Seedling establishment of Acacia tortilis and Hyphaene compressa in the Turkwel riverine forest, Kenya

This study reports on the spatial and temporal patterns of seedling establishment in the Turkwel riverine forest, Kenya. Seedlings of the dominant tree species Acacia tortilis and Hyphaene compressa were mapped and monitored to assess the underlying causes of seedling recruitment and mortality. The broad‐scale distribution of A. tortilis was not correlated with any environmental variables, while H. compressa seedlings were confined to flood‐exposed sites in the arid downstream section of the floodplain. One year of monitoring showed that seedling recruitment of A. tortilis was evoked by prolonged rainfall, while seedling mortality was caused by desiccation, browsing and trampling. In contrast, seedling recruitment and mortality of H. compressa was largely unaffected by rainfall and livestock, probably due to the high moisture requirements of seeds and the tolerance of seedlings to disturbance. There were no effects of soils, light, or seedling density on the establishment of A. tortilis and H. compressa seedlings. This study demonstrates the importance of parallel mapping and monitoring of riverine seedlings in order to understand patterns and processes of forest regeneration in arid and semi‐arid floodplains.     

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.     

Fragmentation in eucalypt woodlands promotes nest‐tree occupancy by a despotic species,the noisy miner (Manorina melanocephala)

The effects of habitat fragmentation as a threat to biodiversity are well known; decreased connectivity can potentially influence population processes and dynamics, resulting in smaller, isolated populations that may not function optimally. However, fragmentation may also increase the amount of edge or ecotone habitat available to open country species, benefiting their populations and enabling them to dominate remnant habitats. Noisy miners (Manorina melanocephala) are one such species, occupying eastern‐Australian eucalypt woodlands. They are considered a ‘despotic’ species, in that their presence negatively impacts woodland avifauna biodiversity due to their aggressive exclusion of other taxa from occupied areas. Despite this well‐known impact, little information exists on the patterns of nest‐tree occupancy by noisy miners within eucalypt woodlands. In the current study, we explored the patterns of nest‐tree occupancy by noisy miners across two successive years, aiming to identify preferences for breeding areas relative to vegetation structure. Our results show that both habitat fragmentation and the characteristics of individual eucalypt trees in an area influenced nest‐tree occupancy. Noisy miners constructed nests in trees near the edge of woodland patches more often than expected. Moreover, the nest tree chosen was a eucalypt that was significantly smaller than randomly selected trees from the surrounding area. The results highlight the importance of habitat management measures that may reduce the suitability of woodland patches as nesting sites for this species, in order to mitigate the severe effects of this despotic edge specialist.     

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.     

Vegetation-environment relations of a Middle Zambezi floodplain

Detrended correspondence analysis was used to study the relationships between environmental factors and the species composition of vegetation on Zambezi River alluvium downstream of the Kariba hydroelectric dam. Grass, sedge and woody species were recorded in 73 stands in Mana Pools National Park, Zimbabwe. Grass and sedge species composition was related to the soil moisture regime, as indexed by soil texture and flooding frequency. The first woody plant ordination axis was related to a stand development gradient; Acacia albida was a pioneer species on lowlying sandbanks and the woody species richness of stands increased with their height above the Zambezi River. Two-way indicator species analysis identified 7 vegetation types which could be separated on the basis of their topsoil texture and flooding frequency. The types were: sandbanks; young A. albida woodland; A. albida woodland; A. albida dominated mixed woodland; mixed riverine woodland with understory; mixed riverine woodland; and grassland on clay soils.     

