John T. Rotenberry

Combined effects of landscape composition and heterogeneity on farmland avian diversity

Abstract Conserving biodiversity on farmland is an essential element of worldwide efforts for reversing the global biodiversity decline. Common approaches involve improving the natural component of the landscape by increasing the amount of natural and seminatural habitats (e.g., hedgerows, woodlots, and ponds) or improving the production component of the landscape by increasing the amount of biodiversity‐friendly crops. Because these approaches may negatively impact on economic output, it was suggested that an alternative might be to enhance the diversity (compositional heterogeneity) or the spatial complexity (configurational heterogeneity) of land cover types, without necessarily changing composition. Here, we develop a case study to evaluate these ideas, examining whether managing landscape composition or heterogeneity, or both, would be required to achieve conservation benefits on avian diversity in open Mediterranean farmland. We surveyed birds in farmland landscapes of southern Portugal, before (1995–1997) and after (2010–2012) the European Unions Common Agricultural Policy (CAP) reform of 2003, and related spatial and temporal variation in bird species richness to variables describing the composition, and the compositional and configurational heterogeneity, of the natural and production components of the landscape. We found that the composition of the production component had the strongest effects on avian diversity, with a particularly marked effect on the richness of farmland and steppe bird species. Composition of the natural component was also influential, mainly affecting the richness of woodland/shrubland species. Although there were some effects of compositional and configurational heterogeneity, these were much weaker and inconsistent than those of landscape composition. Overall, we suggest that conservation efforts in our area should focus primarily on the composition of the production component, by striving to maximize the prevalence of biodiversity‐friendly crops. This recommendation probably applies to other areas such as ours, where a range of species of conservation concern is strongly associated with crop habitats.

Alternative Reproductive Tactics Arising from a Continuous Behavioral Trait: Callers versus Satellites in Field Crickets

Alternative reproductive tactics may arise when natural enemies use sexual signals to locate the signaler. In field crickets, elevated costs to male calling due to acoustically orienting parasitoid flies create opportunity for an alternative tactic, satellite behavior, where noncalling males intercept females attracted to callers. Although the caller-satellite system in crickets that risk detection by parasitoids resembles distinct behavioral phenotypes, a male’s propensity to behave as caller or satellite can be a continuously variable trait over several temporal scales, and an individual may pursue alternate tactics at different times. We modeled a caller-satellite-parasitoid system as a spatially explicit interaction among male and female crickets using individual-based simulation. Males varied in their propensity to call versus behave as a satellite from one night to the next. We varied mortality, density, sex ratio, and female mating behavior, and recorded lifetime number of mates as a function of a male’s probability of calling (vs. acting as a satellite) along a gradient in parasitism risk. Frequently, the optimal behavior switched abruptly from being pure caller (call every night) to pure satellite (never call) as parasitism rate increased. However, mixed strategies prevailed even with high parasitism risk under conditions of higher background mortality rate, decreasing density, increasing female-biased sex ratio, and increasing female choosiness. In natural populations, high parasitoid pressure alone would be unlikely to yield fixation of pure satellite behavior.

Using beta diversity to inform agricultural policies and conservation actions on Mediterranean farmland

Summary Spatial variation in species composition (β-diversity) is an important component of farmland biodiversity, which together with local richness (α-diversity) drives the number of species in a region (γ-diversity). However, β-diversity is seldom used to inform conservation, due to limited understanding of its responses to agricultural management, and lack of clear links between β-diversity changes and conservation outcomes. We explored the value of β-diversity to guide conservation on farmland, by quantifying the contribution of bird α- and β-diversity to γ-diversity variation in low- and high-intensity Mediterranean farmland, before (1995–1997) and after (2010–2012) the Common Agricultural Policy reform of 2003. We further related β-diversity to landscape heterogeneity, and assessed the conservation significance of β-diversity changes. In 1995–1997, bird diversity was highest in low-intensity farmland, where it further increased in 2010–2012 due to a strong positive contribution of α-diversity to γ-diversity. In high-intensity farmland, diversity converged over time to much the same values of low-intensity farmland, with strong positive contributions of both α- and β-diversity. These patterns were largely consistent for total, farmland and species of European conservation concern assemblages, and less so for steppe birds. Beta diversity increased with landscape heterogeneity, particularly related to spatial gradients from agricultural to natural habitats in low-intensity farmland, and from annual to permanent crops (olive groves) in high-intensity farmland. The first gradient was associated with the replacement of steppe birds of high conservation concern by more generalist species, while the second was associated with the replacement between species with lower or higher affinity for woodland and shrubland habitats. Synthesis and applications. In low-intensity farmland, spatial variation in species composition (β-diversity) was largely stable over time, reflecting a positive conservation outcome related to persistence of landscape heterogeneity patterns required by endangered steppe bird species. In contrast, β-diversity in high-intensity farmland was favoured by increases in landscape heterogeneity driven by olive grove expansion, contributing to enhancement of total bird diversity. Overall, our results stress the value of β-diversity to understand impacts of agricultural policies and conservation actions, but also highlight the need to evaluate β-diversity changes against specific conservation goals.

