Variation in migratory behavior influences regional genetic diversity and structure among American Kestrel populations (Falco sparverius) in North America

Birds employ numerous strategies to cope with seasonal fluctuations in high-quality habitat availability. Long distance migration is a common tactic; however, partial migration is especially common among broadly distributed species. Under partial migration systems, a portion of a species migrates, whereas the remainder inhabits breeding grounds year round. In this study, we identified effects of migratory behavior variation on genetic structure and diversity of American Kestrels (Falco sparverius), a widespread partial migrant in North America. American Kestrels generally migrate; however, a resident group inhabits the southeastern United States year round. The southeastern group is designated as a separate subspecies (F. s. paulus) from the migratory group (F. s. sparverius). Using mitochondrial DNA and microsatellites from 183 and 211 individuals, respectively, we illustrate that genetic structure is stronger among nonmigratory populations, with differentiation measures ranging from 0.060 to 0.189 depending on genetic marker and analysis approach. In contrast, measures from western North American populations ranged from 0 to 0.032. These findings suggest that seasonal migratory behavior is also associated with natal and breeding dispersal tendencies. We likewise detected significantly lower genetic diversity within nonmigratory populations, reflecting the greater influence of genetic drift in small populations. We identified the signal of population expansion among nonmigratory populations, consistent with the recent establishment of higher latitude breeding locations following Pleistocene glacial retreat. Differentiation of F. s. paulus and F. s. sparverius reflected subtle differences in allele frequencies. Because migratory behavior can evolve quickly, our analyses suggest recent origins of migratory American Kestrel populations in North America.     

The influence of sample size on the determination of population trends in the vulnerable Lesser Kestrel Falco naumanni overwintering in South Africa

Reliable measures of population sizes of endangered and vulnerable species are difficult to achieve because of high variability in population sizes and logistic constraints on sample sizes, yet such measures are crucial for the determination of the success of conservation and management strategies aimed at curbing declines in population size. In this study, we use bootstrap confidence intervals to assess the effects of sample size on the reliable determination of roost size in the vulnerable Lesser Kestrel Falco naumanni overwintering at a large communal roost in South Africa. Using results from a unique five-season daily survey, we show that the currently recommended number of replicate sample counts (international action plan for the Lesser Kestrel) of three censuses per season is too low to reliably assess roost size. Bootstrap confidence intervals asymptote at 40–60 censuses per season, with 50 being the modal value. We recommend 50 censuses per season as the optimal sample size. We were also able to show that the population size of Lesser Kestrels at this roost significantly increased over each of the first four seasons of the study period and decreased slightly in the final season.     

Postfledging dispersal in the endangered Lesser Kestrel Falco naumanni

Much is known about the breeding period of the threatened Lesser Kestrel, but little of the postfledging period. Here, I examine the postfledging dispersal of juvenile Lesser Kestrels, based on all ringing recoveries of Lesser Kestrels ringed as nestlings in Spain and recovered before September (migration) of the same year. Of 28 juveniles recovered, 19 (67.9%) dispersed and their median recovery distance was 210 km (range 41–578 km), with significantly northern directions. No difference was found in the dispersal distance between individuals ringed as nestlings in mid-north and mid-south Spain, nor was any correlation found between the recovery distance and ringing latitude. These findings for postfledging dispersal in juvenile Lesser Kestrels seem in accordance with predictions of a mental map hypothesis for homing, but they do not exclude food as a potential factor influencing such movements. However, these postfledging movements cannot be explained by the hypothesis of gathering information for identifying a future breeding site.     

