Novel Insights Into the Hyperaccumulation Syndrome in Pycnandra (Sapotaceae).

The discovery of nickel hyperaccumulation, in Pycnandra acuminata, was the start of a global quest in this fascinating phenomenon. Despite recent advances in the physiology and molecular genetics of hyperaccumulation, the mechanisms and tolerance of Ni accumulation in the most extreme example reported to date, P. acuminata, remains enigmatic. We conducted a hydroponic experiment to establish Ni tolerance levels and translocation patterns in roots and shoots of P. acuminata, and analyzed elemental partitioning to gain insights into Ni regulation. We combined a phylogeny and foliar Ni concentrations to assess the incidence of hyperaccumulation within the genus Pycnandra. Hydroponic dosing experiments revealed that P. acuminata can resist extreme Ni concentrations in solution (up to 3,000 µM), and dosing at 100 µM Ni was beneficial to growth. All plant parts were highly enriched in Ni, but the latex had extreme Ni concentrations (124,000 µg g-1). Hyperaccumulation evolved independently in only two subgenera and five species of the genus Pycnandra. The extremely high level of Ni tolerance is posited to derive from the unique properties of laticifers. The evolutionary and ecological significance of Ni hyperaccumulation in Pycnandra is discussed in light of these findings. We suggest that Ni-rich laticifers might be more widespread in the plant kingdom and that more investigation is warranted.

Nickel hyperaccumulation in New Caledonian Hybanthus (Violaceae) and occurrence of nickel-rich phloem in Hybanthus austrocaledonicus.

BACKGROUND AND AIMS: Hybanthus austrocaledonicus (Violaceae) is a nickel (Ni) hyperaccumulator endemic to New Caledonia. One of the specimens stored at the local herbarium had a strip of bark with a remarkably green phloem tissue attached to the sheet containing over 4 wt% Ni. This study aimed to collect field samples from the original H. austrocaledonicus locality to confirm the nature of the green 'nickel-rich phloem' in this taxon and to systematically assess the occurrence of Ni hyperaccumulation in H. austrocaledonicus and Hybanthus caledonicus populations. METHODS: X-ray fluorescence spectroscopy scanning of all collections of the genus Hybanthus (236 specimens) was undertaken at the Herbarium of New Caledonia to reveal incidences of Ni accumulation in populations of H. austrocaledonicus and H. caledonicus. In parallel, micro-analytical investigations were performed via synchrotron X-ray fluorescence microscopy (XFM) and scanning electron microscopy with X-ray microanalysis (SEM-EDS). KEY RESULTS: The extensive scanning demonstrated that Ni hyperaccumulation is not a characteristic common to all populations in the endemic Hybanthus species. Synchrotron XFM revealed that Ni was exclusively concentrated in the epidermal cells of the leaf blade and petiole, conforming with the majority of (tropical) Ni hyperaccumulator plants studied to date. SEM-EDS of freeze-dried and frozen-hydrated samples revealed the presence of dense solid deposits in the phloem bundles that contained >8 wt% nickel. CONCLUSIONS: The occurrence of extremely Ni-rich green phloem tissues appears to be a characteristic feature of tropical Ni hyperaccumulator plants.

In situ observations of the basal angiosperm Amborella trichopoda reveal a long fruiting cycle overlapping two annual flowering periods.

Amborella trichopoda is the sole living angiosperm species belonging to the sister lineage of all other extant flowering plants. In the last decade, the species has been the focus of many phylogenetic, genomic and reproductive biology studies, bringing new highlights regarding the evolution of flowering plants. However, little attention has been paid to in situ A. trichopoda populations, particularly to their fruiting cycle. In this study, an A. trichopoda population was observed during three annual flowering cycles. Individuals and branches were labeled in order to monitor the fruiting cycle precisely, from the flowering stage until the abscission of the fruit. Fruit exocarp was green during the first 9 months following flowering, turned red when the next flowering started a year later then remained on the branch during another year, between fruit ripping and abscission. Presence of fruits with two stages of maturity on shrubs was always noticed. Germination tests showed that seeds acquired their germination capacity 1 year after flowering, when fruits changed color. A. trichopoda's fruiting cycle is a long process overlapping two annual flowering periods. These results introduce a new model for flowering and fruiting cycles. The availability of mature seeds on shrubs for more than 1 year is likely to maximize opportunities to be dispersed, thus promoting the survival of this basal angiosperm.

