Demographic genetics of the American beech, Fagus grandifolia. II. Genet substructure of populations for the Blue Ridge, Piedmont and the Great Smoky Mountains

Populations of American beech in Virginia and the Great Smoky Mountain National Park in Tennessee and North Carolina were investigated for demographic genetic substructurings. Two Virginia populations, one on the Blue Ridge (WG1) and the other on the Piedmont (WG2) occur over an elevational gradient of several hundreds meters. One of the Great Smoky Mountain populations (GS1) was in a 'beech gap' and the other (GS2) in a 'cove forest' along a creek. The populations in the Great Smoky Mountain National Park were only separated by a few hundred meters in elevation, but both on the same physiographic province. The populations had two growth forms. Trees produced extensive root suckers at WG1, GS1 and GS2, but WG2 had no root suckers and all individuals had obviously been established from seeds. A total of 1335 shoots were mapped at the four sites, their size measured [diameter at breast height (DBH) or diameter at ground height (DGH)], and genotypes were determined for each locus using allozyme analysis. F_IS among five different size-classes revealed an excess of homozygotes in WG1, GS1 and GS2, and an excess of heterozygotes in WG2. The offshoot formation from root suckers obviously contributed to the abundance of intermediate size-classes in WG1, GS1 and GS2. Exceedingly localized patchiness of different multilocus genotypes reveals genetic clustering of shoots that have obviously originated from root suckers in WG1, GS1 and GS2. The Piedmont population (WG2), on the other hand, showed loose localization of genetically related trees at a scale of 35–40 m in area, suggesting broader ranges of pollen and seed dispersal. The data are discussed in the light of the differences in growth form and mode of reproduction, and also in relation to the post-glacial migration and the current geographic distribution of the species.     

Local differentiation of Albanian provenances and chorology of beech species in balcan area

Abstract

Some quantitative characters of different Albanian beech provenances are analysed in order to study their local differentiation. The results obtained by discriminant analysis showed the formation of two principal groups. Biza provenance was discriminated from both groups. Principal components and cluster analysis give a good evidence on the similarities among Albanian and Bulgarian beech populations. The results suggest that all area of Balcan is the contact zone of Fagus sylvatica L. and Fagus orientalis Lypski.     

Morphology,Anatomy and Histochemistry of Fagus grandifolia Ehrh. (North American Beech) Ectomycorrhizas

Abstract: Five distinct ectomycorrhizal morphotypes were recognized on root systems of Fagus grandifolia collected in a maple-beech woodlot. Three morphotypes showed extensive root branching whereas two had limited root branching. One morphotype, a bright orange, smooth type, was studied in detail anatomically. A compact mantle with few emanating extraradical hyphae covered the root apex of each mycorrhizal lateral. A Hartig net extended around epidermal and hypodermal cells. Roots were mostly diarch and a single-layered endodermis and a biseriate hypodermis with the inner hypodermis having suberized walls were present in all specimens examined. The mantle, and to a lesser extent the Hartig net, stored lipids, polysaccharides and proteins.     

Chloroplast DNA phylogeography of Fagus crenata (Fagaceae) in Japan

 CpDNA variation in Japanese beech, Fagus crenata (Fagaceae), was studied in 45 populations distributed throughout the species' range. Two cpDNA regions were sequenced: the non-coding region between the trnL (UAA) 5′exon and trnF (GAA), and the trnK region (including matK). Thirteen distinct cpDNA haplotypes were recognized and each haplotype was found to be geographically structured. Two major clades (I and II+III) were revealed in phylogenetic analyses among the haplotypes using F. sylvatica as an outgroup. The haplotypes of Clade I were distributed mainly along the Japan Sea side of the Japanese Archipelago, while those of Clade II+III occurred chiefly along the Pacific Ocean side. Consequently, the distribution of the two major cpDNA clades suggests that there were two migration routes in the history of F. crenata; one along the Japan Sea and the other along the Pacific Ocean side of the Japanese Islands. Received March 19, 2001 Accepted November 22, 2001     

Bioclimate and growth history affect beech lifespan in the Italian Alps and Apennines

