Effect of nitrogen availability on forest understorey cover and its consequences for tree regeneration in the Austrian limestone Alps

The impact of airborne nitrogen on tree regeneration has attracted increasing interest as deposition loads are by far exceeding natural amounts. Eutrophication may have both direct and indirect effects on the regeneration of forest trees. Here, we focus on potential indirect effects, mediated by nitrogen-dependent patterns in forest understoreys. Using data from a long-term monitoring site in a montane forest ecosystem of the Northern Limestone Alps, we first evaluated correlations between understorey cover of herbs and graminoids and nitrogen availability, as indicated by soil C/N ratios. Then, recruitment and growth rates of the four major tree species of this system (Acer pseudplatanus, Fraxinus excelsior, Fagus sylvatica and Picea abies) were related to understorey cover by means of linear and generalized linear mixed models, taking covariates (soil pH value, radiation, microrelief properties) and spatially varied seed input into account. Nitrogen availability is positively related to graminoid and herb cover. Herb cover had a negative effect on A. pseudoplatanus recruitment, while graminoids had a positive effect on F. sylvatica. Browsing damage on juvenile trees was reduced when graminoid cover was higher We conclude that under specific abiotic and biotic conditions, like on steep, erosion prone slopes, or under high levels of herbivore pressure, understorey vegetation may facilitate tree seedlings rather than compete with them. This highlights the complexity of the factors controlling tree regeneration and the difficulty to predict the net effects of the excess deposition of airborne nitrogen.     

Effects of position, understorey vegetation and coarse woody debris on tree regeneration in two environmentally contrasting forests of north-western Patagonia: a manipulative approach

Aim To investigate the differential effects of position within gaps, coarse woody debris and understorey cover on tree seedling survival in canopy gaps in two old‐growth Nothofagus pumilio (Poepp. & Endl.) Krasser forests and the response of this species to gaps in two forests located at opposite extremes of a steep rainfall gradient. Location Nahuel Huapi National Park, at 41° S in north‐western Patagonia, Argentina. Methods In both study sites, seedlings were transplanted to experimental plots in gaps in three different positions, with two types of substrate (coarse woody debris or forest floor), and with and without removal of understorey vegetation. Survival of seedlings was monitored during two growing seasons. Soil moisture and direct solar radiation were measured once in mid‐summer. Seedling aerial biomass was estimated at the end of the experiment. Results Mid‐summer soil water potential was lowest in the centre of gaps, in plots where the understorey had been removed, and highest at the northern edges of gaps. Direct incoming radiation was highest in gap centres and southern edges, and lowest at northern edges. Seedling mortality was highest in gap centres, in both sites. Coarse woody debris had a positive effect on seedling survival during summer in the mesic forest and during winter in the xeric forest. The removal of understorey cover had negative effects in gap centres during summer. Seedling final aerial biomass was positively affected by understorey removal and by soil substrate in both sites. In the dry forest gaps, seedling growth was highest in northern edges, whereas it was highest in gap centres in the mesic forest. Overall growth was positively related to survival in the xeric forest, and negatively related in the mesic forest. Main conclusions Survival and growth were facilitated by the shade of gap‐surrounding trees only in the xeric forest. Understorey vegetation of both forests facilitated seedling survival in exposed microsites but competed with seedling growth. Nurse logs were an important substrate for seedling establishment in both forests; however, causes of this pattern differed between forests. Water availability positively controls seedling survival and growth in the xeric forest while in the mesic forest, survival and growth are differentially controlled by water and light availability, respectively. These two contrasting old‐growth forests, separated by a relatively short distance along a steep rainfall gradient, had different yet unexpected microenvironmental controls on N. pumilio seedling survival and growth. These results underscore the importance of defining microscale limiting factors of tree recruitment in the context of large‐scale spatial variation in resources.     

