Contrasting demographics of tropical savanna and temperate forest eucalypts provide insight into how savannas and forests function. A case study using Corymbia clarksoniana from north‐eastern Australia

Eucalypts (Eucalyptus spp. and Corymbia spp.) dominate many communities across Australia, including frequently burnt tropical savannas and temperate forests, which receive less frequent but more intense fires. Understanding the demographic characteristics that allow related trees to persist in tropical savannas and temperate forest ecosystems can provide insight into how savannas and forests function, including grass–tree coexistence. This study reviews differences in critical stages in the life cycle of savanna and temperate forest eucalypts, especially in relation to fire. It adds to the limited data on tropical eucalypts, by evaluating the effect of fire regimes on the population biology of Corymbia clarksoniana, a tree that dominates some tropical savannas of north‐eastern Australia. Corymbia clarksoniana displays similar demographic characteristics to other tropical savanna species, except that seedling emergence is enhanced when seed falls onto recently burnt ground during a high rainfall period. In contrast to many temperate forest eucalypts, tropical savanna eucalypts lack canopy‐stored seed banks; time annual seed fall to coincide with the onset of predictable wet season rain; have very rare seedling emergence events, including a lack of mass germination after each fire; possess an abundant sapling bank; and every tropical eucalypt species has the ability to maintain canopy structure by epicormically resprouting after all but the most intense fires. The combination of poor seedling recruitment strategies, coupled with characteristics allowing long‐term persistence of established plants, indicate tropical savanna eucalypts function through the persistence niche rather than the regeneration niche. The high rainfall‐promoted seedling emergence of C. clarksoniana and the reduction of seedling survival and sapling growth by fire, support the predictions that grass–tree coexistence in savannas is governed by rainfall limiting tree seedling recruitment and regular fires limiting the growth of juvenile trees to the canopy.     

Savanna tree influence on understory vegetation and soil nutrients in northwestern Kenya

Abstract. Contrary to observations and models in which trees and herbaceous plants are viewed as competitors, we found that trees in an African savanna have positive impacts on herbaceous biomass production and composition, and on soil nutrient status. In the Turkana District of northwestern Kenya, we investigated vegetation and soil gradients along equi-angular transects radiating from the boles of individual Acacia tortilis trees. Total herbaceous biomass averaged 260 ± 17(se) g/m² at the bole and declined to 95 ± 8 g/m² in the tree interspaces. Soil organic carbon and total nitrogen concentrations were greatest (0.72 % and 0.083 %, respectively) in shallow soils near the bole and declined rapidly toward the interspaces and with increasing depth. Transects were also established between tree pairs to assess effects of differential canopy proximities. Grass production averaged 220 ± 21 g / m² below overlapping canopies, 150 ± 15 g / m² under individual canopies, and 95 ± 8 g / m² in interstitial areas. Detrended correspondence analysis revealed that shifts in species composition were correlated with distance from tree bole out to the edge of the canopy. Species response, in terms of relative cover, to increasing distance from the bole, seemed to fall into five general classes: 1) greatest at the bole, 2) increasing with distance from the bole, 3) greatest in the mid canopy zone, 4) least at the bole and 5) no response. Trees did not influence herbaceous compositionbeyondtree canopies. It is assumed that shade cast by the tree canopy with subsequent reductions of understory water stress and temperature and increased nutrient concentrations may be the most important factors affecting understory soil and vegetation.     

