Seasonal dynamics of arboreal spider diversity in a temperate forest

Measuring and estimating biodiversity patterns is a fundamental task of the scientist working to support conservation and inform management decisions. Most biodiversity studies in temperate regions were often carried out over a very short period of time (e.g., a single season) and it is often-at least tacitly-assumed that these short-term findings are representative of long-term general patterns. However, should the studied biodiversity pattern in fact contain significant temporal dynamics, perhaps leading to contradictory conclusions. Here, we studied the seasonal diversity dynamics of arboreal spider communities dwelling in 216 European beeches (Fagus sylvatica L.) to assess the spider community composition in the following seasons: two cold seasons (I: November 2005-January 2006; II: February-April) and two warm seasons (III: May-July; IV: August-October). We show that the usually measured diversity of the warm season community (IV: 58 estimated species) alone did not deliver a reliable image of the overall diversity present in these trees, and therefore, we recommend it should not be used for sampling protocols aimed at providing a full picture of a forest's biodiversity in the temperate zones. In particular, when the additional samplings of other seasons (I, II, III) were included, the estimated species richness nearly doubled (108). Community I possessed the lowest diversity and evenness due to the harsh winter conditions: this community was comprised of one dominant species together with several species low in abundance. Similarity was lowest (38.6%) between seasonal communities I and III, indicating a significant species turnover due to recolonization, so that community III had the highest diversity. Finally, using nonparametric estimators, we found that further sampling in late winter (February-April) is most needed to complete our inventory. Our study clearly demonstrates that seasonal dynamics of communities should be taken into account when studying biodiversity patterns of spiders, and probably forest arthropods in general.     

Soil seed bank composition and diversity in a managed temperate deciduous forest

Little is known about the influence of forest management on the interaction between seed bank and aboveground vegetation. We surveyed seed banks and vegetation in 10 forest stands under similar abiotic conditions but submitted either to a coppice-with-standards treatment (n=5) or to a selective-cutting system (n=5). We analyzed species composition and diversity, community ecological profile, and distribution of taxa among different life forms, strategy, morphology and functional type categories. A total of 2085 seedlings (8296 seedsm^–2) germinated-corresponding to 28 species, among which Juncus effusus was the most abundant. Fifty-seven percent of the species were also recorded in the aboveground vegetation, the dominant species being Rubus fruticosus agg., but only 28% of the aboveground species were present in the seed bank. Our results suggest that (1) vernal geophytes and shade-tolerant perennials, which group most true forest species, are not incorporated in the seed bank, (2) parent plants of most seeds were present either in the stand in an earlier dynamic stage or apart from the stand and long-distance dispersed, (3) as expected, early-successional species are well represented in the seed bank, (4) forestry vehicles seem to be a major means of dispersion for stress-tolerant species normally found in forest lanes and wheel tracks. We conclude that seed banks contain species that have a potentially negative impact on the true forest flora and, thus, forest management should minimize soil disturbance and retain remnants of old-coppice woods to conserve disturbance-sensitive true forest species.     

Plant species diversity in abandoned coppice forests in a temperate deciduous forest area of central Japan

We investigated plant species diversity as it related to stand structure and landscape parameters in abandoned coppice forests in a temperate, deciduous forest area of central Japan, where Fagus crenata was originally dominant. The species occurring in the study plots were classified into habitat types based on a statistical analysis of their occurrence bias in particular habitats (e.g., primary forest, coniferous plantation) in the landscape studied. The relationships between stand structure, which reflected the gradient of management, and forest floor plant species diversity (H and J) and richness (number of species per unit area) were not significant. However, these factors did influence the forest floor plant composition of the different types of habitat. According to the multiple regression analysis, species diversity and the richness of forest floor plants was affected by landscape parameters rather than by stand structure. For trees, species richness was mainly affected by the relative dominance of F. crenata, which is one of the stand structure parameters that decreases with intensive management. This is probably because many of the tree species that are characteristic of coppice forests increase after F. crenata have been eliminated by management; these species are not dominant in the original forest, where they are suppressed by F. crenata, the shade-tolerant dominant species. The species diversity (H and J) of trees was positively correlated with some landscape parameters, including the road density around the study plot, which may be associated with the intensity of management activity. The number of disturbance-tolerant species increased with increasing road density. Stand structure mainly affected disturbance-intolerant forest floor plant species and disturbance-tolerant tree species. Thus, the species diversity responses differed between forest floor plants and trees. The impact of forest management on species diversity was more prominent for forest floor plants.     

