Allometric density responses in butterflies: the response to small and large patches by small and large species

Species are differentially affected by habitat fragmentation as a consequence of differences in mobility, area requirements, use of the matrix, and responses to edges. A quantitative understanding of these differences is essential not only for conservation biology but also for basic ecological theory. Here, we examine density responses by butterflies to patch size and use a quantitative theory on the scaling of population density with patch size to interpret results. Theory suggests that the density distribution of mobile species along a patch size gradient should depend on the scaling of net migration rates, whereas the density distribution of less mobile species should depend more on local growth. Using data from 11 localities in three European countries, we calculated the slope in the relationship between patch size and population density. These slopes were evaluated in relation to butterfly traits and matrix composition. As estimates of butterfly mobility we used both wing span and expert mobility rankings. The slope of the density–area relationship changed as predicted with wing span and the association of species to grasslands. Large and highly mobile species had a negative slope, similarly for grassland specialists and generalist species, and the slope matched quantitative predictions based on the scaling of net migration rates. Small and less mobile grassland specialists had a slope that was less negative than the slope of large and mobile grassland specialists, whereas the slope did not change with size for generalist species. These analyses suggest that the variability in response among butterfly species to patch size could be explained by accounting for body size/mobility and habitat associations among species. A caveat is that edge effects are not explicitly included in the model analysis, and future research should aim to combine area and edge effects in a common theoretical framework.     

Population density and area: the role of between- and within-patch processes

The relationship between population density and the size of host plant patches was investigated for the red milkweed beetle Tetraopestetraophthalmus inhabiting unmanipulated patches of Asclepias syriaca. The resource concentration hypothesis proposes that density-area patterns, specifically that of increasing herbivore density with patch size, are primarily a function of movement between host plant patches. This research investigated the degree to which movement accounted for density-area patterns. Poisson regression analysis of beetle abundance versus milkweed patch size revealed that beetle density tended to increase with patch size. The pattern of density and patch size resulted from local reproduction and residence time. The density of emerging beetles tended to increase with patch size while emigration rates were unrelated to patch size. Immigration rates were constant with patch size for male beetles, and decreased with patch size for female beetles. Net flux of beetles (immigration – emigration) did not vary with patch size for male beetles and decreased with patch size for female beetles. Comparisons are made between this system and previously studied systems where movement plays a significant role in forming density area patterns. Additionally, several hypotheses are presented which may account for greater in situ recruitment and residence time in large patches. Received: 23 February 1996 / Accepted: 8 January 1997     

Testing the role of patch openness as a causal mechanism for apparent area sensitivity in a grassland specialist

Area sensitivity, species being disproportionately present on larger habitat patches, has been identified in many taxa. We propose that some apparently area-sensitive species are actually responding to how open a habitat patch is, rather than to patch size. We tested this hypothesis for Bobolinks (Dolichonyx oryzivorus) by comparing density and occupancy to a novel openness index, patch area, and edge effects. Bobolink density and occupancy showed significant relationships with openness, but logistic models based on an openness occupancy threshold had greater explanatory power. Thresholds remained approximately consistent from June through August, and shifted to be more open in September. Variance partitioning supported the openness index as unique and relevant. We found no relationships between measures of body condition (body mass, body size, circulating corticosterone levels) and either openness or area. Our findings have implications for studies of area sensitivity, especially with regards to inconsistencies reported within species: specifically, (1) whether or not a study finds a species to be area sensitive may depend on whether small, open sites were sampled, and (2) area regressions were sensitive to observed densities at the largest sites, suggesting that variation in these fields could lead to inconsistent area sensitivity responses. Responses to openness may be a consequence of habitat selection mediated by predator effects. Finally, openness measures may have applications for predicting effects of habitat management or development, such as adding wind turbines, in open habitat.     

Effects of aerial dispersal, habitat specialisation, and landscape structure on spider distribution across fragmented grey dunes

Species distribution patterns have been explained by Hutchinson's niche theory, metapopulation theory and source-sink theory. Empirical verification of this framework, however, remains surprisingly scant. In this paper, we test the hypothesis that landscape characteristics (patch size and connectivity), aerial dispersal ability and niche breadth interact in explaining distribution patterns of 29 spider species inhabiting fragmented grey dunes. Distribution patterns only depended on aerial dispersal potential, and the interaction between patch connectivity and area. Niche breadth, measured as the degree of habitat specialisation in the total coastal dune system, did not contribute to the observed distribution patterns. Additional variation in patch occupancy frequency was strongly species-dependent and was determined by different responses to the degree of patch connectivity for ballooning dispersal. Results from this study suggest that dispersal ability largely affects our perception of a species "fundamental niche", and that source-sink and metapopulation dynamics may have a major impact on the distribution of species. From a conservation point of view, specialised (and hence intrinsically rare) species can be predicted to become rarer if fragmentation increases and connectivity decreases. This study is, to our knowledge, one of the few linking species distribution (and not patch occupancy, species diversity or richness) to landscape ecological (patch connectivity and area) and auto-ecological (niche breadth, dispersal potential) features.     

