Changes in Habitat Use at Rain Forest Edges Through Succession: a Case Study of Understory Birds in the Brazilian Amazon

Primary tropical rain forests are being rapidly perforated with new edges via roads, logging, and pastures, and vast areas of secondary forest accumulate following abandonment of agricultural lands. To determine how insectivorous Amazonian understory birds respond to edges between primary rain forest and three age classes of secondary forest, we radio‐tracked two woodcreepers (Glyphorynchus spirurus, N = 17; Xiphorhynchus pardalotus, N = 18) and a terrestrial antthrush (Formicarius colma, N = 19). We modeled species‐specific response to distance to forest edge (a continuous variable) based on observations at varying distances from the primary‐secondary forest interface. All species avoided 8–14‐yr‐old secondary forest. Glyphorynchus spirurus and F. colma mostly remained within primary forest <100 m from the young edge. Young F. colma rarely penetrated >100 m into secondary forest 27–31 yr old. Young Formicarius colma and most G. spirurus showed a unimodal response to 8–14‐yr‐old secondary forest, with relative activity concentrated just inside primary forest. After land abandonment, G. spirurus was the first to recover to the point where there was no detectable edge response (after 11–14 yr), whereas X. pardalotus was intermediate (15–20 yr), and F. colma last (28–30 yr +). Given the relatively quick recovery by our woodcreeper species, new legislation on protection of secondary forests > 20‐yr old in Brazil's Pará state may represent a new opportunity for conservation and management; however, secondary forest must mature to at least 30 yr before the full compliment of rain forest‐dependent species can use secondary forest without adverse edge effects.     

Distributions of lizard species across edges delimiting open‐forest and sand‐mined areas

Abstract The distributions of lizards across habitat edges delimiting open‐forest and regenerating sand‐mined areas as a function of distance from the edge were studied at Tomago, New South Wales, Australia. Pitfall‐trapping was used to survey lizards across the northern edges of four forest fragments, to determine if lizards displayed characteristic responses across the edge, and whether these could be explained by the different habitat conditions. At each site, 11 equally spaced drift fences (each parallel to the edge) were arranged in a transect running perpendicular to the edge, and stretching 50 m into each habitat type. Captures of Amphibolurus muricatus (Agamidae) decreased substantially across the edge from the mine‐path to the forest so that it was identified as a mine‐path specialist lizard species. Captures of two skink species decreased across the mine‐path before reaching the edge, and were not caught (Ctenotus taeniolatus) or were seldom caught (Ctenotus robustus) in the forest, so they were identified as mine‐path specialist, edge avoiding, lizard species. Captures of Lampropholis delicata (Scincidae) increased across edges into the forest, consistent with the expectation for a forest specialist. Regression analyses indicated the responses to edges of three lizard species (A. muricatus, C. robustus and C. taeniolatus) were negatively correlated with canopy cover (probably due to its influence on temperature, as captures of A. muricatus and C. robustus were also correlated positively with mean daily temperature). In addition, the response of C. robustus correlated negatively with a vegetation factor (dense, even vegetation in the first 50 cm from ground level). The response of L. delicata correlated positively with understorey height. We have identified edge response strategies for four species of lizards across edges delimiting temperate open‐forest and mined areas, and identified habitat and microclimate variables that may have driven these responses.     

Edge effects across boundaries between natural and restored jarrah (Eucalyptus marginata) forests in south‐western Australia

