Border and ecotone detection by vegetation composition along forest‐savanna transects in Ivory Coast

Abstract. Question: How do properties of different vegetation components vary along ecotones of semi‐deciduous forest islands, and can the depth of edge influence (DEI) of the components be detected using a novel combination of analyses? Location: Comoé National Park (CNP), NE Ivory Coast. Methods: Along eight transects at semi‐deciduous forest islands tree individuals > 20 cm DBH were mapped. At one transect, tree and shrub individuals down to 1 cm DBH were measured and cover of species was estimated. Split moving window dissimilarity analysis (SMWDA) and moving window regression analysis (MWRA) were combined to detect statistical significance of borders in multivariate vegetation data along continuous transects, to determine the width of associated ecotones, and, thus, the DEI towards the forest interior. Results: For trees > 20 cm DBH, a distinct boundary formation was detected, dominated by the semi‐fire resistant tree species Anogeissus leiocarpus. The median of DEI towards the forest interior was 55 m. Ecotone detection with all species present revealed an interlocked sequence of ecotones for grasses, herbs, woody climbers, shrubs and trees, with each of these ecotones being narrower than the overall ecotone. DEI ranged from 10 m for grasses up to 120 m for trees and shrubs. Conclusions: The coherent set of analyses applied proved to be an objective method for detecting borders and the width of associated ecotones. The patterns found may be explained by successional processes at the forest‐savanna border. The DEI measured for the forest islands in the nearly undisturbed semi‐natural system of the CNP is of relevance to concepts of core‐area analysis and the protection of forest interior species in semi‐deciduous forests in tropical West Africa.     

Core area analysis at semi-deciduous forest islands in the Comoé National Park,NE Ivory Coast

For the protection of forest-interior species in both natural forest islands and anthropogenic forest fragments knowledge on the size of forest-core areas is a central issue. In an intact mosaic of semi-deciduous forests and savanna in the Comoé National Park 31 forest islands were selected (2.1–146.1 ha). Values for the depth-of-edge influence (DEI) of the study area recently published range from 0 m up to nearly 150 m. Thus, core-area analysis was carried out for this range in 5 m steps. For a DEI of 55 m—e.g. computed for tree-species composition of large trees—half of the total forest area can be considered as core area, but only 9 of the studied forest islands still contained a relative core area (rCA) of more than 50%. From non-linear regression it was estimated that for a DEI of 55 m an rCA of 50% can be expected for forest islands with a size of 36.6 ± 7.6 ha. This value increased exponentially with increasing DEI. The GIS-based core-area analysis presented in this paper proved to be suitable to give a well interpretable overview on rCA with respect to varying DEI, and we recommend to incorporate this type of analysis in existing GIS-tools. As the presented study is the first sound core area analysis at forest islands in West Africa, data contribute to a better understanding of this field of ecology that is of high relevance for planners and decision makers to protect biodiversity.     

Sharp transitions in microclimatic conditions between savanna and forest in New Caledonia: Insights into the vulnerability of forest edges to fire

Fires are one of the main causes of forest loss in the tropics. Understanding the dynamic edge effects is critical for managing fires and protecting forests. We measured and analysed trends in microclimatic conditions (air temperature, relative humidity and vapour pressure deficit) over 7 months along three transects extending from core savanna areas to core forest areas. We tested two hypotheses: (i) that the forest edge is subject to microclimatic edge effects, and (ii) that the depth of these edge effects increases during dry periods. Sharp changes in each microclimatic variable were consistently observed between savanna and forest throughout the study period. Microclimatic transitions took place within 5 m outside the forest boundary. Drought levels increased homogenously throughout the forest and were not disproportionately severe in the vicinity of the forest edge. We suggest that these results were related to the fact that the studied period was abnormally humid due to a La Niña episode, and that under such conditions the vulnerability of the forest edge to savanna fires is relatively low. Relatively wet conditions in the savanna close to the forest edge may promote forest expansion by limiting fire spread. Prescribed fires during humid years could reduce fuel loads in savanna without affecting the forest edge, which would prevent fires during the dry years associated with El Niño episodes from having severe impacts.     

