Geopora Species Research Articles

<p><strong><em>Geopora ramila sp. nov.</em> (Pezizales, Pyronemataceae) evidenced by morphological characters and phylogenetic analyses in Iran</strong></p>

A new Geopora species (Pyronemataceae), Geopora ramila was described and illustrated from the soil, under or in the vicinity of Helianthemum ledifolium var. ledifolium annual plant in Fars province, Iran. Morphologically, G. ramila is similar to G. pinyonensis and G. arenicola but distinguished from both by a combination of morphological characters including, color and size of ascocarps, size and shape of ascospores, habit and associated host. The ribosomal DNA internally transcribed spacer (rDNA ITS) sequence of the new species (Acc. No. MT108930 to MT108934) showed 87.82% identity with G. pinyonensis in the BLAST search in GenBank. ITS-based phylogenetic analysis clearly supports G. ramila is a new and distinctive species lacking close relatives among described species of Geopora.

Symbiotic interactions above treeline of long‐lived pines: Mycorrhizal advantage of limber pine (Pinus flexilis) over Great Basin bristlecone pine (Pinus longaeva) at the seedling stage

Abstract In response to contemporary changes in climate, many tree species are shifting upslope to find favorable habitat. In the case of obligate ectomycorrhizal species, seedling growth above upper treeline depends on fungal spore availability. In the mountain ranges of the Great Basin, a recent shift in tree species stratification has been recorded, with limber pine (LP, Pinus flexilis) leapfrogging above the ancient bristlecone pine (BCP, Pinus longaeva) forest and establishing above current treeline. We compared the ability of LP and BCP to interact with soil spore banks collected at different microhabitats (next to dead trees, young live trees or in a treeless control) above current treeline in the White Mountains of California. We found an ectomycorrhizal fungal spore bank community composed of 15 species that was dominated by an undescribed and a hitherto unsequenced species of Geopora and Rhizopogon, respectively. This represents a much richer community than was found previously in this system. While both LP and BCP were able to establish ectomycorrhiza, LP was twice as likely to do so, and when comparing only seedlings that were colonized, its root system was colonized to a three‐fold greater extent. BCP seedlings grown on soils collected under young live trees were much more likely to be colonized compared to soils from the other two microhabitats. Synthesis. These differences in ectomycorrhizal receptivity might help to explain why LP is currently establishing at higher rates above the BCP treeline. Furthermore, it is possible that LP saplings above treeline can provide ectomycorrhizal facilitation for BCP seedlings, enabling the subsequent shift of BCP above treeline.

Geopora ahmadii sp. nov. from Pakistan

A new species, Geopora ahmadii, is described and illustrated based on material from Punjab, Pakistan. This species is characterized by sessile, cup- to saucer-shaped, partly immersed apothecia with whitish to grayish hymenial surfaces; broad ellipsoid, mostly uniguttulate ascospores; and brown excipular hairs. Phylogenetic analyses of the nrDNA ITS region with maximum parsimony, maximum likelihood, and Bayesian inference methods reveal that G. ahmadii is distinct from other described Geopora species. A collection previously identified as Geopora arenosa from Rawalakot, Pakistan, likely represents a second locality of G. ahmadii.

An elusive ectomycorrhizal fungus reveals itself: a new species of Geopora (Pyronemataceae) associated with Pinus edulis

Species of the genus Geopora are important ectomycorrhizal associates that can dominate the communities of some plant taxa, such as pinyon pine (Pinus edulis), a widespread tree of the western United States. Several members of the genus Geopora are known only from ectomycorrhizal root tips and thus have not been described formally. The sporocarps of some Geopora species occur infrequently because they depend on wet years for sporulation. In addition, Geopora sporocarps can be small and may be hypogeous at some developmental stage, limiting the opportunities for describing their morphology. Using molecular and morphological data, we have described a new species of fungus, Geopora pinyonensis, which produced ascocarps after unusually high precipitation at a northern Arizona site in summer 2012. Based on analysis of the ITS and nuLSU regions of the rDNA, G pinyonensis is a new species of Geopora. It has small sporocarps and ascospores relative to other members of the genus; however, these morphological features overlap with other species. Using rDNA data from sporocarps and ectomycorrhizal root tips, we show that the sporocarps correspond to an abundant species of ectomycorrhizal fungus associated with pinyon pines that is increasing in abundance in drought-affected landscapes and may promote drought tolerance.

