Coevolution in an Amazonian hummingbird-plant community

Relationships between hummingbirds and their food plants are often considered to have arisen through coevolution. However, while it is certainly the case that hummingbirds and plants are coevolved, it is not always clear to what extent coevolution has shaped community structure and the morphology of the birds and plants. Here I examine the hummingbird-plant community of a lowland Amazonian rainforest in southeastern Colombia and test hypotheses concerning coevolution and community structure. To determine the strength of interrelationships and the degree to which character displacement has occurred, the distributions of three floral traits (flowering phenology, nectar production and corolla length) and two hummingbird traits (culmen length and foraging ecology) have been statistically tested against random null models. Although the hummingbird-plant community at Matamatá is diverse and is highly likely to have a long shared evolutionary history, there is little evidence for pairwise coevolution. Instead, the community appears to have evolved through diffuse coevolution, resulting in guilds of hummingbirds and plants.     

Nectar provision attracts hummingbirds and connects interaction networks across habitats

Many ecosystems have been modified by humans, creating novel habitats that include human-provided resources. Gardens adjacent to native habitats may affect plant–pollinator interactions by altering the determinants of interactions and species specialization. Here, we characterized a network comprising plants and hummingbirds interacting in a birdwatching garden with human-provided resources (nectar feeders and exotic plants) and adjacent Andean cloud forest in Colombia. Specifically, we investigated the proportion of hummingbirds visiting feeders and native/exotic plants to evaluate the connection between the habitats and the ecological determinants of the interaction network. Hummingbirds relied heavily on artificial nectar feeders in the garden, leaving the natural cloud forest for resources. Morphological matching was the single most important predictor of the observed pairwise interactions, for both hummingbirds and plants. At the species level, longer flowering phenology and a higher amount of sugar in nectar led to a higher degree for plants (i.e. the number of visiting hummingbird species). In contrast, a longer floral corolla was associated with lower specialization. Abundance was the best predictor of the number of partners for hummingbirds. The garden created for birdwatching attracted most, but not all, hummingbird species beyond their natural cloud forest habitat. Interestingly, the most frequently visited plants in the garden were native, especially the endemic and endangered tree Zygia lehmannii (Fabaceae). Our results show that some ecological mechanisms determining interactions in natural communities still hold in intensively modified habitats. Furthermore, a compromise between conservation and hummingbirds’ attraction to birding lodges/gardens is possible, for instance by favouring native and endemic plant species that are highly attractive for pollinators.     

An Assemblage of Hummingbird-pollinated Flowers in a Montane Forest in Southeastern Brazil

Relationships between ornithophilous flowers and hummingbirds have been little studied in southern South America, where hummingbird species richness is low. We studied an ornithophilous flower assemblage and the hummingbird pollinators in a montane forest in southeastern Brazil. Twenty-three native hummingbird-pollinated plant species in 21 genera and 14 families were observed. Bromeliaceae, Fabaceae, Gesneriaceae, and Lobeliaceae are represented by more than one species within the assemblage. Flower shapes vary from narrow tube to bowl-shape, but tubular flowers prevail. The variety of flower shapes and sizes results in diverse pollen placement on the body parts of hummingbird visitors, although pollen is deposited mostly on the bill. Sugar concentration in nectar averages 22.1%, and nectar volume per flower averages 16.9 μl. The plant populations bloom for one month to year-round, and their flowering approaches the steady-state pattern. Four flower subsets may be defined within the assemblage, each subset related to the bill size and foraging habits of the most frequent bird visitor. Of the six species of hummingbirds recorded at the study site, four are common and largely resident. The four hummingbirds differ in bill size, body mass, and favoured foraging sites, attributes which reflect their favoured flower subsets. One hermit and one trochiline hummingbird share most of the flower species they use, these two birds being the major pollinators within the flower assemblage. This montane forest community may be viewed as medium-rich in ornithophilous flower species and poor in hummingbird species.     