Model highlights likely long‐term influences of mesobrowsers versus those of elephants on woodland dynamics

The potential long‐term influences of mesobrowsers versus those of savannah elephants on woodland dynamics have not been explored. This may be a critical omission especially in southern African savannahs, where efforts to preserve existing woodlands are typically directed at elephant management. We describe a simple browse–browser model, parameterized from an extensive review of the literature and our own data, including quantitative assessment of impala impact, from the study site, iMfolozi Park, South Africa. As there is a paucity of species‐specific demographic data on savannah woody species, we modelled, in a novel approach, functional groups of plant species typical of Acacia woodlands. Outputs suggest that over the long term (100 years), low‐to‐moderate densities of impala will have a similar impact on woodland structure, in terms of density of adult trees, as low‐to‐moderate densities of elephant. Further, the outputs highlight the apparently strong synergistic effect impala and elephant impacts combined have on woodland dynamics, suggesting that reduction or removal of either impala or elephant will radically reduce long‐term destruction of savannah woodlands. Recorded changes in adult tree numbers in iMfolozi broadly supported the model's outputs.     

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

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

Flood‐deposited wood creates regeneration niches for riparian vegetation on a semi‐arid South African river

Question : The formation of large woody debris (LWD) piles during floods has significant impacts on riparian succession through pioneering plants often establishing in association with wood. We assess the importance of LWD for seed regeneration of riparian plants after a century‐scale flood disturbance in a semi‐arid environment. Location : The Sabie River within Kruger National Park in the semi‐arid northeast of South Africa. Methods : Our approach was to quantify the riparian soil seed bank, to record the frequency of establishment of riparian plants in woody debris piles, and to conduct experimental out‐plantings of common riparian trees in plots with and without LWD. Results : We found the abundance and diversity of seedlings were higher in soils taken from wood piles than from open reference areas, and most seedlings were herbaceous species. Surveys indicated that numbers of seedlings recorded within woody debris were significantly greater than in open reference areas or within established vegetation. Seedling establishment in various cover‐types also varied for different riparian tree species. Experimental out‐planting of seedlings of two riparian tree species (Philenoptera violacea and Combretum erythrophyllum) revealed that, after 433 days, planted seedlings survived only in woody debris piles. Conclusion : LWD formed after a large flood creates heterogeneous patches that may influence post‐disturbance regeneration of riparian vegetation by providing a variety of environmental niches for seedlings establishment. We suspect that higher seedling survival in LWD is due to increased moisture (particularly in the dry season) and nutrients, and protection from seasonal flooding and herbivory.     

Bat Response to Woodland Restoration within Urban Forest Fragments

In urban environments, woodland areas are typically fragmented and subject to invasive species encroachment, woody overgrowth, and natural succession. In response to negative impacts, conservationists and land managers have implemented restoration strategies to enhance the integrity of woodlands. Because woodland habitat is important for bats (Order Chiroptera), alterations to forest structures may affect how bats utilize forest fragments in urban environments. We evaluated relationships among restoration efforts, microhabitat characteristics and overall bat activity, and interspecific variation among bats in response to woodland characteristics. We monitored bats in nine woodland forest preserves representing various stages of restoration within the Chicago metropolitan area in 2004 and 2005. Overall bat activity was positively related to prescribed burning, invasive species removal, and small tree density (7.7‐20 cm dbh) and inversely related to shrub density and clutter at 0‐6 m heights. There was interspecific variation in response to alterations in woodland structure, with Lasiurus borealis (L. borealis) positively associated with small and medium (20.1‐33 cm) tree densities and inversely related to clutter at 0‐9 m; Myotis spp. positively associated with canopy cover, clutter at 6‐9 m, and small and medium tree densities; and Lasionycteris noctivagans (L. noctivagans) positively associated with more open forests. Eptesicus fuscus (E. fuscus) activity was not strongly associated with any measured vegetation variable. Our results demonstrate bats positively respond to some forms of woodland restoration in urban landscapes. However, species‐specific responses to vegetation differed and should be taken into consideration when developing management plans.     

