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.     

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.     

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.     

Seed longevity and fire: germination responses of an exotic perennial herb in NW Patagonian grasslands (Argentina)

Fire affects grassland composition by selectively influencing recruitment. Some exotic species can increase their abundance as a consequence of fire-stimulated seed germination, but response may depend on seed age. Rumex acetosella L. (Polygonaceae, sheep's sorrel) is a cosmopolitan herb that has invaded NW Patagonia's grasslands. This species forms persistent soil seed banks and increases after disturbances, particularly fire. We studied how fire and seed longevity influence R. acetosella germination. In 2008, we conducted laboratory experiments where we exposed different-aged seeds (up to 19 years old) to heat, smoke, charcoal, ash and control treatments. Total percentage germination and mean germination time depended on both seed age and fire treatment. Germination of younger seeds decreased with increasing temperature. There was no general pattern in germination responses of different-aged seeds to smoke, charcoal and ash. While smoke improved the germination of fresh seeds, charcoal decreased germination. Germination of untreated seeds was negatively correlated with seed age, and mean germination time increased with seed age. In most treatments, fresh seeds had lower germination than 1-5-year-old seeds, indicating an after-ripening requirement. Smoke stimulates R. acetosella germination, causing successful recruitment during post-fire conditions. Fresh seeds are particularly responsive to fire factors, possibly because they have not experienced physical degradation and are more receptive to environmental stimuli. Knowing the colonisation potential from the soil seed bank of this species during post-fire conditions will allow us to predict their impact on native communities.     

Evidence for physiological seed dormancy cycling in the woody shrub Asterolasia buxifolia and its ecological significance in fire‐prone systems

• Dormancy cycling is a key mechanism that contributes to the maintenance of long‐term persistent soil seed banks, but has not been recorded in long‐lived woody shrub species from fire‐prone environments. Such species rely on seed banks and dormancy break as important processes for post‐fire recruitment and recovery.
• We used germination experiments with smoke treatments on fresh seeds and those buried for 1 year (retrieved in spring) and 1.5 years (retrieved the following late autumn) to investigate whether Asterolasia buxifolia, a shrub from fire‐prone south‐eastern Australia with physiologically dormant seeds, exhibited dormancy cycling.
• All seeds had an obligation for winter seasonal temperatures and smoke to promote germination, even after ageing in the soil. A high proportion of germination was recorded from fresh seeds. but germination after the first retrieval was significantly lower, despite high seed viability. After the second retrieval, germination returned to the initial level. This indicates a pattern of annual dormancy cycling; one of the few observations, to our knowledge, for a perennial species. Additionally, A. buxifolia’s winter temperature and smoke requirements did not change over time, highlighting the potential for seeds to remain conditionally dormant (i.e. restricted to a narrow range of germination conditions) for long periods.
• For physiologically dormant species, such as A. buxifolia, we conclude that dormancy cycling is an important driver of successful regeneration, allowing seed bank persistence, sometimes for decades, during fire‐free periods unsuitable for successful recruitment, while ensuring that a large proportion of seeds are available for recruitment when a fire occurs.

     

Fire season and intensity affect shrub recruitment in temperate sclerophyllous woodlands

The season in which a fire occurs may regulate plant seedling recruitment because of: (1) the interaction of season and intensity of fire and the temperature requirements for seed release, germination and growth; (2) post-fire rainfall and temperature patterns affecting germination; (3) the interaction of post-fire germination conditions and competition from surrounding vegetation; and (4) the interaction of post-fire germination conditions and seed predators and/or seedling herbivores. This study examined the effects of different fire intensities and fire seasons on the emergence and survival of shrubs representing a range of fire response syndromes from a summer rainfall cool climate region. Replicated experimental burns were conducted in two seasons (spring and autumn) in 2 consecutive years and fuel loads were increased to examine the effects of fire intensity (low intensity and moderate intensity). Post-fire watering treatments partitioned the effects of seasonal temperature from soil moisture. Higher intensity fires resulted in enhanced seedling emergence for hard-seeded species but rarely influenced survival. Spring fires enhanced seedling emergence across all functional groups. Reduced autumn recruitment was related to seasonal temperature inhibiting germination rather than a lack of soil moisture or competition. In Mediterranean-type climate regions, seedling emergence has been related to post-fire rainfall and exposure of seeds to seed predators. We think a similar model may operate in temperate summer rainfall regions where cold-induced dormancy over winter exposes seeds to predators for a longer time and subsequently results in recruitment failure. Our results support the theory that the effect of fire season is more predictable where there are strong seasonal patterns in climate. In this study seasonal temperature rather than rainfall appears to be more influential.     

