Effects of vole fluctuations on the population dynamics of the barn owl <Emphasis Type="Italic">Tyto alba</Emphasis>

Many predator species feed on prey that fluctuates in abundance from year to year. Birds of prey can face large fluctuations in food abundance i.e. small mammals, especially voles. These annual changes in prey abundance strongly affect the reproductive success and mortality of the individual predators and thus can be expected to influence their population dynamics and persistence. The barn owl, for example, shows large fluctuations in breeding success that correlate with the dynamics in voles, their main prey species. Analysis of the impact of fluctuations in vole abundance (their amplitude, peaks and lows, cycle length and regularity) with a simple predator prey model parameterized with literature data indicates population persistence is especially affected by years with low vole abundance. In these years the population can decline to low owl numbers such that the ensuing peak vole years cannot be exploited. This result is independent of the length and regularity of vole fluctuations. The relevance of this result for conservation of the barn owl and other birds of prey that show a numerical response to fluctuating prey species is discussed.     

Dynamics of the shallow-water fish assemblage of the Patos Lagoon estuary (Brazil) during cold and warm ENSO episodes

In southern Brazil, cold ( La Niña ) and warm ( El Niño ) episodes of the El Niño Southern Oscillation (ENSO) phenomenon cause drought and high rainfall, respectively. The low precipitation and freshwater outflow associated with La Niña during 1995–1996 were associated with an increase in the abundance of marine species in the Patos Lagoon estuary. During the 1997–1998 El Niño , high precipitation and river discharge were associated with low abundance of marine species in the estuary. ANOVA results showed a higher abundance during La Niña than El Niño for estuarine resident (RES) and estuarine dependent (DEP) fishes. During La Niña catch per unit of effort (CPUE) of RES increased from the marine to estuarine area, but during El Niño CPUE increased at the marine area and diminished during summer and autumn in some estuarine sites. DEP fishes had an opposite abundance pattern. During La Niña , these fishes were abundant at the coastal marine area and along some estuarine sites, but during El Niño , CPUE remained almost the same at the marine area but dropped along some estuarine sites. These different abundance patterns for dominant fish groups yielded a positive interaction between stations and climatic events. With higher river discharge and the consequent decline of dominant euryhaline fishes, such as Mugil platanus and Atherinella brasiliensis , freshwater species increased in abundance and richness in the shallow waters of the stuary. The ENSO phenomenon influences precipitation and estuarine salinity in southern Brazil and thereby seems to have a strong influence on recruitment, immigration, and emigration dynamics of fish species living within and adjacent to estuarine habitats.     

Ecological effects of El Niño in terrestrial ecosystems of western South America

I make a summary review of how El Niño/Southern Oscillation (ENSO) determines peculiar atmospheric and oceanographic conditions in western South America, thus affecting precipitation patterns in adjacent land masses, with cascading effects on marine and terrestrial plants, on sea and land birds, and on marine and terrestrial mammals. With regard to terrestrial ecosystems, I discuss the following biotic responses to El Niño-driven precipitation: 1) aboveground vegetation flushes immediately among herbs but not among shrubs. 2) The seed bank is quickly replenished of ephemeral seeds, but perennial seeds recover one year later. 3) Small rodents Irrupt within months of El Niño arrival, but larger ones take a full year to increase. 4) Predator numbers lag one year behind their mammal prey, with smaller predators responding more quickly. Considering these responses, I offer a simplified model of El Niño-driven bottom-up control in terrestrial ecosystems of western South America. Apart from the direct links already described, there is a weak feedback loop between the plant compartments (vegetation and seeds) and their herbivores: primary productivy is the driving force, and is little affected by herbivory. Another weak feedbaek loop links herbivores and their predators: the latter seem to just "surf" over prey levels, skimming excess prey.     

