Pavel Drozd

Low host specificity of herbivorous insects in a tropical forest

Two decades of research have not established whether tropical insect herbivores are dominated by specialists or generalists. This impedes our understanding of species coexistence in diverse rainforest communities. Host specificity and species richness of tropical insects are also key parameters in mapping global patterns of biodiversity. Here we analyse data for over 900 herbivorous species feeding on 51 plant species in New Guinea and show that most herbivorous species feed on several closely related plant species. Because species-rich genera are dominant in tropical floras, monophagous herbivores are probably rare in tropical forests. Furthermore, even between phylogenetically distant hosts, herbivore communities typically shared a third of their species. These results do not support the classical view that the coexistence of herbivorous species in the tropics is a consequence of finely divided plant resources; non-equilibrium models of tropical diversity should instead be considered. Low host specificity of tropical herbivores reduces global estimates of arthropod diversity from 31 million (ref. 1) to 4–6 million species. This finding agrees with estimates based on taxonomic collections, reconciling an order of magnitude discrepancy between extrapolations of global diversity based on ecological samples of tropical communities with those based on sampling regional faunas.

Why Are There So Many Species of Herbivorous Insects in Tropical Rainforests

Despite recent progress in understanding mechanisms of tree species coexistence in tropical forests, a simple explanation for the even more extensive diversity of insects feeding on these plants has been missing. We compared folivorous insects from temperate and tropical trees to test the hypothesis that herbivore species coexistence in more diverse communities could reflect narrow host specificity relative to less diverse communities. Temperate and tropical tree species of comparable phylogenetic distribution supported similar numbers of folivorous insect species, 29.0 ± 2.2 and 23.5 ± 1.8 per 100 square meters of foliage, respectively. Host specificity did not differ significantly between community samples, indicating that food resources are not more finely partitioned among folivorous insects in tropical than in temperate forests. These findings suggest that the latitudinal gradient in insect species richness could be a direct function of plant diversity, which increased sevenfold from our temperate to tropical study sites.

The global distribution of diet breadth in insect herbivores

Significance Dietary specialization determines an organism’s resource base as well as impacts on host or prey species. There are important basic and applied reasons to ask why some animals have narrow diets and others are more generalized, and if different regions of the Earth support more specialized interactions. We investigated site-specific host records for more than 7,500 species of insect herbivores. Although host specialists predominate, the proportion of specialists is affected by the diversity of hosts and shifts globally, supporting predictions of more exclusive tropical interactions. These results not only affect our understanding of the ecology of food webs, but also have implications for how they respond to environmental change, as well as for ecosystem management and restoration. Understanding variation in resource specialization is important for progress on issues that include coevolution, community assembly, ecosystem processes, and the latitudinal gradient of species richness. Herbivorous insects are useful models for studying resource specialization, and the interaction between plants and herbivorous insects is one of the most common and consequential ecological associations on the planet. However, uncertainty persists regarding fundamental features of herbivore diet breadth, including its relationship to latitude and plant species richness. Here, we use a global dataset to investigate host range for over 7,500 insect herbivore species covering a wide taxonomic breadth and interacting with more than 2,000 species of plants in 165 families. We ask whether relatively specialized and generalized herbivores represent a dichotomy rather than a continuum from few to many host families and species attacked and whether diet breadth changes with increasing plant species richness toward the tropics. Across geographic regions and taxonomic subsets of the data, we find that the distribution of diet breadth is fit well by a discrete, truncated Pareto power law characterized by the predominance of specialized herbivores and a long, thin tail of more generalized species. Both the taxonomic and phylogenetic distributions of diet breadth shift globally with latitude, consistent with a higher frequency of specialized insects in tropical regions. We also find that more diverse lineages of plants support assemblages of relatively more specialized herbivores and that the global distribution of plant diversity contributes to but does not fully explain the latitudinal gradient in insect herbivore specialization.