The assembly of plants used as nectar sources by hummingbirds in a Cerrado area of Central Brazil

Studies on hummingbird–plant interactions commonly use a pollination approach emphasizing mutualistic relationships. But floral resources are often used opportunistically by these birds. Plant–pollinator assemblies and pollination sustainability will depend both on the well-adapted plants and other potential floral resources. The Cerrado, Neotropical savannas of Central Brazil, has ca. 7.5 % of its flora supposedly adapted to hummingbird pollination. But detailed information about flowers effectively used by hummingbirds at community level is still lacking. Hence, we recorded all plant species visited by hummingbirds, to determine how these nectariferous flowers were distributed in time and space in different plant formations of a Cerrado area, and also the hummingbird species that visit them. The study was conducted between March 2007 and December 2008 in the Panga Ecological Station. Data regarding flowering phenology, floral morphology and visitation were collected monthly. Forty-six nectariferous species from 39 genera and 17 families were recorded, most with annual flowering dynamics and tubular flowers. But only 21 species had a combination of traits fitting classic ornithophilous syndrome. For the remaining species hummingbird visitation was ascertained from observations at the study site or other sites in the region. Eight hummingbird species occurred in the area and were recorded visiting directly 36 plant species. The study area presented a relatively low number of ornithophilous plants, but a great habitat diversity and many non-ornithophilous plants that hummingbirds used as nectar sources. Therefore, in the studied Cerrado, the diversity of environments and nectariferous plants favour the maintenance of resident and migrant hummingbirds.     

Effect of artificial feeders on pollen loads of the hummingbirds of Cerro de la Muerte, Costa Rica

Although sugar-water feeders are commonly used by enthusiasts to attract hummingbirds, little is known about how they affect hummingbird behavior and flower use. We studied the highland hummingbird assemblage of Cerro de La Muerte, Costa Rica, both at a site with permanent feeders (La Georgina Restaurant) and further from it. We examined how feeder use and monopolization affected seasonal changes in pollen loads during four sampling periods, including dry and wet seasons, from 2003-2005. We expected that species monopolizing the feeders would carry little or no pollen whatsoever, and would have pollen loads characterized by low floral diversity, in contrast with species less dependent on feeders. We obtained pollen samples from 183 individuals of four hummingbird species captured around the feeders using mist nets, which were compared with a pollen reference collection of plants with a pollination syndrome by hummingbirds. The same methods were implemented at a site 3km away from the feeders. Feeder usage was quantified by counting the number of times hummingbirds drank from the feeders in periods of 4min separated by 1min. The effects of hummingbird species and season on pollen load categories were assessed using a nominal logistic regression. The alpha species at the site, the Fiery-throated Hummingbird (Panterpe insignis), dominated the feeders during the dry season. Meanwhile, in the wet season, feeder usage was more evenly distributed across species, with the exception of the Volcano Hummingbird, Selasphorus flammula, which occupies the last place in the dominance hierarchy. Pollen loads of hummingbirds captured near feeders were low in abundance (more than 50% of captured individuals had zero or low pollen loads), and low in species richness (96% of the hummingbirds with pollen from only one plant genus, Centropogon). Overall pollen loads increased during the dry season coinciding with peaks in flower availability, although the majority of captured hummingbirds carried no pollen. Mist nets located 3km from La Georgina returned few captures (one-to-three specimens) per sampling date, contrasting with observations made before feeders were present. These results suggest that sugar-water feeders gather hummingbirds in over considerable distances drawing them away from flowers. The competitive and antagonistic pattern shown between feeders and flowers indicate that natural pollination system could be significantly altered. Supplementing hummingbirds with food seems likely to interfere with pollination networks already stressed by many anthropogenic effects.     

Floral resources and hummingbirds on an island of flooded forest in Central Brazil

Nectar availability for hummingbirds was studied for two and a half years in an island of flooded forest in Uberlândia, MG, in the Cerrado region, the Neotropical savanna areas in Central Brazil. Flowering individuals of ten plant species from eight families that offered resources potentially used by hummingbirds were recorded along transects crisscrossing the area, which were surveyed every other week. The number of flowers was estimated for each plant. The volume and concentration of nectar were measured for a sample of flowers of each species, and the data were used to estimate the caloric content available per flower and the general resource availability in the area throughout the study period. Focal observations of hummingbird visits were carried out on blooming individuals of all plant species. Hummingbird flowers occurred throughout the study period between January 2001 and June 2003. Only three of the studied plants did not present clear ornithophilous features. Six hummingbird species were recorded in the area, but total nectar availability in the area was seldom sufficient for the daily energy requirements of a single bird. Therefore, it can be inferred from the study that these hummingbirds need to move among the mosaic of different plant physiognomies in the region in order to obtain enough energy for their survival. These movements may also contribute to increasing gene flow and regeneration of plant populations isolated in forest patches as the one studied here.     