Post‐fire understorey regeneration in boreal Pinus sylvestris forest sites with different fire histories

Abstract. We studied the characteristics of understorey regeneration on two sites with different fire history in a mature Pinus sylvestris forest in eastern Finland. The study area was a 4‐ha plot, which was divided into two parts based on fire history analysis. In one part the last fire event was a stand‐replacing fire in the early 19^th century, after which the whole stand regenerated, while the other part of the study plot was subsequently burnt by a surface fire in 1906. Understorey P. sylvestris individuals were much more abundant in the area of the 1906 burn compared to the old burn. In both areas the size frequency distribution of living trees was bimodal, with frequency peaks at the < 5 cm and 30–150 cm height classes. In the old burn small understorey trees were mainly associated with microsites created by treefall disturbances while in the 1906 burn most small understorey trees occurred on vegetation‐covered microsites. This indicates that with increasing time since last fire establishment of new understorey trees becomes more restricted by the availability of microsites created by treefall disturbances. In both areas the proportion of vigorous small understorey trees was highest on decayed wood. In the older burn uprooted pits and mounds also had a significant proportion of healthy small understorey trees, while the majority of trees classified as seriously weakened or dying were growing on microhabitats characterized by undisturbed vegetation. Ripley's K‐function analyses showed that spatial distribution of understorey trees was clustered in both areas in all microsite types and clustering at small scales was most pronounced in understorey trees growing in uprooted spots or in association with decayed wood. The bivariate analysis showed a significant repulsion effect between large trees and understorey trees at intermediate spatial scales, indicating that competition had an effect on understorey tree distribution and this effect was more pronounced in the younger burn. The analysis suggests that in Pinus sylvestris forests the abundance, quality and spatial pattern of understorey tree population may vary considerably as a function of disturbance history.     

The influence of Prosopis canopies on understorey vegetation: Effects of landscape position

Abstract. The influence of canopy trees and shrubs on under‐storey plants is complex and context‐dependent. Canopy plants can exert positive, negative or neutral effects on production, composition and diversity of understorey plant communities, depending on local environmental conditions and position in the landscape. We studied the influence of Prosopis velutina (mesquite) on soil moisture and nitrogen availability, and understorey vegetation along a topographic gradient in the Sonoran Desert. We found significant increases in both soil moisture and N along the gradient from desert to riparian zone. In addition, P. velutina canopies had positive effects, relative to open areas, on soil moisture in the desert, and soil N in both desert and intermediate terrace. Biomass of understorey vegetation was highest and species richness was lowest in the riparian zone. Canopies had a positive effect on biomass in both desert and terrace, and a negative effect on species richness in the terrace. The effect of the canopy depended on landscape position, with desert canopies more strongly influencing soil moisture and biomass and terrace canopies more strongly influencing soil N and species richness. Individual species distributions suggested interspecific variation in response to water‐ vs. N‐availability; they strongly influence species composition at both patch and landscape position levels.     

Effects of fire and selective logging on the tree species composition of lowland dipterocarp forest in East Kalimantan, Indonesia

Tree species composition (diameter at breast height (dbh) 10 cm) was studied in primary, selectively logged and heavily burnt forests in East Kalimantan, Indonesia. The number of trees, tree species, and the Fishers's- diversity index were determined for the first 15 years (burnt forest) and 25 years (selectively logged forest) after disturbance. Additionally the population structure of six common and typical Macaranga pioneer tree species was compared through time between selectively logged, burnt and primary forest. Both selectively logged and burnt forest showed a significant reduction in number of trees and tree species per surface area directly after disturbance. Fire especially affected dominant tree species, while for selective logging the opposite was observed. In selectively logged forest the number of trees, tree species and the Fishers's- index reached pre-disturbance levels within c. 15 years. For burnt forest, only the number of trees recovered to pre-disturbance levels. The number of tree species stayed constant after disturbance, while the Fishers's- index decreased. The six studied Macaranga pioneer tree species seedlings were present in all forest types. Their density seems to be unrelated to light levels in the forest understorey but strongly related to the number of mature parent trees. Their sapling densities were strongly related to light levels in the forest understorey. The studied Macaranga species formed an important part of both under- and over-storey in burnt forest 15 years after disturbance, while they were almost absent in the understorey and only moderately common in the overstorey of selectively logged forest.     