Patch Size and Isolation Predict Plant Species Density in a Naturally Fragmented Forest

Studies of the effects of patch size and isolation on plant species density have yielded contrasting results. However, much of the available evidence comes from relatively recent anthropogenic forest fragments which have not reached equilibrium between extinction and immigration. This is a critical issue because the theory clearly states that only when equilibrium has been reached can the number of species be accurately predicted by habitat size and isolation. Therefore, species density could be better predicted by patch size and isolation in an ecosystem that has been fragmented for a very long time. We tested whether patch area, isolation and other spatial variables explain variation among forest patches in plant species density in an ecosystem where the forest has been naturally fragmented for long periods of time on a geological scale. Our main predictions were that plant species density will be positively correlated with patch size, and negatively correlated with isolation (distance to the nearest patch, connectivity, and distance to the continuous forest). We surveyed the vascular flora (except lianas and epiphytes) of 19 forest patches using five belt transects (50×4 m each) per patch (area sampled per patch = 0.1 ha). As predicted, plant species density was positively associated (logarithmically) with patch size and negatively associated (linearly) with patch isolation (distance to the nearest patch). Other spatial variables such as patch elevation and perimeter, did not explain among-patch variability in plant species density. The power of patch area and isolation as predictors of plant species density was moderate (together they explain 43% of the variation), however, a larger sample size may improve the explanatory power of these variables. Patch size and isolation may be suitable predictors of long-term plant species density in terrestrial ecosystems that are naturally and anthropogenically fragmented.     

Migration of the clouded Apollo butterfly Parnassius mnemosyne in a network of suitable habitats – effects of patch characteristics

The clouded Apollo Parnassius mnemosyne is a food plant specialist with short but frequent movements between habitat patches. The short average dispersal distances suggest that the probability of colonisation of vacant patches decreases rapidly as the distance between the source and target patches increases, which means that a dense habitat network is needed for the conservation of the species. Both emigration rate and the number of immigrants varied among patches and were not affected only by isolation but also by several other patch characteristics. The model that explained most of the variation in emigration rates among patches included patch area and the number of conspecifics. The area and the population density of the target patch had significant effects on the number of arriving immigrants. Thus, the colonisation of vacant patches is dependent on these patch characteristics. Generally, emigration rates were lower and residence times longer in large patches with many conspecifics. Butterfly density was the most important single factor explaining the variation in the number of immigrants among patches, although the positive effect of the area of the target patch was also significant. As a consequence of the marked positive density dependence caused by conspecific attraction, small patches with higher than average butterfly density, receive more immigrants than could be expected based on the patch area only. Due to conspecific attraction, per capita immigration rates are higher in small than large patches. Thus, immigration may have a more significant effect on the local dynamics of small than large populations.     

Female ornamentation, parental quality, and competitive ability in the rock sparrow

The evolution of female ornaments is poorly understood. Recent evidence suggests not only that female ornaments may be genetic correlates of selection on males but may also have evolved through male mate choice and/or through female–female aggressive interactions. In the rock sparrow, Petronia petronia, both sexes have a carotenoid-based yellow patch that is sexually selected by both sexes. The benefits that male may gain from choosing an attractive female remain unidentified. Both parents participate in caring for the young, so there should be mutual mate choice because males and females should both benefit from choosing a good parent (good parent hypothesis; GPH). Moreover, it has already been demonstrated that the yellow patch in males is also a badge of status (armament). Therefore, the yellow patch could also serve as both ornament and armament in females (dual utility hypothesis; DUH). We investigated the hypothesis that male and female yellow patch size signals parental quality in the field. We tested by an experiment in captivity the signal function of the yellow patch in female–female aggressive interactions for access to food. Yellow patch size correlated with paternal, but not maternal, feeding rates. Thus, this study supports the hypothesis that yellow patch dimension signals male parental quality, but there is no evidence for the GPH to explain female ornamentation. In the experiment females with relatively large yellow patches had earlier access to food than those with small patches. These results seem to suggest that a sexually selected carotenoid-feather signal may be used in female–female competition, in agreement with the DUH. Males may benefit from choosing well ornamented females because these may be superior competitors.     

