Evolution in the High Andes: The Phylogenetics of Muscisaxicola Ground-Tyrants

Phylogenetic relationships within the genus Muscisaxicola, a primarily Andean group of tyrant-flycatchers, were studied using complete sequences of the mitochondrial genes COII and ND3. Relationships among Muscisaxicola species were found to differ substantially from those of previous views, suggesting convergence in traditional avian taxonomic characters within the genus. The 11 species of large, gray, “typical” Muscisaxicola flycatchers (including M. grisea, newly restored to species status) formed a distinct clade, consisting of two major groups: a clade of 6 species breeding primarily in the central Andes and a clade of 5 species breeding primarily in the southern Andes. The other 2 species traditionally placed in this genus, M. fluviatilis, an Amazonian species, and M. maculirostris, were both rather divergent genetically from the typical species, although M. maculirostris may be the sister taxon to the typical clade. The patterns of sympatry exhibited by Muscisaxicola species in the high Andes appear to be the consequence of speciation and secondary contact within regions of the Andes, rather than a result of dispersal between regions. Diversification of the typical Muscisaxicola species appears to have occurred during the middle and late Pleistocene, suggesting generally that taxa of the high Andes and Patagonia may have been greatly influenced by mid-to-late Pleistocene events. There were likely several independent developments of migration within this genus, but migration is probably ancestral in the southern clade, with subsequent loss of migration in two taxa.     

Evolution in the deep sea: Biological traits, ecology and phylogenetics of pelagic copepods

Deep-sea biodiversity has received increasing interest in the last decade, mainly focusing on benthic communities. In contrast, studies of zooplankton in the meso- to bathypelagic zones are relatively scarce. In order to explore evolutionary processes in the pelagic deep sea, the present study focuses on copepods of two clausocalanoid families, Euchaetidae and Aetideidae, which are abundant and species-rich in the deep-sea pelagic realm. Molecular phylogenies based on concatenated-portioned data on 18S, 28S and internal transcribed spacer 2 (ITS2), as well as mitochondrial cytochrome c oxidase subunit I (COI), were examined on 13 species, mainly from Arctic and Antarctic regions, together with species-specific biological traits (i.e. vertical occurrence, feeding behaviour, dietary preferences, energy storage, and reproductive strategy). Relationships were resolved on genus, species and even sub-species levels, the latter two established by COI with maximum average genetic distances ranging from ?5.3% at the intra-specific, and 20.6% at the inter-specific level. There is no resolution at a family level, emphasising the state of Euchaetidae and Aetideidae as sister families and suggesting a fast radiation of these lineages, a hypothesis which is further supported by biological parameters. Euchaetidae were similar in lipid-specific energy storage, reproductive strategy, as well as feeding behaviour and dietary preference. In contrast, Aetideidae were more diverse, comprising a variety of characteristics ranging from similar adaptations within Paraeuchaeta, to genera consisting of species with completely different reproductive and feeding ecologies. Reproductive strategies were generally similar within each aetideid genus, but differed between genera. Closely related species (congeners), which were similar in the aforementioned biological and ecological traits, generally occurred in different depth layers, suggesting that vertical partitioning of the water column represents an important mechanism in the speciation processes for these deep-sea copepods. High COI divergence between Arctic and Antarctic specimens of the mesopelagic cosmopolitan Gaetanus tenuispinus and the bipolar Aetideopsis minor suggest different geographic forms, potentially cryptic species or sibling species. On the contrary, Arctic and Antarctic individuals of the bathypelagic cosmopolitans Gaetanus brevispinus and Paraeuchaeta barbata were very similar in COI sequence, suggesting more gene flow at depth and/or that driving forces for speciation were less pronounced in bathypelagic than at mesopelagic depths.     

Three native tuber foods of the high Andes

Potatoes, anues, ocas and ullucus fed the pre- Columbian Incas and still feed their modern descendants, but only the potato has traveled to other parts of the world. Who knows what rivalry the potato might have had if the other three had received the same attention in efforts toward improvement?     