Overstorey Control of Understorey Species Composition in a Near-natural Temperate Broadleaved Forest in Denmark

Little is known about the importance of the forest overstorey relative to other factors in controlling the spatial variability in understorey species composition in near-natural temperate broadleaved forests. We addressed this question for the 19 ha ancient forest Suserup Skov (55°22′ N, 11°34′ E) in Denmark, one of the few old-growth temperate broadleaved forest remnants in north-western Europe, by inventorying understorey species composition and environmental conditions in 163 100 m² plots. We use unconstrained and constrained ordinations, variation partitioning, and Indicator Species Analysis to provide a quantitative assessment of the importance of the forest overstorey in controlling understorey species composition. Comparison of the gradients extracted by unconstrained and constrained ordinations showed that the main gradients in understorey species composition in our old-growth temperate broadleaved forest remnant are not caused by variability in the forest overstorey, but are related to topography and soil, edge effects, and unknown broad-scale factors. Nevertheless, overstorey-related variables uniquely accounted for 15% of the total explained variation in understorey species composition, with the pure overstorey-related (R_po), topography and soil (R_pt), edge and anthropogenic disturbance effects (R_pa), and spatial (R_ps) variation fractions being of equal magnitude. The forward variable selection showed that among the overstorey-related variables understorey light availability and to a lesser extent vertical forest structure were most important for understorey species composition. No unique influence of overstorey tree species identity could be documented. There were many indicator species for high understorey light levels and canopy gap centres, but none for medium or low light or closed canopy. Hence, no understorey species behaved as obligate shade plants. Our study shows that, the forest overstorey has a weak control of understorey species composition in near-natural broadleaved forest, in contrast to results from natural and managed forests comprising both conifer and broadleaved species. Nevertheless, >20% of the understorey species found were indicators of high light conditions or canopy openings. Hence, variability in canopy structure and understorey light availability is important for maintaining understorey species diversity.     

Tree invasion in managed tropical forests facilitates endemic species

Aim To determine whether different abundances of introduced species of Cinchona (Rubiaceae) affect species composition and facilitate species richness in managed tropical forests, to test whether any facilitative effects on understorey species depend on forest type, and to investigate whether facilitative effects can be attributed to the ‘substitutive facilitation model’. Location Makawao Forest Reserve on Maui, Hawai’i, USA. Methods Cinchona species (Cinchona pubescens and Cinchona calisaya) were mapped within various forest types. In three forest types (ageing Eucalyptus and Pinus plantations, and near‐natural Acacia koa forests), we analysed environmental parameters (e.g. canopy cover, litter cover, pH value and soil depth) and the species composition of Cinchona‐invaded and non‐invaded plots; data were compared based on Cinchona cover and forest types. Habitat modelling for several endemic species and tree ferns was carried out to test whether Cinchona cover is an important variable for the probability of occurrence of these endemics. Results Cinchona species have naturalized mainly in Eucalyptus and Pinus plantations and Acacia koa forests and here add an additional shrub layer. In contrast to other studies, we revealed facilitative effects of Cinchona on native species within all forest types. Species richness is about 20% higher in invaded plots than in non‐invaded plots, and these show a nearly 50% higher proportion of endemic species, including tree ferns. The proportion of endemics even increases with increasing Cinchona cover. For several endemics, Cinchona is found to be an important variable for the probability of occurrence, and the removal of Cinchona cover as an explanatory variable lowers the model fit. In addition to Cinchona, variables delineating vegetation structure and light availability have a strong effect on the model fit. Main conclusions In the structurally simplified Hawaiian forests studied, Cinchona facilitated endemic species in accordance with the ‘substitutive facilitation model’. This contrasts with the results of an earlier study in the naturally treeless Galápagos highlands, which revealed a sharp decrease in the abundance of endemics under Cinchona canopy. These results illustrate that, through the same structural change (addition of a vegetation layer), an invasive species may exert divergent effects across different ecosystem types. The facilitation of endemic understorey species by invasive tree species in managed forests leads to a dilemma in conservation but also to new perspectives for ecosystem restoration.     

