Temporal dynamics in the composition of bird communities along a gradient of farmland restoration.

Revegetation plantings are a key activity in farmland restoration and are commonly assumed to support biotic communities that, with time, replicate those of reference habitats. Restoration outcomes, however, can be highly variable and difficult to predict; hence there is value in quantifying restoration success to improve future efforts. We test the expectation that, over time, revegetation will restore bird communities to match those in reference habitats; and assess whether specific planting attributes enhance restoration success. We surveyed birds in 255 sites in south-east Australia, arranged along a restoration gradient encompassing three habitat types: unrestored farmland (paddocks), revegetation plantings (comprising a chronosequence up to 52 years old) and reference habitats (remnant native vegetation). Surveys were undertaken in 2006/2007 and again in 2019, with data used to compare bird assemblages between habitat types. We also determined whether, in the intervening 12 years, bird communities in revegetation had shifted toward reference habitats on the restoration gradient. Our results showed that each habitat contained a unique bird community and that, over time, assemblages in revegetation diverged away from those in unrestored farmland and converged toward those in reference habitats. Two planting attributes influenced this transition: the bird assemblages of revegetation were more likely to have diverged away from those of unrestored farmland (with scattered mature trees) 12 years later if they were located in areas with more surrounding tree cover, and were mostly ungrazed by livestock (compared with grazed plantings). Our results highlight three key ways in which revegetation contributes to farmland restoration: (1) by supporting richer and more diverse bird assemblages than unrestored farmland, (2) by enhancing beta diversity in rural landscapes through the addition of a unique bird community, and (3) by shifting bird assemblages toward those found in reference habitats over time. However, revegetation plantings did not replicate reference habitats by ~40-50 years in our region, and complete convergence may take centuries. These findings have implications for environmental offset programs and mean that effective conservation in farmland environments depends on the retention and protection of natural and seminatural habitats as a parallel management strategy to complement restoration.

Differential effects of ecosystem engineering by the superb lyrebird Menura novaehollandiae and herbivory by large mammals on floristic regeneration and structure in wet eucalypt forests.

Ecosystem engineers that modify the soil and ground-layer properties exert a strong influence on vegetation communities in ecosystems worldwide. Understanding the interactions between animal engineers and vegetation is challenging when in the presence of large herbivores, as many vegetation communities are simultaneously affected by both engineering and herbivory. The superb lyrebird Menura novaehollandiae, an ecosystem engineer in wet forests of south-eastern Australia, extensively modifies litter and soil on the forest floor. The aim of this study was to disentangle the impacts of engineering by lyrebirds and herbivory by large mammals on the composition and structure of ground-layer vegetation. We carried out a 2-year, manipulative exclusion experiment in the Central Highlands of Victoria, Australia. We compared three treatments: fenced plots with simulated lyrebird foraging; fenced plots excluding herbivores and lyrebirds; and open controls. This design allowed assessment of the relative impacts of engineering and herbivory on germination rates, seedling density, vegetation cover and structure, and community composition. Engineering by lyrebirds enhanced the germination of seeds in the litter layer. After 2 years, more than double the number of germinants were present in "engineered" than "non-engineered" plots. Engineering did not affect the density of seedlings, but herbivory had strong detrimental effects. Herbivory also reduced the floristic richness and structural complexity (<0.5 m) of forest vegetation, including the cover of herbs. Neither process altered the floristic composition of the vegetation within the 2-year study period. Ecosystem engineering by lyrebirds and herbivory by large mammals both influence the structure of forest-floor vegetation. The twofold increase in seeds stimulated to germinate by engineering may contribute to the evolutionary adaptation of plants by allowing greater phenotypic expression and selection than would otherwise occur. Over long timescales, engineering and herbivory likely combine to maintain a more-open forest floor conducive to ongoing ecosystem engineering by lyrebirds.

The functional roles of psyllid abundance and assemblage on bird-associated forest defoliation.

