Phytophthora quercetorum sp. nov., a novel species isolated from eastern and north-central USA oak forest soils

Isolates belonging to an undescribed Phytophthora species were frequently recovered during an oak forest soil survey of Phytophthora species in eastern and north-central USA in 2004. The species was isolated using an oak leaf baiting method from rhizosphere soil samples collected from Quercus rubra, Q. macrocarpa, and Q. phellos. This species is formally described as P. quercetorum. It is homothallic and has aplerotic oogonia and paragynous antheridia. It produces papillate sporangia (occasionally bipapillate) of ovoid-elongated shapes. Its temperature optimum for growth is ca 22.5 °C with the upper limit of ca 32.5 °C. P. quercetorum differs from the morphologically related P. quercina in producing distinct submerged colony-patterns, different growth-temperature requirements, and oogonial shapes and sizes. Phylogenetic analyses using seven nuclear loci supported P. quercetorum as a novel species within clade 4, closely related to P. arecae, P. palmivora, P. megakarya, and P. quercina.     

Comparative water relations of four Mediterranean oak species

The water relations and responses of two evergreen (Quercus ilex L. and Q. suber L.) and two deciduous (S. afares Pomel. and Q. faginea Will.) Quercus species were studied under experimental conditions. Two-year old seedlings grown in 30 l. pots were subjected to a drying period during which stomatal conductance, pre-dawn potential and minimum foliar potential were measured.The results shows that, for all species, the daily course of stomatal conductance agrees with the patterns proposed by Hinckley et al. (1978 & 1983). Concurrent with the species responses to short-term variation in water availability, it was found that pre-dawn leaf water potential controlled the maximum daily leaf conductance. There was a strong correlation between pre-dawn leaf potential and maximum daily conductance as described by the reciprocal function g_{srmax for}=(-0.47+2.61._p)^-1 the evergreen oaks and g_{srmax for}=(-1.94+7.39._p)^-1 for the deciduous species. These differences between the two groups may partialy explain their geograhic distributions, and suggest general questions concerning the mechanisms which optimize water-use efficiency in Mediterranean oak species.     

The regeneration of pioneer tree species under browsing pressure of Sika deer in an evergreen oak forest

The effects of Sika deer (Cervus nippon) browsing on the regeneration of pioneer species were studied in relation to canopy gaps in a warm temperate evergreen oak forest in Kasugayama, Nara City. Four study sites, three in canopy gaps and one under a closed canopy, were selected and each divided into fenced and unfenced plots. Under the closed canopy, seedlings of all the pioneer species died irrespective of browsing pressure. However, in the canopy gap sites, seedlings of the pioneer species could establish and grow well. The seedling survival ratio in the fenced plots in the canopy gaps was >60% 1 year after germination. However, in the unfenced plots, only <20% of the seedlings survived 1 year, with all dying within 3 years after germination. Thus, the regeneration of pioneer trees in this forest was strongly inhibited by deer browsing. Successful regeneration of a pioneer,Zanthoxylum ailanthoides, occurred for several years even after two major wind disturbances during the past 90 years. This may be due to less browsing pressure from the deer.     

Differential response of two Central-European oak species to single and combined stress factors

We studied the effects of repeated defoliation and drought, which are considered to be most effective in triggering oak decline, on the Central-European oak species Quercus robur L. and Quercus petraea [Matt.] Liebl. Three-year-old saplings were subjected to artificial defoliation in two consecutive years and to drought stress, applied singly or in combination. Differences in the morphological, anatomical and physiological responses to these stress factors among treatments and between species were tested in a 3-year experiment. Drought stress lowered the pre-dawn and the afternoon leaf-water potentials in both species, irrespective of additional defoliation. The relative water content of the leaves was unaffected by drought stress; but, in Q. robur, it decreased in trees subjected to defoliation and to the combination of drought stress and defoliation. When repeated defoliation and drought stress were applied within the same growing season, the leaf-to-air difference in the partial pressure of water vapour (w) explained most of the variation in the daily integral of the stomatal conductance to water vapour (g _{s day sum}) in Q. petraea; however, in Q. robur, the hydraulic conductance of the twigs was the most powerful predictor variable. In the same year, the actual hydraulic conductance was correlated with g _{s day sum} in Q. robur, but not in Q. petraea. The studied parameters of gas exchange and biomass production revealed that Q. robur recovered more rapidly from stress than did Q. petraea. We hypothesize that this is due to Q. roburs lower responsiveness to the environmental variable w and its higher responsiveness to internal hydraulic conductance compared to Q. petraea. This revised version was published online in May 2005 with corrections to Table 4.     

