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.     

Reproductive potential and seedling establishment of the invasive alien tree Schinus molle (Anacardiaceae) in South Africa

Schinus molle (Peruvian pepper tree) was introduced to South Africa more than 150 years ago and was widely planted, mainly along roads. Only in the last two decades has the species become naturalized and invasive in some parts of its new range, notably in semi‐arid savannas. Research is being undertaken to predict its potential for further invasion in South Africa. We studied production, dispersal and predation of seeds, seed banks, and seedling establishment in relation to land uses at three sites, namely ungrazed savanna once used as a military training ground; a savanna grazed by native game; and an ungrazed mine dump. We found that seed production and seed rain density of S. molle varied greatly between study sites, but was high at all sites (384 864–1 233 690 seeds per tree per year; 3877–9477 seeds per square metre per year). We found seeds dispersed to distances of up to 320 m from female trees, and most seeds were deposited within 50 m of putative source trees. Annual seed rain density below canopies of Acacia tortillis, the dominant native tree at all sites, was significantly lower in grazed savanna. The quality of seed rain was much reduced by endophagous predators. Seed survival in the soil was low, with no survival recorded beyond 1 year. Propagule pressure to drive the rate of recruitment: densities of seedlings and sapling densities were higher in ungrazed savanna and the ungrazed mine dump than in grazed savanna, as reflected by large numbers of young individuals, but adult : seedling ratios did not differ between savanna sites. Frequent and abundant seed production, together with effective dispersal of viable S. molle seed by birds to suitable establishment sites below trees of other species to overcome predation effects, facilitates invasion. Disturbance enhances invasion, probably by reducing competition from native plants.     

Winners and losers: tropical forest tree seedling survival across a West African forest–savanna transition

Forest encroachment into savanna is occurring at an unprecedented rate across tropical Africa, leading to a loss of valuable savanna habitat. One of the first stages of forest encroachment is the establishment of tree seedlings at the forest–savanna transition. This study examines the demographic bottleneck in the seedlings of five species of tropical forest pioneer trees in a forest–savanna transition zone in West Africa. Five species of tropical pioneer forest tree seedlings were planted in savanna, mixed/transition, and forest vegetation types and grown for 12 months, during which time fire occurred in the area. We examined seedling survival rates, height, and stem diameter before and after fire; and seedling biomass and starch allocation patterns after fire. Seedling survival rates were significantly affected by fire, drought, and vegetation type. Seedlings that preferentially allocated more resources to increasing root and leaf starch (starch storage helps recovery from fire) survived better in savanna environments (frequently burnt), while seedlings that allocated more resources to growth and resource‐capture traits (height, the number of leaves, stem diameter, specific leaf area, specific root length, root‐to‐shoot ratio) survived better in mixed/transition and forest environments. Larger (taller with a greater stem diameter) seedlings survived burning better than smaller seedlings. However, larger seedlings survived better than smaller ones even in the absence of fire. Bombax buonopozense was the forest species that survived best in the savanna environment, likely as a result of increased access to light allowing greater investment in belowground starch storage capacity and therefore a greater ability to cope with fire. Synthesis: Forest pioneer tree species survived best through fire and drought in the savanna compared to the other two vegetation types. This was likely a result of the open‐canopied savanna providing greater access to light, thereby releasing seedlings from light limitation and enabling them to make and store more starch. Fire can be used as a management tool for controlling forest encroachment into savanna as it significantly affects seedling survival. However, if rainfall increases as a result of global change factors, encroachment may be more difficult to control as seedling survival ostensibly increases when the pressure of drought is lifted. We propose B. buonopozense as an indicator species for forest encroachment into savanna in West African forest–savanna transitions.     

Is prescribed fire a suitable management tool to reduce shrub encroachment in palm savannas?

