Ultramafic Soils Research Articles

Mycorrhizal status of Cyperaceae from New Caledonian ultramafic soils: effects of phosphorus availability on arbuscular mycorrhizal colonization of Costularia comosa under field conditions

Plants from the Cyperaceae family (sedges), usually considered as non-mycorrhizal, constitute almost exclusively the herbaceous stratum of the ultramafic maquis in New Caledonia. These plants are pioneers and are important for the ecological restoration of mined areas. Costularia comosa, one of the most common sedges in this environment, was grown under field conditions on ultramafic soil, fertilized or not with phosphate and/or nitrogen. Results showed that the addition of phosphate to the soil induced a clear increase in mycorrhizal colonization of C. comosa and an increase in arbuscule abundance, reflecting the establishment of a functional mycorrhizal symbiosis. Significant positive correlations were found among mycorrhizal parameters and plant or soil phosphorus concentrations. Nitrogen fertilization did not affect mycorrhizal colonization of C. comosa. The improvement in mycorrhizal colonization by phosphate fertilization did not influence significantly nickel concentrations in the roots and shoots of plants. This study demonstrated that phosphate fertilization of ultramafic soil improved mycorrhizal colonization of C. comosa, with formation of a functional symbiosis under field conditions.

Arbuscular mycorrhizal fungi from New Caledonian ultramafic soils improve tolerance to nickel of endemic plant species

In order to improve knowledge about the role of arbuscular mycorrhizal fungi (AMF) in the tolerance to heavy metals in ultramafic soils, the present study investigated the influence of two Glomus etunicatum isolates from New Caledonian ultramafic maquis (shrubland), on nickel tolerance of a model plant species Sorghum vulgare, and of two ultramafic endemic plant species, Alphitonia neocaledonica and Cloezia artensis. In a first step, plants were grown in a greenhouse, on sand with defined concentrations of Ni, to appreciate the effects of the two isolates on the alleviation of Ni toxicity in controlled conditions. In a second step, the influence of the AMF on A. neocaledonica and C. artensis plants grown in a New Caledonian ultramafic soil rich in extractable nickel was investigated. Ni reduced mycorrhizal colonization and sporulation of the fungal isolates, but the symbionts increased plant growth and adaptation of endemic plant species to ultramafic conditions. One of the two G. etunicatum isolates showed a stronger positive effect on plant biomass and phosphorus uptake, and a greater reduction in toxicity symptoms and Ni concentration in roots and shoots. The symbionts seemed to act as a barrier to the absorption of Ni by the plant and reduced root-to-shoot Ni translocation. Results indicate the potential of selected native AMF isolates from ultramafic areas for ecological restoration of such degraded ecosystems.

A new endemic species of <i>Trigonachras</i> (Sapindaceae) from Sabah, Malaysia (Borneo)

A new species of Trigonachras from Borneo is described and illustrated. Trigonachras postardanjeisin is endemic to the ultramafic soils of the Tawai Forest Reserve in Sabah, Malaysia (Borneo). It is morphologically close to the widespread Trigonachras acuta but differs by its glabrous inflorescence and infrutescence axes and fruits. The two species also differ by their ecology and distribution. Line drawings are provided for the new taxa, along with discussions of its morphological affinities and preliminary risk of extinction assessment.

Rhizosphere bacteria ofCostulariaspp. from ultramafic soils in New Caledonia: diversity, tolerance to extreme edaphic conditions, and role in plant growth and mineral nutrition