Causes of population declines of the Lesser Kestrel Falco naumanni in Israel

We studied the ecology of the Lesser Kestrel Falco naumanni , a globally threatened species, to identify the factors causing its gradual decline in Israel, in order to stabilize and possibly restore its numbers. Lesser Kestrels in Israel breed in colonies, both adjacent to human settlements (rural and urban) and in the countryside, where they are found especially on cliffs. In this study, observations of Lesser Kestrels were carried out in three different breeding areas: (1) an urban colony in Jerusalem, (2) a rural colony in the Alona district and (3) a cliff in the Judean desert (open landscape colony) about 10 km east of Jerusalem. The number of fledglings per nest in Jerusalem (1.91) was lower than in Alona (2.44) and the cliff (3.16). As this lower productivity was associated neither with the clutch size, with hatching success, which were similar in all three regions (about four eggs per clutch, and 80% hatched, respectively), nor with egg fertility, it probably reflects factors operating during the nestling phase. We suggest that the two main factors limiting food availability and causing nestling deaths in Jerusalem are the relatively long flight distances between the breeding and hunting sites, and the use of pesticides in the city parks and lawns. Stochastic events superimposed on such factors, such as the drought of 1999, which markedly reduced productivity, may cause pronounced fluctuations eventually affecting long-term population persistence.     

Reproductive performance of Eurasian Kestrel Falco tinnunculus in an agricultural landscape with a mosaic of land uses

There are serious concerns about the environmental and ecological degradation caused by modern agriculture and its impact on animal populations. There is therefore a need to assess the reproductive performance of free‐living animals in agricultural landscapes. We undertook a 4‐year study on the reproductive biology of Eurasian Kestrel Falco tinnunculus in the Parma district of Italy. Kestrels breeding in areas of intensive agriculture delayed egg‐laying and their offspring were in poorer condition compared with those breeding in areas of temporary or permanent grasslands. However, egg volume and fledging success were similar among sites. The availability of temporary and permanent grasslands may limit the negative impacts of intensive agriculture on the breeding activity of Kestrels.     

7.—CORRESPONDENCE

Pepler, D., Martin, M., Fouche, F. &; VAN Hensbergen, B. 1994. Radionuclide analysis of Lesser Kestrels Falco naumanni and the origins of populations overwintering in southern Africa. Ostrich 65: 122–126.

The Lesser Kestrel Falco naumanni has recently shown major population declines throughout its breeding range in the Palearctic Region. In order to propose integrated conservation measures, it is important to establish the origins of the population overwintering in Africa. Nineteen Lesser Kestrels collected from released over Europe after the explosion at Chernobyl in 1986. Results of the radio-activity analysis show that these birds contain levels of ¹³⁴CS and ¹³⁷Cs that is below the detection threshold of the instrumentation. The measured activity of ¹³⁷Cs is a factor 57 lower than the “expected” level of 0,460 Bq/g ash. It is suggested that these birds originate from uncontaminated regions.     

Individual variation in migratory movements and winter behaviour of Iberian Lesser Kestrels Falco naumanni revealed by geolocators

The population decline of the Lesser Kestrel Falco naumanni has been the subject of studies across its Western Palaearctic breeding range, but little is known about its use of pre‐migratory areas or African wintering quarters. We used geolocators to describe the temporal and spatial patterns of Portuguese Lesser Kestrel migration and wintering behaviour. Data on the complete migration were obtained from four individuals and another three provided further information. Prior to southward migration, Lesser Kestrels showed two different behaviours: northward‐orientated movements to Spain and movements in the proximity of the breeding area. Autumn migration took place mostly in late September; spring departures occurred mainly in the first half of February. Wintering grounds included Senegal, Mauritania and Mali, with individuals overlapping considerably in Senegal. Movements registered within the wintering grounds suggest itinerant behaviour in relation to local flushes of prey. During spring migration, birds crossed the Sahara Desert through Mauritania, Western Sahara and Morocco before passing over the Mediterranean to reach Portugal. Autumn migration lasted 4.8 ± 1.1 days, and spring migration lasted 4.1 ± 0.3 days. The mean daily flight range varied between approximately 300 and 850 km for an entire journey of around 2500 km. Effective protection of roosting sites in both pre‐migratory and wintering areas and maintaining grasshopper populations in Sahelian wintering quarters appear crucial in preserving this threatened migratory raptor across its African–Eurasian flyway. There was no evidence of any deleterious effects of fitting birds with loggers.     

