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.     

Genetic variability, body characteristics and reproductive parameters of neighbouring rural and urban common kestrel (Falco tinnuculus) populations

We tested the genetic and ecological differences between neighbouring urban and rural populations of common kestrels (Falco tinnuculus) in southern Bohemia. The aims were to (1) assess the genetic variability of the studied kestrel populations using microsatellite markers, (2) check the genetic relatedness of individuals within the urbanization gradient, and (3) compare possible gradients of body characteristics and reproductive parameters on the urbanization gradient. The mean expected allelic polymorphism did not differ among the studied populations, which were not genetically separated (F _ST  = 0.0003, P = 0.781). Also, an individual assignment test did not show a separation of these populations. Urban kestrels that bred in the city centre were indicatively more related than others, and no relationship was found in the rural kestrel population. Kestrel females were heavier towards the city centre, but males did not show this relationship. Nest distance from the city centre had no significant effect on any of the tested reproductive parameters. Our results do not support the notion of genetic differentiation between rural and urban kestrels, but revealed trends in body characteristics and genetic relatedness along the urbanization gradient.     

Patterns of Orthoptera abundance and lesser kestrel conservation in arable landscapes

The lesser kestrel Falco naumanni experienced a marked decline during the second half of the 20th century due to changes in land use that influenced breeding success by reducing the abundance and quality of prey. However, the factors governing spatial and temporal variation of prey abundance around lesser kestrel colonies has not yet been investigated. We sampled Orthoptera abundance in the main crop types and edge habitats surrounding six lesser kestrel colonies in southern Spain. Samplings focused on Orthoptera because they constitute the main prey during the nestling period. Only those Orthoptera species that are known to be preyed by lesser kestrels were considered in this study. We found differences in prey density among localities, and crop types. Semi-natural habitats such as grasslands, fallow land, and field margins held the highest densities. However, prey abundance showed a complex pattern that was not possible to explain solely on the basis of crop composition around colonies. Factors determining productivity in individual fields like soil type and productivity or biocide input, and mean size of agricultural fields contributed to explain this complex pattern of prey abundance. Our results highlight the key role of semi-natural and edge habitats in farmed landscapes as prey reservoirs and corridors. Higher conservation priorities for these habitats are suggested to benefit foraging lesser kestrels, but many other farmland species that also experienced steep population declines due to decreasing food supply resulting from modern agriculture.     

Evidence of prey depletion around lesser kestrel Falco naumanni colonies and its short term negative consequences

Predation may reduce prey numbers in such extent that prey may be depleted, which has negative effects on predator populations. Prey depletion would be more likely when the number of predators increase and/or concentrate their activity in a certain area, as is the case of colonial birds. As a matter of fact, food depletion due to intraspecific competition is considered a major cost of coloniality, and several studies have shown indirect evidence of this. However, no direct measures of food depletion have been provided along with its consequences for the fitness of the colony inhabitants. We carried out a field study with the lesser kestrel Falco naumanni, a raptor that breeds in colonies ranging from two to dozens of pairs. During the nestling period we sampled the main prey of the kestrels around different sized colonies at increasing distances. At the same time, we recorded hunting distances and prey delivery rates to the nest. In addition, we monitored the reproductive success in colonies of different sizes. Lesser kestrels feed their nestlings mainly with grasshoppers and these prey became depleted through the season in the surroundings of the large colonies. Prey depletion made kestrels fly longer distances to forage and prey delivery rates to the nest decreased. Lower feeding rates were not compensated by bringing larger prey, hence, the net amount of energy provided to the chicks decreased with the date in large colonies. By contrast, none of this occurred around small colonies, where both prey abundance and hunting distance remained constant throughout the season. As a consequence, the seasonal decline in the reproductive success (number of fledglings and fledgling body condition) was greater the larger the colony. Thus, these results evidence that food depletion and its fitness costs are related to colony size, as they are suffered by the kestrels breeding in large colonies but not by those settled at small ones. Finally, the consequences of prey depletion on the demographic dynamics and the regulation of colony size are discussed.     