Shrub encroachment in grasslands is a worldwide problem that has many ecological consequences, transforming previously open environments into dense forests. Disruption of natural fire regimes is one of the main causes of shrub encroachment, and the use of prescribed fire is a common strategy used to restore these ecosystems. In this study, we provide information about how a palm tree savanna under a process of shrub encroachment responds to the reintroduction of fire. We describe the effects of a first fire event on vegetation composition and structure using an experimental approach. We examine a species‐specific response to the fire. After one prescribed fire event applied to four study areas of 16 ha each, we analyzed the change in vegetation physiognomy and composition in burned and control plots for 1 year. Low‐intensity prescribed fire decreased height and cover of most shrub species and increased herbaceous vegetation cover over time. We classified shrub and herbaceous species response to fire according to the time they became present and their phenological characteristics. Our results can help stakeholders to determine if prescribed fire is helpful at reducing shrub encroachment in short term in similar ecosystems, considering how plant community responds to the reintroduction of fire after decades of fire suppression.     

Seed Rain Produced by Bats and Birds in Forest Islands in a Neotropical Savanna1

The high degree of isolation of forest “islands” relative to “continental” forested areas creates a naturally fragmented landscape in the savanna ecosystem. Because fragmentation can affect the intensity and quality of biological interactions (e.g., seed dispersal) we examined the abundance and species richness of seed rain produced by birds and bats in three different parts of forest islands (center, edge, and exterior) located at the Estación Biológica del Beni, Bolivia. Despite the fact that we found higher species density of seeds in the seed rain at the center of forest islands, when comparing species richness corrected for observed differences in density, species richness was higher at the edge of islands. The three parts of the islands did not differ in total number of seeds. Three genera (Byrsonima, Ficus, and Piper) contributed the most seeds to the seed rain. We found differences in the abundance of dispersed seeds probably because of the variation related with the disturbance line, where the savanna matrix interacts with the forest islands. Carollia perspicillata, Carollia brevicauda, and Sturnira lilium were the bats that contributed most to seed dispersal within forest islands, and Schistochlamys melanopis and Tyranneutes stolzmanni were the most important birds. The movement of seeds produced by bats and birds within forest islands of the savanna is crucial to assure the continuity of ecological process and dynamics of these forest islands.     

Germinable soil seed banks in a tropical savanna: seasonal dynamics and effects of fire.

Abstract The germinable soil seed bank of a tropical eucalypt savanna of north‐eastern Australia was found to be dominated by grasses and forbs, with seed bank density ranging from 58 to 792 seeds per square metre, from a total of 53 species. Late dry season fires and the fire‐related cues, heat shock and smoke, broke the seed dormancy of a range of tropical savanna species. Heat shock promoted the germination of the species groups natives, exotics, subshrubs, ephemeral and twining perennial forbs, and the common species Indigofera hirsuta, Pycnospora lutescens and Triumfetta rhomboidea. Exposure to smoke at ambient temperature promoted germination from the soil seed bank of the species groups combined natives, upright perennial forbs and grasses, as well as the common grasses Digitaria breviglumis and Heteropogon triticeus. The germinable soil seed bank varied seasonally, increasing from the mid wet season (February) and early dry season (May) to a maximum in the late dry season (October). The effect of recent fire history on soil seed bank dynamics was limited to the immediate release of some seed from dormancy; a reduction in seed densities of subshrubs and monocots, other than grasses, in recently burnt savanna; and enhanced seed density of the ephemeral I. hirsuta in the year following fire. The seed banks of most savanna species were replenished in the year following burning.     

森林种子雨研究进展与展望

种子雨阶段是植物更新的关键环节,它连接着繁殖生产与植物后续生活史阶段,对群落结构有着重要的影响.虽然早在19世纪中叶达尔文就认识到种子扩散的重要性,然而对种子雨的广泛研究于20世纪80年代初才开始.本文聚焦于森林木本植物种子雨研究,首先介绍了种子雨监测方法,包括收集器的布置,种子雨的收集、分离和鉴定.然后综述了种子雨的4个主要研究方向:种子产量的时间和空间变化格局(包括季节变化、年际变化和空间变化)、增补限制及其在物种多样性维持中的作用、验证负密度效应假说、种子雨与其他生活史阶段(土壤种子库、幼苗、幼树及母树)的比较.未来还需要加强对种子雨的长期监测,开展增补限制的跨纬度比较研究,探讨植物早期更新阶段负密度效应沿纬度梯度的变化规律,加强数学模型以及分子标记和稳定同位素技术等新手段的运用.     