Hoffmannoscypha, a novel genus of brightly coloured, cupulate Pyronemataceae closely related to Tricharina and Geopora

The rare apothecial, cupulate fungus Geopora pellita (Pyronemataceae) is characterized by a uniquely bright yellow-orange excipulum. We here re-examine its affiliations by use of morphological, molecular phylogenetic and ultrastructural analyses. G. pellita appears as phylogenetically rather isolated, being the sister group of a clade comprising Phaeangium, Picoa, the majority of the Tricharina species, and the remaining Geopora species. Based on its phylogenetic position and its unique combination of morphological characters, we assign G. pellita to Hoffmannoscypha, gen. nov., as H. pellita, comb. nov. As in a previous study, analyses of both large subunit (LSU) and internal transcribed spacer (ITS) ribosomal DNA suggest that the remaining genus Geopora is paraphyletic, with the hypogeous, ptychothecial type species more closely related to Picoa and Phaeangium than to the greyish-brownish cupulate and apothecial Geopora spp., indicating that the latter should be reassigned to the genus Sepultaria. The current study also shows that ITS confirm LSU data regarding the polyphyly of Tricharina.

Linking mycorrhizas to sporocarps: a new species, Geopora cercocarpi, on Cercocarpus ledifolius (Rosaceae)

Mycorrhizal assemblages characterized by molecular data frequently differ from collections of mycorrhizal sporocarps at the same site. Geopora species are frequent mycobionts of ectomycorrhizal roots, but except for G. cooperi they are rarely identified to species by molecular methods. Among the mycobionts of ectomycorrhizas with Cercocarpus ledifolius (Rosaceae) was a fungal species with a 91% BLAST match to G. arenicola. To determine the species of Geopora we surveyed for hypogeous sporocarps under C. ledifolius at sites in southern Oregon where the Geopora mycorrhizas had been collected and identified by DNA sequences of the ITS region. We found sporocarps of a Geopora species with 100% BLAST match to the mycorrhizas. Morphological characters of a white hymenium, inrolled entire margin and large spores, along with a hypogeous habit and a mycorrhizal host of C. ledifolius, distinguished these specimens from previously described species. Here we describe a new species, Geopora cercocarpi.

Genea mexicana, sp. nov., and Geopora tolucana, sp. nov., new hypogeous Pyronemataceae from Mexico, and the taxonomy of Geopora reevaluated

Two new species of hypogeous Pezizales in the Pyronemataceae are described from montane cloud forests in Mexico. Genea mexicana can be recognised by ITS sequence analysis and its distinct spore and seta dimensions; Geopora tolucana by ITS sequence analysis and its light brown hymenium, broadly ellipsoid ascospores and host preference. It belongs to the group of Geopora cooperi Harkn., which appears to be a rather heterogeneous assemblage, comprising a number of cryptic species. Our results indicate that the genus Geopora is non-monophyletic, occurring in two distinct phylogenetic clusters comprising species with either epigeous apothecial or hypogeous ptychothecial ascomata. Moreover, molecular divergence of Geopora is much higher than that of the neighbouring genera. Accordingly, we propose to reassign the cupulate apothecial Geopora species to the genus Sepultaria.

Ectomycorrhizas of Cercocarpus ledifolius (Rosaceae)

Woody species in the Rosaceae form ectomycorrhizal associations, but the fungal symbionts are unknown. The species of fungi determine whether host plants are isolated from other ectomycorrhizal species in the plant community or linked with other trees through mycorrhizal networks. In this study we identified the fungi that form ectomycorrhizas with Cercocarpus ledifolius (curl-leaf mountain mahogany). • Soil samples were collected under canopies of C. ledifolius. Ectomycorrhizas were described by morphology and by DNA sequences of the ITS region. Host species were confirmed by rbcL sequences. • Sixteen species of fungi were identified from ectomycorrhizas of Cercocarpus ledifolius. The ectomycorrhizal community was distinguished by the presence of a Geopora species situated in the G. arenicola clade and by the absence of Rhizopogon, suilloids, and Sebacinales. Of the species on C. ledifolius, two also occurred on trees of Quercus garryana var. breweri and four on Arctostaphylos sp. • The presence of fungal species in common with other ectomycorrhizal hosts shows that C. ledifolius, Q. garryana var. breweri, and Arctostaphylos species could be linked by a mycorrhizal network, allowing them to exchange nutrients or to share inoculum for seedling roots and new fine roots. Single-host fungi limited to C. ledifolius may improve resource acquisition and reduce competition with other ectomycorrhizal hosts. The finding of a Geopora species as a frequent mycobiont of C. ledifolius suggests that this fungus might be appropriate for inoculating seedlings for habitat restoration.

Four ectomycorrhizae of Pyronemataceae (Pezizomycetes) on Chinese Pine (Pinus tabulaeformis): morpho-anatomical and molecular-phylogenetic analyses

