Response of eucalypt species to drought

The response of three eucalypt species (Eucalyptus pulchella, Eucalyptus coccifera and Eucalyptus delegatensis) to a severe drought in the summer of 1982/83 was examined at Snug Plains, south-eastern Tasmania. Few large differences in leaf water potential (Ψ_l) or stomatal conductance (gs) were apparent even at the height of the drought when both Ψ_l and soil water potentials (Ψ_s) reached ca. — 4.5 MPa. However, E. pulchella maintained a higher relative water content (RWC) in its leaves than E. coccifera and E. delegatensis, and showed less severe crown damage. After the first light rains substantial interspecific differences in Ψ_l and gs occurred. Eucalyptus pulchella restored normal Ψ_l, gs and RWC more rapidly than the other two species and, even for severely droughted trees, crown growth commenced via epicormic buds near the ends of its branches while for E. delegatensis and E. coccifera crown regeneration was via epicormic buds arising from stems and larger branches. This resulted in a change in dominance in certain stands and showed that E. pulchella was more drought-resistant than E. coccifera, which was in turn more resistant than E. delegatensis. This conclusion was confirmed during competition experiments using potted seedlings. However, potted seedlings differed from mature field trees by maintaining moderate gs at high vapour pressure deficits and closing stomata at Ψ_l below ca. — 2.0 MPa. Substantial variation in the severity of drought symptoms was observed over short distances. This variation appeared to be determined by the moisture-holding capacity of the soil and the biomass of the stand. Although differences in the rooting patterns of seedlings were evident, field measurements of Ψ_l and Ψ_s suggested that all three species were exploiting the same water resource. In contrast to previous studies, the results suggest that large interspecific differences in tissue hydration and crown damage may be present, even though differences in Ψ_l, gs and characteristics of the root system may appear small.     

The relationships among eucalypt forest,grassland and rainforest in a highland area in north-eastern Tasmania

Changes in the vegetation of a highland area of north-eastern Tasmania are described and related to a change 150 years ago from Aboriginal to settled land use. Expansion of rainforest dominated by Nothofagus into areas previously dominated by Eucalyptus, and of Eucalyptus forest on to grassland that occupied former rain - forest areas is accounted for by local differences in frequency and severity of fires since about 1835. Successional relationships amongst 12 vegetation types are defined, and discussed in relation to three models of vegetational succession.     

The functional morphology of juvenile plants tolerant of strong summer drought in shaded forest understories in southern Spain

It has been hypothesized that plants cannot tolerate combined shade and drought, as a result of morphological trade-offs. However, numerous plant species are reportedly widespread in shaded forest understories that face drought, whether seasonal or occasional. We studied juveniles of six plant species that cope with strong summer drought in the understoreys of mixed Quercus forests in southern Spain: the tall-shrubs Phillyrea latifolia and Viburnum tinus, the perennial herb Rubia peregrina, the small shrub Ruscus aculeatus, and climbers Hedera helix and Smilax aspera. All of these species persist in evergreen shade (c. 3% daylight). Two other species were studied as comparators, Ruscus hypoglossum, less tolerant of drought, and Ceratonia siliqua, less tolerant of shade. Morphological and chemical variables relevant to shade and drought tolerance were measured for juveniles in a range of sizes, and also for the leaves of mature plants. The species converge in features that confer tolerance of shade plus drought by reducing demand for resources. Demand for water is reduced through a moderate to high below-ground mass fraction and low to moderate specific leaf area (respectively 0.22–0.52 and 112–172 cm² g^–1 at 1.00 g total dry mass). Demand for both irradiance and water is reduced through a low to moderate foliar nitrogen concentration and long-lived, physically protected leaves (2 yr). The species also converge in features that confer tolerance of either low irradiance or drought through specialized capture of resource, without precluding the other tolerance. These features include deep roots relative to shoot size, moderately higher specific leaf area in shade (1.2–2.0 × that in sun) and higher chlorophyll:nitrogen ratio in shade. Foliar chlorophyll per unit mass was higher in shade, but chlorophyll was not necessarily synthesized in greater amounts; rather, it was higher apparently due to shade effects on structural features linked with specific leaf area. In contrast, N per unit mass was higher in sun leaves independently of specific leaf area. Despite these convergences, the species diverge considerably in their root mass allocation and architecture, leaf saturated water content, density of stomata and guard cell size. No single narrowly defined functional type is needed for tolerance of shade plus drought.     

