Frequent fuel‐reduction burning: the role of logs and associated leaf litter in the conservation of ant biodiversity

Frequent low‐intensity fires are used in management of Australian forests to reduce fuel loads and protect natural resources and human property. Low‐intensity fires are typically patchy and unburned litter microhabitats are often associated with large objects such as logs, which may act as refuges both for vertebrate and for invertebrate fauna. The aim of this study was to determine whether ants were using unburned leaf litter microhabitats associated with logs as a refuge after fire. The study was carried out in Bulls Ground State Forest, New South Wales, Australia, where experimentally burned and unburned sites had previously been established. Species richness and abundance of ants in leaf litter did not differ between habitats adjacent to logs and away from logs, in burned and unburned sites. Fifteen of the 42 ant species were found in all four habitats, and contributed 94% of total ant abundance. Every habitat had a group of unique species, which together made up 30% of the total species richness. There was also a distinct group of species that was not found in the leaf litter associated with the burned/open habitat. However, as 45% of all species were found in low abundance (less than 10 individuals), care must be taken in inferring patterns for these groups. When functional groups were used to assess community structure, ‘cryptic’ species were found to be common in all habitats, whereas ‘subordinate Camponotini’ were found in burned habitats only. This study indicates that in an area where frequent burning is applied on a broad scale, preserving a range of microhabitats, including those associated with retained logs, may make a substantial contribution to conserving ant biodiversity.     

Comparison of Post‐mining Rehabilitation with Reference Ecosystems in Monsoonal Eucalypt Woodlands,Northern Australia

Rehabilitation of post‐mining lands frequently aims to create “self‐sustaining” systems. Where native vegetation is the designated post‐mining land use, it is generally assumed that rehabilitation that is similar to local native ecosystems is more likely to be sustainable. I compared landscape functionality, plant community composition, and vegetation structure in (1) reference sites representing pre‐mining native forest; (2) reference sites representing potential landscape analogues for the post‐mining landscape; and (3) a 23‐year chronosequence of post‐mining rehabilitation on the Weipa bauxite plateau, Cape York Peninsula, Australia. The trends across the post‐mining chronosequence indicate that vegetation growth is rapid in the first 5–8 years, and then slows with mean height approaching an asymptote after approximately 15 years. Landscape function indices showed a response that coincided with vegetation growth. Vegetation composition was significantly different from reference native forest. Most importantly, from the perspective of creating self‐sustaining ecosystems, the contribution of local framework species to vegetation in rehabilitation was significantly lower than in reference native forest. I discuss the results in relation to theoretical models of succession and conclude that without management intervention, differences between post‐mining rehabilitation and native forest are likely to be persistent.     

The Fire Refuge Value of Patches of a Fire‐Sensitive Tree in Fire‐prone Savannas: Callitris intratropica in Northern Australia

Patches of fire‐sensitive vegetation often occur within fire‐prone tropical savannas, and are indicative of localized areas where fire regimes are less severe. These may act as important fire refugia for fire‐sensitive biota. The fire‐sensitive tree Callitris intratropica occurs in small patches throughout the fire‐prone northern Australian savannas, and is widely seen as an indicator of low‐severity fire regimes and of good ecosystem health. Here, we address the question: to what extent do Callitris patches act as refuges for other fire‐sensitive biota, and therefore play a broader conservation role? We contrast floral and faunal species composition between Callitris patches and surrounding eucalypt savanna, using three case studies. In the first case study, a floristic analysis of 47 Callitris patches across Western Australia's Kimberley region showed that woody species in these patches were overwhelmingly widespread, fire‐tolerant savanna taxa. No species of special conservation concern occurred disproportionately within Callitris patches. Similarly, there was no concentration of fire‐sensitive fauna or flora in five Callitris patches in the East Kimberley. Finally, there was no difference in ant species composition among 12 Callitris patches and surrounding eucalypt savannas in Kakadu National Park, Northern Territory, and there were no fire‐sensitive ant species in Callitris patches. Our three case studies from throughout the northwestern Australia provide no evidence that Callitris patches act as important refuges for fire‐sensitive flora or fauna within fire‐prone eucalypt savannas. This calls into question the notion that Callitris is a strong indicator of general ecosystem health.     