Edge effects are a widespread and ubiquitous ecological phenomenon, yet they remain poorly studied across edges between restored and natural forests. To address this lack of knowledge, we studied vertebrate communities across edges between 3‐year old restored mine‐pits and adjacent unmined forest in the jarrah (Eucalyptus marginata) forest of south‐western Australia. We found that mammal communities showed no edge response but reptile communities did. Overall reptile abundance and Morethia obscura abundance were higher in unmined forest along edges, Egernia napoleonis abundance was lower in unmined forest along edges, while Pogona minor abundance was lower in restored mine‐pits along edges. Predictive models were unable to predict species edge responses, due to the lack of knowledge of the ecology of jarrah forest reptiles, but proved useful in identifying potential ecological mechanisms behind observed edge responses and suggested that potential mechanisms were likely different for each species. Our study is the first to show edge responses in both habitats forming the edge between restored and natural forests, emphasizing the importance of studying both habitats forming the edge. Our results also suggest that, despite being poorly studied, edge responses are common across edges between restored and natural forest and result from a variety of ecological mechanisms. An increased understanding of the ecological mechanisms driving edge responses across edges between restored and natural forests will improve our ability to integrate restored areas into cross‐landscape management and, ultimately, improve our ability to manage landscapes for biodiversity conservation.     

Edge effects as the principal cause of area effects on birds in fragmented secondary forest

Bird communities in tropical forests are strongly affected by both patch area and habitat edges. The fact that both effects are intrinsically confounded in space raises questions about how these two widely reported ecological patterns interact, and whether they are independent or simply different spatial manifestations of the same phenomenon. Moreover, do small patches of secondary forest, in landscapes where the most sensitive species have gone locally extinct, exhibit similar patterns to those previously observed in fragmented and continuous primary forests? We addressed these questions by testing edge‐related differences in vegetation structure and bird community composition at 31 sites in fragmented and continuous landscapes in the imperilled Atlantic forest of Brazil. Over a two‐year period, birds were captured with mist nets to a standardized effort of 680 net‐hours at each site (～22 000 net‐hours resulting in 3381 captures from 114 species). We found that the bird community in patches of secondary forest was degraded in species composition compared to primary continuous forest, but still exhibited a strong response to edge effects. In fragmented secondary forests, edge and area effects also interacted, such that the magnitude of edge to interior differences on bird community composition declined markedly with patch size. The change in bird species composition between forest interiors and edges was similar to the change in community composition between large and small patches (because species had congruent responses to edge and area), but after controlling for edge effects community composition was no longer affected by patch area. Our results show that although secondary forests hold an impoverished bird community, ecological patterns such as area and edge effects are similar to those reported for primary forests. Our data provide further evidence that edge effects are the main drivers of area effects in fragmented landscapes.     

Regeneration pattern of primary forest species across forest-field gradients in the subtropical Mountains of Southwestern China

Evergreen broad-leaved forest is now gradually degraded and fragmented, and there is an increase in the amount of habitat edges as a result of long-term human activity. However, the role of edges in the regeneration of primary forest species is poorly understood. After 20 years of the edge creation, we analyzed primary forest species distribution and abundance, and changes in floristic composition, vegetation structure across forest-field gradients in Ailao Mountain, SW China. Our results revealed that there was a higher abundance and richness of primary species, late secondary species and thorny lianas at the distances 0–50 m than at the distances more than 50 m from the edge into the forest exterior. At the distances >50 m, no individuals of dominant canopy trees Lithocarpus xylocarpus, Castanopsis wattii, and L. jingdongensis were found, whereas the abundance of early pioneer shrub species and herbaceous cover was significantly greater. The richness of primary species showed a decrease with increasing distances from the forest edge to the exterior, particularly of medium-seeded primary species showing a drastic decrease. Moreover, no large-seeded primary species occurred at the distances >60 m. This study indicates that the forest edge as a buffer zone may be in favor of primary species regeneration. A dense shrub and herb layer, and seed dispersal may be the major factors limiting the forest regeneration farther from the forest edge. Therefore, to facilitate forest recovery processes, management should give priority to the protection of buffer zones of this forest edge.     

