多样化松林中昆虫群落多样性特征

马尾松和湿地松是我国南方的2种主要松树。通过对6种不同林分结构下的马尾松林和湿地松林内昆虫群落调查与多样性指数分析,表明2种松树内的昆虫种类和数量无显著差异,混交林中的昆虫群落的种类和数量比纯林多,尤其以捕食天敌类群的种类和数量更为明显。整个昆虫群落和植食类群多样性指数以湿地松林内较大,而天敌(捕食类群和寄生类群)多样性指数则以马尾松林较高。从不同林分结构下昆虫多样性的比较来看,混交林内昆虫群落多样性指数波动较小,明显地高于纯林。但不同林分结构下昆虫多样性随水平分布和垂直分层格局而变化,松树北面和东面各样地之间的昆虫群落多样性指数差异显著,而南、西面之间差异较小;树冠层各样地之间的差异达极显著水平,而枯枝落叶层和树干层之间差异不显著。由此,还进一步讨论了混交林中昆虫群落稳定性问题。     

Influence of watering and trenching ponderosa pine on a pine sawfly

Summary Neodiprion autumnalis (Smith) larvae were caged for two successive years on root-trenched, watered, and untreated ponderosa pine (Pinus ponderosa Doug. ex Laws.) to determine effects of host moisture stress on larval feeding. Levels of moisture stress (as measured by the Scholander pressure chamber) differed significantly among treatment levels during 1984 and 1985 larval feeding periods. Differences in larval feeding success were not detected in 1984. In 1985, however, larvae on trenched (stressed) trees clipped and rejected more foliage, consumed more needles, had lower pupal weights, lower survival, and a longer feeding period than larvae on watered or untreated trees. Frass production did not differ among treatment levels. The length of the feeding period was shorter for larvae on watered trees than for larvae on untreated trees, but other measures of feeding success did not significantly differ between watered and untreated trees.     

Soil seed banks in Pinus ponderosa forests in Arizona: Clues to site history and restoration potential

Question: How does the relationship between the viable soil seed bank species composition and the above‐ground vegetation in northern Arizona Pinus ponderosa forests differ under varying historical land use disturbances (low, intermediate, high)? Is above‐ground vegetation correlated to the viable soil seed bank immediately following soil disturbance from restoration thinning treatments? Location: Northern Arizona, USA. Methods: Soil seed bank samples were taken along replicated transects and collected with a 5‐cm diameter bulk density hammer. Samples included a 5 ‐cm diameter O‐horizon sample (at varying depths) plus the underlying mineral soil to a depth of 5 cm. The seedling emergent method was used to quantify seed bank species composition and density. The herbaceous and shrub plant community was quantified along the same transects using the point intercept method. Results: Early‐successional or ruderal species were common in the soil seed bank at all three disturbance sites. Non‐native species, notably Verbascum thapsus, were more numerous (up to 940 seeds/m²) under high disturbance with overgrazing and logging, and less common or absent under low disturbance. Most viable seeds were found in the O‐horizon and the upper 5 cm of mineral soil; there was little correlation between species in the soil seed bank and the above‐ground vegetation. Conclusions: We recommend that restoration plans be geared toward minimizing activities, such as severe soil disturbance, that may promote the spread of non‐native invasive species, and that manual seeding be explored as an option to restore plant species diversity and abundance.     

First use of pheromones to detect phenology patterns and density relationships of pine sawflies in German forests

1 Monitoring studies of pine sawflies with pheromone traps were performed for the first time in Germany. Pheromone traps baited with species‐specific pheromone substances were installed in pine forests at different locations in Bavaria, Brandenburg and Lower Saxony during two years. 2 It was possible to track the flight phenology of Diprion pini, Gilpinia pallida and Neodiprion sertifer reliably and to get information about the number of generations of these species in 1997 and 1998. 3 A clear relationship between trap catch and population density could not be detected, but qualitative changes in trap catch caused by different density levels were observed. 4 For D. pini, trap catches were different among endemic populations of different forest types. Furthermore, catches of males reflected the results from the regular cocoon collections by foresters during the previous winter. 5 For N. sertifer, trap catches in endemic populations were well separated from trap catches on sites with higher sawfly densities. However, no significant correlation between trap catch and sawfly density or defoliation level could be found. These results suggest that the efficacy of the pheromone traps probably varied with biological features (sex ratio, density level, immigration) of the particular population.     

