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.     

Tests of associational defence provided by hairy plants for glabrous plants of Arabidopsis halleri subsp. gemmifera against insect herbivores

1. Trichome‐producing (hairy) and trichomeless (glabrous) plants of Arabidopsis halleri subsp. gemmifera were investigated to test whether plant resistance to herbivory depends on the plants' phenotypes and/or the phenotypes of neighbouring plants (associational effects). 2. A common garden experiment was conducted in which the relative frequency of hairy and glabrous plants was manipulated. Two species of leaf‐chewing insects (larvae of a white butterfly and a cabbage sawfly) were found less often on hairy plants than on glabrous plants. By contrast, the numbers of aphids and flea beetles did not differ significantly between hairy and glabrous plants. For none of these insects did abundance depend on the frequency of the two plant morphs. 3. A field survey was conducted in two natural populations of A. halleri. In the first population, a species of white butterfly was the dominant herbivore, and hairy plants incurred less leaf damage than glabrous plants across 2 years. By contrast, in the other population, where flea beetles were dominant, there were no consistent differences in leaf damage between the two types of plants. In neither of the two populations was any evidence found of associational effects. 4. This study did not provide any conclusive evidence of associational effects of anti‐herbivore resistance, but it was discovered that trichomes can confer resistance to certain herbivores. Given the results of previous work by the authors on associational effects against a flightless leaf beetle, such associational effects of the trichome dimorphism of A. halleri were herbivore‐specific.     

Landscape controls on the timing of spring,autumn, and growing season length in mid‐Atlantic forests

The timing of spring leaf development, trajectories of summer leaf area, and the timing of autumn senescence have profound impacts to the water, carbon, and energy balance of ecosystems, and are likely influenced by global climate change. Limited field‐based and remote‐sensing observations have suggested complex spatial patterns related to geographic features that influence climate. However, much of this variability occurs at spatial scales that inhibit a detailed understanding of even the dominant drivers. Recognizing these limitations, we used nonlinear inverse modeling of medium‐resolution remote sensing data, organized by day of year, to explore the influence of climate‐related landscape factors on the timing of spring and autumn leaf‐area trajectories in mid‐Atlantic, USA forests. We also examined the extent to which declining summer greenness (greendown) degrades the precision and accuracy of observations of autumn offset of greenness. Of the dominant drivers of landscape phenology, elevation was the strongest, explaining up to 70% of the spatial variation in the onset of greenness. Urban land cover was second in importance, influencing spring onset and autumn offset to a distance of 32 km from large cities. Distance to tidal water also influenced phenological timing, but only within ~5 km of shorelines. Additionally, we observed that (i) growing season length unexpectedly increases with increasing elevation at elevations below 275 m; (ii) along gradients in urban land cover, timing of autumn offset has a stronger effect on growing season length than does timing of spring onset; and (iii) summer greendown introduces bias and uncertainty into observations of the autumn offset of greenness. These results demonstrate the power of medium grain analyses of landscape‐scale phenology for understanding environmental controls on growing season length, and predicting how these might be affected by climate change.     

Divergent phenological response to hydroclimate variability in forested mountain watersheds

