Stem growth and phenology of two tropical trees in contrasting soil conditions

Background and aims

Phenological variations in tropical forests are usually explained by climate. Nevertheless, considering that soil water availability and nutrient content also influence plant water status and metabolism, soil conditions may also be important in the regulation of plant reproductive and vegetative activities over time. We investigated whether phenological patterns and stem growth differ in trees growing in two types of soil that display contrasting water and nutrient availability, namely, Gleysol (moist and nutrient-poor) and Cambisol (drier and nutrient-rich).

Methods

Phenological observations (flushing, leaf fall, flowering and fruiting) and stem diameter growth were recorded for 120 trees fitted with fixed dendrometer bands, at 15 days intervals, for 1 year. Two species of contrasting deciduousness were investigated: Senna multijuga (semi-deciduous) and Citharexylum myrianthum (deciduous).

Results

Both species were seasonal in all phenophases, regardless of soil type. However, frequency, mean date and intensity of phenophases varied according to soil type. Girth increment of C. myrianthum was four times greater in Cambisol than in Gleysol, whereas the type of soil had no significant effect on that of S. multijuga.

Conclusions

These results show that soil characteristics also play an important role in determining phenological patterns and growth and must be considered when analysing phenological patterns in tropical forests.     

Leaf phenology is associated with soil water availability and xylem traits in a tropical dry forest

In tropical dry forests, spatial heterogeneity in soil water availability is thought to determine interspecific differences in key components of resource use strategies, such as leaf phenology and xylem function. To understand the environmental drivers of variation in leaf phenology and xylem function, we explored the relation of soil water potential to topographic metrics derived from a digital elevation model. Subsequently, we compared nine xylem hydraulic, mechanical and storage traits in 18 species in three phenological classes (readily deciduous, tardily deciduous, and evergreen) in the dry tropical forest of Chamela, Mexico. Soil water potential was negatively correlated with elevation, insolation and water flow accumulation. Evergreen species characterized low-elevation moist sites, whereas deciduous species dominated hills and dry sites. Overall, evergreen species had lower xylem specific conductivity than deciduous species, and tardily deciduous species were different from readily deciduous and evergreen species in five of eight xylem traits. In dry tropical forests, water availability promotes divergence in leaf phenology and xylem traits, ranging from low wood density, evergreen species in moist sites to a combination of low wood density, readily deciduous species plus high wood density, tardily deciduous species in dry sites.     

Do litter manipulations affect leaf functional traits of savanna woody plants?

Plant litter is the layer composed of dead plant material that covers soil surfaces in terrestrial ecosystems. It is an important pool of essential nutrients for soil and plants, serving also as a protective layer on the soil surface. In this study, we investigated the effects of litter addition and removal on leaf functional traits of woody Neotropical savanna trees. We measured maximum photosynthesis (A _max), stomatal conductance (g _s), leaf transpiration (E), intrinsic water use efficiency (IWUE), specific leaf area (SLA), and chlorophyll content (CCI) in 15 species belonging to three different phenological groups (evergreen, briefly deciduous, and deciduous species) that were subjected to three distinct litter availability treatments (addition, removal, and control plots) in a Neotropical Savanna site in Brazil. Although SLA and CCI differed among phenology groups, they were not affected by the litter treatments. In contrast, when considered at the community level, we found that the availability of litter affected the leaf traits linked with the water status of the plants (E, g _s and IWUE). Plants in the litter removal plots exhibited lower g _s and E (25 % of reduction in comparison with control group) but higher IWUE, while plants in the litter addition plots had a 10 % decrease in IWUE but a 12 % increase in g _s and E compared with plants in control plots. Savanna woody plants responded promptly to litter manipulation by adjusting leaf water loss, which suggests that in the short term, changes in the amount of litter in Cerrado ecosystems can affect the soil water availability to the plant community.     

