Climatic conditions during outward migration affect apparent survival of an arctic top predator,the peregrine falcon Falco peregrinus

In long‐lived species, population growth rate is highly sensitive to changes in adult survival. Despite the growing concerns regarding recent climate changes, few studies have investigated the effect of climatic conditions on survival in long‐lived wildlife that are either resident or breed in the Arctic. In this study, we evaluated the effect of climate across the annual life cycle (breeding, outward migration, wintering, and inward migration) on apparent annual survival of arctic‐breeding peregrine falcons. From 1982 to 2008, peregrine falcons breeding near Rankin Inlet, Nunavut, Canada were monitored, in part, to assess apparent annual survival (the product of true survival and site fidelity) using re‐observations of marked individuals. Our study indicated that apparent annual survival of adult peregrine falcons was correlated with indices of climatic conditions during outward migration (i.e., flight from the Arctic breeding grounds). These climatic indices (fall NAO of the current year and fall NAO with a lag of one year) explained 35% of the temporal variation in apparent annual survival of peregrine falcons. Our results suggest that this top‐predator is vulnerable to weather‐related environmental conditions encountered during fall migration.     

Hematological, protein electrophoresis and cholinesterase values of free-living nestling peregrine falcons in Spain

Protein electrophoresis, hematological and cholinesterase values were determined in 32 nestling free-living peregrine falcons (Falco peregrinus) (15- to 27-days-old) in order to establish normal reference values for this population. The following values (mean +/- SD) were observed: prealbumin 0.31 +/- 0.04 g/dl, albumin 1.25 +/- 0.06 g/dl, alpha1 and alpha2-globulin 0.23 +/- 0.02 and 0.16 +/- 0.02 g/dl respectively, beta-globulin 1.02 +/- 0.05 g/dl, gamma-globulin 0.060 +/- 0.08 g/dl, total protein 3.79 +/- 0.18 g/dl, 21.26 +/- 1.30 white blood cells/microl (1 x 10(3)), 2.17 +/- 0.07 red blood cells/microl (1 x 10(6)), packed cell volume 37.58 +/- 0.82%, hemoglobin 20.96 +/- 0.29 g/dl, heterophils 61.14 +/- 2.50% and cholinesterase 1,184 +/- 75 IU/L. There were no difference in any of these parameters among males and females. The hematological values obtained could be considered as representative values in free-living nestling peregrine falcons.     

Landscape heterogeneity drives intra‐population niche variation and reproduction in an arctic top predator

While intra‐population variability in resource use is ubiquitous, little is known of how this measure of niche diversity varies in space and its role in population dynamics. Here we examined how heterogeneous breeding environments can structure intra‐population niche variation in both resource use and reproductive output. We investigated intra‐population niche variation in the Arctic tundra ecosystem, studying peregrine falcon (Falco peregrinus tundrius, White) breeding within a terrestrial‐marine gradient near Rankin Inlet, Nunavut, Canada. Using stable isotope analysis, we found that intra‐population niches varied at the individual level; we examined within‐nest and among‐nest variation, though only the latter varied along the terrestrial‐marine gradient (i.e., increased among‐nest variability among birds nesting within the marine environment, indicating higher degree of specialization). Terrestrial prey species (small herbivores and insectivores) were consumed by virtually all falcons. Falcons nesting within the marine environment made use of marine prey (sea birds), but depended heavily on terrestrial prey (up to 90% of the diet). Using 28‐years of peregrine falcon nesting data, we found a positive relationship between the proportion of terrestrial habitat surrounding nest sites and annual nestling production, but no relationship with the likelihood of successfully rearing at least one nestling reaching 25 days old. Annually, successful inland breeders raised 0.47 more young on average compared to offshore breeders, which yields potential fitness consequences for this long‐living species. The analyses of niche and reproductive success suggest a potential breeding cost for accessing distant terrestrial prey, perhaps due to additional traveling costs, for those individuals with marine nest site locations. Our study indicates how landscape heterogeneity can generate proximate (niche variation) and ultimate (reproduction) consequences on a population of generalist predator. We also show that within‐individual and among‐individual variation are not mutually exclusive, but can simultaneously arise and structure intra‐population niche variation.     

