Bird diversity along elevational gradients in the Andes of Colombia: area and mass effects

Aim The decrease in species richness with increasing elevation is a widely recognized pattern. However, recent work has shown that there is variation in the shape of the curve, such that both negative monotonic or unimodal patterns occur, influenced by a variety of factors at local and regional scales. Discerning the shape of the curve may provide clues to the underlying causes of the observed pattern. At regional scales, the area of the altitudinal belts and mass effects are important determinants of species richness. This paper explores the relationship between bird species richness, elevation, mass effects and area of altitudinal zones for birds in tropical mountains. Location The three Andean ranges of Colombia and the peripheral mountain ranges of La Macarena and Santa Marta. Methods Lists of bird species were compiled for altitudinal belts in eastern and western slopes of the three Andean Cordilleras and for La Macarena and Santa Marta. The area of the altitudinal belts was computed from digital elevation models. The effect of area was analysed by testing for differences among altitudinal belts in the slopes and intercepts of the species‐area relationships. Mass effects were explored by separately analysing two sets of species: broadly distributed species, i.e. lowland species whose distributions extend into the Andes, and tropical Andean species, i.e., species that evolved in the Andes. Results Plotting total number of species in each altitudinal belt revealed a decline in species richness with elevation. In slopes with a complete elevational gradient from lowlands to mountain peaks, the decrease was monotonic. In internal Andean slopes where the lower elevational belts are truncated, there was a peak at mid elevations. There was a linear relationship between number of species and area of the altitudinal belts. When controlling for area, there were no differences in the number of species among altitudinal belts (500–2600 m), except for the two upper‐elevation zones (2600–3200 and > 3200 m), which had lower species richness. Diversity of widely distributed species declined monotonically with elevation, whereas tropical Andean species exhibited a mid‐elevation peak. Main conclusions A large proportion of the variation in species richness with elevation was explained by area of the altitudinal belts. When controlling for area, species richness remained constant up to 2600 m and then decreased. This pattern contrasts with a previously reported hump‐shaped pattern for Andean birds. Diversity patterns of widely distributed species suggested that immigration of lowland species inflates diversity of lower elevational belts through mass effects. This influence was particularly evident in slopes with complete altitudinal gradients (i.e. connected to the lowlands). Tropical Andean species, in contrast, were more diverse in mid‐elevational belts, where speciation rates are expected to be higher. The influence of these species was more prevalent in internal Andean slopes with no connection to the lowlands. The decline of species richness at high elevations may be related to higher extinction rates and lower resource levels.     

Assemblage structure and dynamics of terrestrial birds in the southwest Amazon: a camera‐trap case study

Habitat spatial distribution, seasonal variation, and activity patterns influence changes in vertebrate assemblages over time. Terrestrial birds play major roles in the dynamics of tropical forests, but there are few effective methods to study these species due to their cryptic coloration and elusive behavior. We used camera‐trap data collected during 16 mo (February 2017–June 2018) to describe the terrestrial avifauna in southeastern Peru, assess to what extent the composition of terrestrial avifauna changes among seasons and across two major habitats (terra firme and floodplain forests), and determine daily activity patterns of terrestrial birds. We used overlap analyses to examine temporal co‐occurrence between ecologically similar and sympatric species. Camera traps recorded 16 species, including eight species in the family Tinamidae. Capture rates were highest for Pale‐winged Trumpeters (Psophia leucoptera; Psophiidae) and Gray‐fronted Doves (Leptolila rufaxilla; Columbidae). Species composition did not differ between habitats or seasons, and capture rates between habitats only differed for White‐throated Tinamous (Tinamus guttatus). Overlaps of activity patterns were high between ecologically similar species and species found in terra firme habitats (White‐throated Tinamous and Cinereous Tinamous, Crypturellus cinereus) and in both habitat types (Pale‐winged Trumpeters and Gray‐fronted Doves). Low numbers of captures of possibly locally rare or less abundant species hindered a complete analysis of spatial and seasonal patterns of terrestrial bird assemblages. We suggest a greater sampling effort and greater spatial replication to better understand the spatial and seasonal dynamics of the terrestrial avifauna. Further studies that assess the mechanisms that allow the coexistence of sympatric tinamous would be valuable, both in our study area and elsewhere. The use of camera traps in long‐term monitoring projects proved to be an effective tool for monitoring terrestrial birds, identifying cryptic and often rare animals to species level, and providing valuable ecological information at species and community levels.     

