Simultaneous effects of age and territory quality on fecundity in Bonelli's Eagle Hieraaetus fasciatus

Several factors that interact with a bird's age are likely to affect breeding performance. Because of habitat heterogeneity, individuals of different subpopulations may experience different probabilities of breeding, depending on the habitat that they occupy. The aim of this work is to test the hypothesis that age and territory quality independently affect breeding performance. We analysed data on breeding biology and fecundity at 298 breeding sites of Bonelli's Eagle Hieraaetus fasciatus in Andalusia from 1980 to 2000. Our data confirmed that age and territory quality simultaneously affect reproductive output. After controlling for the effect of year and age, breeding performance varied among territories. Independently of territory quality, the age of breeding birds affected the number of fledged young and variance in productivity.     

Population size,breeding performance and territory quality of Bonelli's Eagle Hieraaetus fasciatus in eastern Spain

Capsule A five-year monitoring study is described of overall population stability and differential reproductive success in relation to habitat heterogeneity.

Aims To assess the effect of the altitude and orientation on laying date and breeding performance, and analyse the effect of territory quality as a likely factor that could be regulating the population.

Methods We monitored a population of 28–33 pairs, from 2002 to 2006, counting a total of 131 breeding attempts.

Results Territories located at lower altitude showed higher mean fecundity than those located at higher altitude. The mean laying date was February 18 ± 16 days. Laying date was positively correlated with nest altitude, the coastal pairs laying earlier than those in mountainous regions. Pairs located at lower altitudes showed higher mean fecundity than those located at higher altitudes. There was no preference in mean orientation either in breeding performance or in relationship to nest altitude. We did not find a difference in breeding performance between territories classed as being at high density and those classed as being at low density.

Conclusion The population has remained stable since the first national census was conducted 17 years ago. Our results could be explained in the light of the Habitat Heterogeneity Hypothesis. We suggest a differential reproductive success in relation to habitat heterogeneity.     

Empirical support for a despotic distribution in a California spotted owl population

Territorial species, such as the spotted owl (Strix occidentalis),are predicted to follow an ideal despotic distribution. However,debate exists on whether wild populations actually meet theassumptions of an ideal distribution, such as perfect perceptualabilities (i.e., the ability to recognize high- and low-qualitysites without error). Because this hypothesis has importantlife history ramifications for spotted owls, we investigatedwhether occupancy rates of California spotted owl (S. o. occidentalis)territories in the San Bernardino Mountains of southern Californiapositively correlated with a qualitative "potential fitness"(denoted by _pf) estimated from survival and reproduction ofterritorial owls. Spotted owls in our study tended to occupyterritories with the highest _pf, supporting the assumption ofideal perceptual abilities within this population. However,this relationship was noisy, and we suggest that some individualsdo not assess site quality accurately because of perceptuallimitations, prey dynamics, and large territory sizes. Furthermore,dispersal processes, high survival rates, and long life spansof spotted owls may be other key factors preventing some individualsfrom selecting sites of the highest quality and, consequently,our ability to precisely estimate _pf.     

Factors influencing the decline of a Bonelli's eagle Hieraaetus fasciatus population in southeastern Spain: demography, habitat or competition?

We investigated three possible causes of territory desertion among Bonelli's eagles Hieraaetus fasciatus in Murcia (southeastern Spain): low demographic parameters, low habitat quality and competition with Golden eagles Aquila chrysaetos. From 1983 to 1997, we surveyed a Bonelli's eagle population. Abandoned and occupied territories were compared to find differences in demographic parameters (flight rate, productivity and mortality) or habitat characteristics. Mortality was significantly higher in abandoned territories. Abandoned territories also had larger areas of forest and extensive agriculture, while occupied territories had more shrublands. Competition with Golden eagles was not a determinant of territorial abandonement but interacted with human persecution of the species. Management implications are discussed.     

