Andean Grasslands Research Articles

Bacterial diversity in high Andean grassland soils disturbed with Lepidium meyenii crops evaluated by metagenomics.

Soil quality is usually determined by its physical-chemical characteristics without taking into account the bacterial communities that play a fundamental role in the chemical decomposition of plant nutrients. In this context, the objective of the study was to evaluate bacterial diversity in high Andean grassland soils disturbed with Lepidium meyenii cultivation under different gradients of use (first, second and third use) and crop development (pre-sowing, hypocotyl development and post-harvest). The sampling was carried out in the Bombón plateau in the central Andes of Peru, during the rainy and low water seasons, by the systematic method based on a specific pattern assigned in a geometric rectangular shape at a depth of 0 - 20 cm. The characterization of the bacterial communities was carried out through the metagenomic sequencing of the 16S rRNA. 376 families of bacteria were reported, of which it was determined that there was a significant change in bacterial composition and distribution in relation to use pressure. There were no major changes due to the development of Lepidium meyenii. The families most sensitive to use pressure and soil poverty indicators were Verrucomicrobiaceae, Acidobacteraceae and Aakkermansiaceae.

Evolutionary diversification in the hyper-diverse montane forests of the tropical Andes: radiation ofMacrocarpaea(Gentianaceae) and the possible role of range expansion

AbstractThe humid mid-elevation montane forests (MMF) of the tropical Andes harbour high levels of plant species diversity, contributing to the exceptional overall diversity of the tropical Andean biodiversity hotspot. However, little is known about the diversification dynamics of MMF plant lineages compared to lineages in other Andean biomes. Here, we use Macrocarpaea (Gentianaceae) to investigate patterns of plant diversification in the MMF, using molecular dating with fossils and secondary calibration (from a family-wide phylogenetic analysis of Gentianaceae). We sequenced 76 of 118 recognized Macrocarpaea spp. for six markers to reconstruct a time-calibrated phylogenetic tree and infer the historical biogeography of Macrocarpaea using maximum-likelihood methods implemented in BioGeoBEARS, estimating diversification rates through time and among lineages with BAMM. We document a rapid radiation of Macrocarpaea in Andean MMF coinciding with rapid colonization and range expansion across the entire distribution of the genus in the Andes starting 7.2 Mya. Our results support allopatric founder-event speciation as the dominant process contributing to geographical phylogenetic structure across the genus. We propose that establishment of the MMF in the late Miocene, when the Andes attained critical elevation to modify regional climates, provided large new areas of suitable habitat for Macrocarpaea to quickly colonize and expand through repeated founder-events. We suggest that this wave of colonization and range expansion triggered rapid diversification, and, as the MMF became progressively occupied, the diversification rate slowed. Our study also supports the idea that MMF plant radiations are older and more slowly evolving than the quickly evolving lineages in the recent Andean high-elevation grasslands.

Andean Grassland Species: Net Aerial Primary Productivity, Density, Ecomorphological Indices, and Soil Characteristics

Andean Grassland Species: Net Aerial Primary Productivity, Density, Ecomorphological Indices, and Soil Characteristics

Effects of chestnut bark (Castanea spp.) tannin extracts on selectivity, dry matter intake, weight gain, and enteric methane emission from llamas (Lama glama) under grazing conditions in the high Andean grasslands

