Efrén Cázares

Mycorrhiza-plant colonization patterns on a subalpine glacier forefront as a model system of primary succession

Lyman glacier in the North Cascades Mountains of Washington has a subalpine forefront characterized by a well-developed terminal moraine, inconspicuous successional moraines, fluting, and outwash. These deposits were depleted of symbiotic fungi when first exposed but colonized by them over time after exposure. Four major groups of plant species in this system are (1) mycorrhiza-independent or facultative mycotrophic, (2) dependent on arbuscular mycorrhizae (AM) (3) dependent on ericoid mycorrhiza (ERM) or ectomycorrhizae (EM), and (4) colonized by dark-septate (DS) endophytes. We hypothesized that availability of mycorrhizal propagules was related to the success of mycorrhiza-dependent plants in colonizing new substrates in naturally evolved ecosystems. To test this hypothesis roots samples of 66 plant species were examined for mycorrhizal colonization. The plants were sampled from communities at increasing distances from the glacier terminus to compare the newest communities with successively older ones. Long established, secondary successional dry meadow communities adjacent to the glacier forefront, and nearby high alpine communities were sampled for comparison. DS were common on most plant species on the forefront. Nonmycorrhizal plants predominated in the earlier successional sites, whereas the proportion of mycorrhizal plants generally increased with age of community. AM were present, mostly at low levels, and nearly absent in two sites of the forefront. ERM were present in all species of Ericaceae sampled, and EM in all species of Pinaceae and Salicaceae. Roots of plants in the long established meadow and heath communities adjacent to the forefront and the high alpine community all had one or another of the colonization types, with DS and AM predominating.

Historical biogeography and diversification of truffles in the Tuberaceae and their newly identified southern hemisphere sister lineage.

Truffles have evolved from epigeous (aboveground) ancestors in nearly every major lineage of fleshy fungi. Because accelerated rates of morphological evolution accompany the transition to the truffle form, closely related epigeous ancestors remain unknown for most truffle lineages. This is the case for the quintessential truffle genus Tuber, which includes species with socio-economic importance and esteemed culinary attributes. Ecologically, Tuber spp. form obligate mycorrhizal symbioses with diverse species of plant hosts including pines, oaks, poplars, orchids, and commercially important trees such as hazelnut and pecan. Unfortunately, limited geographic sampling and inconclusive phylogenetic relationships have obscured our understanding of their origin, biogeography, and diversification. To address this problem, we present a global sampling of Tuberaceae based on DNA sequence data from four loci for phylogenetic inference and molecular dating. Our well-resolved Tuberaceae phylogeny shows high levels of regional and continental endemism. We also identify a previously unknown epigeous member of the Tuberaceae – the South American cup-fungus Nothojafnea thaxteri (E.K. Cash) Gamundí. Phylogenetic resolution was further improved through the inclusion of a previously unrecognized Southern hemisphere sister group of the Tuberaceae. This morphologically diverse assemblage of species includes truffle (e.g. Gymnohydnotrya spp.) and non-truffle forms that are endemic to Australia and South America. Southern hemisphere taxa appear to have diverged more recently than the Northern hemisphere lineages. Our analysis of the Tuberaceae suggests that Tuber evolved from an epigeous ancestor. Molecular dating estimates Tuberaceae divergence in the late Jurassic (∼156 million years ago), with subsequent radiations in the Cretaceous and Paleogene. Intra-continental diversification, limited long-distance dispersal, and ecological adaptations help to explain patterns of truffle evolution and biodiversity.

Spore dispersal of ectomycorrhizal fungi on a glacier forefront by mammal mycophagy

Fecal pellets of small mammals (chipmunks, marmots and pikas), mountain goats and deer col? lected from the forefront of the rapidly receding Ly- man Glacier in the North Cascade Mountains of Wash? ington contained spores of both hypogeous and epigeous, ectomycorrhizal fungi. The forefront has been colonized by ectomycorrhizal hosts in the Pina- ceae and Salix spp. Animal mycophagy thus provides inoculum for diversifying the populations of mycor? rhizal fungi for early successional plants in the newly developing soils in this climatically stressful habitat.

