Historical Assembly of Andean Tree Communities.

Patterns of species diversity have been associated with changes in climate across latitude and elevation. However, the ecological and evolutionary mechanisms underlying these relationships are still actively debated. Here, we present a complementary view of the well-known tropical niche conservatism (TNC) hypothesis, termed the multiple zones of origin (MZO) hypothesis, to explore mechanisms underlying latitudinal and elevational gradients of phylogenetic diversity in tree communities. The TNC hypothesis posits that most lineages originate in warmer, wetter, and less seasonal environments in the tropics and rarely colonize colder, drier, and more seasonal environments outside of the tropical lowlands, leading to higher phylogenetic diversity at lower latitudes and elevations. In contrast, the MZO hypothesis posits that lineages also originate in temperate environments and readily colonize similar environments in the tropical highlands, leading to lower phylogenetic diversity at lower latitudes and elevations. We tested these phylogenetic predictions using a combination of computer simulations and empirical analyses of tree communities in 245 forest plots located in six countries across the tropical and subtropical Andes. We estimated the phylogenetic diversity for each plot and regressed it against elevation and latitude. Our simulated and empirical results provide strong support for the MZO hypothesis. Phylogenetic diversity among co-occurring tree species increased with both latitude and elevation, suggesting an important influence on the historical dispersal of lineages with temperate origins into the tropical highlands. The mixing of different floras was likely favored by the formation of climatically suitable corridors for plant migration due to the Andean uplift. Accounting for the evolutionary history of plant communities helps to advance our knowledge of the drivers of tree community assembly along complex climatic gradients, and thus their likely responses to modern anthropogenic climate change.

Strategies of diaspore dispersal investment in Compositae: the case of the Andean highlands.

BACKGROUND AND AIMS: Understanding diaspore morphology and how much a species invests on dispersal appendages is key for improving our knowledge of dispersal in fragmented habitats. We investigate diaspore morphological traits in high-Andean Compositae and their main abiotic and biotic drivers and test whether they play a role in species distribution patterns across the naturally fragmented high-Andean grasslands. METHODS: We collected diaspore trait data for 125 Compositae species across 47 tropical high-Andean summits, focusing on achene length and pappus-to-achene length ratio, with the latter as a proxy of dispersal investment. We analysed the role of abiotic (temperature, elevation and latitude) and biotic factors (phylogenetic signal and differences between tribes) on diaspore traits and whether they are related to distribution patterns across the Andes, using phylogenomics, distribution modelling and community ecology analyses. KEY RESULTS: Seventy-five percent of the studied species show small achenes (length <3.3 mm) and 67% have high dispersal investment (pappus length at least two times the achene length). Dispersal investment increases with elevation, possibly to compensate for lower air density, and achene length increases towards the equator, where non-seasonal climate prevails. Diaspore traits show significant phylogenetic signal, and higher dispersal investment is observed in Gnaphalieae, Astereae and Senecioneae, which together represent 72% of our species. High-Andean-restricted species found across the tropical Andes have, on average, the pappus four times longer than the achene, a significantly higher dispersal investment than species present only in the northern Andes or only in the central Andes. CONCLUSIONS: Small achenes and high diaspore dispersal investment dominate among high-Andean Compositae, traits typical of mostly three tribes of African origin; but traits are also correlated with the environmental gradients within the high-Andean grasslands. Our results also suggest that diaspore dispersal investment is likely to shape species distribution patterns in naturally fragmented habitats.

Understanding climate change impacts on biome and plant distributions in the Andes: Challenges and opportunities.

