Searching for Altitudinal Zonation: Species Distribution and Vegetation Composition in the Superpáramo of Volcán Iliniza, Ecuador

Altitudinal variation of the zonal superpáramo vegetation was studied between 4300 and 4630 m to test a possible occurrence of a fine altitudinal zonation within the superpáramo belt. A rectangular grid of 1 m² sample plots was established; 25 replicate plots separated by a 3 m space were located along a 100 m long transect parallel to the contours, and there were 17 such transects separated by 20 m of altitude. Species were scored using a 7-grade cover scale and basic environmental data were gathered for each sample. Major changes occur over a short altitudinal range, at around 4400 m, which corresponds to a transition between the lower and upper superpáramo. Species richness sharply declines but species turnover (per altitude) increases along the altitudinal gradient. The correlation between richness and bare ground or rock cover is negative, but the correlation to rocks becomes positive above 4500 m. Species from lower altitudes tend to have narrower altitudinal range, although a large number of species appear to be indifferent to altitude. Direct ordination analyses indicate that high-altitude species show stronger correlation to environmental variables, especially rock, than species from lower altitudes. TWINSPAN cluster analysis delimited 15 groups of samples. There is a change in the clustering pattern along the altitudinal gradient from a horizontal (i.e., within altitude) to vertical (i.e., across altitude) arrangement of the cluster groups, although this pattern is partly obscured at the highest altitudes due to a large number of empty samples. MANOVA tests for samples from adjacent altitudinal levels indicate two distinct altitudinal breaks at lower altitudes, corresponding to the Loricaria-belt in lower superpáramo and the transition between lower and upper superpáramo, while no indication of a zonation was found in upper superpáramo.     

A multi-species assessment of post-dispersal seed predation in the central Chilean Andes

BACKGROUND AND AIMS: Post-dispersal seed predation in alpine communities has received little attention despite evidence that seeds removed by granivores can decrease plant recruitment into ecosystems. Moreover, few studies have assessed the effects of removal of seeds of a range of species after dispersal on the seeds remaining in ecosystems. A comparison was made of the magnitude of seed removal by ants and birds of nine different shrubby-, herbaceous- and cushion-plant species in the central Chilean Andes in order to assess the interactions between birds, ants and wind, and the types of seeds. METHODS: A total of 324 soil-covered plates, each containing 50 seeds of one species, were placed in the field at an altitude of 2700 m and assigned to one of four treatments: control, exclusion of ants, birds, and both. The design also allowed the effects of wind to be assessed. Seed removal from plates was monitored over 20 d. KEY RESULTS: Mean accumulative seed removal by granivores averaged over all nine species combined was 25%. However, large differences between species were evident, with limited seed removal (3-11%) in three herbaceous species (Alstroemeria pallida, Sisyrinchium arenarium, Pozoa coriacea), moderate (18-33%) in five species, including a shrub (Chuquiraga oppositifolia), two herbs (Taraxacum officinale, Rhodophiala rhodolirion), and two cushion-plants (Laretia acaulis, Azorella monantha), and substantial (78%) in the shrub Anarthrophyllum cumingii. The magnitudes of losses caused by birds compared with ants did not differ for the majority of species, although removal by birds was greater than by ants in A. cumingii, and smaller for C. oppositifolia. CONCLUSIONS: Post-dispersal seed removal is shown to be an important cause of decreased potential plant species recruitment into alpine ecosystems. The substantial differences in the magnitude of seed losses to ants and birds demonstrate the need for evaluation of seed removal on a wide range of species in any given ecosystem.     

The population genetics of Quechuas, the largest native South American group: autosomal sequences, SNPs, and microsatellites evidence high level of diversity

Elucidating the pattern of genetic diversity for non-European populations is necessary to make the benefits of human genetics research available to individuals from these groups. In the era of large human genomic initiatives, Native American populations have been neglected, in particular, the Quechua, the largest South Amerindian group settled along the Andes. We characterized the genetic diversity of a Quechua population in a global setting, using autosomal noncoding sequences (nine unlinked loci for a total of 16 kb), 351 unlinked SNPs and 678 microsatellites and tested predictions of the model of the evolution of Native Americans proposed by (Tarazona-Santos et al.: Am J Hum Genet 68 (2001) 1485-1496). European admixture is <5% and African ancestry is barely detectable in the studied population. The largest genetic distances were between African versus Quechua or Melanesian populations, which is concordant with the African origin of modern humans and the fact that South America was the last part of the world to be peopled. The diversity in the Quechua population is comparable with that of Eurasian populations, and the allele frequency spectrum based on resequencing data does not reflect a reduction in the proportion of rare alleles. Thus, the Quechua population is a large reservoir of common and rare genetic variants of South Amerindians. These results are consistent with and complement our evolutionary model of South Amerindians (Tarazona-Santos et al.: Am J Hum Genet 68 (2001) 1485-1496), proposed based on Y-chromosome data, which predicts high genomic diversity due to the high level of gene flow between Andean populations and their long-term effective population size.     