The dynamics of three shrub species in a fire-prone temperate savanna: the interplay between the seed bank, seed rain and fire regime

A model was developed to assess how the seed rain and fire regime affect seed bank dynamics and seedling establishment of three native shrub species (Acanthostyles buniifolius, Baccharis pingraea and Baccharis dracunculifolia) with different regeneration strategies, in temperate South American savanna. Seed bank and seed rain were quantified for each species under different fire regimes, and their relative roles in regeneration were evaluated. All species had short-term persistent seed banks and high annual variability in seed production. A high proportion of seeds deposited in the seed rain produced seedlings after fire; few entered the soil seed bank. Fire killed a high proportion of the seeds in the soil seed bank. Seedlings derived from the seed rain had a higher probability of surviving for 2 years than seedlings emerging from the soil seed bank. In the absence of fire, establishment depended on germination both from the seed rain and the soil seed bank, whereas with annual fire, establishment was primarily dependent on germination of seeds arriving in the annual seed rain, regardless of species’ regeneration strategies. These results help to explain changes in the vegetation of South American temperate savannas as a result of changes in fire regime and grazing management during the last 50 years. By revealing the crucial roles of the soil seed bank and seed rain in regeneration, this study provides vital information for the development of appropriate management practices to control populations of shrub species with different regeneration strategies in South American temperate savannas.     

Seed Rain and Seed Limitation in a Planted Gallery Forest in Brazil

With seeds collected monthly during one year from 53 1‐m² seed traps, we investigated the seed rain and seed limitation in a gallery forest planted in 1994 in SE Brazil. Contrasting animal‐ (zoochorous) and wind‐dispersed (anemochorous) plants we investigated (1) which aspects of the composition and structure of the vegetation influence the abundance and species richness of the seed rain; (2) if such influences differ between zoochorous and anemochorous seeds; (3) if the abundance and richness of the seed rain sampled under zoochorous and nonzoochorous plant species differ; and (4) if seed limitation (given by the proportion of sites to which seeds were not dispersed) differs between zoochorous and anemochorous plant species, and also between species that have been planted and those that further colonized the area (colonists). Seed rain was intense and dominated by anemochorous species. The overall seed rain was not influenced by the vegetation parameters we analyzed (canopy height and cover, plant size, abundance, and richness) or by the plant species above the seed trap. The abundance and richness of zoochorous seeds in a given spot was influenced by the abundance and richness of zoochorous plants in its immediate vicinity. Seed limitation was higher for anemochorous than zoochorous species and higher for planted than for colonist species. We concluded with recommendations for the initial establishment of a planted forest, including the homogeneous distribution of zoochorous plants to permit a spatially homogeneous zoochorous seedfall, which will likely enhance the chances of survival and successful establishment of seeds.     

Post‐Establishment Seedling Success in the Brazilian Cerrado: A Comparison of Savanna and Forest Species1

Due to frequent fire, low nutrient availability, and prolonged drought, tropical savanna is a stressful environment for the survival and growth of woody plant seedlings. To understand why forest species do not succeed in this environment while savanna species are able to persist, the effects of fire and woody cover on seedlings of these two functional groups were investigated in the Brazilian Cerrado. Seedlings were established in experimental plots under three densities of woody cover, in sites protected from fire and sites to be subjected to fire. There was a clear difference in the ability of savanna and forest species to survive fire. None of the three forest species were able to survive fire during the first two years of life, whereas eight of the nine savanna species were able to resprout following fire. The small seed size of the ninth savanna species, Miconia albicans, predisposed its seedlings to be sensitive to fire, because there was a strong positive correlation between seed size and survivorship. Savanna species were less dependent on woody cover than were forest species, which exhibited higher growth and survival under tree canopies than in open grassland. The low rates of establishment and survival of forest trees in savanna, combined with high sensitivity to fire, appear sufficient to prevent the expansion of forest into savanna under current fire regimes in the Cerrado.     

