Fluctuating ungulate density shapes tree recruitment in natural stands of the Białowieża Primeval Forest,Poland

Question: What are the main driving factors in 70 years of natural dynamics in tree recruitment in the Bia?owie?a National Park? Location: Bia?owie?a National Park, Poland, is one of the least disturbed temperate, lowland forest systems in Europe. Methods: We tested whether fluctuations in large herbivore populations, changes in climate and openness of the forest explained compositional dynamics. Tree recruitment (to size class DBH≥5 cm) was measured on permanent transects (in total, 14.9 ha) six times between 1936‐2002. These data were related to existing data on ungulate density, climatic parameters and estimates of forest openness collected during the same period. Results: Total recruitment of all tree species combined was negatively correlated with total ungulate density and red deer density. The variation in response between species was related to the preferences of herbivores; the more preferred forage species (especially Carpinus betulus) were positively and the less preferred species negatively related to herbivore density. Total tree recruitment rates were not related to climatic parameters and openness of the forest. Only Alnus glutinosa recruitment was significantly related to climatic parameters, and Ulmus glabra related to forest openness, but there were no predictable patterns in recruitment among species in relation to these factors. Conclusion: The present study indicated that changes in large herbivore density have played an important role in driving patterns in tree recruitment and species composition during the last 70 years in Bia?owie?a National Park. In contrast to other studies, increasing herbivore numbers were associated with higher recruitment of preferred and browsing‐tolerant species. Periodical crashes in ungulate numbers, whether human‐induced or caused by natural factors, may offer windows of opportunity for regeneration of a range of tree species and facilitate more diverse and dynamic forest development.     

Millipede (Diplopoda) communities in an arboretum: Influence of tree species and soil properties

The paper deals with the influence of tree species on millipede communities (Diplopoda). The research was carried out in nine sites in the Borová hora arboretum (Zvolen town, Central Slovakia). Each studied site represents a monoculture of one of nine tree species: Betula pubescens Ehrh., Pinus sylvestris L., Larix decidua Mill., Carpinus betulus L., Abies alba Mill., Picea abies (L.) Karst., Alnus incana (L.) Moench, Populus nigra L., Ulmus laevis Pall. Millipedes were collected by pitfall trapping during vegetation periods in 2008-2011. In total, 1064 individuals of 17 species and 7 families were obtained. The results of research confirmed (i) an influence of tree species on the composition of millipede communities, (ii) a significant influence of soil nitrogen on the species richness and biodiversity, and (iii) an impact of soil pH on the species composition of these terrestrial invertebrate communities. In terms of total dynamic activity and species richness of millipedes, the most favourable conditions were revealed in the forest stands of Alnus incana, Populus nigra, Ulmus laevis and Carpinus betulus. On the contrary, the least favourable biotopes were (from both points of view) the forest stands of Betula pubescens and Larix decidua.     

Compositional dynamics of natural forests in the Bialowieza National Park,northeastern Poland

Abstract. This paper presents results of a long-term study on natural forest dynamics in the Bia?owieza National Park (BNP), northeastern Poland. Five permanent sample areas were used, each consisting of a transect of varying width (40 - 60 m) and length (200 - 1380 m). The total sample area is 14.9 ha. The study covers the period 1936–1992. During this period measurements were made on five occasions at approximately 10-yr intervals. On each measurement date all trees with DBH > 5 cm were identified and their spatial location, diameter, crown condition and position in the canopy determined. During the study period the stands underwent noticeable changes, mainly in terms of tree species composition. The major change was a quantitative increase of the late-successional species: Tilia cordata and Carpinus betulus, also to a lesser degree Fraxinus excelsior and, in the last period, of the early successional Alnus glutinosa. Declining species included both early- and late-succession species. Among the latter group, Picea abies ranked first. This species lost much of its importance during the last few decades. P. abies was followed by Pinus sylvestris which is an important component of the climax vegetation under the conditions prevailing in Bialowieza, at least on more oligotrophic sites. Still, this species has not been able to regenerate during the whole study period. Some other late-succession species, Acer platanoides and Quercus robur, were also amongst the declining species. Although the basal area of Q. robur increased, its population was getting older and the process of natural regeneration was markedly impeded. All typical pioneer, short-lived species: Betula pendula and B. pubescens, Salix caprea and Populus tremula also decreased, which was probably caused by a lack of major disturbances during the study period. In general, the results obtained for the semi-natural conditions of Bialowieza during the 56-yr observation period suggest a rather high compositional instability of the forest stands there. A more precise identification of the role of particular factors in the observed stand dynamics is difficult because of the paucity of appropriate historical and environmental data which refer directly to the study plots; moreover, the data are generally incompatible and of different resolution.     

