Marcell K. Peters

Molecular species identification of Central European ground beetles (Coleoptera: Carabidae) using nuclear rDNA expansion segments and DNA barcodes

BackgroundThe identification of vast numbers of unknown organisms using DNA sequences becomes more and more important in ecological and biodiversity studies. In this context, a fragment of the mitochondrial cytochrome c oxidase I (COI) gene has been proposed as standard DNA barcoding marker for the identification of organisms. Limitations of the COI barcoding approach can arise from its single-locus identification system, the effect of introgression events, incomplete lineage sorting, numts, heteroplasmy and maternal inheritance of intracellular endosymbionts. Consequently, the analysis of a supplementary nuclear marker system could be advantageous.ResultsWe tested the effectiveness of the COI barcoding region and of three nuclear ribosomal expansion segments in discriminating ground beetles of Central Europe, a diverse and well-studied invertebrate taxon. As nuclear markers we determined the 18S rDNA: V4, 18S rDNA: V7 and 28S rDNA: D3 expansion segments for 344 specimens of 75 species. Seventy-three species (97%) of the analysed species could be accurately identified using COI, while the combined approach of all three nuclear markers provided resolution among 71 (95%) of the studied Carabidae.ConclusionOur results confirm that the analysed nuclear ribosomal expansion segments in combination constitute a valuable and efficient supplement for classical DNA barcoding to avoid potential pitfalls when only mitochondrial data are being used. We also demonstrate the high potential of COI barcodes for the identification of even closely related carabid species.

Forest Fragmentation and Selective Logging Have Inconsistent Effects on Multiple Animal-Mediated Ecosystem Processes in a Tropical Forest

Forest fragmentation and selective logging are two main drivers of global environmental change and modify biodiversity and environmental conditions in many tropical forests. The consequences of these changes for the functioning of tropical forest ecosystems have rarely been explored in a comprehensive approach. In a Kenyan rainforest, we studied six animal-mediated ecosystem processes and recorded species richness and community composition of all animal taxa involved in these processes. We used linear models and a formal meta-analysis to test whether forest fragmentation and selective logging affected ecosystem processes and biodiversity and used structural equation models to disentangle direct from biodiversity-related indirect effects of human disturbance on multiple ecosystem processes. Fragmentation increased decomposition and reduced antbird predation, while selective logging consistently increased pollination, seed dispersal and army-ant raiding. Fragmentation modified species richness or community composition of five taxa, whereas selective logging did not affect any component of biodiversity. Changes in the abundance of functionally important species were related to lower predation by antbirds and higher decomposition rates in small forest fragments. The positive effects of selective logging on bee pollination, bird seed dispersal and army-ant raiding were direct, i.e. not related to changes in biodiversity, and were probably due to behavioural changes of these highly mobile animal taxa. We conclude that animal-mediated ecosystem processes respond in distinct ways to different types of human disturbance in Kakamega Forest. Our findings suggest that forest fragmentation affects ecosystem processes indirectly by changes in biodiversity, whereas selective logging influences processes directly by modifying local environmental conditions and resource distributions. The positive to neutral effects of selective logging on ecosystem processes show that the functionality of tropical forests can be maintained in moderately disturbed forest fragments. Conservation concepts for tropical forests should thus include not only remaining pristine forests but also functionally viable forest remnants.

Complementary ecosystem services provided by pest predators and pollinators increase quantity and quality of coffee yields

Wild animals substantially support crop production by providing ecosystem services, such as pollination and natural pest control. However, the strengths of synergies between ecosystem services and their dependencies on land-use management are largely unknown. Here, we took an experimental approach to test the impact of land-use intensification on both individual and combined pollination and pest control services in coffee production systems at Mount Kilimanjaro. We established a full-factorial pollinator and vertebrate exclosure experiment along a land-use gradient from traditional homegardens (agroforestry systems), shaded coffee plantations to sun coffee plantations (total sample size = 180 coffee bushes). The exclusion of vertebrates led to a reduction in fruit set of ca 9%. Pollinators did not affect fruit set, but significantly increased fruit weight of coffee by an average of 7.4%. We found no significant decline of these ecosystem services along the land-use gradient. Pest control and pollination service were thus complementary, contributing to coffee production by affecting the quantity and quality of a major tropical cash crop across different coffee production systems at Mount Kilimanjaro.

Seed dispersal by ants: are seed preferences influenced by foraging strategies or historical constraints?