Soil functional responses to ecological restoration treatments in frequent‐fire forests of the western United States: a systematic review

We investigated general effects of ecological restoration treatments on soil function in frequent‐fire forests of the western United States using a systematic review methodology. We searched numerous publication databases for original research papers and used well‐defined criteria developed a priori to select papers for review. We used meta‐analysis and qualitative summaries to compare reported responses of macronutrients, nitrogen cycling, and soil respiration among tree thinning (thin), prescribed fire (burn), and thinning plus prescribed fire treatments (composite). Results of meta‐analysis showed that mean differences in macronutrients were consistently higher in composite treatments (standardized using controls) when compared to thin‐only and burn‐only treatments. Mean responses related to nitrogen cycling showed similar patterns, with significant increases detected in composite treatments for all nitrogen cycling variables (mineralization, ammonification, and nitrification) and insignificant responses for the majority of the burn‐only and thin‐only treatments. Mean difference in response for soil respiration following composite treatments showed increases as compared to the controls, and no significant differences were detected in response to burn‐ and thin‐only treatments. While soil function, nutrient cycling, and soil respiration differed among treatments, the most significant effects were observed for nitrogen and carbon responses, net mineralization and nitrification, ammonium availability, and soil respiration rate, which experienced the greatest increase following treatments that were both thinned and burned.     

Inter‐ and Intra‐Regional Variability of Leaf Litter Breakdown in Reference Headwater Streams of Northern Spain: Atlantic versus Mediterranean Streams

The main goal of this study was to examine the natural variability of alder (Alnus glutinosa (L.) Gaertn.) leaf processing and explore its potentiality as a functional indicator to assess the ecological status of Spanish headwater streams. Breakdown of leaf litter was studied during autumn‐winter in reference headwater streams of two regions of northern Spain: the Basque Country (on the Atlantic) and Catalonia (on the Mediterranean). Spring experiments were also carried out in the Atlantic region in order to study seasonal changes. Leaf mass loss rates were slightly higher in Catalonian streams. Temperature was not the main factor for explaining differences between the two regions. In the Atlantic one, however, water temperature determined the spatial and seasonal variability of leaf litter processing. Because of the high natural variability in breakdown rates, our results highlight the difficulty in detecting moderate impairments on stream functioning through the analysis of leaf litter decomposition if this process is not accompanied by the study of other factors affecting it. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Negative plant‐phyllosphere feedbacks in native Asteraceae hosts – a novel extension of the plant‐soil feedback framework

Over the past 25 years, the plant‐soil feedback (PSF) framework has catalyzed our understanding of how belowground microbiota impact plant fitness and species coexistence. Here, we apply a novel extension of this framework to microbiota associated with aboveground tissues, termed ‘plant‐phyllosphere feedback (PPFs)’. In parallel greenhouse experiments, rhizosphere and phyllosphere microbiota of con‐ and heterospecific hosts from four species were independently manipulated. In a third experiment, we tested the combined effects of soil and phyllosphere feedback under field conditions. We found that three of four species experienced weak negative PSF whereas, in contrast, all four species experienced strong negative PPFs. Field‐based feedback estimates were highly negative for all four species, though variable in magnitude. Our results suggest that phyllosphere microbiota, like rhizosphere microbiota, can potentially mediate plant species coexistence via negative feedbacks. Extension of the PSF framework to the phyllosphere is needed to more fully elucidate plant‐microbiota interactions.     

Simple three‐pool model accurately describes patterns of long‐term litter decomposition in diverse climates

As atmospheric CO₂ increases, ecosystem carbon sequestration will largely depend on how global changes in climate will alter the balance between net primary production and decomposition. The response of primary production to climatic change has been examined using well‐validated mechanistic models, but the same is not true for decomposition, a primary source of atmospheric CO₂. We used the Long‐term Intersite Decomposition Experiment Team (LIDET) dataset and model‐selection techniques to choose and parameterize a model that describes global patterns of litter decomposition. Mass loss was best represented by a three‐pool negative exponential model, with a rapidly decomposing labile pool, an intermediate pool representing cellulose, and a recalcitrant pool. The initial litter lignin/nitrogen ratio defined the size of labile and intermediate pools. Lignin content determined the size of the recalcitrant pool. The decomposition rate of all pools was modified by climate, but the intermediate pool's decomposition rate was also controlled by relative amounts of litter cellulose and lignin (indicative of lignin‐encrusted cellulose). The effect of climate on decomposition was best represented by a composite variable that multiplied a water‐stress function by the Lloyd and Taylor variable Q₁₀ temperature function. Although our model explained nearly 70% of the variation in LIDET data, we observed systematic deviations from model predictions. Below‐ and aboveground material decomposed at notably different rates, depending on the decomposition stage. Decomposition in certain ecosystem‐specific environmental conditions was not well represented by our model; this included roots in very wet and cold soils, and aboveground litter in N‐rich and arid sites. Despite these limitations, our model may still be extremely useful for global modeling efforts, because it accurately (R²=0.6804) described general patterns of long‐term global decomposition for a wide array of litter types, using relatively minimal climatic and litter quality data.     