Thermal extremes at high elevations in the Andes: Physiological ecology of frogs

1. 1. Some anurans live in the tropical Andes above 3000 m, a cold and variable environment.

2. 2. Coping with high-elevation climate includes shifts in activity patterns, avoidance of extreme temperatures, and thermoregulation by heliothermy and thigmothermy.

3. 3. Behavior alone does not allow for full compensation of low environmental temperatures.

4. 4. Modifications of metabolic physiology allow high-elevation frogs to be active at low temperatures. However at warm temperatures their thermal physiology is similar to that of low-elevation forms.

     

Evolution of host specialization in the Adelgidae (Insecta: Hemiptera) inferred from molecular phylogenetics

The Adelgidae form a small group of insects in the Aphidoidea. They are cyclically parthenogenetic with host alternating, multiple-generation complex life cycles and are restricted to certain host genera in the Pinaceae. Species that host alternate always have Picea as the primary host where sexual reproduction and gall formation occur, and another genus in the Pinaceae as the secondary host where a series of parthenogenetic generations are produced. Other species that do not host alternate complete their entire life cycle on one host and only reproduce parthenogenetically. We studied relationships within Adelgidae using DNA sequences from the mitochondrial COI, COII, and cytb genes, and the nuclear EF1alpha gene. Analysis of the combined data resulted in a well-resolved phylogeny in which the major adelgid clades correspond neatly to their association with secondary host genera. Specialization on each secondary host genus occurred only once and was followed by diversification on the host genus. Molecular dating of divergence times in the Adelgidae suggest that diversification among host genera occurred in the Late Cretaceous and Early Tertiary when the Pinaceae genera were diverging. It is not clear, however, whether the Adelgidae and Pinaceae co-diversified because the relationships among the Pinaceae genera are not fully resolved. We discuss implications for adelgid taxonomy, life cycle evolution, and evolution of the interaction between adelgids and their host plants.     

Synchronous fire activity in the tropical high Andes: an indication of regional climate forcing

Global climate models suggest enhanced warming of the tropical mid and upper troposphere, with larger temperature rise rates at higher elevations. Changes in fire activity are amongst the most significant ecological consequences of rising temperatures and changing hydrological properties in mountainous ecosystems, and there is a global evidence of increased fire activity with elevation. Whilst fire research has become popular in the tropical lowlands, much less is known of the tropical high Andean region (>2000masl, from Colombia to Bolivia). This study examines fire trends in the high Andes for three ecosystems, the Puna, the Paramo and the Yungas, for the period 1982–2006. We pose three questions: (i) is there an increased fire response with elevation? (ii) does the El Niño‐ Southern Oscillation control fire activity in this region? (iii) are the observed fire trends human driven (e.g., human practices and their effects on fuel build‐up) or climate driven? We did not find evidence of increased fire activity with elevation but, instead, a quasicyclic and synchronous fire response in Ecuador, Peru and Bolivia, suggesting the influence of high‐frequency climate forcing on fire responses on a subcontinental scale, in the high Andes. ENSO variability did not show a significant relation to fire activity for these three countries, partly because ENSO variability did not significantly relate to precipitation extremes, although it strongly did to temperature extremes. Whilst ENSO did not individually lead the observed regional fire trends, our results suggest a climate influence on fire activity, mainly through a sawtooth pattern of precipitation (increased rainfall before fire‐peak seasons (t‐1) followed by drought spells and unusual low temperatures (t0), which is particularly common where fire is carried by low fuel loads (e.g., grasslands and fine fuel). This climatic sawtooth appeared as the main driver of fire trends, above local human influences and fuel build‐up cyclicity.     

