Tree functional diversity affects litter decomposition and arthropod community composition in a tropical forest

Disturbance can alter tree species and functional diversity in tropical forests, which in turn could affect carbon and nutrient cycling via the decomposition of plant litter. However, the influence of tropical tree diversity on forest floor organisms and the processes they mediate are far from clear. We investigated the influence of different litter mixtures on arthropod communities and decomposition processes in a 60‐year‐old lowland tropical forest in Panama, Central America. We used litter mixtures representing pioneer and old growth tree species in experimental mesocosms to assess the links between litter types, decomposition rates, and litter arthropod communities. Overall, pioneer species litter decomposed most rapidly and old growth species litter decomposed the slowest but there were clear non‐additive effects of litter mixtures containing both functional groups. We observed distinct arthropod communities in different litter mixtures at 6 mo, with greater arthropod diversity and abundance in litter from old growth forest species. By comparing the decay of different litter mixtures in mesocosms and conventional litterbags, we demonstrated that our mesocosms represent an effective approach to link studies of litter decomposition and arthropod communities. Our results indicate that changes in the functional diversity of litter could have wider implications for arthropod communities and ecosystem functioning in tropical forests.     

Leaf litter arthropod responses to tropical forest restoration

Soil and litter arthropods represent a large proportion of tropical biodiversity and perform important ecosystem functions, but little is known about the efficacy of different tropical forest restoration strategies in facilitating their recovery in degraded habitats. We sampled arthropods in four 7‐ to 8‐year‐old restoration treatments and in nearby reference forests. Sampling was conducted during the wet and dry seasons using extractions from litter and pitfall samples. Restoration treatments were replicated in 50 × 50‐m plots in four former pasture sites in southern Costa Rica: plantation – trees planted throughout the plot; applied nucleation/islands – trees planted in patches of different sizes; and natural regeneration – no tree planting. Arthropod abundance, measures of richness and diversity, and a number of functional groups were greater in the island treatment than in natural regeneration or plantation treatments and, in many cases, were similar to reference forest. Litter and pitfall morphospecies and functional group composition in all three restoration treatments were significantly different than reference sites, but island and plantation treatments showed more recovery than natural regeneration. Abundance and functional group diversity showed a much greater degree of recovery than community composition. Synthesis and applications: The less resource‐intensive restoration strategy of planting tree islands was more effective than tree plantations in restoring arthropod abundance, richness, and functional diversity. None of the restoration strategies, however, resulted in similar community composition as reference forest after 8 years of recovery, highlighting the slow rate of recovery of arthropod communities after disturbance, and underscoring the importance of conservation of remnant forests in fragmented landscapes.     

Disparate effects of plant genotypic diversity on foliage and litter arthropod communities

Intraspecific diversity can influence the structure of associated communities, though whether litter-based and foliage-based arthropod communities respond to intraspecific diversity in similar ways remains unclear. In this study, we compared the effects of host-plant genotype and genotypic diversity of the perennial plant, Solidago altissima, on the arthropod community associated with living plant tissue (foliage-based community) and microarthropods associated with leaf litter (litter-based community). We found that variation among host-plant genotypes had strong effects on the diversity and composition of foliage-based arthropods, but only weak effects on litter-based microarthropods. Furthermore, host-plant genotypic diversity was positively related to the abundance and diversity of foliage-based arthropods, and within the herbivore and predator trophic levels. In contrast, there were minimal effects of plant genotypic diversity on litter-based microarthropods in any trophic level. Our study illustrates that incorporating communities associated with living foliage and senesced litter into studies of community genetics can lead to very different conclusions about the importance of intraspecific diversity than when only foliage-based community responses are considered in isolation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

