Modelling the recovery of an annual savanna grass following a fire-induced crash

Abstract The native annual Sorghum populations of the Australian wet-dry tropics are highly resilient to dry season fires. During the early wet season, however, fires that occur after the new grass population has emerged can cause catastrophic population crashes. We examined savanna plots that had been burnt in this way, and compared them with adjacent unburnt plots. We found that Sorghum densities in the burnt plots were lower on average by a factor of 10, but that some fires had reduced the density only to one-third of the unburnt plots. It is not clear whether these differences relate directly to site or seasonal factors, or to differences in the way the burning was carried out. Other vegetation components responded to the fires differently: forbs (dicotyledonous herbs) increased in cover, while perennial grasses, woody plants, and overall species richness, were not significantly affected. The amount of leaf litter declined. A population model for Sorghum based on the demography of unburnt populations predicted that they should recover from a wet season burn, taking 7–16 years to return to normal densities. However, the actual field populations did not seem to be recovering, suggesting that wet season fires not only lower densities, but may also fundamentally change population processes in these annual grasses.     

Fire season affects size and architecture of Colophospermum mopane in southern African savannas

Wildfires may be started naturally by lightning or artificially by humans. In the savanna regions of southern Africa, lightning fires tend to occur at the start of the wet season, during October and November, while anthropogenic fires are usually started during the dry season, between July and August. A long-term field manipulation experiment initiated in the Kruger National Park in 1952 was used to explore whether this seasonal divergence affects tree abundance, spatial pattern, size and architecture. After 44 years of prescribed burning treatments that simulated the seasonal incidence of lightning and anthropogenic fires, mean densities of the locally-dominant shrub, Colophospermum mopane, were 638 and 500 trees ha^–1 respectively. Trees in burnt plots had aggregated distributions while trees in unburnt plots had random distributions. Significant differences (p < 0.001) were recorded in a range of morphological parameters including tree height, canopy diameter, mean stem circumference and number of stems. The incidence of resprouting also differed significantly between treatments, with burnt trees containing a high proportion of coppiced stems. The differences in tree size and architecture between the mid-dry season and early-wet season burning plots suggest that anthropogenic fires applied during July and August cannot substitute for a natural lightning fire regime. Anthropogenic fire yields a landscape that is shorter, more scrubby and populated by numerous coppiced shrubs than the landscape generated by natural lightning fire conditions.     

Effect of fire regime on plant abundance in a tropical eucalypt savanna of north‐eastern Australia

Abstract Changes in plant abundance within a eucalypt savanna of north‐eastern Australia were studied using a manipulative fire experiment. Three fire regimes were compared between 1997 and 2001: (i) control, savanna burnt in the mid‐dry season (July) 1997 only; (ii) early burnt, savanna burnt in the mid‐dry season 1997 and early dry season (May) 1999; and (iii) late burnt, savanna burnt in the mid‐dry season 1997 and late dry season (October) 1999. Five annual surveys of permanent plots detected stability in the abundance of most species, irrespective of fire regime. However, a significant increase in the abundance of several subshrubs, ephemeral and twining perennial forbs, and grasses occurred in the first year after fire, particularly after late dry season fires. The abundance of these species declined toward prefire levels in the second year after fire. The dominant grass Heteropogon triticeus significantly declined in abundance with fire intervals of 4 years. The density of trees (>2 m tall) significantly increased in the absence of fire for 4 years, because of the growth of saplings; and the basal area of the dominant tree Corymbia clarksoniana significantly increased over the 5‐year study, irrespective of fire regime. Conservation management of these savannas will need to balance the role of regular fires in maintaining the diversity of herbaceous species with the requirement of fire intervals of at least 4‐years for allowing the growth of saplings >2 m in height. Whereas late dry season fires may cause some tree mortality, the use of occasional late fires may help maintain sustainable populations of many grasses and forbs.     