Decline of an arctic top predator: synchrony in colony size fluctuations, risk of extinction and the subpolar gyre

Analysis of synchrony in population fluctuations is a central topic in ecology. It can help identify factors that regulate populations, and also the scales at which these factors exert their influence. Using long-term data from seven Brünnich’s guillemot colonies in Svalbard, Norway, we determined that year to year population fluctuations were synchronized in six of the seven colonies. The seventh colony was located farther away and in a different oceanographic system. Moreover, all seven colonies have declined significantly since the late 1990s following a very similar pattern. If the rate of population decline does not change, Brünnich’s guillemots in Svalbard have a high probability of becoming quasi-extinct within the next 50 years. The high synchrony between the different colonies could further increase this risk of extinction. Our results indicate that environmental forcing plays a role in the colony size fluctuation of Brünnich’s guillemot (i.e., a Moran effect). These fluctuations are well explained by changes in the subpolar gyre in the region where Brünnich’s guillemots overwinter. This subpolar gyre weakened in the mid-1990s, leading to a warming of the North Atlantic. Our study indicates that this basin-scale shift in the subpolar gyre is closely related to the decline in Brünnich’s guillemot in Svalbard. Our results suggest that the causal mechanism linking changes in oceanographic conditions in the North Atlantic and Brünnich’s guillemot population dynamics are likely mediated, at least partly, by changes in recruitment.     

Environmental stability increases relative individual specialisation across populations of an aquatic top predator

The concept of the niche has long been a central pillar in ecological theory, with a traditional focus on quantifying niches at the species or population level. However, the importance of individual‐level niche variation is increasingly being recognised, with a strong focus on individual specialisation. While examples illustrating the contribution of the individual niche to whole population niche structure are accumulating rapidly, surprisingly little is known about the conditions that shape the differences between these two potentially divergent components. Though theory predicts that stability should influence the extent of such intra‐specific specialisation, we know of no previous study that has investigated its role in individual specialisation, and the differentiation between individual‐ and population niches. Here, we studied the diet of individuals from multiple populations of an aquatic top‐predator, Salmo trutta, inhabiting contrasting stable, groundwater fed and unstable, surface water fed pre‐alpine streams. Based on stomach content analysis, we found that individuals living in stable environments displayed a higher degree of specialisation than those in unstable environments, with the between‐individual component of niche width being approximately twice as high in the former. We subsequently validated these results by evidence gained from stable isotope analysis of muscle tissue. As such, we reveal that environmental stability can significantly influence individual niches within populations, leading to increased specialisation.     

Prey availability influences habitat tolerance: an explanation for the rarity of peregrine falcons in the tropics

The density and productivity of peregrine falcon Falco peregri nus populations correlate positively with distance from the Equator, while habitat specificity increases with proximity to the Equator. Low peregrine densities in the tropics may he a result of competition with similar conveners (e.g. the lanner falcon F. biarmicus in Africa), which replace them in many areas. Alternatively, tropical peregrines may he limited by resource deficiencies that do not affect their close relatives. Data from peregrine and lanner populations in South Africa support the resource deficiency hypothesis, and there is no evidence to suggest direct competition between the two species. In areas where prey are not spatially or temporally concentrated, or otherwise particularly vulnerable to attack, morphological and behavioural specializations of peregrines probably restrict them to optimal foraging conditions. The relative dynamics of.Arctic and temperate vs tropical prey populations is suggested as an important factor determining peregrine distribution globally. Populations of other widespread hut particularly specialized avian predators (e.g. osprey Pandion haliaetus) may he similarly controlled. Food limitation (in terms of a dearth of particularly vulnerable prey) in the tropics has resulted in specialization and rarity in peregrines and generalization and relative abundance in similar congeners.     

Individual benefits of nestling begging: experimental evidence for an immediate effect, but no evidence for a delayed effect

The evolutionary stability of honest signalling by offspring is thought to require that begging displays be costly, so the costs and benefits of begging--and whether they are experienced individually or by the whole brood--are crucial to understanding the evolution of begging behaviour. Begging is known to have immediate individual benefits (parents distribute more food to intensely begging individuals) and delayed brood benefits (parents increase provisioning rate to the brood), but the possibility of delayed individual benefits (previous begging affects the current distribution of food) has rarely, if ever, been researched. We did this using playback of great tit Parus major chick begging and a control sound from either side of the nest. Male parents fed chicks close to the speaker more when great tit chick begging, but not other stimuli, was played back. In contrast, there was no effect of playback at the previous visit on the chicks that male parents fed. We have thus demonstrated an immediate individual benefit to begging, but found no evidence of a delayed individual benefit in this species.     

