Negative Correlation between Ant and Spider Abundances in the Canopy of a Bornean Tropical Rain Forest

In tropical rain forests, high canopy trees have diverse and abundant populations of ants and spiders. However, accessing high trees and their fauna remains difficult; thus, how ants and spiders interact in the canopy remains unclear. To better understand the interspecific interactions between these two dominant arthropod groups, we investigated their spatial distributions at the canopy surface in a tropical rain forest in Borneo. We sampled ants and spiders six times between 2009 and 2011 by sweeping with an insect net at the tree crown surfaces of 190 emergent or tall (≥20 m in height) trees. We collected 438 ant individuals belonging to 94 species and 1850 spider individuals (1630 juveniles and 220 adults) belonging to 142 morphospecies (adults only) from a total of 976 samples. The fact that we collected four times more spider individuals than ant individuals suggests that fewer ants forage at the tree crown surface than previously thought. The number of spider individuals negatively correlated with the number of ant individuals and the number of ant species, indicating significant exclusivity between ant and spider spatial distributions at the tree crown surface. Niche‐overlap between the two taxa confirmed this observation. Although our data do not address the causes of these spatial distributions, antagonistic interspecific interactions such as interference behaviors and intra‐guild predation are ecological mechanisms that give rise to exclusive spatial distributions.     

Phylogenetic Structure of Tree Species across Different Life Stages from Seedlings to Canopy Trees in a Subtropical Evergreen Broad-Leaved Forest

Investigating patterns of phylogenetic structure across different life stages of tree species in forests is crucial to understanding forest community assembly, and investigating forest gap influence on the phylogenetic structure of forest regeneration is necessary for understanding forest community assembly. Here, we examine the phylogenetic structure of tree species across life stages from seedlings to canopy trees, as well as forest gap influence on the phylogenetic structure of forest regeneration in a forest of the subtropical region in China. We investigate changes in phylogenetic relatedness (measured as NRI) of tree species from seedlings, saplings, treelets to canopy trees; we compare the phylogenetic turnover (measured as β_NRI) between canopy trees and seedlings in forest understory with that between canopy trees and seedlings in forest gaps. We found that phylogenetic relatedness generally increases from seedlings through saplings and treelets up to canopy trees, and that phylogenetic relatedness does not differ between seedlings in forest understory and those in forest gaps, but phylogenetic turnover between canopy trees and seedlings in forest understory is lower than that between canopy trees and seedlings in forest gaps. We conclude that tree species tend to be more closely related from seedling to canopy layers, and that forest gaps alter the seedling phylogenetic turnover of the studied forest. It is likely that the increasing trend of phylogenetic clustering as tree stem size increases observed in this subtropical forest is primarily driven by abiotic filtering processes, which select a set of closely related evergreen broad-leaved tree species whose regeneration has adapted to the closed canopy environments of the subtropical forest developed under the regional monsoon climate.     

The vertical foliage distributions of six understory tree species in a Chamaecyparis obtusa Endl. forest

Summary The relationships between the amounts of foliage and heights of trees were studied for the dominant understory tree species, including three evergreen and three deciduous species, in a secondary forest of Chamaecyparis obtusa Endl. The relationships showed two phases: leaf increasing and stationary phases. In the leaf-increasing phase, the height growth allowed these species to expand the canopy by increasing the number of leaves. In the stationary phase, the number of leaves was relatively constant number irrespective of tree height from 160 to 400 cm. The number of leaves in the stationary phase represents the maximum number of leaves that can be supported by trees under shady conditions. From the analyses of vertical distributions of leaves in six species, mono- and multi-layer foliage distributions were detected. Two evergreen species, Eurya japonica and Cleyera japonica, showed multi-layer foliage distributions, whereas three deciduous species, Lyonia ovalifolia, Rhododendron reticulatum and Vaccinium hirtum, and one evergreen species, Pieris japonica, showed mono-layer foliage distributions. The relationships between the weights of non-photosynthetic and photosynthetic organs of the six species were examined. The proportion of non-photosynthetic organs increased with tree height. The understory species attained the stationary phase and were maintained by minimizing their investment in non-photosynthetic organs, i.e. their height growth was arrested by the shady conditions under the crown trees.     