The effects of fire on the frillneck lizard (Chlamydosaurus kingii) in northern Australia

Abstract A population of frillneck lizards, Chlamydosaurus kingii, was monitored by radio telemetry and mark-recapture techniques between April 1991 and April 1994, as part of a landscape-scale fire experiment, in Kakadu National Park, Northern Territory. The study aimed to investigate both the short- and longer-term effects of fire on a lizard species in a tropical savanna where fires are frequent and often annual. Frillneck lizards are able to survive fires that occur in the first few months of the dry season by remaining perched in trees. A high level of mortality (29%) occurred during late dry-season fires, along with changes in their behavioural response to fire: sheltering in either larger trees or hollow termite mounds. Food is more accessible after fires due to the removal of ground vegetation. This is reflected in greater volume and diversity of prey in stomach contents after fires. This increase is more pronounced after late dry-season fires, possibly due to increased accessibility of prey caused by more complete vegetation removal. Frillneck lizards show an overall preference for trees with a dense canopy cover located in an area with a low density of grass. Fire has an effect on this relationship. Frillneck lizards in habitat unburnt for a number of years tend to perch in trees with a smaller canopy, whereas lizards in annually burnt habitat perch in trees with a dense canopy. Volume and composition of lizard stomach contents was broadly similar among fire treatments over a 2 year period, although termites were more predominant in stomach contents of lizards in unburnt habitat. Wet-season body condition is lower in lizards from unburnt habitat, although the reason for this is unclear. These results demonstrate the importance of different fire intensities and regimes on the ecology of a lizard species in a tropical savanna.     

Seeds of the annual grasses Schizachyrium spp. as a food resource for tropical granivorous birds

Seed dynamics of the annual tropical grasses Schizachyrium fragile (R. Br.) A. Camus, S. pachyarthron C. Gardner and S. pseudeulalia (Hosok.) S.T. Blake were studied with the aim of documenting fluxes in granivore food resources. In S. fragile, seed production began in the early dry season, and seed output was primarily influenced by seedling survival. Following seed-fall, there were 651 S. fragile seeds/m² (393 kernels/m²⁾ and 1014 S. pachyarthron seeds/m² (593 kernels/m²) across the study area, with a combined kernel biomass of 14.1 × 10³ g/ha. Seed densities remained stable through the dry season, then declined rapidly after wet season rain. Initial wet season rains of up to 25 mm, 40 mm and 50 mm resulted in seed-bank germination of < 5%, 57%, and 93%, respectively. Some seeds were buried by rain and resurfaced or germinated later, but about 30% was lost. Widespread and abrupt depletion of the seed-bank is likely to occur at the start of about 8% of wet seasons on central Cape York Peninsula, leaving little seed, either for subsequent germination or as food for granivores. Burning early in the dry season, when most seeds were still on the plants, reduced seed densities by 85%. The proportion of seeds with sound kernels was reduced in areas burnt by dry season fires, both directly after the fires and, subsequently, as a result of preferential granivore activity. Areas burnt in the dry season were thus depleted of seed earlier than were unburnt areas. Food availability for granivores should therefore be optimized by fire regimes that include a range of burning histories, including fires in both early dry and early wet seasons as well as keeping other areas unburnt.     

Impacts of changed fire regimes on tropical riparian vegetation invaded by an exotic vine

Abstract Riparian habitats are highly important ecosystems for tropical biodiversity, and highly threatened ecosystems through changing disturbance regimes and weed invasion. An experimental study was conducted to assess the ecosystem impacts of fire regimes introduced for the removal of the exotic woody vine, Cryptostegia grandiflora, in tropical north‐eastern Australian woodlands. Experimental sites in subcatchments of the Burdekin River, northern Queensland, Australia, were subjected to combinations of early wet‐season and dry‐season fires, and single and repeated fires, with an unburnt control. Woody vegetation was sampled using permanent quadrats to record and monitor plants species, number and size‐class. Sampling was conducted pre‐fire in 1999 and post‐fire in 2002. All fire regimes were effective in reducing the number and biomass of C. grandiflora shrubs and vines. Few woodland or riparian species were found to be fire‐sensitive and community composition did not change markedly under any fire regime. The more intense dry‐season fires impacted the structure of non‐target vegetation, with large reductions in the number of sapling trees (<5 cm d.b.h.) and reductions in the largest tree size‐class and total tree basal area. Unexpectedly, medium‐sized canopy trees (10–30 cm d.b.h.) appear to have been significantly benefited by fires, with decreases in number of trees of this size‐class in the absence of fire. Although the presence of C. grandiflora as a vine in riparian forest canopies changed the nature and intensity of crown combustion patterns, this did not lead to the initiation of a self‐perpetuating weed–fire cycle, as invaders were unable to take advantage of gaps caused by fire. Low intensity, early wet‐season burning, or early dry‐season burning, is recommended for control of C. grandiflora in order to minimize the fire intensity and risk of the loss of large habitat trees in riparian habitats.     

