Whole-ecosystem experimental manipulations of tropical forests.

Tropical forests are highly diverse systems involving extraordinary numbers of interactions between species, with each species responding in a different way to the abiotic environment. Understanding how these systems function and predicting how they respond to anthropogenic global change is extremely challenging. We argue for the necessity of 'whole-ecosystem' experimental manipulations, in which the entire ecosystem is targeted, either to reveal the functioning of the system in its natural state or to understand responses to anthropogenic impacts. We survey the current range of whole-ecosystem manipulations, which include those targeting weather and climate, nutrients, biotic interactions, human impacts, and habitat restoration. Finally we describe the unique challenges and opportunities presented by such projects and suggest directions for future experiments.This review was initiated during a symposium on ‘The effects of large scale manipulations of tropical forests on arthropod assemblages’ at the INTECOL 2013 congress, London 18–23 August 2013. T.M.F. is funded by the Australian Research Council (DP140101541), T.M.F. and R.M.E. by Yayasan Sime Darby, TMF and Y.B. by the project Biodiversity of Forest Ecosystems (CZ.1.07/2.3.00/20.0064) co-financed by the European Social Fund and the state budget of the Czech Republic, and T.M.F. Y.B. and V.N. by the Czech Science Foundation (GACR 14-32302S, 14-36098G, 14-04258S respectively). Y.B. is also supported by the Sistema Nacional de Investigacio´n of Panama. E.C.T. is supported by funds from PT SMART Research Institute and the Isaac Newton Trust, Cambridge. R.M.E. is supported by European Research Council Project number 281986. We are grateful to Maureen Fayle, Andrew Hector, Jan Leps, Scott Miller, Kalsum M. Yusah, Paul Craze, and two anonymous reviewers for advice during the drafting of the manuscript, and Jennifer Balch for additional information regarding her burning experiments.This is the final published version. It first appeared at http://www.cell.com/trends/ecology-evolution/abstract/S0169-5347%2815%2900069-5

Scientists' warning on climate change and insects

Climate warming is considered to be among the most serious of anthropogenic stresses to the environment, because it not only has direct effects on biodiversity, but it also exacerbates the harmful effects of other human-mediated threats. The associated consequences are potentially severe, particularly in terms of threats to species preservation, as well as in the preservation of an array of ecosystem services provided by biodiversity. Among the most affected groups of animals are insects—central components of many ecosystems—for which climate change has pervasive effects from individuals to communities. In this contribution to the scientists' warning series, we summarize the effect of the gradual global surface temperature increase on insects, in terms of physiology, behavior, phenology, distribution, and species interactions, as well as the effect of increased frequency and duration of extreme events such as hot and cold spells, fires, droughts, and floods on these parameters. We warn that, if no action is taken to better understand and reduce the action of climate change on insects, we will drastically reduce our ability to build a sustainable future based on healthy, functional ecosystems. We discuss perspectives on relevant ways to conserve insects in the face of climate change, and we offer several key recommendations on management approaches that can be adopted, on policies that should be pursued, and on the involvement of the general public in the protection effort.</p

Comparison of rainforest butterfly assemblages across three biogeographical regions using standardized protocols

Insects, like most other organisms, are more diverse in tropical than in temperate regions, but standardized comparisons of diversity among tropical regions are rare. Disentangling the effects of ecological, evolutionary, and biogeographic factors on community diversity requires standardized protocols and long-term studies. We compared the abundance and diversity of butterflies using standardised ‘Pollard walk’ transect counts in the understory of closed-canopy lowland rainforests in Panama (Barro Colorado Island, BCI), Thailand (Khao Chong, KHC) and Papua New Guinea (Wanang, WAN). We observed 1792, 1797 and 3331 butterflies representing 128, 131 and 134 species during 230, 231 and 120 transects at BCI, KHC and WAN, respectively. When corrected for length and duration of transects, butterfly abundance and species richness were highest at WAN and KHC, respectively. Although high butterfly abundance at WAN did not appear to result from methodological artefacts, the biological meaning of this observation remains obscure. The WAN site appeared as floristically diverse as KHC, but supported lower butterfly diversity. This emphasizes that factors other than plant diversity, such as biogeographic history, may be crucial for explaining butterfly diversity. The KHC butterfly fauna may be unusually species rich because the site is at a biogeographic crossroads between the Indochinese and Sundaland regions. In contrast, WAN is firmly within the Australian biogeographic region and relatively low species numbers may result from island biogeographic processes. The common species at each of the three sites shared several traits: fruit and nectar feeders were equally represented, more than half of common species fed on either epiphytes or lianas as larvae, and their range in wing sizes was similar. These observations suggest that Pollard walks in different tropical rainforests target similar assemblages of common species, and, hence, represent a useful tool for long-term monitoring of rainforest butterfly assemblages.Organismic and Evolutionary Biolog

The role of herbivorous insects and pathogens in the regeneration dynamics of Guazuma ulmifolia in Panama

