Shrub facilitation drives tree establishment in a semiarid fog‐dependent ecosystem

Questions

The exceptional occurrence of tall rain forest patches on foggy coastal mountaintops, surrounded by extensive xerophytic shrublands, suggests an important role of plant–plant interactions in the origin and persistence of these patches in semi‐arid Chile. We asked whether facilitation by shrubs can explain the growth and survival of rain forest tree species, and whether shrub effects depend on the identity of the shrub species itself, the drought tolerance of the tree species and the position of shrubs in regard to wind direction.

Location

Open area–shrubland–forest matrix, Fray Jorge Forest National Park, Chile.

Methods

We recorded survival after 12 years of a ~3600 tree saplings plantation (originally ~30‐cm tall individuals) of Aextoxicon punctatum, Myrceugenia correifolia and Drimys winteri placed outside forests, beneath the shrub Baccharis vernalis, and in open (shrub‐free) areas. We assessed the effects of neighbouring shrubs and soil humidity on survival and growth along a gradient related to the direction of fog movement.

Results

B. vernalis had a clear facilitative effect on tree establishment and survival since, after ~12 years, saplings only survived beneath the shrub canopy. Long‐term survival strongly depended on tree species identity, drought tolerance and position along the soil moisture gradient, with higher survival of A. punctatum (>35%) and M. correifolia (>14%) at sites on wind‐ and fog‐exposed shrubland areas. Sites occupied by the shrub Aristeguietia salvia were unsuitable for trees, presumably due to drier conditions than under B. vernalis.

Conclusions

Interactions between shrubs and fog‐dependent tree species in dry areas revealed a strong, long‐lasting facilitation effect on planted tree's survival and growth. Shrubs acted as benefactors, providing sites suitable for tree growth. Sapling mortality in the shrubland interior was caused by lower soil moisture, the consequence of lower fog loads in the air and thus insufficient facilitation. While B. vernalis was a key ecosystem engineer (nurse) and intercepted fog water that dripped to trees planted underneath, drier sites with A. salvia were unsuitable for trees. Consequently, nurse effects related to water input are strongly site and species specific, with facilitation by shrubs providing a plausible explanation for the initiation of forest patches in this semi‐arid landscape.     

Rain Forest Islands in the Chilean Semiarid Region: Fog-dependency, Ecosystem Persistence and Tree Regeneration

Tree presence in semiarid ecosystems is generally constrained by insufficient annual rainfall. However, in semiarid Chile, rainforest patches dominated by Aextoxicon punctatum are unexpectedly found on coastal mountaintops (450–600 m) at 30°S, surrounded by a xerophytic vegetation matrix that receives only 147 mm of annual precipitation. It has been proposed that these forests persist as a result of fog-water inputs. If so, then because fog-water deposition is spatially heterogeneous and shows strong edge effects, the potential environmental gradient created by the direction of fog input should determine forest structure and tree regeneration patterns. To investigate this hypothesis, we measured fog inputs, forest structural attributes (age and size distribution, basal area, and coarse woody debris), and tree regeneration in three different habitats: the windward edges (WE), leeward edges (LE), and the interior (center) of rainforest patches varying in area from 0.2 to 22 ha. Mean fog-water input was estimated from passive collectors over 1 year in WE and LE of patches. Tree regeneration was greater in the WE and forest interior (FI) and decreased toward the LE of patches, following a marked pattern of decline in fog inputs. Older trees and coarse woody debris were concentrated in the FI and LE of patches. Tree regeneration and patch structure appear to be largely controlled by fog-input direction and edge effects. We propose that forest patches may be slowly growing toward the incoming fog edge, while dying at the opposite edge.     