El Niño events,the lean versus fat scenario,and long‐term guild dynamics of vertebrate predators in a South American semiarid ecosystem

Abstract Predator assemblages are complex systems in which asynchrony in the dynamics of resources and consumers, and the idiosyncratic perception of environmental conditions by the predators may obscure the detection of expected patterns. We disentangle the specific effects of these variables on the guild structure of a vertebrate predatory assemblage in a semiarid ecosystem of western South America. Over 16 years, this system faced dramatic fluctuations in prey availability associated with four El Niño events. After controlling for other sources of variation, we tested if increased resource availability is associated with higher niche overlaps, as expected from the lean/fat scenario. We determined the existence of two trophic guilds of predators (specialized mammal‐eaters and omnivorous species with emphasis on arthropods) and found that they responded to increased productivity both at the guild and whole assemblage levels. However, the population response of arthropod prey (almost simultaneous) and of different small mammal prey (delayed by 1 or 2 years) to productivity imposed a degree of asynchrony in prey availability and in the response of predators. This resulted in the between‐guilds exchange of predator species depending on mammal prey scarcity or abundance. As a consequence, the observed pattern was an apparent lack of response at the assemblage level. Despite differences in the perception of prey levels by predators, we conclude that each one of them responded accordingly to theoretical predictions following a simple rule: if prey resources are not limiting, predators behave opportunistically converging over the most abundant resources, thus increasing niche overlap; if prey shortages occur, predators specialize on those prey resources that they gather most efficiently, thus lowering niche overlap; if resources become even scarcer, all predators converge again upon the few prey resources still available, thus increasing overlap – out of necessity.     

The relationship between intra–guild diet overlap and breeding in owls in Israel

Even though intra-guild predators frequently prey on the same species, it is unclear whether diet overlap between two predators is a source of interspecific competition or whether predators simply use the same abundant prey resource. We measured the extent to which the diets of barn owls (Tyto alba) and long-eared owls (Asio otus) in Israel overlap and examined whether yearly differences in diet overlap correlate with barn owl breeding success. Pianka’s index of niche overlap was positively related to barn owl population size but not to its breeding success. The number of breeding barn owls was higher when long-eared owls consumed more rodents, suggesting that diet overlap most likely increased when rodents became more abundant. Therefore, in Israel, when these two owl species prey more often on rodents, their diets are more similar and interspecific competition is reduced. Unlike sympatric populations in Europe, in years when rodents are less abundant in Israel long-eared owls switch to hunting alternative prey (e.g., birds), perhaps to avoid competition with barn owls.     

Using scats of a generalist carnivore as a tool to monitor small mammal communities in Mediterranean habitats

Owl pellets have long been used to analyze communities of small mammals, while analogous analyses of faeces of mammal carnivores are not available. We demonstrate that common genet (Genetta genetta) scats can be used as a reliable method to sample small mammal communities and to monitor their variations. We have compiled data on 6350 small mammal remains of 18 species found in scats from 51 different latrines in a 1200 km² area of northeastern Spain. Genet scats sampled effectively 95.6% of the small mammal species ranging in size from 2.7 to 385 g. Spatial patterns of diet composition along environmental gradients of elevation, climate and land-use matched expected changes in small mammal communities along these gradients according to ecological requirements of prey species. Frequencies of occurrence of prey in genet scats were strongly correlated with frequencies of occurrence in barn owl (Tyto alba) pellets. Genet scats included two forest species not preyed upon by owls, whereas only one species was not preyed upon by genets. Forests species were more frequent in genet than in barn owl diets after correcting for environmental effects, whereas the opposite was true for open-habitat and synanthropic species. Scats of generalist carnivores can be used to estimate the spatial patterns of distribution and abundance of small mammal communities. Genet scats in fact overcome some of the limitations of more traditional sampling methods (live-trapping and owl diets), as genets were less selective and their diets reflect more accurately changes in community composition.     

Population fluctuations of the house mouse in a Peruvian loma and the functional response of burrowing owls

Abstract In this study, we show that the house mouse (Mus musculus) is the most abundant small mammal in the National Reserve of Lachay in central Peru, and that its large population fluctuations are independent of seasonality. Also, we found that M. musculus is the main small mammal prey of burrowing owls (Athene cunicularia) in Lachay, and that owls respond functionally to mouse abundance. In addition, vegetation cover seems to have a strong effect on small mammal predation by burrowing owls, and possibly other predators such as eagles and foxes. We propose the hypothesis that burrowing owls in arid environments can survive several months eating only arthropods, but that their reproduction is synchronized with a highly nutritious diet, provided by small mammal consumption. The principal prediction of this alternative prey hypothesis is now supported by our data from a tropical ecosystem.     