Evolution into and out of the Andes: a Bayesian analysis of historical diversification in Thamnophilus antshrikes

The Andean uplift played important roles in the historical diversification of Neotropical organisms, both by producing new high-elevation habitats that could be colonized and by isolating organisms on either side of the mountains. Here, we present a molecular phylogeny of Thamnophlius antshrikes, a clade of 30 species whose collective distribution spans nearly the entirety of lowland habitats in tropical South America, the eastern slope foothills of the Andes, and the tepuis of northern South America. Our goal was to examine the role of the Andes in the diversification of lowland and foothill species. Using parsimony and Bayesian ancestral state reconstructions of a three-state distribution character (lowland-restricted, lowland-to-highland, highland-restricted), we found that the Andes were colonized twice independently and the tepuis once from lowland-restricted ancestors. Over the entire evolutionary history of Thamnophilus, the highest transition rates were between highland-restricted and lowland-to-highland distributions, with extremely low rates into and out of lowland-restricted distributions. This pattern suggests lowland-restricted distributions are limited not by physiological constraints, but by other forces, such as competition. These results highlight the need for additional comparative studies in elucidating processes associated with the colonization of high-elevation habitats and the differentiation of populations within them.     

Presence of cushion plants increases community diversity in the high equatorial Andes

Cushion plants are a common growth form in the equatorial páramo vegetation and their surfaces are often colonized by other plants. This paper analyzes the effect of the cushion plants on the community diversity at 4650 m on the eastern slope of the Iliniza volcano in Ecuador. Ninety sample plots of 1 m² size were located in the study area and were divided into 25 subplots in which presence and abundance of plant species was recorded. The community diversity was expressed as species richness, Simpson's diversity index, and evenness. Correlation between the cushion species and the composition of the colonists was measured with the CCA ordination analysis, correlation between the cushion cover and community diversity was measured by means of correlation analysis. Randomized species–area curves were used to compare richness of plant communities with and without the cushions. A total of 32 species were found including five cushion plants. Most species preferred to grow on the cushion surface whereas only a few species were able to colonize open ground. Species richness and Simpson's index were significantly correlated to the cushion area but no correlation was found for evenness. The cushions were usually composed of more than one species which hampered the examination of the cushion–colonist specific relationships. Nevertheless, cushions of Azorella and Arenaria seemed to provide more favorable habitat for colonization than the other cushion species. Comparison with an earlier study made on Iliniza indicates that the presence of the cushions significantly increases the richness of the plant community.     

Tropical high Andean drylands: species diversity and its environmental determinants in the Central Andes

Diversity determinants have mostly been evaluated in high diversity areas, leaving behind regions with less species diversity such as drylands. Here we aim to analyze the patterns of plant diversity in tropical drylands in the southern Central Andes, and determine the importance of water, energy, and environmental heterogeneity as diversity determinants of the arid and semi-arid adapted flora. We examined the distribution of 645 native species from lowlands to 6000 m.a.s.l. in the north-western region of Argentina (NWA) and define hotspots of diversity within each NWA ecoregion. Diversity is concentrated in regions of middle elevation with intermediate values of water and energy, at the transition between arid and semi-arid regions. Furthermore, we showed that in tropical drylands energy input is as fundamental for plant diversity as water input is and, we found that the effects of these variables varied with elevation and, also with aridity. Water variables had the strongest effect on the flora in the arid high Andean ecoregions, where an increase in precipitation during the growing season stimulated species diversity. Energy only became more important than water when the arid adapted flora entered the low and semi-arid regions where energy increments reduce species diversity. Our analysis provides strong quantitative support for climate variables as the main determinants of plant diversity across different ecoregions of the southern Central Andes. Given the present climate change events, knowing how these variables affect the distribution of the arid adapted flora is crucial for planning strategies for achieve their present and future conservation.     

