Tracing water uptake by jarrah (Eucalyptus marginata) trees using natural abundances of deuterium

 Seasonal change in the δ²H content of water from twig sap, soil, rainfall and groundwater were measured to determine the water sources accessed by jarrah (Eucalyptus marginata) trees at three sites in Western Australia with differing soils and depths to water table. During winter and spring the main contributor to the water uptake of the trees was stored water in the surface layers of the soil replenished by predominantly winter rainfall. With the onset of summer drought jarrah became more reliant on water from deeper down the profile. There was no clear evidence that jarrah could tap water from groundwater more than 14 m deep in deep sands. Defining the source of water for trees in deep lateritic soils using stable isotopes is hampered by the uniform deuterium profiles down most of the unsaturated zone and into the groundwater. There was a limited response in the δ²H values of sapwater in twigs to changes in the δ²H of the upper layers of the deep sand following input of rainfall in autumn. The damped response was related to the small variation in the δ²H composition of rainfall in most events during the year and the mixing in the tree of water extracted from different locations in the soil profile. Received: 21 August 1995 / Accepted: 3 December 1995     

The soil seed bank and understorey regeneration in Eucalyptus regnans forest,Victoria

Abstract The soil seed bank and its relation to the extant vegetation in a Eucalyptus regnans F. Muell. forest in the Central Highlands of Victoria were examined. The average seed density was 430 germinable seeds m^?2 to a depth of 2 cm. There was a polynomial regression relationship between the density and species richness of seeds in soil and forest age (0. 6–54 years). Species richness was not significantly different among soil depths (0- 2 , 2- 5 , 5–10 and 10–20 cm) in the forest stand of 54 years old. More seeds germinated from the 5–10 cm depth than from the other depths. Forbs accounted for 73% of the total germinable seeds and there was no germination of E. regnans. The number of species, particularly woody plant species, germinating from the soil seed bank were significantly lower than in the extant vegetation. However, almost all species present in the soil seed bank were present in the vegetation. The soil seed bank provides an important source for the rapid regeneration of understorey vegetation following clear-cutting and slash-burning in the E. regnans forest. The rapid understorey establishment may play an important role in protecting soil from erosion, in nutrient conservation, replacement and redistribution. The soil seed bank may also be a necessary source of maintaining genetic diversity in the forest over the long term.     

巨桉(Eucalyptus grandis)人工林土壤动物群落对采伐干扰的初期响应

为了解采伐干扰对巨桉(Eucalyptus grandis)人工林土壤动物的影响,于2005～2007年采用手捡法和干湿漏斗法对四川省洪雅县巨桉人工林采伐干扰前和采伐干扰后土壤动物群落进行调查,采伐干扰前四季共获土壤动物4775头,分属4门11纲27目,采伐干扰后获土壤动物4640头,分属4门10纲23目.采伐干扰后秋季(10月)、冬季(1月)和春季(4月)土壤动物个体数较采伐干扰前分别上升224头、368头和443头,而夏季(7月)下降了1170头.采伐干扰前巨桉人工林土壤动物个体数与类群数季节变化趋势基本一致,而采伐干扰后土壤动物类群数变化受土壤动物个体数变化的影响较小.与采伐干扰前相比,采伐干扰后枯落物层各季节土壤动物密度均低于采伐前,0～5 cm土层土壤动物密度与干扰前呈相反的变化趋势,其波动较5～10 cm和10～15 cm土层大.采伐干扰后土壤动物类群数四季共减少4个,采伐干扰前后土壤动物优势类群类群数波动小,采伐后常见类群类群数上升(除冬季外),稀有类群对采伐干扰较敏感,其类群数下降幅度较大.采伐干扰前后同功能种团均以杂食性和腐食性土壤动物为主,两者百分比之和均在89%以上.密度-类群指数(DG)和多样性指数(H′)的变化趋势相似,采伐干扰前后的变化趋势相反,表明采伐干扰对巨桉人工林土壤动物季节变化有较大影响.     

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.     

