Temporal changes in the dominance of tree functional traits,but no changes in species diversity during woody plant encroachment in a Brazilian savanna

Questions : Woody encroachment in savannas has been associated with changing taxonomic composition and ecosystem function. Interestingly, there is little understanding of how encroachment impacts plant functional diversity and how those changes relate to plant demography, a crucial mediator between taxonomic composition and ecosystem function. Location : Southeastern Brazil. Methods: Using a landscape scale fire suppression experiment in a diverse Brazilian savanna, we quantify how change in species composition over seven years impacted vegetative and reproductive tree functional diversity as determined by new recruits, dead and surviving trees. Results: Over seven years, tree above-ground biomass increased by 15%, while total species richness did not change. Despite minor changes, species composition remained overall similar (82%), with few species contributing significantly to plot dissimilarity over time. There were small changes in vegetative traits, where the community-weighted mean increased in maximum tree height (↑ 2.1%) and specific leaf area (↑ 5.3%), and decreased in wood density (↓ 1.3%) and bark thickness (↓ 9.4%). Changes in reproductive traits were larger than in vegetative traits, with an increase in the prevalence of monoecy (↑ 32.6%), dioecy (↑ 44.2%), large seeds (↑ 20.3%), animal-mediated seed dispersal (↑ 4.9%) and pollination by very small insects (↑ 45.5%), and a decrease in the prevalence of hermaphroditism (↓ 9%), small seeds (6.8%) and pollination by small insects (12.5%). The overall decrease in bark thickness and increase in monoecy and dioecy were mainly driven by characters of the new recruits, while the overall increase in specific leaf area (SLA) and decrease in small seeds appeared largely determined by the loss of trees possessing those traits. Conclusions: Encroachment leads to changes that are likely increasing ecosystem vulnerability to fire and drought. Further, the compositional changes observed appear to drive marked change in reproductive traits, indicating increasing dependence on animals for dispersal and reproduction. Understanding post-hoc encroachment impacts in an era of widespread pervasive encroachment is fundamental to reconciling ecosystem functions such as nutrient cycling and pollination services as there is a loss of species with open ecosystem life-history strategies. Among savannas, there remains an urgent need to understand relationships between woody cover and ecosystem function to determine thresholds in woody cover promoting resilient savanna ecosystems.     

Long‐term livestock exclusion facilitates native woody plant encroachment in a sandy semiarid rangeland

相似文献   

青藏高原木本植物扩张对生长季地表昼夜温度的影响

木本植物扩张或灌丛化是全球性的生态环境问题。近年来青藏高原发生了大规模的木本植物扩张。然而木本植物在青藏高原扩张的时空分布特征及其对局地地表温度(LST)如何影响尚不清楚。基于MODIS土地覆盖产品识别出青藏高原木本植物扩张的空间分布,并利用移动窗口搜索法,探究其对生长季昼夜LST的影响规律及成因。结果表明,2001至2018年木本植物扩张的范围和程度均整体呈增加的趋势。在2018年,木本植物扩张使生长季白天LST降低(2.60±0.34)℃,夜间LST增加(0.94±0.22)℃,净效应使日均LST降低(0.83±0.24)℃。产生这种现象的原因是蒸散发增加((+13.46±6.65)mm/a)等引发的降温效应超过了以反照率减少(-0.031±0.003)为代表的增温效应。气候背景对该影响的空间分布具有相当的控制作用,即降水主导着白天LST的改变,但气温在夜间LST变化中占据更重要的地位。总体上,在气温越低、降水率越高、高程越低的地方发生的木本植物扩张更倾向于降低局地LST。与同一年中越湿润的地方越倾向于降温“相悖”的是,在不同的水文年,更干旱的年份对白天LST具有更强的降温作用,这...     

