Journal of Biogeography (J. Biogeogr.) (2007) 34, 1193–1206

ORIGINAL
ARTICLE

Phylogeography of the white-tailed
eagle, a generalist with large dispersal
capacity
F. Hailer1*, B. Helander2, A. O. Folkestad3, S. A. Ganusevich4, S. Garstad5,
P. Hauff6, C. Koren7, V. B. Masterov8, T. Nygård9, J. A. Rudnick10,
S. Shiraki11, K. Skarphedinsson12, V. Volke13, F. Wille14 and C. Vilà11

Department of Evolutionary Biology, Uppsala
University, Norbyvägen 18d, 75236 Uppsala,Sweden, 2Contaminant Research Group,
Swedish Museum of Natural History, Box
50007, 10405 Stockholm, Sweden, 3Norwegian
Sea-Eagle Project, Eiksund, 6065 Ulsteinvik,
Norway, 4Field Research Group for the Kola
North, Krasnostudencheskiy pr., 21-45,
Moscow 127422, Russia, 5Postb. 10, 7994 Leka,
Norway, 6Lindenallé 5, 19073 Neu Wandrum,
Germany, 7Hagebyv. 39, 9404 Harstad,
Norway, 8Department of Vertebrate Zoology,
Biological Faculty of the Moscow State
University, Leninskie Gory 1-12, 119992
Moscow, Russia, 9Norwegian Institute for
Nature Research, Tungasletta 2, 7485
Trondheim, Norway, 10Department of Forestry
and Natural Resources, Purdue University, 195
Marsteller Street, West Lafayette, IN 479072033, USA, 11Wildlife Section, Nature
Conservation Department, Hokkaido Instituteof Environmental Sciences, Kita-19 Nishi-12
Kita-ku, Sapporo, 060-0819, Japan, 12Icelandic
Institute of Natural History, Hlemmur 3, 105
Reykjavik, Iceland, 13Eagle Club, Vahtra 5,93813 Kuressaare, Estonia and 14Thorsvej 18,4180 Sorö, Denmark*Correspondence: Frank Hailer, Genetics
Program, National Zoological Park, National
Museum of Natural History, Smithsonian
Institution, 3001 Connecticut Ave., NW,
Washington, DC 20008, USA.
E-mail: frashai@gmx.net;
frank.hailer@ebc.uu.se

ABSTRACT

Aim Late Pleistocene glacial changes had a major impact on many boreal andtemperate taxa, and this impact can still be detected in the present-day
phylogeographic structure of these taxa. However, only minor effects are expectedin species with generalist habitat requirements and high dispersal capability. One
such species is the white-tailed eagle, Haliaeetus albicilla, and we therefore tested
for the expected weak population structure at a continental level in this species.
This also allowed us to describe phylogeographic patterns, and to deduce Ice Agerefugia and patterns of postglacial recolonization of Eurasia.
Location Breeding populations from the easternmost Nearctic (Greenland) andacross the Palaearctic (Iceland, continental Europe, central and eastern Asia, andJapan).
Methods Sequencing of a 500 base-pair fragment of the mitochondrial DNA
control region in 237 samples from throughout the distribution range.
Results Our analysis revealed pronounced phylogeographic structure. Overall,
low genetic variability was observed across the entire range. Haplotypes clusteredin two distinct haplogroups with a predominantly eastern or western distribution,and extensive overlap in Europe. These two major lineages diverged during thelate Pleistocene. The eastern haplogroup showed a pattern of rapid population
expansion and colonization of Eurasia around the end of the Pleistocene. The
western haplogroup had lower diversity and was absent from the populations ineastern Asia. These results suggest survival during the last glaciation in two
refugia, probably located in central and western Eurasia, followed by postglacial
population expansion and admixture. Relatively high genetic diversity wasobserved in northern regions that were ice-covered during the last glacial
maximum. This, and phylogenetic relationships between haplotypes encounteredin the north, indicates substantial population expansion at high latitudes. Areas ofglacial meltwater runoff and proglacial lakes could have provided suitable habitats
for such population growth.
Main conclusions This study shows that glacial climate fluctuations had asubstantial impact on white-tailed eagles, both in terms of distribution anddemography. These results suggest that even species with large dispersal
capabilities and relatively broad habitat requirements were strongly affected by
the Pleistocene climatic shifts.
Keywords
Control region, Eurasia, Falconiformes, Haliaeetus albicilla, mtDNA, population
expansion, postglacial colonization.

ª 2007 The Authors
Journal compilation ª 2007 Blackwell Publishing Ltd

www.blackwellpublishing.com/jbi
doi:10.1111/j.1365-2699.2007.01697.x

1193

F. Hailer et al.
INTRODUCTION
The multiple glacial advances and retreats that occurred
during the Pleistocene not only altered the landscape
topography of the northern hemisphere, but also had adramatic influence on the abundance and distribution ofliving forms. Most species underwent dramatic reductions in
numbers and range (summarized in Hewitt, 2000), and many
lineages went extinct (e.g. Shapiro et al., 2004). For many
temperate animal and plant species from Europe, peninsulason the Mediterranean coast acted as glacial refugia (Taberlet
et al., 1998; Hewitt, 2000). Conversely, some Arctic species
appear to have reacted in an opposite manner to the
Pleistocene climatic shifts, with contraction into northern
refugia during warm periods and major population expansions during cold periods (Flagstad & Roed, 2003; Dalénet al., 2005).
Taxa with low mobility and narrow habitat requirements are
expected to have been the most strongly affected by the drastic
habitat changes accompanying the glacial cycles, while thegenetic structure of species with higher dispersal capability that
exploit a broader ecological niche may have been lessinfluenced by climatic changes. For example, the grey wolf
(Canis lupus, Linnaeus, 1758) shows very little phylogeographic structure in mitochondrial DNA (mtDNA) at a continental
level (Vilà et al., 1999; but see Sharma et al., 2004).
Another species group with high dispersal potential aswell as broad habitat and dietary requirements is that ofeagles in the genus Haliaeetus. The three largest species inthis genus occur in the northern hemisphere, including thewhite-tailed eagle H. albicilla (Linnaeus, 1758), its North
American sister species the bald eagle, H. leucocephalus,
Linnaeus, 1766, and the south-east Asian Steller’s sea eagle,
H. pelagicus, Pallas, 1811 (Wink et al., 1996).
Among these three, the white-tailed eagle has the widest
distribution: it occurs from Greenland and Iceland in the west,
throughout Europe, northern and central Asia, to the Pacificcoast and Japan in the east. Breeding habitats are mostly incoastal and freshwater regions from the Arctic to thesubtropics. Prey taken in these regions are fish and waterfowl,
but in drier areas medium-sized mammals are common fooditems (Katzner, 2002). White-tailed eagles also feed on carrion,
especially in winter. Nests are built in trees as well as on cliffs
or on the ground. Except in some northern populations,studied territorial pairs are mainly sedentary (Glutz vonBlotzheim et al., 1971; Helander & Stjernberg, 2003). Younger
birds are vagrant: long-distance wandering behaviour has beenrecorded for juveniles, for example from northern Europe
down to Bulgaria (Glutz von Blotzheim et al., 1971). Despite
this, ringing data from Europe suggest strong philopatry, withindividuals typically settling to breed close to their natal area
(Helander, 2003), a pattern in accordance with variation at
mitochondrial DNA (mtDNA) and autosomal microsatellite
markers in north European populations (Hailer et al., 2006).
Nevertheless, over long time-scales white-tailed eagles havebeen shown to be capable of long-distance dispersal and1194