Decline ofPotentillo albae-Quercetum phytocoenoses associated with the invasion ofCarpinus betulus

The study area in the Bialowieza primeval forest-size 28000 m², divided into quadrats of 100 m², comprised two patches ofPotentillo albae-Quercetum as distinguished by phytosociological methods, and a fragment of an adjacent very largeTilio-Carpinetum stand. The directional variation in the distribution ofCarpinus betulus trunk diameter values in the southern part of the study area indicates invasion of this oak forest byCarpinus. This process probably started 1962–1968. The invasion ofCarpinus betulus caused considerable deterioration in light conditions of the ground layer, as deduced from a decrease in Ellenberg light index values. The phytosociological division of the study area correlates with variation in the spatial pattern ofCarpinus mean trunk diameter distribution.Quadrat groups were distinguished by association analysis. They were described phytosociologically using systematic group values for species, constituting the characteristic combination for thePotentillo albae-Quercetum. Areas occupied by still olderCarpinus individuals correspond to successive stages of oak forest regression. The deterioration of light conditions caused recession of thermo-and heliophilous oak-forest species and decline ofPotentillo albae-Quercetum phytocoenoses. However, this was not accompanied by the expansion pansion of species ofCarpinus forests.     

A comparison of symbiotic nitrogen fixation in Scotland inAlnus glutinosa andAlnus rubra

Summary Alnus glutinosa andAlnus rubra growing in the field in Scotland show specific nitrogenase activities of the same order of magnitude. The period of maximum potential nitrogenase activity coincides with that of maximum growth in late Spring and Summer. It is suggested that the retention of nitrogenase activity into the Autumn when growth has virtually ceased may be important as a contribution to the nitrogenous reserves of the tree.Bioassay of different Scottish soils, all collected from the locality of natural stands ofAlnus glutinosa, showed wide variation in the nodulation of seedlings, although generally a soil poor for nodulation ofAlnus glutinosa generally gave poor nodulation ofAlnus rubra. Soils of pH 4.5 to 6.5, best suited for growth and nitrogen fixation of the two species, often gave nodules showing highest specific nitrogen fixing activity. Young (2 to 3 year old) plants in glasshouse or controlled environment cabinet, inoculated withAlnus glutinosa endophyte, differed from mature field grown plants, however, sinceAlnus rubra required a much larger (up to 2.5 times) mass of root nodules to fix a unit quantity of N. Microscopic comparison of the nodules of glasshouse plants showed that the proportion of cells containing the vesicular (nitrogen fixing) form of the endophyte was only slightly lower inAlnus rubra than inAlnus glutinosa and it is suggested that the differences in specific nitrogen fixing activity between the two species may reflect some incompatibility of function of theAlnus glutinosa endophyte when in symbiosis withAlnus rubra.     

Study of the contribution of the shoot and/or root ofAlnus sp. in the compatibility between the host and a Sp+ Frankia strain using a grafting technique

Two alder species,Alnus glutinosa (L.) Gaertn. andAlnus incana (L) Moench, were inoculated with a Sp⁺ Frankia homogenate obtained fromA. incana root nodules. This inoculum formed effective nodules on the original host plant and ineffective nodules onA. glutinosa. Grafts between the two alder species were made to determine which part of the plant is involved in this phenomenon. The results obtained indicate that the compatibility between Alnus andFrankia is restricted to the root system.     

