Andreas Floren

Arthropod diversity in a tropical forest

Assessing Creepy Crawlies Arthropods are the most diverse group of terrestrial animal species, yet estimates of the total number of arthropod species have varied widely, especially for tropical forests. Basset et al. (p. 1481, see the cover) now provide more reliable estimates of total arthropod species richness in a tropical rainforest in Panama. Intensive sampling of a half hectare of forest yielded just over 6000 arthropod species. Scaling up this result to the whole forest suggests that the total species diversity lies between 17,000 and 40,000 species. Total arthropod species richness in a tropical rainforest can be best predicted by plant diversity. Most eukaryotic organisms are arthropods. Yet, their diversity in rich terrestrial ecosystems is still unknown. Here we produce tangible estimates of the total species richness of arthropods in a tropical rainforest. Using a comprehensive range of structured protocols, we sampled the phylogenetic breadth of arthropod taxa from the soil to the forest canopy in the San Lorenzo forest, Panama. We collected 6144 arthropod species from 0.48 hectare and extrapolated total species richness to larger areas on the basis of competing models. The whole 6000-hectare forest reserve most likely sustains 25,000 arthropod species. Notably, just 1 hectare of rainforest yields >60% of the arthropod biodiversity held in the wider landscape. Models based on plant diversity fitted the accumulated species richness of both herbivore and nonherbivore taxa exceptionally well. This lends credence to global estimates of arthropod biodiversity developed from plant models.

The Importance of Primary Tropical Rain Forest For Species Diversity: An Investigation Using Arboreal Ants as an example

The arboreal ant communities of a primary lowland rain forest and three differently disturbed forests that lie close together forming an anthropogenic disturbance gradient were collected with insecticide fogging. Combined samples from all trees (87 foggings) comprised 153,504 ants sorted to 331 morphospecies. The primary forest ant fauna was characterized by high species richness and 53 foggings were necessary to collect communities representatively. Another 63 species of ants were found in the disturbed forests indicating a large regional species pool that might exceed 420 species of arboreal ants. Anthropogenic disturbance caused a change in the taxonomic composition, diversity and structure of ant communities. Community size was a predictor of species richness in the severely disturbed forest types but not in the old secondary or primary forest. Ant abundance had declined significantly in the disturbed forests and only 10% of the primary forest’s species were collected in the most disturbed forest type. In each of the secondary forests a change in the frequency distribution of species was observed and a small number of species had gained numerical dominance. Analysis of species associations indicates that the strength of species interactions changed with the degree of forest disturbance. These changes were still clearly recognizable after 40 years of forest regeneration despite optimal conditions for colonization from the adjacent primary forest, demonstrating that the time scale needed for forest recovery after anthropogenic disturbance is very long.

The influence of anthropogenic disturbances on the structure of arboreal arthropod communities

In the framework of our research, aimed at understanding the processes structuring tropical arthropod communities, we investigated the changes occurring in tree crown communities of forests of different disturbance levels. These were a mixed dipterocarp primary lowland rain forest in Kinabalu National Park (in Sabah, Malaysia) and, some kilometres away, three forests of regeneration periods 5, 15, and 40 years following a clear-cut. These disturbed forest sites were adjacent to one another and merged into mature forest. From each forest at least ten individuals of one tree species were sampled using the fogging method. In the primary forest relative proportions of some arthropod taxa differed on the ordinal and familial level significantly within trees. The dominance of Formicidae was characteristic as was the almost complete lack of less mobile arthropods such as Lepidoptera larvae. In the five-year- old forest, differences in relative proportions among most taxa had almost disappeared. Formicidae abundances had declined drastically which coincided with an increase of Lepidoptera larvae. With progressing forest succession, arthropod communities increasingly converged on the pattern of primary forest, and total ant abundance as well as diversity increased significantly. Ant communities in the most disturbed forest were of low structural complexity, and to a large degree predictable in species arrangement, but became more and more unpredictable as the complexity of the forest increased. Several species of Coleoptera and non-formicine Hymenoptera occurred in high numbers in the youngest forest, contrasting with the mature forest where all species were typically rare. These changes may indicate a change in the structuring mechanisms from predominantly deterministic processes in disturbed forests to stochastic processes in mature forest.

