Exploring the floristic diversity of tropical Africa.

BACKGROUND: Understanding the patterns of biodiversity distribution and what influences them is a fundamental pre-requisite for effective conservation and sustainable utilisation of biodiversity. Such knowledge is increasingly urgent as biodiversity responds to the ongoing effects of global climate change. Nowhere is this more acute than in species-rich tropical Africa, where so little is known about plant diversity and its distribution. In this paper, we use RAINBIO - one of the largest mega-databases of tropical African vascular plant species distributions ever compiled - to address questions about plant and growth form diversity across tropical Africa. RESULTS: The filtered RAINBIO dataset contains 609,776 georeferenced records representing 22,577 species. Growth form data are recorded for 97% of all species. Records are well distributed, but heterogeneous across the continent. Overall, tropical Africa remains poorly sampled. When using sampling units (SU) of 0.5°, just 21 reach appropriate collection density and sampling completeness, and the average number of records per species per SU is only 1.84. Species richness (observed and estimated) and endemism figures per country are provided. Benin, Cameroon, Gabon, Ivory Coast and Liberia appear as the botanically best-explored countries, but none are optimally explored. Forests in the region contain 15,387 vascular plant species, of which 3013 are trees, representing 5-7% of the estimated world's tropical tree flora. The central African forests have the highest endemism rate across Africa, with approximately 30% of species being endemic. CONCLUSIONS: The botanical exploration of tropical Africa is far from complete, underlining the need for intensified inventories and digitization. We propose priority target areas for future sampling efforts, mainly focused on Tanzania, Atlantic Central Africa and West Africa. The observed number of tree species for African forests is smaller than those estimated from global tree data, suggesting that a significant number of species are yet to be discovered. Our data provide a solid basis for a more sustainable management and improved conservation of tropical Africa's unique flora, and is important for achieving Objective 1 of the Global Strategy for Plant Conservation 2011-2020. In turn, RAINBIO provides a solid basis for a more sustainable management and improved conservation of tropical Africa's unique flora.

Copper and cobalt accumulation in plants: a critical assessment of the current state of knowledge.

This review synthesizes contemporary understanding of copper-cobalt (Cu-Co) tolerance and accumulation in plants. Accumulation of foliar Cu and Co to > 300 µg g-1 is exceptionally rare globally, and known principally from the Copperbelt of Central Africa. Cobalt accumulation is also observed in a limited number of nickel (Ni) hyperaccumulator plants occurring on ultramafic soils around the world. None of the putative Cu or Co hyperaccumulator plants appears to comply with the fundamental principle of hyperaccumulation, as foliar Cu-Co accumulation is strongly dose-dependent. Abnormally high plant tissue Cu concentrations occur only when plants are exposed to high soil Cu with a low root to shoot translocation factor. Most Cu-tolerant plants are Excluders sensu Baker and therefore setting nominal threshold values for Cu hyperaccumulation is not informative. Abnormal accumulation of Co occurs under similar circumstances in the Copperbelt of Central Africa as well as sporadically in Ni hyperaccumulator plants on ultramafic soils; however, Co-tolerant plants behave physiologically as Indicators sensu Baker. Practical application of Cu-Co accumulator plants in phytomining is limited due to their dose-dependent accumulation characteristics, although for Co field trials may be warranted on highly Co-contaminated mineral wastes because of its relatively high metal value.

Multiscale assemblage of an ectomycorrhizal fungal community: the influence of host functional traits and soil properties in a 10-ha miombo forest.

Ectomycorrhizal fungi (EMF) are highly diversified and dominant in a number of forest ecosystems. Nevertheless, their scales of spatial distribution and the underlying ecological processes remain poorly understood. Although most EMF are considered to be generalists regarding host identity, a preference toward functional strategies of host trees has never been tested. Here, the EMF community was characterised by DNA sequencing in a 10-ha tropical dry season forest-referred to as miombo-an understudied ecosystem from a mycorrhizal perspective. We used 36 soil parameters and 21 host functional traits (FTs) as candidate explanatory variables in spatial constrained ordinations for explaining the EMF community assemblage. Results highlighted that the community variability was explained by host FTs related to the 'leaf economics spectrum' (adjusted R(2) = 11%; SLA, leaf area, foliar Mg content), and by soil parameters (adjusted R(2) = 17%), notably total forms of micronutrients or correlated available elements (Al, N, K, P). Both FTs and soil generated patterns in the community at scales ranging from 75 to 375 m. Our results indicate that soil is more important than previously thought for EMF in miombo woodlands, and show that FTs of host species can be better predictors of symbiont distribution than taxonomical identity.

