Status and challenges for conservation of small mammal assemblages in South America.

South America spans about 44° latitude, covers almost 18 million km(2) , and is second only to Africa in continental mammal species richness. In spite of this richness, research on the status of this fauna and on the nature and magnitude of contemporary threats remains limited. Distilling threats to this diverse fauna at a continental scale is challenging, in part because of the limited availability of rigorous studies. Recognizing this constraint, we summarize key threats to small mammals in South America, emphasizing the roles of habitat loss and degradation, direct persecution, and the increasing threat of climate change. We focus on three regional 'case studies': the tropical Andes, Amazonia and adjacent lowland regions, and the southern temperate region. We close with a brief summary of recent findings at our long-term research site in north-central Chile as they pertain to projected threats to this fauna. Habitat alteration is a pervasive threat that has been magnified by market forces and globalization (e.g. extensive agricultural development in Amazonia), and threatens increasing numbers of populations and species. Climate change poses even greater threats, from changes in rainfall and runoff regimes and resulting changes in vegetative structure and composition to secondary influences on fire dynamics. It is likely that many changes have yet to be recognized, but existing threats suggest that the future may bring dramatic changes in the distribution of many mammal taxa, although it is not clear if key habitat elements (vegetation) will respond as rapidly as climatic factors, leading to substantial uncertainty. Climate change is likely to result in 'winners' and 'losers' but available information precludes detailed assessment of which species are likely to fall into which category. In the absence of long-term monitoring and applied research to characterize these threats more accurately, and to develop strategies to reduce their impacts, managers already are being faced with daunting challenges. As the line between 'pure' and 'applied' research blurs in the face of converging interests of scientists and society we hope that solutions to these critical issues will be incorporated in addressing anticipated conservation crises.

Extreme climatic events change the dynamics and invasibility of semi-arid annual plant communities.

Extreme climatic events represent disturbances that change the availability of resources. We studied their effects on annual plant assemblages in a semi-arid ecosystem in north-central Chile. We analysed 130 years of precipitation data using generalised extreme-value distribution to determine extreme events, and multivariate techniques to analyse 20 years of plant cover data of 34 native and 11 exotic species. Extreme drought resets the dynamics of the system and renders it susceptible to invasion. On the other hand, by favouring native annuals, moderately wet events change species composition and allow the community to be resilient to extreme drought. The probability of extreme drought has doubled over the last 50 years. Therefore, investigations on the interaction of climate change and biological invasions are relevant to determine the potential for future effects on the dynamics of semi-arid annual plant communities.

Bottom-up control of consumers leads to top-down indirect facilitation of invasive annual herbs in semiarid Chile.

The abundance of exotic plants is thought to be limited by competition with resident species (including plants and generalist herbivores). In contrast, observations in semiarid Chile suggest that a native generalist rodent, the degu (Octodon degus), may be facilitating the expansion of exotic annual plants. We tested this hypothesis with a 20-year data set from a World Biosphere Reserve in mediterranean Chile. In this semiarid environment, rainfall varies annually and dramatically influences cover by both native and exotic annual plants; degu population density affects the composition and cover of exotic and native annual plants. In low-rainfall years, cover of both native and exotic herbs is extremely low. Higher levels of precipitation result in proportional increases in cover of all annual plants (exotic and native species), leading in turn to increases in degu population densities, at which point they impact native herbs in proportion to their greater cover, indirectly favoring the expansion of exotic plants. We propose that bottom-up control of consumers at our site results in top-down indirect facilitation of invasive annual herbs, and that this pattern may be general to other semiarid ecosystems.

Effects of more frequent and prolonged El Niño events on life-history parameters of the degu, a long-lived and slow-reproducing rodent.

