Strategic light manipulation as a restoration strategy to reduce alien grasses and encourage native regeneration in Hawaiian mesic forests

Question: Is there a light level at which alien grass biomass is reduced while still supporting growth and survival of native woody species, allowing for native species regeneration in abandoned pastures? Location: Island of Hawaii, USA. Methods: In a two‐part study we examined the effect of light availability on common native woody and alien grass species found in secondary forests in Hawaii. A field survey was conducted to examine the relationship between light availability and canopy type (open pasture, planted canopy and secondary forest) on understory grass biomass and litter accumulation. We then experimentally manipulated light levels to determine the effect of light availability on growth and survival of six native woody species and three alien grasses. Low‐light (5%), medium‐light (10%) and high‐light (20‐30%) treatments were created using shade structures erected beneath the existing secondary koa canopy. Results: In the field survey, alien grass biomass was greatest under the open pasture and lowest in the secondary forest. There was a positive correlation between understory light availability and alien grass biomass. In the experimental study, large reductions in relative growth rates were documented for all of the grass species and four of the six woody species under the lowest light level. Although growth at 5% light is substantially reduced, survival is still high (84‐100%), indicating that these species may persist under closed canopy. Conclusion: Low‐light conditions result in the greatest reduction in alien grass biomass while creating an environment in which native woody species can grow and survive.     

Effects of exotic annual grass litter and local environmental gradients on annual plant community structure

Exotic annual grasses have been introduced into many semi-arid ecosystems worldwide, often to the detriment of native plant communities. The accumulation of litter from these grasses (i.e. residual dry biomass) has been demonstrated to negatively impact native plant communities and promote positive feedbacks to exotic grass persistence. More targeted experiments are needed, however, to determine the relative impact of exotic grass litter on plant community structure across local environmental gradients. We experimentally added exotic grass litter to annual forb-dominated open woodland communities positioned along natural canopy cover gradients in southwest Western Australia. These communities are an important component of this region’s plant biodiversity hotspot and are documented to be under threat from exotic annual grasses. After a one-year treatment period, we measured the effects of exotic grass litter, soil properties, and canopy cover on native and exotic species richness and abundance, as well as common species’ biomass and abundances. Plant community structure was more strongly influenced by soil properties and canopy cover than by grass litter. Total plant abundances per plot, however, were significantly lower in litter addition plots than control plots, a trend driven by native species. Exotic grass litter was also associated with lower abundances of one very common native species: Waitzia acuminata. Our results suggest that exotic grass litter limits the establishment of some native species in this system. Over multiple years, these subtle impacts may contribute substantially to the successful advancement of exotic species into this system, particularly in certain microenvironments.     

Impact of invasion of Acer platanoides on canopy structure and understory seedling growth in a hardwood forest in North America

Invasion by exotic plant species is known to affect native communities and ecosystems, but the mechanisms of the impacts are much less understood. In a field study, we examined the effects of a tree invader, Acer platanoides (Norway maple, NM), on canopy structure and seedling growth in the understory of a North American deciduous forest. The experimental site contains a monospecific patch of A. platanoides and a mixed patch of A. platanoides with its native congener, A. rubrum (red maple, RM). In the study, we examined canopy characteristics of three types of trees in the forests, i.e., RM trees in the mixed forest, NM trees in the mixed forest, and NM trees in its monospecific patch. Height growth and biomass production of RM and NM seedlings under intact canopies and newly created gaps of the three types of trees were followed for two growing seasons. We found that removal of half of the canopy from focal trees increased canopy openness and light transmission to the forest floor, but to a greater extent under NM trees than under RM trees. Seedlings of these two Acer species varied greatly in biomass production under canopies of the same type of trees and in their responses to canopy opening. For example, seedlings of the exotic NM grown under the native RM trees in the mixed forests increased biomass production by 102.4% compared to NM seedlings grown under conspecific trees. The native RM seedlings grown under NM trees, however, reduced biomass production by 23.5% compared to those grown under conspecific trees. It was also observed that RM was much more responsive in biomass production to canopy opening than NM. For instance, total seedling biomass increased by 632.2% in RM, but by only 134.6% in NM in response to the newly created gaps. In addition, we found that NM seedlings allocated a greater portion of biomass below-ground as canopy openness increased, whereas the same trend was not observed in RM seedlings. Our results thus demonstrated that invasion of NM significantly altered canopy structure and community dynamics in the hardwood forest. Because the exotic NM seedlings are able to grow well under the native RM trees, but not vice versa, NM will likely expand its distribution in the forests and make it an ever increasingly serious tree invader in its non-native habitats, including North America.     

