Wild mockingbirds distinguish among familiar humans.

Although individuals of some species appear able to distinguish among individuals of a second species, an alternative explanation is that individuals of the first species may simply be distinguishing between familiar and unfamiliar individuals of the second species. In that case, they would not be learning unique characteristics of any given heterospecific, as commonly assumed. Here we show that female Northern Mockingbirds (Mimus polyglottos) can quickly learn to distinguish among different familiar humans, flushing sooner from their nest when approached by people who pose increasingly greater threats. These results demonstrate that a common small songbird has surprising cognitive abilities, which likely facilitated its widespread success in human-dominated habitats. More generally, urban wildlife may be more perceptive of differences among humans than previously imagined.

Changing patterns of nest predation and predator communities along a tropical elevation gradient.

Tropical montane communities host the world's highest beta diversity of birds, a phenomenon usually attributed to community turnover caused by changes in biotic and abiotic factors along elevation gradients. Yet, empirical data on most biotic factors are lacking. Nest predation is thought to be especially important because it appears to be common and can change selective pressures underlying life history traits, which can alter competitive interactions. We monitored 2538 nests, 338 of which had known nest predators, to evaluate if nest predation changes along a tropical elevational gradient. We found that nest predation decreased with elevation, reflecting the loss of lowland predators that do not tolerate colder climates. We found different "super" nest predators at each elevation that accounted for a high percentage of events, suggesting that selection pressures exerted by nest predator communities may be less diffuse than has been hypothesized, at least for birds nesting in the understory.

Habitat fragmentation alters the distance of abiotic seed dispersal through edge effects and direction of dispersal.

Habitat loss and fragmentation are leading causes of species declines, driven in part by reduced dispersal. Isolating the effects of fragmentation on dispersal, however, is daunting because the consequences of fragmentation are typically intertwined, such as reduced connectivity and increased prevalence of edge effects. We used a large-scale landscape experiment to separate consequences of fragmentation on seed dispersal, considering both distance and direction of local dispersal. We evaluated seed dispersal for five wind- or gravity-dispersed, herbaceous plant species that were planted at different distances from habitat edges, within fragments that varied in their connectivity and shape (edge-to-area ratio). Dispersal distance was affected by proximity and direction relative to the nearest edge. For four of five species, dispersal distances were greater further from habitat edges and when seeds dispersed in the direction of the nearest edge. Connectivity and patch edge-to-area ratio had minimal effects on local dispersal. Our findings illustrate how some, but not all, landscape changes associated with fragmentation can affect the key population process of seed dispersal.

Differentiation during fig ontogeny suggests opposing selection by mutualists.

Dioecy allows separation of female and male functions and therefore facilitates separate co-evolutionary pathways with pollinators and seed dispersers. In monoecious figs, pollinators' offspring develop inside the syconium by consuming some of the seeds. Flower-stage syconia must attract pollinators, then ripen and attract seed dispersers. In dioecious figs, male ("gall") figs produce pollen but not viable seeds, as the pollinators' larvae eat all seeds, while female ("seed") figs produce mostly viable seeds, as pollinators cannot oviposit in the ovules. Hence, gall and seed figs are under selection to attract pollinators, but only seed figs must attract seed dispersers. We test the hypothesis that seed and gall syconia at the flower stage will be similar, while at the fruiting stage they will differ. Likewise, monoecious syconia will be more similar to seed than gall figs because they must attract both pollinators and seed dispersers. We quantified syconium characteristics for 24 dioecious and 11 monoecious fig species and recorded frugivore visits. We show that seed and gall syconia are similar at the flower stage but differ at the fruit stage; monoecious syconia are more similar to seed syconia than they are to gall syconia; seed and gall syconia differentiate through their ontogeny from flower to fruit stages; and frugivores visit more monoecious and seed syconia than gall syconia. We suggest that similarity at the flower stage likely enhances pollination in both seed and gall figs and that differentiation after pollination likely enhances attractiveness to seed dispersers of syconia containing viable seeds. These ontogenetic differences between monoecious and dioecious species provide evidence of divergent responses to selection by pollinators and seed dispersers.

Ongoing accumulation of plant diversity through habitat connectivity in an 18-year experiment.