Spatial and temporal soil moisture resource partitioning by trees and grasses in a temperate savanna, Arizona, USA

Stable isotope analysis was used to determine sources of water used by coexisting trees and grasses in a temperate savanna dominated by Quercus emoryi Torr. We predicted that (1) tree seedlings and bunchgrasses utilize shallow sources of soil water, (2) mature savanna trees use deeper sources of water, and (3) trees switch from shallow to deep water sources within 1 year of germination. We found that Q. emoryi trees, saplings, and seedlings (about 2 months, 1 year, and 2 years old), and the dominant bunchgrass [Trachypogon montufari (H.B.K.) Nees.] utilized seasonally available moisture from different depths within the soil profile depending on size/age relationships. Sapling and mature Q. emoryi acquired water from >50 cm deep, 2-month-old seedlings utilized water from <15 cm, and 1- and 2-year-old seedlings and grasses used water from between 20 cm and 35 cm. This suggests that very young seedlings are decoupled from grasses in this system, which may facilitate germination and early establishment of Q. emoryi within extant stands of native grasses. The potential for subsequent interaction between Q. emoryi and native grasses was evidenced by similar patterns of soil water use by 1- and 2-year-old seedlings and grasses. Q. emoryi seedlings did not switch from shallow to deep sources of soil water within 2 years of germination: water use by these seedlings apparently becomes independent of water use by grasses after 2 years of age. Finally, older trees (saplings, mature trees) use water from deeper soil layers than grasses, which may facilitate the stable coexistence of mature trees and grasses. Potential shifts in the seasonality of precipitation may alter interactions between woody plants and grasses within temperate savannas characterized by bimodal precipitation regimes: reductions in summer precipitation or soil moisture may be particularly detrimental to warm-season grasses and seedlings of Q. emoryi. Received: 21 November 1996 / Accepted: 2 May 1997     

Roles of dominant understory Sasa bamboo in carbon and nitrogen dynamics following canopy tree removal in a cool‐temperate forest in northern Japan

To clarify the role of dense understory vegetation in the stand structure, and in carbon (C) and nitrogen (N) dynamics of forest ecosystems with various conditions of overstory trees, we: (i) quantified the above‐ and below‐ground biomasses of understory dwarf bamboo (Sasa senanensis) at the old canopy‐gap area and the closed‐canopy area and compared the stand‐level biomasses of S. senanensis with that of overstory trees; (ii) determined the N leaching, soil respiration rates, fine‐root dynamics, plant area index (PAI) of S. senanensis, and soil temperature and moisture at the tree‐cut patches (cut) and the intact closed‐canopy patches (control). The biomass of S. senanensis in the canopy‐gap area was twice that at the closed‐canopy area. It equated to 12% of total biomass above ground but 41% below ground in the stand. The concentrations of NO₃^? and NH₄⁺ in the soil solution and soil respiration rates did not significantly change between cut and control plots, indicating that gap creation did not affect the C or N dynamics in the soil. Root‐length density and PAI of S. senanensis were significantly greater at the cut plots, suggesting the promotion of S. senanensis growth following tree cutting. The levels of soil temperature and soil moisture were not changed following tree cutting. These results show that S. senanensis is a key component species in this cool‐temperate forest ecosystem and plays significant roles in mitigating the loss of N and C from the soil following tree cutting by increasing its leaf and root biomass and stabilizing the soil environment.     

Savannas after afforestation: Assessment of herbaceous community responses to wildfire versus native tree planting

Afforestation and fire exclusion are pervasive threats to tropical savannas. In Brazil, laws limiting prescribed burning hinder the study of fire in the restoration of Cerrado plant communities. We took advantage of a 2017 wildfire to evaluate the potential for tree cutting and fire to promote the passive restoration of savanna herbaceous plant communities after destruction by exotic tree plantations. We sampled a burned pine plantation (Burned Plantation); a former plantation that was harvested and burned (Harvested & Burned); an unburned former plantation that was harvested, planted with native trees, and treated with herbicide to control invasive grasses (Native Tree Planting); and two old-growth savannas which served as reference communities. Our results confirm that herbaceous plant communities on post-afforestation sites are very different from old-growth savannas. Among post-afforestation sites, Harvested & Burned herbaceous communities were modestly more similar in composition to old-growth savannas, had slightly higher richness of savanna plants (3.8 species per 50-m²), and supported the greatest cover of native herbaceous plants (56%). These positive trends in herbaceous community recovery would be missed in assessments of tree cover: whereas canopy cover in the Harvested & Burned site was 6% (less than typical of savannas of the Cerrado), the Burned Plantation and Native Tree Planting supported 34% and 19% cover, respectively. By focusing on savanna herbaceous plants, these results highlight that tree cutting and fire, not simply tree planting and fire exclusion, should receive greater attention in efforts to restore savannas of the Cerrado.     

