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The Andes mountain range in South America has a high level of endemism and is a major source of ecosystem services. The Andes is increasingly threatened by anthropogenic disturbances that have allowed the establishment of non-native plants, mainly in the lower elevation areas. However, synergies between climate change and anthropogenic pressure are promoting the spread of non-native plants to higher elevation areas. In this article, we evaluate and identify the main non-native plants invading Andean ecosystems, and assess their taxonomic families, growth forms and distribution patterns. Based on a systematic literature review, we identified the importance of climatic and anthropogenic factors as drivers of non-native species establishment in Andean ecosystems and the main impacts of non-native plants in the Andes. We then identified research gaps across each biogeographic region in the Andes. Finally, we highlight key elements to better tackle the problem of non-native plant invasions in Andean ecosystems, including the need for a systematic monitoring of invasion patterns and spread (e.g. MIREN protocol) and a common policy agenda across international borders for the prevention and management of non-native plants in this highly vulnerable region. [ABSTRACT FROM AUTHOR]

Background: Soil microorganisms are in constant interaction with plants, and these interactions shape the composition of soil bacterial communities by modifying their environment. However, little is known about the relationship between microorganisms and native plants present in extreme environments that are not affected by human intervention. Using high-throughput sequencing in combination with random forest and co-occurrence network analyses, we compared soil bacterial communities inhabiting the rhizosphere surrounding soil (RSS) and the corresponding bulk soil (BS) of 21 native plant species organized into three vegetation belts along the altitudinal gradient (2400–4500 m a.s.l.) of the Talabre–Lejía transect (TLT) in the slopes of the Andes in the Atacama Desert. We assessed how each plant community influenced the taxa, potential functions, and ecological interactions of the soil bacterial communities in this extreme natural ecosystem. We tested the ability of the stress gradient hypothesis, which predicts that positive species interactions become increasingly important as stressful conditions increase, to explain the interactions among members of TLT soil microbial communities. Results: Our comparison of RSS and BS compartments along the TLT provided evidence of plant-specific microbial community composition in the RSS and showed that bacterial communities modify their ecological interactions, in particular, their positive:negative connection ratios in the presence of plant roots at each vegetation belt. We also identified the taxa driving the transition of the BS to the RSS, which appear to be indicators of key host-microbial relationships in the rhizosphere of plants in response to different abiotic conditions. Finally, the potential functions of the bacterial communities also diverge between the BS and the RSS compartments, particularly in the extreme and harshest belts of the TLT. Conclusions: In this study, we identified taxa of bacterial communities that establish species-specific relationships with native plants and showed that over a gradient of changing abiotic conditions, these relationships may also be plant community specific. These findings also reveal that the interactions among members of the soil microbial communities do not support the stress gradient hypothesis. However, through the RSS compartment, each plant community appears to moderate the abiotic stress gradient and increase the efficiency of the soil microbial community, suggesting that positive interactions may be context dependent. [ABSTRACT FROM AUTHOR]

Cushion plants are specialized keystone species of alpine environments that can have a positive effect on ecosystem structure and function. However, we know relatively little about how cushion plants regulate the diversity and composition of soil microbial communities, major drivers of soil processes and ecosystem functioning. Identifying what factors drive the diversity and composition of soil microbial communities in high-elevation ecosystems is also fundamental to predict how global changes will affect their conservation and the services and functions they provide. Thus, we sampled four sites along the southern Andes following the vegetation belt of Azorella cushion species. The field sites spread along a latitudinal gradient and had contrasting levels of aridity, UV-B radiation, mean temperature and soil properties. Overall, Azorella, as well as aridity and UV-B radiation, were the major drivers of the distribution, composition and diversity of soil microbial communities in the studied ecosystems of the Chilean Andes. UV-B radiation affected particularly soil fungi, while soil properties such as pH, total C and N content, essential predictors of microbial diversity globally, had a much lower effect on the composition of soil microbial communities. Understanding the factors driving the structure and composition of microbial communities, particularly the role of cushion plants and the feedbacks between plant, climate and soil is of uttermost importance for the preservation of the functionality of high-elevation ecosystems threatened by climate change. [ABSTRACT FROM AUTHOR]

