Spatial distribution patterns of two predominant emergent trees in a tropical rainforest in Sarawak,Malaysia

Spatial distribution patterns of two emergent tropical rainforest tree species (Dryobalanops aromatica & D. lanceolata) were examined in where they were dominant (17–20% of total basal area of canopy trees) in Sarawak, East Malaysia. Newly established seedlings (< 2 years old) were restricted to areas < 40 m from mother trees for both species, suggesting a limited seed dispersal. Seedling (< 1 cm in dbh) density was highest around conspecific adults ( 30 cm in dbh). Negative spatial patterns were observed between larger juveniles (1–5 cm in dbh) and conspecific adults for both species; the most dense populations of sapling (1–5 cm in dbh) and poles (5–30 cm in dbh) were found at a distance of 15–20 m from the nearest conspecific adult. Seedlings of both species were distributed randomly with respect to light conditions evaluated by a forest floor diffuse site factor and a canopy closure index. Saplings of both species, and poles of D. lanceolata, were distributed under more open conditions than expected from spatially random distributions, and from average light conditions of all species of the same size classes. Possible mechanisms for the observed distribution patterns and intermediate canopy dominance of Dryobalanops were discussed from the viewpoints of gap-dynamics and distance-dependent mortality.     

Hedgerows as habitat corridors for forest herbs in central New York, USA

1 Samples from 32 hedgerows ( c.  6 m wide, with full-grown trees) in central New York included 39 forest herb taxa, comprising nearly 70% of the forest herb taxa found in adjacent forest samples.
2 We sampled three types of hedgerow. Two types were attached to forest: remnant hedgerows ( n  = 14), and regenerated hedgerows ( n  =  11 ) that had grown up spontaneously between open fields in the last 50 years. There were no significant differences between remnant and regenerated hedgerows in the richness or abundance of forest herbs, presumably indicating colonization of regenerated hedgerows. Such colonization implies that hedgerows serve a corridor function.
3 The species composition of forest herbs in hedgerows attached to forest stands showed a strong affinity with that of the adjacent stand, both for remnant and regenerated hedgerows.
4 There was a distance effect within hedgerows. Richness of forest herbs and similarity of composition to forest declined with distance along the hedgerow from forest, implying colonization from the adjacent attached stand.
5 The third type of hedgerow sampled, isolated remnants ( n  = 7), was not lower in richness or abundance of forest herbs than hedgerows attached to forest.     

Distribution patterns of tree species in an evergreen broadleaved forest in eastern China

Ecological assembly rules in evergreen broad-leaved forest are far from clear understanding. Spatial dispersion of individuals in a species is central in ecological theory. We analyzed the spatial patterns as well as associations between adult and juvenile of each tree species in a 5-ha subtropical evergreen broad-leaved forest plot in eastern China. Out of the 74 species occurring with more than 10 individuals, 88.4% of these species are aggregated. Most of them are aggregated from small to large scales. Spatial distributions of some species corre-spond with topography. Many bad dispersed species in the Baishanzu exhibit a highly aggregated distribution at small scales. These suggest that environmental heterogeneity and/or dispersal limitation may be the most important mechanisms that control the distribution patterns of these species. Our observations of the aggregations of abundant species basically support the hypothesis that dispersal limitation decreases as the number of reproductive trees increases. The rest species are randomly distributed, with less than 10 individuals. For most common species, spatial aggregation is weaker in larger diameter classes, and the distance between adults is larger than that between juveniles and adults, suggesting that density-dependence works on loosing aggregation and excluding conspecific juveniles away from adult trees. However, the density-dependent effect is not strong enough to eliminate all seedlings near adult trees and to result in a regular distribution of trees; thus the density dependence is usually masked by the refuge effect.     

Distribution patterns of tree species in an evergreen broadleaved forest in eastern China

Ecological assembly rules in evergreen broad-leaved forest are far from clear understanding. Spatial dispersion of individuals in a species is central in ecological theory. We analyzed the spatial patterns as well as associations between adult and juvenile of each tree species in a 5-ha subtropical evergreen broad-leaved forest plot in eastern China. Out of the 74 species occurring with more than 10 individuals, 88.4% of these species are aggregated. Most of them are aggregated from small to large scales. Spatial distributions of some species correspond with topography. Many bad dispersed species in the Baishanzu exhibit a highly aggregated distribution at small scales. These suggest that environmental heterogeneity and/or dispersal limitation may be the most important mechanisms that control the distribution patterns of these species. Our observations of the aggregations of abundant species basically support the hypothesis that dispersal limitation decreases as the number of reproductive trees increases. The rest species are randomly distributed, with less than 10 individuals. For most common species, spatial aggregation is weaker in larger diameter classes, and the distance between adults is larger than that between juveniles and adults, suggesting that density-dependence works on loosing aggregation and excluding conspecific juveniles away from adult trees. However, the density-dependent effect is not strong enough to eliminate all seedlings near adult trees and to result in a regular distribution of trees; thus the density dependence is usually masked by the refuge effect.     