Altitudinal distribution of several eucalypt species in relation to other environmental factors in southern New South Wales

Four problems associated with studying the altitudinal distribution of eucalypt species are examined: the lack of specific physiological relationship between altitude and plant growth, the influence of other environmental factors, the availability of suitable data and the need for statistical analysis. Presence/absence data for eucalypt species were obtained from several sources. Probability of occurrence in 100 m zones is determined for E. maculata, E. muellerana, E. fastigata, E. sieberi, E. dalrympleana and E. pauciflora. The influence of other factors is demonstrated for several species using direct gradient analysis. Aspect is important for E. fastigata and E. rossii in addition to altitude and rainfall. The statisical model used was the logit-linear model: log (p/I– p) = linear function of environmental variables where p is the expected probability of being present for a given combination of environmental variables. Two examples are presented. E. dalrympleana can be predicted from altitude, rainfall, radiation index (measure of aspect) and an interaction term between altitude and aspect. E. rossii presence is predicted by altitude, rainfall, radiation index and geology. Altitude is transformed into an estimate of mean annual temperature which is shown to clarify some overlaps of species distribution. It is concluded that use of data collected for other purposes can be used in a generalized linear model for presence data to show the complex correlations which exist between the altitudinal distribution of some eucalypts and other environmental factors.     

Hydraulic failure and tree dieback are associated with high wood density in a temperate forest under extreme drought

Catastrophic hydraulic failure will likely be an important mechanism contributing to large‐scale tree dieback caused by increased frequency and intensity of droughts under global climate change. To compare the susceptibility of 22 temperate deciduous tree and shrub species to hydraulic failure during a record drought in the southeastern USA, we quantified leaf desiccation, native embolism, wood density, stomatal conductance and predawn and midday leaf water potential at four sites with varying drought intensities. At the two driest sites, there was widespread leaf wilting and desiccation, and most species exhibited predawn leaf water potentials of ≤3 MPa and >60% loss of xylem conductivity in branches. Although species with high wood density were more resistant to cavitation, they had higher levels of native embolism and greater canopy dieback than species with low wood density. This unexpected result can be explained by the failure of species with dense wood to avert a decline in water potential to dangerous levels during the drought. Leaf water potential was negatively correlated with wood density, and the relationship was strongest under conditions of severe water deficit. Species with low wood density avoided catastrophic embolism by relying on an avoidance strategy that involves partial drought deciduousness, higher sensitivity of stomata to leaf water potential and perhaps greater rooting depth. These species therefore maintained water potential at levels that ensured a greater margin of safety against embolism. These differences among species may mediate rapid shifts in species composition of temperate forests if droughts intensify due to climate change.     

Fire-mediated dieback and compositional cascade in an Amazonian forest

The only fully coupled land-atmosphere global climate model predicts a widespread dieback of Amazonian forest cover through reduced precipitation. Although these predictions are controversial, the structural and compositional resilience of Amazonian forests may also have been overestimated, as current vegetation models fail to consider the potential role of fire in the degradation of forest ecosystems. We examine forest structure and composition in the Arapiuns River basin in the central Brazilian Amazon, evaluating post-fire forest recovery and the consequences of recurrent fires for the patterns of dominance of tree species. We surveyed tree plots in unburned and once-burned forests examined 1, 3 and 9 years after an unprecedented fire event, in twice-burned forests examined 3 and 9 years after fire and in thrice-burned forests examined 5 years after the most recent fire event. The number of trees recorded in unburned primary forest control plots was stable over time. However, in both once- and twice-burned forest plots, there was a marked recruitment into the 10-20cm diameter at breast height tree size classes between 3 and 9 years post-fire. Considering tree assemblage composition 9 years after the first fire contact, we observed (i) a clear pattern of community turnover among small trees and the most abundant shrubs and saplings, and (ii) that species that were common in any of the four burn treatments (unburned, once-, twice- and thrice-burned) were often rare or entirely absent in other burn treatments. We conclude that episodic wildfires can lead to drastic changes in forest structure and composition, with cascading shifts in forest composition following each additional fire event. Finally, we use these results to evaluate the validity of the savannization paradigm.     