DIVERSITY OF ARBOREAL SPIDERS IN PRIMARY AND DISTURBED TROPICAL FORESTS

Abstract This study investigates how arboreal spider communities in SE-Asian primary lowland rain forests change after anthropogenic disturbance. Two types of secondary forests were distinguished: 1) forests adjacent to each other, which finally merged into primary forest and 2) forests that were isolated by at least 10 km from the primary forest. Three forests of different age were investigated from each type and compared with undisturbed primary forest. All disturbed forests had been used some years for agriculture and were then left between 5 and 50 years to regenerate naturally. Spiders from at least seven trees per forest type were collected using insecticidal knockdown fogging and sorted to species or morphospecies level. Spiders represented between 5–10% of all canopy arthropods. A similar number of spiders were collected per square meter from all trees. However, communities in the primary forest differed greatly in their alpha- and beta-diversity and in community structure from those in the disturbed forest types. Diversity was high in the regenerating forests connected to the primary forest and approximated the conditions of the primary forest during the course of forest succession. In contrast, the isolated forests were of low diversity and communities showed little change during forest regeneration. These results indicate the importance of a species-source from which disturbed forests can be recolonized. However, even under optimal conditions this process needed decades before spider communities became similar to those of the primary forest. With no species-source available, spider diversity changed little during 50 years of forest regeneration. In the isolated forest we observed a drastic turnover from forest species towards species characteristic of open vegetation and shrubs. Our results give an indication of how large a loss in diversity can be expected in isolated forest fragments.

Arthropod Distribution in a Tropical Rainforest: Tackling a Four Dimensional Puzzle

Quantifying the spatio-temporal distribution of arthropods in tropical rainforests represents a first step towards scrutinizing the global distribution of biodiversity on Earth. To date most studies have focused on narrow taxonomic groups or lack a design that allows partitioning of the components of diversity. Here, we consider an exceptionally large dataset (113,952 individuals representing 5,858 species), obtained from the San Lorenzo forest in Panama, where the phylogenetic breadth of arthropod taxa was surveyed using 14 protocols targeting the soil, litter, understory, lower and upper canopy habitats, replicated across seasons in 2003 and 2004. This dataset is used to explore the relative influence of horizontal, vertical and seasonal drivers of arthropod distribution in this forest. We considered arthropod abundance, observed and estimated species richness, additive decomposition of species richness, multiplicative partitioning of species diversity, variation in species composition, species turnover and guild structure as components of diversity. At the scale of our study (2km of distance, 40m in height and 400 days), the effects related to the vertical and seasonal dimensions were most important. Most adult arthropods were collected from the soil/litter or the upper canopy and species richness was highest in the canopy. We compared the distribution of arthropods and trees within our study system. Effects related to the seasonal dimension were stronger for arthropods than for trees. We conclude that: (1) models of beta diversity developed for tropical trees are unlikely to be applicable to tropical arthropods; (2) it is imperative that estimates of global biodiversity derived from mass collecting of arthropods in tropical rainforests embrace the strong vertical and seasonal partitioning observed here; and (3) given the high species turnover observed between seasons, global climate change may have severe consequences for rainforest arthropods.

Forest management and regional tree composition drive the host preference of saproxylic beetle communities

Summary 1. Among saproxylic beetles, many early colonizers prefer particular host species. Ranking of preferred hosts of local saproxylic beetle communities is critical for effective dead-wood management in forests, but is rarely done because experiments with numerous tree species are labour and cost intensive. 2. We analysed the host preference of local saproxylic beetle communities on logs of 13 tree species in relation to management (unmanaged and managed beech stands, conifer plantations on natural beech sites) in three regions of Germany during the most critical period for host specificity, that is the first two years after harvesting. Hosts were ranked quantitatively based on the ordinal ranking of hosts by single beetle species, which in turn was based on the variation in beetle abundance. First, we employed a Bradley–Terry model in which ranking was derived from paired comparisons of host trees. Then, a model-based recursive partitioning of the Bradley–Terry model tested whether host preference of beetle communities is affected by stand management, region and decay progress of dead wood. 3. Our results indicated that beetle communities overall avoided logs of Fraxinus, Pseudotsuga, Larix and Tilia, and Carpinus ranked highest in preference. Carpinus also ranked highest for communities of broadleaf specialists; Picea abies ranked highest for communities of conifer specialists. Model-based recursive partitioning revealed that ranking of local hosts in conifer stands differed from that of broadleaf stands, and that ranking of hosts in broadleaf stands differed between regions, but only in the first year for both. 4. Synthesis and applications. Early-colonizing saproxylic beetle communities vary locally in their choice of host trees. Therefore, forest managers should focus on the enrichment of dead wood of regional tree species and tree species of the local stand to successfully promote earlycolonizing beetle.

