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.     

Long‐term fire exclusion and ant community structure in an Australian tropical savanna: congruence with vegetation succession

Aim To examine the extent to which succession from tropical savanna to rain forest in the long‐term absence of fire is matched by successional changes in ant communities. This is done by describing ant community responses to 23 years of fire exclusion in a northern Australian tropical savanna, with a particular focus on the extent of colonization by specialist rain forest taxa. Location Solar Village, near Darwin in Australia's Northern Territory. Methods Ants were sampled within 12 plots located inside (‘unburnt’– protected from fire for 23 years) and outside (burnt every 1–2 years) Solar Village in ridge and slope habitat dominated by Eucalyptus spp. The litter, ground‐foraging and arboreal faunas were sampled separately, using Berlese funnels, unbaited pitfall traps and baited pitfall traps attached to tree trunks, respectively. Each species was assigned a forest‐association score ranging from 0 (open savanna species) to 3 (specialist forest species) based on their known habitat preferences in the region. Results A total of 85 ant species from 35 genera were recorded, with multivariate analysis demonstrating distinct litter, ground and arboreal communities. Ant communities also varied substantially with topographic position, which interacted strongly with fire exclusion. A total of 72 species were recorded in burnt habitat, compared with only 45 in unburnt, and the number of ant species records was also about twice as high in burnt compared with unburnt habitat. Fire exclusion has resulted in a dramatic increase in forest‐associated taxa (those occurring in forest and denser, but rarely open, savanna), with such species representing 51% of species records in unburnt habitat compared with 19% in burnt. However, only five specialist forest species were recorded, representing < 1% of total ant records. Main conclusions Fire exclusion at Solar Village has markedly increased the prevalence of forest‐associated ant species, but has led to only very minor incursions by specialist rain forest ant taxa. These responses match very closely those of the vegetation.     

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.     

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.     

Anomalies in grasstree fire history reconstructions for south‐western Australian vegetation

Abstract A new fire history for south‐western Australian sclerophyll forests was proposed recently based on grasstree (Xanthorrhoea preissii ) records that were interpreted to show a high frequency (3–5 years) ‘pre‐European burning regime’. Such a fire regime appears incompatible with the long‐term survival of many fire‐killed woody taxa. We investigated the local fire history in a small area of the northern sand‐plain shrub‐lands of south‐western Australia using 15 grasstrees, examining individual grasstree records in detail and comparing this with the decadal or averaged approach used in the original research, and with fire histories reconstructed from satellite images for the period since 1975. Results lead us to question the utility of the proposed grasstree fire history record as a tool for understanding past fire regimes for two reasons: First, inconsistencies in fire histories among individual grasstrees were considerable – some individuals were not burnt by known fires, while some apparently were burned many times during periods when others were not burned at all. Second, the grasstree record indicates a possible increase in patchiness of fires since 1930, while contemporary evidence and interpretations of the nature of Aboriginal (pre‐European) fire regimes would suggest the opposite. We believe that further research is needed to identify to what extent the grasstree method for reconstruction of fire histories can be used to re‐interpret how fire operated in many highly diverse ecosystems prior to European settlement of Australia.     

Structure and environmental relationships of insectivorous bat assemblages in tropical Australian savannas

Abstract Patterns in the composition of assemblages of microbat species sampled during the late dry season (the ‘build‐up’) in north Australian savannas were assessed against a range of environmental factors as well as four a priori defined habitat types (riparian, escarpments, coastal and woodlands). Distinct species assemblages were most strongly associated with topographic and climatic variables. There were also limited associations with vegetation structure, fire and local roost potential but no associations with insects or water availability. Total species diversity at sample sites was associated with distance to rivers and rainfall. In general, species assemblages were not clearly defined and the number of significant environmental associations was relatively few. We compare these associations with those reported for bat assemblages elsewhere in Australia.     