Diameter Growth of Juvenile Trees after Gap Formation in a Bolivian Rain Forest: Responses are Strongly Species‐specific and Size‐dependent

We evaluated growth responses to gap formation for juvenile individuals of three canopy rain forest species: Peltogyne cf. heterophylla, Clarisia racemosa and Cedrelinga catenaeformis. Gaps were formed during selective logging operations 7 yr before sampling in a Bolivian rain forest. We collected wood samples for tree‐ring analyses at different distances to the stump (<10, 10–40 and >40 m) and from trees with different diameters (5–30 cm diameter at breast height [dbh]). Tree‐rings width was measured in at least two radii and converted to average diameter growth. Changes in 7‐yr median diameter growth before and after selective logging were analyzed. Diameter growth rates significantly increased by 0.7–0.8 mm/yr after gap formation for P. heterophylla and C. catenaeformis, but not for C. racemosa. We applied a multiple regression analysis to explain variation in growth responses of P. heterophylla and C. catenaeformis by distance to logging gap and tree size. For P. heterophylla we found that growth increase occurring close to logging gaps was strongest for large juvenile trees (20–25 cm dbh) and almost absent in small juveniles. For C. catenaeformis, variation in growth responses was not related to tree size or distance to gaps. Our results show that growth responses to gap formation strongly differ across species and tree sizes. This finding calls for caution in the interpretation of growth releases in tree‐ring series, as gap formation does not necessarily invoke growth responses and if such growth responses occur, their strength is species‐ and size specific.     

Understorey vegetation development in North Finnish Picea forests after disturbance: re‐analysis of Sirén's data

Abstract. Sirén (1955) studied understorey species composition, tree stand properties and humus‐layer thickness in 64 unlogged forest stands on topographically and pedologically comparable sites. The stands were of even age (6 – 300 yr), stocked with the first or second tree generation after wildfire. The view of Sirén and several authors after him, that the vegetation of old‐growth boreal Picea forests is homogeneous on a broad scale, was examined by applying, in parallel, the partial variants of two ordination methods (DCA and PCA) to Sirén's vegetation data. Two main vegetation gradients were found: a major gradient running from recently burnt plots with prominence of pioneer species to plots with stand age > 100 yr, a well stocked tree layer and a thick humus layer, dominance of feather‐mosses and ample occurrence of shade‐tolerant as well as light‐preferring vascular plant species, and a second gradient along which first‐ and second‐generation plots segregate. The more prominent element of Betula trees in first‐ than in second‐generation stands < 100 yr contributed to the latter. A minor third gradient related to humus‐layer thickness was recovered by partial DCA only. The main vegetation gradient reappeared in separate ordinations of data from 47 mature forest stands (> 100 yr), but without being correlated with forest age. Variation among mature‐forest stands in the importance of pioneer species is considered mainly to be brought about by fine‐scale disturbance processes such as tree uprooting. Increasing importance of factors operating on within‐stand scales [development of a varied gap structure and stronger gradients in tree influence (radiation at ground level), soil moisture, soil depth and nutrient availability] with time is also reflected in the second and third mature‐forest ordination axes. Possible implications of the results for conservation of biological diversity and monitoring of changes in boreal forests are discussed.     

Contrasting Demographic Structure of Short‐ and Long‐lived Pioneer Tree Species on Amazonian Forest Edges

Although tropical forests have been rapidly converted into human‐modified landscapes, tree species response to forest edges remains poorly examined. In this study, we addressed four pioneer tree species to document demographic shifts experienced by this key ecological group and make inferences about pioneer response to forest edges. All individuals with dbh ≥ 1 cm of two short‐lived (Bellucia grossularioides and Cecropia sciadophylla) and two long‐lived species (Goupia glabra and Laetia procera) were sampled in 20 1‐ha forest edge plots and 20 1‐ha forest interior plots in Oiapoque and Manaus, Northeast and Central Amazon, respectively. As expected, pioneer stem density with dbh ≥ 1 cm increased by around 10–17‐fold along forest edges regardless of species, lifespan, and study site. Edge populations of long‐lived pioneers presented 84–94 percent of their individuals in sapling/subadult size classes, whereas edge populations of short‐lived pioneers showed 56–97 percent of their individuals in adult size classes. These demographic biases were associated with negative and positive net adult recruitment of long‐ and short‐lived pioneers, respectively. Our population‐level analyses support three general statements: (1) native pioneer tree species proliferate along forest edges (i.e., increased density), at least in terms of non‐reproductive individuals; (2) pioneer response to edge establishment is not homogeneous as species differ in terms of demographic structure and net adult recruitment; and (3) some pioneer species, particularly long‐lived ones, may experience population decline due to adult sensitivity to edge‐affected habitats.     