Ecological distribution of homobaric and heterobaric leaves in tree species of Malaysian lowland tropical rainforest

Tree species can generally be classified into two groups, heterobaric and homobaric leafed species, according to whether bundle-sheath extensions (BSEs) are found in the leaf (heterobaric leaf) or not (homobaric leaf). In this study, we study whether the leaf type is related to the growth environment and/or life form type, even in a tropical rain forest, where most trees have evergreen leaves that are generally homobaric. Accordingly, we investigated the distribution of leaf morphological differences across different life forms of 250 tree species in 45 families in a tropical rainforest. In total, 151 species (60%) in 36 families had homobaric leaves, and 99 species (40%) in 21 families had heterobaric leaves. We found that the proportion of heterobaric and homobaric leaf species differed clearly across taxonomic groups and life form types, which were divided into five life form types by their mature tree heights (understory, subcanopy, canopy, and emergent species) and as canopy gap species. Most understory (94%) and subcanopy (83%) species such as Annonaceae had homobaric leaves. In contrast, heterobaric leaf trees appeared more frequently in the canopy species (43%), the emergent species (96%) (such as Dipterocarpaceae), and the canopy gap species (62%). Our results suggest that tree species in the tropical rainforest adapt to spatial differences in the environmental conditions experienced at the mature height of each tree species, such as light intensity and vapor pressure difference, by having differing leaf types (heterobaric or homobaric) because these types potentially have different physiological and/or mechanical functions.     

Comparative height crown allometry and mechanical design in 22 tree species of Kuala Belalong rainforest, Brunei, Borneo

In rainforests, trunk size, strength, crown position, and geometry of a tree affect light interception and the likelihood of mechanical failure. Allometric relationships of tree diameter, wood density, and crown architecture vs. height are described for a diverse range of rainforest trees in Brunei, northern Borneo. The understory species follow a geometric model in their diameter-height relationship (slope, β = 1.08), while the stress-elasticity models prevail (β = 1.27-1.61) for the midcanopy and canopy/emergent species. These relationships changed with ontogeny, especially for the understory species. Within species, the tree stability safety factor (SSF) and relative crown width decreased exponentially with increasing tree height. These trends failed to emerge in across-species comparisons and were reversed at a common (low) height. Across species, the relative crown depth decreased with maximum potential height and was indistinguishable at a common (low) height. Crown architectural traits influence SSF more than structural property of wood density. These findings emphasize the importance of applying a common reference size in comparative studies and suggest that forest trees (especially the understory group) may adapt to low light by having deeper rather than wider crowns due to an efficient distribution and geometry of their foliage.     

Composition and Dynamics of Functional Groups of Trees During Tropical Forest Succession in Northeastern Costa Rica

We compared the functional type composition of trees ≥10 cm dbh in eight secondary forest monitoring plots with logged and unlogged mature forest plots in lowland wet forests of Northeastern Costa Rica. Five plant functional types were delimited based on diameter growth rates and canopy height of 293 tree species. Mature forests had significantly higher relative abundance of understory trees and slow-growing canopy/emergent trees, but lower relative abundance of fast-growing canopy/emergent trees than secondary forests. Fast-growing subcanopy and canopy trees reached peak densities early in succession. Density of fast-growing canopy/emergent trees increased during the first 20 yr of succession, whereas basal area continued to increase beyond 40 yr. We also assigned canopy tree species to one of three colonization groups, based on the presence of seedlings, saplings, and trees in four secondary forest plots. Among 93 species evaluated, 68 percent were classified as regenerating pioneers (both trees and regeneration present), whereas only 6 percent were classified as nonregenerating pioneers (trees only) and 26 percent as forest colonizers (regeneration only). Slow-growing trees composed 72 percent of the seedling and sapling regeneration for forest colonizers, whereas fast-growing trees composed 63 percent of the seedlings and saplings of regenerating pioneers. Tree stature and growth rates capture much of the functional variation that appears to drive successional dynamics. Results further suggest strong linkages between functional types defined based on adult height and growth rates of large trees and abundance of seedling and sapling regeneration during secondary succession.
Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp     