The present study's objective was to evaluate the effects of chestnut bark (Castanea spp.) tannin extracts on selectivity, organic matter intake, dry matter intake, weight gain, and enteric methane emissions from llamas as a mitigation strategy. The study was carried out from August to September 2016 at the Experimental Centre IVITA-Marangani-Cusco-Peru (4200 m above sea level). Twenty young female llamas (4.1 ± 0.34 years old) with an average live weight of 99.2 ± 5.21 kg at the beginning of the experiment were used. The llamas were allocated to two treatments (control and tannin). Organic matter intake was calculated from organic matter excretion and organic matter digestibility, the dry matter intake was calculated by using percentage of ash content of the diet, and methane emissions were determined using the SF6 tracer technique over seven days, while the llamas grazed 9 h per day on a native pasture dominated by Festuca rigescens and Calamagrostis amoena. The tannin group was provided with tannin extract derived from chestnut bark at a dose of 12 g/day orally before grazing. The selectivity in both groups was similar (P > 0.05), consuming 91.5 % of grasses and graminoids, 84.5 % of leaves and stems and 61.5 % of green matter. The control group's feed intake was 1614.64 g/day, while that of the Tannin group was 1624.70 g/day (P > 0.05). Weight gain at 31 days was similar for both groups (P > 0.05) (5.7 and 5.2 kg for the control and tannin groups, respectively). Enteric methane emissions were 27.35 ± 1.573 g/day for the control group and 19.32 ± 1.057 g/day for the tannin group (P < 0.001); showing the same tendency when methane emissions were compared according to llama live weight with 0.24 ± 0.012 and 0.17 ± 0.008 g/kg LW/day for the control and tannin group, respectively, (P < 0.001). It was concluded that the use of tannins in the present study reduced methane emissions by approximately 30 % without affecting selectivity, Dry matter intake, and weight gain in llamas.

Combined impacts of climate and land use change and the future restructuring of Neotropical bat biodiversity

Forecasting the effects of global change on biodiversity is necessary to anticipate the threats operating at different scales in space and time. Climate change may create unsuitable environmental conditions, forcing species to move to persist. However, land-use changes create barriers that limit the access of some species to future available habitats. Here, we project the impacts of climate and land-use change on 228 Neotropical bat species by forecasting changes in environmental suitability, while accounting for the effect of habitat type specialization and simulating dispersal across suitable patches. We also identify the most vulnerable ecoregions and those that may offer future stable refugia. We further investigate potential functional changes by analysing the response of different trophic guilds. We found that the range contraction of habitat specialists, especially frugivores, was more frequent and stronger under all simulated scenarios. Projected changes differ markedly across ecoregions. While the Amazon region is likely to undergo high turnover rates in bat composition, the Andean grassland, Cerrado and Chaco might experience the greatest losses. The expansion of habitat generalists, which forage in open areas and commonly establish large colonies in manmade structures, coupled with the range contraction of habitat specialists is projected to homogenize bat communities across the Neotropics. Overall, dispersal will likely be the key for the future of Neotropical bat diversity. Therefore, safeguarding the refugia highlighted here, by expanding and connecting the existing network of protected areas, for example, may allow species to move in response to global change.

Managing environmental diversity in the eastern foothills of the Andes: pre-Columbian agrarian landscapes in the El Alto-Ancasti mountain range

ABTRACT In this paper we review the growing evidence of anthropogenic landscapespresent in the semi-deciduous neotropical forest biomes of eastern NW Argentina,which have remained relatively neglected in favour of arid to semi-arid western Andean regions. The evidence gathered in de El Alto-Ancasti provides animportant case study where multidisciplinary methodologies have beenapplied to sites that document the emergence and variability in food productionstrategies across the eastern Andean forests and grasslands of NWArgentina. We discuss evidence offarming structures from archaeological surveys, plant management from phy-tolith analysis, and the tempo and nature of settlement from archaeological excavations undertaken at a variety of sites in the El Alto Ancasti mountainrange. We suggest that the communities that inhabited this region during thefirst millennium AD (ca. 1500–1000 BP) established a strategy of ‘overlappingpatchworks’ of food production that were able to contend with considerableseasonal variability. We argue that, through the use of cross-channelling, low river areas, erosion control techniques and the establishment of mesothermal crops,including maize, legumes, and tubers, throughout the region, these societies adopted flexible strategies to adapt to life in a region prone to climatic change

Impact of climate change on the Andean distribution of Poa scaberula (Poaceae)