Aim: Climate change is expected to impact mountain biodiversity by shifting species ranges and the biomes they shape. The extent and regional variation in these impacts are still poorly understood, particularly in the highly biodiverse Andes. Regional syntheses of climate change impacts on vegetation are pivotal to identify and guide research priorities. Here we review current data, knowledge and uncertainties in past, present and future climate change impacts on vegetation in the Andes. Location: Andes. Taxon: Plants. Methods: We (i) conducted a literature review on Andean vegetation responses to past and contemporary climatic change, (ii) analysed future climate projections for different elevations and slope orientations at 19 Andean locations using an ensemble of model outputs from the Coupled Model Intercomparison Project 5, and (iii) calculated changes in the suitable climate envelope area of Andean biomes and compared these results to studies that used species distribution models. Results: Future climatic changes (2040-2070) are projected to be stronger at high-elevation areas in the tropical Andes (up to 4°C under RCP 8.5), while in the temperate Andes temperature increases are projected to be up to 2°C. Under this worst-case scenario, temperate deciduous forests and the grasslands/steppes from the Central and Southern Andes are predicted to show the greatest losses of suitable climatic space (30% and 17%-23%, respectively). The high vulnerability of these biomes contrasts with the low attention from researchers modelling Andean species distributions. Critical knowledge gaps include a lack of an Andean wide plant checklist, insufficient density of weather stations at high-elevation areas, a lack of high-resolution climatologies that accommodates the Andes' complex topography and climatic processes, insufficient data to model demographic and ecological processes, and low use of palaeo data for distribution modelling. Main conclusions: Climate change is likely to profoundly affect the extent and composition of Andean biomes. Temperate Andean biomes in particular are susceptible to substantial area contractions. There are, however, considerable challenges and uncertainties in modelling species and biome responses and a pressing need for a region-wide approach to address knowledge gaps and improve understanding and monitoring of climate change impacts in these globally important biomes.

Hydroclimate and vegetation variability of high Andean ecosystems.

Mountain ecosystems are sensitive to climate fluctuations; however, the scarcity of instrumental data makes necessary the use of complementary information to study the effect of climate change on these systems. Remote sensing permits studying the dynamics of vegetation productivity and wetlands in response to climate variability at different scales. In this study we identified the main climate variables that control vegetation dynamics and water balance in Cumbres Calchaquíes, NW Argentina. For this, we built annual time series from 1986 to 2019 of Soil Adjusted Vegetation Index (SAVI, to quantify spare vegetation productivity), lake area, and snow-ice cover of peatlands, as indicators of mountain productivity and hydrology. We used a decompose function to explore trend, seasonality and random signal of the three-time series, and explored for significant changes in the mean value of consecutive periods. We used correlational analysis to explore their associations with climate records at local, regional, and global scales. The results showed that, SAVI and hydrological indicators presented different fluctuation patterns more pronounced since 2012, when they showed divergent trends with increasing SAVI and decreasing lake area and snow-ice cover. The three indicators responded differently to climate; SAVI increased in warmer years and lake area reflected the water balance of previous years. Snow-ice cover of peatlands was highly correlated with lake area. La Niña had a positive effect on lake area and snow-ice cover and a negative on SAVI, while El Niño had a negative effect on SAVI. Fluctuations of lake areas were synchronized with lake area in the nearby Argentinian puna, suggesting that climate signals have regional extent. The information provided by the three hydroclimate indicators is complementary and reflects different climate components and processes; biological processes (SAVI), physical processes (snow ice cover) and their combination (lake area). This study provides a systematic accessible replicable tool for mountain eco-hydrology long-term monitoring.

Mature Andean forests as globally important carbon sinks and future carbon refuges.

It is largely unknown how South America's Andean forests affect the global carbon cycle, and thus regulate climate change. Here, we measure aboveground carbon dynamics over the past two decades in 119 monitoring plots spanning a range of >3000 m elevation across the subtropical and tropical Andes. Our results show that Andean forests act as strong sinks for aboveground carbon (0.67 ± 0.08 Mg C ha-1 y-1) and have a high potential to serve as future carbon refuges. Aboveground carbon dynamics of Andean forests are driven by abiotic and biotic factors, such as climate and size-dependent mortality of trees. The increasing aboveground carbon stocks offset the estimated C emissions due to deforestation between 2003 and 2014, resulting in a net total uptake of 0.027 Pg C y-1. Reducing deforestation will increase Andean aboveground carbon stocks, facilitate upward species migrations, and allow for recovery of biomass losses due to climate change.

Elevation and latitude drives structure and tree species composition in Andean forests: Results from a large-scale plot network.