Patch occupancy, population density and dynamics in a fragmented red squirrel Sciurusvulgaris population

We studied population dynamics of red squirrels in a group of small forest fragments, that cover only 6.5% of the total study area (4664 ha) and where distances to the nearest source population were up to 2.2 km. We tested effects of patch size, quality and isolation and supplementary feeding on patch occupation during 1995–99. Larger patches and patches with supplementary feeding had a higher probability of being occupied. No patch <3.5 ha was ever occupied. No effects of isolation were found, suggesting that the forest habitat in the study area is not sufficiently fragmented to influence red squirrel distribution across patches. For medium sized patches (3.7–21 ha), that were occupied some years, there was an increase in patch occupation over the years, even though overall population size tended to decrease. These patches had a high turnover, especially of males. Patches in which the squirrel population went extinct were recolonized within a year. For patches that were at least some years occupied, squirrel density depended on patch quality only. No effects of patch size, isolation and winter temperature on population density were found. These data suggest that in our study area habitat fragmentation has no effect on local squirrel density and that the random sample hypothesis explains the distribution pattern across patches.     

Testing the resource concentration hypothesis with tarnished plant bug on strawberry: density of hosts and patch size influence the interaction between abundance of nymphs and incidence of damage

Abstract. 1. Attributes of patches with strawberry plants were manipulated experimentally to assess the impact of patch size and host density on the abundance of tarnished plant bug (Hemiptera: Miridae) and incidence of fruit damage.
2. The density of nymphs per inflorescence increased with patch size and host density for some but not all generations of plant bug, providing partial support for the resource concentration hypothesis, while emphasising the importance of replicating experiments across several generations of herbivore.
3. Increasing density of nymphs with increasing patch size and host density did not translate into a high incidence of damage, which may be due to the relatively low feeding impact of nymphs in patches with numerous fruits and to the relatively high perimeter-to-area ratio in small patches resulting in a relatively high proportion of damaged fruits.
4. The decreasing proportion of damaged fruits with increasing density of fruits per patch suggests that nymphs exhibit a saturating functional response to density of strawberry fruits. Functional responses of herbivores may be a critical yet overlooked component that influences interactions between insects and their host plant. In particular, concentrated plant resources may result in increased numbers of herbivores yet sill have a negligible influence on plant damage and/or fitness.     

Why mammalian lineages respond differently to sexual selection: metabolic rate constrains the evolution of sperm size

The hypothesis that sperm competition should favour increases in sperm size, because it results in faster swimming speeds, has received support from studies on many taxa, but remains contentious for mammals. We suggest that this may be because mammalian lineages respond differently to sexual selection, owing to major differences in body size, which are associated with differences in mass-specific metabolic rate. Recent evidence suggests that cellular metabolic rate also scales with body size, so that small mammals have cells that process energy and resources from the environment at a faster rate. We develop the 'metabolic rate constraint hypothesis' which proposes that low mass-specific metabolic rate among large mammals may limit their ability to respond to sexual selection by increasing sperm size, while this constraint does not exist among small mammals. Here we show that among rodents, which have high mass-specific metabolic rates, sperm size increases under sperm competition, reaching the longest sperm sizes found in eutherian mammals. By contrast, mammalian lineages with large body sizes have small sperm, and while metabolic rate (corrected for body size) influences sperm size, sperm competition levels do not. When all eutherian mammals are analysed jointly, our results suggest that as mass-specific metabolic rate increases, so does maximum sperm size. In addition, species with low mass-specific metabolic rates produce uniformly small sperm, while species with high mass-specific metabolic rates produce a wide range of sperm sizes. These findings support the hypothesis that mass-specific metabolic rates determine the budget available for sperm production: at high levels, sperm size increases in response to sexual selection, while low levels constrain the ability to respond to sexual selection by increasing sperm size. Thus, adaptive and costly traits, such as sperm size, may only evolve under sexual selection when metabolic rate does not constrain cellular budgets.     