The effects of fire on woody plant encroachment are exacerbated by succession of trees of decreased palatability

The ingression of woody plants into the grassy layer of savannas and grasslands has become a global concern. The increase of woody plants has been primarily attributed over grazing, fire and more recently to the increase of atmospheric CO₂. We used long-term observations and analyses to assess changes in woody vegetation in Ithala Game Reserve (IGR), South Africa. Textural analysis of aerial photographs was used to detect changes in woody vegetation, from 1943 to 2007 in Ithala Game Reserve (IGR), South Africa. Daily rainfall data from 1905 to 2009 were used in a time-series analysis to determine if rainfall patterns have changed. The time-series analysis showed that the low magnitude (0–10 mm) rainfall events decreased from 1916 to 2009 and high magnitude rainfall events increased (10–20 and >20 mm). The mean annual rainfall increased from ～700 to ～850 mm from the 1930s to the 2000s. This change in rainfall was a key factor in the increase in woody vegetation from 1943 to 2009. We also used field data from the same reserve collected over 30 years to assess the increases in tree cover. Tree cover and density increased significantly by 32.5% and 657.9 indiv ha^?1 respectively, over 64 years. Before the proclamation of IGR in 1972, increases in woody vegetation from 1943 were non-significant. After the proclamation of IGR, herbivore population numbers and spatial distribution influenced the accumulation of grassy biomass required to fuel fires. In areas with reduced fuel loads, the consequential suppression of fire accelerated the rate of woody plant invasion into savannas. The increase in woody vegetation coincided with a decrease in palatable (e.g. Acacia gerrardii and Acacia davyi) and an increase in unpalatable woody plants. The avoidance of the unpalatable trees (e.g. Euclea and Searsia species) by large mammalian herbivores has allowed these trees to increase in density relatively unhindered.     

A patch-dynamics approach to savanna dynamics and woody plant encroachment – Insights from an arid savanna

The coexistence of woody and grassy plants in savannas has often been attributed to a rooting-niche separation (two-layer hypothesis). Water was assumed to be the limiting resource for both growth forms and grasses were assumed to extract water from the upper soil layer and trees and bushes from the lower layers. Woody plant encroachment (i.e. an increase in density of woody plants often unpalatable to domestic livestock) is a serious problem in many savannas and is believed to be the result of overgrazing in ‘two-layer systems’. Recent research has questioned the universality of both the two-layer hypothesis and the hypothesis that overgrazing is the cause of woody plant encroachment.

We present an alternative hypothesis explaining both tree–grass coexistence and woody plant encroachment in arid savannas. We propose that woody plant encroachment is part of a cyclical succession between open savanna and woody dominance and is driven by two factors: rainfall that is highly variable in space and time, and inter-tree competition. In this case, savanna landscapes are composed of many patches (a few hectares in size) in different states of transition between grassy and woody dominance, i.e. we hypothesize that arid savannas are patch-dynamic systems. We summarize patterns of tree distribution observed in an arid savanna in Namibia and show that these patterns are in agreement with the patch-dynamic savanna hypothesis. We discuss the applicability of this hypothesis to fire-dominated savannas, in which rainfall variability is low and fire drives spatial heterogeneity.

We conclude that field studies are more likely to contribute to a general understanding of tree–grass coexistence and woody plant encroachment if they consider both primary (rain and nutrients) and secondary (fire and grazing) determinants of patch properties across different savannas.     

The resprouting response of co‐occurring temperate woody plant and grass species to elevated [CO2]: An insight into woody plant encroachment of grasslands

The phenomenon of woody plant thickening in grasslands has been observed globally and is likely to have widespread ecological consequences. It has been proposed that woody plant thickening is driven in part by rising atmospheric [CO₂] enhancing the resprouting ability of woody plants relative to grasses so they respond more strongly after disturbances such as herbivory and fire. The aim of this study was to examine the CO₂ effect on the resprouting ability of 16 co‐occurring temperate woody plant and grass species (eight species from each growth form). Plants were grown in a controlled glasshouse experiment under ambient (400 ppm) and elevated [CO₂] (600 ppm) for 14 weeks after which their resprouting ability was measured. Root non‐structural carbohydrate (NSC_mass) and nitrogen (N_mass) storage was used as proxies to measure the resprouting ability of woody plants while for the grasses it was measured directly. We found that both the woody plants (22% on average; P = 0.003) and grasses (20% on average; P = 0.003) produced more biomass under elevated [CO₂]. Despite the woody plants not allocating additional carbon to belowground storage under elevated [CO₂], they had significantly greater root NSC_mass (23% on average; P = 0.007) due to increased root biomass production (8% on average; P = 0.007). In contrast, root N_mass of the woody plants did not differ between CO₂ treatments (P = 0.373). Surprisingly, the resprouting ability of the grasses did not significantly differ between the CO₂ treatments (P = 0.067). These results provide evidence that the differing resprouting response of woody plants and grasses under elevated [CO₂] may be contributing to woody plant encroachment of grasslands worldwide.     