The role of fallow in habitat use by the Lesser Kestrel during the post-fledging period: inferring potential conservation implications from the abolition of obligatory set-aside

The maintenance of fallows has been shown to prevent the loss of farmland biodiversity caused by agricultural intensification. These are mainly introduced as part of both obligatory and voluntary set-aside schemes. However, the obligatory set-aside has recently been abolished by the Common Agricultural Policy Health Check. In this study, we examine the role of fallow in fine-grained habitat use by a threatened farmland bird (Lesser Kestrel) during summer in northwestern Spain. To analyze Lesser Kestrel occurrence, we used generalized linear models, a theoretic-information approach and a hierarchical partitioning analysis. The best AIC-based models explaining occurrence of Lesser Kestrels showed that fallow was the more important habitat type followed, to a lesser extent, by dry cereal stubble and field margin. In contrast, irrigated crops negatively influenced occurrence. Heterogeneity of crop mosaic was not important in explaining occurrence of Lesser Kestrel. Fallows, like dry cereal stubbles and field margins, seem to be suitable for foraging given the abundance of high food resources and their availability due to shorter vegetation cover. The abolition of the obligatory set-aside could reduce the total surface of fallow land (approximately 40.9%), likely affecting habitat use by Lesser Kestrel through an increase of other non-preferred crops (e.g., irrigated crops) or by decreasing food resources. Agri-environment schemes focusing on the maintenance of low-intensive farming systems with a mosaic of crops and semi-natural habitats interspersed should be promoted in premigratory areas to maintain Lesser Kestrel.     

Landscape and weather determinants of prey availability: implications for the Lesser Kestrel Falco naumanni

Spatial and temporal variation in prey abundance have been shown to impact the time of breeding and breeding success of birds. Understanding the ecological requirements of preferred prey can help develop management measures to improve food supply for target species. For the colonial Lesser Kestrel Falco naumanni, mole crickets Gryllotalpa spp. are one of the most important prey items during the mate‐feeding period. Lesser Kestrel colonies with higher mole cricket consumption had earlier egg‐laying dates, suggesting that differences between individuals in the time of breeding could be caused by differences in the diet. Moreover, the mean number of mole crickets in pellets was significantly correlated with clutch size (in one of the studied years) and egg volume. Thus, the impact of environmental variables and land use on mole crickets is likely to be relevant to Lesser Kestrel conservation. Weekly consumption of mole crickets was higher following an increase in either precipitation or minimum temperature values. Furthermore, mole cricket consumption was higher in colonies surrounded by higher quality soils and in wetter areas and years. Predicted probability of mole cricket occurrence in surveyed watercourse margins suggested a positive relationship between soil penetrability and mole cricket occurrence. Among variables that might be the target of management, the presence of riparian vegetation positively influenced the occurrence of mole crickets, whilst tillage and sowing of streambeds were revealed as the most important threats. We suggest that the maintenance of native vegetation in the margins of watercourses could improve soil resilience to erosion, increase water retention, soil penetrability and fertility, and provide a food supply and shelter for mole crickets. Overall, the implementation of such recommendations is likely to benefit other farmland species known to consume mole crickets, including several endangered species.     

The decline of Common Kestrels Falco tinnunculus in a forested area of northern England: the role of predation by Northern Goshawks Accipiter gentilis

We have previously documented the decline of the Common Kestrel Falco tinnunculus over a 23‐year period in a large coniferous forest in northern England. Kestrels fed predominantly on Field Voles Microtus agrestis, which were most abundant in young plantations (1–11 years old). Over the 23 years, voles remained abundant in the study area, but their numbers fluctuated cyclically. Here we consider whether the decline of Kestrels was linked to predation by Northern Goshawk Accipiter gentilis. Goshawks first bred in the study area in 1973 and increased until 1989, after which numbers stabilized. We use a number of approaches to explore the role of Goshawk predation, all of which are correlative, but independent. First, there was a significant negative relationship between Kestrel and Goshawk numbers after controlling for a decline in vole habitat. Short‐eared Owls Asio flammeus, which also hunt by day, declined over the same period as Kestrels. Second, numbers of Tawny Owl Strix aluco and Long‐eared Owl Asio otus did not decline as Goshawk numbers increased. These two species are also vole‐dependent, but active by night, and less vulnerable to Goshawk attack. Third, six species of raptor comprised 4.5% of 5445 Goshawk prey items during the breeding season, but more Kestrels were killed than the combined total of all other raptors. Goshawks not only killed many adult Kestrels in early spring, prior to breeding, when it would have most impact on population levels, but there was also a temporal trend for predation on Kestrels to be inversely density‐dependent. Finally, we estimated that Goshawks removed more Kestrels than were recorded each spring in the study area. We interpreted this as indicating that immigrant Kestrels were being removed continually, mostly before they could breed. We conclude that the decline of Kestrels (and possibly Short‐eared Owls) was mainly due to predation by Goshawks. This study provides some of the strongest evidence yet of the role of predation in the hierarchical structuring of raptor communities.     

