Resolved phylogeny and biogeography of the root pathogen Armillaria and its gasteroid relative, Guyanagaster.

BACKGROUND: Armillaria is a globally distributed mushroom-forming genus composed primarily of plant pathogens. Species in this genus are prolific producers of rhizomorphs, or vegetative structures, which, when found, are often associated with infection. Because of their importance as plant pathogens, understanding the evolutionary origins of this genus and how it gained a worldwide distribution is of interest. The first gasteroid fungus with close affinities to Armillaria-Guyanagaster necrorhizus-was described from the Neotropical rainforests of Guyana. In this study, we conducted phylogenetic analyses to fully resolve the relationship of G. necrorhizus with Armillaria. Data sets containing Guyanagaster from two collecting localities, along with a global sampling of 21 Armillaria species-including newly collected specimens from Guyana and Africa-at six loci (28S, EF1α, RPB2, TUB, actin-1 and gpd) were used. Three loci-28S, EF1α and RPB2-were analyzed in a partitioned nucleotide data set to infer divergence dates and ancestral range estimations for well-supported, monophyletic lineages. RESULTS: The six-locus phylogenetic analysis resolves Guyanagaster as the earliest diverging lineage in the armillarioid clade. The next lineage to diverge is that composed of species in Armillaria subgenus Desarmillaria. This subgenus is elevated to genus level to accommodate the exannulate mushroom-forming armillarioid species. The final lineage to diverge is that composed of annulate mushroom-forming armillarioid species, in what is now Armillaria sensu stricto. The molecular clock analysis and ancestral range estimation suggest the most recent common ancestor to the armillarioid lineage arose 51 million years ago in Eurasia. A new species, Guyanagaster lucianii sp. nov. from Guyana, is described. CONCLUSIONS: The armillarioid lineage evolved in Eurasia during the height of tropical rainforest expansion about 51 million years ago, a time marked by a warm and wet global climate. Species of Guyanagaster and Desarmillaria represent extant taxa of these early diverging lineages. Desarmillaria represents an armillarioid lineage that was likely much more widespread in the past. Guyanagaster likely evolved from a gilled mushroom ancestor and could represent a highly specialized endemic in the Guiana Shield. Armillaria species represent those that evolved after the shift in climate from warm and tropical to cool and arid during the late Eocene. No species in either Desarmillaria or Guyanagaster are known to produce melanized rhizomorphs in nature, whereas almost all Armillaria species are known to produce them. The production of rhizomorphs is an adaptation to harsh environments, and could be a driver of diversification in Armillaria by conferring a competitive advantage to the species that produce them.

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.

New species of Auritella (Inocybaceae) from Cameroon, with a worldwide key to the known species.

Two new species in the genus Auritella (Inocybaceae) are described as new from tropical rainforest in Cameroon. Descriptions, photographs, line drawings, and a worldwide taxonomic key to the described species of Auritella are presented. Phylogenetic analysis of 28S rDNA and rpb2 nucleotide sequence data suggests at least five phylogenetic species that can be ascribed to Auritella occur in the region comprising Cameroon and Gabon and constitute a strongly supported monophyletic subgroup within the genus. Phylogenetic analysis of ITS data supports the conspecificity of numerous collections attributed to the two new species as well as the monophyly of Australian species of Auritella. This work raises the known number of described species of Auritella to thirteen worldwide, four of which occur in tropical Africa, one in tropical India, and eight in temperate and tropical regions of Australia. This is the first study to confirm an ectomycorrhizal status of Auritella using molecular data.

New species of Elaphomyces (Elaphomycetaceae, Eurotiales, Ascomycota) from tropical rainforests of Cameroon and Guyana.