Determinants of woody encroachment and cover in African savannas

Savanna ecosystems are an integral part of the African landscape and sustain the livelihoods of millions of people. Woody encroachment in savannas is a widespread phenomenon but its causes are widely debated. We review the extensive literature on woody encroachment to help improve understanding of the possible causes and to highlight where and how future scientific efforts to fully understand these causes should be focused. Rainfall is the most important determinant of maximum woody cover across Africa, but fire and herbivory interact to reduce woody cover below the maximum at many locations. We postulate that woody encroachment is most likely driven by CO₂ enrichment and propose a two-system conceptual framework, whereby mechanisms of woody encroachment differ depending on whether the savanna is a wet or dry system. In dry savannas, the increased water-use efficiency in plants relaxes precipitation-driven constraints and increases woody growth. In wet savannas, the increase of carbon allocation to tree roots results in faster recovery rates after disturbance and a greater likelihood of reaching sexual maturity. Our proposed framework can be tested using a mixture of experimental and earth observational techniques. At a local level, changes in precipitation, burning regimes or herbivory could be driving woody encroachment, but are unlikely to be the explanation of this continent-wide phenomenon.     

Response of fishes and aquatic habitats to sand-bed stream restoration using large woody debris

Effects of habitat rehabilitation of Little Topashaw Creek, a sinuous, sand-bed stream draining 37 km² in northwest Mississippi are described. The rehabilitation project consisted of placing 72 large woody debris structures along eroding concave banks and planting 4000 willow cuttings in sandbars. Response was measured by monitoring flow, channel geometry, physical aquatic habitat, and fish populations. Initially, debris structures reduced high flow velocities at concave bank toes, preventing further erosion and inducing deposition. Physical response during the first year following construction included creation of sand berms along eroding banks and slight increases in base flow water width and depth. Fish collections showed assemblages typical of incising streams within the region, but minor initial responses to debris addition were evident. Progressive failure of the structures and renewed erosion were observed during the second year after construction.     

Disturbances by desert rodents are more strongly associated with spatial changes in soil texture than woody encroachment

Background and Aims

Soil texture is an important determinant of ecosystem structure and productivity in drylands, and may influence animal foraging and, indirectly, plant community composition.

Methods

We measured the density and composition of surface disturbances (foraging pits) of small, soil-foraging desert vertebrates in shrubland and grasslands, both with coarse- and fine-textured soils. We predicted that the density and functional significance of disturbances would be related more to differences in texture than shrub encroachment.

Results

Soil texture had a stronger influence on animal foraging sites than shrub encroachment. There were more disturbances, greater richness and abundance of trapped seed, and greater richness of germinating plants on coarse- than fine-textured soils. Pits in coarse soils trapped 50 % more litter than those in finer soils. Apart from slightly more soil removal and greater litter capture in shrubland pits, there were no effects of encroachment.

Conclusions

Although the process of woody encroachment has been shown to have marked effects on some ecosystem properties, it is likely to have a more subordinate effect on surface disturbances and therefore their effects on desert plant communities than soil texture. Our results highlight the importance of animal activity in shaping desert plant communities, and potentially, in maintaining or reinforcing shrub dominant processes.     

Rapid and unpredictable changes of the G‐matrix in a natural bird population over 25 years

Knowledge of the genetic variances and covariances of traits (the G ‐matrix) is fundamental for the understanding of evolutionary dynamics of populations. Despite its essential importance in evolutionary studies, empirical tests of the temporal stability of the G ‐matrix in natural populations are few. We used a 25‐year‐long individual‐based field study on almost 7000 breeding attempts of the collared flycatcher (Ficedula albicollis) to estimate the stability of the G‐matrix over time. Using animal models to estimate G for several time periods, we show that the structure of the time‐specific G‐matrices changed significantly over time. The temporal changes in the G‐matrix were unpredictable, and the structure at one time period was not indicative of the structure at the next time period. Moreover, we show that the changes in the time‐specific G‐matrices were not related to changes in mean trait values or due to genetic drift. Selection, differences in acquisition/allocation patterns or environment‐dependent allelic effects are therefore likely explanations for the patterns observed, probably in combination. Our result cautions against assuming constancy of the G ‐matrix and indicates that even short‐term evolutionary predictions in natural populations can be very challenging.     