Variation in evergreen and deciduous species leaf phenology in Assam, India

In the present study phenological activities such as leaf and shoot growth, leaf pool size and leaf fall were observed for 3 years (March 2007–March 2010) in 19 tree species (13 evergreen and 6 deciduous species) in a wet tropical forest in Assam, India. The study area receives total annual average rainfall of 2,318 mm of which most rain fall (>70 %) occurs during June–September. Both the plant groups varied significantly on most of the shoot and leaf phenology parameters. In general, growth in deciduous species initiated before the evergreen species and showed a rapid shoot growth, leaf recruitment and leaf expansion compared to evergreen species. Leaf recruitment period was significantly different between evergreen (4.2 months) and deciduous species (6.8 months). Shoot elongation rate was also significantly different for evergreen and deciduous species (0.09 vs. 0.14 cm day^?1 shoot^?1). Leaf number per shoot was greater for deciduous species than for evergreen species (34 vs. 16 leaves). The average leaf life span of evergreen species (328 ± 32 days) was significantly greater than that of deciduous species (205 ± 16 days). The leaf fall in deciduous species was concentrated during the winter season (Nov–Feb), whereas evergreens retained their leaves until the next growing season. Although the climate of the study area supports evergreen forests, the strategies of the deciduous species such as faster leaf recruitment rate, longer leaf recruitment time, faster shoot elongation rate during favorable growing season and short leaf life span perhaps allows them to coexist with evergreen species that have the liberty to photosynthesize round the year. Variations in phenological strategies perhaps help to reduce the competition among evergreen and deciduous species for resources in these forests and enable the coexistence of both the groups.     

The spatial pattern of leaf phenology and its response to climate change in China

Leaf phenology has been shown to be one of the most important indicators of the effects of climate change on biological systems. Few such studies have, however, been published detailing the relationship between phenology and climate change in Asian contexts. With the aim of quantifying species’ phenological responsiveness to temperature and deepening understandings of spatial patterns of phenological and climate change in China, this study analyzes the first leaf date (FLD) and the leaf coloring date (LCD) from datasets of four woody plant species, Robinia pseudoacacia, Ulmus pumila, Salix babylonica, and Melia azedarach, collected from 1963 to 2009 at 47 Chinese Phenological Observation Network (CPON) stations spread across China (from 21° to 50° N). The results of this study show that changes in temperatures in the range of 39–43 days preceding the date of FLD of these plants affected annual variations in FLD, while annual variations in temperature in the range of 71–85 days preceding LCD of these plants affected the date of LCD. Average temperature sensitivity of FLD and LCD for these plants was ?3.93 to 3.30 days °C^?1 and 2.11 to 4.43 days °C^?1, respectively. Temperature sensitivity of FLD was found to be stronger at lower latitudes or altitude as well as in more continental climates, while the response of LCD showed no consistent pattern. Within the context of significant warming across China during the study period, FLD was found to have advanced by 5.44 days from 1960 to 2009; over the same period, LCD was found to have been delayed by 4.56 days. These findings indicate that the length of the growing season of the four plant species studied was extended by a total of 10.00 days from 1960 to 2009. They also indicate that phenological response to climate is highly heterogeneous spatially.     

Linking above- and belowground phenology of hybrid walnut growing along a climatic gradient in temperate agroforestry systems

Background and aims

Plant phenology is a sensitive indicator of plant response to climate change. Observations of phenological events belowground for most ecosystems are difficult to obtain and very little is known about the relationship between tree shoot and root phenology. We examined the influence of environmental factors on fine root production and mortality in relation with shoot phenology in hybrid walnut trees (Juglans sp.) growing in three different climates (oceanic, continental and Mediterranean) along a latitudinal gradient in France.

Methods

Eight rhizotrons were installed at each site for 21 months to monitor tree root dynamics. Root elongation rate (RER), root initiation quantity (RIQ) and root mortality quantity (RMQ) were recorded frequently using a scanner and time-lapse camera. Leaf phenology and stem radial growth were also measured. Fine roots were classified by topological order and 0–1 mm, 1–2 mm and 2–5 mm diameter classes and fine root longevity and risk of mortality were calculated during different periods over the year.

