Conundrums in mixed woody–herbaceous plant systems

Aims To identify approaches to improve our understanding of, and predictive capability for, mixed tree–grass systems. Elucidation of the interactions, dynamics and determinants, and identification of robust generalizations that can be broadly applied to tree–grass systems would benefit ecological theory, modelling and land management. Methods A series of workshops brought together scientific expertise to review theory, data availability, modelling approaches and key questions. Location Ecosystems characterized by mixtures of herbaceous and woody plant life‐forms, often termed ‘savannas’, range from open grasslands with few woody plants, to woodlands or forests with a grass layer. These ecosystems represent a substantial portion of the terrestrial biosphere, an important wildlife habitat, and a major resource for provision of livestock, fuel wood and other products. Results Although many concepts and principles developed for grassland and forest systems are relevant to these dual life‐form communities, the novel, complex, nonlinear behaviour of mixed tree–grass systems cannot be accounted for by simply studying or modelling woody and herbaceous components independently. A more robust understanding requires addressing three fundamental conundrums: (1) The ‘treeness’ conundrum. What controls the relative abundance of woody and herbaceous plants for a given set of conditions at given site? (2) The coexistence conundrum. How do the life‐forms interact with each other? Is a given woody–herbaceous ratio dynamically stable and persistent under a particular set of conditions? (3) The net primary productivity (NPP) conundrum. How does NPP of the woody vegetation, the herbaceous vegetation, and the total ecosystem (woody + herbaceous) change with changes in the tree–grass ratio? Tests of the theory and conceptual models of determinants of mixed woody–herbaceous systems have been largely site‐ or region‐specific and have seldom been broadly or quantitatively evaluated. Cross‐site syntheses based on data and modelling are required to address the conundrums and identify emerging patterns, yet, there are very few data sets for which either biomass or NPP have been quantified for both the woody and the herbaceous components of tree–grass systems. Furthermore, there are few cross‐site comparisons spanning the diverse array of woody–herbaceous mixtures. Hence, initial synthesis studies should focus on compiling and standardizing a global data base which could be (1) explored to ascertain if robust generalizations and consistent patterns exist; and (2) used to evaluate the performance of savanna simulation models over a range of woody–herbaceous mixtures. Savanna structure and productivity are the result of complex and dynamic interactions between climate, soils and disturbances, notably fire and herbivory. Such factors are difficult to isolate or experimentally manipulate in order to evaluate their impacts at spatial and temporal scales appropriate for assessing ecosystem dynamics. These factors can, however, be evaluated with simulation models. Existing savanna models vary markedly with respect to their conceptual approach, their data requirements and the extent to which they incorporate mechanistic processes. Model intercomparisons can elucidate those approaches most suitable for various research questions and management applications. Conclusion Theoretical and conceptual advances could be achieved by considering a broad continuum of grass–shrub–tree combinations using data meta‐analysis techniques and modelling.     

The invasive grass,Melinis minutiflora,inhibits tree regeneration in a Neotropical savanna

Abstract Exotic grasses are becoming increasingly abundant in Neotropical savannas, with Melinis minutiflora Beauv. being particularly invasive. To better understand the consequences for the native flora, we performed a field study to test the effect of this species on the establishment, survival and growth of seedlings of seven tree species native to the savannas and forests of the Cerrado region of Brazil. Seeds of the tree species were sown in 40 study plots, of which 20 were sites dominated by M. minutiflora, and 20 were dominated by native grasses. The exotic grass had no discernable effect on initial seedling emergence, as defined by the number of seedlings present at the end of the first growing season. Subsequent seedling survival in plots dominated by M. minutiflora was less than half that of plots dominated by native species. Consequently, at the end of the third growing season, invaded plots had only 44% as many seedlings as plots with native grasses. Above‐ground grass biomass of invaded plots was more than twice that of uninvaded plots, while seedling survival was negatively correlated with grass biomass, suggesting that competition for light may explain the low seedling survival where M. minutiflora is dominant. Soils of invaded plots had higher mean Ca, Mg and Zn, but these variables did not account for the higher grass biomass or the lower seedling survival in invaded plots. The results indicate that this exotic grass is having substantial effects on the dynamics of the tree community, with likely consequences for ecosystem structure and function.     