6.—DATA OF BIRDS' EGGS IN MY COLLECTION

Van Zyl, A.J, Jenkins, A.R. &; Allan, D.G. 1994. Evidence for seasonal movements by Rock Kestrels Falco tinnunculus and Lanner Falcons F. biarmicus in South Africa. Ostrich 65:111-121.

Patterns of seasonal movement by Rock Kestrels Falco tinnunculus and Lanner Falcons F. biarmicus in South Africa were determined using data from the Southern African Bird Atlas Project, road transect counts, ringing recoveries and other sources. Both species were found to be partial, facultative migrants, and Lanner Falcon movements involved mostly immature birds. Rock Kestrels were found to concentrate in the southwest of the country in the breeding season, and to move north and east in the non-breeding season. Lanner Falcons concentrated in the east of the country during the breeding season, and moved into the west in the non-breeding season. The movement by both species into the arid northwest of the country is probably associated with dramatic increases in prey abundance, in turn associated with summer rah events. Lanner Falcon movement into the southern Cape apparently coincides with cereal farming practices, affecting the availability of prey (granivorous birds and rodents). Rock Kestrel movement into the east of the aunt is probably to exploit increases in insect abundance mated with summer rain. The pattern of Rock Kestrel movements suggests that it does not compete for food with the Lesser Kestrel F. naumanni. a Palearctic migrant to the east of the country.     

Increased hunting effort buffers against vole scarcity in an urban Kestrel Falco tinnunculus population

Capsule In years with low vole abundance birds visited hunting grounds more frequently and for longer.

Aims To describe diet composition, hunting behaviour, habitat choice and reproductive success of urban Kestrels during changing vole abundance.

Methods For five years, we studied the hunting effort of Kestrels in a medium-sized city during the breeding season. Pitfall traps were used for determining vole abundance. Kestrel diet composition was determined from pellet analyses. The number of eggs and offspring was recorded during at least two consecutive visits for each nest and each breeding stage.

Results In contrast to larger European cities, the Common Vole Microtus arvalis was a key part of the Kestrels' diet and did not fluctuate significantly according to vole availability. Reproductive success was quite high and stable throughout years with different vole abundance. In years of low vole abundance, the arrival frequency at hunting grounds and time spent there increased. During years with a low vole population, Kestrels had less hunting success and the rate of successful visits decreased. Therefore, Kestrels probably had to change hunting grounds more frequently. During low vole years Kestrels used less demanding techniques, e.g. perching, despite the lower success of these hunting techniques, to avoid extremely high energetic costs.

Conclusions An increase in hunting helps to maintain a proper diet and consequently reproductive success. Vole abundance did not change dramatically during the study period, as reported by studies from western and northern Europe. The proportion of ruderal habitats on the city periphery is higher than in more monotonous farmland habitats. Ruderal habitats can be important when Kestrels look for mammals other than voles, especially during vole scarcity.     

Evidence of increasing Lesser Kestrel Falco naumanni populations in Morocco (North Africa): a confirmation of ‘Least Concern’ conservation status

Capsule The number of Lesser Kestrel Falco naumanni breeding pairs in Morocco has varied significantly and positively over thirteen years (1998–2010), between six colonies and over the 13-year period at each colony. The mean growth rate was λ?=?1.13?±?0.02. This positive trend is consistent with the current Least Concern conservation status of the species in the IUCN Red List.     