Cloud frequency correlates to plant species composition in the high Andes of Ecuador

Two data sets representing atmospheric moisture are available for the high Andes of Ecuador, (i) cloud frequency obtained from weather satellites, and (ii) interpolated rainfall estimates obtained from global climate observations. We analyzed their correlation to vascular plant species composition at 18 Ecuadorian superpáramo study sites. Of particular interest was whether cloud frequency could be used as a proxy for precipitation. Cloud frequency had distinct seasonal and spatial variation among the sites. The spatial gradient of cloudiness was strongest during June–August and weakest around the equinoctials. Interpolated rainfall estimates also showed seasonal and spatial variation among the sites, but there was no correlation between them and cloudiness. Cloud frequency during June–August was the only significant variable in the canonical correspondence analysis (CCA) of the species composition, whereas rainfall estimates, geology, presence of glaciers, and size and altitudinal range of the superpáramos were not significant. When the spatial components were filtered out from the species composition data by employing partial CCA, cloud frequency during December–February became the only significant variable. Our results suggest that cloud frequency data may be a useful tool in mountain ecology research, serving as an indicator of habitat humidity when exact precipitation data are lacking.     

Traditional livestock activities modify the spatial behavior of small wildcats in the high Andes

Human activities lead to declines in species’ abundance and diversity, as well as loss of habitat quality from overexploitation. The effects of livestock grazing increase on small felids are poorly understood, primarily due to the low detectability of these species. The grazing of domestic camelids, such as the llama (Lama glama) and the alpaca (Vicugna pacos), is the main traditional source of income for the inhabitants of the Andes, where the Andean cat (Leopardus jacobita) and Pampas cat (L. colocolo) are also present. In this study, we employ camera trap data to evaluate the association of llama and alpaca grazing with occupancy by the Andean and Pampas cat in Sajama National Park. In each of 36 sampling stations, we installed a camera trap, a grid of traps for rodents and two habitat transects along a gradient of intensity of livestock grazing. Occupancy models showed that negative effect of camelid abundance was stronger for the Andean cat than the Pampas cat. For both species, occupancy was also associated positively to the abundance of prey, proximity to water bodies, and vegetation. Given that livestock activity is a way of life for human populations in the Andes, we suggest that mechanisms should be implemented to compensate for the negative effects of livestock on Andean and Pampas cat populations.     

Biomass allocation in herbaceous plants under grazing impact in the high semi-arid Andes

Among the plant traits that affect performance, vitality and herbivore resistance in rangeland vegetation, biomass partitioning ranks top, commonly outweighing processes at single leaves (e.g., photosynthesis, respiration). We explored the allometry of a broad sample of herbaceous species from two high elevation sites in the Andes in order to explain biomass partitioning under harsh climatic conditions and risk of biomass losses under strong camelid grazing pressure. We combined data from NW-Argentina and W-Bolivia from elevations between 4200 and 4250 m in a landscape dominated by tall Festuca orthophylla tussocks and a drought driven seasonality (rainfall only between November and March). Across 10–20 taxa per region we found less investment in leaves with a mean leaf mass fraction of only 11% and instead a massive storage in below-ground compartments (rhizomes, tap roots), particularly at the colder Argentinean site. Though grazing pressure was much greater in Bolivia, the foliage mass fraction was there larger than in Argentina. The inter-tussock space in these open, dry plains (‘pajonal’) was dominated by rosette forming species with a below-ground shoot apex and massive tap roots (70% of all species), rendering these species less sensitive to grazing and trampling. The storage organs of these species represented more than 50% of total biomass. Llamas, which represented the main vertebrate herbivore in these open plains at the Bolivian site, preferred non tap-root herbs and species with low leaf nitrogen concentration. Palatable forbs for llamas (22% of all species at the Bolivian site) only survived when nested (facilitated) in the rigid, tall Festuca orthophylla tussocks or thorny shrubs. In conclusion, these extremely high elevation rangeland herbs invest preferentially in structures for persistence (K-strategy) rather than maximizing carbon gain.     