Variations in resistance to canopy disturbances and their interactions with the spatial structure of major species in a cool‐temperate forest

Question: How does typhoon‐related disturbance (more specifically, disturbance in the understorey due to tree‐fall and branch‐fall) affect different species mortality rates in a vertically well‐structured forest community? Location: Cool‐temperate, old‐growth forest in the Daisen Forest Reserve, Japan. Methods: We investigated the canopy dynamics and mortality rate trends of trees ≥5 cm diameter at breast height in a 4‐ha study plot, and analysed the effects of tree diameter and spatial structure on the mortality risks for major tree species in the understorey. Results: Significant differences were found in the mortality rates and proportions of injured dead stems between census periods, which were more pronounced in the understorey than in the canopy. Acer micranthum, which showed increased mortality during typhoon disturbance periods, had a clumped distribution. In contrast, Acer japonicum and Viburnum furcatum, which showed similar mortality rates between census periods, had a loosely clumped spatial distribution and a negative association with canopy trees, respectively. In the understorey stems of Acanthopanax sciadophylloides and Fagus crenata, whose spatial distribution patterns depended on canopy gaps, significant increases in mortality rates were observed only during severe typhoon‐related disturbance periods. Conclusions: The sensitivity of trees to typhoon‐related canopy disturbance is more pronounced in the lower layers of vertically structured forest communities. Differences in mortality patterns generated through the combined effects of spatial variation in disturbance regime and species‐specific spatial distribution patterns (spatial aggregation, association with canopy trees, and canopy gap dependency) contribute to the co‐existence of understorey species in forest communities that are subject to typhoon‐related disturbance.     

Response of floodplain understorey species to environmental gradients and tree invasion: a functional trait perspective

Plants are connected to habitats by functional traits which are filtered by environmental gradients. Since tree species composition in the forest canopy can influence ecosystem processes by changing resource availability, litter accumulation, and soil nutrient content, we hypothesised that non-native invasive trees can establish new environmental filters on the understorey communities. In the hardwood floodplain forests in Northern Italy, the invasive trees Robinia pseudoacacia L. and Prunus serotina Ehrh. are the dominant canopy species. We used trait data assembled from databases and iterative RLQ analysis to identify a parsimonious set of functional traits responding to environmental variables (soil, light availability, disturbance, and stand structure) and the dominant native and invasive canopy species. Then, RLQ and fourth-corner analysis was conducted to investigate the joint structure between macro-environmental variables and species traits and functional groups were identified. The trait composition of the herb-layer was significantly related to the main environmental gradients and the presence of the invaders in the canopy showed significant relationships with several traits. In particular, the presence of P. serotina may mitigate or even erase the effect of disturbances, maintaining a stable forest microclimate and thus favouring ‘true’ forest species, while R. pseudoacacia may slow down forest succession and regeneration by establishing new stable associations with a graminoid-dominated understorey. The impact of the two invasive trees on herb layer composition appears to differ, indicating that different management and control strategies may be needed.     

Understorey gaps influence regeneration dynamics in subtropical coastal dune forest

Dominant understorey species influence forest dynamics by preventing tree regeneration at the seedling stage. We examined factors driving the spatial distribution of the monocarpic species Isoglossa woodii, a dominant understorey herb in coastal dune forests, and the effect that its cover has on forest regeneration. We used line transects to quantify the area of the forest understorey with I. woodii cover and with gaps in the cover. Paired experimental plots were established in semi-permanent understorey gaps with I. woodii naturally absent and in adjacent areas with I. woodii present to compare plant community composition, soil, and light availability between the two habitats. Isoglossa woodii was widespread, covering 65–95% of the understorey, while gaps covered the remaining 5–35% of the area. The spatial distribution of this species was strongly related to tree canopy structure, with I.␣woodii excluded from sites with dense tree cover. Seedling establishment was inhibited by low light availability (<1% of PAR) beneath I.␣woodii. When present, I. woodii reduced the density and species richness of tree seedlings. The tree seedling community beneath I. woodii represented a subset of the seedling community in gaps. Some species that were found in gaps did not occur beneath I. woodii at all. There were no significant differences between the sapling and canopy tree communities in areas with I. woodii gaps and cover. In the coastal dune forest system, seedling survival under I. woodii is dependent on a species’ shade tolerance, its ability to grow quickly during I. woodii dieback, and/or the capacity to regenerate by re-sprouting and multi-stemming. We propose a general conceptual model of forest regeneration dynamics in which the abundant understorey species, I. woodii, limits local tree seedling establishment and survival but gaps in the understorey maintain tree species diversity on a landscape scale.     