Rhizosphere bacteria were isolated from Costularia spp., pioneer sedges from ultramafic soils in New Caledonia, which is a hotspot of biodiversity in the South Pacific. Genus identification, ability to tolerate edaphic constraints, and plant-growth-promoting (PGP) properties were analysed. We found that 10(5) colony-forming units per gram of root were dominated by Proteobacteria (69%) and comprised 21 genera, including Burkholderia (28%), Curtobacterium (15%), Bradyrhizobium (9%), Sphingomonas (8%), Rhizobium (7%), and Bacillus (5%). High proportions of bacteria tolerated many elements of the extreme edaphic conditions: 82% tolerated 100 μmol·L(-1) chromium, 70% 1 mmol·L(-1) nickel, 63% 10 mmol·L(-1) manganese, 24% 1 mmol·L(-1) cobalt, and 42% an unbalanced calcium/magnesium ratio (1/16). These strains also exhibited multiple PGP properties, including the ability to produce ammonia (65%), indole-3-acetic acid (60%), siderophores (52%), and 1-aminocyclopropane-1-carboxylate (ACC) deaminase (39%); as well as the capacity to solubilize phosphates (19%). The best-performing strains were inoculated with Sorghum sp. grown on ultramafic substrate. Three strains significantly enhanced the shoot biomass by up to 33%. The most successful strains influenced plant nutrition through the mobilization of metals in roots and a reduction of metal transfer to shoots. These results suggest a key role of these bacteria in plant growth, nutrition, and adaptation to the ultramafic constraints.

New Caledonia has a depauperate subterranean ant fauna, despite spectacular radiations above ground

Ants are often a target of tropical biodiversity assessment because of their ecological dominance and value as indicators of ecosystem health, but high microhabitat specificity, patchy distribution and cryptic habits of many species make effective sampling problematic. Although tropical ant faunas have long been known to show strong vertical stratification, only recently has it been recognised that this can include a high diversity of subterranean species that are poorly sampled using traditional methods. Global diversity patterns of subterranean ants and their responses to above-ground disturbance remain largely unknown. We describe ants collected in 360 subterranean traps distributed across 15 sites representing contrasting soil types (volcano-sedimentary and ultramafic) in New Caledonia, a recognised Global Biodiversity hotspot. New Caledonia has a diverse above-ground ant fauna that includes spectacular radiations of both Gondwanan and Indo-Malayan genera from all above-ground layers (litter, epigaeic and arboreal), and so it might be expected also to harbour a subterranean ant fauna of high biogeographic and conservation significance. We show that New Caledonia supports an extremely depauperate subterranean ant fauna, especially on ultramafic soils, with only two cryptobiotic species recorded in subterranean traps, and a trap success rate up to an order of magnitude lower than in comparable Australian studies. Our results show that there is an uncoupling of ant diversity above- and below- ground, such that a high diversity and conservation significance of ants above ground is not necessarily matched below.

Tracking nickel‐adaptive biomarkers in Pisolithus albus from New Caledonia using a transcriptomic approach

The fungus Pisolithus albus forms ectomycorrhizal (ECM) associations with plants growing on extreme ultramafic soils, which are naturally rich in heavy metals such as nickel. Both nickel-tolerant and nickel-sensitive isolates of P. albus are found in ultramafic soils in New Caledonia, a biodiversity hotspot in the Southwest Pacific. The aim of this work was to monitor the expression of genes involved in the specific molecular response to nickel in a nickel-tolerant P. albus isolate. We used pyrosequencing and quantitative polymerase chain reaction (qPCR) approaches to investigate and compare the transcriptomes of the nickel-tolerant isolate MD06-337 in the presence and absence of nickel. A total of 1,071,375 sequencing reads were assembled to infer expression patterns of 19,518 putative genes. Comparison of expression levels revealed that 30% of the identified genes were modulated by nickel treatment. The genes, for which expression was induced most markedly by nickel, encoded products that were putatively involved in a variety of biological functions, such as the modification of cellular components (53%), regulation of biological processes (27%) and molecular functions (20%). The 10 genes that pyrosequencing analysis indicated were induced the most by nickel were characterized further by qPCR analysis of both nickel-tolerant and nickel-sensitive P. albus isolates. Five of these genes were expressed exclusively in nickel-tolerant isolates as well as in ECM samples in situ, which identified them as potential biomarkers for nickel tolerance in this species. These results clearly suggest a positive transcriptomic response of the fungus to nickel-rich environments. The presence of both nickel-tolerant and nickel-sensitive fungal phenotypes in ultramafic soils might reflect environment-dependent phenotypic responses to variations in the effective concentrations of nickel in heterogeneous ultramafic habitats.