Comparison of lepidopteran larval communities among tree species in a temperate deciduous forest, Japan

Abstract.  1. The enormous diversity of phytophagous insects in forest canopies is hypothesised to be supported by the number of herbivorous species per host tree species or host specificity. It is therefore necessary to examine the effect of host plant species on compositional changes in the herbivore communities.
2. The lepidopteran larval communities were examined in the canopies of 10 tree species in a temperate deciduous forest of Japan. The phylogeny and leaf flush phenology of host plant species were taken into account as factors affecting the herbivore community assembly.
3. Examination of seasonal changes in the larval community structures on each tree species showed that larval species richness, abundance, and evenness decreased significantly from spring to summer. Larval species richness and abundance were characterised by family-level phylogenetic differences among tree species, whereas evenness was determined at a higher taxonomic level.
4. Compositional changes in the larval communities among tree species showed a remarkable pattern, with a phylogenetic effect at a high taxonomic level in spring, similar to evenness, but a phenological effect in summer. This suggests that host specificity could support the lepidopteran larval diversity in spring.
5. These results suggest that the differences in host utilisation of the herbivore, which reflects the phylogenetic effect of the host plants, can be important as a factor affecting the diversity of lepidopteran larval communities in temperate forests.     

Additive apportioning of lepidopteran and coleopteran species diversity across spatial and temporal scales in a cool–temperate deciduous forest in Japan

Abstract.  1. Forest entomofauna retain high diversity, and examining beta diversity, or species turnover, among assemblages in a forest community is vital to elucidate the source of this diversity.
2. Under the DIVERSITAS in Western Pacific and Asia–International Biodiversity Observation Year (DIWPA–IBOY) project for simultaneously documenting biodiversity throughout the Western Pacific and Asian Region, 892 lepidopteran species (51 742 specimens) and 355 coleopteran species (11 633 specimens) were collected in 2001 by light traps in a cool–temperate forest in northern Japan.
3. This study evaluated the beta diversity of lepidopteran and coleopteran communities by ecological categories (i.e. trap location, forest strata, sampling days, and months), and assessed the habitat preferences of lepidopteran and coleopteran species.
4.  anova -like additive apportioning models were used to quantify the beta diversity among the categories. The models simultaneously provide assessments of whether species distributions are biased in favour of particular habitats.
5. Significantly high beta diversity was observed among months for both Lepidoptera and Coleoptera. The category of months corresponded fairly well to the number of specialist species detected in the category, although a remarkably large number of significant specialists in Coleoptera were observed on strata.
6. The high beta diversity and number of specialist species among strata in both communities indicate that stratification between canopy and ground, and seasonal variation, played major roles in species composition and the rich entomofauna in the forest. Highly mobile adults were influenced by the vertical spatial scale, as previously suggested for larvae.     

Interactions between two co-dominant, invasive plants in the understory of a temperate deciduous forest

Negative interactions between non-indigenous and native species has been an important research topic of invasion biology. However, interactions between two or more invasive species may be as important in understanding biological invasions, but they have rarely been studied. In this paper, we describe three field experiments that investigated interactions between two non-indigenous plant species invasive in the eastern United States, Lonicera japonica (a perennial vine) and Microstegium vimineum (an annual grass). A press removal experiment conducted within a deciduous forest understory community indicated that M. vimineum was a superior competitor to L. japonica. We tested the hypothesis that the competitive success of M. vimineum was because it overgrew, and reduced light available to, L. japonica, by conducting a separate light gradient experiment within the same community. Shade cloth that simulated the M. vimineum canopy reduced the performance of L. japonica. In a third complementary experiment, we added experimental support hosts to test the hypothesis that the competitive ability of L. japonica is limited by support hosts, onto which L. japonica climbs to access light. We found that the abundance of climbing branches increased with the number of support hosts. Results of this experiment indicate that these two invasive species compete asymmetrically for resources, particularly light.     

