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.     

西双版纳热带雨林林窗空间分布格局及其特征数与林窗下植物多样性的相关性

林窗作为森林群落中一种重要的干扰方式, 对林下物种构成有着重要的影响。开展林窗空间格局及其特征指数与林下植物多样性关系研究对于探讨林窗对林下生物多样性的影响有重要意义, 有助于进一步了解群落动态, 在物种多样性保护方面也具有指导作用。本研究在西双版纳热带雨林地区随机选取3块大小为1 ha的热带雨林为研究样地, 采用轻小型六旋翼无人机搭载Sony ILCE-A7r可见光传感器, 分别获取各个样地的高清数字影像, 结合数字表面高程模型以及各个样地的地形数据用以确定各样区的林窗分布格局, 并进一步提取出各林窗的景观格局指数。结合地面样方基础调查数据, 对各样地各林窗下植物多样性情况进行统计, 旨在分析热带雨林林窗空间分布格局以及林窗下植物多样性对各林窗空间格局特征的响应情况。研究表明, 西双版纳州热带雨林林窗呈大而分散的空间分布, 林窗空间格局特征指数如林窗形状复杂性指数、林窗面积都与林下植物多样性呈显著正相关关系。在面积小的林窗下, 较之林窗形状复杂性因子, 林窗面积大小对林下植物多样性影响更显著; 在面积达到一定程度后, 相对于面积因子, 林窗形状复杂性指数对林下植物多样性影响更显著, 各样地林窗皆趋于向各自所处样地顶极群落发展。     

Canopy Openness Enhances Diversity of Ant–Plant Interactions in the Brazilian Amazon Rain Forest

In closed‐canopy tropical forest understory, light availability is a significant determinant of habitat diversity because canopy structure is highly variable in most tropical forests. Consequently, variation in canopy cover affects the composition and distribution of plant species via creating variable light environments. Nevertheless, little is known about how variation in canopy openness structures patterns of plant–animal interactions. Because of the great diversity and dominance of ants in tropical environments, we used ant–plant interactions as a focal network to evaluate how variation in canopy cover influences patterns of plant–insect interactions in the Brazilian Amazon rain forest. We observed that small increases in canopy openness are associated with increased diversity of ant–plant interactions in our study area, and this change is independent of plant or ant species richness. Additionally, we found smaller niche overlap for both ants and plants associated with greater canopy openness. We hypothesize that enhanced light availability increases the breadth of ant foraging sources because variation in light availability gives rise to plant resources of different quality and amounts. Moreover, greater light availability promotes vegetative growth in plants, creating ant foraging ‘bridges’ between plants. In sum, our results highlight the importance of environmental heterogeneity as a determinant of ant–plant interaction diversity in tropical environments.     

Vertical stratification of ichneumonid wasp communities: the effects of forest structure and life‐history traits

Parasitoid wasp communities of the canopy of temperate forests are still largely unexplored. Very little is known about the community composition of parasitoids between canopy and understory and how much of this difference is related to forest structure or parasitoid biological strategies. In this study we investigated upon the difference in the community composition of the parasitic wasps Ichneumonidae between canopy and understory in a lowland temperate forest in northern Italy. We used general linear models to test whether parasitic strategy modifies species vertical stratification and the effect of forest structure. We also tested differences in β‐diversity between canopy and understory traps and over time within single forest layers. We found that stand basal area was positively related to species richness, suggesting that the presence of mature trees can influence local wasp diversity, providing a higher number of microhabitats and hosts. The ichneumonid community of the canopy was different from that of the understory, and the β‐diversity analysis showed higher values for the canopy, due to a higher degree of species turnover between traps. In our analyses, the vertical stratification was different between groups of ichneumonids sharing different parasitic strategies. Idiobiont parasitoids of weakly or deeply concealed hosts were more diverse in the understory than in the canopy while parasitoids of spiders were equally distributed between the two layers. Even though the ichneumonid community was not particularly species‐rich in the canopy of the temperate forests, the extension of sampling to that habitat significantly increased the number of species recorded.     

