Bird Study

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The origins of Great Spotted Woodpeckers

Dendrocopos major colonizing Ireland revealed by
mitochondrial DNA

Allan D. Mcdevitt , Łukasz Kajtoch , Tomasz D. Mazgajski , Ruth F. Carden ,

Ilaria Coscia , Christian Osthoff , Richard H. Coombes & Faith Wilson

To cite this article: Allan D. Mcdevitt , Łukasz Kajtoch , Tomasz D. Mazgajski , Ruth F. Carden ,
Ilaria Coscia , Christian Osthoff , Richard H. Coombes & Faith Wilson (2011) The origins of Great
Spotted Woodpeckers Dendrocopos�major colonizing Ireland revealed by mitochondrial DNA, Bird
Study, 58:3, 361-364, DOI: 10.1080/00063657.2011.582619

To link to this article: https://doi.org/10.1080/00063657.2011.582619

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Bird Study (2011) 58, 361– 364

SHORT REPORT

The origins of Great Spotted Woodpeckers

Dendrocopos major colonizing Ireland
revealed by mitochondrial DNA

ALLAN D. MCDEVITT1∗ , ŁUKASZ KAJTOCH2, TOMASZ D. MAZGAJSKI3, RUTH F. CARDEN4,

∗
ILARIA COSCIA5, CHRISTIAN OSTHOFF6, RICHARD H. COOMBES7 and FAITH WILSON8
1
Mammal Research Institute, Polish Academy of Sciences, Białowieża, Poland, 2Institute of Systematics and
Evolution of Animals, Polish Academy of Sciences, Slawkowska 17 Street, 31-016 Krakow, Poland, 3Museum
and Institute of Zoology, Polish Academy of Sciences, Wilcza 64 Street, 00-679 Warsaw, Poland, 4National
Museum of Ireland – Natural History, Merrion Street, Dublin 2, Ireland, 5Institute of Biological, Environmental and
Rural Sciences (IBERS), Edward Llwyd Building, Aberystwyth University, Aberystwyth, Ceredigion SY23 3DA, UK,
6
Carrigmore, Glenealy, Co. Wicklow, Ireland, 767 Delgany Park, Killincarrig, Co. Wicklow, Ireland and
8
Kilnamanagh Farm, Kilnamanagh Beg, Glenealy, Co. Wicklow, Ireland

Capsule Although necessarily based on a small number of samples, comparisons of molecular data from
the newly established Great Spotted Woodpecker populations in Ireland with those in Britain and continen-
tal Europe revealed that Britain was the more likely source area of the Irish populations.

Great Spotted Woodpeckers Dendrocopos major were,
until recently, absent as a breeding species on the
island of Ireland (Winkler et al. 1995). The species is
known from (unconfirmed) archaeological records from
two bones in the Edenvale Cave complex in County
(Co.) Clare (Scharff et al. 1906), with woodland loss
thought to be responsible for the disappearance of
several bird species (including D. major) in Ireland
(Yalden & Carthy 2004). Occasional, solitary birds
have been seen/found in Ireland (Stendall 1936) but
these typically occur during the winter months outside
of the breeding season and specimens exist in the
museum collections in the National Museum of
Ireland, Natural History Division (NMINH) in Dublin.
Sightings of birds during the breeding season have
increased in recent years in Ireland and territorial drum-
ming behaviour has been increasingly observed. Breeding
was confirmed in Northern Ireland in 2007 (McComb
et al. 2010). Concurrently, sightings and observations
of territorial behaviour and courtship increased in a sep-
arate area in Co. Wicklow in the east of the Republic of
Ireland with breeding first confirmed in the county in
May 2009 (McComb et al. 2010). A total of 18 nesting
∗
Correspondence authors. Emails: amcdev@gmail.com; wilson@es
atclear.ie
attempts have been documented in Co. Wicklow in
the last 2 years, potentially producing up to 50+ nestlings
in the county (F. Wilson et al. unpubl. data).
Two schools of thought currently exist on the origins
of these two newly breeding populations in Ireland. One
is that the species arrived from Ireland’s nearest land-
mass, Britain, as the result of recent and rapid increases
in the normally sedentary population there (Risely
et al. 2008, McComb et al. 2010). Alternatively, the
species could have arrived from Scandinavia because
the Scandinavian birds are known to cross the North
Sea when irruptive migrations occur (triggered by poor
crops of pine/spruce seed and density (Lindén et al.
2011)), with individuals straying up to 3000 km
(Winkler et al. 1995, Coulson & Odin 2007). To inves-
tigate this, we compared control region sequences (mito-
chondrial DNA (mtDNA)) from modern and museum
specimens in Ireland to those from Britain and continen-
tal Europe in a phylogeographic analysis to decipher the
origins of the newly established populations and past
solitary vagrants in Ireland. We used control region
sequences instead of previously utilized mitochondrial
markers because cytochrome b and NADH dehydrogen-
ase genes (Zink, Drovetski et al. 2002, Garcia-del-Rey
et al. 2007) were previously shown to be relatively unin-
formative in the European range of D. major.