Variation in a sparrow's reproductive success with rainfall: food and predator-mediated processes

From 1997 to 1999, we monitored the reproductive success of individual rufous-crowned sparrows (Aimophila ruficeps) in coastal sage scrub habitat of southern California, USA. Annual reproductive output of this ground-nesting species varied strongly with annual variation in rainfall, attributed to the El Niño-Southern Oscillation. Birds fledged 3.0 young per breeding pair in 1997, when rainfall was near the long-term mean, 5.1 offspring per pair in 1998, a wet El Niño year, and 0.8 fledglings per pair in 1999, a dry La Niña year. Variation in many components of reproductive output was consistent with the hypothesis that food availability was positively correlated with rainfall. However, the factor most responsible for the high reproductive output in 1998 was low early season nest predation which, combined with favorable nesting conditions, enabled more pairs to multiple-brood. Cool, rainy El Niño conditions may have altered the activity of snakes, the main predator of these nests, in the early season of 1998. Overall, more of the annual variation in fecundity was attributable to variation in within-season components of reproductive output (mean number of nests fledged per pair) than to within-nest components (mean brood size). Annual variation in rufous-crowned sparrow fecundity appears to be driven primarily by food resource-mediated processes in La Niña years and by predator-mediated processes in El Niño years.     

Population dynamics of a polychaete during three El Niño events: disentangling biotic and abiotic factors

The macrobenthic community in shallow soft-bottom areas in the Bay of Ancón, Peru, is characterised by low biodiversity due to low oxygen concentrations. During El Niño events, higher temperature and higher concentrations of dissolved oxygen induce a temporary increase in biodiversity. However, the structure and dynamics of the emerging communities vary strongly among events. The reasons for this variation are poorly understood, in particular the relative influence of abiotic vs biotic factors. To disentangle the roles of abiotic and biotic factors, population models based on detailed monitoring data of three El Niño events were developed focused on the population dynamics of one species in the community, the polychaete Sigambra bassi , which showed different responses in different El Niño events. Calculated and observed population dynamics are compared using root mean square deviation (RMSD). The results show that S. bassi abundance can be determined by abiotic environmental conditions. Besides, three biotic factors improved model performance in different El Niño events: negative density dependence in larval settlement, lower carrying capacity in the presence of other species and positive density dependence in adult survival. The results demonstrate how population models can be used to decode information hidden in long-term monitoring data of abiotic and biotic community parameters about factors driving population dynamics.     

Role of prey and intraspecific density dependence on the population growth of an avian top predator

Exploring predator–prey systems in diverse ecosystems increases our knowledge about ecological processes. Predator population growth may be positive when conspecific density is low but predators also need areas with prey availability, associated with competition, which increases the risk of suffering losses but stabilises populations. We studied relationships between European rabbits Oryctolagus cuniculus (prey) and adult eagle owls Bubo bubo (predators) in south-western Europe. We assessed models explaining the predator population growth and stability. We estimated the abundance of rabbits and adult eagle owls during three years in eight localities of central-southern Spain. We explored models including rabbit and adult eagle owl abundance, accounting for yearly variations and including the locality as a random variable. We found that population growth of adult eagle owls was positive in situations with low conspecific abundance and tended to be negative but approaching equilibrium in situations of higher conspecific abundance. Population growth was also positively related to previous summer rabbit density when taking into account eagle owl conspecific abundance, possibly indicating that rabbits may support recruitment. Furthermore, abundance stability of adult eagle owls was positively related to previous winter–spring rabbit density, which could suggest predator population stabilisation through quick territory occupation in high-quality areas. These results exemplify the trade-off between prey availability and abundance of adult predators related to population growth and abundance stability in the eagle owl–rabbit system in south-western Europe. Despite rabbits have greatly declined during the last decades and eagle owls locally specialise on them, eagle owls currently have a favourable conservation status. As eagle owls are the only nocturnal raptor with such dependence on rabbits, this could point out that predators may overcome prey decreases in areas with favourable climate and prey in the absence of superior competitors with similar foraging mode.     