Juggling carbon: allocation patterns of a dominant tree in a fire-prone savanna

In frequently burnt mesic savannas, trees can get trapped into a cycle of surviving fire-induced stem death (i.e. topkill) by resprouting, only to be topkilled again a year or two later. The ability of savanna saplings to resprout repeatedly after fire is a key component of recent models of tree–grass coexistence in savannas. This study investigated the carbon allocation and biomass partitioning patterns that enable a dominant savanna tree, Acacia karroo, to survive frequent and repeated topkill. Root starch depletion and replenishment, foliage recovery and photosynthesis of burnt and unburnt plants were compared over the first year after a burn. The concentration of starch in the roots of the burnt plants (0.08 ± 0.01 g g⁻¹) was half that of the unburnt plant (0.16 ± 0.01 g g⁻¹) at the end of the first growing season after topkill. However, root starch reserves of the burnt plants were replenished over the dry season and matched that of unburnt plants within 1 year after topkill. The leaf area of resprouting plants recovered to match that of unburnt plants within 4–5 months after topkill. Shoot growth of resprouting plants was restricted to the first few months of the wet season, whereas photosynthetic rates remained high into the dry season, allowing replenishment of root starch reserves. ¹⁴C labeling showed that reserves were initially utilized for shoot growth after topkill. The rapid foliage recovery and the replenishment of reserves within a single year after topkill implies that A. karroo is well adapted to survive recurrent topkill and is poised to take advantage of unusually long fire-free intervals to grow into adults. This paper provides some of the first empirical evidence to explain how savanna trees in frequently burnt savannas are able to withstand frequent burning as juveniles and survive to become adults. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of fire on the bird communities of tropical woodlands and open forests in northern Australia

Birds were censused at 24 plots which had been subjected to 14 years of four experimental fire regimes (fire-exclusion, annual early burn, annual late burn and biennial burn) in Eucalyptus open forest and woodland at Munmarlary, Northern Territory. In addition, short-term responses of birds to fire at a savannah woodland site near Katherine were examined. At this site patches of the study area were burnt in April, June and August 1987, and birds were censused monthly over the period April to October 1987. Many bird species were attracted to areas that had been recently burnt. These were mostly granivorous species, and omnivorous and carnivorous species which fed on the ground. Influxes of these species occurred because fires increased accessibility to food by clearing the ground of its previous dense grass layer. Succession of bird species to long-unburnt areas was relatively limited, although species which fed or nested in the shrubby understorey occurred at greater densities in such areas. Frugivorous birds may become more common in areas protected from fire, thereby driving succession to monsoon vine forests from Eucalyptus forests, though, in terms of current fire regime, this process may be prohibitively slow. The hot fires of the mid to late Dry season may be less beneficial to granivores than the cooler fires typical of the early Dry. Accordingly, recent changes in the fire regime may have contributed to the decline of some bird species.     

Endophytic fungi assemblages from 10 <Emphasis Type="Italic">Dendrobium</Emphasis> medicinal plants (Orchidaceae)

Dendrobium is the largest genus of tropical epiphytic orchid, some of which are traditional Chinese medicinal plants. The therapeutic components varied significantly among species. Endophytic microbes (fungi) hidden in medicinal plants may play an important effect on the overall quality of herb. Investigation of fungal composition in host plants is the first step toward elucidating the relationship endophyte-therapeutic content of herbal medicine. In this study, 401 culturable fungal endophytes were isolated and identified from 10 species of medicinal Dendrobium based on morphological and molecular techniques. The results showed that endophytic fungi from Dendrobium plants exhibited high biodiversity (37 genera, about 80 species). Acremonium, Alternaria, Ampelomyces, Bionectria, Cladosporium, Colletotrichum, Fusarium, Verticillium and Xylaria were the dominant fungal endophytes. Tropical epiphytic orchids appear to vary in degree of host specificity in their endophytic fungi.     

Fire severity influences the post-fire habitat structure and abundance of a cool climate lizard