<Emphasis Type="Italic">Hedychium gardnerianum</Emphasis> Invasion into Hawaiian Montane Rainforest: Interactions Among Litter Quality,Decomposition Rate,and Soil Nitrogen Availability

Few studies have examined the invasion of understory species into closed-canopy forests and, despite inter-specific differences in litter quality and quantity between understory and dominant canopy trees, the influence of understory invasions on soil nitrogen (N) cycling remains unknown. This paper examines litter quality and decomposition of kahili ginger (Hedychium gardnerianum), an invasive understory herb, to determine the influence of this species on N cycling in a Hawaiian montane rainforest. To examine the potential feedback between increased soil N availability and litter decomposition, litter from the invasive ginger, a native tree, and native tree fern was collected from unfertilized and fertilized plots and decomposed in a reciprocal transplant design. Hedychium litter decomposed faster than litter from the two native species. Across species, decomposition rates were negatively correlated with litter lignin content. Despite rapid decomposition rates of Hedychium litter, soil nitrogen availability and rates of net mineralization in the soil were similar in invaded and uninvaded plots. Nitrogen cycling at this site may be more strongly influenced by native species, which contribute the most to overall stand biomass. A negative effect of fertilization on the decomposition of Hedychium litter suggests that a negative feedback between litter quality and soil N availability may exist over longer timescales.     

Successful restoration of Metrosideros polymorpha (ʻōhiʻa) is possible in forest sites with active Rapid ‘Ōhiʻa Death infections

Rapid ?ōhi?a Death (ROD), caused by the fungal pathogen Ceratocystis, is killing large numbers of ?ōhi?a trees (Metrosideros polymorpha) in Hawai?i. ?ōhi?a are a dominant tree in Hawaiian forests, have a range that goes from arid to wet forest climates, and are important for endangered species habitat and ecosystem function. To test whether actively planting ?ōhi?a seedlings is a viable restoration strategy in areas with high ROD mortality, we planted ?ōhi?a in a ROD‐affected forest and crossed this with weeding and fencing treatments to compare ROD mortality to other stressors. We also tested for viable Ceratocystis spores in soils around planting areas. We found that seedlings were more likely to die in unweeded and unfenced treatments than controls. Although viable Ceratocystis spores were found in soil, none of the 41 dead seedlings tested positive for Ceratocystis. This indicates that competition from exotic plants and exotic feral ungulate damage are more likely to kill seedlings than ROD within the first year after planting.     

Shrubs as ecosystem engineers in a coastal dune: influences on plant populations,communities and ecosystems

Question: How do two shrubs with contrasting life‐history characteristics influence abundance of dominant plant taxa, species richness and aboveground biomass of grasses and forbs, litter accumulation, nitrogen pools and mineralization rates? How are these shrubs – and thus their effects on populations, communities and ecosystems – distributed spatially across the landscape? Location: Coastal hind‐dune system, Bodega Head, northern California. Methods: In each of 4 years, we compared vegetation, leaf litter and soil nitrogen under canopies of two native shrubs –Ericameria ericoides and the nitrogen‐fixing Lupinus chamissonis– with those in adjacent open dunes. Results: At the population level, density and cover of the native forb Claytonia perfoliata and the exotic grass Bromus diandrus were higher under shrubs than in shrub‐free areas, whereas they were lower under shrubs for the exotic grass Vulpia bromoides. In contrast, cover of three native moss species was highest under Ericameria and equally low under Lupinus and shrub‐free areas. At community level, species richness and aboveground biomass of herbaceous dicots was lower beneath shrubs, whereas no pattern emerged for grasses. At ecosystem level, areas beneath shrubs accumulated more leaf litter and had larger pools of soil ammonium and nitrate. Rates of nitrate mineralization were higher under Lupinus, followed by Ericameria and then open dune. At landscape level, the two shrubs – and their distinctive vegetation and soils – frequently had uniform spatial distributions, and the distance separating neighbouring shrubs increased as their combined sizes increased. Conclusions: Collectively, these data suggest that both shrubs serve as ecosystem engineers in this coastal dune, having influences at multiple levels of biological organization. Our data also suggest that intraspecific competition influenced the spatial distributions of these shrubs and thus altered the distribution of their effects throughout the landscape.     

