Factors facilitating and hindering the invasion of exotic conifer in Isla Victoria

Las especies invasoras constituyen una seria amenaza a la biodiversidad y producen cambios a gran escala en las comunidades nativas y alterando los procesos de los ecosistemas. En la actualidad, los bosques Andino-Patagónicos exhiben una diversidad amplia de especies introducidas, tanto terrestres como acuáticas. Muchas de estas especies poseen un potencial invasor elevado, al tiempo que constituyen la base de diversas actividades de importancia económica para la región, como, por ejemplo, los cotos de caza de ciervos y jabalíes,la pesca deportiva de salmónidos y las forestaciones con coníferas introducidas. La situación descripta para la Patagonia brinda un escenario ideal para probar y/o generar nuevas hipótesis sobre la ecología de las invasiones. En un campo más práctico, determinar el impacto y conocer las relaciones entre las especies nativas y exóticas permitirán establecer prioridades en las acciones de manejo, así como también mitigar los efectos no deseados o inesperados como consecuencia de las tareas de erradicación o control. Presentamos aquí una síntesis de losestudios propios llevados a cabo para entender el patrón de invasión de pináceas encontrado en un bosque nativo de Nothofagus/Austrocedrus en Isla Victoria, Parque Nacional Nahuel Huapi. Entre los mecanismos estudiados encontramos que algunos de ellos la facilitan (e.g., ciervos introducidos), mientras que otros la retrasan (depredación de semillas, baja presencia de micorrizas) o no tienen un efecto sustancial en la invasión (presión de propágulos). Estos resultados y otros antecedentes disponibles a la fecha de ésta y otras regiones del mundo, permiten sugerir estrategias de manejo para impedir o retrasar una invasión masiva de pináceas, las cuales deberían posiblemente estar enfocadas a evitar nuevas introducciones, reducir las poblaciones de los herbívoros exóticos y mantener los ecosistemas nativos con bajos niveles de impacto antrópico. Al mismo tiempo, consideramos importante emprender estudios específicamente diseñados para analizar los efectos de las tareas de erradicación y/o control sugeridas.Invasive species constitute a serious threat to biodiversity, producing wide-scale changes in native communities and altering ecosystem processes. The Andean-Patagonian forests currently exhibit a wide diversity of terrestrial as well as aquatic introduced species. Many of these introduced species possess a high invasive potential, while at the same time have economical relevance to the region, such as sport fishing and hunting, and pine plantations. The described situation for Patagonia offers an ideal scenario to test and generate new hypotheses about the ecology of tree invasions. In a more practical domain, determining the impact and knowing the relationship between native and exotic species will permit the establishment of priorities in management actions, as well as also avoiding or mitigating the undesirable or unexpected effects that arise from eradication or control tasks. Here, we present a synthesis of our own studies carried out to understand the factors driving the pattern of pine invasion in the native Nothofagus/Austrocedrus forests on Isla Victoria, Nahuel Huapi National Park. Among studied factors, we found that some of them facilitate the invasion (e.g, introduced herbivores), while others can delay (e.g, low presence of mycorrhizae fungi, seed predation) or have neutral effects on the invasions (e.g., propagule pressure). These results and others currently available from this region and elsewhere allow us to suggest management actions to prevent or hinder a major pine invasion, which should be focused on avoiding new introductions, reducing the abundance of introduced large mammals and preserve native ecosystems with low levels of anthropic disturbances. At the same time, we consider important to start well designed studies to analyze the effects of these management actions.Fil: Relva, Maria Andrea. 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 Reg.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 Reg.universidad Bariloche. Instituto de Investigaciones En Biodiversidad y Medioambiente; Argentin

Non-native conifer plantations in Patagonia: risks of planting without proper management