Q 2011 British Trust for Ornithology

362 A.D. McDevitt et al.

A total of 41 modern Irish, British and continental

European individuals (Table 1; Fig. 1a) were collected
and subjected to molecular analysis. Shed feathers were
collected non-invasively from fledged nests in Ireland,
and blood and feather samples were also collected from
captured individuals (under license) in Ireland, Britain
and continental Europe. Tissue samples were also
taken from individuals found dead or from individuals
recently deposited in museum collections. All modern
material was collected during the breeding season,
thereby reducing potential sampling of birds on
migration. Furthermore, DNA extraction was attempted
on tissue and feather samples taken from two isolated
individuals found in Ireland in an undisclosed location
in 1880 (tissue; NMINH: 1880.785.1) and in Co.
Offaly (Fig. 1a) in 1950 (primary feather; NMINH:
1950.22.1). These were taken from individuals kept in
the NMINH in Dublin. Details of the molecular
methods and analyses are given in the supplementary
material to this paper, which are available via the

Table 1. Map number (see Fig. 1a), locality name and country of
origin for individuals (n) used in this study. Haplotypes (with the
number of individuals in parentheses) in each locality are indicated
(see Fig. 1b for haplotypes).

Map
no. Locality Country n Haplotypes

1 Co. Wicklow Rep. of 5 H2, H4(2),

Ireland H6(2)
2 Co. Offaly (1950) Rep. of 1 H1
Ireland
– Unknown (1880) Rep. of 1 – Figure 1. (a) Sampling localities of Dendrocopus major in Europe.
Ireland See Table 1 for corresponding numbers and names of localities; (b)
3 Co. Down Northern 1 H1 Median-joining network of D. major haplotypes in Europe. Shading
Ireland corresponds to region of sampling and size to number of individuals
4 Blyth Bridge Scotland 1 H3 belonging to each haplotype; vertical bars represent mutational steps
5 North Berwick Scotland 1 H4 when greater than one and black squares represent missing
– Unknown Scotland 1 H4 haplotypes.
6 Hertfordshire (three England 6 H2(5), H5
sites)
7 Norfolk England 4 H1, H2,
Supplementary Content tab of the article’s online page
H5(2) at http://dx.doi.org/10.1080/00063657.2011.582619.
8 Aberystwyth Wales 1 H3 The control region fragment was successfully ampli-
9 Ouardye Ringing Belgium 2 H7, H8 fied in 42 out of the 43 individuals. Unfortunately, the
Station
10 Frankfurt Germany 1 H1
museum specimen from an undisclosed location in
11 Tägerwilerwald (two Switzerland 9 H1(7), H3(2) Ireland dated to 1880 failed to generate a fragment. Thir-
sites) teen polymorphic nucleotide sites were identified from
12 Ostfold (three sites) Norway 4 H1(2), H9, the 802 base pair (bp) fragment, which resulted in a
H10
13 Mazury District Poland 1 H9
total of 12 unique haplotypes (GenBank Accession
14 Krakow Poland 1 H11 Nos.: JF758647–JF758658) among the 42 individuals.
15 Sudeten Mountains Poland 1 H1 Overall nucleotide diversity was 0.003 + 0.00029 sd
16 Slonsk Poland 1 H1 and haplotype diversity was 0.834 + 0.043 sd. Overall
17 Carpathian Poland 1 H12
Mountains
control region diversity was low and this low level of
differentiation was found on a continental scale in

