Beyond soil water potential: An expanded view on isohydricity including land-atmosphere interactions and phenology.

Over the past decade, the concept of isohydry or anisohydry, which describes the link between soil water potential (ΨS ), leaf water potential (ΨL ), and stomatal conductance (gs ), has soared in popularity. However, its utility has recently been questioned, and a surprising lack of coordination between the dynamics of ΨL and gs across biomes has been reported. Here, we offer a more expanded view of the isohydricity concept that considers effects of vapour pressure deficit (VPD) and leaf area index (AL ) on the apparent sensitivities of ΨL and gs to drought. After validating the model with tree- and ecosystem-scale data, we find that within a site, isohydricity is a strong predictor of limitations to stomatal function, though variation in VPD and leaf area, among other factors, can challenge its diagnosis. Across sites, the theory predicts that the degree of isohydricity is a good predictor of the sensitivity of gs to declining soil water in the absence of confounding effects from other drivers. However, if VPD effects are significant, they alone are sufficient to decouple the dynamics of ΨL and gs entirely. We conclude with a set of practical recommendations for future applications of the isohydricity framework within and across sites.

Savanna woody encroachment is widespread across three continents.

Tropical savannas are a globally extensive biome prone to rapid vegetation change in response to changing environmental conditions. Via a meta-analysis, we quantified savanna woody vegetation change spanning the last century. We found a global trend of woody encroachment that was established prior the 1980s. However, there is critical regional variation in the magnitude of encroachment. Woody cover is increasing most rapidly in the remaining uncleared savannas of South America, most likely due to fire suppression and land fragmentation. In contrast, Australia has experienced low rates of encroachment. When accounting for land use, African savannas have a mean rate annual woody cover increase two and a half times that of Australian savannas. In Africa, encroachment occurs across multiple land uses and is accelerating over time. In Africa and Australia, rising atmospheric CO2 , changing land management and rainfall are likely causes. We argue that the functional traits of each woody flora, specifically the N-fixing ability and architecture of woody plants, are critical to predicting encroachment over the next century and that African savannas are at high risk of widespread vegetation change.

Does leaf shedding protect stems from cavitation during seasonal droughts? A test of the hydraulic fuse hypothesis.

During droughts, leaves are predicted to act as 'hydraulic fuses' by shedding when plants reach critically low water potential (Ψplant ), thereby slowing water loss, stabilizing Ψplant and protecting against cavitation-induced loss of stem hydraulic conductivity (Ks ). We tested these predictions among trees in seasonally dry tropical forests, where leaf shedding is common, yet variable, among species. We tracked leaf phenology, Ψplant and Ks in saplings of six tree species distributed across two forests. Species differed in their timing and extent of leaf shedding, yet converged in shedding leaves as they approached the Ψplant value associated with a 50% loss of Ks and at which their model-estimated maximum sustainable transpiration rate approached zero. However, after shedding all leaves, the Ψplant value of one species, Genipa americana, continued to decline, indicating that water loss continued after leaf shedding. Ks was highly variable among saplings within species and seasons, suggesting a minimal influence of seasonal drought on Ks . Hydraulic limits appear to drive diverse patterns of leaf shedding among tropical trees, supporting the hydraulic fuse hypothesis. However, leaf shedding is not universally effective at stabilizing Ψplant , suggesting that the main function of drought deciduousness may vary among species.

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.

Patterns of compound-leaf form and deciduous-leaf habit across forests in China: Their association and key climatic factors.

Leaf form (compound vs. simple) and habit (evergreen vs. deciduous) are key functional traits of trees to adapt to various climates and are vital in determining plant response to climate change. However, their association and climatic determinants remain uncertain, especially in East Asian forests in the largest monsoon region on earth. To fill these knowledge gaps, we compiled a dataset comprising 42 intact forests and over 2200 angiosperm tree species across China (spanning 30 latitudes and 47 longitudes). The geographical and climatic patterns of leaf form and habit were analyzed. The association between compound leaf and deciduousness was tested for tropical, subtropical and temperate climatic zones. We found that both the percentage of compound leaf (CT%) and deciduous tree species (DT%) increased with latitude and decreased with mean annual precipitation (MAP). For all forests, DT% was negatively related to mean annual temperature (MAT), whereas CT% was not. Nevertheless, both DT% and CT% increased with increasing MAT in the tropics, possibly owing to the high vapor pressure deficits (VPD) and canopy water deficits associated with high temperatures. A positive linear relationship between CT% and DT% was found across all forests and within different climatic zones except for temperate, and the intercept of the regression line was significantly higher in the tropics than in the subtropics. Overall, as supported by principal component analysis, deciduousness was negatively associated with both temperature and precipitation, while CT negatively with precipitation only across zones and positively with temperature in the tropics. Different relationships in different climatic zones suggest potentially different selective forces. Our findings provide novel insights into the linkage between leaf form and habit, as well as how climate shapes the landscape of broadleaf forests, which has important implications regarding the response of forest composition to climate change.

