On global change, direct and indirect interactions, and the structure of ecological communities: theoretical and empirical tests

Human induced global change (climate change, CO2 enrichment, nitrogen deposition, habitat degradation and biological invasions) is the most serious threat to biodiversity. Understanding how ecosystems will respond to different components of global change, and how these responses will affect key ecological processes, has become essential in contemporary ecology. For example, several studies have shown that exotic invasive species have negative impacts on the composition of communities, habitat structure and ecosystem processes. Particularly, exotic species may have negative effects on species interactions due to local extinctions, competition and/or replacement of interactions. Despite the large body of research demonstrating the negative effects of exotic species on native communities, clear responses of the effect of invasive species on seed dispersal mutualisms are scarcely documented or have focused on only a relatively few invasive species. In this dissertation I used exotic species to (1) re-evaluate the inclusion of facilitation in niche theory, (2) study the establishment success and invasion rates of exotic species (i.e. tens rule), and (3) determine how exotic species has influenced community structure by altering the interaction between plants and animals, specifically, plant-seed dispersers mutualisms. In these studies, I established that interactions with exotic facilitators could increase the size of the realized niche by increasing the spatial distribution (reducing dispersal limitations), or by modifying the physical and chemical conditions of the habitats, without altering the size of fundamental niche of native species. I found that the lack of information about failed species introductions and the tendency to report species that have become invasive more than those that have not results in an overestimation of establishment success and invasion rates. Additionally, I found that a suite of exotic vertebrate herbivores and exotic invertebrates lead to a cascade of linked extinctions by disrupting native plant-seed disperser mutualism. Together, my results highlight the negative effect of exotic invasive species and the importance of focusing on interactions among species to understand the causes and consequences of species loss from ecosystems

Invasive trees rely more on mycorrhizas, countering the ideal-weed hypothesis

The ideal-weed hypothesis predicts that invasive plants should be less dependent on mutualisms. However, evidence in favor of or against this hypothesis comes mainly from observational studies. Here, we experimentally tested this hypothesis using a two-factor greenhouse experiment, comparing the seedling growth response of different Pinus species (varying in invasiveness) to ectomycorrhizal fungal inoculation. Most species showed no response until they were 6 mo old, at which point inoculation increased growth between 10 and 260% among the different species. This growth response was higher for species with lower seed mass, higher dispersal ability, higher Z score (a proxy for invasiveness) and higher number of naturalized regions, all of which correspond to higher invasiveness. Our results show that timing is a crucial factor when comparing mycorrhizal dependency of different species. Dependence on mutualistic microorganisms could be part of a strategy in which invasive species produce smaller seeds, in greater number, which can disperse further, but where seedlings are more reliant on mycorrhizas to improve access to water, nutrients, and protection from pathogens. Our results suggest that reliance on mutualisms may enhance, rather than limit, nonnative species in their ability to spread, establish, and colonize.Fil: Moyano, Jaime. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Rodriguez Cabal, Mariano Alberto. University Of Vermont. Rubenstein School.; Estados Unidos. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Nuñez, Martin Andres. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina. University of Houston; Estados Unido

Do not come late to the party: initial success of nonnative species is contingent on timing of arrival of co-occurring nonnatives

Invasions by multiple nonnative species threaten native communities worldwide. We know that interactions among nonnative species influence nonnative success. However, we know relatively less about the influence of community assembly history on the interactions within nonnative species and thereby invasion success. To investigate this, we transplanted seedlings of two highly invasive shrubs, Rosa rubiginosa (hereafter ‘rose’) and Cytisus scoparius (hereafter ‘broom’), at two different times into mesocosm communities of native and nonnative species. We conducted priority and delay treatments that consisted of the early and late arrival of the invasive shrubs, respectively. We gave full priority/delay to each invasive shrub (rose early/late arrival, broom early/late arrival) and simultaneous priority/delay (simultaneous early/late arrival). We predicted that if assembly history were important, the invasive shrubs will benefit from early-arriving and will be disadvantaged by late-arriving and that arriving before the co-invader shrub will be more beneficial than arriving before the rest of the community. We also predicted that assembly history treatments that gave an advantage to invasive shrubs will more negatively affect native species than nonnative species. We found that the invasive shrubs did not benefit by early-arrival, but they were hindered by the early-arrival of the co-invader. The rose paid a high cost for late-arrival, but the broom was only impaired when its late-arrival implied arriving after the rose. Contrary to our predictions, natives paid a lower cost than nonnatives by arriving late. In general, our mesocosm experiment showed that the success of invasive species depended more on not arriving later than other invaders than on arriving early in the community. We suggest that community assembly history modulates the sign and strength of nonnative species interactions whose consideration might improve management practices.Fil: Torres, Agostina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Rodriguez Cabal, Mariano Alberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Nuñez, Martin Andres. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentin

Highly invasive tree species are more dependent on mutualisms

Why some species become invasive while others do not remains an elusive question. It has been proposed that invasive species should depend less on mutualisms, because their spread would then be less constrained by the availability of mutualistic partners. We tested this idea with the genus Pinus, whose degree of invasiveness is known at the species level (being highly and negatively correlated with seed size) and which forms obligate mutualistic associations with ectomycorrhizal fungi (EMF). Mycorrhizal dependence is defined as the degree to which a plant needs the mycorrhizal fungi to show the maximum growth. In this regard, we use plant growth response to mycorrhizal fungi as a proxy for mycorrhizal dependence. We assessed the responsiveness of Pinus species to EMF using 1206 contrasts published on 34 species, and matched these data with data on Pinus species invasiveness. Surprisingly, we found that species which are more invasive depend more on mutualisms (EMF). Seedling growth of species with smaller seeds benefited more from mutualisms, indicating a higher dependence. A higher reliance on EMF could be part of a strategy in which small-seeded species produce more seeds that can disperse further, and these species are likely to establish only if facilitated by mycorrhizal fungi. On the contrary, big seeded species showed a lower dependence on EMF, which may be explained by their tolerance to stressful conditions during establishment. However, the limited dispersal of larger seeds may limit the spread of these species. We present strong evidence against a venerable belief in ecology that species that rely more on mutualisms are less prone to invade, and suggest that in certain circumstances greater reliance on mutualists can increase spread capacity.Fil: Moyano, Jaime. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Rodriguez Cabal, Mariano Alberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Nuñez, Martin Andres. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentin

Secondary invasions hinder the recovery of native communities after the removal of nonnative pines along a precipitation gradient in Patagonia

The removal of nonnative species can lead to re-invasion by nonnative species, especially in communities with multiple co-occurring invaders. Biotic and abiotic conditions shape community structure, reducing the predictability of nonnative management. We evaluated plant community recovery after the removal of nonnative pines with an emphasis on the effect of environmental conditions on the nonnative species response. We compared clearcuts (where pine plantations were removed), pine plantations, and native communities along a precipitation gradient in Patagonia. Nonnative richness and cover were higher in clearcuts compared to native communities along nearly the entire precipitation gradient, with the exception of the harshest sites. Compared to native communities, invasion resistance was lower in clearcuts in the wetter sites. Native richness and cover were lower in clearcuts relative to native communities along the gradient. Species composition in clearcuts diverged in similarity from native communities towards the wetter sites. Plantations showed an extremely lower richness and cover compared to both clearcuts and native communities. Our study highlights that clearcutting is an ineffective strategy to manage nonnatives aimed at restoring native communities and elucidates the importance of environmental context in management approaches. Taken together, our findings reinforce the important consideration of both the biotic and abiotic context of nonnative management.Fil: Torres, Agostina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Alarcón, Pablo Angel Eduardo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Rodriguez Cabal, Mariano Alberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Nuñez, Martin Andres. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentin

The disruption of a keystone interaction erodes pollination and seed dispersal networks