Nurse effect of the native cushion plant Azorella monantha on the invasive non-native Taraxacum officinale in the high-Andes of central Chile

Positive interactions among native plant species are common in alpine habitats, particularly those where one species (nurse plant) generates microclimatic conditions that are more benign than the surrounding environment, facilitating the establishment of other species. Nonetheless, these microclimatic conditions could facilitate the establishment of non-native species as well. A conspicuous component of the alien alpine flora of the central Chilean Andes is the perennial herb Taraxacum officinale agg. (dandelion). In contrast to other alien species that are restricted to human-disturbed sites, T. officinale is frequently observed growing within native plant communities dominated by cushion plants. In this study we evaluated if T. officinale is positively associated with the cushion plant Azorella monantha. Via seedling survival experiments and gas-exchange measurements we also assessed the patterns of facilitation between cushions and dandelions, and explore the potential mechanisms of invasion by dandelions. T. officinale grows spatially positively associated with cushions of A. monantha. Survival of seedlings, as well as their net-photosynthetic rates and stomatal conductance, were higher within cushions than in open areas away from them, suggesting that the microclimatic modifications generated by this native cushion facilitates the establishment and performance of a non-native invasive species. Our results, as well as other recent studies, highlight the role of native communities in facilitating rather than constraining non-native plant invasions, particularly in stressful habitats such as alpine environments.     

Landscape diversity patterns and endemism of Araceae in Ecuador

Araceae is one of the largest herb families in tropical America, but its patterns of diversity and endemism are poorly known. We used predictive distribution modelling in GIS to study Araceae richness on a landscape scale in Ecuador. Modelling was based on georeferenced herbarium collections with humidity and mean annual temperature as climatic variables. Variation partitioning using multiple regression showed that humidity and altitude were main factors in explaining Araceae diversity patterns. Endemism was less well explained by present climatic factors. Unlike common diversity patterns of herbs or epiphytes, Araceae richness was highest in the eastern (Amazon) lowland rain forest with a secondary centre on the Andean foothills of northwestern Ecuador. The peak in endemism was on the western slopes of the Andes, corresponding to areas that have been severely affected by human activities and deforestation. Eastern lowland (Amazonian) forests were poor in endemic Araceae.     

Altitudinal distribution limits of aquatic macroinvertebrates: an experimental test in a tropical alpine stream

1. Temperature and oxygen are recognised as the main drivers of altitudinal limits of species distributions. However, the two factors are linked, and both decrease with altitude, why their effects are difficult to disentangle. 2. This was experimentally addressed using aquatic macroinvertebrates; larvae of Andesiops (Ephemeroptera), Claudioperla, (Plecoptera), Scirtes (Coleoptera) and Anomalocosmoecus (Trichoptera), and the amphipod Hyalella in an Ecuadorian glacier‐fed stream (4100–4500 m a.s.l.). The following were performed: (i) quantitative benthic sampling at three sites to determine altitudinal patterns in population densities, (ii) transplants of the five taxa upstream of their natural altitudinal limit to test the short‐term (14 days) effect on survival, and (iii) in situ experiments of locomotory activity as a proxy for animal response to relatively small differences in temperature (5 °C vs. 10 °C) and oxygen saturation (55% vs. 62%). 3. The transplant experiment reduced survival to a varying degree among taxa, but Claudioperla survived well at a site where it did not naturally occur. In the in situ experiment, Scirtes and Hyalella decreased their activity at lower oxygen saturation, whereas Andesiops and Anomalocosmoecus did so at a low temperature. The decrease in activity from a high to a low temperature and oxygen for the five taxa was significantly correlated with their mortality in the transplant experiment. 4. Together the present experiments indicate that even relatively small differences in temperature and oxygen may produce effects explaining ecological patterns, and depending on the taxon, either water temperature or oxygen saturation, without clear interacting effects, are important drivers of altitudinal limits.     

