Plant diversity and composition along páramo elevation and disturbance
gradients within Podocarpus National Park, Ecuador
AUTHOR: Eric Creeden, University of Idaho, Moscow, ID 83843
COMMITTEE/PI’s: David Tank, Alex Fremier, University of Idaho, Moscow, ID 83843
Abstract
The Andean páramo ecosystem is a unique and valuable ecosystem in the Andean highlands of
South America. Páramo ecosystems provide ecosystem services of carbon storage, water
filtration and provision, and human use, while housing remarkably high levels of vegetative
biodiversity. An assessment of plant community diversity and composition utilizing community
ecology and phylogenetic approaches will be conducted along páramo elevation and disturbance
gradients within Podocarpus National Park (PNP), Ecuador. Objectives of the study will be to:
1) assess how vegetative diversity, composition, and traits differ along elevation and disturbance
gradients; 2) integrate community ecology and phylogenetic methods to address structure and
adaptability of páramo ecosystems; and 3) determine if páramo vegetative communities in PNP
have seen changes in the last 20 years. Potential insights into the survivability and adaptability
of páramo ecosystems to modifications from anthropogenic disturbance and/or global climate
change pressures are desired outcomes of the study.
Keywords: páramo, Ecuador, biodiversity, gradient, community ecology, phylogenetics,
disturbance, climate change
PROBLEM STATEMENT: Ecuador’s páramo ecosystems are under a variety of threats.
Ecological research in understudied ecosystems such as the Andean páramos may provide insight
into how ecosystems are able to adapt or react to development and/or global climate change
pressures.
INTRODUCTION
The Andean páramo ecosystem is a unique and valuable ecosystem in the Andean
highlands of South America. Levels of endemic plant diversity and avifaunal diversity are
remarkably high given the small land area of this particular ecosystem. In addition to supporting
high levels of biodiversity, páramo ecosystems provide ecosystem services of carbon storage,
water filtration and provision, and provide resources for indigenous and local human
communities. The Andean páramo ecosystems have significant social and cultural importance as
well as ecological importance to local communities. Human and resource development within
the southern region of Ecuador threaten the survival and functioning of the páramo ecosystem, as
well as the quality of resources it provides, notably water. Podocarpus National Park (PNP)
houses many of southern Ecuador’s river headwaters, and supplies water to many local
communities.
Research will be conducted to assess the ecology and watershed management of the
Andean páramo in southern Ecuador as part of a collaborative effort between the University of
Idaho (UI) and the Universidad Técnica Particular de Loja (UTPL). Research concerning the
ecology of the Andean páramo will focus around four topics: 1) plant species diversity and
composition across an elevation gradient; 2) development of a plant reference collection for diet
of the Andean bear (Tremarctos ornatus); 3) evaluation of Andean bear foraging on Bromeliads;
and 4) social and cultural importance of the páramo ecosystem.
This proposal focuses primarily on assessing plant diversity and composition of a páramo
ecosystem across an elevation gradient within PNP, Ecuador. Assessment of plant community
diversity and composition utilizing community ecology and phylogenetic approaches will be
used, and may provide insight into survivability and adaptability of páramo ecosystems to
modifications from anthropogenic disturbance and/or global climate change pressures.
BACKGROUND LITERATURE REVIEW
Community assemblages of plant organisms at a particular location are determined by a
multitude of interacting processes that can be explained with both community ecology and
evolutionary biology approaches. Linkages between the two approaches have recently emerged
with advancements in DNA analysis. Plot-level studies collecting information on plant species
traits and abundance estimates have been used frequently to characterize vegetative assemblages
of areas and determine underlying processes of species and environmental interactions. The
páramo of southern Ecuador is a tropical montane ecosystem that exists along gradients of
elevation and disturbance. Gradients have frequently been used in scientific studies and the
páramo may be an ideal place for such analyses (Malhi et al. 2010). Anthropogenic disturbance
from grazing and fire are common practices at lower elevation páramo areas within the tropical
Andes and near PNP, and have occurred for centuries (Luteyn 1999; Keating 2000; Niemann and
Behling 2008). Global environmental change has emerged as a potential threat to páramo
ecosystems, and predictions of future climate within the tropical Andes show significant
warming trends (Intergovernmental Panel on Climate Change 2007; Urrutia and Vuille 2009).
