Evolutionary and Ecological Processes Underlying Geographic Variation in Innate Bird Songs.

AbstractEcological and evolutionary processes underlying spatial variation in signals involved in mate recognition and reproductive isolation are crucial to understanding the causes of population divergence and speciation. Here, to test hypotheses concerning the causes of song divergence, we examine how songs of two sister species of Atlantic Forest suboscine birds with innate songs, the Pyriglena fire-eye antbirds, vary across their ranges. Specifically, we evaluated the influence of isolation by distance and introgressive hybridization, as well as morphological and environmental variation, on geographic variation in male songs. Analyses based on 496 male vocalizations from 63 locations across a 2,200-km latitudinal transect revealed clinal changes in the structure of songs and showed that introgressive hybridization increases both the variability and the homogenization of songs in the contact zone between the two species. We also found that isolation by distance, morphological constraints, the environment, and genetic introgression independently predicted song variation across geographic space. Our study shows the importance of an integrative approach that investigates the roles of distinct ecological and evolutionary processes that influence acoustic signal evolution.

Acoustic monitors and direct observations provide similar but distinct perspectives on bird assemblages in a lowland forest of eastern Ecuador.

Bird communities in lowland Neotropical forests exhibit temporal and spatial variation in species composition and abundance at multiple scales. Detecting and explaining such variation requires adequate methods for sampling those bird communities but counting birds in highly diverse lowland forests of the Neotropics can be particularly challenging. Point counts are one of the most frequently used methods for counting birds in tropical forests but inter- and intra-observer variability in detecting and identifying sounds may cause problems. Acoustic monitors (passive acoustic monitors; autonomous recording units) provide an alternative and potentially effective method to sample bird communities by acting, in effect, as "point counts", recording vocalizations at a given point for a set time. I used acoustic monitors to examine patterns of species richness, spatial distribution, and community composition of birds in a lowland forest in eastern Ecuador, one of the most diverse regions on earth. I deployed monitors at 25 locations, each separated by at least 200 m, on each of two 100-ha plots (Harpia, Puma) at Tiputini Biodiversity Station during January-February, 2013-2017. Monitors were set to record for 10 min followed by a 5-min break, from 0545 h to 0810 h (10 recording periods/morning). Recordings were later reviewed to identify species; no attempt was made to distinguish individuals or to estimate distance. Results were compared with contemporaneous direct observations along transects on the same plots. A total of 214 species were identified from recordings on both plots, combined, with slightly more on Harpia (208) than on Puma (188). Number per year ranged from 142 on Harpia in 2016 to 161 on Puma in 2015. Number per point was ~45 with an overall range of 29-68. Number of species detected in recordings was similar to but somewhat less than the number recorded during direct observations. Number of species recorded increased rapidly from the first period (0545-0555 h) to the third (0615-0625 h) but showed little subsequent change. Most species were recorded at relatively few points; the four most widely distributed species were the same on both plots (Patagioenas plumbea, Xiphorhynchus guttatus, Capito aurita, Ramphastos tucanus), all of which are relatively loud canopy or subcanopy species. Ordinations based on species composition illustrated differences between plots based on both recordings and direct observations; similarly, patterns of species composition differed between methods. Acoustic monitors can be an effective tool for sampling bird communities and may be particularly effective and efficient for sampling loud species with distinctive songs. Nonetheless, results from monitors may provide different perspectives on species composition when compared to direct observations. Which method is preferred likely will depend on the specific objectives of individual studies.

Ecological niche differentiation in Chiroxiphia and Antilophia manakins (Aves: Pipridae).