Deleterious effects of habitat fragmentation and benefits of connecting fragments could be significantly underestimated because changes in colonization and extinction rates that drive changes in biodiversity can take decades to accrue. In a large and well-replicated habitat fragmentation experiment, we find that annual colonization rates for 239 plant species in connected fragments are 5% higher and annual extinction rates 2% lower than in unconnected fragments. This has resulted in a steady, nonasymptotic increase in diversity, with nearly 14% more species in connected fragments after almost two decades. Our results show that the full biodiversity value of connectivity is much greater than previously estimated, cannot be effectively evaluated at short time scales, and can be maximized by connecting habitat sooner rather than later.

Breeding latitude predicts timing but not rate of spring migration in a widespread migratory bird in South America.

Identifying the processes that determine avian migratory strategies in different environmental contexts is imperative to understanding the constraints to survival and reproduction faced by migratory birds across the planet.We compared the spring migration strategies of Fork-tailed Flycatchers (Tyrannus s. savana) that breed at south-temperate latitudes (i.e., austral migrants) vs. tropical latitudes (i.e., intratropical migrants) in South America. We hypothesized that austral migrant flycatchers are more time-selected than intratropical migrants during spring migration. As such, we predicted that austral migrants, which migrate further than intratropical migrants, will migrate at a faster rate and that the rate of migration for austral migrants will be positively correlated with the onset of spring migration.We attached light-level geolocators to Fork-tailed Flycatchers at two tropical breeding sites in Brazil and at two south-temperate breeding sites in Argentina and tracked their movements until the following breeding season.Of 286 geolocators that were deployed, 37 were recovered ~1 year later, of which 28 provided useable data. Rate of spring migration did not differ significantly between the two groups, and only at one site was there a significantly positive relationship between date of initiation of spring migration and arrival date.This represents the first comparison of individual migratory strategies among conspecific passerines breeding at tropical vs. temperate latitudes and suggests that austral migrant Fork-tailed Flycatchers in South America are not more time-selected on spring migration than intratropical migrant conspecifics. Low sample sizes could have diminished our power to detect differences (e.g., between sexes), such that further research into the mechanisms underpinning migratory strategies in this poorly understood system is necessary.

Landscape heterogeneity is key to forecasting outcomes of plant reintroduction.

Conservation and restoration projects often involve starting new populations by introducing individuals into portions of their native or projected range. Such efforts can help meet many related goals, including habitat creation, ecosystem service provisioning, assisted migration, and the reintroduction of imperiled species following local extirpation. The outcomes of reintroduction efforts, however, are highly variable, with results ranging from local extinction to dramatic population growth; reasons for this variation remain unclear. Here, we ask whether population growth following plant reintroductions is governed by variation at two scales: the scale of individual habitat patches to which individuals are reintroduced, and larger among-landscape scales in which similar patches may be situated in landscapes that differ in matrix type, soil conditions, and other factors. Quantifying demographic variation at these two scales will help prioritize locations for introduction and, once introductions take place, forecast population growth. This work took place within a large-scale habitat fragmentation experiment, where individuals of two perennial forb species were reintroduced into eight replicate ~50-ha landscapes, each containing a set of five ~1-ha patches that varied in their degree of isolation (connected by habitat corridors or unconnected) and edge-to-area ratio. Using data on individual growth, survival, reproductive output, and recruitment collected one to two years after reintroduction, we developed models to forecast population growth, then compared forecasts to observed population sizes, three and six years later. Both the type of patch (connected and unconnected) and identity of the landscape to which individuals were reintroduced had effects on forecasted population growth rates, but only variation associated with landscape identity was an accurate predictor of subsequently observed population growth rates. Models that did not include landscape identity had minimal forecasting ability, revealing the key importance of variation at this scale for accurate prediction. Of the five demographic rates used to model population dynamics, seed production was the most important source of forecast error in population growth rates. Our results point to the importance of accounting for landscape-scale variation in demographic models and demonstrate how such models might assist with prioritizing particular landscapes for species reintroduction projects.

Disentangling fragmentation effects on herbivory in understory plants of longleaf pine savanna.