Q 2011 British Trust for Ornithology, Bird Study, 58, 361 –364

previous studies using different mtDNA genes; cyto-
chrome b, ND2 and ND3 (Zink, Drovetski et al. 2002,
Garcia-del-Rey et al. 2007) and in other woodpecker
species (Zink, Rohwer et al. 2002, Pons et al. 2011). It
is worth noting that Rutkowski et al. (2008) demon-
strated greater variability in eastern Europe using a
larger dataset from Poland based on 379 bp of the
control region. This is in agreement with Voous’
(1947) proposed postglacial recolonization routes for
the species in Europe, with several formerly separated
refugial populations meeting in the east.
Haplotype H1 was the most common haplotype and
was found in 15 individuals, including individuals in
Ireland (museum specimen from 1950 and Northern
Ireland), England, Norway, Germany, Switzerland and
Poland (Table 1; Fig. 1b). Four haplotypes were found
in Ireland (Cos Offaly (1950), Wicklow and Northern
Ireland; Table 1, Fig. 1b), five from Britain and eight
in continental Europe (Norway, Belgium, Germany,
Switzerland and Poland). Haplotype H3 was found in
individuals from Scotland, Wales and Switzerland.
Therefore, there is evidence of continental influence on
Britain’s population, with these two haplotypes being
shared with continental Europe. This corroborates evi-
dence from ringing data (Clark et al. 2004, 2007)
and wing measurements (Coulson & Odin 2007, Smith
2010) that dispersal from Scandinavia to Britain occurs
in certain years when irruptions occur in Scandinavia.
Regarding the modern samples from Ireland, eight
polymorphic sites were found, resulting in four diverse
haplotypes found in six individuals. Nucleotide diversity
was 0.00457 + 0.00069 sd. Three of the haplotypes
found were shared with those in England and continen-
tal Europe (H1), England (H2) and Scotland (H4). The
other haplotype (H6) was found in two Irish individuals
and was the most divergent haplotype found in the
network, differing from the nearest British haplotypes
by 4 bp (Fig. 1b). This haplotype was more closely
related to individuals from Britain than to the continent
(Fig. 1b), so it is highly likely that this haplotype is
present on mainland Britain but was not found in this
study owing to limited sampling. All this points strongly
to a British origin for the established Irish populations
and it appears as though the separate breeding groups
in the Republic of Ireland and Northern Ireland were
founded from different areas (Fig. 1b), but material
from only a single individual from Northern Ireland
was available to the study. The diverse nature of the hap-
lotypes found in Ireland may suggest that the separate
Irish breeding populations have been founded from
multiple localities within Britain, rather than a single
Origins of Great Spotted Woodpeckers in Ireland 363
source. The key to why the normally sedentary British
birds have dispersed to Ireland may be because of the
fact that the British population has increased
dramatically in recent decades (increasing 133% and
196% in England and Wales respectively (Risely et al.
2008)), so despite their reluctance to venture great dis-
tances, the pressure of numbers may have triggered a
range expansion of British birds across the relatively
narrow Irish Sea. The species has been increasingly
sighted in the Isle of Man recently, with nests discovered
in 2009 and 2010 (C. Sharpe, pers. comm.). This, in
conjunction with an increase in Ireland’s forest cover
(reaching about 10% cover (Forest Service, Department
of Agriculture, Fisheries and Food 2007) after being as
low as 2% (McCracken 1971)), as well as the continuing
maturation of these (D. major requires a minimum tree
diameter of about 50 cm for nesting and roosting
(Mazgajski 2002; Kosiński & Winiecki 2004)), make
conditions more favourable for future and further
expansion of the species. Previous solitary visitors to
Ireland were assumed to be of Scandinavian origin
(Coulson & Odin 2007) and the fact that the museum
specimen from 1950 shares its haplotype with the main
haplotype found in continental Europe (although also
found in one British individual) corroborates this to a
certain extent. It is clear that further examination of
museum samples, along with more data from modern
specimens, is needed to investigate this fully.
The diverse origins of the populations may bode well
for the retention and proliferation of genetic diversity in
Ireland. However, there is no doubt that future analyses
incorporating more samples and molecular markers
would be beneficial for more accurately pinpointing
the exact source populations and giving further insights
into the levels of genetic diversity present in the Irish
populations. The natural colonization and establishment
of the species is clearly a significant event and efforts
should now focus on maintaining and ensuring that
the Great Spotted Woodpecker populations continue
to expand and thrive in Ireland.
ACKNOWLEDGEMENTS
We are extremely grateful to Tom Kealy, Ken Smith, David
Fuller, Chris Murphy, Ann Fitzpatrick, Gilberto Pasinelli,
Glauco Camenisch, Rolf Keidel, Jo Ranke, Phillippe Sche-
pens, Przemek Chylarecki and Magorzata Bujoczek for provid-
ing samples for analysis and Oran O’Sullivan for sound
recording equipment. Thanks to Chris Sharpe of Manx Bird-
Life for information on birds on the Isle of Man. Licenses for
this work were provided by National Parks and Wildlife
Q 2011 British Trust for Ornithology, Bird Study, 58, 361 –364
364 A.D. McDevitt et al.
Service, Ireland and the British Trust for Ornithology. Thanks
to Andrew Kitchener and Nigel Monaghan for allowing access
to or providing museum samples from the National Museums
of Scotland, Edinburgh and the National Museum of Ireland,
Natural History Division, Dublin, respectively. We also thank
Stefano Mariani for use of the molecular lab in University
College Dublin, Alisha Goodbla and Carlotta Sacchi for tech-
nical assistance, Brian Bourke for advice on DNA extraction
from museum specimens and the two anonymous reviewers
for their comments on the manuscript. This research was
funded by the Heritage Council, Ireland (Research Grant
Scheme No. R00464, 2010).
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