Species richness and soil properties in Pinus ponderosa forests: A structural equation modeling analysis

Question: How are the effects of mineral soil properties on understory plant species richness propagated through a network of processes involving the forest overstory, soil organic matter, soil nitrogen, and understory plant abundance? Location: North‐central Arizona, USA. Methods: We sampled 75 0.05‐ha plots across a broad soil gradient in a Pinus ponderosa (ponderosa pine) forest ecosystem. We evaluated multivariate models of plant species richness using structural equation modeling. Results: Richness was highest at intermediate levels of understory plant cover, suggesting that both colonization success and competitive exclusion can limit richness in this system. We did not detect a reciprocal positive effect of richness on plant cover. Richness was strongly related to soil nitrogen in the model, with evidence for both a direct negative effect and an indirect non‐linear relationship mediated through understory plant cover. Soil organic matter appeared to have a positive influence on understory richness that was independent of soil nitrogen. Richness was lowest where the forest overstory was densest, which can be explained through indirect effects on soil organic matter, soil nitrogen and understory cover. Finally, model results suggest a variety of direct and indirect processes whereby mineral soil properties can influence richness. Conclusions: Understory plant species richness and plant cover in P. ponderosa forests appear to be significantly influenced by soil organic matter and nitrogen, which are, in turn, related to overstory density and composition and mineral soil properties. Thus, soil properties can impose direct and indirect constraints on local species diversity in ponderosa pine forests.     

Influence of overstory composition on understory colonization by native species in plantations on a degraded tropical site

Abstract. Patterns of understory colonization by native and naturalized trees and shrubs were evaluated in 4.5-year-old plantations of three exotic tree species, Casuarina equisetifolia, Eucalyptus robusta, and Leucaena leucocephala, on a degraded coastal grassland site with reference to overstory composition and understory environmental conditions. 19 secondary forest species were established in the plantation understories (with a total area of 0.52 ha), while no natural regeneration occurred in unplanted, though protected, control areas. The majority of these species (90 %) and the total seedling population (97 %) were zoochorous, indicating the importance of frugivorous bats and particularly birds as facilitators of secondary forest species colonization. Understory species richness and seedling densities were affected significantly by overstory composition, the most abundant regeneration occurring beneath Leucaena and least under Casuarina. Understory colonization rates within mixed-species stands were intermediate between those of single-species stands of the trees comprising their overstories. Significant negative correlations were found between understory species richness and seedling density, and forest floor depth and dry mass, especially for small-seeded ornithochorous species. Higher colonization rates near the peripheries of plantation plots relative to plot interiors were due in part to roosting site preferences by frugivores, particularly bats. The study results indicate that overstory species selection can exert a significant influence on subsequent patterns of colonization by secondary forest species and is an important consideration in the design of plantations for ‘catalyzing’ succession on deforested, degraded sites.     

Secondary Succession and Indigenous Management in Semideciduous Forest Fallows of the Amazon Basin1

To the discussion on secondary succession in tropical forests, we bring data on three under‐addressed issues: understory as well as overstory changes, continuous as opposed to phase changes, and integration of forest succession with indigenous fallow management and plant uses. Changes in vegetation structure and species composition were analyzed in secondary forests following swidden agriculture in a semideciduous forest of Bolivian lowlands. Twenty‐eight fallows, stratified by four successional stages (early = 1–5 yr, intermediate = 6–10 yr, advanced = 12–20 yr, and older = 22–36 yr), and ten stands of mature forests were sampled. The overstory (plants ≥5 cm diameter at breast height [DBH]) was sampled using a 20 × 50 m plot and the understory (plants <5 cm DBH) in three nested 2 × 5 m subplots. Semistructured interviews provided information on fallow management. Canopy height, basal area, and liana density of the overstory increased with secondary forest age. The early stage had the lowest species density and diversity in the overstory, but the highest diversity in the understory. Species composition and abundance differentiated mature forests and early successional stage from other successional stages; however, species showed individualistic responses across the temporal gradient. A total of 123 of 280 species were useful with edible, medicinal, and construction plants being the most abundant for both over‐ and understories. Most of Los Gwarayo preferred mature forests for making new swidden, while fallows were valuable for crops, useful species, and regenerating timber species.     