Community Ecology and Evolutionary Biology (Phylogenetics)
The structure and function of plant communities in specific locations are dependent on
many spatial and temporal processes. Plant communities can be defined as groups of interacting
populations that coexist in space and time (Gurevitch et al. 2002). Much debate has occurred
over the past century regarding the relative magnitude of influence that biotic (emergent
properties), abiotic (environmental), and random (stochastic) factors play in determining
community structure (Gurevitch et al. 2002). Emergent properties result through interactions
such as competition or mutualism within a community (Gurevitch et al. 2002), while
environmental factors may incorporate climate and soil properties. Relationships within plant
communities show scale dependencies in both space and time (Webb et al. 2002; Johnson and
Stinchcombe 2007; Cavenders-Bares et al. 2009). The evolution of dominant scientific
paradigms and philosophies in plant ecology has progressed in time from more deterministic and
reductionist (Clementsian) viewpoints to more holistic (Gleasonian) viewpoints that incorporate
random processes governing plant community structure (Gurevitch et al. 2002). Current
scientific ideologies of plant diversity and composition emphasize that plant communities are the
result of many interacting processes and should not be classified by a single approach. The
fields of community ecology and evolutionary biology provide differing and complimentary
methodologies for investigating plant community structure and function.
Community ecology attempts to describe and analyze the interactions of populations
within a specific area with their environments. Johnson and Stinchcombe (2007) and CavendersBares et al. (2009) note that most community ecology studies ignore genetic variation and
evolutionary change under the assumption that evolutionary processes do not act on the short
time scales of community ecology studies. Niche-related processes (competitive exclusion),
neutral processes (dispersal and disturbance), and historical factors (biogeography) are three
dominant processes that determine community assembly, diversity, and composition (Cavenders
et al. 2009). Plant herbivore and plant pathogen interactions may also impact community
diversity by influencing coevolution of varying traits between plant species and their natural
enemies (Herms and Mattson 1992; Cavenders-Bares et al. 2009). Trait based approaches of
describing community assemblages and plant form and function also exist, and have shown
applicability for use along ecological gradients (Ackerly and Cornwell 2007). Specific leaf area,
leaf size, wood density, and maximum height were found to covary across a soil moisture
availability gradient in California (Ackerly and Cornwell 2007).
Evolutionary biology is the study of genetic variation and the methods of genetic and
phenotypic change within populations (Johnson and Stinchcombe 2007). Phylogenetic studies
can determine the relatedness and evolutionary history of species within a community.
Cavenders-Bares et al. (2009) note that phylogenetics can help to resolve arguments and
challenge classical ideas in community ecology, as well as predict future ecosystem properties.
Phylogenetic clustering indicates that related species are located closer to each other than would
be expected by random chance, and that environmental filters may drive community structure
(Webb et al. 2002; Johnson and Stinchcombe 2007). Phylogenetic overdispersion implies that
related species are located farther away from each other than would be expected by random
chance, and that interspecific competition or convergent evolution may be the driving forces of
community structure (Webb et al. 2002; Johnson and Stinchcombe 2007). Cavenders-Bares et
al. (2009) stated the utility and promise of using phylogenetics to predict future community
assemblages and for the potential use of phylogenetics for conservation related efforts.
Methods of utilizing both approaches to address scientific inquiries are currently at the
forefront of the scientific literature, but may be difficult to integrate (Webb et al. 2002; Johnson
and Stinchcombe 2007; Cavenders-Bares et al. 2009). Processes of community ecology may
shape evolutionary biology, and evolutionary biology in turn can influence community ecology
dynamics (Johnson and Stinchcombe 2007; Cavenders-Bares et al. 2009). Understanding the
direction and magnitude of these interacting processes may provide novel and complementary
information regarding plant communities. The use of phylogenetics in association with
community ecology, however, allows for the expansion of community ecology questions to
larger spatial and temporal scales (Cavenders-Bares et al. 2009). This study attempts to integrate
the two fields through field analysis of community ecology and trait data along an elevational
gradient, and analysis of phylogenetic structure of species along that gradient. Webb et al.
(2002) stated that understanding phylogenetic structures across gradients shows promise for
work in community ecology.