Environmental correlates for tree occurrences, species distribution and richness on a high-elevation tropical island.

High-elevation tropical islands are ideally suited for examining the factors that determine species distribution, given the complex topographies and climatic gradients that create a wide variety of habitats within relatively small areas. New Caledonia, a megadiverse Pacific archipelago, has long focussed the attention of botanists working on the spatial and environmental ranges of specific groups, but few studies have embraced the entire tree flora of the archipelago. In this study we analyse the distribution of 702 native species of rainforest trees of New Caledonia, belonging to 195 genera and 80 families, along elevation and rainfall gradients on ultramafic (UM) and non-ultramafic (non-UM) substrates. We compiled four complementary data sources: (i) herbarium specimens, (ii) plots, (iii) photographs and (iv) observations, totalling 38 936 unique occurrences distributed across the main island. Compiled into a regular 1-min grid (1.852 × 1.852 km), this dataset covered ∼22 % of the island. The studied rainforest species exhibited high environmental tolerance; 56 % of them were not affiliated to a substrate type and they exhibited wide elevation (average 891 ± 332 m) and rainfall (average 2.2 ± 0.8 m year(-1)) ranges. Conversely their spatial distribution was highly aggregated, which suggests dispersal limitation. The observed species richness was driven mainly by the density of occurrences. However, at the highest elevations or rainfalls, and particularly on UM, the observed richness tends to be lower, independently of the sampling effort. The study highlights the imbalance of the dataset in favour of higher values of rainfall and of elevation. Projected onto a map, under-represented areas are a guide as to where future sampling efforts are most required to complete our understanding of rainforest tree species distribution.

Mining in New Caledonia: environmental stakes and restoration opportunities.

New Caledonia is a widely recognised marine and terrestrial biodiversity hot spot. However, this unique environment is under increasing anthropogenic pressure. Major threats are related to land cover change and include fire, urban sprawling and mining. Resulting habitat loss and fragmentation end up in serious erosion of the local biodiversity. Mining is of particular concern due to its economic significance for the island. Open cast mines were exploited there since 1873, and scraping out soil to access ores wipes out flora. Resulting perturbations on water flows and dramatic soil erosion lead to metal-rich sediment transport downstream into rivers and the lagoon. Conflicting environmental and economic aspects of mining are discussed in this paper. However, mining practices are also improving, and where impacts are inescapable ecological restoration is now considered. Past and ongoing experiences in the restoration of New Caledonian terrestrial ecosystems are presented and discussed here. Economic use of the local floristic diversity could also promote conservation and restoration, while providing alternative incomes. In this regard, Ecocatalysis, an innovative approach to make use of metal hyperaccumulating plants, is of particular interest.

Phytoextraction from mine spoils: insights from New Caledonia.

Increasing pressure on mineral resources has drawn research efforts into innovative supply and recycling. Metal-rich biomass produced in phytoextraction recently proved an interesting starting material for green chemistry. It allows the production of new catalysts, referred to as ecocatalysts. Ecocatalysts provide increased yields in chemical production and increased regio- and chemo-selectivity, which result in high added value. This new approach to using metal-rich biomass could spur the development of phytoextraction, a technique considered promising for long, yet without credible economic outlets. In this regard, metallophyte biodiversity hotspots, such as New Caledonia, are of particular interest for biomass supply. Potential phytoextraction from mine spoils using two species endemic to New Caledonia is discussed here. Geissois pruinosa, a hypernickelophore, and Grevillea exul, a Mn accumulator, were selected for these original experiments. The results presented here 20 months after plantation of young trees from a nursery show the interest of the approach. Mean Ni concentrations of up to 1513 mg kg(-1) are reported in G. pruinosa, as well as 2000 mg kg(-1) Mn in G. exul. Concentrations of Ni and Mn in the leaves of each species appear to be correlated with leaf age. Plantation of these species may also ensure mine reclamation, and experiments were conducted with the principles of ecological restoration in mind adding a further dimension to the approach.

Leaf-age and soil-plant relationships: key factors for reporting trace-elements hyperaccumulation by plants and design applications.