Influence of light and soil moisture on Sierran mixed-conifer understory communities

Sierra Nevada forests have high understory species richness yet we do not know which site factors influence herb and shrub distribution or abundance. We examined the understory of an old-growth mixed-conifer Sierran forest and its distribution in relation to microsite conditions. The forest has high species richness (98 species sampled), most of which are herbs with sparse cover and relatively equal abundance. Shrub cover is highly concentrated in discrete patches. Using overstory tree cover and microsite environmental conditions, four habitats were identified; tree cluster, partial canopy, gap, and rock/shallow soil. Herb and shrub species were strongly linked with habitats. Soil moisture, litter depth and diffuse light were the most significant environmental gradients influencing understory plant distribution. Herb cover was most strongly influenced by soil moisture. Shrub cover is associated with more diffuse light, less direct light, and sites with lower soil moisture. Herb richness is most affected by conditions which influence soil moisture. Richness is positively correlated with litter depth, and negatively correlated with direct light and shrub cover. Disturbance or management practices which change forest floor conditions, shallow soil moisture and direct light are likely to have the strongest effect on Sierran understory abundance and richness.     

Forest structure and tree species composition of the understory of two central Amazonian várzea forests of contrasting flood heights

We investigated species composition, distribution, and forest structure of understory trees (≥1 m height, <10 cm diameter at breast height) in two late-successional várzea forests subject to contrasting levels of inundation within the Mamirauá Sustainable Development Reserve, western Brazilian Amazon, and compared it with the overstory flora at the same study sites. In total, 1486 individuals and 116 woody species were recorded on an area totaling 3140 m². Individual densities and tree species richness were considerably higher in the high várzea than in the low várzea, which suggests that the heights and durations of the annual inundations are the main factor limiting species regeneration. In addition, approximately one third of the recorded species with densities ≥8 individuals showed regular or random spatial distribution patterns, which suggests that floodwaters act on dispersal strategies and species establishment.Independent of the forest type, floristic similarity between the understory and the overstory amounted to approximately 35%, and to approximately 10% when compared to other understory inventories in Amazonian várzea. Although the inventoried area of the understory amounted to only 16% of that of the overstory, species richness accounted for approximately 52-56% of that of the overstory. The results indicate that the understory flora of várzea forests is distinct and that it significantly increases local tree species richness. The understory flora of várzea forests therefore should be addressed in floristic inventories that provide the basis for regional and/or basin-wide estimations of tree diversity.     

Conservation value of timber quality versus associated non-timber quality stands for understory diversity in Nothofagus forests

Conservation strategies of forested landscapes must consider biodiversity of the included site types, i.e. timber-quality forests and associated non-timber-quality stands. The objectives were to characterize forest overstory structure in timber-quality versus associated non-timber-quality stands; and to compare their understory communities. Six forest types were sampled in Nothofagus forests of Tierra del Fuego (Argentina): two timber-quality N. pumilio forests, and four associated non-timber-quality stands (edge, N. antarctica, wetlands and streamside forests). Overstory structure and understory vegetation (species richness, frequencies, cover and biomass) were characterized during spring and summer seasons. Analysis of variance and multivariates were carried out. Overstory structure differed across the site types, with higher tree size, canopy closure and tree volume in timber-quality stands. Fifty-one understory plant species were observed, but understory variables varied with site types, especially wetlands (highest native and exotic richness, cover and biomass, and 25% of exclusive species). Site types were grouped in three: N. antarctica stands, streamside stands and the other N. pumilio forests according to multivariate analysis. Forty three percent of plants were distributed in all site types, and all timber-quality forest understory species were present in some associated non-timber-quality stands. Timber-quality N. pumilio forests have a marginal value for understory conservation compared to associated non-timber-quality stands, because these last include all the plants observed in timber-quality forests and also possess many exclusive species. Therefore, protection of associated non-timber-quality stands during forest management planning could increase understory conservation at landscape level, and these could be better reserves of understory diversity than retentions of timber-quality stands.     