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.     

Herbaceous vegetation responses to experimental fire in savannas and forests depend on biome and climate

Fire–vegetation feedbacks potentially maintain global savanna and forest distributions. Accordingly, vegetation in savanna and forest ecosystems should have differential responses to fire, but fire response data for herbaceous vegetation have yet to be synthesized across biomes. Here, we examined herbaceous vegetation responses to experimental fire at 30 sites spanning four continents. Across a variety of metrics, herbaceous vegetation increased in abundance where fire was applied, with larger responses to fire in wetter and in cooler and/or less seasonal systems. Compared to forests, savannas were associated with a 4.8 (±0.4) times larger difference in herbaceous vegetation abundance for burned versus unburned plots. In particular, grass cover decreased with fire exclusion in savannas, largely via decreases in C₄ grass cover, whereas changes in fire frequency had a relatively weak effect on grass cover in forests. These differential responses underscore the importance of fire for maintaining the vegetation structure of savannas and forests.     

Self‐thinning and Tree Competition in Savannas

This paper examines the feasibility of applying self‐thinning concepts to savannas and how competition with herbaceous vegetation may modify self‐thinning patterns among woody plants in these ecosystems. Competition among woody plants has seldom been invoked as a major explanation for the persistence of herbaceous vegetation in mixed tree–grass ecosystems. On the contrary, the primary resource‐based explanations for tree–grass coexistence are based on tree–grass competition (niche‐separation) that assumes that trees are inferior competitors unless deeper rooting depths provide them exclusive access to water. Alternative nonresource‐based hypotheses postulate that trees are the better competitors, but that tree populations are suppressed by mortality related to fire, herbivores, and other disturbances. If self‐thinning of woody plants can be detected in savannas, stronger evidence for resource‐limitation and competitive interactions among woody plants would suggest that the primary models of savannas need to be adjusted. We present data from savanna sites in South Africa to suggest that self‐thinning among woody plants can be detected in low‐disturbance situations, while also showing signs that juvenile trees, more so than adults, are suppressed when growing with herbaceous vegetation in these ecosystems. This finding we suggest is evidence for size‐asymmetric competition in savannas.     

Reptile and Amphibian Responses to Restoration of Fire‐Maintained Pine Woodlands

Fire‐maintained woodlands and savannas are important ecosystems for vertebrates in many regions of the world. These ecosystems are being restored by forest managers, but little information exists on herpetofaunal responses to this restoration in areas dominated by shortleaf pine (Pinus echinata). We compared habitat characteristics and herpetofaunal communities in restored pine woodlands to relatively unmanaged, second‐growth forests in the Ouachita Mountains of western Arkansas, USA. We found woodland restoration with periodic burning affected species differently; some species benefited, some species appeared negatively affected, but most species did not respond clearly either way. Overall reptile captures were significantly (p = 0.041) greater in pine‐woodlands than in unrestored forest; one species of snake and three species of lizards were captured more often in woodlands than unrestored forests. Among anurans, we found no significant difference in captures between woodlands and unrestored forests for any species. Among salamanders, we captured western slimy salamanders (Plethodon albagula) almost exclusively in unrestored forest, but captures of other species did not differ between the two treatments. Historically, the Ouachita region likely consisted of a mosaic that included both fire‐maintained habitats (woodlands, savannas, and prairies) and areas of denser forest on mesic sites that were less likely to burn. Consequently, landscapes that retain both open woodlands and denser, less‐intensely burned forest (in the form of unharvested greenbelts or separate stands) would likely promote and maintain a greater diversity of herpetofauna.     