A new process for nickel ammonium disulfate production from ash of the hyperaccumulating plant Alyssum murale

The extraction of nickel (Ni) from ultramafic soils by phytomining can be achieved using Alyssum murale cultures. This study presents a new process for the valorization of Ni accumulated by this plant through the production of a Ni ammonium disulfate salt (Ni(NH4)2(SO4)2.6H2O). The process comprises an initial leaching of the ashes of A. murale with a sulphuric acid solution (1.9M H2SO4, T=95°C, t=240min, TS=150gashL−1), producing a leachate rich in Ni (10.2gNiL−1; 96% Ni solubilisation), Mg, P, K, Fe, Ca and Al. The pH of the acid leachate is increased to 5.0 with NaOH (5M), followed by an evaporation step which produced a purified solution rich in Ni (21.3gNiL−1) and an iron hydroxide precipitate. The cold crystallization (T=2°C, t=6h) of this solution by the stoichiometric addition (×1.2) of ammonium sulfate generates a Ni ammonium disulfate salt, containing 13.2% Ni, that is potentially valuable to industry.

Revision of Pichonia (Sapotaceae) in New Caledonia

Pichonia Pierre (Sapotaceae, Chrysophylloideae) is a small genus of 10 species in Australasia, of which seven are endemic to New Caledonia. We revise the genus for New Caledonia and describe one new species (P. grandiflora), resurrect another (P. dubia) and make two new combinations because of nomenclatural priority (P. balansae, P. deplanchei). P. balansae has been known for decades as P. calomeris, a name that in fact has never been validly published. The members are mainly found in Grande Terre, the main island of New Caledonia, but two species extend to Belep Islands in the north and to Isle of Pines in the south of the archipelago. Most grow in maquis vegetation at low altitudes, on ultramafic soils, preferably serpentinite. The exceptions are P. balansana, confined to limestone areas, and P. dubia that is a large canopy tree of the humid forest on ultramafic soil. Pichonia is distinguished by a character combination of an areolate higher venation, staminodes, a single-seeded fruit, plano-convex cotyledons and absence of endosperm. Because of being restricted to ultramafic soils, they are subsequently sensitive to present and future mining activities in New Caledonia. Hence, preliminary IUCN Red List assessments for all members are provided. P. balansana, P. daenikeri and P. lecomtei are all naturally uncommon, do not occur in any protected area and are proposed the IUCN status of Vulnerable. The herein described species P. grandiflora is rare in nature, known only from the Boulinda–Paéoua–Kopéto Mountains, and is in urgent need of protection. It is therefore assigned a preliminary status of Endangered.

Differential Regulation of Serine Acetyltransferase Is Involved in Nickel Hyperaccumulation in Thlaspi goesingense

When growing in its native habitat, Thlaspi goesingense can hyperaccumulate 1.2% of its shoot dry weight as nickel. We reported previously that both constitutively elevated activity of serine acetyltransferase (SAT) and concentration of glutathione (GSH) are involved in the ability of T. goesingense to tolerate nickel. A feature of SAT is its feedback inhibition by l-cysteine. To understand the role of this regulation of SAT by Cys on GSH-mediated nickel tolerance in T. goesingense, we characterized the enzymatic properties of SATs from T. goesingense. We demonstrate that all three isoforms of SAT in T. goesingense are insensitive to inhibition by Cys. Further, two amino acids (proline and alanine) in the C-terminal region of the cytosolic SAT (SAT-c) from T. goesingense are responsible for converting the enzyme from a Cys-sensitive to a Cys-insensitive form. Furthermore, the Cys-insensitive isoform of SAT-c confers elevated resistance to nickel when expressed in Escherichia coli and Arabidopsis thaliana, supporting a role for altered regulation of SAT by Cys in nickel tolerance in T. goesingense.