Age‐related Crown Thinning in Tropical Forest Trees

Gap dynamics theory proposes that treefall gaps provide high light levels needed for regeneration in the understory, and by increasing heterogeneity in the light environment allow light‐demanding tree species to persist in the community. Recent studies have demonstrated age‐related declines in leaf area index of individual temperate trees, highlighting a mechanism for gradual changes in the forest canopy that may also be an important, but less obvious, driver of forest dynamics. We assessed the prevalence of age‐related crown thinning among 12 tropical canopy tree species sampled in lowland forests in Panama and Puerto Rico (total N = 881). Canopy gap fraction of individual canopy tree crowns was positively related to stem diameter at 1.3 m (diameter at breast height) in a pooled analysis, with 10 of 12 species showing a positive trend. Considered individually, a positive correlation between stem diameter and canopy gap fraction was statistically significant in 4 of 12 species, all of which were large‐statured canopy to emergent species: Beilschmiedia pendula, Ceiba pentandra, Jacaranda copaia, and Prioria copaifera. Pooled analyses also showed a negative relationship between liana abundance and canopy gap fraction, suggesting that lianas could be partially obscuring age‐related crown thinning. We conclude that age‐related crown thinning occurs in tropical forests, and could thus influence patterns of tree regeneration and tropical forest community dynamics.     

Responses of crown development to canopy openings by saplings of eight tropical submontane forest tree species in Indonesia: a comparison with cool-temperate trees

BACKGROUND AND AIMS: Growth in trunk height in canopy openings is important for saplings. How saplings increase height growth in canopy openings may relate to crown architectural constraints. Responses of crown development to canopy openings in relation to trunk height growth were studied for saplings (0.2-2.5 m tall) of eight tropical submontane forest tree species in Indonesia. The results of this study were also compared with those of temperate trees in northern Japan. METHODS: The crown architecture differed among the eight tropical species, i.e. they had sparsely to highly developed branching structures. Crown allometry was compared among the eight species in each canopy condition (closed canopy or canopy openings), and between closed canopy and canopy openings within a species. A general linear regression model was used to analyse how each species increases height growth rate in canopy openings. Crown allometry and its plasticity were compared between tropical and temperate trees by a nested analysis of covariance. KEY RESULTS: Tropical submontane trees had responses similar to cool-temperate trees, showing an increase in height in canopy openings, i.e. taller saplings of sparsely branched species increase height growth rates by increasing the sapling leaf area. Cool-temperate trees have a wider crown projection area and a smaller leaf area per crown projection area to avoid self-shading within a crown compared with tropical submontane trees. Plasticity of the crown projection area is greater in cool-temperate trees than in tropical submontane trees, probably because of the difference in leaf longevity. CONCLUSIONS: This study concluded that interspecific variation in the responses of crown development to canopy openings in regard to increasing height related to the species' branching structure, and that different life-forms, such as evergreen and deciduous trees, had different crown allometry and plasticity.     

达维台风对海南尖峰岭热带山地雨林群落的影响

 海南岛地处我国台风的频发地带, 台风是影响海南岛热带雨林的一个重要的自然干扰因子。该文对2005年达维台风前后海南尖峰岭热带山地雨林天然次生林2 600 m²固定样地的物种组成、胸高断面积、植株数目、风倒木的影响因子、物种多样性、生物量变化及碳储量归还进行比较分析。结果显示, 台风后森林内产生大量的风倒木、断枝和落叶, 导致群落郁闭度减少、透光性增强, 并产生大量的林窗。台风后群落的组成结构发生显著变化, 受损株数达514株(占总株数26.1%); 其中风倒木比例较大, 达206株(占总株数10.5%)。方差分析结果显示, 受台风影响产生的风倒木与未明显受损树木的胸径面积、树高和木材密度均无显著差异; 但与树种组成有较紧密的关联, 对乔木层、幼树层和下木层造成的损害存在种类上的明显差异。台风产生的损害可以分为直接性损害和间接性损害两种: 1)直接性损害主要作用于乔木层, 造成大径级植株严重受损; 部分树种重要值降低, 变成伴生种, 或者次优势种和原有优势种成为共同优势种; 2)间接性损害主要通过风倒木作用于幼树层和下木层, 产生的倒木、断枝和落叶压倒其它植株, 表现为小面积范围内的个体死亡, 甚至有种类消失; 部分优势种的个体数减少较多, 但占该种群所有个体数目的比例较小。乔木层的Shannon-Wiener和Simpson多样性指数均略有下降, 而幼树层和下木层均略有上升, 但3个林层均表现为不同程度的种类消失和一些种类的个体数目减少。此次台风还造成该样地至少占台风前总生物量10.42%的生物量归还给林地, 并对森林生态系统碳循环产生重大影响。     