colonization, as documented by the colonization of Iceland,
Greenland, and Hawaii (population today extinct; Fleischer
et al., 2000). Moreover, the fossil record indicates that thewhite-tailed eagle colonized northern latitudes very soon after
glacial retreat (Ericson & Tyrberg, 2004), compatible with its
being a habitat generalist and indicating high dispersal
potential.
Given this flexibility in habitat and food choice, the whitetailed eagle offers a good opportunity to test for the expected
weak population structure at a continental level in a generalist
species with high dispersal potential. We therefore utilized
genetic markers to deduce the effect of the Pleistocene climatic
shifts on this species. We chose mtDNA as a marker because itis especially suitable for the detection of phylogeographic
structure within animal species (Avise, 2000).
MATERIAL AND METHODS
Study populations, sampling and DNA extraction
A total of 237 white-tailed eagle individuals from 11 breeding
populations throughout the range of the species were analysed(sampling locations are indicated in Fig. 1; sample sizes foreach population are shown in Tables 1 and 2). For 228 of theindividuals we took blood samples of nestlings (n ¼ 209) orcollected moulted feathers (n ¼ 19) from breeding adults inclose vicinity to their nests. This ensured that we did not mix
local breeders with vagrants or dispersing animals in theanalysis, which is important when studying a species in which
wintering individuals can be found several hundreds tothousands of kilometres away from their original breeding
population (Helander & Stjernberg, 2003). We sampled only
one offspring per breeding pair, to avoid inclusion of close
relatives, at least with regard to the present generation.
Furthermore, in order to survey for additional haplotypes wesampled nine individuals whose natal origin could not besafely assigned to any of the 11 regions: one adult foundinjured in southern Sweden during the winter season, twonestlings from the Baltic island of Gotland (situated between
Sweden and the Baltic States, two well-sampled regions), one
feather found in north-eastern Poland during late winter,one adult individual found dead in Kazakhstan during
summer, and four presumably unrelated individuals from azoo in Kazakhstan. These samples were not included in the
population-level analyses.
Blood samples were stored in EDTA/SSC buffer and kept
frozen until treatment in the laboratory, feather samples
were kept at room temperature under dry and dark
conditions. DNA from blood was extracted using a standard
phenol-chloroform procedure after digestion with proteinase
K (Sambrook et al., 1989). For feather quills we used the
DNeasy Tissue Kit (Qiagen, Hilden, Germany) and followed
the protocol of Horváth et al. (2005). DNA from feathersfrom Kazakhstan was extracted using an ammonium acetate precipitation protocol, as described in Rudnick et al.
(2005).

Journal of Biogeography 34, 1193–1206ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd

White-tailed eagle mtDNA phylogeography

Gr

Ice

Nor
Swe

Lap

Amur

Jap

Kola

Ger
Est

Kaz

Figure 1 Locations and haplotype frequencies of studied white-tailed eagle populations. Haplotype colours correspond to those in Fig. 2,
and the positions of population names indicate approximate sampling locations. Locality codes are explained in Table 1. For northern
Europe, limits of the ice sheets during the last glacial maximum (LGM) reconstructed by Svendsen et al. (2004) have been superimposed on
the map (white line; ice margins in other regions are not shown). Inside the glacial limit, younger (about 14,000 yr bp) ice-dammed lakes in
the Baltic Sea depression and around Lake Onega are shown in light blue (following Mangerud et al. 2004).

Polymerase chain reaction (PCR) amplification andDNA sequencing
In order to find a suitable marker for the present study weinitially amplified and sequenced 400 base pairs (bp) of the
mtDNA cytochrome b (Cyt-B) gene in a total of 32
individuals (21 from Sweden, four from Greenland, four
from Germany and three from eastern Russia). This recovered four variable sites defining four haplotypes (data notshown). Given this restricted amount of sequence variation,
we subsequently focused on the non-coding control region
instead.
The complete mtDNA control region (> 1500 bp, including
two heteroplasmic tandem repeats of 11 bp each that hampered exact length estimation) was amplified in a few
individuals from different geographic origins as described in
Hailer et al. (2006). Next, a 544-bp region spanning domains I
and II that contained most of the control-region variability wastargetted using the primers Hal-HVR1F (5¢-CCCCCCCTATGTATTATTGT-3¢) and Hal-HVR1R (5¢-TCTCAGTGAAGAGCGAGAGA-3¢), both located within the control region. PCR
reactions were carried out in 10-lL volumes containing
approximately 15 ng of genomic DNA, 0.3 lm of each primer,
0.2 mm of each dNTP, 0.25 units of HotStarTaq DNA
polymerase (Qiagen) in 1· HotStarTaq (Qiagen) reaction
buffer containing Tris-Cl, KCl, (NH4)2SO4 and 1.5 mm MgCl2.
PCR was performed in a PTC-225 instrument (MJ Research,
Watertown, USA) using the following thermal profile: 15 min
at 95C prior to 36 cycles of 30 s at 56C, 30 s at 72C and 30 s
at 95C; followed by a final 1-min step at 56C and an
extension step of 10 min at 72C. PCR products were cleaned
using the ExoSAP enzyme kit (Amersham Biosciences,
Uppsala, Sweden), and DNA sequencing of both strands was
performed using the original PCR primers and the DYEnamic
ET Terminator kit (Amersham Biosciences). Sequencing
reactions were cleaned using AutoSeq plates (Amersham
Biosciences) and run on a MegaBACE 1000 (Amersham
Biosciences) capillary instrument according to the manufacturer’s recommendations. Electropherograms were assembled,
checked manually, and aligned using sequencher 4.1.4 (GeneJournal of Biogeography 34, 1193–1206ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd

1195

F. Hailer et al.
Table 1 Variable sites and absolute frequencies of the mtDNA control-region haplotypes in the 11 study populations.
Site no.