Mountain watersheds are primary sources of freshwater, carbon sequestration, and other ecosystem services. There is significant interest in the effects of climate change and variability on these processes over short to long time scales. Much of the impact of hydroclimate variability in forest ecosystems is manifested in vegetation dynamics in space and time. In steep terrain, leaf phenology responds to topoclimate in complex ways, and can produce specific and measurable shifts in landscape forest patterns. The onset of spring is usually delayed at a specific rate with increasing elevation (often called Hopkins' Law; Hopkins, 1918), reflecting the dominant controls of temperature on greenup timing. Contrary with greenup, leaf senescence shows inconsistent trends along elevation gradients. Here, we present mechanisms and an explanation for this variability and its significance for ecosystem patterns and services in response to climate. We use moderate‐resolution imaging spectro‐radiometer (MODIS) Normalized Difference Vegetation Index (NDVI) data to derive landscape‐induced phenological patterns over topoclimate gradients in a humid temperate broadleaf forest in southern Appalachians. These phenological patterns are validated with different sets of field observations. Our data demonstrate that divergent behavior of leaf senescence with elevation is closely related to late growing season hydroclimate variability in temperature and water balance patterns. Specifically, a drier late growing season is associated with earlier leaf senescence at low elevation than at middle elevation. The effect of drought stress on vegetation senescence timing also leads to tighter coupling between growing season length and ecosystem water use estimated from observed precipitation and runoff generation. This study indicates increased late growing season drought may be leading to divergent ecosystem response between high and low elevation forests. Landscape‐induced phenological patterns are easily observed over wide areas and may be used as a unique diagnostic for sources of ecosystem vulnerability and sensitivity to hydroclimate change.     

Modification of Vegetative Phenology in a Tropical Semi‐deciduous Forest by Abnormal Drought and Rain1

The control of vegetative phenology in tropical trees is not well understood. In dry forest trees, leaf abscission may be enhanced by advanced leaf age, increasing water stress, or declining photoperiod. Normally, it is impossible to dissect the effects of each of these variables because most leaves are shed during the early dry season when day length is near its minimum and leaves are relatively old. The 1997 El‐Niño Southern Oscillation caused a ten‐week long, severe abnormal drought from June to August in the semi‐deciduous forests of Guanacaste, Costa Rica. We monitored the effect of this drought on phenology and water status of trees with young leaves and compared modifications of phenology in trees of different functional types with the pattern observed during the regular dry season. Although deciduous trees at dry sites were severely water stressed (Ψstem < ‐7MPa) and their mesic leaves remained wilted for more than two months, these and all other trees retained all leaves during the abnormal drought. Many trees exchanged leaves three to four months earlier than normal during the wet period after the abnormal drought and shed leaves again during the regular dry season. Irrigation and an exceptional 70 mm rainfall during the mid‐dry season 1998/1999 caused bud break and flushing in all leafless trees except dormant stem succulents. The complex interactions between leaf age and water stress, the principal determinants of leaf abscission, were found to vary widely among trees of different functional types.     

Seasonal Foliage Changes in the Eastern Amazon Basin Detected from Landsat Thematic Mapper Satellite Images1

Seasonal changes in tropical forests are difficult to measure from the ground, especially in areas of high species diversity and low phenological synchrony. Satellite images, which integrate individual tree canopies and cover a large spatial extent, facilitate tests for stand-level canopy phenology. Variability in near-infrated radiance (TM bands 4 and 5) of several distinct vegetation types was used to detect seasonal changes in a series of three Landsat Thematic Mapper (TM) images from the wet season to the dry season in Marabá, Brazil (eastern Amazon basin). Despite different atmospheric and instrumental conditions among the images, spectral changes were distinguishable. A phenological process (leaf aging, leaf drop, water stress) was determined from the spectral changes for each vegetation type. Changes in the spectral properties suggest that during the dry season, upland terra firme forest increased the rate of leaf exchange and some riparian vegetation was deciduous. Terra firme forest that had been altered by penetration of fires from nearby pastures increased in leaf biomass over a 14-month period. This study shows that a time series of images can provide information on temporal changes in primary vegetation and guide field studies to investigate seasonal changes that may not be detectable from the ground.     

Associational effects and the maintenance of polymorphism in plant defense against herbivores: review and evidence