Fire‐related cues and germination from the soil seed bank of senescent remnants of mallee vegetation on Eastern Kangaroo Island

Plant communities dominated by narrow‐leaved mallee (Eucalyptus cneorifolia) are almost entirely confined to north‐eastern Kangaroo Island, South Australia, an area which has been extensively cleared for agriculture. Consequently, surviving examples consist mostly of small remnants which are thought to be senescent due to the exclusion of fire. This senescence is associated with the loss of many native understory species. Prescribed burns have been suggested as a management tool to stimulate the restoration of native plants from the soil seed bank; however, no seed bank studies have previously been conducted on Kangaroo Island and the seed bank literature usually focuses on particular species rather than on plant communities. We conducted an experiment to investigate the effects of the fire‐related cues heat and smoke on the germination of plants from the seed bank in soil sampled from 10 long‐ungrazed narrow‐leaved mallee sites on Kangaroo Island. Eighty trays of soil were monitored in a controlled glasshouse for five months after being subjected to heat and/or smoke treatments. The overall number of native, but not exotic, plant species germinating from the soil seed bank was significantly increased by all three fire‐related treatments (heat, smoke and heat plus smoke) compared with the control (no fire‐related treatment). Different plant life forms exhibited varying responses to heat and smoke treatments. The results of this study illustrate that the application of fire‐related treatments to soil seed banks in controlled glasshouse conditions can stimulate the recruitment of native species, including several species of conservation concern. These findings also indicate the potential of using these treatments for the ex situ germination of fire dependent species for revegetation purposes and indicate aspects of prescribed burns that may be important for restoring different components of native vegetation.     

Effects of Fire on Seedling Diversity and Plant Reproduction (Sexual vs. Vegetative) in Neotropical Savannas Differing in Tree Density

Little is known about the effects of fire on the structure and species composition of Neotropical savanna seedling communities. Such effects are critical for predicting long‐term changes in plant distribution patterns in these ecosystems. We quantified richness and density of seedlings within 144 plots of 1 m² located along a topographic gradient in long‐unburned (fire protected since 1983) and recently burned (September 2005) savannas in Brazil. These savannas differ in tree density and canopy cover. Sites along the gradient, however, did not differ in species composition prior to the fire. In recently burned savannas we also evaluated the contribution of vegetative reproduction relative to sexual reproduction by quantifying richness and density of root suckers. Finally, we tested seed tolerance to pulses of high temperatures—similar to those occurring during fires on the soil surface and below—of five dominant savanna tree species. Seedlings were more abundant and diverse in unburned than in burned savannas. Seedling species composition differed among unburned and burned savannas probably reflecting early differences in root: shoot biomass allocation patterns. In recently burned savannas, root suckers were more abundant and diverse than seedlings. Relatively long exposures (>10 min) of temperatures of 90 °C reduced seed germination in all studied species suggesting a negative effect of fire on germination of seeds located at or aboveground level. Because vegetative reproduction contributes more than sexual reproduction in burned environments, frequent fires are likely to cause major shifts in species composition of Neotropical savanna plant communities, favoring clonally produced recruits along tree density/topographic gradients.     

