Jill Jankowski

Squeezed at the top: Interspecific aggression may constrain elevational ranges in tropical birds

Tropical montane species are characterized by narrow elevational distributions. Recent perspectives on mechanisms maintaining these restricted distributions have emphasized abiotic processes, but biotic processes may also play a role in their establishment or maintenance. One historically popular hypothesis, especially for birds, is that interspecific competition constrains ranges of closely related species that replace each other along elevational gradients. Supporting evidence, however, is based on patterns of occurrence and does not reveal potential mechanisms. We experimentally tested a prediction of this hypothesis in two genera of tropical songbirds, Catharus (Turdidae) and Henicorhina (Troglodytidae), in which species have nonoverlapping elevational distributions. Using heterospecific playback trials, we found that individuals at replacement zones showed aggressive territorial behavior in response to songs of congeners. As distance from replacement zones increased, aggression toward congener song decreased, suggesting a learned component to interspecific aggression. Additionally, aggressive responses in Catharus were asymmetric, indicating interspecific dominance. These results provide experimental evidence consistent with the hypothesis that interspecific competitive interactions restrict ranges of Neotropical birds. Our results also underscore the need to consider biotic processes, such as competition, when predicting how species ranges will shift with climate change. Asymmetric aggression could be particularly important. For example, if warming in montane landscapes allows upslope range expansion by dominant competitors, then high-elevation subordinate species could be forced into progressively smaller mountaintop habitats, jeopardizing viability of their populations.

Beta diversity along environmental gradients: implications of habitat specialization in tropical montane landscapes

1. Understanding how species in a diverse regional pool are spatially distributed with respect to habitat types is a longstanding problem in ecology. Tropical species are expected to be specialists along environmental gradients, and this should result in rapid compositional change (high beta diversity) across landscapes, particularly when alpha diversity is a small fraction of regional diversity. Corollary challenges are then to identify controlling environmental variables and to ask whether species cluster into discrete community types along a gradient. 2. We investigated patterns of avian species distributions in the Tilarán mountains of Costa Rica between 1000 m and 1700 m elevation where a strong moisture gradient exists. High beta diversity was found with both auditory counts adjusted for detectability and extensive capture data, revealing nearly complete change in community composition over a few kilometres on the Pacific slope. As predicted, this beta diversity was roughly twice as high as on temperate mountainsides. 3. Partial Mantel analyses and canonical correspondence analysis indicate that change in species composition is highly correlated with change in moisture (and correlated epiphyte cover) at different distances from the continental divide on the Pacific slope. Altitude was not a good predictor of change in species composition, as species composition varies substantially among sites at the same elevation. 4. Detrended correspondence analysis and cluster analysis revealed a zone of rapid transition separating a distinct cloud forest community from rainshadow forest. On the Caribbean slope, where a shallower moisture gradient was predicted to result in lower beta diversity, we found lower rates of compositional change and more continuous species turnover. 5. Results suggest that habitat specialization of birds is likely a strong ecological force generating high beta diversity in montane landscapes. Despite overall rapid rates of species turnover, zones of relatively coherent composition could be identified. 6. Landscapes with such high beta diversity are common in the tropics, although little studied. They offer high benefit/cost opportunities for conservation, particularly as climate change threatens to alter the species composition of communities of habitat specialists.

Urban mockingbirds quickly learn to identify individual humans.

Practically all animals are affected by humans, especially in urban areas. Although most species respond negatively to urbanization, some thrive in human-dominated settings. A central question in urban ecology is why some species adapt well to the presence of humans and others do not. We show that Northern Mockingbirds (Mimus polyglottos) nesting on the campus of a large university rapidly learn to assess the level of threat posed by different humans, and to respond accordingly. In a controlled experiment, we found that as the same human approached and threatened a nest on 4 successive days, mockingbirds flushed from their nest at increasingly greater distances from that human. A different human approaching and threatening the nest identically on the fifth day elicited the same response as the first human on the first day. Likewise, alarm calls and attack flights increased from days 1–4 with the first human, and decreased on day 5 with the second human. These results demonstrate a remarkable ability of a passerine bird to distinguish one human from thousands of others. Also, mockingbirds learned to identify individual humans extraordinarily quickly: after only 2 30-s exposures of the human at the nest. More generally, the varying responses of mockingbirds to intruders suggests behavioral flexibility and a keen awareness of different levels of threat posed by individuals of another species: traits that may predispose mockingbirds and other species of urban wildlife to successful exploitation of human-dominated environments.