Edge influence on plant litter biomass in forest and savanna in the Brazilian cerrado

Edge influence, characterized by differences in ecosystem characteristics between the edge and the interior of remnants in fragmented landscapes, affects a variety of organisms and ecosystem processes. An important feature that may be affected by edges is the amount of plant litter, which provides important habitat for a large variety of organisms and influences ecological processes such as fire dynamics. We studied edge influence on plant litter and fine woody debris in the cerrado of São Paulo state, south‐eastern Brazil. We collected, sorted, dried and weighed plant litter along 180 m‐long transects perpendicular to three savanna and eleven forest edges adjacent to different anthropogenic land uses, with four to five transect per edge. There tended to be less biomass of the finer portions of fine woody debris at both savanna and forest edges. Graminoid litter at savanna edges was greater than in the corresponding interior areas, whereas other litter portions were either unaffected by edges or did not show consistent patterns in either savanna or forest. Edge influence was usually restricted to the first 20 m from the edge, was not influenced by edge characteristics and exhibited no clear differences between savanna and forest areas. Several mechanisms may have led to the variable patterns observed including variation in the plant community, plant architecture, and invasive species. The edge‐related variation in plant litter may putatively lead to, for example, increased fire frequency and intensity at the savanna edges and altered trophic dynamics at forest edges; the mechanisms and consequences of this edge influence should be addressed in future studies.     

Spatial patterns in species–area relationships and species distribution in a West African forest–savanna mosaic

Aim To investigate the relationship between the slope z of the species–area relationship (SAR) and the intensity of spatial patterns in species number and dissimilarity for woody plants with different modes of seed dispersal. According to island theory we expect, for any given archipelago, steeper slopes and more pronounced spatial patterns for groups of less dispersive species. Location Ivory Coast, West Africa. Methods In a West African forest–savanna mosaic we collected presence–absence data for woody plant species in 49 forest islands. The parameters of the SARs were fitted by nonlinear regressions and then compared for plant species aggregated according to their mode of seed dispersal. We used the Mantel test to calculate the intensity of spatial patterns in species number, i.e. residual deviation from SAR, and species dissimilarity. Results The z‐value for bird‐dispersed species was lower (0.11) than that for wind‐dispersed species (0.27), with mammal‐dispersed species taking an intermediate value (0.16). This result suggests that, as a group, bird‐dispersed species are better colonizers. The spatial pattern in species number as well as species similarity was more pronounced for bird‐ compared with wind‐dispersed species. Main conclusions The standard interpretation of the theory of island biogeography claims that shallow slopes in the SAR imply low isolation of islands, i.e. good dispersal abilities of species. The results of our study appear to contradict this statement. The contradiction can eventually be resolved by a more detailed account of the colonization process, i.e. by distinguishing between dispersal and consecutive establishment of populations.     

Diversity and distribution of murid rodent populations between forest and derived savanna sites within south western Nigeria

Southward encroachment of the derived savanna zone with its attendant denudation of the rainforest in Nigeria adversely affects the diversity and distribution of mammals, particularly murid rodents. This study was carried out to establish the identities of murid rodent populations and to compare their diversity between forest and derived savanna sites within south western Nigeria. Identification of captured specimens was carried out by multivariate statistics of body and skull measurements, and also by analysis of certain discrete characters. Six species were identified: Praomys tullbergi, Hylomyscus stella, Mastomys natalensis, Arvicanthis rufinus, Lemniscomys striatus and Lophuromys sikapusi. Forest sites possessed a higher Shannon’s diversity index (1.685) than the derived savanna sites (0.978), containing a higher number of taxa and larger abundance of murid rodents. This underscores the negative impact that encroachment of the derived savanna has upon diversity of animals in areas that were once forest.     

Early gap successional pathways in a Fagus-Acer forest preserve: pattern and determinants

Abstract. Many theories of forest succession imply that terrestrial plant community composition within a region tends to converge toward a climax community. That is, given similar climatic and edaphic conditions, succession at different sites within an area will lead to comparable species compositions, a pattern referred to as successional convergence. In this study, we examine changes in plant composition within forest canopy gaps over a 17-yr period to identify potential patterns of successional convergence and to ascertain the factors controlling the successional pathway. To do so, we: (1) sampled 36 forest canopy gaps in Hueston Woods Nature Preserve in 1977, 1981, 1985, 1989 and 1993, (2) evaluated changes in the similarity of gap composition over this period, and (3) examined gap composition in each year as a function of variables describing gap habitat, seed source proximity, and disturbance history. Results indicated an initial pattern of successional divergence, with gaps exhibiting increased dissimilarity over the first 10–12 years of succession. We attribute this initial period of divergence to the effects of differential seed inputs from edge individuals and heterogeneity of available light due to differences in gap size. Recent surveys, however, indicated that gap composition has become more similar as competition within gaps has become more intense. In these samples, gap composition is closely linked to site conditions, including slope, soil conditions, and site exposure. Finally, while these patterns may suggest equilibrium-oriented dynamics, non-equilibrium processes such as repeat disturbances are also evident at Hueston Woods and will likely play an important role in determining future successional pathways.     