Plumage development and visual communication in the Greater Kestrel Falco rupicoloides near Pretoria,South Africa

Kemp, A. C. 1999. Plumage development and visual communication in the Greater Kestrel Falco rupicoloides near Pretoria, South Africa. Ostrich 70 (3&4): 220–224.

I trapped, marked, sexed and examined moult and softparts on 101 free-living Greater Kestrels Falco rupicoloides near Pretoria, South Africa, during 1975–1988.1 collected and plucked one kestrel to determine the number and mass of its feather tracts, as an index of materials invested in moult. I also examined moult on 72 museum specimens from southern Africa and observed plumage and softpart changes in four captive birds. During 1975–1979,1 made focal observations on free-living adult kestrels (males 181 h, females 262 h) and recorded monthly samples of their visual communication behaviours to the nearest minute.

Greater Kestrels can be aged for three successive plumage and softpart stages (juvenile, post-juvenile, adult). Plumage changes with potential signal effects involved only small investments at the post-juvenile and first adult moults (3.7% and 11% of total plumage mass respectively). Changes in colouration and communication behaviour correlated with relevant stages of the annual cycle, possibly to miminze conflict with territorial adults and age of first breeding. Remex and rectrix moult starts on average much later in relation to egglaying for Greater and some tropical kestrels than for some temperate species. This and other individual variation in timing may result for variations in nutritional status.     

Effect of nest-site microclimatic conditions on nesting success in the Lesser Kestrel Falco naumanni

Capsule: Microclimatic conditions in the nests of the Lesser Kestrel Falco naumanni, particularly the percentage of time of extremely low humidity, affect breeding success.

Aim: To study the effect of within-nest temperature and humidity on nest productivity, and the correlation between nest productivity and the order of dates on which nests were occupied by the parents. To compare microclimatic conditions in the nest, breeding success and order of occupation between nests under tile roofs and in artificial nest boxes.

Methods: Three different Lesser Kestrel colonies were monitored in Israel – one rural, one urban and one in an open country habitat. Data loggers were placed in 39 nests for the entire breeding period to measure temperature and humidity. The number of fledglings was recorded for each nest, as well as the date of occupation.

Results: Full microclimatic data from 35 nests suggest that percentage of time of extremely low humidity is the major predictor of nest productivity – low humidity is negatively correlated with nest productivity. Sites of more successful nests were occupied earlier. Considering only successful nests, the urban colony had the lowest breeding success of the three colonies. There was no significant difference in mean productivity between nests in roofs and nest boxes, but nests in roofs were occupied earlier.

Conclusion: Nest microclimate, particularly low humidity, affects nesting success in addition to colony location.     

Nestbox provisioning in a rural population of Eurasian Kestrels: breeding performance,nest predation and parasitism

The breeding biology of the Eurasian Kestrel Falco tinnunculusin nestboxes in farmland was studied to test for differences between artificial and natural sites. We report on the direct effect of nestbox provisioning on some life-history traits and how nestbox use affects nest predation and parasitism. Five types of nest-sites were available: nestboxes on poles and trees (artificial sites), stick nests on trees, stick nests on pylons and holes in buildings (‘natural’ sites). The Kestrel population increased from 23 pairs in 1993 (prior to nestbox installation) to 55 in 1998 as nestboxes were provided. In general, pairs breeding in trees started to lay later than those nesting in nestboxes on poles or in building holes, but this difference was probably associated with habitat quality rather than nest type. Differences in clutch size were found between nest-sites in some years, and were associated with laying date and, probably, with variation in territory quality. Using only data from successful nests, pairs breeding in nestboxes produce more fledglings than those in building holes or pylons. The frequency of nest predation was higher in natural sites than in nestboxes. The number of fledglings from pairs breeding in nestboxes was higher than from those breeding in old stick nests in trees when all nests were considered. Nestbox provisioning had no effect on the occurrence of the ectoparasite Carnus hemapterus, but chicks from nestboxes showed higher intensity of infection. Our results suggest that nestbox provisioning increases reproductive success and the frequency of nest predation or intensity of parasite infestation in Kestrels.     

