The Lichenologist 48(3): 209–227 (2016) © British Lichen Society, 2016
doi:10.1017/S0024282915000572
Phylogenetic relationships among reindeer lichens of North America
Sarangi N. P. ATHUKORALA, Raquel PINO-BODAS, Soili STENROOS,
Teuvo AHTI and Michele D. PIERCEY-NORMORE
Abstract: Cladonia is one of the largest lichen-forming ascomycete genera. It was formerly divided into
ten sections, three of which, Crustaceae (Cladina), Tenues, and Impexae, are called the reindeer lichens.
While previous studies have elucidated the relationships between species and sections, they often
examined only one or a few specimens of each species in the analysis. This study examined the
monophyly of selected members of sections Crustaceae, Tenues, and Impexae and their relationships
in the genus Cladonia using the internal transcribed spacer region of the nuclear ribosomal DNA
(ITS rDNA) and the mitochondrial small subunit gene of the mitochondrial ribosomal DNA (mtSSU).
The phylogenetic tree contained four clades, two representing species in section Impexae, one
representing species that belong to sections Crustaceae and Tenues, and one clade with C. arbuscula and
related species. Five of 22 species, C. pycnoclada, C. stellaris, C. evansii, C. ciliata and C. subtenuis,
showed monophyly in the phylogenetic tree; some of these 5 species have been shown previously to be
monophyletic. The thallus branching pattern was interpreted as an important heritable character using
the mtSSU network. Three duplets of paraphyletic species were further examined using ITS rDNA
haplotype networks and AMOVA analysis. The results for the species duplets showed some mixing of
haplotypes but the AMOVA analysis provided support for species separation within the duplets. While
the evidence supports distinct species, further study is needed to conclusively show separate species in
these duplets.
Keywords: AMOVA, Cladina, haplotype network, ITS rDNA, monophyly, mtSSU, phylogeny
Accepted for publication 12 December 2015
Introduction
Cladonia is the largest genus in the lichen-
forming fungal family Cladoniaceae, consist-
ing of c. 459 accepted species (T. Ahti,
August 2015, unpublished data). Based on
morphology and secondary chemistry, Ahti
(2000) divided the neotropical species of
Cladonia into seven taxonomic sections
and three sections were recognized in the
segregate genus Cladina (Ahti 2000). The
division was applicable to most species in the
world. The group called Cladina (known as
reindeer lichens) is most abundant in the
coniferous belt of the Northern Hemisphere
and in the Nothofagus regions in the Southern
Hemisphere, but is also known in sandy
areas of the south-eastern United States
and elsewhere, as well as at high altitudes in
many mountain ranges. While the lack of
competitive ability of lichens with plants is
well known, the reindeer lichens have
adapted better than almost all other lichens
to the terrestrial niches uninhabited by
vascular plants and bryophytes. Some species,
such as Cladonia arbuscula, C. rangiferina,
C. stygia and C. stellaris, are important
components of northern ecosystems where
they provide vast areas of ground cover
(Auclair & Rencz 1982; Shaver & Chapin
1991) and form a major component of the
winter food for caribou and reindeer (Svihus
& Holand 2000; den Herder et al. 2003).
Knowledge of their species status would inform
ecosystem management and maintenance of
biodiversity.
S. N. P. Athukorala and M. D. Piercey-Normore
(corresponding author): Department of Biological
Sciences, University of Manitoba, Winnipeg,
Manitoba, Canada, R3T 2N2. Email: Michele.Piercey-
Normore@umanitoba.ca
R. Pino-Bodas, S. Stenroos and T. Ahti: Botanical
Museum, Finnish Museum of Natural History,
P.O. Box 7, FI-00014 University of Helsinki, Finland.