Plant size and season of burn affect flowering and fruiting of the grasstree Xanthorrhoea preissii

Flowering and fruiting were assessed on 14 populations of the grasstree, Xanthorrhoea preissii Endl., occurring in the Darling Range near Perth, Western Australia. Independent of site, season of burn or year of flowering, there was a strong relationship between plant height, which varied from 0.1 to over 2 m, and the incidence of postfire flowering, which varied from 1% (winter burn) to 75% (summer burn) of grasstrees present. There was no relationship between inflorescence dimensions, or flower or fruit production on a spike basis, and plant size/age (height). When standardized for height, spring‐burnt populations produced 40% as many inflorescences as autumn‐burnt populations and 20% as many as summer‐burnt populations. Inflorescences produced by spring‐burnt plants were moderately smaller than those by summer–autumn‐burnt plants. Fruit density per spike in autumn‐burnt plants was 80% of that in spring–summer‐burnt plants. The net effect was an average of 70 000 fruits produced per 100 summer‐burnt plants, 22 000 in autumn‐burnt plants, and 14 000 in spring‐burnt plants. Ecophysiological explanations of these results and their implications for population dynamics have yet to be explored.     

Error in the inference of fire history from grasstrees

Abstract A method for constructing fire histories has recently been proposed for fire‐prone southern Australia based on the pattern of coloured bands in the remnant leaf‐bases on stems of grasstrees (Xanthorrhoea species; Xanthorrhoeaceae). In the absence of alternative high‐resolution techniques extending into the premodern period, this method has been utilized to construct fire histories for forest, woodland and shrubland ecosystems, principally in south‐western Australia. However, the technique has not been validated against known fire histories spanning more than one fire interval. Here we compare fire records from 100 grasstrees with a 30‐year record (1973–2002) of fire data derived from satellite imagery in a region of sandplain shrubland vegetation near Eneabba in south‐western Australia. Fires occurred in eight of the 30 years of the satellite record, with sampled grasstrees burning between zero and four times. The grasstree and satellite records agreed in terms of the overall incidence of fires experienced over the 30‐year period, with the grasstree record matching the satellite record significantly better than chance. However, comparison of the grasstree and satellite records found substantial error in the rate of both false positives and false negatives. Grasstrees failed to identify fire in 83% of fire occurrences identified by the satellite record, down to 53% if an error of ±2 years in the attribution of year of fire was allowed. A similar proportion of grasstree fire incidents were not matched in the satellite record (false positives). The rate of false positives increased with time before present, suggesting a temporal bias in the grasstree record. It is clear that the grasstree record does reflect fire history to a degree, but that it contains at least as many false as true fire records and may tend towards over‐reporting the incidence of fire in the past.     

Leaf flushing during the dry season: the paradox of Asian monsoon forests

Aim Most deciduous species of dry monsoon forests in Thailand and India form new leaves 1–2 months before the first monsoon rains, during the hottest and driest part of the year around the spring equinox. Here we identify the proximate causes of this characteristic and counterintuitive ‘spring‐flushing’ of monsoon forest trees. Location Trees of 20 species were observed in semi‐deciduous dry monsoon forests of northern Thailand with a 5–6‐month‐long severe dry season and annual rainfall of 800–1500 mm. They were growing on dry ridges (dipterocarp–oak forest) or in moist gullies (mixed deciduous–evergreen forest) at 680–750 m altitude near Chiang Mai and in a dry lowland stand of Shorea siamensis in Uthai Thani province. Methods Two novel methods were developed to analyse temporal and spatial variation in vegetative dry‐season phenology indicative of differences in root access to subsoil water reserves. Results Evergreen and leaf exchanging species at cool, moist sites leafed soon after partial leaf shedding in January–February. Drought‐resistant dipterocarp species were evergreen at moist sites, deciduous at dry sites, and trees leafed soon after leaf shedding whenever subsoil water was available. Synchronous spring flushing of deciduous species around the spring equinox, as induced by increasing daylength, was common in Thailand's dipterocarp–oak forest and appears to be prevalent in Indian dry monsoon forests of the Deccan peninsula with its deep, water‐storing soils. Main conclusions In all observed species leafing during the dry season relied on subsoil water reserves, which buffer trees against prolonged climatic drought. Implicitly, rainfall periodicity, i.e. climate, is not the principal determinant of vegetative tree phenology. The establishment of new foliage before the summer rains is likely to optimize photosynthetic gain in dry monsoon forests with a relatively short, wet growing season.     

