Plant biodiversity and conservation of tropical semi-evergreen forest in the Shervarayan hills of Eastern Ghats, India

Species diversity, density, population structure and dispersion patterns of all trees and lianas (30cm gbh) were inventoried in a tropical semi-evergreen forest in the Shervarayan hills of Eastern Ghats, south India. Such data are necessary for ecosystem conservation of the under-studied Eastern Ghats, as extensive forests here have already been converted to coffee and orange plantations and the landscape changed due to aluminium ore mining and quarrying. Four 1-ha plots were established in Sanyasimalai (SM) reserve forest of the Shervarayan hills, one plot (SM1) located close to mining and quarrying area, two other contiguous plots (SM2 and SM3) located in selective felling area and the fourth (SM4) in a relatively undisturbed forest. These are 1 to 4km apart in the same semi-evergreen forest tract. In the four study plots a total of 3260 stems (mean density 815ha–1) covering 80 species in 71 genera and 44 plant families were recorded. Species richness was greatest in the undisturbed plot SM4 (50), while lowest (33) in the selectively felled site SM2. The forest stand (SM4) was also denser (986 stemsha–1) and more voluminous (basal area 44.3m2ha–1 as compared with the site mean of 35m2ha–1) than the other plots. Four trees, Chionanthus paniculata, Syzygium cumini, Canthium dicoccum and Ligustrum perrottetii dominated the stand, collectively contributing to >50% of the total density. Species richness and stand density decreased with increasing tree girths. The forest stand contained a growing population, but there was considerable variation in basal area distribution between the plots. Trends in species population structure varied, particularly for selective-felled species. Most species exhibited clumped dispersion of individuals both at 0.25ha and 1-ha scales. Variation in plant diversity and abundance are related to site attributes and human impacts.     

Effects of selective logging on tree species diversity and composition of Bornean tropical rain forests at different spatial scales

Reduced-impact logging (RIL) is known to be beneficial in biodiversity conservation, but its effects on tree diversity remain unknown. Pattern of tree diversity following disturbance usually varies with spatial scale of sampling (i.e., plot size). We examined the impacts of RIL on species richness and community composition of tree species at different spatial scales, and the scale (plot size) dependency of the two metrics; species richness versus community similarity. One 2-ha and three to four 0.2-ha plots were established in each of primary, RIL, and conventionally logged (CL) forest in Sabah, Malaysia. Species richness (the number of species per unit number of stems) was higher in the RIL than in the CL forest at both scales. The relationship between species richness and logging intensity varied with plot size. Species richness was greater in the RIL than in the primary forest at the 2-ha scale, while it was similar between the two forests at 0.2-ha scale. Similarly, species richness in the CL forest demonstrated a greater value at the 2-ha scale than at the 0.2-ha scale. Greater species richness in the two logged forests at the 2-ha scale is attributable to a greater probability of encountering the species-rich, small patches that are distributed heterogeneously. Community composition of the RIL forest more resembled that of the primary forest than that of the CL forest, regardless of plot size. Accordingly, species richness is a scale-dependent metric, while community similarity is a more robust metric to indicate the response of tree assemblage to anthropogenic disturbance.     

Liana Abundance,Diversity, and Distribution on Barro Colorado Island,Panama

Lianas are a key component of tropical forests; however, most surveys are too small to accurately quantify liana community composition, diversity, abundance, and spatial distribution – critical components for measuring the contribution of lianas to forest processes. In 2007, we tagged, mapped, measured the diameter, and identified all lianas ≥1 cm rooted in a 50-ha plot on Barro Colorado Island, Panama (BCI). We calculated liana density, basal area, and species richness for both independently rooted lianas and all rooted liana stems (genets plus clones). We compared spatial aggregation patterns of liana and tree species, and among liana species that varied in the amount of clonal reproduction. We also tested whether liana and tree densities have increased on BCI compared to surveys conducted 30-years earlier. This study represents the most comprehensive spatially contiguous sampling of lianas ever conducted and, over the 50 ha area, we found 67,447 rooted liana stems comprising 162 species. Rooted lianas composed nearly 25% of the woody stems (trees and lianas), 35% of woody species richness, and 3% of woody basal area. Lianas were spatially aggregated within the 50-ha plot and the liana species with the highest proportion of clonal stems more spatially aggregated than the least clonal species, possibly indicating clonal stem recruitment following canopy disturbance. Over the past 30 years, liana density increased by 75% for stems ≥1 cm diameter and nearly 140% for stems ≥5 cm diameter, while tree density on BCI decreased 11.5%; a finding consistent with other neotropical forests. Our data confirm that lianas contribute substantially to tropical forest stem density and diversity, they have highly clumped distributions that appear to be driven by clonal stem recruitment into treefall gaps, and they are increasing relative to trees, thus indicating that lianas will play a greater role in the future dynamics of BCI and other neotropical forests.     