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.     

Macroecological patterns of resource use in resident and migratory hummingbirds

Hummingbirds (Family Trochilidae) are key pollinators in several biodiversity hotspots, including the California Floristic Province in North America. Relatively little is known about how hummingbird diets change throughout the year, especially with regard to how migratory hummingbirds affect resident hummingbirds at stopover sites. In this study, we examine how hummingbird species, migratory status, sex, geographic region and local plant diversity influence floral resource use before, during, and after an influx of migratory hummingbirds (primarily Rufous hummingbirds, Selasphorus rufus) across California. We expected distinct floral resource use based upon species’ migratory status (resident vs. migrant), sex, sampling period, and geographic region. We employed DNA metabarcoding to detect plant DNA in hummingbird fecal samples to analyze diet diversity, composition, overlap, and interaction networks. We found significant effects of sex, sampling period, and migratory status on the alpha and beta diversity of plant taxa present in fecal samples. Analyses of Anna's hummingbirds (Calypte anna) alone revealed that female fecal samples contained higher plant species richness. In addition to hummingbird-pollinated plants, fecal samples also contained non-ornithophilous plants and species of agricultural importance. Diet overlap and plant-pollinator network analyses revealed high overlap in plant taxa used between hummingbird species, and networks were more connected, less nested, and less specialized than null models. DNA metabarcoding is minimally invasive and provides a detailed view of hummingbird diet, permitting large-scale studies. Insights into hummingbird diets are especially valuable given the logistical difficulties of directly observing floral visitation and foraging across broad temporal and spatial scales.     

Pollination Ecology and Effect of Nectar Removal in Macleania bullata (Ericaceae)1

The floral syndrome of Macleania bullataYeo (Ericaceae) reflects its adaptation to hummingbird pollination. Its flowers, however, are subject to high levels of nectar robbing. I examined the floral visitor assemblage of M. bullata in a tropical montane wet forest in southwestern Colombia, focusing on the behavior of the visitors. I also tested for the presence of nocturnal pollination and the effects of nectar removal on new nectar production. The principal floral visitors were the nectar robbing hummingbirds Ocreatus underwoodii (19.1% of visits) and Chlorostilbon mellisugus (18.9%). Only two species of long–billed hummingbirds visited the flowers of M. bullata as “legitimate” pollinators: Coeligena torquata (14.7% of visits) and Doryfera ludoviciae (14.3%). The remaining visits constituted nectar robbing by bees, butterflies, and other species of hummingbirds. Nocturnal pollination took place, although fruit set levels were 2.4 times higher when only diurnal pollination was allowed as opposed to exclusively nocturnal pollination. Nectar robbers removed floral nectar without pollinating the flower. Treatments of experimental nectar removal were carried out to examine if flowers synthesize more nectar after nectar removal. Nectar removal increased the total volume of nectar produced by each flower without affecting sugar concentration. Thus, nectar robbing can impose a high cost to the plants by forcing them to replace lost nectar.     

Hummingbird diversity, food niche characters, and assemblage composition along a latitudinal precipitation gradient in the Bolivian lowlands

As for many other taxa, hummingbird diversity declines away from the equator, but the causes for this decline are still disputed and might involve, among others, climatic factors or the availability of food resources. Because hummingbirds are one of the classical examples for plant–animal coevolution, it has been proposed that the diversity of hummingbird assemblages might depend on the diversity of food plants available. We tested this hypothesis by studying the hummingbird assemblages and their food plants for 1 year at six sites along a 660-km-long transect in Bolivian lowland forests extending from the southernmost Amazonian rain forests to dry Chaco forests. Hummingbird diversity was higher in the northern three sites as compared to the southern ones, with an abrupt decline in species numbers and a corresponding change in taxonomic composition at the boundary from evergreen to drought deciduous forests. Hummingbird diversity and abundance were only weakly correlated to climatic factors or to the diversity of humming-visited flowers, but strongly to the seasonal abundance of flowers. The overlap in nectar diet between hummingbird species depended on the number of plant species: when numerous species were available, the hummingbirds segregated by feeding preferences, but when few flowers were available, all hummingbirds fed on the same plants. We conclude that the local diversity of hummingbird species is not primarily determined by the diversity of food plants, but rather by the abundance of flowers available at any given point in time.     