Many plant species have evolved defense traits against herbivores. Associational effects (AEs) refer to a kind of apparent interaction where the herbivory risk to a focal plant species depends on the composition of other plant species in a neighborhood. Despite ample evidence for AEs between different plant species, this point of view has rarely been applied to polymorphism in defense traits within a plant species. The purpose of this review is to highlight an overlooked role of conspecific AEs in maintaining polymorphism in antiherbivore defense. First, I present a general review of AE between plant species and its role in the coexistence of plant species. This viewpoint of AE can be applied to genetic polymorphism within a plant species, as it causes frequency‐ and density‐dependent herbivory between multiple plant types. Second, I introduce a case study of conspecific AEs in the trichome‐producing (hairy) and glabrous plants of Arabidopsis halleri subsp. gemmifera. Laboratory and semi‐field experiments illustrated that AEs against the brassica leaf beetle Phaedon brassicae mediate a minority advantage in defense and fitness between hairy and glabrous plants. Combined with a statistical modeling approach, field observation revealed that conspecific AEs can maintain the trichome dimorphism via negative frequency‐dependent selection in a plant population. Finally, I discuss spatial and temporal scales at which AEs contribute to shaping genetic variation in antiherbivore defense in a plant metapopulation. Based on the review and evidence, I suggest that AEs play a key role in the maintenance of genetic variation within a plant species.     

Evidence for facultative deciduousness in Colophospermum mopane in semi‐arid African savannas

Leaf phenology dictates the time available for carbon assimilation, transpiration and nutrient uptake in plants. Understanding the environmental cues that control phenology is therefore vital for predicting climate‐related changes to plant and ecosystem function. In contrast to temperate systems, and to a lesser degree, tropical forest systems, the cues initiating leaf drop in tropical savannas are poorly studied. We investigated the cues for leaf fall in a tropical monodominant arid savanna species, Colophospermum mopane, using an irrigation experiment. We tracked soil moisture, solar radiation, air temperature, leaf water status, leaf health and leaf carbon balance through the dry season in both irrigated and control plants. Water was the primary cue driving leaf loss of C. mopane rather than temperature or light. Trees watered throughout the dry season retained their canopies. These leaves remained functional and continued photosynthesis throughout the dry season. Leaf carbon acquisition rates did not decline with leaf age but were affected by soil moisture availability and temperature. Leaf loss did not occur when leaf carbon gain was zero, or when a particular leaf carbon threshold was reached. Colophospermum mopane is facultatively deciduous as water availability determines leaf drop in this widespread arid savanna species. Obligate deciduosity is not the only successful strategy in climates with a long dry season.     

Frequency-dependent herbivory by a leaf beetle,Phaedon brassicae,on hairy and glabrous plants of Arabidopsis halleri subsp. gemmifera

Frequency-dependent prey choice by natural enemies may influence the coexistence of multiple prey types, but little is known about whether frequency-dependent foraging choice occurs in herbivory on plants showing resistance polymorphism within a single population. Here we examined frequency-dependent foraging by a crucifer-feeding leaf beetle, Phaedon brassicae, on trichome-producing (hairy) and trichomeless (glabrous) plants coexisting within a natural population of the perennial herb Arabidopsis halleri subsp. gemmifera. Larvae of P. brassicae fed on hairy leaves showed slower growth than those fed on glabrous leaves. Although adult beetles consumed similar amounts of leaves when they were fed either hairy or glabrous leaves in no-choice conditions, our choice experiment showed that adult beetles fed at less than the proportionally expected level on hairy leaves compared to glabrous leaves when the hairy leaves were less or equally abundant. Both types of leaves were consumed at the proportionally expected levels when the hairy leaves were more abundant than the glabrous leaves. In a natural population, the leaf damage on the hairy plants was negatively correlated with the local proportion of the glabrous plants in a 1-m diameter patch across 2 years, while correlations between the leaf damage on the glabrous plants and their proportion differed between the 2 years. Additionally, we found five glucosinolates in leaves of A. halleri, but their accumulation did not differ between hairy and glabrous plants. Our experimental results indicate that hairy plants incur less herbivory by P. brassicae when glabrous plants are abundant. The field pattern provides evidence suggestive of frequency-dependent herbivory acting on hairy plants. The present study highlights one of the putative mechanisms of maintaining plant resistance polymorphism.     