Post‐fire pickings: Large herbivores alter understory vegetation communities in a coastal eucalypt forest

Fire and herbivores alter vegetation structure and function. Future fire activity is predicted to increase, and quantifying changes in vegetation communities arising from post‐fire herbivory is needed to better manage natural environments. We investigated the effects of post‐fire herbivory on understory plant communities in a coastal eucalypt forest in southeastern Australia. We quantified herbivore activity, understory plant diversity, and dominant plant morphology following a wildfire in 2017 using two sizes of exclosures. Statistical analysis incorporated the effect of exclusion treatments, time since fire, and the effect of a previous prescribed burn. Exclusion treatments altered herbivore activity, but time since fire did not. Herbivory reduced plant species richness, diversity, and evenness and promoted the dominance of the most abundant plants within the understory. Increasing time since fire reduced community diversity and evenness and influenced morphological changes to the dominant understory plant species, increasing size and dead material while decreasing abundance. We found the legacy effects of a previous prescribed burn had no effect on herbivores or vegetation within our study. Foraging by large herbivores resulted in a depauperate vegetation community. As post‐fire herbivory can alter vegetation communities, we postulate that management burning practices may exacerbate herbivore impacts. Future fire management strategies to minimize herbivore‐mediated alterations to understory vegetation could include aggregating management burns into larger fire sizes or linking fire management with herbivore management. Restricting herbivore access following fire (planned or otherwise) can encourage a more diverse and species‐rich understory plant community. Future research should aim to determine how vegetation change from post‐fire herbivory contributes to future fire risk.     

林分因子对神木垒不同森林群落林下植物多样性的影响

为探究神木垒不同森林群落林下植物多样性的差异,本研究采用典型样地法,以夹金山神木垒的5种主要森林群落：云杉林、丽江云杉林、红杉林、针阔混交林、阔叶林为研究对象,对不同森林群落林下植物物种组成和物种多样性进行比较,并对林分因子和林下植物多样性进行冗余分析,确定影响林下植物多样性的主要林分因子,为当地森林经营管理提供理论依据。结果表明：（1）在研究区内共记录林下植物147种,隶属于61科,108属;云杉林群落林下植物的科属种组成最丰富。（2）各类型群落的H值、H"值、D值、JSW值均为：草本层＞灌木层;灌木层多样性最高的群落为云杉林群落,草本层多样性最高的群落为丽江云杉林群落,针阔混交林群落、阔叶林群落林下植物多样性较差。（3）平均枝下高与林分密度是影响灌木层物种多样性的主要林分因子（P<0.01）,平均枝下高与灌木层的D值、H值、H"值呈负相关关系,林分密度与灌木层4个多样性指数均呈正相关关系;平均枝下高是影响草本层物种多样性的主要林分因子（P<0.01）,平均枝下高与草本层H值、H"值、JSW值呈正相关关系。本研究认为,云杉林群落与丽江云杉林群落的林下植物多样性水平较高,平均枝下高与林分密度是影响神木垒不同森林群落林下植物多样性的主要林分因子。     

Comparative thermal adaptation in cicadas (Hemiptera: Cicadidae) inhabiting Mediterranean ecosystems

The thermal responses of cicadas inhabiting the Mediterranean ecosystems in Europe, North America, South Africa, and Australia are investigated. A total of 37 species and two subspecies from 17 genera representing eight tribes and three subfamilies of cicadas are investigated. The analysis includes species that are restricted to the Mediterranean ecosystem as well as those which also inhabit additional environments. The data suggest that cicadas adapt to the climate type regardless of particular types of plants within the various Mediterranean communities. Similarly, cicada thermal responses are independent of body size or taxonomic affinities. There is a wider range of body temperatures for the maximum voluntary tolerance temperature than for heat torpor or minimum flight temperatures. This diversity seems to be determined by the subdivision of the habitat used and the behavior of the species. All species possess relatively elevated heat torpor temperatures adapting to the general thermal characteristics of the Mediterranean ecosystem. The data suggest that cicadas adapt to the Mediterranean climate type regardless of the diversity of particular types of plants within the various communities, of body size or of taxonomic position.     