Population

Haplotype 005 008 009 041 092 108 172 177 192 201 494 497 Gr Ice Nor Ger Swe Lap Est Kola Kaz Amur Jap Count
A01
A02
A03
B01
B02
B03
B04
B05
B06
B07
B08
B09
C01
Total

T
.
.
.
.
.
.
.
.
.
.
.
C

T
.
.
C
C
C
C
.
C
C
C
C
.

C
.
.
T
T
T
T
T
T
T
T
T
T

T
A
.
.
.
.
.
.
.
.
.
.
.

A
.
.
.
.
.
.
.
G
.
.
.
.

T
.
.
C
.
C
C
C
C
C
C
C
.

C
.
.
.
.
.
.
.
.
.
T
.
.

A
.
.
.
.
.
.
.
.
G
.
.
.

G
.
.
A
A
A
A
A
A
A
A
.
A

C
.
.
.
.
.
T
T
.
.
.
.
.

C
.
.
T
T
T
T
T
T
T
T
T
.

G
.
T
A
A
.
A
A
A
A
A
A
.

7

7

1

19

32

4
12

17
1

12

3
1

1

3

1

2

20

5
1

4

13

1

2
2
1

3

1
1
2

2
1
1

18
3

7

1

1

22

8

(1)*
7
2

8

26

33

18

6
44

22

12

10

25

86
14
20
72
6
2
4
2
2
5
7
2
6
228

Gr: Greenland, Ice: Iceland, Nor: Norway, Ger: Germany, Swe: Swedish coast, Lap: Swedish Lapland, Est: Estonia, Kola: Kola Peninsula, north-west
Russia, Kaz: Kazakhstan, Amur: Amur river, eastern Russia, Jap: Japan.
*Data from an individual found dead in Kazakhstan (not certified to be a local breeder) are included in brackets, but not included in the count.

PopulationnNH

H (±SE)p (±SE)

Frequency of
haplogroup A

Greenland
Iceland
NorwayGermany
LaplandSwedenEstonia
Kola peninsula
Kazakhstan
Amur
Japan
Overall

8
26
33
18
22
44
12
10
25
22
8
228

2
2
2
3
6
4
5
7
4
3
2
13

0.250 ± 0.180
0.409 ± 0.083
0.061 ± 0.056
0.523 ± 0.112
0.667 ± 0.092
0.640 ± 0.038
0.818 ± 0.070
0.933 ± 0.062
0.657 ± 0.071
0.325 ± 0.117
0.250 ± 0.180
0.746

0.00050 ± 0.00071
0.00082 ± 0.00087
0.00073 ± 0.00080
0.00345 ± 0.00236
0.00686 ± 0.00406
0.00661 ± 0.00385
0.00706 ± 0.00435
0.00507 ± 0.00336
0.00371 ± 0.00245
0.00068 ± 0.00078
0.00050 ± 0.00071
0.00680 ± 0.00012

1.00
1.00
0.97
0.89
0.55
0.41
0.33
0.10
0.12
0
0
0.53

Codes, Ann Arbor, MI, USA). After removal of primer
sequences and some additional bases close to the primers, thisyielded a 500-bp fragment for analysis.
Several lines of evidence indicate that we did not sequence a
nuclear copy of the mitochondrial control region (a numt).
First, we found only three individuals with double peaks (i.e.
potential heterozygote positions, confirmed by resequencing of
new PCR product) among all analysed electropherograms. All
three instances were from blood samples. Given the overallhigh haplotype diversity (H ¼ 0.746), and assuming Hardy–
Weinberg equilibrium, we would expect 177 (0.746 · 237)
heterozygotes among the 237 analysed individuals if oursequences were from nuclear autosomal inserts. Thus, heteroplasmy is a much more likely explanation for the observed
double peaks. Second, we obtained identical sequences fromthe same individual using a variety of PCR primers. Third, allhaplotypes obtained from feather samples were also encountered
1196

Table 2 Estimates of within-population
variability of partial control-region sequencesof white-tailed eagle mtDNA. Sample size (n),number of unique haplotypes (NH), haplotype
diversity (H), nucleotide diversity (p), and thefrequency of group A haplotypes are reported.

in blood samples, compatible with an identical genomic origin.
Hardened feather quills have earlier been suggested to beparticularly good sources of mitochondrial DNA sequences(Sorenson & Quinn, 1998). Fourth, the observation of atransition–transversion ratio around 7 (see Table 1 and Fig. 2)
is typical of mitochondrial rather than nuclear DNA (Nei &Kumar, 2000). Fifth, identical sequences were obtained on
amplifying fragment sizes between 416 and 1990 bp. Since
nuclear insertions of mtDNA tend to be of rather restricted
length (Sorenson & Quinn, 1998), this also suggests that theanalysed fragment is mitochondrial.
Data analyses
dnasp 4.10 (Rozas et al., 2003) was used to determine Tajima’s
D (Tajima, 1989), calculated based on the total number ofmutations in the alignment. To measure within-population

Journal of Biogeography 34, 1193–1206ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd

White-tailed eagle mtDNA phylogeography
A03
B08
B09
497
172

B02

C01

108

192

497

5

177

A01

B0 1
192

9

8

494
108
92

497

B07

B03
201

41

B06

B04
8

A02
B05

Figure 2 Unrooted statistical parsimony network of Eurasian white-tailed eagle mtDNA control-region haplotypes. Circle area is proportional to haplotype frequency. Dashes indicate inferred mutational steps, and numbers refer to the corresponding sites in the alignment.
Small black circles denote inferred intermediate haplotypes. Haplotype colours correspond to those in Fig. 1.