Global climate change (GCC) can have profound effects on species whose ecology is governed primarily by climatic factors. The ecology of small mammals inhabiting semiarid Chile is strongly affected by the El Niño Southern Oscillation (ENSO). During La Niña events in this area, dry conditions prevail and species may disappear from the thorn-scrub habitat. Conversely, El Niño events bring high rainfall, and associated pulses of food trigger small-mammal population increases. We used capture-mark-recapture to study responses of the degu (Octodon degus), a dominant small mammal, to variation in rainfall over 18 years. In response to a recent trend toward wetter conditions, degus reached record-high densities and maintained more stable numbers in the area. Underlying mechanisms involved variation in adult survival, juvenile persistence, and fecundity linked to rainfall changes during consecutive years (i.e., rainfall phases). During prolonged droughts, degus had low survival and produced fewer offspring, with low persistence. Following high rainfall, these parameters reversed; consecutive wet years resulted in further increases. Weak declines in fecundity and adult survival and high persistence of juveniles explained delayed responses to deteriorating conditions in initial dry years. If GCC leads to increased frequency of El Niño events, we anticipate greater numerical dominance of degus in semiarid Chile and possible range expansion. Furthermore, degus have strong impacts on other small mammal and some plant species, are important prey species, and are agricultural pests and disease reservoirs. Hence, GCC has the potential to dramatically influence their ecology in northern Chile and to have cascading effects on other components of this system.

Population dynamics of two sympatric rodents in a variable environment: rainfall, resource availability, and predation.

Precipitation plays an important role in the dynamics of species found in arid and semiarid environments. However, population fluctuations generally are driven by a combination of multiple factors whose relative contribution may vary through time and among species. We monitored fluctuations of species in three trophic levels for >17 years at a semiarid community in north-central Chile. The region is strongly affected by the El Niño Southern Oscillation, resulting in high variation in rainfall that triggers dramatic changes in food resource availability, with strong effects on upper trophic levels. We focused our analyses on the role played by endogenous and exogenous (climatic) factors on the dynamics of two important rodent species in the community, Octodon degus and Phyllotis darwini. We documented population fluctuations of several orders of magnitude in response to wet and dry episodes of different strength and duration. P. darwini reached similar maximum densities, regardless of the duration of high-rainfall events, whereas O. degus showed additive effects of multiple wet years. Time series diagnostic tools revealed oscillations with a 5-year periodicity in rainfall, which may be the cause of the same periodicity and a weak second-order signal observed in the rodent dynamics. However, the dynamics of both rodent species were dominated by strong first-order processes, suggesting an important role of direct density dependence. Intraspecific competition, expressed as the ratio of rodent density/rainfall (or food resources) explained more than two-thirds of the variation in the population rate of change, whereas less than one-third was explained by lagged rainfall (or food resources). We detected no significant effects of predation. Our results contribute to a growing number of examples of dynamics governed by the combined effect of density dependence and climatic forcing. They also reveal strong bottom-up regulation that may be common in other arid environments.

El Niño effects on soil seed bank dynamics in north-central Chile.

The soil seed bank was monitored in four 75x75 m plots over 6 years (1990-1995) in an arid thorn scrub community in north-central Chile. Sixty-six species were identified. Total seed densities ranged from 2,000 to 42,000/m(2). Average mass of shrub seeds was significantly greater than that of other growth-forms. Between 70 and 90% of the seeds were less than 1 mg, with those in the 0.51-1.00 mg size class being most numerous. Seed densities were highly variable between years as well as within years, but were also closely associated with plant cover patterns and rainfall regime. Higher seed densities were found in wet years, and in samples taken in early summer and early autumn (i.e., after seed set); the lowest seed densities were in late winter (i.e., after annual plant germination). The annual plant species with the highest cover were also the most abundant in the soil seed bank and exhibited the largest seed density fluctuations. In general, seed densities were 5- to 10-fold higher during the 1991-1992 El Niño/southern oscillation (ENSO) years than non-ENSO years, showing the importance of this phenomenon for seed bank replenishment in the arid region of Chile.

Spatial distribution of soil nutrients and ephemeral plants underneath and outside the canopy of Porlieria chilensis shrubs (Zygophyllaceae) in arid coastal Chile.

Soil nutrients and density and biomass of annual plants underneath and outside the canopy of Porlieria chilensis shrubs were measured at the end of the growing season in a protected arid coastal site in Chile. Levels of soil nitrogen, phosphorus and organic matter were significantly higher underneath than outside the canopies of shrubs. Almost 4 times as many plants occurred outside than underneath shrubs, but no significant differences in total aboveground biomass were found. Several species had higher densities and/or biomass outside rather than underneath shrubs, whereas others showed the oppsite trend. Species richness was lower underneath P. chilensis canopy. The spatial microdistribution of ephemeral species may be explained by differential water and nutrient requirements. Comparison of the patterns observed in our protected site versus surrounding unprotected areas supports the generalization that man, by removing shrubs and trees, has changed a previous heterogeneous spatial distribution of nutrients to a more homogenous one.