Impacts of Agriculture on the Diet and Productivity of Mackinder's Eagle Owls (Bubo capensis mackinderi ) in Kenya

Land conversion for agriculture is an increasing threat to biodiversity conservation, but its ecological effects on African birds is practically unknown. We investigated the impacts of agriculture on the diet and productivity of a small, disjunct population of Mackinder's eagle owls ( Bubo capensis mackinderi  ) in central Kenya. Owl diet was determined by analysis of pellets and other remains and compared to small mammal populations estimated by live trapping in two habitats. Small mammal abundance was low and averaged 7.4 small mammals/ha in farms and 0.5 small mammals/ha in grassland. Owls consumed a wide diversity of prey. The majority were mammals (87%) followed by birds (7%) and insects (5%). The percentage of small mammals in owl diet correlated positively with the relative abundance of small mammals during monthly trapping sessions. Diet composition did not influence owl breeding success. Farming activities affected owl diet composition through crop production. The amount of maize, peas, and carrots growing in farms was correlated with the abundance of Mastomys sp. and Procavia sp. in the owl's diet. Agricultural activities had a large effect on Mackinder's eagle owl diet by increasing the abundance of certain small-mammal prey and attracting owl prey to farms, though farming practices harmful to owls were observed.     

Numerical and functional response of predators to a long-term decline in mammalian prey at a semi-arid Neotropical site

Summary Occurrence and diet of ten carnivorous predators (four falconiforms, four owls, and two foxes), and population levels of their mammalian prey, were monitored over 45 months at a semi-arid site in north-central Chile. Early in this period, small mammals irrupted and then declined markedly to a density 7% of that at peak. All four falconiforms (Buteo polyosoma, Falco sparverius, Geranoaetus melanoleucus, Parabuteo unicinctus) and one owl (Tyto alba) responded numerically to the decline in mammalian prey by virtually abandoning the study site. The three other owls (Athene cunicularia, Bubo virginianus, Glaucidium nanum) and the two foxes (Pseudalopex culpaeus and P. griseus) remained. With few exceptions, throughout the study predators maintained species-specific preferences among small mammal species regardless of the absolute and proportional abundance of these prev. At no time did the two prey species most responsible for the irruption (the rodents Phyllotis darwini and Akodon olivaceus) occur in predators' diets out of proportion to their estimated relative abundance in the field. Predators were clearly unable to prevent the irruption from occurring. Given the absence of a clear functional response to the most irruptive species, predators seemed unlikely to have been responsible for the observed crash. At present, however, predators may be prolonging the crash and delaying the return of small-mammal populations to typical densities.     

El Niño Southern Oscillation influences the abundance and movements of a marine top predator in coastal waters

Large‐scale climate modes such as El Niño Southern Oscillation (ENSO) influence population dynamics in many species, including marine top predators. However, few quantitative studies have investigated the influence of large‐scale variability on resident marine top predator populations. We examined the effect of climate variability on the abundance and temporary emigration of a resident bottlenose dolphin (Tursiops aduncus) population off Bunbury, Western Australia (WA). This population has been studied intensively over six consecutive years (2007–2013), yielding a robust dataset that captures seasonal variations in both abundance and movement patterns. In WA, ENSO affects the strength of the Leeuwin Current (LC), the dominant oceanographic feature in the region. The strength and variability of the LC affects marine ecosystems and distribution of top predator prey. We investigated the relationship between dolphin abundance and ENSO, Southern Annular Mode, austral season, rainfall, sea surface salinity and sea surface temperature (SST). Linear models indicated that dolphin abundance was significantly affected by ENSO, and that the magnitude of the effect was dependent upon season. Dolphin abundance was lowest during winter 2009, when dolphins had high temporary emigration rates out of the study area. This coincided with the single El Niño event that occurred throughout the study period. Coupled with this event, there was a negative anomaly in SST and an above average rainfall. These conditions may have affected the distribution of dolphin prey, resulting in the temporary emigration of dolphins out of the study area in search of adequate prey. This study demonstrated the local effects of large‐scale climatic variations on the short‐term response of a resident, coastal delphinid species. With a projected global increase in frequency and intensity of extreme climatic events, resident marine top predators may not only have to contend with increasing coastal anthropogenic activities, but also have to adapt to large‐scale climatic changes.     