Effect of season of burning and removal of herbaceous cover on seedling emergence in a eucalypt savanna of north‐eastern Australia

Abstract Seedling emergence in a eucalypt savanna of north‐eastern Australia was documented over a 12‐month period, between May 1999 and May 2000. Seedling emergence for grasses, forbs and subshrubs was found to mainly occur in a brief pulse at the start of the wet season following fire or the removal of grass biomass. Only a minor number of tree and shrub seedlings were detected overall. Burning, or cutting away the grass layer in unburnt savanna, in both the early (i.e. May) and the late (i.e. October) dry seasons significantly increased seedling emergence over undisturbed savanna that had been unburnt for 3 years. Removing the grass layer in unburnt savanna, during either the early or the late dry season, triggered similar seedling densities to savanna burnt in the early dry season. Late dry season fires promoted the greatest seedling density. We attribute this to the higher intensity, late dry season fires releasing a greater proportion of seed from dormancy, coupled with the higher density of soil seed reserves present in the late dry season.     

Influence of conspecific and heterospecific adults on riparian tree species establishment during encroachment of a humid palm savanna

Woody plant encroachment of savanna ecosystems has been related to altered disturbance regimes, mainly fire suppression and herbivore exclusion. In contrast, neighbourhood interactions among resident and colonising woody species have received little attention, despite their likely influence on the pattern and rate of tree establishment. We examined how resident palm trees (Butia yatay) and established adults of two riparian forest tree species (Allophylus edulis and Sebastiania commersoniana) influenced seed arrival and seedling performance of the latter two species in a humid savanna of east-central Argentina. Seed traps and seedlings of both riparian species were placed in herbaceous openings, and beneath palm, conspecific and heterospecific adult trees in two unburned savanna patches, and were monitored for 2 years. Only seeds of the bird-dispersed Allophylus arrived in palm microsites, yet survival of Allophylus seedlings near adult palms was limited by animal damage through trampling and burrowing, a non-trophic mechanism of apparent competition. Seeds of both riparian species dispersed into conspecific microsites, although adult trees selectively reduced growth of conspecific seedlings, a pattern consistent with the “escape hypothesis”. Further, survival of Sebastiania increased in the moister Allophylus microsites, suggesting a one-way facilitative interaction between woody colonisers. Our results indicate that dispersal facilitation by resident savanna trees may be critical to riparian species invasion after fire suppression. Distance-dependent effects of conspecific and heterospecific adult trees could contribute to shape the subsequent dynamics of woody seedling establishment. Overall, we show that indirect interactions can play a prominent role in savanna encroachment by non-resident woody species.     

Comparative effects of desiccation,heat shock and high temperatures on seed germination of savanna and forest tree species

Although forest and savanna biomes predominate in tropics regions, the factors that control their distribution remain unclear. South American savannas occur in regions that are considered warm and humid enough to support forests, indicating that agents other than climate determine the occurrence of one or the other physiognomy. Herbivory, fire and water deficit have been considered environmental filters that limit the forest species encroachment in savanna physiognomies, but the effects of these filters on the capability of these species to recruit from seeds remain poorly understood. In this study we investigated how stress factors characteristic of savanna environments, such as soil desiccation, heat shocks and high temperatures affect the survival and germination of seeds from savanna and forest tree species. We found that desiccation (to 5%) reduced the germination percentage of forest seeds, but had no effect on the germination of savanna seeds. Forest seeds were less tolerant to heat shocks of 140°C and 200°C, and showed lower germination percentage at temperatures of 35 and 40°C, when compared with savanna seeds. Savanna seeds presented longer germination times and higher germination variance than forest seeds, indicating a risk‐spreading germination strategy among savanna species. The low tolerance of forest seeds to desiccation, heat shock and high temperatures may explain the low recruitment of forest trees into savanna physiognomies. Climate change models predict lower soil moisture, higher temperatures and higher fires frequency for South America biomes. Our results suggest that savanna species are likely to be more capable of withstanding the effects of these changes than forest species.     

