On the importance of root traits in seedlings of tropical tree species.

Plant biomass allocation may be optimized to acquire and conserve resources. How trade-offs in the allocation of tropical tree seedlings depend on different stressors remains poorly understood. Here we test whether above- and below-ground traits of tropical tree seedlings could explain observed occurrence along gradients of resources (light, water) and defoliation (fire, herbivory). We grew 24 tree species occurring in five African vegetation types, varying from dry savanna to moist forest, in a glasshouse for 6 months, and measured traits associated with biomass allocation. Classification based on above-ground traits resulted in clusters representing savanna and forest species, with low and high shoot investment, respectively. Classification based on root traits resulted in four clusters representing dry savanna, humid savanna, dry forest and moist forest, characterized by a deep mean rooting depth, root starch investment, high specific root length in deeper soil layers, and high specific root length in the top soil layer, respectively. In conclusion, tree seedlings in this study show root trait syndromes, which vary along gradients of resources and defoliation: seedlings from dry areas invest in deep roots, seedlings from shaded environments optimize shoot investment, and seedlings experiencing frequent defoliation store resources in the roots.

On the relationship between fire regime and vegetation structure in the tropics.

We assessed data from 11 experiments examining the effects of the timing and/or frequency of fire on tropical forest and/or savanna vegetation structure over one decade or more. The initial 'control treatment' in many such cases consisted of previously cleared land. This is as opposed to natural vegetation subject to some sort of endogenous fire regime before the imposition of fire treatments. Effects of fire on fractional foliar cover are up to 10-fold greater when clearing pre-treatments are imposed. Moreover, because many of the 'classic' fire trials were initialised with applied management questions in mind, most have also used burning regimes much more frequent and/or severe than those occurring in the absence of human activity. Once these factors are taken into account, our modelling analysis shows that nonanthropogenic fire regimes serve to reduce canopy vegetative cover to a much lower extent than has previously been argued to be the case. These results call into question the notion that fire effects on tropical vegetation can be of a sufficient magnitude to maintain open-type savanna ecosystems under climatic/soil regimes otherwise sufficient to give rise to a more luxurious forest-type vegetation cover.

Response of bats to light with different spectra: light-shy and agile bat presence is affected by white and green, but not red light.

Artificial light at night has shown a remarkable increase over the past decades. Effects are reported for many species groups, and include changes in presence, behaviour, physiology and life-history traits. Among these, bats are strongly affected, and how bat species react to light is likely to vary with light colour. Different spectra may therefore be applied to reduce negative impacts. We used a unique set-up of eight field sites to study the response of bats to three different experimental light spectra in an otherwise dark and undisturbed natural habitat. We measured activity of three bat species groups around transects with light posts emitting white, green and red light with an intensity commonly used to illuminate countryside roads. The results reveal a strong and spectrum-dependent response for the slow-flying Myotis and Plecotus and more agile Pipistrellus species, but not for Nyctalus and Eptesicus species. Plecotus and Myotis species avoided white and green light, but were equally abundant in red light and darkness. The agile, opportunistically feeding Pipistrellus species were significantly more abundant around white and green light, most likely because of accumulation of insects, but equally abundant in red illuminated transects compared to dark control. Forest-dwelling Myotis and Plecotus species and more synanthropic Pipistrellus species are thus least disturbed by red light. Hence, in order to limit the negative impact of light at night on bats, white and green light should be avoided in or close to natural habitat, but red lights may be used if illumination is needed.

Artificial night lighting inhibits feeding in moths.

One major, yet poorly studied, change in the environment is nocturnal light pollution, which strongly alters habitats of nocturnally active species. Artificial night lighting is often considered as driving force behind rapid moth population declines in severely illuminated countries. To understand these declines, the question remains whether artificial light causes only increased mortality or also sublethal effects. We show that moths subjected to artificial night lighting spend less time feeding than moths in darkness, with the shortest time under light conditions rich in short wavelength radiation. These findings provide evidence for sublethal effects contributing to moth population declines. Because effects are strong under various types of light compared with dark conditions, the potential of spectral alterations as a conservation tool may be overestimated. Therefore, restoration and maintenance of darkness in illuminated areas is essential for reversing declines of moth populations.

Biome-specific effects of nitrogen and phosphorus on the photosynthetic characteristics of trees at a forest-savanna boundary in Cameroon.

