed weights owing to annual than in 2006. variation in weather conditions. Annual variation in From July to August, fully developed ripe seeds seed germinability and seed production has been from 10 randomly selected cushions were sampled found in long-term investigations of alpine and arctic per site, just before they would have been dispersed plant species (Laine et al. 1995; Zoller et al. 2005). (Table 1). After drying at room temperature, seeds To our knowledge, however, no investigation has were stored at 2°C for five months. Ten filled seeds addressed seed weight variation in a high-alpine plant (i.e., nutlets without coat) were randomly chosen per species along an altitudinal range, and in seasons with cushion and weighed on an electronic analytical different weather conditions. Here, we report on variation in seed weight across Table 1 Characteristics, locations, and sampling dates in 2005 altitudes from 2,435 to 3,055 m a.s.l. in the high- and 2006 for 15 E. nanum sites in southeastern Switzerland, alpine cushion plant E. nanum in two consecutive, but where seeds were collected climatically different growing seasons in the Swiss Site Elevation Longitude/latitude Sampling date Alps. We addressed the following two questions: (1) (m a.s.l.) (E/N) Does seed weight in E. nanum decrease with 2005 2006 increasing altitude? (2) Is annual climatic variation 1 2,435 10°0203800/46°2305600 12/08 25/07 affecting seed weight across altitudes? 2 2,460 9°3901700/46°2502000 13/08 27/07 3 2,600 9°4702000/46°2405700 16/08 29/07 4 2,655 9°3905300/46°2405600 18/08 27/07 Materials and methods 5 2,700 9°4703300/46°2404100 16/08 29/07 6 2,735 9°3604500/46°2502300 24/08 27/07 Eritrichium nanum (L.) Gaud. (Boraginaceae) is a 7 2,790 9°4402600/46°2502000 24/08 27/07 showy, blue-flowered alpine cushion plant. This 8 2,820 9°4401000/46°2904000 27/08 30/07 perennial multi-flowered herb grows in high-alpine 9 2,840 9°3501500/46°2705500 25/08 31/07 habitats of Central Europe, mostly between 2,500 and 10 2,875 9°4405600/46°3009000 27/08 09/08 3,000 m a.s.l. (Schro¨ter, 1926). The flowering season 11 2,905 9°4405700/46°2905900 27/08 09/08 starts in June at low elevations and continues until the 12 2,925 9°4603800/46°3001900 29/08 14/08 end of July at high elevations. Flowers are pollinated 13 2,960 9°4201700/46°3001800 27/08 09/08 by various generalist insects, mostly Diptera (Zoller 14 3,020 9°4309000/46°3001400 27/08 09/08 et al. 2002). Zoller et al. (2002) reported the flowers 15 3,055 9°4707900/46°3005000 29/08 14/08 to be self-compatible, but predominantly outcrossed. 123 Plant Ecol (2010) 207:227–231 229 Table 2 Weather conditions (thawing degree days i.e., sum of degrees over 0°C per day, mean temperature, and days with snowfall in July and August) during the growing seasons of E. nanum of 2005 and 2006 Thawing degree daysa Mean temperature (°C)a Number of days with snowfallb 2005 2006 2005 2006 2005 2006 May 21.9 2 -3.1 -3.6 – – June 73.5 77.9 1 0.3 – – July 79.1 154.5 2 5 7 0 August 36.8 13.8 0.3 -1.4 7 6 a Data from MeteoSwiss station Piz Corvatsch (3,315 m a.s.l; Engadine, Switzerland) b Personal observations at study sites balance (Mettler Toledo MX5, Greifensee). We then 0.7 calculated mean seed weight per site per year, as well 0.6 as the standard errors. 0.5 The effect of elevation on mean seed weight was tested using ordinary least squares (OLS) regressions 0.4 with permutation testing (N = 10,000) implemented 0.3 in the software BLOSSOM (Cade and Richards 0.2 2005). By analyzing two single regressions (sepa- Mean seed weight [mg] 0.1 rately for each year), we explored the effect of 0 elevation on mean seed weight. In order to assess 2400 2500 2600 2700 2800 2900 3000 3100 whether the effect of elevation differed between the Elevation [m a.s.l.] two study years, we investigated the difference in Fig. 1 Relationship between seed weight in E. nanum (mean per slope of the above regressions by testing for a site and year ± S.E.) and elevation. Black circles and black line year 9 elevation interaction. indicate values and linear regression (y =-0.0002x ? 1.0174), respectively, for 2005, and grey squares and grey line indicate those for 2006 (y =-0.0006x ? 1.9701) Results and discussion of the other studies assessing the relationship between Average seed weight in E.
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