Our knowledge about the structure and function of Andean forests at regional scales remains limited. Current initiatives to study forests over continental or global scales still have important geographical gaps, particularly in regions such as the tropical and subtropical Andes. In this study, we assessed patterns of structure and tree species diversity along ~ 4000 km of latitude and ~ 4000 m of elevation range in Andean forests. We used the Andean Forest Network (Red de Bosques Andinos, https://redbosques.condesan.org/) database which, at present, includes 491 forest plots (totaling 156.3 ha, ranging from 0.01 to 6 ha) representing a total of 86,964 identified tree stems ≥ 10 cm diameter at breast height belonging to 2341 identified species, 584 genera and 133 botanical families. Tree stem density and basal area increases with elevation while species richness decreases. Stem density and species richness both decrease with latitude. Subtropical forests have distinct tree species composition compared to those in the tropical region. In addition, floristic similarity of subtropical plots is between 13 to 16% while similarity between tropical forest plots is between 3% to 9%. Overall, plots ~ 0.5-ha or larger may be preferred for describing patterns at regional scales in order to avoid plot size effects. We highlight the need to promote collaboration and capacity building among researchers in the Andean region (i.e., South-South cooperation) in order to generate and synthesize information at regional scale.

Author Correction: Widespread but heterogeneous responses of Andean forests to climate change.

In Fig. 2 of this Article, the positive part of the y axis scale should read 0, 0.02, 0.04 instead of 0, 0.04, 0.02. This has been corrected online.

Widespread but heterogeneous responses of Andean forests to climate change.

Global warming is forcing many species to shift their distributions upward, causing consequent changes in the compositions of species that occur at specific locations. This prediction remains largely untested for tropical trees. Here we show, using a database of nearly 200 Andean forest plot inventories spread across more than 33.5° latitude (from 26.8° S to 7.1° N) and 3,000-m elevation (from 360 to 3,360 m above sea level), that tropical and subtropical tree communities are experiencing directional shifts in composition towards having greater relative abundances of species from lower, warmer elevations. Although this phenomenon of 'thermophilization' is widespread throughout the Andes, the rates of compositional change are not uniform across elevations. The observed heterogeneity in thermophilization rates is probably because of different warming rates and/or the presence of specialized tree communities at ecotones (that is, at the transitions between distinct habitats, such as at the timberline or at the base of the cloud forest). Understanding the factors that determine the directions and rates of compositional changes will enable us to better predict, and potentially mitigate, the effects of climate change on tropical forests.

Vegetation trends over eleven years on mountain summits in NW Argentina.

As global climate change leads to warmer and dryer conditions in the central Andes, alpine plant communities are forced to upward displacements following their climatic niche. Species range shifts are predicted to have major impacts on alpine communities by reshuffling species composition and abundances. Using a standardized protocol, we surveyed alpine plant communities in permanent plots on four high Andean summits in NW Argentina, which range from 4,040 to 4,740 m a.s.l. After a baseline survey in 2006-2008, we resurvey the same plots in 2012, and again in 2017. We found a significant decrease in plant cover, species richness, and diversity across the elevation gradient in the three censuses and a strong decrease in soil temperature along the elevation gradient. We found a high plant community turnover (37%-49%) among censuses, differentiating according to summits and aspects; major changes of community turnover were observed in the lowest summit (49%) and on the northern (47%) and western (46%) aspects. Temporal patterns in community changes were represented by increases in plant cover in the highest summit, in species richness in the lower summit, and in diversity (Shannon index) in the four summits, over time, together with increase in small herbs and non-tussock grasses. We suggest that the observed trend in plant community dynamics responds to short-term temperature and precipitation variability, which is influenced by El Niño Southern Oscillation (ENSO), and due to time lags in plant community response, it may take much longer than one decade for the observed trends to become stables and statistically significant. Our study provides an important foundation for documenting more profound changes in these subtropical alpine plant communities as global climate change continues.