Fluctuations in the abundance of insect herbivores and the severity of damage they cause to their hosts are key factors affecting tree health and trophic interactions. Bell miner associated dieback (BMAD) is a trophic cascade involving eucalypt trees, psyllids and bell miners. Bell miners are a territorial species of bird that reduce insectivorous bird diversity, thereby increasing psyllid abundances. The role of psyllids in BMAD is debated, primarily because psyllid host specificity, genus, defoliation potential and/or lerp composition have been conflated or unexplored. This study documented psyllid communities and canopy structure of four eucalypt hosts (Eucalyptus bridgesiana, E. caliginosa, E. propinqua and E. siderophloia) in BMAD and non-BMAD-affected forests in northern New South Wales. Psyllid abundances were significantly higher in BMAD-affected forests. However, psyllid assemblages did not differ between forest types which contrasts with an influential hypothesis concerning bell miner farming of psyllids that produce sugary lerps (Glycaspis species). Importantly, psyllid communities differed among species of eucalypt host. Hosts supporting higher abundances of Cardiaspina tended to have a stronger negative relationship between canopy health and psyllid abundances than other host trees. Cardiaspina were also found on older leaves and associated with more leaf damage than other psyllids. Unlike most other psyllid genera, Cardiaspina initiate premature foliar senescence leading to defoliation which ultimately changes the age structure of leaves of the canopies of affected trees. Our study supports the functional linkage between psyllid assemblages, their relative abundances and defoliation potential to bottom-up mechanisms behind a bird-associated trophic cascade.

How fire interacts with habitat loss and fragmentation.

Biodiversity faces many threats and these can interact to produce outcomes that may not be predicted by considering their effects in isolation. Habitat loss and fragmentation (hereafter 'fragmentation') and altered fire regimes are important threats to biodiversity, but their interactions have not been systematically evaluated across the globe. In this comprehensive synthesis, including 162 papers which provided 274 cases, we offer a framework for understanding how fire interacts with fragmentation. Fire and fragmentation interact in three main ways: (i) fire influences fragmentation (59% of 274 cases), where fire either destroys and fragments habitat or creates and connects habitat; (ii) fragmentation influences fire (25% of cases) where, after habitat is reduced in area and fragmented, fire in the landscape is subsequently altered because people suppress or ignite fires, or there is increased edge flammability or increased obstruction to fire spread; and (iii) where the two do not influence each other, but fire interacts with fragmentation to affect responses like species richness, abundance and extinction risk (16% of cases). Where fire and fragmentation do influence each other, feedback loops are possible that can lead to ecosystem conversion (e.g. forest to grassland). This is a well-documented threat in the tropics but with potential also to be important elsewhere. Fire interacts with fragmentation through scale-specific mechanisms: fire creates edges and drives edge effects; fire alters patch quality; and fire alters landscape-scale connectivity. We found only 12 cases in which studies reported the four essential strata for testing a full interaction, which were fragmented and unfragmented landscapes that both span contrasting fire histories, such as recently burnt and long unburnt vegetation. Simulation and empirical studies show that fire and fragmentation can interact synergistically, multiplicatively, antagonistically or additively. These cases highlight a key reason why understanding interactions is so important: when fire and fragmentation act together they can cause local extinctions, even when their separate effects are neutral. Whether fire-fragmentation interactions benefit or disadvantage species is often determined by the species' preferred successional stage. Adding fire to landscapes generally benefits early-successional plant and animal species, whereas it is detrimental to late-successional species. However, when fire interacts with fragmentation, the direction of effect of fire on a species could be reversed from the effect expected by successional preferences. Adding fire to fragmented landscapes can be detrimental for species that would normally co-exist with fire, because species may no longer be able to disperse to their preferred successional stage. Further, animals may be attracted to particular successional stages leading to unexpected responses to fragmentation, such as higher abundance in more isolated unburnt patches. Growing human populations and increasing resource consumption suggest that fragmentation trends will worsen over coming years. Combined with increasing alteration of fire regimes due to climate change and human-caused ignitions, interactions of fire with fragmentation are likely to become more common. Our new framework paves the way for developing a better understanding of how fire interacts with fragmentation, and for conserving biodiversity in the face of these emerging challenges.

