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Of Saxifraga Oppositifolia (Saxifragaceae)

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Of Saxifraga Oppositifolia (Saxifragaceae) Oecologia (1998) 114:60±66 Ó Springer-Verlag 1998 Felix Gugerli Effect of elevation on sexual reproduction in alpine populations of Saxifraga oppositifolia (Saxifragaceae) Received: 28 April 1997 / Accepted: 20 October 1997 Abstract Self-compatibility in high arctic and alpine necessarily select for increased sel®ng rates in a prefer- areas is regarded as an adaptation to low pollinator entially outcrossing species like S. oppositifolia. abundance. However, high genetic variability as a con- sequence of outcrossing is, with regard to population Key words Alpine plants á Elevation á Breeding system á persistence, favorable in highly stochastic environments Reproduction á Self-pollination such as tundra habitats. To evaluate these contradictory scenarios, I performed in situ pollination experiments to examine the breeding system of the predominant out- Introduction crosser Saxifraga oppositifolia in ten populations at two dierent elevations in the Swiss Alps. Pollinator limita- The general view about plant reproduction along the tion was detected at both elevations, but fruit set in macrogradient from temperate to arctic regions predicts naturally pollinated ¯owers was only slightly less at the increasing self-compatibility, apomixis, polyploidy, and higher elevation. Increased pollinator limitation at high vegetative reproduction, all of which tend to promote compared with low elevation thus could not be dem- the genetic uniformity of populations (Mosquin 1966). onstrated in this experiment. Hand-crossings yielded Corresponding changes are described for elevational equal mean proportion seed set at both elevations, and gradients in alpine areas, though knowledge about the so did hand-sel®ngs. This constant pattern of the breeding systems is often derived from phenological breeding system in S. oppositifolia indicates selective rather than experimental observations (SchroÈ ter 1926; factors that lead to the maintenance of a high level of Bliss 1962). The same gradient is thought to be recog- outcrossing even in high-elevation populations. Based nized both at the species and at the community level. on sex allocation models, it was expected that a high Observations in Polygonum viviparum conform to this ovule number should be selectively advantageous in a trend, as plants allocate more resources to vegetative plant-pollinator system where chance visitation or self- bulbils relative to ¯owers with increasing latitude and ing play important roles. However, female reproductive elevation (Bauert 1993, 1994b). However, in a survey at oer in terms of ovule number per ¯ower did not change high-arctic northeastern King Christian Island (North- from low to high elevation. Since neither increased west Territories, Canada), only one-third of the species pollinator limitation nor increased seed set in selfed were found to be able to reproduce vegetatively, and ¯owers was found at high compared with low elevation, only three species appeared reasonably successful in the prerequisites for testing the hypothesis were not dispersing their vegetative organs (Bell and Bliss 1980). given. This study contradicts the hypothesis that inimi- To my knowledge, no comparable data are available for cal environmental factors in alpine or arctic habitats alpine areas. In addition, arctic or alpine species usually reported to reproduce exclusively by vegetative means have been found to produce seeds (e.g., Saxifraga cer- nua, S. foliolosa, and P. viviparum; Molau 1992; Bauert F. Gugerli1 1993). Though seed set is rare in these species, it is suf- Institut fuÈ r Systematische Botanik, UniversitaÈ tZuÈrich, ®cient to result in considerable genetic variability within Zollikerstrasse 107, CH-8008 ZuÈ rich, Switzerland some populations (Bauert 1994a; T.M. Gabrielsen and Present address: C. Brochmann, unpublished data). The same holds true 1EidgenoÈ ssische Forschungsanstalt fuÈ r Wald, Schnee und Landschaft, ZuÈ rcherstrasse 111, for the rhizomatous Carex curvula. This species domi- CH-8903 Birmensdorf, Switzerland nates alpine grasslands on silicate bedrock, but so far fax: ++41-1-739-2215; e-mail: [email protected] has never been observed to establish through seeds. 