Intrapopulation variability shaping isotope discrimination and turnover: experimental evidence in arctic foxes

Background

Tissue-specific stable isotope signatures can provide insights into the trophic ecology of consumers and their roles in food webs. Two parameters are central for making valid inferences based on stable isotopes, isotopic discrimination (difference in isotopic ratio between consumer and its diet) and turnover time (renewal process of molecules in a given tissue usually measured when half of the tissue composition has changed). We investigated simultaneously the effects of age, sex, and diet types on the variation of discrimination and half-life in nitrogen and carbon stable isotopes (δ¹⁵N and δ¹³C, respectively) in five tissues (blood cells, plasma, muscle, liver, nail, and hair) of a top predator, the arctic fox Vulpes lagopus.

Methodology/Principal Findings

We fed 40 farmed foxes (equal numbers of adults and yearlings of both sexes) with diet capturing the range of resources used by their wild counterparts. We found that, for a single species, six tissues, and three diet types, the range of discrimination values can be almost as large as what is known at the scale of the whole mammalian or avian class. Discrimination varied depending on sex, age, tissue, and diet types, ranging from 0.3‰ to 5.3‰ (mean  = 2.6‰) for δ¹⁵N and from 0.2‰ to 2.9‰ (mean  = 0.9‰) for δ¹³C. We also found an impact of population structure on δ¹⁵N half-life in blood cells. Varying across individuals, δ¹⁵N half-life in plasma (6 to 10 days) was also shorter than for δ¹³C (14 to 22 days), though δ¹⁵N and δ¹³C half-lives are usually considered as equal.

Conclusion/Significance

Overall, our multi-factorial experiment revealed that at least six levels of isotopic variations could co-occur in the same population. Our experimental analysis provides a framework for quantifying multiple sources of variation in isotopic discrimination and half-life that needs to be taken into account when designing and analysing ecological field studies.     

Effects of land‐use change and rainfall in Sudano‐Sahelian West Africa on the diet and nestling growth rates of an avian predator

Raptor populations in Sudano‐Sahelian West Africa are being severely affected by widespread habitat alteration which depletes prey populations, potentially aggravated by changing rainfall patterns. We studied Grasshopper Buzzards Butastur rufipennis at nests in natural and transformed habitats in the Sudano‐Sahelian region of northern Cameroon to assess the effects of habitat transformation and rainfall on nestling diet and growth. Grasshoppers and small mammals were more frequently taken in natural habitat, whereas lizards were most frequently taken in transformed habitats. These dietary differences reflected differences in prey availability around nests in natural and transformed habitats. Land use was a significant predictor of asymptotic weight: nestlings in natural habitat attained a higher mass than those in transformed habitats, when potentially confounding variables such as hatch order, gender, hatch date, rainfall or the presence of siblings were taken into consideration. These results suggest that body condition at fledging was habitat‐dependent, with potential consequences for subsequent survival. However, we recorded no differences in caloric content of food delivered to nests in natural and transformed habitat, which was possibly related to prey caught during twilight hours. There was a positive relationship between precipitation levels during the nestling phase and nestling growth rate. We predict unfavourable future conditions for nestling growth of raptors in Sudano‐Sahelian savannas as a consequence of continued widespread habitat transformation and diminished rainfall.     

Malar stripe size and prominence in peregrine falcons vary positively with solar radiation: support for the solar glare hypothesis

Many falcons (Falco spp.) exhibit a distinct dark plumage patch below the eye, termed the malar stripe. This stripe is hypothesized to reduce the amount of solar glare reflected into the eyes while foraging, thereby increasing hunting efficiency in bright conditions. Here, we use a novel, global-scale correlative approach to test this ‘solar glare hypothesis'' in peregrine falcons (Falco peregrinus), the most widespread falcon species, using web-sourced photographs from across the species'' global range. We found that the size and prominence of the malar stripe were positively associated with average annual solar radiation, but not with other environmental variables, such as temperature and rainfall. Our results provide the first published evidence for the hypothesis that this plumage feature functions to reduce the amount of solar glare reflected into the falcon''s eyes, thereby improving the ability to pinpoint and target agile prey in bright conditions.     