Sparrowhawk Accipiter nisus presence and winter bird abundance

The decline and subsequent re-colonisation of the Sparrowhawk Accipiter nisus in the UK has allowed an examination of its potential impacts on passerine abundance through analysis of long-term garden bird monitoring data. Weekly counts in gardens in winter have been collected annually since the early 1970s, a period when Sparrowhawks were initially rare but recolonised to a point in the 1990s when they were recorded in the majority of gardens. Change in bird count from year-to-year for 10 common prey species from up to 483 survey sites were analysed in relation to Sparrowhawk re-colonisation, the number of feeding units per site (e.g. bird tables, hanging bird feeders) and, for a subset of the data, minimum temperature. Year-to-year change increased with Sparrowhawk re-colonisation in Great Tit Parus major, Blue Tit Cyanistes caeruleus and Collared Dove Stretopelia decaocto, but decreased in Starling Sturnus vulgaris, Chaffinch Fringilla coelebs and Greenfinch Carduelis chloris. Positive associations may suggest a behavioural response to predators, or an attraction of predators to increased prey caused by confounding effects of improvement in site quality. The significant negative effects detected suggest that there is a temporal matching between increased predation pressure and decreases in bird abundance for certain species. However, there was no strong evidence that Sparrowhawk re-colonisation was linked to long-term declines in population size.

Breeding success of an endangered island endemic kestrel increases with extent of invasion by an alien plant species

The Mauritius Kestrel Falco punctatus, once the rarest kestrel worldwide, became an icon of bird conservation after it recovered from four to six individuals in 1974 to some 800 by 2005 following intense conservation management. Its population however then halved within about a decade prompting a re-evaluation of the IUCN status and up listing of the species in 2014 and an increased conservation attention. Drivers of this new decline are unclear and the influence of habitat structure and diet on breeding success may be important contributors but have received relatively little attention, particularly in the way they may interact to influence production of new fledglings. We address this knowledge gap by studying whether breeding success is influenced by habitat structure (in terms of cover of the invasive Ravenala in native habitats, an alien plant causing strong structural shift in the forests that it invades, and extent of cleared area), diet composition and food pass frequency (as a proxy for food intake) and food quality at 28 nests of a re-introduced kestrel population in south east Mauritius during the 2015–2016 breeding season. The kestrel’s diet comprised native and alien birds, reptiles, insects, and small alien mammals, with a disproportionately high proportion of Phelsuma gecko. A higher frequency of food provisioning and percentage cover of Ravenala both contributed to higher breeding success. Ravenala may increase gecko density or increase gecko detectability and predation by the kestrel, or both, while changed land use (pasture and sugar cane fields) may increase prey diversity in the form of non-forest prey known to be eaten by Kestrels (e.g. alien agamids, small mammals and birds). These prey related influences on breeding suggest that the Bambou mountain range provides a human-generated novel ecosystem altering food availability and increasing the kestrel’s breeding success. However, Ravenala is an invasive alien species harmful to the wider forest biodiversity. Progressive weeding of Ravenala and concurrent re-introduction and augmentation of native palms and Pandanus species which geckos can use at comparable densities to Ravenala, is recommended. This would likely improve the kestrel’s hunting habitat quality and maintain high gecko density or detectability and the vegetation structure required for hunting manoeuvrability and prey availability without the negative consequences of invasive Ravenala.     