Relative importance of host plant patch geometry and habitat quality on the patterns of occupancy,extinction and density of the monophagous butterfly <Emphasis Type="Italic">Iolana iolas</Emphasis>

Habitat fragmentation is a major cause of species rarity and decline because it increases local population extinctions and reduces recolonisation rates of remnant patches. Although two major patch characteristics (area and connectivity) have been used to predict distribution patterns in fragmented landscapes, other factors can affect the occurrence of a species as well as the probability of it becoming extinct. In this paper, we study the spatial structure and dynamics of the butterfly Iolana iolas in a 75-patch network of its host plant (Colutea hispanica) to determine the relative importance of patch area, connectivity and habitat quality characteristics on occupancy, extinction and density over the period 2003–2006. Occupancy in 2003, incidence (proportion of years occupied) and probability of extinction were mostly affected by patch area. Smaller patches were less likely to be occupied because they had a higher probability of extinction, partly due to environmental stochasticity. The density of I. iolas was negatively related to patch area in all study years. Only in 2004 was the density of I. iolas positively influenced by fruit production per plant. Our results suggest that for I. iolas, and probably for other specialist butterflies with clearly delimited resource requirements, metapopulation dynamics can be satisfactorily predicted using only geometric variables because most habitat characteristics are subsumed in patch area. However, this hypothesis should be subject to further testing under diverse environmental conditions to evaluate the extent of its generalisation.     

Mechanisms driving the density–area relationship in a saproxylic beetle

Mechanisms underlying density–area relationships (correlations between population density and patch size) have rarely been tested experimentally. It is often assumed that higher density on large patches is evidence that large patches are high quality (i.e. have greater survival and reproduction), but the same pattern could result from disproportionate movement from small to large patches. Movement-based and within-patch processes must be experimentally separated to show that large patches are indeed of higher quality, but few studies have done so. We experimentally tested movement-based and within-patch hypotheses to explain the positive density–area relationship observed for a saproxylic (decayed wood-dependent) beetle, Odontotaenius disjunctus Illiger (Coleoptera: Passalidae). In separate experiments we quantified (1) immigration into and (2) finite growth rate within logs (=patches) that varied in size and density of conspecific beetles. A log was 11.7-fold [95 % confidence interval (CI) 3.4–40.8) and 10.5-fold (95 % CI 2.7–40.9) more likely to contain a new immigrant if it was large or contained a conspecific pair of beetles, respectively. Neither log size nor conspecific density was associated with changes in finite growth rate that would lead to higher density: decreased log size and increased conspecific density reduced finite growth rate in direct proportion to the loss in available resources per mating pair. We conclude that movement behavior rather than habitat quality is responsible for the positive density–area relationship for O. disjunctus. An important implication of our results is that population density is an imperfect indicator of habitat quality.     

Consequences for a host–parasitoid interaction of host-plant aggregation, isolation, and phenology

Abstract.  1. Spatial habitat structure can influence the likelihood of patch colonisation by dispersing individuals, and this likelihood may differ according to trophic position, potentially leading to a refuge from parasitism for hosts.
2. Whether habitat patch size, isolation, and host-plant heterogeneity differentially affected host and parasitoid abundance, and parasitism rates was tested using a tri-trophic thistle–herbivore–parasitoid system.
3.  Cirsium palustre thistles ( n = 240) were transplanted in 24 blocks replicated in two sites, creating a range of habitat patch sizes at increasing distance from a pre-existing source population. Plant architecture and phenological stage were measured for each plant and the numbers of the herbivore Tephritis conura and parasitoid Pteromalus elevatus recorded.
4. Mean herbivore numbers per plant increased with host-plant density per patch, but parasitoid numbers and parasitism rates were unaffected. Patch distance from the source population did not influence insect abundance or parasitism rates. Parasitoid abundance was positively correlated with host insect number, and parasitism rates were negatively density dependent. Host-plant phenological stage was positively correlated with herbivore and parasitoid abundance, and parasitism rates at both patch and host-plant scales.
5. The differential response between herbivore and parasitoid to host-plant density did not lead to a spatial refuge but may have contributed to the observed parasitism rates being negatively density dependent. Heterogeneity in patch quality, mediated by variation in host-plant phenology, was more important than spatial habitat structure for both the herbivore and parasitoid populations, and for parasitism rates.     

