Heike Kappes

Evolution of endemism on a young tropical mountain

Tropical mountains are hot spots of biodiversity and endemism, but the evolutionary origins of their unique biotas are poorly understood. In varying degrees, local and regional extinction, long-distance colonization, and local recruitment may all contribute to the exceptional character of these communities. Also, it is debated whether mountain endemics mostly originate from local lowland taxa, or from lineages that reach the mountain by long-range dispersal from cool localities elsewhere. Here we investigate the evolutionary routes to endemism by sampling an entire tropical mountain biota on the 4,095-metre-high Mount Kinabalu in Sabah, East Malaysia. We discover that most of its unique biodiversity is younger than the mountain itself (6 million years), and comprises a mix of immigrant pre-adapted lineages and descendants from local lowland ancestors, although substantial shifts from lower to higher vegetation zones in this latter group were rare. These insights could improve forecasts of the likelihood of extinction and ‘evolutionary rescue’ in montane biodiversity hot spots under climate change scenarios.

Response of ground-dwelling beetle (Coleoptera) assemblages to giant knotweed (Reynoutria spp.) invasion

Giant knotweeds (Reynoutria spp.) are highly productive and aggressive invaders in riparian wetlands of Europe and North America. We sampled ground-dwelling beetles by pitfall traps from six sites comparing monotypic Reynoutria stands with the invaded native Urtica-dominated stands. Three sites were located in a semi-natural softwood forest and three sites were on a ruderal embankment. The analyses are based on a total of 13,244 individuals from 218 species. Location and site significantly influenced beetle assemblages. Moreover, there were pronounced differences between vegetation stands. The monotypic Reynoutria stands exhibited lower beetle abundance, species richness and rarefaction diversity irrespective of location. However, the negative effect on species richness, abundance and assemblage similarities were stronger on the transformed ruderal embankment than in the semi-natural softwood forest. Reynoutria invasion seems to influence microclimatic conditions. We found a higher abundance of silvicolous and a lower abundance of xerophilous ground beetles in the Reyountria stands than in the Urtica-dominated stands. Feeding guilds reacted differently to Reynoutria invasion that reduced the abundance of predators and herbivores but enhanced that of detritivores. Detritivores assumingly profit from the perennial presence of the large quantities of Reynoutria litter. We conclude that highly productive invaders pauperise the arthropod fauna and alter link strengths in trophic cascades shifting primary producer-based food webs to detritus-based food webs.

Emergence of Coleoptera from deadwood in a managed broadleaved forest in central Europe

We studied Coleoptera emerging from deadwood in a broadleaved forest that has been managed with nature-oriented forestry since 1990 and in which an accumulation of 10 m3 deadwood/ha has been achieved. We sampled a total of 11 m3 of deadwood from oak and beech trees and yielded 32,477 individuals pertaining to 342 species. The number of species emerging from oak deadwood was significantly higher than that from beech. Significantly more species emerged from deadwood with medium size (15–21 cm) diameter than from smaller (5–7 cm) or larger (40–60 cm) diameters. The stage of decay did not influence the number of species during the first 6 years of decomposition. Species richness strongly increased with the amount of deadwood sampled. Species–area regressions yielded z-values of 0.81 for oak and 0.87 for beech deadwood. The beetle assemblage was characterized by a low number of xylophagous species and a high number of mobile zoophagous species. However, two-thirds of the individuals sampled pertained to nine species of Scolytidae. This is why Shannon–Wiener diversity (H) and evenness-indices (E) greatly decreased with increasing number of scolytids. The species and individuals yielded from the sun-exposed deadwood outnumbered those from shaded areas. Namely the scolytid Taphrorychus bicolor reached remarkably high densities in the sun-exposed beech deadwood (>800 individuals/m2). In contrast, oak deadwood never harboured such high numbers of scolytids. Felling of moribund trees had decreased the number of scolytids but it had also considerably reduced the number of rare and endangered species.