Results

Root growth was not synchronous with leaf phenology in any climate or either year, but was synchronous with stem growth during the late growing season. A distinct bimodal pattern of root growth was observed during the aerial growing season. Mean RER was driven by soil temperature measured in the month preceding root growth in the oceanic climate site only. However, mean RER was significantly correlated with mean soil water potential measured in the month preceding root growth at both Mediterranean (positive relationship) and oceanic (negative relationship) sites. Mean RIQ was significantly higher at both continental and Mediterranean sites compared to the oceanic site. Soil temperature was a driver of mean RIQ during the late growing season at continental and Mediterranean sites only. Mean RMQ increased significantly with decreasing soil water potential during the late aerial growing season at the continental site only. Mean root longevity at the continental site was significantly greater than for roots at the oceanic and Mediterranean sites. Roots in the 0–1 mm and 1–2 mm diameter classes lived for significantly shorter periods compared to those in the 2–5 mm diameter class. First order roots (i.e. the primary or parents roots) lived longer than lateral branch roots at the Mediterranean site only and first order roots in the 0–1 mm diameter class had 44.5% less risk of mortality than that of lateral roots for the same class of diameter.

Conclusions

We conclude that factors driving root RER were not the same between climates. Soil temperature was the best predictor of root initiation at continental and Mediterranean sites only, but drivers of root mortality remained largely undetermined.

     

Do the phenology and functional stem attributes of woody species allow for the identification of functional groups in the semiarid region of Brazil?

The phenology of tree species in environments that are subject to strong climatic seasonality is mainly determined by water availability, which may vary as a function of wood density. The relationship among phenology, water potential, wood density and the capacity of water storage in the stem were determined for woody species of caatinga vegetation (dry forest) in the semiarid region of NE Brazil. Leaf flush and fall, flowering and fruiting events were recorded over a 31-month period, and the water potential was measured over a two-year period. These data were related to precipitation, water availability in the soil and photoperiod. Seven deciduous species exhibited low wood density (DLWD,?<0.5?g?cm^?3), high capacity of water storage in the stem (until 250?% of the dry weight) and high water potential during the year, as opposed to 15 deciduous species that showed high wood density (DHWD,?≥0.5?g?cm^?3). Leaf flush, flowering and the fruiting of DHWD species were related to precipitation, whereas these phenological events occurred at the end of the dry season and/or the beginning of the rainy season for DLWD species and were related to the photoperiod. The two evergreen species showed variations of water potential that were intermediate between those of DHWD and DLWD deciduous species, leaf flush during the dry season and flowering at the end of dry season. These results suggest the existence of three functional groups: evergreen species, DHWD deciduous species and DLWD deciduous species.     

Eastern deciduous tree seedlings advance spring phenology in response to experimental warming,but not wetting,treatments

Changing global climate, particularly rising temperatures, has been linked through observations with advanced spring phenology in temperate regions. We experimentally tested if regional climate change predictions of increased temperature and precipitation alter the spring phenology of eastern US tree seedlings. This study reports the results of a 3-year-field experiment designed to study the responses of eastern deciduous tree species planted in a post-harvest environment to a 2 °C increase in temperature and a 20 % increase in precipitation. Species were monitored for timing of germination and leaf out in four treatment combinations (ambient, warmed, irrigated, and warmed + irrigated) on 16 plots located in a recently harvested central Pennsylvania forest. The 2 °C warming advanced day of seed germination by an average of 2 weeks and seedling leaf out by 10 days among all species (both p < 0.001). However, increased precipitation did not result in a significant change in spring phenology. Species responded uniquely to treatments, with germination advancing in three of five species in response to warming and leaf out advancing in six of six species. Southern species projected to expand northward into the study region with rising temperatures did not show responses to warming treatments that would provide them an advantage over current resident species. Timing of germination and leaf out varied among years of the experiment, most likely driven by year-to-year variability in spring temperatures. The climate change experiment highlighted the potential of a moderate 2 °C temperature increase to advance spring phenology of deciduous tree seedlings by up to 2 weeks, with a lack of a phenological response to a 20 % increase in precipitation.     