Does attraction to conspecifics explain the patch‐size effect? An experimental test

Recent theory suggests that attraction to conspecifics during habitat selection can be one potential, yet untested, mechanism for animal sensitivity to habitat fragmentation. The least flycatcher Empidonax minimus , a highly territorial migratory bird, has previously been shown to be attracted to conspecifics and sensitive to patch size by avoiding small patches of riparian forest in Montana, USA. I used a large-scale field experiment in this region to test the conspecific attraction hypothesis for explaining sensitivity to patch size, and I supplemented this experiment by estimating whether vegetation structure, nest predation, or nest parasitism rates could better explain patterns of sensitivity to patch size. Vegetation structure did not vary consistently with patch size, based on a random sample of patches across 150  km of the Madison and Missouri Rivers, Montana. Nest predation and parasitism rates by brown-headed cowbirds Molothrus ater also did not vary with patch size during the experiment. However, when conspecific cues were simulated across a gradient of patch sizes, flycatchers settled in all patches – and their sensitivity to patch size vanished – providing strong support for the conspecific attraction hypothesis. These results provide the first experimental evidence that attraction to conspecifics can indeed help explain area sensitivity in nature and highlight how understanding the role of animal behavior in heterogeneous landscapes can aid in interpreting pressing conservation issues.     

Immigration of olfactory searching insects into host plant patches: testing scaling rules for olfactory information

Herbivorous insects are commonly faced with host plants being distributed in scattered patches across a landscape. Immigration rates into habitat patches may strongly depend on the sensory cues used in the patch location process, and immigration rates of insects can be predicted based on the scaling of sensory cues. Here, we tested recent estimates of the scaling of olfactory information to patch size, which predicts a scaling coefficient ζ ≈ ?0.5 (A ^ζ , where A = patch size, ζ = scaling coefficient). We predicted that immigration rates of olfactory searching insects into patches of different sizes should scale according to the estimated slope. We investigated attraction of the weevils Cionus tuberculosus and Cionus scrophulariae to odors from figwort Scrophularia nodosa and quantified immigration rates of weevils into differently sized patches. We also investigated oviposition rates of the sawfly Tenthredo scrophulariae. The slope in the regression between density and patch size for herbivores was then compared with the predicted scaling coefficient. Using olfactometers, we found that weevils were attracted to figwort odors. Weevil densities were significantly affected by patch size, and the slope in the relationship between density and patch size was ζ = ?0.53. The slope in the relationship between larval densities of sawflies and patch size was less negative with a slope of ζ = ?0.15, indicating differences in search behavior compared with the weevils. The density–patch size relationship for the weevils closely matched the predicted slope and supported the previous estimations of the scaling of olfactory information to patch size.     

Applying the resource dispersion hypothesis to a fission–fusion society: A case study of the African lion (Panthera leo)

The relationship between the spatiotemporal distribution of resources and patterns of sociality is widely discussed. While the resource dispersion hypothesis (RDH) was formulated to explain why animals sometimes live in groups from which they derive no obvious benefits, it has also been successfully applied to species that benefit from group living. Some empirical tests have supported the RDH, but others have not, so conclusions remain equivocal and further research is required to determine the extent to which RDH predictions hold in natural systems. Here, we test four predictions of the RDH in an African lion population in the context of their fission–fusion society. We analyzed data on group composition of GPS‐collared lions and patterns of prey availability. Our results supported the first and second predictions of the RDH: Home range size (a) was independent of group size and (b) increased with distance between encounters with prey herds. Nonetheless, the third and fourth RDH predictions were not supported: (c) The measure of resource heterogeneity and (d) resource patch richness measured through prey herd size and body size had no significant effect on lion group size. However, regarding the fourth prediction, we added an adaptation to account for dynamics of fission–fusion society and found that the frequency of pride fission increased as group size increased. Our data set restricted us from going on to explore the effect of fission–fusion dynamics on the relationship between group size and patch richness. However, this should be investigated in future studies as including fission–fusion dynamics provides a more nuanced, realistic appreciation of lion society. Our study emphasizes the importance of understanding the complexity of a species' behavioral ecology within the framework of resource dispersion. Whatever larger theoretical framework may emerge to explain lion society, incorporating fission–fusion dynamics should allow the RDH to be refined and improved.     