Molecular evolution, adaptive radiation, and geographic diversification in the amphiatlantic family Rapateaceae: evidence from ndhF sequences and morphology

Rapateaceae (16 genera, approximately 100 species) is largely restricted to the tepuis and sandplains of the Guayana Shield in northern South America, with Maschalocephalus endemic to West Africa. The family has undergone extensive radiation in flower form, leaf shape, habit, and habitat. To analyze the evolution of these distributions and traits, we derived a molecular phylogeny for representatives of 14 genera, based on sequence variation in the chloroplast-encoded ndhF gene. The lowland subfamily Rapateoideae is paraphyletic and includes the largely montane subfamily Saxofridericioideae as a monophyletic subset. Overall, the morphological/anatomical data differ significantly from ndhF sequences in phylogenetic structure, but show a high degree of concordance with the molecular tree in three of four tribes. Branch lengths are consistent with the operation of a molecular clock. Maschalocephalus diverges only slightly from other Monotremae: it is the product of relatively recent, long-distance dispersal, not continental drift--only its habitat atop rifted, nutrient-poor sandstones is vicariant. The family appears to have originated approximately 65 Mya in inundated lowlands of the Guayana Shield, followed by: (1) wide geographic spread of lowland taxa along riverine corridors; (2) colonization of Amazonian white-sand savannas in the western Shield; (3) invasion of tepui habitats with frequent speciation, evolution of narrow endemism, and origin of hummingbird pollination in the western Shield; and (4) reinvasion of lowland white-sand savannas. The apparent timing of speciation in the Stegolepis alliance about 6-12 Mya occurred long after the tepuis began to be dissected from each other as the Atlantic rifted approximately 90 Mya. Given the narrow distributions of most montane taxa, this suggests that infrequent long-distance dispersal combined with vicariance accounts for speciation atop tepuis in the Stegolepis alliance.     

Do Tree Canopies Enhance Perennial Grass Restoration in California Oak Savannas?

Scattered trees in grass‐dominated ecosystems often act as islands of fertility with important influences on community structure. Despite the potential for these islands to be useful in restoring degraded rangelands, they can also serve as sites for the establishment of fast growing non‐native species. In California oak savannas, native perennial grasses are rare beneath isolated oaks and non‐native annual grasses dominate. To understand the mechanisms generating this pattern, and the potential for restoration of native grasses under oaks, we asked: what are the effects of the tree understory environment, the abundance of a dominant non‐native annual grass (Bromus diandrus), and soils beneath the trees on survival, growth, and reproduction of native perennial grass seedlings? We found oak canopies had a strong positive effect on survival of Stipa pulchra and Poa secunda. Growth and reproduction was enhanced by the canopy for Poa but negatively impacted for Stipa. We also found that Bromus suppressed growth and reproduction in Stipa and Poa, although less so for Stipa. These results suggest the oak understory may enhance survival of restored native perennial grass seedlings. The presence of exotic grasses can also suppress growth of native grasses, although only weakly for Stipa. The current limitation of native grasses to outside the canopy edge is potentially the result of interference from annual grasses under oaks, especially for short‐statured grasses like Poa. Therefore, control of non‐native annual grasses under tree canopies will enhance the establishment of S. pulchra and P. secunda when planted in California oak savannas.     

Land management affects grass biomass in the Eucalyptus tetrodonta savannas of monsoonal Australia

Abstract We surveyed herbaceous biomass across the range of Eucalyptus tetrodonta savannas in north‐western Australia. Sample sites (n = 211) were stratified within four broad geographical regions characterized by different mixes of land management regimes. Grasses dominated (87% mean) the herbaceous biomass. After controlling for climatic and edaphic gradients, herbaceous biomass was highest in the Greater Darwin region (2.2 t ha⁻¹) which is managed predominantly by Europeans, and least under semi‐traditional Aboriginal management in Arnhem Land region (1.1 t ha⁻¹). In the drier Gulf of Carpentaria and Kimberley regions, where a mix of Aboriginal, conservation and pastoral land uses occurs, fuel loads were higher than in Arnhem Land region but still considerably lower than around Darwin. Sarga was recorded in all regions except the Gulf of Carpentaria and had the highest biomass in Darwin (0.88 t ha⁻¹) and lowest biomass in the Kimberley (0.54 t ha⁻¹). The proportion of herbaceous biomass made up of perennial grasses was least in Darwin (17%) and greatest in the Gulf (77%) regions. We suggest that climate, soils and land management account for differences between the drier pastoral regions of the Gulf of Carpentaria and the Kimberley and the wet Greater Darwin region relative to the Arnhem Land region. The high frequency, and larger spatial scale, of fires in the Greater Darwin region relative to the Arnhem Land region underpins the contrasting trends in total herbaceous biomass and abundance of flammable annual grasses.     