Productivity and mineral cycling in an oak coppice forest 2. Annual net production of the forest

Annual net production was estimated in the secondary coppice forest near Tokyo, which was dominated by a deciduous oak,Quercus serrata Thunb. Lateral growth of stems and old branches was directly estimated by examining the annual rings for 35 shoots in a clear-cut quadrat of 10m×10m. Phytomasses of current organs were also weighed in the quadrat. Preharvest losses of current organs were determined by twelve 0.5 m² litter traps for fine litter and twelve 6 m² quadrats for woody litter. Branch production was also assessed indirectly by use of the stem-branch allometry and death of branches. The results of the indirect method were in sufficient agreement with the result of the direct one. Grazing loss of leaves from the canopy was estimated directly from the loss in leaf area and indirectly from the animal faeces caught by the litter traps. Net production of the canopy trees was 149 kg a⁻¹ year⁻¹, in which leaf production was 36.9 kg. Animals grazed about 14% of the leaf area by the end of the growing season. True consumption of leaves by animals was 7.6% of leaf production or 10% of leaf mass. Production of undergrowth, mainly a dwarf bamboo,Pleioblastus chino Makino, was 28 kg a⁻¹ year⁻¹, being 15% of the total stand production. Productivity of this forest was significantly higher than that of cool-temperate deciduous broadleaf forests.     

Effects of environmental conditions on isoprene emission from live oak

Live-oak plants (Quercus virginiana Mill.) were subjected to various levels of CO₂, water stress or photosynthetic photon flux density to test the hypothesis that isoprene biosynthesis occurred only under conditions of restricted CO₂ availability. Isoprene emission increases as the ambient CO₂ concentration decreased, independent of the amount of time that plants had photosynthesized at ambient CO₂ levels. When plants were water-stressed over a 4-d period photosynthesis and leaf conductance decreased 98 and 94%, respectively, while isoprene emissions remained constant. Significant isoprene emissions occurred when plants were saturated with CO₂, i.e., below the light compensation level for net photosynthesis (100 mol m^-2 s^-1). Isoprene emission rates increased with photosynthetic photon flux density and at 25 and 50 mol m^-2 s^-1 were 7 and 18 times greater than emissions in the dark. These data indicate that isoprene is a normal plant metabolite and not — as has been suggested — formed exclusively in response to restricted CO₂ or various stresses.Abbreviation PPFD photosynthetic photon flux density     

Importance of fragmentation-tolerant species as seed dispersers in disturbed landscapes

Forest fragmentation can negatively affect plants if animal seed-dispersers become locally extinct in fragments. We conducted a 2-year experiment to evaluate the importance of tree squirrels (Sciurus) as seed dispersers for Quercus, Carya, and Juglans, and to assess dispersal consequences in patches where fragmentation-sensitive eastern gray squirrels (Sciurus carolinensis) are absent. We accounted for fates of ∼15,700 seeds from five tree species in four exclosure treatments at 18 fragments during a high (2003–2004) and low seed (2004–2005) year. Two treatments excluded Sciurus to mimic disperser loss. We sampled nut-tree seedling density at 259 sites across eight watersheds, half of which were too fragmented to support S. carolinensis, but supported fragmentation-tolerant fox squirrels (Sciurus niger). Autumn-to-spring seed survival was low (∼1%) for all species during low seed production. During high seed production, survival was higher for Juglans nigra (20%) and Carya ovata (16%) than for three Quercus species (∼4% for Quercus palustris and Quercus rubra in two exclosure types; ∼1% for Quercus alba in all treatments). Survival of J. nigra, C. ovata, and Q. rubra was ≥2.1–7.7 times higher for seeds in exclosures that Sciurus could access. Seed displacement distance was higher in the low seed than the seed-rich year, but the proportion of seeds surviving to greater distances was higher in seed-rich years for all seed types except Q. rubra. This affirms the importance of masting to seed survival and dispersal, but also suggests an advantage to trees of producing seed in non-mast years. Seedling densities were comparable in watersheds with and without S. carolinensis. These results demonstrate the importance of tree squirrels as dispersers of nut-bearing trees, but suggest that fragmentation may not disrupt dispersal of certain species if losing S. carolinensis from disturbed landscapes is compensated for by fragmentation-tolerant fox squirrels (S. niger).     

