Soil Organic Carbon and Water Retention after Conversion of Grasslands to Pine Plantations in the Ecuadorian Andes

Tree plantations in the high elevations of the tropics constitute a growing land use, but their effect on ecosystem processes and services is not well known. We examined changes in soil organic carbon (C) and water retention in a chronosequence of Pinus radiata stands planted in páramo grasslands in Cotopaxi province, Ecuador. Water retention at 10, 33, and 1,500 kPa declined with stand age, with soils in the oldest pine stands retaining 39%, 55%, and 63% less water than grassland soils at the three pressures tested. Soil organic C in the 0–10-cm depth also declined with stand age, from 5.0 kg m^–2 in grasslands to 3.5 kg m^–2 in 20–25-year-old pine stands (P < 0.001); at greater depth in the A horizon, C contents decreased from 2.8 to 1.2 kg m^–2 (P = 0.047). There were no significant differences among age classes in the AC and C horizons (P = 0.15 and P = 0.34, respectively), where little or no weathering of the primary material has occurred. Inputs of C may be affected by the significantly higher carbon–nitrogen (C:N) ratio of the litter under older pine stands (P = 0.005), whereas outputs are influenced by substrate quality as well as soil environmental factors. Soil ratios at the 0–10 cm depth were significantly higher in grasslands and young pine stands (P < 0.001), whereas carbon–phosphorous (C:P) ratios at 0–10-cm depth followed a similar but not significant trend. However, there was no significant difference in short-term decomposition rates (P = 0.60) when the soils were incubated under uniform temperature and moisture conditions. In páramo ecosystems, where high soil moisture plays an important role in retarding decomposition and driving high C storage, the loss of water retention after afforestation may be the dominant factor in C loss. These results suggest that soil C buildup and water retention respond rapidly to changes in biota and need to be assessed with regard to implications for C sequestration and watershed management.     

Biogeography, geographical variation and taxonomy of the Andean hummingbird genusMetallura GOULD, 1847

Summary Biogeography and taxonomy of the High-Andean hummingbird genusMetallura were analysed on the basis of geographical variation of morphometric and plumage colour characters. The genusMetallura is considered to consist of nine species and 15 subspecies. Within the speciesMetallura tyrianthina seven subspecies are recognized (smaragdinicollis [includingperuviana],septentrionalis, quitensis, tyrianthina, districta, oreopola, chloropogon).M. tyrianthina andM. iracunda are considered to be sister species and to be closely related toM. phoebe and to the members of theM. aeneocauda superspecies. The latter consists of six species as follows:M. aeneocauda (with subspeciesaeneocauda andmalagae),M. eupogon, M. theresiae (with subspeciestheresiae andparkeri),M. odomae, M. baroni andM. williami (with subspecieswilliami, atrigularis, primolina, recisa).M. tyrianthina andM. iracunda exhibit plesiomorphic character states (dimorphism, colour of throat feathers, short bill), that link the genusMetallura to its sister genusChalcostigma. The disjunct distribution of similar phenotypes, and the progression of morphological characters suggest fairly recent allopatric speciation processes in allMetallura taxa. High-Andean habitat fragmentations due to climatic changes during Late-Pliocene and Pleistocene presumably caused geographical isolation and differentiation of mostMetallura populations. The south-north phenotype progression regarding chin and gorget coloration ofM. tyrianthina females, paralleled by a transition from weak to strong sexual dimorphism, suggest for the genusMetallura a centre of origin in the Andes of north-eastern Bolivia or south-eastern Peru.Dedicated to Hans Hudde, Essen, for his friendship and support.     

Topographic controls on the distribution of tree islands in the high Andes of south‐western Ecuador

Aim To evaluate the hypothesis that geomorphometric parameters of upper montane Andean environments have an important influence on the regional fire ecology and consequently play a role in the spatial distribution of ‘remnant’ tree islands dominated by Polylepis. Location A glacial landscape located between 3600 and 4400 m elevation in Cajas National Park, south‐western Ecuador. Methods The eigenvalue ratio method was used to evaluate the regional geomorphometric parameters of a 30‐m digital elevation model for Cajas National Park. The landscape character was evaluated by quantifying the topographic roughness, organization, and gradient. This information was used to determine the spatial correlations between terrain characteristics and the distribution of tree islands in the region. Results We demonstrate a strong spatial correlation between areas of high topographic roughness and gradient, and the locations of the major tree islands. We find that there is a distinctive relationship between the topographic roughness and organization in the vicinity of the tree islands (e.g. increased upslope roughness and decreased topographic grain strength) that substantiates the notion that the tree islands are located in relatively inaccessible topography. Main conclusions In the northern and central Andes, the location of Polylepis‐dominated ‘forest islands’ has been shown to be a function of climate, terrain characteristics, and anthropogenic disturbances. Although the relative importance of various ecological factors has been debated, it remains clear that fires have exerted a strong influence on these ecosystems. Other authors have noted that tree islands are more likely to occur at the base of cliffs, above moist areas, and in other areas where fires do not burn frequently. Our results corroborate these observations, and demonstrate that the occurrence of Polylepis patches is strongly correlated with specific combinations of terrain features. Although we do not discount the importance of other factors in determining the spatial position and areal extent of these forests, we demonstrate strong support for fire‐related hypotheses.     