Density dependence is prevalent in a heterogeneous subtropical forest

Although negative conspecific density dependence among neighbours is widely studied, the general prevalence of the effects is still poorly understood due to a lack of studies from zonal forests other than the tropics. In addition, the detection of density dependence may be confounded by the influence of habitat heterogeneity. Here we examined the spatial distributions of 47 common tree species (diameter at breast height≥1  cm) using the pair-correlation function g(r) in a fully mapped 24-ha subtropical forest in China. We first investigated whether habitat heterogeneity influenced tree distributions, and then examined the conspecific tree patterns and density dependence after removing the effects of habitat heterogeneity. We found that the forest plot exhibited strong large-scale heterogeneity in the distribution of both large adult trees of different growth forms and individual species. After the habitat heterogeneity was accounted for, 39 of the 47 species (83.0%) were found to exhibit density dependence predominantly at close distances among neighbors. Our findings highlight density dependence as a prevalent mechanism for regulating the population spatial structure of most tree species in the species-rich subtropical forest studied here. Furthermore, the occurrence of density dependence is closely associated with species abundance and the strength of conspecific aggregation at local scales. Abundant species with high strength of conspecific aggregation tend to show density dependence.     

Evolutionary history and distance dependence control survival of dipterocarp seedlings

One important hypothesis to explain tree-species coexistence in tropical forests suggests that increased attack by natural enemies near conspecific trees gives locally rare species a competitive advantage. Host ranges of natural enemies generally encompass several closely related plant taxa suggesting that seedlings should also do poorly around adults of closely related species. We investigated the effects of adult Parashorea malaanonan on seedling survival in a Bornean rain forest. Survival of P. malaanonan seedlings was highest at intermediate distances from parent trees while heterospecific seedlings were unaffected by distance. Leaf herbivores did not drive this relationship. Survival of seedlings was lowest for P. malaanonan , and increased with phylogenetic dissimilarity from this species, suggesting that survival of close relatives of common species is reduced. This study suggests that distance dependence contributes to species coexistence and highlights the need for further investigation into the role of shared plant enemies in community dynamics.
Ecology Letters (2010) 13: 51–59     

Tree distribution pattern and fate of juveniles in a lowland tropical rain forest – implications for regeneration and maintenance of species diversity

Three analyses of species diversity in a lowland dipterocarp forest were conducted to examine whether the nature of forest community dynamics are determined by density-dependent recruitment and mortality of saplings with a data set obtained in a 50 ha plot in Pasoh Forest Reserve. The first analysis examined whether sapling density varied as a function of distance from the nearest conspecific adult. The second analysis assessed the relationship between the spatial distribution patterns of saplings and adult trees. A third analysis examined sapling recruitment and mortality based on data from 2 censuses, taken in 1985 and 1990. Four hundred forty-four species (each with more than 100 individuals) out of the total of 814 species recorded in the plot, were chosen for the analyses. Of these selected species, 56 species showed significant reduction in sapling densities close to the conspecific adults. Within this group, 11 species were in the emergent layer (29.0% of the total species in this layer), 17 were in the canopy layer (10.5%), 18 were in the understory layer (11.3%), and 10 were in treelet and shrub layer (11.8%). In contrast, the sapling densities of 53 species decreased with increasing distance from conspecific adults; 2 of these species were in the emergent layer (5.2% of the total species in this layer), 14 were in the canopy layer (8.6%), 21 were in the understory layer (13.2%), and 16 were in the treelet and shrub layer (18.8%). The saplings of 35 of the 444 total selected species were clumped, while adults were regularly or randomly distributed. Of the remaining species, in 183 species (41.2%), the distributions of both adults and saplings were clumped. Thus, these 2 analyses do not support the prediction that most of the species of lowland tropical forests fail to produce new adults in their vicinity and as a result of this, adult trees are more regularly distributed than their conspecific juveniles (Janzen 1970). In the third analysis, the recruitment of saplings of species in the emergent and canopy layers increased significantly and in proportion with mortality, suggesting that the dominant species suffer higher mortality than do less common species. This trend is not so apparent in the understory, and the treelet and shrub layers. The results imply that a dynamic equilibrium process, which prevents competitive exclusion and maintains space for minor species, may be active among the species in the upper layers (particularly the emergent layer); however, such a dynamic equilibrium condition is not due exclusively to the reduced recruitment of saplings near conspecific adults, and the dynamic equilibrium condition is not prevalent among the lower story species.     