It is unclear whether the frequently observed increase in non-structural carbohydrates (NSC) in plants exposed to low temperatures or drought reflects a higher sensitivity of growth than photosynthesis in such conditions (i.e. sink limitation), or a prioritization of carbon (C) allocation to storage. Alpine areas in Mediterranean-type climate regions are characterized by precipitation increases and temperature decreases with elevation. Thus, alpine plants with wide elevational ranges in Mediterranean regions may be good models to examine these alternative hypotheses. We evaluated storage and growth during experimental darkness and re-illumination in individuals of the alpine plant Phacelia secunda from three elevations in the Andes of central Chile. We hypothesized that storage is prioritized regarding growth in plants of both low- and high elevations where drought and cold stress are greatest, respectively. We expected that decreases in NSC concentrations during darkness should be minimal and, more importantly, increases in NSC after re-illumination should be higher than increases in biomass. We found that darkness caused a significant decrease in NSC concentrations of both low- and high-elevation plants, but the magnitude of the decrease was lower in the latter. Re-illumination caused higher increase in NSC concentration than in biomass in both low- and high-elevation plants (1.5- and 1.9-fold, respectively). Our study shows that C allocation in Phacelia secunda reflects ecotypic differences among elevation provenances and suggests that low temperature, but not drought, favours C allocation to storage over growth after severe C limitation. [ABSTRACT FROM AUTHOR]

Freezing temperatures and summer droughts shape plant life in Mediterranean high-elevation habitats. Thus, the impacts of climate change on plant survival for these species could be quite different to those from mesic mountains. We exposed 12 alpine species to experimental irrigation and warming in the Central Chilean Andes to assess whether irrigation decreases freezing resistance, irrigation influences freezing resistance when plants are exposed to warming, and to assess the relative importance of irrigation and temperature in controlling plant freezing resistance. Freezing resistance was determined as the freezing temperature that produced 50 % photoinactivation [lethal temperature (LT)] and the freezing point (FP). In seven out of 12 high-Andean species, LT of drought-exposed plants was on average 3.5 K lower than that of irrigated plants. In contrast, most species did not show differences in FP. Warming changed the effect of irrigation on LT. Depending on species, warming was found to have (1) no effect, (2) to increase, or (3) to decrease the irrigation effect on LT. However, the effect size of irrigation on LT was greater than that of warming for almost all species. The effect of irrigation on FP was slightly changed by warming and was sometimes in disagreement with LT responses. Our data show that drought increases the freezing resistance of high-Andean plant species as a general plant response. Although freezing resistance increases depended on species-specific traits, our results show that warmer and moister growing seasons due to climate change will seriously threaten plant survival and persistence of these and other alpine species in dry mountains. [ABSTRACT FROM AUTHOR]

Abstract: The level and mechanism of resistance to damage from freezing temperatures is related to the ambient temperatures experienced by plants. High-elevation habitats exhibit a thermal gradient in the air–soil profile, exposing ground-level species to more rigorous thermal microhabitats than well aboveground species. Such gradients could explain an inverse relationship between freezing temperature damage (LT50) and plant height, such that leaves of smaller plants tolerate freezing conditions, while leaves of taller plants avoid them. However, microclimatic data have not been included in previous studies evaluating those patterns. On the other hand, a proposed trade-off between plant functional traits and freezing resistance suggest that leaves of growth-forms with higher intensity of growth are less freezing resistant than those with lower growth intensity. In this work, we determined whether the ability to resist freezing temperatures is related to plant height or growth-form in 37 high-elevation species from the Central Chilean Andes. We proposed that (1) plants with contrasting heights are exposed to different thermal conditions during freezing events, (2) the level and the mechanism of freezing resistance are inversely related to plant height, and that (3) the level of freezing resistance varied between plants with low- and high-growth intensities. We found that high-Andean species of different heights are exposed to different thermal conditions depending on the distance from the ground. However, neither level nor mechanisms of freezing resistance were related to plant height. Leaves of both short and tall plants showed similar LT50 and their main freezing resistance mechanism was tolerance. Moreover, leaves of growth-forms with high- and low-growth intensity resisted similar freezing temperatures. Our results are discussed in relation to environmental conditions that characterize the Central Chilean Andes. [Copyright &y& Elsevier]

Alpine habitats have been proposed as particularly sensitive to climate change. Shorter snow cover could expose high-elevation plants to very low temperatures, increasing their risk of suffering damage by freezing, hence decreasing their population viability. In addition, a longer and warmer growing season could affect the hardening process on these species. Thus, understanding the ability of these species to withstand freezing events under warmer conditions is essential for predicting how alpine species may respond to future climate changes. Here we assessed the freezing resistance of 11 species from the central Chilean Andes by determining their low temperature damage (LT50) and freezing point (FP) after experimental warming in the field. Plants were exposed during two growing seasons to a passive increase in the air temperature using open top chambers (OTCs). OTCs increased by ca. 3 K the mean air and soil daytime temperatures, but had smaller effects on freezing temperatures. Leaf temperature of the different species was on average 5.5 K warmer inside OTCs at midday. While LT50 of control plants ranged from −9.9 to −22.4, that of warmed plants ranged from −7.4 to −17.3°C. Overall, high-Andean species growing inside OTCs increased their LT50 ca. 4 K, indicating that warming decreased their ability to survive severe freezing events. Moreover, plants inside OTCs increased the FP ca. 2 K in some studied species, indicating that warming altered processes of ice crystal formation. Resistance of very low temperatures is a key feature of high-elevation species; our results suggest that current climate warming trends will seriously threaten the survival of high-elevation plants by decreasing their ability to withstand severe freezing events. [ABSTRACT FROM AUTHOR]