Regeneration patterns and persistence of the fog-dependent Fray Jorge forest in semiarid Chile during the past two centuries

The persistence of rainforest patches at Fray Jorge National Park (FJNP) in semiarid Chile (30°40′S), a region receiving approximately 147 mm of annual rainfall, has been a source of concern among forest managers. These forests are likely dependent on water inputs from oceanic fog and their persistence seems uncertain in the face of climate change. Here, we assessed tree radial growth and establishment during the last two centuries and their relation to trends in climate and canopy disturbance. Such evaluation is critical to understanding the dynamics of these semiarid ecosystems in response to climate change. We analyzed forest structure of six forest patches (0.2–22 ha) in FJNP based on sampling within 0.1 ha permanent plots. For the main canopy species, the endemic Aextoxicon punctatum (Aextoxicaceae), we used tree‐ring analysis to assess establishment periods, tree ages, growing trends and their relation to El Niño Southern Oscillation (ENSO), rainfall, and disturbance. The population dynamics of A. punctatum can be described by a continuous regeneration mode. Regeneration of A. punctatum was sensitive to different canopy structures. Growth release patterns suggest the absence of large scale human impact. Radial growth and establishment of A. punctatum were weakly correlated with rainfall and ENSO. If water limits forests patch persistence, patches are likely dependent on the combination of fog and rain water inputs. Forest patches have regenerated continuously for at least 250 years, despite large fluctuations in rainfall driven by ENSO and a regional decline in rainfall during the last century. Because of the positive influence on fog interception, forest structure should be preserved under any future climate scenario. Future research in FJNP should prioritize quantifying the long‐term trends of fog water deposition on forests patches. Fog modeling is crucial for understanding the interplay among physical drivers of water inputs under climate change.     

Strong hydraulic segmentation and leaf senescence due to dehydration may trigger die-back in Nothofagus dombeyi under severe droughts: a comparison with the co-occurring Austrocedrus chilensis

Key message

Total leaf hydraulic dysfunction during severe drought could lead to die-back in N. dombeyi , while hydraulic traits of A. chilensis allow it to operate far from the threshold of total hydraulic failure.

Abstract

Die-back was observed in South America temperate forests during one of the most severe droughts of the 20th century (1998–1999). During this drought Austrocedrus chilensis trees survived, whereas trees of the co-occurring species (Nothofagus dombeyi) experienced symptoms of water stress, such as leaf wilting and abscission, before tree die-back occurred. We compared hydraulic traits of these two species (a conifer and an angiosperm species, respectively) in a forest stand located close to the region with records of N. dombeyi mass mortality. We asked whether different hydraulic traits exhibited by the two species could help explain their contrasting survivorship rates. Austrocedrus chilensis had wide leaf safety margins, which appear to be the consequence of relatively high leaf-and-stem capacitance, large stored water use, strong stomatal control and ability to recover from embolism-induced loss of leaf hydraulic capacity. On the other hand, N. dombeyi even though had a stem hydraulic threshold of ?6.7 MPa before reaching substantial hydraulic failure (P₈₈), leaf P₈₈ occurred at leaf water potentials of only ?2 MPa, which probably are reached during anomalous droughts. Massive mortality in N. dombeyi appears to be the result of the total loss of leaf hydraulic conductance leading to leaf dehydration and leaf drop. Drought occurs during the summer and it is highly likely that N. dombeyi cannot recover its photosynthetic surface to produce carbohydrates required to avoid tissue injury in the winter season with subfreezing temperatures. Strong hydraulic segmentation in N. dombeyi does not seem to have an adaptive value to survive severe droughts.     

Effects of herbivory and patch size on tree seedling survivorship in a fog-dependent coastal rainforest in semiarid Chile

The landscape (matrix) surrounding habitat fragments critically affects the biodiversity of those fragments due to biotic interchange and physical effects. However, to date, there have been only a limited number of studies on plant–animal interactions in fragmented landscapes, particularly on how tree seedling herbivory is affected by fragmentation. We have examined this question in a fog-dependent mosaic of rainforest fragments located on coastal mountaintops of semiarid Chile (30°S), where the effects of the surrounding semiarid matrix and forest patch size (0.1–22 ha) on tree seedling survival were simultaneously addressed. The rainforest is strongly dominated by the endemic evergreen tree species Aextoxicon punctatum (Olivillo, approx. 80% of basal area). To assess the magnitudes and causes of Olivillo seedling mortality, we set up a field experiment where 512 tree seedlings of known age were transplanted into four forest fragments of different sizes in four 1.5 × 3-m plots per patch; one-half of each plot was fenced off with chicken wire to exclude small mammals. The plots were monitored for 22 months. Overall, 50% of the plants died during the experiment. The exclusion of small mammals from the plots increased seedling survival by 25%, with the effect being greater in smaller patches where matrix-dwelling herbivores are more abundant. This experiment highlights the important role of the surrounding matrix in affecting the persistence of trees in forest fragments. Because herbivores from the matrix cause greater tree seedling mortality in small patches, their effects must be taken into account in forest conservation–restoration plans.     