Regulation and acclimation of leaf gas exchange in a piñon–juniper woodland exposed to three different precipitation regimes

Leaf gas‐exchange regulation plays a central role in the ability of trees to survive drought, but forecasting the future response of gas exchange to prolonged drought is hampered by our lack of knowledge regarding potential acclimation. To investigate whether leaf gas‐exchange rates and sensitivity to drought acclimate to precipitation regimes, we measured the seasonal variations of leaf gas exchange in a mature piñon–juniper Pinus edulis–Juniperus monosperma woodland after 3 years of precipitation manipulation. We compared trees receiving ambient precipitation with those in an irrigated treatment (+30% of ambient precipitation) and a partial rainfall exclusion (?45%). Treatments significantly affected leaf water potential, stomatal conductance and photosynthesis for both isohydric piñon and anisohydric juniper. Leaf gas exchange acclimated to the precipitation regimes in both species. Maximum gas‐exchange rates under well‐watered conditions, leaf‐specific hydraulic conductance and leaf water potential at zero photosynthetic assimilation all decreased with decreasing precipitation. Despite their distinct drought resistance and stomatal regulation strategies, both species experienced hydraulic limitation on leaf gas exchange when precipitation decreased, leading to an intraspecific trade‐off between maximum photosynthetic assimilation and resistance of photosynthesis to drought. This response will be most detrimental to the carbon balance of piñon under predicted increases in aridity in the southwestern USA.     

Anomalies in grasstree fire history reconstructions for south‐western Australian vegetation

Abstract A new fire history for south‐western Australian sclerophyll forests was proposed recently based on grasstree (Xanthorrhoea preissii ) records that were interpreted to show a high frequency (3–5 years) ‘pre‐European burning regime’. Such a fire regime appears incompatible with the long‐term survival of many fire‐killed woody taxa. We investigated the local fire history in a small area of the northern sand‐plain shrub‐lands of south‐western Australia using 15 grasstrees, examining individual grasstree records in detail and comparing this with the decadal or averaged approach used in the original research, and with fire histories reconstructed from satellite images for the period since 1975. Results lead us to question the utility of the proposed grasstree fire history record as a tool for understanding past fire regimes for two reasons: First, inconsistencies in fire histories among individual grasstrees were considerable – some individuals were not burnt by known fires, while some apparently were burned many times during periods when others were not burned at all. Second, the grasstree record indicates a possible increase in patchiness of fires since 1930, while contemporary evidence and interpretations of the nature of Aboriginal (pre‐European) fire regimes would suggest the opposite. We believe that further research is needed to identify to what extent the grasstree method for reconstruction of fire histories can be used to re‐interpret how fire operated in many highly diverse ecosystems prior to European settlement of Australia.     

Climatic influences on fire regimes along a rain forest-to-xeric woodland gradient in northern Patagonia, Argentina

ABSTRACT. Influences of annual climatic variation on fire occurrence were examined along a rainfall gradient from temperate rainforest to xeric woodlands in northern Patagonia, Argentina. Fire chronologies were derived from fire scars on trees and related to tree-ring proxy records of climate over the period 1820–1974. Similarly, fire records of four Patagonian national parks for the period 1940–1988 were compared to instrumental weather data. Finally, the influences of broad-scale synoptic weather patterns on fire occurrence in northern Patagonia were explored.
Fire in Nothofagus rainforests is highly dependent on drought during the spring and summer of the same year in which fires occur and is less strongly favoured by drought during the spring of the previous year. The occurrence of fire in dry vegetation types near the steppe ecotone is less dependent on drought because even during years of normal weather fuels are thoroughly desiccated during the dry summer. In xeric Austrocedrus woodlands, fire occurrence and spread are promoted by droughts during the fire season and also appear to be favoured by above-average moisture conditions during the preceding 1 to 2 growing seasons which enhances fuel production. Thus, in the xeric woodlands fire is not simply dependent on drought but is favoured by greater climatic variability over time scales of several years.
Fire activity in northern Patagonia is greatly influenced by the intensity and latitudinal position of the subtropical high pressure cell of the southeast Pacific. Greater fire activity is associated with a more intense and more southerly located high pressure cell which blocks the influx of Pacific moisture into the continent. Although long-term changes in fire occurrence along the rainforest-to-xeric woodland gradient have been greatly influenced by human activities, annual variation in fire frequency and extent is also strongly influenced by annual climatic variation.     