Trees improve grass quality for herbivores in African savannas

The tree–grass interactions of African savannas are mainly determined by varying rainfall patterns and soil fertility. Large savanna trees are known to modify soil nutrient conditions, but whether this has an impact on the quality of herbaceous vegetation is unclear. However, if this were the case, then the removal of trees might also affect the structure and quality of the grass layer. We studied the impact of large nitrogen- and non-nitrogen fixing trees on the sub-canopy (SC) grass layer in low- and high-rainfall areas of differing soil fertility in eastern and southern Africa. We compared the structure and nutrient levels of SC grasses with those outside the canopy. Grass leaf nitrogen and phosphorus contents beneath tree canopies were elevated at all study sites and were up to 25% higher than those outside the canopy in the site of lowest rainfall and soil fertility. Grass leaf fibre and organic matter (OM) contents were slightly enhanced beneath tree canopies. At the site of highest rainfall and soil fertility, grasses beneath the canopy had significantly lower ratios of stem:leaf biomass and dead:living leaf material. Grass species composition differed significantly, with the highly nutritious Panicum spp. being most abundant underneath tree crowns. In the two drier study sites, soil nitrogen and OM contents were enhanced by 30% beneath trees. N-fixation capacity of trees did not contribute to the improved quality of grass under the canopy. We conclude that trees improve grass quality, especially in dry savannas. In otherwise nutrient-poor savanna grasslands, the greater abundance of high-quality grass species with higher contents of N and P and favourable grass structure beneath trees could attract grazing ungulates. As these benefits may be lost with tree clearance, trees should be protected in low fertility savannas and their benefits for grazing wildlife recognised in conservation strategies.     

Leaf diffusion resistance pattern in an oak-hornbeam forest

Four tree, five shrub, and ten herbaceous species growing naturally in an oak-hornbeam forest were used for simultaneous study of the leaf diffusive resistances in the course of several summer days. Absolute minima of the stomatal resistance in the sun tree, the shrub, and the herbaceous species leaves were 1.7 to 6.2 s cm^-1, 6.1 to 10.8 s cm^-1, and 4.8 to 9.7 (17.3 inConvallaria majalis leaves) s cm^-1, respectively. Minimum daily leaf resistances in the course of a day were noted earlier in the morning in sun leaves of large trees than in shade leaves of other species. Stomata were fully opened later in the morning and they began to close sooner in the afternoon in usual shade leaves of the plants in the interior of the forest canopy than those in sun leaves in active surfaces of the canopy (tops of tree crowns). The relatively large differences in leaf resistances found among investigated species may be explained by differences in leaf anatomy (stomata frequency and size) and in ambient leaf or plant environment caused by leaf (plant) position in different vertical layers.     

Gap characteristics and gap regeneration in primary evergreen broad-leaved forests of western Japan