Structural heterogeneity and tree spatial patterns in an old-growth deciduous lowland forest in Cantabria, northern Spain

Old-growth deciduous forests in western Europe, for the most part, consist of small tracts that often may be atypical due to human disturbance, poor soil productivity or inaccessibility. In addition, very little information on tree age distributions, structural heterogeneity and tree spatial patterns appears to be available for west-European forests. Characterization of the structural features of tree populations in these old-growth stands can provide the basis to design conservation plans and also inform on how present forests might look in the absence of human interference. Four old-growth stands in a deciduous forest in the Cantabrian lowlands, northern Spain, were surveyed to determine forest structure and spatial patterns. Live and dead trees were identified, measured and mapped, and live trees were cored for age estimation. Structural heterogeneity was analyzed by means of the spatial autocorrelation of tree diameter, height and age, and the uni- and bivariate spatial patterns of trees were analyzed. The dominant species, Fagus sylvatica and Quercus robur, showed reverse-J shaped size distributions but discontinuous age distributions, with maximum ages of 255–270 yr. Tree ages suggested that the forest was largely modified by past changes in forest-use, especially by temporal variation in grazing intensity. Spatial autocorrelation revealed that former parkland stands were heterogeneous with respect to tree height only, while high forest stands were composed of patches of even-aged and even-sized trees. Young trees were clumped at varying distances and establishment occurred preferentially in canopy gaps, except for Ilex aquifolium that mainly occurred beneath mature Quercus trees. Surviving trees became less intensely clumped in the dominant species, and more strongly clumped in understorey ones, which may have been due to the effects of intraspecific competition and of canopy trees on tree survival, respectively. The spatial associations between species varied within the forest, probably as a consequence of specific establishment preferences and competitive interactions.     

Influence of conspecific and heterospecific adults on riparian tree species establishment during encroachment of a humid palm savanna

Woody plant encroachment of savanna ecosystems has been related to altered disturbance regimes, mainly fire suppression and herbivore exclusion. In contrast, neighbourhood interactions among resident and colonising woody species have received little attention, despite their likely influence on the pattern and rate of tree establishment. We examined how resident palm trees (Butia yatay) and established adults of two riparian forest tree species (Allophylus edulis and Sebastiania commersoniana) influenced seed arrival and seedling performance of the latter two species in a humid savanna of east-central Argentina. Seed traps and seedlings of both riparian species were placed in herbaceous openings, and beneath palm, conspecific and heterospecific adult trees in two unburned savanna patches, and were monitored for 2 years. Only seeds of the bird-dispersed Allophylus arrived in palm microsites, yet survival of Allophylus seedlings near adult palms was limited by animal damage through trampling and burrowing, a non-trophic mechanism of apparent competition. Seeds of both riparian species dispersed into conspecific microsites, although adult trees selectively reduced growth of conspecific seedlings, a pattern consistent with the “escape hypothesis”. Further, survival of Sebastiania increased in the moister Allophylus microsites, suggesting a one-way facilitative interaction between woody colonisers. Our results indicate that dispersal facilitation by resident savanna trees may be critical to riparian species invasion after fire suppression. Distance-dependent effects of conspecific and heterospecific adult trees could contribute to shape the subsequent dynamics of woody seedling establishment. Overall, we show that indirect interactions can play a prominent role in savanna encroachment by non-resident woody species.     