Diversity and seasonal variation of ground and understory spiders from a tropical mountain cloud forest

We made intensive samplings to study the seasonal response of spiders across different forest strata (ground and understory) in a tropical mountain cloud forest from Mexico. We sampled spiders from ten plots in six sampling events during the dry and rainy season, to analyze their abundance, structure (distribution of abundance among species), diversity and the response of the five dominant species at each stratum. Results demonstrated that seasonal patterns of spider communities differed among strata, revealing a complex spatiotemporal dynamic. Abundance, structure, diversity of ground spiders, as well as the responses of four dominant species at this stratum, showed low seasonal variations. In contrast, a strong seasonal variation was observed for the understory assemblage, with lowest abundance and highest diversity in the rainy season, and different assemblage structures for each season. Seasonal patterns of each assemblage seem linked to the responses of their dominant species. We found high co‐occurrence among most of the ground dominant species with similar habitat use and with multivoltine patterns, contrasting with low co‐occurrence among most of the understory dominant species with similar habitat use and univoltine patterns. Our results showed that the spiders’ assemblages of tropical mountain cloud forest (opposed to what is found in temperate and boreal forests) increase their species richness with the height, and that their responses to seasonal change differ between strata. Management programs of these habitats should consider the spatial and temporal variations found here, as a better understanding of their ecological dynamics is required to support their sustainable management.     

Understory Plant Community Composition Is Associated with Fine-Scale Above- and Below-Ground Resource Heterogeneity in Mature Lodgepole Pine (Pinus contorta) Forests

Understory plant communities play critical ecological roles in forest ecosystems. Both above- and below-ground ecosystem properties and processes influence these communities but relatively little is known about such effects at fine (i.e., one to several meters within-stand) scales, particularly for forests in which the canopy is dominated by a single species. An improved understanding of these effects is critical for understanding how understory biodiversity is regulated in such forests and for anticipating impacts of changing disturbance regimes. Our primary objective was to examine the patterns of fine-scale variation in understory plant communities and their relationships to above- and below-ground resource and environmental heterogeneity within mature lodgepole pine forests. We assessed composition and diversity of understory vegetation in relation to heterogeneity of both the above-ground (canopy tree density, canopy and tall shrub basal area and cover, downed wood biomass, litter cover) and below-ground (soil nutrient availability, decomposition, forest floor thickness, pH, and phospholipid fatty acids (PLFAs) and multiple carbon-source substrate-induced respiration (MSIR) of the forest floor microbial community) environment. There was notable variation in fine-scale plant community composition; cluster and indicator species analyses of the 24 most commonly occurring understory species distinguished four assemblages, one for which a pioneer forb species had the highest cover levels, and three others that were characterized by different bryophyte species having the highest cover. Constrained ordination (distance-based redundancy analysis) showed that two above-ground (mean tree diameter, litter cover) and eight below-ground (forest floor pH, plant available boron, microbial community composition and function as indicated by MSIR and PLFAs) properties were associated with variation in understory plant community composition. These results provide novel insights into the important ecological associations between understory plant community composition and heterogeneity in ecosystem properties and processes within forests dominated by a single canopy species.     

Does plant diversity increase top–down control of herbivorous insects in tropical forest?

Higher trophic level interactions are key mediators of ecosystem functioning in tropical forests. A rich body of theory has been developed to predict the effects of plant diversity on communities at higher trophic levels and the mechanisms underlying such effects. The 'enemies hypothesis’ states that predators exert more effective top–down control of herbivorous insects with increasing plant diversity. Support for this hypothesis has been found in temperate forests and agroecosystems, but remains understudied in tropical forests. We compared incidence of attacks of different natural enemies using artificial caterpillars in a tropical forest landscape and investigated the role of plant community structure (i.e. species richness, composition and density), and the role of forest fragmentation (i.e. patch size, edge distance and canopy openness) on predation intensity. Plant community effects were tested with respect to three vegetation strata: trees, saplings and herbs. Observed predation was substantially due to ants. Predation rates increased with plant species richness for trees and herbs. Density of saplings, herb cover and herb species composition were important factors for predation. No significant patterns were found for fragmentation parameters, suggesting that forest fragmentation has not altered predation intensity. We conclude that in tropical forests, top–down control of herbivorous insects in the understory vegetation is affected by a combination of plant diversity, plant species composition and structural features of the plant community.     