Effects of meteorological parameters and plantdistribution on Chenopodiaceae-Amaranthaceae, Quercus and Olea airborne pollen concentrationsin the atmosphere of Cartagena (Spain)

We have studied the effects of various meteorologicalparameters and plant distribution on the airborneconcentrations of three different pollen types:Chenopodiaceae-Amaranthaceae, Quercus and Olea, in the city of Cartagena, situated in SouthwestEurope. The aerobiological study has been carried outfrom March 1993 to March 1997, using a Hirst-typeLanzoni 2000 trap, situated on the roof of the citytrain station. Meteorological data were supplied bythe Environmental Service of Cartagena's Town Hall.Daily pollen concentrations were standardised.Correlation analysis showed the importance ofmeteorological data on airborne pollen concentrations.On the other hand, ANOVA analyses depicted theinfluence of vegetation distribution in relation tosampler location and wind directions. Finally,multiple regression analyses allowed us to developcorrelation models for the three different pollentypes considered, accounting for 32%(Chenopodiaceae-Amaranthaceae), 19% (Quercus),and 26% (Olea) of the variation in the airbornepollen concentration.     

Seasonal variation in leaf chemistry of the coast live oak Quercus agrifolia and implications for the California oak moth Phryganidia californica

Summary The perennial foliage of the California coast live oak (Quercus agrifolia Nee) permits herbivores to feed on this oak species throughout the year. Patterns of herbivory for a two-year period on Q. agrifolia were observed in relation to seasonal and age-related changes in the nutritional and defensive characteristics of leaves. Nitrogen and phosphorus contents were higher in new leaves compared to mature foliage. Structural compounds (e.g., cellulose) in leaves rapidly increased with age. Concentrations of tatal phenolics (Folin-Denis) and astringency were higher in new foliage, and concentrations of condensed tannins gradually increased as the leaves matured. Peaks of herbivore damage were observed in June and in September–October, and were caused by outbreaks of the California oak moth (Phryganidia californica). P. californica, a bivoltine oak specialist, exhibited feeding preferences in June for old leaves over emerging leaves, and showed no preferences for leaf classes in September. These results suggest that P. californica is adapted to survive on nutritionally poor foliage and to circumvent quantitative defenses such as condensed tannins.     

Edge effects in a tropical montane forest mosaic: experimental tests of post-dispersal acorn removal

Forest edges created by scattered-patch clear-cutting have become common in tropical montane cloud forests in the highlands of Chiapas, Mexico. It was hypothesised that forest edges may influence regeneration of oak species, which are canopy dominants in these forests, by affecting the activities of small mammal species. Acorns of different oak species varying in germination timing were offered to predators and/or dispersers at different positions along replicated forest edges during 2 consecutive years. We investigated the effects of (1) edge type (hard and soft), (2) distance from the edge (0, 15, 30, 45 and 60 m inside forest fragments) and (3) oak species, on the rate of acorn removal mainly by small mammals. During a non-masting year, acorn removal was affected by the interaction of edge type and distance from the edge (P<0.05), with acorn removal being highest near hard edges compared to adjacent forest interiors. As predicted, acorn removal was greater along soft (100%) than along hard edges (82%), but this pattern was recorded only during the non-masting year. This study partly supports previous studies of rodents preferentially consuming acorns with early germination rather than acorns exhibiting dormancy, however these patterns may change with variation in acorn abundance. These results suggest that patch clear-cutting affects regeneration processes within forest fragments by influencing the activities of small mammals, but the nature of this effect also depends on acorn abundance and the characteristics of the forest edge created.     