Soil seed bank dynamics of fire-prone wooded grassland, woodland and dry forest ecosystems in Ethiopia

In order to reveal the role of soil seed banks in vegetation recovery after fire in savanna, the spatial distribution and temporal changes in the soil seed banks of regularly burning savanna in Gambella, western Ethiopia, was studied. The seedling emergence technique was employed to determine the species composition and density of the soil seed bank of six sites ranging in fire severity from wooded grassland with frequent fires over woodland with intermediate fire frequency to forest with absence of fires. Species composition and density of seeds in the soil were compared between seasons, depths and sites with different types of standing vegetation. Fourteen plant species were recorded in the soil seed bank from the grassland and woodland sites and 6 from the dry forests; 60 % of the taxa in the soil seed bank were annuals and 40 % were perennials. The soil seed banks were largely dominated by graminoids and 48–97 % of the soil seed bank in the grasslands and woodlands was of a single grass species, Hyparrhenia confinis , which was absent from the dry forests. The soil seed pools ranged from less than 100 to 4700 seeds per m² depending upon the season. The soil seed bank of graminoids was nearly empty after the onset of the rainy season whereas seeds of broadleaved herbs and woody species able to germinate were still found after this time. Floristic composition, representation of life forms and density of seeds in soil did not correspond closely with that of the standing vegetation, but within graminoids there was a strong similarity between the soil seed bank and the standing vegetation. The current fire regime of Ethiopian savanna woodlands appears to maintain the dominance of graminoids over broadleaved herbs and woody plants both as seeds in the soil and in the standing vegetation.     

A comparison of seed banks across a sand dune successional gradient at Lake Michigan dunes (Indiana,USA)

In habitats where disturbance is frequent, seed banks are important for the regeneration of vegetation. Sand dune systems are dynamic habitats in which sand movement provides intermittent disturbance. As succession proceeds from bare sand to forest, the disturbance decreases. At Indiana Dunes National Lakeshore, we examined the seed banks of three habitat types across a successional gradient: foredunes, secondary dunes, and oak savanna. There were differences among the types of species that germinated from each of the habitats. The mean seed bank density increased across the successional gradient by habitat, from 376 to 433 to 968 seeds m⁻², but with foredune and secondary dune seed bank densities being significantly lower than the savanna seed bank density. The number of seeds germinated was significantly correlated with soil organic carbon, demonstrating for this primary successional sequence that seed density increases with stage and age. The seed bank had much lower species richness than that of the aboveground vegetation across all habitats. Among sites within a habitat type, the similarity of species germinated from the seed banks was very low, illustrating the variability of the seed bank even in similar habitat types. These results suggest that restoration of these habitats cannot rely on seed banks alone.     

Smoke interacts with fire history to stimulate soil seed bank germination in Mediterranean woodlands

Aims Fire has important consequences on vegetation dynamics. In fire-prone areas, natural selection favors plant species, characterized by a large soil seed bank, and that their germination is stimulated by fire. Although seed germination stimulated by fire heat is common in the eastern Mediterranean Basin, only little is known about germination stimulation by smoke. We examined the interactive effect of aerosol smoke and fire history on the germinable soil seed bank (GSSB) community in eastern Mediterranean woodlands.     

Fire and flood: Soil‐stored seed bank and germination ecology in the endangered Carrington Falls Grevillea (Grevillea rivularis,Proteaceae)