中亚热带喀斯特常绿落叶阔叶林空间结构特征

喀斯特常绿落叶阔叶林是中国西南喀斯特地区非地带性顶极森林群落,该研究基于相邻木关系的角尺度、混交度和大小比数3个结构参数,对木论国家级自然保护区常绿落叶阔叶林25 hm^(2)大型固定监测样地整体、上木层、下木层及其优势物种的空间结构特征进行了分析,以期揭示其森林空间结构现状,为喀斯特植被恢复重建提供依据。结果表明:(1)样地群落整体为轻度聚集分布格局,接近随机分布,物种高度混交且个体间大小分化程度相近,上木层呈现随机分布、强度混交和中庸偏亚优势状态,下木层呈现聚集分布、强度混交和中庸偏劣势状态。(2)样地内优势物种大多处于轻度聚集分布、强度混交且中庸偏劣势的状态。(3)林木大小比数与胸径、树高均呈显著负相关关系,混交度与胸径、树高均呈显著正相关关系,随林木的生长发育,树种优势度逐渐增大,物种多样性逐渐增强。研究认为,木论常绿落叶阔叶林空间结构尚未完全稳定,正处于演替中后期,具有发育成顶极群落的潜力。     

Spatial and temporal variations in photosynthetic capacity of a temperate deciduous-evergreen forest

Key message

The understory evergreen trees showed maximal photosynthetic capacity in winter, while the overstory deciduous trees showed this capacity in spring. The time lag in productive ecophysiologically active periods between deciduous overstory and evergreen understory trees in a common temperate forest was clearly related to the amount of overstory foliage.

Abstract

In temperate forests, where deciduous canopy trees and evergreen understory trees coexist, understory trees experience great variation in incident radiation corresponding to canopy dynamics represented by leaf-fall and leaf-out. It is generally thought that changes in the light environment affect understory plants’ ecophysiological traits. Thus, to project and estimate annual energy, water, and carbon exchange between forests and the atmosphere, it is necessary to investigate seasonal variation in the ecophysiological activities of both evergreen trees in the understory and deciduous trees that make up the canopy/overstory. We conducted leaf-scale gas-exchange measurements and nitrogen content analyses for six tree species along their heights throughout a complete year. Photosynthetic capacity as represented by the maximum carboxylation rate (V _cmax25) and photosynthetic nitrogen use efficiency (PNUE) of deciduous canopy trees peaked immediately after leaf-out in late May, declined and stabilised during the mid-growing season, and drastically decreased just before leaf-fall. On the other hand, the timing of lowest V _cmax25 and PNUE for evergreen understory trees coincided with that of the highest values for canopy trees. Furthermore, understory trees’ highest values appeared just before canopy tree leaf-out, when incident radiation in the understory was highest. This implies that failing to consider seasonal variation in leaf ecophysiological traits for both canopy and understory trees could lead to serious errors in estimating ecosystem productivity and energy balance for temperate forests.

     

Comparison of Herbivores and Herbivory in the Canopy and Understory for Two Tropical Tree Species1