Habitat fragmentation affects species and their interactions through intertwined mechanisms that include changes to fragment area, shape, connectivity and distance to edge. Disentangling these pathways is a fundamental challenge of landscape ecology and will help identify ecological processes important for management of rare species or restoration of fragmented habitats. In a landscape experiment that manipulated connectivity, fragment shape, and distance to edge while holding fragment area constant, we examined how fragmentation impacts herbivory and growth of nine plant species in longleaf pine savanna. Probability of herbivory in open habitat was strongly dependent on proximity to forest edge for every species, increasing with distance to edge in six species (primarily grasses and annual forbs) and decreasing in three species (perennial forbs and a shrub). In the two species of perennial forbs, these edge effects were dependent on fragment shape; herbivory strongly decreased with distance to edge in fragments of two shapes, but not in a third shape. For most species, however, probability of herbivory was unrelated to connectivity or fragment shape. Growth was generally determined more strongly by leaf herbivory than by distance to edge, fragment shape, or connectivity. Taken together, these results demonstrate consistently strong edge effects on herbivory, one of the most important biotic factors determining plant growth and demography. Our results contrast with the generally inconsistent results of observational studies, likely because our experimental approach enabled us to tease apart landscape processes that are typically confounded.

Connectivity from a different perspective: comparing seed dispersal kernels in connected vs. unfragmented landscapes.

Habitat fragmentation can create significant impediments to dispersal. A technique to increase dispersal between otherwise isolated fragments is the use of corridors. Although previous studies have compared dispersal between connected fragments to dispersal between unconnected fragments, it remains unknown how dispersal between fragments connected by a corridor compares to dispersal in unfragmented landscapes. To assess the extent to which corridors can restore dispersal in fragmented landscapes to levels observed in unfragmented landscapes, we employed a stable-isotope marking technique to track seeds within four unfragmented landscapes and eight experimental landscapes with fragments connected by corridors. We studied two wind- and two bird-dispersed plant species, because previous community-based research showed that dispersal mode explains how connectivity effects vary among species. We constructed dispersal kernels for these species in unfragmented landscapes and connected fragments by marking seeds in the center of each landscape with 'IN and then recovering marked seeds in seed traps at distances up to 200 m. For the two wind-dispersed plants, seed dispersal kernels were similar in unfragmented landscapes and connected fragments. In contrast, dispersal kernels of bird-dispersed seeds were both affected by fragmentation and differed in the direction of the impact: Morella cerifera experienced more and Rhus copallina experienced less long-distance dispersal in unfragmented than in connected landscapes. These results show that corridors can facilitate dispersal probabilities comparable to those observed in unfragmented landscapes. Although dispersal mode may provide useful broad predictions, we acknowledge that similar species may respond uniquely due to factors such as seasonality and disperser behavior. Our results further indicate that prior work has likely underestimated dispersal distances of wind-dispersed plants and that factors altering long-distance dispersal may have a greater impact on the spread of species than previously thought.

Gut passage and secondary metabolites alter the source of post-dispersal predation for bird-dispersed chili seeds.

Plants can influence the source and severity of seed predation through various mechanisms; the use of secondary metabolites for chemical defense, for example, is well documented. Gut passage by frugivores can also reduce mortality of animal-dispersed seeds, although this mechanism has gained far less attention than secondary metabolites. Apart from influencing the severity of seed predation, gut passage may also influence the source of seed predation. In Bolivia, we compared impacts of these two mechanisms, gut passage and secondary metabolites, on the source of seed predation in Capsicum chacoense, a wild chili species that is polymorphic for pungency (individual plants either produce fruits and seeds containing or lacking capsaicinoids). Using physical exclosures, we isolated seed removal by insects, mammals, and birds; seeds in the trials were from either pungent or non-pungent fruits and were either passed or not passed by seed-dispersing birds. Pungency had little influence on total short-term seed removal by animals, although prior work on this species indicates that capsaicin reduces mortality caused by fungi at longer time scales. Gut passage strongly reduced removal by insects, altering the relative impact of the three predator types. The weak impact of pungency on short-term predation contrasts with previous studies, highlighting the context dependence of secondary metabolites. The strong impact of gut passage demonstrates that this mechanism alone can influence which seed predators consume seeds, and that impacts of gut passage can be larger than those of secondary metabolites, which are more commonly acknowledged as a defense mechanism.