Tree diversity and distribution in undisturbed and human-impacted sites of tropical wet evergreen forest in southern Western Ghats, India

The composition, abundance, population structure and distribution patterns of all woody species ( 30 cm gbh) were investigated in an undisturbed and two adjacent human-impacted sites of a tropical wet evergreen forest in Kalakad National Park, Western Ghats, south India. Three 1-ha plots were established, one each in (i) an undisturbed site (named site UD), (ii) in a site selectively felled 35 years ago (site SF – small stems felled leaving the large trees (as shade) for developing it into a cardamom estate, on the failure of which the site was abandoned) and (iii) a frequently disturbed site (site FD – round woods logged for use in ovens for curing cardamom). These sites are 1 to 3 km apart in the same wet evergreen forest. In the three study plots a total of 2150 stems (mean density 716 ha^–1) covering 122 species in 89 genera and 41 families were enumerated. Species richness was greatest (85 species ha^–1) in the undisturbed site UD, intermediate (83) in SF and lowest (80) in FD. Tree density was greatest (855 stems ha^–1) in site SF, intermediate (720) in UD and lowest (575) in FD. The forest stand was exceptionally voluminous in site UD (basal area 94.64 m² ha^–1), intermediate (66.9 m²) in SF and least (61.7 m²) in FD, due to tree removal for fuel in the latter sites. Species composition and abundance patterns markedly varied between the three sites. In UD and SF, primary forest species (Cryptocarya bourdillonii , Cullenia exarillata Myristica dactyloides etc.) occurred in greater density. In FD heliophilic secondary forest species (Elaeocarpus venustus, Litsea wightiana, Viburnum punctatum and Vitex altissima) were abundant, while these were absent in UD and SF. The species–area curve did not reach an asymptote in any of the sites on the 1-ha scale. The stand population structure was clearly reverse J shaped in UD and SF, while small stems were 2- to 3-fold fewer in FD. Most trees exhibited clumped distribution of individuals on the 1-ha scale. Variation in the kind and richness of species and their abundance is related to human interference and the need for forest conservation is emphasized.     

Tree population changes in a tropical dry evergreen forest of south India over a decade (1992–2002)

Changes in species composition and density of trees >10 cm gbh in a tropical dry evergreen forest in Puthupet, south India are interpreted for the period between 1992 and 2002. A 1-ha plot was inventoried in 1992 and was recensused in 2002. During the 10-year interval tree taxa diversity as well as stand density increased, but the basal area value decreased. Tree species richness increased by 21% (from 24 to 29 species) by an addition of eight species and local extinction of three species. The tree density increased just by eight individuals (from 1330 stems ha⁻¹ in 1992 to 1338 ha⁻¹ in 2002), but the basal area decreased by 8% (from 37.5 to 34.5 m² ha⁻¹). Many species (11 numbers) have increased in abundance rather than decreased. Many surviving species seem to have considerable stability in abundance at the local scale. The density of smaller stems (10 29 cm gbh) increased by 15.3%, while that of the larger trees decreased drastically (81.6%). Ninety percent of the missing stems were from the middlestorey of the forest. Tree density changes among the three ecological guilds revealed a decrease in stem density and an increase in basal area in the lowerstorey; while the middlestorey exhibited a reverse trend. Family-wise, tree density changes revealed that the majority of families (67%) showed an increase in stem density. Long-term studies on tree population changes are essential to estimate tree mortality and recruitment rates, which will provide a greater insight in tropical forest dynamics.     