Ecology and Study of Mountainous Tropical Regions
High altitude, tropical regions are less studied and have higher biodiversity than their
temperate region counterparts, and can enhance understanding of ecosystem ecology (Malhi et
al. 2010). Because of high biodiversity and species turnover, tropical montane regions have
relatively narrower thermal plant niches than other areas, potentially making changes from
temperature easier to detect (Malhi et al. 2010). Crimmins et al. (2011) showed that water
balance, not temperature, was the principal factor defining plant species distributions in
California. Heterogeneity of montane sites including microclimate, topography, disturbance
history, soil diversity and different limitations to plant success such as water, energy,
temperature, and light may all play a role in determining species distributions in a particular area
Keating 2000; Korner 2007; Malhi et al. 2010; Crimmins et al. 2011).
Mountainous regions generally show steep gradients in elevation and other biophysical
characters such as precipitation and temperature. Malhi et al. (2010) listed five reasons for
studying gradients in high elevation tropical regions including: 1) they are intrinsically
interesting and understudied; 2) high species diversity exists along gradients; 3) elevation
transects are good for testing environmental controls; 4) they provide information about past
climates; and 5) they are good for monitoring ongoing and future impacts of atmospheric change
in the tropics. Elevation gradients may be used to increase understanding regarding
temperature’s role on biodiversity, ecology, ecosystem function, and global change response
(Malhi et al. 2010). Korner (2007) showed the utility of using gradients for testing biotic
responses to geophysical influences, but stressed the necessity of discerning geophysical
phenomena such as temperature gradients, from coincidental local phenomena such as fire, land
use, and drought, and from other gradients present. Korner (2007) stated that altitudinal
gradients impacted little by moisture gradients, such as humid mountains, may be particularly
important for gradient studies in ecology. Páramo ecosystems are located in humid, tropical
areas, and provide ideal locations for gradient analyses.
Páramo Ecosystems
Páramo ecosystems are present in the tropical mountains of South and Central America,
Africa, and Asia, and are found between timberline and snowline (Lauer 1981, Luteyn 1999).
They are typically found at elevations between 3,000 and 4,000 m, but can extend upwards of
4,500 m (Lauer 1981; Keating 1999; Sklenář and Jørgenson 1999). Páramo ecosystems are
distributed most extensively in Latin America in the countries of Ecuador, Columbia, and
Venezuela (Lauer 1981). Sklenář and Jørgenson (1999) found that the distribution patterns of
páramo plants in Ecuador were dependent on altitudinal range and distance between mountains.
Species with broad altitudinal ranges that potentially facilitated migration were generally found
on more mountains than species with narrow altitudinal distributions (Sklenář and Jørgenson
1999). Luteyn (1999) described Andean páramos as areas characterized by unique geographic,
geologic, climatic, physiognomic, and floristic features. Soils are relatively young and may
show high complexity in terms of rock composition, volcanic origin, structure and type (Luteyn
1999).
Three main types of páramo can be distinguished based on elevation including
subpáramo, páramo proper or grass páramo, and superpáramo, and may be dominated by shrubs
or grasses (Lauer 1981; Keating 1999; Luteyn 1999; Sklenář and Jørgenson 1999). Shrub páramo
are generally located in southern Ecuador and show some of the highest levels of endemism
within the country (Keating 2008). Ecuadorian páramos are generally dominated by tussock
grasses, but other growth forms include stem, basal, and acaulescent rosettes, cushions and mats,
upright and prostrate shrubs, and erect, prostrate, and trailing herbs (Ramsay 1997). Vegetation
show many morphological and physiological adaptations to tolerate harsh conditions including
high elevation, low temperatures, high ultraviolet (UV) exposure, freeze-thaw cycles, and high
winds (Luteyn 1999). Across an altitudinal gradient in southern Ecuador, Wilcke et al. (2008)
found that organic material, root/shoot ratio, and C/N ratio increasing with increasing altitude,
while pH and macronutrient (N, P, K, Ca, Mg) concentrations decreased with increasing altitude.