Influences of the biophysical environment on blister rust and mountain pine beetle,and their interactions,in whitebark pine forests

Aim To understand how the biophysical environment influences patterns of infection by non‐native blister rust (caused by Cronartium ribicola) and mortality caused by native mountain pine beetles (Dendroctonus ponderosae) in whitebark pine (Pinus albicaulis) communities, to determine how these disturbances interact, and to gain insight into how climate change may influence these patterns in the future. Location High‐elevation forests in south‐west Montana, central Idaho, eastern and western Oregon, USA. Methods Stand inventory and dendroecological methods were used to assess stand structure and composition and to reconstruct forest history at sixty 0.1‐ha plots. Patterns of blister rust infection and mountain pine beetle‐caused mortality in whitebark pine trees were examined using nonparametric Kruskal–Wallis ANOVA, Mann–Whitney U‐tests, and Kolmogorov–Smirnov two‐sample tests. Stepwise regression was used to build models of blister rust infection and mountain pine beetle‐related mortality rates based on a suite of biophysical site variables. Results Occurrence of blister rust infections was significantly different among the mountain ranges, with a general gradient of decreasing blister rust occurrence from east to west. Evidence of mountain pine beetle‐caused mortality was identified on 83% of all dead whitebark pine trees and was relatively homogenous across the study area. Blister rust infected trees of all ages and sizes uniformly, while mountain pine beetles infested older, larger trees at all sites. Stepwise regressions explained 64% and 58% of the variance in blister rust infection and beetle‐caused mortality, respectively, indicating that these processes are strongly influenced by the biophysical environment. More open stand structures produced by beetle outbreaks may increase the exposure of surviving whitebark pine trees to blister rust infection. Main conclusions Variability in the patterns of blister rust infection and mountain pine beetle‐caused mortality elucidated the fundamental dynamics of these disturbance agents and suggests that the effects of climate change will be complex in whitebark pine communities and vary across the species’ range. Interactions between blister rust and beetle outbreaks may accelerate declines or facilitate the rise of rust resistance in whitebark pine depending on forest conditions at the time of the outbreak.     

阔叶红松林林隙结构与树种多样性关系研究

对阔叶红松林林隙与林下不同层次树种多样性进行比较,探讨林隙大小及发育阶段与树种多样性间的关系.结果表明,林隙与林下树种多样性存在显著差异（P<0.01）.与林下相比,林隙更新层树种多样性、丰富度和均匀度增大,生态优势度减小（P<0.01）;演替层树种多样性、丰富度和均匀度减小,生态优势度增大（P<0.0001）.林隙中不同层次树种多样性随着林隙大小呈现出相反的变化趋势.长白山阔叶红松林林隙演替层树种多样性和丰富度总体上呈单峰形变化,中等大小林隙（100～250 m²）中树种多样性和丰富度较高,生态优势度较小.而更新层树种多样性和均匀度在≥250 m²和<100 m²的林隙中最大,在200～250 m²的林隙中最小;生态优势度在200～250 m²的林隙中达到最大.更新层和演替层树种对林隙面积大小反应不同,有利于更新层幼苗建立的林隙面积并不是演替层幼树发育和成活的最适面积;随林隙年龄的增加,更新层和演替层树种多样性在不同层次上呈互补关系.     

The spatial distribution of birds and carabid beetles in pine plantation forests: the role of landscape composition and structure

Aim To evaluate the joint and independent effects of spatial location, landscape composition and landscape structure on the distribution patterns of bird and carabid beetle assemblages in a mosaic landscape dominated by pine plantation forests. Location A continuous 3000‐ha landscape mosaic with native maritime pine Pinus pinaster plantations of different ages, deciduous woodlands and open habitats, located in the Landes de Gascogne forest of south‐western France. Methods We sampled breeding birds by 20‐min point counts and carabid beetles by pitfall trapping using a systematic grid sampling of 200 points every 400 m over the whole landscape. Explanatory variables were composed of three data sets derived from GIS habitat mapping: (1) spatial variables (polynomial terms of geographical coordinates of samples), (2) landscape composition as the percentage cover of the six main habitats, and (3) landscape structure metrics including indices of fragmentation and spatial heterogeneity. We used canonical correspondence analysis with variance partitioning to evaluate the joint and independent effects of the three sets of variables on the ordination of species assemblages. Moran's I correlograms and Mantel tests were used to assess for spatial structure in species distribution and relationships with separate landscape attributes. Results Landscape composition was the main factor explaining the distribution patterns of birds and carabids at the mesoscale of 400 × 400 m. Independent effects of spatial variables and landscape structure were still significant for bird assemblages once landscape composition was controlled for, but not for carabid assemblages. Spatial distributions of birds and carabids were primarily influenced by the amount of heathlands, young pine plantations, herbaceous firebreaks and deciduous woodlands. Deciduous woodland species had positive responses to edge density, while open habitat species were positively associated with mean patch area. Main conclusions Forest birds were favoured by an increase in deciduous woodland cover and landscape heterogeneity, but there was no evidence for a similar effect on carabid beetles. Fragmentation of open habitats negatively affected both early‐successional birds and carabids, specialist species being restricted to large heathlands and young plantations. Several birds of conservation concern were associated with mosaics of woodlands and grasslands, especially meadows and firebreaks. Conserving biodiversity in mosaic plantation landscapes could be achieved by the maintenance of a significant amount of early‐successional habitats and deciduous woodland patches within a conifer plantation matrix.     