Species distribution models are useful for identifying the ecological characteristics that may limit a species' geographic range and for inferring patterns of speciation. Here, we test a hypothesis of niche conservatism across evolutionary time in a group of manakins (Aves: Pipridae), with a focus on Chiroxiphia boliviana, and examine the degree of ecological differentiation with other Chiroxiphia and Antilophia manakins. We tested whether allopatric sister species were more or less similar in environmental space than expected given their phylogenetic distances, which would suggest, respectively, ecological niche conservatism over time or ecologically mediated selection (i.e. niche divergence). We modeled the distribution of nine manakin taxa (C. boliviana, C. caudata, C. lanceolata, C. linearis, C. p. pareola, C. p. regina, C. p. napensis, Antilophia galeata and A. bokermanni) using Maxent. We first performed models for each taxon and compared them. To test our hypothesis we followed three approaches: (1) we tested whether C. boliviana could predict the distribution of the other manakin taxa and vice versa; (2) we compared the ecological niches by using metrics of niche overlap, niche equivalency and niche similarity; and (3) lastly, we tested whether niche differentiation corresponded to phylogenetic distances calculated from two recent phylogenies. All models had high training and test AUC values. Mean AUC ratios were high (>0.8) for most taxa, indicating performance better than random. Results suggested niche conservatism, and high niche overlap and equivalency between C. boliviana and C. caudata, but we found very low values between C. boliviana and the rest of the taxa. We found a negative, but not significant, relationship between niche overlap and phylogenetic distance, suggesting an increase in ecological differentiation and niche divergence over evolutionary time. Overall, we give some insights into the evolution of C. boliviana, proposing that ecological selection may have influenced its speciation.

Variation in Neotropical river otter (Lontra longicaudis) diet: Effects of an invasive prey species.

Predation is one of the main barriers that exotic species may face in newly colonized areas and may help stop or control the potential negative impacts of invasive species in the environment. We evaluated if the consumption of an invasive prey (armored catfish: Pterygoplichtys sp.) affects the dietary niche breadth and trophic level of a native predator (Neotropical river otter: Lontra longicaudis) in northern Guatemala. We examined otter scats from three rivers: two where the invasive armored catfish occurred and one without the invasive fish. Samples were collected two and seven years after the first report of the catfish in the area. We performed gross scat analysis and stable isotope analyses of nitrogen and carbon of fecal matter. Where the invasive armored catfish occurred, it was the main prey item for L. longicaudis. Particularly in the river outside of protected areas seven years after the first report of the catfish, where it accounted for 49% of the otter diet. Concordance was found between the two techniques to estimate dietary niche breadth and trophic level. The dietary niche breath of otters was narrower seven years after the invasion in comparison to two years after the invasion in both invaded rivers, but the extent of the reduction was lesser inside the protected area. Finally, the trophic level of otters also showed a reduction related to the occurrence of the armored catfish in their diet.

Annual and spatial variation in composition and activity of terrestrial mammals on two replicate plots in lowland forest of eastern Ecuador.

Terrestrial mammals are important components of lowland forests in Amazonia (as seed dispersal agents, herbivores, predators) but there are relatively few detailed studies from areas that have not been affected by human activities (e.g., hunting, logging). Yet, such information is needed to evaluate effects of humans elsewhere. We used camera traps to sample medium to large-sized terrestrial mammals at a site in lowland forests of eastern Ecuador, one of the most biologically rich areas in the world. We deployed cameras on two study plots in terra firme forest at Tiputini Biodiversity Station. Sixteen cameras were arranged 200 m apart in a 4 × 4 grid on each plot. Cameras were operated for  60 days in January-March, 2014-2017, for a total of 3,707 and 3,482 trap-days on the two plots (Harpia, Puma). A total of 28 species were recorded; 26 on Harpia and 25 on Puma. Number of species recorded each year was slightly greater on Harpia whereas overall capture rates (images/100 trap-days) were higher on Puma. Although most species were recorded on each plot, differences in capture rates meant that yearly samples on a given plot were more similar to each other than to samples on the other plot. Images of most species showed a clumped distribution pattern on each plot; Panthera onca was the only species that did not show a clumped distribution on either plot. Images at a given camera location showed no evidence of autocorrelation with numbers of images at nearby camera locations, suggesting that species were responding to small-scale differences in habitat conditions. A redundancy analysis showed that environmental features within 50 or 100 m of camera locations (e.g., elevation, variation in elevation, slope, distance to streams) accounted for significant amounts of variation in distribution patterns of species. Composition and relative importance based on capture rates were very similar to results from cameras located along trails at the same site; similarities decreased at increasing spatial scales based on comparisons with results from other sites in Ecuador and Peru.

Host associations and turnover of haemosporidian parasites in manakins (Aves: Pipridae).