Patterns of Richness and Abundance in a Tropical African Leaf‐litter Herpetofauna1

I compared species richness and habitat correlates of leaf‐litter herpetofaunal abundance in undisturbed and selectively logged forests, and an abandoned pine plantation in Kibale National Park, Uganda. I sampled 50 randomly located 25 m² litter plots in each area during the wet and dry seasons in 1997. Ten anuran, five lizard, and three snake species were captured in plots over the study. Assemblage composition was most similar at logged and unlogged sites. The logged forest herpetofauna had higher species richness and abundance than the unlogged forest, but diversity was greater in the unlogged forest due to greater evenness. In contrast, the pine plantation site had the highest richness, abundance, and evenness of the three study sites, but species composition was distinct from the other areas. Herpetofaunal densities were significantly lower in all three areas during the dry season than in the wet season. During the dry season, soil moisture, litter mass, topography, shrub cover, and number of fallen logs were significant positive predictors of herpetofaunal presence in litter plots, but only soil moisture was significant in the wet season. The interaction of moisture and topography appears to be important in determining seasonal patterns of litter herpetofaunal distribution. Comparison of litter herpetofaunal studies across the tropics have shown that mid‐elevation faunas generally support fewer species than lowland faunas. Compared with other tropical mid‐elevation litter faunas, Kibale supports an intermediate number of species, but at lower densities than observed at any other mid‐elevation site reported in the literature.     

Effects of diversity loss on ecosystem function across trophic levels and ecosystems: A test in a detritus‐based tropical food web

Abstract We investigated the effects of biodiversity loss across trophic levels and across ecosystems (terrestrial to aquatic) on ecosystem function, in a detritus‐based tropical food web. Diversities of consumers (stream shredders) and resources (leaf litter) were experimentally manipulated by varying the number of species from 3 to 1, using different species combinations, and the effects on leaf breakdown rates were examined. In single‐species shredder treatments, leaf diversity loss affected breakdown rates, but the effect depended on the identity of the leaves remaining in the system: they increased when the most preferred leaf species remained, but decreased when this species was lost (leaf preferences were the same for all shredders). In multi‐species shredder assemblages, breakdown rates were lower than expected from single‐species treatments, suggesting an important role of interspecific competition. This pattern was also evident when oneleaf species was available but not with higher leaf diversity, suggesting that lowered leaf diversity promotes competitive interactions among shredders. The influence of diversity and identity of species across trophic levels and ecosystems on stream functioning points to complex interactions that may well be reflected in other types of ecosystem.     

Ants on the Move: Resource Limitation of a Litter‐nesting Ant Community in Costa Rica1

The leaf litter of tropical wet forests is replete with itinerant ant nests. Nest movement may help ants evade the constraints of stress and disturbance and increase access to resources. I studied how nest relocation and environmental factors may explain the density, size, and growth of leaf litter ant nests. I decoupled the relationships among litter depth, food abundance, and nest availability in a 4‐mo manipulation of food and leaf litter in a community of litter‐nesting ants in a lowland wet forest in Costa Rica. Over 4 mo, 290 1 m² treatment and control plots were sampled without replacement. Nest densities doubled in response to food supplementation, but did not decrease in response to litter removal or stress (from litter trampling). The supplementation of food increased the utilization of less favored nesting materials. In response to food supplementation and litter trampling, arboreal ants established nests in the litter, and growth rates of the most common ants (Pheidole spp.) increased. Colony growth was independent of colony size and growth rates of the most abundant ants. In general, I conclude that litter‐nesting ant density is driven primarily by food limitation, that nest relocation behavior significantly affects access to resource and the demographic structure of this community, and that nest fission may be a method to break the growth–reproduction trade‐off.     