Predictably simple: assemblages of caterpillars (Lepidoptera) feeding on rainforest trees in Papua New Guinea

Predictability in the composition of tropical assemblages of insect herbivores was studied using a sample of 35 952 caterpillars (Lepidoptera) from 534 species, feeding on 69 woody species from 45 genera and 23 families in a lowland rainforest in Papua New Guinea. Caterpillar assemblages were strongly dominated by a single species (median 48% of individuals and 49% of biomass). They were spatially and temporally constant (median normalized expected species shared (NESS) similarity between assemblages from the same host was greater than or equal to 0.85 for three sites 8–17 km apart as well as for three four–month periods of the year). Further, the median presence of species was 11 months per year. Assemblages on hosts from different families and genera were virtually disjunct (NESS similarity less than 0.05) as the caterpillars were mostly specialized to a single plant family (77% of species) and, within families, to a single genus (66% of species), while capable of feeding on multiple congeneric hosts (89% of species). The dominance of caterpillar assemblages by a small number of specialized species, which also exhibited low spatial and temporal variability, permitted robust and reliable estimates of assemblage composition and between–assemblage similarity from small samples, typically less than 300 individuals per host plant. By contrast, even considerably larger samples were insufficient for estimates of species richness. A sample of 300 individuals was typically obtained from 1050 m2 of foliage sampled during 596 tree inspections (i.e. a particular tree sampled at a particular time) in the course of 19 sampling days (median values from 69 assemblages). These results demonstrate that, contrary to some previous suggestions, insect herbivore assemblages in tropical rainforests have a predictable structure and, as such, are amenable to study.

Higher predation risk for insect prey at low latitudes and elevations

Risky in the tropics It is well known that diversity increases toward the tropics. Whether this increase translates into differences in interaction rates among species, however, remains unclear. To simplify the problem, Roslin et al. tested for predation rates by using a single approach involving model caterpillars across six continents. Predator attack rates were higher toward the equator, but only for arthropod predators. Science, this issue p. 742 Like diversity, predation rates among insects increase toward the equator and at lower altitudes. Biotic interactions underlie ecosystem structure and function, but predicting interaction outcomes is difficult. We tested the hypothesis that biotic interaction strength increases toward the equator, using a global experiment with model caterpillars to measure predation risk. Across an 11,660-kilometer latitudinal gradient spanning six continents, we found increasing predation toward the equator, with a parallel pattern of increasing predation toward lower elevations. Patterns across both latitude and elevation were driven by arthropod predators, with no systematic trend in attack rates by birds or mammals. These matching gradients at global and regional scales suggest consistent drivers of biotic interaction strength, a finding that needs to be integrated into general theories of herbivory, community organization, and life-history evolution.

Dynamic formation of asexual diploid and polyploid lineages: multilocus analysis of Cobitis reveals the mechanisms maintaining the diversity of clones.

Given the hybrid genomic constitutions and increased ploidy of many asexual animals, the identification of processes governing the origin and maintenance of clonal diversity provides useful information about the evolutionary consequences of interspecific hybridization, asexuality and polyploidy. In order to understand the processes driving observed diversity of biotypes and clones in the Cobitis taenia hybrid complex, we performed fine-scale genetic analysis of Central European hybrid zone between two sexual species using microsatellite genotyping and mtDNA sequencing. We found that the hybrid zone is populated by an assemblage of clonally (gynogenetically) reproducing di-, tri- and tetraploid hybrid lineages and that successful clones, which are able of spatial expansion, recruit from two ploidy levels, i.e. diploid and triploid. We further compared the distribution of observed estimates of clonal ages to theoretical distributions simulated under various assumptions and showed that new clones are most likely continuously recruited from ancestral populations. This suggests that the clonal diversity is maintained by dynamic equilibrium between origination and extinction of clonal lineages. On the other hand, an interclonal selection is implied by nonrandom spatial distribution of individual clones with respect to the coexisting sexual species. Importantly, there was no evidence for sexually reproducing hybrids or clonally reproducing non-hybrid forms. Together with previous successful laboratory synthesis of clonal Cobitis hybrids, our data thus provide the most compelling evidence that 1) the origin of asexuality is causally linked to interspecific hybridization; 2) successful establishment of clones is not restricted to one specific ploidy level and 3) the initiation of clonality and polyploidy may be dynamic and continuous in asexual complexes.

CLONAL TURNOVER VERSUS CLONAL DECAY: A NULL MODEL FOR OBSERVED PATTERNS OF ASEXUAL LONGEVITY, DIVERSITY AND DISTRIBUTION

Abstract Phylogenetic and phylogeographic studies suggest that a majority of asexual organisms are evolutionarily recent offshoots of extant sexual taxa and that old clonal lineages tend to be isolated from their sexual and younger asexual counterparts. These observations have often been interpreted as support for the long-term disadvantages of asexuality resulting from the mechanisms of clonal decay. Although clonal decay is likely to be an important mechanism that limits the temporal and spatial distribution of asexual lineages, we argue here that contemporary phylogenetic analyses, which are mostly restricted to simple comparisons of “recent” and “ancient” clones, need to be tested against an appropriate null model of neutrality. We use computer simulations to show that many empirical observations of the distribution of asexuality do not in fact reject a null model of the neutral turnover of clones spawned by sexual relatives. In particular, neutral clonal turnover results in qualitatively similar pattern of clonal spatial distribution and age structure, as does a process that includes clonal decay. Although there are important quantitative differences between predictions made by the two models, we show that published empirical data are still inadequate to distinguish between them. Further work on sexual-asexual complexes is therefore required before clonal turnover can be rejected as a parsimonious explanation of the spatial distribution and age structure of asexual lineages.

