Primary production dynamics and climate variability: ecological consequences in semiarid Chile

Increase in rainfall variability has important consequences for organisms in arid and semiarid regions around the world. In South American and Australian deserts, the El Niño/Southern Oscillation (ENSO) phenomenon greatly influences rainfall patterns, and therefore the dynamics of plant communities. However, the field data needed to assess the effect of climate change on vegetational patterns is difficult to obtain because of the large spatial scale required for such studies. Normalized Difference Vegetation Index (NDVI) characteristics allow the use of several indexes related to vegetational structure. Due to its direct relationship with primary productivity, it is possible to obtain several measures of annual productivity. These include annual plant yield, annual maximum yield, onset of 'greening-up' and senescence phases, length of the 'green' season, vegetation peak, and therefore, the periods when more or less food is available for herbivores. After verification with ground-truth measures, we used NDVI data from two semiarid localities in north-central Chile (Fray Jorge and Aucó) to determine the relationship between rainfall patterns and vegetation cover and productivity related to El Niño phenomenon. With this information we gauge the influence of climatic processes on primary productivity in western South America, an area subject to strong climate variability. We predict significant variation in Chilean semiarid regions due to climate change, affecting mainly the extent and timing of annual growth season of vegetation, and also including a shorter and delayed greening-up season. Also, we predict that important decreases in rainfall levels will not have strong effects on primary production in these semiarid ecosystems.     

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.     

Effects of large-scale climatic fluctuations on survival and production of young in a Neotropical migrant songbird, the yellow warbler Dendroica petechia

Migrant songbirds are vulnerable to changes in climatic conditions on both the breeding and wintering grounds. For North American Neotropical migrants, the El Niño/Southern Oscillation (ENSO), via its effects on global precipitation and temperature, modulates the productivity of their temperate and tropical terrestrial ecosystems used during the course of their annual cycle. We evaluated how a densely nesting population of yellow warblers Dendroica petechia in a riparian forest in southern Manitoba, Canada, responded to the El Niño/Southern Oscillation (ENSO) between 1992 and 2001. Standardized mist netting was used to estimate apparent annual survival of adults and production of young. Both adult survival and the production of young were positively correlated with the Southern Oscillation Index (SOI). More specifically, values of both these demographic parameters were lowest during El Niño years and highest during La Niña years. These findings demonstrate the influence of climate on populations of Neotropical migrants in North America. The more frequent El Niños predicted to result from future global climate change could negatively affect populations of yellow warblers and other Neotropical migrants breeding in this region.     

Comparing aquatic and terrestrial grazing ecosystems: is the grass really greener?

‘Grazing ecosystem’ is typically used to describe terrestrial ecosystems with high densities of mammalian herbivores such as the Serengeti in East Africa or the Greater Yellowstone Ecosystem in North America. These abundant, large herbivores determine plant community dynamics and ecosystem processes. The general concepts that define grazing ecosystems also aptly describe many aquatic ecosystems, including coral reefs, seagrass beds, and lakes, where herbivores such as parrotfishes, turtles, and zooplankton have strong impacts on ecosystem processes. Here, I compare the ecology of grazing ecosystems in search of common concepts that transcend the terrestrial‐aquatic boundary. Specifically, I evaluate: 1) the feedbacks between herbivory and primary production, 2) the roles of herbivore richness and facilitation, 3) how predators and diet quality shape patterns of herbivory, and 4) how altering herbivory mediates alternative states.     

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.     

