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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; Argentina Fil: 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; Argentina Fil: 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; Argentina 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 Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina Fil: 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; Argentina

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; Argentina 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 Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina Fil: 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

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 los estudios 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.

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.

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; Argentina Fil: Lacey, Eileen Anne. University of California at Berkeley; Estados Unidos 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 Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina

Invasive species are a major threat to native communities and ecosystems worldwide. One factor frequently invoked to explain the invasiveness of exotic species is their release in the new habitat from control by natural enemies (enemy-release hypothesis). More recently, interactions between exotic species have been proposed as a potential mechanism to facilitate invasions (invasional melt- down hypothesis). We studied the effects of intro- duced deer on native plant communities and exotic plant species on an island in Patagonia, Argentina using five 400 m 2 exclosures paired with control areas in an Austrocedrus chilensis native forest stand. We hypothesized that introduced deer modify native understory composition and abundance and facilitate invasion of introduced tree species that have been widely planted in the region. After 4 years of deer exclusion, native Austrocedrus and exotic Pseudots- uga menziesii tree sapling abundances are not differ- ent inside and outside exclosures. However, deer browsing has strongly inhibited growth of native tree saplings (relative height growth is 77% lower with deer present), while exotic tree sapling growth is less affected (relative height growth is 3.3% lower). Deer significantly change abundance and composition of native understory plants. Cover of native plants in exclosures increased while cover in controls remained constant. Understory composition in exclosures after only 4 years differs greatly from that in controls, mainly owing to the abundance of highly-browsed native species. This study shows that introduced deer can aid the invasion of non-native tree species through negatively affecting native plant species.

Natural disturbances such as fires, windstorms, floods, and herbivory often act on plant communities, affecting their structure and the abundance and composition of their species. Most research has focused on the effects of single disturbances on plant communities whereas the synergistic effects of several disturbances have received less attention. In this study, we evaluated how timing and severity of tree mortality modified plant use by introduced deer and early post-mortality successional trajectories in northern Patagonian conifer forests. We sampled understory composition and deer use in Austrocedrus chilensis (cipres de la cordillera) forest stands undergoing varying timing and severity of forest mortality as reconstructed using dendroecological techniques. In addition, we evaluated the effect of fallen logs on plant composition and deer use of plants by monitoring areas of massive dieback where fallen logs had been removed for fire hazard reduction, and nearby control areas not subjected to such removal. Stepwise regression analyses showed that history and severity of tree mortality strongly influence plant composition and deer use of plants. For deer use (with pellet counts and browsing index as response variables), results showed a positive relationship with degree of stand mortality and a negative relationship with cover of fallen logs. Similarly, cover of unpalatable shrub species was explained by canopy mortality history, whereas cover of palatable shrub species was positively associated with severity of canopy mortality. In areas where fallen logs had been removed, pellet counts were six times higher than those in control areas. Though total shrub species cover was similar between log removal and control areas, proportion of unpalatable shrubs increased in areas where fallen logs had been removed. In conclusion, deer use of plants was strongly limited by tall fallen logs, allowing palatable species to establish and grow. Fallen log removal accelerated deer entrance and changed understory composition toward more browse-resistant and unpalatable species. These results underscore the importance of considering the dynamics (timing, severity, and extent) of fallen woody debris influencing understory herbivory and post-disturbance succession. In addition, experimental results underpin the importance of maintaining snags and large woody debris in disturbed landscapes where salvage logging is a routine procedure.

How interactions between exotic species affect invasion impact is a fundamental issue on both theoretical and applied grounds. Exotics can facilitate establishment and invasion of other exotics (invasional meltdown) or they can restrict them by re-establishing natural population control (as predicted by the enemy- release hypothesis). We studied forest invasion on an Argentinean island where 43 species of Pinaceae, including 60% of the world's recorded invasive Pinaceae, were introduced c. 1920 but where few species are colonizing pristine areas. In this area two species of Palearctic deer, natural enemies of most Pinaceae, were introduced 80 years ago. Expecting deer to help to control the exotics, we conducted a cafeteria experiment to assess deer preferences among the two dominant native species (a conifer, Austrocedrus chilensis, and a broadleaf, Nothofagus dombeyi) and two widely introduced exotic tree species (Pseudotsuga menziesii and Pinus ponderosa). Deer browsed much more intensively on native species than on exotic conifers, in terms of number of individuals attacked and degree of browsing. Deer preference for natives could potentially facilitate invasion by exotic pines. However, we hypothesize that the low rates of invasion currently observed can result at least partly from high densities of exotic deer, which, despite their preference for natives, can prevent establishment of both native and exotic trees. Other factors, not mutually exclusive, could produce the observed pattern. Our results underscore the difficulty of predicting how one introduced species will effect impact of another one.

Interactions between exotic plants and animals can play a major role in determining success or failure of plant introductions. Seed predation has been seen as important in explaining biotic resistance to plant invasion, but this hypothesis has rarely been tested. We studied seed predation on exotic forest plants on an island in Patagonia, Argentina where 43 pine species, including 60% of the world’s known invasive Pinaceae, were introduced ca. 80 years ago, but where exotics attain relatively high densities only near the original plantings. To test if seed predation limits exotic conifer establishment in this area, we compared seed predation in areas close to plantations (colonized by exotics) and far from them (not invaded). Seeds of exotics were preferred over seeds of native species, possibly because exotic seeds are bigger. Predation was more intense in areas far from plantations than in areas close to them, substantially reducing the chances of exotic seed establishment. Using automatic cameras, we found that both rodents and birds preyed on exotic seeds. This study suggests that native seed predators can be an important component of biological resistance to plant invasion.

