Syntaxonomy and phytogeographical differentiation of the Fagus woods in the Southwest Balkan Peninsula

Abstract. Up to now the syntaxonomy of the Fagus woods of the southern Balkans is far less known than that of beech communities in the northwestern part of the Balkan Peninsula and in central and western Europe. A set of 607 phytosociological relevés from southeastern Serbia, the Former Yugoslav Republic of Macedonia, and northern and central Greece was subjected to numerical classification and ordination. These results were used for a syntaxonomic revision of Fagus, Fagus‐Abies and Fagus‐Acer woodland communities of the southwestern Balkans. The Doronico columnae‐Fagenion (seven associations from Serbia, the F.Y.R.O.M., and high altitudes in northern Greece) and the Doronico orientalis‐Fagenion (four associations from northern and central Greece) were distinguished. The results of ordination by Canonical Variates Analysis (CVA) showed a very distinct phytogeographical pattern of differentiation between community types. The floristic composition of Fagus woods changes markedly along the north‐south gradient. The share of Subbalkan, Balkan, Submediterranean and Mediterranean species increases significantly towards the south, while the Central‐European, European, Euro‐Siberian, Euro‐Asiatic and Circumpolar species decrease in the same direction. The syntaxonomic differentiation of the Fagus woodlands and their classification into regional suballiances reflects this gradient. With the rise in altitude, the number of Balkan, Mediterranean, Submediterranean, European and Euro‐Asiatic species declines.     

Tree physiological responses to above-ground herbivory directly modify below-ground processes of soil carbon and nitrogen cycling

Above‐ground herbivory is ubiquitous in terrestrial ecosystems, yet its impacts on below‐ground processes and consequences for plants remain ambiguous. To examine whether physiological responses of individual trees may potentially modify soil nutrient availability, we subjected Fagus sylvatica L. (European beech) and Abies alba Mill. (silver fir) to simulated foliar herbivory over two growing seasons. Above‐ground herbivory enhanced N mineralization and inorganic N availability in the soil. The total input of C from the plant roots to the soil is not known; however, carbon sequestration in the soil, measured using stable isotopic techniques, was unaffected by herbivory. Fagus responded to herbivory by producing larger leaves, with increased photosynthetic capacity and N content, which largely compensated for the loss of biomass; Abies exhibited no such response. We conclude that despite large interspecific differences in the growth response, tree physiological responses to foliar herbivory are capable of directly modifying soil biological processes.     

Structure,dynamics and dendroecology of an old‐growth Fagus forest in the Apennines

Abstract. Question: Which are the structural attributes and the history of old‐growth Fagus forest in Mediterranean montane environments? What are the processes underlying their structural organization? Are these forests stable in time and how does spatial scale affect our assessment of stability? How do these forests compare to other temperate deciduous old‐growth forests? Location: 1600–1850 m a.s.l., Fagus forest near the tree line, central Apennines, Italy. Methods: An old‐growth Fagus forest was studied following historical, structural and dendroecological approaches. History of forest cover changes was analysed using aerial photographs taken in 1945, 1954, 1985 and 1994. The structural analysis was carried out in the primary old‐growth portion of the forest using 18 circular and two rectangular plots. Dendroecological analyses were conducted on 32 dominant or co‐dominant trees. Results: These primary old‐growth Fagus remnants consist of four patches that escaped logging after World War II. Both living and dead tree components are within the range of structural attributes recognized for old‐growth in temperate biomes. Dendroecological analyses revealed the roles of disturbance, competition and climate in structuring the forest. We also identified a persistent Fagus community in which gap‐phase regeneration has led to a mono‐specific multi‐aged stand at spatial scales of a few hectares, characterized by a rotated‐sigmoid diameter distribution. Conclusion: Even at the relatively small spatial scale of this study, high‐elevation Apennine Fagus forests can maintain structural characteristics consistent with those of old‐growth temperate forests. These results are important for managing old‐growth forests in the Mediterranean montane biome.     