Foraging by an avian ecosystem engineer extensively modifies the litter and soil layer in forest ecosystems.

Ecosystem engineers physically modify their environment, thereby altering habitats for other organisms. Increasingly, "engineers" are recognized as an important focus for conservation and ecological restoration because their actions affect a range of ecosystem processes and thereby influence how ecosystems function. The Superb Lyrebird Menura novaehollandiae is proposed as an ecosystem engineer in forests of southeastern Australia due to the volume of soil and litter it turns over when foraging. We measured the seasonal and spatial patterns of foraging by Lyrebirds and the amount of soil displaced in forests in the Central Highlands, Victoria. We tested the effects of foraging on litter, soil nutrients and soil physical properties by using an experimental approach with three treatments: Lyrebird exclusion, Lyrebird exclusion with simulated foraging, and non-exclusion reference plots. Treatments were replicated in three forest types in each of three forest blocks. Lyrebirds foraged extensively in all forest types in all seasons. On average, Lyrebirds displaced 155.7 Mg/ha of litter and soil in a 12-month period. Greater displacement occurred where vegetation complexity (<50 cm height) was low. After two years of Lyrebird exclusion, soil compaction (top 7.5 cm) increased by 37% in exclusion plots compared with baseline measures, while in unfenced plots it decreased by 22%. Litter depth was almost three times greater in fenced than unfenced plots. Soil moisture, pH, and soil nutrients showed no difference between treatments. The enormous extent of litter and soil turned over by the Superb Lyrebird is unparalleled by any other vertebrate soil engineer in terrestrial ecosystems globally. The profound influence of such foraging activity on forest ecosystems is magnified by its year-round pattern and widespread distribution. The disturbance regime that Lyrebirds impose has implications for diverse ecosystem processes including decomposition and nutrient cycling, the composition of litter- and soil-dwelling invertebrate communities, the shaping of ground-layer vegetation patterns, and fire behavior and post-fire ecosystem recovery. Maintaining Lyrebird populations as a key facilitator of ecosystem function is now timely and critical as unprecedented wildfires in eastern Australia in summer 2019-2020 have severely burned ~12 million ha of forest, including ~30% of the geographic range of the Superb Lyrebird.

Animal movements in fire-prone landscapes.

Movement is a trait of fundamental importance in ecosystems subject to frequent disturbances, such as fire-prone ecosystems. Despite this, the role of movement in facilitating responses to fire has received little attention. Herein, we consider how animal movement interacts with fire history to shape species distributions. We consider how fire affects movement between habitat patches of differing fire histories that occur across a range of spatial and temporal scales, from daily foraging bouts to infrequent dispersal events, and annual migrations. We review animal movements in response to the immediate and abrupt impacts of fire, and the longer-term successional changes that fires set in train. We discuss how the novel threats of altered fire regimes, landscape fragmentation, and invasive species result in suboptimal movements that drive populations downwards. We then outline the types of data needed to study animal movements in relation to fire and novel threats, to hasten the integration of movement ecology and fire ecology. We conclude by outlining a research agenda for the integration of movement ecology and fire ecology by identifying key research questions that emerge from our synthesis of animal movements in fire-prone ecosystems.

Do multiple fires interact to affect vegetation structure in temperate eucalypt forests?