The Janzen–Connell model of tropical forest tree diversity predicts that seedlings and young trees growing close to conspecific adults should experience higher levels of damage and mortality from herbivorous insects, with the adult trees acting as either an attractant or source of the herbivores. Previous research in a seasonal forest showed that this pattern of distance‐dependent herbivory occurred in the early wet season during the peak of new leaf production. I hypothesized that distance‐dependent herbivory may occur at this time because the new foliage in the canopy attracts high numbers of herbivores that are limited to feeding on young leaves. As a consequence, seedlings and saplings growing close to these adults are more likely to be discovered and damaged by these herbivores. In the late wet season, when there is little leaf production in the canopy, leaf damage is spread more evenly throughout the forest and distance dependence disappears. I tested three predictions based on this hypothesis: (1) the same species of insect herbivores attack young and adult trees of a given plant species; (2) herbivore densities increase on adult trees during leaf production; and (3) herbivore densities in the understory rise during the course of the wet season. Censuses were conducted on adults and saplings of two tree species, raribea asterolepis and Alseis blackiana. Adults and saplings of both species had largely the same suite of chewing herbivore species. On adults of Q. asterolepis, the density of chewing herbivores increased 6–10 times during leaf production, but there was no increase in herbivore density on adults of A. blackiana. Herbivore densities increased 4.5 times on A. blackiana saplings and 8.9 times on Q. asterolepis saplings during the wet season, but there were no clear trends on the adults of either species. These results suggest that the potential of adult trees as a source of herbivores on saplings depends on the value of new leaves to a tree species' herbivores, which may differ across tree species.     

Experimental suppression of ants foraging on rainforest vegetation in New Guinea: testing methods for a whole‐forest manipulation of insect communities

1. Ants are extremely abundant in lowland tropical forests where they are important predators, plant mutualists, and herbivores. Their complex role in tropical plant–insect food webs can be best assessed by experimental manipulation of their abundance. Historically, ant exclusion experiments have had a small‐scale focus, such as single trees. Here, we test a new ‘whole‐forest' method of ant exclusion, using treated canopy bait stations, in a diverse tropical rainforest in New Guinea. 2. We conducted a 10‐month manipulative experiment in primary and secondary rainforests. In each forest type, a 625 m² treatment plot was isolated from the surrounding forest and 135 bait stations treated with fipronil, S‐methoprene, and hydramethylnon were placed in trees to suppress ants. Ant activity was monitored in the forest canopy and understorey with an additional 65 stations in treatment and control plots. 3. We achieved a dramatic decline in ant abundance in treatment plots compared with controls in both forest types, with an average decrease in ant numbers per station of 82.4% in primary and 91.2% in secondary forest. In particular, native dominant species Oecophylla smaragdina, Anonychomyrma cf. scrutator in primary forest, and invasive Anoplolepis gracilipes in secondary forest were greatly affected. In contrast, Tapinoma melanocephalum flourished in treatment plots, perhaps benefiting from reduced competition from other ant species. 4. Our study demonstrates that it is possible to selectively eradicate most of the foraging ants in a structurally complex tropical forest. We propose whole‐forest manipulation as a novel tool for studying the role of ants in shaping plant–insect food webs.     

冠层结构和光环境的时空变化对紫耳箭竹种群特征的影响

选择金佛山国家自然保护区内落叶阔叶林、常绿落叶阔叶混交林、常绿阔叶林3种典型群落类型,研究冠层结构和光环境特征,以及林下优势种--紫耳箭竹的种群特征.结果表明: 随着落叶阔叶林→常绿落叶阔叶混交林→常绿阔叶林的演替,Shannon多样性指数、Simpson优势度指数和Pielou指数呈增加趋势,表明群落趋于稳定发展的状态;冠层结构特征也发生了显著改变,冠层开度和平均叶倾角减小,叶面积指数增加,冠层的消光能力增强,林下光照水平降低.上层林冠是造成林型郁闭的主要原因,其中冠层厚度和冠层面积是2个主要的影响因素.冠层结构与林下光照指标显著相关,对林下散射光的影响最大.冠层开度、林下光照条件均随着生长季的到来而下降,而叶面积指数呈现增长的趋势,峰值出现在6、7月;平均叶倾角在春季达到最大值,在夏季为最小值.紫耳箭竹的生长与冠层结构和光环境密切相关,其在光照适中的常绿落叶阔叶混交林中生长得最好,分株粗壮、密度大(29.69±1.68株·m^-2),地下茎拓展能力强;落叶阔叶林中的强光环境可能造成土壤水分缺失,从而对其生长产生影响;而在常绿阔叶林的低光环境下紫耳箭竹分株矮小,密度小(5.80±1.16株·m^-2),克隆扩展能力降低.在森林结构演变的过程中,冠层结构发生了明显的改变,显著影响林下光环境,过度的低光环境对紫耳箭竹种群的更新和发展有限制作用.     