61 Nevertheless, dierent clones could be delimited by In each of ®ve dierent regions in the central Alps of means of RAPDs (Steinger et al. 1996). These inves- southeastern Switzerland, I examined the breeding sys- tigations demonstrate that reproductive strategies such tem of S. oppositifolia at two elevations diering by 320± as vegetative spreading may also conserve extant genetic 455m. Saxifraga oppositifolia shows great morphological diversity. variation in relation to the wide range of dierent hab- At high elevations and latitudes, reduced pollinator itat types in arctic and alpine environments (Crawford abundance because of inimical abiotic conditions (e.g., et al. 1995; Kaplan 1995), which demonstrates its po- low temperatures; Heinrich 1993), and decreased length tential for local adaptation. It may therefore be expected of the growing season (Bliss 1971; Ellenberg 1988) are that populations of S. oppositifolia are also able to adapt recognized as constraints which promote adaptations for to selective pressure on its breeding system. Saxifraga self-pollination such as self-compatibility or homogamy oppositifolia depends on pollinator visits for seed set in (MuÈ ller 1881; Bliss 1962; Richards 1986). Such a strat- the Arctic as well as in the Alps (Kevan 1972; Ti- egy should ensure fast pollination in the little time khmenev 1984; StenstroÈ m and Molau 1992; Gugerli available for seed development. Despite these expecta- 1997b). Reduced pollinator activity should accordingly tions, S. oppositifolia, the most abundant species at the lead to decreased reproductive output due to low fruit very northern limit of growth of higher plants, which and seed set (see Anderson and Beare 1983). also grows in the upper nival zone in the Alps (Jones and I investigated the eect of elevation on the fruit set of Richards 1956; Bay 1992; Kaplan 1995), is a protogy- naturally pollinated S. oppositifolia, testing the hypoth- nous, entomophilous, predominantly outcrossing, and esis that fruit set is lower in high- than in low-elevation diploid plant (Kevan 1972; Tikhmenev 1984; StenstroÈ m populations. If pollinator activity is reduced at high and Molau 1992). Furthermore, populations of sexually elevations compared with low elevations, this constraint reproducing plant species in arctic and alpine areas, i.e., should select for higher self-compatibility. As a con- S. oppositifolia, Silene acaulis,orPrimula spp., indeed sequence, reduced inbreeding depression should be display high genetic variability, indicating considerable detectable. Current theory suggests that inbreeding outcrossing rates for these species (Miller et al. 1994; depression in outcrossers is mainly expressed in early Abbott et al. 1995; Gabrielsen et al. 1997; own, un- stages of the life cycle such as seed formation (Husband published data). These observations suggest that the and Schemske 1996). I therefore wanted to know various selective constraints may act antagonistically, whether the seed set of self-pollinated compared with and that the general predictions about plant reproduc- cross-pollinated ¯owers of S. oppositifolia increases from tion in arctic and alpine environments need reconsider- low- to high-elevation populations, which would even- ation. tually lead to a shift towards self-pollination in the Winn and Gross (1993) summarize results about the breeding system of S. oppositifolia. eect of elevation or latitude on seed mass or size. They A higher number of ovules ( female reproductive point out that these variables react inconsistently to the oer; Urbanska 1989) relative to a ®xed number of de- environmental change along elevational or latitudinal posited pollen grains should enhance fertilization success, gradients. Empirical investigations into the in¯uence of given that (1) seed set is independent of the degree of increasing elevation or latitude on the breeding systems incompatibility, and (2) more ovules per ¯ower increases of plant species are very scarce. Cruden (1972) found the genetic diversity among ovules (due to meiotic re- that proportion seed set in naturally bee-pollinated combination) and thus the coincidence of female and male Lamiaceae is lower in high-elevation than in mid- gametes compatible for fertilization. An increased sel®ng elevation populations in Mexico. Similarly, seed set in rate also leads to a higher allocation to female relative to Trifolium pallescens and T. thalii decreases from low- to male function, whereas reduced pollen transfer proba- high-elevation populations in the Alps (Hilligardt 1993), bility, e.g., because of low pollinator activity, should but proportion seed set in the annual Gentianella counteract this trend (Brunet and Charlesworth 1995). In caucasea does not dier among alpine to subnival pop- a comparison of two closely related species, Gugerli ulations in the Caucasus (Akhalkatsi and Wagner 1996). (1997b) shows that geitonogamous sel®ng seems to be It is not known in these studies, however, whether seed more important in Saxifraga bi¯ora than in S. oppositi- set resulted from self or cross pollen. Experimental folia. Concordantly, S. bi¯ora has more ovules per fruit pollinations with Pedicularis lapponica indicate a change than does S. oppositifolia. Thus I hypothesized that ovule in the breeding system from low to high elevation in number per fruit is higher at high than
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