When site factors reduce growth rates, tree lifespan tends to increase. This study investigates processes leading to such inverse relationship in Fagus sylvatica stands distributed along two elevation gradients, with an emphasis on climatic response, suppression periods, and growth trends. Dendrochronological records from old‐growth beech populations sampled at different elevations within two different bioclimatic regions (Alps vs. Apennines), were used to investigate factors that control tree lifespan. Differences between old‐growth (12) and nearby managed (15) stands were used to assess effects of silvicultural practices on maximum age. Logging reduced tree lifespan not only by removing older trees, but also by reducing the number of years beech individuals spent in the shaded understory. Tree lifespan and growth rates were affected by climate (spring–summer temperature) and were inversely related to one another along elevation gradients. The greatest lifespan was observed in old‐growth high‐mountain populations, and was related not only to slower growth due to a shorter growing season, but also to multidecadal periods of growth suppression during the initial development stages in the understory (i.e., slower growth rates at the youngest cambial ages). Past unfavorable climatic periods (in this case, the Little Ice Age) also helped increase tree lifespan. Using a linear model, we estimated a reduction in beech lifespan of 23 ± 5 years for each degree of warming. Basal area increment of trees with the maximum observed lifespan showed an increasing trend over time. Because growth of old (>300 years) trees has increased in the Alps, while it has recently declined in the Apennines, different bioclimatic regions can have opposite responses to global climatic change. In the next decades, if warming continues, beech lifespan could be reduced in the Alps by faster growth and in the Apennines by drought‐induced mortality.     

Raman spectroscopy of pigments and oxalates in situ within epilithic lichens: Acarospora from the Antarctic and Mediterranean

The effects of ambient and elevated ozone levels on growth and photosynthesis of beech ( Fagus sylvatica ) were studied by exposing seedlings in open-top chambers for one growing season to three treatments: charcoal-filtered (CF), non-filtered (NF) and non-filtered air with addition of ozone (30 ppb ozone) on clear days for 8–10 h d⁻¹ (NF +). Ambient levels were relatively low and accumulated to an AOT40 (accumulated exposure over a threshold of 40 ppb) of 4055 ppb h (for the period 23 Apr–30 Sept). The NF + chambers received an AOT40 of 8880 ppb h. Throughout the growing season we measured growth and photosynthetic properties. The treatments did not cause strong effects: measurements of gas exchange (light-saturated assimilation rate, CO₂ and light-response curves) and chlorophyll fluorescence showed slight and mostly non-significant reductions of several parameters. No significant differences were found for growth, though in the NF + treatment (AOT40 8880 ppb h) the relative growth rate for diameter increment was at times reduced by 12% compared with the control treatment.     

Development and polymorphism of microsatellite markers for Fagus crenata and the closely related species, F. japonica

 We have developed microsatellite markers (SSRs) applicable to Fagus crenata using the RAHM method and investigated their polymorphisms. We also applied the SSRs in an analysis of a closely related species, F. japonica. Here we describe the isolation and characterization of nine polymorphic microsatellite markers, of which eight are applicable to both species. Among 30 individuals of each of F. crenata and F. japonica we detected a total of 79 and 77 alleles, respectively, with an average of 9.9 and 8.6 alleles per locus. The mean expected heterozygosity (He) was 0.615 (range: 0.216–0.925) in F. crenata and 0.660 in F. japonica (range: 0.259–0.827). The He values were considerably higher than those previously found for isozymes. Paternity exclusion probabilities for multiple loci, calculated over all loci, were extremely high (0.999 and 0.998 in F. crenata and F. japonica, respectively): sufficiently high to study pollen flow in both species. Received: 5 December 1998 / Accepted: 28 December 1998     

Effects of Sika deer on the regeneration of aFagus crenata forest on Kinkazan Island,northern Japan

Kinkazan Island, a sanctuary island that has been conserved for over 100 years, is inhabited by a Sika deer population of high density (60 km⁻²). AFagus crenata forest, a climax forest of this region, was composed of only large sized trees (30–60 cm diameter breast height) and lacked young trees. However, many saplings were found inside a deer proof fence, indicating this forest had a potential to produce seedlings. They were tentatively protected by shrubs ofLeucothoe grayana var.glabra, an upalatable ericaceous shrub, but did not exceed the shrub height. Heavy browsing by the deer suppressed recruitment of saplings ofF. crenata. The forest will be reduced without population control of the deer.     

A new scenario for the quaternary history of European beech populations: palaeobotanical evidence and genetic consequences

Here, palaeobotanical and genetic data for common beech (Fagus sylvatica) in Europe are used to evaluate the genetic consequences of long-term survival in refuge areas and postglacial spread. Four large datasets are presented, including over 400 fossil-pollen sites, 80 plant-macrofossil sites, and 450 and 600 modern beech populations for chloroplast and nuclear markers, respectively. The largely complementary palaeobotanical and genetic data indicate that: (i) beech survived the last glacial period in multiple refuge areas; (ii) the central European refugia were separated from the Mediterranean refugia; (iii) the Mediterranean refuges did not contribute to the colonization of central and northern Europe; (iv) some populations expanded considerably during the postglacial period, while others experienced only a limited expansion; (v) the mountain chains were not geographical barriers for beech but rather facilitated its diffusion; and (vi) the modern genetic diversity was shaped over multiple glacial-interglacial cycles. This scenario differs from many recent treatments of tree phylogeography in Europe that largely focus on the last ice age and the postglacial period to interpret genetic structure and argue that the southern peninsulas (Iberian, Italian and Balkan) were the main source areas for trees in central and northern Europe.