Interpatch movement and edge effects: the role of behavioral responses to the landscape matrix

Animal interpatch movement and spatial distribution are known to be influenced substantially by the composition of the landscape matrix, but little is known about the underlying mechanisms. In previous mark–recapture experiments we have found that the rates of emigration and immigration for the planthopper Prokelisia crocea are greater within a matrix composed of the introduced grass smooth brome (Bromus inermis) than a mudflat matrix. Additionally, census data indicated that individuals aggregate near the edge of host‐plant patches (prairie cordgrass; Spartina pectinata) bordered by mudflat, but not in patches bordered by nonhost grasses such as brome. Here, we investigate the mechanistic basis of these matrix effects by tracking the individual movements of planthoppers released at the edge of brome‐ and mudflat‐bordered cordgrass patches, and within homogeneous habitats of each type (cordgrass, brome, and mudflat). We found that patch edges bordered by brome were three times more permeable to emigration than mudflat‐bordered edges. Also, planthoppers exhibited no tendency to avoid edges by moving away (i.e. towards the patch interior). Within homogeneous habitats, comparison of the fractal dimension of movement paths revealed that movement was more linear in mudflat than in brome or cordgrass. In addition, planthoppers exhibited greater step lengths (distance moved per 10‐min interval), shorter residency times (duration of pauses between movements), and greater rates of net linear displacement in mudflat than brome and cordgrass. We attribute the planthopper's distributional patterns within patches to the lower permeability of mudflat than nonhost grass edges and the absence of edge–avoidance behavior. Contrary to conventional wisdom that low‐resistance matrix types (e.g. those that promote high displacement rates) enhance interpatch dispersal rates, dispersal success may be higher in brome matrix because tortuous movement through this matrix increases the planthopper's rate of encounter with cordgrass patches.     

Landscape context outweighs local habitat quality in its effects on herbivore dispersal and distribution

Past studies with spatially structured herbivore populations have emphasized the primacy of intrinsic factors (e.g., patch quality), patch geometry (e.g., patch size and isolation), and more recently landscape context (e.g., matrix composition) in affecting local population abundance and dispersal rate. However, few studies have examined the relative importance of each factor, or how they might interact to affect herbivore abundance or dispersal. Here, we performed a factorial field experiment to examine the independent and interactive effects of patch quality (plant biomass, leaf protein, leaf phenolics) and matrix composition [mudflat or non-host grass (Bromus inermis)] on planthopper (Prokelisia crocea) emigration from host-plant patches (prairie cordgrass, Spartina pectinata). In addition, a field survey was conducted to examine the relative importance of patch quality, geography, and matrix composition on planthopper occupancy and density. In the experiment, we found that rates of emigration from low and intermediate quality patches were, on average, 21% percent higher for patches embedded in brome than mudflat. In contrast, the emigration rate was unaffected by matrix composition in nutrient-rich patches. Within matrix types, plant quality had little effect on emigration. In the survey, planthopper density and the patch occupancy rate of planthoppers increased nonadditively with increasing patch size and the percentage of the surrounding matrix composed of mudflat. This study suggests that landscape-level factors, such as the matrix, may be more important than factors intrinsic to the patches.     

Patch edges and insect populations

Responses of insect populations may be related to patch size and patch edge responses, but it is not clear how to identify these rapidly. We used a random-walk model to identify three qualitative responses to edges: no edge effect (the null model), reflecting edges and absorbing edges. Interestingly, no edge effect meant that abundance was lower at edges than in the center of patches, and reflecting edges have similar abundance at edges and centers. We then characterized several insect species’ response within maize plots to patch edges and patch size, using a simple, quick, qualitative experiment. Coleomegilla maculata and Trichogramma spp. were the only organisms that responded to patch size and edges as patch theory and the null edge model would predict. Ostrinia nubilalis larvae and possibly Rhopalosiphum maidis and eggs of Chrysopa spp. responded to patch size and edges as predicted by an attracting edge model. Estimation of predation rates suggested that the spatial distribution of these species might be determined by predators. Edge effects or the lack thereof relative to patch size may be rapidly determined for arthropod species, which could lead to understanding the mechanism(s) underlying these effects. This information may be useful in reaction diffusion models through a scaling-up approach to predict population structure of species among patches in a landscape. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Patch dynamics of a native grass in relation to the spread of invasive smooth brome (Bromus inermis)