Functional coordination between branch hydraulic properties and leaf functional traits in miombo woodlands: implications for water stress management and species habitat preference

We investigated functional coordination between branch hydraulic properties and leaf functional traits among nine miombo woodlands canopy tree species differing in habitat preference and phenology. Specifically, we were seeking to answer the question: are branch hydraulic properties coordinated with leaf functional traits linked to plant drought tolerance in seasonally dry tropical forests and what are the implications for species habitat preference? The hydraulic properties investigated in this study were stem area specific hydraulic conductivity (K _S), Huber value (H _v), and xylem cavitation vulnerability (??₅₀). The leaf functional traits measured were specific leaf area (SLA), leaf dry matter content (LDMC), and mean leaf area (MLA). Generalists displayed significantly (P?<?0.05) higher cavitation resistance (??₅₀) and SLA, but lower sapwood specific hydraulic conductivity (K _S), leaf specific conductivity (K _L), MLA, and LDMC than mesic specialists. Although MLA was uncorrelated with ??₅₀, we found significant (P?<?0.05) positive and negative correlations between plant hydraulic properties and leaf functional traits linked to plant drought tolerance ability, indicating that the interactions between branch hydraulics and leaf functional traits related to plant drought tolerance ability may influence tree species habitat preference in water-limited ecosystems.     

Amazonia trees have limited capacity to acclimate plant hydraulic properties in response to long‐term drought

The fate of tropical forests under future climate change is dependent on the capacity of their trees to adjust to drier conditions. The capacity of trees to withstand drought is likely to be determined by traits associated with their hydraulic systems. However, data on whether tropical trees can adjust hydraulic traits when experiencing drought remain rare. We measured plant hydraulic traits (e.g. hydraulic conductivity and embolism resistance) and plant hydraulic system status (e.g. leaf water potential, native embolism and safety margin) on >150 trees from 12 genera (36 species) and spanning a stem size range from 14 to 68 cm diameter at breast height at the world's only long‐running tropical forest drought experiment. Hydraulic traits showed no adjustment following 15 years of experimentally imposed moisture deficit. This failure to adjust resulted in these drought‐stressed trees experiencing significantly lower leaf water potentials, and higher, but variable, levels of native embolism in the branches. This result suggests that hydraulic damage caused by elevated levels of embolism is likely to be one of the key drivers of drought‐induced mortality following long‐term soil moisture deficit. We demonstrate that some hydraulic traits changed with tree size, however, the direction and magnitude of the change was controlled by taxonomic identity. Our results suggest that Amazonian trees, both small and large, have limited capacity to acclimate their hydraulic systems to future droughts, potentially making them more at risk of drought‐induced mortality.     

Drought and cold spells trigger dieback of temperate oak and beech forests in northern Spain

Dieback in temperate forests is understudied, despite this biome is predicted to be increasingly affected by more extreme climate events in a warmer world. To evaluate the potential drivers of dieback we reconstructed changes in radial growth and intrinsic water-use efficiency (iWUE) from stable isotopes in tree rings. Particularly, we compared tree size, radial-growth trends, growth responses to climate (temperature, precipitation, cloudiness, number of foggy days) and drought, and changes in iWUE of declining and non-declining trees showing contrasting canopy dieback and defoliation. This comparison was done in six temperate forests located in northern Spain and based on three broadleaved tree species (Quercus robur, Quercus humilis, Fagus sylvatica). Declining trees presented lower radial-growth rates than their non-declining counterparts and tended to show lower growth variability, but not in all sites. The growth divergence between declining and non-declining trees was significant and lasted more in Q. robur (15–30 years) than in F. sylvatica (5–10 years) sites. Dieback was linked to summer drought and associated atmospheric patterns, but in the wettest Q. robur sites cold spells contributed to the growth decline. In contrast, F. sylvatica was the species most responsive to summer drought in terms of growth reduction followed by Q. humilis which showed coupled changes in growth and iWUE as a function of tree vigour. Low growth rates and higher iWUE characterized declining Q. robur and F. sylvatica trees. However, declining F. sylvatica trees became less water-use efficient close to the dieback onset, which could indicate impending tree death. In temperate forests, dieback and growth decline can be triggered by climate extremes such as dry and cold spells, and amplified by climate warming and rising drought stress.     