Parasites of the genera Plasmodium and Haemoproteus (Apicomplexa: Haemosporida) are a diverse group of pathogens that infect birds nearly worldwide. Despite their ubiquity, the ecological and evolutionary factors that shape the diversity and distribution of these protozoan parasites among avian communities and geographic regions are poorly understood. Based on a survey throughout the Neotropics of the haemosporidian parasites infecting manakins (Pipridae), a family of Passerine birds endemic to this region, we asked whether host relatedness, ecological similarity and geographic proximity structure parasite turnover between manakin species and local manakin assemblages. We used molecular methods to screen 1343 individuals of 30 manakin species for the presence of parasites. We found no significant correlations between manakin parasite lineage turnover and both manakin species turnover and geographic distance. Climate differences, species turnover in the larger bird community and parasite lineage turnover in non-manakin hosts did not correlate with manakin parasite lineage turnover. We also found no evidence that manakin parasite lineage turnover among host species correlates with range overlap and genetic divergence among hosts. Our analyses indicate that host switching (turnover among host species) and dispersal (turnover among locations) of haemosporidian parasites in manakins are not constrained at this scale.

Seasonal patterns in δ2 H values of multiple tissues from Andean birds provide insights into elevational migration.

Elevational migration is a widespread phenomenon in tropical avifauna but it is difficult to identify using traditional approaches. Hydrogen isotope (δ2 H) values of precipitation decrease with elevation so δ2 H analysis of multiple bird tissues with different isotopic incorporation rates may be a reliable method for characterizing seasonal elevational migration. Here we compare δ2 H values in metabolically inert (feathers and claws) and metabolically active (whole blood) tissues to examine whether an upslope migration occurs prior to the breeding season in the Yungas Manakin (Chiroxiphia boliviana). We compare results from C. boliviana with data from a known elevational migrant, the Streak-necked Flycatcher (Mionectes striaticollis). Opposite to our expectations, tissue δ2 H values increased over time, largely reflecting seasonal patterns in precipitation δ2 H rather than elevational effects; linear mixed-effects models with strongest support included ordinal date, tissue type, and elevation. This seasonal increase in precipitation δ2 H is a general phenomenon in both tropical and temperate mountain ranges. We use these data to propose a hypothetical framework that predicts different patterns in tissue δ2 H values collected in different seasons from residents and elevational migrants. This framework can serve as a reference for future studies that assess elevational migration in birds and other animals.

Resource predictability and specialization in avian malaria parasites.

We tested the hypothesis that avian haemosporidian (malaria) parasites specialize on hosts that can be characterized as predictable resources at a site in Amazonian Ecuador. We incorporated host phylogenetic relationship and relative abundance in assessing parasite specialization, and we examined associations between parasite specialization and three host characteristics - abundance, mass and longevity - using quantile regression, phylogenetic logistic regression and t-tests. Hosts of specialist malaria parasite lineages were on average more abundant than hosts of generalist parasite lineages, but the relationship between host abundance and parasite specialization was not consistent across analyses. We also found support for a positive association between parasite specialization and host longevity, but this also was not consistent across analyses. Nonetheless, our findings suggest that the predictability of a host resource may play a role in the evolution of specialization. However, we also discuss two alternative explanations to the resource predictability hypothesis for specialization: (i) that interspecific interactions among the parasites themselves might constrain some parasites to a specialist strategy, and (ii) that frequent encounters with multiple host species, mediated by blood-sucking insects, might promote generalization within this system.

Enigmatic declines in bird numbers in lowland forest of eastern Ecuador may be a consequence of climate change.

Bird populations have declined in many parts of the world but most of those declines can be attributed to effects of human activities (e.g., habitat fragmentation); declines in areas unaffected by human activities are not common. We have been sampling bird populations at an undisturbed site in lowland forest of eastern Ecuador annually since 2001 using a combination of mist nets and direct observations on two 100-ha plots. Bird numbers fluctuated on both plots during the first 8 years but did not show a consistent pattern of change. Since about 2008, numbers of birds on both plots have declined; capture rates in 2014 were ∼40% less than at the start of the study and observation rates were ∼50% less. Both understory and canopy species declined in abundance. Overall, insectivores showed the most pronounced declines but declines varied among trophic groups. The period from 2008 onward also was a period of stronger La Niña events which, at this study site, are associated with increased rainfall. The mechanism for the declines is not known but likely reflects a combination of reduced reproductive success coupled with reduced survival associated with changing climate.