Small montane cloud forest fragments are important for conserving tree diversity in the Ecuadorian Andes

Montane tropical cloud forests, with their complex topography, biodiversity, high numbers of endemic species, and rapid rates of clearing, are a top global conservation priority. However, species distributions at local and landscape scales in cloud forests are still poorly understood, in part because few regions have been surveyed. Empirical work has focused on species distributions along elevation gradients, but spatial variation among forests at the same elevation is less commonly investigated. In this study, the first to compare tree communities across multiple Andean cloud forests at similar elevations, we surveyed trees in five ridge‐top forest reserves at the upper end of the ‘mid‐elevation diversity bulge’ (1900–2250 masl) in the Intag Valley, a heavily deforested region in the Ecuadorian Andes. We found that tree communities were distinct in reserves located as close as 10 to 35 km apart, and that spatially closer forests were not more similar to one another. Although larger (1500 to 6880 ha), more intact forests contained significantly more tree species (108–120 species/0.1 ha) than smaller (30 to 780 ha) ones (56–87 species/0.1 ha), each reserve had unique combinations of more common species, and contained high proportions of species not found in the others. Results thus suggest that protecting multiple cloud forest patches within this narrow elevational band is essential to conserve landscape‐level tree diversity, and that even small forest reserves contribute significantly to biodiversity conservation. These findings can be applied to create management plans to conserve and restore cloud forests in the Andes and tropical montane cloud forests elsewhere.     

Reproductive traits associated with species turnover of amphibians in Amazonia and its Andean slopes

Assembly of ecological communities is important for the conservation of ecosystems, predicting perturbation impacts, and understanding the origin and loss of biodiversity. We tested how amphibian communities are assembled by neutral and niche‐based mechanisms, such as habitat filtering. Species richness, β‐diversities, and reproductive traits of amphibians were evaluated at local scale in seven habitats at different elevation and disturbance levels in Wisui Biological Station, Morona‐Santiago, Ecuador, on the foothills of the Cordillera del Kutukú; and at regional scale using 109 localities across evergreen forests of Amazonia and its Andean slopes (0–3,900 m a.s.l.). At local scale, species composition showed strong differences among habitats, explained mainly by turnover. Reproductive modes occurred differently across habitats (e.g., prevalence of direct developers at high elevation, where breeding in ground level water disappears). At regional scale, elevation was the most important factor explaining the changes in species richness, reproductive trait occurrences, and biotic dissimilarities. Species number in all groups decreased with elevation except for those with lotic tadpoles and terrestrial reproduction stages. Seasonality, annual precipitation, and relative humidity partially explained the occurrence of some reproductive traits. Biotic dissimilarities were also mostly caused by turnover rather than nestedness and were particularly high in montane and foothill sites. Within lowlands, geographic distance explained more variability than elevation. Habitat filtering was supported by the different occurrence of reproductive traits according to elevation, water availability, and breeding microhabitats at both scales, as well as other assembly mechanisms based in biotic interactions at local scale. Human‐generated land use changes in Amazonia and its Andean slopes reduce local amphibian biodiversity by alteration of primary forests and loss of their microhabitats and the interaction network that maintains their unique amphibian assemblages with different reproductive strategies.     