Morphological and anatomical characters of four ectomycorrhizae with affinities to the genera Humaria, Geopora, and Trichophaea of Pyronemataceae (Pezizomycetes, Ascomycota) on Chinese Pine (Pinus tabulaeformis) are described. The ectomycorrhizae are yellowish brown to brown, and have pseudoparenchymatous outer mantle layers and partially warty emanating hyphae with thick walls and without clamps. Intrahyphal hyphae are present, and no rhizomorphs are formed. The four ectomycorrhizae are distinguishable by differences in cell shape of outer mantle layers and the presence of cystidia. Ectomycorrhizae of a possible Humaria species (Pinirhiza humarioides) lack cystidia and have irregularly inflated cells on the outer mantle layer that are connected with thin septa. The two ectomycorrhizae showing probable affinities to Geopora species (“P. daqingensis” and “P. geoporoides”) possess row-like arranged cells in the outer mantle layer and cell heaps, and differ by the presence or absence of cystidia as well as by the structure of the inner mantle layers. Ectomycorrhizae likely having been formed by a Trichophaea species (“P. trichophaeoides”) have oval to polygonal cells and no cystidia. The possible taxa affiliations were assessed by molecular-phylogenetic analyses of the internal transcribed spacer (ITS) and partial large subunit (LSU) nrDNA. Morphological and anatomical characters are discussed against the background of the LSU phylogeny.

The Occurrence of Geopora arenosa in the British Isles

Summary. The occurrence of Geopora arenosa in the British Isles is confirmed by examination of two British collections from Scotland and comparison with type material of the species. The British record of this name is discussed and a revised key to the known British species of Geopora provided. Peziza arenosa Fuckel (the basionym of G. arenosa) is lectotypified to stabilise the concept of the species. Ascospore drawings from relevant collections are provided to facilitate recognition of British species of Geopora. Five species of Geopora Harkn. were recognised from Britain by Yao & Spooner (1996a) who also discussed all other names in the genus reported from Great Britain and Ireland. Whilst that paper was in press, a new collection was received at Kew which confirmed the presence in Britain of G. sepulta (Fr.) Korf. & Burds. This was reported by Yao & Spooner (1996b), with a key to the six British species then known. Based on examination of Fries's type material, the species concept of G. sepulta held by British authors (Cooke 1876; Phillips 1887; Massee 1895) was shown to be incorrect, and to represent a species with smaller spores (Yao & Spooner 1996a). Previous British records of G. sepulta were considered doubtful (Yao & Spooner 1996b) until the recent collection confirmed its presence here. Further collections of this species are now known from Britain. A similar situation applies to Geopora arenosa (Fuckel) S. Ahmad. Apparently, this was first recorded as British under the name Sepultaria arenosa (Fuckel) Boud. sensu Seaver by Dennis (1960); this was repeated in subsequent editions of his book (Dennis 1968, 1978). Cannon et al. (1985) list this taxon under Geopora without further reference, but probably on the basis of Dennis' work. It was not accepted as British by Yao & Spooner (1996a, b), because the species concept presented by Seaver (1928) and Dennis (1960, 1968, 1978) apparently differs from that represented by the type material. The spore size was given as 23.0 - 25.0 x 13.0 18.0 pm by Seaver (1928) and as 20.0 - 25.0 x 10.0 - 14.0 pm by Dennis (1960, 1968, 1978), considerably different from that of the type material (27.0 - 30.5 x 13.5 15.0 pm; Yao & Spooner 1996a). However, two recent collections from Scotland display similar spore size and fruiting season to those of the type and are here considered as evidence for the occurrence of G. arenosa in the British Isles. A description based on these collections is given below together with a revised key to the hitherto known British species of Geopora. The type material of Peziza arenosa Fuckel preserved in K has been examined and a lectotype is here selected for the

Notes on British species of Geopora

Five species of Geopora are recognized from Great Britain. Notes on the nomenclature and taxonomy of these species and of those others in the British checklist are presented and lectotypes for both Peziza sumneriana and P. lanuginosa var. sumneri are also designated.

Geopora sepulta (Pezizales) in Britain, with a Key to British Species of the Genus

Geopora sepulta is recognised as a British species based on a recent collection from Kent. This species is characterised by large, subterranean apothecia up to 45 mm diam. and large broadly ellipsoid spores (22.0 -)24.0 27.5(30.0) x 15.0 18.0(19.0) pm. The main characters taken from an isotype specimen and British material of G. sepulta are compared with those of the isotype of G. arenicola. A key to distinguish the six known British species of Geopora is also provided.

A Revision of the Genus Hydnocystis (Tuberales) and of the Hypogeous Species of Geopora (Pezizales)

Hydnocystis Tul. & Tul. is shown to be a monotypic genus containing only H. piligera Tul. & Tul. in the family Geneaceae of the Tuberales. Sepultaria (Cooke) Boud. (Pezizales) is shown to be congen...

A Revision of the Genus Hydnocystis (Tuberales) and of the Hypogeous Species of Geopora (Pezizales)

SUMMARYHydnocystis Tul. & Tul. is shown to be a monotypic genus containing only H. piligera Tul. & Tul. in the family Geneaceae of the Tuberales. Sepultaria (Cooke) Boud. (Pezizales) is shown to be congeneric with the older genus Geopora Harkness. Two new forms and a new subspecies, all hypogeous, are proposed for Geopora. Two species of Sepultaria and several in Hydnocystis are transferred to Geopora. Hydnocystis convolute McAlpine is transferred and renamed Peziza jactata Burdsall & Korf.