Diversity and composition of tropical butterflies along an Afromontane agricultural gradient in the Jimma Highlands,Ethiopia

Afromontane landscapes are typically characterized by a mosaic of smallholder farms and the biodiversity impacts of these practices will vary in accordance to local management and landscape context. Here, we assess how tropical butterfly diversity is maintained across an agricultural landscape in the Jimma Highlands of Ethiopia. We used transect surveys to sample understory butterfly communities within degraded natural forest, semi‐managed coffee forest (SMCF), exotic timber plantations, open woodland, croplands and pasture. Surveys were conducted in 29 one‐hectare plots and repeated five times between January and June 2013. We found that natural forest supports higher butterfly diversity than all agricultural plots (measured with Hill's numbers). SMCF and timber plantations retain relatively high abundance and diversity, but these metrics drop off sharply in open woodland, cropland and pasture. SMCF and timber plantations share the majority of their species with natural forest and support an equivalent abundance of forest‐dependent species, with no increase in widespread species. There was some incongruence in the responses of families and sub‐families, notably that Lycaenidae are strongly associated with open woodland and pasture. Adult butterflies clearly utilize forested agricultural practices such as SMCF and timber plantations, but species diversity declines steeply with distance from natural forest suggesting that earlier life‐stages may depend on host plants and/or microclimatic conditions that are lost under agricultural management. From a management perspective, the protection of natural forest remains a priority for tropical butterfly conservation, but understanding functioning of the wider landscape mosaic is important as SMCF and timber plantations may act as habitat corridors that facilitate movement between forest fragments.     

Vertical stratification of an avian community in New Guinean tropical rainforest

Vertical stratification of avian communities has been studied in both temperate and tropical forests; however, the majority of studies used ground-based methods. In this study we used ground-to-canopy mist nets to collect detailed data on vertical bird distribution in primary rain forest in Wanang Conservation Area in Papua New Guinea (Madang Province). In total 850 birds from 86 species were caught. Bird abundance was highest in the canopy followed by the understory and lowest in the midstory. Overall bird diversity increased towards the canopy zone. Insectivorous birds represented the most abundant and species-rich trophic guild and their abundances decreased from the ground to canopy. The highest diversity of frugivorous and omnivorous birds was confined to higher vertical strata. Insectivorous birds did not show any pattern of diversity along the vertical gradient. Further, insectivores preferred strata with thick vegetation, while abundance and diversity of frugivores increased with decreasing foliage density. Our ground-to-canopy (0–27 m) mist netting, when compared to standard ground mist netting (0–3 m), greatly improved bird diversity assessment and revealed interesting patterns of avian community stratification along vertical forest strata.     

Forest structure predicts species richness and functional diversity in Amazonian mixed-species bird flocks

Secondary forest has the potential to act as an important habitat for biodiversity and restoring ecological benefits. Functional diversity, which includes morphological and behavioral traits that mediate species interactions with the surrounding environment, relates to the resilience of ecosystems. To assess the relationship between habitat structural differences in primary and secondary forest and the resultant differences in functional diversity of avian species, we followed 11 mixed-species flocks at the Biological Dynamics of Forest Fragments Project, near Manaus, Brazil. We used remote sensing LiDAR to assess which three-dimensional forest structural features are most closely associated with variation in species richness and functional diversity in secondary and primary tropical forest flocks. The species richness of flocks in primary forest increased in areas with higher elevation and higher leaf area density in the understory and subcanopy but was not correlated with habitat structure in secondary forest. Functional diversity increased at lower elevations and with a denser subcanopy in both primary forest and secondary forest but only increased with greater understory leaf area density in primary forest. Together, these results indicate that a dense subcanopy and understory can be important for mixed-species flocks and that flock richness and functional diversity can be predicted by vegetation structure.     