Relationships between the trace-elements (TE) content of plants and associated soil have been widely investigated especially to understand the ecology of TE hyperaccumulating species to develop applications using TE phytoextraction. Many studies have focused on the possibility of quantifying the soil TE fraction available to plants, and used bioconcentration (BC) as a measure of the plants ability to absorb TE. However, BC only offers a static view of the dynamic phenomenon of TE accumulation. Accumulation kinetics are required to fully account for TE distributions in plants. They are also crucial to design applications where maximum TE concentrations in plant leaves are needed. This paper provides a review of studies of BC (i.e. soil-plant relationships) and leaf-age in relation to TE hyperaccumulation. The paper focuses of Ni and Mn accumulators and hyperaccumulators from New Caledonia who were previously overlooked until recent Ecocatalysis applications emerged for such species. Updated data on Mn hyperaccumulators and accumulators from New Caledonia are also presented and advocate further investigation of the hyperaccumulation of this element. Results show that leaf-age should be considered in the design of sample collection and allowed the reclassification of Grevillea meisneri known previously as a Mn accumulator to a Mn hyperaccumulator.

Cryptic adaptive radiation in tropical forest trees in New Caledonia.

The causes of the species richness of tropical trees are poorly understood, in particular the roles of ecological factors such as soil composition. The nickel(Ni)-hyperaccumulating tree genus Geissois (Cunoniaceae) from the South-west Pacific was chosen as a model of diversification on different substrates. Here, we investigated the leaf element compositions, spatial distributions and phylogeny of all species of Geissois occurring on New Caledonia. We found that New Caledonian Geissois descended from a single colonist and diversified relatively quickly into 13 species. Species on ultramafic and nonultramafic substrates showed contrasting patterns of leaf element composition and range overlap. Those on nonultramafic substrates were largely sympatric but had distinct leaf element compositions. By contrast, species on ultramafic substrates showed similar leaf element composition, but occurred in many cases exclusively in allopatry. Further, earlier work showed that at least three out of these seven species use different molecules to bind Ni. Geissois qualifies as a cryptic adaptive radiation, and may be the first such example in a lineage of tropical forest trees. Variation in biochemical strategies for coping with both typical and adverse soil conditions may help to explain the diversification and coexistence of tropical forest trees on similar soil types.

Conservation priorities in a biodiversity hotspot: analysis of narrow endemic plant species in New Caledonia.

New Caledonia is a global biodiversity hotspot facing extreme environmental degradation. Given the urgent need for conservation prioritisation, we have made a first-pass quantitative assessment of the distribution of Narrow Endemic Species (NES) in the flora to identify species and sites that are potentially important for conservation action. We assessed the distributional status of all angiosperm and gymnosperm species using data from taxonomic descriptions and herbarium samples. We characterised species as being NES if they occurred in 3 or fewer locations. In total, 635 of the 2930 assessed species were classed as NES, of which only 150 have been subjected to the IUCN conservation assessment. As the distributional patterns of un-assessed species from one or two locations correspond well with assessed species which have been classified as Critically Endangered or Endangered respectively, we suggest that our distributional data can be used to prioritise species for IUCN assessment. We also used the distributional data to produce a map of "Hotspots of Plant Narrow Endemism" (HPNE). Combined, we used these data to evaluate the coincidence of NES with mining activities (a major source of threat on New Caledonia) and also areas of conservation protection. This is to identify species and locations in most urgent need of further conservation assessment and subsequent action. Finally, we grouped the NES based on the environments they occurred in and modelled the habitat distribution of these groups with a Maximum Entropy Species Distribution Model (MaxEnt). The NES were separable into three different groups based primarily on geological differences. The distribution of the habitat types for each group coincide partially with the HPNE described above and also indicates some areas which have high habitat suitability but few recorded NES. Some of these areas may represent under-sampled hotspots of narrow endemism and are priorities for further field work.

The metal hyperaccumulators from New Caledonia can broaden our understanding of nickel accumulation in plants.

While an excess of metals such as zinc, cadmium or nickel (Ni) is toxic for most plants, about 500 plant species called hyperaccumulators are able to accumulate high amounts of these metals. These plants and the underlying mechanisms are receiving an increasing interest because of their potential use in sustainable biotechnologies such as biofortification, phytoremediation, and phytomining. Among hyperaccumulators, about 400 species scattered in 40 families accumulate Ni. Despite this wide diversity, our current knowledge of the mechanisms involved in Ni accumulation is still limited and mostly restricted to temperate herbaceous Brassicaceae. New Caledonia is an archipelago of the tropical southwest pacific with a third of its surface (5500 km(2)) covered by Ni-rich soils originating from ultramafic rocks. The rich New Caledonia flora contains 2145 species adapted to these soils, among which 65 are Ni hyperaccumulators, including lianas, shrubs or trees, mostly belonging to the orders Celastrales, Oxalidales, Malpighiales, and Gentianales. We present here our current knowledge on Ni hyperaccumulators from New Caledonia and the latest molecular studies developed to better understand the mechanisms of Ni accumulation in these plants.