Effects of parents and Brown-headed Cowbirds (Molothrus ater) on nest predation risk for a songbird.

Nest predation limits avian fitness, so ornithologists study nest predation, but they often only document patterns of predation rates without substantively investigating underlying mechanisms. Parental behavior and predator ecology are two fundamental drivers of predation rates and patterns, but the role of parents is less certain, particularly for songbirds. Previous work reproduced microhabitat-predation patterns experienced by Yellow Warblers (Setophaga petechia) in the Mono Lake basin at experimental nests without parents, suggesting that these patterns were driven by predator ecology rather than predator interactions with parents. In this study, we further explored effects of post-initiation parental behavior (nest defense and attendance) on predation risk by comparing natural versus experimental patterns related to territory density, seasonal timing of nest initiation, and nest age. Rates of parasitism by Brown-headed Cowbirds (Molothrus ater) were high in this system (49% nests parasitized), so we also examined parasitism-predation relationships. Natural nest predation rates (NPR) correlated negatively with breeding territory density and nonlinearly (U-shaped relationship) with nest-initiation timing, but experimental nests recorded no such patterns. After adjusting natural-nest data to control for these differences from experimental nests other than the presence of parents (e.g., defining nest failure similarly and excluding nestling-period data), we obtained similar results. Thus, parents were necessary to produce observed patterns. Lower natural NPR compared with experimental NPR suggested that parents reduced predation rates via nest defense, so this parental behavior or its consequences were likely correlated with density or seasonal timing. In contrast, daily predation rates decreased with nest age for both nest types, indicating this pattern did not involve parents. Parasitized nests suffered higher rates of partial predation but lower rates of complete predation, suggesting direct predation by cowbirds. Explicit behavioral research on parents, predators (including cowbirds), and their interactions would further illuminate mechanisms underlying the density, seasonal, and nest age patterns we observed.

Alternative Reproductive Tactics in Context: How Demography, Ecology, and Behavior Affect Male Mating Success

Exploitation of sexual signals by predators or parasites increases costs to signalers, creating opportunities for establishment of alternative reproductive tactics (ARTs). In field crickets, males calling may attract acoustically orienting parasitoid flies. Alternatively, males behaving as satellites forgo calling and attempt to intercept females attracted to callers. We modeled the contribution of calling versus satellite behavior to male reproductive success in the larger context of variation in ecology (parasitism rate, background mortality), demography (density, sex ratio), and female behavior (phonotaxis, mating choosiness). Male mating success was most influenced by number of females (standardized effect size 0.42), followed by female choosiness (0.33), background mortality (−0.31), number of males (−0.28), and parasitism rate (−0.21). The smallest effects were phonotaxis (0.10) and satellite behavior (−0.09). Although satellite behavior ameliorated negative effects of parasitism, its comparative effect was slight. ARTs seem most likely to evolve and persist when a single selection pressure on signaling is particularly strong.

Elevation and latitude interact to drive life-history variation in precocial birds: a comparative analysis using galliformes.