Predator presence may benefit: kestrels protect curlew nests against nest predators

We studied whether the presence of breeding kestrels (Falco tinnunculus) affected nest predation and breeding habitat selection of curlews (Numenius arquata) on an open flat farmland area in western Finland. We searched for nests of curlews from an area of 6 km² during 1985–1993. For each nest found, we recorded the fate of the nest, and the distance to the nearest kestrel nest and to the nearest perch. We measured the impact of breeding kestrels on nest predation by constructing artificial curlew nests in the vicinity of ten kestrel nests in 1993. Curlew nests were closer to kestrel nests than expected from random distribution, eventhough kestrels fed on average 5.5% of curlew chick production. Predation risk by kestrels was lower than predation risk by corvids and other generalist predators, which predated 9% of curlew nests surviving farming practices and an unknown proportion of chicks. Artificial nest experiment showed that nest predation was lower close to kestrel nests than further away suggesting that the breeding association of curlews and kestrels was a behavioural adaptation against nest predation. Thus, the presence of a predator may sometimes be beneficial to prey, and prey animals have behavioural adaptations to these situations.     

Identification of trophic interactions between <Emphasis Type="Italic">Macrolophus caliginosus</Emphasis> (Heteroptera: Miridae) and <Emphasis Type="Italic">Myzus persicae</Emphasis> (Homoptera: Aphididae) using real time PCR

Understanding food web interactions in native or agricultural ecosystems is an important step towards establishing sustainable pest management strategies. While the role of generalist predators as biological control agents is increasingly appreciated, the study of trophic interactions between individual predator species and their prey provides practical difficulties. Recently, different approaches have been suggested to determine prey items from predator guts using molecular methods. Macrolophus caliginosus is a generalist predator active in herbaceous agro-ecosystems. We developed a system to identify the DNA of its prey after ingestion, using Myzus persicae as a model. Esterase (MpEST) and cytochrome oxidase I (MpCOI) genes were targeted in the aphid, while M. caliginosus COI gene was used as control for predator DNA. Real time PCR proved to be specific and sensitive enough to detect prey DNA upon ingestion after feeding experiments. The system provided a linear amplification response with only 10 fg of prey genomic DNA as template. The detection system of MpCOI gene was more sensitive than MpEST, while the detection period was similar for both genes. Possibilities for using the system in ecological and biosafety studies with regard to sustainable pest management are discussed.

Consumption of aphids by spiders and the effect of additional prey: evidence from microcosm experiments

Spiders are common generalist predators in agroecosystems and have been suggested to lower herbivore abundance in crops. It is not clear, however, if spiders can effectively suppress pest populations, and if so, by what mechanisms. In a microcosm experiment, we examined the consumption of the bird cherry-oat aphid, Rhopalosiphum padi L. (Homoptera: Aphididae), a pest species in wheat fields, by three spider species that differ in their hunting methods. We then tested the effect of additional prey type on the ability of erigonid spiders to reduce aphids. In a 48-h experiment Mermessus denticulatus (Banks) (Araneae: Linyphiidae; Erigoninae) consumed more aphids than did Enoplognatha gemina Bosmans and Van Keer (Araneae: Theridiidae) and Bathyphantes cf. extricatus (O·P.-Cambridge) (Araneae: Linyphiidae; Linyphiinae). This difference may be due to the ability of erigonids to forage actively on the vegetation in addition to using their webs to catch prey. In a 7-week experiment, we provided springtails (Collembola) in high and low densities as additional prey to mated erigonids, prior to aphid introduction. Spiders in the low-density springtail treatment built more webs on the vegetation, and caused a 50% reduction in aphid populations. There were significantly fewer aphids in the low-density springtail treatment, but not in the high-density treatment, in comparison to the control (high-density springtails without spiders). The results suggest that additional prey density affects predatory interactions between M. denticulatus and R. padi and that erigonids, which occur in high densities in wheat fields in the Negev desert, may be involved in aphid suppression in these agroecosystems.

Reproductive responses of birds to experimental food supplementation: a meta-analysis

Introduction

Food availability is an important environmental cue for animals for deciding how much to invest in reproduction, and it ultimately affects population size. The importance of food limitation has been extensively studied in terrestrial vertebrate populations, especially in birds, by experimentally manipulating food supply. However, the factors explaining variation in reproductive decisions in response to food supplementation remain unclear. By performing meta-analyses, we aim to quantify the extent to which supplementary feeding affects several reproductive parameters in birds, and identify the key factors (life-history traits, behavioural factors, environmental factors, and experimental design) that can induce variation in laying date, clutch size and breeding success (i.e., number of fledglings produced) in response to food supplementation.

