Hollow occurrence and abundance varies with tree characteristics and among species in temperate woodland Eucalyptus

Tree hollows are a critical but diminishing resource for a wide range of fauna around the world. Conservation of these fauna depends on sustainable management of tree species that produce the hollows on which they depend. This study addressed the need for empirical data about intraspecific and interspecific variation in hollow occurrence and abundance in woodland trees in Australia. We measured and performed hollow surveys on 1817 trees of seven species of woodland Eucalyptus in central‐western New South Wales, Australia. Trees were surveyed at 51 one‐hectare sites and about 30% of trees surveyed had multiple stems. Generalized linear mixed models that accounted for nestedness of stems within trees and trees within sites detected a significant amount of variation in hollow occurrence and abundance. Models for individual tree stems of live trees showed hollow probability and abundance increased with diameter at breast height (DBH) and with increasing senescence (form). Stems of Eucalyptus microcarpa Maiden had a higher probability of having hollows than similar DBH stems of Eucalyptus camaldulensis Dehnh., Eucalyptus melliodora A.Cunn. ex Schauer or Eucalyptus populnea ssp. bimbil L.A.S.Johnson & K.D.Hill. Dead stems in live trees were more likely to have hollows than live stems of similar DBH. Each stem in a multi‐stemmed tree had a lower probability of hollow occurrence and lower abundance of hollows than single‐stemmed trees of similar DBH. For stems of dead trees, hollow occurrence and abundance increased with DBH and differed depending on stage of senescence. A comparison of our data with other studies indicates regional variation of hollow abundances within tree species.     

Overstorey and juvenile response to thinning and drought in a jarrah (Eucalyptus marginata Donn ex Sm.) forest of southwestern Australia

Aims

Forest thinning is expected to affect tree water use and carbon assimilation, but the related influence from climate variability is little known. Recent forest thinning in the Wungong catchment coincided with a record dry year following the thinning, which provides a rare opportunity to understand the climate influence on the thinning effect.

Methods

A field experiment was conducted to examine changes before and after thinning, especially the rainfall, soil moisture, leaf water status, tissue isotope signature (¹³?C and ¹⁵?N) and N concentration of overstorey and understorey juvenile trees of Eucalyptus marginata (Donn ex Sm.).

Results

Despite the post-thinning drought, surface soil was moister and juvenile jarrah plants were less water stressed, attributable to reduced rain interception and transpiration as a result of less canopy cover. The overstorey was under stress but mainly due to drought rather than by thinning. The concentration of N declined in both tree stems and juvenile leaves along with available N in soil, suggesting a soil N limitation. No treatment effects were detected from leaf relative water content and tissue isotope signature (¹³?C and ¹⁵?N).

Conclusions

The drought effects were superimposed over the thinning effects on overstorey growth, with stemwood δ¹³C being a major indicator of water stress. The water relations and carbon assimilation of understorey juveniles were however dependent more on topsoil moisture, and the wetter soil during the year following thinning enhanced growth activity and hence the depletion of ¹³?C (more negative δ¹³C) in juvenile leaves.     

Digestive plasticity as a response to woodland fragmentation in roe deer

Digestive plasticity, which refers to changes in digestive features due to changes in both internal and external environmental conditions, is a crucial factor for understanding the ability of species to cope with environmental changes. In Europe, agricultural intensification and the loss of forests have been major challenges for original forest dwellers, however some species, such as the European roe deer (Capreolus capreolus), have been able to successfully colonize these new habitats. In this work, we investigated the adaptation of some digestive features of roe deer to the agricultural landscapes. We assessed whether changes in local landscape structure influenced the mass of both reticulorumen (RR) and distal fermentation chamber (DFC) of 47 juvenile and adult roe deer inhabiting an agro-ecosystem in southwest France. Woodland cover had a clear effect on diet quality (estimated by the rate of gas production of digestive contents) and DFC weight of animals. In fact, deer from the most forested landscapes showed heavier DFCs and fed on poorer quality diet (lower gas production) than their counterparts from the most open landscapes. RR mass was less influenced by the landscape openness, being the age of animals the main factor for understanding the variations of this digestive feature in our study area. We can conclude that colonizing agricultural landscapes increases the access to highly energetic and digestive resources.     