Drought effects on the dynamics of leaf production and senescence in field-grown Medicago arborea and Medicago citrina

Forage shrub production in the Mediterranean region is frequently limited by soil water availability. To study plant responses to water deficit under such conditions is important for improving crop management and for selecting better yielding forage shrub species. Pre-dawn leaf water potential (Ψ_pd), plant leaf area (PLA), leaf area per stem (LA_s), leaf appearance rate (LAR₁;), leaf senescence rate (LSR), individual leaf area (LA) and maximal leaf elongation rate (LER) were studied throughout the year for Medicago arborea (MA) and Medicago citrina (MC) under irrigated (control) and low rainfall field conditions, at the experimental field site of the University of the Balearic Islands in Spain. With irrigation, the highest LA and LER were observed in autumn and spring and the lowest in winter and summer. LAR; was similar for both species in autumn and winter. Throughout the spring, LAR₁ was higher for MC compared to MA. PLA was similar for both species during the autumn, winter and spring seasons; however, during the summer PLA of MA was significantly reduced by 53%. This decline was attributed to higher leaf senescence during seed maturity. As a consequence, MC maintained higher leaf area (∼ 5 m² plant⁻¹) than MA (3 m² plant⁻¹). Under natural field conditions, soil water deficit increased from February to late August. The main effect of water stress was a marked reduction in LAR₁, LA and LER reflected in lower LA_s and PLA. Leaf area was severely reduced for both species during the summer, but much more intensively in MA, which developed full leaf senescence. Thus, MC maintained higher PLA than MA (0.5 m² compared to 0.0 m²). Throughout the year, but especially in the driest months, MC was superior to MA in leaf growth parameters and PLA.     

夏玉米三叶期持续干旱下不同叶位叶片含水量变化及其与光合作用的关系

植物干物质的累积依赖于群体光合速率,而群体光合速率又与单叶的光合能力密切有关。叶片光合作用与其含水量密切相关,目前关于不同叶位叶片含水量对持续干旱的响应及其与光合作用的关系还未见报道。以华北夏玉米郑单958为材料,设置6个不同灌水处理,模拟不同灌溉量下持续干旱对夏玉米不同叶位叶片生理特性的影响,分析夏玉米顶部开始的第一、三、五叶位叶片的水分变化及其与净光合速率的关系。结果表明:夏玉米不同叶位的叶片最大含水量不同,且随干旱进程的推进叶片含水量的变化速率也不同,第一叶的叶片含水量下降速率高于第三、第五叶,第一叶的最大含水量高于第三、五叶,且可进行光合产物积累的叶片含水量下限随叶位的增加而增大。同时,第一叶的叶片含水量与土壤水分呈显著相关,且与净光合速率的相关性也非常强。第一叶可进行光合产物积累的叶片水分下限(净光合速率为零时的叶片含水量)最小,表明其耐旱性最强,对干旱具有指导意义。研究结果可为提高冠层光合作用模拟的准确性及夏玉米干旱发生发展的监测预警提供参考。     

Environmental control of flowering periodicity in Costa Rican and Mexican tropical dry forests