variability, arlequin 3.0 (Excoffier et al., 2005) and dnasp
were used to calculate haplotype diversity (H) and nucleotide
diversity (p, based on uncorrected genetic distances p).
To verify that our comparisons of population genetic variability(p and H) were not affected by sample size, we employed abootstrap resampling procedure using a macro in Microsoft
excel: eight or ten sequences (corresponding to the sample
sizes from Greenland, Japan and the Kola peninsula) were
sampled with replacement from each of the more extensively
sampled populations 1000 times, and p and H were calculated
for each resampling. From that we calculated the mean acrossresamplings and the 95% confidence intervals (CI) (percentile
method) of p and H.
A statistical parsimony network (Templeton et al., 1992) ofthe nucleotide sequences was constructed using the programtcs 1.21 (Clement et al., 2000) with the default setting of 95%
parsimony connection limit. Compared with bifurcating trees,
networks are better suited for the typically shallow intraspecies phylogenies in which divergence is low and ancestral
haplotypes may still exist in the population (Posada &Crandall, 2001).
We used the software modeltest 3.7 (Posada & Crandall,1998) to identify the model of sequence evolution that best fits
the data. The suggested model was more complex than any ofthe models available in the software used for subsequentcalculations. However, Nei & Kumar (2000) show that, forsequence divergences as small as those observed in this study,
complex models of sequence evolution do not greatly modify
distance estimates. To estimate the divergence time between
major phylogenetic clades, we applied the Tamura & Nei(1993) distance correction when calculating the ‘net average
distances between groups’ using mega 3 (Kumar et al., 2004).
Standard errors were estimated with 1,000 bootstrap replicatesacross sites. The obtained divergence value, which is corrected

for within-clade diversity to mimic ancestral polymorphism(Nei, 1987), was then divided by the divergence rate (two times
the mutation rate l). To our knowledge, independently
calibrated estimates of the mtDNA control-region divergence
rate have not been published for raptors. We therefore used thedivergence rate for the combined hypervariable regions I and II
of 14.8% per site per million years estimated by Wenink et al.
(1996) for Calidris alpina. We also considered a range of otherplausible rates (5–20%, see Brito, 2005) since the rate is known
to vary between species (Garcia-Moreno, 2004) and between
different parts of the mtDNA control region (Ingman &Gyllensten, 2001). Comparisons of sequence divergence amongwhite-tailed eagles sequenced for both Cyt-B and the control
region indicated that the control region evolved at least 3–4
times faster (data not shown). Therefore, we did not consider a
2% rate estimate in our analyses.
To investigate phylogeographic structure in the data set, wepartitioned the amount of genetic variation into components
within and between regions by performing an analysis ofmolecular variance (amova; Excoffier et al., 1992) as implemented in arlequin. For this analysis we used Tamura–Neicorrected distances between sequences. Significance of thecovariance components was assessed using a permutation
procedure, thus avoiding dependence on normality of the data
(Excoffier et al., 1992). We also calculated KXY, the divergence
between groups of sequences as measured by the uncorrected
average number of nucleotide substitutions per site between
populations (Nei, 1987), using dnasp. The obtained pairwise
distances between populations were used for a neighbourjoining (NJ) analysis in mega 3. The resulting tree is
influenced by many population genetic factors, including
mutation, drift and migration. Therefore, it depicts presentday overall similarity of populations, but does not necessarilyreflect ancestry relationships.

Journal of Biogeography 34, 1193–1206ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd

1197

F. Hailer et al.
The presence of limited gene flow can be indicated by apattern of isolation by distance across regions. We determined
the geographical coordinates of the approximate distribution
midpoints for each of the sampled populations. The geod
program (US Geological Service) was used to calculatedistances assuming a spherical Earth surface. Following Rousset(1997), we then plotted UST/(1 ) UST) against the log of thegeographic distances using the ibdws software (Jensen et al.,2005). The same program was used to investigate statisticalsignificance using Mantel tests with 10,000 randomizations.
To investigate the demographic history of white-tailed eagle
populations, we tested for signals of sudden population
expansion using the mismatch distribution approach fully
implemented in arlequin. The mismatch distribution is
generally multimodal in samples from populations that are in
demographic equilibrium, and unimodal when drawn from a
population that has undergone a recent demographic expansion (Slatkin & Hudson, 1991; Rogers & Harpending, 1992).
The demographic parameters Q0, Q1 and s [Q ¼ 2 u Ne,f,where Ne,f is the female effective population size before (Q0)and after (Q1) a single instantaneous expansion, and u is the
haplotype mutation rate; s ¼ 2 u t, where t is the time since
population expansion] were estimated with a generalized nonlinear least-squares approach. Deviation from the assumed
model was tested by parametric bootstrapping (Schneider &Excoffier, 1999) using a coalescent algorithm modified from
Hudson (1990). The 95% confidence intervals of the demographic parameters were estimated with 5000 replicates of thesame bootstrapping procedure.
We used arlequin and dnasp to calculate Fu’s (1997) test
statistic FS and Fu & Li’s (1993) D* and F* for each haplogroupand for the whole data set combined. These tests show
different degrees of sensitivity to deviation from neutrality
caused by demography or selection. Population growth (orgenetic hitchhiking, which can produce a similar signal) can bedetected from the patterns of significance of these tests: given a
population expansion, FS is expected to deviate significantly
from the null expectation, while D* and F* are less sensitive topopulation growth and usually do not show significant
deviation. Non-significant values of D* and F* justify exclusionof background selection (Ramos-Onsins & Rozas, 2002). In
addition, we used the maximum likelihood coalescent-based
approach implemented in lamarc 2.0.2 (Kuhner et al., 1998)
to estimate g, a parameter of population growth.
Following Roman & Palumbi (2003) and the formula for Q
given above, we used p (the expected value of Q; Nei & Kumar,2000) to determine the current genetic effective population sizeof females. For this calculation, we used the estimate of l by
Wenink et al. (1996) and multiplied it by the generation timeof white-tailed eagles (around 14.5 years; author calculations
based on data from Struwe-Juhl, 2003).
RESULTSIn the 500-bp sequences analysed in the 237 white-tailed eagle
samples we discovered 12 variable sites that defined 13 distinct
1198