Increasing Himalayan tahr and decreasing chamois densities in the eastern Southern Alps, New Zealand: evidence for interspecific competition

There is anecdotal evidence that increasing densities of Himalayan tahr (Hemitragus jemlahicus) are associated with declining densities of chamois (Rupicapra rupicapra) in the Southern Alps, New Zealand. To examine this phenomenon, densities of tahr and chamois were measured at 53 sites within their sympatric range in the eastern Southern Alps during 1978–1979. In sites where only one species was present, tahr density was significantly higher than chamois density (P=0.032), probably reflecting species differences in sociality. Chamois density was higher in catchments without tahr than in those with tahr (P=0.012). Similarly, tahr density was significantly higher at sites without chamois than at those with chamois (P=0.033). Sites with both species present (termed sympatric) were significantly larger than those with only chamois or tahr (P<0.001). Following the prohibition of aerial hunting of tahr in 1983, 16 of the 17 sites where tahr and chamois were sympatric during 1978–1979 were recounted during 1991–1996. There was a 6-fold increase in the mean density of tahr between the two counts (P=0.001), whereas chamois density had declined significantly (P=0.006). Chamois persisted at only three sites, two of which had the highest chamois densities in 1978–1979. This is evidence that increasing densities of tahr exclude chamois from all but the `best' habitats. We conclude that intensive aerial hunting of tahr during 1967–1983 reduced tahr densities such that chamois could co-exist with tahr. Received: 20 March 1997 / Accepted: 8 September 1997     

Effects of fire season on flowering of forbs and shrubs in longleaf pine forests

Summary Effects of variation in fire season on flowering of forbs and shrubs were studied experimentally in two longleaf pine forest habitats in northern Florida, USA. Large, replicated plots were burned at different times of the year, and flowering on each plot was measured over the twelve months following fire. While fire season had little effect on the number of species flowering during the year following fire, fires during the growing season decreased average flowering duration per species and increased synchronization of peak flowering times within species relative to fires between growing seasons. Fires during the growing season also increased the dominance of fall flowering forbs and delayed peak fall flowering. Differences in flowering resulting from variation in fire season were related to seasonal changes in the morphology of clonal forbs, especially fall-flowering composites. Community level differences in flowering phenologies indicated that timing of fire relative to environmental cues that induced flowering was important in determining flowering synchrony among species within the ground cover of longleaf pine forests. Differences in fire season produced qualitatively similar effects on flowering phenologies in both habitats, indicating plant responses to variation in the timing of fires were not habitat specific.     

Influence of landscape and vegetation characteristics on herpetofaunal assemblages in Gulf Coastal Plain pine forests