Effects of fire and drought in a tropical eucalypt savanna colonized by rain forest

Aim This study documents the effects of multiple fires and drought on the woody structure of a north Australian savanna never grazed by domestic stock. Location The study was conducted in a 500 ha pocket of Eucalyptus‐dominated savanna surrounded by a late Quaternary lava flow. The flow is known as the Great Basalt Wall, located c. 50 km northeast of Charters Towers in semi‐arid north‐eastern Australia. This region was exposed to the largest 5‐year rainfall deficit on record between 1992 and 1996. Methods All individual woody plants were tagged within a 1.56 ha plot. Species were segregated into their habitat affinities (rain forest, ecotone, savanna) and regeneration strategy (resprouter, seeder). The survivorship of plants within these categories was analysed in relation to fire intensity from the first fire, and to each of four fires lit between 1996 and 2001. Results Before the first fire, the plot contained thirty‐one tree species including twenty‐one typical of the surrounding dry rain forest. These rain forest species were represented by small individuals and constituted <1% of the total basal area of woody plants. The basal area of savanna trees was 7.5 m² ha^?1 at the commencement of monitoring, although 31% had recently died and others had major crown damage. Further death of the drought debilitated savanna trees was substantial during the first year of monitoring and the basal area of live savanna trees declined to 1.1 m² ha^?1 after 5 years. Most species from both rain forest and savanna were classified as resprouters and are capable of regenerating from underground organs after fire. Species without this ability (rain forest seeders and ecotone seeders) were mostly eliminated after the first two consecutive fires. Among resprouters, survivorship declined as fire intensity increased and this was more pronounced for rain forest than for savanna species. Repeated burning produced a cumulative effect of decreasing survivorship for rain forest resprouters relative to savanna resprouters. Main conclusions The study provides evidence that savanna and rain forest trees differ in fire susceptibility and that recurrent fire can explain the restricted distribution of rain forest in the seasonally arid Australian tropics. The time of death of the savanna trees is consistent with the regional pattern after severe drought, and highlights the importance of medium term climate cycles for the population dynamics of savanna tree species and structure of Australian savannas.     

Monsoon rain forest seedling dynamics, northern Australia: contrasts with regeneration in eucalypt-dominated savannas

Aim To explore: (1) the relative influences of site conditions, especially moisture relations, on pathways and rates of monsoon rain forest seedling and sapling regeneration, especially of canopy dominants, in northern Australia; and (2) contrasts between regeneration syndromes of dominant woody taxa in savannas and monsoon rain forest. Location Four monsoon rain forest sites, representative of regional major habitat and vegetation types, in Kakadu National Park, northern Australia. Methods A decadal study involved: (1) initial assessment over 2.5 years to explore within‐year variability in seed rain, dormant seed banks and seedling (< 50 cm height) dynamics; and (2) thereafter, monitoring of seedling and sapling (50 cm height to 5 cm d.b.h.) dynamics undertaken annually in the late dry season. On the basis of observations from this and other studies, regeneration syndromes of dominant monsoon rain forest taxa are contrasted with comparable information for dominant woody savanna taxa, Eucalyptus and Corymbia especially. Results Key observations from the monsoon rain forest regeneration dynamics study component are that: (1) peak seed rain inputs of rain forest taxa were observed in the wet season at perennially moist sites, whereas inputs at seasonally dry sites extended into, or peaked in, the dry season; (2) dormant soil seed banks of woody rain forest taxa were dominated by pioneer taxa, especially figs; (3) longevity of dormant seed banks of woody monsoon rain forest taxa, including figs, was expended within 3 years; (4) seedling recruitment of monsoon rain forest woody taxa was derived mostly from wet season seed rain with limited inputs from soil seed banks; (5) at all sites rain forest seedling mortality occurred mostly in the dry season; (6) rain forest seedling and sapling densities were consistently greater at moist sites; (7) recruitment from clonal reproduction was negligible, even following unplanned low intensity fires. Main conclusions By comparison with dominant savanna eucalypts, dominant monsoon rain forest taxa recruit substantially greater stocks of seedlings, but exhibit slower aerial growth and development of resprouting capacity in early years, lack lignotubers in mesic species, and lack capacity for clonal reproduction. The reliance on sexual as opposed to vegetative reproduction places monsoon rain forest taxa at significant disadvantage, especially slower growing species on seasonally dry sites, given annual–biennial fires in many north Australian savannas.     