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.     

Arboreal ants as key predators in tropical lowland rainforest trees

Ants numerically dominate the canopy fauna of tropical lowland rain forests. They are considered to be key predators but their effects in this regard have only rarely been studied on non-myrmecophytes. A conspicuously low abundance of less mobile, mainly holometabolous arthropods like Lepidoptera larvae corresponds with ant dominance, while hemimetabolous highly mobile nymphs occur regularly and in large numbers in the trees. This is in contrast to the temperate regions where ants are mostly lacking on trees and holometabolous larvae are frequent. In this study we experimentally measured ant predation in the trees by offering caterpillars as baits. Fifty-four ant species were tested, of which 46 killed caterpillars and carried them away to their nests while only eight species ignored the offered larvae. Insecticidal knockdown fogging of ten trees after finishing the prey experiments showed that on average 85% of ant individuals per tree were predacious. With the analysis of another 69 foggings and meticulous observations in many other trees this suggests that arboreal ants are responsible for the low abundance of less mobile arthropods in tropical lowland rain forest canopies. Ant predation was significantly lower in a disturbed forest indicating that human disturbance induces a change in the functional interactions in these ecosystems.     

Insects and light interact to mediate vine colonization of fast growing Microberlinia bisulcata tree seedlings in gaps of an African rain forest

Vines thrive in lowland tropical forests, yet the biotic factors underlying their colonization of host tree seedlings and saplings remain surprisingly understudied. Insect herbivores presumably could influence this process, especially where disturbance has opened the canopy (i.e., gaps)—temporary areas of higher primary productivity favoring the recruitment of vines and trees and invertebrates in forests—but their impact on vine colonization has never been experimentally tested. Using data from an insect herbivore exclusion (mesh-netting cages) experiment conducted in an African rain forest (Korup, Cameroon), I logistically modeled the probability of vines colonizing seedlings of three co-dominant species (Microberlinia bisulcata vs. Tetraberlinia bifoliolata and T. korupensis) in paired shaded understory and sunny gap locations (41 blocks across 80 ha, starting n = 664 seedlings) in a 1–2-yr period (2008–2009). Vine colonization occurred almost exclusively in gaps, occurring on 16% of seedlings there. Excluding herbivores in gaps doubled colonization of the light-demanding and faster growing M. bisulcata but had negligible effects on the two shade-tolerant, slower growing and less palatable Tetraberlinia species, which together were twice as susceptible to vines under natural forest gap conditions (controls). When protected from herbivores in gaps, more light to individual seedlings strongly increased vine colonization of M. bisulcata whereas its well-lit control individuals supported significantly fewer vines. These results suggest vines preferably colonize taller seedlings, and because light-demanding tree species grow faster in height with more light, they are more prone to being colonized in gaps; however, insect herbivores can mediate this process by stunting fast growing individuals so that colonization rates becomes more similar between co-occurring slow and fast growing tree species. Further influencing this process might be associational resistance or susceptibility to herbivores linked to host species’ leaf traits conferring shade-tolerant ability as seedlings or saplings. A richer understanding of how vines differentially influence forest regeneration and species composition may come from investigating vine–tree–herbivore interactions across light gradients, ideally via long-term studies and intercontinental comparisons. Abstract in French is available with online material.     

Anti-herbivore effects of an ant species,Crematogaster difformis,inhabiting myrmecophytic epiphytes in the canopy of a tropical lowland rainforest in Borneo

Ants are believed to regulate herbivorous insects in the canopy of tropical rainforests, but few studies have empirically investigated the anti-herbivore effects of the ants there. We examined the anti-herbivore effects of the ant species Crematogaster difformis, which territorializes a large area of the crown of emergent canopy trees and inhabits the myrmecophytic epiphytes, Lecanopteris sp. and Platycerium sp., which grow in the crown, by performing an ant-exclusion experiment in the field. The average proportion of leaf area loss, the proportion of damaged leaves, and the proportion of leaves with ≥50% leaf area loss were all significantly higher on experimentally ant-excluded branches than on ant-attended branches at 3 months from the beginning of the ant-exclusion treatment. These results suggest that C. difformis regulates not only herbivorous insects that potentially feed on its host epiphytes but also those that could feed on leaves of emergent canopy trees that harbor the epiphytes.     