Diversity, ecology, and conservation of truffle fungi in forests of the Pacific Northwest

Forests of the Pacific Northwest have been an epicenter for the evolution of truffle fungi with over 350 truffle species and 55 genera currently identified. Truffle fungi develop their reproductive fruit-bodies typically belowground, so they are harder to find and study than mushrooms that fruit aboveground. Nevertheless, over the last five decades, the Corvallis Forest Mycology program of the Pacific Northwest Research Station has amassed unprecedented knowledge on the diversity and ecology of truffles in the region. Truffle fungi form mycorrhizal symbioses that benefit the growth and survival of many tree and understory plants. Truffle fruit-bodies serve as a major food souce for many forest-dwelling mammals. A few truffle species are commercially harvested for gourmet consumption in regional restaurants. This publication explores the biology and ecology of truffle fungi in the Pacific Northwest, their importance in forest ecosystems, and effects of various silvicultural practices on sustaining truffle populations. General management principles and considerations to sustain this valuable fungal resource are provided.

Vesicular endophytes in roots of the Pinaceae

Vesicles and hyphae typical of vesicular-arbuscular mycorrhizae (VAM) were common in seedlings of Pseudotsuga menziesii, Abies lasiocarpa and Tsuga mertensiana growing in openings where herbaceous hosts of these fungi were common. Seedlings of A. lasiocarpa, Tsuga heterophylla, and T. mertensiana growing under closed forest canopies also had vesicles but at a much lower incidence than seedlings in the openings. The Pinaceae are generally assumed to be ectomycorrhizal, but Glomus-type colonizations occurred on the same seedlings as the ectomycorrhizae. The ecological significance of abundant VAM-type endophytes in otherwise ectomycorrhizal hosts deserves comprehensive study.

Vesicular mycorrhizal colonization of seedlings of Pinaceae and Betulaceae after spore inoculation with Glomus intraradices

Abstract Although Pinaceae and Betulaceae have been reported to contain Glomus–type root endophytes, its ecological importance and the conditions influencing this symbiosis are poorly understood. Seedlings of Abies lasiocarpa, Alnus rubra, Pinus contorta, Pinus ponderosa, Pseudotsuga menziesii, and Tsuga heterophylla were inoculated with Glomus intraradices to determine the vesicular-arbuscular mycorrhizae (VAM) development and responsiveness of these hosts. The role of companion VAM host plants on mycorrhizal colonization and nutrient uptake by Pseudotsuga menziesii was also examined by growing seedlings of Pseudotsuga menziesii in dual culture with VAM hosts Thuja plicata or Calamagrostis rubescens. After 8 weeks, no seedlings were colonized. At 16 weeks, 8 of 17 Thuja plicata seedlings grown with Pseudotsuga menziesii and all 18 inoculated Thuja plicata seedlings grown alone were colonized with vesicles and hyphae. Two of 17 inoculated Pseudotsuga menziesii seedlings grown in dual culture with Thuja plicata were colonized with abundant vesicles and hyphae. No ectomycorrhizal seedlings grown in monoculture were colonized. At 9 months, all 10 Calamagrostis rubescens and all 10 inoculated Pseudotsuga menziesii seedlings grown in dual culture were colonized by vesicles and hyphae. Two of 10 inoculated Pseudotsuga menziesiiand 1 of 10 inoculated Pinus ponderosa seedlings grown in monoculture were similarly colonized. The mean phosphorus content in the needles of colonized Pseudotsuga menziesii seedlings grown with Calamagrostis rubescens was about twice as high as in noncolonized Pseudotsuga menziesiiseedlings grown with Calamagrostis rubescens. Tissue nitrogen did not differ between these treatments. The results show that Glomus intraradices colonization of Pinaceae is most successful when a VAM host is present, although some vesicular colonization of Pinaceae occurred in the absence of a VAM host.

