A conceptual model of vegetation dynamics for the unique obligate‐seeder eucalypt woodlands of south‐western Australia

Models of vegetation dynamics framed as testable hypotheses provide powerful tools for predicting vegetation change in response to contemporary disturbances or climate change. Synthesizing existing information and applying new data, we develop a conceptual model of vegetation states and transitions for the previously overlooked woodlands dominated by obligate‐seeder eucalypts of dry to semi‐arid south‐western Australia. These comprise the largest extant temperate woodland globally, are uniquely dominated by a high diversity of obligate‐seeder eucalypts (55 taxa), but are under threat from wildfire. Our conceptual model incorporates four critical ecological processes that also distinguish obligate‐seeder woodlands from temperate woodlands dominated by resprouting eucalypts: (i) a lack of well‐protected epicormic buds results in major disturbances (prominently fire) being stand‐replacing. The pre‐disturbance tree cohort is killed, followed by dense post‐disturbance recruitment from seed shed from a serotinous seed bank; (ii) competition between saplings leads to self‐thinning over a multi‐century timeframe, with surviving individuals having great longevity (regularly >400 years); (iii) multiple processes limit recruitment in the absence of stand‐replacement disturbances, leading to frequent single‐cohort stands. However, unlike the few other obligate‐seeder eucalypt communities, trickle recruitment in very long‐unburnt stands can facilitate indefinite community persistence in the absence of stand‐replacement disturbances; and (iv) discontinuous fuels, relatively low understorey flammability (low grass and often high chenopod cover) and topographic barriers to fire (salt lakes) allow mature woodlands to persist for centuries without burning. Notably though, evidence suggests that flammability peaks at intermediate times since fire, establishing a ‘flammability bottleneck’ (or landscape fire trap) through which regenerating woodlands must pass. Our model provides a framework to support management to conserve obligate‐seeder eucalypt woodlands. Research into reasons for exceptional tree heights relative to ecosystem productivity, the evolution of diverse and dominant obligate‐seeder eucalypts, the paucity of grass, and the recent spatial distribution of fires, will further inform conservation management.     

Revegetation,restoration and reptiles in rural landscapes: Insights from long‐term monitoring programmes in the temperate eucalypt woodlands of south‐eastern Australia

Over the past decade, there has been a concerted effort to better understand the distribution and abundance of reptiles in agricultural landscapes and to specifically evaluate their response to revegetation (tree and shrub plantings) and habitat restoration in the wheat‐sheep belt of south‐eastern Australia. This article reviews the response of reptiles to revegetation and woodland management and provides ten insights and lessons that can be applied to help improve reptile conservation in temperate eucalypt woodlands and fragmented agricultural landscapes in Australia. The review focuses primarily on revegetation programmes conducted by Landcare and Greening Australia, and management interventions funded by Local Land Services in NSW and Catchment Management Authorities in Victoria.     

Characteristics of woods used recently and historically by Lesser Spotted Woodpeckers Dendrocopos minor in England

Lesser Spotted Woodpecker Dendrocopos minor numbers have declined greatly in England since the early 1980s for reasons that are not yet fully understood. It has been suggested that the species’ decline may be linked to the increase in Great Spotted Woodpeckers Dendrocopos major, changes in woodland habitat quality (such as deadwood abundance) and landscape‐scale changes in tree abundance. We tested some of these hypotheses by comparing the characteristics of woods in southern England where the species is still relatively numerous with those of woods used in the 1980s before the major decline. In each time period, habitat, predator and landscape information from woods known to be occupied by Lesser Spotted Woodpeckers was compared with those found to be unoccupied during surveys. Before the main period of decline, Lesser Spotted Woodpeckers used oak‐dominated, mature, open woods with a large amount of standing deadwood. Habitat use assessed from recent data was very similar, the species being present in mature, open, oak‐dominated woodlands. There was a strong relationship between wood use probability and the extent of woodland within a 3‐km radius, suggesting selection for more heavily wooded landscapes. In recent surveys, there was no difference in deadwood abundance or potential predator densities between occupied and unoccupied woods. Habitat management should focus on creating and maintaining networks of connected woodlands in areas of mature, open woods. Finer‐scale habitat selection by Lesser Spotted Woodpecker within woodlands should be assessed to aid development of beneficial management actions.     