Gap characteristics and regeneration in gaps were studied in some primary evergreen broad-leaved forests of the warm temperate zone in western Japan. Total observed 161 gaps covered 15.7% of the total land area of 8.23 ha. Gap density was 19. 6 gaps ha⁻¹ and mean gap size was 80.3 m². Smaller gaps (<80 m²) were much more frequent than larger ones, and gaps larger than 400 m² were rare. Gaps created by the death or the injury of single trees were 79.5%. Canopy trees died most often with broken trunks and not so often by uprooting or leaving standing-dead. Different types of gap regeneration behaviour were recognized among the major canopy tree species, though gap regeneration of the common evergreen broad-leaved tree species did not clearly depend on a species-specific gap size.Distylium racemosum, which occurred in equal importance (about 25%) in the canopy layer of each study stand, regenerates in gaps from saplings recruited before gap creation and can replace not only its own gaps but also gaps of other species. Therefore, it can be considered a typical climax species in this forest type of western Japan.Persea thunbergii, which can reproduce vegetatively, showed a similar type of gap regeneration behaviour.Castanopsis cuspidata can replace itself with relatively higher frequency by means of vegetative reproduction (stump sprouting) after gap creation.Quercus acuta andQuercus salicina did not regenerate under the current gap-disturbance regime. Though the frequency of uprooting is low, soil disturbance by uprooting seems to be important for the perpetuation of the pioneer tree species,Fagara ailanthoides, which recruits from buried seeds in the soil     

Canopy disturbance and gap partitioning promote the persistence of a pioneer tree population in a near‐climax temperate forest of the Qinling Mountains,China

An unresolved question of temperate forests is how pioneer tree species persist in mature forests. In order to understand the responsible mechanisms, we investigated a near‐climax mixed temperate forest dominated by Betula albosinensis in the Qinling Mountains of China. Through establishing four 50 m × 50 m plots, we examined the canopy disturbance characteristics and its effects on tree recruitments. We further test the intra‐ and interspecific effects on the recruitment of B. albosinensis. The obtained data demonstrated canopy disturbance was frequent but most small‐sized. The canopy gaps are caused mainly by adult B. albosinensis by snapping. The regeneration of coexistent tree species shows a distinct preference for gap size. B. albosinensis were clumped at the juvenile stage and small scales. B. albosinensis juveniles were positively associated with B. utilis juveniles and negatively associated with the conspecific and B. utilis large trees. In addition, B. albosinensis juveniles showed negative associations with contemporary other tree species. Our results suggested that canopy disturbance caused by canopy trees and gap partitioning among the coexistent tree species are important for the persistence of the mixed forest. As a main gapmaker, B. albosinensis appear to develop a self‐perpetuating life‐history trait and allow them to persist.     

Structure and regeneration of canopy species in an old-growth evergreen broad-leaved forest in Aya district,southwestern Japan

The population structure and regeneration of canopy species were studied in a 4 ha plot in an old-growth evergreen broad-leaved forest in the Aya district of southwestern Japan. The 200 m × 200 m plot contained 50 tree species, including 22 canopy species, 3,904 trees (dbh5 cm) and a total basal area of 48.3 m²/ha. Forty one gaps occurred within the plot, and both the average gap size (67.3 m²) and the total area of gap to plot area (6.9%) were small. Species found in the canopy in the plot were divided into three groups (A, B, C) based on size and spatial distribution patterns, and density in each tree size. Group A (typical species: Distylium racemosum, Persea japonica) showed a high density, nearly random distribution and an inverse J-shaped size distribution. Species in group B (Quercus salicina, Quercus acuta, Quercus gilva) were distributed contagiously with conspicuous concentration of small trees (<5 cm dbh) around gaps. However, the species in this group included few trees likely to reach the canopy in the near future. Group C included fast-growing pioneer and shade intolerant species (e.g. Cornus controversa, Carpinus tschonoskii, Fagara ailanthoides), which formed large clumps. Most gaps were not characterized by successful regeneration of group B and C but did appear to accelerate the growth of group A. Group B species appear to require long-lived or large gaps while group C species require large, catastrophic disturbances, such as landslides, for regeneration.     