Limited Edge Effects Along a Burned‐Unburned Bornean Forest Boundary Seven Years after Disturbance

Large parts of the everwet tropics have been burned, leaving many unburned–burned forest edges. Here we studied a Bornean forest edge to determine: (1) how unburned and burned forest differ in vegetation structure, diversity, composition and plant functional traits 7 yr after fire, and (2) if these variables showed significant edge effects. Environmental and inventory data from 120 plots (0.01 ha each), covering both sides of a ～1.3 km forest boundary were sampled. Differences in vegetation structure, diversity, composition and plant functional traits were analyzed in relation to disturbance type (Mann–Whitney tests) and edge distance (partial correlation analysis that controlled for confounding effects of elevation, slope and fire intensity). Seven years after fire, burned forest differed significantly from unburned forest in most measured variables while few significant edge effects were detected, i.e., there existed a sharp delimitation between the two forest types. The regeneration of the burned forest depended almost entirely on in situ recruitment with little input of late successional species from the neighboring old growth forest. On the other hand, old growth forest showed few signs of edge degradation. A possible explanation for these results might be related to the absence of a mast fruiting event during these first 7 yr of forest recovery, resulting in low levels of late successional species seed input into the burned forest, combined with the quick development of a closed canopy in the burned forest by early successional species that shielded the unburned forest from adverse edge effects.     

Butterfly Response to Microclimatic Conditions Following Ponderosa Pine Restoration

Efforts to restore ponderosa pine ecosystems to open, park‐like conditions that predominated prior to European‐American settlement result in altered stand structure and increased landscape heterogeneity, potentially altering habitat suitability for invertebrates and other forest organisms. We examined the responses of two butterfly species, Colias eurytheme and Neophasia menapia, to microclimatic changes at structural edges created by experimental restoration treatments in northern Arizona. We monitored microclimate, including air temperature, light intensity, and vapor pressure deficit (VPD), on several mornings during butterfly releases. We placed adult butterflies at east‐ and west‐facing edges approximately one half‐hour before dawn to determine their behavioral response to microclimatic differences between east‐ and west‐facing edges. After sunrise, all three microclimatic variables were higher at east‐facing edges, and the difference in microclimate between the two edge orientations increased through early morning. For both species, butterflies placed at east‐facing edges flew earlier than butterflies at west‐facing edges. Colias eurytheme, an open‐habitat species, tended to move toward the treated forest during initial flight, while movements of Neophasia menapia, a forest‐dwelling species, did not differ from random flight. Our results indicate that butterflies respond to microclimatic factors associated with restoration treatments, while responses to structural changes in habitat vary among species, based on habitat and food plant preferences. These changes in forest structure and microclimate may affect the distribution of many mobile invertebrates in forested landscapes undergoing restoration treatments.     

Forest edges have high conservation value for bird communities in mosaic landscapes

A major conservation challenge in mosaic landscapes is to understand how trait‐specific responses to habitat edges affect bird communities, including potential cascading effects on bird functions providing ecosystem services to forests, such as pest control. Here, we examined how bird species richness, abundance and community composition varied from interior forest habitats and their edges into adjacent open habitats, within a multi‐regional sampling scheme. We further analyzed variations in Conservation Value Index (CVI), Community Specialization Index (CSI) and functional traits across the forest‐edge‐open habitat gradient. Bird species richness, total abundance and CVI were significantly higher at forest edges while CSI peaked at interior open habitats, i.e., furthest from forest edge. In addition, there were important variations in trait‐ and species‐specific responses to forest edges among bird communities. Positive responses to forest edges were found for several forest bird species with unfavorable conservation status. These species were in general insectivores, understorey gleaners, cavity nesters and long‐distance migrants, all traits that displayed higher abundance at forest edges than in forest interiors or adjacent open habitats. Furthermore, consistently with predictions, negative edge effects were recorded in some forest specialist birds and in most open‐habitat birds, showing increasing densities from edges to interior habitats. We thus suggest that increasing landscape‐scale habitat complexity would be beneficial to declining species living in mosaic landscapes combining small woodlands and open habitats. Edge effects between forests and adjacent open habitats may also favor bird functional guilds providing valuable ecosystem services to forests in longstanding fragmented landscapes.     