Competition for safe real estate,not food,drives density‐dependent juvenile survival in a large herbivore

Density‐dependent competition for food reduces vital rates, with juvenile survival often the first to decline. A clear prediction of food‐based, density‐dependent competition for large herbivores is decreasing juvenile survival with increasing density. However, competition for enemy‐free space could also be a significant mechanism for density dependence in territorial species. How juvenile survival is predicted to change across density depends critically on the nature of predator–prey dynamics and spatial overlap among predator and prey, especially in multiple‐predator systems. Here, we used a management experiment that reduced densities of a generalist predator, coyotes, and specialist predator, mountain lions, over a 5‐year period to test for spatial density dependence mediated by predation on juvenile mule deer in Idaho, USA. We tested the spatial density‐dependence hypothesis by tracking the fate of 251 juvenile mule deer, estimating cause‐specific mortality, and testing responses to changes in deer density and predator abundance. Overall juvenile mortality did not increase with deer density, but generalist coyote‐caused mortality did, but not when coyote density was reduced experimentally. Mountain lion‐caused mortality did not change with deer density in the reference area in contradiction of the food‐based competition hypothesis, but declined in the treatment area, opposite to the pattern of coyotes. These observations clearly reject the food‐based density‐dependence hypothesis for juvenile mule deer. Instead, our results provide support for the spatial density‐dependence hypothesis that competition for enemy‐free space increases predation by generalist predators on juvenile large herbivores.     

Sequential settlement and site dependence in a migratory raptor

Habitat selection models, such as the ideal free, ideal despotic,site-dependent, and conspecific cuing models, are of great importanceto behavioral ecologists given their capability to predict habitatdistributions and to link individual behavior to populationprocesses. However, there have been relatively few field testsof their predictions. We tested the 4 models by studying theprocess of sequential settlement on territory in 2 distant populationsof a migratory raptor, the black kite Milvus migrans. Resultswere mainly consistent with the site-dependent model: on arrival,kites settled on progressively lower quality territories, andearlier arriving individuals were older, larger, and in betterbody condition than later arriving ones, leading to a state-dependentarrival sequence also predicted by a previous theoretical modelof settlement pattern. Occupation of superior territories bysuperior phenotypes resulted in cascading advantages for earlierarriving individuals in terms of subsequent reproductive performance.At the population level, the populations expanded/retractedfrom lower quality sites during population increases/declines.The above scenario was consistent across the 2 populations,and a review of the literature uncovered a remarkably consistentpicture of state-dependent arrival, progressive monopolizationof best quality sites, and cascading effects on subsequent breedingperformance. We propose as a general paradigm of sequentialsettlement the following process: 1) arrival date is a reliablesurrogate of phenotype quality, 2) early-arriving individualshave preferential access to the best quality sites and partners,3) the above conditions cascade into a number of benefits ultimatelyrelated to higher fitness for earlier arriving individuals.     

Territorial occupancy and breeding performance in a migratory raptor do not follow ideal despotic distribution patterns

It has commonly been argued that many territorial species select their breeding sites following an ideal despotic distribution model, in which the most productive, high-quality territories are more frequently occupied. Theoretical and empirical studies have shown that this occupancy pattern may have population regulatory consequences, leading to density dependence in heterogeneous habitats. During a 9-year research project in a forested area of south-eastern Spain, we tested some of the predictions of the ideal despotic distribution model and the site-dependent population regulation theory in a migratory raptor species, the booted eagle Hieraaetus pennatus . Contrary to the predictions of the despotic model, our results showed that the temporal pattern of territorial occupation did not differ from randomness, and that the territory occupancy rate was not significantly related to the reproductive parameters considered. At population level, the breeding variables were density independent, suggesting the absence of site-dependent regulation. In addition, we were unable to find significant differences in the habitat characteristics between high-quality and low-quality territories, classified according to the criteria of both occupancy frequency and average productivity. Overall, our results suggest that booted eagles select their territories at random, probably due to the lack of strong environmental heterogeneity, and that occupancy rate is not a good measure of territory quality for the population studied.     

Interacting effects of temperature and density on individual growth performance in a wild population of brown trout

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Density‐dependent effects on reproductive performance in a recovering population of White‐tailed Eagles Haliaeetus albicilla