Copper and cobalt mobility in soil and accumulation in a metallophyte as influenced by experimental manipulation of soil chemical factors.

The influence of Fe oxides, Mn oxides and organic matter (OM) on the Cu and Co mobility in soil and accumulation in the metallophyte Anisopappus chinensis (Ac), as compared with Helianthus annuus (Ha), was experimentally investigated. Growth and accumulation response when increasing the exchangeable Cu and Co concentrations in soil were also investigated. Plants were cultivated on soil where concentrations of Cu, Co, Fe oxides, Mn oxides and OM content were varied according to 36 treatments. The OM supply decreased the Cu mobility and increased the Co mobility, resulting in decreasing the foliar Cu of Ac and increasing the foliar Co of Ha. The Fe oxides supply could increase the Cu accumulation for Ac, but was not verified for Ha. Compared with Ha, Ac increasingly accumulated Cu and Co without negative effect on plant growth while increasing Cu and Co mobility to phytotoxic concentrations. The results revealed promising perspectives for the use of Ac in Cu-contaminated environment phytoremediation applications.

RAINBIO: a mega-database of tropical African vascular plants distributions.

The tropical vegetation of Africa is characterized by high levels of species diversity but is undergoing important shifts in response to ongoing climate change and increasing anthropogenic pressures. Although our knowledge of plant species distribution patterns in the African tropics has been improving over the years, it remains limited. Here we present RAINBIO, a unique comprehensive mega-database of georeferenced records for vascular plants in continental tropical Africa. The geographic focus of the database is the region south of the Sahel and north of Southern Africa, and the majority of data originate from tropical forest regions. RAINBIO is a compilation of 13 datasets either publicly available or personal ones. Numerous in depth data quality checks, automatic and manual via several African flora experts, were undertaken for georeferencing, standardization of taxonomic names and identification and merging of duplicated records. The resulting RAINBIO data allows exploration and extraction of distribution data for 25,356 native tropical African vascular plant species, which represents ca. 89% of all known plant species in the area of interest. Habit information is also provided for 91% of these species.

Phytostabilisation of copper-contaminated soil in Katanga: an experiment with three native grasses and two amendments.

This study evaluates the feasibility of using the grass species Rendlia altera, Monocymbium ceresiiforme, Cynodon dactylon, and amendments (compost and lime) for the phytostabilisation of soils contaminated by Cu in the province of Katanga (Democratic Republic of Congo). Species were grown on control and Cu-contaminated plots (artificially contaminated with 2,500 mg kg(-1) Cu) unamended (NA), amended with 4.5 kg compost m(-2) or 0.2 kg lime m(-2). R. altera was also grown on contaminated plots amended with 22.5 kg compost m(-2) or 1 kg lime m(-2). Plant survival, growth, and reproduction were monitored for two years. Cu-concentration in leaves of R. altera and M. ceresiiforme were analysed. pH and extractable Cu (0.01 M CaCl2) in soil were analysed in April 2007 and 2008. Results showed that R. altera seems to be the best candidate because of its highest survival on NA, followed by M. ceresiiforme, while liming was necessary to ensure survival of C. dactylon. Lime increased plant reproduction and reduced Cu accumulation in leaves compared to compost. However, higher survival and number of spikes of R. altera obtained in experiment 2 with 22.5 kg compost m(-2) suggest that lime x compost interactions should be investigated in further studies.

Impacts of alien invasive plants on soil nutrients are correlated with initial site conditions in NW Europe.

Alien invasive plants are capable of modifying ecosystem function. However, it is difficult to make generalisations because impacts often appear to be species- and site-specific. In this study, we examined the impacts of seven highly invasive plant species in NW Europe (Fallopia japonica, Heracleum mantegazzianum, Impatiens glandulifera, Prunus serotina, Rosa rugosa, Senecio inaequidens, Solidago gigantea) on nutrient pools in the topsoil and the standing biomass. We tested if the impacts follow predictable patterns, across species and sites or, alternatively, if they are entirely idiosyncratic. To that end, we compared invaded and adjacent uninvaded plots in a total of 36 sites with widely divergent soil chemistry and vegetation composition. For all species, invaded plots had increased aboveground biomass and nutrient stocks in standing biomass compared to uninvaded vegetation. This suggests that enhanced nutrient uptake may be a key trait of highly invasive plant species. The magnitude and direction of the impact on topsoil chemical properties were strongly site-specific. A striking finding is that the direction of change in soil properties followed a predictable pattern. Thus, strong positive impacts (higher topsoil nutrient concentrations in invaded plots compared to uninvaded ones) were most often found in sites with initially low nutrient concentrations in the topsoil, while negative impacts were generally found under the opposite conditions. This pattern was significant for potassium, magnesium, phosphorus, manganese and nitrogen. The particular site-specific pattern in the impacts that we observed provides the first evidence that alien invasive species may contribute to a homogenisation of soil conditions in invaded landscapes.