The sequestrate false truffles Elaphomyces favosus, E. iuppitercellus, and E. labyrinthinus spp. nov. are described as new to science from the Dja Biosphere Reserve, Cameroon. Elaphomyces adamizans sp. nov. is described as new from the Pakaraima Mountains of Guyana. The Cameroonian species are the first Elaphomyces taxa to be formally described from Africa, occurring in lowland Guineo-Congolian tropical rainforests dominated by the ectomycorrhizal (ECM) canopy tree Gilbertiodendron dewevrei (Fabaceae subfam. Caesalpinioideae). The Guyanese species is the third to be discovered in lowland tropical South America, occurring in forests dominated by the ECM trees Pakaraimaea dipterocarpacea (Dipterocarpaceae) and Dicymbe jenmanii (Fabaceae subfam. Caesalpinioideae). Macromorphological, micromorphological, habitat, and DNA sequence data are provided for each new species. Molecular and morphological data place these fungi in Elaphomycetaceae (Eurotiales, Ascomycota). Unique morphological features are congruent with molecular delimitation of each of the new species based on a phylogenetic analysis of the rDNA ITS and 28S loci across the Elaphomycetaceae. The phylogenetic analysis also suggests that a common ancestor is shared between some Elaphomyces species from Africa and South America, and that species of the stalked, volvate genus Pseudotulostoma may be nested in Elaphomyces.

Kombocles bakaiana gen. sp. nov. (Boletaceae), a new sequestrate fungus from Cameroon.

Kombocles bakaiana gen. sp. nov. is described as new to science. This sequestrate, partially hypogeous fungus was collected around and within the stilt root system of an ectomycorrhizal (ECM) tree of the genus Uapaca (Phyllanthaceae) in a Guineo-Congolian mixed tropical rainforest in Cameroon. Molecular data place this fungus in Boletaceae (Boletales, Agaricomycetes, Basidiomycota) with no clear relationship to previously described taxa within the family. Macro- and micromorphological characters, habitat, and DNA sequence data are provided. Unique morphological features and a molecular phylogenetic analysis of 304 sequences across the Boletales justify the recognition of the new taxa. Kombocles bakaiana is the fourth sequestrate Boletaceae described from the greater African tropics, and the first to be described from Cameroon.

Mixed-forest species establishment in a monodominant forest in central Africa: implications for tropical forest invasibility.

BACKGROUND: Traits of non-dominant mixed-forest tree species and their synergies for successful co-occurrence in monodominant Gilbertiodendron dewevrei forest have not yet been investigated. Here we compared the tree species diversity of the monodominant forest with its adjacent mixed forest and then determined which fitness proxies and life history traits of the mixed-forest tree species were most associated with successful co-existence in the monodominant forest. METHODOLOGY/PRINCIPAL FINDINGS: We sampled all trees (diameter in breast height [dbh]≥10 cm) within 6×1 ha topographically homogenous areas of intact central African forest in SE Cameroon, three independent patches of G. dewevrei-dominated forest and three adjacent areas (450-800 m apart). Monodominant G. dewevrei forest had lower sample-controlled species richness, species density and population density than its adjacent mixed forest in terms of stems with dbh≥10 cm. Analysis of a suite of population-level characteristics, such as relative abundance and geographical distribution, and traits such as wood density, height, diameter at breast height, fruit/seed dispersal mechanism and light requirement-revealed after controlling for phylogeny, species that co-occur with G. dewevrei tend to have higher abundance in adjacent mixed forest, higher wood density and a lower light requirement. CONCLUSIONS/SIGNIFICANCE: Our results suggest that certain traits (wood density and light requirement) and population-level characteristics (relative abundance) may increase the invasibility of a tree species into a tropical closed-canopy system. Such knowledge may assist in the pre-emptive identification of invasive tree species.

Does lowering glutamine synthetase activity in nodules modify nitrogen metabolism and growth of Lotus japonicus?

A cDNA encoding cytosolic glutamine synthetase (GS) from Lotus japonicus was fused in the antisense orientation relative to the nodule-specific LBC3 promoter of soybean (Glycine max) and introduced into L. japonicus via transformation with Agrobacterium tumefaciens. Among the 12 independent transformed lines into which the construct was introduced, some of them showed diminished levels of GS1 mRNA and lower levels of GS activity. Three of these lines were selected and their T(1) progeny was further analyzed both for plant biomass production and carbon and nitrogen (N) metabolites content under symbiotic N-fixing conditions. Analysis of these plants revealed an increase in fresh weight in nodules, roots and shoots. The reduction in GS activity was found to correlate with an increase in amino acid content of the nodules, which was primarily due to an increase in asparagine content. Thus, this study supports the hypothesis that when GS becomes limiting, other enzymes (e.g. asparagine synthetase) that have the capacity to assimilate ammonium may be important in controlling the flux of reduced N in temperate legumes such as L. japonicus. Whether these alternative metabolic pathways are important in the control of plant biomass production still remains to be fully elucidated.