Evolutionary islands in the Andes: persistence and isolation explain high endemism in Andean dry tropical forests

Aim The tropical Andes are a world biodiversity hotspot. With diverse biomes and dramatic, geologically recent mountain uplift, they offer a system to study the relative contributions of geological and biome history to species richness. There are preliminary indications that historical species assembly in the Andes has been influenced by physiographical heterogeneity and that distinct biomes have evolved in relative isolation despite physical proximity. Here we test this ‘Andean biotic separation hypothesis’ by focusing on the low‐elevation, seasonally dry tropical forest (SDTF) biome to determine whether patterns of plant diversification within the SDTF differ from those in mid‐ and high‐elevation biomes. Location Tropical Andes, South America. Methods Densely sampled time‐calibrated phylogenies for five legume genera (Amicia, Coursetia, Cyathostegia, Mimosa and Poissonia) containing species endemic to the Andean SDTF biome were used to investigate divergence times and levels of geographical structure. Geographical structure was measured using isolation‐by‐distance methods. Meta‐analysis of time‐calibrated phylogenies of Andean plant groups was used to compare the pattern and tempo of endemic species diversification between the major Andean biomes. Results Long‐term persistence of SDTF in the Andes is suggested by old stem ages (5–27 Ma) of endemic genera/clades within genera, and deep divergences coupled with strong geographical structure among and within species. Comparison of species diversification patterns among different biomes shows that the relatively old, geographically confined pattern of species diversification in SDTF contrasts with the high‐elevation grasslands that show rapid and recent radiations driven by ecological opportunities. Main conclusions The SDTF biome has a long history in the Andes. We suggest that the diverse SDTF flora has been assembled gradually over the past c. 19 Ma from lineages exhibiting strong phylogenetic niche conservatism. These patterns suggest that Andean SDTFs have formed stable and strongly isolated ‘islands’ despite the upheavals of Andean uplift. Indeed, the Andean SDTFs may represent some of the most isolated and evolutionarily persistent continental plant communities, similar in many respects to floras of remote oceanic islands.     

Cryptic speciation on the high seas; global phylogenetics of the copepod family Eucalanidae

Few genetic data are currently available to assess patterns of population differentiation and speciation in planktonic taxa that inhabit the open ocean. A phylogenetic study of the oceanic copepod family Eucalanidae was undertaken to develop a model zooplankton taxon in which speciation events can be confidently identified. A global survey of 20 described species (526 individuals) sampled from 88 locations worldwide found high levels of cryptic diversity at the species level. Mitochondrial (16S rRNA, CO1) and nuclear (ITS2) DNA sequence data support 12 new genetic lineages as highly distinct from other populations with which they are currently considered conspecific. Out of these 12, at least four are new species. The circumglobal, boundary current species Rhincalanus nasutus was found to be a cryptic species complex, with genetic divergence between populations unrelated to geographic distance. 'Conspecific' populations of seven species exhibited varying levels of genetic differentiation between Atlantic and Pacific basins, suggesting that continental landmasses form barriers to dispersal for a subset of circumglobal species. A molecular phylogeny of the family based on both mitochondrial (16S rRNA) and nuclear (ITS2, 18S rRNA) gene loci supports monophyly of the family Eucalanidae, all four eucalanid genera and the 'pileatus' and 'subtenuis' species groups.     

Ribosomal RNA phylogenetics: the third dimension

With integration of secondary structures, ribosomal genes have once again become very popular for phylogenetic analyses. This additional source of information to the nucleotide sequence provides a massive boost for taxonomic inferences. Herein, we propose that in the near future a further benefit for phylogenetics with such genes will be very likely by inclusion of the third dimension. For the first time, we determined the tertiary structure of the ribosomal internal transcribed spacer 2 for Chlamydomonas rheinhardtii by application of two different in silico prediction algorithms. We compared these methods with focus on phylogenetic usability. Further, we determined the tertiary structures for closely related green algae to provide a small phylogenetic example. The results suggest that the tertiary structure inherits evolutionary information observable neither within the sequence nor in the secondary structure.