Photosynthesis/nutrient relationships of proximally growing forest and savanna trees were determined in an ecotonal region of Cameroon (Africa). Although area-based foliar N concentrations were typically lower for savanna trees, there was no difference in photosynthetic rates between the two vegetation formation types. Opposite to N, area-based P concentrations were-on average-slightly lower for forest trees; a dependency of photosynthetic characteristics on foliar P was only evident for savanna trees. Thus savanna trees use N more efficiently than their forest counterparts, but only in the presence of relatively high foliar P. Along with some other recent studies, these results suggest that both N and P are important modulators of woody tropical plant photosynthetic capacities, influencing photosynthetic metabolism in different ways that are also biome specific. Attempts to find simple unifying equations to describe woody tropical vegetation photosynthesis-nutrient relationships are likely to meet with failure, with ecophysiological distinctions between forest and savanna requiring acknowledgement.

Co-limitation of photosynthetic capacity by nitrogen and phosphorus in West Africa woodlands.

Photosynthetic leaf traits were determined for savanna and forest ecosystems in West Africa, spanning a large range in precipitation. Standardized major axis fits revealed important differences between our data and reported global relationships. Especially for sites in the drier areas, plants showed higher photosynthetic rates for a given N or P when compared with relationships from the global data set. The best multiple regression for the pooled data set estimated V(cmax) and J(max) from N(DW) and S. However, the best regression for different vegetation types varied, suggesting that the scaling of photosynthesis with leaf traits changed with vegetation types. A new model is presented representing independent constraints by N and P on photosynthesis, which can be evaluated with or without interactions with S. It assumes that limitation of photosynthesis will result from the least abundant nutrient, thereby being less sensitive to the allocation of the non-limiting nutrient to non-photosynthetic pools. The model predicts an optimum proportionality for N and P, which is distinct for V(cmax) and J(max) and inversely proportional to S. Initial tests showed the model to predict V(cmax) and J(max) successfully for other tropical forests characterized by a range of different foliar N and P concentrations.

Plant-soil feedback of native and range-expanding plant species is insensitive to temperature.

Temperature change affects many aboveground and belowground ecosystem processes. Here we investigate the effect of a 5 degrees C temperature increase on plant-soil feedback. We compare plant species from a temperate climate region with immigrant plants that originate from warmer regions and have recently shifted their range polewards. We tested whether the magnitude of plant-soil feedback is affected by ambient temperature and whether the effect of temperature differs between these groups of plant species. Six European/Eurasian plant species that recently colonized the Netherlands (non-natives), and six related species (natives) from the Netherlands were selected. Plant-soil feedback of these species was determined by comparing performance in conspecific and heterospecific soils. In order to test the effect of temperature on these plant-soil feedback interactions, the experiments were performed at two greenhouse temperatures of 20/15 degrees C and 25/20 degrees C, respectively. Inoculation with unconditioned soil had the same effect on natives and non-natives. However, the effect of conspecific conditioned soil was negative compared to heterospecific soil for natives, but was positive for non-natives. In both cases, plant-soil interactions were not affected by temperature. Therefore, we conclude that the temperature component of climate change does not affect the direction, or strength of plant-soil feedback, neither for native nor for non-native plant species. However, as the non-natives have a more positive soil feedback than natives, climate warming may introduce new plant species in temperate regions that have less soil-borne control of abundance.

Growth form and seasonal variation in leaf gas exchange of Colophospermum mopane savanna trees in northwest Botswana.

We investigated differences in physiological and morphological traits between the tall and short forms of mopane (Colophospermum mopane (Kirk ex Benth.) Kirk ex J. Léonard) trees growing near Maun, Botswana on a Kalahari sandveld overlying an impermeable calcrete duricrust. We sought to determine if differences between the two physiognomic types are attributable to the way they exploit available soil water. The tall form, which was located on deeper soil than the short form (5.5 versus 1.6 m), had a lower leaf:fine root biomass ratio (1:20 versus 1:6), but a similar leaf area index (0.9-1.0). Leaf nitrogen concentrations varied between 18 and 27 mg g(-1) and were about 20% higher in the tall form than in the short form. Maximum net assimilation rates (A sat) occurred during the rainy seasons (March-April 2000 and January-February 2001) and were similar in the tall and short forms (15-22 micromol m(-2) s(-1)) before declining to less than 10 micromol m(-2) s(-1) at the end of the rainy season in late April. As the dry season progressed, A sat, soil water content, predawn leaf water potential (Psi pd) and leaf nitrogen concentration declined rapidly. Before leaf abscission, Psi pd was more negative in the short form (-3.4 MPa) than in the tall form (-2.7 MPa) despite the greater availability of soil water beneath the short form trees. This difference appeared attributable to differences in root depth and density between the physiognomic types. Stomatal regulation of water use and carbon assimilation differed between years, with the tall form having a consistently more conservative water-use strategy as the dry season progressed than the short form.

