The vertical distribution of small flying insects in the lowland rain forest of Zaire

To monitor the vertical distribution of flying insects in the semi-deciduous rain forest of Zaire, two types of trap were hung at four vertical levels. Large insects were excluded. The entire catch was analysed. There was a marked concentration of flying insects within and above the canopy, over a wide range of orders. The most abundant taxa showed the greatest tendency to occur at these upper levels. Within the size range collected, biomass in general reflected abundance, but was influenced by the relatively high proportion of small insects in and above the canopy. There was a strong similarity in the relative abundance of orders between trap types and sites, but total numbers varied greatly. This variation and the biases inherent in these trapping methods are discussed and we conclude that the bulk of the small flying insects of the rain forest are concentrated in the upper canopy.     

Importance of the understory stratum to entomofaunal diversity in a temperate deciduous forest

The vertical stratification of lepidopteran and coleopteran communities in a cool-temperate deciduous forest in Japan was examined to evaluate the hypothesis of an expected uniform distribution of mobile flying insects between the canopy and understory of temperate forests. Lepidopteran and coleopteran insects were trapped using light traps at three sites in each of the canopy and understory for three consecutive nights each month from April to October 2001. For Lepidoptera, species richness, abundance, and family richness were significantly higher in the understory than in the canopy. For Coleoptera, only abundance was larger in the canopy relative to the understory; species and family richness did not differ between the strata. The beta diversity of the lepidopteran community was larger between the strata than among sites, but the coleopteran community showed an inverse pattern. These results imply the presence of vertical stratification within the lepidopteran community, but not within the coleopteran community, in the temperate forest. The understory contributes more than the canopy to lepidopteran diversity in the temperate forest, although this stratification may be relatively weak because, in contrast to the situation in tropical forests, the canopy and understory assemblages share many species.     

Seasonality and vertical structure of light-attracted insect communities in a dipterocarp forest in Sarawak

Nocturnal flying insects were collected monthly for 13 months using ultra violet light-traps set at various vertical levels in a weakly-seasonal, tropical lowland dipterocarp forest in Sarawak, Malaysia. Abundance, faunal composition, size distribution and guild structure of these samples were analyzed with respect to temperal and vertical distributions. The nocturnal flying insect community in the canopy level was highly dominated by fig wasps (84%) in individual number, and by scarabaeid beetles (28%) in weight. A principal component analysis on monthly catches detected non-random, seasonal trends of insect abundance. The first two principal trends were an alternation of wetter (September to January) and less wet seasons (February to August) and an alternation between the least wet (January to March) and the other seasons. Many insect groups were less abundant in the least wet season than the other seasons, whilst inverse patterns were found in Scarabaeidae and Tenebrionidae. Significantly positive and negative correlations between monthly catch and rainfall were detected only in ovule-feeders and in phloem-feeders, respectively. Delayed, significant negative correlations between monthly catch and 1–3 month preceding rainfall were more frequently detected in phytophages, phloem-feeders, seed-feeders, wood-borers and scavengers. The peak in abundance along vertical levels were found at the canopy level (35 m) for phloem-, ovule-, seed-, root-, fungal-feeders and nectar collectors, at an upper subcanopy level (25 m) for scavengers and aquatic predators, and at a middle subcanopy level (17 m) for ants. Catches at the emergent level (45 m) did not exceed those at the canopy level.     

Vertical distribution of fruit‐feeding butterflies with evidence of sex‐specific differences in a Tanzanian forest

Tropical forests accommodate rich species diversity, particularly among insects. Habitat heterogeneity along the vertical gradient extending from the forest understorey to the tree canopy influences diversity. The vertical distribution of forest insects is poorly understood across Africa, most especially eastern Africa. Food‐baited traps were used to study the vertical stratification of adult fruit‐feeding nymphalid butterflies in Mtai Forest Reserve, north‐eastern Tanzania. Traps were located in the forest canopy and understorey. A total of 277 individuals of 24 species were captured. Species composition differed by trap locations: 33% of the species captured were found in both the canopy and understorey strata; however, significantly more species were captured in the understorey (54%) than canopy (13%). Males were significantly more abundant than females and captured in both strata. A greater proportion of females were captured in the understorey than the canopy. The time of day affected capture rates, with more individuals caught in the afternoon; however, there was no association between the time period and the sex of individuals captured in canopy versus understorey locations. Understanding how the sexes of butterflies vary in understorey versus canopy offers new biological insights into the vertical stratification of insects.     