Understanding the mechanisms that shape density‐dependent processes and population dynamics is often essential for species conservation. Two key mechanisms of density‐dependent reductions in reproductive performance are a limited access to foraging habitats (the habitat heterogeneity hypothesis) and territorial aggression towards conspecifics (the interference competition hypothesis) at high population densities. Disentangling the relative importance of these mechanisms within populations below their carrying capacity is important for the evaluation of the success of conservation measures. However, relatively few studies have attempted to quantify the relative importance of both mechanisms for the reproductive performance of a population. Many raptor populations are ideal model systems to investigate density‐dependent effects because they are currently recovering from human‐induced reductions during the last decades. Using a 14‐year dataset, we combined analyses of individual reproductive performance with a mechanistic population model to investigate early signs of density‐dependent regulation in a population of White‐tailed Eagles Haliaeetus albicilla in north‐east Germany. We found a negative effect of the number of neighbouring breeding pairs and a positive effect of water surface area (as a proxy for the availability of favourable foraging habitat) on breeding success and on the average number of nestlings. The mean nearest neighbour distance between breeding pairs has decreased, and the mean distance of nests to the nearest water body has increased over the last 14 years. Moreover, the population model indicates that even though the population is still growing, carrying capacity could be reached at about 500–950 territorial pairs. These results suggest that the selection of nesting sites is determined by a trade‐off between the distance to favourable foraging habitat and the distance to neighbouring breeding pairs. To avoid increasing competition with conspecifics, due to continued population growth, breeding pairs seem to select increasingly suboptimal habitats. Therefore, our results suggest that the habitat heterogeneity and interference competition hypotheses are not necessarily mutually exclusive as mechanisms of density‐dependent population regulation, but can determine the reproductive performance of a raptor population simultaneously. Thus, a future decline in breeding success does not necessarily reflect a decrease in habitat quality but may rather be a consequence of density‐dependent mechanisms. This information may be useful for the interpretation of population trends and for the development of appropriate management strategies for recovering raptor populations.     

Linking warming effects on phenology,demography, and range expansion in a migratory bird population

Phenological changes in response to climate change have been recorded in many taxa, but the population‐level consequences of these changes are largely unknown. If phenological change influences demography, it may underpin the changes in range size and distribution that have been associated with climate change in many species. Over the last century, Icelandic black‐tailed godwits (Limosa limosa islandica) have increased 10‐fold in numbers, and their breeding range has expanded throughout lowland Iceland, but the environmental and demographic drivers of this expansion remain unknown. Here, we explore the potential for climate‐driven shifts in phenology to influence demography and range expansion. In warmer springs, Icelandic black‐tailed godwits lay their clutches earlier, resulting in advances in hatching dates in those years. Early hatching is beneficial as population‐wide tracking of marked individuals shows that chick recruitment to the adult population is greater for early hatched individuals. Throughout the last century, this population has expanded into progressively colder breeding areas in which hatch dates are later, but temperatures have increased throughout Iceland since the 1960s. Using these established relationships between temperature, hatching dates and recruitment, we show that these warming trends have the potential to have fueled substantial increases in recruitment throughout Iceland, and thus to have contributed to local population growth and expansion across the breeding range. The demographic consequences of temperature‐mediated phenological changes, such as the advances in lay dates and increased recruitment associated with early hatching reported here, may therefore be key processes in driving population size and range changes in response to climate change.     

The effects and limits of territoriality on population regulation in grey red-backed voles,Clethrionomys rufocanus bedfordiae

The effects of breeding territoriality on the stability of grey red-backed vole (Clethrionomys rufocanus bedfordiae) populations were investigated on a control grid and a grid on which the voles were fed, in an outdoor enclosure in Hokkaido, Japan. Vole populations were monitored by live trapping from 1984 to 1986: (1) Population density was 2–7 times greater on the experimental grid to which food was added than on the control grid. Reproductive output was more closely associated with the difference in density between grids than survival or dispersal (immigration and emigration) rates. (2) The number of adult females and pregnancy rate of the experimental population were significantly greater than those of the control one. The difference in the number of adult females between the populations was greater than that in pregnancy rate. (3) The proportion of successful litters and the number of weanlings per litter were not significantly different between the control and experimental population. (4) Adult females held territories on both the control and experimental grid; they were spaced out more than would be expected from random occupation. The territories overlapped more on the experimental grid than on the control grid. (5) Mean territory size of adult females on the experimental grid was about half of that on the control grid. The territory size was correlated negatively with population density. (6) The proportion of trap sites that were used by adult females was significantly greater on the experimental grid than on the control grid. This suggests that adult females on the experimental grid used the area more extensively. This factor, in association with territory size and overlapping of territory, was also important in causing the difference in the number of adult females between the grids. (7) These results call into question the hypothesis that territoriality stabilizes the density in populations of Clethrionomys.     