High moon brightness and low ambient temperatures affect sloth predation by harpy eagles.

BACKGROUND: Climate plays a key role in the life histories of tropical vertebrates. However, tropical forests are only weakly seasonal compared with temperate and boreal regions. For species with limited ability to control core body temperature, even mild climatic variation can determine major behavioural outcomes, such as foraging and predator avoidance. In tropical forests, sloths are the arboreal vertebrate attaining the greatest biomass density, but their capacity to regulate body temperature is limited, relying on behavioural adaptations to thermoregulate. Sloths are largely or strictly nocturnal, and depend on crypsis to avoid predation. The harpy eagle (Harpia harpyja) is a sloth-specialist and exerts strong top-down control over its prey species. Yet the role of environmental variables on the regulation of predator-prey interactions between sloths and harpy eagles are unknown. The harpy eagle is considered Near Threatened. This motivated a comprehensive effort to reintroduce this species into parts of Mesoamerica. This effort incidentally enabled us to understand the prey profile of harpy eagles over multiple seasons. METHODS: Our study was conducted between 2003 and 2009 at Soberanía National Park, Panamá. Telemetered harpy eagles were seen hunting and feeding on individual prey species. For each predation event, field assistants systematically recorded the species killed. We analysed the effects of climatic conditions and vegetation phenology on the prey species profile of harpy eagles using generalised linear mixed models. RESULTS: Here we show that sloth predation by harpy eagles was negatively affected by nocturnal ambient light (i.e. bright moonshine) and positively affected by seasonally cool temperatures. We suggest that the first ensured low detectability conditions for sloths foraging at night and the second posed a thermally unsuitable climate that forced sloths to forage under riskier daylight. We showed that even moderate seasonal variation in temperature can influence the relationship between a keystone tropical forest predator and a dominant prey item. Therefore, predator-prey ecology in the tropics can be modulated by subtle changes in environmental conditions. The seasonal effects shown here suggest important demographic consequences for sloths, which are under top-down regulation from harpy eagle predation, perhaps limiting their geographic distribution at higher latitudes.

Geographical and climatic gradients of evergreen versus deciduous broad-leaved tree species in subtropical China: Implications for the definition of the mixed forest.

Understanding climatic influences on the proportion of evergreen versus deciduous broad-leaved tree species in forests is of crucial importance when predicting the impact of climate change on broad-leaved forests. Here, we quantified the geographical distribution of evergreen versus deciduous broad-leaved tree species in subtropical China. The Relative Importance Value index (RIV) was used to examine regional patterns in tree species dominance and was related to three key climatic variables: mean annual temperature (MAT), minimum temperature of the coldest month (MinT), and mean annual precipitation (MAP). We found the RIV of evergreen species to decrease with latitude at a lapse rate of 10% per degree between 23.5 and 25°N, 1% per degree at 25-29.1°N, and 15% per degree at 29.1-34°N. The RIV of evergreen species increased with: MinT at a lapse rate of 10% per °C between -4.5 and 2.5°C and 2% per °C at 2.5-10.5°C; MAP at a lapse rate of 10% per 100 mm between 900 and 1,600 mm and 4% per 100 mm between 1,600 and 2,250 mm. All selected climatic variables cumulatively explained 71% of the geographical variation in dominance of evergreen and deciduous broad-leaved tree species and the climatic variables, ranked in order of decreasing effects were as follows: MinT > MAP > MAT. We further proposed that the latitudinal limit of evergreen and deciduous broad-leaved mixed forests was 29.1-32°N, corresponding with MAT of 11-18.1°C, MinT of -2.5 to 2.51°C, and MAP of 1,000-1,630 mm. This study is the first quantitative assessment of climatic correlates with the evergreenness and deciduousness of broad-leaved forests in subtropical China and underscores that extreme cold temperature is the most important climatic determinant of evergreen and deciduous broad-leaved tree species' distributions, a finding that confirms earlier qualitative studies. Our findings also offer new insight into the definition and distribution of the mixed forest and an accurate assessment of vulnerability of mixed forests to future climate change.

Retention of stored water enables tropical tree saplings to survive extreme drought conditions.