Convergent structural responses of tropical forests to diverse disturbance regimes

Size frequency distributions of canopy gaps are a hallmark of forest dynamics. But it remains unknown whether legacies of forest disturbance are influencing vertical size structure of landscapes, or space-filling in the canopy volume. We used data from LiDAR remote sensing to quantify distributions of canopy height and sizes of 434 501 canopy gaps in five tropical rain forest landscapes in Costa Rica and Hawaii. The sites represented a wide range of variation in structure and natural disturbance history, from canopy gap dynamics in lowland Costa Rica and Hawaii, to stages and types of stand-level dieback on upland Mauna Kea and Kohala volcanoes. Large differences in vertical canopy structure characterized these five tropical rain forest landscapes, some of which were related to known disturbance events. Although there were quantitative differences in the values of scaling exponents within and among sites, size frequency distributions of canopy gaps followed power laws at all sites and in all canopy height classes. Scaling relationships in gap size at different heights in the canopy were qualitatively similar at all sites, revealing a remarkable similarity despite clearly defined differences in species composition and modes of prevailing disturbance. These findings indicate that power-law gap-size frequency distributions are ubiquitous features of these five tropical rain forest landscapes, and suggest that mechanisms of forest disturbance may be secondary to other processes in determining vertical and horizontal size structure in canopies.     

Canopy structure and vertical patterns of photosynthesis and related leaf traits in a deciduous forest

Canopy structure and light interception were measured in an 18-m tall, closed canopy deciduous forest of sugar maple (Acer saccharum) in southwestern Wisconsin, USA, and related to leaf structural characteristics, N content, and leaf photosynthetic capacity. Light attenuation in the forest occurred primarily in the upper and middle portions of the canopy. Forest stand leaf area index (LAI) and its distribution with respect to canopy height were estimated from canopy transmittance values independently verified with a combined leaf litterfall and point-intersect method. Leaf mass, N and A _max per unit area (LMA, N/area and A _max/area, respectively) all decreased continuously by over two-fold from the upper to lower canopy, and these traits were strongly correlated with cumulative leaf area above the leaf position in the canopy. In contrast, neither N concentration nor A _max per unit mass varied significantly in relation to the vertical canopy gradient. Since leaf N concentration showed no consistent pattern with respect to canopy position, the observed vertical pattern in N/area is a direct consequence of vertical variation of LMA. N/area and LMA were strongly correlated with A _max/area among different canopy positions (r²=0.81 and r²=0.66, respectively), indicating that vertical variation in area-based photosynthetic capacity can also be attributed to variation in LMA. A model of whole-canopy photosynthesis was used to show that observed or hypothetical canopy mass distributions toward higher LMA (and hence higher N/area) in the upper portions of the canopy tended to increase integrated daily canopy photosynthesis over other LMA distribution patterns. Empirical relationships between leaf and canopy-level characteristics may help resolve problems associated with scaling gas exchange measurements made at the leaf level to the individual tree crown and forest canopy-level.     

