Hong-Zhang Zhou

Community assembly during secondary forest succession in a Chinese subtropical forest

Subtropical broad-leaved forests in southeastern China support a high diversity of woody plants. Using a comparative study design with 30 × 30 m plots (n = 27) from five successional stages ( 1 m in height in each plot and counted all woody recruits (bank of all seedlings ≤1 m in height) in each central 10 × 10 m quadrant of each plot. In addition, we measured a number of environmen...

Designing forest biodiversity experiments: general considerations illustrated by a new large experiment in subtropical China

Summary 1. Biodiversity–ecosystem functioning (BEF) experiments address ecosystem-level consequences of species loss by comparing communities of high species richness with communities from which species have been gradually eliminated. BEF experiments originally started with microcosms in the laboratory and with grassland ecosystems. A new frontier in experimental BEF research is manipulating tree diversity in forest ecosystems, compelling researchers to think big and comprehensively. 2. We present and discuss some of the major issues to be considered in the design of BEF experiments with trees and illustrate these with a new forest biodiversity experiment established in subtropical China (Xingangshan, Jiangxi Province) in 2009/2010. Using a pool of 40 tree species, extinction scenarios were simulated with tree richness levels of 1, 2, 4, 8 and 16 species on a total of 566 plots of 25� 8 9 25� 8m each. 3. The goal of this experiment is to estimate effects of tree and shrub species richness on carbon storage and soil erosion; therefore, the experiment was established on sloped terrain. The following important design choices were made: (i) establishing many small rather than fewer larger plots, (ii) using high planting density and random mixing of species rather than lower planting density and patchwise mixing of species, (iii) establishing a map of the initial ‘ecoscape’ to characterize site heterogeneity before the onset of biodiversity effects and (iv) manipulating tree species richness not only in random but also in trait-oriented extinction scenarios. 4. Data management and analysis are particularly challenging in BEF experiments with their hierarchical designs nesting individuals within-species populations within plots within-species compositions. Statistical analysis best proceeds by partitioning these random terms into fixed-term contrasts, for example, species composition into contrasts for species richness and the presence of particular functional groups, which can then be tested against the remaining random variation among compositions. 5. We conclude that forest BEF experiments provide exciting and timely research options. They especially require careful thinking to allow multiple disciplines to measure and analyse data jointly and effectively. Achiev

Predator diversity and abundance provide little support for the enemies hypothesis in forests of high tree diversity.

Predatory arthropods can exert strong top-down control on ecosystem functions. However, despite extensive theory and experimental manipulations of predator diversity, our knowledge about relationships between plant and predator diversity—and thus information on the relevance of experimental findings—for species-rich, natural ecosystems is limited. We studied activity abundance and species richness of epigeic spiders in a highly diverse forest ecosystem in subtropical China across 27 forest stands which formed a gradient in tree diversity of 25–69 species per plot. The enemies hypothesis predicts higher predator abundance and diversity, and concomitantly more effective top-down control of food webs, with increasing plant diversity. However, in our study, activity abundance and observed species richness of spiders decreased with increasing tree species richness. There was only a weak, non-significant relationship with tree richness when spider richness was rarefied, i.e. corrected for different total abundances of spiders. Only foraging guild richness (i.e. the diversity of hunting modes) of spiders was positively related to tree species richness. Plant species richness in the herb layer had no significant effects on spiders. Our results thus provide little support for the enemies hypothesis—derived from studies in less diverse ecosystems—of a positive relationship between predator and plant diversity. Our findings for an important group of generalist predators question whether stronger top-down control of food webs can be expected in the more plant diverse stands of our forest ecosystem. Biotic interactions could play important roles in mediating the observed relationships between spider and plant diversity, but further testing is required for a more detailed mechanistic understanding. Our findings have implications for evaluating the way in which theoretical predictions and experimental findings of functional predator effects apply to species-rich forest ecosystems, in which trophic interactions are often considered to be of crucial importance for the maintenance of high plant diversity.