mtDNA haplotypes (Table 1). The sequences have been
deposited in the EMBL database (accession numbers
AM156933 to AM156945). Tajima’s D for the whole data set
was 1.389 (P > 0.10), compatible with neutral evolution of the
DNA sequences. Haplotype relationships could be deducedwith the exception of one loop, as shown in the statisticalparsimony network (Fig. 2). No insertions or deletions wereobserved. Two of the 16 inferred substitutions were transversions, and there were at least three sites with multiple
mutations.
All sequences except haplotype C01 clustered into one of
two haplogroups, referred to as A and B (Fig. 2). Haplotypes
A01 and B01 occupied the central position in each of the two
haplogroups and were the most frequent haplotypes in the
total data set, altogether occurring in almost 70% of thestudied individuals. Within most populations, one of these two
was the predominant haplotype (Fig. 1 and Table 1). Amongthe haplotypes found at lower frequencies, three (B02, B04,
B06) were distributed over large geographic areas, occurring inboth the Baltic Sea region and central or eastern Asia.
Haplogroups A and B were found to be admixed over large
geographic areas (Fig. 1, Table 1). The two haplogroups
showed an east–west cline in their respective frequencies(Table 2). Sequences from haplogroup B had a 100% occurrence in the two easternmost populations (Amur and Japan),low to intermediate frequencies in Kazakhstan and around theBaltic Sea, and were largely absent from the populations
adjacent to the Atlantic Ocean (Greenland, Iceland andNorway). Conversely, haplogroup A went from 100% frequency in Greenland and Iceland, through varying frequencies
around the Baltic, to being rare (12%) in Kazakhstan andabsent from the two easternmost populations.
Within-population variability
The number of haplotypes per population ranged from two inGreenland, Iceland, Norway and Japan to seven on the Kolapeninsula (Table 2). Genetic variability (haplotype and nucleotide diversity) was low on the extremes of the distribution
range (Greenland and Japan), and was highest in the Baltic
region. Overall, haplotype and nucleotide diversities showedsimilar patterns. Nucleotide diversity was lowest in Greenland,
Iceland, Norway, Amur and Japan, and highest in Estonia,
Sweden and Lapland. Nucleotide diversity was strongly
affected by the degree of admixture between clades A and B,
as illustrated by the large haplotype but relatively lower
nucleotide diversity in Kola. Resampling 1000 times of eightindividuals per population (sample size from Greenland andJapan) revealed that the low nucleotide diversity observed inGreenland and Japan was outside the 95% CI of all European
populations except Germany and Norway. Resampling 1000
times of ten individuals (sample size from the Kola peninsula)from the other eight populations with larger sample sizes(Table 2) revealed that the high haplotype diversity observedin Kola was outside the 95% CI for all other populations and,
therefore, not likely to be an artefact of restricted sample size.

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White-tailed eagle mtDNA phylogeography
The net average distance between groups A and B was
0.0098 ± 0.0039 (±SD). Assuming a divergence rate of 14.8%
per site per million years (Wenink et al., 1996), the divergenceof the two clades was estimated to have occurred
66,200 ± 26,400 years ago. Divergence rates between 5 and
20% yielded average estimates between 49,000 and
196,000 years ago. The assumption that substitution rates
may be higher on shorter than on longer time-scales (see Ho &Larson, 2006) would support a more recent date for thedivergence. The origin of lineages A and B was thus confidently
placed in the late Pleistocene.

Greenland
Iceland
NorwayGermany
Lapland
Swedish coast
Estonia
Kola peninsula
Kazakhstan
Amur
Japan
0.5

Population structure
Overall, white-tailed eagle populations exhibited clear differences in their genetic composition. UST across all populations
was 0.512 (P < 0.001). Pairwise UST values between populations were significantly larger than zero in most cases(Table 3).
The neighbour-joining tree (Fig. 3) based on the mean
number of pairwise nucleotide differences between populations (KXY) roughly reflected the geographic locations of the
populations. Atlantic populations clustered closely together, as
did the Asian populations. The central and north European
populations were located between these clusters. However,genetic distances between European populations were in somecases very small and in other cases substantial, indicatingcomplex differentiation patterns on a smaller spatial scale. For
example, the sample from Germany appeared geneticallysimilar to the Atlantic ones, and the Kola population seemedsimilar to Asian populations.
amova results (Table 4) confirmed the presence of phylogeographic structure in our data. Grouping white-tailed eagle
populations according to large-scale geographic regions (Atlantic islands, Europe, Asia; grouping 5) yielded a high andsignificant UCT value, indicating strong geographic differentiation. In contrast, separating Greenland from the remaining

Figure 3 Neighbour-joining tree of populations of the Eurasian
white-tailed eagle, based on pairwise KXY distances between
mtDNA control-region haplotypes.

populations (grouping 2), which would reflect current taxonomical segregation into two recognized subspecies, H. albicillagroenlandicus and H. a. albicilla, did not explain much of thevariance and was non-significant. Groupings 3 and 4 indicatedthat Greenland and Iceland were less differentiated from theremaining populations than Japan and Amur were from allothers. The best grouping we explored was number 7. This,
however, was an a posteriori grouping based on the results of theneighbour-joining population tree (Fig. 3).
A plot of genetic differentiation in relation to geographic
distance (Fig. 4) indicated a weak but significant correlation(Z ¼ 1726.40, r ¼ 0.424, one-sided P < 0.001). This correlation was largely the result of strong differentiation between thegeographically most distant populations, namely Greenlandand Iceland compared with Amur and Japan. The exclusion ofGreenland, Iceland, Amur and Japan (but still including
Kazakhstan) yielded a non-significant correlation (Z ¼ 142.41,
r ¼ 0.248, one-sided P > 0.15).
The individuals we had left unassigned to any breeding
populations generally carried haplotypes found in individuals

Table 3 Genetic differentiation between white-tailed eagle populations. Pairwise UST values based on Tamura–Nei distances are shown
below the diagonal. Corresponding significance as assessed by 2024 permutations is indicated by a plus (P < 0.05) or minus (nonsignificant)above the diagonal. More conservatively, to account for multiple testing, values in bold indicate significance at the 0.05 level after sequential
Bonferroni correction.
Greenland
Greenland
Iceland
NorwayGermany
LaplandSwedenEstonia
Kola
Kazakhstan
Amur
Japan

0.473
)0.028
0.236
0.258
0.310
0.510
0.724
0.746
0.947
0.960

Iceland

NorwayGermany

LaplandSwedenEstonia

Kola

Kazakhstan

Amur

Japan

)

)
+

+
+
+

+
+
+
+

+
+
+
+
)

+
+
+
+
)
)

+
+
+
+
+
+
)

+
+
+
+
+
+
+
)

+
+
+
+
+
+
+
+
+

+
+
+
+
+
+
+
)
)
)

0.566
0.495
0.423
0.428
0.661
0.811
0.800
0.937
0.940

0.346
0.380
0.390
0.666
0.824
0.809
0.939
0.943

0.269
0.305
0.452
0.637
0.683
0.837
0.798

)0.010
0.042
0.195
0.298
0.470
0.387

Journal of Biogeography 34, 1193–1206ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd

0.021
0.154
0.237
0.365
0.320

)0.008
0.108
0.307
0.204

0.024
0.105
0.048

0.101
0.052

)0.0061199

F. Hailer et al.
Table 4 Analysis of molecular variance (amova) describing the partitioning of mitochondrial DNA haplotype variation across a range ofconceivable population (pop) groupings.