Understory vegetation indicates historic fire regimes in ponderosa pine‐dominated ecosystems in the Colorado Front Range

Question: Can current understory vegetation composition across an elevation gradient of Pinus ponderosa‐dominated forests be used to identify areas that, prior to 20th century fire suppression, were characterized by different fire frequencies and severities (i.e., historic fire regimes)? Location: P. ponderosa‐dominated forests in the montane zone of the northern Colorado Front Range, Boulder and Larimer Counties, Colorado, USA. Methods: Understory species composition and stand characteristics were sampled at 43 sites with previously determined fire histories. Indicator species analyses and indirect ordination were used to determine: (1) if stands within a particular historic fire regime had similar understory compositions, and (2) if understory vegetation was associated with the same environmental gradients that influence fire regime. Classification and regression tree analysis was used to ascertain which species could predict fire regimes. Results: Indicator species analysis identified 34 understory species as significant indicators of three distinct historic fire regimes along an elevation gradient from low‐ to high‐elevation P. ponderosa forests. A predictive model derived from a classification tree identified five species as reliable predictors of fire regime. Conclusions: P. ponderosa‐dominated forests shaped by three distinct historic fire regimes have significantly different floristic composition, and current understory compositions can be used as reliable indicators of historical differences in past fire frequency and severity. The feasibility demonstrated in the current study using current understory vegetation properties to detect different historic fire regimes, should be examined in other fire‐prone forest ecosystems.     

Post-fire recovery of ant communities in Submediterranean Pinus nigra forests

This study analyzes the variations in the structure and composition of ant communities in burned Pinus nigra forests in central Catalonia (NE Spain). Pinus nigra forests do not recover after fire, changing to shrublands and oak coppices. For this reason, we suggest that ant communities of burned P. nigra forests will change after fire, because the post‐fire scenario, in particular with the increase of open areas, is different to the unburned one, and more favourable for some species than for others. In four locations previously occupied by P. nigra forests where different fires occurred 1, 5, 13 and 19 yr before the sampling, we sampled the structure and composition of ant communities with pitfall traps, tree traps and net sweeping in unburned plots and in plots affected by canopy and understory fire. The results obtained suggest that canopy and understory fire had little effect on the structure of ant communities. Thus, many variables concerning ant communities were not modified either by fire type (understory or canopy fire) or by time since fire. However, a number of particular species were affected, either positively or negatively, by canopy fire: three species characteristic of forest habitats decreased after fire, while eight species characteristic of open habitats increased in areas affected by canopy fire, especially in the first few years after fire. These differences in ant community composition between burned and unburned plots imply that the maximum richness is achieved when there is a mixture of unburned forests and areas burned with canopy fire. Moreover, as canopy cover in P. nigra forests burned with canopy fire is not completed in the period of time studied, the presence of the species that are characteristic of burned areas remains along the chronosequence studied, while the species that disappear after fire do not recover in the period of time considered. Overall, the results obtained indicate that there is a persistent replacement of ant species in burned P. nigra forests, as is also the case with vegetation.     

Changes in riparian forest composition along a sedimentation rate gradient

Riparian forests are highly valued for maintaining water quality through the retention of sediments and nutrients. They also provide some of the most diverse and species-rich habitats in the world. What is largely unknown, however, is how sediment deposition affects plant community composition in these forests. The objective of this study was to examine changes in plant community composition across a gradient of increasing rates of sedimentation in riparian forests in the southeastern Coastal Plain, USA. Seventeen plots were established within riparian forests receiving between 0 and 5.5 cm year⁻¹ of sediment deposits. Species density and biomass estimates were collected annually from 2002 to 2006 for overstory and mid-story plant species within each plot. Percent cover and nested frequency of understory plant species were determined annually during 2004–2006. Measures of community composition in the understory, mid-story, and overstory layers of forests were compared to changes in environmental factors associated with increased sedimentation. In the understory, annual, exotic, and upland species had higher importance values in plots receiving high sediment deposition. The densities of shade-intolerant and N-fixing species in the mid-story also increased with increasing sedimentation rates. Increased overstory mortality was associated with high sedimentation rates, though increases in understory light levels in these gaps were not the main driver of understory species changes. Edaphic factors, such as soil texture, moisture, and temperature, were significantly correlated to species composition in all three forest layers, suggesting that changes in soil physical structure due to sedimentation may drive community-level changes in these forests.     