Differences between forest type and vertical strata in the diversity and composition of hymenopteran families and mymarid genera in northeastern temperate forests

Most insects' assemblages differ with forest type and show vertical stratification. We tested for differences in richness, abundance and composition of hymenopteran families and mymarid genera between sugar maple (Acer saccharum) and white pine (Pinus strobus) stands and between canopy and understory in northeastern temperate forests in Canada. We used flight interception traps (modified malaise traps) suspended in the canopy and the understory in a split-split block design, with forest type as the main factor, forest stratum as the first split factor, and collection bottle location as the second split factor. Hymenopteran families and mymarid genera differed in their diversity depending on forest type and stratum. Both family and genera richness were higher in maple than in pine forests, whereas family richness was higher in the canopy and top bottles and generic richness was higher in the understory and bottom bottles. Multivariate analysis separated samples by forest type, vegetation stratum, and bottle location. Family composition showed 77% similarity between forest types and 73% between the canopy and understory. At the lower taxa level, mymarid genera showed only 47% similarity between forest types and 40% between forest strata, indicating vertical stratification and relatively high beta-diversity. Our study suggests that hymenopteran diversity and composition is strongly dependent on forest type and structure, making flying members of this order particularly vulnerable to forest management practices. It also shows that insect assemblage composition (especially at low-taxon levels), rather than relative abundance and richness, is the community attribute most sensitive to forest type and vertical stratification.     

The biodiversity of arthropods from Australian rainforest canopies: General introduction,methods, sites and ordinal results

Abstract The arthropod assemblages occurring in the canopies of tropical, subtropical and cool temperate sites have been sampled using a pyrethrum knockdown technique. Details of the techniques used and the climate and vegetation of the areas studied are presented together with an analysis of the distribution of individual arthropods across Orders. An approach using generalized linear modelling partitioned the variance in numbers among sites within forest types and across the three forest types. The effects of both these components were significant. The differences between the ordinal signatures of each forest type are discussed and a number of hypotheses proposed to account for these differences, based on knowledge of the biology of the groups concerned. For the tropical and subtropical sites a comparison was made between samples collected in the low to mid-canopy with ones collected in the high canopy. Numbers of both insects and non-insects collected differed significantly with height in the subtropical forest and the distribution of insects across Orders was also significantly different in this forest type. In the tropical forests numbers of insects differed significantly between the two strata but neither the numbers of non-insects nor the ordinal profiles of either insects or non-insects were shown to be significantly different.     

Between-forest variation in vertical stratification of drosophilid populations

Abstract 1. The spatial distribution pattern of forest‐dwelling drosophilid flies was compared among species and among four forests with markedly different vertical foliage structures, including secondary and primary forests, with special reference to vertical stratification. 2. All 20 drosophilid species analysed showed vertically stratified distribution patterns, which were detected statistically in at least one forest site during the 2‐year study period. As a whole, the vertical distribution patterns were stable, with or without stratification, over the 2 years. 3. The ratio of species with stratified distribution to the total number of species changed in a similar trend with the vertical foliage complexity of forests, both showing a large gap between secondary and primary forests, even for species common to all four forests. 4. Many species, most of which were tree‐sap feeders, showed highly predictable patterns of vertical stratification across forests and years. This was associated strongly with regular preference for microhabitat, mostly for the canopy, suggesting that the unvarying nature of canopy environments in any forest is very important in producing, maintaining, and promoting the vertical stratification in organism abundance, and contributes to the ubiquity of the pattern. 5. Some species with a wider range of food resources than other species, most of which showed no clear preference for the canopy, were characterised by large between‐forest differences in vertical distribution patterns and tended to show non‐stratified patterns in secondary forests, contributing to the lower ratio of species with stratified distribution in the two secondary forests than in the primary forests. 6. The role of forest structure in organising the environment is discussed; it is suggested that vertical complexity of foliage structure affects the prevalence of vertical stratification in animal communities indirectly through the vertical heterogeneity of environmental conditions.     

Ecological impacts of tropical forest fragmentation: how consistent are patterns in species richness and nestedness?