Abstract Seed germination is dependent on the interaction between the dormancy state of a seed and the presence of favourable environmental conditions. Thus, the spectacular pulse of seedling recruitment in many Australian vegetation communities following disturbances such as fire can be attributed to changes in microsite conditions and/or the dormancy‐breaking effect of the disturbance on accumulated seed banks. Grevillea rivularis is a threatened species endemic to the area immediately above Carrington Falls in the NSW Southern Highlands. Most of the population is confined to the riparian vegetation zone in woodland and heath, and is therefore subject to periodic disturbance from fire and flood. For this species, a pulse of seedling recruitment has been recorded after fire, flood and mechanical soil disturbance. The aims of this study were to examine the density and vertical distribution of the soil‐stored seed bank and to investigate the role of heat and scarification as cues for germination of fresh and soil‐stored seed. There was a large seed bank under the canopies of established individuals (194 ± 73 seeds m^?2) and most seeds were found in the 0–2 cm and leaf‐litter layers of the soil profile. The germination response of soil‐stored and fresh seed was examined using a hierarchical series of laboratory experiments. Seeds of G. rivularis showed marked dormancy polymorphism. Thirty‐six percent of soil‐stored seed germinated without treatment, whereas no untreated fresh seeds germinated. Scarification or heating caused significant germination of dormant soil‐stored seed, but only scarification resulted in germination of dormant fresh seeds. These results highlight important differences in the dormancy state of soil‐stored and fresh seed. Thus, being a riparian species in a fire‐prone environment, the dormancy mechanisms in seeds of G. rivularis suit this species to disturbance by both fire and flood.     

Burial increases seed longevity of two Artemisia tridentata (Asteraceae) subspecies

? Premise of the study: Seed longevity and persistence in soil seed banks may be especially important for population persistence in ecosystems where opportunities for seedling establishment and disturbance are unpredictable. The fire regime, an important driver of population dynamics in sagebrush steppe ecosystems, has been altered by exotic annual grass invasion. Soil seed banks may play an active role in postfire recovery of the foundation shrub Artemisia tridentata, yet conditions under which seeds persist are largely unknown. ? Methods: We investigated seed longevity of two Artemisia tridentata subspecies in situ by retrieving seed bags that were placed at varying depths over a 2 yr period. We also sampled naturally dispersed seeds in litter and soil immediately after seed dispersal and before flowering in subsequent seasons to estimate seed persistence. ? Key results: After 24 mo, seeds buried at least 3 cm below the soil surface retained 30-40% viability whereas viability of seeds on the surface and under litter declined to 0 and < 11%, respectively. The density of naturally dispersed seeds in the seed bank was highly heterogeneous both spatially and temporally, and attrition varied significantly by region. ? Conclusions: Our study suggests that Artemisia tridentata has the potential to form a short-term soil seed bank that persists longer than has been commonly assumed, and that burial is necessary for seed longevity. Use of seeding techniques that promote burial of some seeds to aid in formation of a soil seed bank may increase restoration potential.     

Reproductive potential and seed fate of Sclerocarya birrea subsp. caffra (marula) in the low altitude savannas of South Africa

Even though Sclerocarya birrea subsp. caffra (marula) is a well-studied, keystone tree species with high ecological, commercial and cultural value, significant gaps in our understanding of its reproductive biology exist, particularly the factors limiting fruit and seed production, seed fate and the persistence of the seed bank. Therefore, a detailed quantitative assessment of these factors was conducted at five sites in the low altitude savannas of South Africa. Sites varied with respect to fire regime, large mammals, geology and rainfall. Most sites showed male-biased secondary sex ratios and the minimum fruiting stem diameter ranged between 7.1 and 15.7 cm across sites. Sites with higher levels of disturbance (fire and large browsers) had trees producing fruit at larger minimum stem diameters than sites with lower levels of disturbance. Fruit production was highly variable between individuals, within and between sites, and from year to year. Variability in fruit production across years at one site was greater than the variability across sites in one year, indicating that drivers such as weather, insect herbivory, fire and predator numbers, which vary annually, play a greater role than more constant drivers such as mammalian herbivory, soil types or long-term rainfall. No significant relationship was found between environmental variables (rainfall and temperature) and annual fruit production, indicating a trade-off between vegetative growth and reproduction between years. Since marula fruits are large and heavy, the species relies primarily on mammalian dispersal agents such as the African elephant, which have also been shown to increase the germination rate. However, rodents also appear to play a significant role in seed dispersal. Seed predation rates tend to be highest in areas of low disturbance (no fire and no large browsers). While marula has a small persistent seed bank, recruitment appears to be reliant on the current season's fruit crop. This study provides a detailed quantitative assessment of important reproductive and seed fate vital rates for future population modelling.     