Effects of population density on forest structure and species richness and diversity of trees in Kampong Thom Province, Cambodia

This study examined differences in stand structure, tree species richness, and tree species diversity in relation to population density in Kampong Thom Province, Cambodia. Tree data were obtained from a 1997 forest inventory involving 60 clusters (540 plots) systematically distributed over 30% of the provincial forest area. Spatially referenced population data were obtained from the 1998 national population census. The average number of trees per cluster was 356/ha, the average basal area, 23 m²/ha, the average stand volume, 217 m³/ha, and the average aboveground biomass, 273 Mg/ha for all trees of DBH 10 cm and larger. The average species richness per cluster was 37 species, while average species diversity was measured as 0.916 using Simpson’s index and 2.98 by Shannon’s index. Significant negative correlations were generally found between population density surrounding clusters and tree density, basal area, stand volume, aboveground biomass, and species richness and diversity for three examined diameter classes (DBH of 10–30, ≥30, and ≥10 cm). As the distance from clusters for calculating population density increased, the correlation levels increased up to 5 or 7 km, depending on the variables and diameter class, and then stayed relatively constant for stand structure variables and decreased for species richness and diversity. The results indicate that evidence of disturbance was more pronounced at higher population density up to around 5 to 7 km. We suggest that introduction of greater controls on human disturbance should be a high priority for resource management and conservation in Kampong Thom Province and, presumably, Cambodia as a whole.     

Native tree regeneration in native tree plantations: understanding the contribution of Araucaria angustifolia to biodiversity conservation in the threatened Atlantic Forest in Argentina

Deforestation is a global process that has strongly affected the Atlantic Forest in South America, which has been recognised as a threatened biodiversity hotspot. An important proportion of deforested areas were converted to forest plantations. Araucaria angustifolia is a native tree to the Atlantic Forest, which has been largely exploited for wood production and is currently cultivated in commercial plantations. An important question is to what extent such native tree plantations can be managed to reduce biodiversity loss in a highly diverse and vulnerable forest region . We evaluated the effect of stand age, stand basal area, as a measure of stand density, and time since last logging on the density and richness of native tree regeneration in planted araucaria stands that were successively logged over 60 years, as well as the differences between successional groups in the response of plant density to stand variables. We also compared native tree species richness in planted araucaria stands to neighbouring native forest. Species richness was 71 in the planted stands (27 ha sampled) and 82 in native forest (18 ha sampled) which approximate the range of variation in species richness found in the native forests of the study area. The total abundance and species richness of native trees increased with stand age and time since last logging, but ecological groups differed in their response to such variables. Early secondary trees increased in abundance with stand age 3–8 times faster than climax or late secondary trees. Thus, the change in species composition is expected to continue for a long term. The difference in species richness between native forest and planted stands might be mainly explained by the difference in plant density. Therefore, species richness in plantations can contribute to local native tree diversity if practices that increase native tree density are implemented.     

Insights into regional patterns of Amazonian forest structure,diversity, and dominance from three large terra-firme forest dynamics plots

We analyze forest structure, diversity, and dominance in three large-scale Amazonian forest dynamics plots located in Northwestern (Yasuni and Amacayacu) and central (Manaus) Amazonia, to evaluate their consistency with prevailing wisdom regarding geographic variation and the shape of species abundance distributions, and to assess the robustness of among-site patterns to plot area, minimum tree size, and treatment of morphospecies. We utilized data for 441,088 trees (DBH ≥1 cm) in three 25-ha forest dynamics plots. Manaus had significantly higher biomass and mean wood density than Yasuni and Amacayacu. At the 1-ha scale, species richness averaged 649 for trees ≥1 cm DBH, and was lower in Amacayacu than in Manaus or Yasuni; however, at the 25-ha scale the rankings shifted, with Yasuni < Amacayacu < Manaus. Within each site, Fisher’s alpha initially increased with plot area to 1–10 ha, and then showed divergent patterns at larger areas depending on the site and minimum size. Abundance distributions were better fit by lognormal than by logseries distributions. Results were robust to the treatment of morphospecies. Overall, regional patterns in Amazonian tree species diversity vary with the spatial scale of analysis and the minimum tree size. The minimum area to capture local diversity is 2 ha for trees ≥1 cm DBH, or 10 ha for trees ≥10 cm DBH. The underlying species abundance distribution for Amazonian tree communities is lognormal, consistent with the idea that the rarest species have not yet been sampled. Enhanced sampling intensity is needed to fill the still large voids we have in plant diversity in Amazon forests.     