Understanding the impacts of global change on ecological communities is a major challenge in modern ecology. The gain or loss of particular species and the disruption of key interactions are both consequences and drivers of global change that can lead to the disassembly of ecological networks. We examined whether the disruption of a hummingbird–mistletoe–marsupial mutualism by the invasion of non-native species can have cascading effects on both pollination and seed dispersal networks in the temperate forest of Patagonia, Argentina. We focused on network motifs, subnetworks composed of a small number of species exhibiting particular patterns of interaction, to examine the structure and diversity of mutualistic networks. We found that the hummingbird–mistletoe–marsupial mutualism plays a critical role in the community by increasing the complexity of pollination and seed dispersal networks through supporting a high diversity of interactions. Moreover, we found that the disruption of this tripartite mutualism by non-native ungulates resulted in diverse indirect effects that led to less complex pollination and seed dispersal networks. Our results demonstrate that the gains and losses of particular species and the alteration of key interactions can lead to cascading effects in the community through the disassembly of mutualistic networks.Fil: Vitali, Agustín. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Sasal, Yamila. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Vazquez, Diego P.. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mendoza. Instituto Argentino de Investigaciones de las Zonas Áridas. Provincia de Mendoza. Instituto Argentino de Investigaciones de las Zonas Áridas. Universidad Nacional de Cuyo. Instituto Argentino de Investigaciones de las Zonas Áridas; ArgentinaFil: Miguel, María Florencia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mendoza. Instituto Argentino de Investigaciones de las Zonas Áridas. Provincia de Mendoza. Instituto Argentino de Investigaciones de las Zonas Áridas. Universidad Nacional de Cuyo. Instituto Argentino de Investigaciones de las Zonas Áridas; ArgentinaFil: Rodriguez Cabal, Mariano Alberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina. University Of Vermont. Rubenstein School.; Estados Unido

The interplay between propagule pressure, seed predation and ectomycorrhizal fungi in plant invasion

There are many hypotheses aiming to explain invasion success, but evaluating individual hypotheses in isolation may hinder our ability to understand why some species invade and others fail. Here we evaluate the interaction between propagule pressure, seed predation and missed mutualism in the invasion success of the pine, Pinus ponderosa. We evaluated the independent and interactive effects of propagule pressure and seed predation at increasing distances from a pine plantation. Additionally, because pines are obligate mutualists with ectomycorrhizal fungi (EMF) and pine invasions fail in the absence of their EMF symbionts, we evaluated EMF availability through a growth chamber bioassay. In this bioassay we measured root colonization by EMF with soil samples collected from the different distances from the plantation. We found that propagule pressure overwhelms seed predation only at the edge of the pine plantation, while seed predation overcomes propagule pressure at 25 m and further distances from the plantation. We also found that EMF root colonization decreases with distance from the plantation. However, pine roots were colonized up to 200 m from the plantation, suggesting that EMF may not be hindering invasion, at least not on the scale of this experiment. Taken together our results demonstrate that seed predation may be limiting the invasion of P. ponderosa in the study region as propagule pressure only overcomes seed predation at the plantation edge. Here we provide evidence of how strong biotic resistance can suppress an invasion, regardless of the variation in propagule pressure and the availability of mutualists.Fil: Moyano, Jaime. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Chiuffo, Mariana Cecilia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Policelli, Nahuel. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Nuñez, Martin Andres. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Rodriguez Cabal, Mariano Alberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentin

Diaspore traits specialized to animal adhesion and sea current dispersal are positively associated with the naturalization of European plants across the world

Understanding what drives non-native species naturalization (the establishment of a self-sustainable population outside its native range) is a central question in invasion science. Plant capacity for long-distance dispersal (LDD) is likely to influence the spread and naturalization of non-native species differently according to their introduction pathways. These pathways include intentional introductions (for economic use, e.g. for agriculture), unintentional introductions (e.g. seed contaminants), plant dispersal via human infrastructures (e.g. roads) and plant spread from an adjacent region where the species was previously introduced. We tested the relationship between sets of LDD traits (syndromes) of 10 308 European plant species and their global naturalization incidence (i.e. whether a species has become naturalized or not) and extent (i.e. the number of regions where a species has become naturalized) using the most comprehensive database of naturalized plants worldwide (GloNAF). Diaspore traits allowed the identification of four traditional LDD syndromes, namely those with specializations for dispersal by: wind (anemochorous), animal ingestion (endozoochorous), attached to animals (epizoochorous) and sea currents (thalassochorous). These evolutionary specializations have been historically interpreted by biologists even though actual dispersal is not always related to diaspore syndromes. We found that while epizoochorous and thalassochorous traits are positively associated with global plant naturalization incidence, anemochorous and endozoochorous traits show a negative relationship. Species' residence time outside their native range, their economic use and presence of epizoochorous traits (such as hooks, hairs and adhesive substances) are positively associated with global naturalization extent. Furthermore, we found that plant economic use reduces the influence of LDD syndromes on the naturalization incidence of intentionally introduced plants. While the success of non-native plants is influenced by a broad array of species- and context-specific factors, LDD syndromes play an important role in this context depending on the economic use of plants.Fil: Moyano, Jaime. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Essl, Franz. Universidad de Viena; AustriaFil: Heleno, Ruben. Universidad de Coimbra; PortugalFil: Vargas, Pablo. Consejo Superior de Investigaciones Científicas. Real Jardín Botánico; EspañaFil: Nuñez, Martin Andres. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina. University Of Houston; Estados UnidosFil: Rodriguez Cabal, Mariano Alberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina. University of Vermont; Estados Unido