Large areas of tropical forest now exist as remnants scattered across agricultural landscapes, and so understanding the impacts of forest fragmentation is important for biodiversity conservation. We examined species richness and nestedness among tropical forest remnants in birds (meta-analysis of published studies) and insects (field data for fruit-feeding Lepidoptera (butterflies and moths) and ants). Species-area relationships were evident in all four taxa, and avian and insect assemblages in remnants typically were nested subsets of those in larger areas. Avian carnivores and nectarivores and predatory ants were more nested than other guilds, implying that the sequential loss of species was more predictable in these groups, and that fragmentation alters the trophic organization of communities. For butterflies, the ordering of fragments to achieve maximum nestedness was by fragment area, suggesting that differences among fragments were driven mainly by extinction. In contrast for moths, maximum nestedness was achieved by ordering species by wing length; species with longer wings (implying better dispersal) were more likely to occur at all sites, including low diversity sites, suggesting that differences among fragments were driven more strongly by colonization. Although all four taxa exhibited high levels of nestedness, patterns of species turnover were also idiosyncratic, and thus even species-poor sites contributed to landscape-scale biodiversity, particularly for insects.     

Comparing four simple,inexpensive methods for sampling forest arthropod communities

Terrestrial arthropods are diverse, and quantifying their availability to consumers is important for understanding both consumer and insect distribution, abundance, and communities. However, characterizing arthropod communities in complex forest ecosystems is challenging. We compared arthropod communities in a wet‐limestone forest in Jamaica during the dry season sampled by four methods: branch clips, sweep netting, and sticky traps applied to tree trunks and hanging free of vegetation. We found no effect of relative height in the canopy for the two methods that could be used at different heights, i.e., hanging sticky traps and branch clips. In addition, the arthropod community sampled changed over time (season) for sweep nets and branch clips. We also found that branch clips and sweep nets sampled more arthropod taxa than the two sticky‐trap methods. In addition, branch clips and sweep nets sampled more ants and spiders than the two sticky‐trap methods, whereas collar sticky traps on tree trunks sampled more bark lice (Psocoptera), and hanging sticky traps more flies (Diptera) than the other methods. Percentages of flying insects and strong‐flying insects sampled did not differ between sweep netting and branch clipping, but a higher percentage of both groups were captured with collar and hanging sticky traps. Because we found that the different methods sampled different subsets of the arthropod community, both taxonomically and in terms of aerial versus non‐aerial taxa, investigators should choose the arthropod sampling methods that most closely align with their focal species and study questions. For example, investigators might use collar traps for studies of bark gleaners, hanging sticky traps for aerial foragers, and branch clips or sweep nets for foliage gleaners. Alternatively, if a focal species is known to prefer certain prey items, investigators may instead select a method that effectively samples those prey taxa. Finally, for some studies, using multiple sampling methods may be the best option.     

Stratification and diel activity of arthropods in a lowland rainforest in Gabon

The abundance, activity and species richness of arthropods, particularly of insect herbivores, were investigated in the upper canopy and understorey of a lowland rainforest at La Makandé, Gabon. In total 14 161 arthropods were collected with beating, flight interception and sticky traps, from six canopy sites, during the day and at night, from mid-January to mid-March 1999. The effects of stratum were most important, representing between 40 and 70% of the explained variance in arthropod distribution. Site effects represented between 20 and 40% of the variance and emphasized the néed for replication of sampling among canopy sites. Time effects (diel activity) explained a much lower percentage of variance (6–9%). The density and abundance of many arthropod taxa and species were significantly higher in the upper canopy than in the understorey. Arthropod activity was also higher during the day than at night. In particular, insect herbivores were 2.5 times more abundant and twice as speciose in the upper canopy than in the understorey, a probable response to the greater and more diverse food resources in the former stratum. Faunal overlap between the upper canopy and understorey was low. The most dissimilar herbivore communities foraged in the understorey at night and the upper canopy during the day. Further, a taxonomic study of a species-rich genus of herbivore collected there (Agrilus , Coleoptera Buprestidae) confirmed that the fauna of the upper canopy was different, diverse and very poorly known in comparison to that of the understorey. Herbivore turnover between day and night was rather high in the upper canopy and no strong influx of insect herbivores from lower foliage to the upper canopy was detected at night. This suggests that insect herbivores of the upper canopy may be resident and well adapted to environmental conditions there.     