Aim We analyse the proximate causes of the large variation in flowering periodicity among four tropical dry forests (TDF) and ask whether climatic periodicity or biotic interactions are the ultimate causes of flowering periodicity. Location The four TDFs in Guanacaste (Costa Rica), Yucatan, Jalisco and Sonora (Mexico) are characterized by a 5–7 month long dry season and are located along a gradient of increasing latitude (10–30°N). Methods To dissect the differences in flowering periodicity observed at the community level, individual tree species were assigned to ‘flowering types’, i.e. groups of species with characteristic flowering periods determined by similar combinations of environmental flowering cues and vegetative phenology. Results Large variation in the fraction of species and flowering types blooming during the dry and wet season, respectively, indicates large differences in the severity of seasonal drought among the four forests. In the dry upland forests of Jalisco, flowering of leafless trees remains suppressed during severe seasonal drought and is triggered by the first rains of the wet season. In the other forests, leaf shedding, exceptional rainfall or increasing daylength cause flowering of many deciduous species at various times during the dry season, well before the summer rains. The fraction of deciduous species leafing out during the summer rains and flowering when leafless during the dry season is largest in the Sonoran TDF. Main conclusions In many wide‐ranging species the phenotypic plasticity of flowering periodicity is large. The distinct temporal separation of spring flowering on leafless shoots and subsequent summer flushing represents a unique adaptation of tree development to climates with a relatively short rainy season and a long dry season. Seasonal variation in rainfall and soil water availability apparently constitutes not only the proximate, but also the ultimate cause of flowering periodicity, which is unlikely to have evolved in response to biotic adaptive pressures.     

Distribution and leaf growth of Thalassodendron pachyrhizum den Hartog in Southern Western Australia

The seagrass Thalassodendron pachyrhizum den Hartog grows on limestone reef platforms. Monthly leaf biomass was measured over 2 years and showed a strong seasonal variation with maximum biomass of 500 g m⁻². This seagrass loses all its leaves except for a bud and this characteristic was used to obtain a conservative estimate of productivity by change in standing stock. Leaf growth during the growing season was 6.6 mg Cg⁻¹ day⁻¹. Leaf length frequencies showed that new leaves formed during autumn (March–April). They grew from autumn until spring (November) and began to senesce in summer, followed by leaf fall in late summer (February–March).The growth of rhizome shoots “invading” free substratum space and the growth of new stems was measured for a 300-day period; about 9 leaves were produced in this period.     

Field-simulated droughts affect elemental leaf stoichiometry in Mediterranean forests and shrublands

This study evaluated the change induced by the year season and by experimentally induced drought on foliar element stoichiometry of the predominant woody species (Quercus ilex and Erica multiflora) in two Mediterranean ecosystems, a forest and a shrubland. This study is based in two long-term (11 yr) field experiments that simulated drought throughout the annual cycle.The effects of experimental droughts were significant but weaker than the changes produced by ontogeny and seasonality. Leaf N and P concentrations were higher in spring (the main growing season) in E. multiflora and, in Q. ilex in autumn (a period of additional growth). Leaf N:P ratios were lower in spring. In Q. ilex, the highest leaf K concentrations and leaf K:P ratios, and the lowest leaf C:K and N:K ratios, occurred in summer, the season when water stress was greatest. In E. multiflora, leaf K concentrations and K:P ratios were highest, and leaf C:K and N:K ratios were lowest in the plants from the drought-treated plots.The plant capacity to change K concentrations in response to seasonality and to drought is at least as great as the capacity to change N and P concentrations. The results underscore the importance of K and its stoichiometry relative to C, N and P in dry environments. These results indicate first, that N:P ratio shifts are not uniquely related to growth rate in Mediterranean plants but also to drought, and second, that there is a need to take into account K in ecological stoichiometry studies of terrestrial plants.     

Gas exchange of Agropyron desertorum: diurnal patterns and responses to water vapor gradient and temperature

