Network structure of avian mixed-species flocks decays with elevation and latitude across the Andes.

Birds in mixed-species flocks benefit from greater foraging efficiency and reduced predation, but also face costs related to competition and activity matching. Because this cost-benefit trade-off is context-dependent (e.g. abiotic conditions and habitat quality), the structure of flocks is expected to vary along elevational, latitudinal and disturbance gradients. Specifically, we predicted that the connectivity and cohesion of flocking networks would (i) decline towards tropical latitudes and lower elevations, where competition and activity matching costs are higher, and (ii) increase with lower forest cover and greater human disturbance. We analysed the structure of 84 flock networks across the Andes and assessed the effect of elevation, latitude, forest cover and human disturbance on network characteristics. We found that Andean flocks are overall open-membership systems (unstructured), though the extent of network structure varied across gradients. Elevation was the main predictor of structure, with more connected and less modular flocks upslope. As expected, flocks in areas with higher forest cover were less cohesive, with better defined flock subtypes. Flocks also varied across latitude and disturbance gradients as predicted, but effect sizes were small. Our findings indicate that the unstructured nature of Andean flocks might arise as a strategy to cope with harsh environmental conditions. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

High Foraging Fidelity and Plant-Pollinator Network Dominance of Non-native Honeybees (Apis mellifera) in the Ecuadorian Andes.

Pollinator foraging fidelity (i.e., consistent and repeated visitation to a particular plant species or area) is poorly understood for most bee species, but is important information for both the conservation of plant and pollinator species and the ecosystem services they provide to humans. We used plant-pollinator surveys and mark-recapture of floral-visiting Hymenoptera to study the foraging fidelity and species interaction network properties of a plant-pollinator community in the tropical Andes of southern Ecuador. After marking 92 bees visiting six plant taxa along four 100-m transects between July 16th and July 31st of 2019, only honeybees were resighted at a recapture rate of 47.7% (41/86). During our surveys, we observed nine bee and two wasp taxa feeding from the flowers of 10 morphospecies of plants, and we found significantly low network nestedness and significantly high network-level specialization. Specialization (d') was also significant for honeybees and bumblebees and for three plant taxa. Overall, our findings indicated that feral, non-native honeybees in this region dominated the local plant-pollinator network, yet this species is acting as a specialist forager at the individual level. Our results suggest that honeybees may be replacing the pollination services of some native bees and wasps in the region, but more research is needed to determine the effectiveness of honeybee pollination for the local plants.

Temporal stability in species richness but reordering in species abundances within avian assemblages of a tropical Andes conservation hot spot.

As the pace of environmental change increases, there is an urgent need for quantitative data revealing the temporal dynamics of local communities in tropical areas. Here, we quantify the stability of avian assemblages in the highly threatened, but poorly studied, Andean biodiversity hot spot. We evaluated the temporal variation in species richness and community composition of local bird assemblages in three habitat types (native forest, introduced forest, native shrub) using a unique, relatively long-term data series from Cajas National Park and Mazán Reserve in the southern Andes of Ecuador. We sampled birds with mist nets using a standardized protocol over 11 years, from 2006 to 2016. Species richness remained stable over time across habitats, but community composition changed in the native forest. In particular, we observed taxonomic reordering in the native forest, in which the evenness in the distribution of abundances of taxa decreased over time. This finding is consistent with other studies where species richness remained constant over time while community composition changed. Our study highlights the value of long-term studies in the tropical Andes as we show that species composition of birds in a montane forest is changing, consistent with global trends in biodiversity change.

La tasa acelerada del incremento en el cambio ambiental crea la necesidad urgente de datos cuantitativos que expongan la dinámica temporal de comunidades locales en áreas tropicales. En este manuscrito, cuantificamos la estabilidad de ensambles de aves un área altamente amenazada pero poco estudiada, el hot­spot de biodiversidad Andino. Evaluamos la variación temporal en riqueza y composición del ensamble de aves local en tres tipos de hábitat (bosque nativo, bosque introducido, matorrales nativos) usando una serie de datos de largo plazo relativo, obtenidos en el Parque Nacional Cajas y la reserva Mazan, en los Andes del sur del Ecuador. Las aves se estudiaron con redes de neblina por medio de protocolos estandarizados por 11 años, desde 1996 al 2016. La riqueza de especies permaneció estable en el tiempo en todos los hábitats, pero la composición de especies cambió en el bosque nativo. En particular, observamos un reorden taxonómico en el bosque nativo, en el cual existió una diminución en la equitatividad de las distribuciones de las especies en el tiempo. Este resultado es consistente con otros estudios que encuentran que la riqueza de especies permanece constante en el tiempo mientras que la composición de la comunidad cambia. Nuestro trabajo resalta el valor de los estudios de largo tiempo en los Andes tropicales al demostrar que la composición de aves está cambiando, y concuerda con los patrones globales de cambio de la biodiversidad.

Biodiversity and ecosystem functions depend on environmental conditions and resources rather than the geodiversity of a tropical biodiversity hotspot.