The effects of invasive species on the patch dynamics (establishment, growth, and local extinction) of native species are not well studied, owing to the need for relatively fine-scale data on the distribution of species. Within the prairie pothole region of the United States and Canada, the grass, Bromus inermis (smooth brome) has become established by invading disturbed prairies, and through repeated introductions for soil retention and animal graze. In this study, the impact of smooth brome on the patch dynamics of a dominant native grass species, Spartina pectinata (prairie cordgrass), was assessed using fine-scale (sub-meter) mapping of the distribution of cordgrass and brome in three prairie fragments from 2000 to 2006. Using GIS spatial analyses, we determined that cordgrass patch growth was two times greater in areas not invaded by smooth brome versus areas that were heavily infested with smooth brome. Among sites and time periods, there was a consistent significant negative relationship between the amount of smooth brome surrounding a patch of cordgrass and the growth of that cordgrass patch. The probability of establishment of a new patch of cordgrass averaged 1.3 times higher in areas of low brome coverage (<25%) than areas of high brome coverage (>75%). Conversely, existing cordgrass patches were 7.8 times more likely to go extinct in areas of high than low brome coverage. This is one of only a few field studies to provide evidence of the negative impact of smooth brome on native flora and hopefully will serve as justification for the development of a formal management plan to limit the distribution of this species in tallgrass prairie ecosystems.     

The role of the physical structure of Spartina densiflora Brong. in structuring macroinvertebrate assemblages

In Patagonian rocky salt marshes, the presence of the austral cordgrass Spartina densiflora provide habitat for diverse faunal assemblages. Two different mechanisms may influence the distribution and abundance patterns of these associated organisms: those generated by the biological properties of cordgrasses and those caused or mediated by the physical structure supplied by the plants. The aims of this study were: (1) to determine the effect of cordgrasses on macroinvertebrate assemblages dominating a rocky marsh and (2) to determine how much of this effect is caused by the physical structure supply by the plants. In order to achieve these objectives, we conducted two field manipulative experiments. In the first one, we manipulated the presence of defaunated transplants of cordgrass, and in the second one, we did the same with plastic cordgrass mimics simulating the architecture of Spartina densiflora. In both experiments, sessile and mobile assemblages were separately examined to evaluate whether they respond in the same way or not. Abundance and richness of mobile and sessile organisms were higher in transplant and mimic cordgrass plots compared to the controls, indicating that the presence of cordgrasses, either transplanted or artificial, has a positive effect on sessile and mobile assemblages. Furthermore, we found that the composition of mobile and sessile macroinvertebrates assemblages did not differ between transplants and mimics. Therefore, the physical structure of cordgrass was found to be the major factor influencing macroinvertebrates assemblages in the studied Patagonia rocky salt marshes. Within the period of 1 year, mobile and sessile fauna colonized several times more abundantly transplant and artificial cordgrass than the control unvegetated plots, highlighting the key role of cordgrasses in optimizing the colonization rate of macroinvertebrate communities in this rocky bottom environment. Since Spartina densiflora is invading different regions worldwide, our study may help to predict its potential effect on the invaded communities.     

Estimation of the diffusion rate and crossing probability for biased edge movement between two different types of habitat

One of the fundamental goals of ecology is to examine how dispersal affects the distribution and dynamics of insects across natural landscapes. These landscapes are frequently divided into patches of habitat embedded in a matrix of several non-habitat regions, and dispersal behavior could vary within each landscape element as well as the edges between elements. Reaction–diffusion models are a common way of modeling dispersal and species interactions in such landscapes, but to apply these models we also need methods of estimating the diffusion rate and any edge behavior parameters. In this paper, we present a method of estimating the diffusion rate using the mean occupancy time for a circular region. We also use mean occupancy time to estimate a parameter (the crossing probability) that governs one type of edge behavior often used in these models, a biased random walk. These new methods have some advantages over other methods of estimating these parameters, including reduced computational cost and ease of use in the field. They also provide a method of estimating the diffusion rate for a particular location in space, compared to existing methods that represent averages over large areas. We further examine the statistical properties of the new method through simulation, and discuss how mean occupancy time could be estimated in field experiments.     