Few Mendelian genes underlie the quantitative response of a forest tree, Eucalyptus globulus, to a natural fungal epidemic

Foliar fungal pathogens from the genus Mycosphaerella affect eucalyptus in natural forests and plantations worldwide. QTL analysis was conducted to dissect the genetic control of resistance in Eucalyptus globulus to a natural infection by Mycosphaerella leaf disease, using a clonally replicated outbred F2 family (112 genotypes) planted in a field trial. Two major QTL, with high LOD support (20.2 and 10.9) and high genomewide significance, explained a large proportion (52%) of the phenotypic variance in the severity of damage by Mycosphaerella cryptica, which may be indicative of oligogenic control. Both QTL were validated in a second F2 family and one was validated in a third F2 family. The mean values of different genotype classes at both major QTL argue for Mendelian inheritance with resistance dominant over susceptibility. There were strong correlations between the levels of Mycosphaerella damage in related genetic material planted in three widely separated locations in Tasmania. These findings together provide evidence that the genes controlling resistance to Mycosphaerella damage are stable in different genetic backgrounds and across different environments.     

Landscape drivers of genomic diversity and divergence in woodland Eucalyptus

Spatial genetic patterns are influenced by numerous factors, and they can vary even among coexisting, closely related species due to differences in dispersal and selection. Eucalyptus (L'Héritier 1789; the “eucalypts”) are foundation tree species that provide essential habitat and modulate ecosystem services throughout Australia. Here we present a study of landscape genomic variation in two woodland eucalypt species, using whole‐genome sequencing of 388 individuals of Eucalyptus albens and Eucalyptus sideroxylon. We found exceptionally high genetic diversity (π ≈ 0.05) and low genome‐wide, interspecific differentiation (F_ST = 0.15) and intraspecific differentiation between localities (F_ST ≈ 0.01–0.02). We found no support for strong, discrete population structure, but found substantial support for isolation by geographic distance (IBD) in both species. Using generalized dissimilarity modelling, we identified additional isolation by environment (IBE). Eucalyptus albens showed moderate IBD, and environmental variables have a small but significant amount of additional predictive power (i.e. IBE). Eucalyptus sideroxylon showed much stronger IBD and moderate IBE. These results highlight the vast adaptive potential of these species and set the stage for testing evolutionary hypotheses of interspecific adaptive differentiation across environments.     

Spatial distribution of fine root biomass in a remnant Eucalyptus tereticornis woodland in Eastern Australia

Plant Ecology - In forests, the majority of fine roots are located within the upper soil horizons, and fine root biomass decreases with depth. We evaluated spatial patterns in the distribution of...     

Demographic response to shading and defoliation in two woodland orchids

Many woodland herbs are long-lived, clonal geophytes that have evolved life histories favoring survival over reproduction. We examined the life history responses of natural populations of two woodland orchid species,Cypripedium calceolus andCephalanthera longifolia to defoliation and heavy shading conducted early in the growing seasons of 2002 and 2003. We asked whether, in view of the importance of growth for the survival of geophytes, treated plants were more likely to exhibit reduced flowering than reduced vegetative growth in the seasons following treatment. We also asked whether plants would suffer reduced ramet performance. Both treatments led to significant declines in flower number per ramet, number of leaves per ramet, and mean ramet height relative to controls inCypripedium. However, inCephalanthera, only shaded plants exhibited significant declines in flower number per ramet, and only defoliated plants exhibited declines in mean ramet height. The number of ramets per plant did not decline relative to controls in either species. Thus, these orchids, especiallyCypripedium, appeared to allocate resources preferentially to vegetative growth functions over sexual reproduction. Per-plant variation in leaf and flower number per ramet, as well as in mean ramet height, consistently declined in response to treatment, significantly so in the case of mean ramet height, suggesting that ramets became more similar within genets. These results suggest both similarities and differences in the ways in whichCephalanthera andCypripedium mobilize resources in response to stress.     