The distribution of deer biomass in North America supports the hypothesis of exploitation ecosystems

The hypothesis of exploitation ecosystems (EEH) predicts that, along a productivity gradient in terrestrial environments, predators will regulate herbivores at a relatively constant density whenever primary productivity exceeds 700 g m⁻² y⁻¹; under this threshold, or if predators are absent, forage production determines herbivore density. I tested EEH using the pattern of deer biomass distribution over North America, the dominant family of large herbivores. Deer biomass increased from the High Arctic to the north of the boreal forest and remained in the same range southward within the gray wolf range; for the same latitude, deer biomass increased by a factor of 5 in the absence of wolves. South of the wolf range, there existed a clear relationship between actual evapotranspiration, a proxy of primary productivity, and deer biomass. Highest deer densities occurred in the south-east of the continent where only white-tailed deer are present. The observed pattern lends support to EEH and suggests that the removal of large predators in southern North America may have imposed an unprecedented pressure on plants eaten by deer.     

A cross-ecosystem comparison of the strength of trophic cascades

Although trophic cascades (indirect effects of predators on plants via herbivores) occur in a wide variety of food webs, the magnitudes of their effects are often quite variable. We compared the responses of herbivore and plant communities to predator manipulations in 102 field experiments in six different ecosystems: lentic (lake and pond), marine, and stream benthos, lentic and marine plankton, and terrestrial (grasslands and agricultural fields). Predator effects varied considerably among systems and were strongest in lentic and marine benthos and weakest in marine plankton and terrestrial food webs. Predator effects on herbivores were generally larger and more variable than on plants, suggesting that cascades often become attenuated at the plant–herbivore interface. Top‐down control of plant biomass was stronger in water than on land; however, the differences among the five aquatic food webs were as great as those between wet and dry systems.     

EVOLUTION OF SPRINT SPEED IN AFRICAN SAVANNAH HERBIVORES IN RELATION TO PREDATION

Predator–prey arms races are widely speculated to underlie fast speed in terrestrial mammals. However, due to lack of empirical testing, both the specificity of any evolutionary coupling between particular predator and prey species, and the relevance of alternative food‐based hypotheses of speed evolution, remain obscure. Here I examine the ecological links between the sprint speed of African savannah herbivores, their vulnerability to predators, and their diet. I show that sprint speed is strongly predicted by the vulnerability of prey to their main predators; however, the direction of the link depends on the hunting style of the predator. Speed increases with vulnerability to pursuit predators, whereas vulnerability to ambush predators is associated with particularly slow speed. These findings suggest that differential vulnerability to specific predators can indeed drive interspecific variation in speed within prey communities, but that predator hunting style influences the intensity and consistency with which selection on speed is coupled between particular species.     

More than a barrier: The complex effects of ecotone vegetation type on terrestrial consumer consumption of an aquatic prey resource

Community structure and dynamics can be influenced by resource transfers between ecosystems, yet little is known about how boundary structure determines both the magnitude of exchanges and their effects on recipient and donor communities. Aquatic and terrestrial ecosystems are often linked by resource fluxes and riparian vegetation is commonly affected by anthropogenic alterations to land use or river hydrological regime. I investigated whether shrubs at the freshwater–terrestrial interface alter the supply, distribution and importance of aquatic prey resources to terrestrial consumers. Shrubs were predicted to alter the larval community composition of aquatic insects and the emergence of winged adults, thus affecting aquatic prey subsidies to terrestrial consumers. In addition, shrubs were hypothesized to alter the microclimatic suitability of the riparian zone for adult aquatic insects, act as a physical barrier to their dispersal and affect terrestrial community composition, particularly the abundance and type of predators that could benefit from the aquatic prey resource. Stable isotope dietary analyses and a survey of shrub‐dominated and open grassland riparian habitats revealed that larval densities of aquatic insects (EPTM: Ephemeroptera, Plecoptera, Trichoptera and Megaloptera) were higher in shrub than grassland habitats; however, reduced emergence and lateral dispersal in shrub areas led to lower densities of adults. The temperature and relative humidity of the riparian zone did not differ between the habitats. Ground‐active terrestrial invertebrate communities had a higher proportion of cursorial spiders in grassland, coinciding with greater abundances of aquatic prey. Aquatic prey contribution to cursorial spider diet matched adult aquatic insect abundances. Overall, riparian shrubs reduced the magnitude, or at least altered the timing, of cross‐ecosystem subsidy supply, distribution and use by consumers through mechanisms operating in both the aquatic and terrestrial ecosystems. Thus, the structure of ecosystem boundaries has complex effects on the strength of biological interactions between adjacent systems.     