Dormant Season Prescribed Fire as a Management Tool for the Control of <Emphasis Type="Italic">Salix caroliniana</Emphasis> Michx. in a Floodplain Marsh

Expansion of woody species into herbaceous wetlands is a serious concern in wetland management. Prescribed fire is often used as a tool to manage woody species, although many species resprout after fire making control problematic. In this study, we assessed the usefulness of repeated dormant season fires for controlling Salix caroliniana (Michx.) in a floodplain marsh in Florida. Salix is a common shrub in southeastern marshes that resprouts prolifically after fire. We compared stem basal area, stem density, and cover of Salix in three adjacent sites in a floodplain marsh in east central Florida. One site was burned once in February 1997, another site was burned in February 1997 and then again in March 1999 and one site was left unburned. At the unburned site, Salix stem basal area, stem density, and cover increased over the course of the study. In the two burned sites, the first fire destroyed large diameter stems and stimulated production of sprouts. As a result, stem basal area and cover decreased but stem density remained unchanged. The second fire caused a decline in stem density and a further decline in cover. Changes in understory species composition and cover could not be attributed to the fires. Our results suggest that dormant season fires are effective in reducing Salix cover and basal area, and that repeated fires have greater effects than a single fire.     

A Comparative Study of Leaf Breakdown of Three Native Tree Species in a Slowly‐Flowing Headwater Stream in the Colombian Andes

The dynamics of leaf breakdown in a headwater Colombian stream were evaluated for the native tree species Myrsine guianensis, Cupania latifolia and Nectandra lineatifolia using coarse and fine mesh litter bags. Ten bags of each species (five of each mesh size) were retrieved from the stream at 1, 8, 15, 30, 60 and 120 days. k values ranged from 0.0008 to 0.0058 day^–1 and density of macroinvertebrates from 35 to 55 individuals per leaf bag, peaking at day 8. Myrsine guianensis degraded more rapidly than the other species for both coarse and fine mesh bags. This species and Nectandra lineatifolia presented differences in k values between coarse and fine mesh bags, suggesting that macroinvertebrates influenced the decay rate. Despite the low densities of macroinvertebrates found, shredders represented 12.7% of individuals and 50 to 68% of the invertebrate biomass in bags, indicating that this functional feeding group was an important component of fauna associated with litter breakdown in this first order tropical stream. (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Linking Geographical Information Systems and Observational and Experimental Studies to Determine Optimal Seedling Microsites of an Endangered Plant in a Subtropical Urban Fire‐Adapted Ecosystem

Understanding microsite requirements is critical for restoring sustainable rare plant populations and creating meaningful management plans that will enhance native species’ population viability. Natural areas bordered by urban communities have restricted size and constrained management options that confound goals of preserving biodiversity. Using geographical information systems and observational and experimental studies, we determined that current microsites required for seedling establishment within existing habitat of the federally endangered Crenulate leadplant are few and spatially restricted, resulting in little wild seedling recruitment. Experimental and wild seedlings had highest germination rates in litter layers 1–2 cm in depth and survived the longest in litter 0.5–2 cm deep. Fire suppression and aggressive native and exotic plants have accumulated litter up to 33 cm near some of the wild adult Crenulate leadplant. Only 9% of habitat occupied by this taxon at its largest population is conducive to seedling establishment. Similarly, 30% is conducive to seedling establishment in the second largest population. Fire plays a critical role in South Florida ecosystem dynamics and the restoration and preservation of its landscape and biodiversity. When fire is not possible, other management is needed. Manual litter and downed debris removal are recommended at both population sites to improve the probability of Crenulate leadplant seedling establishment and population persistence.     

Variations of leaf longevity in tropical moist forests predicted by a trait‐driven carbon optimality model

Leaf longevity (LL) varies more than 20‐fold in tropical evergreen forests, but it remains unclear how to capture these variations using predictive models. Current theories of LL that are based on carbon optimisation principles are challenging to quantitatively assess because of uncertainty across species in the ‘ageing rate:’ the rate at which leaf photosynthetic capacity declines with age. Here, we present a meta‐analysis of 49 species across temperate and tropical biomes, demonstrating that the ageing rate of photosynthetic capacity is positively correlated with the mass‐based carboxylation rate of mature leaves. We assess an improved trait‐driven carbon optimality model with in situLL data for 105 species in two Panamanian forests. We show that our model explains over 40% of the cross‐species variation in LL under contrasting light environment. Collectively, our results reveal how variation in LL emerges from carbon optimisation constrained by both leaf structural traits and abiotic environment.     

