Plant-plant interactions vary with different mycorrhizal fungus species

Because different species of mycorrhizal fungi have different effects on the growth of particular plant species, variation in mycorrhizal fungus species composition could cause changes in the strength of plant-plant interactions. Results are presented from a growth chamber experiment that compared the strength of interactions among seedlings of ponderosa pine (Pinus ponderosa) when the pines were colonized by two different groups of ectomycorrhizal fungi in the genus Rhizopogon. Plant density effects differed between the two groups of mycorrhizal fungi: plant growth was low regardless of density when plants were colonized with pine-specific Rhizopogon species, while plant growth declined with plant density when plants were colonized by Rhizopogon species having a broader host range. This result parallels results from previous studies showing that plant interactions are more antagonistic with mycorrhizal fungi than without, implying that plant responsiveness to beneficial mycorrhizal fungi declines with increasing plant density. If such effects are prevalent in plant communities, then variation in mycorrhizal fungus community composition is predicted to have a density-dependent effect on plants.     

Below-ground interactions with arbuscular mycorrhizal shrubs decrease the performance of pinyon pine and the abundance of its ectomycorrhizas

Few studies have examined how below-ground interactions among plants affect the abundance and community composition of symbiotic mycorrhizal fungi. Here, we combined observations during drought with a removal experiment to examine the effects of below-ground interactions with arbuscular mycorrhizal (AM) shrubs on the growth of pinyon pines (Pinus edulis), and the abundance and community composition of their ectomycorrhizal (EM) fungi. Shrub density was negatively correlated with pinyon above- and below-ground growth and explained 75% of the variation in EM colonization. Consistent with competitive release, pinyon fine-root biomass, shoot length and needle length increased with shrub removal. EM colonization also doubled following shrub removal. EM communities did not respond to shrub removal, perhaps because of their strikingly low diversity. These results suggest that below-ground competition with AM shrubs negatively impacted both pinyons and EM fungi. Similar competitive effects may be observed in other ecosystems given that drought frequency and severity are predicted to increase for many land interiors.     

Variation in woody plant mortality and dieback from severe drought among soils, plant groups, and species within a northern Arizona ecotone

Vegetation change from drought-induced mortality can alter ecosystem community structure, biodiversity, and services. Although drought-induced mortality of woody plants has increased globally with recent warming, influences of soil type, tree and shrub groups, and species are poorly understood. Following the severe 2002 drought in northern Arizona, we surveyed woody plant mortality and canopy dieback of live trees and shrubs at the forest–woodland ecotone on soils derived from three soil parent materials (cinder, flow basalt, sedimentary) that differed in texture and rockiness. Our first of three major findings was that soil parent material had little effect on mortality of both trees and shrubs, yet canopy dieback of trees was influenced by parent material; dieback was highest on the cinder for pinyon pine (Pinus edulis) and one-seed juniper (Juniperus monosperma). Ponderosa pine (Pinus ponderosa) dieback was not sensitive to parent material. Second, shrubs had similar mortality, but greater canopy dieback, than trees. Third, pinyon and ponderosa pines had greater mortality than juniper, yet juniper had greater dieback, reflecting different hydraulic characteristics among these tree species. Our results show that impacts of severe drought on woody plants differed among tree species and tree and shrub groups, and such impacts were widespread over different soils in the southwestern U.S. Increasing frequency of severe drought with climate warming will likely cause similar mortality to trees and shrubs over major soil types at the forest–woodland ecotone in this region, but due to greater mortality of other tree species, tree cover will shift from a mixture of species to dominance by junipers and shrubs. Surviving junipers and shrubs will also likely have diminished leaf area due to canopy dieback.     

Effectiveness of bifenthrin (Onyx) and carbaryl (Sevin SL) for protecting individual, high-value conifers from bark beetle attack (Coleoptera: Curculionidae: Scolytinae) in the Western United States