Summary The response of leaf gas exchange to environmental variables were measured at different levels of drought stress for Agropyron desertorum, a naturalized perennial bunchgrass of the semiarid shrub steppes of western North America. Leaf conductance (stomatal plus boundary layer) was more sensitive to changes in water vapor gradient than to changes in leaf temperature. Assimilation was sensitive to both temperature and vapor gradient, and also appeared to be affected by conductance and high transpiration rates. The magnitudes of both assimilation and conductance decreased with increased drought conditions. Diurnal patterns of gas exchange were measured during 3 growing seasons. For a typical spring day with moderate leaf temperature and vapor gradient, diurnal patterns were similar for plants at different levels of soil water availability. Assimilation was relatively constant during most of the day, but conductance decreased during the afternoon. Total daily carbon gain was decreased to a lesser extent than daily water loss as soil water was depleted. Consequently, the ratio of daily carbon gain to daily water loss, i.e. daily water use efficiency, increased with decreased soil water content for diurnals under spring conditions. Diurnal patterns of assimilation for a typical summer day with high leaf temperature and vapor gradient differend from those for a spring day. An afternoon decrease in assimilation was typical during a summer day. Daily carbon gain, water use, and water use efficiency for summer diurnals decreased only under severe drought conditions. Almost complete recovery of assimilation and conductance occurred if leaf microclimate was ameliorated during the afternoon of either spring or summer diurnals. Thus, conditions responsible for a midday depression in assimilation during a single day did not have persistent effects on leaf gas exchange. Daily carbon gain of a typical summer day was restricted by leaf microclimate during the afternoon, but daily water use efficiency was not relatively increased by the amelioration of leaf microclimate.     

Increased evapotranspiration demand in a Mediterranean climate might cause a decline in fungal yields under global warming

Wild fungi play a critical role in forest ecosystems, and its recollection is a relevant economic activity. Understanding fungal response to climate is necessary in order to predict future fungal production in Mediterranean forests under climate change scenarios. We used a 15‐year data set to model the relationship between climate and epigeous fungal abundance and productivity, for mycorrhizal and saprotrophic guilds in a Mediterranean pine forest. The obtained models were used to predict fungal productivity for the 2021–2080 period by means of regional climate change models. Simple models based on early spring temperature and summer–autumn rainfall could provide accurate estimates for fungal abundance and productivity. Models including rainfall and climatic water balance showed similar results and explanatory power for the analyzed 15‐year period. However, their predictions for the 2021–2080 period diverged. Rainfall‐based models predicted a maintenance of fungal yield, whereas water balance‐based models predicted a steady decrease of fungal productivity under a global warming scenario. Under Mediterranean conditions fungi responded to weather conditions in two distinct periods: early spring and late summer–autumn, suggesting a bimodal pattern of growth. Saprotrophic and mycorrhizal fungi showed differences in the climatic control. Increased atmospheric evaporative demand due to global warming might lead to a drop in fungal yields during the 21st century.     

Drought and changes in leaf orientation for two California chaparral shrubs: Ceanothus megacarpus and Ceanothus crassifolius

Summary Predawn xylem pressure potentials were measured on two California chaparral shrubs, Ceanothus megacarpus and Ceanothus crassifolius, throughout the winter and spring growing season and into the summer drought. On the days xylem pressure potentials were measured, leaf orientation measurements were made on a population of marked leaves from the same shrubs. Predawn xylem pressure potentials decreased from -0.1 MPa in both species to -7.8 and -6.6 MPa in C. megacarpus and C. crassifolius, respectively, between May and August, 1981. Leaf inclinations became more vertical during this period with the greatest change observed in C. crassifolius. This change in leaf inclination was reversible, and, in the late winter and early spring, one year old leaves became more horizontal. Leaf azimuths were random and did not change seasonally. Simulations of solar radiation interception indicated that the increase in leaf inclination associated with summer drought reduced the absorption of solar radiation in August by 6% for C. megacarpus and 20% for C. crassifolius. Standard leaf energy budget calculations suggest that steep leaf inclinations would result in slightly lower leaf temperatures and transpiration rates under summer conditions.     

Different seasonal feeding activity of diverse herbivores in two temperate forests

The aim of the present study was to understand the effects of abiotic conditions on seasonal feeding activity of diverse herbivores on the same oak tree species in two different forests. We tracked changes in herbivore feeding activities on an oak tree species (Quercus serrata) in two localities: a low elevation small hillock forest patch (Muan, MN) and a middle elevation mountain forest patch (Mt. Jirisan, JR). A total of five sites were selected in each of two forest localities. Data for leaf expansion, leaf chemical qualities, leaf damage ratio, and numbers of lepidopteran caterpillars were collected during spring (May) and summer (July to August), 2012. Leaf expansion rate was higher at the low hillock forest than the mid‐mountain forest from spring to summer. Nitrogen and carbon content decreased seasonally at both localities. Lepidopteran larval diversity was high in the mid‐mountain forest, and two‐way ANOVA showed that species richness of lepidopteran larvae was significantly affected by the interaction between season and locality. Leaf damage by all herbivores was higher in the low hillock forest than the mid‐mountain forest in spring, but was higher in the mid‐mountain forest in summer. Relative proportion of general herbivores increased from spring to summer in the mid‐mountain forest, but not in the low hillock forest. Canonical Correspondence Analysis (CCA) ordination showed that altitude‐ and season‐related variables were significant species and environment interaction factors. Our data indicate that locality and temperature disproportionally affected the feeding activities of diverse herbivores in two different temperate forests.     

