Plant functional diversity in tropical Andean páramos

ABSTRACT

Background: Tropical high mountains present extreme daily temperature variations, frequent high air evaporative demands and seasonal differences in soil water availability. Plants have adapted to these conditions through different avoidance-tolerance mechanisms. This review focuses on plant-growth forms and their adaptive strategies.

Aims: This integrated review of páramo plant traits aims at contributing to understanding the functioning of plant-growth forms and their significance on ecosystem properties under environmental climate and land-use changes.

Methods: Plant responses are presented along avoidance-tolerance gradients considering three main aspects: freezing resistance, water relations and gas exchange characteristics. Results from 45 herbaceous and 42 woody species along elevational gradients in the Venezuelan high Andes were analysed.

Results: Leaf supercooling is the common avoidance response of woody plants to night-time freezing temperatures, while herbaceous plants tolerate frost. Trees and caulescent rosettes maintain more positive leaf water potentials under water deficit conditions compared to more tolerant herbaceous species. All plant growth-forms showed strong stomatal control under dry-season conditions.

Conclusions: Páramo plant growth-forms may be separated according to an avoidance-tolerance gradient in response to water deficit and low temperature resistance. Woody growth-forms tend to avoid both freezing and water stress, while herbaceous forms tolerate frost and resist an unfavourable water status. Grasses and cushion plants are at the tolerant extreme of the gradient and coincide in that both reach the highest elevations in the páramo. Andean giant rosettes are freezing avoidant, particularly susceptible to water deficit and the most vulnerable, of all growth-forms, to changing environmental conditions.     

Ecohydrology of the Venezuelan páramo: water balance of a high Andean watershed

ABSTRACT

Background: The páramo provides key ecosystem services, including regulation and provision of water. To understand the underlying functions, an ecosystem approach is necessary.

Aims: We quantified the combined effect of vegetation and soils (integrated topographic and vegetation units – TVU) on the hydrological balance of a Venezuelan páramo micro-watershed and analyse its hydrological response to intra- and interannual rainfall variability.

Methods: Data (2008–2016) from meteorological stations of TVUs and of a streamflow station was used to calculate watershed level hydrologic balances. We quantified the impact of the TVUs outputs by calculating evapotranspiration under non-standard conditions (ETc adj).

Result: Evapotranspiration of wetlands and tarns was high, exceeding annual precipitation. Shrubland had low evapotranspiration. Recharge of páramo reservoirs (soils, wetlands, tarns) occurred when monthly rainfall exceeded 90 mm. In dry years there were lower water yields with less effective hydrological regulation. In average years the differences between input and output in watershed balances were very small.

Conclusions: The high and constant evapotranspiration of the wetlands and tarns (due to permanent water availability) suggests they could maintain streamflow during dry periods. Their high evapotranspiration rates are compensated by low rates in shrublands units, reducing the mean total evapotranspiration of the watershed. The watershed balances suggest a limited regulatory capacity in these relatively dry páramos with no volcanic soils.     

Ecological research in the tropical alpine ecosystems of the Venezuelan páramo: past,present and future

ABSTRACT

Background: Tropical mountain ecosystems of the Northern Andes have long fascinated researchers because of the unique conditions associated with cold climates in equatorial latitudes. More than six decades have elapsed since the beginning of systematic ecological research in the Venezuelan páramos, making them one of the best-studied tropical alpine regions in the world.

Aims: We review the conceptual development and state of the art of ecological research in the Venezuelan páramos, with emphasis on environmental and plant ecology research, presenting a general framework for the studies included in this special issue.

Methods: We provide a historical sketch of the periods that have marked ecological studies in the Venezuelan páramos. Then, we synthesise research on environmental drivers, plant population and community ecology, ecosystem functioning, the response of the páramo to climate change and human disturbance; we finally consider agroecology and conservation.

Results and conclusions: This review demonstrates the significant contributions made to alpine ecology in key areas such as biodiversity/ecosystem function changes during succession, nutrient cycling, species interactions and socio-ecological research. We indicate the need to develop a more integrated view of the links between evolutionary processes, functional diversity, community dynamics and ecosystem services both in natural and human-impacted areas.     