Intraspecific variability of specific leaf area fosters the persistence of understorey specialists across a light availability gradient

• Forest understorey plants are sensitive to light availability, and different species groups can respond differently to changing light conditions. A plant trait tightly linked to light capture is specific leaf area (SLA). Studies considering the relative role of within‐ and among‐species SLA variation across different species groups (e.g. specialists and generalists) are rarely implemented in temperate forest understories varying in their maturity.
• We examined community‐level SLA patterns of beech forest understories along a light availability gradient, and for habitat specialists and generalists separately. We then disentangled and quantified the contribution of intraspecific trait variability and interspecific trait differences in shaping SLA patterns.
• We revealed that the increase in community‐level SLA with decreasing light availability was primarily driven by beech forest specialists (and, to a lesser extent, by forest generalists), and this pattern was mainly determined by specialists’ high intraspecific variability. Community‐level SLA was therefore formed by different responses at different organizational levels, i.e. within and among species, and for separate species groups.
• This study provides insights into factors shaping the shade tolerance strategy in beech forest understorey plants; specialists persistence under putative less favourable conditions (i.e. high irradiation) may be fostered by their ability to adjust their light capture strategies intraspecifically.

     

Crown structure and leaf area of the understorey species Prunus serotina

A detailed biometrical study of the exotic understorey invader Prunus serotina (Ehrh.) was carried out in a mixed coniferous forest stand in northern Belgium. Based on detailed destructive measurements of eight selected model trees, allometric relations of tree height, crown projected area, woody and leaf dry mass and leaf area on tree diameter at breast height (DBH) were derived. The scaling-up procedure from the tree to the stand level was done using the frequency distribution of DBH obtained at the selected experimental plot. The vertical and radial distributions of the tree foliage were estimated by the “cloud” technique. The vertical profile of leaf area showed a bimodal distribution pattern with maxima at heights of 4 and 6 m above the ground. The leaf area index (LAI) of the understorey Prunus serotina as estimated by the described up-scaling procedure (5.1) was significantly higher than the LAI (2.6) as measured by a plant canopy analyser and was also significantly higher than the LAI of the overstorey species Scots pine (1.5–3.0). The LAI of a neighbouring Rhododendron understorey reached only 1.25. This study emphasises the importance of an exotic understorey species in the total leaf area of mixed coniferous forests which might have important implications for the energy and mass exchanges of the entire forest.     

Spatial pattern and process in forest stands within the Virginia piedmont

Abstract. Question: Underlying ecological processes have often been inferred from the analysis of spatial patterns in ecosystems. Using an individual‐based model, we evaluate whether basic assumptions of species’life‐history, drought‐susceptibility, and shade tolerance generate dynamics that replicate patterns between and within forest stands. Location: Virginia piedmont, USA. Method: Model verification examines the transition in forest composition and stand structure between mesic, intermediate and xeric sites. At each site, tree location, diameter, and status were recorded in square plots ranging from 0.25 to 1.0 ha. Model validation examines the simulated spatial pattern of individual trees at scales of 1–25 m within each forest site using a univariate Ripley's K function. Results: 7512 live and dead trees were surveyed across all sites. All sites exhibit a consistent, significant shift in pattern for live trees by size, progressing from a clumped understorey (trees ± 0.1 m in diameter) to a uniform overstorey (trees > 0.25 m). Simulation results reflect not only the general shift in pattern of trees at appropriate scales within sites, but also the general transition in species composition and stand structure between sites. Conclusions: This shift has been observed in other forest ecosystems and interpreted as a result of competition; however, this hypothesis has seldom been evaluated using simulation models. These results support the hypothesis that forest pattern in the Virginia piedmont results from competition involving species’life‐history attributes driven by soil moisture availability between sites and light availability within sites.     

El Niño droughts and their effects on tree species composition and diversity in tropical rain forests