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.     

Occurrence and specialization of vessels in Xyridales

The occurrence and variation among vessels in available parts of 41 species in 16 genera of Rapateaceae and of 20 species in the four genera of Xyridaceae were determined. The vessels in Xyridaceae are more specialized in all organs of the plant than they are in Rapateaceae. Simple and scalariform perforation plates occur in the inflorescence axes and leaves of nearly all species of Xyridaceae but only scalariform plates occur in these organs of Rapateaceae – infrequently vessels are lacking in stems and leaves, at least in early metaxylem. Vessels in roots and stems of Mono–tremeae are most specialized (simple and scalariform plates) among tribes of Rapateaceae, with those in Rapateeae intermediate, and those in Schoenocephalieae and Saxofridericieae most primitive (only scalariform perforation plates). Brief comments are made about vessels as possible indicators of relationships with other families.     

Reproductive phenology of two co‐occurring Neotropical mountain grasslands

Aim

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

Location

Serra do Cipó, Minas Gerais, Brazil.

Methods

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

Results

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

Conclusions

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

Browsing herbivores improve the state and functioning of savannas: A model assessment of alternative land‐use strategies

Changing climatic conditions and unsustainable land use are major threats to savannas worldwide. Historically, many African savannas were used intensively for livestock grazing, which contributed to widespread patterns of bush encroachment across savanna systems. To reverse bush encroachment, it has been proposed to change the cattle‐dominated land use to one dominated by comparatively specialized browsers and usually native herbivores. However, the consequences for ecosystem properties and processes remain largely unclear. We used the ecohydrological, spatially explicit model EcoHyD to assess the impacts of two contrasting, herbivore land‐use strategies on a Namibian savanna: grazer‐ versus browser‐dominated herbivore communities. We varied the densities of grazers and browsers and determined the resulting composition and diversity of the plant community, total vegetation cover, soil moisture, and water use by plants. Our results showed that plant types that are less palatable to herbivores were best adapted to grazing or browsing animals in all simulated densities. Also, plant types that had a competitive advantage under limited water availability were among the dominant ones irrespective of land‐use scenario. Overall, the results were in line with our expectations: under high grazer densities, we found heavy bush encroachment and the loss of the perennial grass matrix. Importantly, regardless of the density of browsers, grass cover and plant functional diversity were significantly higher in browsing scenarios. Browsing herbivores increased grass cover, and the higher total cover in turn improved water uptake by plants overall. We concluded that, in contrast to grazing‐dominated land‐use strategies, land‐use strategies dominated by browsing herbivores, even at high herbivore densities, sustain diverse vegetation communities with high cover of perennial grasses, resulting in lower erosion risk and bolstering ecosystem services.     

Diversity of germination strategies and seed dormancy in herbaceous species of campo rupestre grasslands

The effects of fire on the vegetation vary across continents. However, in Neotropical fire‐prone grasslands, the relationship between fire and seed germination is still poorly understood, while their regeneration, especially after strong anthropogenic disturbance, is challenging for their conservation. In the present study, we assessed diversity of germination strategies in 15 dominant herbaceous species from Neotropical altitudinal grasslands (locally known as campos rupestres). We exposed seeds to several fire‐related treatments. We also compared germination between regularly and post‐fire fruiting species. Finally, we investigated the diversity of dormancy classes aiming at better understanding the biogeography and phylogeny of seed dormancy. Germination strategies varied among families. Velloziaceae and Xyridaceae produced non‐dormant, fast‐germinating seeds. Cyperaceae and Poaceae showed an extremely low or null germination due to a high proportion of unviable or embryo‐less seeds. The seeds of campo rupestre grasslands are fire resistant, but there is no evidence that fire triggers germination in this fire‐prone ecosystem. Although heat and charred wood did not promote germination, smoke enhanced germination in one grass species and decreased the mean germination time and improved synchrony in Xyridaceae and Velloziaceae. Fire had a positive effect on post‐fire regeneration by stimulating fruit set in some Cyperaceae and Poaceae species. These species produced faster germinating seeds with higher germination percentage and synchrony compared to regularly fruiting Cyperaceae and Poaceae species. This strategy of dispersion and regeneration seems to be an alternative to the production of seeds with germination triggered by fire. Physiological dormancy is reported for the first time in several clades of Neotropical plants. Our data help advance the knowledge on the role of fire in the regeneration of Neotropical grasslands.     