Reciprocal specialization in multihost malaria parasite communities of birds: a temperate-tropical comparison.

How specialization of consumers with respect to resources varies with respect to latitude is poorly understood. Coexistence of many species in the tropics might be possible only if specialization also increases. Alternatively, lower average abundance of more diverse biotic resources in the tropics might force consumers to become more generalized foragers. We examine levels of reciprocal specialization in an antagonistic system-avian malaria-to determine whether the number of host species used and/or parasite lineages harbored differ between a temperate and a tropical assemblage. We evaluate the results of network analysis, which can incorporate both bird and parasite perspectives on specialization in one quantitative index, in comparison to null models. Specialization was significantly greater in both sample sites than predicted from null models. We found evidence for lower per-host species parasite diversity in temperate compared to tropical birds. However, specialization did not differ between the tropical and temperate sites from the parasite perspective. We supplemented the network analysis with estimates of specialization that incorporate phylogenetic relationships of associates and found no differences between sites. Thus, our analyses indicate that specialization within an antagonistic host-parasite (resource-consumer) system varies little between tropical and temperate localities.

Yasuní--a hotspot for jaguars Panthera onca (Carnivora: Felidae)? Camera-traps and jaguar activity at Tiputini Biodiversity Station, Ecuador.

Jaguars (Panthera onca) are the largest predator in lowland forests of Amazonia but there have been few studies on their occurrence and activity in such forests. Here, we used camera traps to document the occurrence and activity of jaguars within a local area (-650ha) of lowland forest of Eastern Ecuador, over two sample periods (2005-2008, 7 222 trap days; 2010-2012, 6 199 trap days). We accumulated 151 independent photos of jaguars (189 total photographs) that represented 21 different individuals, including 11 males (114 photographs), seven females (32 photographs), and three that could not be assigned to a sex. Individual jaguars varied in the number of months they were recorded in the area; ten were photographed in only one month; five were photographed over periods of 8 to 22 months; and five from 45 to 81 months. Capture rates across all camera stations averaged 10.6/1 000 trap days; capture rates did not differ between the two sample periods. Male jaguars were more active during the day (06:00am-18:00pm; 71% of photographs), whereas females were equally active during the day and night. Monthly activity was variable but showed no consistent pattern. Although the study area is much smaller than typical home ranges of jaguars, the area is clearly visited by a large number of different individuals, some of whom repeatedly visit the area, indicating that it forms part of their home range. Other individuals likely were simply passing through the area. Based on the number of jaguars recorded during this study, it is clear that the region is an important area for conservation. Continued protection will be needed to ensure that populations ofjaguars and other species remain viable.

Yasuní - a hotspot for jaguars Panthera onca (Carnivora: Felidae)? Camera-traps and jaguar activity at Tiputini Biodiversity Station, Ecuador / Yasuní - Un hotspot para jaguares Panthera onca (Carnivora: Felidae)? Cámaras trampa y actividad del jaguar en la Estación de Biodiversidad Tiputini, Ecuador

Jaguars (Panthera onca) are the largest predator in lowland forests of Amazonia but there have been few studies on their occurrence and activity in such forests. Here, we used camera traps to document the occurrence and activity of jaguars within a local area (~650ha) of lowland forest of Eastern Ecuador, over two sample periods (2005-2008, 7 222 trap days; 2010-2012, 6 199 trap days). We accumulated 151 independent photos of jaguars (189 total photographs) that represented 21 different individuals, including 11 males (114 photographs), seven females (32 photographs), and three that could not be assigned to a sex. Individual jaguars varied in the number of months they were recorded in the area; ten were photographed in only one month; five were photographed over periods of 8 to 22 months; and five from 45 to 81 months. Capture rates across all camera stations averaged 10.6/1 000 trap days; capture rates did not differ between the two sample periods. Male jaguars were more active during the day (06:00am-18:00pm; 71% of photographs), whereas females were equally active during the day and night. Monthly activity was variable but showed no consistent pattern. Although the study area is much smaller than typical home ranges of jaguars, the area is clearly visited by a large number of different individuals, some of whom repeatedly visit the area, indicating that it forms part of their home range. Other individuals likely were simply passing through the area. Based on the number of jaguars recorded during this study, it is clear that the region is an important area for conservation. Continued protection will be needed to ensure that populations of jaguars and other species remain viable. Rev. Biol. Trop. 62 (2): 689-698. Epub 2014 June 01.