The independent effects of heat,smoke and ash on emergence of seedlings from the soil seed bank of a heathy Eucalyptus woodland in Grampians (Gariwerd) National Park,western Victoria

Abstract The independent effects of smoke, ash, and wet and dry heat treatments on seedling emergence from the soil seed bank were tested for soils from fire-prone heathy woodlands in western Victoria. A total of 763 individuals from 56 species were recorded from the surface soil samples (which covered a total area of 1 m²). Both species richness and density of seedlings was greater for smoke- and heat-treated samples than for controls and ash-treated samples. However, only the density differences were significant. Mean seed bank densities for the smoke and heat treatments ranged from 855 ± 70m^?2 to 1080 ± 58 m^?2 and are similar to estimates obtained elsewhere in Australia for heat-treated soils from dry sclerophyll communities. Of the 56 species recorded, 46 occurred in the smoke and heat treatments but only 33 in the control and ash treatments. The sudden increase in surface soil pH, exchangeable cations and extractable phosphorus which was associated with the ash treatment did not act as a trigger for germination in any of the species recorded here. Chemical constituents from smoke do appear to provide a stimulus separate from the effects of heat, but were not identified with any particular taxa.     

Decomposition of litter from three major plant species of jarrah (Eucalyptus marginata Donn ex Sm.) forest in relation to site fire history and soil type

Rates of weight loss and release of N, P, K, S, Ca, Mg, Na, and Cl from litter of several species in jarrah (E. marginata Donn ex Sm.) forest were measured in relation to site fire history and soil type. Weight loss from leaf litter decreased in the order jarrah > marri (E. calophylla R. Br. ex Lindl.) > Banksia grandis Willd. After 18 months on the forest floor senesced leaves of jarrah, marri and Banksia had lost 45%, 42% and 19%, respectively, of their original weight. Although greatest rates of decomposition occurred on a site burnt 3 y previously by an intense autumn fire and slowest rates on a site which had not been burnt for 8 y, the differences between burn sites were small in comparison with the total weight loss from decomposing litter. The order of release of nutrients from decomposing eucalypt litter was P<N<Ca<S<Mg<Cl<K<Na. There appears to be only slow release of N and P from the litter layer of these forests in the period between successive control burns. Fresh jarrah leaves, which are similar in chemical composition to leaf litter falling after crowns have been scorched by intense fire, decompose rapidly in comparison with senescent leaf tissue. Release of nutrients, particularly N and P, is also more rapid from fresh leaves than from leaf litter. Rates of decomposition of green leaves differed between soil types in the order reddish gravels > dark sandy duplex soil > yellow gravels. These differences may be related to the higher nutrient status of the reddish gravel soils.     

Response of plant species and life form diversity to variable fire histories and biomass in the jarrah forest of south‐west Australia

Frequent fires reduce the abundance of woody plant species and favour herbaceous species. Plant species richness also tends to increase with decreasing vegetation biomass and cover due to reduced competition for light. We assessed the influence of variable fire histories and site biomass on the following diversity measures: woody and herbaceous species richness, overall species richness and evenness, and life form evenness (i.e. the relative abundance or dominance among six herbaceous and six woody plant life forms), across 16 mixed jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) forest stands in south‐west Australia. Fire frequency was defined as the total number of fires over a 30‐year period. Overall species richness and species evenness did not vary with fire frequency or biomass. However, there were more herbaceous species (particularly rushes, geophytes and herbs) where there were fewer shrubs and low biomass, suggesting that more herbaceous species coexist where dominance by shrubs is low. Frequently burnt plots also had lower number and abundance of shrub species. Life form evenness was also higher at both high fire frequency and low biomass sites. These results suggest that the impact of fire frequency and biomass on vegetation composition is mediated by local interactions among different life forms rather than among individual species. Our results demonstrate that measuring the variation in the relative diversity of different woody and herbaceous life forms is crucial to understanding the compositional response of forests and other structurally complex vegetation communities to changes in disturbance regime such as increased fire frequency.     