The response of Alaskan arctic tundra to experimental warming: differences between short- and long-term responses

Global climate models predict continued rapid warming for most of the Arctic throughout the next century. To further understand the response of arctic tundra to climate warming, four sites in northern Alaska were warmed for five to seven consecutive growing seasons using open‐top chambers. Sites were located in dry heath and wet meadow communities near Barrow (71°18′N, 156°40′W) and Atqasuk (70°29′N, 157°25′W). Change in plant community composition was measured using a point frame method. During the period of observation, species richness declined in control plots by up to 2.7 species plot^?1. Responses to warming varied by site but similar trends included increased canopy height (?0.1 to 2.3 cm) and relative cover of standing dead plant matter (1.5–6.0%) and graminoids (1.8–5.8%) and decreased species diversity (0.1–1.7 species plot^?1) and relative cover of lichens (0.2–9.1%) and bryophytes (1.4–4.6%) (parentheses enclose the range of average values for the sites). The response to warming was separated into an initial short‐term response assessed after two growing seasons of warming and a secondary longer‐term response assessed after an additional three to five growing seasons of warming. The initial responses to warming were similar in the four sites, while the secondary responses varied by site. The response to warming was greater at Barrow than Atqasuk because of a greater initial response at Barrow. However, the long‐term response to warming was projected to be greater at Atqasuk because of a greater secondary response at Atqasuk. These findings show that predictions of vegetation change due to climate warming based on manipulative experiments will differ depending on both the duration and plant community on which the study focuses.     

Correlates of seasonal change in the body condition of an Arctic top predator

Climate‐driven sea ice loss has led to changes in the timing of key biological events in the Arctic, however, the consequences and rate of these changes remain largely unknown. Polar bears (Ursus maritimus) undergo seasonal changes in energy stores in relation to foraging opportunities and habitat conditions. Declining sea ice has been linked to reduced body condition in some subpopulations, however, the specific timing and duration of the feeding period when bears acquire most of their energy stores and its relationship to the timing of ice break‐up is poorly understood. We used community‐based sampling to investigate seasonality in body condition (energy stores) of polar bears in Nunavut, Canada, and examined the influence of sea ice variables. We used adipose tissue lipid content as an index of body condition for 1,206 polar bears harvested from 2010–2017 across five subpopulations with varying seasonal ice conditions: Baffin Bay (October–August), Davis Strait and Foxe Basin (year‐round), Gulf of Boothia and Lancaster Sound (August–May). Similar seasonal patterns were found in body condition across subpopulations with bears at their nadir of condition in the spring, followed by fat accumulation past break‐up date and subsequent peak body condition in autumn, indicating that bears are actively foraging in late spring and early summer. Late season feeding implies that even minor advances in the timing of break‐up may have detrimental effects on foraging opportunities, body condition, and subsequent reproduction and survival. The magnitude of seasonal changes in body condition varied across the study area, presumably driven by local environmental conditions. Our results demonstrate how community‐based monitoring of polar bears can reveal population‐level responses to climate warming in advance of detectable demographic change. Our data on the seasonal timing of polar bear foraging and energy storage should inform predictive models of the effects of climate‐mediated sea ice loss.     

The use of genetics for the management of a recovering population: temporal assessment of migratory peregrine falcons in North America

Background

Our ability to monitor populations or species that were once threatened or endangered and in the process of recovery is enhanced by using genetic methods to assess overall population stability and size over time. This can be accomplished most directly by obtaining genetic measures from temporally-spaced samples that reflect the overall stability of the population as given by changes in genetic diversity levels (allelic richness and heterozygosity), degree of population differentiation (F _ST and D _EST), and effective population size (N _e). The primary goal of any recovery effort is to produce a long-term self-sustaining population, and these genetic measures provide a metric by which we can gauge our progress and help make important management decisions.