Isla Victoria (Nahuel Huapi National Park, Argentina), a large island dominated by native Nothofagus and Austrocedrus forest, has old plantations of many introduced tree species, some of which are famed invaders of native ecosystems elsewhere. There are also large populations of introduced deer and shrubs that may interact in a complex way with the introduced trees, as well as a recently arrived population of wild boar. Long-standing concern that the introduced trees will invade and transform native forest may be unwarranted, as there is little evidence of progressive invasion, even close to the plantations, despite over 50 years of opportunity. Introduced and native shrubs allow scattered introduced trees to achieve substantial size in abandoned pastures, but in almost all areas neither the trees nor the shrubs appear to be spreading beyond these sites. These shrub communities may be stable rather than successional, but the technology for restoring them to native forest is uncertain and probably currently impractical. Any attempt to remove the exotic tree seedlings and saplings from native forest would probably create the very conditions that would favor colonization by exotic plants rather than native trees, while simply clear-cutting the plantations would be unlikely to lead to regeneration of Nothofagus or Austrocedrus. The key to maintaining native forest is preventing catastrophic fire, as several introduced trees and shrubs would be favored over native dominant trees in recolonization. Deer undoubtedly interact with both native and introduced trees and shrubs, but their net effect on native forest is not yet clear, and specific management of deer beyond the current hunting by staff is unwarranted, at least if preventing tree invasion is the goal. The steep terrain and shallow soil make the recently arrived boar a grave threat to the native forest. Eradication is probably feasible and should be attempted quickly.

In the forests of Austrocedrus chilensis (D.Don) Florin et Boutleje (Cupressaceae) it is frequent to observe numerous mutilated and dwarfed saplings which have been browsed by domestic cattle and exotic deer. One way to assess the effects of browsing on plant growth is by simulating this process through defoliation. This study includes results from four years of artificial browsing at different intensities on A. chilensissaplings. Our objective was to evaluate the possible compensatory growth by A. chilensis. To that end we measured the effects of simulated browsing on the growth of saplings kept under two different experimental conditions, with and without water stress. Treatments were applied at the end of winter of 1993 and were repeated on 1994, 1995 and 1996. Treatments consisted in pruning and cutting. Neither cutting effects nor factors interactions were found. Pruning resulted in a reduction of saplings biomass of 44% and was only significant at the more intensive level. Watering produced saplings with five times more biomass, but the general response of A. chilensis to simulated browsing was the lack of compensation regardless the intensity of pruning or the quality of site.

Austrocedrus chilensis (D. Don) et Boutleje (Cupressaceae) is an endemic conifer of northern Patagonia, Argentina and south-central Chile, which has been affected by browsing by livestock and introduced deer (especially red deer). In northern Patagonia, we studied the effects of livestock and deer on the densities and heights of Austrocedrus saplings and on the structure and floristic composition of the vegetation in which this species occurs under a variety of different browsing regimes. Twenty-seven stands were sampled within the `Compact' forest type (i.e., dense and pure) of Austrocedrus in both national reserves and on unprotected properties. Principal components analysis (PCA) indicates that greater abundance of Austrocedrus saplings is associated with lower stand basal areas as expected for this relatively shade-intolerant species. Although browsing does not directly reduce the abundance of saplings, it has an important influence in terms of reducing height growth and producing deformed stems. Multiple correspondence analysis (MCA) indicates that saplings between 0.5 and 1 m in height were more frequently browsed than other height classes; saplings shorter than 0.3 m in height were not greatly influenced by browsing. Saplings of intermediate heights (0.5–1 m) were scarce at sites lacking any current browsing, but this may reflect release of saplings from earlier browsing and subsequent inhibition of continued recruitment into intermediate height classes. High browsing indexes occur beneath canopy gaps, which are also sites of higher sapling densities. In the present study, sites characterized by heavy browsing also have a high cover of spiny shrubs (Berberis spp.) and other unpalatable woody species (e.g., Lomatia hirsuta). Where the preferred browse species, Schinus patagonicus, is abundant, Austrocedrus is less influenced by browsing. However, as browsing reduces the abundance of S. patagonicus, Austrocedrus becomes a preferred species. Austrocedrus stands that are in a phase of fine-scale gap-phase regeneration are the most susceptible to inhibition of tree regeneration by livestock and deer browsing. In such stands, long-term persistence of a forest cover requires protection from large browsing animals.

ISSN 1948-6596 news and update Warren, M., Robertson, M. P., & Greeff, J. M. (2010) A comparative approach to understanding factors limiting abundance patterns and distributions in a fig tree-fig wasp mutualism. Ecography, 33, Whittaker, R. J., Willis, K. J., & Field, R. (2001) Scale and species richness: towards a general, hierarchical theory of species diversity. Journal of Biogeogra- phy, 28, 453-470. Wilson, R. J., Davies, Z. G., & Thomas, C. D. (2010) Link- ing habitat use to range expansion rates in frag- mented landscapes: a metapopulation ap- proach. Ecography, 33, 73-82. Tim Newbold UNEP World Conservation Monitoring Centre, Cambridge, UK e-mail: Tim.Newbold@unep-wcmc.org http://www.unep-wcmc.org/aboutWCMC/dsp/ dynamicstaff.aspx?id=169T journal compilation © 2010 The International Biogeography Society — frontiers of biogeography 2.2, 2010