Aquatic insects indicate terrestrial habitat degradation: changes in taxonomical structure and functional diversity of dragonflies in tropical rainforest of East Kalimantan

As a group of freshwater invertebrates, dragonflies (Odonata) are commonly used as ecological indicators of freshwater ecosystems. Despite earlier studies suggesting that adult odonates may be good indicators for complex changes in a landscape, the utility of odonates as suitable indicators to indicate health of non-aquatic (forest) habitats remains poorly understood. This study analyses the adult dragonfly assemblage pattern against spatial and temporal disturbance characteristics in Indonesias Sungai Wain Protection Forest. The core of this reserve comprises one of the few remaining fragments of primary rain forest along the East Kalimantan coast, whereas the rest of the reserve is covered by secondary forest, scrub, grassland, and farmland. Adult dragonfly assemblages at individual sampling sites were analysed in relation to (1) their intensity, (2) frequency of human-caused disturbances, and (3) the time since the last such disturbance, while controlling random variables (type of aquatic and terrestrial habitat) were removed. This study tests the effect of these factors on (1) species richness, (2) proportion of Zygoptera, (3) proportion of forest specialists, and (4) proportion of Borneos endemics. The human-induced disturbances in the rain forest resulted in pronounced changes in the taxonomical composition and functional diversity of the odonate fauna. Results reported here demonstrate that gradual changes in the odonate assemblages correspond to the degree of anthropogenic influences on forest environments. Adult odonates comprise an appropriately sensitive and versatile indicator group for identifying changes in terrestrial forest environments as well as in freshwater habitats.

Response to Comment on "Why Are There So Many Species of Herbivorous Insects in Tropical Rainforests?"

Norton and Didham suggest that differences in plant abundance between tropical and temperate forests may influence the host specificity of herbivores in these forests. We agree in principle but show that this is likely only for very rare plant species in tropical forests. Studies of herbivores hosted by rare plant species would help our understanding of tropical plant-insect interactions.

Do clones degenerate over time? Explaining the genetic variability of asexuals through population genetic models

BackgroundQuest for understanding the nature of mechanisms governing the life span of clonal organisms lasts for several decades. Phylogenetic evidence for recent origins of most clones is usually interpreted as proof that clones suffer from gradual age-dependent fitness decay (e.g. Mullers ratchet). However, we have shown that a neutral drift can also qualitatively explain the observed distribution of clonal ages. This finding was followed by several attempts to distinguish the effects of neutral and non-neutral processes. Most recently, Neiman et al. 2009 (Ann N Y Acad Sci.:1168:185-200.) reviewed the distribution of asexual lineage ages estimated from a diverse array of taxa and concluded that neutral processes alone may not explain the observed data. Moreover, the authors inferred that similar types of mechanisms determine maximum asexual lineage ages in all asexual taxa. In this paper we review recent methods for distinguishing the effects of neutral and non-neutral processes and point at methodological problems related with them.Results and DiscussionWe found that contemporary analyses based on phylogenetic data are inadequate to provide any clear-cut answer about the nature and generality of processes affecting evolution of clones. As an alternative approach, we demonstrate that sequence variability in asexual populations is suitable to detect age-dependent selection against clonal lineages. We found that asexual taxa with relatively old clonal lineages are characterised by progressively stronger deviations from neutrality.ConclusionsOur results demonstrate that some type of age-dependent selection against clones is generally operational in asexual animals, which cover a wide taxonomic range spanning from flatworms to vertebrates. However, we also found a notable difference between the data distribution predicted by available models of sequence evolution and those observed in empirical data. These findings point at the possibility that processes affecting clonal evolution differ from those described in recent studies, suggesting that theoretical models of asexual populations must evolve to address this problem in detail.ReviewersThis article was reviewed by Isa Schön (nominated by John Logsdon), Arcady Mushegian and Timothy G. Barraclough (nominated by Laurence Hurst).