Douglas A. Kelt

Extreme climatic events shape arid and semiarid ecosystems

6 Climatic changes associated with the El Nino Southern Oscillation (ENSO) can have a dramatic impact on ter- restrial ecosystems worldwide, but especially on arid and semiarid systems, where productivity is strongly lim- ited by precipitation. Nearly two decades of research, including both short-term experiments and long-term studies conducted on three continents, reveal that the initial, extraordinary increases in primary productivity percolate up through entire food webs, attenuating the relative importance of top-down control by predators, providing key resources that are stored to fuel future production, and altering disturbance regimes for months or years after ENSO conditions have passed. Moreover, the ecological changes associated with ENSO events have important implications for agroecosystems, ecosystem restoration, wildlife conservation, and the spread of disease. Here we present the main ideas and results of a recent symposium on the effects of ENSO in dry ecosystems, which was convened as part of the First Alexander von Humboldt International Conference on the El Nino Phenomenon and its Global Impact (Guayaquil, Ecuador, 16-20 May 2005).

The Ecology and Macroecology of Mammalian Home Range Area

Although many studies employ allometric relationships to demonstrate possible dependence of various traits on body mass, the relationship between home range size and body mass has been perhaps the most difficult to understand. Early studies demonstrated that carnivorous species had larger home ranges than herbivorous species of similar mass. These studies also argued that scaling relations (e.g., slopes) of the former were steeper than those of the latter and explained this in terms of the distribution of food resources, which are more uniformly distributed for most herbivores than for carnivores. In contrast to these studies, we show that scaling relations of home ranges for carnivorous mammals do not differ significantly from those of herbivorous and omnivorous species and that all three exhibit slopes that are significantly steeper than predicted on the basis of energetic requirements. We also demonstrate that home range size is constrained to fit within a polygonal constraint space bounded by lines representing energetic and/or biophysical limitations, which suggests that the log‐linear relationship between home range area and mass may not be the appropriate function to compare against the energetically predicted slopes of 0.75 or 1.0. It remains unclear, however, why the slope of the relationship between home range area and body mass, whether based on raw data or on constraint lines, always exceeds that predicted by the energetic needs hypothesis.

Thirteen Years of Shifting Top-Down and Bottom-Up Control

Abstract For 13 years, we studied the role of biotic interactions, including predation, herbivory, and interspecific competition, in a semiarid thorn scrub community in north-central Chile. Using a large-scale field manipulation, we monitored changes in small mammals, plants, and vertebrate predators. We documented important “top-down” predation effects on some small-mammal species, and small-mammal effects on some plant species resulting from experimental exclusions. However, periodic El Niño Southern Oscillation (ENSO) events caused several high rainfall episodes during this interval, resulting in large “bottom-up” increases in both plants and animals. Therefore, we suggest that instead of exclusive top-down or bottom-up control, this system undergoes shifting control, with relatively greater importance of biotic interactions in wet years and of resource limitation in dry ones. Because intervals between ENSO events are long and responses are slow, long-term studies are essential for understanding such effects in semiarid or arid systems.

Community Structure of Desert Small Mammals: Comparisons Across Four Continents

Presence/absence data for the small-mammal species at sites in seven deserts were analyzed for evidence of similarity in community structure. The deserts studied were located in North and South America (268 and 118 sites, respectively), Australia (245 sites), Israel (54 sites), and greater Eurasia (Thar, 15 sites; Turkestan, 36 sites; Gobi, 98 sites). Patterns observed in all deserts included: (1) low a diversity (2-4 species per site); (2) high P diversity (species turnover between sites); and (3) local coexistence of 20-30% of the species in the regional pool. Additionally, the number of species with which a species co-occurred increased with the number of sites at which that species occurred. Although these results suggested that some features of community structure were similar across deserts, other aspects, especially trophic structure, differed widely. Deserts in the northern hemisphere possessed more granivores, and the Turkestan Desert more folivores, than other deserts. Carnivorous small mammals were most strongly represented in Australia, and omnivores in South America, Australia, and the Thar. The structure of desert small-mammal communities is strongly influenced by historical factors; different taxonomic groups with distinctive trophic adaptations proliferate in different desert regions where they are subject to some common structuring processes of community assembly.