Wood anatomy and carbon‐isotope discrimination support long‐term hydraulic deterioration as a major cause of drought‐induced dieback

Hydraulic impairment due to xylem embolism and carbon starvation are the two proposed mechanisms explaining drought‐induced forest dieback and tree death. Here, we evaluate the relative role played by these two mechanisms in the long‐term by quantifying wood‐anatomical traits (tracheid size and area of parenchyma rays) and estimating the intrinsic water‐use efficiency (iWUE) from carbon isotopic discrimination. We selected silver fir and Scots pine stands in NE Spain with ongoing dieback processes and compared trees showing contrasting vigour (declining vs nondeclining trees). In both species earlywood tracheids in declining trees showed smaller lumen area with thicker cell wall, inducing a lower theoretical hydraulic conductivity. Parenchyma ray area was similar between the two vigour classes. Wet spring and summer conditions promoted the formation of larger lumen areas, particularly in the case of nondeclining trees. Declining silver firs presented a lower iWUE than conspecific nondeclining trees, but the reverse pattern was observed in Scots pine. The described patterns in wood anatomical traits and iWUE are coherent with a long‐lasting deterioration of the hydraulic system in declining trees prior to their dieback. Retrospective quantifications of lumen area permit to forecast dieback in declining trees 2–5 decades before growth decline started. Wood anatomical traits provide a robust tool to reconstruct the long‐term capacity of trees to withstand drought‐induced dieback.     

Hydraulic characteristics and leaf water use efficiency in trees from tropical montane habitats

This study was carried out in pioneer and successional forest tree species in a lower montane tropical forest with seasonal rains. We tested whether pioneer species feature high hydraulic conductance allowing them to use water profusely at leaf level. Conversely, forest species may have relatively low hydraulic conductance accompanied with better control over water use. This may lead in turn to pioneer species being at a relatively higher risk of shoot water potential falling below the threshold value at which cavitations occur compared to forest. Specific hydraulic conductance ( K _s) measured during the wet season was comparable between pioneers and forest species. During drought, K _s was significantly reduced, and species of both plant groups responded to this by modifying the relationship between conducting area and leaf area (Huver value), such that leaf specific conductivity ( K _l) was unaffected. Thus, leaf area seemed to be adjusted to maintain constant hydraulic sufficiency during drought. Pioneer species were more efficient in conducting water to their leaves but had low control over water use compared to forest species. A trade-off between water transport and leaf water use efficiency was suggested. These ecophysiological differences may have an impact on the performance of the species occupying contrasting habitats. Nonetheless, drought-induced embolisms occurred in trees growing in both open and forest habitats. Overall, during drought, adjustment of leaf area occurred in order to maintain a homeostasis of some physiological traits (leaf-specific conductivity and carbon assimilation).     

Allocation to leaf area and sapwood area affects water relations of co-occurring savanna and forest trees

Water availability is a principal factor limiting the distribution of closed-canopy forest in the seasonal tropics, suggesting that forest tree species may not be well adapted to cope with seasonal drought. We studied 11 congeneric species pairs, each containing one forest and one savanna species, to test the hypothesis that forest trees have a lower capacity to maintain seasonal homeostasis in water relations relative to savanna species. To quantify this, we measured sap flow, leaf water potential (Ψ_L), stomatal conductance (g _s), wood density, and Huber value (sapwood area:leaf area) of the 22 study species. We found significant differences in the water relations of these two species types. Leaf area specific hydraulic conductance of the soil/root/leaf pathway (G _t) was greater for savanna species than forest species. The lower G _t of forest trees resulted in significantly lower Ψ_L and g _s in the late dry season relative to savanna trees. The differences in G _t can be explained by differences in biomass allocation of savanna and forest trees. Savanna species had higher Huber values relative to forest species, conferring greater transport capacity on a leaf area basis. Forest trees have a lower capacity to maintain homeostasis in Ψ_L due to greater allocation to leaf area relative to savanna species. Despite significant differences in water relations, relationships between traits such as wood density and minimum Ψ_L were indistinguishable for the two species groups, indicating that forest and savanna share a common axis of water-use strategies involving multiple traits.     