Restoration of Midwestern U.S. Savannas: One Size Does Not Fit All

Lowland savannas are a rare variant of Midwestern United States savanna occurring on alluvial soils, for which reference information is sparse. To evaluate the appropriateness of using upland savanna as a surrogate source of reference information for lowland savanna, we studied a pre‐Euro‐American lowland savanna using original U.S. Public Land Survey data and other historical records. Historical vegetation was reconstructed and compared among upland savannas, lowland savannas, and lowland forests; we also evaluated potential disturbance dynamics maintaining these systems. We found that all three communities were dominated by members of the genus Quercus but also had extensive representation by many other tree species, especially notable for savannas in this region. There were no clear size–density relationships for species in the genus Quercus, indicating that these historical savannas were not characterized exclusively by large, scattered oak trees but rather by trees of many oak species and nonoak species in a wide range of size classes. Both upland and lowland savannas also contained a substantial shrub component. We found no evidence that lowland savannas were maintained by flooding, although the uneven‐aged canopy structure suggested that periodic disturbance occurred. Restoration of lowland savanna in this region should include provisions for maintaining nonoak species and shrubs, with disturbance timed to maintain an uneven‐aged canopy structure. Although the appropriateness of historical data in the face of climate change may be questionable, in this region, a warmer climate may actually help promote the “oak parkland” that was present from 8,000 BP up to Euro‐American settlement.     

Fine Root Distribution in Dehesas of Central-Western Spain

A Dehesa is a structurally complex agro-silvo-pastoral system where at least two strata of vegetation, trees and herbaceous plants coexist. We studied the root distribution of trees (Quercus ilex L.) and herbaceous plants, in order to evaluate tree and crops competition and complementarity in Dehesas of Central Western Spain. 72 soil cores of 10 cm diameter (one to two metre deep) were taken out around 13 trees. Seven trees were intercropped with Avena sativa L. and six trees were in a grazed pasture dominated by native grasses. Soil coring was performed at four distances from the tree trunks, from 2.5 (beneath canopy) till 20 m (out of the canopy). Root length density (RLD) of herbaceous plants and trees was measured using the soil core-break method. Additionally, we mapped tree roots in 51 profiles of 7 recently opened road cuts, located between 4 and 26 m of distance from the nearest tree. The depth of the road cuts varied between 2.5 and 5.5 m. Herbaceous plant roots were located mostly in the upper 30 cm, above a clayey, dense soil layer. RLD of herbaceous plants decreased exponentially with depth until 100 cm depth. Holm-oak showed a much lower RLD than herbs (on average, 2.4 versus 23.7 km m⁻³, respectively, in the first 10 cm of the soil depth). Tree RLD was surprisingly almost uniform with depth and distance to trees. We estimated a 5.2 m maximum depth and a 33 m maximum horizontal extension for tree roots. The huge surface of soil explored by tree roots (even 7 times the projection of the canopy) could allow trees to meet their water needs during the dry Mediterranean summers. The limited vertical overlap of the two root profiles suggests that competition for soil resources between trees and the herbaceous understorey in the Dehesa is probably not as strong as usually assumed.     

Composition,structure and diversity of cove forest stands in the Great Smoky Mountains: a patch dynamics perspective

Abstract. Cove forests of the Great Smoky Mountains are North American examples of old-growth temperate forest. Ecological attributes of seven stands were studied using one 0.6 - 1.0 ha plot per stand. Stand basal area (39 - 55 m²/ha) and biomass (326 - 471 Mg/ha) were high for temperate deciduous forest. Density ranged from 577 to 1075 stems/ha. All stands had a mixture of deciduous canopy species. Only rarely did a single species comprise more than half of the stand by density, basal area or biomass. Shade-intolerant species were present at low levels (1 - 5 % of total stand density). A wide range of stem diameters was characteristic of most species. However, some species lacked small stems, indicating discontinuous regeneration. Stands tended to have 10 - 20 tree species per ha and at least five species had biomass levels > 10 Mg/ha, indicating high evenness. Canopy gaps covered 10 % of the total area (2 - 21 % by stand). Gaps and conspecific patches of canopy trees > 0.05 ha in size were infrequent. Spatial analyses revealed a variety of patterns among species at inter-tree distances of 1 to 25 m. When all species were combined, juveniles showed aggregation, and adults were often hyperdispersed. Analyses for individual species confirmed that the mosaic of canopy species is influenced by non-random spatial processes. Adults of several species were aggregated at distances > 10 m. Juveniles of all major species exhibited aggregation. Several species exhibited regeneration near conspecific adults. This pattern suggested limited mobility for such species within the shifting mosaic. A diverse patchwork resulted despite the fact that many species did not exhibit segregation of adults and juveniles. Further understanding of patch dynamics and the potential for compositional steady state in cove forests requires long-term study with spatial data.     