Node-by-node disassembly of a mutualistic interaction web driven by species introductions

Interaction webs summarize the diverse interactions among species in communities. The addition or loss of particular species and the alteration of key interactions can lead to the disassembly of the entire interaction web, although the nontrophic effects of species loss on interaction webs are poorly understood. We took advantage of ongoing invasions by a suite of exotic species to examine their impact in terms of the disassembly of an interaction web in Patagonia, Argentina. We found that the reduction of one species (a host of a keystone mistletoe species) resulted in diverse indirect effects that led to the disassembly of an interaction web through the loss of the mistletoe, two key seed-dispersers (a marsupial and a bird), and a pollinator (hummingbird). Our results demonstrate that the gains and losses of species are both consequences and drivers of global change that can lead to underappreciated cascading coextinctions through the disruption of mutualisms.Fil: Rodriguez Cabal, Mariano Alberto. University Of Tennessee; Estados Unidos. University Of British Columbia; CanadáFil: Barrios Garcia Moar, Maria Noelia. University Of Tennessee; Estados Unidos. University Of British Columbia; CanadáFil: Amico, Guillermo Cesar. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Patagonia Norte. Instituto de Investigación en Biodiversidad y Medioambiente; ArgentinaFil: Aizen, Marcelo Adrian. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Patagonia Norte. Instituto de Investigación en Biodiversidad y Medioambiente; ArgentinaFil: Sanders, Nathan J.. University Of Tennessee; Estados Unidos. Natural History Museun Of Copenhague; Dinamarc

Multiple plant traits shape the genetic basis of herbivore community assembly

1. Community genetics research has posited a genetic basis to the assembly of ecological communities. For arthropod herbivores in particular, there is strong support that genetic variation in host plants is a key factor shaping their diversity and composition. However, the specific plant phenotypes underlying herbivore responses remain poorly explored for most systems. 2. We address this knowledge gap by examining the influence of both genetic and phenotypic variation in a dominant host-plant species, Salix hookeriana, on its associated arthropod herbivore community in a common garden experiment. Specifically, we surveyed herbivore responses among five different arthropod feeding guilds to 26 distinct S. hookeriana genotypes. Moreover, we quantified the heritability of a suite of plant traits that determine leaf quality (e.g. phenolic compounds, trichomes, specific leaf area, C : N) and whole-plant architecture, to identify which traits best accounted for herbivore community responses to S. hookeriana genotype. 3. We found that total herbivore abundance and community composition differed considerably among S. hookeriana genotypes, with strong and independent responses of several species and feeding guilds driving these patterns. We also found that leaf phenolic chemistry displayed extensive heritable variation, whereas leaf physiology and plant architecture tended to be less heritable. Of these traits, herbivore responses were primarily associated with leaf phenolics and plant architecture; however, different herbivore species and feeding guilds were associated with different sets of traits. Despite our thorough trait survey, plant genotype remained a significant predictor of herbivore responses in most trait association analyses, suggesting that unmeasured host-plant characteristics and/or interspecific interactions were also contributing factors. 4. Taken together, our results support that the genetic basis of herbivore community assembly occurs through a suite of plant traits for different herbivore species and feeding guilds. Still, identifying these phenotypic mechanisms requires measuring a broad range of plant traits and likely further consideration of how these traits affect interspecific interactions.Fil: Barbour, Matthew A.. University Of British Columbia; CanadáFil: Rodriguez Cabal, Mariano Alberto. University Of British Columbia; Canadá. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Patagonia Norte. Instituto de Investigación en Biodiversidad y Medioambiente; Argentina. Universidad Nacional del Comahue; ArgentinaFil: Wu, Elizabeth T.. Humboldt State University; Estados UnidosFil: Julkunen Tiitto, Riitta. University of Eastern Finland; FinlandiaFil: Ritland, Carol E.. University Of British Columbia; CanadáFil: Miscampbell, Allyson E.. University Of British Columbia; CanadáFil: Jules, Erik S.. Humboldt State University; Estados UnidosFil: Crutsinger, Gregory M.. University Of British Columbia; Canad