Tree mortality due to an El Niño flood along the lower Tana River, Kenya

This study investigated forest tree mortality as a result of the extremely strong 1997–1998 El Niño flood in Tana River, Kenya, directly tested how mortality varied in relation to tree species and diameter at breast height (DBH), and indirectly tested how mortality varied in relation to floodwater depth. The study forest was under 2 m of water for approximately 71 days, from December 1997 until February 1998. Twenty-five liana, subcanopy tree, and canopy tree species were selected based on their importance in the diet of the Tana River mangabey ( Cercocebus galeritus Peters), a critically endangered primate endemic to the Tana River. Reproductive-sized individuals of these species were enumerated in 16.25 ha. I also enumerated trees killed by the El Niño flood, recorded their location in the forest, and measured their DBH. Ninety-two trees in fifteen species were killed by the flood. There were significant differences in mortality by species and by 0.25 ha quadrat. There were negative correlations between number of dead and DBH, number of dead and distance from the river, and per cent of quadrat killed and distance from the river. There appear to have been few long-term consequences of the El Niño flood in the forest or for the mangabeys.     

Density‐dependence in common tree species in a tropical dry forest in Mudumalai,southern India

Abstract. Density‐dependence in tree population dynamics has seldom been examined in dry tropical forests. Using long‐term data from a large permanent plot, this study examined 16 common species in a dry tropical forest in southern India for density‐dependence. Employing quadrat‐based analyses, correlations of mortality, recruitment and population change with tree densities were examined. Mortality in 1–10 cm diameter trees was largely negatively correlated with conspecific density, whereas mortality in > 10 cm diameter trees was positively correlated. Mortality was, however, largely unaffected by the basal area and abundance of heterospecific trees. Recruitment was poor in most species, but in Lagerstroemia microcarpa (Lythraceae), Tectona grandis (Verbenaceae) and Cassia fistula (Fabaceae), species that recruited well, strong negative correlations of recruitment with conspecific basal area and abundance were found. In a few other species that could be tested, recruitment was again negatively correlated with conspecific density. In Lagerstroemia, recruitment was positively correlated with the basal area and abundance of heterospecific trees, but these correlations were non‐significant in other species. Similarly, although the rates of population change were negatively correlated with conspecific density they were positive when dry‐season ground fires occurred in the plot. Thus, the observed positive density‐dependence in large‐tree mortality and the negative density‐dependence in recruitment in many species were such that could potentially regulate tree populations. However, repeated fires influenced density‐dependence in the rates of population change in a way that could promote a few common species in the tree community.     

Host specificity of ambrosia and bark beetles (Col., Curculionidae: Scolytinae and Platypodinae) in a New Guinea rainforest

Abstract.  1. Bark and ambrosia beetles are crucial for woody biomass decomposition in tropical forests worldwide. Despite that, quantitative data on their host specificity are scarce.
2. Bark and ambrosia beetles (Scolytinae and Platypodinae) were reared from 13 species of tropical trees representing 11 families from all major lineages of dicotyledonous plants. Standardised samples of beetle-infested twigs, branches, trunks, and roots were taken from three individuals of each tree species growing in a lowland tropical rainforest in Papua New Guinea.
3. A total of 81 742 beetles from 74 species were reared, 67 of them identified. Local species richness of bark and ambrosia beetles was estimated at 80–92 species.
4. Ambrosia beetles were broad generalists as 95% of species did not show any preference for a particular host species or clade. Similarity of ambrosia beetle communities from different tree species was not correlated with phylogenetic distances between tree species. Similarity of ambrosia beetle communities from individual conspecific trees was not higher than that from heterospecific trees and different parts of the trees hosted similar ambrosia beetle communities, as only a few species preferred particular tree parts.
5. In contrast, phloeophagous bark beetles showed strict specificity to host plant genus or family. However, this guild was poor in species (12 species) and restricted to only three plant families (Moraceae, Myristicaceae, Sapindaceae).
6. Local diversity of both bark and ambrosia beetles is not driven by the local diversity of trees in tropical forests, since ambrosia beetles display no host specificity and bark beetles are species poor and restricted to a few plant families.     