Thermal acclimation of shoot respiration in an Arctic woody plant species subjected to 22 years of warming and altered nutrient supply

Despite concern about the status of carbon (C) in the Arctic tundra, there is currently little information on how plant respiration varies in response to environmental change in this region. We quantified the impact of long‐term nitrogen (N) and phosphorus (P) treatments and greenhouse warming on the short‐term temperature (T) response and sensitivity of leaf respiration (R), the high‐T threshold of R, and associated traits in shoots of the Arctic shrub Betula nana in experimental plots at Toolik Lake, Alaska. Respiration only acclimated to greenhouse warming in plots provided with both N and P (resulting in a ~30% reduction in carbon efflux in shoots measured at 10 and 20 °C), suggesting a nutrient dependence of metabolic adjustment. Neither greenhouse nor N+P treatments impacted on the respiratory sensitivity to T (Q₁₀); overall, Q₁₀ values decreased with increasing measuring T, from ~3.0 at 5 °C to ~1.5 at 35 °C. New high‐resolution measurements of R across a range of measuring Ts (25–70 °C) yielded insights into the T at which maximal rates of R occurred (T_max). Although growth temperature did not affect T_max, N+P fertilization increased T_max values ~5 °C, from 53 to 58 °C. N+P fertilized shoots exhibited greater rates of R than nonfertilized shoots, with this effect diminishing under greenhouse warming. Collectively, our results highlight the nutrient dependence of thermal acclimation of leaf R in B. nana, suggesting that the metabolic efficiency allowed via thermal acclimation may be impaired at current levels of soil nutrient availability. This finding has important implications for predicting carbon fluxes in Arctic ecosystems, particularly if soil N and P become more abundant in the future as the tundra warms.     

Temperature and precipitation controls over leaf‐ and ecosystem‐level CO2 flux along a woody plant encroachment gradient

Conversion of grasslands to woodlands may alter the sensitivity of CO₂ exchange of individual plants and entire ecosystems to air temperature and precipitation. We combined leaf‐level gas exchange and ecosystem‐level eddy covariance measurements to quantify the effects of plant temperature sensitivity and ecosystem temperature responses within a grassland and mesquite woodland across seasonal precipitation periods. In so doing, we were able to estimate the role of moisture availability on ecosystem temperature sensitivity under large‐scale vegetative shifts. Optimum temperatures (T_opt) for net photosynthetic assimilation (A) and net ecosystem productivity (NEP) were estimated from a function fitted to A and NEP plotted against air temperature. The convexities of these temperature responses were quantified by the range of temperatures over which a leaf or an ecosystem assimilated 50% of maximum NEP (Ω₅₀). Under dry pre‐ and postmonsoon conditions, leaf‐level Ω₅₀ in C₃ shrubs were two‐to‐three times that of C₄ grasses, but under moist monsoon conditions, leaf‐level Ω₅₀ was similar between growth forms. At the ecosystems‐scale, grassland NEP was more sensitive to precipitation, as evidenced by a 104% increase in maximum NEP at monsoon onset, compared to a 57% increase in the woodland. Also, woodland NEP was greater across all temperatures experienced by both ecosystems in all seasons. By maintaining physiological function across a wider temperature range during water‐limited periods, woody plants assimilated larger amounts of carbon. This higher carbon‐assimilation capacity may have significant implications for ecosystem responses to projected climate change scenarios of higher temperatures and more variable precipitation, particularly as semiarid regions experience conversions from C₄ grasses to C₃ shrubs. As regional carbon models, CLM 4.0, are now able to incorporate functional type and photosynthetic pathway differences, this work highlights the need for a better integration of the interactive effects of growth form/functional type and photosynthetic pathway on water resource acquisition and temperature sensitivity.     

