Soil temperature and depth of legume germination during early and late dry season fires in a tropical eucalypt savanna of north‐eastern Australia

Abstract Temperatures that significantly increase seed germination of some tropical legumes (i.e. 80–100°C) were documented in the topsoil during the passage of early (May) and late (October) dry season fires in a tropical eucalypt savanna of north‐eastern Australia. Elevated temperatures penetrated at least 30 mm into the soil during the higher‐intensity, late dry season fires, but were only detected at 10 mm during the early dry season fires. The depth from which germination of two native legume forbs Galactia tenuiflora and Indigofera hirsuta occurred was positively related to the temperature elevation in the topsoil and was greater after late compared with early dry season fires. A broader range in germination depth, resulting in higher seedling densities, was recorded for I hirsuta after late dry season fires. These results suggest that seedling emergence of native leguminous forbs is likely to occur at a greater density after late rather than early dry season fires in tropical eucalypt savannas of north‐eastern Australia. Therefore, the season of burning, as a result of its relationship to fire intensity, can influence species composition through its effect on seed germination.     

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.     

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.     

The impact of experimental fire regimes on seed production in two tropical eucalypt species in northern Australia

Abstract This study investigated the effect of three experimental fire regimes on the fecundity, ovule development and seedfall of two common wet-dry tropical savanna eucalypts, Eucalyptus minima and Eucalyptus tetrodonta, in northern Australia. Both species flower early in the dry season and ovule development occurs during the dry season. This coincides with a period of frequent fires. The three fire regimes considered were applied for four years between 1990 and 1994. These regimes were (i) Unburnt, (ii) Early, fires lit early in the dry season, and (iii) Late, fires lit late in the dry season. The treatments were applied to nine catchments (15–20 km²) with each fire regime replicated three times. Fire intensity typically increases as the dry season proceeds. Therefore, early dry season fires generally differ from late dry season fires in both their intensity and their timing in relation to the reproductive phenology of the eucalypts. Late dry season burning significantly reduced the fecundity of both species, whereas Early burning had no significant effect. Ovule success was significantly reduced by the Early burning for both species. The Late burning significantly reduced ovule success in E. tetrodonta, but not in E. miniata. The results suggest that fire intensity and fire timing may both be important determinants of seed supply. Fire intensity may be a determinant of fecundity, whereas fire timing in relation to the reproduction phenology may have a significant impact on ovule survival. Both fire regimes resulted in a substantial reduction in seed supply compared with the Unburnt treatment. This may have a significant impact on seedling regeneration of these tropical savanna eucalypts.     

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.     

内蒙古典型草原群落实生苗动态对刈割和放牧干扰的响应

 该文通过刈割和放牧样地的实生苗动态观测,试图揭示刈割和放牧干扰对冰草(Agropyron cristatum)、克氏针茅(Stipa krylovii)和糙隐子 草(Cleistogenes squarrosa )等 3种禾草实生苗出生、存活和死亡季节动态的影响,以及这种效应是如何受气温和降水影响的。利用有重复测 量的方差分析,比较了不同刈割留茬高度(0、5和15 cm)与放牧处理之间在实生苗密度、死亡率和出现率的时间动态方面的差异。结果表明：3 种禾草的实生苗密度自5月下旬至8月中旬一直呈增加态势。刈割在一定程度上抑制了克氏针茅的实生苗出现率,原因可能是相对高大的克氏针 茅在前一年刈割时会被去掉相当多的生殖部分,其种子数量必然受到影响,导致实生苗出现率的降低。同时,刈割对相对低矮的糙隐子草实生 苗有正向效应 ,即促进其实生苗的出现,可能的原因是刈割使植被高度降低、植被稀疏,植被冠层以下至地面的光照较好、地温较高,同时位 于下层的糙隐子草受到的高大植株的竞争压力减小。生长季早期和晚期实生苗死亡率较高,这与这两个时期的干旱和低温有关,而自6月中旬至 7月初之间的死亡率小高峰也是先前的干旱少雨导致的。3种禾草实生苗的大规模出现几乎都在5～7月,这与该时期相对适宜的气温、地温和水 分条件有关。对实生苗更新相关过程 (如种子萌发、休眠、种子库存量等)进行更加深入细致的研究,将进一步揭示种子生产和有性繁殖在群落 过程中的作用。     

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.     