Grasslands play an important ecological role and are the dominant landscape in the whole extent of the Andes and high elevation regions of South America. Climate change models indicate a strong impact on mountain systems. The climatic conditions of mountain systems are greatly influenced by topography and elevation. Poa scaberula Hook. f. is an Andean grass and its habitat may be threatened by climate change and human practices. Species distribution models, using bioclimatic variables, are often used to predict the future distribution and environmental requirements of species. In this study we prepared two bioclimatic models for P. scaberula using the Maxent algorithm, the first based on the inclusion of the variable elevation (E +) and the second on the exclusion of elevation (E-) to identify the main environmental variables for habitat suitability shift, to examine changes in the extent of the area of suitable habitat under current and future climate scenarios, and to project and quantify the spatial pattern of shifts in the areal extent of suitable habitat under future climate conditions. We then compared both projections and evaluate the contribution of the elevation variable in the performance of the models. We observed that the inclusion of elevation decreased the performance of the models. In models with omission of the elevation mean cold hardiness, temperature seasonality and annual mean temperature emerge as the critical factors shaping habitat suitability for P. scaberula. Under the low and higher concentration greenhouse gas emissions scenario, the range of the species may decrease as global warming intensifies. The information gained from this study should be an useful reference for development biogeographic studies and the impact of global warming on regions of high elevations, and contribute to implement conservation and management strategies for Andean grassland and high elevation grasslands.

Lead transfer in the soil-root-plant system in a highly contaminated Andean area.

Lead (Pb) is highly toxic heavy metal that is detrimental to the food system. There are large mining and metallurgical companies in the central highlands of Peru that have been active for almost a century and contribute to air, water, and soil pollution, affecting food quality and causing damage to the environment and human health. Our study, conducted in 2018, assessed the content and transfer of lead in the soil-root-plant system in the high Andean grasslands in a geographical area near the metallurgical complex of La Oroya. Lead levels were measured in 120 samples of top soil (0–20 cm), roots, and grass shoots by flame atomic absorption spectroscopy. No significant differences were found between the soil pH, organic matter content, and lead among the samples evaluated (P > 0.05). Mean Pb concentrations decreased in the order of soil > root > shoot (P < 0.01) (212.36 ± 38.40, 154.65 ± 52.85 and 19.71 ± 2.81 mg/kg, respectively). The soil-to-root Pb bioconcentration factor, root-to-shoot translocation factor, and soil-to-shoot bioaccumulation factor values were 0.74 ± 0.26, 0.14 ± 0.06 and 0.10 ± 0.03, respectively. Lead in the soil was 3.03 times higher than the maximum limit for agricultural soil, and was 1.97 times higher than the value limit for fodder. Our findings are important and show that soils and pasture in this geographical area have high Pb levels due to metallurgical emissions that have been occurring since 1922. Such pollution negatively impacts health and the socio-economic status of the exposed populations.

Spatial distribution of Poa scaberula (poaceae) along the andes

Mountains support a great diversity of species and habitat types. Grasslands are the dominant landscape in the Andes and play an important ecological role. However, they are threatened by many factors, including climate change and human activities. The spatial distribution of species that compose, and the ecological and evolutionary factors that provide for the spatial biodiversity patterns, are little known. The largest Poa L. (Poaceae) genera are widely diversified and distributed in the Andes. In particular, P. scaberula Hook. f. shows great environmentally mediated phenotypic plasticity, and is distributed from North America to the tip of South America. However, the impact of environmental variables has on the spatial distribution of this species, remain largely unknown. Using high-resolution climatic data, herein we modeled the current suitable habitat for P. scaberula and identified the main climatic variables that best predict its potential distribution. In addition, we assess the species status in the predicted habitats through herbarium data and relate it with species distribution models. The models showed that P. scaberula has a suitable habitat of ca. 162.747 km2 along the Andes and high elevation regions. The most influential variables with a 68.5% contribution to the distribution of the species, particularly high elevation areas, included mean cold hardiness, water vapor pressure and temperature seasonality. The areas of greatest suitability with the highest occurrence of the species were identified geographically by the models. The present study provides useful information that can assist in the identification of areas where the species is most sensitive to different variables, including climate change and human activities and contributes in assessing the conservation status of Andean grassland at a regional scale.