Tree regeneration on rotten wood and on soil in old-growth stand

Forest regeneration on soil and on decaying wood was studied in natural mixed stand of Facus sylvatica L., Abies alba Mill. and Picea abies Karst. in Babia Góra National Park, Western Carpathians.Downed wood, divided into five decay classes covered around 6% of the forest floor. Among seedlings, Fagus and Abies codominated, while Picea was less numerous. The average seedling density on the soil with herb layer (240 ind./100 m²) was higher than on the logs, even on the strongly decayed ones (177 ind./100 m²). However, the density of Abies and Picea seedlings was higher on the rotten wood than on soil. Seedling survival of all species was better on the logs, especially in conifers. Because of the total dominance of Fagus among saplings, the presence of Abies and Picea in the next generation of canopy trees can strongly depend upon their regeneration on decaying wood.     

Spatial pattern dynamics over 10 years in a conifer/broadleaved forest,northern Japan

The present study investigated stand dynamics during 10-year period in a conifer/broadleaved mixed forest in Hokkaido, northern Japan, focusing on spatially dependent recruitment, mortality and growth of two growth-form groups, deciduous broadleaved species and the dominant evergreen conifer Abies sachalinensis. The stand-level basal area was maintained over the 10-year period, while a compositional equilibrium at the individual species level was not confirmed. Univariate and bivariate spatial analyses revealed clustering of many of the constituent species. The absence of single-species patches suggested an ambiguous mosaic formed by co-occurrence of Abies and broadleaved trees. The trend towards an aggregated distribution of Abies and broadleaved trees was caused by spatially dependent recruitment rather than mortality. New recruits of broadleaved species were spatially associated with surviving broadleaved trees, while this was not the case for Abies. The degree of competitive effects on growth was not consistent over the 10-year period. Abies showed between-groups competition, but not within-group competition. In contrast, we found asymmetric competitions between the broadleaved trees. Our results suggest that Abies is not sufficiently competitively dominant to exclude broadleaved trees, and that the co-occurrence of the two growth-form groups might be maintained.     

Predicting tree death for Fagus sylvatica and Abies alba using permanent plot data

Question: How well can mortality probabilities of deciduous trees(Fagus sylvatica) and conifers (Abies alba) be predicted using permanent plot data that describe growth patterns, tree species, tree size and site conditions? Location: Fagus forests in the montane belt of the Jura folds (Switzerland). Method: Permanent plot data were used to develop and validate logistic regression models predicting survival probabilities of individual trees. Backward model selection led to a reduced model containing the growth‐related variable ‘relative basal area increment’ (growth‐dependent mortality) and variables not directly reflecting growth such as species, size and site (growth‐independent mortality). Results: The growth‐mortality relationship was the same for both species (growth‐dependent mortality). However, species, site and tree size also influenced mortality probabilities (growth‐independent mortality). The predicted survival probabilities of the final model were well calibrated, and the model showed an excellent discriminatory power (area under the receiver operating characteristic curve = 0.896). Conclusion: Mortality probabilities of Fagus sylvatica and Abies alba can be predicted with high discriminatory power using a well calibrated logistic regression model. Extending this case study to a larger number of tree species and sites could provide species‐ and site‐specific tree mortality models that allow for more realistic projections of forest succession.     

Intraspecific facilitation: a missing process along increasing stress gradients – insights from simulated shrub populations