A significant proportion of the mortality of rainforest trees occurs during early life stages (seeds and seedlings), but mortality agents are often elusive. Our study investigated the role of herbivorous insects and pathogens in the early regeneration dynamics of Guazuma ulmifolia (Malvaceae), an important tree species in agroforestry in Central America. We reared pre-dispersal insect seed predators from G. ulmifolia seeds in Panama. We also carried out an experiment, controlling insects and pathogens using insecticide and/or fungicide treatments, as well as seed density, and compared survivorship of G. ulmifolia seeds and seedlings among treatments and relative to untreated control plots. We observed (1) high pre-dispersal attack (92%) of the fruits of G. ulmifolia, mostly by anobiine and bruchine beetles; (2) negligible post-dispersal attack of isolated seeds by insects and pathogens; (3) slow growth and high mortality (> 95%) of seedlings after 14 weeks; (4) low insect damage on seedlings; and (5) a strong positive correlation between seedling mortality and rainfall. We conclude that for G. ulmifolia at our study site the pre-dispersal seed stage is by far the most sensitive stage to insects and that their influence on seedling mortality appears to be slight as compared to that of inclement weather. Thus, the regeneration of this important tree species may depend on effective primary dispersal of seeds by vertebrates (before most of the seed crop is lost to insects), conditioned by suitable conditions in which the seedlings can grow

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.1012SolVin-Solvay SASTRIUnited Nations Environment ProgrammeSmithsonian Institution (Walcott Fund)European Science FoundationGlobal Canopy ProgrammeCzech Science foundation GACR grant [14-36098G]European Social FundCzech Ministry of Education [CZ.1.07/2.3.00/20.0064]U.S. National Science Fundation [DEB-0841885]Australian Research Council [FT100100040]Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)"Investissement d'Avenir'' grant [ANR-10-LABX-25-01]Norwegian Research CouncilGrant Agency of the Czech Republic [14-36098G

Insect assemblages attacking seeds and fruits in a rainforest in Thailand

Insect seed predators are important agents of mortality for tropical trees, but little is known about the impact of these herbivores in rainforests. During three years at Khao Chong (KHC) in southern Thailand we reared 17,555 insects from 343.2 kg or 39,252 seeds/fruits representing 357 liana and tree species. A commented list of the 243 insect species identified is provided, with details about their host plants. We observed that: (1) about 43% of identified species can be considered pests. Most were seed eaters, particularly on dry fruits. (2) About 19% of parasitoid species (all Opiinae) for which we could determine whether their primary insect host was a pest or not (all Bactrocera spp. breeding in fruits) can be considered beneficials. (3) The seeds/fruits of about 28% of the plant species in this forest were free of attack. Phyllanthaceae, Rubiaceae, and Meliaceae were attacked relatively infrequently; in contrast, Annonaceae, Fabaceae, Sapindaceae, and Myristicaceae were more heavily attacked. There was no apparent effect of plant phylogeny on rates of attack but heavily attacked tree species had larger basal area in the KHC plot than rarely attacked tree species. (4) Insects reared from fleshy fruits were more likely to exhibit relatively stable populations compared to insects reared from dry fruits, but this was not true of insects reared from dipterocarps, which appeared to have relatively stable populations throughout the study period. We tentatively conclude that insects feeding on seeds and fruits have little effect on observed levels of host abundance in this forest

Thermoregulatory ability and mechanism do not differ consistently between neotropical and temperate butterflies

Climate change is a major threat to species worldwide, yet it remains uncertain whether tropical or temperate species are more vulnerable to changing temperatures. To further our understanding of this, we used a standardised field protocol to (1) study the buffering ability (ability to regulate body temperature relative to surrounding air temperature) of neotropical (Panama) and temperate (the United Kingdom, Czech Republic and Austria) butterflies at the assemblage and family level, (2) determine if any differences in buffering ability were driven by morphological characteristics and (3) used ecologically relevant temperature measurements to investigate how butterflies use microclimates and behaviour to thermoregulate. We hypothesised that temperate butterflies would be better at buffering than neotropical butterflies as temperate species naturally experience a wider range of temperatures than their tropical counterparts. Contrary to our hypothesis, at the assemblage level, neotropical species (especially Nymphalidae) were better at buffering than temperate species, driven primarily by neotropical individuals cooling themselves more at higher air temperatures. Morphology was the main driver of differences in buffering ability between neotropical and temperate species as opposed to the thermal environment butterflies experienced. Temperate butterflies used postural thermoregulation to raise their body temperature more than neotropical butterflies, probably as an adaptation to temperate climates, but the selection of microclimates did not differ between regions. Our findings demonstrate that butterfly species have unique thermoregulatory strategies driven by behaviour and morphology, and that neotropical species are not likely to be more inherently vulnerable to warming than temperate species

A highly-resolved food web for insect seed predators in a species-rich tropical forest

The top-down and indirect effects of insects on plant communities depend on patterns of host use, which are often poorly documented, particularly in species-rich tropical forests. At Barro Colorado Island, Panama, we compiled the first food web quantifying trophic interactions between the majority of co-occurring woody plant species and their internally-feeding insect seed predators. Our study is based on more than 200,000 fruits representing 478 plant species, associated with 369 insect species. Insect host-specificity was remarkably high: only 20% of seed predator species were associated with more than one plant species, while each tree species experienced seed predation from a median of two insect species. Phylogeny, but not plant traits, explained patterns of seed predator attack. These data suggest that seed predators are unlikely to mediate indirect interactions such as apparent competition between plant species, but are consistent with their proposed contribution to maintaining plant diversity via the Janzen-Connell mechanism