En la última década se realizaron en la región patagónica una gran cantidad de investigacionesevaluando el impacto ecológico de las plantaciones y analizado los modos de producción con el fin de aumentarla cosecha y reducir su impacto. Si bien las plantaciones brindan beneficios para las economías regionales, a lafecha no está clara la magnitud de esta contribución. Por otro lado, los problemas ambientales que producen lasplantaciones son varios; en particular, las invasiones de ecosistemas aledaños es uno de los más importantesen el largo plazo. Otro problema relacionado a las plantaciones, aunque incipiente aún en la región, es elaumento de los incendios forestales. Las plantaciones acumulan gran cantidad de combustible seco e inflamable,especialmente cuando no son manejadas. Además, muchas especies o variedades de pinaceas están adaptadasal fuego, por lo que no sólo podrían cambiar el régimen y el tipo de incendios, sino también retroalimentarsepositivamente, ocasionando altos costos sociales, económicos y ecológicos. Consideramos que estos problemaspodrían ser minimizados en gran parte con la aplicación de buenas prácticas forestalesIn the last decade in Patagonia, a significant number of research studies evaluated the environmental impact of plantations and analyzed the modes of production in order to increase the yield and reduce its impact. While plantations provide benefits for regional economies, at the present the extent of this contribution is unclear. Furthermore, there are several environmental problems produced by plantations, with the invasions of adjacent ecosystems being one of the most damaging impacts in the long term. Another problem related to the plantations, although still incipient in the region, is the increment of forest wildfires. Plantations accumulate large amounts of dry and flammable fuel, particularly when they are not properly managed. Also many species or varieties of Pinaceae are adapted to fire, so they could not only change the regime and type of fires, but also display positive feedbacks causing high social, economic and ecological costs. We believe that these problems could be minimized to a great extent with the application of good forestry practices.Fil: Raffaele, Estela. 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: Nuñez, Martin Andres. 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: Relva, Maria Andrea. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Patagonia Norte. Instituto de Investigación En Biodiversidad y Medioambiente; Argentina. Universidad Nacional del Comahue; Argentin

Sheep feeding preference as a tool to control pine invasion in Patagonia: influence of foliar toughness, terpenoids and resin content

Herbivores modulate the structure and composition of plant communities, including plant invasions. This is conditioned by plant palatability which can be reduced by its chemical or physical traits. The effects that ungulates browsing has on pine invasions are variable and the empirical evidence on the causes of this variability is scarce. We experimentally explored how sheep browsing preference varies between seedlings of pine species with different invasiveness; Pinus contorta (high invasiveness), P. ponderosa (medium invasiveness), P. radiata (medium invasiveness) and P. jeffreyi (low invasiveness). Secondly, we quantified anti-herbivory chemical compounds and physical traits of these species and related them with sheep preference observed. The browsing incidence of P. contorta was 68%, P. ponderosa 58%, P. radiata 29%, and P. jeffreyi 84%. Among anti-herbivory traits analyzed, α-pinene concentration had a negative effect on the probability of a terminal bud being browsed and on browsing intensity. Meanwhile, foliar toughness was negatively related to browsing intensity and water concentration was positively related to browsing intensity. Also, the most invasive species, P. contorta, was highly damaged. Thus, sheep herbivory could be slowing pine invasion rate; suggesting that could be considered a tool to control early invasions, especially for this particular species.Fil: Zamora Nasca, Lucía Belén. 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: Dimarco, Romina Daniela. Instituto Nacional de Tecnología Agropecuaria. Centro Regional Patagonia Norte. Estación Experimental Agropecuaria San Carlos de Bariloche. Instituto de Investigaciones Forestales y Agropecuarias Bariloche. - Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones Forestales y Agropecuarias Bariloche; ArgentinaFil: Nassini, Daniela. Comisión Nacional de Energía Atómica. Gerencia Complejo Tecnológico Pilcaniyeu; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; ArgentinaFil: Alvear, Pablo 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: Mayoral, Ariel Alejandro. 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: Relva, Maria Andrea. 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

Litter and soil properties are not altered by invasive deer browsing in northern Patagonia forests.