Data-driven techniques for modelling the gross primary production of the páramo vegetation using climate data: Application in the Ecuadorian Andean region

As one of the main areas of carbon cycle and climate change studies, water and CO₂ relations are of great significance for estimation of gross primary production (GPP). Various biogeochemical process-based models have been set up to estimate the GPP based on mathematical representation of biological, physiological and ecological processes. However, they ended up increasing the complexity and computational processing power due to the large number of physical equations that need to be solved. Computational time becomes an important matter in the simulation of multiple scenarios using models for long periods of time (e.g. climate projections). Data driven surrogate models have proven to be a useful tool for environmental modelling especially when ecological and climatic co-variates are large. The advantages of Data Driven Models (DDM) are: the possibility of adding new independent variables even if their understanding is weak, and short computational time to run. The aim is to explore the ability of DDMs to replicate a biochemical model calculating GPP. This study evaluates the performance of four surrogate DDMs, namely linear regression method (LRM), model tree (MT), instance-based learning (IBL) and artificial neural network (ANN). A simple empirical and semi-empirical relationship between GPP and climatic variables are studied. Input variable selection (IVS) methods were used to decide on the most relevant and potential environmental model inputs and then followed by a two-step approach which included a model-free and a model-based technique. Data from the highlands (páramo ecosystem) in the Ecuadorian Andean Region from 12-year time-series (2000 − 2011) were used to evaluate the models at various time frames and at different altitudes. The GPP time series data for the same period were derived from an earlier study using the biomodel BIOME-BGC (BioGeochemical Cycles), which is a comprehensive physical based model used in different analysis of carbon fluxes around the world. So-called IBL (nearest neighbour method) showed a great capability to reproduce the GPP when data was aggregated to monthly time frame. The computational time used to evaluate the time series with IBL as the selected DDM is shorter with enough accuracy for using it in multi-model runs.     

Peatland carbon stocks and accumulation rates in the Ecuadorian páramo

The páramo is a high altitude tropical Andean ecosystem that contains peatlands with thick horizons of carbon (C) dense soils. Soil C data are sparse for most of the páramo, especially in peatlands, which limits our ability to provide accurate regional and country wide estimates of C storage. Therefore, the objective of our research was to quantify belowground C stocks and accumulation rates in páramo peatland soils in two regions of northeastern Ecuador. Peatland soil cores were collected from Antisana Ecological Reserve and Cayambe-Coca National Park. We measured soil C densities and ¹⁴C dates to estimate soil accumulation rates. The mean peatland soil depth across both regions was 3.8 m and contained an estimated mean C storage of 1282 Mg ha^?1. Peatlands older than 3000 cal. year BP had a mean long-term C accumulation rate of 26 g m^?2 year^?1, with peatlands younger than 500 cal. year BP displaying mean recent rates of C accumulation of 134 g m^?2 year^?1. These peatlands also receive large inputs of mineral material, predominantly from volcanic deposition, that has created many interbedded non-peat mineral soil horizons that contained 48 % of the soil C. Because of large C stocks in Ecuadorian mountain peatlands and the potential disturbance from land use and climate change, additional studies are need to provide essential baseline assessments and estimates of C storage in the Andes.     

The impact of roads on the avifauna of páramo grasslands in Cajas National Park,Ecuador