Lithology- and climate-controlled soil aggregate-size distribution and organic carbon stability in the Peruvian Andes

Abstract. Recent studies indicate that climate change influences soil mineralogy by altering weathering processes and thus impacts soil aggregation and organic carbon (SOC) stability. Alpine ecosystems of the Neotropical Andes are characterized by high SOC stocks, which are important for sustaining ecosystem services. However, climate change in the form of altered precipitation patterns can potentially affect soil aggregation and SOC stability with potentially significant effects on the soil's ecosystem services. This study aimed to investigate the effects of precipitation and lithology on soil aggregation and SOC stability in the Peruvian Andean grasslands, and it assessed whether occlusion of organic matter (OM) in aggregates controls SOC stability. For this, samples were collected from soils on limestone and soils on acid igneous rocks from two sites with contrasting precipitation levels. We used a dry-sieving method to quantify aggregate-size distribution and applied a 76 d soil incubation with intact and crushed aggregates to investigate SOC stability's dependence on aggregation. SOC stocks ranged from 153±27 to 405±42 Mg ha−1, and the highest stocks were found in the limestone soils of the wet site. We found lithology rather than precipitation to be the key factor regulating soil aggregate-size distribution, as indicated by coarse aggregates in the limestone soils and fine aggregates in the acid igneous rock soils. SOC stability estimated by specific SOC mineralization rates decreased with precipitation in the limestone soils, but only minor differences were found between wet and dry sites in the acid igneous rock soils. Aggregate destruction had a limited effect on SOC mineralization, which indicates that occlusion of OM in aggregates played a minor role in OM stabilization. This was further supported by the inconsistent patterns of aggregate-size distribution compared to the patterns of SOC stability. We propose that OM adsorption on mineral surfaces is the main OM stabilization mechanism controlling SOC stocks and stability. The results highlight the interactions between precipitation and lithology on SOC stability, which are likely controlled by soil mineralogy in relation to OM input.

Lithology controlled soil organic carbon stabilization in an alpine grassland of the Peruvian Andes

Alpine grasslands of the Neotropical Andes have high soil organic carbon (SOC) stocks and provide crucial ecosystem services. However, stability of the SOC in these grasslands is not well-studied. Having insights into SOC stability contributes to a better understanding of ecosystem vulnerability and maintaining of ecosystem services. The objectives were to get a first insight into organic matter (OM) stabilization in soils from different bedrocks of Andean alpine grasslands near Cajamarca, Peru (7° 11″ S, 78° 35″ W) and how this controls SOC stocks. Samples were collected from soils formed on limestone and acid igneous rocks. Stabilization mechanisms of OM were investigated using selective extraction methods separating active Fe, Al and Ca fractions and determined SOC stocks. In both soil types, the results showed important contributions of complexation with and/or adsorption on Fe and Al (oxides) to OM stabilization. Exclusively in the limestone soils, Ca induced OM stabilization by promoting the formation of Ca2+ bridges between OM and mineral surfaces. Furthermore, no evidence showed that OM stabilization was controlled by crystalline Fe oxides, clay contents, allophones, Al toxicity or aggregate stability. Limestone soils had significantly higher SOC stocks (405 ± 42 Mg ha−1) compared to the acid igneous rock soils (226 ± 6 Mg ha−1), which is likely explained by OM stabilization related to Ca2+ bridges in addition to the stabilization related to Fe and Al (oxides) in the limestone soils. Our results suggest a shift from OM stabilization dominated by Fe and Al (oxides) to that with the presence of Ca-related cation bridges, with increasing pH values driven by lithology.

Composition and floral diversity in Andean grasslands in natural post-harvest restoration with Lepidium meyenii Walpers

The Andean grassland ecosystems undergo natural and anthropogenic degradation processes. The change of land use for agricultural use is the greatest threat, with a great loss of biodiversity followed by a very slow process of revegetalization. The objective was to assess the richness, abundance and diversity, alpha and beta, in areas of two-, three-, five-, six- and eight years of post-harvest abandonment of Lepidium meyenni Walpers. Ten affected areas were selected for agrostological evaluation, through four linear transects of interception points with 100 records each, applied on the edge and inside the affected area, inside and outside the affected areas, as well as samples composed of soils for each area. Composite soil samples were collected from each transect and study area to analyze their physical and chemical properties. The data were analyzed using the generalized linear mixed model with Rstudio v 5.3.2, and the multivariate analysis of canonical correspondence between biological and environmental variables, using CANOCO v.1.4 software. A significant difference was found between floral composition and alpha diversity according to abandonment time and seasonal periods. The greatest richness and abundance was observed between five- and eight years of abandonment, due to the greater presence of perennial species. Linear correspondence of alpha richness and diversity with organic matter, nitrogen, soil phosphorus and abandonment time was observed.