Assessing the origin of Neotropical mountain dung beetle assemblages (Scarabaeidae: Scarabaeinae): the comparative influence of vertical and horizontal colonization

Aims The fauna of mountains and their surrounding regions are likely to be influenced principally by two biological processes: horizontal colonization along similar altitudinal levels by elements originating from lineages inhabiting higher latitudes; and vertical colonization by lineages from the same latitude, but at lower altitudes. We examine whether the expected patterns derived from the latter process can be observed in mountain dung beetle assemblages. Specifically, we study the variation in species composition and richness with altitude in five regions spanning elevation gradients, analysing whether the altitudinal rates of change in the number of species and genera differ, and whether beta‐diversity scores for adjacent sites in each altitudinal gradient are different for species and genera. Location Eastern Cordillera of the Colombian Andes. Methods Field work was carried out in 1997–99 at 27 sites in five regions with elevation gradients, with 10–32 pitfall traps placed in each site. For each altitudinal level the numbers of species and genera were analysed with respect to altitude, and the slope of the linear regression between these variables was calculated. The slope of the curve of the altitude against the cumulative number of species and genera was also calculated for each altitudinal gradient to describe the compositional change between adjacent sites (beta diversity). Species and generic slopes were compared using analysis of covariance. The turnover of species along each altitudinal gradient was measured using presence/absence data and Cody's beta‐diversity index between adjacent pairs of sites. A cluster analysis was used to detect faunistically homogeneous groups of localities. Results Species richness always decreased with altitude, although the slopes did not differ significantly from zero. The number of genera also decreased with increasing altitude, but generally at a significantly slower rate than for species. Variation in the species beta‐diversity scores between altitudinal levels did not follow a homogeneous pattern in the different regions. Two main altitudinal groups of sites with a boundary c. 1500–1750 m a.s.l. can be detected with respect to faunistic similarity. Low‐ and mid‐altitude sites are inhabited by all of the genera (19) and 80% of all species collected. Eight genera and 61 species (c. 60% of the total) are unable to inhabit high‐altitude sites, and only 20 species appear to be exclusive to these high‐altitude environments (> 2000 m a.s.l.). Main conclusions The dominant processes explaining dung beetle composition in the high north‐eastern Andean mountains are probably those of vertical colonization. The limited role of horizontal colonization processes, or colonization from northern or southern lineages, could be a consequence of the isolation and recent geological origin of these mountains.     

Phylogeny,diversification patterns and historical biogeography of euglossine orchid bees (Hymenoptera: Apidae)

The orchid bees constitute a clade of prominent insect pollinators distributed throughout the Neotropical region. Males of all species collect fragrances from natural sources, including flowers, decaying vegetation and fungi, and store them in specialized leg pockets to later expose during courtship display. In addition, orchid bees provide pollination services to a diverse array of Neotropical angiosperms when foraging for food and nesting materials. However, despite their ecological importance, little is known about the evolutionary history of orchid bees. Here, we present a comprehensive molecular phylogenetic analysis based on ～4.0 kb of DNA from four loci [cytochrome oxidase (CO1), elongation factor 1‐α (EF1‐α), arginine kinase (ArgK) and RNA polymerase II (Pol‐II)] across the entire tribe Euglossini, including all five genera, eight subgenera and 126 of the approximately 200 known species. We investigated lineage diversification using fossil‐calibrated molecular clocks and the evolution of morphological traits using disparity‐through‐time plots. In addition, we inferred past biogeographical events by implementing model‐based likelihood methods. Our dataset supports a new view on generic relationships and indicates that the cleptoparasitic genus Exaerete is sister to the remaining orchid bee genera. Our divergence time estimates indicate that extant orchid bee lineages shared a most recent common ancestor at 27–42 Mya. In addition, our analysis of morphology shows that tongue length and body size experienced rapid disparity bursts that coincide with the origin of diverse genera (Euglossa and Eufriesea). Finally, our analysis of historical biogeography indicates that early diversification episodes shared a history on both sides of Mesoamerica, where orchid bees dispersed across the Caribbean, and through a Panamanian connection, thus reinforcing the hypothesis that recent geological events (e.g. the formation of the isthmus of Panama) contributed to the diversification of the rich Neotropical biota. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 552–572.     