National parks are an important tool for conserving biodiversity, particularly in areas of high biodiversity and endemism such as the tropical Andes. However, national parks often face a variety of stressors related to recreation, road construction and illegal extraction of natural resources. Unfortunately, the influence of these stressors for biodiversity is rarely well documented. Cajas National Park in Ecuador is no exception. Despite being traversed by the Cuenca-Molleturo-Naranjal road, effects of the road construction on biodiversity have not been determined. We therefore assessed the influence of road proximity on bird species richness and abundance as well as composition of bird habitat groups in Cajas National Park using transect walks at 25 m and 250 m distance to the road (overall 18 transects, each 1 km length). In total, we recorded 1110 individuals of 28 páramo bird species. Overall species richness did not differ between transects near and far from the road. Nevertheless, the average abundance of shrubby páramo species was significantly higher far from the road than near the road (Far = 36, Near = 25). Moreover, we found a tendency towards differences in the composition of bird habitat groups between transects near and far from the road. One aspect potentially driving the observed patterns was the increasing proportion of planted non-native woody tree species within páramo grassland near the road, which may have caused reduced abundances of shrubby páramo bird species there. While roads represented a clear impact on the composition of bird species in the páramo, the major effect seems to be driven by the introduction of non-native plant species along the roadside. In order to reduce the impact of roads to a minimum, we suggest that park managers should control the introduction of such plant species.     

First report on generalized pollination systems in Melastomataceae for the Andean páramos

Melastomataceae is a megadiverse family with records of transitions from specialized to generalized pollination systems for several species. These transitions are associated with the colonization of new, unpredictable and/or impoverished pollinator habitats or habitats where specialized pollinators are scarce (e.g., in highland environments). The bee species diversity is low in highlands. Therefore, autonomous breeding systems such as apomixis and self-pollination emerge in these environments. In this paper, we studied the floral traits associated with the generalization of pollination systems and registered the floral visitors of two species in the Colombian Andes: Miconia cataractae and M. elaeoides. We investigated the breeding system of M. elaeoides. Both species presented small flowers, short anthers of medium pore size, and nectar-producing stomata on the base of the anthers. Miconia cataractae produced an average of 1.62 μl nectar/flower, a sugar concentration of 6.78%, whereas M. elaeoides produced 0.09 μl nectar/flower, a sugar concentration of 6.13%. We recorded a wide diversity of pollinators for both species, mainly insects from the orders Hymenoptera and Diptera. Miconia elaeoides presented a mixed breeding system and was also capable of setting fruits by apomixis. We conclude that flower and anther morphology, combined with nectar production, thus represent convergent traits resulting in a generalist pollination system shared by M. cataractae and M. elaeoides. Here, we presented the first generalist pollination system recorded for Miconia (and the Melastomataceae) in the Andes, the first report for a species from the small-pored section Amblyarrhena, and the first report for a species from the large-pored section Cremanium in Colombia.     

Three new species of Lysipomia (Lobeliaceae) endemic to the páramos of southern Ecuador

Three species of Lysipomia, L. caespitosa, L. cylindrocarpa, and L. speciosa, from a tropical alpine ecosystem called páramo, are described, illustrated, and compared to related species. Notes on morphology, especially floral presentation, and general ecology are given where possible. A published key to the species in Ecuador is revised to include the three new species plus L. multiflora McVaugh, a species previously known only from northern Peru.     

Involvement of the vacuolar processing enzyme γVPE in response of Arabidopsis thaliana to water stress

Plant vacuoles play several roles in controlling development, pathogen defence, and stress response. γVPE is a vacuolarlocalised cysteine protease with a caspase-1 like activity involved in the activation and maturation of downstream vacuolar hydrolytic enzymes that trigger hypersensitive cell death and tissue senescence. This work provides evidence that γVPE is strongly expressed in Arabidopsis guard cells and is involved in water stress response. The γvpe knock-out mutants showed reduced stomatal opening and an increased resistance to desiccation suggesting a new role of γVPE in control of stomatal movements.     

Influence of habitat modification by livestock on páramo bird abundance in southern Andes of Ecuador

Livestock grazing on natural grasslands is widespread with negative consequences to biodiversity. In the High Andes, páramo grassland is a distinctive ecosystem where the influence of livestock grazing on páramo birds is poorly documented. We assessed the influence of habitat modification of páramo grassland related to livestock grazing on bird habitat guilds in the southern Andes of Ecuador. We recorded birds occurring along transects located in areas which showed a gradient (low to high) of grazing pressure. We found a decrease in abundance of páramo specialists in transects with more grazing pressure. We interpret this habitat modification as loss of key habitat necessary for habitat-specialized birds.     