Results

Food supplementation produced variable but mostly positive effects across reproductive parameters in a total of 201 experiments from 82 independent studies. The outcomes of the food effect were modulated by environmental factors, e.g., laying dates advanced more towards low latitudes, and food supplementation appeared not to produce any obvious effect on bird reproduction when the background level of food abundance in the environment was high. Moreover, the increase in clutch size following food addition was more pronounced in birds that cache food, as compared to birds that do not. Supplementation timing was identified as a major cause of variation in breeding success responses. We also document the absence of a detectable food effect on clutch size and breeding success when the target species had poor access to the feed due to competitive interactions with other animals.

Conclusions

Our findings indicate that, from the pool of bird species and environments reviewed, extra food is allocated to immediate reproduction in most cases. Our results also support the view that bird species have evolved different life-history strategies to cope with environmental variability in food supply. However, we encourage more research at low latitudes to gain knowledge on how resource allocation in birds changes along a latitudinal gradient. Our results also emphasize the importance of developing experimental designs that minimise competition for the supplemented food and the risk of reproductive bottle-necks due to inappropriate supplementation timings.

     

Realistic threshold policy with hysteresis to control predator–prey continuous dynamics

This paper introduces a threshold policy with hysteresis (TPH) for the control of one-predator one-prey models. The models studied are the Lotka–Volterra and Rosenzweig–MacArthur two species density-dependent predator–prey models and the Arditi–Ginzburg nondimensional ratio-dependent model. The proposed policy (TPH) changes the dynamics of the system in such a way that a bounded oscillation is achieved confined to a region that does not allow extinction of either species. The policy can be designed by a suitable choice of so called virtual equilibrium points in a simple and intuitive manner.

Birth Order Influences Reproductive Measures in Australians

We examine the relationship between birth order and reproductive behaviors in a sample of Australian residents, accounting for personality, personal achievements, and family structure. Using generalized linear models and survival analyses we build predictive models for each reproductive measure and test those models on an independent data subset. Compared with functional firstborns (middle-borns more than 5 years younger than their next older sibling), male middle-borns and last-born females had younger ages of first sexual intercourse, and middle-born females had a younger age at first pregnancy. There was no difference in females’ age at first birth. Male middle-borns tended to have an older average age at first birth. Controlling for age, both male and female middle-borns had fewer children. Overall, middle-borns differ more from functional firstborns than do last-borns. Given the significant but small effect sizes demonstrated in this study, we suggest that developmental characteristics that may facilitate middle-borns’ success within the natal family carry slight fitness costs during the reproductive years.

The Behavioral Ecology of Shellfish Gathering in Western Kiribati,Micronesia 1: Prey Choice

Focusing on contemporary shellfish exploitation among several atoll communities in Kiribati, Micronesia, this paper examines the relationship between human foragers and their invertebrate prey via the prey choice or diet breadth model derived from optimal foraging theory. Shellfish, like many other reef organisms, are relatively sedentary and predictable, but these characteristics make them susceptible to over-harvesting. The research reveals that shellfish gatherers are foraging in a manner that matches the predictions of optimal foraging theory. The work adds to our understanding of optimal foraging decisions in atoll settings by critically evaluating the depiction of atoll dwellers as conservationists.