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.     

内蒙古典型草原群落实生苗动态对刈割和放牧干扰的响应

该文通过刈割和放牧样地的实生苗动态观测,试图揭示刈割和放牧干扰对冰草(Agropyron cristatum)、克氏针茅(Stipa krylovii)和糙隐子 草(Cleistogenes squarrosa )等 3种禾草实生苗出生、存活和死亡季节动态的影响,以及这种效应是如何受气温和降水影响的。利用有重复测 量的方差分析,比较了不同刈割留茬高度(0、5和15 cm)与放牧处理之间在实生苗密度、死亡率和出现率的时间动态方面的差异。结果表明：3 种禾草的实生苗密度自5月下旬至8月中旬一直呈增加态势。刈割在一定程度上抑制了克氏针茅的实生苗出现率,原因可能是相对高大的克氏针 茅在前一年刈割时会被去掉相当多的生殖部分,其种子数量必然受到影响,导致实生苗出现率的降低。同时,刈割对相对低矮的糙隐子草实生 苗有正向效应 ,即促进其实生苗的出现,可能的原因是刈割使植被高度降低、植被稀疏,植被冠层以下至地面的光照较好、地温较高,同时位 于下层的糙隐子草受到的高大植株的竞争压力减小。生长季早期和晚期实生苗死亡率较高,这与这两个时期的干旱和低温有关,而自6月中旬至 7月初之间的死亡率小高峰也是先前的干旱少雨导致的。3种禾草实生苗的大规模出现几乎都在5～7月,这与该时期相对适宜的气温、地温和水 分条件有关。对实生苗更新相关过程 (如种子萌发、休眠、种子库存量等)进行更加深入细致的研究,将进一步揭示种子生产和有性繁殖在群落 过程中的作用。     

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.     

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.     

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.     

Effects of fire regimen on two species of epiphytic orchids in tropical savannas of the Northern Territory

The effect of four fire regimens (early dry season annual, late dry season annual, early dry season biennial, unburnt) on the numbers of epiphytic orchids in a savanna community in Kakadu National Park was investigated. These fire regimens had been maintained on I ha plots for 16 years. Two species of epiphytic orchid were present — Cymbidium canaliculatum and Dendrobium affine. The numbers of Cymbidium plants were insufficient to enable statistical analysis, but plants were present in the unburnt, early annual and early biennial fire treatments. Dendrobium occurred in substantial numbers irrespective of fire treatment, but the extent of colonization of all available host trees was greater in the unburnt than the burnt treatments. The lower numbers of Dendrobium plants in burnt treatments was due to a lower proportion of potential host trees being colonized and a lower number of orchids on each colonized tree. The proportion of small trees colonized was markedly lower in burnt treatments. Frequent fires late in the dry season may prevent recruitment of the main host species. Eucalyptus tectifica. The survival of orchids despite frequent fires depended on the availability of relatively protected sites for colonization and the ability of the orchids to withstand some fire damage.     

Protected from fire,but not from harm: seedling emergence of savanna grasses is constrained by burial depth

ABSTRACT

Background

The Cerrado is a fire-prone Neotropical savanna and grass seeds are the main component of the soil seed bank. Although grass seeds are presumed to tolerate heat pulses during fires, the assumption that these seeds will be recruited may be an overestimation, since it does not account for factors that limit seedling emergence from the soil.     

