Knowledge of the manner in which genetic variation within a tree species affects associated communities and ecosystem processes across its entire range is important for understanding how geographic mosaics of genetic interactions might develop and support different communities. While numerous studies have investigated the community and ecosystem consequences of genetic variation at the hybrid cross type or genotype level within a species, none has investigated the community-level effects of intraspecific genetic variation across the geographic range of a widespread species. This is the scale at which geographic mosaics of coevolution are hypothesized to exist. Studies at this level are particularly important for foundation tree species, which typically support numerous microbial, fungal, plant, and animal communities. We studied genetic variation across eight geographical races of the forest tree Eucalyptus globulus representing its natural distribution across southeastern Australia. The study was conducted in a 15-year-old common garden trial based on families derived from single-tree open-pollinated seed collections from the wild. Neutral molecular genetic variation within E. globulus was also assessed and compared with genetic divergence in the phenotypic and community traits. Three major findings emerged. First, we found significant genetically based, hierarchical variation in associated communities corresponding to geographical races of E. globulus and families within races. Second, divergence in foliar communities at the racial level was associated with genetically based divergence in specific leaf morphological and chemical traits that have known defensive functions. Third, significant positive correlations between canopy community dissimilarity and both neutral molecular genetic and leaf quantitative genetic dissimilarity at the race level supported a genetic similarity rule. Our results argue that genetic variation within foundation tree species has the potential to be a significant driver of the geographical mosaics of variation typical of forest communities, which could have important ecological and evolutionary implications.
The contamination of native-eucalypt gene pools via exotic pollen is of concern as (i) pollen dispersal is believed to be much more widespread than seed dispersal, (ii) reproductive barriers are often weak between closely related species, (iii) European settlement has already had a major impact on Australia's eucalypt woodlands and mallee, (iv) there has been a rapid expansion of eucalypt plantations and restoration plantings in Australia and (v) Australia is the custodian of an internationally important genetic resource. Pollen flow between plantation and native eucalypt species has already been reported and implementation of strategies to minimise the risk and consequences of genetic pollution is important if Australian forestry is to be considered sustainable. The risks associated with the introduction of non-native species, provenances and hybrids include direct effects on the gene pool through genetic pollution as well as indirect effects on dependent biodiversity. In many cases, the risk of genetic pollution will be small due to strong barriers to hybridisation between distantly related species, differences in flowering time or poor fitness of hybrids. There is no risk of hybridisation between species from the different major eucalypt genera and/or subgenera (e.g. symphyomyrts, monocalypts, eudesmids, bloodwoods and angophora). The main plantation species are symphyomyrts and within this subgenus, the probability of successful hybridisation generally decreases with increasing taxonomic distance between species. The planting of non-local provenances or improved material within the range of native populations has the potential to have an impact on local gene pools to varying degrees, indicating the requirement for the adoption of management strategies to reduce this risk. Naturally small or remnant populations are at particular risk. A framework for assessment of the risk of genetic pollution is developed herein.
While habitat fragmentation is a central issue in forest conservation studies in the face of broad-scale anthropogenic changes to the environment, its effects on contemporary mating patterns remain controversial. This is partly because of the inherent variation in mating patterns which may exist within species and the fact that few studies have replication at the landscape level. To study the effect of forest fragmentation on contemporary mating patterns, including effective pollen dispersal, we compared four native populations of the Australian forest tree, Eucalyptus globulus. We used six microsatellite markers to genotype 1289 open-pollinated offspring from paired fragmented and continuous populations on the island of Tasmania and in Victoria on mainland Australia. The mating patterns in the two continuous populations were similar, despite large differences in population density. In contrast, the two fragmented populations were variable and idiosyncratic in their mating patterns, particularly in their pollen dispersal kernels. The continuous populations showed relatively high outcrossing rates (86-89%) and low correlated paternity (0.03-0.06) compared with the fragmented populations (65-79% and 0.12-0.20 respectively). A greater proportion of trees contributed to reproduction in the fragmented (de/d>or= 0.5) compared with the continuous populations (de/d = 0.03-0.04). Despite significant inbreeding in the offspring of the fragmented populations, there was little evidence of loss of genetic diversity. It is argued that enhanced medium- and long-distance dispersal in fragmented landscapes may act to partly buffer the remnant populations from the negative effects of inbreeding and drift.
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