A review of climate-driven mismatches between interdependent phenophases in terrestrial and aquatic ecosystems

Mismatches in phenology between mutually dependent species, resulting from climate change, can have far-reaching consequences throughout an ecosystem at both higher and lower trophic levels. Rising temperatures, due to climate warming, have resulted in advances in development and changes in behaviour of many organisms around the world. However, not all species or phenophases are responding to this increase in temperature at the same rate, thus creating a disruption to previously synchronised interdependent key life-cycle stages. Mismatches have been reported between plants and pollinators, predators and prey, and pests and hosts. Here, we review mismatches between interdependent phenophases at different trophic levels resulting from climate change. We categorized the studies into (1) terrestrial (natural and agricultural) ecosystems, and (2) aquatic (freshwater and marine) ecosystems. As expected, we found reports of 'winners' and 'losers' in each system, such as earlier emergence of prey enabling partial avoidance of predators, potential reductions in crop yield if herbivore pests emerge before their predators and possible declines in marine biodiversity due to disruption in plankton-fish phenologies. Furthermore, in the marine environment rising temperatures have resulted in synchrony in a previously mismatched prey and predator system, resulting in an abrupt population decline in the prey species. The examples reviewed suggest that more research into the complex interactions between species in terrestrial and aquatic ecosystems is necessary to make conclusive predictions of how climate warming may impact the fragile balances within ecosystems in future.     

Historical biogeography of the Isthmus of Panama

About 3 million years ago (Ma), the Isthmus of Panama joined the Americas, forming a land bridge over which inhabitants of each America invaded the other—the Great American Biotic Interchange. These invasions transformed land ecosystems in South and Middle America. Humans invading from Asia over 12000 years ago killed most mammals over 44 kg, again transforming tropical American ecosystems. As a sea barrier, the isthmus induced divergent environmental change off its two coasts—creating contrasting ecosystems through differential extinction and diversification. Approximately 65 Ma invading marsupials and ungulates of North American ancestry, and xenarthrans of uncertain provenance replaced nearly all South America's non‐volant mammals. There is no geological evidence for a land bridge at that time. Together with rodents and primates crossing from Africa 42 to 30 Ma, South America's mammals evolved in isolation until the interchange's first heralds less than 10 Ma. Its carnivores were ineffective marsupials. Meanwhile, North America was invaded by more competitive Eurasian mammals. The Americas had comparable expanses of tropical forest 55 Ma; later, climate change confined North American tropical forest to a far smaller area. When the isthmus formed, North American carnivores replaced their marsupial counterparts. Although invaders crossed in both directions, North American mammals spread widely, diversified greatly, and steadily replaced South American open‐country counterparts, unused to effective predators. Invading South American mammals were less successful. South America's birds, bats, and smaller rainforest mammals, equally isolated, mostly survived invasion. Its vegetation, enriched by many overseas invaders, remained intact. This vegetation resists herbivory effectively. When climate permitted, South America's rainforest, with its bats, birds and mammals, spread to Mexico. Present‐day tropical American vegetation is largely zoned by trade‐offs between exploiting well‐watered settings versus surviving droughts, exploiting fertile versus coping with poor soil, and exploiting lowland warmth versus coping with cooler altitudes. At the start of the Miocene, a common marine biota extended from Trinidad to Ecuador and western Mexico, which evolved in isolation from the Indo‐Pacific until the Pleistocene. The seaway between the Americas began shoaling over 12 Ma. About 10 Ma the land bridge was briefly near‐complete, allowing some interchange of land mammals between the continents. By 7 Ma, the rising sill had split deeper‐water populations. Sea temperature, salinity and sedimentary carbon content had begun to increase in the Southern Caribbean, but not the Pacific. By 4 Ma, the seaway's narrowing began to extinguish Caribbean upwellings. By 2 Ma, upwellings remained only along Venezuela; Caribbean plankton, suspension‐feeding molluscs and their predators had declined sharply, largely replaced by bottom‐dwelling corals and calcareous algae and magnificent coral reefs. Closing the seaway extinguished the Eastern Pacific's reef corals (successors recolonized from the Indo‐Pacific 6000 years ago), whereas many molluscs of productive waters that once thrived in the Caribbean now survive only in the Eastern Pacific. The present‐day productive Eastern Pacific, with few, small coral reefs and a plankton‐based ecosystem contrasts with the Caribbean, whose clear water favours expansive coral reefs and bottom‐dwelling primary producers. These ecosystems reflect the trade‐off between fast growth and effective defence with attendant longevity. Overfishing with new technologies during the last few centuries, however, has caused population crashes of ever‐smaller marine animals, devastating Caribbean ecosystems.     