Effects of exotic invasive trees on nitrogen cycling: a case study in Central Spain

We assess the hypothesis that rates of nitrogen transformations in the soil are altered upon replacement of native by exotic trees, differing in litter properties. Ailanthus altissima and Robinia pseudoacacia, two common exotic trees naturalized in the Iberian Peninsula, were compared with the native trees Ulmus minor and Fraxinus angustifolia, respectively. Naturally senesced leaves of each species were collected and C:N ratio, N and lignin content assessed. We prepared 64 litter bags per species and left them to decompose, below the canopy of the same species and below the canopy of the paired species. Dry mass, N concentration and N pool of the remaining litter were assessed after 5 and 7 months. Soil samples were collected three times during the experiment to assess soil moisture, organic matter, pH, potential mineralization rates and mineral N pools. Mineral N availability was assessed three times in the field by using ion-exchange resin-impregnated membranes. Ailanthus litter decomposed faster than Ulmus litter, probably due to the higher toughness of the latter. In spite of its high N content, Robinia litter decomposed slower than Fraxinus one, probably due to its high lignin content. In both cases, litter decomposition was faster below the exotic than the native canopies. The release of N per unit of initial litter mass was higher under both invaded situations (Ailanthus below Ailanthus and Robinia below Robinia) than under the native ones. However, soils collected below native and exotic trees neither differed in potential N mineralization rate nor in mineral N. This may be attributed to a quick plant uptake of released N and/or to a high organic matter accumulation in the soil previous to invasion that can exert a tighter control on soil N transformations than the current exotic litter.     

Hemlock Woolly Adelgid in New England Forests: Canopy Impacts Transforming Ecosystem Processes and Landscapes

Exotic insect pests may strongly disrupt forest ecosystems and trigger major shifts in nutrient cycling, structure, and composition. We examined the relationship between these diverse effects for the hemlock woolly adelgid (HWA, Adelges tsugae Annand) in New England forests by studying its impacts on local canopy processes in stands differing in infestation levels and linking these impacts to shifts in canopy nutrient cycling and stand and landscape effects. HWA initiated major changes in canopy biomass and distribution. Whereas uninfested trees exhibit a significant decline in canopy biomass from the center to the periphery and a positive correlation between total needle litter and estimated biomass, infested trees have significantly less total canopy biomass, produce less new foliage, shed relatively more needles, and exhibit no correlation between litter and canopy biomass. Foliar N content of infested trees was 20%–40% higher than reference trees, with the strongest increase in young foliage supporting the highest densities of HWA. Foliar %C was unaffected by HWA or foliar age. Epiphytic microorganisms on hemlock needles exhibited little variation in abundance within canopies, but colony-forming units of bacteria, yeast, and filamentous fungi were 2–3 orders of magnitude more abundant on medium and heavily infested than uninfested trees. Throughfall chemistry, quantity, and spatial pattern were strongly altered by HWA. Throughfall exhibits a strong gradient beneath uninfested trees, decreasing in volumes from the canopy periphery to the trunk by more than 45%. The amount of throughfall beneath infested trees exhibits no spatial pattern, reaches 80%–90% of the bulk precipitation, and is characterized by significantly higher concentrations of nitrogen compounds, dissolved organic carbon, and cations. Across the southern New England landscape there is a strong south-to-north gradient of decreasing hemlock tree and sapling mortality and understory compositional change that corresponds to the duration of infestation. Regionally, black birch (Betula lenta L.) is profiting most from hemlock decline by significantly increasing in density and cover. These findings suggest that it is necessary to study the connections between fast/small-scale processes such as changes in nutrient cycling in tree canopies and slow/integrative processes like shifts in biogeochemieal cycling and compositional changes at forest stands and landscapes to better understand the effects of an exotic pest species like HWA on forest ecosystem structure and function.     