Interpreting tree responses to thinning and fertilization using tree-ring stable isotopes

? Carbon sequestration has focused renewed interest in understanding how forest management affects forest carbon gain over timescales of decades, and yet details of the physiological mechanisms over decades are often lacking for understanding long-term growth responses to management. ? Here, we examined tree-ring growth patterns and stable isotopes of cellulose (δ(13)C(cell) and δ(18)O(cell)) in a thinning and fertilization controlled experiment where growth increased substantially in response to treatments to elucidate physiological data and to test the dual isotope approach for uses in other locations. ? δ(13)C(cell) and δ(18)O(cell) results indicated that fertilization caused an increase in intrinsic water-use efficiency through increases in photosynthesis (A) for the first 3 yr. The combination treatment caused a much larger increase in A and water-use efficiency. Only the thinning treatments showed consistent significant increases in δ(18)O(cell) above controls. Changes in canopy microclimate are the likely drivers for δ(18)O(cell) increases with decreases in relative humidity and increases in leaf temperature associated with thinning being the most probable causes. ? Tree-ring isotopic records, particularly δ(13)C(cell), remain a viable way to reconstruct long-term physiological mechanisms affecting tree carbon gain in response to management and climate fluctuations.     

The response of sap flow to pulses of rain in a temperate Australian woodland

In water-limited systems, pulses of rainfall can trigger a cascade of plant physiological responses. However, the timing and size of the physiological response can vary depending on plant and environmental characteristics, such as rooting depth, plant size, rainfall amount, or antecedent soil moisture. We investigated the influence of pulses of rainfall on the response of sap flow of two dominant evergreen tree species, Eucalyptus crebra (a broadleaf) and Callitris glaucophylla (a needle leaved tree), in a remnant open woodland in eastern Australia. Sap flow data were collected using heat-pulse sensors installed in six trees of each species over a 2 year period which encompassed the tail-end of a widespread drought. Our objectives were to estimate the magnitude that a rainfall pulse had to exceed to increase tree water use (i.e., define the threshold response), and to determine how tree and environmental factors influenced the increase in tree water use following a rainfall pulse. We used data filtering techniques to isolate rainfall pulses, and analysed the resulting data with multivariate statistical analysis. We found that rainfall pulses less than 20 mm did not significantly increase tree water use (P > 0.05). Using partial regression analysis to hold all other variables constant, we determined that the size of the rain event (P < 0.05, R ² = 0.59), antecedent soil moisture (P < 0.05, R ² = 0.29), and tree size (DBH, cm, P < 0.05, R ² = 0.15), all significantly affected the response to rainfall. Our results suggest that the conceptual Threshold-Delay model describing physiological responses to rainfall pulses could be modified to include these factors. We further conclude that modelling of stand water use over an annual cycle could be improved by incorporating the T-D behaviour of tree transpiration. Responsible Editor: Stephen S.O. Burgess     

Effects of bird predation on canopy arthropods in wandoo Eucalyptus wandoo woodland

Abstract: As top predators, birds may have significant effects on arthropod abundances and affect the trophic structure of arthropod communities through predation of lower order predators (e.g. spiders) and by competition for prey. We investigated the effects of bird predation on canopy arthropods in south‐western Australia by using plastic bird mesh to exclude insectivorous birds from the foliage of wandoo Eucalyptus wandoo saplings. Exclosure resulted in an increase in the number of herbivorous and predatory arthropods. Total arthropods (with and without ants), spiders, adult Coleoptera, and larval Lepidoptera were significantly more abundant on meshed than unmeshed saplings. All size‐classes of arthropods, taxa grouped, were more abundant on meshed than unmeshed saplings, but with no evidence of a disproportionate increase of the largest arthropods on meshed saplings. All size‐classes of spiders increased in abundance on saplings from which birds were excluded. There were significant differences in the total abundance of arthropods (with and without ants), spiders (Araneae), sucking bugs (Homoptera), adult beetles (Coleoptera), larval moths (Lepidoptera), and wasps and ants (Hymenoptera) for both unmeshed and meshed saplings between sample periods. These seasonal patterns of abundance and differences between sample periods appeared to be determined by seasonal weather patterns, with the lowest numbers associated with drier and hotter conditions in summer and autumn than in winter and spring. The conclusion reached is that eucalypt forest birds have limited effects on temporal variation in canopy arthropod abundances, but depress abundances, and affect the size and trophic composition of the fauna. Given the cascading effects of birds as predators on arthropods, successful conservation management of eucalypt ecosystems, including plantations and revegetation, should be planned to maximize bird numbers and diversity.     