In recent years many field studies have been conducted to assess the relative importance of facilitation and competition in structuring vegetation communities in different environments. Herein, we present a simulation model which systematically explores the relative importance of intra‐specific facilitation and competition between adult shrubs and seedlings for spatial pattern formation. A grid‐based simulation model was constructed and calibrated using data collected in the field from Sarcopoterium spinosum populations in Israel to simulate population dynamics along a rainfall gradient. A series of simulation experiments was conducted in which manipulations of seedling survival probabilities were carried out to assess the relative importance of these processes in generating spatial patterns. Increased survival probabilities of first‐year shrubs in open areas were used to simulate competition effects, while increased survival probabilities in the vicinity of shrubs were used to simulate facilitation effects. Simulation results were then compared to shrub spatial patterns observed in the field. The results indicate that facilitation is not an important process in generating intra‐specific spatial patterns. Rather, in mesic environments with high precipitation, competition is the dominant process generating spatial patterns, resulting in regular spacing of shrubs, similarly to the patterns observed in the field (L(h) values<0). In arid sites, where precipitation values are lower, and stress conditions are higher, the dominant process generating spatial patterns was random mortality due to drought conditions. The resulting spatial patterns in this case are random (L(h)～0), whereas observed field populations exhibited clumped patterns (L(h)>0). We conclude that as stress conditions increase, the importance of intraspecific neighborhood interactions decrease whereas the importance of environmental factors increase in dictating intra‐specific spatial pattern formation. Consequently in mesic environments intra‐specific competition among adults determines the emerging patterns, while intraspecific facilitation is a negligible process.     

Tree mortality and effects of release from competition in an old‐growth Fagus‐Abies‐Picea stand

Abstract. In a montane mixed Fagus‐Abies‐Picea forest in Babia Gora National Park (southern Poland), the dynamics of an old‐growth stand were studied by combining an 8‐yr annual census of trees in a 1‐ha permanent sample plot with radial increments of Abies and Picea growing in the central part of the plot. The mortality among the canopy trees was relatively high (10% in 8 yr), but the basal area increment of surviving trees slightly exceeded the losses caused by tree death. DBH increment was positively correlated with initial diameter in Abies and Picea, but not in Fagus. For individual trees smaller than the median height, basal area increment was positively related to the basal area of old snags and the basal area of recently deceased trees in their neighbourhood, but negatively related to the basal area of live trees. Dendrochronological analysis of the past growth patterns revealed numerous periods of release and suppression, which were usually not synchronized among the trees within a 0.3 ha plot. The almost normal distribution of canopy tree DBH and the small number of young individuals in the plot indicated that stand dynamics were synchronized over a relatively large area and, hence, were consistent with the developmental phase concept. On the other hand, the lack of synchronization among periods of growth acceleration in individual mature Abies and Picea trees conforms more closely to the gap‐dynamics paradigm.     

The evolutionary history of Fagus in western Eurasia: Evidence from genes, morphology and the fossil record

 Fagus (beech) is among the most abundant and economically important genera of broad-leaved trees in northern hemisphere temperate forests. The number of modern taxa present in Europe and Asia Minor has long been a matter of debate and up to five species have been recognised. To resolve taxonomic and phylogenetic relationships we conducted morphological and molecular genetic analyses in western Eurasiatic taxa and evaluated palaeontological evidence. To place our findings from western Eurasiatic populations in a broader context additional East Asiatic and North American species of the same subgenus Fagus as well as two species of the subgenus Engleriana were included in our study. The morphological features exhibited in western Eurasiatic populations of Fagus show a west-east gradient that is characterised by strongly overlapping variability between geographical races. Fagus populations from Asia Minor exhibit an even higher variability that is also reflected in their genetic variability of nuclear rDNA internal transcribed spacer (ITS) sequences. The intraspecific genetic variability recorded here is in conflict with previous ITS studies in Fagus. The high amount of ITS polymorphism within Fagus from western Eurasia along with the clinal variation observed for morphological characters suggest the presence of only a single species, Fagus sylvatica L., in Europe and Asia Minor. Previously recognised taxa such as F. orientalis Lipsky and Fagus moesiaca (Maly) Czeczott should therefore be treated as synonyms of Fagus sylvatica. Although species belonging to the subgenus Engleriana were genetically distinct from species of the subgenus Fagus, relationships within the subgenus Fagus could not be clearly resolved. A reason for this could be the low rate of diversification in Fagus during the early phase of range expansion of the genus in the Oligocene period as indicated by the uniformity of leaf and cupule/nut fossils. This may account for the low overall ITS divergence and the high degree of polymorphism encountered in the subgenus Fagus and points to a late differentiation of western Eurasiatic and eastern Asiatic species. Area disruptions during the Pleistocene and the post-glacial recolonisation of western Europe appear to have caused the west-east gradient that is apparent in modern Fagus of western Eurasia but absent in Late Tertiary ancestors of Fagus sylvatica. Received June 22, 2001 Accepted February 25, 2002     