Removal of invasive shrubs alters light but not leaf litter inputs in a deciduous forest understory

The establishment and spread of non‐native, invasive shrubs in forests poses an important obstacle to natural resource conservation and management. This study assesses the impacts of the physical removal of a complex of woody invasive shrub species on deciduous forest understory resources. We compared leaf litter quantity and quality and understory light transmittance in five pairs of invaded and removal plots in an oak‐dominated suburban mature forest. Removal plots were cleared of all non‐native invasive shrubs. The invasive shrubs were abundant (143,456 stems/ha) and diverse, dominated by species in the genera Ligustrum, Viburnum, Lonicera, and Euonymus. Annual leaf litter biomass and carbon inputs of invaded plots were not different from removal plots due to low leaf litter biomass of invasive shrubs. Invasive shrub litter had higher nitrogen (N) concentrations than native species; however, low biomass of invasive litter led to low N inputs by litter of invasive species compared to native. Light transmittance at the forest floor and at 2 m was lower in invaded plots than in removal plots. We conclude that the removal of the abundant invasive shrubs from a native deciduous forest understory did not alter litter quantity or N inputs, one measure of litter quality, and increased forest understory light availability. More light in the forest understory could facilitate the restoration of forest understory dynamics.     

Do Tree Canopies Enhance Perennial Grass Restoration in California Oak Savannas?

Scattered trees in grass‐dominated ecosystems often act as islands of fertility with important influences on community structure. Despite the potential for these islands to be useful in restoring degraded rangelands, they can also serve as sites for the establishment of fast growing non‐native species. In California oak savannas, native perennial grasses are rare beneath isolated oaks and non‐native annual grasses dominate. To understand the mechanisms generating this pattern, and the potential for restoration of native grasses under oaks, we asked: what are the effects of the tree understory environment, the abundance of a dominant non‐native annual grass (Bromus diandrus), and soils beneath the trees on survival, growth, and reproduction of native perennial grass seedlings? We found oak canopies had a strong positive effect on survival of Stipa pulchra and Poa secunda. Growth and reproduction was enhanced by the canopy for Poa but negatively impacted for Stipa. We also found that Bromus suppressed growth and reproduction in Stipa and Poa, although less so for Stipa. These results suggest the oak understory may enhance survival of restored native perennial grass seedlings. The presence of exotic grasses can also suppress growth of native grasses, although only weakly for Stipa. The current limitation of native grasses to outside the canopy edge is potentially the result of interference from annual grasses under oaks, especially for short‐statured grasses like Poa. Therefore, control of non‐native annual grasses under tree canopies will enhance the establishment of S. pulchra and P. secunda when planted in California oak savannas.     

Individual Canopy‐tree Species Effects on Their Immediate Understory Microsite and Sapling Community Dynamics

Canopy trees are largely responsible for the environmental heterogeneity in the understory of tropical and subtropical species‐rich forests, which in turn may influence sapling community dynamics. We tested the effect of the specific identity of four cloud forest canopy trees on total solar radiation, canopy openness, soil moisture, litter depth, and soil temperature, as well as on the structure and dynamics of the sapling community growing beneath their canopies. We observed significant effects of the specific identity of canopy trees on most understory microenvironmental variables. Soil moisture was higher and canopy openness lower beneath Cornus disciflora. In turn, canopy openness and total solar radiation were higher beneath Oreopanax xalapensis, while the lowest soil moisture occurred beneath Quercus laurina. Moreover, Chiranthodendron pentadactylon was the only species having a positive effect on litter depth under its canopy. In spite of these between‐species environmental differences, only C. pentadactylon had significant, negative effects on sapling density and species richness, which may be associated to low seed germination and seedling establishment due to an increased litter depth in its vicinity. The relevance of the specific identity of canopy trees for natural regeneration processes and species richness maintenance depends on its potential to differentially affect sapling dynamics through species‐specific modifications of microenvironmental conditions.     

Fire, native species, and soil resource interactions influence the spatio-temporal invasion pattern of Bromus tectorum

Bromus tectorum (cheatgrass) is an invasive annual that occupies perennial grass and shrub communities throughout the western United States. Bronus tectorum exhibits an intriguing spatio‐temporal pattern of invasion in low elevation ponderosa pine Pinus ponderosa/bunchgrass communities in western Montana where it forms dense rings beneath solitary pines following fire. This pattern provides a unique opportunity to investigate several indirect effects of native vegetation that influence the invasion pattern of B. tectorum, and specifically how native species, disturbance, and soil resources interact to influence the spatio‐temporal pattern of invasion. We established four replicate field sites, each containing burned‐tree, burned‐grass, unburned‐tree, and unburned‐grass sampling locations, and initiated a series of field sampling and greenhouse experiments utilizing these locations. The objective of our first greenhouse experiment was to identify whether belowground factors contributed to the pattern of B. tectorum biomass observed in these field locations. This experiment generated a B. tectorum biomass response that was nearly identical to the invasion pattern observed in the field, suggesting further investigation of belowground factors was necessary. We measured resin‐sorbed NH₄⁺ and NO₃⁻ during one generation of B. tectorum, and measured a suite of P fractions through a sequential extraction procedure from these soils. These data revealed that a resource island of high N and P exists beneath pine trees. Through a second greenhouse experiment, we determined that N limited B. tectorum biomass in tree soil, whereas P limited biomass in bunchgrass soil. Finally, through a germination experiment we determined that pine litter strongly inhibited B. tectorum germination. These data suggest B. tectorum is regulated by P in bunchgrass soil, and by N and inhibition by pine litter beneath trees, effects that are likely alleviated by fire. These data demonstrate the combined role of direct and indirect interactions between native and invasive species in regulating biological invasions.     