Influence of leaf trichomes on boundary layer conductance and gas‐exchange characteristics in Metrosideros polymorpha (Myrtaceae)

The amount of trichomes on the leaves of Metrosideros polymorpha varies enormously, ranging from 0 to ca 150 g/m² across environmental gradients on the island of Hawaii. Pubescent individuals are abundant in dry areas or on young lava flows, whereas glabrous individuals are abundant in wet areas or on developed soils. To understand the adaptive advantages of pubescent individuals in arid environments, we addressed the following questions: (1) whether leaf trichomes increase the boundary layer resistance to gas diffusion, which in turn reduces the transpiration rate and increases water‐use efficiency (WUE); and (2) whether pubescent individuals have other associated leaf and shoot traits that have adaptive significance in arid environments. We made detailed ecophysiological measurements on M. polymorpha in three populations in habitats that varied in aridity. We found a large allocation of leaf mass to trichomes, up to 33 percent at the arid site, but our analyses showed that trichomes had small effects (1–9%) on gas exchange and negligible effects on WUE, suggesting the trichomes may have roles beyond increasing WUE. However, pubescent individuals did have higher Rubisco amount and a lower leaf‐to‐ sapwood area ratio, which are considered adaptive in arid environments. These results suggest that pubescent individuals of M. polymorpha are indeed adapted to arid environments with changes in a suite of traits. The adaptive significance of the enormous variation in amounts of trichomes remains unclear and may be related to functions other than increasing boundary layer resistance.     

Functional differences between summer and winter season rain assessed with MODIS‐derived phenology in a semi‐arid region

Questions: We asked several linked questions about phenology and precipitation relationships at local, landscape, and regional spatial scales within individual seasons, between seasons, and between year temporal scales. (1) How do winter and summer phenological patterns vary in response to total seasonal rainfall? (2) How are phenological rates affected by the previous season rainfall? (3) How does phenological variability differ at landscape and regional spatial scales and at season and inter‐annual temporal scales? Location: Southern Arizona, USA. Methods: We compared satellite‐derived phenological variation between 38 distinct 625‐km² landscapes distributed in the northern Sonoran Desert region from 2000 to 2007. Regression analyses were used to identify relationships between landscape phenology dynamics in response to precipitation variability across multiple spatial and temporal scales. Results: While both summer and winter seasons show increases of peak greenness and peak growth with more precipitation, the timing of peak growth was advanced with more precipitation in winter, while the timing of peak greenness was advanced with more precipitation in summer. Surprisingly, summer maximum growth was negatively affected by winter precipitation. The spatial variations between summer and winter phenology were similar in magnitude and response. Larger‐scale spatial and temporal variation showed strong differences in precipitation patterns; however the magnitudes of phenological spatial variability in these two seasons were similar. Conclusions: Vegetation patterns were clearly coupled to precipitation variability, with distinct responses at alternative spatial and temporal scales. Disaggregating vegetation into phenological variation, spanning value, timing, and integrated components revealed substantial complexity in precipitation‐phenological relationships.     

Variation in the phenology of photosynthesis among eastern white pine provenances in response to warming

In higher‐latitude trees, temperature and photoperiod control the beginning and end of the photosynthetically active season. Elevated temperature (ET) has advanced spring warming and delayed autumn cooling while photoperiod remains unchanged. We assessed the effects of warming on the length of the photosynthetically active season of three provenances of Pinus strobus L. seedlings from different latitudes, and evaluated the accuracy of the photochemical reflectance index (PRI) and the chlorophyll/carotenoid index (CCI) for tracking the predicted variation in spring and autumn phenology of photosynthesis among provenances. Seedlings from northern, local and southern P. strobus provenances were planted in a temperature‐free‐air‐controlled enhancement (T‐FACE) experiment and exposed to ET (+1.5/3°C; day/night). Over 18 months, we assessed photosynthetic phenology by measuring chlorophyll fluorescence, gas exchange, leaf spectral reflectance and pigment content. During autumn, all seedlings regardless of provenance followed the same sequence of phenological events with the initial downregulation of photosynthesis, followed by the modulation of non‐photochemical quenching and associated adjustments of zeaxanthin pool sizes. However, the timing of autumn downregulation differed between provenances, with delayed onset in the southern provenance (SP) and earlier onset in the northern relative to the local provenance, indicating that photoperiod at the provenance origin is a dominant factor controlling autumn phenology. Experimental warming further delayed the downregulation of photosynthesis during autumn in the SP. A provenance effect during spring was also observed but was generally not significant. The vegetation indices PRI and CCI were both effective at tracking the seasonal variations of energy partitioning in needles and the differences of carotenoid pigments indicative of the stress status of needles. These results demonstrate that PRI and CCI can be useful tools for monitoring conifer phenology and for the remote monitoring of the length of the photosynthetically active season of conifers in a changing climate.     