Plant biodiversity inventory and conservation of two tropical dry evergreen forests on the Coromandel coast, south India

Species diversity, population structure, abundance and dispersion patterns of all woody plants 10cm gbh were inventoried in two 1-ha plots of tropical dry evergreen (sacred grove or temple) forests at Kuzhanthaikuppam (KK) and Thirumanikkuzhi (TM) on the Coromandel coast of south India. Site KK is a stunted forest (average tree height ca 6 m) and TM a tall forest (average tree height ca 10 m). A total of 54 species (in 47 genera and 31 families) were recorded. Species richness and stand density were 42 and 38 species and 1367 and 974 individuals ha–1 respectively for the sites KK and TM. About 50% of the total species were common to both the sites. Site TM is twofold more voluminous (basal area 29.48 m2 ha–1) than KK (basal area 15.44 m2 ha–1). Nearly one third of the individuals are multi-stemmed in the low-statured site KK whereas one fourth of the tree density is multi-stemmed in TM. Species abundance pattern varied between the two sites. The abundance of three species in KK and two species in TM is pronounced. Memecylon umbellatum, the most abundant species contributing to one third of total stand density in KK, is least represented in TM. Species richness, density and diversity indices decreased with increasing girth threshold. Most species exhibited clumped dispersion of individuals both at 0.25 and 1-ha scales. Population structure for girth frequency is an expanding one for both the sites, except for basal area distribution in KK. Variations in plant diversity and abundance are related to site attributes and human impacts. In the light of habitat uniqueness, species richness and sacred grove status, the need for conservation is emphasized.     

Tree diversity and population structure in undisturbed and human-impacted stands of tropical wet evergreen forest in Arunachal Pradesh, Eastern Himalayas, India

Tree species richness, tree density, basal area, population structure and distribution pattern were investigated in undisturbed, mildly disturbed, moderately disturbed and highly disturbed stands of tropical wet evergreen forests of Arunachal Pradesh. The forest stands were selected based on the disturbance index (the basal area of the cut trees measured at ground level expressed as a fraction of the total basal area of all trees including felled ones): (i) undisturbed stand (0% disturbance index), (ii) mildly disturbed (20% disturbance index), (iii) moderately disturbed (40% disturbance index), and (iv) highly disturbed stand (70% disturbance index). Tree species richness varied along the disturbance gradient in different stands. The mildly disturbed stand showed the highest species richness (54 of 51 genera). Species richness was lowest (16 of 16 genera) in the highly disturbed stand. In the undisturbed stand, 47 species of 42 genera were recorded while in the moderately disturbed stand 42 species of 36 genera were found. The Shannon–Wiener diversity index for tree species ranged from 0.7 to 2.02 in all the stands. The highest tree diversity was recorded in the undisturbed stand and the lowest in the highly disturbed stand. The stands differed with respect to the tree species composition at the family and generic level. Fagaceae, Dipterocarpaceae and Clusiaceae dominated over other families and contributed 53% in the undisturbed, 51% in the mildly disturbed, 42% in the moderately disturbed and 49% in the highly disturbed forest stands to the total density of the respective stand. Stand density was highest (5452 stems ha^–1) in the undisturbed stand, followed by the mildly disturbed stand (5014), intermediate (3656) in the moderately disturbed stand and lowest (338) in the highly disturbed stand. Dominance, calculated as the importance value index of different species, varied greatly across the stands. The highest stand density and species richness were represented in the medium girth class (51–110 cm) in all the stands. In the undisturbed stand, the highest density was found in the 111–140 cm girth class, while in the mildly disturbed stand the 51–80 cm girth range recorded the highest density. About 55, 68 and 52% species were found to be regenerating in the undisturbed, mildly disturbed and moderately disturbed stands, respectively. No regeneration was recorded in the highly disturbed stand. Variation in species richness, distribution pattern and regeneration potential is related to human interference and the need for forest conservation is emphasized.     