It is largely accepted that large mammalian herbivores can indirectly influence ecosystem properties by changing the quantity and quality of litter inputs to soil through selective browsing on plant communities. However, idiosyncratic effects (positive, neutral and negative) have been found for different vegetation types, herbivory and soil properties. In this study we tested whether browsing by exotic deer introduced into Patagonian forests 90 years ago alters physical, chemical and biological soil properties and litter quantity and quality. As in many terrestrial ecosystems, N is the main nutrient that limits plant growth in Patagonia. Consequently, any disturbance that alters the N cycle, such as changes in the type or intensity of herbivory, is expected to affect these forest ecosystems. We compared soil and total litter from inside and outside five 7-yr old exclosures located on Isla Victoria, Parque Nacional Nahuel Huapi, Argentina. Despite introduced deer has significantly affected the composition, abundance and structure of the understory vegetation, we found no differences between browsed and unbrowsed areas in the physical (bulk density, moisture), chemical (pH, base cations, organic C and total N) and biological (potential microbial respiration and net N mineralization) soil properties. This could be attributable to the high capacity of volcanic soils to stabilize organic matter, buffering disturbance-induced changes. However, the quantity and quality (C, N and C/N ratio) of total litter were also not different between browsed and unbrowsed areas. Although non-significant differences were found between treatments in both compartments, litter and soil, most variables showed a slight trend toward higher values in unbrowsed areas. This suggests that 7 years of browsing exclusion would be not enough to detect changes induced by browsing, particularly in highly stable volcanic soils.Fil: Relva, Maria Andrea. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Patagonia Norte. Instituto de Investigación En Biodiversidad y Medioambiente; Argentina. Universidad Nacional del Comahue. Centro Regional Universitario Bariloche. Laboratorio de Ecotono; ArgentinaFil: Castán, Elisa. Universidad Nacional del Comahue. Centro Regional Universitario Bariloche; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Patagonia Norte. Instituto de Investigación en Biodiversidad y Medioambiente; ArgentinaFil: Mazzarino, Maria Julia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Patagonia Norte. Instituto de Investigación En Biodiversidad y Medioambiente; Argentina. Universidad Nacional del Comahue. Centro Regional Universitario Bariloche; Argentin

Impacts of domestic cattle on forest and woody ecosystems in southern South America

There is a long lasting debate on the effects of domestic cattle grazing on natural ecosystems worldwide. Cattle are generally assumed to have negative effects on forest conservation; however, several studies also report positive and neutral effects. We aimed to investigate the available evidence for positive, negative and neutral effects of cattle grazing on forest and woody ecosystems of southern South America. We conducted a peer-review literature search using the ISI Web of Knowledge and Scopus databases to identify studies dealing with cattle impacts for nature conservation. We compiled a database of 211 cases from 126 original publications. A reduced number of forest ecosystems (Patagonian forest, Chaco and Monte) concentrated ~ 85% of the reported study cases. The hierarchical cluster analysis to group cases based on cattle effects, ecological variables and ecosystems reported that negative effects (~ 66% of cases) were mostly informed for vegetation variables and mainly occur in Patagonian forest and Chaco; positive effects (~ 16%) were mostly informed for Monte (no particular variable associated), while neutral effects (~ 18%) were mostly informed for fauna-related variables and Uruguayan savanna. Our study suggests that grazing effects by cattle on southern South America forests are not homogeneous and depend on the particular forest ecosystem considered as well as on the forest attribute measured. Different cattle effects found can be partially explained by differences in grazing history and different ecosystems productivity. It is vital to improve our understanding of cattle–forest interactions to guide synergies between sustainable management and forest conservation.Fil: Mazzini, Flavia. Universidad Nacional de Jujuy. Instituto de Ecorregiones Andinas. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Salta. Instituto de Ecorregiones Andinas; ArgentinaFil: Relva, Maria Andrea. 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: Malizia, Lucio Ricardo. Universidad Nacional de Jujuy. Instituto de Ecorregiones Andinas. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Salta. Instituto de Ecorregiones Andinas; Argentin

Patterns of use and damage by exotic deer on native plant communities in northwestern Patagonia

Ungulate overabundance can strongly alter composition and structure of forest communities through selective damage independently of the herbivory history of the recipient system. In the early 1900s, red and fallow deer (Cervus elaphus and Dama dama, respectively) were introduced to northwestern Patagonia. We studied patterns of relative habitat use, damage, and species selection by introduced deer populations in three major plant communities that have the longest history of invasion in the region. We also evaluated community structure and composition characteristics as possible explanatory factors for the observed patterns of browsing, fraying, and bark-stripping. At the species level, exotic deer browsed more than expected on two evergreen species (Austrocedrus chilensis and Schinus patagonicus) and two spiny species (Colletia hystrix and Dasyphyllum diacanthoides), while one dominant shrubland tree (Lomatia hirsuta) was selected for fraying and bark-stripping. Browsing was the more widespread damage in all plant communities, while fraying and bark-stripping occurred at low incidence. Furthermore, species identity was found as the main driver of deer damage over plant community type. Finally, variation in damage and in habitat use was explained by community structure and composition characteristics. Bark-stripping and fraying are best predicted by community composition, whereas browsing and habitat use are best predicted by structural properties including understory cover, cover of fallen logs, and tree density. The patterns of damage and the association with community structure characteristics reported here provide insight for developing management strategies.Fil: Barrios Garcia Moar, Maria Noelia. University of Tennessee; 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; Argentina. Universidad Nacional del Comahue. Centro Regional Universitario Bariloche. Laboratorio de Ecotono; ArgentinaFil: Relva, Maria Andrea. 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. Universidad Nacional del Comahue. Centro Regional Universitario Bariloche. Laboratorio de Ecotono; ArgentinaFil: Kitzberger, Thomas. 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. Universidad Nacional del Comahue. Centro Regional Universitario Bariloche. Laboratorio de Ecotono; Argentin