Exotic Grass Invasions: Applying a Conceptual Framework to the Dynamics of Degradation and Restoration in Australia’s Tropical Savannas

Plant invasions can cause severe degradation of natural areas. The ability of an ecosystem to recover autogenically from degradation following weed control is in part determined by the type and magnitude of changes to both biotic and abiotic processes caused by the invasion and how these interact with structural and functional components of the ecosystem. Recently, a number of conceptual frameworks have been proposed to describe the dynamics of degradation and regeneration in degraded ecosystems. We assessed the utility of one of these frameworks in describing the degradation and restoration potential of Australia’s tropical savannas following exotic grass invasion. First, we identified easily measured structural characteristics of putative states. We found that a continuous cover of the exotic grasses Gamba grass (Andropogon gayanus Kunth.) and Perennial mission grass (Pennisetum polystachion (L.) Schult.) under an intact tree canopy was a common state with an understorey characterized by reduced species richness and abundance and a change in the relative contribution of functional groups. Further degradation led to a state where the canopy was severely reduced and the impacts on the understorey were more severe. In both states, the seed bank was substantially less degraded than the understorey vegetation. Guided by the framework, we combined our study with other studies to construct a conceptual model for degradation in exotic grass‐invaded savannas.     

Holocene global warming and the origin of the Neotropical Gran Sabana in the Venezuelan Guayana

Aim The assumedly anomalous occurrence of savannas and forest–savanna mosaics in the Gran Sabana – a neotropical region under a climate more suitable for tropical rain forests – has been attributed to a variety of historical, climatic, and anthropogenic factors. This paper describes a previously undocumented shift in vegetation and climate that occurred during the early Holocene, and evaluates its significance for the understanding of the origin of the Gran Sabana vegetation. Location A treeless savanna locality of the Gran Sabana (4°30′–6°45′ N and 60°34′–62°50′ W), in the Venezuelan Guayana of northern South America, at the headwaters of the Caroní river, one of the major tributaries of the Orinoco river. Methods Pollen and charcoal analysis of a previously dated peat section spanning from about the Pleistocene/Holocene boundary until the present. Results Mesothermic cloud forests dominated by Catostemma (Bombacaceae) occupied the site around the Pleistocene/Holocene boundary. During the early Holocene, a progressive but relatively rapid trend towards savanna vegetation occurred, and eventually the former cloud forests were replaced by a treeless savanna. Some time after the establishment of savannas, a marked increase in charcoal particles indicates the occurrence of the first local fires. Main conclusions The occurrence of cloud forests at the Pleistocene/Holocene boundary contradicts the historical hypothesis according to which the Gran Sabana is a relict of the hypothetical widespread savannas that have been assumed to have dominated the region during the last glaciation. The first local fires recorded in the Holocene were on savanna vegetation, which is against the hypothesis of fire as the triggering factor for the establishment of these savannas. Climate change, in the form of global warming and a persistently drier climate, emerges as the most probable cause for the forest–savanna turnover.     

Complex Effects of Prescribed Fire on Restoring the Soil Water Content in a High-Elevation Riparian Meadow, Arizona