Tropical ant communities are in long-term equilibrium

Communities change with time. Studying long-term change in community structure permits deeper understanding of community dynamics, and allows us to forecast community responses to perturbations at local (e.g. fire, secondary succession) and global (e.g. desertification, global warming) spatial scales. Monitoring efforts exploring the temporal dynamics of indicator taxa are therefore a critical part of conservation agendas. Here, the temporal dynamics of the Otongachi leaf litter ant community, occurring in a cloud forest in coastal Ecuador, were explored. By sampling this community six times over eleven years, I assessed how the ant fauna caught by Winkler traps (more diverse and cryptic fauna) and caught by pitfall traps (larger, more mobile fauna) changed over time. The Otongachi leaf litter ant community was dynamic. Although species richness in the community remained constant, temporal turnover of species was high: on average, 51% of the ant species in Winkler traps, and 56% of those in pitfall traps, were replaced with other ant species from one year to the other. Shifts in the rank abundance of species in the community were also large across the eleven years and, on average, shifts in the rank abundance of species collected by Winkler traps doubled those occurring in pitfall traps from one census to the other. In spite of these trends, the Otongachi ant fauna showed no (Winkler) or weak (pitfall) evidence of directional change (towards a new community). Thus, this tropical ant community can be divided in two community compartments. The Winkler compartment composed by a more diverse and cryptic ant fauna appears to be resilient and stable in time. The pitfall compartment composed by larger and more mobile ants may be prone to respond to disturbance. This study suggests that 1) species appearing/disappearing from a site may be rather the rule, difficult to separate from responses to ecological stress. 2) Conclusions made in short-term studies, or studies comparing two (e.g. before and after) snapshots of a community, should thus be revisited. Finally, 3) the ant fauna caught by pitfall traps (a rather simple and cheap survey method) is the most likely community compartment to indicate ecological perturbation. This study adds to the growing evidence that using ants as ecological indicators should incorporate long-term temporal dynamics.     

Cedrela nebulosa: A novel species for dendroclimatological studies in the montane tropics of South America

Up to now, the development of dendrochronological records from tropical regions in South America has been limited to the lowlands with emphasis in the Amazon basin. In this contribution, we present the first chronology of Cedrela nebulosa, a species that develops in the tropical mountainous regions of South America. We collected samples from trees in Monobamba district in Peru, analysed the anatomical features that determine the growth rings, and processed following the methods commonly used in dendrochronology. The 133-years chronology covering the 1883–2015 period, showed large correlation between series. In order to determine the climatic variables that control tree growth, we performed correlation analyses between tree-growth and local and regional precipitation and temperature records. We found that precipitation triggers tree growth at the beginning of the spring season but temperature seems to be the main control in annual growth. Also, C. nebulosa chronology present coherent variations with Multivariate Enso Index (MEI) and Pacific Ocean sea surface temperatures during summer months. This climate-sensitive tree-ring record indicates good potential for dendroclimatic studies and provides an opportunity to reconstruct climatic variations in montane forests of the tropical Andes.     

The elevational gradient of Andean plant endemism: varying influences of taxon-specific traits and topography at different taxonomic levels

Aim Little is known about the elevational gradient of plant endemism. It is mostly assumed that patterns are determined by topographical factors such as area of elevational belts and degree of habitat fragmentation, but comparative studies of different plant taxa along the same elevational gradient are lacking. The aim was to compare the elevational patterns of plant endemism of the entire flora and selected families and genera in a search for commonalities. Methods The elevational patterns of endemism for the entire Ecuadorean vascular plant flora, for twenty‐seven selected families, and for twenty‐four selected genera based on the Catalogue of Vascular Plants of Ecuador were analysed. Results Elevational patterns of endemism were non‐random at all taxonomic levels but there was no common elevational pattern. Rather, the study groups showed a wide variety of independent patterns at all taxonomic levels. Most groups had hump‐shaped patterns with maxima at different elevations and mostly at the same or at higher elevations than the maxima of species richness. The overall flora showed highest endemism in the narrowest and most fragmented elevational belts, presumably because of the consequent fragmentation of species populations. Main conclusions Patterns of endemism appear to be influenced both by taxon‐specific ecological traits (e.g. life form, reproduction, dispersal, demography, spatial population structure, competitive ability) in their specific interaction with historical processes and by environmental factors such as topographical fragmentation. The degree to which these influences become visible along the elevational gradient are determined by which combination of species is analysed: for a given genus or family, taxon‐specific traits dominate the patterns, for the entire flora taxon‐specific patterns are blurred by averaging and the signal of topography emerges. Beyond the elevational gradient as such, this study shows that the frequently stated assumption that given biogeographical settings lead to similar patterns of endemism among different taxa is wrong.