Does forest loss affect the communities of trap-nesting wasps (Hymenoptera: Aculeata) in forests? Landscape vs. local habitat conditions

We investigated changes in the communities of trap-nesting Hymenoptera in forests in relation to forest loss on a landscape scale and understory conditions on a local habitat scale. Two specific questions were addressed. (1) Do the communities change with degrees of forest loss? (2) Do the communities change with varying local environmental conditions of understory habitats? The study was made in a landscape characterized by distributed forest patches within intensively managed agricultural surroundings. We deployed trap-nests at eight randomly selected sites in forests in summer. To quantify forest loss, the amount of forest coverage was calculated using GIS. To indicate local habitat conditions, the species richness of understory flowering plants was used. All together, 12 species of wasps and no bees were captured. Regression analyses showed that both abundance and species richness of the wasps were not significantly related to forest coverage. However, abundance of trap-nesting wasps was significantly related to species richness of understory plants, but species richness of the wasps was not significantly related to the plants. These results suggest that communities of trap-nesting wasps in forests are influenced more by the local habitat conditions than by forest loss.     

Upper-montane plant invasions in the Hawaiian Islands: Patterns and opportunities

In the Hawaiian Islands, massive volcanoes have created extreme elevation gradients, resulting in environments ranging from nearly tropical to alpine, spread across a distance of only a few dozen kilometers. Although the Hawaiian Islands are widely recognized for opportunities to study lowland tropical forest invasions, less attention has been paid to invasions of Hawaii's upper-montane forest, sub-alpine and alpine environments. This study synthesizes current knowledge of plant naturalization in upper-montane environments of the Hawaiian Islands in order to (1) determine whether patterns of tropical versus temperate species invasion change with elevation, and (2) evaluate whether upper-montane invaders are having significant impacts on native plant communities. A total of 151 naturalized plant species have been recorded at 2000 m or higher. Most species (93%) are herbaceous, and over half (52%) are native to Europe/Eurasia. Twenty-one species (14%) are reported to be disruptive in native plant communities, mainly by forming dense stands that appear to inhibit recruitment of natives, but also by altering vegetation structure or causing changes in ecosystem processes. Fourteen species (9%) were first recorded within the past 30 years, indicating that new invasions of upper-montane habitats are ongoing. At 1200 m elevation, only 38% of naturalized species are temperate in origin, but the proportion of temperate species increases linearly with elevation up to 3000 m (alpine habitat), where all naturalized species are temperate in origin and over 80% are native to Europe/Eurasia. Declining temperature along the elevation gradient probably drives this pattern. The extreme elevation gradients in the Hawaiian Islands provide specific opportunities for comparative studies on the ecology and evolution of temperate invaders while also creating a unique field environment for understanding interactions between temperate and tropical species.     

Avian Habitat Preference in Tropical Forest Restoration in Southern Costa Rica

An important question for tropical forest restoration is whether degraded lands can be actively managed to attract birds. We censused birds and measured vegetation structure at 27 stations in young (6–9‐yr old) actively and passively restored pasture and old growth forest at Las Cruces Biological Station in southern Costa Rica. During 481 10‐min point counts, we detected a high diversity—186 species—of birds using the restoration area. Surprisingly, species richness and detection frequency did not differ among habitats, and proportional similarity of bird assemblages to old growth forest did not differ between restoration treatments. Bird detection frequency was instead explained by exotic grass cover and understory stem density—vegetation structures that were not strongly impacted by active restoration. The similarity of bird assemblages in actively and passively restored forest may be attributed to differential habitat preferences within and among feeding guilds, low structural contrast between treatments, or the effect of nucleation from actively restored plots into passively restored areas. Rapid recovery of vegetation in this recently restored site is likely due to its proximity to old growth forest and the lack of barriers to effective seed dispersal. Previous restoration studies in highly binary environments (i.e., open pasture vs. tree plantation) have found strong differences in bird abundance and richness. Our data contradict this trend, and suggest that tropical restoration ecologists should carefully consider: (1) when the benefits of active restoration outweigh the cost of implementation; and (2) which avian guilds should be used to measure restoration success given differential responses to habitat structure.     