The importance of temperate woodland in travelling stock reserves for vertebrate biodiversity conservation

Summary Travelling stock reserves have well recognised and important commercial, cultural and other values in the pastoral and agricultural regions of Australia, but their conservation values remain poorly known. In general, many areas of woodlands within travelling stock reserves have not been subjected to the same high levels of degradation pressure as similar temperate woodlands on private land. In this study, we compared several measures of vertebrate biota in 217 permanent field sites located in travelling stock reserves and woodland remnants located almost exclusively on private land in the Murray Riverina and South West Slopes bioregions in southern New South Wales. We found that in comparison with temperate woodland remnants on private land, temperate woodland in travelling stock reserves tended to support more species of birds, more species of declining birds, and a greater abundance of arboreal marsupials. Temperate woodland in travelling stock reserves was more likely to be occupied by particular individual species of birds of conservation concern. However, such tenure differences were not consistent between bioregions. For example, the Rufous Whistler (Pachycephala rufiventris) and the Red‐capped Robin (Petroica goodenovii ) were significantly more likely to occupy temperate woodland in travelling stock reserves than temperate woodland on private land in the Murray Riverina but such effects were not apparent in the South West Slopes. Similarly, while the Grey‐crowned Babbler (Pomatostomus temporalis) was significantly more likely to occupy temperate woodland in travelling stock reserve sites in the South West Slopes, we found the opposite effect in Murray Riverina. We demonstrated that travelling stock reserves have important conservation values for some taxa and individual species. Because of the paucity of large ecological reserves in the Murray Riverina and South West Slopes bioregions, temperate woodlands located in travelling stock reserves should be considered as an important resource that contributes to a portfolio of vegetation assets in formerly woodland‐dominated, but now extensively cleared, agricultural landscapes.     

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.     

Restoring grassy woodland diversity through direct seeding: Insights from six ‘best‐practice’ case studies in southern Australia

Ecological restoration of grassy woodland ecosystems is now a significant landscape‐scale conservation objective throughout southern Australia. Technological improvements in direct seeding are now sufficiently well‐advanced to examine whether cost‐effective restoration of grassy woodlands is feasible. Consideration of six ‘best practice case studies shows substantial evidence of success. Further refinement of direct seeding techniques, in combination with native seed production systems, however, will be required into the future to meet the scale of woodland conservation targets and restore ecological function.     

Broad‐scale patterns in plant diversity vary between land uses in a variegated temperate Australian agricultural landscape

Plant diversity is threatened in many agricultural landscapes. Our understanding of patterns of plant diversity in these landscapes is mainly based on small‐scale (<1000 m²) observations of species richness. However, such observations are insufficient for detecting the spatial heterogeneity of vegetation composition. In a case‐study farm on the North‐West Slopes of New South Wales, Australia, we observed species richness at four scales (quadrat, patch, land use and landscape) across five land uses (grazed and ungrazed woodlands, native pastures, roadsides and crops). We applied two landscape ecological models to assess the contribution of these land uses to landscape species richness: (i) additive partitioning of diversity at multiple spatial scales, and (ii) a measure of habitat specificity – the effective number of species that a patch contributes to landscape species richness. Native pastures had less variation between patches than grazed and ungrazed woodlands, and hence were less species‐rich at the landscape scale, despite having similar richness to woodlands at the quadrat and patch scale. Habitat specificity was significantly higher for ungrazed woodland patches than all other land uses. Our results showed that in this landscape, ungrazed woodland patches had a higher contribution than the grazed land uses to landscape species richness. These results have implications for the conservation management of this landscape, and highlighted the need for greater consensus on the influence of different land uses on landscape patterns of plant diversity.     

Distinct ecosystem types respond differentially to grazing exclosure

Here, we evaluate the ecosystem functioning and the ecosystems services supply of different vegetation types (grasslands, shrublands and woodlands) under contrasting management regimes by comparing a protected area with the surrounding landscape, which has been subjected to human disturbance in the Eastern Hills of Uruguay. We propose, based on functional attributes and vegetation physiognomy, a State and Transition Model for the dynamics of the grassland–woodland mosaic. We used remote sensing techniques to: (i) develop a land‐cover map of the study area based on supervised Landsat imagery classification, and (ii) compare attributes of the ecosystem functioning (productivity and seasonality) and service supply derived from the Normalized Difference Vegetation Index (NDVI) images provided by the moderate resolution imaging spectroradiometer (MODIS) sensor. The land‐cover map showed that grasslands and shrublands were the most extensive land covers in the study area. These vegetation types presented higher productivity, seasonality and ecosystem service supply, outside the protected area than inside it. On the other hand, woodlands showed higher productivity, ecosystem service supply and lower seasonality inside the protected area than outside of it. Two axes represented the grassland–woodland mosaic dynamic: (i) the mean annual and (ii) the intra‐annual coefficient of variation of the NDVI. Our results highlight that conservation of grasslands, shrublands and woodlands require different management strategies based on particular disturbance regimes like moderate grazing and controlled burns. Moderate disturbances may help to preserve ecosystem services provisioning in grasslands and shrublands. On the contrary, woodland conservation requires a more rigorous regime of protection against disturbances.     