Biomass and nutrient distribution in two eucalypt forest ecosystems

The distribution of biomass and nutrients (N, P, K, Na, Mg, and Ca) among components of a Eucalyptus regnans forest and a mixed Eucalyptus obliqua-Eucalyptus dives forest near Melbourne in southern Victoria have been determined and are discussed. Both forests were found to have relatively low root biomass. Trees and soil accounted for more than 80% of each nutrient present in both ecosystems. The results of nutrient distribution studies in Australian eucalypt forests and in temperate forests elsewhere in the world are compared. These comparisons suggest that the above-ground living biomass in eucalypt forests contains no abnormally high or low quantities of macronutrients in relation to coniferous or deciduous hardwood forests. Detailed comparison among ecosystems is rendered difficult by the variable treatment given to roots and soils.     

The effect of regeneration burning upon the nutrient status of soil in two forest types in southern Tasmania

In two forest types in southern Tasmania, eucalypt rainforest (mixed forest) and eucalypt dry sclerophyll forest, surface soils (0–10 cm) from stands that had been clear-felled and burned between 1976 and 1979 were compared with those from uncut, unburned stands. Factors compared were total organic C, N, P, K, Mg, Ca, Zn, Mn; pH; exchangeable Ca, Mg, and K; cation exchange capacity; extractable P; soil phosphate buffering capacity; and N-mineralisation rates. Sampling started in April 1979 and ended in October 1980. Within each forest type, soils from burned coupes had higher mean values for pH, exchangeable cations, percent base saturation, and nitrate-N produced during aerobic incubation, and had lower mean values for exchangeable acidity and ammonium-N produced during aerobic incubation than soils from unburned coupes. In mixed forest only, soils from burned coupes had higher mean values for extractable P and soil phosphate buffering capacity, and lower mean values for total organic C than those of unburned coupes. There were only small differences between burned and unburned soils in cation exchange capacity and ammonium-N produced during anaerobic incubation. For each burned coupe in mixed forest, with increase in time since burning there was a decrease in pH, an increase in exchangeable acidity, and a decrease in rate of production of nitrate: no changes were detected in other factors. It is concluded that, for clay soils developed on dolerite, the nutritional status of soil in both forest types is probably improved by burning. The improvement lasts for more than 4 years in mixed forest and more than two years in dry sclerophyll forest. Only minor leaching of nutrients to below 10 cm in depth is likely to occur in either type.     

Response of soil chemistry to forest dieback after bark beetle infestation

We evaluated changes in the chemistry of the uppermost soil horizons in an unmanaged spruce forest (National Park Bohemian Forest, Czech Republic) for 3 years after dieback caused by a bark beetle infestation, and compared these changes with a similar undisturbed forest area. The soils below the disturbed forest received 2–6 times more elements via litter fall compared to the unaffected plot. The subsequent decomposition of litter and reduced nutrient uptake by trees resulted in a steep increase in soil concentrations of soluble N (NH₄-N, organic-bound N) and P forms in the disturbed plot. The average concentrations of NH₄-N and soluble reactive P increased from 0.8 to 4.4 mmol kg^?1 and from 0.04 to 0.9 mmol kg^?1, respectively, in the uppermost soil horizon. Decomposition of litter at the disturbed plot elevated soil concentrations of Ca²⁺, Mg²⁺ and K⁺, which replaced Al³⁺ and H⁺ ions from the soil sorption complex. Consequently, soil concentrations of exchangeable base cations increased from 120 to 200 meq kg^?1, while exchangeable Al³⁺ and H⁺ decreased 66 and 50 %, respectively, and soil base saturation increased from 40 to 70 %. The Al³⁺ liberation did not elevate concentrations of ionic Al in the soil solution, because most of the liberated Al³⁺ was rapidly complexed by dissolved organic carbon (DOC) and transformed to DOC–Al complexes. The chemical parameters investigated at the unaffected plot remained stable during the study.     