Long-term dynamics of Asplanchna priodonta in Lake Windsborn with comments on the diet

We evaluate trophic interactions between the rotifer Asplanchna priodonta and the plankton community of Lake Windsborn (Eifel District, Germany). The long-term population dynamics of this species and of the two most abundant zooplankton species, Keratella cochlearis and Bosmina longirostris, were analyzed in relation to hydrochemical and climatic variates. The maximum abundance of A. priodonta was limited to the summer months with temperature being the only significant predictor of abundance. Intrinsic rate of growth and abundance of the three species were independent from each other suggesting the absence of trophic interactions. In Lake Windsborn, A. priodonta was a grazer rather than a predator. Its diet included high proportions of colonial algae and cyanobacteria which cannot be ingested by the other abundant zooplankton species, whereas zooplanktonic food were rarely found. Thus, our study supports the hypothesis that A. priodonta is an opportunistic feeder whose trophic position entirely depends on the structure of the plankton community.

Litter-dwelling beetles in primeval forests of Central Europe: Does deadwood matter?

We investigated the distribution pattern of litter dwelling beetles in four primeval forests of the Western Carpathians. The forests are situated in two mountain ranges and are either southerly exposed oak forests or northerly exposed beech forests. Beetles were extracted from leaf litter of plots close to coarse woody debris (c-CWD) and distant from coarse woody debris (d-CWD). We collected 2946 individuals of 172 species. Plots close to CWD usually were hot spots of species richness and beetle density, which could be increased two-fold and five-fold, respectively, when compared to d-CWD sites. The influence of CWD within each forest type was stronger than that of any other environmental factor. The c-CWD plots were characterised by both a higher presence of common species and a higher number of rare species. Especially, zoophagous and mycetophagous beetles were enhanced. The assemblage wide carrying capacity at c-CWD sites varied in respect to the gradients of temperature and moisture within a forest. A longer gradient provided a higher carrying capacity. We assume that the influence of downed deadwood on litter dwelling beetles will also be effective in commercial forests and stop the downward spiral with continuing loss of species.

On the growth of bivalve gills initiated from a lobule-producing budding zone.

The growth of bivalve gills proceeds at the posterior end of the gill from a meristem-like budding zone, that is, an undifferentiated terminal organ, which continuously proliferates new gill elements in growing bivalves. In representatives of protobranch, filibranch, and eulamellibranch gills (13 species from Protobranchia, Pteriomorphia, Palaeoheterodonta, and Heterodonta), the first growth steps demonstrate a uniform basic pattern. The budding zone produces either transverse folds that split after a transition zone into parallel pairs of lobules (which themselves later differentiate into the inner and outer demibranchs), or it produces the lobules directly, without first forming a transition zone. The lobules elongate, differentiate into lobes, and transform into leaflet-like structures (protobranchs) or into filaments (filibranchs and eulamellibranchs). The filaments represent the differentiated outer margins of each lobe, of which the central tissue (interlamellar septum) becomes incised or fenestrated, or transformed by tissue junctions. A distally located main growth zone for each lobe is suggested. With regard to the delayed onset of the differentiation of the outer demibranch in juvenile unionids, an additional temporary growth zone for filaments is suggested to exist at the anterior end of the outer demibranch.

Urban summer heat load: meteorological data as a proxy for metropolitan biodiversity

Regional climate models forecast that the incidence of extreme heat waves will increase in Central Europe, and the associated intensification of urban heat islands causes concerns about human health and biodiversity. We investigated species diversity of terrestrial gastropods across an urban gradient in a metropolitan area. We considered 34 sites of different degrees of urbanization and of different thermal stress classes (TSC) that were derived from a classification of urban heat load based on the physiological equivalent temperature (PET). A total of 31 species were recorded with 0–21 species per site (mean: 4.4 ± 5.2 species per site). Alpha diversity was best explained by TSC, bush cover and perennial vegetation cover, and it decreased with increasing thermal stress. Overall, thermal stress predicted snail and slug species richness better than urban cover (ANOVA(analysis of variance); TSC: F = 10.0, p < 0.001; urbanization: F = 3.9, p = 0.018), and the proportional loss in species richness was higher for native species than for introduced species. The results indicate that climatic stress contributes to the impoverishment of biodiversity in urban areas. We propose that TSC and/or PET are useful indicators for environmental stress levels in biodiversity studies in natural and anthropogenically transformed landscapes.