Basal metabolism in tropical birds: latitude, altitude, and the ‘pace of life’

Summary nLife history varies across latitudes, with the ‘pace of life’ being ‘slower’ in tropical regions. Because life history is coupled to energy metabolism via allocation tradeoffs and links between performance capacity and energy use, low metabolic intensity is expected in tropical animals. Low metabolism has been reported for lowland tropical birds, but it is unclear if this is due to ‘slow’ life history or to a warm, stable environment. nWe measured basal metabolic rates (BMR) of 253 bird species across a 2·6xa0km altitude gradient in Peru. We predicted higher BMR at high altitude due to lower temperatures leading to elevated thermoregulatory costs. We also tested for BMR differences between widely separated tropical regions (Peru and Panama), and between tropical- and temperate-breeding birds. nWe found no effect of altitude on BMR in Peruvian species and no difference in BMR between Peruvian and Panamanian birds, suggesting that BMR in Neotropical birds is consistent and independent of environmental temperature. In a data set encompassing more than 500 species, tropical birds had significantly lower BMR than temperate-breeding birds. nIn contrast to several recent analyses, we found higher BMR in passerine birds than in non-passerines, independent of breeding latitude. nBreeding latitude affects BMR, but diversity in avian life history within and between temperate and tropical regions may explain some of the residual variation in BMR after accounting for body mass and breeding latitude. Future studies of links between life history, metabolism and environmental factors might benefit from examining these variables within individual species as well as across broad geographic contrasts.

Urbanized landscapes favored by fig‐eating birds increase invasive but not native juvenile strangler fig abundance

Propagule pressure can determine the success or failure of invasive plant range expansion. Range expansion takes place at large spatial scales, often encompassing many types of land cover, yet the effect of landscape context on propagule pressure remains largely unknown. Many studies have reported a positive correlation between invasive plant abundance and human land use; increased propagule pressure in these landscapes may be responsible for this correlation. We tested the hypothesis that increased rates of seed dispersal by fig-eating birds, which are more common in urban habitats, result in an increase in invasive strangler fig abundance in landscapes dominated by human land use. We quantified abundance of an invasive species (Ficus microcarpa) and a native species (F. aurea) of strangler fig in plots spanning the entire range of human land use in South Florida, USA, from urban parking lots to native forest. We then compared models that predicted juvenile fig abundance based on distance to adult fig seed sources and fig-eating bird habitat quality with models that lacked one or both of these terms. The best model for juvenile invasive fig abundance included both distance to adult and fig-eating bird habitat terms, suggesting that landscape effects on invasive fig abundance are mediated by seed-dispersing birds. In contrast, the best model for juvenile native fig abundance included only presence/absence of adults, suggesting that distance from individual adult trees may have less effect on seed limitation for a native species compared to an invasive species undergoing range expansion. However, models for both species included significant effects of adult seed sources, implying that juvenile abundance is limited by seed arrival. This result was corroborated by a seed addition experiment that indicated that both native and invasive strangler figs were strongly seed limited. Understanding how landscape context affects the mechanisms of plant invasion may lead to better management techniques. Our results suggest that prioritizing removal of adult trees in sites with high fig-eating bird habitat may be the most effective method to control F. microcarpa abundance.

Integrating Biogeography with Contemporary Niche Theory

There is no consensus on when biotic interactions impact the range limits of species. Starting from MacArthurs use of invasibility to understand how biotic interactions influence coexistence, here we examine how biotic interactions shape species distributions. Range limits emerge from how birth, death, and movement rates vary with the environment. We clarify some basic issues revolving around niche definitions, illustrated with simple resource-consumer theory. We then highlight two different avenues for linking community theory and range theory; the first based on calculating the effects of biotic interactions on range limits across scales and landscape configurations, and the second based on aggregate measures of diffuse interactions and network strength. We conclude with suggestions for a future research agenda.

Do thermoregulatory costs limit altitude distributions of Andean forest birds

Summary nAlong tropical mountains, species often occupy narrow altitude ranges. Numerous biotic and abiotic factors have been proposed as determinants of altitude occupancy. nWe measured several aspects of thermal physiology of 215 bird species across a 2·6-km altitude gradient in the Peruvian Andes. We predicted that highland species would show adaptation to the colder high-altitude climate and that energy costs of thermoregulation might limit upslope dispersal of lowland natives. nWe found reductions in thermal conductance, body temperature and lower critical temperature in highland birds compared with lowland species. These combine to make highland natives more resistant to heat loss. nWe did not find convincing evidence that acute thermal limits or energy costs of thermoregulation constrained altitude distributions. Heat-budget models predicted low-to-moderate long-term costs at native altitudes. Costs increased for lowland natives modelled in the highland climate, but for all but a few species, costs remained within putative expenditure limits. nAlthough we did not test heat tolerances, we measured all species at temperatures similar to the hottest air temperatures at the lowland site. There was no evidence that high lowland temperatures preclude downslope movements of highland birds. nWhile thermal tolerances probably do not directly determine altitude occupancy by most species, the additional energy cost of thermoregulation experienced by lowland species moving upslope may trade off against investment in important life-history components such as breeding, and thereby affect altitude range limits.