Soil–atmosphere exchange of greenhouse gases in a Eucalyptus marginata woodland, a clover-grass pasture, and Pinus radiata and Eucalyptus globulus plantations

Soils provide the largest terrestrial carbon store, the largest atmospheric CO₂ source, the largest terrestrial N₂O source and the largest terrestrial CH₄ sink, as mediated through root and soil microbial processes. A change in land use or management can alter these soil processes such that net greenhouse gas exchange may increase or decrease. We measured soil–atmosphere exchange of CO₂, N₂O and CH₄ in four adjacent land‐use systems (native eucalypt woodland, clover‐grass pasture, Pinus radiata and Eucalyptus globulus plantation) for short, but continuous, periods between October 2005 and June 2006 using an automated trace gas measurement system near Albany in southwest Western Australia. Mean N₂O emission in the pasture was 26.6 μg N m⁻² h⁻¹, significantly greater than in the natural and managed forests (< 2.0 μg N m⁻² h⁻¹). N₂O emission from pasture soil increased after rainfall events (up to 100 μg N m⁻² h⁻¹) and as soil water content increased into winter, whereas no soil water response was detected in the forest systems. Gross nitrification through ¹⁵N isotope dilution in all land‐use systems was small at water holding capacity < 30%, and under optimum soil water conditions gross nitrification ranged between < 0.1 and 1.0 mg N kg⁻¹ h⁻¹, being least in the native woodland/eucalypt plantation < pine plantation < pasture. Forest soils were a constant CH₄ sink, up to −20 μg C m⁻² h⁻¹ in the native woodland. Pasture soil was an occasional CH₄ source, but weak CH₄ sink overall (−3 μg C m⁻² h⁻¹). There were no strong correlations (R < 0.4) between CH₄ flux and soil moisture or temperature. Soil CO₂ emissions (35–55 mg C m⁻² h⁻¹) correlated with soil water content (R < 0.5) in all but the E. globulus plantation. Soil N₂O emissions from improved pastures can be considerable and comparable with intensively managed, irrigated and fertilised dairy pastures. In all land uses, soil N₂O emissions exceeded soil CH₄ uptake on a carbon dioxide equivalent basis. Overall, afforestation of improved pastures (i) decreases soil N₂O emissions and (ii) increases soil CH₄ uptake.     

山坡退化林地林木生长对微地形人工干预的响应

为探究水土保持微地形人工干预措施(等高反坡阶)对坡面退化林地林木生长的影响,在昆明市松华坝水源区内选取典型山坡退化林地,通过动态观测和对比分析不同样地林木胸径、树高、新枝生长量及样地叶面积指数(LAI)的变化差异,再结合土壤水分有效性分析以探讨差异原因.结果表明:等高反坡阶样地云南松胸径和树高的最大值和均值均大于对照样地(CK),退化林群落中小径阶和低矮的植株占更大比例,表明等高反坡阶样地中优势木新生植株和退化严重植株的生长潜力得到激发,不同样地云南松植株新枝平均枝长和枝径的年变化差异率达到了72.4%和39.1%.等高反坡阶样地云南松植株新枝生长量和样地LAI的变化及新枝生长的速度大于CK.土壤含水率和新枝生长量与LAI均呈显著相关,对水分有效性分析后得到等高反坡阶样地易效水比例(64.2%)整体高于CK(54.7%),5月至9月雨季中,等高反坡阶样地易效水存在时间更长,这有利于林木的生长,进而改变退化林地植物群落结构.     

An analysis of the vegetation pattern in a semi-arid Eucalyptus populnea woodland in north-west New South Wales

Shrub invasion of Eucalyptus populnea woodland is of concern to the pastoral industry. As part of an ecosystem study, different analyses were used to identify spatial pattern and associations in the tree and shrub species, with the aim of identifying what factors were most influential on the ecosystem. The results suggested that there were no strong edaphic or topographic influences on the distribution of plants. The principal influence on the arrangement of trees and shrubs was the large Eucalyptus trees, which were randomly arranged, and the canopy of which covered 15% of the area. It is concluded that any disturbance of the large Eucalyptus trees would have intense effects on the whole ecosystem. Qualitative and quantitative measurements of the plant species in sample quadrats were subjected to a variety of pattern analyses including association analysis (DIVINF), correlation analysis, principal components analysis and several classification programs including POLYDIV and MULCLAS. A criticism of such analyses is that positive results are both inevitable and not subject to statistical proof, whilst their strength is that they can simplify complicated sets of data. In this case the patterns suggested by the results were visually apparent and no additional insights were achieved. The dangers inherent in relying on a single pattern analysis were revealed when the groups of quadrats produced by the different analyses were mapped. Even when the groupings derived from different analyses were based on a similar species content, their spatial arrangement was dissimilar.     