The role of succession in the evolution of flammability

Fire-prone ecosystems contain plants that are both fire-adapted and flammable. It has been hypothesized that these plants were under selection to become more flammable, but it is unclear whether this could be adaptive for an individual plant. We propose arrested succession as a robust mechanism that supports the evolution of flammability in surface fire ecosystems without the need to invoke group selection or additional fitness benefits. We used the natural history of lodgepole pine (Pinus ponderosa) forests, longleaf pine (Pinus palustris) forests, and tall grass prairies to create a general mathematical model of surface fire ecosystems and solved for the evolutionarily stable strategy (ESS) level of flammability. In our model, fires always kill understory plants and only sometimes kill overstory plants. Thus, more flammable plants suffer increased mortality due to fires, but also more frequently arrest succession by clearing their understory of late successional competitors. Increased flammability was selected for when the probability of an overstory plant dying from an individual fire was below a maximum threshold and the rate of succession relative to fires was above a minimum threshold. Future studies can test our model predictions and help resolve whether or not plants have been selected to be more flammable.     

Development of Tree Regeneration in Fern-dominated Forest Understories After Reduction of Deer Browsing

Dennstaedtia punctilobula (hay‐scented fern) can act as a native invasive species in forests in eastern North America where prolonged deer browsing occurs in stands with partially open overstory canopies. Ferns dominate the understory with a 60‐cm tall canopy, with little regeneration of native tree species. It has been hypothesized that, once established, ferns may continue to inhibit tree regeneration after deer browsing has been reduced. To test this hypothesis, we documented the pattern of recovery of the tree seedling understory in plantations of Pinus strobus (white pine) and Pinus resinosa (red pine) on the Quabbin Reservation watershed protection forest in central Massachusetts, where after 40 years of intensive deer browsing the deer herd was rapidly reduced through controlled hunting. Dense fern understories occur on nearly 4,000 ha of the predominantly oak–pine forest. Three years after deer herd reduction, stands with the highest density fern cover (77% of plots with>90% cover) had significantly fewer seedlings at least 30 cm in height, compared with stands with lower fern density, and those seedlings consisted almost entirely of Betula lenta (black birch) and white pine. Height growth analysis showed that black birch and white pine grew above the height of the fern canopy in 3 and 6 years, respectively. In contrast, two common species, Fraxinus americana (white ash) and Quercus rubra (red oak), grew beneath the dense fern cover for 5 years with height growth less than 5 cm/yr after the first year. A study of spring phenology indicated that the ability of black birch to grow through the fern canopy might have been due to its early leaf development in spring before the fern canopy was formed, in contrast to oak and ash with delayed leaf development. Thus, the ferns showed differential interference among species with seedling development after reduction of deer browse.     

Changes in the understory during 14 years following catastrophic windthrow in two Minnesota forests

Abstract. We studied the effects of windthrow on the understory plant species composition of a pine forest (dominated by Pinus strobus) and an oak forest (dominated by Quercus ellipsoidalis). We recorded the presence of vascular plant species in randomly located quadrats in the two forests, and in three microsite types associated with tipup mounds (pit, old soil and new soil) in the pine forest at irregular intervals over the course of 14 years. The understories of the two forests remained distinct throughout the study. The frequency of occurrence of a number of forest floor species considerably increased; few species decreased. The disturbance specialists Rubus idaeus and Polygonum cilinode increased in frequency throughout the study in the pine forest, but are beginning to decline in the less disturbed oak forest. Annuals and biennials preferentially colonized the disturbed soil of microsites on tipups, and declined in frequency after about 7 yr. Both forests have increased in understory species richness, but have not changed substantially in the distribution of growth forms. Despite early differences in species composition, microsite types associated with tipup mounds became more similar through time. Although small in magnitude, there was a directional change in understory composition at both forests, with no apparent sign of a return to pre‐disturbance conditions.     