Phenology of temperate trees in tropical climates

Several North American broad-leaved tree species range from the northern United States at 47°N to moist tropical montane forests in Mexico and Central America at 15–20°N. Along this gradient the average minimum temperatures of the coldest month (T _Jan), which characterize annual variation in temperature, increase from –10 to 12°C and tree phenology changes from deciduous to leaf-exchanging or evergreen in the southern range with a year-long growing season. Between 30 and 45°N, the time of bud break is highly correlated with T _Jan and bud break can be reliably predicted for the week in which mean minimum temperature rises to 7°C. Temperature-dependent deciduous phenology—and hence the validity of temperature-driven phenology models—terminates in southern North America near 30°N, where T _Jan>7°C enables growth of tropical trees and cultivation of frost-sensitive citrus fruits. In tropical climates most temperate broad-leaved species exchange old for new leaves within a few weeks in January-February, i.e., their phenology becomes similar to that of tropical leaf-exchanging species. Leaf buds of the southern ecotypes of these temperate species are therefore not winter-dormant and have no chilling requirement. As in many tropical trees, bud break of Celtis, Quercus and Fagus growing in warm climates is induced in early spring by increasing daylength. In tropical climates vegetative phenology is determined mainly by leaf longevity, seasonal variation in water stress and day length. As water stress during the dry season varies widely with soil water storage, climate-driven models cannot predict tree phenology in the tropics and tropical tree phenology does not constitute a useful indicator of global warming.     

Source water, phenology and growth of two tropical dry forest tree species growing on shallow karst soils

Seasonally dry tropical forests are dominated by deciduous and evergreen tree species with a wide range of leaf phenology. We hypothesized that Piscidia piscipula is able to extend leaf senescence until later in the dry season due to deeper and more reliable water sources than Gymnopodium floribundum, which loses leaves earlier in the dry season. Physiological performance was assessed as timing of leaf production and loss, growth, leaf water potential, depth of water uptake determined by stable isotopes, and leaf stable isotopic composition of carbon (δ¹³C) and oxygen (δ¹⁸O). P. piscipula took water primarily from shallow sources, whereas G. floribundum took water from shallow and deep sources. The greatest variation in water sources occurred during the onset of the dry season, when G. floribundum was shedding old leaves and growing new leaves, but P. piscipula maintained its leaves from the previous wet season. P. piscipula showed greater relative growth rate, greater leaf expansion rates, and more negative predawn and midday water potentials than G. floribundum. P. piscipula also exhibited greater leaf organic δ¹³C and lower δ¹⁸O values, indicating that the decrease in photosynthetic carbon isotope discrimination was associated with greater stomatal conductance and greater photosynthesis. Our results indicate that the contrasting early and late dry season leaf loss phenology of these two species is not simply determined by rooting depth, but rather a more complicated suite of characteristics based on opportunistic use of dynamic water sources, maximizing carbon gain, and maintenance of water potential during the dry season.     

Reproductive phenology of two co‐occurring Neotropical mountain grasslands

Aim

Climate tends to explain phenological variations in tropical ecosystems. However, water availability and nutrient content in soil strongly affect plant communities, especially those on old, climatically buffered, infertile landscapes (OCBILs), and may impact these ecosystems’ plant reproductive phenology over time. Here, we compare the reproductive phenology of sandy and stony tropical grasslands, two co‐occurring herbaceous communities of the campo rupestreOCBILs. We asked whether flowering, fruiting and dispersal are seasonal in both grasslands, and whether these phenophases differ due to variations in soil properties. We also asked whether the phenological strategies and the number of flowers and fruits differ between these two grasslands as soil conditions vary.

Location

Serra do Cipó, Minas Gerais, Brazil.

Methods

The phenology of herbaceous species of sandy and stony grasslands was monitored monthly over two consecutive years.