Effects of fragmentation on parasite burden (nematodes) of generalist and specialist small mammal species in secondary forest fragments of the coastal Atlantic Forest,Brazil

Parasites are considered to play an important role in the regulation of wild animal populations. We investigated parasite burden of gastrointestinal nematodes and body condition in specialist and generalist small mammal species in secondary forest fragments in the highly endangered coastal Atlantic Forest. We hypothesized that body condition decreases with increasing parasite load and that parasite burden increases with increasing fragmentation in specialist species but not in generalist species as a consequence of differing responses to fragmentation effects. Investigated species were Akodon montensis, Oligoryzomys nigripes, and Delomys sublineatus (rodents) and the marsupials Marmosops incanus and Gracilinanus microtarsus. Prevalence of parasites was high in all species except for the arboreal G. microtarsus, presumably because of decreased infection probability. No correlation was found between body condition and parasite load in any of the species. Contrary to our expectations, body condition of the specialists D. sublineatus and M. incanus increased in both species with increasing fragmentation. In D. sublineatus, parasite burden increased and body condition decreased in fragments with relatively high density probably due to increased contact rates and facilitation of infection with nematodes. In all generalist species, low or no correlation between parasite burden and fragmentation was detected, suggesting little effect of fragmentation on population health.     

Effects of fragmentation on forest structure and litter dynamics in Atlantic rainforest in Pernambuco, Brazil

This study evaluates the effects of fragmentation on the spatial and temporal dynamics of small litterfall production in Atlantic rainforest in Pernambuco State, Brazil. Litterfall was collected for 24 months at two 0.2 ha sites, located in the forest edge zone and the forest interior, within a rainforest patch of about 300 ha. Structural parameters of both forest sites were recorded. Litter was sorted into six fractions (foliage, twigs, buds/flowers, fruits/seeds, peduncles, rest), dried and weighed. The interior forest plot contained 314 live trees with a dbh 5 cm and a stand basal area of 41.8±8.7 m², whereas the forest edge contained 211 live trees and a stand basal area of 23.4±3.6 m². Total small litterfall was extraordinarily high and totalled 12.62±4.73 t ha⁻¹ yr⁻¹ in interior forest and 14.74±2.78 t ha⁻¹ yr⁻¹ in forest edge. High litterfall rates are probably due to a pronounced periodicity, edge effects alter litterfall strongly.     

Montane forest ecotones moved downslope in northeastern USA in spite of warming between 1984 and 2011

Ecotones are transition zones that form, in forests, where distinct forest types meet across a climatic gradient. In mountains, ecotones are compressed and act as potential harbingers of species shifts that accompany climate change. As the climate warms in New England, USA, high‐elevation boreal forests are expected to recede upslope, with northern hardwood species moving up behind. Yet recent empirical studies present conflicting findings on this dynamic, reporting both rapid upward ecotonal shifts and concurrent increases in boreal species within the region. These discrepancies may result from the limited spatial extent of observations. We developed a method to model and map the montane forest ecotone using Landsat imagery to observe change at scales not possible for plot‐based studies, covering mountain peaks over 39 000 km². Our results show that ecotones shifted downward or stayed stable on most mountains between 1991 and 2010, but also shifted upward in some cases (13–15% slopes). On average, upper ecotone boundaries moved down ?1.5 m yr^?1 in the Green Mountains, VT, and ?1.3 m yr^?1 in the White Mountains, NH. These changes agree with remeasured forest inventory data from Hubbard Brook Experimental Forest, NH, and suggest that processes of boreal forest recovery from prior red spruce decline, or human land use and disturbance, may swamp out any signal of climate‐mediated migration in this ecosystem. This approach represents a powerful framework for evaluating similar ecotonal dynamics in other mountainous regions of the globe.     

Long‐term variability and rainfall control of savanna fire regimes in equatorial East Africa

Fires burning the vast grasslands and savannas of Africa significantly influence the global carbon cycle. Projecting the impacts of future climate change on fire‐mediated biogeochemical processes in these dry tropical ecosystems requires understanding of how various climate factors influence regional fire regimes. To examine climate–vegetation–fire linkages in dry savanna, we conducted macroscopic and microscopic charcoal analysis on the sediments of the past 25 000 years from Lake Challa, a deep crater lake in equatorial East Africa. The charcoal‐inferred shifts in local and regional fire regimes were compared with previously published reconstructions of temperature, rainfall, seasonal drought severity, and vegetation dynamics to evaluate millennial‐scale drivers of fire occurrence. Our charcoal data indicate that fire in the dry lowland savanna of southeastern Kenya was not fuel‐limited during the Last Glacial Maximum (LGM) and Late Glacial, in contrast to many other regions throughout the world. Fire activity remained high at Lake Challa probably because the relatively high mean‐annual temperature (~22 °C) allowed productive C₄ grasses with high water‐use efficiency to dominate the landscape. From the LGM through the middle Holocene, the relative importance of savanna burning in the region varied primarily in response to changes in rainfall and dry‐season length, which were controlled by orbital insolation forcing of tropical monsoon dynamics. The fuel limitation that characterizes the region's fire regime today appears to have begun around 5000–6000 years ago, when warmer interglacial conditions coincided with prolonged seasonal drought. Thus, insolation‐driven variation in the amount and seasonality of rainfall during the past 25 000 years altered the immediate controls on fire occurrence in the grass‐dominated savannas of eastern equatorial Africa. These results show that climatic impacts on dry‐savanna burning are heterogeneous through time, with important implications for efforts to anticipate future shifts in fire‐mediated ecosystem processes.     