不同演替阶段热带森林地表凋落物和土壤节肢动物群落特征

为了解不同演替阶段热带森林土壤节肢动物群落结构特征及其与地表凋落物的关系, 2001年9月采用样线调查法对西双版纳23年次生林、35年次生林、季节雨林地表凋落物及其中的土壤节肢动物进行了调查。所获数据表明, 地表凋落物数量(现存量干重)和质量(N和C/N)总体上表现为35年次生林最好, 23年次生林次之; 蜱螨目和弹尾目为3林地地表凋落物土壤节肢动物群落优势类群, 膜翅目蚂蚁、马陆目、鞘翅目、双翅目和半翅目为常见类群。土壤节肢动物个体密度和个体相对密度均表现为35年次生林>季节雨林>23年次生林。群落的丰富度指数以季节雨林最高, 多样性和均匀度指数显示为23年次生林最高, 35年次生林的优势度指数最高, 3林地土壤节肢动物群落类群组成相似性达到较好水平。相关分析表明, 3种不同演替阶段热带森林土壤节肢动物个体密度与林地地表凋落物现存量呈正相关, 而现存凋落物N元素储量与土壤节肢动物的相关性仅表现在23年次生林和季节雨林。研究认为, 热带森林土壤节肢动物群落的发展与森林植被演替密切相关, 其群落个体数量和多样性受森林地表凋落物数量、质量的调控, 但其他环境因素如捕食效应、人为干扰等影响亦不可忽视。     

Stable baselines of temporal turnover underlie high beta diversity in tropical arthropod communities

While the high species diversity of tropical arthropod communities has often been linked to marked spatial heterogeneity, their temporal dynamics have received little attention. This study addresses this gap by examining spatio‐temporal variation in the arthropod communities of a tropical montane forest in Honduras. By employing DNA barcode analysis and Malaise trap sampling across 4 years and five sites, 51,596 specimens were assigned to 8,193 presumptive species. High beta diversity was linked more strongly to elevation than geographic distance, decreasing by 12% when only the dominant species were considered. When sampling effort was increased by deploying more traps at a site, beta diversity only decreased by 2%, but extending sampling across years decreased beta diversity by 27%. Species inconsistently detected among years, likely transients from other settings, drove the low similarity in species composition among traps only a few metres apart. The dominant, temporally persistent species substantially influenced the cyclic pattern of change in community composition among years. This pattern likely results from divergence–convergence dynamics, suggesting a stable baseline of temporal turnover in each community. The overall results establish that large sample sizes are necessary to reveal species richness, but are not essential for quantifying beta diversity. This study further highlights the need for standardized methods of sampling and species identification to generate the comparative data required to evaluate biodiversity change in space and time.     

Metabarcoding as a tool for investigating arthropod diversity in Nepenthes pitcher plants

The biodiversity of tropical forests consists primarily of small organisms that are difficult to detect and characterize. Next‐generation sequencing (NGS) methods can facilitate analyses of these arthropod and microbial communities, leading to a better understanding of existing diversity and factors influencing community assembly. The pitchers of carnivorous pitcher plants often house surprisingly discrete communities and provide ideal systems for analysis using an NGS approach. The plants digest insects in order to access essential nutrients while growing in poor soils; however, the pitchers are also home to communities of living organisms, called inquilines. Certain arthropods appear to have coevolved with their pitcher plant hosts and are not found in other environments. We used Illumina amplicon sequencing of 18S rDNA to characterize the eukaryotes in three species of Nepenthes (Nepenthaceae) pitcher plants – N. gracilis, N. rafflesiana and N. ampullaria – in each of three different parks in Singapore. The data reveal an unexpected diversity of eukaryotes, significant differences in community diversity among host species, variation in host specificity of inquilines and the presence of gregarine parasites. Counts of whole inquiline arthropods from the first collection year were roughly correlated with scaled 18S sequence abundances, indicating that amplicon sequencing is an effective means of gauging community structure. We barcoded a subset of the dipteran larvae using COI primers, and the resulting phylogenetic tree is mostly congruent with that found using the 18S locus, with the exception of one of five morphospecies. For many 18S and COI sequences, the best BLASTn matches showed low sequence identity, illustrating the need for better databases of Southeast Asian dipterans. Finally, networks of core arthropods and their host species were used to investigate degree of host specificity across multiple hosts, and this revealed significant specialization of certain arthropod fauna.     