Dimorphism in trichome production of <Emphasis Type="Italic">Persicaria lapathifolia</Emphasis> var. <Emphasis Type="Italic">lapathifolia</Emphasis> and Its multiple effects on a leaf beetle

Polymorphisms in plants are main factors that determine the diversity of associated animal communities and their population dynamics. Typically, Persicaria lapathifolia var. lapathifolia (Polygonaceae) has no trichomes on leaf surfaces (glabrous type), but a hairy type does sometimes occur. Based on a cultivation experiment, the presence or absence of trichomes is clarified to be under genetic control. To reveal the defensive function of trichomes against herbivores, laboratory experiments were conducted using a major herbivore, Galerucella grisescens (Coleoptera: Chrysomelidae). In both choice and no-choice feeding tests, the glabrous type was significantly more consumed by G. grisescens adults, while the hairy type was not consumed. In the hairy leaf treatment, larval duration tended to become longer, the adult body weight became significantly lower, and adults laid significantly more eggs than in the glabrous leaf treatment. Hairy leaves contained significantly more total phenolics and condensed tannins than glabrous leaves, suggesting that the hairy type allocates more resources for physical and chemical defence. Because no significant differences in leaf consumption were detected in the feeding experiment using powdered host leaves, G. grisescens seems to have adapted to the chemical defences of P. lapathifolia var. lapathifolia. These results clearly indicate that leaf trichomes of P. lapathifolia var. lapathifolia effectively act as a physical defence against G. grisescens.     

Fine‐scale frequency differentiation along a herbivory gradient in the trichome dimorphism of a wild Arabidopsis

Geographic variation is commonly observed in plant resistance traits, where plant species might experience different selection pressure across a heterogeneous landscape. Arabidopsis halleri subsp. gemmifera is dimorphic for trichome production, generating two morphs, trichome‐producing (hairy) and trichomeless (glabrous) plants. Trichomes of A. halleri are known to confer resistance against the white butterfly, cabbage sawfly, and brassica leaf beetle, but not against flea beetles. We combined leaf damage, microclimate, and microsatellite loci data of 26 A. halleri populations in central Japan, to explore factors responsible for fine‐scale geographic variation in the morph frequency. We found that hairy plants were less damaged than glabrous plants within populations, but the among‐site variation was the most significant source of variation in the individual‐level damage. Fixation index () of a putative trichome locus exhibited a significant divergence along population‐level damage with an exception of an outlier population, inferring the local adaptation to herbivory. Notably, this outlier was a population wherein our previous study reported a balancing role of the brassica leaf beetle Phaedon brassicae on the morph frequency. This differentiation of the trichome locus was unrelated to neutral genetic differentiation (evaluated by of microsatellite loci) and meteorological factors (including temperature and solar radiation). The present findings, combined with those of our previous work, provide suggestive evidence that herbivore‐driven divergence and occasional outbreak of a specific herbivore have jointly contributed to the ecogeographic pattern in the frequency of two morphs.     