Tolerance of Ceriodaphnia quadrangula and Diaphanosoma brachyurum (Crustacea: Cladocera) to experimental soft water acidification

AbstractWe analysed in how far the decrease of pH, that is part of the ongoing restoration of the softwater Lake Windsborn (conductivity below 30 μS cm−1), may in future influence the occurrence of the two cladoceran species Ceriodaphnia quadrangula and Diaphanosoma brachyurum. In the field, the abundance of Ceriodaphnia was positively correlated with lake water pH, whereas there was no correlation between abundance and pH for D. brachyurum. Experiments on the tolerance against acidification included dynamic (24 h) and static tests (24, 48 h, 30 d), and were conducted with acidified lake water. C. quadrangula tolerated a slight acidification to pH 5.2, but not pH 4.8, whereas the NOEC value seems to be between pH 4.2 and 4.5 for D. brachyurum. Differences between the experimental NOEC values and field data may be explained by diurnal pH fluctuations and the low ion content of Lake Windsborn which puts an additional physiological challenge to its inhabitants.

Differences in dispersal- and colonization-related traits between taxa from the freshwater and the terrestrial realm

The aquatic and terrestrial realms differ in many physical properties that not only require specific physiological adaptations but also cause differences in dispersal options. We thus expect that life-history traits related to dispersal and colonization are under selection pressure because freshwater habitats are more isolated and thus more difficult to reach. We compared traits from European databases of three taxonomic groups along the passive–active dispersal gradient: plants (Plantes), snails (Mollusca: Gastropoda: Prosobranchia et Pulmonata) and hoverflies (Diptera: Syrphidae), all of which have both terrestrial and freshwater species (plants and snails) or early life stages (hoverflies). Aquatic taxa seem to be more successful long-distance dispersers than are terrestrial taxa. Our analysis also revealed lower numbers of seeds or eggs produced in the aquatic habitats. However, aquatic taxa often allocate resources to offspring guarding (vegetative propagules in plants, egg capsules in snails) and breeding-site selection (syrphids). Colonization of the aquatic realm is reinforced by increases in life span (plants), clonal spread (plants), shorter generation times (snails), selfing ability (marginal effect in pulmonate snails) or paedogenesis (two incidences in hoverflies, needs further studies). Probably, the variety of strategies reflects the different evolutionary backgrounds that elicit different combinations of trade-offs, but all traits also might increase invasibility of species.

Complex Responses of Songbirds to Soil Acidification of Managed Beech Forests in Central Europe

A bstractMany forest ecosystems have been extensively degraded by acidifying pollutants over the last decades. There is also widespread concern about population decline of a number of songbirds due to acidification-driven calcium limitation. We hypothesize that the carrying capacity of forests decreases with reduced soil calcium content. We analyzed songbird assemblage/habitat relationships by territory mapping at 30 sites in a total of five managed beech forests. The sites harbored 38 songbird species. A canonical correspondence analysis showed that soil acidification, represented by soil calcium content, had a significant influence on the species assemblages of the 14 most frequent songbirds. The vertical complexity of the canopy additionally influenced the assemblages. Multiple regressions revealed that the density of the territories of song thrush (Turdus philomelos) and nuthatch (Sitta europaea) were positively correlated to calcium content. Blackbirds (Turdus merula) seemed to compensate for the negative effect of acidification by selecting breeding sites close to forest edges to forage in less acidified agricultural habitats. In contrast, the territorial density of robin (Erithacus rubecula), chaffinch (Phylloscopus collybita) and coal tit (Parus ater) increased with increasing soil acidification. Contrary to our hypothesis, the overall carrying capacity of songbird was not reduced in forests with acidified soils. However, the nesting success of song thrush and blackbird, as determined by the number of fledglings, was higher in the forest with the highest calcium content when compared to the forest with the lowest calcium content. We conclude that some species are severely affected by acidification, whereas others seem to have evolved successful strategies to meet their calcium demand even in acidified habitats.