Short‐term changes affecting regeneration of Fagus crenata after the simultaneous death of Sasa kurilensis

Abstract. Question: The aim of the present study is to determine whether seed/seedling predation will increase and Fagus survival will decline with the recovery of the Sasa cover. Methods: We examined Fagus crenata regeneration for seven years in an old‐growth Fagus‐Sasa forest near Lake Towada, northern Japan, by examining the effects of simultaneous death of Sasa, tree canopy gap formation, mast seeding of Fagus and seed and seedling predation by rodents on the survival of Fagus seeds and current year seedlings. We established four types of sites differing in forest canopy (closed or gap) and Sasa status (dead or alive) following the simultaneous flowering and death of Sasa kurilensis (dwarf bamboo) in 1995. Results: Fallen Fagus seed was abundant in 1997 and 2000 (mast years). In sites with alive Sasa, survival from the first growing season was low due to high seed and seedling predation. In sites with dead Sasa, seed survival under the canopy was high for both mast years, but in gaps it varied between years. Seedling survival was highest in canopy gaps with dead Sasa (gap‐dead) in 1998, because of higher light levels and lower predation by rodents. However, seedling survival in these plots was low in 2001, apparently because rapid Sasa recovery favoured rodent predation. In both mast years, Sasa die‐back had significant positive effects on seed and seedling survival under closed canopies because the seedlings there were more successful in escaping predation. Conclusion: The change in successful sites for the early stage of regeneration of Fagus appears to reflect the combined effects of canopy gap, seed/seedling predation and revegetation of Sasa.     

Implications of 19th century landscape patterns for the recovery of Fagus crenata forests

Questions: What is the effect of the 19th century (pre‐industrialization) landscape pattern on the recovery of climax forests in cool‐temperate mountain areas dominated by Fagus crenata (Japanese beech)? Location: Secondary forests on Mt. Daisen, western Japan. Methods: Vegetation patterns before and after industrialization were obtained from maps drawn in 1898 and 1979. Tree measurements were made in 12 plots in 1997. Correlation between current Fagus crenata dominance and forest edge in the 19th century was analysed using an S‐shaped regression curve. Fagus juvenile density was counted in the plots, and distances from each plot to the five nearest mother trees were measured to determine the dispersal kernel. Results: Secondary grassland covered a substantial area in 1898, whereas forest covered most of the area in 1997. Fagus was dominant in places in the interior forest 100 years ago, and mature Fagus trees were absent in secondary forests that had been grasslands in 1898. The expected number of juveniles decreased to one individual per 100 m² at 43.5 m from the mother tree. Conclusions: The pre‐industrialization landscape greatly affected recovery of Fagus forest. Forests found on the 1898 vegetation map might have acted as refugia for Fagus. The limited dispersal ability of Fagus suggests that it would take many generations (several hundred years) for Fagus forests to recover at the centre of what had been grasslands in the 19th century.     

Distribution pattern and age structure ofAbies firma saplings in a mature mixed forest ofA. firma andFagus japonica

Age and spatial distribution ofAbies firma saplings were analyzed in a mature mixed forest ofA. firma andFagus japonica. There were two major peaks in the age structure ofAbies saplings at around 10 and 75 years of age. Distribution of these age classes of saplings was contagious, while that of all saplings was approximately random. These facts showed that most of the establishment ofA. firma occurs just after a gap is formed, and the frequency at which such establishment occurs is about once every 60 years in a 1600 m² area.     