Biodiversity and ecosystem functions are highly threatened by global change. It has been proposed that geodiversity can be used as an easy-to-measure surrogate of biodiversity to guide conservation management. However, so far, there is mixed evidence to what extent geodiversity can predict biodiversity and ecosystem functions at the regional scale relevant for conservation planning. Here, we analyse how geodiversity computed as a compound index is suited to predict the diversity of four taxa and associated ecosystem functions in a tropical mountain hotspot of biodiversity and compare the results with the predictive power of environmental conditions and resources (climate, habitat, soil). We show that combinations of these environmental variables better explain species diversity and ecosystem functions than a geodiversity index and identified climate variables as more important predictors than habitat and soil variables, although the best predictors differ between taxa and functions. We conclude that a compound geodiversity index cannot be used as a single surrogate predictor for species diversity and ecosystem functions in tropical mountain rain forest ecosystems and is thus little suited to facilitate conservation management at the regional scale. Instead, both the selection and the combination of environmental variables are essential to guide conservation efforts to safeguard biodiversity and ecosystem functions.

Different responses of taxonomic and functional bird diversity to forest fragmentation across an elevational gradient.

Many studies have investigated how habitat fragmentation affects the taxonomic and functional diversity of species assemblages. However, the joint effects of habitat fragmentation and environmental conditions on taxonomic and functional diversity, for instance across elevational gradients, have largely been neglected so far. In this study, we compare whether taxonomic and functional indicators show similar or distinct responses to forest fragmentation across an elevational gradient. We based our analysis on a comprehensive data set of species-rich bird assemblages from tropical montane forest in the Southern Andes of Ecuador. We monitored birds over 2 years in two habitat types (continuous and fragmented forest) at three elevations (i.e., 1000, 2000, and 3000 m a.s.l) and measured nine morphological traits for each bird species on museum specimens. Bird species richness and abundance were significantly higher in fragmented compared to continuous forests and decreased towards high elevations. In contrast, functional diversity was significantly reduced in fragmented compared to continuous forests at low elevations, but fragmentation effects on functional diversity tended to be reversed at high elevations. Our results demonstrate that taxonomic and functional indicators can show decoupled responses to forest fragmentation and that these effects are highly variable across elevations. Our findings reveal that functional homogenization in bird communities in response to fragmentation can be masked by apparent increases in taxonomic diversity, particularly in diverse communities at low elevations.

Spatio-temporal variation in bird assemblages is associated with fluctuations in temperature and precipitation along a tropical elevational gradient.

Understanding the spatial and temporal dynamics of species assemblages is a main challenge in ecology. The mechanisms that shape species assemblages and their temporal fluctuations along tropical elevational gradients are particularly poorly understood. Here, we examined the spatio-temporal dynamics of bird assemblages along an elevational gradient in Ecuador. We conducted bird point counts at three elevations (1000, 2000 and 3000 m) on 18 1-ha plots and repeated the sampling eight times over two years (216 hours in total). For each plot, we obtained data of monthly temperatures and precipitation and recorded the overall resource availability (i.e., the sum of flower, fruit, and invertebrate resources). As expected, bird richness decreased from low to high elevations. Moreover, we found a significant decrease in bird abundance and richness and an increase in evenness between the most and least humid season at each of the three elevations. Climatic factors were more closely related to these temporal fluctuations than local resource availability. While temperature had significant positive effects on the abundance of birds at mid and high elevations, precipitation negatively affected bird abundance at low and mid elevations. Our study highlights that bird assemblages along tropical elevational gradients can show pronounced seasonal fluctuations. In particular, low temperatures and high precipitation seem to impose important constraints on birds. We conclude that potential changes in climate, due to global warming, are likely to affect the spatio-temporal dynamics of bird assemblages along tropical elevational gradients.

Land use change has stronger effects on functional diversity than taxonomic diversity in tropical Andean hummingbirds.

Land use change modifies the environment at multiple spatial scales, and is a main driver of species declines and deterioration of ecosystem services. However, most of the research on the effects of land use change has focused on taxonomic diversity, while functional diversity, an important predictor of ecosystem services, is often neglected. We explored how local and landscape scale characteristics influence functional and taxonomic diversity of hummingbirds in the Andes Mountains in southern Ecuador. Data was collected in six landscapes along a land use gradient, from an almost intact landscape to one dominated by cattle pastures. We used point counts to sample hummingbirds from 2011 to 2012 to assessed how local factors (i.e., vegetation structure, flowering plants richness, nectar availability) and landscape factors (i.e., landscape heterogeneity, native vegetation cover) influenced taxonomic and functional diversity. Then, we analyzed environment - trait relationships (RLQ test) to explore how different hummingbird functional traits influenced species responses to these factors. Taxonomic and functional diversity of hummingbirds were positively associated with landscape heterogeneity but only functional diversity was positively related to native vegetation coverage. We found a weak response of taxonomic and functional diversity to land use change at the local scale. Environment-trait associations showed that body mass of hummingbirds likely influenced species sensitivity to land use change. In conclusion, landscape heterogeneity created by land use change can positively influence hummingbird taxonomic and functional diversity; however, a reduction of native vegetation cover could decrease functional diversity. Given that functional diversity can mediate ecosystem services, the conservation of native vegetation cover could play a key role in the maintenance of hummingbird pollination services in the tropical Andes. Moreover, there are particular functional traits, such as body mass, that increase a species sensitivity to land use change.