Effects of carrion decomposition on litter arthropod assemblages

1. Many organisms contribute to the decomposition of carrion. For arthropods, many studies focus on the necrophagous community, those directly consuming carrion.
2. Necrophagous arthropods use carrion as a shelter or food source. Therefore, carrion generally increases the abundance and biodiversity of necrophagous species. However, it is unclear if carrion has similar effects on detrital communities.
3. This study examines changes in community structure and composition of necrophilous and detrital communities over the course of decomposition.
4. Five pig head carrion were placed at least 7 m apart under cages in temperate mixed forest. Leaf litter was sampled 0 m, 1.5 m, and 3 m from each carrion weekly during summer, and monthly during autumn, until the first frost. Arthropods were extracted from leaf litter by using Berlese funnels.
5. At the carrion site, necrophagous insects increased in abundance, species richness, and Shannon diversity during decomposition, and all decreased after dry decay.
6. Detritus arthropods displayed a sharp increase in abundance during advanced decay, decreasing during dry decay, and a general increase over time for species richness and diversity.
7. In conclusion, carrion can influence the surrounding, non-necrophagous arthropod community, highlighting the need to investigate carrion effects beyond typical necrophagous species to have a more holistic understanding of carrion ecology.

     

Effects of bund crops and insecticide treatments on arthropod diversity and herbivore regulation in tropical rice fields

Ecological engineering using vegetable or flower strips is promoted as a potential pest management strategy in irrigated rice. Farmers in the Philippines often plant rice levees (bunds) with vegetables, particularly string beans (Vigna unguiculata [L.] Walpers) to supplement income, but without considering the potential for pest management. This study examines the effects of planted bunds on rice herbivores and their natural enemies. We compared arthropods in (a) rice fields that had string beans planted on bunds, (b) fields without string beans and without any insecticide applications and (c) fields without string beans but with insecticide treatments (standard practice). Rice yield was similar across all treatments; however, the vegetation strips produced an extra 3.6 kg of fresh string bean pods per metre of bund. There were no apparent increases in major natural enemy groups in fields with string beans compared to fields with conventional bunds. Fields with insecticide treatments had higher damage from leaffolders (Lepidoptera: Pyralidae). The sprayed fields also had lower parasitism of planthopper eggs and fewer predatory dragonflies and damselflies (Odonata). Furthermore, the mortality of planthopper (Delphacidae: Hemiptera) and stemborer (Pyralidae) eggs by parasitoids and predators was density dependent only in the unsprayed fields (with and without string beans). Our results demonstrate that planting string beans on rice bunds improves the productivity of rice farms, but our ecological engineering system did not appreciably affect natural enemy or herbivore abundance; however, chemical insecticides adversely affected pest regulatory ecosystem functions leading to higher pest damage.     

Unique arthropod communities on different host-plant genotypes results in greater arthropod diversity

Studies on the effect of plant-species diversity on various ecological processes has led to the study of the effects of plant-genetic diversity in the context of community genetics. Arthropod diversity can increase with plant-species or plant-genetic diversity (Wimp et al. in Ecol Lett 7:776–780, 2004). Plant diversity effects can be difficult to separate from other ecological processes, for example, complementarity. We asked three basic questions: (1) Are arthropod communities unique on different host-plant genotypes? (2) Is arthropod diversity greater when associated with greater plant-genetic diversity? (3) Are arthropod communities more closely associated with host-plant genetics than the plant neighborhood? We studied canopy arthropods on Populus fremontii trees randomly planted in a common garden. All trees were planted in a homogeneous matrix, which helped to reduce P. fremontii neighborhood effects. One sample was comprised of few P. fremontii genotypes with many clones. A second sample was comprised of many P. fremontii genotypes with few clones. A second data set was used to examine the relationships between the arthropod community with P. fremontii genetic composition and the neighborhood composition of the focal host plant. Unique arthropod communities were associated with different P. fremontii genotypes, and arthropod community diversity was greater in the sample with greater P. fremontii genotypic diversity. Arthropod community similarity was negatively correlated with P. fremontii genetic distance, but arthropod community similarity was not related to the neighborhood of the P. fremontii host plant.     