Short‐term effects of herbicides and a prescribed fire on restoration of a shrub‐encroached pine savanna

Shrub encroachment occurring worldwide in savannas and grasslands has commonly been hypothesized to result from anthropogenically altered environments. Two disturbance‐based approaches to restoration have involved: (1) application of selective herbicides to reduce density/cover of shrubs; (2) reinstatement of natural fire regimes to generate environmental conditions favoring herbaceous species. We studied short‐term responses of native shrubs, vines, and grasses in a Louisiana pine savanna to herbicides followed by a prescribed fire and fire alone. Plots established in the summer, 2013, were hand‐sprayed in the fall with Imazapyr and Triclopyr, Triclopyr alone, or no herbicide, then prescribed burned the following spring. Numbers of species of shrubs and vines at scales of 1 and 100 m², numbers of stems and regrowth of stems produced by six common species of shrubs, and the number of flowering culms of perennial C₄ grasses were assessed postfire in 2014. Compared with fire alone, herbicides followed by fire resulted in (1) small reductions in species richness of shrubs and no effects on vines, (2) fewer stems comprising shrub genets, but similar postfire regrowth of resprouting shrub stems, and (3) fewer flowering culms of C₄ grasses. Underground storage organs of savanna shrubs and vines survived both aboveground disturbances. Thus, single applications of herbicides followed by fires reduced, but did not reverse shrub encroachment and negatively affected grasses. Because effects of herbicides overrode those of prescribed fires, these disturbances did not act synergistically, suggesting that reinstating natural fire regimes should be a priority in restoration of savannas and grasslands.     

Woody structure facilitates invasion of woody plants by providing perches for birds

Woody encroachment threatens prairie ecosystems globally, and thus understanding the mechanisms that facilitate woody encroachment is of critical importance. Coastal tallgrass prairies along the Gulf Coast of the US are currently threatened by the spread of several species of woody plants. We studied a coastal tallgrass prairie in Texas, USA, to determine if existing woody structure increased the supply of seeds from woody plants via dispersal by birds. Specifically, we determined if (i) more seedlings of an invasive tree (Tridacia sebifera) are present surrounding a native woody plant (Myrica cerifera); (ii) wooden perches increase the quantity of seeds dispersed to a grassland; and (iii) perches alter the composition of the seed rain seasonally in prairie habitats with differing amounts of native and invasive woody vegetation, both underneath and away from artificial wooden perches. More T. sebifera seedlings were found within M. cerifera patches than in graminoid‐dominated areas. Although perches did not affect the total number of seeds, perches changed the composition of seed rain to be less dominated by grasses and forbs. Specifically, 20–30 times as many seeds of two invasive species of woody plants were found underneath perches independent of background vegetation, especially during months when seed rain was highest. These results suggest that existing woody structure in a grassland can promote further woody encroachment by enhancing seed dispersal by birds. This finding argues for management to reduce woody plant abundance before exotic plants set seeds and argues against the use of artificial perches as a restoration technique in grasslands threatened by woody species.     