Carbon storage,structure and composition of miombo woodlands in Tanzania’s Eastern Arc Mountains

We determine the aboveground biomass and carbon storage (ABGC) of trees and the herbaceous layer in miombo woodland in the Eastern Arc Mountains (EAM) of Tanzania. In four 1‐ha sample plots in Nyanganje and Kitonga Forests, we measured all trees ≥10 cm diameter alongside height and wood mass density. The plots contained an average of 20 tree species ha^?1 (range 11–29) and 344 stems ha^?1 (range 281–382) with Shannon diversity values of 1.05 and 1.25, respectively. We weighted nine previously published woody savannah allometric models based on whether: (i) the model was derived from the same geographical region; (ii) the model included tree height/wood mass density in addition to stem diameter; and (iii) sample size was used to fit the model. The weighted mean ABGC storage from the nine models range from 13.5 ± 2 to 29.8 ± 5 Mg ha^?1. Measured ABGC storage in the herbaceous layer, using the wet combustion method, adds 0.55 ± 0.02 Mg C ha^?1. Estimates suggest that EAM miombo woodlands store a range of 13–30 Mg ha^?1 of carbon. Although the estimates suggest that miombo woodlands store significant quantities of carbon, caution is required as this is the first estimate based on in situ data.     

Compositional patterns of riparian woodlands in the Rift Valley of northern Kenya

Environmental factors governing the composition of riparian woodlands of the Gregory Rift Valley, northern Kenya were investigated. We collected tree species composition data on elevational transects along four active stream channels and two recently abandoned channels in Turkana District, Kenya and examined variation in riparian woodland species richness and composition at two scales: first, regional differences among channels and second, patterns along individual channels. The major factors governing woodland composition were: geographic location in relation to rainfall distribution, local heterogeneity in stream substrate, fluvial disturbance within individual channels and, in some cases, location along elevation gradients. Species richness increased with increasing rainfall. There were no significant differences in the composition of woodlands associated with active and recently abandoned channel segments. However, the species richness of riparian woodlands along abandoned channel segments seemed to decline with time since abandonment. The species richness of riparian woodlands in Turkana is an order of magnitude lower than that of comparable neotropical dry forests.     

State‐space modeling reveals habitat perception of a small terrestrial mammal in a fragmented landscape

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A reassessment of the fire‐tolerance of some miombo woody species in the Central Province,Zambia

The fire‐tolerance of 21 woody species in a dry miombo woodland in the Central Province, Zambia was evaluated and compared to an existing classification. The evaluation is based on differences in the vertical distribution of canopy volumes over a range of height intervals after repeated exposure of woodlands to fire. Woody species that showed an inverse correlation between canopy volume and tree height were classified as fire‐sensitive, whilst those that showed a positive correlation were classified as fire‐tolerant. A new method of a rapid and simple field evaluation is described, and further applications are discussed.     

Natural regeneration of trees in urban woodlands

Abstract. We studied tree regeneration, a key process for the existence of urban woodlands. We hypothesized that, besides the usual biological factors, anthropogenic ones (fragmentation, wear, pollution etc.) determine the regeneration success of tree species in urban woodlands. To test this hypothesis, within an observational setting, we collected data from 30 urban woodlands in the cities of Helsinki and Vantaa, Finland. We defined the number of living saplings (30–200 cm in height) as an indicator of regeneration success and used regression analysis to test different factors as independent variables. The results showed that different tree species responded differently to urban pressure. The regeneration of Picea abies decreased with increasing fragmentation of the forest landscape, whereas for the other most common (deciduous) species, regeneration increased. Wear, measured as total path area per study site, had a negative effect on regeneration success. An a posteriori examination of the data suggested that coarse woody debris might promote regeneration. We conclude that, although tree regeneration in general is not threatened in urban woodlands in the area we studied, the species composition may gradually change. We discuss some management implications for counteracting the urban pressures on tree regeneration.     