Contributions of woody and herbaceous vegetation to tropical savanna ecosystem productivity: a quasi-global estimate.

To estimate the relative contributions of woody and herbaceous vegetation to savanna productivity, we measured the 13C/12C isotopic ratios of leaves from trees, shrubs, grasses and the surface soil carbon pool for 22 savannas in Australia, Brazil and Ghana covering the full savanna spectrum ranging from almost pure grassland to closed woodlands on all three continents. All trees and shrubs sampled were of the C3 pathway and all grasses of the C4 pathway with the exception of Echinolaena inflexa (Poir.) Chase, a common C3 grass of the Brazilian cerrado. By comparing the carbon isotopic compositions of the plant and carbon pools, a simple model relating soil delta 13C to the relative abundances of trees + shrubs (woody plants) and grasses was developed. The model suggests that the relative proportions of a savanna ecosystem's total foliar projected cover attributable to grasses versus woody plants is a simple and reliable index of the relative contributions of grasses and woody plants to savanna net productivity. Model calibrations against woody tree canopy cover made it possible to estimate the proportion of savanna productivity in the major regions of the world attributable to trees + shrubs and grasses from ground-based observational maps of savanna woodiness. Overall, it was estimated that 59% of the net primary productivity (Np) of tropical savannas is attributable to C4 grasses, but that this proportion varies significantly within and between regions. The C4 grasses make their greatest relative contribution to savanna Np in the Neotropics, whereas in African regions, a greater proportion of savanna Np is attributable to woody plants. The relative contribution of C4 grasses in Australian savannas is intermediate between those in the Neotropics and Africa. These differences can be broadly ascribed to large scale differences in soil fertility and rainfall.

Artificial light at night causes diapause inhibition and sex-specific life history changes in a moth.

Rapidly increasing levels of light pollution subject nocturnal organisms to major alterations of their habitat, the ecological consequences of which are largely unknown. Moths are well-known to be attracted to light at night, but effects of light on other aspects of moth ecology, such as larval development and life-history, remain unknown. Such effects may have important consequences for fitness and thus for moth population sizes. To study the effects of artificial night lighting on development and life-history of moths, we experimentally subjected Mamestra brassicae (Noctuidae) caterpillars to low intensity green, white, red or no artificial light at night and determined their growth rate, maximum caterpillar mass, age at pupation, pupal mass and pupation duration. We found sex-specific effects of artificial light on caterpillar life-history, with male caterpillars subjected to green and white light reaching a lower maximum mass, pupating earlier and obtaining a lower pupal mass than male caterpillars under red light or in darkness. These effects can have major implications for fitness, but were absent in female caterpillars. Moreover, by the time that the first adult moth from the dark control treatment emerged from its pupa (after 110 days), about 85% of the moths that were under green light and 83% of the moths that were under white light had already emerged. These differences in pupation duration occurred in both sexes and were highly significant, and likely result from diapause inhibition by artificial night lighting. We conclude that low levels of nocturnal illumination can disrupt life-histories in moths and inhibit the initiation of pupal diapause. This may result in reduced fitness and increased mortality. The application of red light, instead of white or green light, might be an appropriate measure to mitigate negative artificial light effects on moth life history.

Foliar trait contrasts between African forest and savanna trees: genetic versus environmental effects.

Variations in leaf mass per unit area (Ma) and foliar concentrations of N, P, C, K, Mg and Ca were determined for 365 trees growing in 23 plots along a West African precipitation gradient ranging from 0.29 to 1.62m a-1. Contrary to previous studies, no marked increase in Ma with declining precipitation was observed, but savanna tree foliar [N] tended to be higher at the drier sites (mass basis). Generally, Ma was slightly higher and [N] slightly lower for forest vs savanna trees with most of this difference attributable to differences in soil chemistry. No systematic variations in [P], [Mg] and [Ca] with precipitation or between trees of forest vs savanna stands were observed. We did, however, find a marked increase in foliar [K] of savanna trees as precipitation declined, with savanna trees also having a significantly lower [K] than those of nearby forest. These differences were not related to differences in soil nutrient status and were accompanied by systematic changes in [C] of opposite sign. We suggest an important but as yet unidentified role for K in the adaption of savanna species to periods of limited water availability; with foliar [K] being also an important factor differentiating tree species adapted to forest vs savanna soils within the 'zone of transition' of Western Africa.