Further new endemic taxa of Cunoniaceae from New Caledonia

A description, illustration, distribution map and conservation assessment are given for each of three new species of Cunoniaceae from New Caledonia and plants endemic to the same localities from other genera are mentioned. Cunonia bopopensis Pillon & H. C. Hopkins is known from forest on ultramafic soils at high elevation on Mt Tchingou. This massif offers ecological conditions markedly different from those of its surroundings, which may explain the occurrence of several micro-endemics on this mountain. Some specimens of C. bopopensis were previously identified as C. bernieri Guillaumin but the type of the latter is probably a hybrid and this name is now considered doubtful. Geissois belema Pillon & H. C. Hopkins is endemic to Ile Art in the Belep archipelago, where it occurs in forest on ultramafic soils; it is the only species of Geissois known from Ile Art. Pancheria xaragurensis H. C. Hopkins & Pillon is found largely on the little-explored Forgotten Coast (“Cote Oubliee”) of south-eastern Grande Terre, generally in maquis along streams and in river beds at low elevation on ultramafic substrate. Morphological and ecological variation in the widespread taxa Geissois pruinosa Brongn. & Gris and Weinmannia dichotoma Brongn. & Gris are discussed and the combinations G. pruinosa var. intermedia (Vieill. ex Pamp.) H. C. Hopkins & Pillon and W. dichotoma var. monticola (Daniker) H. C. Hopkins & Pillon are published to accommodate some of this variability.

The effect of mycorrhiza on the growth and elemental composition of Ni-hyperaccumulating plant Berkheya coddii Roessler

The effect of arbuscular mycorrhizal fungi (AMF) on growth and element uptake by Ni-hyperaccumulating plant, Berkheya coddii, was studied. Plants were grown under laboratory conditions on ultramafic soil without or with the AM fungi of different origin. The AM colonization, especially with the indigenous strain, significantly enhanced plants growth and their survival. AMF affected also the elemental concentrations that were studied with Particle-induced X-ray emission (PIXE). AMF (i) increased K and Fe in shoots, Zn and Mn in roots, P and Ca both, in roots and shoots; (ii) decreased Mn in shoots, Co and Ni both, in shoots and roots. Due to higher biomass of mycorrhizal plants, total Ni content was up to 20 times higher in mycorrhizal plants compared to the non-mycorrhizal ones. The AMF enhancement of Ni uptake may therefore provide an improvement of a presently used technique of nickel phytomining.

A new species of Sartidia (Graminae), endemic to ultramafic soils

A species of Sartidia De Winter, first collected by P.J. Muller in 1972 in the Cythna Letty Nature Reserve in Mpumalanga, does not match existing material of Sartidia, a genus comprising four species. The new species, S. dewinteri J. Munday & L. Fish, is most similar to S. jucunda (Schweick.) De Winter but differs in the leaf sheath colour, lower leaf blade surface texture, spikelet length and upper glume length. It differs also from all other species in the genus in the shape and branching of the inflorescence, the relative length of the lateral awns to the median awn, lemma body surface texture, callus shape and hair arrangement, palea shape and distribution. Sartidia dewinteri differs anatomically from S. angolensis and S. vanderystii in that the stereome strands in the leaf blades project partly or almost completely into the first order vascular bundles rather than not at all, and from S. jucunda by having 3 rather than 5 first order vascular bundles in the leaf. Sartidia dewinteri is known only from ultramafic soils of the Barberton Greenstone Belt and is thus considered a serpentine endemic. The IUCN conservation status of S. dewinteri is considered to be Lower Risk — least concern, although some of its ultramafic habitat is under threat from afforestation and the after effects of mining.

New insights into the mycorrhizal status of Cyperaceae from ultramafic soils in New Caledonia

In New Caledonia, a hot spot of biodiversity, plants from the Cyperaceae family are mostly endemic and considered pioneers of the nickel-rich natural serpentine ecosystem. The aim of the study was to highlight the mycorrhizal status of these Cyperaceae and to bring new insights into the role of this symbiosis in plant tolerance to ultramafic soils. Nine Cyperaceae species were studied and presented evidence of root colonization by arbuscular mycorrhizas (AMs), with frequencies ranging from 8% to 57%. The highest level of AM colonization was observed in plants from the endemic dominant genus Costularia. Molecular evidence demonstrated the presence of Glomus sp. inside the roots. In a controlled greenhouse assay, AM inoculation of Costularia comosa grown under ultramafic conditions significantly enhanced plant growth, with an increase in biomass by up to 2.4-fold for shoots and 1.2-fold for roots, and also reduced nickel content in roots by 2.5-fold, as compared with the controls. All these data support our hypotheses (i) that a relationship exists between the mycorrhizal status of Cyperaceae and their habitat, and (ii) that AM have a positive role in plant tolerance to ultramafic soils (mineral nutrition and metal tolerance), suggesting the use of these pioneer plants with AM management as potential tools for nickel mine site rehabilitation in New Caledonia.