Avian mitochondrial DNA sequence divergence across a headwater stream of the Rio Tapajós, a major Amazonian river

Populations of ten Amazonian bird species were sampled on opposite banks of the Rio Teles Pires, a headwater stream of the Rio Tapajós, in the Alta Floresta region, northern Mato Grosso, Brazil. The river is 100–300 m wide in this region. We found a range of genetic differentiation from none to relatively high levels; six of the ten species studied exhibit what appear to be genetic breaks at the river. With one exception, the antbird Hylophylax poecilinota, there is no morphologically recognized differentiation correlating with genetic differentiation. From the perspective of traditional morphology-based taxonomy, the Rio Teles Pires is not a faunal barrier. Rather, contact zones between members of species and subspecies pairs appear more or less randomly distributed in this region, some being located at varying distances to the east, others at varying distances to the west of the Rio Teles Pires, with few following the course of this river itself.

Trophic niche width,offspring condition and immunity in a raptor species

Strategies developed by organisms to maximize foraging efficiency have a strong influence on fitness. The way in which the range of food resources is exploited has served to classify species, populations and individuals from more specialist (narrow trophic niche) to more generalist (broad trophic niche). Recent studies have provided evidence that many of the considered generalist species/populations are actually composed of different specialist individuals (individual specialization). Even the existence of generalism as an adaptive strategy has been questioned. In this study, we investigated the relationship between trophic niche width, individual quality and offspring viability in a population of common kestrel Falco tinnunculus during 4 years. We showed that the diet of kestrels varied significantly among years and that individuals of better quality fed their offspring with a higher diversity of prey species and a higher amount of food. Moreover, body condition and immune response of nestlings were positively correlated with diversity of prey delivered by parents. Our study suggests that generalism has the potential to increase fitness and that broadening the trophic niche may be an adaptive strategy in unpredictable environments.     

Otters and fish farms in the Sado estuary: ecological and socio-economic basis of a conflict

The degree of conflict between otter conservation and fish farming was assessed at Sado estuary (SW Portugal), using ecological (otter visiting rates to fish farms and consumption of commercial fish) and socio-economic (past and current instruments and policies addressing the conflict and a social impact assessment, including a discourse analysis of relevant stakeholders) parameters. The study concerned 14 fish farms producing Sparus aurata, Dicentrarchus labrax, Solea senegalensis and Solea solea. Results indicate high visiting rates in most fish farms (average: 76%), although in only 29%, species stocked were the most consumed prey. Other marine species and freshwater prey were the basis of otter diet in the remaining fish farms. The conflict is quite consensual among most fish farmers, and the results indicate that the perceived conflict by fish farmers has an ecological basis, although there are large gaps between effective and perceived predation. No specific instruments exist in Portugal to address the conflict, but some not specifically targeted can have an effect (e.g. species protection legislation and aquaculture licensing), although with limitations to effectively contribute to its mitigation (e.g. lack of enforcement and supervision). Formulating and assessing solutions is the following step, using a participatory approach to the development and evaluation of mitigation/compensation strategies, capable of providing an effective reconciliation of the conflict.

Prey size, prey nutrition, and food handling by shrews of different body sizes

We tested some predictions relating metabolic constraints offoraging behavior and prey selection by comparing food handlingand utilization in four sympatric shrew species: Sorex minutus(mean body mass = 3.0 g), S. araneus (8.0 g), Neomys anomalus(10.0 g), and N. fodiens (14.4 g). Live fly larvae, mealwormlarvae, and aquatic arthropods were offered to shrews as smallprey (body mass <0.1 g). Live earthworms, snails, and smallfish were offered as large prey (>0.3 g). The larvae werethe high-nutrition food (>8 kJ/g), and the other prey werethe low-nutrition food (<4 kJ/g). The smallest shrew, S.minutus, utilized (ate + hoarded) <30% of offered food,and the other species utilized >48% of food. The largerthe shrew, the more prey it ate per capita. However, highlyenergetic insect larvae composed 75% of food utilized by S.minutus and only >40% of the food utilized by the other species. Thus, inverse relationships appeared between shrewbody mass and mass-specific food mass utilization and betweenshrew body mass and mass-specific food energy utilization:the largest shrew, N. fodiens, utilized the least food massand the least energy quantity per 1 g of its body mass. Also,the proportion of food hoarded by shrews decreased with increase in size of shrew. With the exception of S. araneus, the sizeof prey hoarded by the shrews was significantly larger thanthe size of prey eaten. Tiny S. minutus hoarded and ate smallerprey items than the other shrews, and large N. fodiens hoardedlarger prey than the other shrews.     