Kestrel-Prey Dynamic in a Mediterranean Region: The Effect of Generalist Predation and Climatic Factors

Background

Most hypotheses on population limitation of small mammals and their predators come from studies carried out in northern latitudes, mainly in boreal ecosystems. In such regions, many predators specialize on voles and predator-prey systems are simpler compared to southern ecosystems where predator communities are made up mostly of generalists and predator-prey systems are more complex. Determining food limitation in generalist predators is difficult due to their capacity to switch to alternative prey when the basic prey becomes scarce.

Methodology

We monitored the population density of a generalist raptor, the Eurasian kestrel Falco tinnunculus over 15 years in a mountainous Mediterranean area. In addition, we have recorded over 11 years the inter-annual variation in the abundance of two main prey species of kestrels, the common vole Microtus arvalis and the eyed lizard Lacerta lepida and a third species scarcely represented in kestrel diet, the great white-toothed shrew Crocidura russula. We estimated the per capita growth rate (PCGR) to analyse population dynamics of kestrel and predator species.

Principal Findings

Multimodel inference determined that the PCGR of kestrels was better explained by a model containing the population density of only one prey species (the common vole) than a model using a combination of the densities of the three prey species. The PCGR of voles was explained by kestrel abundance in combination with annual rainfall and mean annual temperature. In the case of shrews, growth rate was also affected by kestrel abundance and temperature. Finally, we did not find any correlation between kestrel and lizard abundances.

Significance

Our study showed for the first time vertebrate predator-prey relationships at southern latitudes and determined that only one prey species has the capacity to modulate population dynamics of generalist predators and reveals the importance of climatic factors in the dynamics of micromammal species and lizards in the Mediterranean region.     

Do native predators benefit from non‐native prey?

Despite knowledge on invasive species’ predatory effects, we know little of their influence as prey. Non‐native prey should have a neutral to positive effect on native predators by supplementing the prey base. However, if non‐native prey displace native prey, then an invader's net influence should depend on both its abundance and value relative to native prey. We conducted a meta‐analysis to quantify the effect of non‐native prey on native predator populations. Relative to native prey, non‐native prey similarly or negatively affect native predators, but only when studies employed a substitutive design that examined the effects of each prey species in isolation from other prey. When native predators had access to non‐native and native prey simultaneously, predator abundance increased significantly relative to pre‐invasion abundance. Although non‐native prey may have a lower per capita value than native prey, they seem to benefit native predators by serving as a supplemental prey resource.     

El Niño events, precipitation patterns, and rodent outbreaks are statistically associated in semiarid Chile

In the last two decades, several researchers have noted rodent population outbreaks in semiarid South America, in association with unusually high precipitation that seemingly concurs with El Niño events. To date, no studies have been conducted to determine the statistical relationships between ENSO (El Niño Southern Oscillation) events, increased precipitation, and rodent irruptions. Here we show that: 1) there is a statistical association between ENSO events and inereased precipitation in the semiarid region of northern Chile: 2) the occurrence of rodent outbreaks in that region is statistically related with the precipitation levels of the same year: 3) the multi-annual patterns of the total annual precipitation levels and population abundance of those rodents during the summer are positively associated. The putative chain of effects seems to start with unusually high rainfall brought by ENSO to semiarid environments, which thus respond with inereased primary productivity (herbage and seeds), which then fuels the rodent outbreaks.     

Assessing effects of radiation on abundance of mammals and predator–prey interactions in Chernobyl using tracks in the snow

To test whether radioactive contamination reduced the abundance of mammals, and whether species differed in susceptibility to radiation, we censused mammals by counting tracks in the snow along 161 100-m line transects around Chernobyl during February 2009. The abundance of mammal tracks was negatively related to level of background radiation, independent of the statistical model, with effects of radiation accounting for a third of the variance. The effect of radiation differed significantly among species. There was a positive relationship between abundance of predators and abundance of prey, modified by the level of background radiation because the number of predators increased disproportionately with the number of prey at high levels of radiation. These findings suggest that predatory mammals aggregate in areas with abundant prey, especially when prey are exposed to high levels of radiation. This study emphasizes the negative effects of level of background radiation on the abundance of mammals and predator–prey interactions.     