Summary The objective of this study was to quantify preferences of ants for seeds of different plant species and to test if these preferences were caused by foraging strategies or by historical constraints. We compared seed removal rates of ten different ant-dispersed plant species found in temperate forests, along forest edges and in grassland. We found significant differences in seed removal rates among the ten species. To test if these differences were caused by foraging strategies we examined the relationship between seed and elaiosome size and seed removal rate. We found that seeds with larger elaiosomes had significantly higher removal rates. To test the historical constraint hypothesis we investigated whether plants from forest, forest edge or grassland habitats were preferred by ant species found in the same habitat type. If historical constraints exist we would expect that ants remove preferentially the seeds of plant species which they are used to handle in the course of their phylogenetic and ontogenetic history. In contrast to this prediction, we found that plants of a certain habitat type were in general not preferred by ants of the same habitat type. These results demonstrate that seed preferences of ants are mainly determined by foraging strategies and not by historical constraints.

EFFECTS OF HABITAT FRAGMENTATION AND DEGRADATION ON FLOCKS OF AFRICAN ANT‐FOLLOWING BIRDS

Tropical rain forests are rapidly cleared, fragmented, and degraded in sub-Saharan Africa; however, little is known about the response of species and even of key ecological groups to these processes. One of the most intriguing (but often neglected) ecological phenomena in African rain forests is the interaction between swarm-raiding army ants and ant-following birds. Similar to their well-known Neotropical representatives, ant-following birds in Africa track the massive swarm raids of army ants and feed on arthropods flushed by the ants. In this study we analyzed the effect of habitat fragmentation and degradation of a mid-altitude Congo-Guinean rain forest in western Kenya on the structure of ant-following bird flocks. Significant numbers of swarm raids were located in all forest fragments and in both undegraded and degraded forest. Fifty-six different species of birds followed army ant raids, forming bird flocks of one to 15 species. We quantitatively differentiated the bird community into five species of specialized ant-followers and 51 species of opportunistic ant-followers. Species richness and size of bird flocks decreased with decreasing size of forest fragments and was higher in undegraded than in degraded habitat. This was caused by the decrease of the species richness and number of specialized ant-followers at swarms, while the group of opportunistic ant-followers was affected little by habitat fragmentation and degradation. The composition of bird flocks was more variable in small fragments and degraded forest, compared to undegraded habitat in large fragments. The effect of habitat fragmentation on flock structure was best explained by the strong decline of the abundance of specialized ant-followers in small forest fragments. To conserve the association of army ants and ant-following birds in its natural state, vast areas of unfragmented and undegraded tropical rain forest are necessary.

Predictors of elevational biodiversity gradients change from single taxa to the multi-taxa community level.

The factors determining gradients of biodiversity are a fundamental yet unresolved topic in ecology. While diversity gradients have been analysed for numerous single taxa, progress towards general explanatory models has been hampered by limitations in the phylogenetic coverage of past studies. By parallel sampling of 25 major plant and animal taxa along a 3.7 km elevational gradient on Mt. Kilimanjaro, we quantify cross-taxon consensus in diversity gradients and evaluate predictors of diversity from single taxa to a multi-taxa community level. While single taxa show complex distribution patterns and respond to different environmental factors, scaling up diversity to the community level leads to an unambiguous support for temperature as the main predictor of species richness in both plants and animals. Our findings illuminate the influence of taxonomic coverage for models of diversity gradients and point to the importance of temperature for diversification and species coexistence in plant and animal communities.

A Preliminary Checklist of the Ants (Hymenoptera: Formicidae) of Kakamega Forest (Kenya)

ABSTRACT A preliminary species checklist of the ants (Hymenoptera: Formicidae) of Kakamega Forest, Western Kenya, is presented. The species list is based on specimens sampled from 1999 until 2009, which are deposited in the ant collection of the Zoological Research Museum Koenig, Bonn, Germany, and the Natural History Museum of Los Angeles County, Los Angeles, United States. The known ant fauna comprises 11 subfamilies with 52 genera and 288 species. This species richness is the second highest reported from the Afrotropical zoogeographical region. The observed ant fauna can be divided into a group of forest habitat species with approximately 180 species and another group, which consists of about 100 species, of open habitat specialists. The greatest part of the ant fauna (204 species or 71%) could be identified to species level but the rest remains unidentifiable or undescribed (84 species or 29%). The preliminary data from the observed ant fauna seems to support the hypothesis that Kakamega Forest is the eastern-most remnant of the former Guineo-Congolian rain forest belt, while the zoogeographical influence from East African Afromontane forests seems to be less significant.