High-value trees, such as those located in residential, recreational, or administrative sites, are particularly susceptible to bark beetle (Coleoptera: Curculionidae: Scolytinae) attack as a result of increased amounts of stress associated with drought, soil compaction, mechanical injury, or vandalism. Tree losses in these unique environments generally have a substantial impact. The value of these individual trees, cost of removal, and loss of esthetics may justify protection until the main thrust of a bark beetle infestation subsides. This situation emphasizes the need for ensuring that effective insecticides are available for individual tree protection. In this study, we assess the efficacy of bifenthrin (Onyx) and carbaryl (Sevin SL) for protecting: ponderosa pine, Pinus ponderosa Dougl. ex. Laws., from western pine beetle, Dendroctonus brevicomis LeConte, in California; mountain pine beetle, Dendroctonus ponderosae Hopkins in South Dakota; and Ips spp. in Arizona; lodgepole pine, Pinus contorta Dougl. ex Loud., from D. ponderosae in Montana; pinyon, Pinus edulis Engelm. in Colorado and Pinus monophylla Torr. and Frem. in Nevada from pinyon ips, Ips confusus (LeConte); and Engelmann spruce, Picea engelmannii Parry ex. Engelm. from spruce beetle, Dendroctonus rufipennis (Kirby) in Utah. Few trees were attacked by Ips spp. in Arizona and that study was discontinued. Sevin SL (2.0%) was effective for protecting P. ponderosa, P. contorta, and P. monophylla for two field seasons. Estimates of efficacy could not be made during the second field season in P. edulis and P. engelmannii due to insufficient mortality in untreated, baited control trees. Two field seasons of efficacy was demonstrated in P. ponderosa/D. brevicomis and P. monophylla for 0.06% Onyx. We conclude that Onyx is an effective individual tree protection tool, but repeated annual applications may be required in some systems if multiyear control is desired.     

The effect of different pine hosts on the sampling of Rhizopogon spore banks in five Eastern Sierra Nevada forests

Our primary goal was to determine whether detection of Rhizopogon internal transcribed spacer (ITS) groups is affected by the pine species used in seedling bioassays. Our secondary goal was to investigate composition of Rhizopogon spore banks in the Eastern Sierra Nevada of California, a previously unsampled region. We used seedlings of Pinus contorta, Pinus jeffreyi, Pinus lambertiana, and Pinus muricata as bioassay plants and identified the Rhizopogon retrieved by internal transcribed spacer (ITS) sequence analysis. We found that each of the pine species retrieved all of the abundant Rhizopogon ITS groups, but there were significant differences among pines in the richness of Rhizopogon ITS groups recovered. Pinus muricata recovered all ITS groups found in this study and was significantly better than P. lambertiana. Rhizopogon communities from the five sampled sites contained six to eight ITS groups per site, with two unique sequence groups and a higher abundance of the Rhizopogon ellenae and Rhizopogon arctostaphyli groups than at previously sampled sites. These results show high cross-receptivity between Rhizopogon and pine species, and regional patterns in spore bank composition.     

Pine invasion impacts on plant diversity in Patagonia: invader size and invaded habitat matter

Conifers, which are widely planted as fast growing tree crops, are invading forested and treeless environments across the globe, causing important changes in biodiversity. However, how small-scale impacts on plant diversity differ according to pine size and habitat context remains unclear. We assessed the effects of different stages of pine invasion on plant communities in forest and steppe sites located in southern Chile. In each site, we sampled plant diversity under and outside the canopy of Pinus contorta individuals, using paired plots. We assessed the relative impact of pine invasion on plant species richness and cover. In both sites, richness and cover beneath pine canopy decreased with increasing pine size (i.e. height and canopy area). A significant negative impact of pines on species richness and plant cover was detected for pines over 4 m in height. The impact of pines on plant richness and cover depended on pine size (i.e. canopy area) and habitat type. Larger pines had more negative impacts than smaller pines in both sites, with a greater impact for a given pine size in the Patagonian steppe compared to the A. araucaria forest. Species composition changed between under and outside canopy plots when pines were 4 m or taller. Pine presence reduced cover of most species. The impacts of pine invasions are becoming evident in forested and treeless ecosystems of southern Chile. Our results suggest that the magnitude of pine invasion impacts could be related to how adapted the invaded community is to tree cover, with the treeless environment more impacted by the invasion.     