A meta‐analysis of the effects of detritus on primary producers and consumers in marine,freshwater, and terrestrial ecosystems

Detritus is a central feature in marine, freshwater, and terrestrial ecosystems. Despite the ubiquity of detritus, ecologists have largely ignored its role in influencing food web structure. We used a meta‐analytic approach to ask three questions about how detritus affects food web structure in a wide variety of ecosystems. First, what is the effect strength of detritus on primary producers, detritivores, herbivores, and predators? Second, what functional role does detritus serve for consumers (energetic, habitat, or both)? Third, how does the effect of detritus on consumers vary between aquatic and terrestrial ecosystems? We found that detritus has strong positive effects on primary producers and consumers in a wide range of ecosystems types. Detritus has a positive direct effect on detritivores by providing both an energetic resource and habitat (refuge from predators). Detritus has equally strong positive effects on herbivores and predators, driven by a positive direct effect of habitat. Detritus has positive effects on consumers in both aquatic and terrestrial ecosystems with 1.7 times stronger effects in terrestrial ecosystems. These results suggest that detritus has strong effects on food‐web structure in a variety of ecosystem types. Even the portion of the food web that is linked most strongly to living plant tissue as its primary energy source is strongly positively affected.     

Ecologically functional landscapes and the role of dingoes as trophic regulators in south‐eastern Australia and other habitats

Large carnivores can play a pivotal role in maintaining healthy, balanced ecosystems. By suppressing the abundances and hence impacts of herbivores and smaller predators, top predators can indirectly benefit the species consumed by herbivores and smaller predators. Restoring and maintaining the ecosystem services that large carnivores provide has been identified as a critical step required to sustain biodiversity and maintain functional, resilient ecosystems. Recent research has shown that Australia's largest terrestrial predator, the Dingo (Canis lupus dingo), has strong effects on ecosystems in arid Australia and that these effects are beneficial for the conservation of small mammals and vegetation. Similarly, there is evidence from south‐eastern Australia that dingoes suppress the abundance of macropods and red Fox (Vulpes vulpes). It is likely that dingoes in south‐eastern Australia also generate strong indirect effects on the prey of foxes and macropods, as has been observed in the more arid parts of the continent. These direct and indirect effects of dingoes have the potential to be harnessed as passive tools to assist biodiversity conservation through the maintenance of ecologically functional dingo populations. However, research is required to better understand dingoes' indirect effects on ecosystems and the development of dingo management strategies that allow for both the preservation of dingoes and protection of livestock.     