Patrones espaciales en comunidades de macroinvertebrados acuáticos de la Puna Argentina / Spatial patterns in communities of aquatic macroinvertebrates of Argentinean Puna

ResumenLos macroinvertebrados son un componente vital en los ecosistemas acuáticos, procesan materia orgánica y sirven de alimento para otros organismos como los peces o anfibios. En los sistemas altoandinos, y más específicamente en las vegas o bofedales de Argentina, el conocimiento de la diversidad acuática es escaso o nulo, lo que ha dificultado la interpretación de patrones espaciales a escalas regionales. Típicamente estos ecosistemas se sitúan entre los 3 200 y 5 200 m.s.n.m. en zonas con precipitaciones por debajo de los 300 mm anuales. En este trabajo se realizó la primera descripción cuantitativa de los patrones espaciales de diversidad y composición de macroinvertebrados acuáticos en siete vegas de la Puna Argentina, y su relación con las características del ambiente, de la vegetación circundante y las características físico-químicas del agua. Se recolectaron 3 131 individuos de macroinvertebrados acuáticos, los cuales fueron identificados en 25 taxa que pertenecen a 22 familias y 24 géneros. Entre las plantas vasculares se registraron un total de 62 especies, correspondiente a 20 familias, las graminoides cespitosas fueron la forma de vida más abundante seguida por plantas en cojín o tapete. A partir de un análisis estadístico de ordenamiento NMDS (Non-Metrical Multidimensional Scaling) se segregaron los sitios de muestreo de cada vega. Las vegas ubicadas más al oeste y al sur y con mayor temperatura del agua se agruparon sobre el eje 1, mientras que las vegas con mayor conductividad, sólidos totales disueltos, y concentración de sales en el agua lo hicieron sobre el eje 2. La temperatura del agua fue siempre mayor que la del aire, y no mostró un patrón claro de disminución con la altitud. La altitud no se correlacionó con la abundancia de los macroinvertebrados ni con la riqueza de las plantas. Las formas de vida de plantas como gramíneas dispersas, hierbas rastreras y postradas (que en muchos casos ingresan al cauce de agua) y plantas acuáticas fueron más abundantes en las vegas 4, 5 y 7 y estuvieron asociadas con ciertos taxa de macroinvertebrados que pertenecen a distintos grupos funcionales tróficos como colectores- filtradores, colectores- recolectores y raspadores. Finalmente, las variables de conductividad, sólidos totales disueltos y concentración de sales fueron más elevadas en la vega 6, asociadas a Bivalvia (colector-filtrador) y Hyalella (colector-recolector). A partir de estos resultados se evidencia que los macroinvertebrados presentaron patrones espaciales de diversidad y composición, siendo en primer lugar más importante su relación con la vegetación circundante y en segundo término con las características físico-químicas del agua en siete vegas de la Puna argentina.

AbstractThe macroinvertebrates are a vital component of freshwater ecosystems as they contribute to the process of organic matter while serving as food for other organisms such as fish and amphibians. Unfortunately, the knowledge of the aquatic diversity is poor in the high Andean systems (between 3 200 and 5 200 m.a.s.l. and rainfall below 300 mm per year), especially in the Argentinean peatbogs, a fact which has made difficult the interpretation of spatial patterns on a regional scale. The present study aimed to describe the composition of aquatic macroinvertebrates in seven peatbogs in the Argentinean Puna, and to analyze for the first time their spatial patterns. For this, we studied the relationship of these organisms with the environment, and obtained information about the surrounding vegetation and water physico-chemical characteristics. A total of 3 131 specimens of aquatic macroinvertebrates were collected, representing 25 taxa belonging to 22 families and 24 genera. In addition, 62 species of vascular plants were recorded, belonging to 20 families. The most abundant life form were the tufted grasses, followed by cushions. By using an NMDS (Non-Metrical Multidimensional Scaling) statistical analysis, the sampling sites were ordered in each peatbog as follows. The peatbogs located furthest West and South with higher water temperature were grouped on axis 1, whereas those with higher conductivity, whole water dissolved solids and salt concentration were grouped on axis 2. The water temperature was higher than air temperature at all times, and we found no association between temperature and altitude. The altitude had no correlation with the abundance of macroinvertebrates or with plant richness. Life forms such as scattered graminoids, trailing and prostrate herbs (in many cases they get into the channel) and aquatic plants were more abundant in peatbogs 4, 5 and 7 and they had a correlation with some macroinvertebrates belonging to functional trophic groups such as filter-collectors, collectors and scrapers. Finally, conductivity, whole dissolved solids and salt concentration had their highest value in peatbog 6, where Bivalvia (filter-collector) and Hyalella (collector) were also found. The results obtained attest that these macroinvertebrates displayed diversity and composition spatial patterns, the most important feature being their relationship with the surrounding vegetation, and to a lesser extent, with the physical and chemical traits of water in seven fertile lowlands in the Argentinean Puna. Rev. Biol. Trop. 64 (2): 747-762. Epub 2016 June 01.