An alpine-breeding songbird can adjust dawn incubation rhythms to annual thermal regimes

ABSTRACT Small-bodied birds engaging in incubation by a single sex experience a tradeoff between incubating to create a buffered thermal environment for their eggs and foraging to meet their own energetic requirements. This tradeoff is intensified in alpine environments, which are characterized by cold and variable conditions. We monitored the incubation rhythms of alpine Horned Larks (Eremophila alpestris) in British Columbia, Canada, across different annual thermal regimes (2005: moderate; 2006: warm; 2010: cold overnight; 2011: cold during the day). In this species, females alone incubated and left their nests to forage at dawn, following 7 hr of nighttime incubation in near-freezing conditions. However, with early morning ambient temperatures still <5°C, this placed embryos at high risk of chilling during incubation recesses. Focusing on behavioral decisions made by females at dawn (06:00–08:00 hours), we examined relationships between incubation rhythms and ambient temperature among years for evidence of variable responses to temperature. In all years, females spent more time off the nest at dawn in warmer temperatures, but in 2010, which was colder overnight, the slope of the line relating attentiveness to ambient temperature was steeper, indicating that females left their nests at colder temperatures compared with other years. In 2010 females also took shorter recesses at cold temperatures. Hatching success remained high in 2010 relative to warm or moderate years; however, overwinter survival of females declined to 48% from 2010 to 2011 compared with 72% in earlier years. When faced with exceptional thermal constraints, alpine Horned Larks made behavioral adjustments to their incubation rhythms and were able to maintain fecundity. However, potential survival costs to females implies a shift in balance of the parent–offspring tradeoff, revealing limits to coping mechanisms of alpine-breeding Horned Larks.

Interspecific interactions and range limits: contrasts among interaction types

There is a great deal of interest in the effects of biotic interactions on geographic distributions. Nature contains many different types of biotic interactions (notably mutualism, commensalism, predation, amensalism, and competition), and it is difficult to compare the effects of multiple interaction types on species’ distributions. To resolve this problem, we analyze a general, flexible model of pairwise biotic interactions that can describe all interaction types. In the absence of strong positive feedback, a species’ ability to be present depends on its ability to increase in numbers when it is rare and the species it is interacting with is at equilibrium. This insight leads to counterintuitive conclusions. Notably, we often predict the same range limit when the focal species experiences competition, predation, or amensalism. Similarly, we often predict the same range margin or when the species experiences mutualism, commensalism, or benefits from prey. In the presence of strong positive density-dependent feedback, different species interactions produce different range limits in our model. In all cases, the abiotic environment can indirectly influence the impact of biotic interactions on range limits. We illustrate the implications of this observation by analyzing a stress gradient where biotic interactions are harmful in benign environments but beneficial in stressful environments. Our results emphasize the need to consider the effects of all biotic interactions on species’ range limits and provide a systematic comparison of when biotic interactions affect distributions.

Relative abundance, habitat use, and long-term population changes of wintering and resident landbirds on St. John, U.S. Virgin Islands

Abstract St. John, U.S. Virgin Islands, is one of the most forested islands in the West Indies and provides an opportunity to conserve both resident birds and wintering neotropical migrants. We conducted double-observer point counts of landbirds in December 2005 and 2006 in Forest Inventory and Analysis (FIA) plots and National Park Service (NPS) trails in Virgin Islands National Park (VINP) to assess population trends of birds in subtropical dry and moist forests. We recorded 2,270 individual birds representing 35 species at 150 point count stations in 2005, and 3,092 individuals of 32 species at 143 of these stations in 2006. The increase in birds per point from 2005 (15.1) to 2006 (21.6) was due to resident species, 17 of which were recorded more frequently in 2006. The 17 species of neotropical migrants composed 11.8% of all registrations in 2005 and 2006. Subtropical moist and dry forest habitats differed strongly in vegetation characteristics and plant species, but no species of birds exhibited a strong affiliation with either habitat type on FIA plots. Data from NPS trails showed that most migrant species were detected more often in moist, mature forest. The resident Bridled Quail-Dove (Geotrygon mystacea) also was correlated with mature forest. Plant and bird species co-occurrence with positive correlations that may carry a signal of preferred frugivory included Guettarda odorata (Rubiaceae) with Bridled Quail-Dove, and Myrciaria floribunda (Myrtaceae) with Pearly-eyed Thrasher (Margarops fuscatus). Migrant species did not exhibit strong long-term changes in relative abundance since founding of VINP in 1957, but four open-country resident species declined significantly between 1957 and 2006 as the forest matured. Forest maturation should continue on St. John, yielding a bright future for most of its landbirds barring catastrophic hurricanes, pathogens, or invasive plants.