Montane Andean rain forests are a global diversity hotspot of geometrid moths

Aim Andean forests are known to be a major diversity hotspot for vascular plants and vertebrates, but virtually nothing is known about the diversity of arthropods. We examined whether montane rain forests in southern Ecuador are also a diversity hotspot for arthropods, and chose geometrid moths as a model group. Location The study area in southern Ecuador (Province Zamora‐Chinchipe, 79° W, 04° S) covers c. 40 km², with 39 collecting sites within an elevational range of 1040–2677 m a.s.l. Thirty‐five of the sites were situated in an area c. 2.5 km². Additional qualitative sampling was carried out in the same area and up to an elevation of 3100 m. Methods Nocturnal moths were collected quantitatively and qualitatively using portable light towers consisting of two 15 W fluorescent tubes, and diurnal moths were collected qualitatively using an insect net. Insects were sampled in six fieldwork periods in the years 1999–2003. As diversity measures, Fisher's alpha of the log‐series distribution as well as eight estimators of total species richness were applied. Results A total of 1266 species were recorded, 63% of which were identified to named species, whereas the remainder are likely to include many undescribed species. Quantitative samples at light towers collected 35,238 specimens representing 1223 species. The extrapolated species number for these data is 1420 (incidence coverage estimator). Twenty‐one additional nocturnal species and 22 exclusively diurnal species were sampled qualitatively at elevations between 1040 and 3100 m. The pooled value of Fisher's alpha for all quantitative samples is 246 ± 3. Main conclusions The diversity of Geometridae documented here is much higher than anywhere else in the world, even without the inclusion of additional species from adjacent lowland rain forests. The number of recorded species in this small area corresponds to more than 6% of the known world fauna of geometrid moths. Our study emphasizes the importance of protecting the remaining montane Andean rain forests. For setting priorities in conservation, more studies on insect diversity are urgently required in other regions of the Andes, since montane forests are being destroyed at an alarming rate.     

Patterns of diversification in the high Andean Ponderacris grasshoppers (Orthoptera: Acrididae: Melanoplinae)

The Andes, the world's longest mountain chain, harbours great taxonomic and ecological diversity. Despite their young age, the tropical Andes are highly diverse due to recent geological uplift. Speciation either followed the orogeny closely or occurred after the Andean uplift, as a result of subsequent climatic changes. Different scenarios have been proposed to explain the diversification of high Andean taxa. The Melanoplinae grasshopper Ponderacris Ronderos & Cigliano is endemic to the eastern slopes of the Andes of Peru and Bolivia, mostly distributed between 1000 and 4000 m above sea level. Diversification in several montane habitats of Bolivia and Peru allows tests via cladistic analysis of distinct possible geographic modes of speciation. Eight species are recognized, with three described here as new with revised diagnostic morphological characters provided: Ponderacris carlcarbonelli sp.n., P. chulumaniensis sp.n. and P. amboroensis sp.n. Cladistic analyses of 15 species (8 ingroup and 7 outgroup) and 38 morphological characters, under equal and implied weighting, confirm the monophyly of Ponderacris. Characters from the external morphology and colour pattern provided less phylogenetic information than did the male abdominal terminalia and phallic complex. Species distributed in the Peruvian Andes constituted a monophyletic group, whereas those from the Bolivian Andes formed a basal paraphyletic grade. Dispersal–vicariance analysis resulted in one ancestral distribution reconstruction indicating that the most recent common ancestor was distributed in the Lower Montane Yungas of Bolivia. Eleven dispersal and one vicariant events are postulated, with a South‐to‐North speciation pattern coincident with progressive Andean uplift. Vicariance could relate to fragmentation of montane forest during the dry intervals of the late Cenozoic. From the Bolivian area, ancestral Peruvian Ponderacris may have dispersed northward, coinciding with the rise of the Andes. Ten of 11 dispersal events occurred at terminal taxa and are likely to be recent. However, diversification of Ponderacris cannot be explained solely by the South‐to‐North speciation hypothesis, but may also include both vicariance and dispersal across barriers influenced by Pleistocene climatic cycles.     

Elevational Ranges of Montane Birds and Deforestation in the Western Andes of Colombia

Deforestation causes habitat loss, fragmentation, degradation, and can ultimately cause extinction of the remnant species. Tropical montane birds face these threats with the added natural vulnerability of narrower elevational ranges and higher specialization than lowland species. Recent studies assess the impact of present and future global climate change on species’ ranges, but only a few of these evaluate the potentially confounding effect of lowland deforestation on species elevational distributions. In the Western Andes of Colombia, an important biodiversity hotspot, we evaluated the effects of deforestation on the elevational ranges of montane birds along altitudinal transects. Using point counts and mist-nets, we surveyed six altitudinal transects spanning 2200 to 2800m. Three transects were forested from 2200 to 2800m, and three were partially deforested with forest cover only above 2400m. We compared abundance-weighted mean elevation, minimum elevation, and elevational range width. In addition to analysing the effect of deforestation on 134 species, we tested its impact within trophic guilds and habitat preference groups. Abundance-weighted mean and minimum elevations were not significantly different between forested and partially deforested transects. Range width was marginally different: as expected, ranges were larger in forested transects. Species in different trophic guilds and habitat preference categories showed different trends. These results suggest that deforestation may affect species’ elevational ranges, even within the forest that remains. Climate change will likely exacerbate harmful impacts of deforestation on species’ elevational distributions. Future conservation strategies need to account for this by protecting connected forest tracts across a wide range of elevations.     