Habitat distribution and predation on a western population of terrestrial Leiopelma (Anura: Leiopelmatidae) in the northern King Country,New Zealand

Abstract

A new population of terrestrial Leiopel‐matid frog was discovered in the Whareorino Forest, northern King Country, New Zealand, in 1991. Searches were carried out from June 1991 to December 1993 to determine the species present and to document variation in external morphology, habitat, and local distribution. These confirmed that a terrestrial frog resembling L. archeyi is present in the area, as well as Hochstetter's frog Leiopelma hochstetteri and the introduced Australian hylid frog Litoria aurea.

In Whareorino Forest, the terrestrial Leiopelma was mostly above 500 m altitude and L. hochstetteri above 350 m. The terrestrial Leiopelma occupies sites under rocks and logs in forest. It also occurs in vegetation, such as crown fern Blechnum discolor, tree fern Cyathea smithii, hook grass Uncinia uncinata, and rice grass Microlaena avenacea. Egg clusters of this frog were found in crown fern and tree fern, as well as under stones.

The terrestrial Leiopelma is susceptible to predation by Litoria aurea and rats. This is the first documented evidence of predation on Leiopelma on the New Zealand mainland. The future of this small remnant Leiopelma population is uncertain, and further investigation of the impact of anuran and mammalian predators is needed.     

Relationships between local population persistence, local abundance and regional occupancy of species: distribution patterns of diatoms in boreal streams

Aims We have two aims: (1) to examine the relationship between local population persistence, local abundance and regional occupancy of stream diatoms and (2) to characterize the form of the species–occupancy frequency distribution of stream diatoms. Location Boreal streams in Finland. There were three spatial extents: (1) across ecoregions in Finland, (2) within ecoregions in Finland, and (3) within a single drainage system in southern Finland. Methods Diatoms were sampled from stones (epilithon), sediment (epipelon) and aquatic plants (epiphyton) in streams using standardized sampling methods. To assess population persistence, diatom sampling was conducted monthly at four stream sites from June to October. The relationships between local population persistence, local abundance and regional occupancy were examined using correlation analyses. Results There was a significant positive relationship between local persistence and abundance of diatoms in epilithon, epipelon and epiphyton. Furthermore, local abundance and regional occupancy showed a significant positive relationship at multiple spatial extents; that is, across ecoregions, within ecoregions and within a drainage system. The relationships between occupancy and abundance did not differ appreciably among impacted and near pristine‐reference sites. The occupancy–frequency distribution was characterized by a large number of satellite species which occurred at only a few sites, whereas core species that occurred at most sites were virtually absent. Main conclusions The positive relationship between local population persistence and abundance suggested that a high local abundance may prevent local extinction or that high persistence is facilitated by a high local cell density. High local persistence and local abundance may also positively affect the degree of regional occupancy in stream diatoms. The results further showed that anthropogenic effects were probably too weak to bias the relationship between occupancy and abundance, or that the effects have already modified the distribution patterns of stream diatoms. The small number of core species in the species–occupancy frequency distribution suggested that the regional distribution patterns of stream diatoms, or perhaps unicellular microbial organisms in general, may not be fundamentally different from those described previously for multicellular organisms, mainly in terrestrial environments, although average global range sizes may differ sharply between these two broad groups of organisms.     

The role of food, weather and climate in limiting the abundance of animals

More and more studies are demonstrating that populations of animals ‐ from herbivores to top predators, vertebrates and invertebrates ‐ are limited by their food, and that the availability of this food is dictated by the weather. Satellite monitoring is revealing how cyclic and quasi‐cyclic climatic patterns, like the El Niño Southern Oscillation and the North Atlantic Oscillation, are driving and synchronising these weather‐driven changes in the supplies of food. Changes in the amount of food available operate to limit the abundance of populations largely through their influence on the survival of the very young: the Achilles heel of all populations. Any individual organism struggles to use whatever resources it can get from a mostly inhospitable environment to maximise the proliferation of its genes. Each level of a food chain is thus dependent upon, and pressing hard against the limits set by the one below. The resulting intra‐ and inter‐specific interactions produce a multitude of complex outcomes, that significantly influence the dynamics of populations, but do not determine their ultimate size. There is no density‐dependent regulation of abundance. Intra‐specific competition does not determine the size of populations, it only decides which few individuals gain access to the limited food. Nor do predators regulate their prey. They, too, are limited by their food, and the abundance and quality of food is dictated by the weather.     