Size distribution and expansion of canopy gaps in a northern Appalachian spruce-fir forest

Canopy gap area/age distributions and growth mechanisms were examined in a virgin subalpine forest in the White Mountains, New Hampshire, USA. The gap area distribution was negative exponential in form. Whithin gap tree ages varied widely in response to stepwise gap expansion caused by windthrow of peripheral trees or death of standing mature Picea rubens at gap edges. As a consequence, the density of small gaps may have been underestimated and the density of large gaps overestimated. The estimates of canopy turnover time, 303 yr, and of patch birth rate on an area basis, 3.3×10^-3 ha new patches/ha land area/yr, were not affected by the gap expansion phenomenon. However, any estimate of patch birth rate as numbers of new patches formed per year would have been too low. Because of increasingly widespread Picea death, the patch area/age distribution of this forest may not currently be in steady-state.     

Convergence of tree water use within an arid-zone woodland

We examined spatial and temporal patterns of tree water use and aspects of hydraulic architecture in four common tree species of central Australia—Corymbia opaca, Eucalyptus victrix, E. camaldulensis and Acacia aneura—to better understand processes that constrain water use in these environments. These four widely distributed species occupy contrasting niches within arid environments including woodlands, floodplains and riparian environments. Measurements of tree water use and leaf water potential were made at two sites with contrasting water table depths during a period of high soil water availability following summer rainfall and during a period of low soil water availability following 7 months of very little rainfall during 2007. There were significant differences in specific leaf area (SLA), sapwood area to leaf area ratios and sapwood density between species. Sapwood to leaf area ratio increased in all species from April to November indicating a decline in leaf area per unit sapwood area. Despite very little rainfall in the intervening period three species, C. opaca, E. victrix and E. camaldulensis maintained high leaf water potentials and tree water use during both periods. In contrast, leaf water potential and water use in the A. aneura were significantly reduced in November compared to April. Despite contrasting morphology and water use strategies, we observed considerable convergence in water use among the four species. Wood density in particular was strongly related to SLA, sapwood area to leaf area ratios and soil to leaf conductance, with all four species converging on a common relationship. Identifying convergence in hydraulic traits can potentially provide powerful tools for scaling physiological processes in natural ecosystems.     

Differences in xylem and leaf hydraulic traits explain differences in drought tolerance among mature Amazon rainforest trees

Considerable uncertainty surrounds the impacts of anthropogenic climate change on the composition and structure of Amazon forests. Building upon results from two large‐scale ecosystem drought experiments in the eastern Brazilian Amazon that observed increases in mortality rates among some tree species but not others, in this study we investigate the physiological traits underpinning these differential demographic responses. Xylem pressure at 50% conductivity (xylem‐P₅₀), leaf turgor loss point (TLP), cellular osmotic potential (π_o), and cellular bulk modulus of elasticity (ε), all traits mechanistically linked to drought tolerance, were measured on upper canopy branches and leaves of mature trees from selected species growing at the two drought experiment sites. Each species was placed a priori into one of four plant functional type (PFT) categories: drought‐tolerant versus drought‐intolerant based on observed mortality rates, and subdivided into early‐ versus late‐successional based on wood density. We tested the hypotheses that the measured traits would be significantly different between the four PFTs and that they would be spatially conserved across the two experimental sites. Xylem‐P₅₀, TLP, and π_o, but not ε, occurred at significantly higher water potentials for the drought‐intolerant PFT compared to the drought‐tolerant PFT; however, there were no significant differences between the early‐ and late‐successional PFTs. These results suggest that these three traits are important for determining drought tolerance, and are largely independent of wood density—a trait commonly associated with successional status. Differences in these physiological traits that occurred between the drought‐tolerant and drought‐intolerant PFTs were conserved between the two research sites, even though they had different soil types and dry‐season lengths. This more detailed understanding of how xylem and leaf hydraulic traits vary between co‐occuring drought‐tolerant and drought‐intolerant tropical tree species promises to facilitate a much‐needed improvement in the representation of plant hydraulics within terrestrial ecosystem and biosphere models, which will enhance our ability to make robust predictions of how future changes in climate will affect tropical forests.     