Effects of precipitation and soil water potential on drought deciduous phenology in the Kalahari

We utilized an ecosystem process model to investigate the influence of precipitation and soil water potential on vegetation phenology in the semi‐arid, drought‐deciduous ecosystems in the Kalahari region of South Africa. The timing of leaf flush was assumed to be the first day during which a rainfall event exceeded that day's estimate of potential evapotranspiration after a defined dry season. Leaf senescence was assumed to be a dynamic feedback between soil water potential and net plant carbon gain and was determined by dynamically modeling the effects of concomitant trends in soil water potential and net primary production on leaf area index (LAI). Model predictions of LAI were compared with satellite‐derived normalized difference vegetation indices (NDVI) for 3 years at two sites along the Kalahari transect. The mean absolute error for the prediction of modeled leaf flush date compared with leaf flush dates estimated from NDVI were 10.0 days for the Maun site and 39.3 days for the Tshane site. Correlations between model predicted 10‐day average LAI and 10‐day composite NDVI for both Maun and Tshane were high (ρ=0.67 and 0.74, respectively, P<0.001), suggesting that this method adequately predicts intra‐annual leaf area dynamics in these dry tropical ecosystems.     

Severe dry winter affects plant phenology and carbon balance of a cork oak woodland understorey

Mediterranean climates are prone to a great variation in yearly precipitation. The effects on ecosystem will depend on the severity and timing of droughts. In this study we questioned how an extreme dry winter affects the carbon flux in the understorey of a cork oak woodland? What is the seasonal contribution of understorey vegetation to ecosystem productivity?We used closed-system portable chambers to measure CO₂ exchange of the dominant shrub species (Cistus salviifolius, Cistus crispus and Ulex airensis), of the herbaceous layer and on bare soil in a cork oak woodland in central Portugal during the dry winter year of 2012. Shoot growth, leaf shedding, flower and fruit setting, above and belowground plant biomass were measured as well as seasonal leaf water potential. Eddy-covariance and micrometeorological data together with CO₂ exchange measurements were used to access the understorey species contribution to ecosystem gross primary productivity (GPP).The herbaceous layer productivity was severely affected by the dry winter, with half of the yearly maximum aboveground biomass in comparison with the 6 years site average. The semi-deciduous and evergreen shrubs showed desynchronized phenophases and lagged carbon uptake maxima. Whereas shallow-root shrubs exhibited opportunistic characteristics in exploiting the understorey light and water resources, deep rooted shrubs showed better water status but considerably lower assimilation rates. The contribution of understorey vegetation to ecosystem GPP was lower during summer with 14% and maximum during late spring, concomitantly with the lowest tree productivity due to tree canopy renewal. The herbaceous vegetation contribution to ecosystem GPP never exceeded 6% during this dry year stressing its sensitivity to winter and spring precipitation.Although shrubs are more resilient to precipitation variability when compared with the herbaceous vegetation, the contribution of the understorey vegetation to ecosystem GPP can be quite variable and will ultimately depend of tree density and canopy cover.     