Component analysis of fire patterns in open eucalypt forest

The understorey vegetation of nine localities with different fire histories from open eucalypt forest near Melbourne, Victoria, was analysed by principal component analysis. Floristically, localities were quite similar; however, structural differences caused mainly by different burning regimes of recent years were more evident. An analysis of presence-absence data displayed a marked discontinuity that was explainable in terms of the timing and intensity of a recent fire. Using unstandardized height data the pattern was related to inter- and intra-locality differences in time since the last fires. An analysis of standardized height data demonstrated a connection between understorey structure and fire frequency. Despite apparent differences in the scatter diagrams obtained a statistical comparison of the analytical results indicated that, in many respects, the ordinations were similar     

Psychosocial stress in pregnancy and its relation to the onset of premature labour.

A modified life events inventory was presented over a four-month period to 132 consecutive women going into spontaneous labour in Hull and Manchester. Three study groups were identified according to the duration of pregnancy. The levels of psychosocial stress in pregnancy were found to be particularly high in the mothers whose babies were born preterm. Stressful events may precipitate preterm labour in some women. The concept of antenatal care may have to be broadened if the incidence of premature labour and resulting perinatal mortality are to be reduced.     

Impact of commercial forest management on geophilous carabid beetles (Coleoptera: Carabidae) in tall,wet Eucalyptus obliqua forest in southern Tasmania

Abstract The impact of silvicultural practices (clearfelling, slash burning and sowing) on carabid beetle communities in tall wet Eucalyptus obliqua forests in southern Tasmania was examined using traplines of pitfalls in a chronosequence of regenerating coupes. Total species richness (n= 18) was modest compared to other temperate forests and was not systematically changed by forest management, although it was considerably lower in 20 year old regrowth than in younger regrowth or old-growth controls, possibly due to habitat simplification. The Shannon-Weiner Index was at a minimum in intermediate aged regeneration. TWINSPAN analysis assisted recognition of beetle communities typical of broad stages in the forest succession, with a major dichotomy between most old-growth sites plus young sites and intermediate plus advanced regeneration sites. Vector fitting of environmental variables in an ordination of the sites by non-metric multidimensional scaling revealed a significant influence for the age of regeneration and litter depth, but not bare ground percentage or soil pH. Survival of carabids in the habitat mosaic created by commercial forestry activity is likely to depend on both reinvasion from edges and survival of individuals in the fire-protected refuges that exist within the coupes. Pioneer species were winged, small in size, and non-endemic whereas the opposite was true of the fauna in the older sites. Carabids in eucalypt forests have good potential as indicators but their seasonality in occurrence demands that sampling be extended over most of the year.     

Provenance variation in morphology and leaflet anatomy of Parkia biglobosa and its relation to drought tolerance

 Measurements of morphological and anatomical characteristics made on 4-month-old seedlings from five provenances of Parkia biglobosa (Jacq.) Benth., grown in a tropical greenhouse under controlled conditions, showed significant differences between the northern provenances, which represented the Sudan savanna zone, and the southern provenances which represented the forest zone of the species distribution. Seedlings of provenances from the savanna zone showed more xerophytic characteristics than those from the forest zone. They were smaller in height, with a greater number of smaller leaflets per pinnae, a lower specific leaflet mass, smaller palisade and guard cells and a lower ratio of palisade:non-palisade tissue. There were also significant differences in the same characteristics, except palisade cell length, between provenances from the easterly and westerly extremes of the species range. Stomatal conductance measured after 2 weeks of droughting also showed that provenances from the savanna zone had higher values, indicating greater physiological activity and reduced stress, compared with provenances from the forest zone. Multivariate discriminant analysis of the morphological and anatomical characteristics showed that 98% of the seedlings were classified correctly with respect to provenance. The squared distances between provenances were closely related to the geographical distances between locations. Yet, not all variation in leaf anatomy could be explained by the difference in climatic conditions from north to south. Also drought tolerance was not always related to morphological and anatomical features. It is possible that clinal variation within the species from east to west may be interacting with climatic differences from north to south. Received: 2 June 1998 / Accepted: 29 June 1998     