Population grouping

Among groups(UCT)

Among populations
within groups

Within populations

1. All in one group
2. [Gr] [all remaining pops]
3. [Gr/Ice] [all remaining pops]
4. [Amur/Jap] [all remaining pops]
5. [Gr/Ice] [Eur1] [Kaz/Amur/Jap]
6. [Gr/Ice] [Eur1/Kaz] [Amur/Jap]
7. [Gr/Ice/Nor/Ger] [Swe/Lap/Est] [Kola/Kaz/Amur/Jap]–
0.038n.s.
0.211n.s.
0.373*
0.401**
0.308*
0.523**

0.512
0.491
0.367
0.267
0.185
0.272
0.053

0.488
0.471
0.422
0.359
0.414
0.421
0.424

1

Eur denotes the northern and central European populations [Nor, Swe, Lap, Kola, Est and Ger], each treated separately within the group.
P > 0.05, *P < 0.05, **P < 0.01, as assessed by 10,100 permutations.

n.s.

Figure 4 Isolation-by-distance plot of Eurasian white-tailed eagle
populations, with genetic distance plotted against the log of geographic distances. Filled circles correspond to comparisons
between the most geographically distant populations (Greenlandor Iceland vs. Amur or Japan).

from neighbouring populations. Interestingly, however, theadult found dead in Kazakhstan carried haplotype B06,
otherwise present in Lapland and the Kola peninsula, but
not previously detected in the Kazakhstan sample.
Demography
Phylogenetic analysis (Fig. 2) revealed the presence of two
distinct clades (A and B) and thus suggested past evolutionof white-tailed eagles in two groups. We therefore investigated population history not on the basis of the current
geographically defined populations, but instead on the basisof the past subdivision and evolutionary divergence of the
major phylogenetic lineages (as in Flagstad & Roed, 2003;
Godoy et al., 2004). Haplogroup B exhibited a frequentcentral haplotype together with several closely related derived
1200

haplotypes at lower frequencies. Such a star-shaped pattern
is a common characteristic of lineages following a demographic expansion (Slatkin & Hudson, 1991). Haplogroup A
also shows a predominant central haplotype. However, only
two other derived haplotypes were encountered within thisclade.
Fu’s (1997) FS statistic was significant for clade B ()5.54,
P < 0.001), but not for clade A (0.80, P > 0.10), while Fu &Li’s (1993) F* and D* were non-significant (P > 0.05) for
both B (F* ¼ 0.38, D* ¼ 1.26) and A (F* ¼ 0.71,
D* ¼ 0.001). Results consistent with this were obtained withthe program lamarc, which yielded a growth parameter ofg ¼ 2830 (95% CI: 669 to 9062) for clade B, but a value forclade A (g ¼ 1172) whose 95% CI included negative values(95% CI: )12 to 8793), thus not excluding population decline
or stasis for A. In summary, these tests indicate a population
expansion in haplogroup B, but give inconclusive results for
haplogroup A.
The mismatch distribution analyses yielded a bimodal
pattern for the total data set (results not shown). For
haplogroup A, the least-squares procedure in arlequin did
not converge when fitting the model to the observed data.
For lineage B, deviation from the sudden expansion model
was not significant (Q0 ¼ 0.000, Q1 ¼ 483.1, P > 0.05). The
peak of the corresponding mismatch distribution was at
s ¼ 0.689 (95% CI: 0.335 to 0.951). Using the formula t ¼ s/
(l 2 l), where l and l represent the sequence length and thesubstitution rate per site and million years, and using thedivergence rate calibration by Wenink et al. (1996) of 14.8%
per bp and million years, we estimated that the expansion of
haplogroup B took place 9311 yr bp (95% CI: 4527 to
12,851). Using divergence rates of between 5% and 20% (seeBrito, 2005) yielded average values for the time since
expansion of between 6890 and 27,560 yr bp – thus aroundthe Pleistocene–Holocene transition across a wide range ofparameters.
Values of nucleotide diversity (p) for the eastern (B) andwestern (A) clade were 0.00120 and 0.00098, corresponding toa genetic effective population size of females (Ne,f) of around
560 and 460 females, respectively.

Journal of Biogeography 34, 1193–1206ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd

White-tailed eagle mtDNA phylogeography
DISCUSSION
Species-level diversity
Our study revealed relatively few haplotypes and a shallow
within-species divergence between the white-tailed eagle
haplogroups. Over a broad range of conceivable mutation
rates, the divergence of the major lineages present in thespecies dates back 50,000 to 200,000 years, pointing to an
intra-species lineage splitting during the late Pleistocene. Thispattern has been observed in several other Northern Hemisphere, especially boreal, species of birds (Lovette, 2005).
Pleistocene climatic shifts are a likely explanation for both the
splitting into an eastern and western group and the recent
diversification of each haplogroup.
Another related large raptor with a wide geographic
distribution is the bearded vulture (Gypaetus barbatus,
Linnaeus, 1758). The white-tailed eagle harbours less overall
nucleotide diversity than the bearded vulture (about 0.7%, ascompared with 2.9% in the bearded vulture; Godoy et al.,2004). The bearded vulture thus appears to have retained
higher effective population sizes than the white-tailed eagle
has. This may indicate a higher sensitivity of white-tailed eagles
to climatic fluctuations, and/or be related to the wider
distribution range of the bearded vulture, which extends tosouthern Africa. Furthermore, the amount of mtDNA diversity
in the white-tailed eagle is similar to that described for controlregion sequences of another raptor, the red kite (Milvus milvus,
Linnaeus, 1758; Roques & Negro, 2005), despite the distribution of the latter being largely restricted to Europe.
Phylogeographic structure and range contraction
during cold periods
MtDNA control-region sequences of the white-tailed eagle
clustered into two distinct major haplogroups with an east–west cline through Eurasia (Fig. 1, Table 2). Such a pattern has
been observed in many other Eurasian and North American
taxa (Hewitt, 2000; Ruokonen et al., 2004; Lovette, 2005) andprobably reflects range contraction into two allopatric refugia,
followed by postglacial re-expansion. The finding of haplotype
C01 in the data set may result from retained ancestralpolymorphism, or indicate the presence of a third refugium.
For species like the white-tailed eagle it is not clear if a
glacial refugium can be envisioned in a similar way as the
traditional refugia of temperate forest species. White-tailed
eagles have a large dispersal capability and vagrant individuals
can cover vast areas. Populations live at rather low densities,
and it is possible that large areas were required to sustain viable
populations during glacial maxima. A glacial refugium for thewhite-tailed eagle may therefore have spanned large areas andvaried spatio-temporally along with climatic and other environmental changes.
The restricted number of region-specific haplotypes prevents us from establishing precise locations for the refugia.
However, our data suggest that the white-tailed eagle survived