Natural 15N abundance of vegetation and soil in the Kapalga savanna,Australia

Abstract The natural abundance of the stable isotope ¹⁵N was measured in different vegetation components and in the soil of a northern Australian savanna. Most of the vegetation was found to be ¹⁵N-depleted compared to atmospheric N₂. Herbaceous legumes, perennial grasses, tree legumes, non-legume trees and annual grasses exhibited mean δ¹⁵N of ? 1.7, ? 0.8, ? 0.7, 0.0 and + 0.3‰, respectively. These results are in good agreement with previous studies. Legumes exhibit slightly negative values, indicating that they are likely to be nitrogen-fixing plants. Non-legume plants have a δ¹⁵N close to zero, which could equally result from non-symbiotic fixation, soil organic matter mineralization, or fresh root litter mineralization. In contrast, soil organic matter was ¹⁵N-enriched. Values of δ¹⁵N increased with depth and were + 2.5, + 5.2 and +6.1‰ in the 0–10, 10–20 and 20–40cm layers, respectively. Soil organic matter δ¹⁵N shows a typical profile of mature soils.     

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

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

Assessing effects of flow alteration on macroinvertebrate assemblages in Australian dryland rivers

1. Possible impacts of water‐resource development on assemblages of freshwater macroinvertebrates were investigated in the upper Darling River and some of its tributaries in north‐western New South Wales (Australia), an arid and semi‐arid region of low relief where alteration of river flows has intensified through expansion of irrigated agriculture. 2. Study sites were grouped into four hydrological regimes resulting from impoundment, flow regulation, water abstraction and natural variation, namely (i) intermittent flow with relatively little hydrological alteration from water‐resource development, (ii) intermittent flow with substantial alteration, (iii) near‐perennial flow with substantial alteration but unimpounded and (iv) near‐perennial flow with substantial alteration plus impoundment by weirs that stabilise water levels. 3. Macroinvertebrates were sampled with three methods (a quantitative cylinder sampler, handnet sampling and baited traps) in three periods with differing hydrology (recessional low flow in June 2003, high flow in March 2004 and increasing flow after drought in December 2004). 4. Taxonomic richness, assemblage composition and catch per unit effort of the crayfish Cherax destructor differed significantly among the site groups, but total macroinvertebrate density and the AUSRIVAS O/E (Australian River Assessment System observed‐over‐expected) index did not. The principal spatial differences were between the intermittent and near‐perennial rivers, and apparent effects of water‐resource development and impoundment were more subtle. Temporal differences in richness, abundance and composition were substantial and appeared to be related mainly to variations in discharge and temperature. 5. Current macroinvertebrate‐based methods for assessing the ‘condition’ or ‘health’ of Australian dryland rivers are inadequate. Such assessments might be improved with (i) reference data that take adequate account of antecedent hydrological conditions, (ii) consideration of long‐term taxonomic richness as well as richness on individual sampling occasions, (iii) evaluation of invertebrate population sizes, (iv) analysis of assemblage data by trait composition and (v) adoption of the genus as the default level of taxonomic resolution.     

Distribution and abundance of large herbivores in a northern Australian tropical savanna: A multi‐scale approach

Australian mammals have exhibited exceptionally high rates of decline since European settlement 230 years ago with much focus on small mammals in northern tropical savannas. In these systems, little scientific attention has been given to the suite of grazing macropods, family Macropodidae, (common wallaroo (Osphranter robustus), antilopine wallaroo (O. antilopinus) and agile wallaby (Notamacropus agilis)). These species may be impacted by feral herbivores and contemporary fire regimes, two threats linked to small mammal declines. A multi‐scale approach using aerial surveys, road surveys and camera trapping was utilised to determine the effects of feral cattle and fire on the distribution and abundance of large macropods in the North Kimberley bioregion. Feral cattle density and biomass exceeded that of macropods regardless of survey technique. Density estimates for cattle were up to 125 times higher (0.3–10.0 km^?2) than estimates for macropods (0.08–0.49 km^?2). Cattle biomass, based on the aerial survey estimates (corrected for perception bias), were 15 and 95 times higher than macropods for infertile (279 vs. 19 kg km^?2) and fertile savannas (518 vs. 5 kg km^?2), respectively. Proximity to the nearest pastoral station was a significant predictor of the aerial sightings of feral cattle (P ≤ 0.05). Abundance and foraging activity of cattle were positively associated (P ≤ 0.05) with recently burnt areas. In contrast, camera trapping showed agile wallaby and wallaroo occurrence and foraging were associated with longer unburnt areas (P ≤ 0.05). Agile wallaby and wallaroo were negatively associated with cattle (P ≤ 0.05) and showed substantial diurnal and seasonal separation consistent with an antagonistic interspecific interaction. Results also suggest that the agile wallaby is the primary prey of the dingo, not wallaroo. Collectively, this study suggests that recent landscape changes such as altered fire regimes and introduced herbivores have negatively impacted large grazing macropod species.     