Recruitment limitation in dry sclerophyll forests: Regeneration requirements and potential density‐dependent effects in Eucalyptus tricarpa (L.A.S. Johnson) L.A.S. Johnson & K.D. Hill (Myrtaceae)

Recruitment limitation in canopy trees is receiving increasing attention as restoration of tree cover for connectivity, biodiversity offsets, carbon‐trading and improved catchment health becomes more prominent. Recruitment limitation is often addressed by examining seed traits and germination requirements. Distance between trees is also often explored as a factor, particularly in agricultural landscapes where forest structure has been altered, with large distances between fragments generating Allee effects linked to pollen limitation. Fewer studies have examined how short distances between trees (high stem densities), which can characterize regenerated forests, might affect recruitment. This study examined recruitment limitation by exploring germination requirements of Eucalyptus tricarpa (red ironbark) and the effects of stem density on seed characters and germination. Eucalyptus tricarpa is a canopy species in the box‐ironbark forests of central Victoria, Australia. During the early European settlement phase in the 1800s these forests were heavily impacted by human disturbances including mining, logging and livestock grazing. In many regenerated stands, now more than 100 years old, current stem density exceeds those documented in 19th century reports and there is little regeneration of key canopy species such as E. tricarpa. In laboratory trials seed germinated at 18°C in darkness, and while viability varied among populations, overall it remained high (91–96%) and did not differ among populations with different stem densities. An in situ germination trial addressing sowing season, ground‐cover, soil‐ripping, canopy‐cover and stem density recorded no germination. High stem density in coppicing box‐ironbark forest was hypothesized to limit resources for reproduction; however, no relationship was found between nearest neighbour distance, allocation to selected reproductive structures or seed viability. The relative insensitivity of reproductive allocation to intraspecific competition (high stem density) may be interpreted as a trait linked to unpredictable environments and consistent with long‐lived species. In terms of management, recruitment limitation in E. tricarpa appears not primarily attributable to seed viability, tested seedbed treatments or stem density so further factors, including fire, now require exploration.     

Conifer seedling recruitment across a gradient from forest to alpine tundra: effects of species,provenance, and site

Background: Seedling germination and survival is a critical control on forest ecosystem boundaries, such as at the alpine–treeline ecotone. In addition, while it is known that species respond individualistically to the same suite of environmental drivers, the potential additional effect of local adaptation on seedling success has not been evaluated.

Aims: To determine whether local adaptation may influence the position and movement of forest ecosystem boundaries, we quantified conifer seedling recruitment in common gardens across a subalpine forest to alpine tundra gradient at Niwot Ridge, Colorado, USA.

Methods: We studied Pinus flexilis and Picea engelmannii grown from seed collected locally at High (3400 m a.s.l.) and Low (3060 m a.s.l.) elevations. We monitored emergence and survival of seeds sown directly into plots and survival of seedlings germinated indoors and transplanted after snowmelt.

Results: Emergence and survival through the first growing season was greater for P. flexilis than P. engelmannii and for Low compared with High provenances. Yet survival through the second growing season was similar for both species and provenances. Seedling emergence and survival tended to be greatest in the subalpine forest and lowest in the alpine tundra. Survival was greater for transplants than for field-germinated seedlings.

Conclusions: These results suggest that survival through the first few weeks is critical to the establishment of natural germinants. In addition, even small distances between seed sources can have a significant effect on early demographic performance – a factor that has rarely been considered in previous studies of tree recruitment and species range shifts.     

Emergence of seedlings of sugar beet (Beta vulgaris L.) as affected by the size,roughness and position of aggregates in the seedbed

Laboratory and field experiments were carried out to quantify the effects of the size and roughness of aggregates placed in the seedling path of sugar beet, in order to help in decision making for soil tillage and sowing operations. Graded aggregates (10, 15, 20, 30, 40, 50, 60 and 70 mm longest axis) were either laid on the soil surface or included in the soil over the seeds. The percent emergence decreased exponentially with aggregate size over 10 mm when the aggregates were included in the seedbed. The result was the same with aggregates laid on the soil surface, but for aggregates over 30 mm (mass>10 g). Aggregates on the soil surface could be lifted by the seedlings until their weight exceeded the seedling emergence force. Larger aggregates or aggregates in the soil layer could not be moved. Seedlings which did not emerge remained blocked in small cavities in the aggregate surface. No seedlings were blocked under smooth aggregates or glass beads. The experimental results fits with a model giving the probability to meet a hole according the distance covered and the diameter and density of holes. The results obtained under controlled conditions were similar to those obtained in field experiments for a wide range of aggregate sizes. These results will be incorporated into a computerised seedbed generator to simulate the effects of seedbed structure on seedling emergence.     