El jaguar (Panthera onca) es el depredador más grande de los bosques bajos de la Amazonía, pero existen pocos estudios sobre su presencia y actividad en los mismos. En este estudio utilizamos cámaras trampa para documentar la presencia y actividad de jaguares dentro de una área de bosque bajo de la Amazonía este de Ecuador (~650 ha) en dos períodos (2005-2008, 7 222 trampas noche; 2010-2012, 6 199 trampas noche). Acumulamos 151 fotos independientes de jaguares (189 fotografías en total) que representaron 21 individuos, incluyendo 11 machos (114 fotos), siete hembras (32 fotos) y tres a los que no se les pudo asignar sexo. Los jaguares individuales variaron en el número de meses en que fueron registrados en el área; diez fueron fotografiados solo en un mes; cinco fueron fotografiados en un periodo entre 8 y 22 meses; y cinco jaguares entre 45 y 81. Las tasas de captura a través de las estaciones de trampeo promediaron 10.6/1 000 trampas noche; las tasas de captura no presentaron diferencias en los dos períodos de muestreo. Los jaguares machos fueron más activos durante el día (06:00am-18:00pm; 71% de las fotografías) mientras que las hembras fueron igualmente activas tanto en el día como en la noche. La actividad mensual fue variable pero no mostró un patrón consistente. Aunque el área de estudio es mucho más pequeña que el rango de vida típico para un jaguar, el área es claramente visitada por un alto número de individuos diferentes, algunos de los cuales visitan repetidamente el lugar, indicando que este forma parte de su rango de vida. Es probable que otros individuos solamente sean transeúntes del área.

Apparent survival rates of forest birds in eastern Ecuador revisited: improvement in precision but no change in estimates.

Knowledge of survival rates of Neotropical landbirds remains limited, with estimates of apparent survival available from relatively few sites and species. Previously, capture-mark-recapture models were used to estimate apparent survival of 31 species (30 passerines, 1 Trochilidae) from eastern Ecuador based on data collected from 2001 to 2006. Here, estimates are updated with data from 2001-2012 to determine how additional years of data affect estimates; estimates for six additional species are provided. Models assuming constant survival had highest support for 19 of 31 species when based on 12 years of data compared to 27 when based on six; models incorporating effects of transients had the highest support for 12 of 31 species compared to four when based on 12 and six years, respectively. Average apparent survival based on the most highly-supported model (based on model averaging, when appropriate) was 0.59 (± 0.02 SE) across 30 species of passerines when based on 12 years and 0.57 (± 0.02) when based on six. Standard errors of survival estimates based on 12 years were approximately half those based on six years. Of 31 species in both data sets, estimates of apparent survival were somewhat lower for 13, somewhat higher for 17, and remained unchanged for one; confidence intervals for estimates based on six and 12 years of data overlapped for all species. Results indicate that estimates of apparent survival are comparable but more precise when based on longer-term data sets; standard error of the estimates was negatively correlated with numbers of captures (rs  = -0.72) and recaptures (rs  = -0.93, P<0.001 in both cases). Thus, reasonable estimates of apparent survival may be obtained with relatively few years of data if sample sizes are sufficient.

Estimates of dietary overlap for six species of Amazonian manakin birds using stable isotopes.

We used stable isotope ratios to determine the metabolic routing fraction of carbon and nitrogen in feathers in addition to faecal analysis to estimate diet overlap of six sympatric species of manakins in the eastern lowland forest of Ecuador. Collectively, all species varied from-23.7 to-32.7 ‰ for δ(13)C, and from 6.0 to 9.9‰ for δ(15)N, with Machaeropterus regulus showing isotopic differences from the other species. We developed a mixing model that explicitly addresses the routing of carbon and nitrogen to feathers. Interestingly, these results suggest a higher proportion of nitrogen and carbon derived from insects than anticipated based on feeding observations and faecal analysis. A concentration-dependent mixing isotopic model was also used to look at dietary proportions. While larvae and arachnids had higher δ(15)N values, these two groups may also be preferred prey of manakins and may be more assimilated into tissues, leading to a potential overestimation of the contribution to diet. This study supports the finding that manakin species, previously thought be primarily frugivorous, contain a significant amount of arthropods in their diet.