The Andean Hapalopsittaca parrots (Psittacidae,Aves): an example of montane‐tropical lowland vicariance

Quintero, E., Ribas, C. C. & Cracraft, J. (2012). The Andean Hapalopsittaca parrots (Psittacidae, Aves): an example of montane‐tropical lowland vicariance. —Zoologica Scripta, 42, 28–43. We describe a phylogenetic and biogeographical pattern connecting high montane biotas to those of the lowlands, as exemplified by the exclusively montane parrot genus Hapalopsittaca and its lowland sister genus Pyrilia, both nested within Tavares et al.’s “amazons and allies” clade. As Hapalopsittaca is the only genus within the “amazons and allies” clade that is exclusively distributed in the Andes, the optimization leads to the inference that the ancestral distribution of the branch leading to Pyrilia + Hapalopsittaca was lowland. Museum specimens were examined to determine basal diagnosably distinct taxonomic units. Based on this analysis, mitochondrial sequences (cyt b and ND2 genes) from 17 individuals, mostly from toe pads, and representing all basal taxa within Hapalopsittaca, were obtained. A divergence‐dating analysis using both nuclear (RAG‐1) and mitochondrial (cyt b) genes was conducted to explore whether the uplift of the Andes coincides temporarily with the origin of this montane group, and thus might be causally linked to its origin. Molecular dating estimates the split between Hapalopsittaca and Pyrilia to have occurred between 6.6 and 8.0 Myr; thus, the timing of this highland/lowland disjunction is consistent with that of the final uplift of the Central Andes, supporting a hypothesis of vicariance due to Andean uplift. These results suggest that the taxonomic assembly of montane biotas may be, at least in part, explained by events of Earth history rather than by long‐distance dispersal and colonization. Diversification within Hapalopsittaca and the origin of current species are more recent in time and probably are related to Pleistocene climatic oscillations, as has been shown in other montane groups.     

Explaining Andean megadiversity: the evolutionary and ecological causes of glassfrog elevational richness patterns

The Tropical Andes are an important global biodiversity hotspot, harbouring extraordinarily high richness and endemism. Although elevational richness and speciation have been studied independently in some Andean groups, the evolutionary and ecological processes that explain elevational richness patterns in the Andes have not been analysed together. Herein, we elucidate the processes underlying Andean richness patterns using glassfrogs (Centrolenidae) as a model system. Glassfrogs show the widespread mid‐elevation diversity peak for both local and regional richness. Remarkably, these patterns are explained by greater time (montane museum) rather than faster speciation at mid‐elevations (montane species pump), despite the recency of the major Andean uplift. We also show for the first time that rates of climatic‐niche evolution and elevational change are related, supporting the hypothesis that climatic‐niche conservatism decelerates species' shifts in elevational distributions and underlies the mid‐elevation richness peak. These results may be relevant to other Andean clades and montane systems globally.     

Biological diversification in a complex region: a spatial analysis of faunistic diversity and biogeography of the Andes of Colombia