Population estimates of the Endangered Hawaiʻi ʻĀkepa (Loxops coccineus) in different habitats on windward Mauna Loa

Endangered Hawai‘i ?ākepas (Loxops coccineus) are endemic to Hawai‘i island, where they occur in five spatially distinct populations. Data concerning the status and population trends of these unique Hawaiian honeycreepers are crucial for assessing the effectiveness of recovery and management actions. In 2016, we used point‐transect distance sampling to estimate the abundance of Hawai‘i ?ākepas in portions of Hawai‘i Volcanoes National Park (HAVO) and the Ka?ū Forest Reserve (KFR) on Mauna Loa volcano. We then compiled the survey data from four other populations to provide a global population estimate. In our HAVO and KFR study area, we mapped habitat classes to determine the population densities in each habitat. Densities were highest (1.03 birds/ha) in open‐canopy montane ?ōhi?a (Metrosideros polymorpha) woodland. In contrast, densities of the largest ?ākepa population on Mauna Kea volcano were highest in closed‐canopy ?ōhi?a and koa (Acacia koa) forest where the species is dependent on nest cavities in tall (> 15 m), large (> 50‐cm diameter at breast height) trees. We surveyed potential nesting habitat in HAVO and KFR and found only one cavity in the short‐stature montane ?ōhi?a woodland and five cavities in the tall‐stature forest. Differences in densities between the Mauna Kea and Mauna Loa populations suggest that Hawai‘i ?ākepas may exhibit different foraging and nesting behaviors in the two habitats. The estimated overall population density in the HAVO and KFR study area was 0.52 birds/ha, which equates to 3663 (95% CI 1725–6961) birds in their 11,377‐ha population range. We calculated a global population of 16,428 (95% CI 10,065–25,198) birds, which is similar to an estimate of 13,892 (95% CI 10,315–17,469) birds made in 1986. Our results suggest that populations are stable to increasing in the two largest populations, but the three other populations are smaller (range = 77–1443 birds) and trends for those populations are unknown.     

The interaction between propagule pressure,habitat suitability and density‐dependent reproduction in species invasion

Seedling recruitment limitations create a demographic bottleneck that largely determines the viability and structure of plant populations and communities, and pose a core restriction on the colonization of novel habitat. We use a shade‐tolerant, invasive grass, Microstegium vimineum, to examine the interplay between seed and establishment limitations – phenomena that together determine recruitment success but usually are investigated individually. We add increasing amounts of seed to microhabitats containing variable levels of leaf litter thickness – with reduced leaf litter simulating disturbance – to investigate whether reduced seed limitation overcomes the establishment limitation posed by litter cover. We do this across gradients in understory light, moisture and temperature, and quantify germination, survival, and then per capita adult biomass and reproduction in order to understand the implications for invasion across the landscape. We find that the combined effects of seed and establishment limitation influence recruitment; however, propagule pressure overwhelms the inhibitory effects of leaf litter thickness. Leaf litter reduces germination by 22–57% and seedling survival by 13–15% from that observed on bare soil. However, density‐dependent reproduction compensates as 1–3 plants can produce far more seeds (approx. 525) than are required for persistence. As such, just a few plants may establish in understory forest habitat and subsequently overwhelm establishment barriers with copious propagule production. These results, for a widespread, invasive plant, are consistent with the emerging perspective for native plants that seed and establishment limitation jointly influence recruitment. The ability for an exotic plant species to compensate for low population densities with high per capita seed production, that then overrides establishment limitations, makes its invasive potential daunting. Further work is required to test if this is a common mechanism underlying plant invasions.     