Modelling fallen branch volumes in a temperate eucalypt woodland: implications for large senescent trees and benchmark loads of coarse woody debris

Fallen branches are a substantial component of coarse woody debris and a key ecological resource. The depletion of stocks of coarse woody debris since European settlement has contributed to the degradation of Australian grassy box woodlands, including the loss of biodiversity. Restoration options for remnant woodlands include the augmentation of coarse woody debris stocks. However, the extensive modification of grassy box woodlands has left few reference sites for establishing benchmarks to guide such restoration. In this paper we demonstrate a method for predicting fallen branch debris loads in the absence of reference sites, using data from a yellow box–red gum woodland. Our methodology is in two stages: first, the total volume of branch debris under individual trees was modelled; and second, these models were applied to groups of trees to predict stand‐level loads of fallen branch debris. Although the models were developed for yellow box–red gum woodlands, the methodology would be applicable to other communities that lack reference sites. Predicted benchmark loads of fallen branch debris for yellow box–red gum woodland were between 7.0 m³ ha^?1 and 11.9 m³ ha^?1. Large senescing trees contributed the bulk of fallen branch debris. Model predictions indicated a 100‐cm diameter at breast height (dbh) tree was 10 times more likely to produce debris than a 50‐cm dbh tree, and if debris was present a 100‐cm dbh tree produced approximately 10 times the volume of branch debris produced by a 50‐cm dbh tree. These results highlight the importance of large senescing trees for the production of fallen branch debris and support the keystone role of large trees within remnant woodlands, and the need to conserve these structures. Our results also support the active management of regrowth woodland stands to facilitate the progression of individual trees to maturity and senescence. In particular, thinning of regrowth stands may promote the growth of retained trees, ensuring they contribute to fallen branch debris stocks with a minimum time lag.     

Vegetation dynamics in a Quercus‐Juniperus savanna: An isotopic assessment

Abstract. Woody plants are increasing in many grassland and savanna ecosystems around the world. As a case in point, the Edwards Plateau of Texas, USA, is a vast region (93 000 km²) in which rapid woody encroachment appears to be occurring. The native vegetation (prior to the Anglo‐European settlement 150–200 yr ago) and the biogeochemical consequences of woody encroachment in this region, however, are poorly understood. To assess these matters we measured plant and soil δ¹³C, soil organic C and soil N content from grasslands and two important woody patch types (mature Quercus virginiana clusters and Juniperus ashei woodlands) in this region. Soil δ¹³C values showed that relative productivity of C₃ species has increased in grassland and both woody habitats in recent times. δ¹³C of SOC in grasslands and Q. virginiana clusters increased with depth from the litter layer to 30 cm (grasslands =?21 to ?13‰Q. virginiana clusters =?27 to ?17‰) and were significantly different between habitats at all depths, indicating that Q. virginiana has been a long‐term component of the landscape. In J. ashei woodlands, soil δ¹³C values (at 20–30 cm depth) near the woodland edge (‐13‰) converged with those of an adjacent grassland (‐13‰) while those from the woodland interior (‐15‰) remained distinct, indicating that the woodland has been present for many years but has recently expanded. Concentrations and densities of SOC and total N were generally greater in woody patches than in grasslands. However, differences in the amount of SOC and N stored beneath the two woody patch types indicates that C and N sequestration potentials are species dependent.     

Woodland dynamics under the influence of elephants and fire in Northern Botswana

Sustained elephant browsing and intense burning could result in the loss of woodlands under conditions where elephant densities are high, such as in northern Botswana. Three woodland types dominated by Acacia erioloba, Baikiaea plurijuga and Colophospermum mopane were monitored in plots and contemporary recruitment rates of woody plants were compared with the associated local elephant densities and fire occurrences. Woodland types differed with respect to structure, extent of elephant damage and the occurrence of fire. Canonical correlations indicated that high extent of fire damage and high elephant densities did not covary within the woodland types investigated. Low tree densities in some woodland types were associated with high elephant densities and new elephant damage to plants increased with high elephant densities during the dry season. Plots with an apparent high fire frequency had lower tree densities and higher cover abundance of shrubs and seedlings.The annual rates of tree recruitment/loss in each woodland type were estimated through a model based on observed seedling recruitment, mortality and reversal to lower height classes due to combinations of fire occurrence and elephant browsing. The model suggested that elephants induce tree loss in woodlands dominated by plant species which are principal food sources. Fire however, seems to have a widespread effect across woodlands which could result in extensive tree loss.     