Influence of forest type and tree species on canopy ants (Hymenoptera: Formicidae) in Budongo Forest, Uganda

The arboreal ant fauna was investigated in Budongo Forest, a seasonal rain forest in Uganda, using the insecticidal fogging technique. Ants were collected from 61 trees, between 7 and 33 m in height, belonging to four tree species. Trees were growing in adjacent plots of forests characterized by different use and structure: an old primary forest, a primary swamp forest along a small river, and a secondary forest where selective logging was carried out for 30 years. A total number of 37,065 ants, belonging to 161 species in 30 genera were collected. Considering the high number of species found only once, the completeness of the canopy ant fauna was relatively high and of relatively similar magnitude as samples from the Neotropics or the Oriental region. Up to 37 ant species on a single tree, with an average of 18.2 species per tree, were found. Forty-four ant species (28.1%) were found only once, less than ten individuals were found for each of 88 species (54.7%), but 64.0% of all individuals belonged to one of five species. Considering the high numerical dominance of a few ant species like a Pheidole sp., Tetramorium aculeatum (Mayr) and a Crematogaster sp., there is some evidence for an ant mosaic in the lower canopy of the Budongo Forest. Individual numbers of ants were strongly correlated with nests in the fogged tree, though the ants were not homogeneously distributed in the tree crowns. Diversity measures that strongly depend on individual numbers such as the Morisita-Horn index or rarefaction methods were calculated, but results were not concordant with those of incidence-based estimates such as jack-knife calculations. Differences in ant species richness and faunal composition between tree species were low, but more significant between forest types. The ant fauna in the secondary forest was less diverse with 12.6% fewer species compared to the primary forest sites. The average number of ant species per tree was significantly lower in the secondary forest (<20% of the species; F=8.03, df=59, P<0.01) than in the undisturbed forest types. Cataulacus, Leptothorax, Tetraponera, and Polyrhachis, which are typical canopy-dwelling ant genera, had a significantly higher diversity and frequency in the two primary forest types (F=4.17, df=53, P<0.05). Secondary forest trees are often younger, lacking dead branches and epiphytes which are important requisites for ant colonization on trees.     

Effects of size,neighbors, and site condition on tree growth in a subtropical evergreen and deciduous broad‐leaved mixed forest,China

Successful growth of a tree is the result of combined effects of biotic and abiotic factors. It is important to understand how biotic and abiotic factors affect changes in forest structure and dynamics under environmental fluctuations. In this study, we explored the effects of initial size [diameter at breast height (DBH)], neighborhood competition, and site condition on tree growth, based on a 3‐year monitoring of tree growth rate in a permanent plot (120 × 80 m) of montane Fagus engleriana–Cyclobalanopsis multiervis mixed forest on Mt. Shennongjia, China. We measured DBH increments every 6 months from October 2011 to October 2014 by field‐made dendrometers and calculated the mean annual growth rate over the 3 years for each individual tree. We also measured and calculated twelve soil properties and five topographic variables for 384 grids of 5 × 5 m. We defined two distance‐dependent neighborhood competition indices with and without considerations of phylogenetic relatedness between trees and tested for significant differences in growth rates among functional groups. On average, trees in this mixed montane forest grew 0.07 cm year^?1 in DBH. Deciduous, canopy, and early‐successional species grew faster than evergreen, small‐statured, and late‐successional species, respectively. Growth rates increased with initial DBH, but were not significantly related to neighborhood competition and site condition for overall trees. Phylogenetic relatedness between trees did not influence the neighborhood competition. Different factors were found to influence tree growth rates of different functional groups: Initial DBH was the dominant factor for all tree groups; neighborhood competition within 5 m radius decreased growth rates of evergreen trees; and site condition tended to be more related to growth rates of fast‐growing trees (deciduous, canopy, pioneer, and early‐successional species) than the slow‐growing trees (evergreen, understory, and late‐successional species).