A non-native invasive grass increases soil carbon flux in a Hawaiian tropical dry forest

Non‐native plants are invading terrestrial ecosystems across the globe, yet little is known about how invasions impact carbon (C) cycling or how these impacts will be influenced by climate change. We quantified the effect of a non‐native C₄ grass invasion on soil C pools and fluxes in a Hawaiian tropical dry forest over 2 years in which annual precipitation was average (Year 1) and ～60% higher than average (Year 2). Work was conducted in a series of forested plots where the grass understory was completely removed (removal plots) or left intact (grass plots) for 3 years before experiment initiation. We hypothesized that grass invasion would: (i) not change total soil C pools, (ii) increase the flux of C into and out of soils, and (iii) increase the sensitivity of soil C flux to variability in precipitation. In grass plots, grasses accounted for 25–34% of litter layer C and ～70% of fine root C. However, no differences were observed between treatments in the size of any soil C pools. Moreover, grass‐derived C constituted a negligible fraction of the large mineral soil C pool (< 3%) despite being present in the system for ≥50 years. Tree litterfall was ～45% lower in grass plots, but grass‐derived litterfall more than compensated for this reduction in both years. Annual cumulative soil‐surface CO₂ efflux (R_soil) was ～40% higher in grass plots in both years, and increased in both treatments by ～36% in the wetter Year 2. Despite minimal grass‐derived mineral soil C, > 75% of R_soil in grass plots was of C₄ (i.e. grass) origin. These results demonstrate that grass invasion in forest ecosystems can increase the flux of C into and out of soils without changing total C pools, at least over the short term and as long as the native tree canopy remains intact, and that invasion‐mediated changes in belowground C cycling are sensitive to precipitation.     

Ecological Restoration of Native Forest at Aratiatia, North Island, New Zealand

Successional pathways in native forest, planted 15–33 years ago on reconstructed surfaces to restore aesthetic values destroyed by hydro‐electric dam construction at Aratiatia, central North Island, New Zealand, were compared with those on similar surfaces left unplanted. Only native species were planted. Classification identified three canopy communities and several ground layer communities with significant inter‐stratum relationships: Pittosporum tenuifolium‐Sophora tetraptera short forest with ground layers dominated by litter; P. tenuifolium‐Kunzea ericoides short forest over adventive grasses on planted sites; and adventive Cytisus scoparius shrubland over grasses on unplanted sites. Planted communities mirror young secondary forests on intact substrates in the district, but have lower density and similar or higher basal area than such forests elsewhere. Established seedlings of seven planted canopy trees, mostly early successional bird‐dispersed species, are reasonably widespread in floristically rich Pittosporum‐Sophora forest. Seedlings of only two species are widespread in floristically poor Pittosporum‐Kunzea forest, and none on unplanted sites. This first large‐scale attempt at ecological restoration in New Zealand, by mass planting of new surfaces with early successional native woody species, has created aesthetically‐pleasing stands of indigenous forest on sites which would otherwise remain in relatively stable adventive shrubland communities for the foreseeable future. Only continued monitoring will show whether further management is necessary and whether natural processes are operating at a level sufficient to ensure that artificially initiated successions will continue along more or less natural pathways.     

Spatial pattern of soil nitrogen availability and its relationship to stand structure in a coniferous-broadleaved mixed forest with a dense dwarf bamboo understory in northern Japan

Natural disturbances create spatial patterns of the ecosystem processes and functions in natural forests. However, how dynamics and the spatial structure of forests relate to soil nitrogen dynamics is not well understood. We examined the spatial relationship between the distributions of canopy and understory species, and soil nitrogen dynamics in a natural coniferous-broadleaved mixed forest with a dense understory of Sasa dwarf bamboo in northern Japan. The O horizon was thick where coniferous litter predominated, and it was thin where broadleaved litter predominated. The soil water content was low in areas with a thick O horizon and a high abundance of coniferous trees. The soil nitrate content was low where the soil water content was low, and the soil nitrate content increased linearly with increasing net nitrification potential. These results suggest that the soil nitrate content under the coniferous canopy was lower because of the low nitrification potential of soil microbes in soils with low water contents. The soil nitrate content and nitrification potential were higher in the canopy gap than under the canopy. Our results suggest that forest structure, specifically the thickness of the forest floor, significantly affects the spatial pattern of the soil water content, thereby creating a spatial pattern of soil nitrogen availability at a relatively small scale with flat topography. The higher nitrification potential under the canopy gap could pose a long-term risk of nitrate leaching because of the suppression of the natural regeneration of canopy species by dense Sasa dwarf bamboo in this forest ecosystem.     