Climatic and biotic velocities for woody taxa distributions over the last 16 000 years in eastern North America

We estimated the latitudinal velocity (km/decade) of northern and southern boundaries of core distributions for 30 woody taxa over the last 16 000 years (biotic velocities) using networks of fossil pollen records, and compared these with climate velocities estimated from CCSM3 simulations. Biotic velocities were faster during periods of rapid temperature change (i.e. 16 to 7 ka) than times of relative stability (i.e. 7 to 1 ka), with a consistent northward movement of northern and southern boundaries. For most taxa, biotic velocities were faster for northern than for southern boundaries between 12 and 7 ka, resulting in expanding distributions. For individual time periods, biotic velocities were as fast or faster than climate velocities calculated using multivariate approaches. These results indicate that climate change paced the rate of distribution shifts in both northern and southern populations while suggesting that northern populations were more sensitive. A similar sensitivity and pacing is expected under 21st century climate change.     

Stochastic losses of fire‐dependent endemic herbs revealed by a 65‐year chronosequence of dispersal‐limited woody plant encroachment

The factors responsible for maintaining diverse groundcover plant communities of high conservation value in frequently burned wet pine savannas are poorly understood. While most management involves manipulating extrinsic factors important in maintaining species diversity (e.g., fire regimes), most ecological theory (e.g., niche theory and neutral theory) examines how traits exhibited by the species promote species coexistence. Furthermore, although many ecologists focus on processes that maintain local species diversity, conservation biologists have argued that other indices (e.g., phylogenetic diversity) are better for evaluating assemblages in terms of their conservation value. I used a null model that employed beta‐diversity calculations based on Raup–Crick distances to test for deterministic herbaceous species losses associated with a 65‐year chronosequence of woody species encroachment within each of three localities. I quantified conservation value of assemblages by measuring taxonomic distinctness, endemism, and floristic quality of plots with and without woody encroachment. Reductions in herb species richness per plot attributable to woody encroachment were largely stochastic, as indicated by a lack of change in the mean or variance in beta‐diversity caused by woody encroachment in the savannas studied here. Taxonomic distinctness, endemism, and floristic quality (when summed across all species) were all greater in areas that had not experienced woody encroachment. However, when corrected for local species richness, only average endemism and floristic quality of assemblages inclusive of herbs and woody plants were greater in areas that had not experienced woody encroachment, due to the more restricted ranges and habitat requirements of herbs. Results suggest that frequent fires maintain diverse assemblages of fire‐dependent herb species endemic to the region. The stochastic loss of plant species, irrespective of their taxonomic distinctness, to woody encroachment suggests that the relevance of niche partitioning or phylogenetic diversity to the management of biodiversity in wet pine savannas is minimal.     

Temporal changes in abundance–occupancy relationships over 40 years

1. Abundance–occupancy (A–O) relationships are widely documented for many organismal groups and regions, and have been used to gain an understanding of regional population and community trends. Monitoring changes in abundance and occupancy over time may be what is required to document changes in conservation status and needs for some species, communities, or areas.
2. We hypothesize that if there is a higher proportion of declining species in one group of species compared with another (e.g., migratory species vs. permanent residents), then a consequence of that difference will be vastly different abundance–occupancy relationships. If this difference persists through time, then the resulting A–O relationships between the groups will continue to diverge.
3. For neotropical migrants, short‐distance migrants, and permanent resident birds of North America, we assess the numbers of declining species over 1969–2009. We further test for differences in the A–O relationship across these three groups, and in rates of change in abundance and occupancy separately.
4. We find significant differences in numbers of declining species across the migratory groups, a significant decline in the A–O relationship for permanent residents, a significant increase for Neotropical migrants, and a nonsignificant decline for short‐distance migrants over the 40 years. Further, abundances are not changing at different rates but occupancies are consistently greater over time for neotropical migrants versus permanent residents, likely driving the changes in A–O relationships observed.
5. In these analyses, we documented changing A–O trends for different groups of species, over a relatively long time period for ecological studies, one of only a few studies to examine A–O relationships over time. Further, we have shown that a temporally unvarying abundance–occupancy relationship is not universal, and we posit that variability in A–O relationships is due to human impacts on habitats, coupled with variation in species' abilities to respond to human impacts.