Migration of herbaceous plant species across ancient–recent forest ecotones in central Belgium

1 We studied the migration of forest plant species using their percentage cover and frequency in 197 plots distributed over 26 transects across ecotones between ancient and recent deciduous forests in the Meerdaal forest complex in central Belgium. The recent forest stands varied in age between 36 and 132 years, and all occurred on silty, well-drained soils.
2 The total cover, number and diversity of field layer species did not differ significantly between ancient and recent forest stands.
3 The number and cover of the ancient forest plant species and of ant-dispersed species correlated positively with the age of the recent forest and negatively with both the duration of its former agricultural land use and the distance to the ancient forest. This implies a slow colonization of the recent forest stands by these species; all species were, however, able to migrate across the ecotones.
4 The cover of four species ( Anemone nemorosa , Lamium galeobdolon , Convallaria majalis and Polygonatum multiflorum ) declined along the transect, suggesting that they are limited by seed dispersal. Their colonization rates, calculated from the occurrence of the farthest individual, ranged from < 0.05 to 1.15 m year⁻¹ and for other measures from < 0.05 to 0.65 m year⁻¹. Anemone and Lamium appeared to colonize the recent forest by establishment of isolated individuals, while Polygonatum and Convallaria expanded populations from existing patches on the border between ancient and recent forest.
5 Several forest species were able to colonize the recent forest rapidly, where some of them even reached a higher abundance, due to the increased availability of colonization sites with a higher nutrient content and a thinner organic layer.     

Synchrony in small mammal populations of montane forest patches in southern India

1. Small mammal populations were studied in montane evergreen forests in the Nilgiris, southern India, using live-trapping from January 1994 to September 1996. Two sites were selected, one with a single large forest patch and adjoining plantations, and the other with several small forest patches separated by grassland.
2. Nine species were recorded, of which eight were trapped in the forest patches, two in the grasslands and five in plantations. Rattus rattus was the most abundant species (2–36 individuals ha⁻¹) followed by Suncus montanus (0–11 individuals ha⁻¹). Densities of seven other species recorded were very low.
3. Synchrony in various population variables – density, biomass, mean weight, density of adults and adult females, and proportion of adults, adult females and sub-adults – was examined for Rattus rattus populations in the forest patches and plantations. Density and biomass were studied in seven other species recorded at these sites. Spearman's rank order correlation was used as a measure of synchrony between the population variables.
4. Within-site synchrony was higher than between-site synchrony in population characters. Synchrony was also higher between plots within the unfragmented site than they were between plots in the fragmented site. Relatively high synchrony in proportions of adults, adult females and juveniles in the forest patches implied that breeding is probably influenced by climate and food availability, which are seasonal in this habitat.
5. Given the small patch sizes (≈1–10 ha) and low population sizes, asynchrony is likely to be an outcome of demographic and environmental stochasticity, and low dispersal rates may impede establishment of synchrony.     

Heterogeneity in soil water and light environments and dispersal limitation: what facilitates tree species coexistence in a temperate forest?

In the present study, we analysed the habitat association of tree species in an old‐growth temperate forest across all life stages to test theories on the coexistence of tree species in forest communities. An inventory for trees was implemented at a 6‐ha plot in Ogawa Forest Reserve for adults, juveniles, saplings and seedlings. Volumetric soil water content (SMC) and light levels were measured in 10‐m grids. Relationships between the actual number of stems and environmental variables were determined for 35 major tree species, and the spatial correlations within and among species were analysed. The light level had no statistically significant effect on distribution of saplings and seedlings of any species. In contrast, most species had specific optimal values along the SMC gradient. The optimal values were almost identical in earlier life stages, but were more variable in later life stages among species. However, no effective niche partitioning among the species was apparent even at the adult stage. Furthermore, results of spatial analyses suggest that dispersal limitation was not sufficient to mitigate competition between species. This might result from well‐scattered seed distribution via wind and bird dispersal, as well as conspecific density‐dependent mortality of seeds and seedlings. Thus, both niche partitioning and dispersal limitation appeared less important for facilitating coexistence of species within this forest than expected in tropical forests. The tree species assembly in this temperate forest might be controlled through a neutral process at the spatial scale tested in this study.     