Woodland biodiversity, palaeo-human ecology and some implications for conservation management

This paper appraises the role of prehistoric human societies in modifying the natural woodland biodiversity of several regions throughout Scotland. It draws attention to the likelihood that, contrary to popular belief, biodiversity was enhanced by interactions between agricultural communities and woodlands. The possible purposefulness of such interactions is discussed. Mindful of the present concern to re-establish ‘native’ woodlands in many parts of northern Britain, we then review three approaches to the identification of native tree types in Scotland, and conclude that insufficient attention in reafforestation schemes is being paid to (a) the palaeoecological record as a record of native woodland or (b) the likely former high taxonomic diversity of woodlands. Both these failings will lead to the creation of new woodlands lacking in species diversity.     

How widespread is woody plant encroachment in temperate Australia? Changes in woody vegetation cover in lowland woodland and coastal ecosystems in Victoria from 1989 to 2005

Aim Encroachment or densification by woody plants affects natural ecosystems around the world. Many studies have reported encroachment in temperate Australia, particularly in coastal ecosystems and grassy woodlands. However, the degree to which published studies reflect broad-scale changes is unknown because most studies intentionally sampled areas with conspicuous densification. We aimed to estimate changes in woody vegetation cover within lowland grassy woodland and coastal ecosystems in Victoria from 1989 to 2005 to determine whether published reports of recent encroachment are representative of broad-scale ecosystem changes. Location All lowland grassy woodland and coastal ecosystems (c. 6.11 × 10⁵ ha) in Victoria, Australia. Four major ecosystems were analysed: Plains woodlands, Herb-rich woodlands, Riverine woodlands and Coastal vegetation. Methods Changes in woody vegetation cover from 1989 to 2005 were assessed based on state-wide vegetation maps and Landsat analyses of woody vegetation cover conducted by the Australian Greenhouse Office’s National Carbon Accounting System. The results show changes in woody cover within mapped patches of native vegetation, rather than changes in the extent of woody vegetation resulting from clearing and revegetation. Results When pooled across all ecosystems, woody vegetation increased by 18,730 ha from 1989 to 2005. Woody cover within Riverine woodlands and within Plains woodlands each increased by >7000 ha. At the patch scale, the mean percentage cover of woody vegetation in each polygon increased by >5% in all four ecosystems: Riverine woodlands (+9.2% on average), Herb-rich woodlands (+7.6%), Plains woodlands (+6.7%) and Coastal vegetation (+5.9%). Regression models relating degree of encroachment to geographic and climatic variables were extremely weak (r² ≤ 0.026), indicating that most variation occurred at local scales rather than across broad geographic gradients. Main conclusions At the scale of observation, woody vegetation cover increased in all lowland woodland and coastal ecosystems over the 16-year period. Thus, published examples of encroachment in selected coastal and woodland patches do appear to reflect widespread increases in woody vegetation cover in these ecosystems. This densification appears to be associated with changes in land management rather than with post-fire vegetation recovery and is likely to be ongoing and long-lasting, with substantial implications for biodiversity conservation and ecosystem services.     

Using historical woodland creation to construct a long‐term,large‐scale natural experiment: the WrEN project

Natural experiments have been proposed as a way of complementing manipulative experiments to improve ecological understanding and guide management. There is a pressing need for evidence from such studies to inform a shift to landscape‐scale conservation, including the design of ecological networks. Although this shift has been widely embraced by conservation communities worldwide, the empirical evidence is limited and equivocal, and may be limiting effective conservation. We present principles for well‐designed natural experiments to inform landscape‐scale conservation and outline how they are being applied in the WrEN project, which is studying the effects of 160 years of woodland creation on biodiversity in UK landscapes. We describe the study areas and outline the systematic process used to select suitable historical woodland creation sites based on key site‐ and landscape‐scale variables – including size, age, and proximity to other woodland. We present the results of an analysis to explore variation in these variables across sites to test their suitability as a basis for a natural experiment. Our results confirm that this landscape satisfies the principles we have identified and provides an ideal study system for a long‐term, large‐scale natural experiment to explore how woodland biodiversity is affected by different site and landscape attributes. The WrEN sites are now being surveyed for a wide selection of species that are likely to respond differently to site‐ and landscape‐scale attributes and at different spatial and temporal scales. The results from WrEN will help develop detailed recommendations to guide landscape‐scale conservation, including the design of ecological networks. We also believe that the approach presented demonstrates the wider utility of well‐designed natural experiments to improve our understanding of ecological systems and inform policy and practice.     