Landscape‐level effects on aboveground biomass of tropical forests: A conceptual framework

Despite the general recognition that fragmentation can reduce forest biomass through edge effects, a systematic review of the literature does not reveal a clear role of edges in modulating biomass loss. Additionally, the edge effects appear to be constrained by matrix type, suggesting that landscape composition has an influence on biomass stocks. The lack of empirical evidence of pervasive edge‐related biomass losses across tropical forests highlights the necessity for a general framework linking landscape structure with aboveground biomass. Here, we propose a conceptual model in which landscape composition and configuration mediate the magnitude of edge effects and seed‐flux among forest patches, which ultimately has an influence on biomass. Our model hypothesizes that a rapid reduction of biomass can occur below a threshold of forest cover loss. Just below this threshold, we predict that changes in landscape configuration can strongly influence the patch's isolation, thus enhancing biomass loss. Moreover, we expect a synergism between landscape composition and patch attributes, where matrix type mediates the effects of edges on species decline, particularly for shade‐tolerant species. To test our conceptual framework, we propose a sampling protocol where the effects of edges, forest amount, forest isolation, fragment size, and matrix type on biomass stocks can be assessed both collectively and individually. The proposed model unifies the combined effects of landscape and patch structure on biomass into a single framework, providing a new set of main drivers of biomass loss in human‐modified landscapes. We argue that carbon trading agendas (e.g., REDD+) and carbon‐conservation initiatives must go beyond the effects of forest loss and edges on biomass, considering the whole set of effects on biomass related to changes in landscape composition and configuration.     

Edge Effects Act Differentially on Multiple Early Regeneration Stages of a Shade‐tolerant Tree Tapirira mexicana

Forest fragmentation is pervasive in tropical landscapes, and one pathway by which fragmentation may negatively impact populations is via edge effects. Early life‐stages are particularly important for species regeneration as they act as bottlenecks, but how edge effects may act differentially on different life‐stages is unknown. This study evaluated edge effects on multiple early life‐stages of a currently common animal‐dispersed, shade‐tolerant tree Tapirira mexicana (Anacardiaceae). The study was conducted in tropical premontane wet forest fragments in a highly deforested region of Costa Rica. The stages assessed were pre‐dispersal predation, primary dispersal, post‐dispersal predation, secondary dispersal, ex situ germination, in situ seed longevity, first and second year seedling abundance, second year seedling survivorship, and basal diameter growth. Results showed that impacts of edge effects were not equal across stages, but were limited to specific stages and times. One stage which may act as a bottleneck for species regeneration was pre‐dispersal predation. Over 60 percent of the seeds were predated by larvae, and predation was higher near the edge than interior habitat. Seeds lost viability within 10 d in the forest. Germination to first year seedling stage was also lower near edges, but such effect was eliminated within a year after that. Primary dispersal, seedling survivorship, and growth were not affected by proximity to edges, and both secondary dispersal and post‐dispersal predation were rare. This study demonstrates that current population abundance may not guarantee future species persistence and the importance of considering multiple life‐stages for a comprehensive assessment of forest fragmentation effects on species regeneration.     

