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Phylogenetic biogeography and taxonomy of disjunctly distributed
bryophytes
1
Jochen HEINRICHS
1
J orn HENTSCHEL
1
Kathrin FELDBERG
1
Andrea BOMBOSCH
2
Harald SCHNEIDER
1
(Department of Systematic Botany, Albrecht von Haller Institute of Plant Sciences, Georg-August-University, D-37073 G ottingen, Germany)
2
(Botany Department, Natural History Museum, London SW7 5BD, UK)
Abstract More than 200 research papers on the molecular phylogeny and phylogenetic biogeography of bryophytes
have been published since the beginning of this millenium. These papers corroborated assumptions of a complex ge-
netic structure of morphologically circumscribed bryophytes, and raised reservations against many morphologically
justied species concepts, especially within the mosses. However, many molecular studies allowed for corrections
and modications of morphological classication schemes. Several studies reported that the phylogenetic structure
of disjunctly distributed bryophyte species reects their geographical ranges rather than morphological disparities.
Molecular data led to new appraisals of distribution ranges and allowed for the reconstruction of refugia and migra-
tion routes. Intercontinental ranges of bryophytes are often caused by dispersal rather than geographical vicariance.
Many distribution patterns of disjunct bryophytes are likely formed by processes such as short distance dispersal,
rare long distance dispersal events, extinction, recolonization and diversication.
Key words bryophytes, cryptic speciation, disjunctions, divergence time estimates, Diversity Arrays Technology,
DNA sequence variation, isozymes, molecular phylogeny.
Bryophytes (liverworts, mosses and hornworts)
comprise the three lineages of land plants with a life
cycle in which the haploid gametophyte is the dom-
inant photosynthetic active generation. In contrast to
other land plants, the sporophyte is unbranched and not
autonomously viable (Schoeld, 1985). Bryophytes dis-
perse frequently both by spores and by propagules that
descend from the gametophyte, or by unspecialized ga-
metophyte fragments with a high potential of regenera-
tion (Correns, 1899). Bryophytes are the progeny of the
rst plants that successfully colonized terrestrial habi-
tats (Qiu, 2008). Their evolution in space and time is
still insufciently known.
In the nineteenth and early twentieth centuries,
bryologists preferred to use a geographical or typo-
logical species concept where species were dened
as largely invariant units. Many species were known
only from type material (e.g., Stephani, 18981925;
Warnstorf, 1911). More recently, authors accepted in-
traspecic morphological variation and lowered numer-
ous local taxa to synonyms of widespread bryophyte
species (Gradstein, 1994; Buck, 1998; Heinrichs, 2002).
Consequently, broad geographical ranges that often span
Received: 11 November 2008 Accepted: 26 February 2009