Methods of isolation and cultivation of Frankia species from actinorhizas

A critical review is given about the isolation and cultivation methods of Frankia species fromAlnus glutinosa root-nodules. The best results so far are obtained with a combination of sucrose (60% w/v)-sedimentation of root-nodule homogenate and subsequent suspension in the top-layer of a doubleagar layer system. The top-layer needs to contain a suitable C-source, in this study often a lipid factor from an alcoholic root-extract and an organic N-source.The isolation and cultivation of Sp(–) and Sp(+) strains fromAlnus glutinosa root nodules and a Frankia from the root-nodules ofMyrica gale is reported. The regular observation of growing colonies appears to be very important for the interpretation of results. The latter was illustrated by the remarkable diauxic growth of the strains isolated fromAlnus glutinosa Sp(+) root nodules.     

Comparative studies of ectomycorrhiza formation in Alnus glutinosa and Pinus resinosa with Paxillus involutus

 Mycorrhiza ontogeny and details of Hartig net and mantle structure were compared in ectomycorrhizas synthesized in growth pouches between the broad host range fungus Paxillus involutus and the tree species European black alder (Alnus glutinosa) and red pine (Pinus resinosa). In Alnus glutinosa, a paraepidermal Hartig net was restricted to the proximal (basal) portion of first-order laterals; the hypodermal layer appeared to be a barrier to fungal penetration. Phi-thickenings were present in some cortical cells but these were not related to lack of fungal ingress into the cortex. The mantle was often present close to the root apex but in many roots it was loosely organized and patchy. In several instances, the mantle formed around the root apex was only temporary; renewed root growth occurred without the formation of a mantle. In Pinus resinosa, the Hartig net developed between cortical cell layers of monopodial and dichotomously branched first–order laterals. Fungal hyphae in the Hartig net exhibited a complex labyrinthine mode of growth. The mantle had a pseudoparenchymatous structure and covered the root, including apices of dichotomously branched roots. The Paxillus–Pinus resinosa interaction had all the characteristics of a compatible ectomycorrhizal association. The Paxillus–Alnus glutinosa interaction, however, showed only aspects of superficial ectomycorrhizas, including the presence of a minimal (sometimes absent) and mostly proximal Hartig net and variable mantle development. Sclerotia were produced in the extraradical mycelium of Paxillus involutus when associated with either Alnus glutinosa or Pinus resinosa. Accepted: 22 October 1998     

Low modularity and specialization in a commensalistic epiphyte–phorophyte network in a tropical cloud forest

Species interactions can shape the structure of natural communities. Such sets of interactions have been described as complex ecological networks, an example of which is the commensal network formed by epiphyte–phorophyte interactions. Vascular epiphytes germinate and grow on phorophytes (support trees), assuming a horizontal distribution (among the phorophyte species) and a vertical distribution (from the base of the tree trunk to the crown of phorophytes, i.e., through ecological zones). Here, we investigated the organization of these structural dimensions of the epiphyte–phorophyte network in a Brazilian tropical montane cloud forest. The analyzed network, comprising 66 epiphyte species and 22 phorophyte species, exhibited a nested structure with a low degree of specialization, a typical pattern for epiphyte–phorophyte networks in forests. The network was slightly modular, with 65% of the species common to three modules, and had vertical structure corresponding to the vertical organization of the phorophytes. The size (diameter at breast height) of phorophyte individuals influenced the network structure, possibly due to the increase in habitat area, the time available for colonization by epiphytes, and a greater number of microenvironments. We found that the distribution of the epiphyte species differed between the phorophyte ecological zones, with greater richness in the lower portions and greater abundance in the upper portions of the phorophytes. The results provide relevant guidance for future research on the characteristics and the vertical and horizontal organization of vascular epiphyte and phorophyte networks. Abstract in Portuguese is available with online material.     