Genera of Scrophulariaceae in the páramos of the Venezuelan Andes

A report on the family Scrophulariaceae in the páramos of the Venezuelan Andes is made, which includes 13 genera. A key to the genera is provided.     

Changes in the patterns of synthesis of ribonucleic acid species in immature rat uterus in response to oestradiol-17β

1. After treatment of immature rats with diethylstilboestrol, the wet weight and RNA content of uterine tissue increased rapidly, reaching a peak at 40hr. After an initial lag of a few hours, the acid-soluble ribose and protein contents also rose to maxima at 40hr. No increase in DNA content occurred until at least 24hr. after treatment. 2. The RNA from immature rat uterus isolated at various times up to 6hr. after administration of oestradiol-17beta was labelled by injecting [(3)H]uridine and [(3)H]guanosine intraperitoneally 30min. before the animals were killed. It was fractionated on columns of kieselguhr coated with methylated serum albumin and the radioactivity in fractions corresponding to transfer RNA, 7s RNA, ribosomal RNA, Q(1)-RNA, Q(2)-RNA and DNA-like RNA was determined. 3. The radioactivity of the whole RNA increased steadily for 6hr. after hormone treatment. The earliest changes occurred in the Q(1)-RNA (ribosomal RNA precursor), whereas at longer time-intervals the radioactivity of the ribosomal RNA, 7s RNA and transfer RNA increased by four- to five-fold. The radioactivity of the DNA-like RNA increased by about 50%, but only at the longer time-intervals. 4. It is concluded that one of the earliest changes in response to oestradiol treatment is a major increase in synthesis of ribosomal RNA followed later by a similar increase in synthesis of transfer RNA and by a much smaller increase in synthesis of DNA-like RNA. The change in synthesis of ribosomal RNA in immature rat uterus may represent one of the most important responses to oestradiol treatment.     

Improving the provision of ecosystem services from urban forest by integrating the species’ potential environmental functions in tree selecting process

Due to the fact that urban environments and population demands are evolving rapidly and species selection is inevitable, it is possible to gain substantial environmental benefit by implementing more effective urban tree planting programs, especially with the aim of increasing the upcoming provision of multiple ecosystem services (ES) through proper species selection. In this paper, we used a new approach to improve the potential of urban trees in optimizing more desirable environmental benefits. This was done by selecting the most appropriate tree species among a vast range of species based on their potential environmental function (both services and disservices) in Tabriz city, Iran. Also, three main planting scenarios (divided to six sub-scenarios) were developed so as to understand the long-term effectiveness of introducing the selected tree species in improving the environmental benefits (both urban forest structure and ES) in comparison with planting the existing tree species. The results indicate that regardless of the quantity of planting, the benefits of introducing the selected trees will be more than planting the existing species. Moreover, as the amount of the annual planting of the recommended species increases, so does the improvement in the projected tree characteristics and ES. This approach creates more opportunities which enable urban forest managers and policymakers to understand the importance of selecting the proper urban tree species when looking for a nature-based solution to promote the wellbeing of the urban population, to create more livable and ecologically sustainable cities and to mitigate urban environmental problems.

     

A new species of Elatine (Elatinaceae) from the Colombian páramos in the northern Andes

Elatine paramoana, a new species from the high Andean region of the Cordillera Oriental in Colombia, is described, illustrated, and compared to its nearest relatives,E. ecuadoriensis andE. fassettiana. Notes on morphological variation and general ecology and a key to the central and northern Andes species are also given.     

How should parents adjust the size of their young in response to local environmental cues?