Old trees contribute bio-available nitrogen through canopy bryophytes

Symbiotic cyanobacteria??bryophyte associations on the forest floor are shown to contribute significantly to stand-level nitrogen budgets through the process of biological nitrogen fixation (BNF), but few studies have considered the role of canopy bryophytes. Given the high biomass of epiphytic bryophytes in many tree species of the North American temperate rain forest, we suggest that canopy bryophytes may contribute substantially to stand-level N dynamics. We confirm the presence of cyanobacteria and measure rates of BNF at three heights (0, 15 and 30 m) in Sitka spruce trees across three watershed estuaries of Clayoquot Sound, British Columbia, Canada. This study is the first to report BNF by cyanobacteria associated with epiphytic and forest floor bryophytes in the coastal temperate rain forest of North America. Cyanobacteria density was significantly greater in epiphytic bryophytes compared to mosses on the forest floor, and rates of BNF were highest at 30 m in the canopy. The majority of total stand-level BNF (0.76 kg N · ha^-1 · yr^-1) occurs in the canopy, rather than on the forest floor (0.26 kg N · ha^-1 · yr^-1). We suggest that BNF by cyanobacterial-bryophyte associations in the canopy of coastal temperate rain forests is a unique source of ecosystem N, which is dependent on large, old trees with high epiphytic bryophyte biomass.     

Intra- and inter-specific variation in canopy photosynthesis in a mixed deciduous forest

 Within the same forest, photosynthesis can vary greatly among species and within an individual tree. Quantifying the magnitude of variation in leaf-level photosynthesis in a forest canopy will improve our understanding of and ability to model forest carbon cycling. This information requires extensive sampling of photosynthesis in the canopy. We used a 22-m-tall, four-wheel-drive aerial lift to reach five to ten leaves from the tops of numerous individuals of several species of temperate deciduous trees in central Massachusetts. The goals of this study were to measure light-saturated photosynthesis in co-occurring canopy tree species under field conditions, and to identify sampling schemes appropriate for canopy tree studies with challenging logistics. Photosynthesis differed significantly among species. Even though all leaves measured were canopy-top, sun-acclimated foliage, the more shade-tolerant species tended to have lower light-saturated photosynthetic rates (P _max) than the shade-intolerant species. Likewise, leaf mass per area (LMA) and nitrogen content (N) varied significantly between species. With only one exception, the shade-tolerant species tended to have lower nitrogen content on an area basis than the intolerant species, although the LMA did not differ systematically between these ecological types. Light-saturated P _max rates and nitrogen content, both calculated on either an area or a mass basis, and the leaf mass to area ratio, significantly differed not only among species, but also among individuals within species (P<0.0001 for both). Differences among species accounted for a greater proportion of variance in the P _max rates and the nitrogen content than the differences among individuals within a species (58.5–78.8% of the total variance for the measured parameters was attributed to species-level differences versus 5.5–17.4% of the variance was attributed to differences between individual trees of a given species). Furthermore, more variation is accounted for by differences among leaves in a single individual tree, than by differences among individual trees of a given species (10.7–30.4% versus 5.5–17.4%). This result allows us to compare species-level photosynthesis, even if the sample size of the number of trees is low. This is important because studies of canopy-level photosynthesis are often limited by the difficulty of canopy access. As an alternative to direct canopy access measurements of photosynthesis, it would be useful to find an ”easy-to-measure” proxy for light-saturated photosynthetic rates to facilitate modeling forest carbon cycling. Across all species in this study, the strongest correlation was between nitrogen content expressed on an area basis (mmol m^–2, N _area) and light-saturated P _max rate (μmol m^–2 s^–1, P _maxarea) (r ²=0.511). However, within a given species, leaf nitrogen was not tightly correlated with photosynthesis. Our sampling design minimized intra-specific leaf-level variation (i.e., leaves were taken only from the top of the canopy and at only one point in the season). This implies that easy-to-measure trends in nitrogen content of leaves may be used to predict the species-specific light-saturated P _max rates. Received: 16 March 1996 / Accepted: 16 August 1996     