Ecology and diversity of leaf litter fungi during early-stage decomposition in a seasonally dry tropical forest

Leaf litter samples of 12 dicotyledonous tree species (belonging to eight families) growing in a dry tropical forest and in early stages of decomposition were studied for the presence of litter fungi. Equal-sized segments of the leaves incubated in moist chambers were observed every day for 30 d for the presence of fungi. Invariably, the fungal assemblage on the litter of each tree species was dominated by a given fungal species. The diversity of fungi present in the litter varied with the tree species although many species of fungi occurred in the litter of all 12 species. A Pestalotiopsis species dominated the litter fungal assemblage of five trees and was common in the litter of all tree species. The present study and earlier studies from our lab indicate that fungi have evolved traits such as thermotolerant spores, ability to utilize toxic furaldehydes, ability to produce cell wall destructuring enzymes and an endophyte-litter fungus life style to survive and establish themselves in fire-prone forests such as the one studied here. This study shows that in the dry tropical forest, the leaf litter fungal assemblage is governed more by the environment than by the plant species.     

Effects of litter removal on arthropod communities in pine plantations

Natural and anthropogenic disturbances can cause abrupt changes in trophic interactions by altering the rate, timing, or composition of organic inputs to ecological systems which in turn can shift patterns of species dominance. We examined the short-term effects of litter removal on soil fauna in pine plantations of three different species (longleaf, Pinus palustris; loblolly, P. taeda; and slash, P. elliottii) using a manipulative experiment, with the goal of examining differences among dominant orders of arthropods and differences among timber types. We sampled arthropods once per month for 6 months immediately following raking, and found that removal of the litter caused significant changes to abundance or presence of five of the nine dominant orders. Reductions in abundances of arthropod orders were most apparent in loblolly pine stands, while increases in abundance were more common in longleaf and slash pine stands. The differential impact among orders suggests that removal of the litter layer is likely to alter trophic interactions by changing the relative abundance of functional groups. Repeated litter removal via raking could have negative repercussions on ecosystem stability. Finally, nutrient additions through fertilization seem unlikely to mitigate the changes imposed on the arthropod community through litter removal.     

Leaf litter identity rather than diversity shapes microbial functions and microarthropod abundance in tropical montane rainforests

In tropical forest ecosystems leaf litter from a large variety of species enters the decomposer system, however, the impact of leaf litter diversity on the abundance and activity of soil organisms during decomposition is little known. We investigated the effect of leaf litter diversity and identity on microbial functions and the abundance of microarthropods in Ecuadorian tropical montane rainforests. We used litterbags filled with leaves of six native tree species (Cecropia andina, Dictyocaryum lamarckianum, Myrcia pubescens, Cavendishia zamorensis, Graffenrieda emarginata, and Clusia spp.) and incubated monocultures and all possible two‐ and four‐species combinations in the field for 6 and 12 months. Mass loss, microbial biomass, basal respiration, metabolic quotient, and the slope of microbial growth after glucose addition, as well as the abundance of microarthropods (Acari and Collembola), were measured at both sampling dates. Leaf litter diversity significantly increased mass loss after 6 months of exposure, but reduced microbial biomass after 12 months of exposure. Leaf litter species identity significantly changed both microbial activity and microarthropod abundance with species of high quality (low C‐to‐N ratio), such as C. andina, improving resource quality as indicated by lower metabolic quotient and higher abundance of microarthropods. Nonetheless, species of low quality, such as Clusia spp., also increased the abundance of Oribatida suggesting that leaf litter chemical composition alone is insufficient to explain variation in the abundances of soil microarthropods. Overall, the results provide evidence that decomposition and microbial biomass in litter respond to leaf litter diversity as well as litter identity (chemical and physical characteristics), while microarthropods respond only to litter identity but not litter diversity.     