From pampa to puna: Biogeography and diversification of a group of Neotropical obligate grassland birds (Anthus:Motacillidae)

The evolution of Neotropical birds of open landscapes remains largely unstudied. We investigate the diversification and biogeography of a group of Neotropical obligate grassland birds (Anthus: Motacillidae). We use a multilocus phylogeny of 22 taxa of Anthus to test the hypothesis that these birds radiated contemporaneously with the development of grasslands in South America. We employ the R package DDD to analyze the dynamics of Anthus diversification across time in Neotropical grasslands, explicitly testing for shifts in dynamics associated with the Miocene development of grasslands, the putative Pleistocene expansion of arid lowland biomes, and Pleistocene sundering of Andean highland grasslands. A lineage-through-time plot revealed increases in the number of lineages, and DDD detected shifts to a higher clade-level carrying capacity during the late Miocene, indicating an early burst of diversification associated with grassland colonization. However, we could not corroborate the shift using power analysis, probably reflecting the small number of tips in our tree. We found evidence of a divergence at similar to 1 Mya between northern and southern Amazonian populations of Anthus lutescens, countering Haffer's idea of Pleistocene expansion of open biomes in the Amazon Basin. We used BioGeoBears to investigate ancestral areas and directionality of colonization of Neotropical grasslands. Members of the genus diversified into, out of, and within the Andes, within-Andean diversification being mostly Pleistocene in origin.

Bacterial communities associated to Chilean altiplanic native plants from the Andean grasslands soils

The rhizosphere is considered the primary place for soil microbiome differentiation and plays a key role in plant survival, especially for those subjected to environmental stress. Using high-throughput sequencing of the 16S rRNA gene, we analyzed and compared soil bacterial communities associated to four of the most abundant high altitude native plant species of the Chilean Andean grasslands. We examined three soil compartments: the rhizosphere (bacteria firmly attached to the roots), the rhizosphere-surrounding soil (bacteria loosely attached to the roots) and the bulk soil (plant-free soil). The rhizosphere microbiome was in all cases the least diverse, exposing that the bulk soil was a more complex environment. Taxonomic analysis revealed an abrupt change between the rhizosphere and the rest of the non-rhizospheric soils. Thus, while rhizobacterial communities were enriched in Proteobacteria (mainly Alphaproteobacteria), Actinobacteria (mostly Blastocatellia) dominated in bulk soils. Finally, we detected certain taxonomic rhizosphere signatures, which could be attributed to a particular genotype. Overall, our results indicate that the thin layer of soil surrounding the roots constitute a distinctive soil environment. This study contributes to expand the knowledge about soil bacterial communities in the Chilean highlands and takes the first step to understand the processes that might lead to the rhizosphere differentiation in that area.

Relationship between vegetation assemblages and modern pollen in semiarid environments of Jujuy, northwestern Argentina

Steppes and grasslands of semiarid environments in the Central Andes are poorly understood in fossil records due to the lack of modern pollen–vegetation studies. This paper is focused on a systematic analysis of the modern pollen–vegetation relationship in three protected areas of the Puna in Jujuy, northwestern Argentina. Vegetation assemblages were described on the basis of their floristic composition in 32 plots. Different multivariate methods, indices of association and simple linear regression models were applied in order to investigate the degree to which the different vegetation assemblages can be distinguished by their pollen spectra. Five plant assemblages were described, which were partially represented by their pollen spectra: mixed and shrubby steppes, scrubland, grassland with shrubs and high Andean grassland. The pollen spectra reflected both altitudinal variations and environmental conditions. In the Puna belt, at 3600 m, shrubs were present in great abundance and their highest pollen percentages were recorded. Both the abundance of Poaceae herbs and their pollen type increased with increasing altitude up to 4220 m, indicating the transition towards the Altoandean belt. Pollen assemblages evidenced the following environmental conditions: (1) the temperature decrease is characterized by taxa adapted to either cold air (<7 °C) or cold soil (Aridisols-Cryids/Inceptisols-Ustepts type) including Poaceae, Frankenia triandra, Apiaceae, Hoffmansegia glauca, Ephedra, Caryophyllaceae; (2) moisture availability in the soil can be discernible by assemblages indicating dry soil (Aridisols-Durids type), such as Baccharis, Fabiana densa, Maihuniopsis boliviana, Fabaceae, Tetraglochin cristatum and Nassauvia axilaris, from those indicating humid soil (Entisols-Psaments type), such as Parastrephia, Astragalus garbancillo and Gentianaceae. Four taxa exhibited both a good association and a significant correlation and could be considered indicative of the steppe (Baccharis and M. boliviana), scrublands (Parastrephia) and grassland (Poaceae) in a fossil record. This work has important implications for interpreting the dynamics of vegetation during the Holocene in the Puna.