The influence of habitat fragmentation on multiple plant-animal interactions and plant reproduction.

Despite broad recognition that habitat loss represents the greatest threat to the world's biodiyersity, a mechanistic understanding of how habitat loss and associated fragmentation affect ecological systems has proven remarkably challenging. The challenge stems from the multiple interdependent ways that landscapes change following fragmentation and the ensuing complex impacts on populations and communities of interacting species. We confronted these challenges by evaluating how fragmentation affects individual plants through interactions with animals, across five herbaceous species native to longleaf pine savannas. We created a replicated landscape experiment that provides controlled tests of three major fragmentation effects (patch isolation, patch shape [i.e., edge-to-area ratio], and distance to edge), established experimental founder populations of the five species to control for spatial distributions and densities of individual plants, and employed structural equation modeling to evaluate the effects of fragmentation on plant reproductive output and the degree to which these impacts are mediated through altered herbivory, pollination, or pre-dispersal seed predation. Across species, the most consistent response to fragmentation was a reduction in herbivory. Herbivory, however, had little impact.on plant reproductive output, and thus we found little evidence for any resulting benefit to plants in fragments. In contrast, fragmentation rarely impacted pollination or pre-dispersal seed predation, but both of these interactions had strong and consistent impacts on plant reproductive output. As a result, our models robustly predicted plant reproductive output (r2 = 0.52-0.70), yet due to the weak effects of fragmentation on pollination and pre-dispersal seed predation, coupled with the weak effect of herbivory on plant reproduction, the effects of fragmentation on reproductive output were generally small in magnitude and inconsistent. This work provides mechanistic insight into landscape-scale variation in plant reproductive success, the relative importance of plant-animal interactions for structuring these dynamics, and the nuanced nature of how habitat fragmentation can affect populations and communities of interacting species.

Habitat fragmentation and its lasting impact on Earth's ecosystems.

We conducted an analysis of global forest cover to reveal that 70% of remaining forest is within 1 km of the forest's edge, subject to the degrading effects of fragmentation. A synthesis of fragmentation experiments spanning multiple biomes and scales, five continents, and 35 years demonstrates that habitat fragmentation reduces biodiversity by 13 to 75% and impairs key ecosystem functions by decreasing biomass and altering nutrient cycles. Effects are greatest in the smallest and most isolated fragments, and they magnify with the passage of time. These findings indicate an urgent need for conservation and restoration measures to improve landscape connectivity, which will reduce extinction rates and help maintain ecosystem services.

Loss of animal seed dispersal increases extinction risk in a tropical tree species due to pervasive negative density dependence across life stages.

Overhunting in tropical forests reduces populations of vertebrate seed dispersers. If reduced seed dispersal has a negative impact on tree population viability, overhunting could lead to altered forest structure and dynamics, including decreased biodiversity. However, empirical data showing decreased animal-dispersed tree abundance in overhunted forests contradict demographic models which predict minimal sensitivity of tree population growth rate to early life stages. One resolution to this discrepancy is that seed dispersal determines spatial aggregation, which could have demographic consequences for all life stages. We tested the impact of dispersal loss on population viability of a tropical tree species, Miliusa horsfieldii, currently dispersed by an intact community of large mammals in a Thai forest. We evaluated the effect of spatial aggregation for all tree life stages, from seeds to adult trees, and constructed simulation models to compare population viability with and without animal-mediated seed dispersal. In simulated populations, disperser loss increased spatial aggregation by fourfold, leading to increased negative density dependence across the life cycle and a 10-fold increase in the probability of extinction. Given that the majority of tree species in tropical forests are animal-dispersed, overhunting will potentially result in forests that are fundamentally different from those existing now.

Landscape corridors can increase invasion by an exotic species and reduce diversity of native species.

Landscape corridors are commonly used to mitigate negative effects of habitat fragmentation, but concerns persist that they may facilitate the spread of invasive species. In a replicated landscape experiment of open habitat, we measured effects of corridors on the invasive fire ant, Solenopsis invicta, and native ants. Fire ants have two social forms: polygyne, which tend to disperse poorly but establish at high densities, and monogyne, which disperse widely but establish at lower densities. In landscapes dominated by polygyne fire ants, fire ant abundance was higher and native ant diversity was lower in habitat patches connected by corridors than in unconnected patches. Conversely, in landscapes dominated by monogyne fire ants, connectivity had no influence on fire ant abundance and native ant diversity. Polygyne fire ants dominated recently created landscapes, suggesting that these corridor effects may be transient. Our results suggest that corridors can facilitate invasion and they highlight the importance of considering species' traits when assessing corridor utility.