Comparative studies on acorn and seedling dynamics of fourQuercus species in an evergreen broad-leaved forest

The dynamics and regeneration probabilities ofQuercus acuta, Q. gilva, Q. salicina, andQ. sessilifolia were studied in a 0.48 ha plot in an old-growth evergreen broad-leaved forest, Kagoshima prefecture, southwestern Japan. The canopy was well closed and had only one small gap. Seventeen tree species occurred in the canopy. The fourQuercus species ranked second to fifth in basal area. Excluding the trees originated from sprouts,Q. acuta and Q. gilva lacked trees at the subcanopy layer, andQ. salicina andQ. sessilifolia had some trees at the same layer. For eachQuercus species, the amount of fallen acorns fluctuated from year to year. The number of current year seedlings was positively correlated with the number of fallen acorns in the previous year. A disproportion between the spatial distribution of fallen acorns and that of established seedlings has been shown. The establishment of seedlings was frequently observed where no acorns had fallen in the previous year. Saplings of four species survived for several years and formed sapling banks under the closed canopy. During the three years of the study, the mortality ofQ. salicina andQ. sessilifolia saplings was lower than that ofQ. acuta andQ. gilva. However, the differences in the average annual height growth of saplings between four species were not significant, and the most of them did not grow much. These results suggest that the saplings of the fourQuercus, especiallyQ. gilva andQ. acuta, have no chance to recruit to the canopy or subcanopy layer within a closed stand, and that some changes in the environment caused by gap formation are required for their recruitment.     

Macaranga and Mallotus species (Euphorbiaceae) as indicators for disturbance in the mixed lowland dipterocarp forest of East Kalimantan (Indonesia)

The indicator value (IV) of Macaranga and Mallotus species (Euphorbiaceae) for different types of disturbance in lowland dipterocarp forest was assessed by counting and identifying all individuals of species of these genera taller than 30 cm in 45 (10 m×300 m) plots at nine locations. Twelve Macaranga and nine Mallotus species were found. The main forest disturbance types (primary forest, secondary forest, selectively logged forest, forest burned once, and repeatedly burned forest used for shifting-cultivation) each had their own set of indicator species. The level of disturbance in the forest types was assessed by measuring nine forest structural parameters. The occurrence of Macaranga and Mallotus species was closely related to the level of disturbance in a forest. Most Macaranga species were characteristic of high disturbance levels, while most Mallotus species preferred intermediate to low levels of disturbance. However, both genera had species at both disturbance extremes. Using multiple regression analysis, combinations of Macaranga and Mallotus species were formed and used to predict the separate forest structural parameters and the general level of disturbance of a forest. The Macaranga and Mallotus species could be grouped into (1) primary forest ‘remnant’ species; (2) generalist pioneer species; and (3) high disturbance pioneer species.     

Determinants of woody species richness in Scot pine and beech forests: climate, forest patch size and forest structure

We analysed patterns of woody species richness in Pinus sylvestris and Fagus sylvatica forests in Catalonia (NE Spain) from forestry inventory databank in relation to climate and landscape structure. Both types of forests are found within the same climatic range, although they have been managed following somewhat different goals. Overall, woody species richness significantly increased when conditions get closer to the Mediterranean ones, with milder temperatures. Differences between the two types of forests arose when comparing the relationship between richness and forest patch size. Woody species richness increased in pine forests with patch size, while the opposite trend was observed in beech forests. This pattern is explained by the different behaviour of structural canopy properties, since leaf area index and canopy cover showed a steeper increase with increasing forest patch size in Fagus forests than in Pinus ones. Accordingly, richness decreased with canopy cover in Fagus plots, but not in Pinus ones. We suggest that these differences would be related to management history, which may have enhanced the preservation of beech stands in larger forest landscape units.     

Predation by vertebrates and invertebrates on the seeds of five canopy tree species of an Amazonian forest