Disturbance Processes and Climate Change in the Tropical Andes
Numerous disturbance process, natural and anthropogenic, are at work within páramo
areas. Landslides and erosional events are common on the steep and wet slopes of the tropical
Andes, and are important sources of landscape heterogeneity and regeneration events. Shrub and
grass páramos are not isolated from anthropogenic disturbance and may frequently be grazed,
burned, or cut (Luteyn 1999; Sklenář and Jørgenson 1999; Keating 2000). Centuries of
anthropogenic disturbance have influenced the boundaries between páramo and the upper
altitudinal limit of timberline, and have also altered the vegetative composition of some páramo
communities (Luteyn 1999). Current disturbance intervals of anthropogenic burning and grazing
are thought to be too short for the maintenance of species diversity within the Andean páramos
(Luteyn 1999). Upper montane forested areas in southern Ecuador near PNP that have
undergone slash and burn forest clearing are often abandoned after bracken fern (Pteridium
arachnoidem) and other bushes invade and make grazing and agriculture difficult (Beck et al.
2008). Threat of invasion by alien species may be higher following disturbance events.
Tropical montane areas such as páramo are likely to see higher than average changes
from climate change because warming may be more pronounced at higher elevations
(Intergovernmental Panel on Climate Change 2007; Urrutia and Vuille 2009). Buytaert et al.
(2011) described humid tropical alpine environments as ecosystems most vulnerable to
environmental change. Feeley and Silman (2010) estimated a 900 m upslope migration of
species within the Andes under a 5 °C increase in mean temperature, and noted the potential for
barriers to species migration from human occupation and land use. A 3 ±1.5 °C increase in
mean temperature was predicted by 2100 for the tropical Andes using IPCC 2007 data by
Buytaert et al. (2011). An upslope migration of species may not be feasible in parts of the
páramo because species may already occupy the full extent of the available habitat, or may be
isolated from nearby páramo areas. Impacts to human and natural communities in the tropical
Andes from climate and environmental change include reduced water supply and flow, losses in
biodiversity, and decreases in soil carbon storage (Buytaert et al. 2010).
This proposal will build upon knowledge gained from Keating (1999; 2008) by collecting
additional data from more plots to assess plant community composition and structure along a
páramo elevation gradient. Data from Keating (1999) was collected from 1991-1993, so data
collected in 2011 by this proposal will provide a 20-year time step in sampling. Repeated
sampling will provide opportunities to assess changes within the studied páramo ecosystem, and
will complement community ecology and phylogenetic techniques used. Changes in sampled
species occurrences, abundances, character traits, and spatial locations may indicate that local or
global processes such as anthropogenic disturbance or climate change are influencing species
assemblages within páramo ecosystems.
OBJECTIVES
1) Assess vegetative diversity, composition, and traits along disturbance and elevational
gradients
2) Integrate community ecology and phylogenetic methods to address structure and
adaptability of páramo ecosystems
3) Determine if páramo vegetative communities in PNP have seen changes in the last 20
years
QUESTIONS
1) How do species presence and abundance correlate with environmental factors across
environmental and disturbance gradients?
a. Hypothesis 1: Species presence and abundance will vary along an elevation
gradient as environmental factors such as wind, water content, humidity, and
temperatures change.
b. Hypothesis 2: Along a disturbance gradient, species presence and abundance will
vary depending on time since disturbance and type of disturbance in addition to
environmental factors.
2) How does a phylogenetic understanding inform community assembly and evolution
across environmental and disturbance gradients?
a. Hypothesis 1: Phylogenetic clustering will be higher than expected at higher
elevations of the gradients because of greater assumed environmental controls.
b. Hypothesis 2: Less clustering, or phylogenetic over-dispersion, is expected at
lower elevations of the gradients.
c. Hypothesis 3: Phylogenetic patterns between the elevation and disturbance
gradients will differ.
3) How do vegetative traits inform patterns of community assemblage?
a. Hypothesis 1: Vegetative traits will vary along elevation transects as
environmental factors such as wind, water content, humidity, and temperature
change.
4) How does vegetative community ecology data compare between the two sampling
periods (1991/2011)?
a. Hypothesis: Community assemblages and traits will differ between the two time
periods due to change processes occurring at the study site.
5) How do spatial patterns of vegetation characteristics vary across gradients?
a. Hypothesis: Spatial patterns of vegetation characteristics such as clustering or
dispersion will provide insight into community assemblage processes.