Phylogeny and Evolution of the Neotropical Radiation of Lachemilla (Rosaceae): Uncovering a History of Reticulate Evolution and Implications for Infrageneric Classification

Reticulate evolution often leads to incongruence between nuclear and plastid phylogenies, and comparisons between them have been used as a first approximation to disentangle patterns of hybridization. Because other processes like incomplete lineage sorting and phylogenetic error also produce similar incongruence patterns, additional sources of evidence must be incorporated. Here we focus on reconstructing the phylogeny of Lachemilla (Rosaceae) using nuclear ribosomal ITS and plastid trnL-FDNAsequences, and explore widespread patterns of cytonuclear discordance in this group. Lachemilla is a highly morphologically variable group of perennial herbs and shrubs, and a nearly ubiquitous member of the diverse Neotropical high-altitude grasslands. Our analyses identified four major clades within Lachemilla that are in part congruent with previous morphological classifications of the group. Furthermore, using multiple sources of evidence, including a procrustean approach to cophylogeny estimation, coalescent-based simulations, phylogenetic networks, chromosome counts, and genome size estimations, we also revealed a large-scale pattern of incongruence between the plastid and nuclear phylogenies in Lachemilla, which is mainly the result of widespread hybridization and polyploidy.We also estimated that the origin of Lachemilla in South America (;14.5 MYA) predates the “rapid-uplift” diversification model that has been suggested for other high species-richness Andean plant clades, but following the formation of the high-elevation Andean grasslands during the last 5 MYA, a rapid accumulation of particular nested lineages has contributed to the ubiquitous presence of Lachemilla in these biomes.

Quantification of rainfall interception in the high Andean tussock grasslands

AbstractThe páramo ecosystem provides most of the water for the tropical Andean highlands in South America. Although the comprehension of this environment has increased lately, there remains an urgent need to quantify the processes involved in the hydrological cycle. Interception loss (IL) is one of the least studied processes in the páramo, and more generally, in grasslands globally. The main objective of this study was to quantify IL at event scale by estimating it indirectly from precipitation (P) and effective rainfall (ER). Furthermore, the following questions were assessed: (a) how much of the P becomes ER? (b) what is the impact on IL calculations of using a rain gauge instead of a disdrometer? (c) which meteorological variables are related to the IL process? and (d) is it possible to estimate IL from meteorological variables? High percentages of IL in relation to P were found (10–100%). The canopy storage capacity of tussock grasses was approximately 2 mm. The disdrometer observations led to more accurate results than the rain gauge observations because only the disdrometer registers light precipitation, horizontal precipitation, and drizzle that increases the amount of P, ER, and IL estimates. Also, we found that IL is more strongly correlated with P; and IL can be estimated with a multiple linear regression (R2 = 0.9) from P and relative humidity for events where 1.7 &lt; P &lt; 8.5 mm. These findings show the important role of IL in the páramo and provide a stepping stone to modelling of water resources.

Evolutionary radiation of earless frogs in the Andes: molecular phylogenetics and habitat shifts in high-elevation terrestrial breeding frogs.