Rising synchrony controls western North American ecosystems

Along the western margin of North America, the winter expression of the North Pacific High (NPH) strongly influences interannual variability in coastal upwelling, storm track position, precipitation, and river discharge. Coherence among these factors induces covariance among physical and biological processes across adjacent marine and terrestrial ecosystems. Here, we show that over the past century the degree and spatial extent of this covariance (synchrony) has substantially increased, and is coincident with rising variance in the winter NPH. Furthermore, centuries‐long blue oak (Quercus douglasii) growth chronologies sensitive to the winter NPH provide robust evidence that modern levels of synchrony are among the highest observed in the context of the last 250 years. These trends may ultimately be linked to changing impacts of the El Niño Southern Oscillation on midlatitude ecosystems of North America. Such a rise in synchrony may destabilize ecosystems, expose populations to higher risks of extinction, and is thus a concern given the broad biological relevance of winter climate to biological systems.     

Food web structure and climate effects on the dynamics of small mammals and owls in semi-arid Chile

Population dynamics of small mammals and predators in semi-arid Chile is positively correlated with rainfall associated with incursions of El Niño (El Niño Southern Oscillation: ENSO). However, the causal relationships between small mammal fluctuations, predator oscillations, and climatic disturbances are poorly understood. Here, we report time series models for three species of small mammal prey and two species of owl predators. The large differences in population fluctuations between the three small mammal species are related to differences in their respective feedback structures. The analyses reveal that per capita growth rate of the leaf-eared mouse is a decreasing function of log density and of log barn owl abundance together with a positive rainfall effect. In turn, per capita population growth rate ( R -function) of the barn owl is a negative function of log barn owl abundance and a positive function of leaf-eared mouse abundance, suggesting a predator–prey interaction. The dramatic population fluctuations exhibited by leaf-eared mouse ( Phyllotis darwini ) are caused by climate effects coupled with a complex food web architecture.     

Geometric factors influencing the diet of vertebrate predators in marine and terrestrial environments

Predator–prey relationships are vital to ecosystem function and there is a need for greater predictive understanding of these interactions. We develop a geometric foraging model predicting minimum prey size scaling in marine and terrestrial vertebrate predators taking into account habitat dimensionality and biological traits. Our model predicts positive predator–prey size relationships on land but negative relationships in the sea. To test the model, we compiled data on diets of 794 predators (mammals, snakes, sharks and rays). Consistent with predictions, both terrestrial endotherm and ectotherm predators have significantly positive predator–prey size relationships. Marine predators, however, exhibit greater variation. Some of the largest predators specialise on small invertebrates while others are large vertebrate specialists. Prey–predator mass ratios were generally higher for ectothermic than endothermic predators, although dietary patterns were similar. Model‐based simulations of predator–prey relationships were consistent with observed relationships, suggesting that our approach provides insights into both trends and diversity in predator–prey interactions.     

Mechanisms of coexistence in diverse herbivore–carnivore assemblages: demographic,temporal and spatial heterogeneities affecting prey vulnerability

Simple models coupling the dynamics of single predators to single prey populations tend to generate oscillatory dynamics of both predator and prey, or extirpation of the prey followed by that of the predator. In reality, such oscillatory dynamics may be counteracted by prey refugia or by opportunities for prey switching by the predator in multi‐prey assemblages. How these mechanisms operate depends on relative prey vulnerability, a factor ignored in simple interactive models. I outline how compositional, temporal, demographic and spatial heterogeneities help explain the contrasting effects of top predators on large herbivore abundance and population dynamics in species‐rich African savanna ecosystems compared with less species‐diverse northern temperate or subarctic ecosystems. Demographically, mortality inflicted by predation depends on the relative size and life history stage of the prey. Because all animals eventually die and are consumed by various carnivores, the additive component of the mortality inflicted is somewhat less than the predation rate. Prey vulnerability varies annually and seasonally, and between day and night. Spatial variation in the risk of predation depends on vegetation cover as well as on the availability of food resources. During times of food shortage, herbivores become prompted to occupy more risky habitats retaining more food. Predator concentrations dependent on the abundance of primary prey species may restrict the occurrence of other potential prey species less resistant to predation. The presence of multiple herbivore species of similar size in African savannas allows the top predator, the lion, to shift its prey selection flexibly dependent on changing prey vulnerability. Hence top–down and bottom–up influences on herbivore populations are intrinsically entangled. Models coupling the population dynamics of predators and prey need to accommodate the changing influences of prey demography, temporal variation in environmental conditions, and spatial variation in the relative vulnerability of alternative prey species to predation. Synthesis While re‐established predators have had major impacts on prey populations in northern temperate regions, multiple large herbivore species typically coexist along with diverse carnivores in African savanna ecosystems. In order to explain these contrasting outcomes, certain functional heterogeneities must be recognised, including relative vulnerability of alternative prey, temporal variation in the risk of predation, demographic differences in susceptibility to predation, and spatial contrasts in exposure to predation. Food shortfalls prompt herbivores to exploit more risky habitats, meaning that top–down and bottom–up influences on prey populations are intrinsically entangled. Models coupling the interactive dynamics of predator and prey populations need to incorporate these varying influences on relative prey vulnerability.     