the last glaciation period in at least two Eurasian regions,
neither of which was likely to have been on the Pacific coast.
The populations in eastern Russia and Japan show low
haplotype and nucleotide diversity and share all their haplotypes with the European populations. This pattern of genetic
diversity is characteristic of postglacially founded populations.
The only extant breeding population of Steller’s sea eagles(Haliaeetus pelagicus) is located on the Pacific coast of Asia. In
areas of sympatry with the white-tailed eagle, competitive
advantage with regard to both nesting sites and prey has been
documented for Steller’s sea eagle (Masterov, 1992). Whether
or not competitive exclusion by Steller’s sea eagles had an
impact on the survival and distribution of the white-tailed
eagle during glacial times is not known.
The western refugium could have been located in western
Europe, possibly along the Atlantic coast. A glacial refugium
somewhere in that region has been postulated for many
species, for example coastal birds (seagulls and eider ducks;
Tiedemann et al., 2004; Liebers et al., 2004), several fishes(e.g. Volckaert et al., 2002) and a seaweed (Provan et al.,2005). Compatible with this, the white-tailed eagle has beenreported as the most abundant diurnal raptor in the
Pleistocene fossil record of the Iberian peninsula (SánchezMarco, 2004; Antonio Sánchez-Marco, personal communication). A glacial refugium in Iberia has been postulated for the
Spanish imperial eagle (Aquila adalberti, C. L. Brehm, 1861
Ferrer & Negro, 2004).
The location of the eastern refugium could be the region
around the Aralo-Caspian and Black Sea basin. Water levels of
the Caspian and Black Sea during the last glacial maximum(LGM) were considerably higher than they are today(Grosswald & Hughes, 2002), implying a larger surface andthus longer coastline. The region has a high degree ofendemism (Dumont, 1998) and has been proposed as glacial
refugium for many other species linked to aquatic habitats, forexample fishes (Bernatchez, 2001; Kotlik et al., 2004), crustaceans (Audzijonyte et al., 2005) and seagulls (Liebers et al.,2004).
The roles of the Danube river system and the Mediterranean
coast as possible glacial refugia for the white-tailed eagle
remain unclear. These alternatives are difficult to assessbecause many historic populations have gone extinct in Spain,
Italy and France, or have recently undergone dramatic declines
in Greece, Albania, Serbia, Croatia and Romania (Helander &Stjernberg, 2003).
The survival of white-tailed eagles in these refugia is very
similar to the scenario proposed by Liebers et al. (2004) for the
herring gull complex, a taxonomic group resembling thewhite-tailed eagle in many ecological features regarding theirpredominant habitat choice and foraging requirements. Also
for the herring gull complex, mtDNA indicates Ice Age survival
in a Western European and one central Eurasian refugium(Liebers et al., 2004). Such congruence of phylogeographic
patterns among different species with similar requirements
supports the importance of ecological factors in shaping
current mtDNA variation.

Journal of Biogeography 34, 1193–1206ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd

1201

F. Hailer et al.
Current population-level diversity

Demography: population expansion out of the
refugia
The fossil record indicates that the white-tailed eagle was an
early postglacial colonizer of northern regions. Bone remains(9000 14C yr bp) have been recovered from southern Sweden(Ericson & Tyrberg, 2004) and from near Stavanger, Norway(7000–8000 yr bp; J Mangerud, University of Bergen, Norway,
personal communication). In accordance with this, resultsfrom the mismatch distribution analysis for the eastern clade(B) indicated a sudden population expansion from the eastern
refugium at or after the latest stages of the Pleistocene(< 30,000 yr bp; average estimate with rate calibration by
Wenink et al., 1996: 9311 yr bp). This demographic growth
may have been triggered by glacier retreat and climate
warming following the LGM (20–15,000 yr bp; Svendsen et al.,2004). After the LGM, climate warming led to an increased
availability of coastal landscapes. Extensive new suitable
habitats appeared where meltwater from retreating glaciers
accumulated, and an abundance of ice-dammed lakes, rivers
and other water systems existed from the Caspian Sea towards
the Baltic region (Mangerud et al., 2004; see also Fig. 1).
The demography of the western white-tailed eagle clade A isdifficult to deduce from our data. While it is possible that thisgroup did not expand as markedly as the eastern group B, orthat it expanded later, a third option is also conceivable.
During the last 150 years, white-tailed eagles have disappeared
from vast parts of their historic distribution range in southern
and western Europe (Helander & Stjernberg, 2003). Especially
if those regions harboured refugial populations, loss ofdiversity during recent centuries may have influenced our
results (see Leonard et al., 2005). We may thus underestimate
the original clade A diversity and miss signals of a postglacial
population expansion.
Our results indicate that Iceland and Greenland werecolonized by white-tailed eagles from northern or western
Europe, possibly via the Faroe islands. Two haplotypes wereencountered in Iceland and Greenland, namely A01 and A03(Table 1). A01 is the presumed western refugial haplotype,while A03 is a derived form currently restricted to Iceland andGreenland and which probably arose during postglacial range
expansion. A similar colonization history has been documented for many other animal taxa now present in Greenland and/or Iceland (Sadler, 1999; Tiedemann et al., 2004; MuñozFuentes et al., 2006), i.e. a predominantly Palaearctic rather
than Nearctic origin.
Demographic responses to climate change have previously
been proposed to vary in timing and intensity at different
latitudes (Hewitt, 2000; Lessa et al., 2003). In the white-bellied
sea eagle (Haliaeetus leucogaster, Gmelin, 1788), which ismainly distributed in tropical and subtropical regions (India
through Southeast Asia to Australia), a major population
expansion was dated to have occurred around 160,000 yr bp(Shepard et al., 2005; also analysing the mtDNA control
region). The authors related this expansion to a period oflowered sea levels, enabling colonization of new habitats.
1202