Genetic variation in Tertiary relics: The case of eastern‐Mediterranean Abies (Pinaceae)

The eastern‐Mediterranean Abies taxa, which include both widely distributed species and taxa with minuscule ranges, represent a good model to study the impacts of range size and fragmentation on the levels of genetic diversity and differentiation. To assess the patterns of genetic diversity and phylogenetic relationships among eastern‐Mediterranean Abies taxa, genetic variation was assessed by eight nuclear microsatellite loci in 52 populations of Abies taxa with a focus on those distributed in Turkey and the Caucasus. Both at the population and the taxon level, the subspecies or regional populations of Abies nordmanniana s.l. exhibited generally higher allelic richness, private allelic richness, and expected heterozygosity compared with Abies cilicica s.l. Results of both the Structure analysis and distance‐based approaches showed a strong differentiation of the two A. cilicica subspecies from the rest as well as from each other, whereas the subspecies of A. nordmanniana were distinct but less differentiated. ABC simulations were run for a set of scenarios of phylogeny and past demographic changes. For A. ×olcayana, the simulation gave a poor support for the hypothesis of being a taxon resulting from a past hybridization, the same is true for Abies equi‐trojani: both they represent evolutionary branches of Abies bornmuelleriana.     

Pollen dispersal and deposition characteristics of Abies alba, Fagus sylvatica and Pinus sylvestris, Roztocze region (SE Poland)

Pinus sylvestris L., Abies alba Mill. and Fagus sylvatica L.—the significant forest forming tree species in Europe are important for palaeoecological interpretations based on the results of pollen analysis of fossil deposits. The potential pollen loading for Pinus sylvestris, Abies alba and Fagus sylvatica was modelled using simulated and actual vegetation maps, measured fall-speed values and pollen productivity estimates from the literature. The influx of fir pollen drops sharply with distance from the pollen source due to the high fall speed and moderate pollen productivity. The vast majority of Abies alba pollen is deposited within less than 50 m of the sampling site and a major proportion within 100 m. For beech the corresponding numbers would be 300 and 1,800 m, and for pine 1,000 and 4,500 m. The observed mean pollen accumulation rate (PAR) values for Pinus and Fagus were ca. 5,800 and 1,100 grains cm⁻² year⁻¹, respectively. In the case of Abies, the mean annual PAR for the whole region is ca. 700 grains cm⁻² year⁻¹. In SE Poland the regional signal is represented by PARs of Abies alba <200 grains cm⁻² year⁻¹ and of Fagus sylvatica <500 grains cm⁻² year⁻¹. The local presence/absence threshold values for Abies alba, Fagus sylvatica and Pinus sylvestris are >1,000 grains cm⁻² year⁻¹, >2,000 grains cm⁻² year⁻¹ and >3,500 grains cm⁻² year⁻¹ respectively.     

Competitive effects of herbaceous vegetation on tree seedling emergence,growth and survival: Does gap size matter?

Question: What is the effect of gap size on the seedling emergence, growth and survival of four common tree species in wooded pastures? Location: A pasture in the Jura mountains, Switzerland. Methods: Seeds were sown in a complete three‐way factorial design with eight blocks in May 2003. Each block consisted of a competition treatment (four gap sizes including zero) and a mowing treatment (mown and unmown). Emergence, survival and total biomass of tree seedlings of three species (Picea abies, Acer pseudoplatanus and Fagus sylvatica) were measured. A fourth species (Abies alba) failed to germinate. Results: Gaps had a positive influence on the early stages of tree development for all species. Larger gaps favoured growth and survival more than small gaps. Seedling growth was higher when vegetation around the openings was mown. Mowing the vegetation at gap size zero enhanced both growth and survival compared to unmown vegetation. Mown gaps larger than zero had increased seedling desiccation but decreased seedling predation. Species showed similar trends in their emergence and growth responses to gap size and mowing treatments but for Picea emergence rate was higher and survivorship was lower than for Acer and Fagus. Conclusions: Gap size does matter for tree seedling success but even in more favourable large gaps only a small percentage of seedlings emerged and survived. The effects of gaps on tree seedling establishment are complex as a result of interactions between biotic and abiotic changes caused by gaps.     