Methodology/Principal Findings

The peregrine falcon in North America (Falco peregrinus tundrius and anatum) was delisted in 1994 and 1999, respectively, and its abundance will be monitored by the species Recovery Team every three years until 2015. Although the United States Fish and Wildlife Service makes a distinction between tundrius and anatum subspecies, our genetic results based on eleven microsatellite loci suggest limited differentiation that can be attributed to an isolation by distance relationship and warrant no delineation of these two subspecies in its northern latitudinal distribution from Alaska through Canada into Greenland. Using temporal samples collected at Padre Island, Texas during migration (seven temporal time periods between 1985–2007), no significant differences in genetic diversity or significant population differentiation in allele frequencies between time periods were observed and were indistinguishable from those obtained from tundrius/anatum breeding locations throughout their northern distribution. Estimates of harmonic mean N _e were variable and imprecise, but always greater than 500 when employing multiple temporal genetic methods.

Conclusions/Significance

These results, including those from simulations to assess the power of each method to estimate N _e, suggest a stable or growing population, which is consistent with ongoing field-based monitoring surveys. Therefore, historic and continuing efforts to prevent the extinction of the peregrine falcon in North America appear successful with no indication of recent decline, at least from the northern latitude range-wide perspective. The results also further highlight the importance of archiving samples and their use for continual assessment of population recovery and long-term viability.     

Genetic and historic evidence for climate-driven population fragmentation in a top cetacean predator: the harbour porpoises in European water

Recent climate change has triggered profound reorganization in northeast Atlantic ecosystems, with substantial impact on the distribution of marine assemblages from plankton to fishes. However, assessing the repercussions on apex marine predators remains a challenging issue, especially for pelagic species. In this study, we use Bayesian coalescent modelling of microsatellite variation to track the population demographic history of one of the smallest temperate cetaceans, the harbour porpoise (Phocoena phocoena) in European waters. Combining genetic inferences with palaeo-oceanographic and historical records provides strong evidence that populations of harbour porpoises have responded markedly to the recent climate-driven reorganization in the eastern North Atlantic food web. This response includes the isolation of porpoises in Iberian waters from those further north only approximately 300 years ago with a predominant northward migration, contemporaneous with the warming trend underway since the ‘Little Ice Age’ period and with the ongoing retreat of cold-water fishes from the Bay of Biscay. The extinction or exodus of harbour porpoises from the Mediterranean Sea (leaving an isolated relict population in the Black Sea) has lacked a coherent explanation. The present results suggest that the fragmentation of harbour distribution range in the Mediterranean Sea was triggered during the warm ‘Mid-Holocene Optimum’ period (approx. 5000 years ago), by the end of the post-glacial nutrient-rich ‘Sapropel’ conditions that prevailed before that time.     

Population status of a cryptic top predator: an island-wide assessment of tigers in Sumatran rainforests

Large carnivores living in tropical rainforests are under immense pressure from the rapid conversion of their habitat. In response, millions of dollars are spent on conserving these species. However, the cost-effectiveness of such investments is poorly understood and this is largely because the requisite population estimates are difficult to achieve at appropriate spatial scales for these secretive species. Here, we apply a robust detection/non-detection sampling technique to produce the first reliable population metric (occupancy) for a critically endangered large carnivore; the Sumatran tiger (Panthera tigris sumatrae). From 2007–2009, seven landscapes were surveyed through 13,511 km of transects in 394 grid cells (17×17 km). Tiger sign was detected in 206 cells, producing a naive estimate of 0.52. However, after controlling for an unequal detection probability (where p = 0.13±0.017; ±S.E.), the estimated tiger occupancy was 0.72±0.048. Whilst the Sumatra-wide survey results gives cause for optimism, a significant negative correlation between occupancy and recent deforestation was found. For example, the Northern Riau landscape had an average deforestation rate of 9.8%/yr and by far the lowest occupancy (0.33±0.055). Our results highlight the key tiger areas in need of protection and have led to one area (Leuser-Ulu Masen) being upgraded as a ‘global priority’ for wild tiger conservation. However, Sumatra has one of the highest global deforestation rates and the two largest tiger landscapes identified in this study will become highly fragmented if their respective proposed roads networks are approved. Thus, it is vital that the Indonesian government tackles these threats, e.g. through improved land-use planning, if it is to succeed in meeting its ambitious National Tiger Recovery Plan targets of doubling the number of Sumatran tigers by 2022.