METHODS
Study Area: Podocarpus National Park, Ecuador
The study area for this project lies within PNP in southern Ecuador, and encompasses
both the Loja and Zamora-Chinchipe provinces. The páramo ecosystems of PNP are generally at
lower elevations than other areas within Ecuador and the South American Andes. As noted by
Luteyn (1999), most Andean páramo ecosystems show a complicated paleohistory of glacial and
interglacial periods, have young and variable soils, cold and humid climates, and have vegetation
that shows morphological and physiological adaptations to high altitude, low temperatures, and
high UV exposure including leaf pubescence, retention of dead leaves, and other freeze tolerant
mechanisms (Luteyn 1999). Local topography is complex, impacting microclimates of
temperature and winds among other factors.
A thorough documentation of floristic composition of the study site was completed by
Keating (2008), with a total of 136 species documented within a 2.4 ha area. This site was
unique because of its high percentage of woody species, low dominance of grasses, and high
number of endemic plant species (Keating 2008). Physiographic features of surrounding areas
that impact species migration, elevation, and disturbance regimes were noted to influence the
community composition of PNP páramo ecosystems (Keating 2008).
Study Design
This study uses sampling methods similar to Keating (1999), which provided a
preliminary reference study within PNP regarding how páramo vegetation changes along an
elevation gradient in southern Ecuador. Objectives of Keating (1999) were to: 1) describe plant
composition and diversity; 2) summarize plant community patterns; and 3) suggest how changes
were related to environmental factors. Keating (1999) found distinct vegetation types within the
páramo study area using ordination techniques, and noted that species diversity remained
relatively constant across the gradient. This study will be build from Keating (1999) by
including disturbance gradients, additional environmental factors of soil metrics, and
incorporating spatial analysis and phylogenetic methodologies.
Sampling Design
A sampling design will be developed to account and test for variations such as slope,
aspect, and topographic position within the gradient study areas. Small changes in these variants
can show large differences in observed frequency and diversity of vegetation at a sampled
location. A stratified sampling design will be used in this study with 3x2 m sample plots
(quadrats) selected on gradients of elevation and disturbance (or both). First, a gradient will be
stratified if significant differences exist in vegetation such as between shrub or grass/herbaceous
dominated páramo. To account for local variation in slope, aspect, topographic variation, plots
will be allocated to each category or combination of categories. The number of plots allocated to
each stratified region will be proportional to each region’s area. The size and shape of a quadrat
can influence the results of an analysis, and rectangular plots may be preferable over square plots
when sampling on an elevation contour (Gurevitch et al. 2002). The longer side of the quadrat
will be arranged perpendicular to the slope at a given location to maintain constant elevation, and
a geospatial location will be recorded at the center of each plot.
Sample plots will be subdivided in 1x1 m squares for ease of analysis and greater
sampling efficiency. As many plots as possible from both elevation and disturbance gradients
will be sampled given the short duration of the field season for which data will be collected. The
two gradients will be paired so that they are as similar as possible except in terms of disturbance.
This will allow for both within and between transect analysis. If field and time constraints exist,
data collected in May-July 2011 may be collected for only one of the gradient types.
Data Collection
1) Community Ecology Data
Data will be collected for grass, herbaceous, woody/shrub, and moss species within each
plot. Frequency and percent cover will be calculated for grass, herbaceous, and woody/shrub
species. Height and percent cover will be documented for grasses and mosses. Species density
will be derived from frequency data and plot area. In addition to vegetative data, microsite and
microclimate information of elevation, slope, aspect, topographic position (rise, flat,
depression/top, bottom of gradient), soil pH, soil depth, soil texture, and percent rock will be
recorded at each plot location. Soil pH will be measured at 3 random locations within each plot.
Soil depth will be measured at 3 random locations within each plot by measuring distance to
rock. Soil texture will be measured at 3 random locations in each plot by hand texturing to
provide inference about soil water holding capacity. Percent cover of rock within each plot will
be estimated ocularly. Following the collection of plot data, community metrics will be
calculated including species richness, diversity, evenness, and dominance. Alpha, beta, and
gamma vegetative species diversity will also be calculated for all plots along both elevation and
disturbance gradients. Lastly, plots will be photographed for future reference.
2) Trait Data
Trait characteristics of leaf size, specific leaf area (SLA), and height will be collected
within each plot for herbaceous and woody/shrub species encountered similar to the study by
Cornwell et al. (2006). Height will be collected within each plot for grass species. Leaf size and
specific leaf area will be calculated for 2 randomly selected individuals per plot of easily
identifiable and common species within the gradient. Leaf area will be calculated in the lab by
overlaying the leaf on a 1cm2 grid and counting the number of overlain squares. Specific leaf
area will be calculated by dividing fresh leaf area by dry leaf mass. Leaves will be measured and
dried in the lab facilities at UTPL. Species height will be measured in the field using a
measurement tape and/or ruler.