Codominance of <Emphasis Type="Italic">Acer saccharum</Emphasis> and <Emphasis Type="Italic">Fagus grandifolia</Emphasis>: the role of <Emphasis Type="Italic">Fagus</Emphasis> root sprouts along a slope gradient in an old-growth forest

We studied how the unusual capacity of mature Fagus grandifolia to form clumps of clonal stems from root sprouts can contribute to its frequent codominance with Acer saccharum in southern Quebec, Canada. In an old-growth forest, the degree of dominance by the two species shifted along topographic gradients spanning a few hundreds of meters, with Fagus more frequent on lower slopes and Acer on upper slopes. The frequency distribution of Fagus stem diameter had an inverse J distribution at all slope positions, which is indicative of continuous recruitment. Acer stem diameter also had an inverse J pattern, except at lower slope positions where size structure was discontinuous. For stems <2 m tall, Fagus regenerated mainly by sprouts at the upper and mid-slopes, while regeneration from seed was more pronounced on the lower slope. This change of regeneration mode affected the spatial pattern of Fagus stems. Understory trees of Fagus were positively correlated with conspecific canopy trees on upper and mid-slopes, but not on lower slopes where Fagus regenerated mainly by seedlings. Understory trees of Acer were positively correlated with conspecific canopy trees only on the mid-slope. There were many Fagus seedlings around Acer canopy trees at the lower slope, suggesting the potential replacement of Acer canopy trees by Fagus. This study suggests that the regeneration traits of the two species changed with slope position and that Fagus patches originating from root sprouts can contribute to the maintenance of Acer–Fagus codominance at the scale of local landscapes.     

Long-term human impact as registered in an upland pollen profile from the southern Black Forest, south-western Germany

A detailed,¹⁴C-dated, pollen profile from Steerenmoos, a raised bog in the uplands of the southern Black Forest (Schwarzwald) is presented. The Late-glacial and early Holocene vegetation history conforms to the known pattern of forest dynamics for that region. At ca. 6100 cal. B.P.,Abies replaced the mixed oak forest, which is in contrast to adjacent regions whereFagus spread beforeAbies. From the Subboreal onwards,Fagus expanded and slowly largely replacedAbies. The mire developed from a fen to a raised bog. The mountain pine (Pinus mugo ssp.rotundata) on the present-day bog surface is a result of medieval burning. Cereal pollen are first recorded in the Neolithic (7600 cal. B.P.) and there is a closed curve forPlantago Lanceolata — a good indicator of human impact — since the Bronze Age (4000 cal. B.P.). On the basis of the cereal pollen record nine human impact phases (HIP) are described. HIP 1 and 2, which are short, date to ca. 7600 and 6700 cal. B.P., respectively, in a mixed oak forest context and are characterized by declines inCorylus, Tilia, Ulmus and bySalix (but no major deforestation) and peaks in charcoal and loss-on-ignition curves. HIP 3 and 4, which are short and weak, date to ca. 6000 and 5300 cal. B.P., respectively, and occur in the context of anAbies alba forest. The Bronze Age and Iron Age HIPs 5-7 are more intense and of longer duration than the Neolithic phases and result in a decline inAbies and an increase inFagus. The early medieval HIP 8, although rather weak, probably finds expression also in an archaeological artefact, namely a dug-out boat from the near-by Schluchsee. Finally, the late Medieval HIP 9 resulted in a major transformation in the landscape. It is argued that the earlier HIPs are not a reflection of distant events in the lowland valleys of the Rhine, Danube or Neckar but reflect more or less local developments.     