Drosera ultramafica (Droseraceae), a new sundew species of the ultramafic flora of the Malesian highlands

Drosera ultramafica, a new montane sundew species endemic to ultramafic soils of the Malesian highlands, is described and illustrated.

Howellanthus dalesianus, Recognition of a New Genus and Species in Tribe Phacelieae (Boraginaceae)

Howellanthus (Constance) Walden and R. Patt. is recognized as a new genus, containing a single species H. dalesianus (J. T. Howell) Walden and R. Patt., based on morphological evidence. Howellanthus dalesianus (Boraginaceae) is a small perennial herb, endemic to northern California and restricted to ultramafic soils. The species was first collected in 1936 by Ella Dales Cantelow and Herbert Clair Cantelow, described by John Thomas Howell as Phacelia dalesiana J. T. Howell, and later placed in the monotypic Phacelia subg. Howellanthus by Lincoln Constance.

Evolutionary lineages of nickel hyperaccumulation and systematics in European Alysseae (Brassicaceae): evidence from nrDNA sequence data.

Nickel (Ni) hyperaccumulation is a rare form of physiological specialization shared by a small number of angiosperms growing on ultramafic soils. The evolutionary patterns of this feature among European members of tribe Alysseae (Brassicaceae) are investigated using a phylogenetic approach to assess relationships among Ni hyperaccumulators at the genus, species and below-species level. Internal transcribed spacer (ITS) sequences were generated for multiple accessions of Alysseae. Phylogenetic trees were obtained for the genera of the tribe and Alyssum sect. Odontarrhena. All accessions and additional herbarium material were tested for Ni hyperaccumulation with the dimethylglyoxime colorimetric method. Molecular data strongly support the poorly known hyperaccumulator endemic Leptoplax (Peltaria) emarginata as sister to hyperaccumulator species of Bornmuellera within Alysseae. This is contrary to current assumptions of affinity between L. emarginata and the non-hyperaccumulator Peltaria in Thlaspideae. The lineage Bornmuellera-Leptoplax is, in turn, sister to the two non-hyperaccumulator Mediterranean endemics Ptilotrichum rupestre and P. cyclocarpum. Low ITS sequence variation was found within the monophyletic Alyssum sect. Odontarrhena and especially in A. murale sensu lato. Nickel hyperaccumulation was not monophyletic in any of three main clades retrieved, each consisting of hyperaccumulators and non-hyperaccumulators of different geographical origin. Nickel hyperaccumulation in Alysseae has a double origin, but it did not evolve in Thlaspideae. In Bornmuellera-Leptoplax it represents an early synapomorphy inherited from an ancestor shared with the calcicolous, sister clade of Mediterranean Ptilotrichum. In Alyssum sect. Odontarrhena it has multiple origins even within the three European clades recognized. Lack of geographical cohesion suggests that accumulation ability has been lost or gained over the different serpentine areas of south Europe through independent events of microevolutionary adaptation and selection. Genetic continuity and strong phenotypic plasticity in the A. murale complex call for a reduction of the number of Ni hyperaccumulator taxa formally recognized.