In this study I investigated the effects of the extreme, 1997/98 El Niño related drought on tree mortality and understorey light conditions of logged and unlogged tropical rain forest in the Indonesian province of East Kalimantan (Borneo). My objectives were to test (1) whether drought had a significant effect on tree mortality and understorey light conditions, (2) whether this effect was greater in logged than in undisturbed forest, (3) if the expected change in tree mortality and light conditions had an effect on Macaranga pioneer seedling and sapling densities, and (4) which (a)biotic factors influenced tree mortality during the drought. The 1997/1998 drought led to an additional tree mortality of 11.2, 18.1, and 22.7% in undisturbed, old logged and recently logged forest, respectively. Mortality was highest in logged forests, due to extremely high mortality of pioneer Macaranga trees (65.4%). Canopy openness was significantly higher during the drought than during the non-drought year (6.0, 8.6 and 10.4 vs 3.7, 3.8 and 3.7 in undisturbed, old logged and recently logged forest, respectively) and was positively correlated with the number of dead standing trees. The increase in light in the understorey was accompanied by a 30 to 300-fold increase in pioneer Macaranga seedling densities. Factors affecting tree mortality during drought were (1) tree species successional status, (2) tree size, and (3) tree location with respect to soil moisture. Tree density and basal area per surface unit had no influence on tree mortality during drought. The results of this study show that extreme droughts, such as those associated with El Niño events, can affect the tree species composition and diversity of tropical forests in two ways: (1) by disproportionate mortality of certain tree species groups and tree size classes, and (2) by changing the light environment in the forest understorey, thereby affecting the recruitment and growth conditions of small and immature trees.     

Establishment of Perennial Shrub and Tree Species in Degraded Eucalyptus salmonophloia (Salmon Gum) Remnant Woodlands: Effects of Restoration Treatments

Woodlands dominated by Eucalyptus salmonophloia (salmon gum) occur throughout the fragmented landscape of the southwestern Australian wheatbelt. These remnants are often degraded by livestock grazing and weed invasion and in many cases there is little or no understorey remaining and little or no regeneration of the dominant tree E. salmonophloia. There is a growing interest in developing techniques for restoring remnant woodlands. This study describes techniques for establishing seedlings of the dominant tree and perennial understorey species in E. salmonophloia (salmon gum) woodlands degraded by livestock grazing. The study tests the hypothesis that, in addition to the exclusion of livestock, management of weeds and reintroduction of plant species, restoration of plant species diversity will require techniques which mimic large‐scale disturbances, reduce soil compaction, and restore soil water infiltration to suitable rates. Five‐month‐old seedlings of the dominant tree E. salmonophloia and four commonly associated woody shrubs (Acacia hemiteles, Atriplex semibaccata, Maireana brevifolia, and Melaleuca pauperiflora) were planted into areas that differed with respect to grazing (–rabbit/ ?livestock and +rabbit/–livestock), tree canopy disturbance (+/–competition with tree canopy) and amelioration of soil compaction (+/–deep ripping). Following three growing seasons and two summers, the exclusion of rabbits had no significant effect on the survival and growth of planted species. As a consequence grazing treatments are pooled for the purposes of presenting the impacts of removing competition with adult trees and soil deep ripping. The removal of competition with adult E. salmonophloia trees significantly improved the survival of E. salmonophloia seedlings but did not improve survival of understorey species. Deep ripping the soil significantly improved the survival of both E. salmonophloia and the shrub A. hemiteles but did not improve the survival of other understorey species. In contrast to seedling survival, the removal of adult E. salmonophloia trees and deep ripping soil significantly increased the growth of all species. The results indicate that increasing levels of intervention will increase the chances of successfully restoring tree and understorey species diversity in degraded E. salmonophloia woodlands.     

Effect of forest canopy composition on soil nutrients and dynamics of the understorey: mixed canopies serve neither vascular nor bryophyte strata

Question: The effect of overstorey composition on above‐ground dynamics of understorey vegetation is poorly understood. This study examines the understorey biomass, production and turnover rates of vascular and non‐vascular plants along a conifer–broadleaf gradient of resource availability and heterogeneity. Location: Canadian boreal forests of northwest Quebec and Ontario. Methods: We sampled mature stands containing various proportions of black spruce (Picea mariana (Mill.) BSP), trembling aspen (Populus tremuloides Michx.) and jack pine (Pinus banksiana Lamb.). Above‐ground biomass of the understorey vegetation was assessed through harvesting; annual growth rates were calculated as the differences between biomass in 2007 and 2008, as estimated by allometric relationships, and turnover rates were estimated as net primary production divided by the biomass in 2007. Results: Higher aspen presence, linked to greater nutrient availability in the forest floor, was generally associated with higher vascular biomass and production in the understorey. This effect was less pronounced in sites of high intrinsic fertility. In contrast, bryophyte biomass was positively associated with conifer abundance, particularly in wet sites of the Quebec study area. Non‐linear responses resulted in total understorey biomass being lower under mixed canopies than under pure aspen or pure conifer canopies. Turnover rates did not differ with overstorey composition. Conclusions: While resource availability is a main driver of understorey productivity, resources as drivers appear to differ with differences in understorey strata components, i.e. vascular versus non‐vascular plants. Resource heterogeneity induced by a mixed canopy had overall negative effects on understorey above‐ground productivity, as this productivity seemed to rely on species adapted to the specific conditions induced by a pure canopy.     