Suitability of some southern and western pines as hosts for the pine shoot beetle, Tomicus piniperda (Coleoptera: Scolytidae)

The pine shoot beetle, Tomicus piniperda (L.), is an exotic pest that has become established in North America. Discovered in Ohio in 1992, it has since been found in at least 13 states and parts of Canada. The beetle can cause significant growth loss in pines, and it represents a potential threat to trees in areas where it has not yet become established. To evaluate this threat to native pines, field and laboratory tests were conducted on several common and important southern and western species to determine whether they are acceptable hosts for T. piniperda. Comparisons with Pinus sylvestris L., Scots pine, a preferred natural host for the beetle, were made where possible. Measurements of beetle attack success on southern pine billets showed that Pinus taeda L., Pinus echinata Miller, Pinus elliottii var. elliottii Engelmann, Pinus palustris Miller, and Pinus virginiana Miller (loblolly, shortleaf, slash, longleaf, and Virginia pine, respectively) and two western pines, Pinus ponderosa Lawson and Pinus contorta Douglas (ponderosa and lodgepole pine, respectively), were acceptable for breeding material, but brood production was highly variable. Among the southern pines, P. taeda and P. echinata were susceptible to shoot feeding by T. piniperda, whereas P. elliottii was highly resistant and P. palustris seemed to be virtually immune. Shoot feeding tests on the western pines were conducted only in the laboratory, but there was moderate-to-good survival of adults feeding on both species. It seems that if T. piniperda is introduced into the south and west it will likely establish and may cause some damage to native pines. P. taeda may be affected more than other southern pines because it is the most abundant species, it is readily attacked for brood production, which can result in moderately large broods, and the beetle survives well during maturation feeding on P. taeda shoots.     

Identities and distributions of the co-invading ectomycorrhizal fungal symbionts of exotic pines in the Hawaiian Islands

Pine species have become invasive throughout the globe and threaten to replace native biota. The threat of pine invasion is particularly pressing in parts of the tropics where there are no native pines. The factors that govern pine invasion are not often well understood. However, key to pine survival is an obligate and mutualistic interaction with ectomycorrhizal fungi. Thus for pines to successfully invade new habitats compatible ectomycorrhizal fungi must already be present, or be co-introduced. The purpose of this study was to examine the community structure of non-native ectomycorrhizal fungi associated with pine invasions in the Hawaiian Islands. To accomplish this we executed a field and greenhouse study and used a molecular ecology approach to identify the fungi associating with invasive pines in Hawai‘i. We show that: (1) ectomycorrhizal fungal species richness in non-native pine plantations is far less than what is found in pine’s native range, (2) there was a significant decrease in average ectomycorrhizal fungal species richness as distance from pine plantations increased and, (3) Suillus species were the dominant fungi colonizing pines outside plantations. The keystone ectomycorrhizal fungal taxa responsible for pine establishment in Hawai‘i are within genera commonly associated with pine invasions throughout the globe. We surmise that these fungi share functional traits such as the ability for long-distance dispersal from plantations and host tree colonization via spore that lead to their success when introduced to new habitats.     

Neighboring trees regulate the root‐associated pathogenic fungi on the host plant in a subtropical forest

Root‐associated fungi and host‐specific pathogens are major determinants of species coexistence in forests. Phylogenetically related neighboring trees can strongly affect the fungal community structure of the host plant, which, in turn, will affect the ecological processes. Unfortunately, our understanding of the factors influencing fungal community composition in forests is still limited. In particular, investigation of the relationship between the phytopathogenic fungal community and neighboring trees is incomplete. In the current study, we tested the host specificity of members of the root‐associated fungal community collected from seven tree species and determined the influence of neighboring trees and habitat variation on the composition of the phytopathogenic fungal community of the focal plant in a subtropical evergreen forest. Using high‐throughput sequencing data with respect to the internal transcribed spacer (ITS) region, we characterized the community composition of the root‐associated fungi and found significant differences with respect to fungal groups among the seven tree species. The density of conspecific neighboring trees had a significantly positive influence on the relative abundance of phytopathogens, especially host‐specific pathogens, while the heterospecific neighbor density had a significant negative impact on the species richness of host‐specific pathogens, as well as phytopathogens. Our work provides evidence that the root‐associated phytopathogenic fungi of a host plant depend greatly on the tree neighbors of the host plant.     