Host-parasitoid extinction and colonization in a fragmented prairie landscape

Few field studies of natural populations have examined the factors influencing local extinctions and colonization of empty habitat patches for a prey species and its predator. In this study, I carried out a census of planthopper (Prokelisia crocea; Hemiptera: Delphacidae) and egg parasitoid (Anagrus columbi; Hymenoptera: Mymaridae) incidence and densities in 147 host-plant patches (Spartina pectinata; Poaceae) over seven planthopper generations in a tall-grass prairie landscape. For both species, the likelihood of going extinct in a patch was related to a number of patch-specific variables: density, temporal variability in density, proportion of hosts parasitized (planthopper only), host-plant density, patch size, patch isolation, and composition of the surrounding matrix. Colonization likelihood was related only to the physical attributes of the patch. There was high patch turnover in this prairie landscape. On average, planthoppers went extinct in 23% of the patches and A. columbi went extinct in 51% of the patches in each generation. For the planthopper, extinction likelihood increased with a decrease in patch size and the proportion of the matrix composed of mudflat. Parasitism of eggs had no effect on the extinction likelihood of local P. crocea populations, suggesting that A. columbi may not play a major role in the patch dynamics of its host. The likelihood of extinction for A. columbi was dependent on factors that spanned three trophic levels. An increase in plant density, decrease in host density and decrease in parasitoid density all increased the likelihood of A. columbi extinction within a patch. The dependency on multiple trophic levels may explain the higher extinction risk for the parasitoid than its host. A. columbi extinction was also affected by the matrix habitat surrounding the patch—the effect was the opposite of that for P. crocea. Finally, vacant patches were colonized at rates of 53% and 34% per generation for the planthopper and parasitoid, respectively. For both species, colonization probabilities decreased with an increase in patch isolation. High host densities in a patch also favored high rates of colonization by A. columbi. I discuss how anthropogenic changes to the prairie landscape can affect the metapopulation dynamics and persistence time of this host-parasitoid interaction.     

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.     

Edge fires drive the shape and stability of tropical forests

In tropical regions, fires propagate readily in grasslands but typically consume only edges of forest patches. Thus, forest patches grow due to tree propagation and shrink by fires in surrounding grasslands. The interplay between these competing edge effects is unknown, but critical in determining the shape and stability of individual forest patches, as well the landscape‐level spatial distribution and stability of forests. We analyze high‐resolution remote‐sensing data from protected Brazilian Cerrado areas and find that forest shapes obey a robust perimeter–area scaling relation across climatic zones. We explain this scaling by introducing a heterogeneous fire propagation model of tropical forest‐grassland ecotones. Deviations from this perimeter–area relation determine the stability of individual forest patches. At a larger scale, our model predicts that the relative rates of tree growth due to propagative expansion and long‐distance seed dispersal determine whether collapse of regional‐scale tree cover is continuous or discontinuous as fire frequency changes.     

Cropland edge,forest succession,and landscape affect shrubland bird nest predation

The effects of habitat edges on nest survival of shrubland birds, many of which have experienced significant declines in the eastern United States, have not been thoroughly studied. In 2007 and 2008, we collected data on nests of 5 shrubland passerine species in 12 early successional forest patches in North Carolina, USA. We used model selection methods to assess the effect of distance to cropland and mature forest edge on nest predation rates and additionally accounted for temporal trends, nest stage, vegetation structure, and landscape context. For nests of all species combined, nest predation decreased with increasing distance to cropland edge, by nearly 50% at 250 m from the cropland edge. Nest predation of all species combined also was higher in patches with taller saplings and less understory vegetation, especially in the second year of our study when trees were 4–6 m tall. Predation of field sparrow (Spizella pusilla) nests was lower in landscapes with higher agricultural landcover. Nest predation risk for shrubland birds appears to be greater near agricultural edges than mature forest edges, and natural forest succession may drive patterns of local extirpation of shrubland birds in early successional forest patches. Thus, we suggest that habitat patches managed for shrubland bird populations should be considerably large or wide (>250 m) when adjacent to crop fields and maintained in structurally diverse early seral stages. © 2011 The Wildlife Society.     

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.     

Habitat fragmentation and extinction thresholds on fractal landscapes

Habitat fragmentation is a potentially critical factor in determining population persistence. In this paper, we explore the effect of fragmentation when the fragmentation follows a fractal pattern. The habitat is divided into patches, each of which is suitable or unsuitable. Suitable patches are either occupied or unoccupied, and change state depending on rates of colonization and local extinction. We compare the behaviour of two models: a spatially implicit patch-occupancy (PO) model and a spatially explicit cellular automaton (CA) model. The PO model has two fixed points: extinction, and a stable equilibrium with a fixed proportion of occupied patches. Global extinction results when habitat destruction reduces the proportion of suitable patches below a critical threshold. The PO model successfully recreates the extinction patterns found in other models. We translated the PO model into a stochastic cellular automaton. Fractal arrangements of suitable and unsuitable patches were used to simulate habitat fragmentation. We found that: (i) a population on a fractal landscape can tolerate more habitat destruction than predicted by the patch-occupancy model, and (ii) the extinction threshold decreases as the fractal dimension of the landscape decreases. These effects cannot be seen in spatially implicit models. Landscape struc-ture plays a vital role in mediating the effects of habitat fragmentation on persistence.     