Aim Understanding large‐scale patterns of beta diversity and endemism is essential for ecoregional conservation planning. We present a study of spatial patterns of faunal diversification and biogeographical relationships in the Andean region of Colombia. This region has a great geomorphological complexity, as it is formed by several mountain ranges with different geologic origins. We hypothesize that this complexity results in a high turnover in species composition among subregions. Location The Andean region of Colombia, including the Santa Marta and Macarena mountain ranges. Methods The region was divided into subregions, represented by the eastern and western slopes of each of the three Andean cordilleras, the Cauca and Magdalena valley bottoms, and the peripheral mountain ranges of Perijá, Macarena and Sierra Nevada de Santa Marta. Species lists for five animal taxa (rodents, bats, birds, frogs and butterflies) were compiled for each subregion and similarities in species composition were determined by cluster analysis. To explore biogeographical relationships, species were classified into one of four distributional categories: endemic, tropical Andean, Andean‐Central American and wide continental distribution. Results The highest species richness in the region was found in the Pacific and eastern versants of the Andes, and the lowest in the Cauca and Magdalena valley bottoms. Inter‐Andean slopes were intermediate in species richness. However, when species richness was calculated per unit area, the most diverse regions were the Santa Marta and Macarena ranges, the Cauca Valley watershed and the Pacific slope. Although each taxonomic group had a different branching pattern, dendrograms indicated five common subregional clusterings: (1) Perijá‐Sierra Nevada, (2) the Pacific slope, (3) the eastern Andean slope, (4) the Cauca and Magdalena valley bottoms, and (5) the inter‐Andean slopes. Clustering patterns of inter‐Andean slopes varied among taxa. In birds, bats and rodents, grouping was by opposite slopes of the same valley, whereas frogs were grouped by mountain ranges and butterflies by valleys and their respective slopes. Seventy‐five per cent of species in all taxa were found in less than five subregions. The fauna of the Magdalena and Cauca valley bottoms was composed mostly of lowland species with wide geographical distributions, whereas the cordilleran fauna was mostly restricted to the tropical Andes. Main conclusions The western and eastern versants of the Andes have the highest species richness, but are also the largest subregions. On a per unit area basis, the peripheral ranges (Santa Marta and Macarena) are the richest, followed by the western portion of the Andes (the Cauca Valley watershed and the Pacific versant). Clustering patterns in dendrograms suggest two major patterns of differentiation of the Andean fauna: one elevational (lowlands vs. highlands) and one horizontal (among ranges and/or slopes). Biogeographical affinities of the inter‐Andean valley bottoms are with the lowland faunas of tropical America. In contrast, Andean faunas diversified locally, resulting in the evolution of a large number of endemic species, particularly among the less vagile taxa. Three different main branches of Andean fauna can be recognized, one confined to the Pacific, another to the eastern (Amazonian‐Llanos) versant of the Andes, and the third one composed by the inter‐Andean slopes of the Cauca and Magdalena valleys. The identification of five main biogeographical units in the Andean region of Colombia has important implications for the conservation of the regional biota. Conservation initiatives that seek to preserve representative samples of the regional biodiversity should take into account the patterns of diversification described here, and the evolutionary processes that gave rise to these patterns.     

Differential high‐altitude adaptation and restricted gene flow across a mid‐elevation hybrid zone in Andean tit‐tyrant flycatchers

The tropical Andes are a global hotspot of avian diversity that is characterized by dramatic elevational shifts in community composition and a preponderance of recently evolved species. Bird habitats in the Andes span a nearly twofold range of atmospheric pressure that poses challenges for respiration, thermoregulation, water balance and powered flight, but the extent to which physiological constraints limit species' elevational distributions is poorly understood. We report a previously unknown hybrid zone between recently diverged flycatchers (Aves, Tyrannidae) with partially overlapping elevational ranges. The southern Anairetes reguloides has a broad elevational range (0–4200 m), while the northern Anairetes nigrocristatus is restricted to high elevations (>2200 m). We found hybrids in central Peru at elevations between ~3100 and 3800 m, with A. nigrocristatus above this elevation and A. reguloides below. We analysed variation in haematology, heart mass, morphometrics, plumage and one mitochondrial and three nuclear loci across an elevational transect that encompasses the hybrid zone. Phenotypic traits and genetic markers all showed steep clines across the hybrid zone. Haemoglobin concentration, haematocrit, mean cellular haemoglobin concentration and relative heart mass each increased at altitude more strongly in A. reguloides than in A. nigrocristatus. These findings suggest that A. nigrocristatus is more resistant than A. reguloides to high‐altitude hypoxic respiratory stress. Considering that the ancestor of the genus is suggested to have been restricted to high elevations, A. reguloides may be secondarily adapted to low altitude. We conclude that differential respiratory specialization on atmospheric pressure combined with competitive exclusion maintains replacement along an elevational contour, despite interbreeding.     