Landbird community composition varies among seasons in a heterogeneous ponderosa pine forest

ABSTRACT.   There is growing recognition of the need to conserve areas used by birds during migration, including forest and upland habitats. Because extensive thinning and burning treatments are planned for ponderosa pine ( Pinus ponderosa ) forests in the southwestern United States, information on the use of these forests by landbirds during migration is needed for conservation planning. We compared species richness among spring, breeding, and fall seasons at 69 points in a ponderosa pine forest to assess changes in landbird communities and the role of different ponderosa pine cover types in habitat selection among seasons. We detected a total of 64 bird species. Bird community similarity was lowest between the breeding and fall seasons and highest between the spring and breeding seasons. Twenty percent of the species detected were present exclusively in the fall and, of these, over half were Neotropical migrants. Only two species (3%) were detected exclusively during the spring. Although we found little difference in bird species similarity among vegetative cover types during the breeding season, forests that contained a deciduous component exhibited higher bird species similarity with each other than with habitats that did not include a deciduous component in spring and fall. In addition, foliage foragers dominated the community in spring and fall, and all Neotropical migrants detected exclusively in fall were found in ponderosa pine forests with a deciduous component. Our results indicate that ponderosa pine forests may be important to migrating or dispersing landbirds in autumn, especially if there is a deciduous component.     

Arthropod community diversity and trophic structure: a comparison between extremes of plant stress

Abstract.  1. Previous studies have shown that plant stress and plant vigour impact the preference and performance of many insect species. Global climate-change scenarios suggest that some regions such as continental interiors may become increasingly subject to severe drought. In combination, these two issues suggest that drought-driven plant stress may impact insect communities on a landscape scale. While there have been many population studies relating plant stress to the life history of individual herbivore species, far less is known about how plant stress affects entire communities.
2. To study the effect of plant stress on arthropod communities, arthropods were sampled from the canopies of pinyon pines ( Pinus edulis ) growing at sites with a history of chronically high environmental stress (e.g. lower water potentials, soil moisture, and reduced growth rates), and those growing under more favourable conditions. Sampling in these environments yielded >59 000 arthropods, representing 287 species from 14 orders and 80 families, and revealed three major community patterns.
3. First, chronic stress significantly altered community composition. Second, trees growing under high stress supported about 1/10th the number of arthropods, and roughly half the species as trees growing under more favourable conditions. Third, of the 33 abundant herbivore species that exhibited a significantly skewed distribution towards either high- or low-stress trees, 73% were skewed with higher numbers on low-stress trees.
4. The pattern of potentially reduced arthropod diversity and abundance on stressed pines observed in this study may further compound the loss of species resulting from the recent, landscape-scale drought-induced mortality of pines in the southwestern USA.     

Positive effect of catastrophic winds on reptile community recovery in pine plantations

The greatest threat to biodiversity is the alteration and destruction of habitats, and one of the causes is tree plantation, which normally are monospecific and unnatural high-density forest stands. In Spain these contain mainly Pinus species, and cover 25% of the forested area. Due to their dense canopy cover, these monocultures harbor poor communities in terms of species richness and abundance of other organisms, such as reptiles. In November 2014, wind storms affected a pine plantation in the western end of the Sierra Nevada Natural Park, totally or partially knocking down pines 65 years old. We have studied the response of the reptile community in the affected plots three and five years after the disturbance. Due to the low thermal quality of pine plantations for reptiles, we hypothesized that reptile community metrics (abundance and species richness) would be positively affected by this perturbation. We found greater richness and density of reptiles in the plots affected by maximum and intermediate disturbance than in the non-affected plots. We conclude that natural catastrophic events such as wind storms can diversify reptile communities in Mediterranean pine plantations thanks to a rapid response of generalist reptile species.     

Defoliation effects on the ectomycorrhizal community of a mixed Pinus contorta/Picea engelmannii stand in Yellowstone Park

Molecular genetic methods were used to determine whether artificial defoliation affects ectomycorrhizal (EM) colonization, EM fungal species richness, and species composition in a mixed Pinus contorta (lodgepole pine)/Picea engelmannii (Engelmann spruce) forest in Yellowstone National Park, Wyoming. All lodgepole pines in three replicate plots were defoliated 50%, while Engelmann spruce were left untreated. This was done to determine how defoliation of one conifer species would affect EM mutualisms of both treated and neighboring, untreated conifers. The results indicated no significant effect on either EM colonization (142.0 EM tips/core in control plots and 142.4 in treatment plots) or species richness (5.0 species/core in controls and 4.5 in treatments). However, the relative abundance of EM of the two tree species shifted from a ratio of approximately 6:1 without treatment (lodgepole EM:spruce EM), to a near 1:1 ratio post-treatment. This shift may be responsible for maintaining total EM colonization and species richness following defoliation. In addition, EM species composition changed significantly post-defoliation; the system dominant, an Inocybe species, was rare in defoliation plots, while Agaricoid and Suilloid species that were rare in controls were dominant in treatments. Furthermore, species of EM fungi associating with both lodgepole pine and Engelmann spruce were affected, which indicates that changing the photosynthetic capacity of one species can affect mycorrhizal associations of neighboring non-defoliated trees.     