Cost of adaptation to a metalliferous environment for Thlaspi caerulescens: a field reciprocal transplantation approach.

Field reciprocal transplantations of two metallicolous populations (Mpops) and two nonmetallicolous populations (NMpops) of Thlaspi caerulescens were performed here to determine the pattern of local adaptation and to assess the cost of adaptation of Mpops to a metalliferous environment (Menv). The role of herbivores as an important selective pressure in the nonmetalliferous environment (NMenv) was also examined. Growth, survival, fitness, life cycle and herbivore consumption were monitored for each transplant for 2 yr. Local adaptation of Mpops to their own environment was clearly demonstrated, as Mpops consistently outperformed NMpops in Menv. In NMenv, no advantage of NMpops over Mpops was detected. However, the fitness of Mpops was generally lower in NMenv than in Menv. Herbivore consumption appeared to be a significant selective pressure for Mpops in NMenv. An imbalance of selective forces between Menv and NMenv probably explains the greater local adaptation of Mpops. Therefore, colonization of NMenv by Mpops appears possible. Although Mpops were able to survive and reproduce in NMenv, they nevertheless expressed a cost attributable in part to their higher susceptibility to herbivores.

Development of Issoria lathonia (Lepidoptera: Nymphalidae) on zinc-accumulating and nonaccumulating Viola species (Violaceae).

The larvae of Issoria lathonia L. feed in natural conditions on several Viola spp., among which are the zinc-accumulating Viola calaminaria (Gingins) Lej. and the nonmetal-accumulating Viola tricolor L. To examine how I. lathonia caterpillars cope with the naturally high foliar zinc concentration of V. calaminaria, we compared the growth of caterpillars reared on leaves varying in zinc concentration. Larvae were fed in controlled conditions with V. calaminaria and V. tricolor grown on noncontaminated soil (i.e., two low-Zn diets) and with V. calaminaria grown on zinc-enriched soil (i.e., one high-Zn diet). Larvae had a higher growth rate when fed with noncontaminated V. calaminaria compared to zinc-enriched V. calaminaria, suggesting that zinc slows down larval growth. However, larvae consumed more leaves of zinc-enriched V. calaminaria (+45%; estimated from fecal mass) compared with noncontaminated V. calaminaria, suggesting that zinc accumulation would not be advantageous to plants. Caterpillars reared on high-zinc leaves regulate their internal zinc concentration through excretion of highly metal-concentrated feces. When kinetics of growth on both low-zinc diets were compared, it appeared that larval development was faster on noncontaminated V. calaminaria than on V. tricolor. This suggests that more nutrients or less feeding inhibitors in V. calaminaria account for fastest growth. Developmental rates on V. tricolor and on zinc-enriched V. calaminaria were similar, despite the high leaf zinc concentration of the latter species. Together with the abundance of V. calaminaria on calamine soils, this may explain why the largest populations of I. lathonia develop on V. calaminaria in Belgium.

Do metal-rich plants deter herbivores? A field test of the defence hypothesis.

Some plant species growing on metalliferous soils are able to accumulate heavy metals in their shoots up to very high concentrations, but the selective advantage of this behaviour is still unknown. The most popular hypothesis, that metals protect plants against herbivores, has been tested several times in laboratory conditions, with contradictory results. We carried out the first large-scale test of the defence hypothesis in eight natural populations of the model Zn hyperaccumulator Thlaspi caerulescens J. and C. Presl (Brassicaceae). In two climatic regions (temperate, Belgium-Luxembourg, and Mediterranean, southern France), we worked in metalliferous and in normal, uncontaminated environments, with plants spanning a wide range of Zn concentrations. We also examined the importance of glucosinolates (main secondary metabolites of Brassicaceae) as antiherbivore defences. When exposed to natural herbivore populations, T. caerulescens suffered lower herbivory pressures in metal-enriched soils than in normal soils, both in Belgium-Luxembourg and in southern France. The trapping of gastropods shows an overall lower population density in metalliferous compared to normal environments, which suggests that herbivory pressure from gastropods is lower on metalliferous soils. In addition, foliar concentration of glucosinolates was constitutively lower in all populations from metal-enriched soils, suggesting that these have evolved towards lower investment in organic defences in response to lower herbivory pressure. The Zn concentration of plants had a protective role only for Belgian metallicolous plants when transplanted in normal soils of Luxembourg. These results do not support the hypothesis that Zn plays a key role in the protection of T. caerulescens against enemies. In contrast, glucosinolates appear to be directly involved in the defence of this hyperaccumulator against herbivores.