Long‐term variability and rainfall control of savanna fire regimes in equatorial East Africa

Fires burning the vast grasslands and savannas of Africa significantly influence the global carbon cycle. Projecting the impacts of future climate change on fire‐mediated biogeochemical processes in these dry tropical ecosystems requires understanding of how various climate factors influence regional fire regimes. To examine climate–vegetation–fire linkages in dry savanna, we conducted macroscopic and microscopic charcoal analysis on the sediments of the past 25 000 years from Lake Challa, a deep crater lake in equatorial East Africa. The charcoal‐inferred shifts in local and regional fire regimes were compared with previously published reconstructions of temperature, rainfall, seasonal drought severity, and vegetation dynamics to evaluate millennial‐scale drivers of fire occurrence. Our charcoal data indicate that fire in the dry lowland savanna of southeastern Kenya was not fuel‐limited during the Last Glacial Maximum (LGM) and Late Glacial, in contrast to many other regions throughout the world. Fire activity remained high at Lake Challa probably because the relatively high mean‐annual temperature (~22 °C) allowed productive C₄ grasses with high water‐use efficiency to dominate the landscape. From the LGM through the middle Holocene, the relative importance of savanna burning in the region varied primarily in response to changes in rainfall and dry‐season length, which were controlled by orbital insolation forcing of tropical monsoon dynamics. The fuel limitation that characterizes the region's fire regime today appears to have begun around 5000–6000 years ago, when warmer interglacial conditions coincided with prolonged seasonal drought. Thus, insolation‐driven variation in the amount and seasonality of rainfall during the past 25 000 years altered the immediate controls on fire occurrence in the grass‐dominated savannas of eastern equatorial Africa. These results show that climatic impacts on dry‐savanna burning are heterogeneous through time, with important implications for efforts to anticipate future shifts in fire‐mediated ecosystem processes.     

The role of seed banks in vegetation dynamics and restoration of dry tropical ecosystems

Abstract. This paper reviews studies on seed banks in tropical dry vegetation and, for comparison, some wet tropical and dry subtropical vegetation. A first general conclusion is that tropical seed banks are smaller than those in temperate ecosystems. Many studies are devoted to only one species, among which are several Acacia species, both tropical and subtropical, some of which reach densities in the seed bank of up to 10000 seeds/m². The lesser importance of the seed bank strategy in tropical species may be related to higher risks of seed loss through higher mortality (fire, predation, pathogens etc.), or to intermittent germination occasions in relation to ephemeral favourable conditions (e.g. rains during the dry season). Regarding vegetation dynamics, the importance of seed banks and regeneration from seeds is most pronounced in larger forest gaps. Examples are discussed of rapid succession to forest of savannas after protection from fire, which is partly due to regeneration from the seed bank. The significance of seed banks in vegetation restoration projects is outlined and the need for additional sowing of seeds of important species underlined.     

Effect of fire on hard-coated Cistaceae seed banks and its influence on techniques for quantifying seed banks