Evidence for indirect effects of plant diversity and composition on net nitrification

Abiotic controls on net nitrification rates are well documented, but the potential effects of plants on this important ecosystem process are poorly understood. We evaluated four structural equation models to determine the relative importance of plant community composition, aboveground herbaceous production, and plant species richness on nitrifier abundance and net nitrification following restoration treatments in a ponderosa pine forest. Model selection criteria indicated that species richness was the best predictor of nitrifier abundance, but a model that included community composition effects also had some support in the data. Model results suggest that net nitrification was indirectly related to plant species richness via a positive relationship between species richness and nitrifier abundance. Community composition was indirectly related to nitrifier abundance through its relationship with species richness. Our model indicates that species-rich plant communities dominated by C₃ graminoids and legumes are associated with soils that have high abundances of nitrifiers. This study highlights the complexity of deciphering effects of ecological treatments on a system response when multiple interacting factors are simultaneously affected. Our results suggest that plant diversity and composition can both respond to forest thinning, prescribed fire and fuel manipulations, and can be factors that might indirectly influence an ecosystem process such as nitrification. Ecological restoration treatments designed to increase plant diversity and alter community composition may have cascading effects on below-ground processes.     

Understorey plant community structure in lower montane and subalpine forests, Grand Canyon National Park, USA

Aim Our objectives were to compare understorey plant community structure among forest types, and to test hypotheses relating understorey community structure within lower montane and subalpine forests to fire history, forest structure, fuel loads and topography. Location Forests on the North Rim of Grand Canyon National Park, Arizona, USA. Methods We measured understorey (< 1.4 m) plant community structure in 0.1‐ha plots. We examined differences in univariate response variables among forest types, used permutational manova to assess compositional differences between forest types, and used indicator species analysis to identify species driving the differences between forest types. We then compiled sets of proposed models for predicting plant community structure, and used Akaike's information criterion (AIC_C) to determine the support for each model. Model averaging was used to make multi‐model inferences if no single model was supported. Results Within the lower montane zone, pine–oak forests had greater understorey plant cover, richness and diversity than pure stands of ponderosa pine (Pinus ponderosa P. & C. Lawson var. scopulorum Engelm.). Plant cover was negatively related to time since fire and to ponderosa pine basal area, and was highest on northern slopes and where Gambel oak (Quercus gambelii Nutt.) was present. Species richness was negatively related to time since fire and to ponderosa pine basal area, and was highest on southern slopes and where Gambel oak was present. Annual forb species richness was negatively related to time since fire. Community composition was related to time since fire, pine and oak basal area, and topography. Within subalpine forests, plant cover was negatively related to subalpine fir basal area and amounts of coarse woody debris (CWD), and positively related to Engelmann spruce basal area. Species richness was negatively related to subalpine fir basal area and amounts of CWD, was positively related to Engelmann spruce basal area, and was highest on southern slopes. Community composition was related to spruce, fir and aspen basal areas, amounts of CWD, and topography. Main conclusions In montane forests, low‐intensity surface fire is an important ecological process that maintains understorey communities within the range of natural variability and appears to promote landscape heterogeneity. The presence of Gambel oak was positively associated with high floristic diversity. Therefore management that encourages lightning‐initiated wildfires and Gambel oak production may promote floristic diversity. In subalpine forests, warm southern slopes and areas with low amounts of subalpine fir and CWD were positively associated with high floristic diversity. Therefore the reduction of CWD and forest densities through managed wildfire may promote floristic diversity, although fire use in subalpine forests is inherently more difficult due to intense fire behaviour in dense spruce–fir forests.     