Elevational plumage divergence in the Rufous-capped Babbler (Cyanoderma ruficeps) on a mountainous island

Environment plays an important role in the evolution of plumage coloration in birds and may also lead to sexual dichromatism if males and females face different selection pressures. Mountains exhibit varying ecological conditions along their elevation gradient that may impose divergent selection on elevationally widespread species, causing intraspecific plumage divergence. For example, UV light environments often vary between montane and lowland habitats, which could potentially cause differences in plumage UV reflection between birds occurring in the two types of habitats. However, few studies have examined the effects of elevation on plumage evolution. In this study, we quantified the plumage coloration of the Rufous-capped Babbler Cyanoderma ruficeps from montane and lowland habitats on a mountainous island, Taiwan. We aimed to examine whether their plumage showed differences associated with changing ecological environments across the elevational gradient. The results supported that the plumage of babblers occupying montane habitats had higher UV-reflectance and brightness than that of lowland birds, corresponding to the higher UV intensity in montane than lowland background light environments. The elevational differences were mainly found across the ventral parts of babblers that had relatively higher levels of UV reflectance compared with their dorsal parts. Alternatively, the brighter plumage, with higher UV-reflectance in montane than lowland birds, might be mediated by physiological adaptation to other ecological factors, such as parasite pressures. The elevational differences in plumage UV-reflectance and brightness were more dramatic in males than in females. However, we found significant sexual dichromatism in different body parts between montane and lowland babblers in which females had brighter or stronger UV-associated coloration than males, suggesting that sexual selection has little impact on babbler plumage. Our study suggests the importance of elevational divergent selection associated with UV light or other ecological environments on avian plumage evolution.     

Large-scale diversity patterns of vascular epiphytes in Neotropical montane rain forests

Aim Epiphytes contribute up to 30% to the number of vascular plant species in certain global biodiversity hotspots, e.g. the Ecuadorian Andes. However, their large scale diversity patterns are still discussed on the base of results from a few, local epiphyte inventories. Consequently, explanatory models on epiphyte diversity concentrate on the impact of local climate on small scale epiphyte species richness. Our aim was to analyse large scale elevational patterns of epiphyte diversity integrating data from different geographic scales. Location Tropical America, with special emphasis on the Ecuadorian Andes. Methods Our study is based on two data sources. First, we analysed the elevational patterns of epiphyte diversity based on the Catalogue of the Vascular Plants of Ecuador and the Libro Rojo de las Plantas Endèmicas del Ecuador. Secondly, the floristic turnover between the epiphyte inventories of seven montane and four lowland study sites in the Neotropics was analysed. Results The floristic turnover between Neotropical montane epiphyte floras is higher than the one between lowland epiphyte floras. Montane study sites located only a few kilometres apart from each other show considerable differences in their epiphyte species inventories. Irrespectively of their similar dispersal mode, the floristic turnover is much higher for orchids than for Pteridophyta. The Orchidaceae are the species richest group in all of the examined 11 Neotropical epiphyte floras. At the larger scale of the Ecuadorian Flora, c. 50% of the species in the elevational zone with maximum epiphyte diversity (between 1000 and 1500 m) are orchids. Elevational patterns of epiphyte diversity strongly reflect patterns of Orchidaceae. Main conclusions Our results support the observation of a ‘mid‐elevation bulge’ of epiphyte diversity by Gentry and Dodson. It has been frequently shown that the high humidity in mid‐elevations is suitable to maintan a high epiphyte species richness. Our findings show that in addition, large scale epiphyte diversity in montane rain forest is increased by the high floristic turnover at local and regional scale. Based on the importance of Orchidaceae for epiphyte diversity, we discuss that speciation processes corresponding to the highly diverse environment are a driving force for endemism, floristic heterogeneity and consequently for large scale epiphyte species richness in montane forests.     

Dispersal,isolation and diversification with continued gene flow in an Andean tropical dry forest

The Andes are the world's longest mountain chain, and the tropical Andes are the world's richest biodiversity hot spot. The origin of the tropical Andean cordillera is relatively recent because the elevation of the mountains was relatively low (400–2500 m palaeoelevations) only 10 MYA with final uplift being rapid. These final phases of the Andean orogeny are thought to have had a fundamental role in shaping processes of biotic diversification and biogeography, with these effects reaching far from the mountains themselves by changing the course of rivers and deposition of mineral‐rich Andean sediments across the massive Amazon basin. In a recent issue of Molecular Ecology, Oswald, Overcast, Mauck, Andersen, and Smith (2017) investigate the biogeography and diversification of bird species in the Andes of Peru and Ecuador. Their study is novel in its focus on tropical dry forests (Figure 1) rather than more mesic biomes such as rain forests, cloud forests and paramos, which tend to be the focus of science and conservation in the Andean hot spot. It is also able to draw powerful conclusions via the first deployment of genomic approaches to a biogeographic question in the threatened dry forests of the New World.     