Spatial and temporal variations in insect‐infested acorn fall in a Quercus liaotungensis forest in North China

Three plots with different aspects and slope characteristics were surveyed in 1999 and 2000 to clarify the spatial and temporal variations in insect-infested acorn fall patterns in a Quercus liaotungensis Koidz. forest in the Dongling Mountain region, North China. There was a significant difference in the proportion of infested acorns in the three plots in a low crop year, but not in a mast year. Within oakwoods on the southeast-facing slope, the insect infestation rate on the upper slope was significantly higher than on the lower slope, but not in the northwest-facing plot. Infestation rate in the low crop year in all three plots was significantly higher than infestation in the mast year. Most of the early fallen acorns had a higher proportion of insect infestation, and in the mast year it was much more obvious than in the low crop year. The proportion of infested acorns in seed bank along the topographic gradient showed a similar decreasing trend with acorn fall time, but the proportion on the upper slope was the highest and the proportion on the lower slope was the lowest. Larval emergence from acorns commenced just after acorns fell from the trees and lasted for 40–50 days, with peak emergence occurring from 24 to 32 days after acorn rain began. We conclude that insect-infested acorn distribution in Q. liaotungensis shows spatial and temporal heterogeneity, and an early drop of infested acorns can be a short-term defensive strategy against insect infestation.

Distribution of ground-dwelling beetle assemblages (Coleoptera) across ecotones between natural oak forests and mature pine plantations in North China

This paper studied edge effects resulting from logging to reforestation on the distribution of ground-dwelling beetles (Coleoptera) across ecotones between natural oak forests and mature pine plantations established after harvesting of natural forests. Using pitfall traps, ground-dwelling beetles were investigated at three replicated plots (ecotones) with three sampling positions of slope (lower, middle and upper) for each plot. Rarefaction estimates of species richness indicated that traps on natural forests and transition zones had more species than mature plantations did, and traps on the middle slope had more species than on the lower and upper slopes did. Results of an ANOVA analysis, which used forest type and slope position as factors and number of species and individuals as the response variables, showed a significant effect of forest type and slope position, and a significant interaction between forest type and slope position. Multivariate analyses (DCA and CCA) showed that beetles of transition zones were more similar to those of natural forests than to those of mature plantations, and that some environmental characteristics, i.e., proportion of broad-leaved trees, canopy cover and elevation (slope position), significantly affected species abundances. We conclude that the logging of natural oak forests and the reforestation of pine plantations can result in subtle variation in the composition and distribution of beetle assemblages at a local scale and such variation should be taken into account when conservation issues are involved.

Distribution of ground‐dwelling beetles (Coleoptera) across a forest‐clearcut ecotone in Wolong Natural Reserve, southwestern China

Abstract The influence of edge effect on ground‐dwelling beetles (Coleoptera) across a forest‐clearcut ecotone was studied in Wolong Natural Reserve, southwestern China. During the field research, a total of 30 739 beetles were collected with pitfall traps along transects, which extending 100 m from the edge into the forest interior and 100 m into the clearcut. Of the collection, Carabidae comprised 92%, Staphylinidae 3%, Curculionidae 2%, and Tenebrionidae 2%, and these four families can be considered as abundant groups. Family richness, Shannon diversity and equitability display a significant decrease from forest interior, edge to clearcut. Based on the family composition and abundance, ground‐dwelling beetles of the forest interior can be separated from those in the clearcut by Principal coordinate analysis ordination, and beetle assemblages in the forest edge were more similar to forest assemblages than to those found in the clearcut by cluster analysis. Seasonal dynamics of family richness showed a monotone peak in the middle season, with a highest value in the forest interior and a lowest value in the clearcut. Family abundance showed two peaks in the middle season, always with more individuals in the clearcut than in the forest interior or in the edge. Multiple linear regression analyses showed that the cover of shrub and litter were the two most important factors in determining family richness, Shannon diversity, equitability and abundance. Our results show that the forest edge and clearcut have obviously different composition and diversity of ground‐dwelling beetles from forest interior at the family level. However, more edges have been formed due to increasing forest fragmentation (clearcutting or logging), so it is necessary to preserve large and intact forest to protect the diversity of ground‐dwelling beetles in Wolong Natural Reserve.

3D visualization of the microstructure of Quedius beesoni Cameron using micro-CT

The investigation of the internal morphology of insects is usually performed using classical microtomy yielding optical micrographs of stained thin sections. The achievement of high-quality cross sections for microtomy is time-consuming and the risk of damaging sections is unavoidable. Moreover, the approach is impractical, in particular when quick acquisition of 3D structural information is required. Recently, X-ray computed microtomography (micro-CT) with a high spatial resolution was considered as a potential tool for the morphological classification of insects. We used micro-CT to investigate Quedius beesoni Cameron at the cellular length scale. This method provides a new powerful and nondestructive approach to obtain 3D structural information on the biological organization of insects. The preliminary images presented in this contribution clearly reveal the endoskeleton and the muscles of the head and the thorax with a full 3D structure. We also reconstructed the 3D structure of the brain of Quedius beesoni Cameron, and this is the first reconstruction in Staphylinidae, which will be a great advancement for morphological and phylogenic research. We claim that both the spatial resolution and the contrast characteristic of micro-CT imaging may fulfill the requirements necessary for zoological insect morphology and phylogeny, in particular, when a classification of a rare and unique insect specimen is required.