In the spring and summer of 2019–2020, the ‘Black Summer’ bushfires burned more than 97 000 km² of predominantly Eucalyptus dominated forest habitat in eastern Australia. The Black Summer bushfires prompted great concern that many species had been imperilled by the fires. Here, we investigate the effects that fire severity had on the habitat and abundance of a cool climate lizard Eulamprus tympanum that was identified as a species of concern because 37% of its habitat was burnt in the Black Summer bushfires. We quantified habitat structure and the abundance of E. tympanum at sites which were unburnt, burnt at low severity and at high severity 10, 15 and 23 months after the fires. Our classification of fire severity based on scorch height and canopy status corresponded well with the Australian Government Google Earth Engine Burnt Area Map (AUS GEEBAM) fire severity layer. Ten months after the fires, sites burnt at high severity had less canopy cover, more bare ground and less fine fuel than sites burnt at low severity or unburnt sites. The abundance of E. tympanum varied with survey occasion and was greatest during the warmest sampling period and lowest during the coolest sampling period. The abundance of E. tympanum was consistently lower on sites burnt at high severity than sites burnt at low severity or unburnt sites. Our findings show that higher severity fires had a greater effect on E. tympanum than low severity fires. Our results suggest that E. tympanum were likely to have persisted in burnt sites, with populations in low severity and unburnt sites facilitating population recovery in areas burnt at high severity. Our results also suggest that wildfire impacts on E. tympanum populations will increase because the frequency and extent of severe fires are expected to increase due to climate change.     

The effect of long‐term repeated burning and fire exclusion on above‐ and below‐ground Blackbutt (Eucalyptus pilularis) forest vegetation assemblages

We used a long‐term fire experiment in south‐east Queensland, Australia, to determine the effects of frequent prescribed burning and fire exclusion on understorey vegetation (<7.5 m) richness and density in Eucalyptus pilularis forest. Our study provided a point in time assessment of the standing vegetation and soil‐stored vegetation at two experimental sites with treatments of biennial burning, quadrennial burning since 1971–1972 and no burning since 1969. Vegetation composition, density and richness of certain plant groups in the standing and soil‐stored vegetation were influenced by fire treatments. The density of resprouting plants <3 m in height was higher in the biennially burnt treatment than in the unburnt treatment, but resprouters 3–7.5 m in height were absent from the biennial burning treatment. Obligate seeder richness and density in the standing vegetation was not significantly influenced by the fire treatments, but richness of this plant group in the seed bank was higher in the quadrennial treatment at one site and in the long unburnt treatment at the other site. Long unburnt treatments had an understorey of rainforest species, while biennial burning at one site and quadrennial burning at the other site were associated with greater standing grass density relative to the unburnt treatment. This difference in vegetation composition due to fire regime potentially influences the flammability of the standing understorey vegetation. Significant interactions between fire regime and site, apparent in the standing and soil‐stored vegetation, demonstrate the high degree of natural variability in vegetation community responses to fire regimes.     

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.     

Observations of responses to re‐introducing fire in a Basalt Plains grassland after the removal of grazing: Implications for restoration

Natural grasslands in southern Australia commonly exist in altered states. One widespread altered state is grassland pasture dominated by cool‐season (C3) native grasses maintained by ongoing grazing. This study explores the consequences of removing grazing and introducing fire as a conservation management tool for such a site. We examined the abundance of two native and three exotic species, across a mosaic of fire regimes that occurred over a three‐year period: unburnt, summer wild‐fire (>2 years previous), autumn management fire (<1 year previously) and burnt in both fires. Given that one aim of conservation management is to increase native species at the expense of exotics, the impacts of the fires were largely positive. Native grasses were at higher cover levels in the fire‐managed vegetation than in the unburnt vegetation. Of the three exotic species, one was consistently at lower density in the burnt plots compared to the unburnt plots, while the others were lower only in those plots burnt in summer. The results show that the response of a species varies significantly between different fire events, and that the effects of one fire can persist through subsequent fires. Importantly, some of the effects were large, with changes in the density of plants of over 100‐fold. Fire is potentially a cost‐effective tool to assist the ecological restoration of retired grassland pastures at large scales.     

Frequent burning promotes invasions of alien plants into a mesic African savanna

Fire is both inevitable and necessary for maintaining the structure and functioning of mesic savannas. Without disturbances such as fire and herbivory, tree cover can increase at the expense of grass cover and over time dominate mesic savannas. Consequently, repeated burning is widely used to suppress tree recruitment and control bush encroachment. However, the effect of regular burning on invasion by alien plant species is little understood. Here, vegetation data from a long-term fire experiment, which began in 1953 in a mesic Zimbabwean savanna, were used to test whether the frequency of burning promoted alien plant invasion. The fire treatments consisted of late season fires, lit at 1-, 2-, 3-, and 4-year intervals, and these regularly burnt plots were compared with unburnt plots. Results show that over half a century of frequent burning promoted the invasion by alien plants relative to areas where fire was excluded. More alien plant species became established in plots that had a higher frequency of burning. The proportion of alien species in the species assemblage was highest in the annually burnt plots followed by plots burnt biennially. Alien plant invasion was lowest in plots protected from fire but did not differ significantly between plots burnt triennially and quadrennially. Further, the abundance of five alien forbs increased significantly as the interval (in years) between fires became shorter. On average, the density of these alien forbs in annually burnt plots was at least ten times as high as the density of unburnt plots. Plant diversity was also altered by long-term burning. Total plant species richness was significantly lower in the unburnt plots compared to regularly burnt plots. These findings suggest that frequent burning of mesic savannas enhances invasion by alien plants, with short intervals between fires favouring alien forbs. Therefore, reducing the frequency of burning may be a key to minimising the risk of alien plant spread into mesic savannas, which is important because invasive plants pose a threat to native biodiversity and may alter savanna functioning.     