The Role of Flower Width in Hummingbird Bill Length‐Flower Length Relationships1

Observations of hummingbirds feeding at flowers longer or shorter than their bills seem to contradict the view that bill lengths of hummingbirds evolved in concert with the lengths of their flowers. Recent experiments, however, indicate that a hummingbird's ability to feed at artificial flowers of different lengths depends on the widths of the flowers. We examined if the broad range of flower lengths visited by many hummingbird species can be explained by the widths of the flowers. We predicted that both short‐ and long‐billed hummingbirds would include long, wide flower species in their diets, but that short‐billed hummingbirds would not include long, narrow flower species because nectar in these species might be beyond the reach of their bills. If so, the slope of the regression for flower width versus flower length should be smaller for flower species visited by longer‐billed hummingbirds relative to those visited by shorter‐billed hummingbirds. Analyses of data sets for some North American and Monteverde hummingbirds and their food plants were consistent with this prediction, and bill lengths were significantly correlated with the slopes of the regressions of flower width versus length for seven hummingbird species. Comparisons of observed flower use by some Monteverde hummingbird species to flower assemblages generated at random suggest that these significant regressions were not simply a result of allometric relationships between flower lengths and widths, but in some cases reflected active choice by the birds. The two hummingbird–flower data sets also differed significandy in the scaling of corolla width relative to corolla length. In particular, the Monteverde data set contained a large number of long, narrow flower species, which we suggest is a consequence of a different floral evolutionary history and association with long‐billed hummingbird species. The evolutionary effects of hummingbirds and their flowers upon one another are more complex than has generally been realized, and a consideration of corolla length jointly with other floral characters may improve our understanding of hummingbird‐flower relationships.     

The role of size and dominance in the feeding behaviour of coexisting hummingbirds

Interspecific competition can strongly influence community structure and limit the distribution and abundance of species. One of the main factors that determine hummingbird community structure is competition for food. The temporal and spatial distribution of nectar has a strong impact on hummingbird assemblages, shaping foraging niches according to hummingbird dominance and foraging strategy. We investigated whether body size and the degree of aggressive dominance influence feeding behaviour of hummingbirds in a temperate forest in northwestern Mexico (El Palmito, Mexico) when winter migrant hummingbirds are present in the community. First, we determined the dominance status of hummingbirds and evaluated the relationship between dominance and body mass, wing disc loading and migratory status. Secondly, we determined how hummingbird species used plant species differently. Thirdly, we examined whether the most dominant hummingbird species defended floral patches with more energy and/or with a larger number of flowers. At each flower patch, hummingbird species, number of hummingbird interactions, feeding time and number of flowers present were recorded. The total number of calories available within each floral patch was also determined. Our results demonstrate that the dominance hierarchy of 13 hummingbird species (migratory and resident) was correlated with body size but not wing disc loading, and that members of the hummingbird community showed a clear separation in resource use (by plant species). Hummingbirds at the top of the dominance hierarchy defended and fed on the best flower patches, defined by the quantity of calories available. Hence, the feeding behaviour of hummingbirds at El Palmito depends on the abundance of plant species used by hummingbirds and on the amount of energy available from each flower patch. Thus, hummingbird body size, aggressive dominance and defence of quality flower patches determines niche partitioning among species.     

Evolutionary transition between bee pollination and hummingbird pollination in Salvia: Comparing means,variances and covariances of corolla traits

Covariation among traits can modify the evolutionary trajectory of complex structures. This process is thought to operate at a microevolutionary scale, but its long‐term effects remain controversial because trait covariation can itself evolve. Flower morphology, and particularly floral trait (co)variation, has been envisioned as the product of pollinator‐mediated selection. Available evidence suggests that major changes in pollinator assemblages may affect the joint expression of floral traits and their phenotypic integration. We expect species within a monophyletic lineage sharing the same pollinator type will show not only similarity in trait means but also similar phenotypic variance‐covariance structures. Here, we tested this expectation using eighteen Salvia species pollinated either by bees or by hummingbirds. Our findings indicated a nonsignificant multivariate phylogenetic signal and a decoupling between means and variance‐covariance phenotypic matrices of floral traits during the evolution to hummingbird pollination. Mean trait value analyses revealed significant differences between bee‐ and hummingbird‐pollinated Salvia species although fewer differences were detected in the covariance structure between groups. Variance‐covariance matrices were much more similar among bee‐ than hummingbird‐pollinated species. This pattern is consistent with the expectation that, unlike hummingbirds, bees physically manipulate the flower, presumably exerting stronger selection pressures favouring morphological convergence among species. Overall, we conclude that the evolution of hummingbird pollination proceeded through different independent transitions. Thus, although the evolution of hummingbird pollination led to a new phenotypic optimum, the process involved the diversification of the covariance structure.     