The chemical exploitation of nickel phytoextraction: an environmental, ecologic and economic opportunity for New Caledonia.

Herein, we explore the outlines of an innovative method based on the chemical recovery of metal-rich biomass produced in phytoextraction technologies. Taking advantage of the adaptive capacity of some New Caledonian plants to hyperaccumulate Ni(2+) cations in their aerial parts, this technique is based on the direct use of metals derived from plants as "Lewis acid" catalysts in organic chemistry. Metallic cations contained in New Caledonian nickel hyperaccumulators are recovered through a simple cost-effective process and serve the preparation of heterogeneous catalysts used in synthetic transformations allowing access to molecules with high added-value. The design of all processes is in line with the principles of green chemistry; it is adapted to the new economic constraints; it offers a new relevant outlet for metal-rich biomass; and it represents an alternative to non-renewable mineral materials.

Wood properties and trunk allometry of co-occurring rainforest canopy trees in a cyclone-prone environment.

PREMISE OF STUDY: New Caledonia commonly experiences cyclones, so trees there are expected to have enhanced wood traits and trunk allometry that confer resistance to wind damage. We ask whether there is evidence of a trade-off between these traits and growth rate among species. METHODS: Wood traits, including density, microfibril angle (MFA), and modulus of elasticity (MOE), ratio of tree height to stem diameter, and growth rate were investigated in mature trees of 15 co-occurring canopy species in a New Caledonian rainforest. KEY RESULTS: In contrast to some studies, wood density did not correlate negatively with growth increment. Among angiosperms, wood density and MOE correlated positively with diameter-adjusted tree height, and MOE correlated positively with stem-diameter growth increment. Tall slender trees achieved high stiffness with high efficiency with respect to wood density, in part by low MFA, and with a higher diameter growth increment but a lower buckling safety factor. However, some tree species of a similar niche differed in whole-tree resistance to wind damage and achieved wood stiffness in different ways. CONCLUSIONS: There was no evidence of a growth-safety trade-off in these trees. In forests that regularly experience cyclones, there may be stronger selection for high wood density and/or stiffness in fast-growing trees of the upper canopy, with the potential growth trade-off amortized by access to the upper canopy and by other plant traits. Furthermore, decreasing wood density does not necessarily decrease resistance to wind damage, resistance being influenced by other characteristics including cell-level traits (e.g., MFA) and whole-plant architecture.

Correlations between leaf toughness and phenolics among species in contrasting environments of Australia and New Caledonia.

BACKGROUND AND AIMS: Plants are likely to invest in multiple defences, given the variety of sources of biotic and abiotic damage to which they are exposed. However, little is known about syndromes of defence across plant species and how these differ in contrasting environments. Here an investigation is made into the association between carbon-based chemical and mechanical defences, predicting that species that invest heavily in mechanical defence of leaves will invest less in chemical defence. METHODS: A combination of published and unpublished data is used to test whether species with tougher leaves have lower concentrations of phenolics, using 125 species from four regions of Australia and the Pacific island of New Caledonia, in evergreen vegetation ranging from temperate shrubland and woodland to tropical shrubland and rainforest. Foliar toughness was measured as work-to-shear and specific work-to-shear (work-to-shear per unit leaf thickness). Phenolics were measured as 'total phenolics' and by protein precipitation (an estimate of tannin activity) per leaf dry mass. KEY RESULTS: Contrary to prediction, phenolic concentrations were not negatively correlated with either measure of leaf toughness when examined across all species, within regions or within any plant community. Instead, measures of toughness (particularly work-to-shear) and phenolics were often positively correlated in shrubland and rainforest (but not dry forest) in New Caledonia, with a similar trend suggested for shrubland in south-western Australia. The common feature of these sites was low concentrations of soil nutrients, with evidence of P limitation. CONCLUSIONS: Positive correlations between toughness and phenolics in vegetation on infertile soils suggest that additive investment in carbon-based mechanical and chemical defences is advantageous and cost-effective in these nutrient-deficient environments where carbohydrate may be in surplus.