Breeding occupancy and success of two hole-nesting passerines: the impact of fragmentation caused by forestry

We studied the distribution and the reproductive success of the pied flycatcher Ficedula hypoleuca and the redstart Phoenieurus phoenicurus in relation to forest patch size, edge type (clearcut vs natural), distance from the forest edge, and vegetation characteristics in a forest-dominated landscape. Breeding performances were recorded in up to 72 forest patches during 1992–1994 for birds breeding in nest-boxes. In the spring, breeding individuals of both species arrived earlier in large forest patches (> 1 ha) than in smaller ones. Pied flycatchers arrived earlier on clearcut edges than natural edges but in the redstart there was no preference for a particular edge type. The territory distance from the forest to open land edge did not affect the arrival dates of either species. In the case of the pied flycatcher, the proportion of unpaired males was highest in patches < 1 ha in size and in the case of the redstart this applied to patches < 5 ha in size. Pairing success was not related to the forest edge type or the nest's distance from the edge. Nest predation was not patch-size nor edge-related for either of the species, but in the combined data for both species nest predation was higher at clearcut edges than at natural edges. Clutch size, brood size and the survival of nestlings to the fiedgling stage (fledgling/egg. %) were independent of the patch size, edge type and nest's "distance from the forest edge.     

Discriminating the Drivers of Edge Effects on Nest Predation: Forest Edges Reduce Capture Rates of Ship Rats (Rattus rattus), a Globally Invasive Nest Predator,by Altering Vegetation Structure

Forest edges can strongly affect avian nest success by altering nest predation rates, but this relationship is inconsistent and context dependent. There is a need for researchers to improve the predictability of edge effects on nest predation rates by examining the mechanisms driving their occurrence and variability. In this study, we examined how the capture rates of ship rats, an invasive nest predator responsible for avian declines globally, varied with distance from the forest edge within forest fragments in a pastoral landscape in New Zealand. We hypothesised that forest edges would affect capture rates by altering vegetation structure within fragments, and that the strength of edge effects would depend on whether fragments were grazed by livestock. We measured vegetation structure and rat capture rates at 488 locations ranging from 0–212 m from the forest edge in 15 forest fragments, seven of which were grazed. Contrary to the vast majority of previous studies of edge effects on nest predation, ship rat capture rates increased with increasing distance from the forest edge. For grazed fragments, capture rates were estimated to be 78% lower at the forest edge than 118 m into the forest interior (the farthest distance for grazed fragments). This relationship was similar for ungrazed fragments, with capture rates estimated to be 51% lower at the forest edge than 118 m into the forest interior. A subsequent path analysis suggested that these ‘reverse’ edge effects were largely or entirely mediated by changes in vegetation structure, implying that edge effects on ship rats can be predicted from the response of vegetation structure to forest edges. We suggest the occurrence, strength, and direction of edge effects on nest predation rates may depend on edge-driven changes in local habitat when the dominant predator is primarily restricted to forest patches.     

Local movement and edge effects on competition and coexistence in ephemeral-patch models

For insects exploiting spatially structured arrays of resource patches (host plants, fungi, carrion, etc.), the distribution of individuals among patches can have important consequences for the coexistence of competitors. In general, intraspecific aggregation of consumer individuals over the landscape of patches stabilizes competition. Oviposition behavior of individual females can generate aggregation of larvae across patches and, therefore, strongly influences the outcome of competition between co-occurring species. We used simulation models to evaluate the consequences (for the coexistence of competitors) of different movement behaviors by females before and between oviposition events. Coexistence times increase when females are more likely to travel among neighboring patches than among distant ones. Coexistence times are also longer when females begin egg laying near the site of their emergence. Preoviposition dispersal is, therefore, destabilizing. We also considered responses by females to edges of resource arrays. Edge effects are generally stabilizing, delaying competitive exclusion by increasing larval aggregation, but different responses to edges have dramatically different effects on coexistence. The longest coexistence times occur when edges are "sticky", such that females encountering an edge tend to remain there.     