Longleaf pine (Pinus palustris) savanna characterized by open-canopy, diverse herbaceous vegetation, and high amounts of bare soil once covered much of the southeastern United States Coastal Plain. The unique structural and vegetative conditions of this ecosystem support endemic reptiles and amphibians that have declined as longleaf pine forests have been lost or degraded. Private working pine (Pinus spp.) forests managed for timber production now occur throughout the southeastern United States and have replaced much of the historical longleaf pine savanna. The examination of herpetofaunal (reptile, amphibian) communities in private working loblolly pine (P. taeda) landscapes, particularly in the western Gulf Coastal Plain is lacking. Using repeated field surveys and hierarchical community occupancy models, we examined occupancy and species richness of herpetofauna across 81 sites spanning gradients of management practices, vegetative conditions, and soil composition in northwestern Louisiana, USA, 2017–2019. Young pine stands (<6 yr) exhibited structural characteristics most similar to mature longleaf pine reference sites (>30 yr), while mid-aged stands (13–26 yr) often featured closed canopy and dense midstory. Vegetation conditions varied widely depending on landscape characteristics and site-specific disturbance regimes. We documented 43 species of herpetofauna, including 9 open-pine-associated species. Occupancy of open-pine-associated herpetofauna was positively associated with open-canopy and understory conditions, and sandy soil area. Sites providing open-canopy conditions were often occupied by open-pine-associated species regardless of overstory type and disturbance method. Overall richness of herpetofauna was greatest at sites with moderate canopy cover outside of sandy soil regions. Working pine landscapes in the western Gulf Coastal Plain can support diverse herpetofaunal assemblages, including open-pine-associated species, when management practices maintain open-canopy conditions on sandy, upland soils. More broadly, our results provide insight into how forest management practices affect herpetofauna and may guide practices that can contribute to conservation value of working pine forests.     

小兴安岭阔叶红松林大径级个体对同种邻木分布的影响

利用小兴安岭凉水自然保护区内10.4 hm²样地的调查数据,统计并分析样地内乔木树种的组成与分布,借助混交度计算方法探究大径级个体周围同种邻木分布情况,并分析其产生的原因及大径级个体在其中发挥的作用,研究结果表明：（1）大径级个体（DBH ≥ 50.0 cm）与其周围5-6 m范围内的邻木的混交程度最低,易形成围绕大径级个体（以下简称中央木）的同种邻木环形结构,该环形结构在一定程度上推动着未来的森林演替与更新;（2）距离中央木15 m范围内的中小径级个体（1级木（1.0 cm ≤ DBH < 5.0 cm）、2级木（5.0 cm ≤ DBH < 10.0 cm）、3级木（10.0 cm ≤ DBH < 30.0 cm））与中央木均保持较高的混交程度,而4级木（30.0 cm ≤ DBH < 50.0 cm）、5级木（DBH ≥ 50.0 cm）与中央木的混交程度明显下降,以5级木最低,这种现象的产生与密度制约及物种共存机制存在密切关系。通过研究不同物种及不同生长阶段（不同径级）的林木分布,揭示了凉水地区阔叶红松林中大径级个体周围邻木分布格局特征及其成因,论证了大径级个体在控制周围林分结构、维持物种多样性及推动森林演替中的重要作用,为阔叶红松林的经营及管理提供了数据支撑。     

Predation risk alters interactions among species: competition and facilitation between ants and nesting birds in a boreal forest

Although interactions between species are often assumed to be fixed, theory and empirical evidence suggest that they may be quite variable, changing in the presence of other species or environmental conditions. The interaction between ants and nesting birds exhibits such variability, ants sometimes being predators of bird nests and other times protectors of them. Hypothesizing that predation risk might be a critical factor in altering the interaction of ants with birds, I investigated the interaction of wood ants Formica aquilonia with nesting birds under different levels of predation risk. In a controlled field experiment, I allowed tits ( Parus major, P. caeruleus, P. ater ) and pied flycatchers ( Ficedula hypoleuca ) to select nest boxes in trees with ants (ant trees) or trees without ants. I found that birds usually nested in trees without ants, apparently to avoid the danger of injury from encounters with ants. Nesting in ant trees occurred mainly in the habitat where risk of predation was highest (along the forest edge), and with the bird taxa that lost nests most frequently in trees without ants (tits). Tits nesting on the forest edge achieved significantly greater nesting success, and fledged significantly more young, in ant trees compared with trees without ants. As the season progressed, ant traffic increased in trees without nesting birds, but decreased in trees with nesting birds, indicating that the outcome of interference competition between ants and nesting birds was reversed under increased predation risk. These results support the idea that predation risk can shift species interactions from predominately competitive processes to predominately facilitative processes.     