The impact of fire on hard-coated Cistaceae (Halimium ocymoides, Cistus ladanifer, and C. salvifolius) soil seed banks in a Mediterranean 'maquis' shrubland, and its effect on seed germinability were studied. The study also contrasts the effectiveness of two widely used techniques for quantifying seed banks, the seedling emergence and the physical separation methods, in relation to fire. The null hypothesis that a massive enhancement of physically-dormant Cistaceae seed germination by fire would make use of the time-consuming physical separation technique unnecessary was tested. Fire reduced Cistaceae seed banks in the 0–2 cm deep soil layer by both seed fire-consumption and lethal temperatures, revealed by the significant decreasing of the seed bank density and by the increase of apparently-intact but soft-unviable seeds, respectively. In contrast, no damage was recorded in the 2–5 cm soil layer. A dramatic seed bank depletion (> 90%) in both soil layers was recorded one year after fire in the burnt area, coinciding with a significant increase of seedling density confined to the first post-fire year. The ecological consequences of this massive post-fire seed bank input are discussed. A germinability test revealed that germination of surviving Cistaceae seeds was significantly enhanced in all cases except for the C. salvifolious seed bank in the deeper soil layer. However, final germination levels (60–75%) did not correspond to the magnitude of seed bank depletion, especially for C. salvifolious, which suggests that other environmental factors not exclusively associated with fire may also be important in softening Cistaceae seeds. Germination enhancement by fire soil-heating was not high enough to reject the physical separation technique, at least in the deeper soil layer. The simultaneous use of both seedling emergence and physical separation is recommended for reliable seed bank estimates when a physically-dormant hard-seeded component can be expected in the soil, as in many Mediterranean ecosystems, regardless of fire occurrence.     

Unusually high‐quality soil seed banks in a Midwestern U.S. oak savanna region: variation with land use history,habitat restoration,and soil properties

An overarching conclusion in the literature is that soil seed banks rarely contain many restoration‐target species and are often liabilities rather than assets to restoration. Our objective was to evaluate composition and spatial variation of seed banks and their potential contributions to restoration, including restoration‐target species such as rare species and those characterizing historical habitats. On 64 sites in a Midwestern U.S. oak savanna landscape, we sampled soil seed banks in seven habitat types (restored oak savannas, oak woodlands, and mesic prairies; unmanaged upland oak and mesic forests; and unmanaged and managed pine plantations). The germinable seed bank was exceptionally rich in restoration‐target species. In total for the 64 sites, seedlings of 127 species emerged from seed bank samples. Of the 101 native species, 56 were restoration‐target species, an unusually high number among seed bank studies. Restoration‐target species in seed banks included 13 threatened or endangered species, in addition to 43 other specialist species associated with high‐quality native habitats or on a floral list thought to characterize historical ecosystems. When analyzed across the 64‐site gradient, seed banks differed among the seven habitat types and varied with historical (1939) land use, recent management activities that restored open‐structured habitats, and biophysical gradients of tree density, soil drainage, and soil texture. While not all restoration‐target species were detected in the seed bank, the unusually high‐quality seed bank is a potential asset to restoration and was partly structured along environmental gradients across the landscape.     

Burning grasses,poor seeds: post-fire reproduction of early-flowering Neotropical savanna grasses produces low-quality seeds

Fire is a key factor triggering ecological processes in old-growth grasslands and savannas and could have strong implications for reproduction via seeds for the herbaceous layer. In the Neotropical savannas, grasses show strong synchronous post-fire flowering, and their reproduction is often considered fire-dependent, with their massive post-fire seed production being suggested as a source of population maintenance. However, literature lacks studies to provide evidence of fire-dependent flowering and no study has assessed the quality of the post-fire seed production. Therefore, we aimed to describe a phenological pattern across early-flowering Neotropical savanna grasses in both recently burnt and unburnt cerrado communities addressing three questions: (1) Do the early-flowering species rely on fire for reproduction via seeds? (2) If no, what are the effects of fire on their reproductive phenology? (3) Does the massive seed production in post-fire cerrado communities lead to high-quality seeds? We recorded the reproductive phenology of nine early-flowering grasses for 17 weeks in unburnt and recently burnt cerrado communities. We collected the seeds, estimated the production of fertile seeds, and tested germination. No species showed a pattern of fire-dependent reproduction. Fire stimulated earlier flowering while reproduction in the unburnt community was related to continuous rainfall. Seed production following fire was of low quality, and no species produced?>?7% fertile seeds. Seed germination remained below 50% for most species. Post-fire seed production of early-flowering species led to poor seed quality, suggesting a constraint to the recruitment of new individuals of early-flowering Neotropical savanna grasses in recently burnt cerrados.

     

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.