Assessing the Impact of Competition on Community Assembly: A Case Study using Small Mammals

Competition has been shown to be an important factor in natural systems. Yet, its effect on community structure remains poorly understood and is often disputed. Null models have been employed by researchers in recent years, but these generally suffer from a lack of power estimation and unclear distinction between statistical significance and biological importance. Models that employ an appropriate alternative hypothesis allow for power estimates and for quantitative evaluation of the effects of competition. We have developed methodologies to (1) allow for testing the significance of competition in natural communities, (2) determine the power of these estimates, and (3) produce maximum likelihood estimates of interaction parameters for these communities. Using these methodologies we show that small mammal communities at the interface between Valdivian temperate rain forest and Patagonian steppe in southern South America appear significantly and importantly structured by competition, although we cannot distinguish between resource competition, interference competition, and apparent competition. Secondarily, we show that the geographic origin of the component species (Valdivian vs. Patagonian) is relatively more important than their habitat affinities in structuring these communities.

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.

ENERGETIC CONSTRAINTS AND THE RELATIONSHIP BETWEEN BODY SIZE AND HOME RANGE AREA IN MAMMALS

Recent theoretical developments to explain the unimodal and asymmetric distribution of body sizes among species in higher taxa have yielded predictions for related demographic and life history traits. In particular, it has been predicted that there is an energetically optimal body size (M*) for terrestrial mammals at ∼100 g, and that the relationships of many biological characteristics will change slope or even sign at this point. We reanalyze a well-known data set on the relationship between home range size and body size in mammals. If the distribution of home range sizes as a function of body size is energetically constrained, then it is reasonable to assume that the lower size limit of a home range for a given body mass will also be constrained. To evaluate this hypothesis, we predicted that the relationship between minimal home range size and body size is nonlinear, and that the smallest home ranges should correspond to species in the vicinity of M*. Our data tentatively support both hypotheses but constitute a clear call for more comprehensive analyses with larger data sets.

Assembly Rules: Desert Rodent Communities Are Structured at Scales from Local to Continental

Ecologists have long argued about the patterns and processes that characterize assemblages of coexisting species. Patterns in space and time can provide clues to underlying mechanistic processes, but often in community ecology even the patterns are debated. Some of the most longstanding and acrimonious debates have concerned patterns that have been interpreted to suggest an important role of competition in the structure and dynamics of ecological communities and the evolution of species. There is little disagreement that interspecific competition can and does affect the local abundance and distribution of species. This has been repeatedly demonstrated by small-scale, manipulative experimental studies of populations in the field (see reviews by Connell 1983; Schoener 1983; Gurevitch et al. 1992). More controversial have been comparative geographic analyses of assemblages that have identified “assembly rules,” patterns of community structure for which competition is assumed to be the primary mechanism (e.g., Diamond 1975; Fox 1987, 1999; Weiher and Keddy 1999). Because the experimental and geographic studies have often been done with different kinds of organisms and habitats, and because their scales and methodologies have been so disparate, it has not been clear how they should be interrelated. How do the processes

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.

Home range and parasite diversity in mammals.

Parasite diversity among and within host species is not solely the result of random processes; rather, it depends on a suite of physiological or ecological host traits as well as environmental factors. Because most macroparasites exhibit life cycles that include infective stages off the definitive host and that rely on host movements for dissemination, parasite acquisition by a host depends largely on hosts being present in a given area where and when infective stages are present. Consequently, host ranging pattern may have a major influence on parasite diversity. Larger home range size is hypothesized to be associated with higher parasite species richness because hosts living in large home ranges should encounter a greater diversity of habitats and other host individuals, which in turn may favor infection by a great diversity of parasite species. By focusing on helminths in wild mammals, we show that an increase in home range area does not lead to an increase in parasite diversity in ungulates and, moreover, that it is associated with a decrease in parasite species richness in carnivores and in glires (rodents and lagomorphs). We also show that home range size is negatively correlated with host density in mammals after correcting both variables for host body mass. We discuss these results from an epidemiological perspective.