Seedling interactions in a tropical forest in Panama

Competition is believed to be a central force limiting local diversity and controlling the structure of plant communities. However, it has been proposed that the stressed understory environment limits total understory plant density to such low levels that competitive exclusion cannot be an important factor limiting the local diversity of understory plants. To evaluate the importance of inter-seedling competition, we performed a seedling competition experiment with five shade-tolerant species in a tropical moist forest in Panama. Three-month-old seedlings were transplanted into the forest singly or with their roots intertwined with a single conspecific or heterospecific seedling in all pairwise species combinations. If competition is important, performance (survival, stem height, and number of leaves after one and six years) would be expected to be lowest with a conspecific neighbor and greatest without a neighbor. The experiment was replicated in five 0.24-m² plots at each of 20 sites in tall secondary forest. To test whether seedling performance differed among treatments we fitted linear mixed models (LMM) and generalized linear mixed models (GLMM), treating species identity and microsite (site and plot) as random effects. The five shade-tolerant study species all experienced good establishment with relatively high survival and growth rates. The neighbor treatment consistently affected seedling performance, but the effect was always very small, both in absolute terms and relative to the much stronger species and microsite effects. Seedlings with a conspecific neighbor consistently performed worse than seedlings with a heterospecific neighbor, but having no neighbor generally did not cause superior performance relative to the other treatments. We conclude that direct competitive interactions are relatively unimportant among understory plants in humid tropical forests.     

Effects of experimental nitrogen additions on plant diversity in an old‐growth tropical forest

Response of plant biodiversity to increased availability of nitrogen (N) has been investigated in temperate and boreal forests, which are typically N‐limited, but little is known in tropical forests. We examined the effects of artificial N additions on plant diversity (species richness, density and cover) of the understory layer in an N saturated old‐growth tropical forest in southern China to test the following hypothesis: N additions decrease plant diversity in N saturated tropical forests primarily from N‐mediated changes in soil properties. Experimental additions of N were administered at the following levels from July 2003 to July 2008: no addition (Control); 50 kg N ha^?1 yr^?1 (Low‐N); 100 kg N ha^?1 yr^?1 (Medium‐N), and 150 kg N ha^?1 yr^?1 (High‐N). Results showed that no understory species exhibited positive growth response to any level of N addition during the study period. Although low‐to‐medium levels of N addition (≤100 kg N ha^?1 yr^?1) generally did not alter plant diversity through time, high levels of N addition significantly reduced species diversity. This decrease was most closely related to declines within tree seedling and fern functional groups, as well as to significant increases in soil acidity and Al mobility, and decreases in Ca availability and fine‐root biomass. This mechanism for loss of biodiversity provides sharp contrast to competition‐based mechanisms suggested in studies of understory communities in other forests. Our results suggest that high‐N additions can decrease plant diversity in tropical forests, but that this response may vary with rate of N addition.     

Fine-scale habitat structure complexity determines insectivorous bird diversity in a tropical forest

Habitat complexity in reforested stands has been acknowledged as a key factor that influences habitat use by birds, being especially critical for habitat disturbance-sensitive species such as tropical understory insectivorous birds. Most studies regarding the relationship between forest structure and species diversity were conducted at the landscape scale, but different diversity patterns may emerge at a finer scale (i.e., within a habitat patch). We examined a tropical reforested area (State of Caldas, Colombia), hypothesizing that insectivorous bird richness, abundance, and foraging guild abundance would increase as intra-habitat complexity increases. We established 40 monitoring plots within a reforested area, measured their structural features, and determined their relationships with species richness, total abundance, and foraging guild abundance, using Generalized Additive Models. We found that the increasing variation in basal area, stem diameter, and number of stems was positively correlated with species richness, total abundance, and foraging guild abundance. Relationships between richness or abundance and structural features were not lineal, but showing curvilinear responses and thresholds. Our results show that heterogeneity on basal area, stem diameter, and the number of stems was more correlated to insectivorous bird richness and abundance than the average of those structural features. Promoting structural variation on reforested areas by planting species with different growth rates may contribute to increase the richness and abundance of a tropical vulnerable group of species such as the understory insectivorous birds.