Effects of seed phosphorus concentration on the emergence and growth of subterranean clover (Trifolium subterraneum)

Subterranean clover seed (Trifolium subterraneum cv. Dalkeith) with phosphorus concentrations of 0.75% (high P seed) and 0.48% (low P seed) and of uniform size (2.0–2.4 mm diameter) was used to measure the effect of seed P concentrations on seedling emergence and growth.Seedling emergence numbers were 35% greater for the high P seed, and this effect was independent of external P supply. High P seed also emerged more quickly than low P seed.Leaf emergence was faster and shoot dry weight was greater for seedlings grown from high compared with low P seed, but only when external P supply was deficient for plant growth. Phosphorus concentrations in the shoots of two-week old seedlings were 32–51% higher for high P seed, although by four weeks plants grown from high and low P seed had similar concentrations of P in their shoots. We suggest that establishing pastures using high P seed would improve both early and late season pasture production.     

Fire frequency and biodiversity conservation in Australian tropical savannas: implications from the Kapalga fire experiment

Abstract Every year large proportions of northern Australia's tropical savanna landscapes are burnt, resulting in high fire frequencies and short intervals between fires. The dominant fire management paradigm in these regions is the use of low‐intensity prescribed fire early in the dry season, to reduce the incidence of higher‐intensity, more extensive wildfire later in the year. This use of frequent prescribed fire to mitigate against high‐intensity wildfire has parallels with fire management in temperate forests of southern Australia. However, unlike in southern Australia, the ecological implications of high fire frequency have received little attention in the north. CSIRO and collaborators recently completed a landscape‐scale fire experiment at Kapalga in Kakadu National Park, Northern Territory, Australia, and here we provide a synthesis of the effects of experimental fire regimes on biodiversity, with particular consideration of fire frequency and, more specifically, time‐since‐fire. Two recurring themes emerged from Kapalga. First, much of the savanna biota is remarkably resilient to fire, even of high intensity. Over the 5‐year experimental period, the abundance of most invertebrate groups remained unaffected by fire treatment, as did the abundance of most vertebrate species, and we were unable to detect any effect of fire on floristic composition of the grass‐layer. Riparian vegetation and associated stream biota, as well as small mammals, were notable exceptions to this general resilience. Second, the occurrence of fire, independent of its intensity, was often the major factor influencing fire‐sensitive species. This was especially the case for extinction‐prone small mammals, which have suffered serious population declines across northern Australia in recent decades. Results from Kapalga indicate that key components of the savanna biota of northern Australia favour habitat that has remained unburnt for at least several years. This raises a serious conservation concern, given that very little relatively long unburnt habitat currently occurs in conservation reserves, with most sites being burnt at least once every 2 years. We propose a conservation objective of increasing the area that remains relatively long unburnt. This could be achieved either by reducing the proportion of the landscape burnt each year, or by setting prescribed fires more strategically. The provision of appropriately long unburnt habitat is a conservation challenge for Australia's tropical savanna landscapes, just as it is for its temperate forests.     

Structure and biomass along an Acacia zanzibarica woodland–savanna gradient in a former ranching area in coastal Tanzania

Questions: What factors influence the density, size and growth form of trees in secondary Acacia zanzibarica woodlands on a former humid savanna rangeland? How does tree density relate to variation in tree foliage and spines, and woody and grass biomass? Location: Tropical coastal Tanzania (former Mkwaja Ranch, now in Saadani National Park). Methods: We surveyed 97 circular plots (4‐m radius) representing a gradient from open savanna to dense woodland. Within each plot, we measured all trees and estimated the biomass of spines. Foliage biomass of tree and grass layers was estimated on three occasions, twice during the wet season and once in the dry season. Soil samples were taken from each plot and analysed for texture and nutrient content. Interrelationships among various variables were investigated using linear multiple regression and mixed effects models. Results: Tree densities were highest on more nutrient‐rich, heavy soils. Spinescence was highest on trees in open savanna. Biomass of tree foliage in the wet season was best explained by numbers of ant nests and tree live‐wood ratio. Foliage biomass in the dry season was less than half that in the wet season and best predicted by grass biomass. Variables related to biomass of the grass layer were strongly influenced by fire; living grass biomass also decreased with increasing tree density. Conclusions: A. zanzibarica is a tree with a high water demand, and the association with heavy soils is probably due to greater availability of water on these sites. Establishment of A. zanzibarica woodlands significantly reduced grazing resources at Mkwaja Ranch. Under post‐ranching conditions, however, fires and soil conditions predominate. The woodlands may, therefore, represent a transient state of woody density in a still resilient humid savanna.     