Greater deciduous shrub abundance extends tundra peak season and increases modeled net CO2 uptake

Satellite studies of the terrestrial Arctic report increased summer greening and longer overall growing and peak seasons since the 1980s, which increases productivity and the period of carbon uptake. These trends are attributed to increasing air temperatures and reduced snow cover duration in spring and fall. Concurrently, deciduous shrubs are becoming increasingly abundant in tundra landscapes, which may also impact canopy phenology and productivity. Our aim was to determine the influence of greater deciduous shrub abundance on tundra canopy phenology and subsequent impacts on net ecosystem carbon exchange (NEE) during the growing and peak seasons in the arctic foothills region of Alaska. We compared deciduous shrub‐dominated and evergreen/graminoid‐dominated community‐level canopy phenology throughout the growing season using the normalized difference vegetation index (NDVI). We used a tundra plant‐community‐specific leaf area index (LAI) model to estimate LAI throughout the green season and a tundra‐specific NEE model to estimate the impact of greater deciduous shrub abundance and associated shifts in both leaf area and canopy phenology on tundra carbon flux. We found that deciduous shrub canopies reached the onset of peak greenness 13 days earlier and the onset of senescence 3 days earlier compared to evergreen/graminoid canopies, resulting in a 10‐day extension of the peak season. The combined effect of the longer peak season and greater leaf area of deciduous shrub canopies almost tripled the modeled net carbon uptake of deciduous shrub communities compared to evergreen/graminoid communities, while the longer peak season alone resulted in 84% greater carbon uptake in deciduous shrub communities. These results suggest that greater deciduous shrub abundance increases carbon uptake not only due to greater leaf area, but also due to an extension of the period of peak greenness, which extends the period of maximum carbon uptake.     

How do leaf wetting events affect gas exchange and leaf lifespan of plants from seasonally dry tropical vegetation?

• Foliar uptake of dew is likely an important mechanism of water acquisition for plants from tropical dry environments. However, there is still limited experimental evidence describing the anatomical pathways involved in this process and the effects of this water subsidy on the maintenance of gas exchange and leaf lifespan of species from seasonally dry tropical vegetation such as the Brazilian caatinga.
• We performed scanning electron, bright‐field and confocal microscopic analyses and used apoplastic tracers to examine the foliar water uptake (FWU) routes in four woody species with different foliar phenology and widely distributed in the caatinga. Leaves of plants subjected to water stress were exposed to dew simulation to evaluate the effects of the FWU on leaf water potentials, gas exchange and leaf lifespan.
• All species absorbed water through their leaf cuticles and/or peltate trichomes but FWU capacity differed among species. Leaf wetting by dew increased leaf lifespan duration up to 36 days compared to plants in the drought treatment. A positive effect on leaf gas exchange and new leaf production was only observed in the anisohydric and evergreen species.
• We showed that leaf wetting by dew is relevant for the physiology and leaf lifespan of plants from seasonally dry tropical vegetation, especially for evergreen species.

     

Water inputs across a tropical montane landscape in Veracruz,Mexico: synergistic effects of land cover,rain and fog seasonality,and interannual precipitation variability

Land‐cover change can alter the spatiotemporal distribution of water inputs to mountain ecosystems, an important control on land‐surface and land‐atmosphere hydrologic fluxes. In eastern Mexico, we examined the influence of three widespread land‐cover types, montane cloud forest, coffee agroforestry, and cleared areas, on total and net water inputs to soil. Stand structural characteristics, as well as rain, fog, stemflow, and throughfall (water that falls through the canopy) water fluxes were measured across 11 sites during wet and dry seasons from 2005 to 2008. Land‐cover type had a significant effect on annual and seasonal net throughfall (NTF <0=canopy water retention plus canopy evaporation; NTF >0=fog water deposition). Forest canopies retained and/or lost to evaporation (i.e. NTF<0) five‐ to 11‐fold more water than coffee agroforests. Moreover, stemflow was fourfold higher under coffee shade than forest trees. Precipitation seasonality and phenological patterns determined the magnitude of these land‐cover differences, as well as their implications for the hydrologic cycle. Significant negative relationships were found between NTF and tree leaf area index (R²=0.38, P<0.002), NTF and stand basal area (R²=0.664, P<0.002), and stemflow and epiphyte loading (R²=0.414, P<0.001). These findings indicate that leaf and epiphyte surface area reductions associated with forest conversion decrease canopy water retention/evaporation, thereby increasing throughfall and stemflow inputs to soil. Interannual precipitation variability also altered patterns of water redistribution across this landscape. Storms and hurricanes resulted in little difference in forest‐coffee wet season NTF, while El Niño Southern Oscillation was associated with a twofold increase in dry season rain and fog throughfall water deposition. In montane headwater regions, changes in water delivery to canopies and soils may affect infiltration, runoff, and evapotranspiration, with implications for provisioning (e.g. water supply) and regulating (e.g. flood mitigation) ecosystem services.     