What Explains Differences in Men’s and Women’s Production?

Researchers commonly use long-term average production inequalities to characterize cross-cultural patterns in foraging divisions of labor, but little is known about how the strategies of individuals shape such inequalities. Here, we explore the factors that lead to daily variation in how much men produce relative to women among Martu, contemporary foragers of the Western Desert of Australia. We analyze variation in foraging decisions on temporary foraging camps and find that the percentage of total camp production provided by each gender varies primarily as a function of men’s average bout successes with large, mobile prey. When men target large prey, either their success leads to a large proportional contribution to the daily harvest, or their failure results in no contribution. When both men and women target small reliable prey, production inequalities by gender are minimized. These results suggest that production inequalities among Martu emerge from stochastic variation in men’s foraging success on large prey measured against the backdrop of women’s consistent production of small, low-variance resources.

The heterospecific habitat copying hypothesis: can competitors indicate habitat quality?

According to the "habitat copying" hypothesis, animals use thereproductive performance of conspecifics to assess habitat suitabilityand choose their future breeding site. This is because conspecificsshare ecological needs and thus indicate habitat suitability.Here, we propose the "heterospecific habitat copying" hypothesis,which states that animals should use public information (i.e.,information derived from the performance of others) from con-and heterospecifics sharing ecological needs. In a correlationalapproach we test some assumptions and predictions of this hypothesiswith a data set from two sympatric bird populations, rollers(Coracias garrulus) and kestrels (Falco tinnunculus), usingthe same nest-boxes and exploiting similar food resources. Sincekestrels are residents and breed earlier, we assumed that theyare dominant over rollers for nest-box acquisition. The environmentappears to be patchy for both species and temporally predictablefor kestrels only. Two results suggest that the use of heterospecificpublic information in breeding habitat selection may be at work:(1) an increase in the reoccupancy probability by kestrels ofprevious roller nests with increasing nest success, and (2)an increase in roller breeding population with increasing localkestrel success. Most of the other observed patterns could beexplained by alternative mechanisms such as natal philopatry,breeding fidelity, conspecific attraction, intraspecific habitatcopying, and the effect of interspecific competition.     

Impact of spatial variation of a crucial prey,the molecricket,on hoopoe territory occupancy and reproduction

Direct benefits accrued from securing a territory of sufficient quality are important determinants of individual fitness and population persistence. Food supply is one of the main factors of animal territory quality, with spatial and temporal variation in prey availability largely dictating reproductive output and thus population dynamics. In a Swiss hoopoe population, molecrickets Gryllotalpa gryllotalpa, the most profitable prey locally, can constitute most of the food biomass delivered to chicks by parents. We first investigated the impact of molecricket prey on hoopoes' fitness‐related traits by quantifying the spatial variation in the food allocation pattern of both male and female parents to chicks across the whole population range; and second, analysed the impact of this prey on current reproduction and, using a 11 yr dataset, on the temporal occupancy rate of each territory. We found considerable but spatially repeatable variation, over the years, of molecricket biomass in the diet provisioned to chicks. This spatial heterogeneity in chicks' diet composition was mirrored both in the history of territory occupancy (2002–2012) and in current reproductive success (2012). Territories with a greater biomass of molecrickets in chicks' diet produced more fledglings in better body condition. Yet, these effects on current reproduction were exclusively demonstrated for male parents, corroborating that paternal provisioning patterns play a predominant role in hoopoe reproductive success. This study demonstrates how a single, very profitable prey species might affect spatial variation in territory settlement and individual reproductive success in a regionally endangered bird species, with potential consequences for its population dynamics and persistence.