The population ecology of Rattus exulans on Tiritiri Matangi Island,and a model of comparative population dynamics in New Zealand

Abstract

On Tiritiri, a small predator-free island in northern New Zealand, kiore (Rattus exulans) were live and snap trapped in grassland and forest. In both habitats, kiore abundance peaked in late summer/autumn. The increase followed a 3 month breeding season during which females produced two to three litters, each averaging 7 young. During the population decline in autumn and winter, animals lost weight. Few bred in the breeding season of their birth and none lived to breed in a second breeding season, so the population consisted of distinct age cohorts. These patterns may relate to a highly seasonal food supply.

Kiore elsewhere in New Zealand show seasonal breeding, but the length of breeding, sexual maturation, and litter size vary. Other studies of kiore in the Pacific show less marked seasonal fluctuations, longer breeding seasons, and smaller litters. We propose a model to explain the variation in rodent demography in New Zealand. The model is based on the seasonal availability of food, along with the modifying influences of predation and dispersal.     

Low twinning rate and seasonal effects on twinning in a fertile population,the Hutterites

This paper analyzes from the mid 18th century to 1987 the birth records of the Dariusleut, one of the three subgroups of the Hutterite population. The aim of this study is to describe several aspects of the twinning rate in a fertile population. The overall rate of twinning was 0.90%:103 twins among all 11492 maternities. The rate peaked at the 7th birth order and at the maternal age of 40 years and over. Until the mid 19th century when the Hutterites lived in Russia, the twinning rate was higher (1.5%), and it decreased during the migration period in the second half of the 19th century (0.7%). After the group had settled in the USA and Canada, the population maintained a twinning rate of 1.0% until 1965. After 1965 the rate decreased to 0.7%, partly due to a decline in fertility among women aged 30 years and over. There was a significant seasonal variation: the twinning rate decreased to 0.5% in May–July compared to 1.2% for the other three seasons during the years up to 1965 (P<0.01), while more recent mothers did not show such a seasonal variation. The incidence of twin births in this population seems to have been influenced by environmental factors, which would change their effect seasonally and secularly.     

A genome scan reveals QTL for growth, fatness, leanness and meat quality in a Duroc-Pietrain resource population

We performed a genome-wide QTL scan for production traits in a line cross between Duroc and Pietrain breeds of pigs, which included 585 F(2) progeny produced from 31 full-sib families genotyped with 106 informative microsatellites. A linkage map covering all 18 autosomes and spanning 1987 Kosambi cM was constructed. Thirty-five phenotypic traits including body weight, growth, carcass composition and meat quality traits were analysed using least square regression interval mapping. Twenty-four QTL exceeded the genome-wide significance threshold, while 47 QTL reached the suggestive threshold. These QTL were located at 28 genomic regions on 16 autosomal chromosomes and QTL in 11 regions were significant at the genome-wide level. A QTL affecting pH value in loin was detected on SSC1 between marker-interval S0312-S0113 with strong statistical support (P < 3.0 x 10(-14)); this QTL was also associated with meat colour and conductivity. QTL for carcass composition and average daily gain was also found on SSC1, suggesting multiple QTL. Seventeen genomic segments had only a single QTL that reached at least suggestive significance. Forty QTL exhibited additive inheritance whereas 31 QTL showed (over-) dominance effects. Two QTL for trait backfat thickness were detected on SSC2; a significant paternal effect was found for a QTL in the IGF2 region while another QTL in the middle of SSC2 showed Mendelian expression.     