Ectomycorrhizal fungi associated with ponderosa pine and Douglas-fir: a comparison of species richness in native western North American forests and Patagonian plantations from Argentina

The putative ectomycorrhizal fungal species registered from sporocarps associated with ponderosa pine and Douglas-fir forests in their natural range distribution (i.e., western Canada, USA, and Mexico) and from plantations in south Argentina and other parts of the world are listed. One hundred and fifty seven taxa are reported for native ponderosa pine forests and 514 taxa for native Douglas-fir forests based on available literature and databases. A small group of genera comprises a high proportion of the species richness for native Douglas-fir (i.e., Cortinarius, Inocybe, and Russula), whereas in native ponderosa pine, the species richness is more evenly distributed among several genera. The comparison between ectomycorrhizal species richness associated with both trees in native forests and in Patagonia (Argentina) shows far fewer species in the latter, with 18 taxa for the ponderosa pine and 15 for the Douglas-fir. Epigeous species richness is clearly dominant in native Douglas-fir, whereas a more balanced relation epigeous/hypogeous richness is observed for native ponderosa pine; a similar trend was observed for Patagonian plantations. Most fungi in Patagonian Douglas-fir plantations have not been recorded in plantations elsewhere, except Suillus lakei and Thelephora terrestris, and only 56% of the fungal taxa recorded in Douglas-fir plantations around the world are known from native forests, the other taxa being new associations for this host, suggesting that new tree + ectomycorrhizal fungal taxa associations are favored in artificial situations as plantations.

Ectomycorrhizal fungi in Lyman Lake Basin: a comparison between primary and secondary successional sites

The results of eight years of study of the ectomycorrhizal macrofungi at the subalpine Lyman Lake Basin (Glacier Peak Wilderness area in the North Cascade Mountains, Washington, USA) are reported. The basin was divided into three sites: the primary successional glacier forefront vs two secondary successional habitats (subalpine parkland and old-growth montane Tsuga mertensiana-Abies amabilis). The 145 collections of ectomycorrhizal fungi obtained represented 68 species, 25 genera, 14 families and 7 orders. The Cortinariaceae was the most species-rich family with 25 species. Cortinarius was the most species-rich genus with 17 species. The two secondary successional sites shared 12.1% of the species; while the primary successional site shared only 2 and 5.1 %, respectively, with the two secondary successional sites. No ectomycorrhizal species occurred on all three sites. The secondary successional sites shared 7 species (Boletus edulis, Elaphomyces granulatus, Hydnotrya variiformis, Rhizopogon subsalmoneus, Rh. vulgaris, Russula silvicola and Thaxterogaster pingue) while the primary successional site shared two species with the parkland (Suillus aeruginascens and Su. cavipes) and only one with the montane site (Inocybe lacera). Fifty-eight species occurred at only one site. These data show that the communities of ectomycorrhizal fungi differ between the primary and secondary successional sites. We hypothesize that fungal life history strategies and habitat characteristics are more important determinants of succession of mycorrhizal fungi than host age or physiology as suggested by the relatively simple earlyand late-stage model.

Occurrence of vesicular-arbuscular mycorrhizae in Pseudotsuga menziesii and Tsuga heterophylla seedlings grown in Oregon Coast Range soils

Abstract Vesicular-arbuscular mycorrhizae (VAM) were common in seedlings of Pseudotsuga menziesii and Tsuga heterophylla grown in a greenhouse soil bioassay in soils collected from the Oregon Coast Range. Although root samples were heavily colonized by ectomycorrhizal fungi (EM), VAM colonization was observed in the cortical cells of both secondary and feeder roots. Vesicles, arbuscules, and hyphae typical of VAM occurred in 48% of 61 P. menziesii and 25% of 57 T. heterophylla seedlings. The ecological significance of VAM presence in the Pinaceae, as well as interactions among VAM, EM, and the plant host, deserve future investigation.

HYPOGEOUS FUNGI FROM NORTHERN MEXICO

Explorations in the mountains of Nuevo Le6n, Coahuila, Durango, and Tamaulipas from 1980 to 1985 yielded 24 species of hypogeous fungi, one being Rhizopogon guzmanii sp. nov. and 17 being first reports from Mexico. Twelve genera are represented: Glomus (Zygomycotina); Elaphomyces, Genea, Hydnobolites, Pachyphloeus, and Tuber (Ascomycotina); Gautieria, Hymenogaster, Leucogaster, Melanogaster, Octavianina, and Rhizopogon (Basidiomycotina).