Dynamics in Central European near‐natural Abies‐Fagus forests: Does the mosaic‐cycle approach provide an appropriate model?

Question: The mosaic‐cycle concept of forest dynamics dominates in Central Europe. According to this concept intermediate‐scale disturbances only accelerate the forest break‐up under existing cycles of forest development. Is such an approach correct, or should new developmental cycles be elaborated for intermediate‐scale disturbances? Location: Near‐natural Abies alba ‐ Fagus sylvatica forests in the ?wiétokrzyski National Park in Central Poland. In these forests intermediate‐scale disturbances occurred between 1970 and 1990. Methods: Data were collected twice in areas surrounding 212 permanent sample points (in 1994 and 2004). Two increment cores were taken from 259 sample Abies trees. The effect of intermediate‐scale disturbances on radial increment of Abies was assessed. Probabilities of stand transition during a 10‐year period between individual stages and phases of development of the mixed forest were calculated. The development stages and phases were arranged into hypothetical succession series of successive changes. Results: In 1994 70 stands and in 2004, 47 stands representing stages and phases containing the older generation formed by trees > 100–150 years were found. Also, in 1994 142 and in 2004, 165 stands representing stages and phases containing the younger generations only, formed by trees < 100‐150 years, were recorded. Stages and phases containing only younger generations do not occur in the existing forest development cycle which does not consider the influence of intermediate‐scale disturbances separately. Two developmental cycles, which take into account the presence of the older generation and the younger generations only (under conditions of the occurrence of intermediate‐scale disturbances), are proposed. Conclusion: The mosaic‐cycle concept of forest dynamics can be used to analyse the dynamics of Central European near‐natural mixed‐species forests, but new developmental cycles should be elaborated for intermediate‐scale disturbances.     

Buxus sirindhorniana sp. nov. (Buxaceae), a bicarpellate species from Thailand

A new bicarpellate Buxus species, Buxus sirindhorniana W. K. Soh, M. von Sternburg, Hodk. & J. Parn., is described from northern Thailand. This is the first reported bicarpellate species of Buxus. The pollen morphology and the degree of pollen viability of the species are reported. DNA was extracted from herbarium samples and the internal transcribed spacer (ITS) region of nuclear ribosomal DNA sequenced. The palynological evidence and the phylogenetic reconstruction support the contention that this new species belongs to the genus Buxus. The morphological diversity of reproductive organs in the Buxaceae, in relation to this new species, is briefly discussed.     

Forecasting the pattern and pace of Fagus forest expansion in Majella National Park,Italy

Question: Can the pattern and pace of spontaneous Fagus forest expansion from 1975 to 2003 be accurately detected with mid‐resolution satellite imagery? Can the historical Fagus expansion be modelled on the basis of environmental predictors? If so, where are the highest probabilities for future Fagus expansion? What are the implications for park management? Location: Majella National Park, Italy, > 1000 m a.s.l.; municipalities of S. Eufemia and Pacentro. Methods: Fagus cover change was detected by overlaying three classified sequential satellite images. Historical Fagus expansion was related to environmental variables using ordinary logistic and autologistic regression models. Fagus expansion probabilities were generated with the best predictive model. Results: From 1975 to 2003 Fagus advanced into abandoned farmland and subalpine pastures from the contiguous, midaltitudinal Fagus forest and from Fagus outliers, at a rate of 1.2 % per year. Substantial spatial and temporal variations in expansion rates were detected. The ordinary and autologistic models based on the single predictor Distance‐from‐Fagus‐1975 forecasted the Fagus expansion well (AUC 0.81 resp. 0.88). Multiple logistic models, including the topo‐climatic and substrate predictors, improved prediction insignificantly. The strong predictive power of proximity to historical Fagus presence is explained by the dispersal biology of Fagus combined with the shading impact of the Fagus canopy at the forest fringe. Conclusion: Decade‐long Fagus expansion patterns might be reliably forecasted by proximity to historical Fagus distribution. Consequences for park management options are outlined.