Population dynamics of three songbird species in a nestbox population in Central Europe show effects of density,climate and competitive interactions

Unravelling the contributions of density‐dependent and density‐independent factors in determining species population dynamics is a challenge, especially if the two factors interact. One approach is to apply stochastic population models to long‐term data, yet few studies have included interactions between density‐dependent and density‐independent factors, or explored more than one type of stochastic population model. However, both are important because model choice critically affects inference on population dynamics and stability. Here, we used a multiple models approach and applied log‐linear and non‐linear stochastic population models to time series (spanning 29 years) on the population growth rates of Blue Tits Cyanistes caeruleus, Great Tits Parus major and Pied Flycatchers Ficedula hypoleuca breeding in two nestbox populations in southern Germany. We focused on the roles of climate conditions and intra‐ and interspecific competition in determining population growth rates. Density dependence was evident in all populations. For Blue Tits in one population and for Great Tits in both populations, addition of a density‐independent factor improved model fit. At one location, Blue Tit population growth rate increased following warmer winters, whereas Great Tit population growth rates decreased following warmer springs. Importantly, Great Tit population growth rate also decreased following years of high Blue Tit abundance, but not vice versa. This finding is consistent with asymmetric interspecific competition and implies that competition could carry over to influence population dynamics. At the other location, Great Tit population growth rate decreased following years of high Pied Flycatcher abundance but only when Great Tit population numbers were low, illustrating that the roles of density‐dependent and density‐independent factors are not necessarily mutually exclusive. The dynamics of this Great Tit population, in contrast to the other populations, were unstable and chaotic, raising the question of whether interactions between density‐dependent and density‐independent factors play a role in determining the (in) stability of the dynamics of species populations.     

Host-specific variation in Ieaf miner population dynamics: effects on density, natural enemies and behaviour of Stilbosis quadricustatella (Lepidoptera: Cosmopterigidae)

Abstract. 1. We compared high and low density populations of a leaf miner ( Stilbosis quadricustatella (Cham.)) on two host oaks to ascertain mechanisms influencing abundance. High density miner populations occurred on sand live oak, Quercus geminata (Fagaceae), found in homogeneous stands at inland and coastal sites. Quercus nigra , water oak, a patchily distributed inland species, supported a low density leaf miner population.
2. Average foliar nitrogen of Q.geminata was significantly lower than that of Q.nigra , and lad mining period on Q.geminata was correspondingly longer than on Q.nigra .
3. The average leaf area of Q.nigra was significantly greater than that of Q.geminata .
4. Parasitism was significantly greater in Q.geminata miner populations and predation was significantly higher in Q.nigra populations. However, parasitism and predation rates were roughly reciprocal so that overall larval mortality did not differ significantly between hosts.
5. In a laboratory experiment, pupal overwintering survivorship did not differ significantly between moist and dry treatments of the sand and loam soil types that typify Qgeminata and Q.nipra habitats, respectively.
6. Within-leaf miner density on Q.geminata significantly affected larval survivorship, parasitism and predation. Leaf area and within-leaf miner density were positively correlated.
7. We detected no host-patch area or isolation effect on miner densities on coastal Qgeminata and there was no significant gradient of local coastal conditions affecting larval survivorship or natural enemies.
8. Coastal leaf miner densities were significantly higher on oak patch edges than in interiors.     

Habitat fragmentation and population features differently affect fruit predation,fecundity and offspring performance in a non‐specialist gypsum plant

• The effects of habitat fragmentation on plant populations are complex, as it might disrupt many ecological processes, including plant reproduction and plant–animal interactions. Gypsum specialist plants may be resilient to fragmentation due to their evolutionary history in fragmented landscapes, but the effects on non‐specialist plants occurring in gypsum are unknown.
• We conducted a study focusing on different aspects of the reproductive cycle of Astragalus incanus subsp. incanus, a plant facultatively linked to gypsum soils. We focused on plant fecundity and pre‐dispersal predation, obtained from field observations, and offspring performance, assessed in a common garden. Beyond fragment size and connectivity, we also considered habitat quality, population size and density and plant size as predictors.
• Fragment size and connectivity had no effect on plant fecundity, but jointly determined fruit predation, while fragment size was positively related to offspring growth. Population density, rather than population size, had a positive effect on predation but negatively affected plant fecundity and offspring performance. Habitat quality reduced both plant fecundity and predation incidence.
• In this non‐specialist species, habitat fragmentation, population features and habitat quality affect different facets of plant performance. Predation was the only process clearly affected by fragmentation variables, fecundity mainly depended on population features and offspring performance and was better explained by mother plant identity. Our results show the need to consider habitat and population features together with fragment size and connectivity in order to assess the effects of fragmentation. Importantly, these effects can involve different aspects of plant reproduction, including plant–animal interactions.