Landscape scale genetic effects of habitat fragmentation on a high gene flow species: Speyeria idalia (Nymphalidae)

Detection of the genetic effects of recent habitat fragmentation in natural populations can be a difficult task, especially for high gene flow species. Previous analyses of mitochondrial DNA data from across the current range of Speyeria idalia indicated that the species exhibited high levels of gene flow among populations, with the exception of an isolated population in the eastern portion of its range. However, some populations are found on isolated habitat patches, which were recently separated from one another by large expanses of uninhabitable terrain, in the form of row crop agriculture. The goal of this study was to compare levels of genetic differentiation and diversity among populations found in relatively continuous habitat to populations in both recently and historically isolated habitat. Four microsatellite loci were used to genotype over 300 individuals from five populations in continuous habitat, five populations in recently fragmented habitat, and one historically isolated population. Results from the historically isolated population were concordant with previous analyses and suggest significant differentiation. Also, microsatellite data were consistent with the genetic effects of habitat fragmentation for the recently isolated populations, in the form of increased differentiation and decreased genetic diversity when compared to nonfragmented populations. These results suggest that given the appropriate control populations, microsatellite markers can be used to detect the effects of recent habitat fragmentation in natural populations, even at a large geographical scale in high gene flow species.     

Experimental test on public information use in the colonial Lesser Kestrel

The formation of colonies and the evolution of coloniality have been suggested to be a by-product of the use of public information to select breeding habitats. In this study we performed an experimental test to investigate whether the Lesser Kestrel (Falco naumanni) uses breeding success of conspecifics as a source of public information to select a breeding colony. We considered four potential cues based on reproductive performance of conspecifics in each colony: mean and variance of breeding success measured in both all pairs (TRS, total reproductive success), and only successful pairs (PRS, partial reproductive success). Both mean PRS and mean TRS could be predictive cues of the future reproductive output at a given site, because they varied among colonies and were autocorrelated from one year to the next. To dissociate any relationship between habitat quality and reproductive success we manipulated the mean brood sizes by transferring chicks among ten colonies. In five colonies, all the broods with at least two nestlings were reduced in one chick and in other five colonies all broods were enlarged by one chick. In the following year, we found that the number of adult immigrant kestrels settled at a colony was only explained by mean PRS in the previous year. We also examined the settlement of yearlings, which lack of information on breeding success in previous years. Unlike adult immigrant kestrels, yearlings tended to settle more frequently in those colonies with low mean PRS in the previous year, probably because of the lower pressure for occupation in those colonies. A multiple regression analysis showed that the growth of colonies was positively dependent on the mean PRS in the experimental year, and negatively on the number of predated nests. This study provides a solid support to the hypothesis that colonial species can use mean breeding success of successful pairs as a source of public information to make settlement decisions.     

Different trends of neighboring populations of Lesser Kestrel: Effects of climate and other environmental conditions

The sensitivity of population trends to the climate and environment is generally considered a species-specific trait. However, evidence that populations may show different responses to the climate and environmental conditions is growing. Whether this differential sensitivity may arise even among neighboring populations remains elusive. We compared the trends of two neighboring populations of the Lesser Kestrel Falco naumanni, using data from a 12-year survey of 158 colonies in Sicily, Italy; the two populations inhabiting a lowland and an highland area, respectively. Population trends were modeled through the TRIM algorithms implemented in R (package rtrim). A reversed U-shaped population trend was observed in the lowland, while the highland population showed oscillations around a stable trend. Sahel rainfall 2 years before each annual survey significantly affected population variation in the lowland, while rainfall in March and an index of primary productivity in the breeding areas affected population variation in the highland. This suggests that the population in the lowland may be limited mainly by winter survival in Sahel, because the lowland may be an optimal breeding area for this species. In contrast, the highland population, which occupies a different part of the climatic niche of the species, may be limited mainly by reproductive output, because rainfall in March and the primary productivity in May could represent prey availability immediately before and during the breeding months. Overall, our findings suggest that population-specific environmental sensitivity might occur even over small (<100 km) geographical scales, highlighting the need for population-specific conservation strategies.