Rhizosphere-driven increase in nitrogen and phosphorus availability under elevated atmospheric CO2 in a mature Eucalyptus woodland

Plant and Soil - Soil degradation is a major global problem; to investigate the potential for recovery of soil biota and associated key functions, soils were monitored during the early years of...     

Elevated CO2 did not affect the hydrological balance of a mature native Eucalyptus woodland

Elevated atmospheric CO₂ concentration (eC_a) might reduce forest water‐use, due to decreased transpiration, following partial stomatal closure, thus enhancing water‐use efficiency and productivity at low water availability. If evapotranspiration (E_t) is reduced, it may subsequently increase soil water storage (ΔS) or surface runoff (R) and drainage (D_g), although these could be offset or even reversed by changes in vegetation structure, mainly increased leaf area index (L). To understand the effect of eC_a in a water‐limited ecosystem, we tested whether 2 years of eC_a (~40% increase) affected the hydrological partitioning in a mature water‐limited Eucalyptus woodland exposed to Free‐Air CO₂ Enrichment (FACE). This timeframe allowed us to evaluate whether physiological effects of eC_a reduced stand water‐use irrespective of L, which was unaffected by eC_a in this timeframe. We hypothesized that eC_a would reduce tree‐canopy transpiration (E_tree), but excess water from reduced E_tree would be lost via increased soil evaporation and understory transpiration (E_floor) with no increase in ΔS, R or D_g. We computed E_t, ΔS, R and D_g from measurements of sapflow velocity, L, soil water content (θ), understory micrometeorology, throughfall and stemflow. We found that eC_a did not affect E_tree, E_floor, ΔS or θ at any depth (to 4.5 m) over the experimental period. We closed the water balance for dry seasons with no differences in the partitioning to R and D_g between C_a levels. Soil temperature and θ were the main drivers of E_floor while vapour pressure deficit‐controlled E_tree, though eC_a did not significantly affect any of these relationships. Our results suggest that in the short‐term, eC_a does not significantly affect ecosystem water‐use at this site. We conclude that water‐savings under eC_a mediated by either direct effects on plant transpiration or by indirect effects via changes in L or soil moisture availability are unlikely in water‐limited mature eucalypt woodlands.     

Elevated carbon dioxide increases soil nitrogen and phosphorus availability in a phosphorus‐limited Eucalyptus woodland

Free‐air CO₂ enrichment (FACE) experiments have demonstrated increased plant productivity in response to elevated (e)CO₂, with the magnitude of responses related to soil nutrient status. Whilst understanding nutrient constraints on productivity responses to eCO₂ is crucial for predicting carbon uptake and storage, very little is known about how eCO₂ affects nutrient cycling in phosphorus (P)‐limited ecosystems. Our study investigates eCO₂ effects on soil N and P dynamics at the EucFACE experiment in Western Sydney over an 18‐month period. Three ambient and three eCO₂ (+150 ppm) FACE rings were installed in a P‐limited, mature Cumberland Plain Eucalyptus woodland. Levels of plant accessible nutrients, evaluated using ion exchange resins, were increased under eCO₂, compared to ambient, for nitrate (+93%), ammonium (+12%) and phosphate (+54%). There was a strong seasonality to responses, particularly for phosphate, resulting in a relatively greater stimulation in available P, compared to N, under eCO₂ in spring and summer. eCO₂ was also associated with faster nutrient turnover rates in the first six months of the experiment, with higher N (+175%) and P (+211%) mineralization rates compared to ambient rings, although this difference did not persist. Seasonally dependant effects of eCO₂ were seen for concentrations of dissolved organic carbon in soil solution (+31%), and there was also a reduction in bulk soil pH (‐0.18 units) observed under eCO₂. These results demonstrate that CO₂ fertilization increases nutrient availability – particularly for phosphate – in P‐limited soils, likely via increased plant belowground investment in labile carbon and associated enhancement of microbial turnover of organic matter and mobilization of chemically bound P. Early evidence suggests that there is the potential for the observed increases in P availability to support increased ecosystem C‐accumulation under future predicted CO₂ concentrations.     