The fine-scale population dynamics of spruce budworm: survival of early instars related to forest condition

Abstract.  1. A lagged, density-dependent relationship between survival of early instars and host-tree condition is revealed during outbreaks of spruce budworm, Choristoneura fumiferana Clem. Persistent damage to hosts leads to deterioration of the stand.
2. Resource limitation affects survival during early-instar dispersal of spruce budworm. Impediments to distinguishing these events with estimates of survival were overcome with a simple model that describes the dispersal and survival processes. The model was used to analyse a recent 15-year population series from Black Sturgeon Lake and two historical datasets from Green River, in Canada.
3. Defoliation-induced damage to the trees resulted in increased losses of spring-emerging larvae that are dispersing in search of feeding sites. Losses were further exacerbated by biotic factors such as maternal fecundity, rates of infection by the pathogen, Nosema fumiferanae , and by weather-related effects on the foraging period.
4. Survival of early-stage budworm larvae in persistent outbreaks declined and the likelihood of other density-related factors such as rate of mortality from natural enemies increased. These results may reconcile outstanding differences in interpretation of the role of the forest resource in spruce budworm population dynamics and point to a common process linking the dynamics of other well-known budworm species.     

Japanese maple (Acer palmatum var. Matsumurae, Aceraceae) recruitment patterns: seeds, seedlings, and saplings in relation to conspecific adult neighbors

We analyzed the spatial patterns among seeds, seedlings, saplings, and conspecific adult trees of the cool-temperate tree species Acer palmatum var. Matsumurae in a conifer-hardwood mixed forest in northern Japan, using two models that consider the influence of each adult within the neighborhood of the offspring. The results showed that recruitment patterns of each stage could be characterized and that significant shifts occur between successive stages. Sound seeds were more widely dispersed than unsound seeds; the mean dispersal distance (MDD) was 41.5 m for sound seeds, but only 12.6 m for unsound seeds. Most seedlings were located near conspecific adult trees, with a MDD of 14.3 m. Saplings, however, were more dispersed away from conspecific adult trees, with an MDD of more than 35 m. Light and gap distributions did not strongly affect the spatial distribution of the offspring; most saplings were located under nonconspecific canopies. These results suggest that the recruitment pattern of Japanese maple offspring is strongly affected by conspecific adult neighbors, rather than by light and gap distributions, with close proximity to conspecific adult trees reducing the growth and survival of seedlings during the transition to saplings.     

Tree population dynamics in Kibale National Park, Uganda 1975–1998

Changes in species composition, stem abundance, and basal area of trees taller than or equal to 10 m in a medium altitude tropical rain forest at the Ngogo study area, Kibale National Park, Uganda are described for the period between 1975 and 1998 ( n  = 23 years). Two enumeration episodes were conducted in 263 plots of 5 m by 50 m during 1975–80 and 1997–98. During this period, species richness decreased by 3% (from 92 to 89). Species diversity (H') also declined slightly from 2.97 to 2.86. The number of trees recorded in the sample plots decreased by 8% (from 2545 to 2329), while basal area decreased from 49.48 m² ha⁻¹ to 48.68 m² ha^−1. However, stem abundance and basal area increased for some species.     

Seed dispersal, seedling establishment and gap partitioning among tropical pioneer trees

1 We examined the abundance and distribution patterns of pioneer seeds in the soil seed bank, and of pioneer seedlings in 53 recently formed gaps, in a 50‐ha forest dynamics plot on Barro Colorado Island (BCI), Panama. The aim was to assess the importance of dispersal limitation (failure of seeds to arrive at all sites suitable for their germination) and establishment limitation (failure of seeds having reached a site to germinate successfully and establish as seedlings) in determining patterns of gap occupancy.
2 The abundance of seeds in the soil seed bank was strongly negatively correlated with seed size, but was not correlated with the abundance of reproductive‐sized adult trees in the plot. In contrast, the abundance of pioneer seedlings > 10 cm height in natural gaps was strongly correlated with adult abundance, but was not correlated with seed size.
3 Seedlings were non‐randomly distributed among gaps, but seedling abundance was not directly related to gap size, and there was no evidence of partitioning of the light environment of gaps by small seedlings. Large differences in growth and mortality rates among species were observed after 1 year, and this may result in the gap size partitioning previously found in saplings of the same species.
4 Seedlings of most species, particularly those with large seeds, were relatively more abundant than expected in gaps close to their conspecific adults. Proximity to reproductives, and by inference dispersal limitation, therefore exerts some effect on seedling distribution. None the less, large differences between seed and seedling abundances for some species, and low seedling occupancy rates in some gaps close to adult conspecifics, suggest that seedling emergence probabilities and species‐specific establishment requirements may also be important determinants of local abundance.     