Non-structural carbohydrate dynamics associated with antecedent stem water potential and air temperature in a dominant desert shrub

Non-structural carbohydrates (NSCs) are necessary for plant growth and affected by plant water status, but the temporal dynamics of water stress impacts on NSC are not well understood. We evaluated how seasonal NSC concentrations varied with plant water status (predawn xylem water potential, Ψ) and air temperature (T) in the evergreen desert shrub Larrea tridentata. Aboveground sugar and starch concentrations were measured weekly or monthly for ~1.5 years on 6–12 shrubs simultaneously instrumented with automated stem psychrometers; leaf photosynthesis (A_net) was measured monthly for 1 year. Leaf sugar increased during the dry, premonsoon period, associated with lower Ψ (greater water stress) and high T. Leaf sugar accumulation coincided with declines in leaf starch and stem sugar, suggesting the prioritization of leaf sugar during low photosynthetic uptake. Leaf starch was strongly correlated with A_net and peaked during the spring and monsoon seasons, while stem starch remained relatively constant except for depletion during the monsoon. Recent photosynthate appeared sufficient to support spring growth, while monsoon growth required the remobilization of stem starch reserves. The coordinated responses of different NSC fractions to water status, photosynthesis, and growth demands suggest that NSCs serve multiple functions under extreme environmental conditions, including severe drought.     

A quantitative comparison of two extremes in chaparral shrub phenology

We quantitatively compared phenology and water relations of a fully deciduous shrub, Styrax officinalis, and an evergreen shrub, Arctostaphylos glauca, in shared microsites in a sandstone outcrop in southern California during a multi-year drought. Pre-dawn xylem pressure potentials, Ψ_pd, were similar for the two species during most months of 2 years, but occasional differences and watering experiment results suggest S. officinalis may have phreatophytic roots that tap water in deep rock cavities and joint traces, while A. glauca may have primarily shallow roots. Neither species varied in maximum or minimum Ψ_pd between years of very different rainfall totals. Twig elongation and leaf production of S. officinalis began earlier during spring, and its leaves matured more quickly and more synchronously than A. glauca. Leaves lived a mean of 180 days for S. officinalis and 849 days for A. glauca. Leaf life spans varied among years in both species. S. officinalis leaf senescence occurred mostly in August and September and was not discernibly related to Ψ_pd or drought avoidance. A. glauca leaf senescence occurred throughout the year, but especially coincided with leaf production. In A. glauca most senescence occurred at the beginning of a leaf cohort's third growing season, but numbers of retained older leaves increased during 4 years of drought. Timing of twig elongation and leaf and flower production appeared to be related to current rainfall, but amount of twig growth and numbers of leaves and flowers produced appeared to be related to rainfall of the previous year or years for S. officinalis and, more complexly, A. glauca. Because of an interrupted pattern of flower production, number of flowers produced by A. glauca may be responsive to rainfall amounts during two environmental periods. Morphological differences, including much higher above ground allocation, many more leaves/twig, lower allocation to stem mass, and longer duration of leaves/year in A. glauca, are probably responsible for A. glauca having >6 times more above ground biomass per plant than S. officinalis. During the multi-year drought S. officinalis changed little, indicating either superb adaptation or growth pattern rigidity, while A. glauca underwent extensive phenological and morphological changes, indicating either stress or adaptive flexibility.     

Effects of the time of drought occurrence within the growing season on growth and survival of Pinus ponderosa seedlings

Key message

A drought event during spring produces a stronger and long lasting decrease in growth of ponderosa pine seedlings than a summer drought event. However, survival is not differentially affected.

Abstract

Although there is certainty about the increasing frequency of extreme climatic events, the consequences of changing patterns of drought events within the growing season on the growth and survival of different species are much less certain. In particular, little knowledge is available on the differential effect on tree seedlings of a drought event at different times within the growing season. The objective of this study was to quantify the effect of a drought event imposed at different times over the growing season on the growth, survival and some related morphological and physiological variables of Pinus ponderosa seedlings from two seed sources. Four treatments were applied: control conditions; spring drought; summer drought and spring plus summer drought (SpSuD). A drought event in spring reduced stem growth and biomass accumulation in ponderosa pine seedlings during the occurrence of the drought and afterwards, even when plant water status had recovered. The lack of growth recovery could not be associated with loss of stem hydraulic conductivity or reduction in stomatal conductance after drought. However, the spring drought did not differentially affect plant survival, as was the case with prolonged drought in the SpSuD treatment. The summer drought event had a significant but much smaller impact on plant growth. Our results suggest different consequences of a drought event in spring or in summer in ponderosa pine seedlings. This knowledge may be relevant to understand and predict tree seedlings responses to changing patterns of drought events within the growing season in the framework of climatic change.