Integrating intraspecific variation in community ecology unifies theories on body size shifts along climatic gradients

1.Physiological and energetic mechanisms have been proposed to constrain body sizes of organisms along climatic gradients; however, these provide contrasting predictions. While Bergmann′s rule predicts increases of body sizes in cooler climates resulting from physiological constraints, energy-based community assembly rules suggest declines in the mean body size of species caused by increased extinction probabilities for large-bodied species in low-energy habitats. 2.We tested these contrasting hypotheses by quantifying trait distributions in bee communities along a 3.6 km elevational gradient at Mt. Kilimanjaro. Traditionally, intra- and interspecific trait shifts along environmental gradients have been investigated in isolation. However, a surge of theoretical approaches and studies on plants demonstrated that the explicit integration of trait variation among and within species can be essential for identifying the mechanisms that shape traits and related ecosystem functions along environmental gradients. We therefore studied variation of body size and related morphological traits at both the intra- and interspecific level. 3.We found support for both physiological constraints and energy-based community assembly rules as drivers of trait distribution in bee communities along elevational gradients, which, however, affected different levels of biotic organization, i.e. the population and community level. While the number of bee species with large body sizes declined with elevation, individuals within species became on average larger, resulting in contrasting trends in morphometric parameters at the community versus population level. 4.Furthermore, body size within bee communities became less variable at higher elevations, largely as a result of a non-random, directive loss of species, but paralleled by a decline in intraspecific variance, suggesting intensified filtering effects with increasing elevation. Similar patterns were found for other functional traits related to the foraging ecology of bees (tongue length, relative forewing length). 5.We conclude that along climatic gradients both physiological and energetic constraints shape trait distributions of pollinators, but at different levels of biological organization. This article is protected by copyright. All rights reserved.

Exploring the Leaf Beetle Fauna (Coleoptera: Chrysomelidae) of an Ecuadorian Mountain Forest Using DNA Barcoding

Background Tropical mountain forests are hotspots of biodiversity hosting a huge but little known diversity of insects that is endangered by habitat destruction and climate change. Therefore, rapid assessment approaches of insect diversity are urgently needed to complement slower traditional taxonomic approaches. We empirically compare different DNA-based species delimitation approaches for a rapid biodiversity assessment of hyperdiverse leaf beetle assemblages along an elevational gradient in southern Ecuador and explore their effect on species richness estimates. Methodology/Principal Findings Based on a COI barcode data set of 674 leaf beetle specimens (Coleoptera: Chrysomelidae) of 266 morphospecies from three sample sites in the Podocarpus National Park, we employed statistical parsimony analysis, distance-based clustering, GMYC- and PTP-modelling to delimit species-like units and compared them to morphology-based (parataxonomic) species identifications. The four different approaches for DNA-based species delimitation revealed highly similar numbers of molecular operational taxonomic units (MOTUs) (n = 284–289). Estimated total species richness was considerably higher than the sampled amount, 414 for morphospecies (Chao2) and 469–481 for the different MOTU types. Assemblages at different elevational levels (1000 vs. 2000 m) had similar species numbers but a very distinct species composition for all delimitation methods. Most species were found only at one elevation while this turnover pattern was even more pronounced for DNA-based delimitation. Conclusions/Significance Given the high congruence of DNA-based delimitation results, probably due to the sampling structure, our study suggests that when applied to species communities on a regionally limited level with high amount of rare species (i.e. ~50% singletons), the choice of species delimitation method can be of minor relevance for assessing species numbers and turnover in tropical insect communities. Therefore, DNA-based species delimitation is confirmed as a valuable tool for evaluating biodiversity of hyperdiverse insect communities, especially when exact taxonomic identifications are missing.

Testing a short nuclear marker for inferring staphylinid beetle diversity in an African tropical rain forest.

Background The use of DNA based methods for assessing biodiversity has become increasingly common during the last years. Especially in speciose biomes as tropical rain forests and/or in hyperdiverse or understudied taxa they may efficiently complement morphological approaches. The most successful molecular approach in this field is DNA barcoding based on cytochrome c oxidase I (COI) marker, but other markers are used as well. Whereas most studies aim at identifying or describing species, there are only few attempts to use DNA markers for inventorying all animal species found in environmental samples to describe variations of biodiversity patterns. Methodology/Principal Findings In this study, an analysis of the nuclear D3 region of the 28S rRNA gene to delimit species-like units is compared to results based on distinction of morphospecies. Data derived from both approaches are used to assess diversity and composition of staphylinid beetle communities of a Guineo-Congolian rain forest in Kenya. Beetles were collected with a standardized sampling design across six transects in primary and secondary forests using pitfall traps. Sequences could be obtained of 99% of all individuals. In total, 76 molecular operational taxonomic units (MOTUs) were found in contrast to 70 discernible morphospecies. Despite this difference both approaches revealed highly similar biodiversity patterns, with species richness being equal in primary and secondary forests, but with divergent species communities in different habitats. The D3-MOTU approach proved to be an efficient tool for biodiversity analyses. Conclusions/Significance Our data illustrate that the use of MOTUs as a proxy for species can provide an alternative to morphospecies identification for the analysis of changes in community structure of hyperdiverse insect taxa. The efficient amplification of the D3-marker and the ability of the D3-MOTUs to reveal similar biodiversity patterns as analyses of morphospecies recommend its use in future molecular studies on biodiversity.