Ecological niche breadth and microhabitat guild structure in temperate Australian reptiles: Implications for natural resource management in endangered grassy woodland ecosystems

Ecological theory predicts that species with narrow niche requirements (habitat specialists) are more vulnerable to anthropocentric disturbances than those with broad niche requirements (habitat generalists). Hence, understanding a species ecological niche and guild membership would serve as a valuable management tool for providing a priori assessments of a species extinction risk. It also would help to forecast a species capacity to respond to land use change, as what might be expected to occur under financial incentive schemes to improve threatened ecological vegetation communities. However, basic natural history information is lacking for many terrestrial species, particularly reptiles in temperate regions of the world. To overcome this limitation, we collated 3527 reptile observations from 52 species across an endangered woodland ecoregion in south‐eastern Australia and examined ecological niche breadth and microhabitat guild structure. We found 30% of species had low ecological niche values and were classified as habitat specialists associated with large eucalypt trees, woody debris, surface rock or rocky outcrops. Cluster analysis separated species into six broad guilds based on microhabitat similarity. Approximately 80% of species belonged to guilds associated with old growth vegetation attributes or non‐renewable litho‐resources such as surface rock or rocky outcrops. Our results suggest that agri‐environment schemes that focus purely on grazing management are unlikely to provide immediate benefits to broad suites of reptiles associated with old growth vegetation and litho‐resources. Our classification scheme will be useful for identifying reptile species that are potentially vulnerable to anthropocentric disturbances and may require alternative strategies for improving habitat suitability and reptile conservation outcomes in grassy woodland ecosystems.     

Remotely‐sensed foliage cover and ground‐measured stand attributes are complimentary when estimating tree hollow abundances across relictual woodlands in agricultural landscapes

Hollows, also known as tree cavities, are critical to the survival of many animal species but are too poorly mapped across landscapes to allow for their adequate consideration in regional planning. Managing cost is important, so we tested whether freely available satellite‐derived foliage projective cover and field‐measured stand attributes could be used separately or combined to predict tree hollow abundance in relictual Australian temperate woodlands. Satellite‐derived foliage projective cover revealed variation in woody vegetation densities both within mapped woodland remnants and cleared areas of the agricultural matrix. Plot‐based field assessment of the actual number of hollows in each one‐hectare site (n = 110 sites) revealed a relationship with foliage cover. Improvement of the model was achieved if site‐based estimates of the proportion of the canopy due to Eucalyptus species and the number of mature trees per hectare were included. Remotely sensed foliage cover can improve on traditional vegetation mapping for predicting hollow‐bearing tree and hollow abundances at landscape scales when managing hollow‐dependent fauna habitat across relictual woodlands in temperate Australian agricultural landscapes. At finer scales, the addition of other predictors is necessary to raise the accuracy of the predicted hollow densities.     

Continental‐scale syntheses of Australian pyromes – misclassification of south‐western eucalypt woodlands misinforms management

Global and continental‐scale analysis of ecological phenomena can offer important insights through the identification of patterns and associations not detectable at smaller scales. However, using proxies for ecological phenomena, such as vegetation mapping for spatially projecting fire regime niches and post‐fire plant responses, require critical examination of predictions to determine utility. Using local studies in south‐western Australia, we demonstrate that while this approach has been largely successful in mallee woodland and shrubland, it has failed in eucalypt woodland, with the consequence that values for a range of fire‐related parameters from the continent‐wide approaches, if adopted in informing management, would result in undesirable conservation outcomes for the world's largest extant temperate woodland.     

Woodland Restoration in the Western Australian Wheatbelt: A Conceptual Framework Using a State and Transition Model

Woodlands dominated by Eucalyptus spp. in temperate southeastern and southwestern Australia have been extensively cleared for agriculture and are often badly degraded by livestock grazing. This has resulted in the loss of biodiversity and widespread land degradation. The continuing decline of these woodlands has become a concern for the conservation of biodiversity, and there is a growing interest among farmers, land managers, and researchers in developing techniques for restoring them. Currently few scientific guidelines exist for undertaking woodland restoration programs. We use a state and transition model to develop hypotheses on restoration strategies for salmon gum (Eucalyptus salmonophloia) woodlands. We consider that this approach provides a suitable framework for organizing knowledge and identifying areas where further information is needed, and hence provides a useful starting point for a restoration program. The model has the potential to provide a tool for land managers with which they can assess the action and effort needed to undertake woodland restoration in agricultural landscapes.     