Stand density and species richness affect carbon storage and net primary productivity in early and late successional temperate forests differently

How stand density and species richness affect carbon (C) storage and net primary productivity (NPP) changes with forest succession is poorly understood. We quantified the C storage of trees and the aboveground NPP in an early successional secondary birch forest (birch forest) and a late successional mixed broadleaf-Korean pine (Pinus koraiensis) forest (mixed forest) in northeastern China. We found that: 1) tree C storage in the mixed forest (120.3 Mg C ha^?1) was significantly higher than that in the birch forest (78.5 Mg C ha^?1), whereas the aboveground NPP was not different between the two forest types; and 2) only stand density had a positive linear relationship with tree C storage and aboveground NPP in the birch forest. In the mixed forest, both tree C storage and aboveground NPP were significantly affected by the combination of the stand density and species richness. The tree C storage to stand density and species richness relationships were hump-shaped. The aboveground NPP increased with increasing stand density, but its relationship to species richness was hump-shaped. We conclude that the effect of stand density and species richness on tree C storage and aboveground NPP was influenced by forest stand succession, and such effects should be considered in studying stand density- and species richness- ecosystem function (e.g., C storage and NPP) relationships in temperate forest ecosystems.     

Impact of human foraging on tree diversity,composition, and abundance in a tropical rainforest

Tropical forest tree communities are structured by a range of large-scale drivers including elevation, certain high-impact anthropogenic activities (e.g., deforestation), and fires. However, low-impact human activities such as foraging may also be subtly but notably altering the composition of tropical forest tree communities. The study assessed the (i) differences in species diversity, patterns of relative abundance, and pairwise beta diversity between trees with edible and inedible fruits and seeds along an elevation gradient, and (ii) impact of human foraging on the forest tree communities in Oban Division of Cross River National Park, Nigeria. Fifteen permanent 40 by 40 m plots were established along an elevational gradient (120–460 m above mean sea level). All trees of 0.1 m diameter at breast height (dbh) and above were measured, identified, and, with the aid of structured questionnaires, classified into those with edible and inedible fruits/seeds. A total of 35 edible species with density of 128 stems/hectare and basal area of 11.99 m²/hectare, and 109 inedible species with density of 364 stems/hectare and basal area of 22.42 m²/hectare were sampled. However, the evenness of edible and inedible species was similar at pooled and plot levels. For inedible species, there was a positive relationship between pairwise beta diversity and elevation, and this was driven mainly by turnover. In contrast, edible species exhibited a non-significant trend between elevation and beta diversity. Thus, the study showed that human foraging of edible fruits may have subtly influenced patterns of species diversity and community structure in this tropical forest.     

Growth and mortality of trees in Amazonian tropical rain forest in Ecuador

Abstract. Structural changes are analysed in four samples representing 4 ha, two line transects and two hectare plots, of Amazonian tropical lowland rain forest in northern Ecuador. Only trees with a DBH ≥ 10 cm were included. A sample of floodplain forest in Añangu represents the largest turnover found in tropical forests (stand half-life = 23 yr). The line transect and hectare plot both of tierra firme forest in Añangu have the same turnover (37 yr) and were balanced for death and in-growth of both individuals and wood (basal area). The 1-ha tierra firme sample in Cuyabeno had a turnover of 67 yr and was in a growing phase. The floodplain line transect in Añangu was in a phase of structural breakdown. However, the floodplain line transect had the largest growth of basal area per tree (23.4 cm²/yr). The tierra firme samples had a growth of 9.6, 10.1, and 13.6 cm²/yr. Most of the dead trees fell with some uprooting in three of the four samples. However, no significant difference in the distribution of mode of death was found between the four samples. Death was independent of topography and the dead trees were randomly distributed. As the trees grow up they occupy more space and larger trees (DBH ≥ 15 cm) become more uniformly distributed, whereas smaller trees (DBH ≤ 15 cm) were randomly distributed. Our study confirms that plots of 1 ha are not sufficient to include representative samples of different stages of forest structure.     