Competition among canopy trees in indigenous forests: An analysis of the ‘additive basal area’ phenomenon

The basal area of indigenous forest plots containing large canopy individuals appears to be larger than plots without them. One explanation for this effect is the avoidance of competition for light due to these large individuals emerging above the rest of the canopy and thus casting relatively little shade. In this way the basal area of these emergent individuals becomes ‘additive’ to that of the ‘rest’ of the individuals on a plot. The ‘additive basal area’ phenomenon was tested for in the Knysna Forest, South Africa by regressing the basal area of focal species versus the total basal area of 0.04‐ha plots, as well as against the basal area of the ‘rest’. Regression analysis suggested weak competition and a stronger additive effect. However, no emergent individuals occurred in the study taxa. A strong impact of the size of the largest individual on total plot basal area was found. It is suggested that the reason for this is that large individuals overcome spatial and packing limitations in forests.     

The symmetry of competitive interactions in mixed Norway spruce,silver fir and European beech forests

Questions

We aim for a better understanding of the different modes of intra‐ and inter‐specific competition in two‐ and three‐species mixed‐forests. How can the effect of different modes of competitive interactions be detected and integrated into individual tree growth models? Are species interactions in spruce–fir–beech forests more associated with size‐symmetric or size‐asymmetric competition? Do competitive interactions between two of these species change from two‐ to three‐species mixtures?

Location

Temperate mixed‐species forests in Central Europe (Switzerland).

Methods

We used data from the Swiss National Forest Inventory to fit basal area increment models at the individual tree level, including the effect of ecological site conditions and indices of size‐symmetric and size‐asymmetric competition. Interaction terms between species‐specific competition indices were used to disentangle significant differences in species interactions from two‐ to three‐species mixtures.

Results

The growth of spruce and fir was positively affected by increasing proportions of the other species in spruce–fir mixtures, but negative effects were detected with increasing presence of beech. We found that competitive interactions for spruce and fir were more related to size‐symmetric competition, indicating that species interactions might be more associated with competition for below‐ground resources. Under constant amounts of stand basal area, the growth of beech clearly benefited from the increasing admixture of spruce and fir. For this species, patterns of size‐symmetric and size‐asymmetric competitive interactions were similar, indicating that beech is a strong self‐competitor for both above‐ground and below‐ground resources. Only for silver fir and beech, we found significant changes in species interactions from two‐ to three‐species mixtures, but these were not as prominent as the effects due to differences between intra‐ and inter‐specific competition.

Conclusions

Species interactions in spruce–fir–beech, or other mixed forests, can be characterized depending on the mode of competition, allowing interpretations of whether they occur mainly above or below ground level. Our outcomes illustrate that species‐specific competition indices can be integrated in individual tree growth functions to express the different modes of competition between species, and highlight the importance of considering the symmetry of competition alongside competitive interactions in models aimed at depicting growth in mixed‐species forests.

     

Concentrations and fluxes of dissolved organic carbon in an age-sequence of white pine forests in Southern Ontario,Canada

We determined concentrations and fluxes of dissolved organic carbon (DOC) in precipitation, throughfall, forest floor and mineral soil leachates from June 2004 to May 2006 across an age-sequence (2-, 15-, 30-, and 65-year-old) of white pine (Pinus strobus L.) forests in southern Ontario, Canada. Mean DOC concentration in precipitation, throughfall, leachates of forest floor, Ah-horizon, and of mineral soil at 1 m depth ranged from ∼2 to 7, 9 to 18, 32 to 88, 20 to 66, and 2 to 3 mg DOC L⁻¹, respectively, for all four stands from April (after snowmelt) through December. DOC concentration in forest floor leachates was highest in early summer and positively correlated to stand age, aboveground biomass and forest floor carbon pools. DOC fluxes via precipitation, throughfall, and leaching through forest floor and Ah-horizon between were in the range of ∼1 to 2, 2 to 4, 0.5 to 3.5, and 0.1 to 2 g DOC m⁻², respectively. DOC export from the forest ecosystem during that period through infiltration and groundwater discharge was estimated as ∼7, 4, 3, and 2 g DOC m⁻² for the 2-, 15-, 30-, and 65-year-old sites, respectively, indicating a decrease with increasing stand age. Laboratory DOC sorption studies showed that the null-point DOC concentration fell from values of 15 to 60 mg DOC L⁻¹ at 0 to 5 cm to <15 mg DOC L⁻¹ at 50 cm. Specific ultraviolet light absorption at 254 nm (SUVA₂₅₄) increased from precipitation and throughfall to a maximum in forest floor and decreased with mineral soil depth. No age-related pattern was observed for SUVA₂₅₄ values. DOC concentration in forest floor soil solutions showed a positive exponential relationship with soil temperature, and a negative exponential relationship with soil moisture at all four sites. Understanding the changes and controls of DOC concentrations, chemistry, and fluxes at various stages of forest stand development is necessary to estimate and predict DOC dynamics on a regional landscape level and to evaluate the effect of land-use change.     