Habitat selection and ranges of tolerance: how do species differ beyond critical thresholds?

Sensitivity to habitat fragmentation often has been examined in terms of thresholds in landscape composition at which a species is likely to occur. Observed thresholds often have been low or absent, however, leaving much unexplained about habitat selection beyond initial thresholds of occurrence, even for species with strong habitat preferences. We examined responses to varying amounts of tree cover, a widely influential measure of habitat loss, for 40 woodland bird species in a mixed woodland/grassland landscape in eastern North Dakota, USA. We used LOESS smoothing to describe incidence for each species at three scales: within 200, 400, and 1200 m around sample locations. For the 200‐m scale, we also calculated the most‐preferred range of tree cover (within which at least half of observations were predicted to occur) for each species. Only 10 of 40 species had occurrence thresholds greater than about 10% tree cover. After initial occurrence, species showed three general patterns: some increased monotonically with tree cover; some increased up to an asymptote; some peaked at intermediate amounts of tree cover and then declined. These patterns approximate selection for interior woodlands and for edge‐rich environments, but incidence plots provide greater detail in landscape‐scale selection than do those categories. For most species, patterns persisted at larger scales, but for some, larger scales had distinctly different patterns than local scales. Preferred ranges of tree cover varied from <20% tree cover (common grackle, Quiscalus quiscula) to >60% (veery, Catharus fuscescens). We conclude that incidence patterns provide more information on habitat selection than do threshold measures for most species: in particular, they differentiate species preferring concentrated woodlands from those preferring mixed landscapes, and they show contrasting degrees of selectiveness. [Correction added on 16 October 2012, after first online publication: the Abstract section has been reworded].     

Vegetation and songbird response to land abandonment: from landscape to census plot

While intensification of human activities and its ecological effects in many natural areas have recently received much attention, land abandonment in marginal areas is still the largely ignored side of a process rooted in the same socioeconomic context. Decreasing human impact in marginal rural areas often triggers a recovery of seminatural vegetation. Over a period of 25 years, we studied the changes in landscape and vegetation structure that followed land abandonment in a traditional Mediterranean mosaic of crops, grasslands, shrublands and woodlands, and assessed their effects on songbird occurrence and distribution. We combined an analysis of vegetation changes based on aerial photo interpretation with an analysis of bird censuses from 1978, 1992 and 2003 at two spatial scales: landscape and census plot (respectively 2800 and 3 ha). The perceived temporal changes in the vegetation were scale dependent. At the landscape scale, open habitats tended to disappear and woodlands matured. The contrasts in vegetation structure that defined habitat patches at the onset of the study tended to disappear. There was an overall shift of the bird community in favour of woodland species. At the scale of the census plot, however, the colonization by woody vegetation of patches formerly characterized by a homogeneous grass cover increased the local diversity of the vegetation, at least temporarily. Of seven species dependent on open habitats, the occurrence rate of five species significantly decreased, whereas it increased for two species: woodlark (Lulula arborea) and melodious warbler (Hippolais polyglotta). This increase was linked to the transitional increase in local vegetation diversity. In patches originally dominated by woodlands, local vegetation diversity decreased as woody vegetation expanded into clearings. The occurrence rate significantly increased for seven species relying on closed woodlands, while it decreased for two woodland species. As most species of high conservation profile in the Mediterranean are tied to open or to heterogeneous transitional habitats, these trends raise questions concerning their persistence in the future.