Heritable variation in the survival of seedlings from Patagonian cypress marginal xeric populations coping with drought and extreme cold

The rear edges of tree species have begun to be perceived as highly valuable for genetic resources conservation and management. In view of expected climatic changes, the responses of trees at their xeric limits may largely be determined by their capacity to cope with augmented environmental variance. We assess the heritability of early survival of Patagonian cypress in two common-garden field tests with contrasting summer water deficits, comprising 140 and 163 open-pollinated families from 10 marginal xeric populations. The first experiment underwent less rigorous conditions than the average mesic, Mediterranean climatic conditions, which were sufficient to reveal additive genetic effects of summer drought on seedling survival. The second trial suffered strong summer water-deficit stress and a winter extreme cold event. In this harsher environment, the heritabilities of survival under summer water-deficit stress were high in all the populations (h ²?=?0.84 on average), while the heritabilities of seasonal, extreme cold survival were moderate or even nil (h ²?=?0.28 on average). We did not find evidence of genetic differentiation among populations in their capabilities to survive droughts and cold extremes. Our results indicate that even when climatic changes were strong enough to cause the extinction of the most threatened populations, heritable variation for traits underlying drought and cold tolerances may allow the marginal xeric edge of cypress to persist under augmented environmental variance, without losing overall genetic diversity.     

Variation in woody plant mortality and dieback from severe drought among soils, plant groups, and species within a northern Arizona ecotone

Vegetation change from drought-induced mortality can alter ecosystem community structure, biodiversity, and services. Although drought-induced mortality of woody plants has increased globally with recent warming, influences of soil type, tree and shrub groups, and species are poorly understood. Following the severe 2002 drought in northern Arizona, we surveyed woody plant mortality and canopy dieback of live trees and shrubs at the forest–woodland ecotone on soils derived from three soil parent materials (cinder, flow basalt, sedimentary) that differed in texture and rockiness. Our first of three major findings was that soil parent material had little effect on mortality of both trees and shrubs, yet canopy dieback of trees was influenced by parent material; dieback was highest on the cinder for pinyon pine (Pinus edulis) and one-seed juniper (Juniperus monosperma). Ponderosa pine (Pinus ponderosa) dieback was not sensitive to parent material. Second, shrubs had similar mortality, but greater canopy dieback, than trees. Third, pinyon and ponderosa pines had greater mortality than juniper, yet juniper had greater dieback, reflecting different hydraulic characteristics among these tree species. Our results show that impacts of severe drought on woody plants differed among tree species and tree and shrub groups, and such impacts were widespread over different soils in the southwestern U.S. Increasing frequency of severe drought with climate warming will likely cause similar mortality to trees and shrubs over major soil types at the forest–woodland ecotone in this region, but due to greater mortality of other tree species, tree cover will shift from a mixture of species to dominance by junipers and shrubs. Surviving junipers and shrubs will also likely have diminished leaf area due to canopy dieback.     

Drought enhances symbiotic dinitrogen fixation and competitive ability of a temperate forest tree

General circulation models project more intense and frequent droughts over the next century, but many questions remain about how terrestrial ecosystems will respond. Of particular importance, is to understand how drought will alter the species composition of regenerating temperate forests wherein symbiotic dinitrogen (N₂)-fixing plants play a critical role. In experimental mesocosms we manipulated soil moisture to study the effect of drought on the physiology, growth and competitive interactions of four co-occurring North American tree species, one of which (Robinia pseudoacacia) is a symbiotic N₂-fixer. We hypothesized that drought would reduce growth by decreasing stomatal conductance, hydraulic conductance and increasing the water use efficiency of species with larger diameter xylem vessel elements (Quercus rubra, R. pseudoacacia) relative to those with smaller elements (Acer rubrum and Liriodendron tulipifera). We further hypothesized that N₂ fixation by R. pseudoacacia would decline with drought, reducing its competitive ability. Under drought, growth declined across all species; but, growth and physiological responses did not correspond to species’ hydraulic architecture. Drought triggered an 80 % increase in nodule biomass and N accrual for R. pseudoacacia, improving its growth relative to other species. These results suggest that drought intensified soil N deficiency and that R. pseudoacacia’s ability to fix N₂ facilitated competition with non-fixing species when both water and N were limiting. Under scenarios of moderate drought, N₂ fixation may alleviate the N constraints resulting from low soil moisture and improve competitive ability of N₂-fixing species, and as a result, supply more new N to the ecosystem.     

