Roland Brandl

Pattern-oriented modelling in population ecology

Abstract Ecological modelling should take its orientation more from real patterns observed in nature, than has been the case up to now, to overcome the deficiencies of the present strategies. Firstly, the orientation towards patterns provides guidelines about the manner and extent of the aggregation of biological information in the model. Modelling thereby loses much of its arbitrariness; secondly, pattern-oriented models are not ‘scale-free’, i.e. they relate explicitly to spatial and temporal scales; and finally, they produce comparative predictions which are better suited for testing than the predictions achieved by models, which are only either complex or generalizing. To demonstrate the strategy of pattern-oriented modelling and its advantages, three examples from population ecology are presented: (1) In a model concerning density dependence and individual variability, the orientation towards a pattern in weight distributions requires explicit inclusion of the within-generation time scale in the model. (2) Orientation towards a pattern in the dispersal capabilities of small organisms leads to a metapopulation model, where environmental correlations about certain distances are taken into account. The survival of the metapopulation depends mainly on the ratio of the correlation distance to the range of dispersal of the organisms. (3) The wave-like pattern of the spread of rabies is reproduced by an extremely simple one-dimensional model, which is based on an extended cellular automaton approach. From this basic model, a description on finer spatial and temporal scales can be developed with the aim of constructing a model which allows for the investigation of spatial barriers against the spread of rabies. A comparison of the three example models shows that the main features of pattern-oriented models are generic. In producing comparative predictions which are related explicitly to scales, pattern-oriented modelling seems to be a strategy well suited to the ‘scaling up’ from population ecology to community and ecosystem ecology.

Loss of functional diversity of ant assemblages in secondary tropical forests

Secondary forests and plantations increasingly dominate the tropical wooded landscape in place of primary forests. The expected reduction of biodiversity and its impact on ecological functions provided by these secondary forests are of major concern to society and ecologists. The potential effect of biodiversity loss on ecosystem functioning depends largely on the associated loss in the functional diversity of animal and plant assemblages, i.e., the degree of functional redundancy among species. However, the relationship between species and functional diversity is still poorly documented for most ecosystems. Here, we analyze how changes in the species diversity of ground-foraging ant assemblages translate into changes of functional diversity along a successional gradient of secondary forests in the Atlantic Forest of Brazil. Our analysis uses continuous measures of functional diversity and is based on four functional traits related to resource use of ants: body size, relative eye size, relative leg length, and trophic position. We find a strong relationship between species and functional diversity, independent of the functional traits used, with no evidence for saturation in this relationship. Recovery of species richness and diversity of ant assemblages in tropical secondary forests was accompanied by a proportional increase of functional richness and diversity of assemblages. Moreover, our results indicate that the increase in functional diversity along the successional gradient of secondary forests is primarily driven by rare species, which are functionally unique. The observed loss of both species and functional diversity in secondary forests offers no reason to believe that the ecological functions provided by secondary forests are buffered against species loss through functional redundancy.

Current Near‐to‐Nature Forest Management Effects on Functional Trait Composition of Saproxylic Beetles in Beech Forests

With the aim of wood production with negligible negative effects on biodiversity and ecosystem processes, a silvicultural practice of selective logging with natural regeneration has been implemented in European beech forests (Fagus sylvatica) during the last decades. Despite this near-to-nature strategy, species richness of various taxa is lower in these forests than in unmanaged forests. To develop guidelines to minimize the fundamental weaknesses in the current practice, we linked functional traits of saproxylic beetle species to ecosystem characteristics. We used continental-scale data from 8 European countries and regional-scale data from a large forest in southern Germany and forest-stand variables that represented a gradient of intensity of forest use to evaluate the effect of current near-to-nature management strategies on the functional diversity of saproxylic beetles. Forest-stand variables did not have a statistically significant effect on overall functional diversity, but they did significantly affect community mean and diversity of single functional traits. As the amount of dead wood increased the composition of assemblages shifted toward dominance of larger species and species preferring dead wood of large diameter and in advanced stages of decay. The mean amount of dead wood across plots in which most species occurred was from 20 to 60 m(3) /ha. Species occurring in plots with mean dead wood >60 m(3) /ha were consistently those inhabiting dead wood of large diameter and in advanced stages of decay. On the basis of our results, to make current wood-production practices in beech forests throughout Europe more conservation oriented (i.e., promoting biodiversity and ecosystem functioning), we recommend increasing the amount of dead wood to >20 m(3) /ha; not removing dead wood of large diameter (50 cm) and allowing more dead wood in advanced stages of decomposition to develop; and designating strict forest reserves, with their exceptionally high amounts of dead wood, that would serve as refuges for and sources of saproxylic habitat specialists.

Association of extinction risk of saproxylic beetles with ecological degradation of forests in Europe.