Ultramafic soils and species sorting in the flora of New Caledonia

Summary1. Island floras are generally characterized by disharmony due to uneven representations of plant groups. This has usually been explained by the unequal capacity of plants to disperse and colonize islands, but it has been suggested that the ecology of the recipient islands may also be important.2. We investigated the phylogenetic structure of the flora of New Caledonia, a biodiversity hotspot in the South‐West Pacific, to examine the phylogenetic pattern of a disharmonic flora and to test two competing hypotheses: disharmony due to uneven dispersability versus disharmony due to uneven ability to adapt to the ultramafic soils that are widespread on the archipelago and constrain plant growth.3. Dispersal did not seem to be a significant factor as plant families with low levels of endemism and thus high effective dispersal capacities tended to be under‐represented.4. We identified several clusters of over‐represented families, such as the COM clade (Celastrales, Oxalidales and Malpighiales), in which nickel hyperaccumulation (a syndrome associated with ultramafic soils) was particularly common. We indeed found a positive association between hyperaccumulation and over‐representation. Therefore, ultramafic soils may have been a major force for species sorting in the flora of New Caledonia.5. Because we think that most colonizers were naïve to ultramafic soils before they reached the archipelago, the over‐represented groups in New Caledonia may have possessed an exaptation to ultramafic soils, which has facilitated their settlement and diversification.6. Synthesis. This study supports that beyond dispersal, ecology is an important factor in determining the success of different plant groups to colonize an island. In the case of New Caledonia, exaptation to ultramafic soils is likely a primary cause of ecological filtering.

A new species of Polygala (Polygalaceae) from ultramafic soils in Sekhukhuneland, South Africa, with notes on its ecology

Polygala sekhukhuniensis Retief, Siebert & A.E.Van Wyk (Polygala; section Polygala; subsection Heterolophus), a new species with a restricted range in Sekhukhuneland, South Africa, is described, illustrated and compared with other members of the genus. It is a dwarf shrub that can be distinguished by its much-branched habit, sparsely flowered inflorescences, pink alae with darker pink veins, brown to black seed testa, and oblate pollen grains with pronounced opercula. Geographically, P. sekhukhuniensis is confined to heavily eroded localized sites, a natural geomorphological feature of some of the highly water-dispersible soils derived from ultramafic rocks in the valleys of the Steelpoort River and its tributaries in the Sekhukhuneland Centre of Plant Endemism. P. sekhukhuniensis is a calciotrophic excluder of heavy metals that accumulates Ca in its leaves. It is ecologically compared with co-occurring species of Polygala on ultramafic-derived soil.

Ultramafic soils from New Caledonia structurePisolithus albusin ecotype

Isolates of ectomycorrhizal Pisolithus albus were sampled from both ultramafic and volcano-sedimentary soils in New Caledonia, a tropical hotspot of biodiversity, to investigate the relationships between genetic diversity and edaphic constraint through tolerance to nickel (Ni). Carpophore description, spore morphology and phylogenetic analysis based on internal transcribed spacer (ITS) rDNA sequences confirmed that all isolates belong to P. albus and are closely related to other Australasian specimens. Using molecular tools, ITS-restriction fragment length polymorphism and amplified fragment length polymorphism markers, we showed the existence of two distinct genetic clusters within P. albus: ultramafic and volcano-sedimentary. Mycelia response to Ni toxicity supports such a population structure. Pisolithus albus from ultramafic soils included isolates with a high diversity of in vitro Ni tolerance, with both Ni-tolerant isolates (average Ni EC(50) at 575 microM) and Ni-sensitive isolates (average Ni EC(50) at 37 microM). In contrast, all isolates from volcano-sedimentary soils were found to be Ni sensitive (average Ni EC(50) at 32 microM). We highlight that (1) P. albus population from ultramafic soils of New Caledonia are genetically structured in ecotype, and that (2) Ni tolerance among ultramafic isolates suggests an adaptive physiological response to Ni toxicity.

A revision of MalesianAustrobuxus(Picrodendraceae/Euphorbiaceae s.l. subfam. Oldfieldioideae)

The Malesian species of the predominantly New Caledonian genus Austrobuxus were revised. Three species are recognized. Austrobuxus nitidus is widespread from south Thailand to Borneo. Characteristic for this species are the very short papillae of the caruncle. The caruncle papillae are much longer in the other two, newly described species. Austrobuxus celebicus sp. nov. is endemic to Sulawesi, where it is found on ultramafic soils, and A. dentatus sp. nov. is an endemic species of western New Guinea. The former is notable for the very small fruits and the latter for the teeth-like glands along the leaf blade margins, unique within the genus.