Plastic bimodal growth in a Mediterranean mixed-forest of Quercus ilex and Pinus halepensis

Mediterranean tree species have evolved to face seasonal water shortages, but may fail to cope with future increases in drought frequency and intensity. We investigated stem radial increment dynamics in two typical Mediterranean tree species, Aleppo pine (Pinus halepensis), a drought-avoiding species, and holm oak (Quercus ilex), a drought-tolerant species, in a mixed forest and on contrasting slope aspects (south- and north-facing). Intra- and inter-annual growth patterns were modelled using the VS-Lite2 model for each tree species and slope-aspect. Both species showed a bimodal growth pattern, with peaks coinciding with favourable conditions in spring and autumn. A bimodal growth pattern is always observed in P. halepensis, while in Q. ilex is facultative, which suggests different strategies adopted by these species to cope with summer drought. More specifically, trees on south-facing slope showed a more evident bimodal pattern and more intra-annual density fluctuations. In recent decades, the intensity of both growth peaks has diminished and drifted away due to the increased summer drought. The VS-Lite2 model reveals a niche partitioning between both species. Differences in growing season’s length and timings of growth peaks in both species are relevant for their coexistence and should be considered for estimating mixed-forest responses under climate change scenarios.     

立地条件和树龄对刺槐和小叶杨叶水力性状及抗旱性的影响

以形成黄土高原“小老树”的2种典型树种刺槐和小叶杨为对象,研究了立地条件(沟谷台地和沟间坡地)和树龄对两种树木叶水力学性质和抗旱性的影响,探讨“小老树”形成的水力生理机制.结果表明: 水分较好的沟谷台地上生长的两种树木的叶最大水力导度(K_max)明显大于水分较差的沟间坡地,叶水力脆弱性(P₅₀)也较高;随树龄增加,两种树木的K_max明显下降,但P₅₀差异不大.台地上生长的两种树木的叶表皮导度和PV曲线参数(膨压损失点时的相对含水量RWC_tlp、膨压损失点时的水势ψ_tlp、饱和含水量时的渗透势ψ_sat)均大于坡地;随树龄增加,两种树木的叶表皮导度显著下降,PV曲线参数出现不同程度的下降.两种树木K_max与ψ_tlp呈显著正相关,P₅₀与PV曲线参数之间存在一定的相关性,表明K_max与抗旱性之间存在一种权衡关系,P₅₀是反映两种树木的抗旱性特征之一.     

Drought stress limits the geographic ranges of two tree species via different physiological mechanisms

Range shifts are among the most ubiquitous ecological responses to anthropogenic climate change and have large consequences for ecosystems. Unfortunately, the ecophysiological forces that constrain range boundaries are poorly understood, making it difficult to mechanistically project range shifts. To explore the physiological mechanisms by which drought stress controls dry range boundaries in trees, we quantified elevational variation in drought tolerance and in drought avoidance‐related functional traits of a widespread gymnosperm (ponderosa pine – Pinus ponderosa) and angiosperm (trembling aspen – Populus tremuloides) tree species in the southwestern USA. Specifically, we quantified tree‐to‐tree variation in growth, water stress (predawn and midday xylem tension), drought avoidance traits (branch conductivity, leaf/needle size, tree height, leaf area‐to‐sapwood area ratio), and drought tolerance traits (xylem resistance to embolism, hydraulic safety margin, wood density) at the range margins and range center of each species. Although water stress increased and growth declined strongly at lower range margins of both species, ponderosa pine and aspen showed contrasting patterns of clinal trait variation. Trembling aspen increased its drought tolerance at its dry range edge by growing stronger but more carbon dense branch and leaf tissues, implying an increased cost of growth at its range boundary. By contrast, ponderosa pine showed little elevational variation in drought‐related traits but avoided drought stress at low elevations by limiting transpiration through stomatal closure, such that its dry range boundary is associated with limited carbon assimilation even in average climatic conditions. Thus, the same climatic factor (drought) may drive range boundaries through different physiological mechanisms – a result that has important implications for process‐based modeling approaches to tree biogeography. Further, we show that comparing intraspecific patterns of trait variation across ranges, something rarely done in a range‐limit context, helps elucidate a mechanistic understanding of range constraints.