For several European populations of the white-tailed eagle,
levels of mtDNA diversity are relatively high, reflecting theadmixture of sequences from divergent clades. However, thefinding of high mtDNA variation in northern populations that
inhabited regions covered by glaciers during the LGM isnoteworthy. Confirmed by all diversity measures, the highest
variability was found in the populations surrounding the Baltic
Sea (Table 2), and not in regions that were unglaciated during
the LGM. Several factors can explain this pattern of highgenetic diversity in northern populations. First, an eradicationof diversity in the south might have occurred. However, thespecies has a long generation time and has been shown to berelatively resilient against loss of genetic diversity, at least
within the time perspective of a few decades (Hailer et al.,2006). Second, the population expansion could have occurred
mainly in the north. Most haplotypes are separated from either
A01 or B01 by just one or two mutational steps. Our dating of
the population expansion of clade B indicates that the age ofthese derived haplotypes postdates the LGM and thus points
towards an origin in the postglacially expanding populations.
Following glacier retreat, the vast water masses arising in
proglacial regions of northern Europe (see above) might haveconstituted highly productive habitats enabling early colonization and pronounced population expansion. In summary,
we propose a scenario of Ice Age survival in southerly regions,followed by rapid postglacial colonization of northern habitats,major population expansion, and generation of a large
proportion of the current mtDNA gene pool of the species.
On a regional scale, despite clear general phylogeographic
structuring of mtDNA diversity (Fig. 3, Table 4), someobserved patterns deviate from the large-scale picture. Comparing the neighbouring populations from Norway and Kolapeninsula, very different patterns of genetic diversity areobserved. While the Norwegian population is largely fixed for
haplotype A01, a variety of different haplotypes were found on
Kola (Table 1). This difference between neighbouring populations is indicative of low gene flow and stresses theimportance of local population history for present-day genetic
variability. Because the Norwegian population has been thelargest in Europe during the last century, this pattern is not theresult of any recent declines (Hailer et al., 2006).
Conservation implications
White-tailed eagle mtDNA suggests the absence of populations
that should be defined as evolutionary significant units (ESUs;
see Moritz, 1994). The two major lineages (A and B) have abroad zone of admixture and are thus largely sympatric today,at least in Europe. However, the present distribution range andrecent history of some populations indicate that they should be
regarded as largely isolated reproductive units.
For Greenland, some authors have proposed a separate
subspecies (H. albicilla groenlandicus; see Glutz von Blotzheim
et al., 1971). The fossil record indicates that the white-tailed

Journal of Biogeography 34, 1193–1206ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd

White-tailed eagle mtDNA phylogeography
eagle colonized Greenland during the Holocene. Salomonsen(1979) suggested that this might have been during the
hypsithermal interval, between 6000 and 4000 yr bp. Given
the absence of well-differentiated haplotypes from Greenland,
and the likely postglacial origin of haplotype A03, our data arecompatible with this recent colonization. However, given thelack of reciprocal monophyly, our data do not lend strong
support to the subspecies distinction. MtDNA structuring andsubspecies distinctions commonly do not coincide for avian
taxa (Zink, 2004), suggesting that ecological and/or morphological distinction may be attained more quickly than mtDNA
lineage sorting (Bulgin et al., 2003). As emphasized byCrandall et al. (2000), ESUs should also be defined on thebasis of ecological characteristics. A series of morphologicalmeasurements (e.g. bill length, wing length and egg size) are
known to vary along a cline from north-west to south-east
throughout Eurasia, which led Glutz von Blotzheim et al.
(1971) to refute the status of white-tailed eagles fromGreenland as a separate subspecies (H. a. groenlandicus).
However, as Salomonsen (1979) pointed out, skeletal measurements of H. a. groenlandicus and H. a. albicilla do not
overlap, possibly supporting the subspecies distinction. The
low mtDNA diversity in Greenland and Iceland, and theunique occurrence of haplotype A03 strongly suggest that these
populations have long been isolated from other white-tailed
eagle populations, and they may today also be isolated fromeach other. Such small and isolated populations deserve special
attention and high conservation priority.
ACKNOWLEDGEMENTS
Sampling of blood was coordinated by the Swedish Society for
Nature Conservation/Project Sea Eagle. We thank Todd
Katzner, Evgeny Bragin, Andreas Marten and Sascha Rösnerfor contributing samples, and Kurt Elmqvist, Robert Franzén,
Robert Stefansson, Finnur Johannsson, Hallgrimur Gunnarsson and Menja von Schmalensee for assistance with thesampling of blood. Martin Jakobsson made the map, and
J. Andrew DeWoody kindly allowed us to use his laboratory
for some analyses. We are grateful to Jennifer Leonard, Øystein
Flagstad, Jan Mangerud, Violeta Muñoz, Todd Katzner, Ülo
Väli, Love Dalén and Jennifer Seddon for insightful discussions
and/or comments on the manuscript. Hans Ellegren is thanked
for discussions and logistic and economic support. This work
was financially supported by Alvin’s foundation, the Sven and
Lilli Lawski foundation and the Knut and Alice Wallenberg
foundation (to F.H.).
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F. Hailer et al.
BIOSKETCH
Frank Hailer did his PhD work in the research group of Carles Vilà and Hans Ellegren, studying conservation genetics of thewhite-tailed eagle.
Jamie Rudnick uses genetic analyses on non-invasively collected feathers to investigate eagle biology in central Asia.
Björn Helander is senior scientist responsible for the monitoring of white-tailed sea eagle reproduction and population trends
within the National Environmental Monitoring Programme under the Swedish Environment Protection Agency. He is also leader ofProject Sea Eagle run by the Swedish Society for Nature Conservation/SNF.
Alv O. Folkestad, Sergey Ganusevich, Steinar Garstad, Peter Hauff, Christian Koren, Vladimir Masterov, Torgeir
Nygård, Saiko Shiraki, Kristinn Skarphedinsson, Veljo Volke and Frank Wille work with various aspects of raptor biology
and contributed samples to this study.

Editor: Brett Riddle

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