Composition,structure and diversity of cove forest stands in the Great Smoky Mountains: a patch dynamics perspective

Abstract. Cove forests of the Great Smoky Mountains are North American examples of old-growth temperate forest. Ecological attributes of seven stands were studied using one 0.6 - 1.0 ha plot per stand. Stand basal area (39 - 55 m²/ha) and biomass (326 - 471 Mg/ha) were high for temperate deciduous forest. Density ranged from 577 to 1075 stems/ha. All stands had a mixture of deciduous canopy species. Only rarely did a single species comprise more than half of the stand by density, basal area or biomass. Shade-intolerant species were present at low levels (1 - 5 % of total stand density). A wide range of stem diameters was characteristic of most species. However, some species lacked small stems, indicating discontinuous regeneration. Stands tended to have 10 - 20 tree species per ha and at least five species had biomass levels > 10 Mg/ha, indicating high evenness. Canopy gaps covered 10 % of the total area (2 - 21 % by stand). Gaps and conspecific patches of canopy trees > 0.05 ha in size were infrequent. Spatial analyses revealed a variety of patterns among species at inter-tree distances of 1 to 25 m. When all species were combined, juveniles showed aggregation, and adults were often hyperdispersed. Analyses for individual species confirmed that the mosaic of canopy species is influenced by non-random spatial processes. Adults of several species were aggregated at distances > 10 m. Juveniles of all major species exhibited aggregation. Several species exhibited regeneration near conspecific adults. This pattern suggested limited mobility for such species within the shifting mosaic. A diverse patchwork resulted despite the fact that many species did not exhibit segregation of adults and juveniles. Further understanding of patch dynamics and the potential for compositional steady state in cove forests requires long-term study with spatial data.     

Diversity of canopy and understorey spiders in north-temperate hardwood forests

1 We characterized and compared diversity patterns of canopy and understorey spiders (Arachnida: Araneae) on sugar maple ( Acer saccharum Marsh.) and American beech ( Fagus grandifolia Ehrh.) in hardwood forests of southern Québec, Canada.
2 We sampled canopies of 45 sugar maple and 45 American beech trees and associated understorey saplings in mature protected forests near Montréal. Samples were obtained by beating the crown foliage at various heights and by beating saplings around each tree.
3 Eighty-two species were identified from 13 669 individuals. Forty-eight species and 3860 individuals and 72 species and 9809 individuals were collected from the canopy and the understorey, respectively.
4 Multivariate analyses (NMDS ordination and NPMANOVA) showed the composition of canopy and understorey assemblages differed significantly, and canopy assemblages differed between tree species. Rank-abundance distribution models fitted to the canopy and understorey data indicated that different mechanisms structure the assemblages in both habitats. Three abundant spider species were significantly more common in the canopy; ten species were collected significantly more often in the understorey.
5 The forest canopy was shown to be an important reservoir for spider diversity in north-temperate forests.     

Species richness of phytophagous beetles in the tropical tree Brosimum utile (Moraceae): the effects of sampling strategy and the problem of tourists

Abstract.  1. Phytophagous beetles on six mature living trees and two dead trees of Brosimum utile (Moraceae) were surveyed during 1 year in a tropical wet forest in Panama. The dead trees were surveyed both as suspended in the canopy and after falling down to the understorey.
2. Canopy access was provided by a construction crane and sampling was performed by beating and hand-collecting. The same amount of time was spent on each tree in order to standardise sampling effort. A list of all species associated with the tree is presented.
3. A total of 3009 individuals representing 364 species were collected. Tourists were excluded from the analyses by recording host associations directly and by probability assessments of host associations based on abundance categories. A total of 2603 individuals and 244 species were associated with the tree. The proportion of tourists in the trees increased with sample size.
4. A single mature living tree had on average 58.5 ± 6.5 species. The local species richness of B. utile was estimated as 2.5 times higher than in a single mature tree; however, a substantial increase in species richness was attained when dead wood habitats were included. Saproxylic species made up 82% of the total material.
5. The investigated habitat types of B. utile constituted distinct, complementary species assemblages. Similarity between saproxylic species of dead suspended wood and dead understorey wood of the same tree was 0.2 (Morisita–Horn index), confirming a prominent vertical stratification among this guild.