Contingent fire effects on granivore removal of exotic woody plant seeds in longleaf pine savannas

Prescribed fire is increasingly used to inhibit woody encroachment into fire-dependent ecosystems, yet its effects on other processes influencing invasion are poorly understood. We investigated how fire influences exotic woody invasion through its effects on granivore activity, and whether these effects depend on the habitat in which seed predation takes place. We quantified seed removal for four species of exotic woody plants (Albizia julibrissin, Elaeagnus umbellata, Melia azedarach and Triadica sebifera) in 17 sites in longleaf pine savanna that varied in time since fire (one or three growing seasons post-fire) in the sandhills region of North Carolina, USA. Within each site, we established paired plots in upland and upland-wetland ecotone communities and presented seeds in depots that allowed either arthropod, or arthropod and small vertebrate access. We found that differences in seed removal with time since fire were contingent on habitat and granivore community. In ecotones, three of four species had higher proportions of seeds removed from plots that were three growing seasons post-fire than plots one growing season post-fire, whereas only T. sebifera showed this pattern in upland habitat. Allowing vertebrate granivores access to seeds enhanced seed removal, and this effect was strongest in ecotone habitat. While granivores removed seeds of all four plant species, removal of E. umbellata was significantly higher than that of the other species, suggesting that granivores exhibited seed selection. These findings suggest that ecotone habitats in this system experience greater seed removal than upland habitats, particularly as time since fire increases, and differences are mainly due to the activity of vertebrate granivores. Such differences in seed removal, together with seed selection, may contribute to variation in exotic woody invasion of longleaf pine savannas.     

Vertebrate‐mediated seed rain and artificial perches contribute to overcome seed dispersal limitation in a Mediterranean old field

Natural regeneration of vegetation is a frequent outcome of land abandonment, although the rate and diversity of such regeneration may be severely restricted by seed dispersal limitation, among other factors. In spite of this, studies aiming to quantify seed rain and test methods to enhance it, such as artificial perches, are still underrepresented in the Mediterranean. In our study, we quantified seed rain density and richness and tested the effects of artificial perches on such rain over a distance gradient on seven Mediterranean island old fields. In each of the seven sites, we positioned three sampling stations, each consisting of 1 seed trap under an artificial perch and 1 as a control on the ground, distributed at 30, 60, and 90 m from natural vegetation remnant. All traps received seeds, suggesting no overall dispersal limitation. Of the 11 seed species found, 10 were fleshy‐fruited and dispersed by vertebrates. Seed traps under perches received significantly higher seed rain of fleshy‐fruited species dispersed by birds, while ground traps received significantly more seeds of the species also dispersed by mammals, especially Rubus ulmifolius. The distance from the seed source was nonsignificant in all cases. Our study demonstrates the key role of vertebrate‐mediated seed dispersal services to overcome dispersal limitation in old fields, as well as the effective contribution of even small artificial perches in contrasting such limitation. The lack of differences over the distance gradient reveal that the upper spatial limit of dispersal limitation was not achieved.     

Fire weather risk differs across rain forest—savanna boundaries in the humid tropics of north‐eastern Australia

Alternative stable state theory has been applied to understanding the control by landscape fire activity of pyrophobic tropical rain forest and pyrophytic eucalypt savanna boundaries, which are often separated by tall eucalypt forests. We evaluate the microclimate of three vegetation types across an elevational gradient and their relative fire risk as measured by McArthur's Forest Fire Danger Index (FFDI). Microclimatic data were collected from rain forest, tall eucalypt forest and savanna sites on eight vegetation boundaries throughout the humid tropics in north Queensland over a 3‐year period and were compared with data from a nearby meteorological station. There was a clear annual pattern in daily FFDI with highest values in the austral winter dry season and lowest values in the austral summer wet season. There was a strong association of the meteorological station FFDI values with those from the three vegetation types, albeit they were substantially lower. The rank order of FFDI values among the vegetation types decreased from savanna, tall eucalypt forest, then rain forest, a pattern that was consistent across each transect. Only very rarely would rain forest be flammable, despite being adjacent to highly flammable savannas. These results demonstrate the very strong effect of vegetation type on microclimate and fire risk, compared with the weak effect of elevation, consistent with a fire–vegetation feedback. This study is the first demonstration of how vegetation type influences microclimate and fire risk across a topographically complex tropical forest–savanna gradient.