Understory Plant Community Composition Is Associated with Fine-Scale Above- and Below-Ground Resource Heterogeneity in Mature Lodgepole Pine (Pinus contorta) Forests

Understory plant communities play critical ecological roles in forest ecosystems. Both above- and below-ground ecosystem properties and processes influence these communities but relatively little is known about such effects at fine (i.e., one to several meters within-stand) scales, particularly for forests in which the canopy is dominated by a single species. An improved understanding of these effects is critical for understanding how understory biodiversity is regulated in such forests and for anticipating impacts of changing disturbance regimes. Our primary objective was to examine the patterns of fine-scale variation in understory plant communities and their relationships to above- and below-ground resource and environmental heterogeneity within mature lodgepole pine forests. We assessed composition and diversity of understory vegetation in relation to heterogeneity of both the above-ground (canopy tree density, canopy and tall shrub basal area and cover, downed wood biomass, litter cover) and below-ground (soil nutrient availability, decomposition, forest floor thickness, pH, and phospholipid fatty acids (PLFAs) and multiple carbon-source substrate-induced respiration (MSIR) of the forest floor microbial community) environment. There was notable variation in fine-scale plant community composition; cluster and indicator species analyses of the 24 most commonly occurring understory species distinguished four assemblages, one for which a pioneer forb species had the highest cover levels, and three others that were characterized by different bryophyte species having the highest cover. Constrained ordination (distance-based redundancy analysis) showed that two above-ground (mean tree diameter, litter cover) and eight below-ground (forest floor pH, plant available boron, microbial community composition and function as indicated by MSIR and PLFAs) properties were associated with variation in understory plant community composition. These results provide novel insights into the important ecological associations between understory plant community composition and heterogeneity in ecosystem properties and processes within forests dominated by a single canopy species.     

Root dynamics influence tree–grass coexistence in an Australian savanna

Both resource and disturbance controls have been invoked to explain tree persistence among grasses in savannas. Here we determine the extent to which competition for available resources restricts the rooting depth of both grasses and trees, and how this may influence nutrient cycling under an infrequently burned savanna near Darwin, Australia. We sampled fine roots <2 mm in diameter from 24 soil pits under perennial as well as annual grasses and three levels of canopy cover. The relative proportion of C₃ (trees) and C₄ (grasses) derived carbon in a sample was determined using mass balance calculations. Our results show that regardless of the type of grass both tree and grass roots are concentrated in the top 20 cm of the soil. While trees have greater root production and contribute more fine root biomass grass roots contribute a disproportional amount of nitrogen and carbon to the soil relative to total root biomass. We postulate that grasses maintain soil nutrient pools and provide biomass for regular fires that prevent forest trees from establishing while savanna trees, are important for increasing soil N content, cycling and mineralization rates. We put forward our ideas as a hypothesis of resource‐regulated tree–grass coexistence in tropical savannas.     

Soil ecosystem function under native and exotic plant assemblages as alternative states of successional grasslands

Old fields often become dominated by exotic plants establishing persistent community states. Ecosystem functioning may differ widely between such novel communities and the native-dominated counterparts. We evaluated soil ecosystem attributes in native and exotic (synthetic) grass assemblages established on a newly abandoned field, and in remnants of native grassland in the Inland Pampa, Argentina. We asked whether exotic species alter soil functioning through the quality of the litter they shed or by changing the decomposition environment. Litter decomposition of the exotic dominant Festuca arundinacea in exotic assemblages was faster than that of the native dominant Paspalum quadrifarium in native assemblages and remnant grasslands. Decomposition of a standard litter (Triticum aestivum) was also faster in exotic assemblages than in native assemblages and remnant grasslands. In a common garden, F. arundinacea showed higher decay rates than P. quadrifarium, which reflected the higher N content and lower C:N of the exotic grass litter. Soil respiration rates were higher in the exotic than in the native assemblages and remnant grasslands. Yet there were no significant differences in soil N availability or net N mineralization between exotic and native assemblages. Our results suggest that exotic grass dominance affected ecosystem function by producing a more decomposable leaf litter and by increasing soil decomposer activity. These changes might contribute to the extended dominance of fast-growing exotic grasses during old-field succession. Further, increased organic matter turnover under novel, exotic communities could reduce the carbon storage capacity of the system in the long term.