1. Although the consequences of facilitation at individual and population levels are well known, the community-level consequences of these processes have received much less attention. In particular, the importance of facilitation in determining richness at the entire community level has seldom been evaluated. 2. In this study, we sampled 11 alpine plant communities along the southern Andes in South America, spanning from tropical (25°S) to sub-antarctic latitudes (55°S). Plant communities were dominated by cushion plants, a particular growth form that acts as a nurse plant for other plant species. Through rarefaction curves, we assessed the effectiveness of community sampling and estimated the number of species present within and outside cushions. Non-metric Multidimensional Scaling ordinations (NMDS) were used to assess differences between the species assemblages growing within and outside cushions. Finally, samples from cushions and open areas were combined in a single matrix accounting for the difference in cover between both microhabitats, and through rarefaction curves we assessed how many more species are added to the community due to the presence of cushions. 3. Samples taken within cushions always contained more species than equivalent samples from open areas. However, the magnitude of this difference varied among communities. NMDS ordination indicated that cushions generate species assemblages structurally different from those found in open areas. Inclusion of samples from cushion and open areas in synthetic analyses – where differences in cover were accounted for – indicated that the presence of cushions consistently increased species richness at the entire community level. The magnitude of these increases in species richness varied with habitat severity, with lower values at both extremes of the environmental severity gradient. 4. Synthesis. Facilitative interactions with cushion nurse plants along the high Andes of southern South America changed plant assemblage structure and increased species richness at the entire community level, indicating that facilitative interactions are pivotal in maintaining the diversity of these harsh environments. [ABSTRACT FROM AUTHOR]

• Predicted increases in the length of the growing season as a result of climate change may more frequently expose high-elevation plants to severe frosts. Understanding the ability of these species to resist frosts during the growing season is essential for predicting how species may respond to changes in temperature regimes. • Here, we assessed the freezing resistance of 24 species from the central Chilean Andes by determining their low temperature damage (LT50), ice nucleation temperature (NT), freezing point (FP) and freezing resistance mechanism (i.e. avoidance or tolerance). • The Andean species were found to resist frosts from −8.2 to −19.5°C during the growing season, and freezing tolerance was the most common resistance mechanism. Freezing resistance (LT50) varied within the growing season, decreasing towards the end of this period in most of the studied species. However, the FP showed the opposite trend. LT50 increased with elevation, whilst FP was lower in plants from lower elevations, especially late in the growing season. • Andean species have the potential to withstand severe freezing conditions during the growing season, and the aridity of this high-elevation environment seems to play an important role in determining this high freezing resistance. [ABSTRACT FROM AUTHOR]

Few studies have compared the response of native and invasive populations under stressful conditions. Furthermore, there is little consensus as to whether a plastic response is related to invasiveness in stressful environments. Exotic species have recently been reported in the high Andes of central Chile, where individuals have to cope with drought and poor soils, in addition to extreme temperatures. We explored if the exotic species Taraxacum officinale (dandelion) has plastic responses to soil moisture and nutrient availability, and whether two sets of alpine populations derived from native and introduced populations can converge to similar plastic responses to environmental constraints. Using a common garden approach, we compared plants grown from seeds collected in alpine populations of its native range (Alps, France) and in alpine populations of its introduced range (Andes, Chile) under a drought experiment, a potassium gradient, and a nitrogen gradient. Plasticity was only found as a response to drought. Moreover, different responses were found between both origins. Andean individuals are drought-resistant, while individuals from the Alps were drought-sensitive. According to the nutrient experiments, Andean dandelions behave as a nitrogen demanding-potassium avoiding species, whereas individuals from the Alps did not show any particular dependency or repulsion tendency to either of these two nutrients. Results suggest that differences in life history traits of both derived sets of populations may have an important role in determining the response of dandelions under the evaluated conditions. However, the relative importance of genetic adaptation in these responses is still unclear. Although T. officinale is a cosmopolite weed, this is the first study that compares individuals coming from its native and invaded range under stressful conditions. [ABSTRACT FROM AUTHOR]