Evolutionary implications of nucleotide sequence relatedness between Alnus nepalensis and Alnus glutinosa and also between corresponding Frankia microsymbionts

Frankia DNAs were isolated directly from root nodules of Alnus nepalensis and Alnus nitida collected from various natural sites in India. For comparison, a nodule sample from Alnus glutinosa was also collected from Tuebingen, Germany. Nucleotide sequence analyses of amplified 16S–23S ITS region revealed that one of the microsymbionts from Alnus nepalensis was closely related to the microsymbiont from Alnus glutinosa. A similar exercise on the host was also carried out. It was found that one sample of Alnus nepalensis was closely related to Alnus glutinosa sequence from Europe. Since both Frankia and the host sequences studied revealed proximity between Alnus glutinosa and Alnus nepalensis, it is hypothesised that the common progenitor of all the alders first entered into an association with Frankia, and the symbiotic association has evolved since.     

Studies on the Holocene history of vegetation in the region of lake Srebarna (northeast Bulgaria)

The results of pollen analysis of two sediment cores from lake Srebarna (northeast Bulgaria) are presented. On the basis of the palynological data and the radiocarbon dates a reconstruction of the past flora and vegetation is made. For the first time a continuous palaeosuccession is established for the area of the Danubian Dobrudza and the Ludogorie district of northeast Bulgaria: a dominance of mixed xero- and mesophilous oak woods with Carpinus betulus, Ulmus, Tilia, Corylus during the Atlantic period; mixed oak woods with increasing importance of Tilia, Ulmus, Acer and a considerable presence of Carpinus betulus and Fagus during the Sub-boreal period; degradation of the woodland vegetation and transition to secondary plant communities with Carpinus orientalis and herbs during the Sub-atlantic period. Received January 5, 2001 / Accepted March 13, 2001     

干旱胁迫对欧洲鹅耳枥幼苗生理生化特征的影响

以一年生欧洲鹅耳枥幼苗为材料,通过盆栽控水试验,设置5个土壤含水量梯度,研究不同梯度干旱胁迫对其幼苗生理生化指标的影响,以明确欧洲鹅耳枥的耐旱特性。结果显示:（1）随着干旱胁迫的加剧,欧洲鹅耳枥幼苗受到损害程度逐渐加重,形态表现出明显的缺水特征,叶片相对含水量和叶绿素总量随着干旱胁迫程度的增加而下降,且胁迫强度越强、时间越久其变化幅度越大。（2）幼苗叶片SOD活性、POD活性、可溶性糖含量和可溶性蛋白含量均随着干旱程度的增加表现出先升高后降低的趋势。（3）随着干旱胁迫程度的增加和干旱时间持续,幼苗叶片MDA含量和相对电导率总体上呈增大趋势,并在干旱末期达到最大值。研究表明,欧洲鹅耳枥幼苗在短期和轻度（田间最大持水量的60%）、中度（田间最大持水量的45%）干旱胁迫下能通过调节自身保护酶活性和渗透调节物质含量来减轻干旱造成的危害;而在重度（田间最大持水量的30%）干旱胁迫下,幼苗自我调节能力受影响,极度（田间最大持水量的15%）干旱胁迫对其造成一定的损害。研究认为,欧洲鹅耳枥具有一定的抗干旱特性,可在园林中推广运用。     

影响广东黑石顶树附生苔藓分布的环境因子

通过对广东省黑石顶自然保护区内5个2500 m2样地内树附生苔藓的调查及有关环境因子的测定,研究了树附生苔藓的分布格局及其与环境因子的关系.树附生苔藓在不同高度的分布存在一定梯度,20 cm高处树附生苔藓的种类数与盖度均大于60 cm及更高处,且其群落优势种的数量组成与后者存在较大差异.不同树种附生苔藓盖度和种数差异较大.基于附生苔藓植物的盖度进行DCA排序及聚类分析将树种分成四组,马尾松(Pinus massoniana)因具有两种特有的网藓(巴西网藓Syrrhopodon prolifer和鞘刺网藓S.armatus)单独一组,福建青冈(Quercus chungii)亦与其余各种的差异均较大,形成一组,其余阔叶树种根据其所处的森林类型分成两组,针阔叶混交林内的阔叶树种和次生阔叶林内的阔叶树种各形成一组.对环境因子及树皮含水量和pH的分析显示,垂直梯度上空气湿度的差异可能是造成附生苔藓在不同高度分布差异的主要影响因子之一,不同树种附生苔藓的差异在一定程度上受树皮pH的影响,而与树皮含水量无关.同一树种上树附生苔藓的分布又在一定程度上受森林类型的影响.     