We studied the pre-germination loss of seeds to invertebrate and vertebrate seed predators of 5 species of Amazonian trees (Astrocaryum macrocalyx—Palmae; Bertholletia excelsa—Lecithydaceae; Calatola venezuelana—Icacinaceae; Dipteryx micrantha—Leguminosae (Papilionoidae); Hymenaea courbaril-Leguminosae (Caesalpinoidae)). These five species were selected from a large tree flora on several criteria. All possess large (3–10 cm) well-protected seeds that might plausibly be attractive to mammalian seed predators. The reproductive biology of three of the species, or close congeners, had been studied elsewhere in the Neotropics (Astrocaryum, Dipteryx, Hymenaea); one is important to the economy of southeastern Peru (Bertholletia); and one, despite large and apparently edible seeds, appeared to suffer no pre-germination loss to predators (Calatola). We conducted the research in mature forests in the Manu National Park of southeastern Peru where mammal densities are unperturbed by human activities. Densities of adult trees of the five species in our area range from very high (>30 per ha: Astrocaryum) to very low (1 per ha: Hymenaea).Loss of seeds to all causes, and to mammalian seed predators in particular, was determined for seeds placed in 2-square meter mammal exclosures and in open controls located at 10 m (near) and 50 m (far) from a large mature individual of the target species (with minor variations in the design for Astrocaryum and Calatola). The exclosures were of two types: impermeable—designed to exclude all mammals, but not invertebrate seed predators, and semipermeable—designed to admit small (<500 g), but not large mammals. Experimental and control plots were stocked with apparently viable seeds during the dry-wet transition period (October–November) and scored one year later.A significant distance effect (higher predation near vs far from a large conspecific adult) was found in only one of the species (Astrocaryum), the only one to be attacked with high frequency by invertebrate seed predators. The absence of any detectable distance effect attributable to mammals suggests that mammals, over the course of a year, thoroughly search the forest floor for seeds. Invertebrates may thus be responsible for most pre-germination distance (density) effects. With respect to the treatments, we found three qualitatively distinct results: seeds of three species (Astrocaryum, Bertholletia, Dipteryx) were significantly protected by the impermeable, but not semipermeable exclosures, implicating small mammals in seed loss; the seeds of one species (Hymenaea) were significantly protected by exclosures of both types, implicating large mammals; and the seeds of one species (Calatola) exhibited 100% survival, whether or not protected by exclosures.The importance of large mammals as seed predators is generally underestimated in these experiments because semipermeable exclosures may serve as foraging reserves for small mammals. Finally, we noted no relationship between the intensity of mammalian seed predation (as suggested by the survival of unprotected seeds) and the abundance of adults of the five species in the environment. The diversity of results obtained for the five species reveals that large-seeded tropical trees may display a wide range of demographic patterns, and points to the likely importance of post-germination bottlenecks in the population biology of many species, even those that may experience severe pre-germination seed loss.     

Copper uptake by some forest tree species from an acid sandy soil

Summary The behaviour of some young conifers and broadleaves on a very acid sandy humuspodzol soil was studied in pot trials. Species differed considerably in foliar copper concentration, reaction to N fertilization and development of Cu deficiency symptoms. Foliar copper concentrations were highest for the broadleaves (esp. oak and beech), and was increased by Cu and N fertilization, the last reaction showing an optimum curve. Cu deficiency symptoms could easily be induced in the conifers but not in the broadleaves.     

Light, nutrients and the growth of herbaceous forest species

The herb layer of forests planted on former agricultural land often differs from that of old-growth forest. This study investigates if the expected increased nutrient availability in the shaded conditions of newly planted forests and the plasticity of the species to adjust their biomass allocation to different levels of light and nutrients help to explain these differences in the herb layers of the two forest types. In a greenhouse experiment biomass distribution and production of two species characteristic for the highly shaded forest floor, Circaea lutetiana and Mercurialis perennis, and two species more common in the forest-edge, Aegopodium podagraria and Impatiens parviflora were studied at different levels of light (2%, 8% and 66% of the full light level) and nutrients (30 and 300 kg N ha^–1 per year). The main factor affecting allocation and biomass production was light availability. Nutrient supply only had a significant effect at the higher light levels. Species were mainly plastic to changes in light and the two species from the forest floor showed to be more rigid in allocation pattern than the species from the forest-edge. So, although the species from the forest-edge were more plastic, they did not profit from the increased nutrient supply because the main factor affecting biomass distribution and production was light availability.     

Interactions between host plants,endophytic fungi,and a phytophagous insect in an oak (Quercus grisea x Q. gambelii) hybrid zone

We examined interactions between host plants, endophytic fungi, and leaf-mining moths (Phyllonorycter sp.) in an oak (Quercus grisea x Q. gambelii) hybrid zone. The community of endophytic fungi and two common endophyte species examined responded to host plant hybridization. Total fungal frequency (TFF) and frequency of Gnomonia cerastis were lowest on hosts resembling Q. grisea, and increased linearly towards those resembling Q. gambelii. In contrast, Coccochorella quercicola was most frequently isolated from Q. grisea-like hosts and decreased in frequency across hybrids towards Q. gambelii. Frequency of G. cerastis and TFF covaried with Phyllonorycter density across the hybrid zone, but direct effects of endophytes on Phyllonorycter density were not detected. Associations between endophytes and unexplained mortality of Phyllonorycter varied according to endophyte species and state of Phyllonorycter development. In the sap-feeding stage, unexplained mortality was negatively associated with TFF and frequencies of Hormonema sp. and Preussia funiculata; whereas, in the tissue-feeding stage, unexplained Phyllonorycter mortality was positively associated with G. cerastis frequency. Three-way interactions between plant hybridization, endophytic fungi, and the insect herbivore were not significant.     