Ecological integrity of upper Warri River,Niger Delta using aquatic insects as bioindicators

Aquatic insects are bioindicators of water quality. Their structure and species composition is used in assessing the ecological integrity of streams and rivers. Their composition and density of the upper Warri River, Niger Delta, Nigeria were assessed and the influence of different physical and chemical variables on their distribution was explored at three designated stations. A total of 57 taxa were recorded with station 2 accounting for the greatest Ephemeroptera–Plecoptera–Trichoptera (EPT) richness. Abundance of the aquatic insects was affected by the nature of the substrate, macrophytes and canopy cover at the various stations examined. Generally, the upper Warri River is a fairly clean water body rich in EPT organisms. Pollution tolerant insect taxa such as chironomids and culicids larvae were only sporadically present.     

Movement and vertical stratification of fruit-feeding butterflies in a managed West African rainforest

We analysed movement parameters and vertical stratification in fruit-feeding butterflies between a control, a thinned and a plantation site within a West African rainforest. Overall, distances moved between traps were largest in the plantation. Movement parameters were generally largest in species feeding on early successional hostplants of gap and margin habitats. In these species, distances between recaptures were significantly shorter in the thinned compared to the control forest. Conversely, forest floor species feeding on climbers and understorey shrubs showed significantly larger movement in the thinned forest. Higher strata species also flew larger distances in the thinned understorey. Including higher strata samples, they were significantly less abundant in the thinned plot, although understorey sampling alone indicated the contrary. Comparing vertical distribution patterns between thinned and control sites indicated a disruption of vertical stratification after thinning. Canopy species seem to fly in the upper strata of the more closed-canopy control forest, whereas they descend more frequently in the forest opened by the thinning management. Understorey sampling might therefore lead to biased conclusions due to differences in vertical distribution between forest plots. This study showed that thinning can affect the restricted-range forest floor butterflies as well as the more widespread canopy butterfly fauna.     

Understorey versus canopy: patterns of vertical stratification and diversity among Lepidoptera in a Bornean rain forest

We studied the vertical distribution of Lepidoptera from a canopy walkway within a dipterocarp rain forest at Kinabalu Park (Borneo) using three different methods: (1) Bait traps to survey fruit-feeding nymphalid butterflies, (2) standardized counts for predominantly flower-visiting butterflies and their potential predators, aerial-hawking birds, and (3) attraction by blacklight for hawk- and tiger moths. There was a distinct decrease in the abundance of fruit-feeding nymphalids towards the canopy, probably due to a reduced and less predictable availability of rotting fruits in higher strata. These constraints might also be responsible for a higher abundance variation in the canopy, and a significant shift in size from larger species in the understorey to smaller ones in the canopy. Changes of microclimate and the conspicuous increase of insectivorous aerial-hawking birds from ground to canopy layer may be responsible for the prominent change in species composition of fruit-feeding nymphalids between 20 and 30 m. Nectar-feeding Lepidoptera showed a reversed abundance pattern. One main factor contributing to the much higher abundance of flower-visiting butterflies and moth taxa in the canopy, such as Sphingidae and some Arctiinae, might be the increase of nectar resources available in upper vegetation layers. A distinctly higher diversity in hawkmoths was also found in the canopy. A higher abundance of insectivorous aerial-hawking birds in the canopy might contribute to the shift in body design of fruit-feeding nymphalids from more slender bodies at lower vegetation layers to stouter ones (i.e. species which are stronger on the wing) in the canopy. Larval resources could play an additional role in specialisation on but a small part of the vertical gradient. This may explain stratification pattern of the nymphalid subfamilies Morphinae and Satyrinae. Monocotyledoneous larval food plants of both taxa, whose flight activity is largely restricted to the understorey, occur mostly in lower vegetation layers. Our observations on a wide taxonomic and ecological range of butterflies and moths indicate that tropical forest canopies hold a distinct and unique Lepidoptera fauna, whose species richness and abundance patterns differ from lower strata. However, the notion of tropical forest canopies as peaks of terrestrial diversity does not hold uniformly for all taxa or guilds.     