Epiphyte differentiation between Quercus petraea and Fraxinus excelsior trees in a maritime area of South West England

A survey was undertaken in S.W. England to study epiphyte differentiation between free-standing oak (Quercus petraea) and ash (Fraxinus excelsior) trunks. Epiphyte cover was measured on 23 trees in 87 15×15 cm quadrats positioned on the north, east, south and west faces of each trunk at a height of 1.52 m. Measurements of 20 environmental factors were made at each quadrat position. Analysis of the vegetation data by reciprocal averaging showed that floristic variation is continuous and corresponds only partially to phorophyte identity. Rank correlation coefficients were calculated between the floristic axis obtained by reciprocal averaging, and individual environmental factors including a variate, obtained by principal components analysis of the environmental data, which expressed the major differences between the oak and ash environments. The most significant correlation was obtained with bark pH. Values of bark pH for oak and ash samples overlapped considerably although the mean for ash (5.1) is higher than for oak (4.7). It is concluded that the epiphytic flora is responding to variations in bark acidity or some closely related factor and not to other, closely phorophyte dependent nutritional factors. Factors influencing bark acidity in oak and ash are briefly discussed.     

Phorophyte specificity and environmental parameters versus stochasticity as determinants for species composition of corticolous crustose lichen communities in the Atlantic rain forest of northeastern Brazil

A transect of 47 mature trees was studied within an Atlantic rain-forest plot in northeastern Brazil to determinate effects of phorophyte specificity and environmental parameters vs. stochasticity on the structure of corticolous, crustose microlichen communities. A total of 150 lichen species was found, most being rare to extremely rare. Multivariate analysis of sample plots indicated subtle phorophyte preferences among certain lichen species, corresponding to differences in bark pH, degree of bark shedding, density and size of bark lenticels, and presence of milk sap. Individual and multiple regressions revealed correlations between lichen species richness; respectively, area cover and bark pH (negative); density and size of bark lenticels (negative); degree of bark shedding (negative); presence of milk sap (positive); and diffuse site factor (positive). No strongly delimited lichen communities were detected, but cluster analysis revealed three main groups and six subgroups with slightly different lichen species composition, each one with characteristic indicator species but with highly variable overall species composition. Beta diversity was high among samples and lacked spatial structure. However, beta diversity was significantly lower among samples belonging to the same tree species, independent of their spatial arrangement. It was concluded that community formation in tropical rain-forest understory lichens subtly correlates with two main environmental factor complexes—phorophyte bark characteristics and microclimate—but is to a large extent determined by the stochastic effects of species dispersal, especially of rare species.     

Competition between Quercus petraea and Carpinus betulus in an ancient wood in England: seedling survivorship

Abstract. Age structures of populations of canopy trees in Wormley Wood are consistent with reports that successional change is occurring, with Carpinus betulus replacing Quercus petraea as the dominant species. TWINSPAN analysis of data from a vegetation survey identifies three communities (described in earlier work), the ‘Bracken’, ‘Bramble’, and‘Bareground’ societies, characterized by increasing prominence of C. betulus and loss of species diversity. An experiment was set up in which C. betulus, Q. petraea and Betula pendula seedlings were explanted into each of the communities. Survival of seedlings was monitored over 860 days and differences investigated. Cohort survivorship differed between species and sites. C. betulus seedlings survived longer than Q. petraea in Bracken and Bramble communities. Species- and site-specific variation in the types and effects of herbivory were found. Herbivory did not appear to be a critical factor in the survival of Q. petraea seedlings. The photosynthetic light response of the seedling species was measured in the field. The light compensation point for Q. petraea seedlings (77 μmol photon m^-2 s^-1) is higher than the maximum available light under the main tree canopies in the wood. In contrast, seedlings of C. betulus have a lower light compensation point (15 μmol photon m² s^-1) and the mature tree casts a deeper shade than Q. petraea. It is suggested that the invasion of the canopy by C. betulus, following the cessation of coppicing, is creating light levels too low for Q. petraea seedling banks to persist.     