Structure and regeneration of canopy species in an old-growth evergreen broad-leaved forest in Aya district,southwestern Japan

The population structure and regeneration of canopy species were studied in a 4 ha plot in an old-growth evergreen broad-leaved forest in the Aya district of southwestern Japan. The 200 m × 200 m plot contained 50 tree species, including 22 canopy species, 3,904 trees (dbh5 cm) and a total basal area of 48.3 m²/ha. Forty one gaps occurred within the plot, and both the average gap size (67.3 m²) and the total area of gap to plot area (6.9%) were small. Species found in the canopy in the plot were divided into three groups (A, B, C) based on size and spatial distribution patterns, and density in each tree size. Group A (typical species: Distylium racemosum, Persea japonica) showed a high density, nearly random distribution and an inverse J-shaped size distribution. Species in group B (Quercus salicina, Quercus acuta, Quercus gilva) were distributed contagiously with conspicuous concentration of small trees (<5 cm dbh) around gaps. However, the species in this group included few trees likely to reach the canopy in the near future. Group C included fast-growing pioneer and shade intolerant species (e.g. Cornus controversa, Carpinus tschonoskii, Fagara ailanthoides), which formed large clumps. Most gaps were not characterized by successful regeneration of group B and C but did appear to accelerate the growth of group A. Group B species appear to require long-lived or large gaps while group C species require large, catastrophic disturbances, such as landslides, for regeneration.     

Effects of anthropogenic disturbance on plant diversity and community structure of a sacred grove in Meghalaya, northeast India

This study analyses the effects of anthropogenic disturbance on plant diversity and community attributes of a sacred grove (montane subtropical forest) at Swer in the East Khasi Hills district of Meghalaya in northeast India. The undisturbed, moderately disturbed and highly disturbed stands were identified within the sacred grove on the basis of canopy cover, light interception and tree (cbh 15 cm) density. The undisturbed forest stand had >40% canopy cover, >50% light interception and a density of 2103 trees per hectare, whereas the highly disturbed stand had <10% canopy cover, <10% light interception and 852 trees per hectare. The moderately disturbed stand occupied the intermediate position with respect to these parameters. The study revealed that the mild disturbance favoured species richness, but with increased degree of disturbance, as was the case in the highly disturbed stand, the species richness markedly decreased. The number of families of angiosperms was highest (63) in the undisturbed stand, followed by the moderately (60) and highly disturbed (46) stands. The families Rubiaceae, Asteraceae and Poaceae were the dominant families in the sacred forest. Rubiaceae was represented by 11, 14 and 10 species in the undisturbed, moderately disturbed and highly disturbed stands, respectively, whilst the family Asteraceae had 16 species in the moderately disturbed stand and 14 species in the highly disturbed stand. The number of families represented by a single species was reduced significantly from 33 in the undisturbed stand to 23 in the moderately and 21 in the highly disturbed stand. The similarity index was maximum (71%) between the undisturbed and moderately disturbed stand and minimum (33%) between the undisturbed and highly disturbed stands. The Margalef index, Shannon diversity index and evenness index exhibited a similar trend, with highest values in the moderately disturbed stand. In contrast, the Simpson dominance index was highest in the highly disturbed stand. There was a sharp decline in tree density and basal area from the undisturbed (2103 trees ha^–1 and 26.9 m² ha^–1) to the moderately disturbed (1268 trees ha^–1 and 18.6 m² ha^–1) and finally to the highly disturbed (852 trees ha^–1 and 7.1 m² ha^–1) stand. Density–girth curves depicted a successive reduction in number of trees in higher girth classes from the undisturbed to the moderately and highly disturbed stands. The log-normal dominance–distribution curve in the undisturbed and moderately disturbed stands indicated the complex and stable nature of the community. However, the short-hooked curve obtained for the highly disturbed stand denoted its simple and unstable nature.     

Tree species richness and density affect parasitoid diversity in cacao agroforestry