Potential negative ecological effects of corridors.

Despite many studies showing that landscape corridors increase dispersal and species richness for disparate taxa, concerns persist that corridors can have unintended negative effects. In particular, some of the same mechanisms that underlie positive effects of corridors on species of conservation interest may also increase the spread and impact of antagonistic species (e.g., predators and pathogens), foster negative effects of edges, increase invasion by exotic species, increase the spread of unwanted disturbances such as fire, or increase population synchrony and thus reduce persistence. We conducted a literature review and meta-analysis to evaluate the prevalence of each of these negative effects. We found no evidence that corridors increase unwanted disturbance or non-native species invasion; however, these have not been well-studied concerns (1 and 6 studies, respectively). Other effects of corridors were more often studied and yielded inconsistent results; mean effect sizes were indistinguishable from zero. The effect of edges on abundances of target species was as likely to be positive as negative. Corridors were as likely to have no effect on antagonists or population synchrony as they were to increase those negative effects. We found 3 deficiencies in the literature. First, despite studies on how corridors affect predators, there are few studies of related consequences for prey population size and persistence. Second, properly designed studies of negative corridor effects are needed in natural corridors at scales larger than those achievable in experimental systems. Third, studies are needed to test more targeted hypotheses about when corridor-mediated effects on invasive species or disturbance may be negative for species of management concern. Overall, we found no overarching support for concerns that construction and maintenance of habitat corridors may result in unintended negative consequences. Negative edge effects may be mitigated by widening corridors or softening edges between corridors and the matrix. Other negative effects are relatively small and manageable compared with the large positive effects of facilitating dispersal and increasing diversity of native species.

The importance of long-distance seed dispersal for the demography and distribution of a canopy tree species.

Long-distance seed dispersal (LDD) is considered a crucial determinant of tree distributions, but its effects depend on demographic processes that enable seeds to establish into adults and that remain poorly understood at large spatial scales. We estimated rates of seed arrival, germination, and survival and growth for a canopy tree species (Miliusa horsfieldii), in a landscape ranging from evergreen forest, where the species' abundance is high, to deciduous forest, where it is extremely low. We then used an individual-based model (IBM) to predict sapling establishment and to compare the relative importance of seed arrival and establishment in explaining the observed distribution of seedlings. Individuals in deciduous forest, far from the source population, experienced multiple benefits (e.g., increased germination rate and seedling survival and growth) from being in a habitat where conspecifics were almost absent. The net effect of these spatial differences in demographic processes was significantly higher estimated sapling establishment probabilities for seeds dispersed long distances into deciduous forest. Despite the high rate of establishment in this habitat, Miliusa is rare in the deciduous forest because the arrival of seeds at long distances from the source population is extremely low. Across the entire landscape, the spatial pattern of seed arrival is much more important than the spatial pattern of establishment for explaining observed seedling distributions. By using dynamic models to link demographic data to spatial patterns, we show that LDD plays a pivotal role in the distribution of this tree in its native habitat.

How fragmentation and corridors affect wind dynamics and seed dispersal in open habitats.

Determining how widespread human-induced changes such as habitat loss, landscape fragmentation, and climate instability affect populations, communities, and ecosystems is one of the most pressing environmental challenges. Critical to this challenge is understanding how these changes are affecting the movement abilities and dispersal trajectories of organisms and what role conservation planning can play in promoting movement among remaining fragments of suitable habitat. Whereas evidence is mounting for how conservation strategies such as corridors impact animal movement, virtually nothing is known for species dispersed by wind, which are often mistakenly assumed to not be limited by dispersal. Here, we combine mechanistic dispersal models, wind measurements, and seed releases in a large-scale experimental landscape to show that habitat corridors affect wind dynamics and seed dispersal by redirecting and bellowing airflow and by increasing the likelihood of seed uplift. Wind direction interacts with landscape orientation to determine when corridors provide connectivity. Our results predict positive impacts of connectivity and patch shape on species richness of wind-dispersed plants, which we empirically illustrate using 12 y of data from our experimental landscapes. We conclude that habitat fragmentation and corridors strongly impact the movement of wind-dispersed species, which has community-level consequences.