The validity of ecogeographical rules is context‐dependent: testing for Bergmann's and Allen's rules by latitude and elevation in a widespread Andean duck

Consistent responses by various organisms to common environmental pressures represent strong evidence of natural selection driving geographical variation. According to Bergmann's and Allen's rules, animals from colder habitats are larger and have smaller limbs than those from warmer habitats to minimize heat loss. Although evidence supporting both rules in different organisms exists, most studies have considered only elevational or latitudinal temperature gradients. We tested for the effects of temperature associated with both elevation and latitude on body and appendage size of torrent ducks (Merganetta armata), a widespread species in Andean rivers. We found a negative relationship between body size and temperature across latitude consistent with Bergmann's rule, whereas there was a positive relationship between these variables along replicate elevational gradients at different latitudes. Limb‐size variation did not support Allen's rule along latitude, nor along elevation. High‐elevation ducks were smaller and had longer wings than those inhabiting lower elevations within a river. We hypothesize that temperature is likely a major selective pressure acting on morphology across latitudes, although hypoxia or air density may be more important along elevational gradients. We conclude that the effect of temperature on morphology, and hence the likelihood of documenting ecogeographical ‘rules’, depends on the environmental context in which temperature variation is examined. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111, 850–862.     

A new genus in the diverse Andean Pedaliodes complex uncovered using target enrichment (Lepidoptera,Nymphalidae)

A new genus of Neotropical Satyrinae butterflies, Viloriodes Pyrcz & Espeland gen. n. is described in the Pedaliodes Butler complex comprising 11–13 genera and more than 400 species. Support for the new genus is provided by a phylogenetic analysis based on target enrichment (TE) data including 618 nuclear loci with a total of 248,940 nucleotides, and the mitochondrial gene cytochrome oxidase subunit 1 (COI). Five species, whose DNA sequences were obtained by TE during this study, form a strongly supported clade sister to the large clade comprising Pedaliodes and four other genera. Complementary COI analysis confirms the monophyly of Viloriodes gen. n., with the above five plus eight other species clustering in highly supported clades in both Bayesian Inference and Maximum Likelihood analyses, and a TE + COI concatenated tree. Based on molecular and morphological data, 30 species are assigned to Viloriodes gen. n. The new genus can be recognized by a set of subtle morphological characteristics of colour patterns and male and female genitalia. An analysis of divergence times indicates that Viloriodes gen. n. and Steromapedaliodes Forster separated around 5.9 Mya. Viloriodes gen. n. has a wider geographic distribution than any other genus of the Pedaliodes complex, being found from central Mexico to northern Argentina and to the Guyana Shield, typically occurring at lower elevations than Pedaliodes.     

Reproductive biology and mating system estimates of two Andean melocacti, Melocactus schatzlii and M. andinus (Cactaceae)

BACKGROUND AND AIMS: The genus Melocactus comprises 36 species of globose cacti with the most derived traits in the Cereeae tribe. It is the proper study system to examine what are the most derived reproductive strategies within that tribe. This study aims to characterize the reproductive biology and to estimate the mating system parameters of two Andean melocacti, Melocactus schatzlii and M. andinus. METHODS: The reproductive attributes of the two species were described, including floral morphology, anthesis patterns, floral rewards, floral visitors and visitation patterns. Levels of self-compatibility and autonomous self-pollination were estimated by hand-pollination experiments. Mating system estimates were obtained by conducting progeny array analyses using isozymes. KEY RESULTS: The flowers of the two species present the typical hummingbird-pollination syndrome. Despite their morphological resemblance, the two species differ in flower size, pollen and ovule production and anthesis pattern. Their main pollinator agents are hummingbirds, four species in M. schatzlii and one species in M. andinus. Both cacti are self-compatible and capable of self-pollination without the aid of pollen vectors. Population-level outcrossing rate was higher for M. schatzlii (t(m)=0.9) than for M. andinus (t(m)=0.4). At the family level, outcrossing rates for most mothers of M. schatzlii were higher (t(m)>0.8) than for M. andinus (t(m)<0.5). CONCLUSIONS: Although the two cacti are capable of selfing, M. schatzlii is a predominantly outcrossing species, while M. andinus behaves as a mixed-mating cactus. Hummingbirds are the only pollinators responsible for outcrossing and gene flow events in these species. In their absence, both melocacti set seeds by selfing. Based on its low population size, restricted distribution in Venezuela, low rates of floral visits, and high levels of inbreeding, M. andinus is considered to be an endangered species deserving further study to define its conservation status.     

Tree-line changes along the Andes: implications of spatial patterns and dynamics

The Andes provide an extensive latitudinal and topographical framework for studying the factors that control the spatial patterns of forests (timberlines) and their species components expressed through the presence of tree growth forms (tree lines). Despite consistent overall similarities in landscape patterns, many processes must be unique, given the dramatic differences in species richness and biophysical constraints along the Andes. In all cases evaluated to date, morphological plasticity is a common trait of plant species that dominate at tree lines. In fact, many changes observed can be related to species-specific traits. Physiological limitations on tree growth form only explain species limits, while disturbances and cyclical climate fluctuations interact to affect many landscape patterns. Over long periods of time, tree lines provide unique habitats and perhaps opportunities for speciation. Understanding the spatial organization of tree-line dynamics is one viable research approach for evaluating the likely past fluxes and possible future changes.