Results

Plants on sandy and stony grasslands flowered and fruited throughout the year. We did not find a distinct seasonal pattern at the community level of either studied grassland. However, flowering, fruiting and seed dissemination occurred in stony grasslands mainly during the rainy season, while sandy grassland species flowered in both seasons and fruited and disseminated seed mainly during the dry season, as observed in other savanna vegetation types in the Cerrado. Flower and fruit production was higher in sandy grasslands than in stony grasslands, which may be linked to higher water retention in sandy grassland soils. In both communities, species of Cyperaceae, Eriocaulaceae and Xyridaceae contributed most to overall production, whereas Poaceae and Velloziaceae, two important families in campo rupestre, barely participated in the reproductive phenology during our 2‐yr survey.

Conclusions

Despite a strong seasonal climate, there was no reproductive seasonal pattern at the community level in campo rupestre. This first investigation of Neotropical grassland phenology indicates that the differences in soil content may constrain the grassland reproductive phenology and restrict reproduction of stony grassland species to the most favourable season. Further studies of grassland phenology are necessary to disentangle the relative importance of soil, climate and other triggers, especially fire.     

Periodicity of leaf growth and leaf dry mass changes in the evergreen and deciduous species of Southern Assam,India

The study described patterns of leaf dry mass change, leaf mass per area (LMA), relative growth rate and leaf life span (LL) for 14 evergreen and 7 deciduous species of a tropical forest of Southern Assam, India. Leaf expansion in both the groups was, in general, completed before June (i.e. well before the onset of monsoon rains). Although leaf dry mass during leaf initiation phase was significantly higher (P < 0.01) in evergreen species than in deciduous species, at the time of full leaf expansion, average leaf dry mass relative to the peak leaf dry mass, realised by the evergreen species was lower (66 %) than for deciduous species (76 %). Leaf dry mass increase in both groups continued after leaf full expansion. Evergreen species had a longer leaf dry mass steady phase than deciduous species (2–6 vs 2–3 months). Average LMA of mature leaves for evergreen species (77.43 g m^?2) was significantly greater than that of deciduous species (48.43 g m^?2). LL ranged from 165 days in Gmelina arborea (deciduous) to 509 days in Dipterocarpus turbinatus (evergreen). LMA was correlated positively with LL, indicating that evergreen species with higher leaf construction cost retain leaves for longer period to pay back. The average leaf dry mass loss before leaf shedding was greater (P < 0.01) for deciduous species (30.29 %) than for evergreen species (18.31 %). Although the cost of leaf construction in deciduous species was lower than for evergreen species, they replace leaves at a faster rate. Deciduous species perhaps compensate the cost involved in faster leaf replacement through higher reabsorption of dry mass during senescence, which they remobilise to initiate growth in the following spring when soil resources remain limiting.     

Does irrigation affect leaf phenology in deciduous and evergreen trees of the savannas of northern Australia?

Abstract Soil moisture was augmented experimentally during two successive dry seasons and the intervening wet season in a humid tropical savanna in Darwin, northern Australia. Leaf phenology was monitored in four common tree species Termmalia ferdinandiana and Planchonia careya (both deciduous species), and Eucalyptus miniata and Eucalyptus tetrodonta (both evergreen species). Irrigation produced consistently significant effects in only T. ferdinandiana. In this species leaf-flush was significantly earlier, canopy decline and leaf-fall were significantly later and the attainment of full canopy was earlier in irrigated compared with non-irrigated trees. Litterfall, and the seasonal patterns of contraction and expansion of stems (a measure of stem water status or storage) were not significantly affected by irrigation in any species. Leaf longevity in the deciduous species was 6–8 months; some eucalypt leaves lived for approximately 1 year, but none lived longer than 18 months. Irrigation had relatively little effect on leaf longevity. While variation in soil moisture is a potentially important cue to both leaf-fall and leaf-flush, stem water status and climatic factors such as vapour pressure deficit may also be important climatic cues to phenological behaviour.     