Trait‐abundance relation in response to nutrient addition in a Tibetan alpine meadow: The importance of species trade‐off in resource conservation and acquisition

In competition‐dominated communities, traits promoting resource conservation and competitive ability are expected to have an important influence on species relative abundance (SRA). Yet, few studies have tested the trait‐abundance relations in the line of species trade‐off in resource conservation versus acquisition, indicating by multiple traits coordination. We measured SRA and key functional traits involving leaf economic spectrum (SLA, specific leaf area; LDMC, leaf dry matter content; LCC, leaf carbon concentration; LNC, leaf nitrogen concentration; LPC, leaf phosphorus concentration; Hs, mature height) for ten common species in all plots subjected to addition of nitrogen fertilizer (N), phosphorus fertilizer (P), or both of them (NP) in a Tibetan alpine meadow. We test whether SRA is positively related with traits promoting plant resource conservation, while negatively correlated with traits promoting plant growth and resource acquisition. We found that species were primarily differentiated along a trade‐off axis involving traits promoting nutrient acquisition and fast growth (e.g., LPC and SLA) versus traits promoting resource conservation and competition ability (e.g., large LDMC). We further found that SRA was positively correlated with plant height, LDMC, and LCC, but negatively associated with SLA and leaf nutrient concentration irrespective of fertilization. A stronger positive height‐SRA was found in NP‐fertilized plots than in other plots, while negative correlations between SRA and SLA and LPC were found in N or P fertilized plots. The results indicate that species trade‐off in nutrient acquisition and resource conservation was a key driver of SRA in competition‐dominated communities following fertilization, with the linkage between SRA and traits depending on plant competition for specific soil nutrient and/or light availability. The results highlight the importance of competitive exclusion in plant community assembly following fertilization and suggest that abundant species in local communities become dominated at expense of growth while infrequent species hold an advantage in fast growth and dispersals to neighbor meta‐communities.     

Post‐burning germination responses of woody invaders in a fire‐prone ecosystem

It has been frequently recognised that there is a positive feedback between plant invasion and fire underlying invasion success in fire‐prone ecosystems. Accordingly, the response of woody alien species germination to fire may have direct implications on their invasiveness in those ecosystems, particularly when fruit ripening occurs in the fire season. Here, we experimentally evaluated the germination response of some of the main woody invaders of the Chaco Serrano dry woodlands (Gleditsia triacanthos, Cotoneaster glaucophyllus, Ligustrum lucidum, Pyracantha angustifolia and Melia azedarach), which fruit in the seasons of highest fire frequency. Seeds were subjected to heat‐shock treatments that simulated a range of heat intensities, and the species were classified according to their germination response as heat sensitive, tolerant or stimulated. Since Gleditsia triacanthos has indehiscent fruits that fall from the plant and might be exposed to flames, its germination response was also assessed of seeds exposed to fruit burning. Germination responses to heat varied among the invasive species. G. triacanthos seeds experienced increased germination under very low and low heat indexes; it was therefore classified as heat stimulated. The other four species showed no change in germination under very low heat indexes and were therefore considered heat tolerant. However, all species were sensitive to high heat as indicated by their significant decline in germination. G. triacanthos would have limited capacity to recruit from seeds following flaming combustion of its fruits. The prevalence of heat‐tolerant rather than heat‐stimulated germination responses suggests that the occurrence of frequent and seasonal fires in this subtropical savanna system might delay rather than boost the expansion of these invasive species in the system. Yet, the presence of heat‐stimulated germination in one of the studied species warns against generalisation, even within the same ecosystem, and further supports the idiosyncratic nature of invasion success. Abstract in Spanish is available with online material.     

Leaf Litter as a Source of Dissolved Organic Carbon in Streams

Dissolved organic carbon (DOC) is an abundant form of organic matter in stream ecosystems. Most research has focused on the watershed as the source of DOC in streams, but DOC also comes from leaching of organic matter stored in the stream channel. We used a whole-ecosystem experimental approach to assess the significance of leaching of organic matter in the channel as a source of DOC in a headwater stream. Inputs of leaf litter were excluded from a forested Appalachian headwater stream for 3 years. Stream-water concentration, export, and instream generation of DOC were reduced in the litter-excluded stream as compared with a nearby untreated reference stream. The proportion of high molecular weight (HMW) DOC (more than 10,000 daltons) in stream water was not altered by litter exclusion. Mean DOC concentration in stream water was directly related to benthic leaf-litter standing stock. Instream generation of DOC from leaf litter stored in the stream channel contributes approximately 30% of daily DOC exports in this forested headwater stream. This source of DOC is greatest during autumn and winter and least during spring and summer. It is higher during increasing discharge than during base flow. We conclude that elimination of litter inputs from a forested headwater stream has altered the biogeochemistry of DOC in this ecosystem. Received 2 September 1997; accepted 27 January 1998.