Geographical distribution of two haplotypes of chloroplast DNA in four oak species (<Emphasis Type="Italic">Quercus</Emphasis>) in Japan

Chloroplast DNA polymorphism in four oak species (Quercus serrata, Q. mongolica var. crispula, Q. dentata and Q. aliena) was studied using collections from a total of 127 localities in Japan and South Korea on the basis of five intergenic spacers (trnD-trnT, trnT-trnL, rps14-psaB, trnS-trnT and trnQ-trnS). Although no variation existed in sequences among the four species, a single nucleotide (T/C) substitution in the trnQ-trnS intergenic spacer was found in all the four species, resulting in two haplotypes (T- and C-type). Phylogenetic analyses of the four species and related species showed that the C-type is derived and even likely of monophyletic origin, while the T-type is ancestral. Geographically, the T-type is widespread from South Korea to Japan, whereas the C-type is restricted to eastern Japan with rare exceptions. Eastern Japan approximately coincides with the distribution range of the boreal conifer forest during the last glacial maximum. Overall evidence suggests that the mutation from T- to C-type occurred in an individual of one of the four oak species and then was transferred to all the species by hybridization in eastern Japan, and that the Kanto District provided individuals with the C-type with a refugium during the last glacial maximum.Electronic Supplementary Material Supplementary material is available in the online version of this article at     

Loss of oak dominance in dry-mesic deciduous forests predicted by gap capture methods

Gap capture methods predict future forest canopy species composition from the tallest trees growing in canopy gaps rather than from random samples of shaded understory trees. We used gap capture methods and a simulation approach to forecast canopy composition in three old oak forests (Quercus spp.) on dry-mesic sites in southern Wisconsin, USA. In the simulation, a gap sapling is considered successful if it exceeds a threshold height of 13–17 m (height of maximum crown width of canopy trees) before its crown center can be overtopped by lateral crown growth of mature trees. The composition of both the tallest gap trees and simulated gap captures suggests that 68–90% of the next generation of canopy trees in the stands will consist of non-Quercus species, particularly Ulmus rubra, Carya ovata and Prunus serotina. Quercus species will probably remain as a lesser stand component, with Quercus alba and Quercus rubra predicted to comprise about 19% of successful gap trees across the three stands. Several methods of predicting future canopy composition gave similar results, probably because no gap opportunist species were present in these stands and there was an even distribution of species among height strata in gaps. Gap trees of competing species already average 11–13 m tall, and mean expected time for these trees to reach full canopy height is only 19 years. For these reasons, we suggest that dominance will shift from oaks to other species, even though late successional species (e.g., Acer and Tilia) are not presently common in the understories of these stands.     

Biomass and production of a naturally regenerated oak forest in southern Korea

Biomass and production of two stands with Quercus variabilis Bl. as the dominant species (stands 1 and 3) and one with Q. mongolica Fisch. as the dominant species (stand 2) were investigated in southern Korea. Stands 1 and 3 naturally occurred on sites with southerly aspects while stand 2 naturally occurred on northerly aspects; stand ages were similar for the three stands (36–38 years old). Total above- and belowground biomass including understory vegetation (Mg ha^–1) was 108.4 for stand 1, 115.6 for stand 2, and 132.0 for stand 3, respectively. Understory vegetation constituted 17.4% of the total biomass in stand 1 but only 3.7–4.5% in stand 2 and stand 3. Roots constituted 20.1–24.6% of the biomass of the overstory vegetation. Although stand 3 showed the highest total biomass, net production was highest in stand 2 at 12.6 (Mg ha^–1 year^–1); net production levels for stands 1 and 3 were 11.7 and 11.1 (Mg ha^–1 year^–1), respectively. It appeared that the differences in site conditions related to aspect influenced the distribution of naturally regenerated oak species within a relatively small area and resulted in differences in biomass and production among the stands.