Fire plays an important role in structuring vegetation in fire-prone regions worldwide. Progress has been made towards documenting the effects of individual fire events and fire regimes on vegetation structure; less is known of how different fire history attributes (e.g., time since fire, fire frequency) interact to affect vegetation. Using the temperate eucalypt foothill forests of southeastern Australia as a case study system, we examine two hypotheses about such interactions: (1) post-fire vegetation succession (e.g., time-since-fire effects) is influenced by other fire regime attributes and (2) the severity of the most recent fire overrides the effect of preceding fires on vegetation structure. Empirical data on vegetation structure were collected from 540 sites distributed across central and eastern Victoria, Australia. Linear mixed models were used to examine these hypotheses and determine the relative influence of fire and environmental attributes on vegetation structure. Fire history measures, particularly time since fire, affected several vegetation attributes including ground and canopy strata; others such as low and sub-canopy vegetation were more strongly influenced by environmental characteristics like rainfall. There was little support for the hypothesis that post-fire succession is influenced by fire history attributes other than time since fire; only canopy regeneration was influenced by another variable (fire type, representing severity). Our capacity to detect an overriding effect of the severity of the most recent fire was limited by a consistently weak effect of preceding fires on vegetation structure. Overall, results suggest the primary way that fire affects vegetation structure in foothill forests is via attributes of the most recent fire, both its severity and time since its occurrence; other attributes of fire regimes (e.g., fire interval, frequency) have less influence. The strong effect of environmental drivers, such as rainfall and topography, on many structural features show that foothill forest vegetation is also influenced by factors outside human control. While fire is amenable to human management, results suggest that at broad scales, structural attributes of these forests are relatively resilient to the effects of current fire regimes. Nonetheless, the potential for more frequent severe fires at short intervals, associated with a changing climate and/or fire management, warrant further consideration.

Historical Maps from Modern Images: Using Remote Sensing to Model and Map Century-Long Vegetation Change in a Fire-Prone Region.

Understanding the age structure of vegetation is important for effective land management, especially in fire-prone landscapes where the effects of fire can persist for decades and centuries. In many parts of the world, such information is limited due to an inability to map disturbance histories before the availability of satellite images (~1972). Here, we describe a method for creating a spatial model of the age structure of canopy species that established pre-1972. We built predictive neural network models based on remotely sensed data and ecological field survey data. These models determined the relationship between sites of known fire age and remotely sensed data. The predictive model was applied across a 104,000 km(2) study region in semi-arid Australia to create a spatial model of vegetation age structure, which is primarily the result of stand-replacing fires which occurred before 1972. An assessment of the predictive capacity of the model using independent validation data showed a significant correlation (rs = 0.64) between predicted and known age at test sites. Application of the model provides valuable insights into the distribution of vegetation age-classes and fire history in the study region. This is a relatively straightforward method which uses widely available data sources that can be applied in other regions to predict age-class distribution beyond the limits imposed by satellite imagery.

Landscape properties mediate the homogenization of bird assemblages during climatic extremes.

Extreme weather events, such as drought, have marked impacts on biotic communities. In many regions, a predicted increase in occurrence of such events will be imposed on landscapes already heavily modified by human land use. There is an urgency, therefore, to understand the way in which the effects of such events may be exacerbated, or moderated, by different patterns of landscape change. We used empirical data on woodland-dependent birds in southeast Australia, collected during and after a severe drought, to document temporal change in the composition of bird assemblages in 24 landscapes (each 100 km2) representing a gradient in the cover of native wooded vegetation (from 60% to < 2%). We examined (a) whether drought caused region-wide homogenization of the composition of landscape bird assemblages, and (b) whether landscape properties influenced the way assemblages changed in response to drought. To quantify change, we used pairwise indices of assemblage dissimilarity, partitioned into components that represented change in the richness of assemblages and change in the identity of constituent species (turnover). There was widespread loss of woodland birds in response to drought, with only partial recovery following drought-breaking rains. Region-wide, the composition of landscape assemblages became more different over time, primarily caused by turnover-related differentiation. The response of bird assemblages to drought varied between landscapes and was strongly associated with landscape properties. The extent of wooded vegetation had the greatest influence on assemblage change: landscapes with more native vegetation had more stable bird assemblages over time. However, for the component processes of richness- and turnover-related compositional change, measures of landscape productivity had a stronger effect. For example, landscapes with more riparian vegetation maintained more stable assemblages in terms of richness. These results emphasize the importance of the total extent of native vegetation, both overall cover and that occurring in productive parts of the landscape, for maintaining bird communities whose composition is resistant to severe drought. While extreme climatic events cannot be prevented, their effects can be ameliorated by managing the pattern of native vegetation in anthropogenic landscapes, with associated benefits for maintaining ecological processes and human well-being.

