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The Effects of Hybridization on the Small-Scale Variation in Seed-Bank Composition of a Rare Plant Species, Erica Ciliaris L

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The Effects of Hybridization on the Small-Scale Variation in Seed-Bank Composition of a Rare Plant Species, Erica Ciliaris L Seed Science Research (2007) 17, 201–210 DOI: 10.1017/S0960258507782855 The effects of hybridization on the small-scale variation in seed-bank composition of a rare plant species, Erica ciliaris L. R.J. Rose* NERC Centre for Ecology and Hydrology, Winfrith Technology Centre, Dorchester, Dorset, DT2 8ZD UK Abstract evolution (Arnold, 1997). Studies have looked at the barriers to gene flow (Grant, 1949; Bouck et al., 2005) or The size and composition of the seed bank of a rare the composition of the resultant vegetation as a result species (Erica ciliaris L.) was analysed. E. ciliaris of gene flow, e.g. the production of hybrid swarms hybridizes with a common relative in southern England (Arnold, 1994). Others have described potential rare (Erica tetralix). The seed banks of these co-occurring plant extinction due to demographic swamping and species were measured at a number of sites with a genetic assimilation by a common congener (Levin range of vegetation types and different management et al., 1996), or predicted species displacement (Wolf histories. Additional sets of samples were taken from et al., 2001). forestry plantations on former heathland sites, where Seed-bank studies in natural or semi-natural these species were known to occur. Relatively few vegetation types are usually related to their use in hybrid seedlings were found in any of the seed-bank habitat re-creation and restoration (Mitchell et al., samples, even though their vegetative abundance 1998; Pywell et al., 2002; Bossuyt and Hermy, 2003; within the sampling areas was equal to that of the pure Allison and Ausden, 2006), or for the conservation of plants. However, the abundance in the vegetation of rare species (Fischer and Matthies, 1998). The relative the two pure species was reflected in the seed-bank fitness of seeds and seedling genotypes of known size on each of the vegetation types, both with and provenance from a hybrid zone has been investigated without burning management. The seed banks from experimentally by Johnston et al. (2003). However, the forestry plantations show that the numbers of there are no studies that examine the effects of seeds of both E. tetralix and the hybrid were depleted, hybridization on the species composition of naturally but that the seed bank of E. ciliaris was not occurring soil seed banks. This study examines the significantly different from that of open heathland seed banks. The long-lived nature of the seed bank small-scale variation in the seed-bank structure of a indicates that there are opportunities for habitat dwarf shrub, heathland vegetation type that is restoration on former heathland sites. characterized by the abundance of two closely related Erica species and their hybrids. Keywords: conservation, disturbance, Erica ciliaris, Erica tetralix and Erica ciliaris, the parent species, Erica tetralix, extinction, forestry, heathland, plant have overlapping geographical distributions. E. ciliaris hybridization, restoration, seed bank is a rare plant in the British Isles (Wigginton, 1999), where it is at the northern limit of its European range in south-west England (Rose et al., 1996). The hybrid Erica £ watsonii is a naturally occurring cross between Erica ciliaris and Erica tetralix. The breeding system of Introduction E. ciliaris and E. tetralix creates an F1 generation of hybrids with low pollen viability and high seed There have been many studies of naturally occurring sterility (Gay, 1957). The hybrid is found in all areas hybrid plant populations. These have concentrated and habitats where the two parent species coexist in mostly on the mechanisms involved in the hybridiz- continental Europe (Neyraut, 1900) and Britain ation process and the role of hybridization in (Perring and Sell, 1968; Stace, 1975). In south-west England, both species and the hybrid co-occur on a series of acidic soils that range from dry, free-draining *Correspondence mineral soil to waterlogged, organic peat soil. These Fax: þ44 1305 213600 soils support heath and mire vegetation, which Email: [email protected] corresponds to the H3 (Ulex minor–Agrostis curtisii 202 R.J. Rose heath), M16 (Ericetum tetralicis wet heath) and M21 seeds of E. cinerea has been measured (Bullock and (Narthecio–Sphagnetum valley mire) plant commu- Clarke, 2000). Their results showed that, although nities of Rodwell (1991). The plants of both species are long-range dispersal and colonization of new sites is most abundant on the wetter heaths (M16), where they possible, most of the seeds produced by a heather are characteristically co-dominant with mixtures of plant fall within a radius of 1 m of its own centre. This Calluna vulgaris, the grass Molinia caerulea and the bog species of heather has a similar growth form and seed mosses Sphagnum compactum and S. tennellum. The size to E. ciliaris and E. tetralix, (see Bannister, 1965, flowering periods for