Genetic diversity and geographical structure of the pitcher plant Nepenthes vieillardii in New Caledonia: A chloroplast DNA haplotype analysis.

Among the many species that grow in New Caledonia, the pitcher plant Nepenthes vieillardii (Nepenthaceae) has a high degree of morphological variation. In this study, we present the patterns of genetic differentiation of pitcher plant populations based on chloroplast DNA haplotype analysis using the sequences of five spacers. We analyzed 294 samples from 16 populations covering the entire range of the species, using 4660 bp of sequence. Our analysis identified 17 haplotypes, including one that is widely distributed across the islands, as well as regional and private haplotypes. The greatest haplotype diversity was detected on the eastern coast of the largest island and included several private haplotypes, while haplotype diversity was low in the southern plains region. The parsimony network analysis of the 17 haplotypes suggested that the genetic divergence is the result of long-term isolation of individual populations. Results from a spatial analysis of molecular variance and a cluster analysis suggest that the plants once covered the entire serpentine area of New Caledonia and that subsequent regional fragmentation resulted in the isolation of each population and significantly restricted seed flow. This isolation may have been an important factor in the development of the morphological and genetic variation among pitcher plants in New Caledonia.

Mass flowering and parental death in the regeneration of Cerberiopsis candelabra (Apocynaceae), a long-lived monocarpic tree in New Caledonia1.

Monocarpy is rare among forest trees, and the selective conditions allowing this life history to evolve are poorly known. Here we examined the regeneration dynamics of a New Caledonian monocarpic tree species, Cerberiopsis candelabra, to better understand how postreproductive mortality and mass flowering might contribute to the success of this strategy. We investigated population size structures and seedling establishment patterns in 18 stands following the 2003 flowering event. We found little evidence of recent recruitment. Instead, population size structures suggested a history of one or more substantial recruitment events followed by recruitment failure, despite multiple flowering events in most populations. The canopy gap created by death of an individual parent appeared generally insufficient to allow seedling establishment, but mass flowering and consequent death of many adults appeared to enhance seedling recruitment by opening more of the canopy. Site means of seedling density below dead parents correlated strongly with light availability and with the proportional density of flowering trees. Therefore, we suggest that mass flowering and gregariousness may have been necessary preconditions for the evolution of monocarpy in this species. However, the degree to which regeneration relies on synchronous flowering and gregariousness vs. infrequent catastrophic disturbances remains uncertain.

Nickel resistance determinants in bradyrhizobium strains from nodules of the endemic New Caledonia legume Serianthes calycina.

Bradyrhizobium strains, isolated in New Caledonia from nodules of the endemic legume Serianthes calycina growing in nickel-rich soils, were able to grow in the presence of 15 mM NiCl2. The genomes of these strains harbored two Ni resistance determinants, the cnr and nre operons. By constructing a cnrA mutant, we demonstrated that the cnr operon determines the high nickel resistance in Bradyrhizobium strains.

Age-size plasticity for reproduction in monocarpic plants.

Empirical and theoretical investigations of monocarpy have usually addressed the question of minimum or threshold sizes for reproduction. However, the range of flowering sizes observed in many monocarpic species is extraordinarily large (well beyond what can be called a "threshold"), and the sizes of flowering and nonflowering plants may overlap greatly. We attempt to explain these reproductive patterns in terms of optimal reaction norms predicted by simple deterministic life history models. We assume that individuals differ in their growth trajectories due to the heterogeneous quality of microsites and ask how the optimal age and size at flowering varies with environmental variation in growth and for different assumptions about fecundity and mortality. Under two very different growth functions (one with no age- or size-related decline in growth rate and another with such a decline as size approaches an asymptote), the optimal reaction norms imply considerable plasticity for size at reproduction, particularly when poor growth is associated with higher mortality or lower asymptotic size. Deterministic models such as these may be more applicable to long-lived than to short-lived monocarps, because fitness potential should be less affected by stochastic variability in yearly growing condition in the former than in the latter. We consider the case of a tropical monocarpic and masting tree species, Cerberiopsis candelabra (Apocynaceae), and show that our model results can account for wide ranges of reproductive size and overlap in size of flowering and nonflowering plants, in accord with observation. We suggest that empirical attention to norms of reaction across growth environments will be a more profitable approach than investigation of size thresholds per se.