The last few centuries have seen an increase in the mean air temperature of the planet, a phenomenon that is called “global warming”. One of the most sensitive habitats to the effects of global warming is the high-elevation mountain environments, because these habitats are characterized by low temperature. Cushion plants are one of the best-adapted growth forms in this habitat, generating more suitable sites for other plants and insects. In the present study, we experimentally evaluated the effects of global warming by open-top chambers on the abundance and interaction of two ladybirds at 3,600 m, growing over cushions of the Azorella monantha species in the Andes of central Chile. Additionally, we measured variation in temperature, water content, and food availability by the presence of open-top chambers as possible mechanisms of spatial segregation between ladybirds. Without open-top chambers, the abundance of native and alien beetles was similar; but with open-top chambers, the native beetle species is spatially segregated by alien species, decreasing in abundance. The open-top chambers increase temperature and food availability, but not water content. We suggest that under the global warming scenario, the native insects will decrease in abundance or become extinct by the presence of alien insects, at least in the high-elevation mountain environments. [ABSTRACT FROM AUTHOR]

We developed multiple a priori hypotheses to link the observed spatial patterns with colonisation processes in the high alpine cushion plant, Azorella madreporica. We conducted this study in the Molina River basin (33°20′ S, 70°16′ W, 3600 m a.s.l.), in the Andes of central Chile, approximately 50 km east of Santiago. We mapped and measured size (as a surrogate for age) of individual cushions in two populations and used a standard spatial analytical tool (semivariograms) to test our alternative a priori hypotheses related to colonisation mode of the cushion species. In both populations, the size distribution of A. madreporica reflected a negative exponential or inverse-J pattern, typical of uneven-aged populations, where most of the cushions belonged to relatively smaller size classes, in effect, a regular success in the establishment of seedlings, where all size classes of cushions were represented in the population. The results were site-specific, where best-fit semivariograms for spatial cushion's size distribution suggested a gradual colonisation in one population and an episodic colonisation in the other population. Microsite distribution proved to be homogeneous at both sites. Thus, the study of the spatial explicit size-age population distribution of an alpine species provides valuable information about the frequency, magnitude and site variation of the reproductive pulses in these harsh environments. [ABSTRACT FROM AUTHOR]

Cushion plants are one of the most common growth forms in alpine habitats. Their low stature, dense canopy, and compact form allow them to decouple their microclimate from the surrounding environment, mitigating the effect of low temperatures and drought, enhancing the survival of other species. In this study, we evaluated the modifications on soil temperature and moisture over an entire growing season by two cushion species (Laretia acaulis and Azorella monantha) in alpine communities located at two different elevations in the central Chilean Andes. Additionally, we performed seedling survival experiments with two native herbaceous species (Hordeum comosum and Erigeron andicola) and the non-native forb Cerastium arvense to assess if seedling survival is higher within cushions than outside them. Our results indicated that cushions ameliorated extreme low and high substrate temperatures, improved soil moisture, and enhanced seedling survival of the three herbaceous plant species evaluated. Our results suggest that microclimatic modifications associated with cushion plants could be important for the establishment and survival of other plant species, both native and non-native, in the high alpine communities of central Chile. [ABSTRACT FROM AUTHOR]

• 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. [ABSTRACT FROM AUTHOR]

• In alpine habitats, positive interactions among plants tend to increase with elevation as a result of altitudinal increase in environmental harshness. However, in mountains located in arid zones, lower elevations are also stressful because of scarce availability of water, suggesting that positive interactions may not necessarily increase with elevation. • Here we analysed the spatial association of plant species with the nurse cushion plant Laretia acaulis at two contrasting elevations, and monitored the survival of seedlings of two species experimentally planted within and outside cushions in the semiarid Andes of central Chile. • Positive spatial associations with cushions were more frequent at lower elevations. Species growing at the two elevations changed the nature of their association with cushions from neutral or negative at higher elevations to positive at lower elevations. Survival of seedlings was higher within cushions, particularly at lower elevations. • The increased facilitation by cushions at lower elevations seems to be related to provision of moisture. This result suggests that cushion plants play a critical role in structuring alpine plant communities at lower elevations, and that climatic changes in rainfall could be very relevant for persistence of plant communities. [ABSTRACT FROM AUTHOR]