Long‐term changes in the vegetation after the cessation of livestock grazing in Eucalyptus marginata (jarrah) woodland remnants

This paper documents changes in the floristic composition of Eucalyptus marginata Donn (jarrah) woodlands over 7 years of recovery from continual, intensive livestock grazing. In remnants of native woodland left after agricultural clearing, which have been subjected to livestock grazing, comparisons were made between the floristics of fenced exclosure plots and open plots that continued to be grazed. The vegetation in nearby remnants, which had not been subjected to livestock grazing, was also surveyed. An initial increase in annual exotic pasture species after grazing relief was only temporary and highly influenced by fluctuations in annual climatic patterns, particularly rainfall distribution and abundance. Subsequent years saw a decrease in exotic annuals in exclosure plots and an increase in native perennials, in a trend towards becoming more floristically similar to the ungrazed sites. Germination of overstorey species was observed in the exclosure plots, however, development of seedlings and saplings was sparse. Results indicate that for jarrah woodland in southwestern Australia, natural regeneration is possible after the removal of livestock, with the return (within 6 years) of native species richness to levels similar to those found in ungrazed vegetation. Re‐establishment of cover, however, appears to take longer. The floristic dynamics are described in terms of a nonequilibrium model. Two vegetation states exist, degraded remnants with an understorey dominated by annual species, and ungrazed vegetation with an understorey dominated by perennial shrubs and herbs. The former state is maintained by continual heavy grazing by livestock. Upon relief from grazing, the vegetation undergoes a transition towards floristic similarity to ungrazed vegetation. After 6 years, vegetation change in the exclosure plots appears to be continuing and therefore it is still in transition.     

沙埋和种子大小对柠条锦鸡儿种子萌发、出苗和幼苗生长的影响

研究了沙埋深度和种子大小对内蒙古毛乌素沙地植被群落中占优势的柠条锦鸡儿(Caragana korshinskii Kom.)种子萌发、出苗、幼苗存活和生长的影响.结果表明,沙埋深度显著影响柠条锦鸡儿的种子萌发率、休眠率、出苗率、幼苗存活率及生物量.在0.5-2cm的浅层沙埋下,种子萌发率、出苗率、幼苗存活率及生物量最高,休眠率最低;沙埋深度≥4 cm时,柠条锦鸡儿的种子萌发率、出苗率、幼苗存活率及生物量随着沙埋深度增加显著降低,而休眠率却显著升高;沙埋深度≥12 cm时,柠条锦鸡儿种子不能够出苗,幼苗也不能够存活.种子大小对柠条锦鸡儿种子萌发率没有显著影响,但对出苗率、幼苗存活率及生物量影响显著.在各个沙埋深度下,不同大小的柠条锦鸡儿种子间的萌发率没有显著差异.当沙埋深度≤6 cm时,不同大小的柠条锦鸡儿种子在同一沙埋深度下的出苗率间没有显著差异;但当沙埋深度≥8 cm时,在同一沙埋深度下,大种子的出苗率显著高于中种子和小种子的出苗率,而中种子和小种子出苗率间没有显著差异.0.5-10 cm的沙埋深度中,除6 cm和8 cm深度下中种子和小种子萌发幼苗的生物量间没有显著差异外,其余深度下,大种子萌发的幼苗的存活率及生物量显著高于同一沙埋深度下中种子萌发的幼苗的存活率及生物量,后者又显著高于小种子萌发的幼苗的存活率及生物量.可能正是种子萌发对沙埋环境的忍耐或响应能力以及种子的多态性提高了柠条锦鸡儿在毛乌素沙地的适合度,为其在流动或半流动沙丘环境中成功定居并形成优势群落奠定了基础.     