The role of reproductive phenology, seedling emergence and establishment of perennial Salix gordejevii in active sand dune fields

BACKGROUND AND AIMS: The function of sexual reproduction of perennials in restoration of vegetation of active dune fields frequently has been underestimated. The objective of this study was to evaluate the role of sexual reproduction of the perennial Salix gordejevii in the revegetation of active dunes. METHODS: Seedling emergence and establishment of S. gordejevii were examined both in controlled experiments (germination at different burial depths with different watering regimes) and in field observations in three dune slacks. The reproductive phenology and soil seed bank of S. gordejevii, the dynamics of soil moisture, the groundwater table and the landform level of three dune slacks were monitored. KEY RESULTS: Seeds of S. gordejevii began maturation on 1 May, and seed dispersal lasted from 8 May to 20 May. Seeds on the soil surface germinated significantly faster than those buried in soil (P<0.05). Seedling emergence was negatively correlated with landform level. When most seedlings emerged, there was a significantly positive correlation between soil moisture and seedling emergence (P<0.01). Rainfall was negatively correlated with seedling emergence. Seedling establishment was significantly and positively correlated with seedling emergence (P<0.05), and 72.3 % of the emergent seedlings were established at the end of the growing season. These results indicated that (a) seeds matured and dispersed before the rainy season; (b) seeds germinated as soon as they contacted a moist surface and relied more on soil moisture than on rainfall; and (c) more seedlings emerged at lower sampling points in dune slacks. CONCLUSIONS: In natural conditions, restoration of active sand dune fields generally commences with revegetation of dune slacks where sexual reproduction of perennials contributes greatly to species encroachment and colonization and hence plays an important role in restoration of active dune fields. Furthermore, aeolian erosion in dune slacks, leading to good soil moisture, facilitates seed germination, seedling emergence and establishment of S. gordejevii.     

Arthropod responses to experimental fire regimes in an Australian tropical savannah: ordinal‐level analysis

Fire is widely used for conservation management in the savannah landscapes of northern Australia, yet there is considerable uncertainty over the ecological effects of different fire regimes. The responses of insects and other arthropods to fire are especially poorly known, despite their dominant roles in the functioning of savannah ecosystems. Fire often appears to have little long‐term effect on ordinal‐level abundance of arthropods in temperate woodlands and open forests of southern Australia, and this paper addresses the extent to which such ordinal‐level resilience also occurs in Australia’s tropical savannahs. The data are from a multidisciplinary, landscape‐scale fire experiment at Kapalga in Kakadu National Park. Arthropods were sampled in the two major savannah habitats (woodland and open forest) using pitfall traps and sweep nets, in 15–20 km² compartments subjected to one of three fire regimes, each with three replicates: ‘early’ (annual fires lit early in the dry season), ‘late’ (annual fires lit late in the dry season), and ‘unburnt’ (fires absent during the five‐year experimental period 1990–94). Floristic cover, richness and composition were also measured in each sampling plot, using point quadrats. There were substantial habitat differences in floristic composition, but fire had no measured effect on plant richness, overall composition, or cover of three of the four dominant species. Of the 11 ordinal arthropod taxa considered from pitfall traps, only four were significantly affected by fire according to repeated‐measures ANOVA . There was a marked reduction in ant abundance in the absence of fire, and declines in spiders, homopterans and silverfish under late fires. Similarly, the abundances of only four of the 10 ordinal taxa from sweep catches were affected by fire, with crickets and beetles declining in the absence of fire, and caterpillars declining under late fires. Therefore, most of the ordinal taxa from the ground and grass‐layer were unaffected by the fire treatments, despite the treatments representing the most extreme fire regimes possible in the region. This indicates that the considerable ordinal‐level resilience to fire of arthropod assemblages that has previously been demonstrated in temperate woodlands and open forests of southern Australia, also occurs in tropical savannah woodlands and open forests of northern Australia.