Factors influencing sapling composition in canopy gaps of a temperate deciduous forest

To detect the factors that affect sapling species composition in gaps, we investigated 55 gaps in an old-growth temperate deciduous forest in Ogawa Forest Reserve, central Japan. Gap size, gap age, gap maker species, topographic location, adult tree composition around gaps, and saplings of tree species growing in the gaps were censused. For gaps 5 m², mean gap size was 70 m² and the maximum was 330 m². Estimated ages of gaps had a tendency to be concentrated in particular periods relating to strong wind records in the past. The sapling composition in gaps was highly and significantly correlated to that under closed canopy, indicating the importance of advance regeneration in this forest. However, some species showed significant occurrence biases in gaps or under closed canopy, suggesting differences in shade tolerance. The result of MANOVA showed that gap size and topography were important factors in determining the sapling composition in gaps. Species of gap makers affected the sapling composition indirectly by influencing gap size. The existence of parent trees around gaps had effects on sapling densities of several species. Gap age did not have clear influences on sapling composition. Variations in gap size and topography were considered as important factors that contribute to maintenance of species diversity in this forest.     

How tree species identity and diversity affect light transmittance to the understory in mature temperate forests

Light is a key resource for plant growth and is of particular importance in forest ecosystems, because of the strong vertical structure leading to successive light interception from canopy to forest floor. Tree species differ in the quantity and heterogeneity of light they transmit. We expect decreases in both the quantity and spatial heterogeneity of light transmittance in mixed stands relative to monocultures, due to complementarity effects and niche filling. We tested the degree to which tree species identity and diversity affected, via differences in tree and shrub cover, the spatiotemporal variation in light availability before, during, and after leaf expansion. Plots with different combinations of three tree species with contrasting light transmittance were selected to obtain a diversity gradient from monocultures to three species mixtures. Light transmittance to the forest floor was measured with hemispherical photography. Increased tree diversity led to increased canopy packing and decreased spatial light heterogeneity at the forest floor in all of the time periods. During leaf expansion, light transmittance did differ between the different tree species and timing of leaf expansion might thus be an important source of variation in light regimes for understory plant species. Although light transmittance at the canopy level after leaf expansion was not measured directly, it most likely differed between tree species and decreased in mixtures due to canopy packing. A complementary shrub layer led, however, to similar light levels at the forest floor in all species combinations in our plots. Synthesis. We find that a complementary shrub layer exploits the higher light availability in particular tree species combinations. Resources at the forest floor are thus ultimately determined by the combined effect of the tree and shrub layer. Mixing species led to less heterogeneity in the amount of light, reducing abiotic niche variability.     

Stand height and cover type complement forest age structure as a biodiversity indicator in boreal and northern temperate forest management

Forest age structure is one of the main indicators of biodiversity in temperate and boreal forests worldwide. This indicator was mainly chosen for the conservation of a subset of rare or sensitive species related to the oldest age classes, not to capture variability across the entire biodiversity spectrum, but is often considered as such. In this study, we analysed alpha and beta diversity in temporary plots of western Quebec, Canada, to consider biodiversity indicators complementary to existing forest age structure targets. Our analysis revealed that considered individually, stand characteristics such as cover type and height are better predictors of changes in site-level contribution to tree beta diversity than age. We also show that plots belonging to different age classes can be similar in terms of tree alpha diversity. Height class was found to have a more significant impact on tree alpha diversity than expected: height was more important than age in coniferous forests, and in deciduous and mixedwood stands it frequently complemented age in explaining the observed diversity patterns. Our results suggest that forest age structure target levels should not be used as the sole indicator of ecosystem sustainability, and that some mature secondary stands can provide significant contributions to biodiversity. We propose that more efficient trade-offs between forest exploitation, ecosystem functioning and environmental conservation can be attained if: (i) forest age structure targets are complemented by cover type and stand height; or (ii) complementary biodiversity indicators of ecosystem sustainability are implemented.     

森林冠层昆虫多样性研究方法

森林冠层是森林昆虫栖息、取食、避敌的重要生境,其中生活着十分丰富的昆虫物种。但由于乔木树体高大,冠层难于接近,在很大程度上限制了冠层昆虫多样性的研究。冠层昆虫类群结构的划分和冠层昆虫取样技术也是冠层昆虫多样性研究的重要基础。文章综述当前冠层接近和冠层昆虫抽样技术的最新进展,并评述冠层昆虫类群结构划分的方法。     

Winter photosynthesis by saplings of evergreen broad-leaved trees in a deciduous temperate forest