Reaction norms of life history traits in response to zinc in Thlaspi caerulescens from metalliferous and nonmetalliferous sites.

* We examined phenotypic plasticity of fitness components in response to zinc (Zn) in the Zn hyperaccumulator, Thlaspi caerulescens. * Two populations from Zn-enriched soils (M) and two populations from normal soils (NM) were grown in pots at three Zn concentrations (0, 1000 and 8000 mg kg(-1) Zn), for an entire life cycle. Growth, Zn accumulation and fitness components were assessed. * Based on vegetative growth, M and NM populations had similar Zn tolerance at 1000 mg kg(-1) Zn. However, reproductive output was markedly decreased in NM at 1000 and 8000 mg kg(-1) Zn. In M populations, Zn did not affect fitness. However, low Zn status enhanced reproductive output in year 1 compared with year 2 and decreased survival after the first flowering season. * M populations are able to achieve equal fitness across a broad range of Zn concentrations in soil by different combinations of fecundity and longevity. No cost of higher tolerance was demonstrated in M populations. Reproductive traits appeared to be a more sensitive indicator of tolerance than vegetative growth.

Thlaspi caerulescens on nonmetalliferous soil in Luxembourg: ecological niche and genetic variation in mineral element composition.

Forty-seven populations of Thlaspi caerulescens in Luxembourg were characterised for population size, soil mineral element composition and other habitat characteristics. Foliar concentrations of eight elements were assessed in 15 populations in the field and in eight populations cultivated in zinc (Zn)-cadmium (Cd)-nickel (Ni)-enriched soil. T. caerulescens favoured stony soil developed on steep, south-facing Emsian shale outcrops. All soil samples were nonmetalliferous. Soil pH ranged from 4.2 to 6.9. Field-growing plants had very high concentrations of heavy metals in the leaves (Zn, 3000-13 000 mg kg(-1); Cd, 11-44 mg kg(-1); Ni, 38-473 mg kg(-1)). Positive soil-plant correlations existed for Zn and Mn. In cultivation, significant genetic variation was found for biomass and six of eight mineral elements. For Cd and Zn, variation range among 48 half-sib families was two-fold (Cd, 183-334 mg kg(-1); Zn, 8030-16 295 mg kg(-1)). Most of the variation occurred among populations, consistent with the selfing mating system of those populations. There was a tight Zn-Cd genetic correlation (r = +0.83, P < 0.0001). The significance of the results to the conservation of T. caerulescens in Luxembourg is briefly discussed.

An experimental investigation of life history and plasticity in two cytotypes of Polygonum aviculare L. subsp. aviculare that coexist in an abandoned arable field.

Polygonum aviculare subsp. aviculare is an annual selfing weed common in abandoned arable fields where it occurs as a widespread hexaploid cytotype (6x=60) and a rarer tetraploid cytotype (4x=40). The basis of phenological differentiation between the two cytotypes observed in a natural population where they coexist was examined in a greenhouse experiment comprising six soil conditions consisting of factorial combinations of two levels of fertility and three pot sizes. The environmental and genetic component of variation in 11 life history and morphological traits was quantified. Even though all traits except life span were plastic the two cytotypes appear to have evolved contrasting life history strategies and it is inferred that this can account for the temporal niche differentiation observed in the abandoned field during the first year of dereliction. Tetraploids are short-lived plants allocating a high proportion of their biomass to reproduction and completing their life cycle before July when the plant cover is sparse. Hexaploids are larger, later flowering, longer lived, plants with a lower reproductive effort and a higher final seed yield; it is inferred that these traits enable the hexaploids to compete successfully with the dense herbaceous layer of summer annuals that develops in the course of the first year of secondary succession. Differentiation in phenotypic plasticity between the two cytotypes was interpreted as indicative of higher opportunism and lower tolerance of poor soils and restricted rooting space in the hexaploid compared to the tetraploid cytotype.