Growth rhythms and productivity of a coastal subtropical eucalypt forest

Growth of plant parts in different species in Eucalyptus signata dominated forest in Queensland occurs at different times. Measurement of increments in root basal diameter combined with previously obtained regressions of root weight on root diameter allows separate estimation of the growth of woody root parts on a seasonal basis for the first time. The growth of lateral and of tap roots as well as of stems and leafy twigs, were separately measured on three tree species every 6 weeks for 2 years. Net primary productivity for the forest is estimated at 26 t ha^-1 yr^-1, of which 66% is aboveground production, and 78% is contributed by a single species. This productivity rate is unexpectedly high, and may reflect a growth stimulus due to release of nutrients following a recent fire.     

When the forest dies: the response of forest soil fungi to a bark beetle-induced tree dieback

Coniferous forests cover extensive areas of the boreal and temperate zones. Owing to their primary production and C storage, they have an important role in the global carbon balance. Forest disturbances such as forest fires, windthrows or insect pest outbreaks have a substantial effect on the functioning of these ecosystems. Recent decades have seen an increase in the areas affected by disturbances in both North America and Europe, with indications that this increase is due to both local human activity and global climate change. Here we examine the structural and functional response of the litter and soil microbial community in a Picea abies forest to tree dieback following an invasion of the bark beetle Ips typographus, with a specific focus on the fungal community. The insect-induced disturbance rapidly and profoundly changed vegetation and nutrient availability by killing spruce trees so that the readily available root exudates were replaced by more recalcitrant, polymeric plant biomass components. Owing to the dramatic decrease in photosynthesis, the rate of decomposition processes in the ecosystem decreased as soon as the one-time litter input had been processed. The fungal community showed profound changes, including a decrease in biomass (2.5-fold in the litter and 12-fold in the soil) together with the disappearance of fungi symbiotic with tree roots and a relative increase in saprotrophic taxa. Within the latter group, successive changes reflected the changing availability of needle litter and woody debris. Bacterial biomass appeared to be either unaffected or increased after the disturbance, resulting in a substantial increase in the bacterial/fungal biomass ratio.     

A comparative study of oak (Quercus, Fagaceae) seedling physiology during summer drought in southern California

Natural recruitment of oaks appears to be declining throughout the northern hemisphere. Summer drought poses a potentially important barrier to oak recruitment in southern California. To evaluate this barrier, we grew evergreen Quercus agrifolia and deciduous Q. lobata from seeds near parental trees. We measured water relations, chlorophyll fluorescence, and gas exchange during these seedlings' fourth and fifth summers and compared them to neighboring adults. Most seedlings had substantially lower values for predawn xylem pressure potential (Ψ(pd)), minimum photosystem II (PSII) quantum efficiency (Φ(PSIIMIN)), maximum quantum efficiency for PSII under dark-adapted leaf conditions (Fv/Fm), and maximum photosynthetic assimilation (Amax), and higher values for maximum nonphotochemical quenching (NPQmax) than did conspecific adults. The high, unvarying Ψ(pd) values of the adults suggest they use perennially available groundwater. Quercus lobata seedlings commonly had lower values for Ψ(pd) than did Q. agrifolia, and values for Ψ(pd) and Φ(PSIIMIN) were significantly related to size in Q. lobata but not in Q. agrifolia. These data suggest important interspecific differences in root architecture. Lower values for Φ(PSIIMIN), Fv/Fm, and higher NPQmax in Q. agrifolia indicate that Q. agrifolia seedlings were usually under more stress than Q. lobata, which typically had higher Amax rates than did Q. agrifolia seedlings. Diurnal photosynthesis curves were quite flat for Q. agrifolia, but they peaked in the morning for Q. lobata. Established seedlings appeared to be under more stress than adults, but this stress did not appear severe enough to cause death. Access to perennially available groundwater may be crucial for the seedling to sapling transition.