Connecting fruit production to seedling establishment in two co-occurring Miconia species: consequences of seed dispersal by birds in upper Amazonia.

This study investigated links between seed production by two species of Miconia (Melastomataceae), whose seeds are dispersed by birds, and later stages of recruitment in lowland forests of eastern Ecuador. Seed dispersal and survival in later stages are crucial for understanding and predicting patterns of plant population dynamics as well as for understanding patterns of diversity in tropical forests. A major goal was to determine if the spatial template of seed deposition established by birds predicted probability of recruitment. We used observational and experimental approaches to compare patterns of recruitment in Miconia fosteri and M. serrulata. We calculated probabilities of transition between successive stages of recruitment for each species in three habitats. The number of plants with fruit, number of fruits removed, and, to a lesser extent, patterns of seed deposition varied between species and among habitats, whereas seed survival, germination, and establishment showed little variation among habitats. The location of seed deposition directly influenced the cumulative probabilities of survival. Among-habitat differences in the probabilities of recruitment set by seed deposition were not modified by later stages, although probability of recruitment was 2.5 times higher for M. serrulata than for M. fosteri after 1 year. The more critical stages for recruitment were seed removal and deposition. Our results from multiple life-cycle stages suggest that habitat associations among plants that reach reproductive maturity become established at early life stages and were mostly a consequence of seed dispersal by birds. These results differ from those obtained in temperate zones and suggest fundamental differences in the importance of recruitment processes. Dispersers, such as manakins, play significant roles in recruitment and population dynamics of M. fosteri, M. serrulata and numerous other understory plants of Neotropical forests. Their role in plant recruitment could be much greater than previously considered in megadiverse tropical forests. Thus, loss of dispersers could have long-term and far-reaching implications for maintenance of diversity.

It takes two to tango: reproductive skew and social correlates of male mating success in a lek-breeding bird.

Variance in reproductive success among individuals is a defining characteristic of many social vertebrates. Yet, our understanding of which male attributes contribute to reproductive success is still fragmentary in most cases. Male-male reproductive coalitions, where males jointly display to attract females, are of particular interest to evolutionary biologists because one male appears to forego reproduction to assist the social partner. By examining the relationship between social behaviour and reproductive success, we can elucidate the proximate function of coalitions in the context of mate choice. Here, we use data from a 4-year study of wire-tailed manakins (Pipra filicauda) to provide molecular estimates of reproductive skew and to test the hypothesis that male-male social interactions, in the context of coordinated displays, positively influence a male's reproductive success. More specifically, we quantify male-male social interactions using network metrics and predict that greater connectivity will result in higher relative reproductive success. Our data show that four out of six leks studied had significant reproductive skew, with success apportioned to very few individuals in each lek. Metrics of male social affiliations derived from our network analysis, especially male connectivity, measured as the number of males with whom the focal male has extended interactions, were strong predictors of the number of offspring sired. Thus, network connectivity is associated with male fitness in wire-tailed manakins. This pattern may be the result of shared cues used by both sexes to assess male quality, or the result of strict female choice for coordinated display behaviour.

Female mate choice across spatial scales: influence of lek and male attributes on mating success of blue-crowned manakins.

Lekking males compete for females within and among leks, yet female choice is expected to work differently at each of these spatial scales. We used paternity analyses to examine how lek versus male attributes influence mate choice in the blue-crowned manakin Lepidothrix coronata. We tested the hypotheses that females prefer (i) to mate at larger leks where a larger number of potential mates can be assessed, (ii) to mate with unrelated or highly heterozygous males expected to produce high-quality offspring, (iii) to mate with males that display at higher rates, and that (iv) display honestly reflects male genetic quality. Our results show that (i) males at larger leks are not more likely to sire young, although females nesting close to small leks travel further to reach larger leks, (ii) siring males are not less related to females or more heterozygous than expected, (iii) within a lek, high-display males are more likely to sire young, and (iv) both male heterozygosity and display rate increased with lek size, and as a result display does not reliably reflect male genetic quality across leks. We suggest that female mate choice in this species is probably driven by a Fisherian process rather than adaptive genetic benefits.