One of the largest and rarest Bebb willow (Salix bebbiana) communities in the United States occurs at Hart Prairie, Arizona. Low recruitment of the willow over the past several decades has been linked to inadequate soil water content for seed germination and seedling establishment. We tested a hypothesis that a prescribed burn would reduce biomass of and evapotranspiration by herbaceous plants, thereby increasing soil water content. Three treatments (unburned control, early‐growing season burned, late‐growing season burned) were applied in year 2001 to replicated plots in fern‐ and grass‐dominated herbaceous communities. Soil water content (0–30 cm) was measured weekly in plots during the 2001, 2002, and 2003 growing seasons. Both early‐ and late‐season burning reduced herbaceous biomass in the fern‐dominated community in 2002 and 2003 and reduced biomass in the grass‐dominated community in 2002 but not in 2003. Soil water content increased for approximately four weeks in 2001 following the early‐season burn, but the early‐season and late‐season burns reduced soil water content in both communities over much of the 2002 and 2003 growing seasons. Thus, early‐season burning may benefit willow seed germination by increasing soil water content immediately following burning but be detrimental to germination in the second and third growing seasons after burning because of drier soil. Large temporal variation in the effect of prescribed burning on soil water content will complicate the use of fire as a restoration tool to manage soil water available for threatened plants such as Bebb willow and for recharge of groundwater.     

When is a ‘forest’ a savanna,and why does it matter?

Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high‐rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree–C₄ grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C₄ grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life‐history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire‐resistant tree species in a C₄ grass‐dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them.     

Grazing and landscape controls on nitrogen availability across 330 South African savanna sites

The availability of nitrogen (N) is an important determinant of ecosystem and community dynamics for grasslands and savannas, influencing factors such as biomass productivity, plant and herbivore composition, and losses of N to waters and the atmosphere. To better understand the controls over N availability at landscape to regional scales, we quantified a range of plant and soil characteristics at each of 330 sites in three regions of South Africa: Kruger National Park (KNP), private game reserves adjacent to KNP (private protected areas – PPAs) and Hluhluwe‐iMfolozi Park (HiP). In comparing regions and sites within regions, grazing appeared to have a strong influence on N availability. Sites in the PPAs adjacent to KNP as well as sodic and alluvial sites in general typically had the highest N availability. The high N availability of these sites was not generally associated with greater potential N mineralization, but instead with less grass biomass and more forb biomass that indicated greater grazing pressure. Whereas sodic sites had a long history of high N availability as evidenced by their high soil δ¹⁵N, the greater N availability in the PPAs over the two parks appeared to be relatively recent. Grazer biomass, average potential mineralization rates and grass biomass for HiP were greater than KNP, yet there were no differences in N availability as indexed by soil and foliar δ¹⁵N between sites in the two parks. Although the short‐term increase in N availability in PPAs is not necessarily deleterious, it is uncertain whether current productivity levels in those ecosystems is sustainable. With differences in management causing herbivore biomass to be 150% greater in the PPAs than the adjacent KNP, changes in plant communities and nitrogen cycling might lead to long‐term degradation of these ecosystems, their ability to sustain herbivore populations, and also serve as an economic resource for the region.     

Plant Community Biomass Shifts in Response to Mowing and Fencing in Invaded Oak Meadows with Non‐Native Grasses and Abundant Ungulates

Many herbaceous meadows are dominated by competitive non‐native grasses and subject to ungulate herbivory, ecological processes that shift the proportional biomass of plant groups in the community. Predicting the outcome of restoration is complicated because herbivory and competition can interact. We examined the relationship between herbivory by native black‐tailed deer and domestic sheep and dominance of non‐native grasses in Garry oak meadows, one of North America's most endangered habitat types. A 3‐year factorial experiment tested the effects of mowing and fencing on plant community biomass, categorized into eight groups by geographic origin (native/non‐native), growth form (annual/perennial), and plant type (forb/grass). To test if the rarity of native plant groups was related to herbivory, we estimated ungulate foraging preferences for each plant group. Mowing and fencing treatments interacted for annual and perennial non‐native grasses. Dominance was shifted from non‐native to native grasses only when both mowing and fencing were applied. Fencing increased the total biomass, whereas mowing had no overall effect; however, fencing alone did not affect any individual plant group. Mowing shifted dominance from grasses to forbs, although both native and non‐native forbs benefited from the increased light availability. We also noted that herbivore fecal pellet densities were greatest in the spring, which coincided with the peak season of their preferred plant group, native perennial forbs. Overall, applying both mowing and fencing was the most effective restoration treatment to increase native plant groups and biomass.