Models of parental investment typically assume that populations are well mixed and homogeneous and have devoted little attention to the impact of spatial variation in the local environment. Here, in a patch‐structured model with limited dispersal, we assess to what extent resource‐rich and resource‐poor mothers should alter the size of their young in response to the local environment in their patch. We show that limited dispersal leads to a correlation between maternal and offspring environments, which favours plastic adjustment of offspring size in response to local survival risk. Strikingly, however, resource‐poor mothers are predicted to respond more strongly to local survival risk, whereas resource‐rich mothers are predicted to respond less strongly. This lack of sensitivity on the part of resource‐rich mothers is favoured because they accrue much of their fitness through dispersing young. By contrast, resource‐poor mothers accrue a larger fraction of their fitness through philopatric young and should therefore respond more strongly to local risk. Mothers with more resources gain a larger share of their fitness through dispersing young partly because their fitness in the local patch is constrained by the limited number of local breeding spots. In addition, when resource variation occurs at the patch level, the philopatric offspring of resource‐rich mothers face stronger competition from the offspring of other local mothers, who also enjoy abundant resources. The effect of limited local breeding opportunities becomes less pronounced as patch size increases, but the impact of patch‐level variation in resources holds up even with many breeders per patch.     

Fire temperatures and postfire plant community dynamics in Ecuadorian grass páramo

Three aspects of the páramo vegetation's response to fires were investigated: the measurement of fire temperatures, general observations of changes in plant communities following fires, and monitoring the fate of individual plants after burning.Fire temperatures were strongly influenced by the physiognomy of the vegetation, dominated by tussocks of Calamagrostis spp. Temperatures were highest amongst the upper leaves of the tussock (sometimes >500°C). The middle levels of the tussock experienced temperatures in excess of 400°C, but in the dense leaf bases temperatures were often below 65°C. On the ground between tussocks, temperatures were variable, whereas 2 cm below ground temperatures failed to reach 65°C.Plant survival depended on the intensity of the fire and the plant's position within the tussock structure. Survival was often the result of high temperature avoidance (with buds shielded by other plant parts or buried beneath the soil surface).Post-fire Calamagrostis tiller mortality rates were high and tussock regrowth was slow. Some other species appear to maintain their populations by exploiting this recovery phase for seedling establishment on tussocks.Between tussocks, changes of occupancy at the level of the individual plants were greater after fire than in control vegetation. Most transitions were random. Those which departed from random often involved gaps and were related to post-fire mortality, regrowth from below-ground parts, colonisation or, in the case of a clonal mat-forming species, to spatial rearrangement of rosettes. Recovery was slower at higher altitude. Recovery was much slower in burned plots when the upper 2 cm of soil was removed (along with buried plant parts) compared with burned plots.Qualitative observations suggest that recovery may consist of a cyclical process, mediated by the serial dominance of several species that are physiognomically important.The frequency of fires determines the amount of fuel accumulated within grass tussocks and some plants may be unable to survive repeated burning. Chance survival of species in unburned patches of vegetation and random colonisation of gaps may be important determinants of subsequent community structure.     

Selective logging in tropical forests decreases the robustness of liana–tree interaction networks to the loss of host tree species

Selective logging is one of the major drivers of tropical forest degradation, causing important shifts in species composition. Whether such changes modify interactions between species and the networks in which they are embedded remain fundamental questions to assess the ‘health’ and ecosystem functionality of logged forests. We focus on interactions between lianas and their tree hosts within primary and selectively logged forests in the biodiversity hotspot of Malaysian Borneo. We found that lianas were more abundant, had higher species richness, and different species compositions in logged than in primary forests. Logged forests showed heavier liana loads disparately affecting slow-growing tree species, which could exacerbate the loss of timber value and carbon storage already associated with logging. Moreover, simulation scenarios of host tree local species loss indicated that logging might decrease the robustness of liana–tree interaction networks if heavily infested trees (i.e. the most connected ones) were more likely to disappear. This effect is partially mitigated in the short term by the colonization of host trees by a greater diversity of liana species within logged forests, yet this might not compensate for the loss of preferred tree hosts in the long term. As a consequence, species interaction networks may show a lagged response to disturbance, which may trigger sudden collapses in species richness and ecosystem function in response to additional disturbances, representing a new type of ‘extinction debt’.