Oxygen Diffusion from the Roots of Woody Species

Application of the‘polarographic’ technique for studying oxygen diffusion from roots has provided preliminary results which establish that oxygen passes through the woody species Salix atrocinerea Brot., Salix fragilis L., Salix repens L., and Myrica gale L., in the gaseous phase as it does in other wetland species. Entry into the shoots occurs through the bark directly above the water table and in the willow cuttings the effective length of shoot for gas intake was the basal three centimetres above the water table. The length of shoot involved was longer in Myrica gale and the roots were of the normal (un-nodulated) type. Trials on Alnus glutinosa (L.) Gaertn. provided no conclusive results and it is thought that this was perhaps because measurements were on nodulated roots only.     

Species deletion in Potentillo albae-Quercetum phytocoenoses reversed by the removal of Carpinus betulus

The studies were conducted in the Biaowiea primeval forest on a study area of 28,000 m², divided into two plots and 280 quadrats. In 1979, all Carpinus betulus individuals were cut off within the experimental plot (E, 7800 m²), whilst the control plot (C, 13000 m²) remained unchanged. Each plot contained a phytocoenose of Potentillo albae-Quercetum and an adjacent community of Tilio-Carpinetum.By 1983, hornbeam invasion into the oak-forest habitat within C plot had resulted in: 1) a decrease in species number by more than 10 per 100 m² on average (c.a. 30%); 2) a twofold greater deletion rate of heliophil oak-forest species than for Fagetalia and Querco-Fagetea; 3) a diminution of the area of oak-forest phytocoenose by more than 100 m² per year; 4) formation of a community with a species combination corresponding to Tilio-Carpinetum. Hornbeam removal had the opposite effect: 1) the number of species increased by more than 100%; 2) the number and frequency of oak forest species rose considerably; 3) those parts of the phytocoenose colonized previously by Carpinus betulus regenerated, and thence the area of Potentillo albae-Quercetum community increased.     

Tree and stand-scale factors affecting richness and composition of epiphytic bryophytes and lichens in deciduous woodland key habitats

Conservation and sustainable forestry are essential in a multi-functional landscape. In this respect, ecological studies on epiphytes are needed to determine abiotic and biotic factors associated with high diversity. The aim of the present study was to evaluate relative sensitivity of conservation targets (epiphytic bryophytes and lichens) in relation to contrasting environmental variables (tree species, tree diameter at breast height, bark crevice depth, pH, tree inclination, pH, forest stand age, area and type) in boreo-nemoral forests. The study was conducted in Latvian 34 woodland key habitat (WKH) boreo-nemoral forest stands. Generalized linear mixed models and canonical correspondence analysis showed that tree species and tree bark pH were the most important variables explaining epiphytic bryophyte and lichen composition and richness (total, Red-listed, WKH indicator species). Forest stand level factors, such as stand size and habitat type, had only minor influence on epiphytic species composition and richness. The results of the present study indicate a need to maintain the diversity of tree species and large trees, particularly Acer platanoides, Carpinus betulus, Fraxinus excelsior, Populus tremula, Tilia cordata, Ulmus glabra and Ulmus laevis in conservation of epiphytic bryophyte and lichen communities in the future.     

Effects of some pure and mixedFrankia strains on seedling growth in differentAlnus species

In greenhouse experiments plants of eightAlnus species, from various parts of the world, and from different taxonomic sections, were inoculated with threeFrankia strains in order to show any possible interaction. Mixtures in equal parts of theseFrankia strains were also tried. The growth of inoculated plants was significantly higher than of the controls, with one of the three strains being superior. Mixtures of strains generally provided higher growth than the best individual strain. No interaction betweenFrankia strains andAlnus species was detected in the young plants 60 days after inoculation. Three clones ofAlnus glutinosa were inoculated with the same pure cultures ofFrankia, without producing any interaction. Inoculation time was studied in one clone and one progeny ofAlnus glutinosa. The best results were obtained with the earlier inoculation (at sowing for the progeny and at transfer to soil for thein vitro-propagated clone). The results are discussed in terms of nursery practice and field experiments for selection in breeding programmes.