Ecto- and arbuscular mycorrhizal symbiosis can induce tolerance to toxic pulses of phosphorus in jarrah (Eucalyptus marginata) seedlings

In common with many plants native to low P soils, jarrah (Eucalyptus marginata) develops toxicity symptoms upon exposure to elevated phosphorus (P). Jarrah plants can establish arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) associations, along with a non-colonizing symbiosis described recently. AM colonization is known to influence the pattern of expression of genes required for P uptake of host plants and our aim was to investigate this phenomenon in relation to P sensitivity. Therefore, we examined the effect on hosts of the presence of AM and ECM fungi in combination with toxic pulses of P and assessed possible correlations between the induced tolerance and the shoot P concentration. The P transport dynamics of AM (Rhizophagus irregularis and Scutellospora calospora), ECM (Scleroderma sp.), non-colonizing symbiosis (Austroboletus occidentalis), dual mycorrhizal (R. irregularis and Scleroderma sp.), and non-mycorrhizal (NM) seedlings were monitored following two pulses of P. The ECM and A. occidentalis associations significantly enhanced the shoot P content of jarrah plants growing under P-deficient conditions. In addition, S. calospora, A. occidentalis, and Scleroderma sp. all stimulated plant growth significantly. All inoculated plants had significantly lower phytotoxicity symptoms compared to NM controls 7 days after addition of an elevated P dose (30 mg P kg^?1 soil). Following exposure to toxicity-inducing levels of P, the shoot P concentration was significantly lower in R. irregularis-inoculated and dually inoculated plants compared to NM controls. Although all inoculated plants had reduced toxicity symptoms and there was a positive linear relationship between rank and shoot P concentration, the protective effect was not necessarily explained by the type of fungal association or the extent of mycorrhizal colonization.     

The effects of a single hot summer fire on soil fauna and on litter decomposition in jarrah (Eucalyptus marginata) forest in Western Australia

An area of jarrah (Eucalyptus marginata Sm) forest was burnt with a hot fire (ca 600 kw/ha) in March 1975. The meso- and microarthropod faunas were sampled in May, June and July 1976 and the decomposer activity of the soil and litter was estimated using buried cotton strips. The growth of jarrah seedlings in pots was measured under different litter and nutrient regimes. Feeding experiments with Eucalyptus, Banksia and Bossiaea litter were carried out on Podykipus sp., a litter millipede common at the site of the fire. Burning reduced the numbers of arthropods, the proportion of juveniles and the proportion of fungal feeders in the micro-arthropod population. The rate of decomposition was also reduced. The seedlings grew most under leaf Utter and millipede faeces and least under leaf ash. Podykipus sp. preferred Bossiaea litter (high nutrient content) to either Eucalyptus or Banksia litter (low nutrient content).     

Arthropod fauna of jarrah (Eucalyptus marginata) foliage in Mediterranean forest of Western Australia: Spatial and temporal variation in abundance,biomass, guild structure and species composition

Abstract A 2 year study (using a branch-clipping technique) of the arthropod fauna of jarrah crowns (14 m above ground level) in a 7500km² area centred on Manjimup, Western Australia, yielded 7461 individuals belonging to 396 species. Lepidoptera, Hemiptera and Araneae dominated the fauna in abundance and biomass but Lepidoptera less so in number of species. Consequently, leaf chewers, sapsuckers and predators were the dominant guilds. Most species showed great spatial and temporal variation in their occurrence, with some 73% being recorded on only one or two of the nine sampling occasions. In addition 77% of species were recorded on four or fewer of the 45 trees sampled. Crowns of these trees tended to have highly dissimilar assemblages of arthropods. Dissimilarity between trees was not predictable from the distance between them. Total species richness was greatest in samples collected in summer. Predators and sapsuckers tended to be the most speciose guilds. Within-stand differences were much less pronounced than between-stand differences: ants were more abundant on jarrah foliage within 2 m of ground level and the pest defoliator Uraba lugens (Lepidoptera, Noc-tuidae) was more abundant on foliage of jarrah pole crowns 14 m above ground. The most abundant arthropod, U. lugens, did not reduce appreciably the biomass or abundance of other arthropod groups present in jarrah pole crowns. This is consistent with the paradigm that competition between species of herbivorous insects is infrequent. After comparison with other studies in jarrah forest, we tentatively conclude that there is no single fixed pattern of organization or predictable assemblage of invertebrates on jarrah foliage.