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.     

A grass–fire cycle eliminates an obligate‐seeding tree in a tropical savanna

A grass–fire cycle in Australian tropical savannas has been postulated as driving the regional decline of the obligate-seeding conifer Callitris intratropica and other fire-sensitive components of the regional flora and fauna, due to proliferation of flammable native grasses. We tested the hypothesis that a high-biomass invasive savanna grass drives a positive feedback process where intense fires destroy fire-sensitive trees, and the reduction in canopy cover facilitates further invasion by grass. We undertook an observational and experimental study using, as a model system, a plantation of C. intratropica that has been invaded by an African grass, gamba (Andropogon gayanus) in the Northern Territory, Australia. We found that high grass biomass was associated with reduced canopy cover and restriction of foliage to the upper canopy of surviving stems, and mortality of adult trees was very high (>50%) even in areas with low fuel loads (1 t·ha⁻¹). Experimental fires, with fuel loads >10 t·ha⁻¹, typical of the grass-invasion front, caused significant mortality due to complete crown scorch. Lower fuel loads cause reduced canopy cover through defoliation of the lower canopy. These results help explain how increases in grass biomass are coupled with the decline of C. intratropica throughout northern Australia by causing a switch from litter and sparse perennial grass fuels, and hence low-intensity surface fires, to heavy annual grass fuel loads that sustain fires that burn into the midstorey. This study demonstrates that changes in fuel type can alter fire regimes with substantial knock-on effects on the biota.     

Weather effects on birds of different size are mediated by long‐term climate and vegetation type in endangered temperate woodlands

Species occurrence is influenced by a range of factors including habitat attributes, climate, weather, and human landscape modification. These drivers are likely to interact, but their effects are frequently quantified independently. Here, we report the results of a 13‐year study of temperate woodland birds in south‐eastern Australia to quantify how different‐sized birds respond to the interacting effects of: (a) short‐term weather (rainfall and temperature in the 12 months preceding our surveys), (b) long‐term climate (average rainfall and maximum and minimum temperatures over the period 1970–2014), and (c) broad structural forms of vegetation (old‐growth woodland, regrowth woodland, and restoration plantings). We uncovered significant interactions between bird body size, vegetation type, climate, and weather. High short‐term rainfall was associated with decreased occurrence of large birds in old‐growth and regrowth woodland, but not in restoration plantings. Conversely, small bird occurrence peaked in wet years, but this effect was most pronounced in locations with a history of high rainfall, and was actually reversed (peak occurrence in dry years) in restoration plantings in dry climates. The occurrence of small birds was depressed—and large birds elevated—in hot years, except in restoration plantings which supported few large birds under these circumstances. Our investigation suggests that different mechanisms may underpin contrasting responses of small and large birds to the interacting effects of climate, weather, and vegetation type. A diversity of vegetation cover is needed across a landscape to promote the occurrence of different‐sized bird species in agriculture‐dominated landscapes, particularly under variable weather conditions. Climate change is predicted to lead to widespread drying of our study region, and restoration plantings—especially currently climatically wet areas—may become critically important for conserving bird species, particularly small‐bodied taxa.