Phylogeography of willow grouse (Lagopus lagopus) in
the Arctic: taxonomic discordance as inferred from
molecular data
JACOB HÖGLUND
1
*, BIAO WANG
1
, THOMAS AXELSSON
2
and MARÍA QUINTELA
1
†
1
Population Biology and Conservation Biology, Department of Ecology and Genetics, Evolutionary
Biology Centre, Uppsala University, Norbyvägen 18D, SE 75236 Uppsala, Sweden
2
Molecular Medicine, Department of Medical Sciences, University Hospital, SE 751 85 Uppsala,
Sweden
Received 21 November 2012; revised 10 March 2013; accepted for publication 10 March 2013
Using independently segregating nuclear single nucleotide polymorphisms (SNPs) and mitochondrial control region
sequences, we found an east–west division among sampled willow grouse Lagopus lagopus subspecies. This division
cut across the range of the subspecies with the largest distribution (lagopus) and thus contradicted existing
taxonomic classifications. Russian Lagopus lagopus lagopus tended to cluster with North American willow grouse
partly classified as other subspecies. Scandinavian willow grouse (L. l. lagopus) clustered with red grouse from
Britain and Ireland (Lagopus lagopus scoticus and Lagopus lagopus hibernicus) but substructuring confirmed the
monophyly of the latter. In North America, we could not detect any major genetic divisions apart from two birds
described as alexandrae from the Heceta Island (Alaska) when using mitochondrial sequences. Other samples from
North America were intermingled regardless of whether they were described as muriei, alexandrae or lagopus.A
specimen described as alexandrae was to some extent distinct when analysing the SNP data. The genetic analyses
indicated some concordance between genetics and taxonomy but not complete congruence. This is particularly
evident for mitochondrial DNA network analyses. We suggest that the taxonomy of this species would benefit by
a careful re-examination of the available evidence for subspecies. It appears as if subspecies status is a poor proxy
for assigning evolutionary significant units and management units in this species. © 2013 The Linnean Society
of London, Biological Journal of the Linnean Society, 2013, 110, 77–90.
ADDITIONAL KEYWORDS: alexandrae – evolutionary significant unit – hibernicus – kamschatkensis –
Lagopus lagopus – muriei – phylogeography – scoticus – willow grouse subspecies.
INTRODUCTION
To take accurate conservation actions, it is important
to correctly identify the taxonomic units relevant for
conservation (Moritz, 1994; Crandall et al., 2000;
Ballard & Whitlock, 2004; Beaumont & Balding,
2004). Many conservation actions still rely on taxo-
nomic subspecies classifications based on morphologi-
cal characters. Although these may be valid in many
instances, they should be confirmed by the genetic
data because classifications based purely on morpho-
logical data may be misleading as a result of pheno-
typic plasticity and clinal variation (Storz, 2002;
Relethford, 2004). Subsequent to the introduction of
genetic techniques in conservation, it has been
common practice to use mitochondrial (mt)DNA
sequences to infer species and other taxonomic rela-
tionships. For taxa to classify as an evolutionary
significant unit (ESU), it has been suggested that
they need to be reciprocally monophyletic for mtDNA
alleles (Moritz, 1994). However, maternally inherited
mtDNA has no (or very low) recombination rates and
is thus inherited as a single linkage group (i.e. as one
gene) (Ballard & Whitlock, 2004). It has long been
known that single gene trees may be discordant with
*Corresponding author. E-mail: jacob.hoglund@ebc.uu.se
†Current address: Department of Animal Biology, Plant
Biology and Ecology, Faculty of Science, University A
Coruña, E-15008 A Coruña, Spain.
Biological Journal of the Linnean Society, 2013, 110, 77–90. With 6 figures
© 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110, 77–90 77
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