The lichen genus Umbilicaria in Ecuador

Nine species of the saxicolous lichen genus Umbilicaria from Ecuador are reported and a key is provided: U. africana, U. aprina, U. cinereorufescens, U. decussata, U. dendrophora, U. haplocarpa, U. leprosa, U. nylanderiana and U. vellea. The species diversity of this genus on the equatorial high mountains of South America is low compared to North America and Eurasia, but similar to that found on high African peaks close to the equator. The species mostly belong to a high‐alpine element with worldwide distribution. Two species belong to an Andean endemic element, viz U. haplocarpa and U. leprosa. The low diversity and low percentage of endemism may reflect the fairly recent uplift of the Andes and the comparatively small geographic extent of the alpine Andean biota. The dense rainforests of Ecuador leave few suitable open rock habitats for the establishment of these light‐craving lichens below the tree‐line (4000–4200 m a.s.l.), and thus their equatorial habitat is almost exclusively restricted to the alpine zone between the tree‐line and the snow‐line (4700–4800 m a.s.l.). The equatorial species mainly reproduce asexually by thalloconidia, and the adaptive significance of this type of reproduction in high altitude habitats is discussed.     

The role of geography and ecology in shaping the phylogeography of the speckled hummingbird (Adelomyia melanogenys) in Ecuador

The Andes of South America contain one of the richest avifaunas in the world, but little is known about how this diversity arises and is maintained. Variation in mitochondrial DNA and morphology within the speckled hummingbird (Adelomyia melanogenys) was used to elucidate the phylogeographic pattern along an Ecuadorian elevational gradient, from the coastal cordillera to the inland Andean montane region. We examined sequence, climatic/remote sensing and morphological data to understand the effects of topography and ecology on patterns of variation. Populations on either side of the Andes are genetically divergent and were separated during a period that corresponds to the final stages of Andean uplift during the Pliocene. Despite isolation, these two populations were found to be morphologically similar suggesting a strong effect of stabilizing selection across ecologically similar Andean cloud forests, as assessed using climatic and remote sensing data. In contrast, little genetic divergence was found between coastal and west-Andean individuals, suggesting recent interruption of gene flow between these localities. However, coastal populations were found to inhabit different habitats compared to Andean populations as shown by climatic and remote sensing variables. Furthermore, coastal individuals had significantly longer bills compared to their montane relatives, indicative of differential directional selection and the influence of habitat differences in shaping phenotypic variation. Results highlight the role of both isolation and ecology in diversification in Ecuadorian montane regions, while suggesting the two may not always act in concert to produce divergence in adaptive traits.     

Topographic and spatial controls of palm species distributions in a montane rain forest, southern Ecuador

The northern Andes harbour a flora that is as species-rich or even richer than the 18-times larger lowland Amazon basin. Gaining an understanding of how the high species richness of the Andean region is generated and maintained is therefore of particular interest. Environmental sorting due to elevational gradients in climate has been emphasized as a driver of vegetation distribution and plant community assembly in tropical mountain areas such as the Andes for two centuries, while alternative mechanisms have been little studied. Here, we investigated the importance of topography and spatial location as factors controlling species distributions in a palm community in a montane rain forest landscape in the Andes of southern Ecuador (1900–2150 m above sea level). Eleven species were present: Aiphanes verrucosa, Ceroxylon parvifrons, Chamaedorea pinnatifrons, Dictyocaryum lamarckianum, Euterpe precatoria, Geonoma densa, Geonoma orbignyana, Geonoma paradoxa, Prestoea acuminata and Wettinia aequatorialis. To study their spatial distribution, forty 250 m² (5 × 50 m²) plots were laid out perpendicular to four paths that were categorized into three areas and two topographic units (ridges and gullies). Mantel tests and indicator species analysis showed that both topography and spatial location imposed strong controls on palm species distributions at the study site. Our results suggest that species distributions in the studied montane forest landscape were partly determined by the species’ habitat requirements, but also by unknown spatial effects. Although a number of possible explanations exist for the latter, such as unmeasured environmental variables and historical disturbance events, we believe dispersal limitation is likely to be involved. Furthermore, although the gully- or ridge-association of some species corresponded to their general elevational ranges in southern Ecuador, this was not the case for other species. Based on such considerations, we conclude that elevational climatic gradients are likely to only form part of the explanation for the topographic effects on palm species distributions at the study site. Other factors must also be involved, notably wind-exposure and hydrology, as discussed for lowland palm communities. Our results show that to understand plant community assembly in the tropical montane forests of the Andes it is too simple to focus just on environmental sorting by elevational climatic gradients.     