An invasive plant-fungal mutualism reduces arthropod diversity

Ecological theory holds that competition and predation are the most important biotic forces affecting the composition of communities. Here, we expand this framework by demonstrating that mutualism can fundamentally alter community and food web structure. In large, replicated field plots, we manipulated the mutualism between a dominant plant ( Lolium arundinaceum ) and symbiotic fungal endophyte ( Neotyphodium coenophialum ). The presence of the mutualism reduced arthropod abundance up to 70%, reduced arthropod diversity nearly 20%, shifted arthropod species composition relative to endophyte-free plots and suppressed the biomass and richness of other plant species in the community. Herbivorous arthropods were more strongly affected than carnivores, and for both herbivores and carnivores, effects of the mutualism appeared to propagate indirectly via organisms occurring more basally in the food web. The influence of the mutualism was as great or greater than previously documented effects of competition and predation on arthropod communities. Our work demonstrates that a keystone mutualism can significantly reduce arthropod biodiversity at a broad community scale.     

Capturing arthropod diversity in complex cave systems

Aim

Identify the optimal combination of sampling techniques to maximize the detection of diversity of cave‐dwelling arthropods.

Location

Central‐western New Mexico; north‐western Arizona; Rapa Nui, Chile.

Methods

From 26 caves across three geographically distinct areas in the Western Hemisphere, arthropods were sampled using opportunistic collecting, timed searches, and baited pitfall trapping in all caves, and direct intuitive searches and bait sampling at select caves. To elucidate the techniques or combination of techniques for maximizing sampling completeness and efficiency, we examined our sampling results using nonmetric multidimensional scaling (NMDS), analysis of similarity (ANOSIM), Wilcoxon signed‐rank tests, species richness estimators and species accumulation curves.

Results

To maximize the detection of cave‐dwelling arthropod species, one must apply multiple sampling techniques and specifically sample unique microhabitats. For example, by sampling cave deep zones and nutrient resource sites, we identified several undescribed cave‐adapted and/or cave‐restricted taxa in the south‐western United States and eight new species of presumed cave‐restricted arthropods on Rapa Nui that would otherwise have been missed. Sampling techniques differed in their detection of both management concern species (e.g., newly discovered cave‐adapted/restricted species, range expansions of cave‐restricted species and newly confirmed alien species) and specific taxonomic groups. Spiders were detected primarily with visual search techniques (direct intuitive searches, opportunistic collecting and timed searches), while most beetles were detected using pitfall traps. Each sampling technique uniquely identified species of management concern further strengthening the importance of a multi‐technique sampling approach.

Main conclusions

Multiple sampling techniques were required to best characterize cave arthropod diversity. For techniques applied uniformly across all caves, each technique uniquely detected between ~40% and 67% of the total species observed. Also, sampling cave deep zones and nutrient resource sites was critical for both increasing the number of species detected and maximizing the likelihood of detecting management concern species.

     