Demography of woody plants in relation to fire: Banksia ericifolia L.f. and Petrophile pulchella (Schrad) R.Br.

Plants of Banksia ericifolia and Petrophile pulchella are sensitive to fire. Changes in population size under different fire regimes were estimated, based on measurements of post-fire seedling emergence, seedling survival, survival and seed production in established plants of differing ages, survival of seeds held in serotinous cones and seed-release in the periods between fire. Seeds were first available at 5 years in P. pulchella and 6 years in B. ericifolia. Exact replacement would be possible when burnt at these ages, if seedling establishment were very high. Low establishment would delay replacement to 13 years of age in both species. Late summer/autumn fires of high intensity favour high establishment. Such fires at 8–10 year intervals would be tolerated without any sustained decline in numbers. Fires at 10–15 year intervals could occur regardless of season or intensity with little risk of a population decline. Large increases in numbers and density would follow fires spaced at 15–30 years. Enough seeds would be available for replacement up to about 50 years in both species. Viable seed-release in unburnt conditions was sufficient to compensate for deaths in stands over 20 years old, even with very low levels of establishment. Two variables accounted for the biggest changes in numbers and density between generations interspersed by fires; namely the age at which a stand is burned and the proportion of seeds which emerge as seedlings. A comparison with other similar species showed similarities in controls on emergence and establishment, lengths of primary juvenile periods and life spans.     

Analyzing the Spatial Structure of Broughtonia cubensis (Orchidaceae) Populations in the Dry Forests of Guanahacabibes,Cuba

A current key issue in ecology is the role of spatial effects on population and community dynamics. In this paper, we test several hypotheses related to spatial structures and coexistence of epiphytic tropical orchid species with special emphasis on the endemic species Broughtonia cubensis. More specifically, we explored the spatial structure of orchid–host plant communities at three different levels of organization (occupied vs. nonoccupied host trees, trees with B. cubensis vs. other orchids, and reproductive vs. nonreproductive B. cubensis plants). We mapped all potential host trees and orchids at three 20 × 20 m plots and applied techniques of spatial point pattern analysis such as mark connection and mark correlation functions to evaluate departures from randomized communities. We found spatial aggregation of trees with epiphytic orchids and segregation between trees with and without epiphytic orchids, and that there was an intraspecific spatial aggregation of B. cubensis in relation to the other seven epiphytic orchid species. Furthermore, we found spatial aggregation of reproductive B. cubensis individuals and segregation between reproductive and nonreproductive individuals on their phorophytes. Thus, orchid–host plant communities show hierarchical spatial structuring with aggregation and segregation at different levels of organization. Our results point to an enhancement of local species in the coexistence of tropical epiphytic orchid communities, by reducing competition through niche differentiation.     

Landscape analysis of Aboriginal fire management in Central Arnhem Land, north Australia