Dispersal in a metapopulation of the bush cricket, Metrioptera bicolor (Orthoptera: Tettigoniidae)

1. I present a stochastic simulation model that describes individual movements of Metrioptera bicolor Philippi in a heterogeneous landscape, consisting of patches of suitable habitat surrounded by a matrix of unprofitable habitats. Although the model is parameterized with information about daily movement behaviour, it can generate spatially explicit predictions about inter-patch dispersal rates for much longer periods, e.g. one generation.
2. Long-term dispersal experiments were conducted to evaluate model predictions. Patch-specific emigration rates and the total distance moved by individuals could be predicted with satisfactory precision. Because of the stochastic nature of the model, it failed to predict which recipient patches emigrating individuals actually chose in a particular situation.
3. Spatially explicit simulations of the movement model were made for the whole natural distribution area of M. bicolor . The results suggest that emigration rates are negatively correlated with patch size. Local populations occurring on small patches may be more prone to extinction than those on large patches, by losing more emigrants than are compensated for by immigration.     

Homogenization techniques for population dynamics in strongly heterogeneous landscapes

An important problem in spatial ecology is to understand how population-scale patterns emerge from individual-level birth, death, and movement processes. These processes, which depend on local landscape characteristics, vary spatially and may exhibit sharp transitions through behavioural responses to habitat edges, leading to discontinuous population densities. Such systems can be modelled using reaction–diffusion equations with interface conditions that capture local behaviour at patch boundaries. In this work we develop a novel homogenization technique to approximate the large-scale dynamics of the system. We illustrate our approach, which also generalizes to multiple species, with an example of logistic growth within a periodic environment. We find that population persistence and the large-scale population carrying capacity is influenced by patch residence times that depend on patch preference, as well as movement rates in adjacent patches. The forms of the homogenized coefficients yield key theoretical insights into how large-scale dynamics arise from the small-scale features.     

Rain Forest Structure at Forest-Pasture Edges in Northeastern Costa Rica

Land-use change in the Sarapiquí region of Costa Rica has resulted in a fragmented forest landscape with abrupt edges between forest and pasture. Forest responses to edge effects vary widely and can significantly affect ecosystem integrity. Our objective was to examine forest structure at 20+ yr old forest-pasture edges in Sarapiquí. Three transects with 0.095-ha plots at seven distances from forest edges were established in each of six forest patches. Stem density, basal area, and aboveground biomass in trees and palms ≥ 10-cm diameter at breast height were measured in all plots. In addition, hemispherical photographs were taken to determine leaf area index, understory light availability, and percent canopy openness. Linear mixed-effects models showed significantly higher tree stem density at forest edges, relative to interiors, a pattern reflected by increased stem density, basal area, and aboveground biomass in small diameter trees (≤ 20 cm) growing near edges. No differences in total tree basal area, aboveground biomass, or hemispherical photograph-derived parameters were detected across the forest edge to interior gradient. The recruitment of small diameter trees following edge creation has contributed to the development of dense vegetation at the forest edge and has aided in the maintenance of similar tree basal area and aboveground biomass between edge and interior environments. These data reflect on the robustness of forest edges in Sarapiquí, a characteristic that will likely minimize future detrimental edge effects and promote a number of high-value environmental services in these forests.     

Seagrass growth and patch dynamics: cross-scale morphological plasticity

By examining the spatial distribution of rhizome morphological characteristics of the seagrass Halodule wrightii, in relation to a seasonal pattern of seagrass patch dynamics, we attempted to derive a mechanistic explanation for the variety of changes exhibited by seagrass patch shapes. Rhizome morphological characteristics (mean internodal distance, branching frequency and biomass) were measured at three spatially-recognized regions (Flood edge, Center, Ebb edge) of 5 seagrass patches, reflecting position relative to hydrodynamic flow. In addition, maps (1 resolution) of the seagrass patches were used to quantify changes in seagrass patch margins across the growing season. Rhizome morphological characteristics varied with spatial position: longer internodal distances were recorded on both edges of the patch relative to patch center, and rhizomes from Flood edges exhibited longer internodes than plants on the Ebb edge of patches. Branching frequency showed no spatially-explicit distribution across the seagrass patches. Patch change analysis indicated a pattern of increase in patch area on the Flood edges of seagrass patches and recession (or no change) on the Ebb edges. Patch margin change was significantly correlated with internodal distances: the more positive the increase in patch seagrass coverage on an edge, the greater the internodal distances.Sediment nutrients were explored as a potential mechanism for the distinct spatial distribution of morphologies found; experimental addition of phosphorus, but not nitrogen, significantly altered the rhizome morphology and biomass, but measurement of ambient sediment nutrient concentrations produced no significant correlations with the in situ distribution of rhizome morphologies. These results suggest that larger-scale landscape characteristics of patch dynamics may be determined by predictable behaviors of small-scale components, but the results do not conclusively describe a mechanism for this system.