拔节期干旱和复水对春玉米物候的影响及其生理生态机制

物候不仅是气候变化的指示指标,也是作物模型的关键参数。现有研究主要关注物候变化与气候环境因子的关系,关于植物物候变化的生理生态机制研究很少。基于春玉米拔节期干旱与不同时间(抽雄期和吐丝期)复水的田间模拟试验分析表明:(1)不同时间复水均使灌浆期延长,乳熟期推迟(9d),表明物候对前期水分胁迫存在记忆。(2)干旱条件下叶片净光合速率(P_n)、蒸腾速率(T_r)、气孔导度(G_s)和相对叶绿素含量(SPAD)均随物候进程呈先降后升再降趋势,且均在抽雄期达到极小值;不同时间复水均使P_n、T_r和G_s在吐丝期达到极大值,而SPAD则在灌浆期达到极大值;叶水势(LWP)随干旱进程整体呈下降趋势,不同时间复水均只是减缓了其下降速度,表明LWP可用于描述物候对前期水分胁迫的记忆。(3)通径分析和决策系数分析表明,P_n是最主要的物候影响因子,而影响LWP的土壤相对湿度(RSWC)则是物候的主要控制因子,物候的变化是由P_n的累积变化引起...     

Differential ecophysiological response of deciduous shrubs and a graminoid to long-term experimental snow reductions and additions in moist acidic tundra, Northern Alaska

Changes in winter precipitation that include both decreases and increases in winter snow are underway across the Arctic. In this study, we used a 14-year experiment that has increased and decreased winter snow in the moist acidic tussock tundra of northern Alaska to understand impacts of variation in winter snow depth on summer leaf-level ecophysiology of two deciduous shrubs and a graminoid species, including: instantaneous rates of leaf gas exchange, and δ¹³C, δ¹⁵N, and nitrogen (N) concentrations of Betula nana, Salix pulchra, and Eriophorum vaginatum. Leaf-level measurements were complemented by measurements of canopy leaf area index (LAI) and depth of thaw. Reductions in snow lowered summer leaf photosynthesis, conductance, and transpiration rates by up to 40 % compared to ambient and deep snow conditions for Eriophorum vaginatum, and reduced Salix pulchra conductance and transpiration by up to 49 %. In contrast, Betula nana exhibited no changes in leaf gas exchange in response to lower or deeper snow. Canopy LAI increased with added snow, while reduced winter snow resulted in lower growing season soil temperatures and reduced thaw depths. Our findings indicate that the spatial and temporal variability of future snow depth will have individualistic consequences for leaf-level C fixation and water flux by tundra species, and that these responses will be manifested over the longer term by changes in canopy traits, depth of thaw, soil C and N processes, and trace gas (CO₂ and H₂O) exchanges between the tundra and the atmosphere.     

Reproductive phenology of coastal plain Atlantic forest vegetation: comparisons from seashore to foothills

The diversity of tropical forest plant phenology has called the attention of researchers for a long time. We continue investigating the factors that drive phenological diversity on a wide scale, but we are unaware of the variation of plant reproductive phenology at a fine spatial scale despite the high spatial variation in species composition and abundance in tropical rainforests. We addressed fine scale variability by investigating the reproductive phenology of three contiguous vegetations across the Atlantic rainforest coastal plain in Southeastern Brazil. We asked whether the vegetations differed in composition and abundance of species, the microenvironmental conditions and the reproductive phenology, and how their phenology is related to regional and local microenvironmental factors. The study was conducted from September 2007 to August 2009 at three contiguous sites: (1) seashore dominated by scrub vegetation, (2) intermediary covered by restinga forest and (3) foothills covered by restinga pre-montane transitional forest. We conducted the microenvironmental, plant and phenological survey within 30 transects of 25 m × 4 m (10 per site). We detected significant differences in floristic, microenvironment and reproductive phenology among the three vegetations. The microenvironment determines the spatial diversity observed in the structure and composition of the flora, which in turn determines the distinctive flowering and fruiting peaks of each vegetation (phenological diversity). There was an exchange of species providing flowers and fruits across the vegetation complex. We conclude that plant reproductive patterns as described in most phenological studies (without concern about the microenvironmental variation) may conceal the fine scale temporal phenological diversity of highly diverse tropical vegetation. This phenological diversity should be taken into account when generating sensor-derived phenologies and when trying to understand tropical vegetation responses to environmental changes.     