Trees generally maintain a small safety margin between the stem water potential (Ψstem) reached during seasonal droughts and the Ψstem associated with their mortality. This pattern may indicate that species face similar mortality risk during extreme droughts. However, if tree species vary in their ability to regulate Ψstem, then safety margins would poorly predict drought mortality. To explore variation among species in Ψstem regulation, I subjected potted saplings of six tropical tree species to extreme drought and compared their responses with well-watered plants and pretreatment reference plants. In the drought treatment, soil water potential reached <-10 MPa, yet three species, Bursera simaruba (L.) Sarg., Cavanillesia platanifolia (Bonpl.) Kunth and Cedrela odorata L. had 100% survival and maintained Ψstem near -1 MPa (i.e., desiccation-avoiding species). Three other species, Cojoba rufescens (Benth.) Britton and Rose, Genipa americana L. and Hymenaea courbaril L. had 50%, 0% and 25% survival, respectively, and survivors had Ψstem <-6 MPa (i.e., desiccation-susceptible species). The desiccation-avoiding species had lower relative water content (RWC) in all organs and tissues (root, stem, bark and xylem) in the drought treatment than in the reference plants (means 72.0-90.4% vs 86.9-97.9%), but the survivors of the desiccation-susceptible C. rufescens had much lower RWC in the drought treatment (44.5-72.1%). Among the reference plants, the desiccation-avoiding species had lower tissue density, leaf-mass fraction and lateral-root surface area (LRA) than the desiccation-susceptible species. Additionally, C. platanifolia and C. odorata had reduced LRA in the drought treatment, which may slow water loss into dry soil. Together, these results suggest that the ability to regulate Ψstem during extreme drought is associated with functional traits that favor retention of stored water and that safety margins during seasonal drought poorly predict survival during extreme drought.

Impacts of artificial reservoirs on floristic diversity and plant functional traits in dry forests after 15 years / Impactos de um reservatório artificial sobre a diversidade florística e traços funcionais de espécies arbóreas de uma Floresta Estacional Decidual após 15 anos do represamento

AbstractDams are of paramount importance to a wide variety of human services and many of their environmental problems are known; however, there are few studies in the world addressing the impacts on the native vegetation previously distant from water bodies which became close to the lakeshore created by a dam. Thus, this paper aims to analyze the responses of a dry forest to a dam after 15 years. For this, 20 random samples of 40 trees were made, 10 close to the lakeshore and 10 distant from it, by applying the central square point method. Close to the dam, we found higher values regarding basal area, number of trees, number of evergreen trees, and zoochoric syndrome, but there were lower values of Shannon’s diversity index. Therefore, the impacts of the dam after 15 years caused several changes to the tree community. The greater basal area close to the dam suggests that water deficit during the dry season was decreased and plants have thicker trunks. On the other hand, this sector had much more zoochoric syndrome and a larger number of evergreen trees than plots which are distant from water, suggesting changes with regard to the community’s ecological functions. Furthermore, structural floristic data shows that the sector close to the dam is less similar to other deciduous forests within the same geographical region than the sector distant from water, thus providing evidence of the impacts of dams on the tree community.

ResumoBarragens são de suma importância para uma ampla variedade de serviços humanos e muitos de seus problemas ambientais já são conhecidos. No entanto, existem poucos estudos no mundo abordando os impactos sobre a vegetação nativa anteriormente distante de corpos d'água, que após a barragem se aproximou do lago criado. Assim, este trabalho tem como objetivo analisar as respostas de uma floresta seca as margens da represa formada por uma barragem após 15 anos. Para isso, 20 amostras aleatórias de 40 árvores foram feitas, sendo 10 perto da margem do lago e 10 distante dele, através da aplicação do método de ponto central. Perto da barragem, encontramos valores mais elevados em relação à área basal, densidade, número de árvores perenifólias e zoocórica, mas houve menores valores de índice de diversidade de Shannon. Portanto, os impactos da barragem depois de 15 anos causou várias mudanças na comunidade de árvores. A maior área basal perto da barragem sugere que o déficit hídrico durante a estação seca foi diminuído, o que aumenta o crescimento em área basal das espécies. Por outro lado, este setor apresentou maior número de indivíduos zoocóricos e número maior de árvores perenifólias do que as parcelas que estão distantes da água, sugerindo mudanças no que diz respeito às funções ecológicas da comunidade. Além disso, os dados florísticos estruturais mostram que o setor perto da barragem é menos similar a outras florestas estacionais deciduais dentro de uma mesma região geográfica que o setor distante da água, proporcionando, assim, evidências dos impactos das barragens sobre a comunidade arbórea.