Effects of time since fire on birds: how informative are generalized fire response curves for conservation management?

Fire is both a widespread natural disturbance that affects the distribution of species and a tool that can be used to manage habitats for species. Knowledge of temporal changes in the occurrence of species after fire is essential for conservation management in fire-prone environments. Two key issues are: whether postfire responses of species are idiosyncratic or if multiple species show a limited number of similar responses; and whether such responses to time since fire can predict the occurrence of species across broad spatial scales. We examined the response of bird species to time since fire in semiarid shrubland in southeastern Australia using data from surveys at 499 sites representing a 100-year chronosequence. We used nonlinear regression to model the probability of occurrence of 30 species with time since fire in two vegetation types, and compared species' responses with generalized response shapes from the literature. The occurrence of 16 species was significantly influenced by time since fire: they displayed six main responses consistent with generalized response shapes. Of these 16 species, 15 occurred more frequently in mid- or later-successional vegetation (> 20 years since fire), and only one species occurred more often in early succession (< 5 years since fire). The models had reasonable predictive ability for eight species, some predictive ability for seven species, and were little better than random for one species. Bird species displayed a limited range of responses to time since fire; thus a small set of fire ages should allow the provision of habitat for most species. Postfire successional changes extend for decades and management of the age class distribution of vegetation will need to reflect this timescale. Response curves revealed important seral stages for species and highlighted the importance of mid- to late-successional vegetation (> 20 years). Although time since fire clearly influences the distribution of numerous bird species, predictive models of the spatial distribution of species in fire-prone landscapes need to incorporate other factors in addition to time since fire.

Birds in agricultural mosaics: the influence of landscape pattern and countryside heterogeneity.

Agricultural environments are critical to the conservation of biota throughout the world. Efforts to identify key influences on the conservation status of fauna in such environments have taken complementary approaches. Many studies have focused on the role of remnant or seminatural vegetation and emphasized the influence on biota of spatial patterns in the landscape. Others have recognized that many species use diverse "countryside" elements within farmland, and emphasize the benefits of landscape heterogeneity for conservation. Here, we investigated the effect of independent measures of both the spatial pattern (extent and configuration) and heterogeneity of elements (i.e., land uses/vegetation types) on bird occurrence in farm-scale agricultural mosaics in southeastern Australia. Birds were sampled in all types of elements in 27 mosaics (each 1 x 1 km) selected to incorporate variation in cover of native vegetation and the number of different element types in the mosaic. We used an information-theoretic approach to identify the mosaic properties that most strongly influenced bird species richness. Subgroups of birds based on habitat requirements responded most strongly to the extent of preferred elements in mosaics. Woodland birds were richer in mosaics with higher cover of native vegetation while open-tolerant species responded to the extent of scattered trees. In contrast, for total species richness, mosaic heterogeneity (richness of element types) and landscape context (cover of native vegetation in surrounding area) had the greatest influence. These results showed that up to 76% of landscape-level variation in richness of bird groups is attributable to mosaic properties directly amenable to management by landowners. Key implications include (1) conservation goals for farm landscapes must be carefully defined because the richness of different faunal components is influenced by different mosaic properties; (2) the extent of native vegetation is a critical influence in agricultural environments because it drives the farm-scale richness of woodland birds and has a broader context effect on total bird richness in mosaics; (3) land-use practices that enhance the heterogeneity of farmland mosaics are beneficial for native birds; and (4) the cumulative effect of even small elements in farm mosaics contribute to the structural properties of entire landscapes.