Suitability of pines and other conifers as hosts for the invasive Mediterranean pine engraver (Coleoptera: Scolytidae) in North America

The invasive Mediterranean pine engraver, Orthotomicus erosus (Wollaston) (Coleoptera: Scolytidae), was detected in North America in 2004, and it is currently distributed in the southern Central Valley of California. It originates from the Mediterranean region, the Middle East, and Asia, and it reproduces on pines (Pinus spp.). To identify potentially vulnerable native and adventive hosts in North America, no-choice host range tests were conducted in the laboratory on 22 conifer species. The beetle reproduced on four pines from its native Eurasian range--Aleppo, Canary Island, Italian stone, and Scots pines; 11 native North American pines--eastern white, grey, jack, Jeffrey, loblolly, Monterey, ponderosa, red, Sierra lodgepole, singleleaf pinyon, and sugar pines; and four native nonpines--Douglas-fir, black and white spruce, and tamarack. Among nonpines, fewer progeny developed and they were of smaller size on Douglas-fir and tamarack, but sex ratios of progeny were nearly 1:1 on all hosts. Last, beetles did not develop on white fir, incense cedar, and coast redwood. With loblolly pine, the first new adults emerged 42 d after parental females were introduced into host logs at temperatures of 20-33 degrees C and 523.5 or 334.7 accumulated degree-days based on lower development thresholds of 13.6 or 18 degrees C, respectively.     

Ectomycorrhizal fungi of exotic pine plantations in relation to native host trees in Iran: evidence of host range expansion by local symbionts to distantly related host taxa

Introduction of exotic plants change soil microbial communities which may have detrimental ecological consequences for ecosystems. In this study, we examined the community structure and species richness of ectomycorrhizal (EcM) fungi associated with exotic pine plantations in relation to adjacent native ectomycorrhizal trees in Iran to elucidate the symbiont exchange between distantly related hosts, i.e. Fagales (Fagaceae and Betulaceae) and Pinaceae. The combination of morphological and molecular identification approaches revealed that 84.6 % of species with more than one occurrence (at least once on pines) were shared with native trees and only 5.9 % were found exclusively on pine root tips. The community diversity of ectomycorrhizal fungi in the pine plantations adjacent to native EcM trees was comparable to their adjacent native trees, but the isolated plantations hosted relatively a species-poor community. Specific mycobionts of conifers were dominant in the isolated plantation while rarely found in the plantations adjacent to native EcM trees. These data demonstrate the importance of habitat isolation and dispersal limitation of EcM fungi in their potential of host range expansion. The great number of shared and possibly compatible symbiotic species between exotic Pinaceae and local Fagales (Fagaceae and Betulaceae) may reflect their evolutionary adaptations and/or ancestral compatibility with one another.     

Fuel-Reduction Treatment Effects on Avian Community Structure and Diversity

Abstract: We assessed responses of the breeding bird community to mechanical thinning and prescribed surface fire, alone and in combination, between 2000 and 2006 in ponderosa pine (Pinus ponderosa) forests in northern Arizona, USA. Fuel-reduction treatments did not affect species richness or evenness, and effects on density of 5 commonly detected species varied among species. Populations of some species, such as the western bluebird (Sialia mexicana), increased following burning treatments, whereas others, such as the mountain chickadee (Poecile gambeli), decreased in response to thinning treatments. Our results also identified a temporal response component, where avian community composition and structure changed synchronously on all treatments over time. Given the modest effects these small-scale fuel-reduction treatments had on avian composition and the specific density responses of particular species, our results suggest that land managers should consider implementing prescribed surface fire after thinning projects, where appropriate.     

Habitat matters: The role of spore bank fungi in early seedling establishment of Florida slash pines

This study investigated broad patterns in communities of ectomycorrhizal fungi from three Florida habitats (sandhills, scrub, and pine rocklands) and the ability of spore bank fungi to associate with Pinus elliottii (slash pine) and Pinus densa (south Florida slash pine). Efforts to replant pines in the endangered pine rocklands are vital to the persistence of this habitat, yet little is known about the ectomycorrhizal fungi communities or how they may differ from those in other pine-dominated habitats in Florida. We used high-throughput amplicon sequencing (HTS) to assess baseline fungal communities and greenhouse bioassays to bait ectomycorrhizal fungi using seedlings. HTS soil data recovered 188 ectomycorrhizal species but only a few subsequently colonized the bioassay seedlings. We recovered 21 ectomycorrhizal species on pine seedlings including common spore bank fungi such as Cenococcum, Suillus, and Tuber, but Rhizopogon species were dominant across all sites and habitats. Habitat type and site were significant variables influencing the community composition of the total soil fungal community, soil ectomycorrhizal community, and the fungi found on seedling root tips. However, we found no significant differences between the ectomycorrhizal communities on seedling roots from the two Pinus species.