Ecomorphological pattern in Bermuda birds: The influence of competition and implications for nature preserves

Summary Previous ecomorphological studies of introduced land birds on oceanic islands have revealed several patterns consistent with the hypothesis that interspecific competition influenced the assembly process of that community. We extend one of these analyses to Bermuda, which differs from the previous islands in its size, latitude and isolation from the mainland. We tested for the presence of morphological overdispersion in surviving introduced passeriforms. Despite a possible predominance of random events in the assembly process due to Bermuda's small size and the presence of migrant species, the surviving passeriforms were morphologically overdispersed. The presence of this pattern supports the hypothesis that the assembly of the Bermuda passeriform community has been influenced by interspecific competition. This marks the third distinct community of island land birds in which this competitive pattern has been identified. This is particularly interesting because certain attributes of Bermuda and its avifauna make it more equivalent to communities in mainland refuges than avian assemblages on islands studied previously. This suggests that interspecific competition may have important effects on the structure of refuge communities.     

Rewiring of interactions in a changing environment: nettle-feeding butterflies and their parasitoids

Climate and land use change can alter the incidence and strength of biotic interactions, with important effects on the distribution, abundance and function of species. To assess the importance of these effects and their dynamics, studies quantifying how biotic interactions change in space and time are needed. We studied interactions between nettle-feeding butterflies and their shared natural enemies (parasitoids) locally and across 500 km latitudinal gradient in Sweden. We also examined the potential impact of the range-expansion of the butterfly Araschnia levana on resident butterflies via shared parasitoids, by studying how parasitism in resident butterflies covaries with the presence or absence of the newly-established species. We collected 6777 larvae of four nettle-feeding butterfly species (Aglais urticae, Aglais io, Ar. levana and Vanessa atalanta), over two years, at 19 sites distributed along the gradient. We documented the parasitoid complex for each butterfly species and measured their overlap, and analysed how parasitism rates were affected by butterfly species assemblage, variations in abundance, time, and the arrival of Ar. levana. Parasitoids caused high mortality, with substantial overlap in the complex of parasitoids associated with the four host butterflies. Levels of parasitism differed significantly among butterflies and were influenced by the local butterfly species assemblage. Our results also suggest that parasitism in resident butterflies is elevated at sites where Ar. levana has been established for a longer period. In our study system, variations in butterfly species assemblages were associated in a predictable way with substantial variations in rates of parasitism. This relationship is likely to affect the dynamics of the butterfly host species, and potentially cascade to the larger number of species with which they interact. These results highlight the importance of indirect interactions and their potential to reorganise ecological communities, especially in the context of shifts in species distributions in a warmer world.     

Effects of water potential and solute on the growth and interactions of two fungal symbionts of the mountain pine beetle

We investigated the effect of water potential (WP) on the growth of, and interaction between, two ophiostomatoid fungi, Grosmannia clavigera and Ophiostoma montium, associated with the mountain pine beetle (Dendroctonus ponderosae). The WP of malt extract agar was amended by adding potassium chloride (KCl) or sucrose. Growth of both fungi decreased with WP on KCl-amended media. Growth of G. clavigera also decreased with WP on sucrose-amended media, although growth was stimulated on these media compared to unamended treatments. Growth of O. montium remained relatively constant on sucrose-amended media, confounding the effect of WP on this species. Both fungi were able to colonize media occupied by the other species, but at a slower rate than on unoccupied media, indicating competition. In most treatments, G. clavigera grew faster than O. montium and colonized a greater area when the two fungi were inoculated concurrently but distant to one another on a Petri dish. However, when each fungus was inoculated adjacent to a 10-d-old well-established colony of the other species, O. montium colonized occupied media more effectively than G. clavigera considering the growth rate of each species alone. Thus, G. clavigera dominated primary (uncolonized) resources on most media, whereas O. montium was more effective in colonizing secondary (occupied) resources. The differential response of the two fungi to sucrose indicates that they may use different carbon sources, or use different carbon sources at different rates, in the tree. Fine-scale resource partitioning, differences in primary and secondary resource capture abilities, and the non-equilibrium dynamics in an attacked tree over time, could all act to promote the co-existence of two unit-restricted dispersers on a discontinuous resource.