Fire-induced shifts in overstory tree species composition and associated understory plant composition in Glacier National Park,Montana

In Rocky Mountain forests, fire can act as a mechanism of change in plant community composition if postfire conditions favor establishment of species other than those that dominated prefire tree communities. We sampled pre and postfire overstory and postfire understory species following recent (1988–2006) stand-replacing fires in Glacier National Park (GNP), Montana. We identified changes in relative density of tree species and groups of species (xerophytes vs. mesophytes and reseeders vs. resprouters) in early succession. Postfire tree seedling densities were adequate to maintain prefire forest structure, but relative densities among species were variously changed. Changes were directly related to individual species’ response to severe fires. Most notably, relative density of the mesophytic resprouter quaking aspen (Populus tremuloides) and the xerophytic reseeder lodgepole pine (Pinus contorta) increased substantially following fire, with a concomitant decline in proportional abundance of other tree species that, in some cases, dominated stands before fire. Trends identified in our study suggest that forest community shifts toward those dominated by lodgepole pine and quaking aspen are occurring in GNP. Cover of understory species was not affected by tree species composition or density. These forest communities will likely change throughout succession with the addition of shade-intolerant species in early seral stages and shade-tolerant species later in succession. However, with increased fire frequency, the lodgepole pine-dominated postfire communities observed in our study may become more common throughout time.     

Mixed-conifer understory response to climate change, nitrogen, and fire

California's Sierra Nevada mountains are predicted to experience greater variation in annual precipitation according to climate change models, while nitrogen deposition from pollution continues to increase. These changes may significantly affect understory communities and fuels in forests where managers are attempting to restore historic conditions after a century of altered fire regimes. The objective of this research was to experimentally test the effects of increasing and decreasing snowpack depth, increasing nitrogen, and applying prescribed fire to mixed-conifer forest understories at two sites in the central and southern Sierra Nevada. Understory response to treatments significantly differed between sites with herb biomass increasing in shrub-dominated communities when snowpack was reduced. Fire was a more important factor in post-treatment species richness and cover than either snowpack addition or reduction. Nitrogen additions unexpectedly increased herbaceous species richness. These varied findings indicate that modeling future climatic influences on biodiversity may be more difficult than additive prediction based on increasing the ecosystem's two limiting growth resources. Increasing snowpack and nitrogen resulted in increased shrub biomass production at both sites and increased herb production at the southern site. This additional understory biomass has the potential to increase fuel connectivity in patchy Sierran mixed-conifer forests, increasing fire severity and size.     

The population biology of Bromus tectorum in forests: effect of disturbance,grazing, and litter on seedling establishment and reproduction

Summary The effect of tree canopy, understory, herbivores, and litter depth on seedling establishment, survival, and reproduction of the alien grass, Bromus tectorum (cheatgrass), was examined in a series of experiments in four forest habitat types in western North America. Higher recruitment, survival, and reproduction on clearcuts, which would be expected if the overstory alone is limiting the distribution of cheatgrass in forests, were not observed. Removing the understory in an otherwise undisturbed Pinus ponderosa forest did, however, increase the emergence of B. tectorum, but plants in these experimentally-created openings were more vulnerable to grazing by small mammals. In contrast, removing the sparse understory in an Abies forest neither enhanced recruitment nor increased the incidence of grazing of B. tectorum seedlings. Regardless of the forest habitat, most grazed plants died before maturity; even fewer grazed plants produced seeds. Litter depth influenced both recruitment and biomass production: both the rate of germination and the size of resultant seedlings were lower on thick litter (6 cm) compared to results on thin litter (1.5 cm). In the more open Pinus ponderosa and Pseudotsuga menziesii forests, cheatgrass colonization may often occur in openings in the understory alone. Colonization in the more shady A. grandis and Thuja plicata forests is unlikely, however, unless the opening extends through both the understory and the overstory. As a result, cheatgrass is unlikely to increase in any of these forests unless the scale and incidence of disturbance increases substantially.     

Do differences in understory light contribute to species distributions along a tropical rainfall gradient?