Densities of Two Frugivorous Primates with Respect to Forest and Fragment Tree Species Composition and Fruit Availability

Conservation of wildlife populations requires extensive knowledge of their habitat requirements, efficient methods to evaluate habitat quality, and an understanding of the value of fragments and edges. Kibale National Park, Uganda has areas that differ in the densities of 2 species of frugivorous monkeys—Cercopithecus mitis and Lophocebus albigena—including one on an edge and forest fragments outside the park that lack both species. We compared the basal area densities of important food trees with primate densities. The density of Cercopithecus mitis correlates most strongly with the basal area density of all types of food trees combined. The density of Lophocebus albigena does not correlate with the basal area densities of any category of food trees or with fruit availability. An index of their density—number of groups seen per km walked—correlates to fruit availability but with marginal significance. Lack of a relationship between the basal area densities of food trees and density of Lophocebus albigena may be the result of a mismatch in scale between the forest area measured and their large home ranges. We compared the unused area of forest to the other areas of the forest and the fragments and found it had higher basal area densities in all food tree categories for both species than the fragments and lower basal area densities of most categories than the other parts of the forest, indicating that the fragments are poor quality and would probably be unused even if dispersal were likely.     

Effect of legume understorey on decomposition and nutrient content of eucalypt forest litter

Summary In Jarrah (Eucalyptus marginata Donn ex Sm.) forest of south-western Australia dense germination and regeneration of the native legumeAcacia Pulchella R. Br. can occur following moderate to high intensity fire. The effect of this legume understorey on rate of decomposition and change in nutrient content ofE. marginata litter was investigated using the mesh bag techniques and by examining four components of forest floor litter representing increasing stages of decomposition. E. marginata leaf litter confined in mesh bags lost 37% of its initial dry weight in the first 8 months on the forest floor and 44% of its initial dry weight after 20 months. During this period weight loss was similar for leaf litter located in forest without legume understorey and for leaf litter placed under dense stands ofA. pulchella. MixingA. pulchella litter withE. marginata litter had no significant effect on rate ofE. marginata litter breakdown. The presence of understorey vegetation had a marked effect on chemical composition of decomposingE. marginata leaves. After 8 and 20 months exposure on the forest floor, leaf litter in mesh bags placed underA. pulchella understorey had significantly (P<0.001) higher concentration and contained significantly (P<0.001) greater amounts of N, P, K, S, Ca and Mg than leaf litter placed in areas without legume understorey. This effect was particularly marked for N and P. In forest without legume understorey the amounts of these two nutrients inE. marginata leaf litter changed little during the first 20 months of decomposition, but forE. marginata leaf litter in mesh bags underA. pulchella there were absolute gains of up to 68% in the amount of N and 109% in the amount of P during this period. This represents accumulation of N and P from sources outside the litter bags. The concentration of N, P, S, Ca and Mg were higher at each of the four stages of decomposition in eucalypt leaf litter collected from the forest floor beneathA. pulchella compared to eucalypt leaf litter collected in forest without understorey. Concentrations of N, P and S increased with stage of decomposition. Levels of these three nutrients in eucalypt litter from under the legume were 1.5 to 2.9 fold higher than in the same component of litter from forest without understorey. The effect of legume understorey on nutrient concentrations in the forest floor and on Cielement ratios in decomposing litter is discussed in relation to long term rates of litter breakdown and net mineralisation of litter nutrients.