[Spatial patterns in communities of aquatic macroinvertebrates of Argentinean Puna]. / Patrones espaciales en comunidades de macroinvertebrados acuáticos de la Puna Argentina.

Spatial patterns in communities of aquatic macroinvertebrates of Argentinean Puna. The macroinvertebrates are a vital component of freshwater ecosystems as they contribute to the process of organic matter while serving as food for other organisms such as fish and amphibians. Unfortunately, the knowledge of the aquatic diversity is poor in the high Andean systems (between 3 200 and 5 200 m.a.s.l. and rainfall below 300 mm per year), especially in the Argentinean peatbogs, a fact which has made difficult the interpretation of spatial patterns on a regional scale. The present study aimed to describe the composition of aquatic macroinvertebrates in seven peatbogs in the Argentinean Puna, and to analyze for the first time their spatial patterns. For this, we studied the relationship of these organisms with the environment, and obtained information about the surrounding vegetation and water physico-chemical characteristics. A total of 3 131 specimens of aquatic macroinvertebrates were collected, representing 25 taxa belonging to 22 families and 24 genera. In addition, 62 species of vascular plants were recorded, belonging to 20 families. The most abundant life form were the tufted grasses, followed by cushions. By using an NMDS (Non-Metrical Multidimensional Scaling) statistical analysis, the sampling sites were ordered in each peatbog as follows. The peatbogs located furthest West and South with higher water temperature were grouped on axis 1, whereas those with higher conductivity, whole water dissolved solids and salt concentration were grouped on axis 2. The water temperature was higher than air temperature at all times, and we found no association between temperature and altitude. The altitude had no correlation with the abundance of macroinvertebrates or with plant richness. Life forms such as scattered graminoids, trailing and prostrate herbs (in many cases they get into the channel) and aquatic plants were more abundant in peatbogs 4, 5 and 7 and they had a correlation with some macroinvertebrates belonging to functional trophic groups such as filter-collectors, collectors and scrapers. Finally, conductivity, whole dissolved solids and salt concentration had their highest value in peatbog 6, where Bivalvia (filter-collector) and Hyalella (collector) were also found. The results obtained attest that these macroinvertebrates displayed diversity and composition spatial patterns, the most important feature being their relationship with the surrounding vegetation, and to a lesser extent, with the physical and chemical traits of water in seven fertile lowlands in the Argentinean Puna.

Large-scale patterns of turnover and Basal area change in Andean forests.

General patterns of forest dynamics and productivity in the Andes Mountains are poorly characterized. Here we present the first large-scale study of Andean forest dynamics using a set of 63 permanent forest plots assembled over the past two decades. In the North-Central Andes tree turnover (mortality and recruitment) and tree growth declined with increasing elevation and decreasing temperature. In addition, basal area increased in Lower Montane Moist Forests but did not change in Higher Montane Humid Forests. However, at higher elevations the lack of net basal area change and excess of mortality over recruitment suggests negative environmental impacts. In North-Western Argentina, forest dynamics appear to be influenced by land use history in addition to environmental variation. Taken together, our results indicate that combinations of abiotic and biotic factors that vary across elevation gradients are important determinants of tree turnover and productivity in the Andes. More extensive and longer-term monitoring and analyses of forest dynamics in permanent plots will be necessary to understand how demographic processes and woody biomass are responding to changing environmental conditions along elevation gradients through this century.