* Here we investigated photosynthetic traits of evergreen species under a deciduous canopy in a temperate forest and revealed the importance of CO2 assimilation during winter for annual CO2 assimilation. * Saplings were shaded by the canopy trees from spring through to autumn, but were less shaded during the winter months. Photosynthetic rates at light saturation (Aarea) were lower during winter than during the growing season. Aarea was higher in Camellia, Ilex and Photinia than in Castanopsis, Cleyera and Quercus during the winter, but differed little during summer and autumn. * Estimated daily CO2 assimilation (Aday) was higher during the winter than during the growing season in Camellia, Ilex and Photinia but was higher than that during the growing season only at the beginning and end of winter in Castanopsis, Cleyera and Quercus. Aday was higher in Camellia, Ilex and Photinia than in Castanopsis, Cleyera and Quercus but differed little among them during the growing season. * These results reveal the importance of winter CO2 assimilation for the growth of Camellia, Ilex and Photinia. Furthermore, differences in annual CO2 assimilation among species are strongly modified by species-specific photosynthetic traits during the winter under deciduous canopy trees.     

Impact of Prunus serotina invasion on understory functional diversity in a European temperate forest

We investigated the relationships between the overstory dominance of Prunus serotina, and the functional diversity of the understory plant communities, in a managed European forest. Vegetation, habitat characteristics and disturbance history were surveyed in 32 invaded stands vs. 32 paired uninvaded stands, after a random stratification. Community specialization and functional diversity indices were compared and a RLQ analysis was run to link species traits to environmental variables. The herb layer of invaded stands exhibited significantly more specialist species and a lower trait diversity compared to uninvaded stands, with respect of species richness and vegetation cover. Light arrival to the forest floor and soil properties explained most of the variation in the RLQ analysis, but 20% of the variation strongly correlated with P. serotina dominance and associated disturbances. Traits characterizing shade-tolerant, short-living ruderals and shade-avoiders (vernal geophytes) were significantly associated to invaded stands, while those associated to light-demanding graminoids characterized uninvaded stands. The establishment of functionally close species (seedlings and saplings of native woody species) tended to be lower in invaded stands. We concluded that the invader was becoming the new ecosystem engineer, first by inducing trait convergence and community specialization, thus promoting traits that enable species to capture resources in the new environment it was creating, and second by reducing the grain of local heterogeneities.     

Vascular diversity patterns of forest ecosystem before and after a 43-year interval under changing climate conditions in the Changbaishan Nature Reserve, northeastern China

The Changbaishan Nature Reserve (CNR) is the largest protected temperate forest in the world. It was established in 1960 to protect the virgin Korean pine (Pinus koraiensis) mixed hardwood forest, a typical temperate forest of northeast China. Studies of vascular diversity patterns on the north slope of the CNR mountainside forest (800–1700 m a.s.l.) were conducted in 1963 and in 2006. The aim of this comparison was to assess the long-term effects of the protected status on plant biodiversity during the intervening 43 years. The research was carried out in three forest types: mixed coniferous and broad-leaved forest (MCBF), mixed coniferous forest (MCF), and sub-alpine coniferous forest (SCF), characterized by different dominant species. The alpha diversity indicted by species richness and the Shannon–Wiener index were found to differ for the same elevations and forest types after the 43-year interval, while the beta diversity indicated by the Cody index depicted the altitudinal patterns of plant species gain and loss. The floral compositional pattern and the diversity of vascular species were generally similar along altitudinal gradients before and after the 43-year interval, but some substantial changes were evident with the altitude gradient. In the tree layer, the dominant species in 2006 were similar to those of 1963, though diversity declined with altitude. The indices in the three forest types did not differ significantly between 1963 and 2006, and these values even increased in the MCBF and MCF. However, originally dominant species, such as Pinus koraiensis, tended to decline, the proportion of broad-leaved trees increased, and the species turnover in the succession layers showed a trend to shift to higher altitudes. The diversity pattern of the understory fluctuated along the altitudinal gradient due to micro-environmental variations. A comparison of the alpha diversity indices among the three forest types reveals that the diversity of the shrub and herb layer decreased, and some rare and medicinal species disappeared. Meteorological records show that climate has changed significantly in this 43-year intervening period, and information collected from another field survey found that the most severe human disturbances to the CNR forests stemmed from the exploitation of Ginseng roots and Korean pine nuts. W. Sang and F. Bai are contributed equally to this research.     