How Old Are the Rove Beetles (Insecta: Coleoptera: Staphylinidae) and Their Lineages? Seeking an Answer with DNA

The phylogeny and related evolutionary history of rove beetles (Coleoptera, Staphylinidae) remain unclear. This study provides phylogenetic analyses for the family based on three genes (mitochondrial COI, nuclear protein-coding wingless and a portion of the ribosomal 28S rDNA) including 2413 bp for 104 taxa representing most major staphylinid lineages. The subfamilies Oxyporinae, Paederinae, Steninae, and Proteininae are all well-supported clades, as evidenced by all three inference methods, namely maximum parsimony, Bayesian inference, and maximum likelihood. From fossils available for calibration, the divergence time of the main lineages in the family is estimated based on an uncorrelated lognormal relaxed molecular clock analysis method. The molecular clock analysis suggests that the family Staphylinidae dates from approximately the Early Triassic epoch and the most lineages of the family started to radiate from the Late Jurassic to the Early Paleogene.

Elevational Gradient in Species Richness Pattern of Epigaeic Beetles and Underlying Mechanisms at East Slope of Balang Mountain in Southwestern China

We report on the species richness patterns of epigaeic beetles (Coleoptera: Carabidae and Staphylinidae) along a subtropical elevational gradient of Balang Mountain, southwestern China. We tested the roles of environmental factors (e.g. temperature, area and litter cover) and direct biotic interactions (e.g. foods and antagonists) that shape elevational diversity gradients. Beetles were sampled at 19 sites using pitfall traps along the studied elevational gradient ranging from 1500 m–4000 m during the 2004 growing season. A total of 74416 specimens representing 260 species were recorded. Species richness of epigaeic beetles and two families showed unimodal patterns along the elevational gradient, peaking at mid-elevations (c. 2535 m), and the ranges of most beetle species were narrow along the gradient. The potential correlates of both species richness and environmental variables were examined using linear and second order polynomial regressions. The results showed that temperature, area and litter cover had strong explanatory power of beetle species richness for nearly all richness patterns, of beetles as a whole and of Carabidae and Staphylinidae, but the density of antagonists was associated with species richness of Carabidae only. Multiple regression analyses suggested that the three environmental factors combined contributed most to richness patterns for most taxa. The results suggest that environmental factors associated with temperature, area and habitat heterogeneity could account for most variation in richness pattern of epigaeic beetles. Additionally, the mid-elevation peaks and the small range size of most species indicate that conservation efforts should give attention to the entire gradient rather than just mid-elevations.

Composition and distribution of ground‐dwelling beetles among oak fragments and surrounding pine plantations in a temperate forest of North China

In this study, we compared ground‐dwelling beetle assemblages (Coleoptera) from a range of different oak fragments and surrounding conifer plantations to evaluate effects of forest size and surrounding matrix habitat in a temperate forest of north China. During 2000, beetles were sampled via pitfall traps within two large oak fragments (ca. 2.0−4.0 ha), two small oak fragments (ca. 0.2−0.4 ha) and two surrounding matrices dominated by pine plantations (>4 ha) in two sites of different aspects. Overall, no significantly negative effects from forest patch size and the surrounding matrix habitat were detected in total species number and abundance of ground‐dwelling beetles. However, compared with small oak patches or pine plantations, more species were associated with an affinity for at least one large oak patch of the two aspects. Multivariate regression trees showed that the habitat type better determined the beetle assemblage structure than patch size and aspect, indicating a strong impact of the surrounding matrix. Linear mixed models indicated that species richness and abundance of all ground‐dwelling beetles or beetle families showed different responses to the selected environmental variables. Our results suggest that more disturbed sites are significantly poorer in oak forest specialists, which are usually more abundant in large oak fragments and decrease in abundance or disappear in small fragments and surrounding matrix habitats. Thus, it is necessary to preserve a minimum size of forest patch to create conditions characteristic for forest interior, rather than the more difficult task of increasing habitat connectivity.