Predator-induced synchrony in population oscillations of coexisting small mammal species

Comprehensive analyses of long-term (1977-2003) small-mammal abundance data from western Finland showed that populations of Microtus voles (field voles M. agrestis and sibling voles M. rossiaemeridionalis) voles, bank (Clethrionomys glareolus) and common shrews (Sorex araneus) fluctuated synchronously in 3 year population cycles. Time-series analyses indicated that interspecific synchrony is influenced strongly by density-dependent processes. Synchrony among Microtus and bank voles appeared additionally to be influenced by density-independent processes. To test whether interspecific synchronization through density-dependent processes is caused by predation, we experimentally reduced the densities of the main predators of small mammals in four large agricultural areas, and compared small mammal abundances in these to those in four control areas (2.5-3 km(2)) through a 3 year small-mammal population cycle. Predator reduction increased densities of the main prey species, Microtus voles, in all phases of the population cycle, while bank voles, the most important alternative prey of predators, responded positively only in the low and the increase phase. Manipulation also increased the autumn densities of water voles (Arvicola terrestris) in the increase phase of the cycle. No treatment effects were detected for common shrews or mice. Our results are in accordance with the alternative prey hypothesis, by which predators successively reduce the densities of both main and alternative prey species after the peak phase of small-mammal population cycles, thus inducing a synchronous low phase.     

Episodic outbreaks of small mammals influence predator community dynamics in an east African savanna ecosystem

Little is known about the dynamics of small mammals in tropical savanna: a critical gap in our understanding of Africa's best known ecosystems. Historical evidence suggested small mammals peak in abundance (outbreak) in Serengeti National Park (SNP), as in agricultural systems. We asked 1) what are bottom–up drivers of small mammals and 2) do predators have top–down effects? We documented dynamics of small mammals, birds of prey, and mammalian carnivores in SNP and agricultural areas. We used climatic fluctuations and differences between unmodified and agricultural systems as perturbations to examine trophic processes, key to understanding responses to climate change and increasing human pressures. Data were derived from intermittent measures of abundance collected 1968–1999, combined with systematic sampling 2000–2010 to construct a 42‐year time series. Data on abundance of black‐shouldered kites (1968–2010), eight other species of rodent‐eating birds (1997–2010), and 10 carnivore species (1993–2010) were also collated. Outbreaks occurred every 3–5 years in SNP, with low or zero abundance between peaks. There was a positive relationship between rainfall in the wet season and 1) small mammal abundance and 2) the probability of an outbreak, both of which increased with negative Southern Oscillation Index values. Rodent‐eating birds and carnivores peaked 6–12 months after small mammals. In agricultural areas, abundance remained higher than in natural habitats. Abundances of birds of prey and mammalian carnivores were extremely low in these areas and not related to small mammal abundance. Small mammals are an important food resource for higher trophic levels in the Serengeti ecosystem. Changes in climate and land use may alter their future dynamics, with cascading consequences for higher trophic levels, including threatened carnivores. Although outbreaks cause substantial damage to crops in agricultural areas, small mammals also play a vital role in maintaining some of the diversity and complexity found in African savanna ecosystems.     

Diet choice and predator—prey dynamics

Summary We compare the dynamics of predator-prey systems with specialist predators or adaptive generalist predators that base diet choice on energy-maximizing criteria. Adaptive predator behaviour leads to functional responses that are influenced by the relative abundance of alternate prey. This results in the per capita predation risk being positively density-dependent near points of diet expansion. For a small set of parameter values, systems with adaptive predators can be locally stable whereas systems with specialist predators would be unstable. This occurs mainly when alternate prey have low enough profitability that predators cannot sustain themselves indefinitely when feeding on alternate prey. Local stability of systems with adaptive predator behaviour is inversely related to the goodness of fit to optimal diet choice criteria. Hence, typical patterns of partial prey preference are more stabilizing than perfect optimal diet selection. Locally stable systems with adaptive predators are often globally unstable, converging on limit cycles for many initial population densities. The small range of parameter combinations and initial population densities leading to stable equilibria suggest that adaptive diet selection is unlikely to be a ubiquitous stabilizing factor in trophic interactions.