Measured and modelled leaf and stand‐scale productivity across a soil moisture gradient and a severe drought

Environmental controls on carbon dynamics operate at a range of interacting scales from the leaf to landscape. The key questions of this study addressed the influence of water and nitrogen (N) availability on Pinus palustris (Mill.) physiology and primary productivity across leaf and canopy scales, linking the soil‐plant‐atmosphere (SPA) model to leaf and stand‐scale flux and leaf trait/canopy data. We present previously unreported ecophysiological parameters (e.g. V_cmax and J_max) for P. palustris and the first modelled estimates of its annual gross primary productivity (GPP) across xeric and mesic sites and under extreme drought. Annual mesic site P. palustris GPP was ～23% greater than at the xeric site. However, at the leaf level, xeric trees had higher net photosynthetic rates, and water and light use efficiency. At the canopy scale, GPP was limited by light interception (canopy level), but co‐limited by nitrogen and water at the leaf level. Contrary to expectations, the impacts of an intense growing season drought were greater at the mesic site. Modelling indicated a 10% greater decrease in mesic GPP compared with the xeric site. Xeric P. palustris trees exhibited drought‐tolerant behaviour that contrasted with mesic trees' drought‐avoidance behaviour.     

Spatial and temporal variations in the timing of leaf replacement in a Quercus cornelius‐mulleri population

Abstract. The phenology of spring leaf replacement was studied in a population of 46 evergreen scrub oaks (Quercus cornelius‐mulleri) at the edge of the Mojave Desert in each of five years over the period 1990–2001. The oaks occupied a site that spanned rocky slopes to sandy bajadas. The site receives variable annual rainfall (estimated 12‐yr average 195 mm; range in study years 67–706 mm). The spatial coordinates of all individuals were recorded, and in April, when leaf replacement was underway, individual replacement phenologies were assessed. Shrub sizes were recorded in three separate years, and in 2001 water potentials were measured. Individuals vary greatly in their timing of leaf replacement within years, and also between years. Many individuals with an early phenology one year are significantly later in the following year, and vice versa. While we detected weak influences on leaf replacement phenology due to shrub size, position within the site, and a genetic component, stronger influences were attributable to the phenology of the shrub in prior years, and to the phenology of neighbours within years. Neighbouring individuals that are close and/or large are significantly disparate in phenology, with one early and the other late. A potential mechanism of local resource depletion associated with costs to an early phenology is discussed.     

Ceropegia thaithongiae sp. nov. (Apocynaceae,Asclepiadoideae) from northern Thailand and preliminary observations on impacts of climate change on its only known population

A new pendulous herb, Ceropegia thaithongiae Kidyoo (Apocynaceae, Asclepiadoideae), is described from northern Thailand. This plant is strictly endemic to Chiang Dao Wildlife Sanctuary and critically endangered owing to it living in an ecosystem affected by frequent natural disturbances. It is here described, illustrated and compared to the similar species, C. mairei. Both species have fusiform rootstocks, 1–2‐flowered cymes, deeply bifid outer corona lobes and linear, hairy inner corona lobes. However, the new species can be distinguished by its glabrous stem, leaf covered with hairs on both surfaces and glabrous corolla tube and lobes.