Experimental transplants reveal strong environmental effects on the growth of non‐vascular epiphytes in Afromontane forests

Transplant studies can provide valuable information on the growth responses of epiphytic bryophytes and lichens to environmental factors. We studied the growth of six epiphyte species at three sites in moist Afromontane forests of Taita Hills, Kenya. With 558 pendant transplants, we documented the growth of four bryophytes and two lichens over 1 yr. The transplants were placed into the lower canopy of one forest site in an upper montane zone, and two forest sites in a lower montane zone. Several pendant moss species grew very well in the cool and humid environment of the upper montane forest, with some transplants more than doubling their biomass during the year. Conversely, all transplanted taxa performed poorly in the lower montane zone, presumably because of the unfavorable combination of ample moisture with excessive warmth and insufficient light which characterizes the lower canopy in dense lower montane forests. The results demonstrate that pendant transplants can be used for monitoring growth of non‐vascular epiphytes in tropical forests. The starting weight of 0.25 g for pendant transplants worked well and can be recommended for future studies.     

Assessment of higher insect taxa as bioindicators for different logging-disturbance regimes in lowland tropical rain forest in Sabah, Malaysia

One of the serious environmental problems since the 1980s has been the conflict between the high rate of deforestation and maintenance of healthy ecosystem services and biological values in tropical forests. There is an urgent demand for setting up an appropriate environmental assessment to keep healthy ecosystem functions and biodiversity along with sustainable forest use based on ecology. In this study, we tried to assess logging-disturbance effects on the abundances of several flying insect groups (higher-taxon approach) in lowland tropical rain forest (Deramakot Forest Reserve, Sabah, Malaysia), while considering seasonal changes and vertical forest stratification. The season was the most important factor affecting the abundances of all the insect groups. Effects of logging disturbance were prominent in the understorey but obscure in the canopy. Changes in physical conditions caused by logging—possibly an increased evaporation due to solar radiation—may have decreased the abundance of desiccation-sensitive insects, especially in the understorey. There are also two probable reasons for the difference between events in the understorey and those in the canopy: (1) noise effects of various physical, environmental factors may have obscured insect responses to logging disturbance in the canopy; (2) higher spatio-temporal variation in quality and quantity of living food resources—such as leaves, flowers and fruits—provided in the canopy may have affected the abundance of their consumer insects independently of logging disturbance. Thus, this study suggests that the abundance of some insect groups at higher-taxon level, especially in the understorey, can be used as bioindicators for assessing effects of logging disturbance on the forest ecosystem.     

Vertical variation in leaf gas exchange parameters for a Southeast Asian tropical rainforest in Peninsular Malaysia

Vertical variation in leaf gas exchange characteristics of trees grown in a lowland dipterocarp forest in Peninsular Malaysia was investigated. Maximum net photosynthetic rate, stomatal conductance, and electron transport rate of leaves at the upper canopy, lower canopy, and forest floor were studied in situ with saturated condition photosynthetic photon flux density. The dark respiration rate of leaves at the various heights was also studied. Relationships among gas exchange characteristics, and also with nitrogen content per unit leaf area and leaf dry matter per area were clearly detected, forming general equations representing the vertical profile of several important parameters related to gas exchange. Numerical analysis revealed that the vertical distribution of gas exchange parameters was well determined showing both larger carbon gain for the whole canopy and at the same time positive carbon gain for the leaves of the lowest layer. For correct estimation of gas exchange at both leaf and canopy scales using multi-layer models, it is essential to consider the vertical distribution of gas exchange parameters with proper scaling coefficients.