To reduce future loss of biodiversity and to allocate conservation funds effectively, the major drivers behind large-scale extinction processes must be identified. A promising approach is to link the red-list status of species and specific traits that connect species of functionally important taxa or guilds to resources they rely on. Such traits can be used to detect the influence of anthropogenic ecosystem changes and conservation efforts on species, which allows for practical recommendations for conservation. We modeled the German Red List categories as an ordinal index of extinction risk of 1025 saproxylic beetles with a proportional-odds linear mixed-effects model for ordered categorical responses. In this model, we estimated fixed effects for intrinsic traits characterizing species biology, required resources, and distribution with phylogenetically correlated random intercepts. The model also allowed predictions of extinction risk for species with no red-list category. Our model revealed a higher extinction risk for lowland and large species as well as for species that rely on wood of large diameter, broad-leaved trees, or open canopy. These results mirror well the ecological degradation of European forests over the last centuries caused by modern forestry, that is the conversion of natural broad-leaved forests to dense conifer-dominated forests and the loss of old growth and dead wood. Therefore, conservation activities aimed at saproxylic beetles in all types of forests in Central and Western Europe should focus on lowlands, and habitat management of forest stands should aim at increasing the amount of dead wood of large diameter, dead wood of broad-leaved trees, and dead wood in sunny areas.

Demographic and random amplified polymorphic DNA analyses reveal high levels of genetic diversity in a clonal violet

We performed demographic and molecular investigations on woodland populations of the clonal herb Viola riviniana in central Germany. We investigated the pattern of seedling recruitment, the amount of genotypic (clonal) variation and the partitioning of genetic variation among and within populations. Our demographic study was carried out in six violet populations of different ages and habitat conditions. It revealed that repeated seedling recruitment takes place in all of these populations, and that clonal propagation is accompanied by high ramet mortality. Our molecular investigations were performed on a subset of three of these six violet populations. Random amplified polymorphic DNA analyses using six primers yielded 45 scorable bands that were used to identify multilocus genotypes, i.e. putative clones. Consistent with our demographic results and independent of population age, we found a large genotypic diversity with a mean proportion of distinguishable genotypes of 0.93 and a mean Simpson’s diversity index of 0.99. Using amova we found a strong genetic differentiation among these violet populations with a ΦST value of 0.41. We suggest that a high selfing rate, limited gene flow due to short seed dispersal distances and drift due to founder effects are responsible for this pattern. Although Viola riviniana is a clonal plant, traits associated with sexual reproduction rather than clonality per se are moulding the pattern of genetic variation in this species.

Cuticular hydrocarbons and aggression in the termite Macrotermes subhyalinus.

Cuticular hydrocarbons are among the prime candidates for nestmate recognition in social insects. We analyzed the variation of cuticular hydrocarbons in the termite species M. subhyalinus in West Africa (Comoë National Park) on a small spatial scale (<1 km). We found considerable variation in the composition of cuticular hydrocarbons among colonies, with four distinct chemical phenotypes. Different phenotypes occurred within each of the four habitats. The difference between these phenotypes is primarily due to unsaturated compounds. A clear correlation between the difference of the hydrocarbon composition and the aggression between colonies was found. This correlation also holds in a multivariate analysis of genetic similarity (measured by AFLPs), morphometric distances (measured by Mahalanobis-distances), as well as geographic distances between colonies. In a more detailed analysis of the correlation between the composition of cuticular hydrocarbons and aggression, we found that no single compound is sufficient to explain variation in aggression between pairings of colonies. Thus, termites seem to use a bouquet of compounds. Multiple regression analysis suggested that many of these compounds are unsaturated hydrocarbons and, thus, may play a key role in colony recognition.

Pattern formation triggered by rare events: lessons from the spread of rabies

Understanding of large–scale spatial pattern formation is a key to successful management in ecology and epidemiology. Neighbourhood interactions between local units are known to contribute to large–scale patterns, but how much do they contribute and what is the role of regional interactions caused by long–distance processes? How much long–distance dispersal do we need to explain the patterns that we observe in nature? There seems to be no way to answer these questions empirically. Therefore, we present a modelling approach that is a combination of a grid–based model describing local interactions and an individual–based model describing dispersal. Applying our approach to the spread of rabies, we show that in addition to local rabies dynamics, one long–distance infection per 14000km2 per year is sufficient to reproduce the wave–like spread of this disease. We conclude that even rare ecological events that couple local dynamics on a regional scale may have profound impacts on large–scale patterns and, in turn, dynamics. Furthermore, the following results emerge: (i) Both neighbourhood infection and long–distance infection are needed to generate the wave–like dispersal pattern of rabies; (ii) randomly walking rabid foxes are not sufficient to generate the wave pattern; and (iii) on a scale of less than 100 km times 100 km, temporal oscillations emerge that are independent from long–distance dispersal.