Pairing automated mark–recapture and social network models to explore the effects of landscape configuration on hummingbird foraging patterns

Landscape changes can alter pollinator movement and foraging patterns which can in turn influence the demographic processes of plant populations. We leveraged social network models and four fixed arrays of five hummingbird feeders equipped with radio frequency identification (RFID) data loggers to study rufous hummingbird (Selasphorus rufus) foraging patterns in a heterogeneous landscape. Using a space-for-time approach, we asked whether forest encroachment on alpine meadows could restrict hummingbird foraging movements and impede resource discovery. We fit social network models to data on 2221 movements between feeders made by 29 hummingbirds. Movements were made primarily by females, likely due to male territoriality and early migration dates. Distance was the driving factor in determining the rate of movements among feeders. The posterior mean effects of forest landscape variables (local canopy cover and intervening forest cover) were negative, but with considerable uncertainty. Finally, we found strong reciprocity in hummingbird movements, indicative of frequent out and back movements between resources. Together, these findings suggest that reciprocal movements by female hummingbirds could help maintain bidirectional gene flow among nearby subpopulations of ornithophilous plants; however, if the distance among meadows increases with further forest encroachment, this may limit foraging among progressively isolated meadows.     

Plant–hummingbird interactions in the West Indies: floral specialisation gradients associated with environment and hummingbird size

Floral phenotype and pollination system of a plant may be influenced by the abiotic environment and the local pollinator assemblage. This was investigated in seven plant–hummingbird assemblages on the West Indian islands of Grenada, Dominica and Puerto Rico. We report all hummingbird and insect pollinators of 49 hummingbird-pollinated plant species, as well as six quantitative and semi-quantitative floral characters that determine visitor restriction, attraction and reward. Using nonmetric multidimensional scaling analysis, we show that hummingbird-pollinated plants in the West Indies separate in floral phenotypic space into two gradients—one associated with the abiotic environment and another with hummingbird size. Plants pollinated by large, long-billed hummingbirds had flowers with long corolla tube, large amounts of nectar and showy orange-red colouration. These attracted few or no insect species, whereas plants pollinated by small, short-billed hummingbirds were frequently pollinated by insects, particularly lepidopterans. The separation of plants related to environmental factors showed that species in the wet and cold highlands produced large amounts of dilute nectar, possessed no or a weak odour, and were associated with few insects, particularly few hymenopterans, compared to plants in the dry and warm lowlands. The most specialised hummingbird-pollinated plants are found in the West Indian highlands where they are pollinated by mainly large, long-billed hummingbirds. At the other extreme, highly generalised plants growing in the dry and warm lowlands are pollinated by small, short-billed hummingbirds and numerous insect species. This illustrates that, even within the hummingbird-pollinated flora, pollination syndrome and the degree of specialisation may vary tremendously depending on pollinator morphology and environment. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Seasonal resource tracking by Amazonian hummingbirds

Many ecological theories predict a close match between food resources and the number and diversity of consumers but comparatively few empirical studies have simultaneously quantified seasonal variation in an entire assemblage and its resource base. Hummingbirds provide an excellent model system to examine this relationship because their primary food resource is conspicuous and easily quantified. Over a 2-year period I used mist-netting and direct observation of birds feeding at flowering plants to record the abundance of 16 species of hummingbird in lowland Amazonian rainforest. Although there was very little annual variation in temperature and daylength, both rainfall and river level showed significant seasonal variation. At the study site this resulted in an annual flood lasting for over 5 months, inundating much of the area to a depth of 2 m. I recorded marked seasonal patterns in flowering phenology and arthropod abundance, which correlated with river level and rainfall, respectively. In turn, these seasonal fluctuations in resource abundance correlated very closely with variation in hummingbird species richness, hummingbird abundance and the total biomass of hummingbirds in the assemblage. This study is the first to demonstrate tracking of seasonal resource variation in equatorial rainforest and provides evidence that hummingbirds are sensitive to resource availability at spatial and temporal scales much greater than previously documented.     