Using historical and experimental data to reveal warming effects on ant assemblages.

Historical records of species are compared with current records to elucidate effects of recent climate change. However, confounding variables such as succession, land-use change, and species invasions make it difficult to demonstrate a causal link between changes in biota and changes in climate. Experiments that manipulate temperature can overcome this issue of attribution, but long-term impacts of warming are difficult to test directly. Here we combine historical and experimental data to explore effects of warming on ant assemblages in southeastern US. Observational data span a 35-year period (1976-2011), during which mean annual temperatures had an increasing trend. Mean summer temperatures in 2010-2011 were ∼ 2.7 °C warmer than in 1976. Experimental data come from an ongoing study in the same region, for which temperatures have been increased ∼ 1.5-5.5 °C above ambient from 2010 to 2012. Ant species richness and evenness decreased with warming under natural but not experimental warming. These discrepancies could have resulted from differences in timescales of warming, abiotic or biotic factors, or initial species pools. Species turnover tended to increase with temperature in observational and experimental datasets. At the species level, the observational and experimental datasets had four species in common, two of which exhibited consistent patterns between datasets. With natural and experimental warming, collections of the numerically dominant, thermophilic species, Crematogaster lineolata, increased roughly two-fold. Myrmecina americana, a relatively heat intolerant species, decreased with temperature in natural and experimental warming. In contrast, species in the Solenopsis molesta group did not show consistent responses to warming, and Temenothorax pergandei was rare across temperatures. Our results highlight the difficulty of interpreting community responses to warming based on historical records or experiments alone. Because some species showed consistent responses to warming based on thermal tolerances, understanding functional traits may prove useful in explaining responses of species to warming.

When condition trumps location: seed consumption by fruit-eating birds removes pathogens and predator attractants.

Seed ingestion by frugivorous vertebrates commonly benefits plants by moving seeds to locations with fewer predators and pathogens than under the parent. For plants with high local population densities, however, movement from the parent plant is unlikely to result in 'escape' from predators and pathogens. Changes to seed condition caused by gut passage may also provide benefits, yet are rarely evaluated as an alternative. Here, we use a common bird-dispersed chilli pepper (Capsicum chacoense) to conduct the first experimental comparison of escape-related benefits to condition-related benefits of animal-mediated seed dispersal. Within chilli populations, seeds dispersed far from parent plants gained no advantage from escape alone, but seed consumption by birds increased seed survival by 370% - regardless of dispersal distance - due to removal during gut passage of fungal pathogens and chemical attractants to granivores. These results call into question the pre-eminence of escape as the primary advantage of dispersal within populations and document two overlooked mechanisms by which frugivores can benefit fruiting plants.

Habitat patch shape, not corridors, determines herbivory and fruit production of an annual plant.

Habitat corridors confer many conservation benefits by increasing movement of organisms between habitat patches, but the benefits for some species may exact costs for others. For example, corridors may increase the abundance of consumers in a habitat to the detriment of the species they consume. In this study we assessed the impact of corridors on insect herbivory of a native plant, Solanum americanum, in large-scale, experimentally fragmented landscapes. We quantified leaf herbivory and assessed fruit production as a proxy for plant fitness. We also conducted field surveys of grasshoppers (Orthoptera), a group of abundant, generalist herbivores that feed on S. americanum, and we used exclosure cages to explicitly link grasshopper herbivory to fruit production of individual S. americanum. The presence of corridors did not increase herbivory or decrease plant fruit production. Likewise, corridors did not increase grasshopper abundance. Instead, patches in our landscapes with the least amount of edge habitat and the greatest amount of warmer "core" area had the highest levels of herbivory, the largest cost to plant fruit production as a result of herbivory, and the most grasshoppers. Thus habitat quality, governed by patch shape, can be more important than connectivity for determining levels of herbivory and the impact of herbivory on plant fitness in fragmented landscapes.