In tropical forests, regional differences in annual rainfall correlate with differences in plant species composition. Although water availability is clearly one factor determining species distribution, other environmental variables that covary with rainfall may contribute to distributions. One such variable is light availability in the understory, which decreases towards wetter forests due to differences in canopy density and phenology. We established common garden experiments in three sites along a rainfall gradient across the Isthmus of Panama in order to measure the differences in understory light availability, and to evaluate their influence on the performance of 24 shade-tolerant species with contrasting distributions. Within sites, the effect of understory light availability on species performance depended strongly on water availability. When water was not limiting, either naturally in the wetter site or through water supplementation in drier sites, seedling performance improved at higher light. In contrast, when water was limiting at the drier sites, seedling performance was reduced at higher light, presumably due to an increase in water stress that affected mostly wet-distribution species. Although wetter forest understories were on average darker, wet-distribution species were not more shade-tolerant than dry-distribution species. Instead, wet-distribution species had higher absolute growth rates and, when water was not limiting, were better able to take advantage of small increases in light than dry-distribution species. Our results suggest that in wet forests the ability to grow fast during temporary increases in light may be a key trait for successful recruitment. The slower growth rates of the dry-distribution species, possibly due to trade-offs associated with greater drought tolerance, may exclude these species from wetter forests.     

Two bromeliad species with contrasting functional traits partition the understory space in a South American xerophytic forest: correlative evidence of environmental control and limited dispersal

We examined the spatial distribution of two bromeliad species with contrasting functional traits in the understory of a xerophytic South American Chaco forest. Aechmea distichantha is a facultative terrestrial species with well-developed phytotelma and short rhizomes, whereas Bromelia serra is a strictly terrestrial species with soil-exploring roots and long rhizomes. Both bromeliads develop colonies on relatively elevated patches in Schinopsis balansae forests. We evaluated the roles of environmental controls, limited dispersal, and interspecific competition as drivers of the different distribution of these bromeliads. We mapped the overstory, understory and topography of 16 forest plots with bromeliads (400 m² each, subdivided in 100 4-m² subplots). We sampled soil characteristics on sectors dominated by each bromeliad species. We used structural equation modeling to assess direct and indirect associations of each bromeliad species cover with environmental conditions, abundance of conspecifics in the vicinity, and local abundance of the other species. A. distichantha cover increased on elevated subplots with high tree/shrub basal area, whereas B. serra cover showed the opposite pattern. In addition, A. distichantha cover was negatively associated with B. serra cover, but not vice versa, and cover of both species increased with the abundance of nearby conspecifics, suggesting that limited vegetative dispersal partly accounted for their distribution. Sectors dominated by A. distichantha had lower soil bulk density and higher organic matter content than those dominated by B. serra. According to our model, influences of competition and limited vegetative dispersal reinforce the association between distribution of these bromeliads and environmental heterogeneity of the forest understory.     

Stand- and plot-level changes in a boreal forest understory community following wildfire

Background: Boreal forest understory plant communities are known to be resilient to fire – the species composition of stands after a fire is quite similar to the pre-fire composition. However, we know little about recovery of individual plants within particular locations in forest stands (i.e. plot-level changes) since we usually do not have pre-fire data for plots.

Aims: We wanted to determine whether species recruited into the same or different locations in a Pinus banksiana stand that experienced a severe wildfire.

Methods: We used pre-existing permanent plots to evaluate the cover of understory after an unplanned wildfire.

Results: Across the entire stand nine of 47 species showed a significant change in cover. The largest change in a plant functional group was in the mosses, with all species present before fire being eliminated. There was no change in species diversity or total cover. At the plot level, species composition showed a much greater change. An average of 47% of the species present in a plot before the fire were absent in the same plot after the fire, and the total species turnover in plots was 88% of the species present before the fire. The plots showed a similar shift in species composition.

Conclusions: These results confirm that boreal forest communities show a high degree of resilience to fire, but within a forest stand species will be found in different locations following fire, potentially exposing them to a different set of biotic and abiotic conditions in these new locations.     

大兴安岭次生林白桦对林下日阴菅及其它主要草本植物的影响

探讨了白桦树冠投影面积对林下日阴菅密度的影响,发现日阴菅密度随白桦树冠投影面积呈双峰型变化。海拔高度、坡向、坡度对于日阴菅随白桦树冠投影面积的变化也有一定程度的影响。白桦荫蔽度、日阴菅地上部生物量和土壤水分含量三者具有较为一致的格局规模,这表明白桦通过改变林下的光照和地表水分而影响了日阴菅的生长。日阴菅在中等尺度上与土壤有机质的格局规模一致。溪荪个体数量与土壤pH值在大、中尺度上均显示一致的格局规模,说明溪荪受pH值的影响。