Forest fragmentation and loss reduce richness,availability, and specialization in tropical hummingbird communities

Hummingbirds are important pollinators of many native Neotropical plants but their abundance and diversity in landscapes dominated by intensive human uses such as agriculture have rarely been examined, despite such land‐uses prevailing in the tropics. We examined how tropical deforestation affects hummingbird community structure in premontane forest patches embedded in a tropical countryside of Coto Brus Canton, Costa Rica. We captured hummingbirds in fourteen landscapes representing a gradient in patch size and forest amount, and tested for the effects of these variables on (1) hummingbird captures at flowers (pollinator availability); (2) species richness; and (3) filtering of functional traits. After accounting for sampling effects, both hummingbird availability and species richness declined by 40% and 50%, respectively, across the gradient in deforestation that we observed (9–66% forest within 1000 m). Focal patch size was the strongest predictor, even after statistically accounting for the amount of forest and matrix composition of landscapes. These reductions in availability and richness were well predicted by functional traits; morphologically specialized species with the capacity to transport long‐distance outcrossed pollen and low functional redundancy within the pollinator network showed the greatest sensitivity to landscape change. We hypothesize that declines in hummingbird availability, diversity, and functional traits are important mechanisms driving the observed pollen limitation of ornithophilous flowers in fragmented tropical landscapes. Efforts to conserve large forest patches and enhance matrix permeability are critical for maintaining forest hummingbird communities and pollination services under current and predicted deforestation regimes.     

Hummingbird diversity in a fragmented tropical landscape in the Chocó biogeographic zone

Forest loss and fragmentation drive widespread declines in biodiversity. However, hummingbirds seem to exhibit relative resilience to disturbance, characterized by increasing abundance alongside declining species richness and evenness. Yet, how widespread this pattern may be, and the mechanisms by which it may occur, remain unclear. To fill in this knowledge gap, we investigated habitat- and site-level patterns of diversity, and community composition of hummingbirds between continuous forest (transects n = 16 in ~3500 ha) and more disturbed surrounding fragments (n = 39, 2.5–48.0 ha) in the Chocó rain forest of northwestern Ecuador. Next, we assessed within-patch and patch-matrix characteristics associated with hummingbird diversity and composition. We found higher hummingbird species richness in forest fragments relative to the continuous forest, driven by increased captures of rare species in fragments. Community composition also differed between continuous forest and fragments, with depressed evenness in fragments. Increased canopy openness and density of medium-sized trees correlated with hummingbird diversity in forest fragments, although this relationship became nonsignificant after applying false discovery rate (p < .01). Higher species richness in fragments and higher evenness in the continuous forest highlight the complex trade-offs involved in the conservation of this ecologically important group of birds in changing Neotropical landscapes. Abstract in Spanish is available with online material.     

Site identity and moss species as determinants of soil microbial community structure in Norway spruce forests across three vegetation zones

Soil microbial community structure was investigated by PLFA-analysis in four spruce forests in Norway. The maximum latitudinal distance between the sites was approximately 350 km. Bilberry Vaccinium myrtillus dominated the forest floor vegetation in the study sites, which were selected because of the vegetation type. Soil samples were taken from all four sites under close to 100% homogeneous ground cover of each of two feathermoss species, i.e. Hylocomium splendens or Pleurozium schreberi, respectively. These mosses are ubiquitous in the boreal forest and constitute an abundant component of the forest floor vegetation over vast areas. Since there are no studies on how these mosses affect soil microbial community structure, our first aim was to investigate the effect of moss species on soil microbial communities. Our second aim was to investigate whether microbial communities differ among geographically separated forest sites with similar vegetation across vegetation zones. Soil microbial community structure differed between the study sites, although they appeared similar in terms of vegetation and abiotic soil conditions. Study site was the most important predictor of the variation in the PLFAs, more important than moss species, although there was a tendency for separation of microbial community structure between the two moss species.