Tree Regeneration and Understory Woody Plants Show Diverse Responses to Forest–Pasture Edges in Costa Rica

ABSTRACT Edge effects along tropical forest–pasture margins are thought to cause a shift toward early successional characteristics of the understory forest vegetation. We tested this idea by sampling vegetation at five forest sites in northeast Costa Rica each of which had edges that were established over 20 yr earlier. Four of these sites had been selectively logged. We sampled woody plants >0.2 and ≤1.3 m height in 54 m² within 0.2 ha plots at edges (N=14), and at 150 m (N=11) and 300 m from edges (N=9). Composition and diversity did not vary with edge distance. Abundance of tree regeneration, mainly of canopy and emergent species, increased at edges. Abundance of lianas and slow‐growing tree species did not differ significantly across the sampling locations. Weighted mean wood density varied little, with a reduction at edges for canopy species. Palms were less abundant at edges, but not less species rich. At edges, these plant assemblages maintain many of the characteristics of forest interior vegetation, though the changes observed may indicate ongoing functional change. Degradation of forest–pasture edges is not a universal feature of tropical forest fragmentation, and forests with high rates of natural turnover might have a high capacity to maintain themselves within forest edges alongside pasture.     

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.     

Secondary Succession and Indigenous Management in Semideciduous Forest Fallows of the Amazon Basin1

To the discussion on secondary succession in tropical forests, we bring data on three under‐addressed issues: understory as well as overstory changes, continuous as opposed to phase changes, and integration of forest succession with indigenous fallow management and plant uses. Changes in vegetation structure and species composition were analyzed in secondary forests following swidden agriculture in a semideciduous forest of Bolivian lowlands. Twenty‐eight fallows, stratified by four successional stages (early = 1–5 yr, intermediate = 6–10 yr, advanced = 12–20 yr, and older = 22–36 yr), and ten stands of mature forests were sampled. The overstory (plants ≥5 cm diameter at breast height [DBH]) was sampled using a 20 × 50 m plot and the understory (plants <5 cm DBH) in three nested 2 × 5 m subplots. Semistructured interviews provided information on fallow management. Canopy height, basal area, and liana density of the overstory increased with secondary forest age. The early stage had the lowest species density and diversity in the overstory, but the highest diversity in the understory. Species composition and abundance differentiated mature forests and early successional stage from other successional stages; however, species showed individualistic responses across the temporal gradient. A total of 123 of 280 species were useful with edible, medicinal, and construction plants being the most abundant for both over‐ and understories. Most of Los Gwarayo preferred mature forests for making new swidden, while fallows were valuable for crops, useful species, and regenerating timber species.     

Short‐term effect of selection cutting in boreal balsam fir forest on cerambycid and scolytid beetles

The ecology and management of boreal forest ecosystems are drawing greater attention worldwide as their importance is being increasingly recognized for carbon sequestration or for harbouring the world's largest remaining intact forests. Selection cuts have been introduced as a more socially acceptable silvicultural method to improve the maintenance of habitat structure and functions as they mimic aspects of boreal forest succession dynamics. Many studies have shown that selection cutting helps maintaining arthropod communities in mature forests, but few have examined the increased risks of damage by bark‐ and wood‐boring insects in boreal forests of eastern North America. We used multidirectional flight‐interception traps to quantify the response of these beetles to 25 and 40% selection cutting in a balsam fir–white birch forest of Québec, Canada. The abundance and species number of both cerambycid and scolytid beetles were 5–6 times larger in selectively cut stands than in controls the year following treatments. Analyses revealed that bark‐ and wood‐boring beetles’ response was mostly associated with increased canopy openness in selectively cut stands (and sun‐exposed locations within them) and residual tree injuries caused by harvesting operations. These conditions attracted beetles such as Trypodendron lineatum (Scolytinae) and Rhagium inquisitor (Cerambycidae), two species known for their ability to attack weakened, dying and dead hosts. Most species were more abundant in selection cuts, except for Evodinus m. monticola (Cerambycidae) whose abundance was strongly reduced after treatment. Some beetles can have detrimental effects on residual trees and thus could reduce timber value, but most species found in treated stands do not represent a high risk for healthy trees. Thus, selection cuts do not seem favourable to the establishment of tree‐killing beetles. However, as they were found more active/abundant after selection cutting, it would be wise to further study their population dynamics over mid‐ and long‐term periods, along with the ecological and economic implications associated with this silvicultural treatment.     