Edaphic constraints on seed germination and emergence of three <Emphasis Type="Italic">Acacia</Emphasis> species for dryland restoration in Saudi Arabia

In situ edaphic factors affecting seed germination and seedling emergence of three framework species of Acacia were investigated with the intent of developing fundamental and scalable restoration capacity for Arabian dryland restoration. Direct seeding represents the most efficient means to restore vegetation at the landscape scale and this study provides insight into edaphic and ecological limitations, as well as effective protocols governing the use of native seeds for restoration in hyper-arid environments. The study was conducted in extant Acacia woodland habitat on conserved land (Thumamah Nature Park) in close proximity to Riyadh, Saudi Arabia. Broad-scale direct seeding using un- and pretreated Acacia gerrardii, A. tortilis, and A. ehrenbergiana seed, and two seed burial depths were implemented across three sites with distinct soil surface characteristics. Eight weeks post-sowing, random samples for each species × seed treatment × burial depth combination were excavated, sieved, and categorized as follows: failed to germinate, germinated but died prior to emerging, or successfully emerged. We show that germination and emergence of Acacia gerrardii, A. tortilis, and A. ehrenbergiana were driven by a three-way interaction among species, site, and seed burial depth. Treating seed with the signaling compound Moddus did not have a definitive effect, positive or negative, on any of the species investigated. Acacia gerrardii was the only species that exhibited widespread emergence, though emergence was not consistent across sites or burial depths. Germination was highest in disturbed soil (up to 69% for A. gerrardii), but very few (<2%) successfully emerged; a greater proportion of germinants in sandy soil emerged (up to 44% for A. gerrardii) even though the overall germination was less. Though species-dependent, a 2-cm sowing depth was most effective in sand; while in disturbed soil, sowing depths of 1 and 2 cm were comparable; and no germination was observed in gravelly clay soil. Sandy soil exhibited rapid water infiltration (107.6 mm min^?1), and post-sowing surface crusting was a non-factor (0.44 kg cm^?2). Disturbed soil exhibited moderate water infiltration (1.46 mm min^?1) and post-sowing surface crusting was double that of sand (0.88 kg cm^?2) and restrictive on seedling emergence. Gravelly clay exhibited extremely poor water infiltration (0.12 mm min^?1), and surface crusting was severe (4.49 kg cm^?2) and an order of magnitude greater than sand. The medium-coarse sand fraction, a key driver of the observed soil surface processes, was greatest in sand (55%) and significantly less and uniform in the disturbed (22%) and gravelly clay (22%) soils. Our findings demonstrate that soil surface characteristics and associated processes can dictate ecological processes at depths as shallow as 1–2 cm, and that soil crusts that slow water infiltration and impede seedling emergence rapidly reconstitute after disturbance; both are important considerations for restoring dryland vegetation.     

The shifting phenological landscape: Within‐ and between‐species variation in leaf emergence in a mixed‐deciduous woodland

Many organisms rely on synchronizing the timing of their life‐history events with those of other trophic levels—known as phenological matching—for survival or successful reproduction. In temperate deciduous forests, the extent of matching with the budburst date of key tree species is of particular relevance for many herbivorous insects and, in turn, insectivorous birds. In order to understand the ecological and evolutionary forces operating in these systems, we require knowledge of the factors influencing leaf emergence of tree communities. However, little is known about how phenology at the level of individual trees varies across landscapes, or how consistent this spatial variation is between different tree species. Here, we use field observations, collected over 2 years, to characterize within‐ and between‐species differences in spring phenology for 825 trees of six species (Quercus robur, Fraxinus excelsior, Fagus sylvatica, Betula pendula, Corylus avellana, and Acer pseudoplatanus) in a 385‐ha woodland. We explore environmental predictors of individual variation in budburst date and bud development rate and establish how these phenological traits vary over space. Trees of all species showed markedly consistent individual differences in their budburst timing. Bud development rate also varied considerably between individuals and was repeatable in oak, beech, and sycamore. We identified multiple predictors of budburst date including altitude, local temperature, and soil type, but none were universal across species. Furthermore, we found no evidence for interspecific covariance of phenology over space within the woodland. These analyses suggest that phenological landscapes are highly complex, varying over small spatial scales both within and between species. Such spatial variation in vegetation phenology is likely to influence patterns of selection on phenology within populations of consumers. Knowledge of the factors shaping the phenological environments experienced by animals is therefore likely to be key in understanding how these evolutionary processes operate.     