Spread of violets in polluted pine forests: morphological and molecular evidence for the ecological importance of interspecific hybridization

Hybridization between plant species occurs frequently but hybrids are often restricted to ecotones or disturbed habitats. In this study we show that introgressive hybrids between the tetraploid Viola riviniana and the diploid V. reichenbachiana invaded pine forests of the Dübener Heide (central Germany), an area affected by calcareous pollutants. The spread of these violet populations was correlated with the impact of pollution on habitat conditions. We compared morphology, cytology and random amplified polymorphic DNA (RAPD) bands among six Viola populations from the Dübener Heide and three populations of each pure species. RAPD analysis using 12 primers revealed 141 scorable bands. We considered bands as species specific if they occurred in at least 75% of the plants in one pure violet species but in none of the other. Seven bands were specific to V. riviniana and 11 bands were specific to V. reichenbachiana. Two plants of a V. reichenbachiana population were identified as hybrids. Of the Viola populations from the Dübener Heide, one was diploid and could be classified as V. reichenbachiana by morphology and RAPD markers. However, the majority of the Dübener Heide populations were tetraploid, and of a more variable morphology than V. riviniana and V. reichenbachiana, showing different combinations of intermediate characters, characters of the pure species and extreme characters. Despite their overall genetic similarity to V. riviniana, these plants could be identified as introgressive hybrids between V. riviniana and V. reichenbachiana by species‐specific RAPD bands. Therefore, we propose that recurrent hybridization and backcrossing resulted in novel genotypes adapted to the changed environment of polluted pine forests.

Aggregative response in bats: Prey abundance versus habitat

In habitats where prey is either rare or difficult to predict spatiotemporally, such as open habitats, predators must be adapted to react effectively to variations in prey abundance. Open-habitat foraging bats have a wing morphology adapted for covering long distances, possibly use information transfer to locate patches of high prey abundance, and would therefore be expected to show an aggregative response at these patches. Here, we examined the effects of prey abundance on foraging activities of open-habitat foragers in comparison to that of edge-habitat foragers and closed-habitat foragers. Bat activity was estimated by counting foraging calls recorded with bat call recorders (38,371 calls). Prey abundance was estimated concurrently at each site using light and pitfall traps. The habitat was characterized by terrestrial laser scanning. Prey abundance increased with vegetation density. As expected, recordings of open-habitat foragers clearly decreased with increasing vegetation density. The foraging activity of edge- and closed-habitat foragers was not significantly affected by the vegetation density, i.e., these guilds were able to forage from open habitats to habitats with dense vegetation. Only open-habitat foragers displayed a significant and proportional aggregative response to increasing prey abundance. Our results suggest that adaptations for effective and low-cost foraging constrains habitat use and excludes the guild of open-habitat foragers from foraging in habitats with high prey abundance, such as dense forest stands.

Impacts of Land Abandonment on Vegetation: Successional Pathways in European Habitats

Changes in traditional agricultural systems in Europe in recent decades have led to widespread abandonment and colonization of various habitats by shrubs and trees. We combined several vegetation databases to test whether patterns of changes in plant diversity after land abandonment in different habitats followed similar pathways. The impacts of land abandonment and subsequent woody colonization on vegetation composition and plant traits were studied in five semi-natural open habitats and two arable habitats in six regions of Europe. For each habitat, vegetation surveys were carried out in different stages of succession using either permanent or non-permanent plots. Consecutive stages of succession were defined on a physiognomic basis from initial open stages to late woody stages. Changes in vegetation composition, species richness, numbers of species on Red Lists, plant strategy types, Ellenberg indicator values of the vegetation, Grime CSR strategy types and seven ecological traits were assessed for each stage of the successional pathway. Abandonment of agro-pastoral land-use and subsequent woody colonization were associated with changes in floristic composition. Plant richness varied according to the different habitats and stages of succession, but semi-natural habitats differed from arable fields in several ecological traits and vegetation responses. Nevertheless, succession occurred along broadly predictable pathways. Vegetation in abandoned arable fields was characterized by a decreasing importance of R-strategists, annuals, seed plants with overwintering green leaves, insect-pollinated plants with hemi-rosette morphology and plants thriving in nutrient-rich conditions, but an increase in species considered as endangered according to the Red Lists. Conversely, changes in plant traits with succession within the initially-open semi-natural habitats showed an increase in plants thriving in nutrient-rich conditions, stress-tolerant plants and plants with sexual and vegetative reproduction, but a sharp decrease in protected species. In conclusion, our study showed a set of similarities in responses of the vegetation in plant traits after land abandonment, but we also highlighted differences between arable fields and semi-natural habitats, emphasizing the importance of land-use legacy.