Seed-deposition and recruitment patterns of Clusia species in a disturbed tropical montane forest in Bolivia

Spatial patterns of seed dispersal and recruitment of fleshy-fruited plants in tropical forests are supposed to be driven by the activity of animal seed dispersers, but the spatial patterns of seed dispersal, seedlings and saplings have rarely been analyzed simultaneously. We studied seed deposition and recruitment patterns of three Clusia species in a tropical montane forest of the Bolivian Andes and tested whether these patterns changed between habitat types (forest edge vs. forest interior), distance to the fruiting tree and consecutive recruitment stages of the seedlings. We recorded the number of seeds deposited in seed traps to assess the local seed-deposition pattern and the abundance and distribution of seedlings and saplings to evaluate the spatial pattern of recruitment. More seeds were removed and deposited at the forest edge than in the interior. The number of deposited seeds decreased with distance from the fruiting tree and was spatially clustered in both habitat types. The density of 1-yr-old seedlings and saplings was higher at forest edges, whereas the density of 2-yr-old seedlings was similar in both habitat types. While seedlings were almost randomly distributed, seeds and saplings were spatially clustered in both habitat types. Our findings demonstrate systematic changes in spatial patterns of recruits across the plant regeneration cycle and suggest that the differential effects of biotic and abiotic factors determine plant recruitment at the edges and in the interior of tropical montane forests. These differences in the spatial distribution of individuals across recruitment stages may have strong effects on plant community dynamics and influence plant species coexistence in disturbed tropical forests.     

Effects of habitat configuration and quality on species richness and distribution in fragmented forest patches near Rome

Question: How important are habitat configuration, quality, history and anthropic disturbance in determining nemoral plant species richness and distribution of fragmented forest patches in a Mediterranean region? Location: Agricultural landscape north of Rome, Italy. Methods: Sixty‐nine woodland patches, identified through a stratified random sampling, were sampled for nemoral plant species. The homogeneity of woodlands was tested through a hierarchical classification of the floristic data and a Mann‐Whitney test of dependent and independent variables. The importance of habitat configuration (area, isolation, shape), quality (soil properties, forest structure, anthropic disturbance) and history (age of woodland) in determining species richness was estimated through a Poisson regression model. Presence‐absence of each species was analysed by logistic regression. Differences among plant life‐trait types (life span, dispersal mode, habitat preference) were analysed by comparing their median β‐values through ANOVA models. Results: Through hierarchical classification, two woodland types were identified that differed in species composition, habitat quality and spatial configuration. Poisson regression showed that habitat configuration and history influenced species richness. Multiple logistic regression resulted in significant fits for 88 species/variable combinations: 38 are habitat quality variables, 25 are habitat configuration variables, and 13 are anthropic factors. Dispersal strategies varied significantly with respect to area, isolation and age, while generalist and specialist species differed according to age of the woodland. Conclusion: Our results show that habitat history and configuration are the key factors determining species richness of woodland. Together with habitat configuration, habitat quality (mainly soil acidity) appeared to influence species composition.     

Arthropods on plants in a fragmented Neotropical dry forest: A functional analysis of area loss and edge effects

Loss and fragmentation of natural ecosystems are widely recognized as the most important threats to biodiversity conservation, with Neotropical dry forests among the most endangered ecosystems. Area and edge effects are major factors in fragmented landscapes. Here, we examine area and edge effects and their interaction, on ensembles of arthropods associated to native vegetation in a fragmented Chaco Serrano forest. We analyzed family richness and community composition of herbivores, predators, and parasitoids on three native plant species in 12 fragments of varying size and at edge/interior positions. We also looked for indicator families by using Indicator Species Analysis. Loss of family richness with the reduction of forest fragment area was observed for the three functional groups, with similar magnitude. Herbivores were richer at the edges without interaction between edge and area effects, whereas predators were not affected by edge/interior position and parasitoid richness showed an interaction between area and position, with a steeper area slope at the edges. Family composition of herbivore, predator, and parasitoid assemblages was also affected by forest area and/or edge/interior situation. We found three indicator families for large remnants and five for edges. Our results support the key role of forest area for conservation of arthropods taxonomic and functional diversity in a highly threatened region, and emphasize the need to understand the interactions between area and edge effects on such diversity.