Structure,diversity, and regeneration of tropical dry deciduous forest of northern India

This study examined the impact of disturbance on the pattern of diversity, forest structure and regeneration of tree species in the Vindhyan dry tropical forest of India. A total of 1500 quadrats distributed over five, 3-ha permanent plots in five sites, differing in degree of disturbance, were used to enumerate and measure the tree species. A total of 65 species with 136,983 individuals were enumerated in the total 15-ha area for stems 30 cm height. The number of species and number of stems ranged from 12 to 50 and 8063–65331 per 3-ha area. The number of species and stems for trees 10 cm dbh ranged from 3 to 28 species, with a mean value of 16 species ha^–1, and from 16 to 477 stems, with a mean value of 256 stems ha^–1, respectively. The adult based PCA ordination indicated uniqueness of sites in terms of species composition and habitat characteristics. PCA ordination also showed uniqueness of sites in terms of seedling composition, but the seedling and adult distributions were not spatially associated. The distinct species composition at the different sites and at the two life-cycle stages on the same site is indicative of marked spatio-temporal dynamics of the dry tropical forest. The density–diameter semi-logarithmic curves ranged from a near linear to an overall concave appearance with a limited plateau in the mid-diameter ranges. The -diversity and its components decreased with increasing disturbance intensity, reflecting enhanced utilization pressure with increasing disturbance. The site-wise and species-wise regression analyses of the number of individuals in different stages of the species revealed that both the level of disturbance and the nature of species strongly affect the regeneration. In conclusion, although the forest is relatively species-poor, the differential species composition on different sites and the temporal dynamics lend a unique level of diversity to the tropical dry deciduous forest.     

Seed demography of three co-occurring Acer species in a Japanese temperate deciduous forest

Abstract. Seed demography of three co-occurring Acer species in an old-growth mixed deciduous forest in Japan was studied. Almost all of the seeds of A. mono germinated in the first spring, while those of A. palmatum var. amoenum showed a delay in germination of almost one year. A. rufinerve showed a rather opportunistic germination habit. Both A. palmatum var. amoenum and A. rufinerve form short-term persistent seed banks, but without input of newly dispersed seeds they may become extinct in about one year. The seed bank for these two species is not as significant as for a typical pioneer species, and the seedling bank is important for all three species. Only a small proportion of the dispersed propagules contained viable embryos, mainly due to pollination failure or abortion (A. mono and A. palmatum var. amoenum), and invertebrate predation (A. rufinerve). For all three species, larger seed crops had a higher percentage of viable seeds. Even for these relatively small, wind-dispersed seeds, the predation pressure was very high. A large part of the dispersed seeds was eaten by wood mice during the first winter (30–80 %). Estimation from the 5-yr average of seed dispersal and seedling emergence showed that only 7–16 % of the dispersed viable seeds succeeded in germinating.     

Short-term responses of the understory to the removal of plant functional groups in the cold-temperate deciduous forest

It is widely believed that functional diversity contributes to the stability of ecosystems. Indeed, greater redundancy among species within functional groups and greater complementarity among functional groups within communities should increase the resistance and resilience of ecosystems. In the present study, we tested for functional group complementarity by examining how the loss of specific functional groups may alter the role that other groups play in ecosystem functions. We removed different functional groups, one at a time, from the understory of three maple-dominated forests in southern Québec (Canada) and followed the understory response over a 2-year period. The experimental design included a control and five removal treatments. Five functional groups were defined: spring-flowering ephemeral species; spring-flowering persistent species; summer-flowering species; fern species; and seedlings and juveniles of woody species. Richness, cover, soil pH and organic matter content were determined after two years of removal. The results of our experiment revealed that richness was significantly lower than what we expected when spring-flowering persistent species or seedlings and juveniles of woody species were removed, suggesting not only direct but also indirect positive effects of both of these groups on understory richness (mostly through effects on summer-flowering species and fern species). Removal of the seedlings and juveniles of woody species and, to a lesser extent, of spring-flowering persistent species and of fern species lead to a decrease in the cover of summer-flowering species, implying a positive effect of the former groups on the latter. The cover–richness relationship in the control and in each one of the five removal treatments was positive and well fitted by a linear regression. Yet, the slope of the relationship differed among treatments, but not between the control and any one of the removal treatments (pair-wise comparisons). Our results suggest that the different functional groups are complementary and that positive interactions predominate over negative ones. Contrary to common belief, understory plants can respond quite rapidly to changes in community functional composition. Although we have not investigated the specific mechanisms responsible for the short-term responses reported here, we suggest that complex intergroup interactions may favour functional diversity and enhance ecosystem functions.