Elevational gradients provide a powerful laboratory for understanding the environmental and ecological drivers of geographic variation in avian life-history strategies. Environmental variation across elevational gradients is hypothesized to select for a trade-off of reduced fecundity (lower clutch size and/or fewer broods) for higher offspring quality (larger eggs and/or increased parental care) in higher elevation species and populations. In birds, a focus on altricial species from north temperate latitudes has prevented an evaluation of the generality of this trade-off, and how it is affected by latitude and intrinsic factors (development mode). We performed a comparative analysis controlling for body size and phylogenetic relationships on a global data set of 135 galliform species to test (i) whether higher elevation precocial species have lower fecundity (smaller clutch and/or fewer broods) and invest more in offspring quality (greater egg mass) and (ii) whether latitude influences the traits involved and/or the trade-off, and (iii) to identify ecological and environmental drivers of life-history variation along elevational gradients. Life-history traits showed significant interaction effects across elevation and latitude: temperate higher elevation species had smaller clutches and clutch mass, larger eggs and shorter incubation periods, whereas more tropical species had larger clutches, eggs and clutch mass, and longer incubation periods as elevation increased. Number of broods and body mass did not vary with elevation or latitude. Latitudinal gradient in clutch size was observed only for low-elevation species. Significantly, an overlooked latitude-by-elevation interaction confounds our traditional view of clutch size variation across a tropical-to-temperate gradient. Across all latitudes, higher elevation species invested in offspring quality via larger eggs but support for reduced fecundity resulting from smaller clutches was found only along temperate elevational gradients; contrary to expectations, tropical high-elevation species showed increased fecundity. Variation in nest predation risk could explain differences between temperate and tropical elevational gradients, but we lack a consistent mechanism to explain why predation risk should vary in this manner. Alternatively, a resource availability hypothesis based on physical attributes that globally differ between elevation and latitude (seasonality in day length and temperature) seems more plausible.

Mismatches between habitat preferences and risk avoidance for birds in intensive Mediterranean farmland

Land use intensification may create habitats that organisms perceive as suitable, but where reproduction or survival is insufficient to maintain self-sustaining populations. Such conditions may qualify as ecological traps, but their existence is often hard to prove. Here, we provide a practical framework to evaluate a potential ecological trap resulting from mismatch between habitat preferences and predation risk, focusing on ground-nesting farmland birds of conservation concern. The framework is based on species-specific associations with safe or unsafe habitat types (i.e. field and landscape types with high or low nest survival), and the occurrence of risk avoidance (i.e. negative responses to predator abundances or to nest failure rates after controlling for habitat effects). Bird densities were far more influenced by field characteristics than landscape context. Corn bunting and fan-tailed warbler were associated with tall swards (safe habitats), and did not show risk avoidance. Tawny pipit and and Galerida larks were associated with short swards (unsafe habitats), with the former avoiding fields with high nest predation rates, and the later avoiding high mongoose abundances. Short-toed lark was associated with fields with short swards and low nest trampling rates. Results suggest that short-toed lark may be the most vulnerable to ecological trapping, because it nests on unsafe habitats and did not show predation risk avoidance. Our approach provides a practical first step to infer vulnerability to a potential ecological trap, though further research is needed to confirm this effect. Management actions increasing nest survival in short sward fields will likely favour grassland bird conservation in intensive Mediterranean farmland.

Social behavior and cooperative breeding in a precocial species: The Kalij Pheasant (Lophura leucomelanos) in Hawaii

ABSTRACT We examined social behavior and documented cooperative breeding in Kalij Pheasants (Lophura leucomelanos) in an introduced population in Hawaii, only the third reported instance of cooperative breeding in Phasianidae. Birds in 31 distinct social groups occupied overlapping home ranges, and group composition remained relatively constant over the 3-yr study period. Each social group contained 1 breeding female and 1 to 6 males. One male was socially dominant in each multiple-male group. Age was the only factor found to determine within-group dominance, suggesting that subordinate males may gain dominance and breeding status by staying in the group. All adults exhibited cooperative behavior, including caring for chicks, agonistic behaviors against conspecific intruders, and vigilance against predators. Average population density was high (3.21 residents per ha), which may lead to habitat saturation in this population. The adult sex ratio was male-biased, with an average M:F ratio of 2.1:1.0. Genetic sex identification of egg samples revealed unbiased primary and secondary sex ratios, which suggests that the bias in adult sex ratio may be caused by differential survival and/or dispersal. Paternity analysis of 13 broods revealed that ~68% of offspring were fathered by the dominant male of the social group, while ~17% were fathered by subordinate males in the group, suggesting that helpers gained some direct benefit by sharing reproduction. We suggest that cooperative breeding may be more common in precocial species than conventionally recognized.