转基因玉米对田间节肢动物群落多样性的影响

通过对转基因耐除草剂(EPSPS)抗虫(Cry1Ab)玉米转化体‘DBN9936’、受体玉米‘DBN318’、常规玉米‘先玉335’和喷施除草剂的转化体‘DBN9936’玉米田中节肢动物种类及数量的调查, 评价转基因玉米对田间节肢动物群落多样性的影响。2015年和2017年我们采用直接观察法、陷阱调查法和剖秆法对田间节肢动物进行调查, 采用聚类分析、物种累积曲线等方法对数据进行分析, 并比较了4个处理玉米田节肢动物群落的Margalef丰富度指数、Shannon-Wiener多样性指数、Simpson多样性指数、Pielou均匀度指数、优势集中性指数和群落相似性指数的差异及其随时间变化的规律。调查期间共记录节肢动物20目80科; 转化体玉米‘DBN9936’ (2015: 10.3 ± 2.6头, 2017: 3.3 ± 1.7头)和喷施除草剂的转化体玉米‘DBN9936’ (2015: 6.0 ± 1.5头, 2017: 17.0 ± 0.6头)上鳞翅目昆虫的数量明显低于受体‘DBN318’ (2015: 20.0 ± 3.2头, 2017: 24.0 ± 6.0头)和‘先玉335’ (2015: 21.0 ± 8.9头, 2017: 26.7 ± 2.0头); 物种累积曲线呈典型的抛物线, 各类玉米田间总体物种丰富度差异较小; 玉米生育期节肢动物调查结果累计数量的功能团组成及其丰富度、多样性、均匀度、优势集中性间均无明显的差异, 各类指数随时间变化的动态趋于一致, 群落间相似性程度较高。转基因玉米‘DBN9936’对鳞翅目害虫有明显的抗性, 对非靶标节肢动物无显著的影响, 对田间节肢动物的群落多样性、均匀度、丰富度、优势集中性等没有明显的影响。     

Assembly mechanisms shaping tropical litter ant communities

Community ecology seeks to unravel the mechanisms that allow species to coexist in space. Some of the contending mechanisms may generate tractable signatures in the amount of trait and phylogenetic dispersion among co‐existing species. When a community presents a pattern with reduced trait or phylogenetic dispersion, mechanisms based on ecological filters are brought into consideration. On the other hand, limiting similarity mechanisms such as competitive exclusion are proposed when communities present patterns of trait or phylogenetic even‐dispersion. The strength of these mechanisms likely varies with the spatial scale of an observed sample. I surveyed species‐rich tropical litter ant communities in a spatially nested design that allowed me to explore the spatial scales, fine (0.25 m²), intermediate (9 m²), and broad (361 m²) at which these mechanisms act. I then assessed the relationship between observed ant communities and potential species pools ranging in size, from plot, site, and island‐wide areas. Patterns of phylogenetic dispersion within ant communities suggested that ant communities were composed of species that were more closely related than expected by a random sampling of phylogenetic pools. The magnitude of phylogenetic ‘clustering’ increased with the size of the species pool but was similar among communities assembled from different spatial scales. Patterns of dispersion of one ecological trait (i.e. body size) within ant communities also showed clustering of body sizes, and most communities were composed of ant species that were smaller than expected by a random sampling of trait pools. Trait clustering increased with the size of the species pool but decreased at broad spatial scales. Together, these results suggest that ecological filters, not interspecific interactions, are structuring tropical ant communities, favoring clades with small worker sizes. The larger dependency on the size of regional pools than on the spatial scale suggests that environmental heterogeneity is greater among than within the study sites.     

Experimental tests of the dependence of arthropod diversity on plant diversity

ABSTRACT Because a diversity of resources should support a diversity of consumers, most models predict that increasing plant diversity increases animal diversity. We report results of a direct experimental test of the dependence of animal diversity on plant diversity. We sampled arthropods in a well-replicated grassland experiment in which plant species richness and plant functional richness were directly manipulated. In simple regressions, both the number of species planted ([Formula: see text] transformed) and the number of functional groups planted significantly increased arthropod species richness but not arthropod abundance. However, the number of species planted was the only significant predictor of arthropod species richness when both predictor variables were included in ANOVAs or a MANOVA. Although highly significant, arthropod species richness regressions had low [Formula: see text] values, high intercepts (24 arthropod species in monocultures), and shallow slopes. Analyses of relations among plants and arthropod trophic groups indicated that herbivore diversity was influenced by plant, parasite, and predator diversity. Furthermore, herbivore diversity was more strongly correlated with parasite and predator diversity than with plant diversity. Together with regression results, this suggests that, although increasing plant diversity significantly increased arthropod diversity, local herbivore diversity is also maintained by, and in turn maintains, a diversity of parasites and predators.