Crop size, plant aggregation, and microhabitat type affect fruit removal by birds from individual melastome plants in the Upper Amazon.

We studied the efficiency (proportion of the crop removed) and quantitative effectiveness (number of fruits removed) of dispersal of Miconia fosteri and M. serrulata (Melastomataceae) seeds by birds in lowland tropical wet forest of Ecuador. Specifically, we examined variation in fruit removal in order to reveal the spatial scale at which crop size influences seed dispersal outcome of individual plants, and to evaluate how the effect of crop size on plant dispersal success may be affected by conspecific fruit abundance and by the spatial distribution of frugivore abundance. We established two 9-ha plots in undisturbed terra-firme understory, where six manakin species (Pipridae) disperse most seeds of these two plant species. Mean levels of fruit removal were low for both species, with high variability among plants. In general, plants with larger crop sizes experienced greater efficiency and effectiveness of fruit removal than plants with smaller crops. Fruit removal, however, was also influenced by microhabitat, such as local topography and local neighborhood. Fruit-rich and disperser-rich patches overlapped spatially for M. fosteri but not M. serrulata, nonetheless fruit removal of M. serrulata was still much greater in fruit-rich patches. Fruit removal from individual plants did not decrease in patches with many fruiting conspecifics and, in fact, removal effectiveness was enhanced for M. fosteri with small crop sizes when such plants were in patches with more conspecifics. These results suggest that benefits of attracting dispersers to a patch balanced or outweighed the costs of competition for dispersers. Spatial pattern of fruit removal, a measure of plant fitness, depended on a complex interaction among plant traits, spatial patterns of plant distribution, and disperser behavior.

Social networks in the lek-mating wire-tailed manakin (Pipra filicauda).

How social structure interacts with individual behaviour and fitness remains understudied despite its potential importance to the evolution of cooperation. Recent applications of network theory to social behaviour advance our understanding of the role of social interactions in various contexts. Here we applied network theory to the social system of lek-mating wire-tailed manakins (Pipra filicauda, Pipridae, Aves). We analysed the network of interactions among males in order to begin building a comparative framework to understand where coordinated display behaviour lies along the continuum from solitary to obligately cooperative dual-male displays in the family Pipridae. Network degree (the number of links from a male to others) ranged from 1 to 10, with low mean and high variance, consistent with the theory for the evolution of cooperation within social networks. We also assessed factors that could predict social and reproductive success of males. Four network metrics, degree, eigenvector centrality, information centrality and reach, some of which assess circuitous as well as the shortest (geodesic) paths of male connectivity, predicted male social rise. The duration of a male's territorial tenure during the 4 years of the study predicted his probability of siring offspring.

Breeding Birds of Isolated Woodlots: Area and Habitat Relationships.

We investigated breeding bird communities of isolated woodlots (1.8-600 ha) in east-central Illinois during three summers (1979-1981) to compare the influence of area and habitat on community structure. Woodlots supported from 9 to 43 species and composition was relatively constant among years. Ecological generalists dominated small woodlots, while more specialized species increased in importance with area. Area accounted for most variation (86-98%) in total species number in each year and the species-area relationship did not change significantly among years. The amount of variance accounted for by area was greater than in previous studies. Neither habitat nor woodlot isolation explained significant additional variation in total species richness after area. Area accounted for most variation in number of species in different migratory and breeding habitat categories, except for short-distance migrants, which correlated most strongly with habitat. Variation in habitat was not related to woodlot area and habitat accounted for additional variation in bird species numbers in most cases. Abundances of one-third to one-half of species examined correlated with woodlot area, but a greater proportion (66-72%) were influenced more strongly by habitat variables. Results from Illinois support previous conclusions that species that breed in forest interior habitat and winter in the tropics are most likely to be adversely affected by a reduction in forest habitat. Results also show that bird communities in isolated tracts of forest are not random assemblages, but rather that species found in smaller woodlots are subsets of species found in larger forests.