Eucalyptus recruitment in degraded woodlands: no benefit from elevated soil fertility

Grasses and forbs compete heavily with young tree seedlings for available resources, greatly reducing tree seedling establishment success. Soil nutrient enrichment associated with agricultural intensification can increase the growth of both herbaceous and woody lifeforms growing in isolation, but may change the balance of competitive advantage when growing together. The effects of nitrogen and phosphorus enrichment on pasture biomass and competition with two Australian grassy woodland trees (Eucalyptus albens and Eucalyptus microcarpa) was investigated in a field plot trial. Soil nutrients increased pasture biomass, but had no measurable effect on tree growth in our experiment. Competition from pasture species, even at low levels, led to high tree seedling mortality and greatly reduced tree seedling growth compared with pasture-free plots. However, when pasture-free plots were excluded from the analysis, tree seedling leaf area was not strongly correlated with herbaceous biomass. Tree seedling establishment was severely restricted even at the lowest levels of pasture biomass. We conclude that increased soil fertility resulted in a competitive advantage to the pasture, and does not improve tree seedling establishment when grown either with or without exotic herbaceous pasture (grassy understorey) species.     

Disturbance of the herbaceous layer after invasion of an eutrophic temperate forest by wild boar

Disturbances of the soil and the tree canopy are crucial factors determining the diversity, composition and biomass of the herbaceous layer in forests. This study presents a detailed account of ground vegetation in permanent plots surveyed before and after invasion of wild boar (Sus scrofa) to a temperate deciduous broadleaf forest. Specifically, we aimed to quantify the effect of wild boar rooting on cover, richness and composition of spring ephemerals, summer green herbs and saplings of woody species in relation to tree canopy cover. Rooting frequency in sample plots increased from 0% in 2010 to 61% in 2013. In heavily rooted plots, the mean cover of spring ephemeral geophytes (mainly Anemone nemorosa, A. ranunculoides and Ranunculus ficaria) decreased from 75% to 39% between 2010 and 2013. Species richness of summer green herbs generally increased between 2010 and 2013 and was additionally positively affected by heavy rooting and low canopy cover. Rooting also caused heterogenization of the herbaceous layer and amplified ongoing compositional changes induced by changing light conditions. Frequency and richness of spring ephemeral and woody species remained unchanged. We conclude that overall species richness of the herbaceous layer may increase in the short‐term as a result of increased plant recruitment and seed dispersal. However, wild boar rooting can greatly reduce the ground cover of spring ephemerals in eutrophic broadleaf forests, thereby threatening their important ecological function. To avoid long‐term losses of characteristic spring flora elements, local population control of wild boar is necessary to reduce abundance and frequency of soil rooting.     

Influence of tree cover on herbaceous layer development and carbon and water fluxes in a Portuguese cork-oak woodland