Spatial variation in vegetation damage relative to primary productivity,small rodent abundance and predation

The relative importance of top‐down and bottom‐up mechanisms in shaping community structure is still a highly controversial topic in ecology. Predatory top‐down control of herbivores is thought to relax herbivore impact on the vegetation through trophic cascades. However, trophic cascades may be weak in terrestrial systems as the complexity of food webs makes responses harder to predict. Alternatively, top‐down control prevails, but the top‐level (predator or herbivore) changes according to productivity levels. Here we show how spatial variation in the occurrence of herbivores (lemmings and voles) and their predators (mustelids and foxes) relates with grazing damage in landscapes with different net primary productivity, generating two and three trophic level communities, during the 2007 rodent peak in northern Norway. Lemmings were most abundant on the unproductive high‐altitude tundra, where few predators were present and the impact of herbivores on vegetation was strong. Voles were most common on a productive, south facing slope, where numerous predators were present, and the impacts of herbivores on vegetation were weak. The impact of herbivores on the vegetation was strong only when predators were not present, and this cannot be explained by between‐habitat differences in the abundance of plant functional groups. We thus conclude that predators influence the plant community via a trophic cascade in a spatial pattern that support the exploitation ecosystems hypothesis. The responses to grazing also differed between plant functional groups, with implications for short and long‐term consequences for plant communities.     

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.     

Density dependence,prey accessibility and prey depletion by fisheries drive Peruvian seabird population dynamics

In marine ecosystems top predator populations are shaped by environmental factors affecting their prey abundance. Coupling top predators’ population studies with independent records of prey abundance suggests that prey fluctuations affect fecundity parameters and abundance of their predators. However, prey may be abundant but inaccessible to their predators and a major challenge is to determine the relative importance of prey accessibility in shaping seabird populations. In addition, disentangling the effects of prey abundance and accessibility from the effects of prey removal by fisheries, while accounting for density dependence, remains challenging for marine top predators. Here, we investigate how climate, population density, and the accessibility and removal of prey (the Peruvian anchovy Engraulis ringens) by fisheries influence the population dynamics of the largest sedentary seabird community (≈ 4 million individuals belonging to guanay cormorant Phalacrocorax bougainvillii, Peruvian booby Sula variegata and Peruvian pelican Pelecanus thagus) of the northern Humboldt Current System over the past half‐century. Using Gompertz state–space models we found strong evidence for density dependence in abundance for the three seabird species. After accounting for density dependence, sea surface temperature, prey accessibility (defined by the depth of the upper limit of the subsurface oxygen minimum zone) and prey removal by fisheries were retained as the best predictors of annual population size across species. These factors affected seabird abundance the current year and with year lags, suggesting effects on several demographic parameters including breeding propensity and adult survival. These findings highlight the effects of prey accessibility and fishery removals on seabird populations in marine ecosystems. This will help refine management objectives of marine ecosystems in order to ensure sufficient biomass of forage fish to avoid constraining seabird population dynamics, while taking into account of the effects of environmental variability.