Ants on oaks: effects of forest structure on species composition

The ant fauna of oak forest canopies in Northern Bavaria was studied by canopy fogging on 45 trees in August 2000 and May 2001. The study focused on a comparison of several different forestry management practices resulting in several types of canopy cover. Forests surveyed were: (1) high forest (high canopy cover), (2) coppice with standards (low canopy cover), (3) forest pasture with mostly solitary trees (very low canopy cover) and (4) transitional forest from former coppice with standards to high forest (approaching high canopy cover). This comprised a full gradient of canopy coverage. On the 45 oak trees sampled, a total of 17 ant species were found. Species composition was dependent on the different forestry management practices. The total number of species and the number of species listed in the Red Data Books of both Germany and Bavaria were much higher in the forest pasture and the coppice with standards, as compared to the high forest. The transitional forest was at an intermediate level. The highest number of ant species was found in the forest pasture. This can be explained by the occurrence of species of open habitats and thermophilous species. In the coppice with standards, forest dwelling and arboricolous species dominated, whereas the high forest showed much lower frequencies of arboricolous species like Temnothorax corticalis, Dolichoderus quadripunctatus and Temnothorax affinis. A multivariate analysis revealed that canopy cover (measured as “shade”, in percentage intervals of canopy cover) was the best parameter for explaining species distribution and dataset variation, and to a lesser extent the amount of dead wood, canopy and trunk diameter. Thus ant fauna composition was mostly driven by structural differences associated to the different forestry management practices. Many ant species clearly preferred the more open and light forest stands of the coppice with standards as compared to the dense and shady high forest.

Diversity and Interactions of Wood-Inhabiting Fungi and Beetles after Deadwood Enrichment

Freshly cut beech deadwood was enriched in the canopy and on the ground in three cultural landscapes in Germany (Swabian Alb, Hainich-Dün, Schorfheide-Chorin) in order to analyse the diversity, distribution and interaction of wood-inhabiting fungi and beetles. After two years of wood decay 83 MOTUs (Molecular Operational Taxonomic Units) from 28 wood samples were identified. Flight Interception Traps (FITs) installed adjacent to the deadwood enrichments captured 29.465 beetles which were sorted to 566 species. Geographical ‘region’ was the main factor determining both beetle and fungal assemblages. The proportions of species occurring in all regions were low. Statistic models suggest that assemblages of both taxa differed between stratum and management praxis but their strength varied among regions. Fungal assemblages in Hainich-Dün, for which the data was most comprehensive, discriminated unmanaged from extensively managed and age-class forests (even-aged timber management) while canopy communities differed not from those near the ground. In contrast, the beetle assemblages at the same sites showed the opposite pattern. We pursued an approach in the search for fungus-beetle associations by computing cross correlations and visualize significant links in a network graph. These correlations can be used to formulate hypotheses on mutualistic relationships for example in respect to beetles acting as vectors of fungal spores.

The spider fauna (Araneae) of tree canopies in the Białowieża Forest

Spider communities of tree-canopies in primary forest sanctuaries and forest plantations of the Bialowieza Forest were sampled using insecticidal knockdown fogging. The communities showed the typical guild-, family- and species-composition of European forests with a dominance of web spiders and woodland-canopy spiders (e. g. Diaea dorsata, Anyphaena accentuata, Paidiscura pallens, Keijia tincta, Theridion varians). Based on 78 fogged trees, 14522 spiders (3936 adults) were sampled and sorted to 89 species from 15 families. They comprised 21% of the known species pool of the Bialowieza Forest (428 species). The importance of the canopy stratum as a habitat for spiders is emphasized. Six species are new records for the Bialowieza Forest. The rarely found Dipoena nigroreticulata Simon, 1879 (Theridiidae) was the only species with a preference for a single forest type. It was recorded from canopies of old oaks in a primary forest and might be an indicator species of old forests close to their natural condition.

Are temperate canopy spiders tree-species specific?

Arboreal spiders in deciduous and coniferous trees were investigated on their distribution and diversity. Insecticidal knock-down was used to comprehensively sample spiders from 175 trees from 2001 to 2003 in the Białowieża forest and three remote forests in Poland. We identified 140 species from 9273 adult spiders. Spider communities were distinguished between deciduous and coniferous trees. The richest fauna was collected from Quercus where beta diversity was also highest. A tree-species-specific pattern was clearly observed for Alnus, Carpinus, Picea and Pinus trees and also for those tree species that were fogged in only four or three replicates, namely Betula and Populus. This hitherto unrecognised association was mainly due to the community composition of common species identified in a Dufrene-Legendre indicator species analysis. It was not caused by spatial or temporal autocorrelation. Explaining tree-species specificity for generalist predators like spiders is difficult and has to involve physical and ecological tree parameters like linkage with the abundance of prey species. However, neither did we find a consistent correlation of prey group abundances with spiders nor could differences in spider guild composition explain the observed pattern. Our results hint towards the importance of deterministic mechanisms structuring communities of generalist canopy spiders although the casual relationship is not yet understood.