Spatial dynamics of regeneration in a conifer/broad‐leaved forest in northern Japan

Abstract. This study deals with stand dynamics over a 6‐yr period in a conifer/broad‐leaved mixed forest in Hokkaido, northern Japan. The annual rates of gap formation and recovery were 81.3 m²/ha and 66.7 m²/ha, respectively and turnover time of the canopy was 125 yr. The recruitment processes of the component species in this cool‐temperate forest were governed by different canopy types: gap, canopy edge and closed canopy. Magnolia obovata regenerated in canopy edges, and Acer mono and Prunus ssiori regenerated in canopy edges and gaps. The results suggested that the mosaic structure made up of closed canopy, canopy edge and gap was related to various regeneration niches. Abies sachalinensis had high mortality rates, initiating gap expansion. The transition probabilities from closed canopy or canopy edge to gap for deciduous broad‐leaved trees were lower than for A. sachalinensis, which implies that the difference in degeneration patterns of conifer and broad‐leaved canopies contributes to the heterogeneity of spatial structure in the mixed forests. Spatial dynamics were determined by a combination of gap expansion by A. sachalinensis (neighbour‐dependent disturbance) and gap formation by deciduous broad‐leaved trees (random disturbance).     

Differential effects of anthropogenic edges and gaps on the reproduction of a forest‐dwelling plant: The role of plant reproductive effort and nectar robbing by bumblebees

Forest fragmentation produces sharp changes in the spatial configuration of remnant forest patches, which include the increasing influence of patch edges on the interior of forest patches (‘edge effects’). Human activities responsible for forest fragmentation tend also to change the internal characteristics of remnant patches, for example, through the creation of gaps by selective logging. While edges and gaps can be expected to cause comparable changes in the micro‐environmental conditions of the forest, their effects on forest‐dwelling species and their interactions are not necessarily comparable. This study compares the effect of forest edges and anthropogenic gaps on the reproductive success of a self‐incompatible epiphytic plant (Mitraria coccinea), mediated by changes in its relationship with mutualists (pollinators) and antagonists (flower larcenists). Mitraria coccinea's flowers are pollinated by hummingbirds and robbed by the bumblebee Bombus dahlbomii. Edges and gaps had comparable positive effects on flower production and fruit set; however, nectar robbing was up to sevenfold higher in patches with numerous gaps and resulted in lower reproductive success (fruit set and total fruit crop). Forest fragmentation studies should therefore avoid treating forest remnants as homogeneous units (i.e. focusing exclusively on their characteristics and connectivity), without taking into account the internal heterogeneity caused by anthropogenic pressures (e.g. gaps and edge effects).     

Species traits and the response of open‐habitat species to forest edge in landscape mosaics

Human driven changes in land‐use have increased the need to understand how landscape structure affects species distribution. We studied how forest edges affected the distribution of birds in grasslands recently encroached by forest patches. We investigated how species’ biological traits influenced their response to vegetation change near forest edges. We censured birds along 300‐m line transects run into the open habitat perpendicularly to forest edges. We recorded habitat variables and landscape context along each transect and characterized edges and forest patches. We recorded 33 bird species in 153 transects for a total of 654 individuals. We analyzed species response to edges with generalized linear mixed models. Habitat preference was prevalent to explain species response to forest edges. The abundance of open‐habitat birds such as skylark Alauda arvensis decreased significantly in the vicinity of edges. This negative response extended within 150 m from the edge. The effect was disproportionately higher in open‐habitat species with high conservation concern. The abundance of species feeding or/and breeding in both forest and open habitat, such as woodlarks Lullula arborea, sharply increased near edges (positive edge response). Abundance of shrub and non‐shrub dependent species increased with distance to edge. The two species groups did no differ in abundance/distance to edge relationship. Intensity of species response to forest edges varied among transects in relation to transect vegetation characteristics. Edge length or aspect, diet and nest height had no direct effect. We discuss the possible role of variation in resources and nest predation risk to explain observed patterns.