Habitat use by colonial tuco-tucos (Ctenomys sociabilis): Specialization, variation, and sociality

Understanding habitat requirements has implications for numerous aspects of a species' biology, including where individuals live and how they behave. Specialization for mesic, resource-rich habitats known as mallines is thought to have favored group living in the colonial tuco-tuco (Ctenomys sociabilis), a subterranean rodent that is endemic to Neuquén Province in southwestern Argentina. To explore the proposed relationship between mallines and sociality in this species in greater detail, we characterized the habitats occupied by C. sociabilis at 3 locations representing the extremes of this species' geographic range. Specifically, plant composition and vegetative structure were characterized for 57 occupied burrow systems distributed across the 3 sampling localities. Our data indicate that C. sociabilis is not restricted to mallines. Although significant variation in vegetation was detected among the 3 study sites, the majority of active burrow systems surveyed at each site occurred in nonmallín habitats. In addition to providing the first species-wide survey of habitat use by C. sociabilis, our data yield new insights into the role of habitat specialization in promoting sociality in this behaviorally unusual species of ctenomyid rodent.Fil: Tammone, Mauro Nicolás. 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: Lacey, Eileen Anne. University of California at Berkeley; Estados UnidosFil: Relva, Maria Andrea. 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

Ungulates can control tree invasions: experimental evidence from nonnative conifers and sheep herbivory

Invasive conifer species are increasingly recognized as a serious problem in many parts of the world, where they are having large ecological and economic impacts. Understanding factors that trigger and can control invasion is key to management. Grazing and browsing by large herbivores have been suggested as a mechanism that may halt conifer invasions, although conflicting results have been reported (i.e. positive, negative or no effect of grazing on invasion). We believe that some of these opposing responses arise due to the absence of well-planned and replicated experiments, since current evidence is mostly observational, and for example, differences in animal densities can produce different results. Thus, in this study, we tested whether large herbivores can control invasion by nonnative conifers and whether the severity of the invasion process would be lessened by increased herbivory intensity. We evaluate experimentally herbivore damage on Pinus contorta, a highly invasive species in many countries of the Southern Hemisphere, under different sheep stocking rates in Patagonia, Argentina. We used four stocking rates, corresponding to 1, 2, 4 and 8 times the recommended sheep herbivory intensity for the study area. The response was not linear but rather presented a threshold. The greater stocking rate, the greater the browsing, higher reduction in seedling height, and decrease of survival of P. contorta. Also, the highest stocking rate damaged and killed 99% of them. This study provides evidence that large domestic herbivores can play a key role in the invasion process and, if managed properly, would provide a tool to help limit conifer invasion.Fil: Zamora Nasca, Lucía Belén. 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: Relva, Maria Andrea. 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

Not one but three: undetected invasive Alnus species in northwestern Patagonia confirmed with cpDNA and ITS sequences

Species of Alnus (alders) have become invaders in several parts of the world. Here we report the presence of three naturalized alien species: A. glutinosa, A. incana and A. rubra from several populations in nature reserves of northwestern Patagonia, an area of remarkably high biodiversity. Alnus glutinosa had been cited previously for Chile and southern Argentina, but A. incana and A. rubra are here reported for the first time. As we found morphological variation within and among the populations of these introduced species that makes their discrimination difficult, we used chloroplast (trnH-psbA) and nuclear ribosomal (ITS) DNA sequences to confirm their identifications from morphological characteristics. Results from nuclear and chloroplast sequence data confirm the morphological tentative identification of the three species and remark the utility of molecular information together with morphology for the detection of introduced species of taxonomically difficult groups. The invasive characteristics of these alien tree species are discussed in relation to the conservation of the nature reserves where they are found.Fil: Calviño, Carolina Isabel. 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: Edwards, Priscila. 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: Fernández, Martina. 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: Relva, Maria Andrea. 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: Ezcurra, 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; Argentin