Facilitation and competition between different vegetation layers may have a large impact on small-scale vegetation development. We propose that this should not only influence overall herbaceous layer yield but also species distribution and understory longevity, and hence the ecosystems carbon uptake capacity especially during spring. We analyzed the effects of trees on microclimate and soil properties (water and nitrate content) as well as the development of an herbaceous community layer regarding species composition, aboveground biomass and net water and carbon fluxes in a cork-oak woodland in Portugal, between April and November 2011.The presence of trees caused a significant reduction in photosynthetic active radiation of 35 mol m⁻² d⁻¹ and in soil temperature of 5 °C from April to October. At the same time differences in species composition between experimental plots located in open areas and directly below trees could be observed: species composition and abundance of functional groups became increasingly different between locations from mid April onwards. During late spring drought adapted native forbs had significantly higher cover and biomass in the open area while cover and biomass of grasses and nitrogen fixing forbs was highest under the trees. Further, evapotranspiration and net carbon exchange decreased significantly stronger under the tree crowns compared to the open during late spring and the die back of herbaceous plants occurred earlier and faster under trees. This was most likely caused by interspecific competition for water between trees and herbaceous plants, despite the more favorable microclimate conditions under the trees during the onset of summer drought.     

Interspecific effects on plant size inequality: evidence from a temperate savanna community

The influences of intraspecific competition on plant size inequality have been well documented, but interspecific effects on this topic remain little understood. Here we examined the effects of canopy shading and fine roots of the trees (Elaeagnus angustifolia) on size inequality of the grasses (Achnatherum splendens) in a temperate savanna community in northwest China. Three study plots of 400 m² were divided into 4-m² quadrats, within each of which (1) canopy shading was quantified by modeling cumulative direct solar radiation (CDSR) and (2) the root effect was quantified using an empirical relationship between tree fine root density (TFRD) and relative distance to tree bases. Morphological traits were measured to represent grass size. Redundancy analysis (RDA) was conducted to examine the relative influences of grass density, CDSR and TFRD on the coefficient of variation of grass size. Results showed that no significant correlation occurred between grass density and grass size inequality. Both CDSR and TFRD had significant negative correlations with grass size inequality, suggesting that canopy shading and the presence of fine roots of trees can, respectively, increase and reduce grass size inequality. Canopy shading and TFRD played competitive roles in determining grass size inequality, where the root effect was a stronger factor than canopy shading. The tree effects can substantially alter the intensity of water stress. In response, size inequality of the grasses could be influenced through size-specific growth/mortality and slowed size divergence. These mechanisms could operate together in the savanna community.     

Fire in sub‐Saharan Africa: The fuel,cure and connectivity hypothesis

Aim

Past analyses of satellite‐based fire activity in tropical savannas support the intermediate fire–productivity hypothesis (IFP), which posits a close correlation with estimates of total net primary productivity in drier savannas and declines towards the extremes. However, these analyses ignore the distinct roles played by herbaceous and woody vegetation in fire ignition and spread. We hypothesize that, as herbaceous vegetation provides the primary fuel, fire activity in African savannas is asymptotically correlated with herbaceous production. Conversely, woody production affects fires indirectly through effects on herbaceous production and its connectivity. In contrast to the IFP, we propose the fuel, cure and connectivity (FCC) conceptual model for tropical fire activity. The FCC model makes explicit the distinct role of herbaceous and woody fuels, avoiding the confounding interpretation of the role of total production, while providing opportunities to quantify fuel curability, effects of trees on herbaceous fuel growth and connectivity, and human management.

Location

Sub‐Saharan Africa (SSA).

Time period

2003–2015.

Major taxa studied

Woody and herbaceous vegetation.

Methods

We used boosted regression tree analysis to test competing models explaining fire activity: (a) aggregate fuel loads; and (b) partitioned woody and herbaceous fuel loads; both derived from MODIS leaf area index.

Results

Herbaceous fuel load was consistently most influential, providing more explanatory power than overall biomass in fire activity. Fuel curability rated second, then human population density (HPD), and woody biomass was least important. We observed an asymptotic relationship between herbaceous fuel load and fire activity consistent with the FCC model; trees promote fires at low densites but suppress fires at higher densities; fires were rare in wetter regions, emphasizing the need for fuel to cure; and fires were concentrated in areas of low human population, underscoring the crucial role of land management.

Conclusions

The proposed FCC framework provides a more nuanced understanding of fire activity in tropical ecosystems, where herbaceous biomass is the key determinant of fire activity.