In Brazil, cacao is mostly planted beneath shade trees. The diversity of shade trees varies from monospecific to highly diverse canopies, characteristic of pristine Atlantic Forest. This study evaluates the relationships between family richness of Hymenoptera-Parasitica and Chrysidoidea, and tree species richness and density, the species richness of herbaceous understorey, and the area and age of the cacao agroforestry system. We sampled 16 cacao agroforestry systems, with canopy diversity ranging from one to 22 tree species per hectare, in three seasons: summer (March), winter (August) and spring (November). Parasitoids were sampled using eight Malaise-Townes traps per site. Tree species richness and density were enumerated within 1 ha at each site, and herbaceous plant species richness was calculated in eight 1 m² plots, within the hectare. The number of parasitoid families increased with tree species richness and density in spring and summer, but decreased in winter. Neither species richness of herbaceous plants nor area and age of the system affected parasitoid family richness. We suggest that the increase of parasitoid diversity with tree species richness and density in warmer seasons reflects increasing heterogeneity and availability of resources. The decrease in parasitoid family number with tree density in winter may be due to local impoverishment of resources, leading to parasitoid emigration to neighbouring forest remnants. This result implies that a higher diversity of shade trees will help to maintain high parasitoid levels and, in consequence, higher levels of natural enemies of cacao pests, particularly in the warmer seasons. This prediction is borne out in the experience of cacao producers. The proper management of shade tree diversity will play a vital role in maintaining the sustainability of cacao agroforestry production systems in the tropics and, concurrently, will maintain high biodiversity values in these locations.     

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.     

Alpha diversity of lianas in a tropical evergreen forest in the Anamalais, Western Ghats, India

Abstract. Species richness, abundance, size-class distribution, climbing mode and spatial patterns of lianas were investigated in a 30-ha permanent plot of tropical evergreen forest at Varagalaiar in the Anamalais, Western Ghats, India. Each hectare was subdivided into 10 m × 10 m quadrats, in which all lianas ≥ 1 cm d.b.h. were measured, tagged and identified. The total liana density was 11, 200 individuals (373 ha–1) and species richness was 75 species, representing 66 genera and 37 families. The richness estimators employed for species and family accumulation curves after 100 times randomization of sample order, have stabilized the curves at 16th and 15th hectares, respectively. A greater proportion of lianas was twiners (55% of species and 44.4% of density) and root climbers (5% of species and 14% of density), and a few were tendril climbers, reflecting the late successional stage of the forest. In the size-class distribution, 82% of abundance and 97% of species richness fell within 1–3 cm diameter threshold. The dominance of succulent diaspore type signifies the faunal dependence of lianas on vertebrate frugivores for dispersal. The diversity, population density and family composition of lianas of our site is compared with those of other tropical forests. The need for biomonitoring of this synusia in the permanent plot for forest functioning is emphasized.     

Thirty Years of Forest Census at Barro Colorado and the Importance of Immigration in Maintaining Diversity

The neutral theory of community ecology can predict diversity and abundances of tropical trees, but only under the assumption of steady input of new species into the community. Without input, diversity of a neutral community collapses, so the theory''s predictions are not relevant unless novel species evolve or immigrate. We derive analytically the species input needed to maintain a target tree diversity, and find that a rate close to per recruit would maintain the observed diversity of 291 species in the Barro Colorado 50-ha tree plot in Panama. We then measured the rate empirically by comparing species present in one complete enumeration of the plot to those present five years later. Over six census intervals, the observed rate of input was to species per recruit, suggesting that there is adequate immigration of novel species to maintain diversity. Species interactions, niche partitioning, or density-dependence, while they may be present, do not appear to enhance tree species richness at Barro Colorado.     

Fine‐scale forest variability and biodiversity in the boreal mixedwood forest

Local spatial variation in species distributions is driven by a mix of abiotic and biotic factors, and understanding such hierarchical variation is important for conservation of biodiversity across larger scales. We sought to understand how variation in species composition of understory vascular plants, spiders, and carabid beetles is associated with concomitant spatial variation in forest structure on a 1‐ha permanent plot in a never‐cut mixedwood forest in central Alberta (Canada). Using correlations among dendrograms produced by cluster analysis we associated data about mapped distribution of all living and dead stems > 1 cm diameter at breast height with distributions of the three focal taxa sampled from regular grids across the plot. Variation in each of these species assemblages were significantly associated with several forest structure variables at various spatial scales, but the scale of the associations varied among assemblages. Variation in species richness and abundance was explained mostly by changes in basal area of trees across the plot; however, other variables (e.g. snag density and tree density) were also important, depending on assemblage. We conclude that fine‐scale habitat variation is important in structuring spatial distribution of the species of the forest floor, even within a relatively homogeneous natural forest. Thus, assessments that ignore within‐stand heterogeneity and management that ignores its maintenance will have limited utility as conservation measures for these taxa, which are major elements of forest biodiversity.