Elevational differentiation accelerates trait evolution but not speciation rates in Amazonian birds

The importance of ecologically mediated divergent selection in accelerating trait evolution has been poorly studied in the most species‐rich biome of the planet, the continental Neotropics. We performed macroevolutionary analyses of trait divergence and diversification rates across closely related pairs of Andean and Amazonian passerine birds, to assess whether the difference in elevational range separating species pairs – a proxy for the degree of ecological divergence – influences the speed of trait evolution and diversification rates. We found that elevational differentiation is associated with faster divergence of song frequency, a trait important for pre‐mating isolation, and several morphological traits, which may contribute to extrinsic post‐mating isolation. However, elevational differentiation did not increase recent speciation rates, possibly due to early bursts of diversification during the uplift of the eastern Andes followed by a slow‐down in speciation rate. Our results suggest that ecological differentiation may speed up trait evolution, but not diversification of Neotropical birds.     

Islands in the sky or squeezed at the top? Ecological causes of elevational range limits in montane salamanders

Many animal and plant taxa reach their highest endemism and species richness in montane regions. The study of elevational range limits is central to understanding this widespread pattern and to predicting the responses of montane species to climate change. Yet, because large‐scale manipulations of the distributions of most species are difficult, the causes of species’ elevational range limits (e.g. competitive interactions, physiological specialization) are poorly understood. Here, we harness the power of new mechanistic approaches to dissect the factors that underlie the elevational replacement of two salamander species in the Appalachian Highlands. Our results challenge the long‐held idea that competitive interactions drive the lower elevational range limits of montane species and that physiological stress prevents low‐elevation species from expanding to high elevations. We show that physiological constraints drive the lower elevational range limit of the montane‐endemic species, Plethodon jordani. Conversely, we find that competition with P. jordani prevents the low‐elevation species, P. teyahalee, from expanding its range to include higher‐elevation habitats. These results are broadly consistent with the biogeography and behavior of other montane species, suggesting that similar mechanisms underlie patterns of elevational zonation across a variety of taxa and montane regions. To the extent that our findings are taxonomically and geographically widespread, these results challenge the idea that competitive release at species’ lower elevational range limits is driving the downslope range shifts exhibit by some montane taxa. Instead, our results raise the sobering possibility that even small changes in climate might cause erosion of the ranges of many high‐elevation species.     

Different responses of avian feeding guilds to spatial and environmental factors across an elevation gradient in the central Himalaya

Although elevational patterns of species richness have been well documented, how the drivers of richness gradients vary across ecological guilds has rarely been reported. Here, we examined the effects of spatial factors (area and mid‐domain effect; MDE) and environmental factors, including metrics of climate, productivity, and plant species richness on the richness of breeding birds across different ecological guilds defined by diet and foraging strategy. We surveyed 12 elevation bands at intervals of 300 m between 1,800 and 5,400 m a.s.l using line‐transect methods throughout the wet season in the central Himalaya, China. Multiple regression models and hierarchical partitioning were used to assess the relative importance of spatial and environmental factors on overall bird richness and guild richness (i.e., the richness of species within each guild). Our results showed that richness for all birds and most guilds displayed hump‐shaped elevational trends, which peaked at an elevation of 3,300–3,600 m, although richness of ground‐feeding birds peaked at a higher elevation band (4,200–4,500 m). The Normalized Difference Vegetation Index (NDVI)—an index of primary productivity—and habitat heterogeneity were important factors in explaining overall bird richness as well as that of insectivores and omnivores, with geometric constraints (i.e., the MDE) of secondary importance. Granivore richness was not related to primary production but rather to open habitats (granivores were negatively influenced by habitat heterogeneity), where seeds might be abundant. Our findings provide direct evidence that the richness–environment relationship is often guild‐specific. Taken together, our study highlights the importance of considering how the effects of environmental and spatial factors on patterns of species richness may differ across ecological guilds, potentially leading to a deeper understanding of elevational diversity gradients and their implications for biodiversity conservation.