each species show similar 1966; Rose et al., 1996). Thus, both E. ciliaris and patterns. The first flowers of E. tetralix open in May, E. tetralix have no specific seed-dispersal mechanisms, and those of E. ciliaris appear in June. The peak and are likely to exhibit locally variable seed-bank flowering is in early July for E. tetralix and early seed densities that are dependent on the above- August for E. ciliaris. Flowering continues into ground vegetation composition. September for E. tetralix and late October for E. ciliaris. On the Dorset heaths, there are patches of Hybrid plants can be found flowering throughout the heathland vegetation with differing proportions flowering period of the two parent species, and they of these two Erica species and the hybrid on a range have an intermediate flowering peak. The majority of of different vegetation types, based on the local flowers of both species and the hybrid are produced hydrological conditions. Heathland vegetation con- between July and September. sists primarily of dwarf shrubs interspersed with The species composition of heathland soil seed grasses, forming a dense vegetation canopy. In mature banks does not always reflect the species composition vegetation stands, there is also a ground layer of of the above-ground vegetation (Pakeman and bryophytes or litter, consisting of dead plant material. Marshall, 1997; Valbuena and Trabaud, 2001). This Under these conditions, seed germination and seed- can be due to a number of biological and environ- ling establishment are rare, and gaps for seedling mental factors, such as: seed production, seed long- growth tend to occur only following disturbance evity, predation, soil conditions and vegetation events, such as a fire, physical disturbance of the soil management. Seed-bank composition can be an by animals or by anthropogenic activities, such as important factor in the species composition of afforestation. Historical information on pre-afforesta- recovering vegetation following extreme events, such tion and pre-fire vegetation and hybrid composition as fire and drought (Valbuena and Trabaud, 2001; enabled this study to compare the relative compo- Odion and Tyler, 2002) or soil disturbance (Britton sition of the Erica spp. in the seed bank with that of the et al., 2000). Many plants of the Ericaceae, especially the existing vegetation types, and to see if this varies with Erica spp., produce seeds that can persist in the soil for soil conditions and site history. The results of this many years, building up a considerable and long- work will help us to understand the current lived seed bank (type IV seed bank of Thompson and distribution of the different genotypes and to predict Grime, 1979; long-term persistent in Thompson et al., the possible changes that may result from different 1997). It is also important to consider the residual seed forms of management. bank when making decisions on the suitability of sites for habitat restoration (Walker et al., 2004a, b). Soil moisture and groundwater depths can have an Materials and methods effect on soil seed-bank survival. By raising the groundwater from 30 cm below the surface to 5 cm The species and study sites below the surface for 3 years, significant differences in both species composition and individual species E. ciliaris and E. tetralix, the two parent species, are numbers in a wet meadow seed bank have been closely related (to sub-genus). They are similar in reported by Bekker et al. (1998). The soil conditions in the growth form, and where they co-occur, they are three vegetation communities in this study varied from partially co-flowering, with the E. tetralix flowering aerobic throughout most of the year on the relatively season peaking in early July and that of E. ciliaris free-draining humid heathland (H3) sites, to seasonally peaking in August. The corollas are of similar shape: inundated wet heathland (M16) and to permanently E. ciliaris tubular–urceolate and E. tetralix ovoid– anaerobic, waterlogged soils of the M21 mire. urceolate; length: E. ciliaris 8–10 mm and E. tetralix In heterogeneous vegetation types, such as those 6–8 mm; and colour: E. ciliaris dark rose and E. tetralix, where there is an intimate mix of both parent plants pink. They share the same floral visitors (in particular and their hybrids, the method of seed dispersal can bumblebees) and are likely to share the same have an effect on the small-scale distribution of seeds mechanisms of cross-pollination (Bannister, 1966; within a seed bank. Local variations in seed densities Rose et al., 1996). related to dispersal have been demonstrated by Eshel This study was carried out on the Purbeck Heaths et al. (2000) for Pinus halepensis. The dispersal of the in Dorset, south-west England. This area contains the Effects of hybridization on seed-bank composition of Erica ciliaris 203 majority of the remaining British sites for E. ciliaris. with a continual seed input for more than 35 years; Within this area, much of its former range has been and disturbed heathland where the seed bank had lost to conifer plantations and some to agricultural been removed or severely reduced by burning 20 conversion.
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