Influence of elevation on canopy transpiration of temperate deciduous forests in a complex mountainous terrain of South Korea

Background and aims

Variations in microclimate and soil characteristics on mountain slopes influence forest structure and function. Precipitation, incoming solar radiation and relative humidity change along a mountain slope. Equally, soil depth and the amount of stored soil moisture vary. The objective of this study was to examine the impacts of these factors on forest water use in mountainous terrains.

Methods

Transpiration of four temperate deciduous forest stands located at different elevations in South Korea was monitored with a sap flow technique throughout the growing season in 2010. The study sites were located on the north slope at 450 m (450 N), 650 m (650 N), and 950 m (950 N). To examine the effect of aspect, an additional site with a southern aspect was studied at 650 m (650S). All the sites were dominated by Quercus species, with leaf area index (L) ranging between 5 ? 6 m² m^?2.

Results

Rainfall increased, while air temperature (T _A ) and daytime vapor pressure deficit (D) decreased with increasing elevation. We did not observe any gradients in solar radiation (R _S ), soil moisture and sap flux density of the individual trees (J _st ) with an elevational gradient. Sapwood area (A _S ), i.e., hydro-active xylem area, and daily maximum tree water use (max TWU) increased non-linearly with increasing diameter at breast height (DBH). Neither A _S nor max TWU varied among tree species or along the elevation. The total annual canopy transpiration (E _C ) was 175, 115, 110, and 90 mm for 450 N, 650 N, 650S, and 950 N, respectively. E _C declined with increasing elevation as a result of decreasing length of the growing season, D, and T _A along the elevation. Significantly (P < 0.001) higher stomatal sensitivity to changes in D was found at the 950 N, leading to lower annual E _C and lower water use efficiency (WUE) at this elevation.

Conclusions

We conclude that differences in E _C exist along the mountain slope studied, corresponding to changing T _A , D, length of the growing season, and stomatal sensitivity to D, which should be considered when establishing forest catchment water balances.     

Vulnerability of phenological synchrony between plants and pollinators in an alpine ecosystem

The relationship between flowering phenology and abundance of bumble bees (Bombus spp.) was investigated using 2 years of phenological data collected in an alpine region of northern Japan. Abundance of Bombus species was observed along a fixed transect throughout the flowering season. The number of flowering species was closely related to the floral resources for pollinators at the community scale. In the year with typical weather, the first flowering peak corresponded to the emergence time of queen bees from hibernation, while the second flowering peak corresponded to the active period of worker bees. In the year with an unusually warm spring, however, phenological synchrony between plants and bees was disrupted. Estimated emergence of queen bees was 10 days earlier than the first flowering date owing to earlier soil thawing and warming. However, subsequent worker emergence was delayed, indicating slower colony development. The flowering season finished 2 weeks earlier in the warm-spring year in response to earlier snowmelt. A common resident species in the alpine environment, B. hypocrita sapporoensis, flexibly responded to the yearly fluctuation of flowering. In contrast, population dynamics of other Bombus species were out of synchrony with the flowering: their frequencies were highest at the end of the flowering season in the warm-spring year. Therefore, phenological mismatch between flowers and pollinators is evident during warm years, which may become more prevalent in a warmer climate. To understand the mechanism of phenological mismatch in the pollination system of the alpine ecosystem, ground temperature, snowmelt regime, and life cycle of pollinators are key factors.