Regional patch dynamics of Cirsium arvense and possible implications for plant‐animal interactions

Abstract. Plant population biology considers the dynamics of plant modules within stands. However, stands themselves may have considerable regional turnover in space and time. These changes in the number, distribution and size of plant stands generate a dynamic spatial pattern with important implications for the spatial and temporal dynamics of phytophagous insects using these plants as a host. During five successive years we studied the regional distribution and patch dynamics of the creeping thistle Cirsium arvense and the distribution of associated populations of the herbivore Urophora cardui (Diptera: Tephritidae), a specialist stem gall former. The study conducted was in a 15 km² heterogeneous, agricultural area in northeastern Bavaria. The distribution of the number of plants per patch was skewed with many more small C. arvense patches than large ones. During the five years of study, there was a 50% increase in the number of C. arvense patches, and a decrease in the mean number of plants per patch (= patch size) to less than half the patch size of the first year. Whilst patch size was randomly distributed in space, patch density showed a consistent, non‐random spatial pattern. Patch density was spatially auto‐correlated, with areas of high or low patch density having a characteristic dimension of ca. 1 km. Patch size was predictable in time and appeared to be regulated by size dependent processes, with the extinction probability of a patch being negatively correlated with its size. Correlated with the decline of C. arvense patch size during the study, the occupancy and total numbers of the herbivore U. cardui had a marked decrease, suggesting that the regional distribution of the stem gall former is not only influenced by patch number but more importantly by the mean patch size. With decreasing patch sizes, U. cardui was faced with an increasingly dynamic landscape due to higher extinction rates of small patches, although the mean distance between host plant patches decreased.     

Density‐dependent mortality in adults,but not juveniles,of stream‐resident brown trout (Salmo trutta)

1. This study investigates when and where density dependence operates on the mortality rates of stream‐resident brown trout Salmo trutta. To this aim, I explored populations in habitats of different quality containing high, low or intermediate densities over broad scales of space and time. The study is based on census data of 170 cohorts quantified from recruitment to the total disappearance at 12 sites in four contrasting tributaries of the Rio Esva drainage (north‐western Spain), over the years 1986–2007. 2. Log₁₀‐transformed survivor density over time highlighted a consistent pattern for the 170 cohorts characterised by the occurrence of only two life stages. An early stage starts at recruitment, lasts about half the lifetime and shows no or negligible mortality. A threshold time at 425–620 days after emergence preceded a second stage of continuous and constant mortality until the final disappearance of the cohorts. Consequently, in all scenarios, mortality only occurred in the adult component and no effect of season, year, age‐class and/or reproductive stage was detected. 3. Substantial spatial and temporal variations typified both recruitment (range R = 0.01–1.62 ind m^?2) and adults’ mortality rates (range Z = 0.03–0.38 day^?1). Nested anova s revealed strong effects of site and year on both recruitment and mortality with sites interspersed along the stream gradients where recruitment and mortality were typically high relative to other sites located either nearby in the same stream or distant in another stream, where both recruitment and mortality rates were typically low or intermediate. 4. Adult mortality rates plotted against recruitment for the 170 cohorts pooled revealed a continuous, positive power relationship that explained 45.3% of variation in mortality rates over the whole range of recruitment values. Similarly, highly significant power relationships were elucidated for site‐specific mortality rates averaged across years and for annual‐specific mortality rates averaged across sites against the corresponding mean recruitment averaged across years and sites, respectively. These relationships support the hypothesis that the operation of density dependence is scale independent and context independent but operates in a continuous manner across all scenarios examined. 5. A chronic effect of density dependence on adult losses induces temporally persistent populations maintained by a low number of spawners. Apparently, the operation of density dependence adjusts the number of spawners to the availability of rearing and spawning habitat. This dynamic process may also help to explain the small effective population size (Ne) recently documented by genetic studies of stream‐living brown trout and other salmonids.     

Testing the importance of patch scale on forest birds

The relationship between population density and habitat area is of central importance to conservation biology, particularly for species dependent on declining habitat. A recent study by Lee et al. in Oikos in 2002 found that the densities of three forest interior bird species (ovenbirds, wood thrushes, and red-eyed vireos) decline with increasing patch size, contradicting many other studies that demonstrate a positive correlation between area and density for these species. The authors' argument is based on a misapplied criticism of the prevailing approach based on density-and on an inappropriate statistical methodology, caused by incorrect specification of the null hypothesis after log–transformation. We use three different methods of testing area-dependence: a corrected log–abundance log–area regression; a similar abundance–area regression; and the prevailing density-area approach. For each species, all three methods agree broadly, suggesting the robustness of each approach. For ovenbirds and wood thrushes, we do not find that population density is negatively correlated to patch size. While our analyses of the red-eyed vireo data find a negative correlation between the two factors, the strength of the correlation is far weaker than that of Lee et al. and may derive from landscape factors unconsidered in the original data. Nevertheless, we do not find positive density–area relationships for any of these forest-interior species, further underscoring the site-specificity of the underlying mechanisms of area- sensitivity.