Climate,landscape, habitat,and woodland management associations with hazel dormouse Muscardinus avellanarius population status

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Microsite factors influencing Eucalyptus regeneration in temperate woodlands

Natural regeneration of Eucalyptus is a major problem for the restoration and preservation of temperate woodlands. South‐eastern Australia experienced a period of above average rainfall from mid‐2010 to early 2011, creating favourable conditions for localised Eucalyptus regeneration. We conducted a case–control study comparing sites where natural regeneration from seed was present and absent. Vegetation cover, soil condition and hydrophobicity, as well as shelter objects and landscape position, were recorded to identify what influences natural regeneration from seed. Using conditional logistic regression, we found the odds of regeneration occurring increased with uphill slope angle and herbaceous cover. Our results suggest that, during periods of high rainfall, stable microsites that capture seed and resources are most likely to support Eucalyptus regeneration.     

Bryophytes as indicators of ancient woodlands in Schleswig-Holstein (Northern Germany)

Ancient woodlands, with their long ecological continuity, frequently harbor a high number of typical, rare and threatened species, and are therefore of particular importance for nature conservation. To pinpoint these habitats, a common application is the use of plants as “ancient woodland indicators”. The occurrence of these particular species allows for evaluating the continuity of woodland cover in time. While lists of ancient woodland vascular plants have been derived for many regions, the identification and use of bryophytes as ancient woodland indicators has been widely neglected. This is a bit surprising because certain woodland bryophytes are very sensitive to varying environmental conditions or changes in land management. It therefore appeared promising to compile an ecologically grounded list of ancient woodland indicator bryophytes for practical use.In this study, we present a set of ancient woodland indicator bryophytes based on the analysis of datasets from the North German federal state of Schleswig-Holstein. To compile this list, we systematically evaluated the bryophyte distribution data from floristic surveys in relation to ancient woodland cover data from state-wide inventories. In this way, we were able to determine ancient woodland bryophytes using consistent and repeatable statistical methods.The presented list of 31 ancient woodland indicator bryophytes is ecologically sound and corresponds well with data from the sparse literature. We could distinguish two groups of ancient woodland indicator bryophytes. The first group is linked to base-rich, semi-natural deciduous woodlands with high soil and air humidity. The second group comprises acidophilic bryophytes that occur not only in acidic beech and oak woods, but also in acidic mixed or coniferous forests on ancient woodland sites. Apart from the ancient woodland indicator bryophytes, we could identify one group of recent woodland bryophytes and four groups of bryophytes that are more or less indifferent with respect to woodland continuity.Finally, we provide recommendations for the application of ancient woodland indicator bryophytes in nature conservation practice. Management suggestions for the conservation of the typical bryophyte diversity of ancient semi-natural woodlands are also given.     

Woodland expansion in South African grassy biomes based on satellite observations (1990–2013): general patterns and potential drivers

Increases in woody plant cover in savanna grassland environments have been reported on globally for over 50 years and are generally perceived as a threat to rangeland productivity and biodiversity. Despite this, few attempts have been made to estimate the extent of woodland increase at a national scale, principally due to technical constraints such as availability of appropriate remote sensing products. In this study, we aimed to measure the extent to which woodlands have replaced grasslands in South Africa's grassy biomes. We use multiseason Landsat data in conjunction with satellite L‐band radar backscatter data to estimate the extent of woodlands and grasslands in 1990 and 2013. The method employed allows for a unique, nationwide measurement of transitions between grassland and woodland classes in recent decades. We estimate that during the 23‐year study period, woodlands have replaced grasslands over ~57 000 km² and conversely that grasslands have replaced woodlands over ~30 000 km², a net increase in the extent of woodland of ~27 000 km² and an annual increase of 0.22%. The changes varied markedly across the country; areas receiving over 500 mm mean annual precipitation showed higher rates of woodland expansion than regions receiving <500 mm (0.31% yr^?1 and 0.11% yr^?1, respectively). Protected areas with elephants showed clear loss of woodlands (?0.43% yr^?1), while commercial rangelands and traditional rangelands showed increases in woodland extent (>0.19% yr^?1). The woodland change map presented here provides a unique opportunity to test the numerous models of woody plant encroachment at a national/regional scale.     

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