• Andean native species are dependent on pollen transport by insects for seed set. • The invasive Taraxacum officinale co-flowers and shared pollinators with natives. • The overlap in pollinators was not related with flower similarity among species. • Taraxacum officinale might alter the reproduction of native species in the Andes. Non-native plant species with a taxonomic affinity to co-occurring natives may perform best as invaders into plant-pollinator networks because their similarity in floral morphologies with natives enabled them to co-opt their pollinators. However, non-native species may impact the reproduction of natives by altering their pollination only if they share pollinators and if natives depend on pollinators for seed-set, premises that are often assumed but seldom tested. The non-native invasive species Taraxacum officinale (Asteraceae) co-exist and co-flower with other asteraceous species in the high elevation of the central Chilean Andes. In this study we determined the dependence on pollinators for seed-set in five co-occurring and co-flowering native asteraceous species and assessed the similarity in insect pollinators between the non-native and this five native species that differ in their floral similarity with the former species. Pollinator exclusion experiments were performed to determine the dependence of pollinators for seed-set in the native species. Characteristics such as capitula size and colour were measured to quantitatively assess the similarity in the floral morphology between T. officinale and the native species. The identity and visitation rates of insect pollinators that visit T. officinale and the five native species were determined. Native species were highly dependent on the transport of pollen by insects to produce seeds. T. officinale shared a moderate to high proportion (34–94%) of insect pollinators with the five asteraceous native species studied, but the level of overlap in pollinators was not related with the floral similarity between the non-native and the native species. The presence of the non-native species T. officinale has the potential to alter the pollinator services to co-flowering native species in the Andes of central Chile, with no higher impacts on those species with the most similar flowers. [ABSTRACT FROM AUTHOR]

Abstract: It has been predicted that subalpine forests will be negatively affected by global warming; however, direct responses to experimental warming have been scarcely examined in these systems. In this study we evaluated the effects of higher temperatures with and without water addition on the survival and growth of recently emerged (small) and large seedlings of the widely distributed species Nothofagus pumilio in subalpine forests of the southern Chilean Andes. We also examined the variations in seedling traits related to carbon balance in order to infer the causal mechanisms of survival and growth responses. Treatments of open top chambers (OTCs) were combined with watering in two locations with differing climates: Antillanca (40°S, humid) and Cerro Castillo (46°S, drier). OTCs increased mean and maximum air temperatures by 0.6°C and 2–3°C, respectively, and decreased soil humidity by 56% in Antillanca and 30% in Cerro Castillo, fulfilling methodological expectations and climate model predictions. After two complete growing seasons, the survival, relative growth rate (RGR), biomass, and a suite of seedling traits were measured and analyzed using mixed-effects models. Warming and warming in combination with watering significantly increased large seedling survival in Cerro Castillo. In Antillanca, warmer conditions increased the height, biomass, and leaf area of small seedlings, and the RGR of large seedlings. In this location, warming also caused lower leaf carbon isotopic composition in both age classes and higher specific leaf area in small seedlings, suggesting whole-plant carbon gain improvements; warming did not produce any drought effects. Our results indicate that warming produces positive effects on the seedling establishment of N. pumilio in the southern Andes, highlighting the importance of site-specific effects in response to climate change in widespread species. Site-specific effects can most likely explain the discrepancies between the results of this study and the predictions outlined by previous studies for these forests. [Copyright &y& Elsevier]

Mountain protected areas provide a range of ecosystem services including conserving biodiversity, while often providing recreation and tourism opportunities. Unfortunately, tourists and pack animals used to transport equipment can damage sensitive alpine vegetation particularly when they leave trails. This study assessed the impacts of disturbance from off trail use on alpine vegetation in a popular park in the Andes. The effect of different levels of disturbance as well as abiotic factors on alpine steppe vegetation was assessed using generalized linear models and ordinations in 91 plots (20 m2) in the popular Horcones Valley that is used to access remote areas in Aconcagua Provincial Park in Argentina. Disturbance off trails resulted in declines in the cover of native plants, including the endemic shrub Adesmia aegiceras but increases in the cover of herbs including the non-native Convolvulus arvensis. Increased disturbance was associated with shifts from stress tolerant species to ruderal plants characterized by more acquisitive traits, including shorter plants with greater Specific Leaf Area. The research demonstrates the severity of impacts from off trail trampling including how trampling favours some species with specific traits over others and why it is important to limit off track use in areas of high conservation value. • Low alpine vegetation can be affected by off trail use from visitors and pack animals. • Off trail disturbance resulted in shifts of species dominance. • Ruderal and non-native species with acquisitive traits were favoured by disturbance. • Limiting off trail use is critical to conserve alpine plant biodiversity. [ABSTRACT FROM AUTHOR]