3) Phylogenetic Data
A composite phylogeny for the study site will be developed using a species list that is
inclusive of all encountered plants within the sampled plots using. Phylogenetic trees will be
developed for both elevational and disturbance gradients, as well as for the entire study area.
Phylogenetic trees will also be developed separately for low and high regions of the gradients to
examine similarities or differences.
Data Analysis
1) Community Ecology Data
A variety of community ecology metrics will be calculated with the collected plot data to
assess vegetative diversity and abundance within the study area. Species richness is determined
by the number of different species within an area and will be recorded. The greatest number of
species within an area can determine the dominance of a particular species at a location, and may
indicate the influence of environmental or competitive interactions within the vegetative
community. Species diversity and evenness will be measured using a variety of metrics including
the Shannon-Weiner Index, Simpson’s Index and Shannon evenness. A Gini coefficient will be
estimated as a measure of species evenness (Gurevitch et al. 2002). The Shannon-Weiner index
may be biased compared to Simpson’s index, but is more sensitive to proportions of rare species
(Gurevitch et al. 2002). Because of its sensitivity to rare species, the Shannon-Weiner index
may be a good measure with the high biodiversity found within páramo ecosystems. In addition
to the afore mentioned indexes, the importance value (IV) is a weighted measure that sums the
relative cover, relative density, and relative frequency of a given species in a particular area, and
will be calculated in the study area. If a nested analysis is performed with the sampling of
quadrats at multiple spatial scales, species area curves will be developed for the plots in the study
area.
2) Trait Data
Trait information of species height and stem density were collected by Keating (1999) for
woody and grass species. Comparisons of species height and stem density will be conducted
between this study and data collected by Keating (1999) to assess if any changes in vegetative
traits have occurred at the study site between the two time periods of sampling. Alpha (within
community) and beta (between community) values of species’ trait values will be calculated
similar to methods developed and used by Cornwell et al. (2006) and Ackerly and Cornwell
(2007). Four pieces of information are needed to conduct this analysis including: 1) list of plots
or communities; 2) list of species sampled in each plot; 3) values of traits for each species in
each plot; and 4) measure of relative abundance of each species (Ackerly and Cornwell 2007). R
source code scripts are provided in the appendices of Ackerly and Cornwell (2007) for this
analysis.
3) Phylogeny Data
Phylogenetic trees will first be developed by using Phylomatic, a web-based tool for
quick tree assembly (Webb and Donoghue 2005; The Phylomatic Project). Phylocom may also
be run for analyzing phylogenetic community structure (Webb et al. 2008). Willis et al. (2008)
successfully used these tools to show phylogenetic patterns and relationships with climate
change. Analysis of phylogenies at both low and high regions of the sampled elevation and
disturbance gradients will allow for comparisons between both region and gradient.
Theoretically, higher phylogenetic conservatism and trait clustering should occur at higher
elevations where environmental controls on species assemblages are predicted to be stronger.
The opposite is predicted to be true at lower elevations. Along a disturbance gradient, higher
disturbance severity should favor early seral or colonizing species (potentially invasive or nonnative species) that are better adapted to disturbed environments. Analysis of phylogenetic traits
of colonizing species may show predictive power for future community assemblages.
RESEARCH RELEVANCE
Research concerning the ecology of the Andean páramo provides a novel way of
integrating ecological and social science research. The páramo ecosystem is a hotspot of
vegetative diversity with high levels of plant endemism. Keystone species of this ecosystem
including the Andean bear are dependent on lower trophic levels for nutrition and habitat, and
are impacted by land use decisions and changes to the páramo ecosystem. By assessing the
social and cultural significance of the páramo ecosystem, management actions may be guided
towards specific and targeted actions that maintain the ecological integrity of the region.
Understanding how plant diversity and composition change across the area may help to inform
feeding habits of local species. The development of a plant reference collection and use of DNA
extraction techniques will advance the work of molecular ecology and conservation genetics at
the UI and UTPL. By implementing a multi-disciplinary collaborative approach to ecological
research, this project intends to extend the scope of traditional research and help to answer both
basic and applied research questions.
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