Dynamic tree aggregation patterns in a species‐poor temperate woodland disturbed by fire

Abstract. Question: How does fire affect the aggregation patterns of trees in a species‐poor oak woodland? Location: East‐central Minnesota, USA. Methods: More than 10 000 trees with DBH > 2 cm (comprising more than 11 000 stems) were monitored in a 16‐ha grid on an annual basis from 1995‐ 2001 in a species‐poor temperate woodland. Different portions of the grid experienced different frequencies of controlled burns. Aggregation indices were calculated for individual species and individual size classes within species. A community‐wide aggregation index was also calculated for different burn units. Spatial data were managed, and many of the aggregation indices calculated using a GIS ArcInfo? (ESRI). Results: Fire initially increased clumping, although repeated fires reduced it, a finding that suggests a corollary to the intermediate‐disturbance hypothesis, the corollary stating that intermediate levels of disturbance are expected to maximize community‐wide patterns of aggregation. Analyses also showed that all species are aggregated at small scales, that the degree of aggregation of a stem type (species or size) declines with distance from individual stems, that the degree of aggregation of large stems is usually less than that of small stems, and that rare species are more aggregated than common species. Findings from this study are consistent with those from similar studies in other temperate and tropical forests, woodlands, and savannas. Conclusion: The spatial patterns of trees in this woodland are dynamic, continually changing in response to the relative strengths of the often opposing forces of competition, which tends to reduce clumping, and disturbance, which, at low and intermediate frequencies, tends to increase it.     

Macrotermes mounds as sites for tree regeneration in a Sudanian woodland (Burkina Faso)

The importance of mounds created by Macrotermes subhyalinus as safe site for tree regeneration was analysed in a savannah woodland of Burkina Faso. Plantlets (height <1.5 m) were sampled and followed over an year in 72 × 4 m² quadrats located on M. subhyalinus mounds and adjacent areas. The mechanisms of regeneration and plantlet mortality were also determined. We identified three regeneration mechanisms: seedlings regenerated by seed (abundant on mounds), sprouts (abundant on adjacent areas) and root suckers (a rare case on both sites). A total of 37 species representing 17 families and 30 genera were found on all quadrats, of which 29 species were found on termite mounds and 22 species on adjacent areas. Species richness and density of plantlets at the 4 m² scale were higher on mounds than in the adjacent area (P < 0.05). Among plantlet categories, seedling density was significantly different among microhabitats (P < 0.001) and across sampling periods (P < 0.01) and, the majority of plantlet individuals appeared within the 0–25 cm height class. The mortality of plantlets and particularly seedling mortality differed significantly between microhabitats (P < 0.01) and between periods (P < 0.01), whereas more than half the variation in the death of Acacia erythrocalyx seedlings (the most abundant species) were related to the density of the live seedlings of the same species (P < 0.001). The observed mortality rate was way below 50%; plantlet density remained higher on mound during sampling periods as compared to the adjacent area. It can thus be concluded that Macrotermes termite mounds are favourable sites for the recruitments of woody plants in savannah woodlands.     

Shifts from competition to facilitation between a foundation tree and a pioneer shrub across spatial and temporal scales in a semiarid woodland

* Theoretical and empirical research has supported the hypothesis that plant-plant interactions change from competition to facilitation with increasing abiotic stress. However, the consistency of such changes has been questioned in arid and semiarid ecosystems. * During a drought in the semiarid south-western USA, we used observations and a field experiment to examine the interactions between juveniles of a foundation tree (Pinyon pine, Pinus edulis) and a common shrub (Apache plume, Fallugia paradoxa) in replicated areas of high and low stress. * The presence of F. paradoxa reduced P. edulis performance at low-stress sites, but had the opposite effect at high-stress sites. However, the intensity of the interactions depended on temporal variation in climate and age of P. edulis. Both above- and below-ground factors contributed to competition, while only above-ground factors contributed to facilitation. * These results support the hypothesis that interactions can change from competition to facilitation as abiotic stress increases in semiarid environments. A shift from competition to facilitation may be important for the recovery of P. edulis and other foundation species that have experienced large-scale mortality during recent droughts.