The loss of hearing structures and loss of advertisement calls in many terrestrial breeding frogs (Strabomantidae) living at high elevations in South America are common and intriguing phenomena. The Andean frog genus Phrynopus Peters, 1873 has undergone an evolutionary radiation in which most species lack the tympanic membrane and tympanic annulus, yet the phylogenetic relationships among species in this group remain largely unknown. Here, we present an expanded molecular phylogeny of Phrynopus that includes 24 nominal species. Our phylogeny includes Phrynopus peruanus, the type species of the genus, and 10 other species for which genetic data were previously unavailable. We found strong support for monophyly of Phrynopus, and that two nominal species—Phrynopus curator and Phrynopus nicoleae—are junior synonyms of Phrynopus tribulosus. Using X-ray computed tomography (CT) imaging, we demonstrate that the absence of external hearing structures is associated with complete loss of the auditory skeletal elements (columella) in at least one member of the genus. We mapped the tympanum condition on to a species tree to infer whether the loss of hearing structures took place once or multiple times. We also assessed whether tympanum condition, body size, and body shape are associated with the elevational distribution and habitat use. We identified a single evolutionary transition that involved the loss of both the tympanic membrane and tympanic annulus, which in turn is correlated with the absence of advertisement calls. We also identified several species pairs where one species inhabits the Andean grassland and the other montane forest. When accounting for phylogenetic relatedness among species, we detected a significant pattern of increasing body size with increasing elevation. Additionally, species at higher elevations tend to develop shorter limbs, shorter head, and shorter snout than species living at lower elevations. Our findings strongly suggest a link between ecological divergence and morphological diversity of terrestrial breeding frogs living in montane gradients.

Espeletia giant rosette plants are reliable biological indicators of time since fire in Andean grasslands

Paramo grasslands in the tropical Andes are fire-prone ecosystems and an understanding of their fire ecology is fundamental to biodiversity conservation and ecosystem management. Fire registers are normally impractical in these remote, cloud-covered landscapes, but Espeletia giant rosette plants have been proposed as biological indicators of time since fire in paramos. Espeletia giant stem rosettes tolerate fire well, protecting apical buds in at the heart of their leaf rosettes, and for some species, germination is known to be enhanced by fire. As the plant grows, its dead leaves remain attached to the stem, but fire removes these and resets the “leaf clock”. This study uses a unique register of fires in one Ecuadorian paramo to assess the robustness of this biological indicator. Dead leaf cover on Espeletia pycnophylla giant rosette plants was measured in fifteen different sites with known fire dates from 2000 to 2014. The growth rates of plants at four different elevations were measured over a 2-year period and used to estimate time since fire based on dead leaf cover in the known sites. Estimates were accurate to ± 2 year. Thus, where fire records are missing, relatively easy measurements of growth rates and dead leaf cover of Espeletia giant rosette plants can provide reliable estimates across a wide range of times since fire. This approach has value for direct investigations into fire ecology but also for studies in which controlling for fire dynamics is necessary to reveal underlying patterns. Therefore, this approach also offers a means to obtain better information on other landscape-scale processes such as the impact of climate change on biodiversity or the provision of ecosystem services.

A new species of Phrynopus (Amphibia, Anura, Craugastoridae) from upper montane forests and high Andean grasslands of the Pui Pui Protected Forest in central Peru.

We describe a new species of Phrynopus from the upper montane forests and high Andean grasslands (puna) of the Pui Pui Protected Forest and its close surroundings (Región Junín, central Peru) and compare it morphologically and genetically with other species of Phrynopus. Phrynopus inti sp. n. is known from four localities outside and two localities inside the Pui Pui Protected Forest between 3350 and 3890 m a.s.l. Studied specimens of the new species are characterized by a snout-vent length of 27.2–35.2 mm in males (n = 6), and 40.4 mm in a single female, by having the skin on dorsum and flanks smooth with scattered tubercles, venter smooth, by lacking a tympanum, and males without vocal slits and nuptial pads. In life, the dorsum is pale grayish brown with or without dark brown blotches, or dorsum blackish brown with small yellow flecks, throat, chest and venter are pale grayish brown with salmon mottling, groin is pale grayish brown with salmon colored flecks, and the iris is golden orange with fine dark brown reticulations. The new species is morphologically most similar to Phrynopus kauneorum and P. juninensis. For the latter we describe the coloration in life for a specimen obtained at the type locality. A molecular phylogenetic analysis based on mitochondrial and nuclear DNA sequences inferred that the new species is most closely related to Phrynopus kauneorum, P. miroslawae, P. tautzorum, and an undescribed species distributed at high elevation in Región Pasco, central Peru.