Aim To describe the spatial and temporal pattern of landscape burning with increasing distance from Aboriginal settlements. Location Central Arnhem Land, a stronghold of traditional Aboriginal culture, in the Australian monsoon tropics. Methods Geographical information system and global positioning system technologies were used to measure spatial and temporal changes in fire patterns over a one decade period in a 100 × 80 km area that included a cluster of Aboriginal settlements and a large uninhabited area. The major vegetation types were mapped and fire activity was assessed by systematic visual interpretation of sequences of cloud‐free Landsat satellite images acquired in the first (May to July) and second (August to October) halves of the 7‐month dry season. Fire activity in the middle and end of one dry season near an Aboriginal settlement was mapped along a 90‐km field traverse. Canopy scorch height was determined by sampling burnt areas beside vehicle tracks. Results Satellite fire mapping was 90% accurate if the satellite pass followed shortly after a fire event, but the reliability decayed dramatically with increasing time since the fire. Thus the satellite mapping provided a conservative index of fire activity that was unable to provide reliable estimates of the spatial extent of individual fires. There was little landscape fire activity in the first half of the dry season, that was mostly restricted to areas immediately surrounding Aboriginal settlements, with burning of both inhabited and uninhabited landscapes concentrated in the second half of the dry season. The mean decadal fire indices for the three dominant vegetation types in the study area were three in the plateau savanna, two in the sandstone and five in the wet savanna. The spatial and temporal variability of Aboriginal burning apparent in the satellite analyses were verified by field traverse surrounding a single settlement. Fires set by Aborigines had low scorch height of tree crowns reflecting low intensity, despite generally occurring late in the dry season. Conclusions Our findings support the idea that Aboriginal burning created a fine‐scale mosaic of burnt and unburnt areas but do not support the widely held view that Aboriginal burning was focused primarily in the first half of the dry season (before July). The frequency and scale of burning by Aborigines appears to be lower compared with European fire regimes characterized by fires of annual or biennial frequencies that burn large areas. The European fire regime appears to have triggered a positive feedback cycle between fire frequency and flammable grass fuels. The widely advocated management objective of burning in the first half of the dry season burning provides one of the few options to control fires once heavy grass fuel loads have become established, however we suggest it is erroneous to characterize such a regime as reflecting traditional Aboriginal burning practices. The preservation of Aboriginal fire management regimes should be a high management priority given the difficulty in breaking the grass‐fire cycle once it has been initiated.     

Effect of fires on soil nutrient availability in an open savanna in Central Brazil

Fire is common in savannas but its effects on soil are poorly understood. We analyzed long-term effects of fire on surface soil of an open Brazilian savanna (campo sujo) in plots submitted to different fire regimes during 18 years. The five fire regimes were: unburned, quadrennial fires in middle dry season, and biennial fires in early, middle or late dry season. Soil was collected during the wet and the middle dry season of 2008, and analyzed for pH, organic matter, total N, potential acidity, exchangeable cations and available P, S, Mn, Cu, Zn and Fe. We applied multivariate analysis to search for patterns related to fire regimes, and to local climate, fuel, and fire behavior. Spearman test was used to establish correlations between soil variables and the multivariate analysis gradient structure. Seasonal differences were tested using t-test. We found evidence of long-term fire effects: the unburned plot was segregated mainly by lower soil pH; the quadrennial plot was also segregated by lower soil pH and higher amount of exchangeable cations; the time of burning during the dry season in biennial plots did not significantly affect soil availability of nutrients. Differences in elements amounts due to the season of soil sampling (wet or dry) were higher than due to the effect of fires. Higher availability of nutrients in the soil during the wet season was probably related to higher nutrient inputs via rainfall and higher microbial activity.     

Water relations of epiphytic and terrestrially-rooted strangler figs in a Venezuelan palm savanna

Water use patterns of two species of strangler fig, Ficus pertusa and F. trigonata, growing in a Venezuelan palm savanna were contrasted in terms of growth phase (epiphyte and tree) and season (dry and wet). The study was motivated by the question of how C3 hemiepiphytes accommodate the marked change in rooting environment associated with a life history of epiphytic establishment followed by substantial root development in the soil. During the dry season, stomatal opening in epiphytic plants occurred only during the early morning, maximum stomatal conductances were 5 to 10-fold lower, and midday leaf water potentials were 0.5–0.8 MPa higher (less negative) than in conspecific trees. Watering epiphytes of F. pertusa during the dry season led to stomatal conductances comparable to those exhibited by conspecific trees, but midday leaf water potentials were unchanged. During the rainy season, epiphytes had lower stomatal conductances than conspecific trees, but leaf water potentials were similar between the two growth phases. There were no differences in ¹³C between the two growth phases for leaves produced in either season. Substrate water availability differed between growth phases; tree roots extended down to the permanent water table, while roots of epiphytic plants were restricted to material accumulated behind the persistent leaf bases of their host palm tree, Copernicia tectorum. Epiphytic substrate moisture contents were variable during both seasons, indicating both the availability of some moisture during the dry season and the possibility of intermittent depletion during the rainy season. Epiphytic strangler figs appear to rely on a combination of strong stomatal control, maintenance of high leaf water potentials, and perhaps some degree of stem water storage to cope with the fluctuating water regime of the epiphytic environment.