Nova Hedwigia 86 1—2 1—21 Stuttgart, February 2008

A morpho-molecular classification of the mosses (Bryophyta)

by

Michael Stech1 and Wolfgang Frey2

1
Nationaal Herbarium Nederland, Universiteit Leiden branch, P.O. Box 9514, 2300 RA Leiden,
The Netherlands. E-mail stech@nhn.leidenuniv.nl
2
Institut für Biologie - Systematische Botanik und Pflanzengeographie, Freie Universität Berlin,
Altensteinstr. 6, D-14195 Berlin, Germany

With 1 figure and 2 tables

Stech, M. & W. Frey (2008): A morpho-molecular classification of the mosses (Bryophyta). Nova
Hedwigia 86: 1-21.
Abstract: Molecular relationships within the Bryopsida are inferred from a new approach, i.e., the
combined analysis of solely non-coding plastid markers (trnLUAA intron, atpB-rbcL spacer, psbA-
trnH spacer). The current state of moss systematics is discussed with respect to the impact of this
approach as well as other recently published molecular phylogenies and phylogenomic inferences.
Taking into account the progress that has been made in the last few years by analysing molecular and
non-molecular data, a new synopsis of the suprageneric classification of the Bryophyta is presented.
Taxonomic changes comprise a re-evaluation of the taxonomy of Takakia (including Takakiophytina
stat. nov.), Gigaspermidae subclass. nov., elevation of subfamilies of Dicranaceae to family level
(Amphidiaceae stat. nov., Dicranellaceae stat. nov., Oncophoraceae stat. nov.), Hypodontiaceae fam.
nov. and Helicophyllales ord. nov.
Key words: Bryophyta, suprageneric classification, molecular phylogeny, trnLUAA intron, atpB-rbcL
spacer, psbA-trnH spacer.

Introduction

Mosses form the second largest of the major land plant lineages. They comprise
approximately 12,800 species (Crosby et al. 1999) and exhibit a high morphological
and ecological diversity. Traditionally, the three groups of bryophytes (mosses,
liverworts and hornworts) were treated as classes of the division Bryophyta. Molecular
data, however, revealed that bryophytes are paraphyletic, with liverworts being sister
to all other extant land plants, and mosses sister to a clade of hornworts and vascular
plants (Qiu et al. 2006). As the three bryophyte groups obviously represent independent
lineages in land plant evolution, it is now widely accepted to treat them as separate
divisions, Bryophyta (mosses), Marchantiophyta (liverworts) and Anthocerotophyta

DOI: 10.1127/0029-5035/2008/0086-0001 0029-5035/08/0086-0001 $ 5.25

© 2008 J. Cramer in der Gebrüder Borntraeger
Verlagsbuchhandlung, D-14129 Berlin · D-70176 Stuttgart
1
(hornworts). In contrast to the latter two groups, mosses are scarcely characterised
in terms of morphological-anatomical synapomorphies. At the molecular level,
however, all three bryophyte lineages are well circumscribed (e.g. Qiu et al. 1998,
Stech et al. 2003, Cox et al. 2004, Shaw & Renzaglia 2004, Groth-Malonek et al.
2005, Forrest et al. 2006, Qiu et al. 2006). A general overview of the present state of
bryophyte phylogeny was recently provided by Renzaglia et al. (2007).
In the 19th and 20th century different and often contradictory classification systems
of mosses were published, which either relied more on gametophytic or on sporophytic
characters (summarised, for example, in Buck 2007). Nowadays the phylogenetic
significance of morphological-anatomical characters can be evaluated based on
molecular phylogenies. The most recent classifications of liverworts (Frey & Stech
2005, He-Nygrén et al. 2006), hornworts (Stotler & Crandall-Stotler 2005, Duff et
al. 2007) and mosses (Goffinet & Buck 2004) already integrated molecular and non-
molecular characters to provide a consensus of the current knowledge. However,
several suprageneric relationships within the mosses remained unresolved or
ambiguous at the time of the Goffinet & Buck (2004) classification, such as ordinal
relationships within the Bryidae and the closest relatives of pleurocarpous mosses.
A number of recent attempts shed more light on these problematic relationships
(e.g. Tsubota et al. 2004, Shaw et al. 2005, Stech & Quandt 2006, Bell et al. 2007,
Goffinet et al. 2007, contributions in Newton & Tangney 2007, Quandt et al. 2007,
this paper). Although some clades of the moss phylogeny are still partly contradictory
or do not receive significant statistical support, these studies seem to converge on a
common hypothesis of moss relationship at suprageneric levels, allowing us to
establish an updated and revised classification.
In the present study a molecular phylogeny of the Bryopsida is presented based on a
new approach, i.e., the combined analysis of solely non-coding plastid markers (trnLUAA
intron, atpB-rbcL spacer, psbA-trnH spacer). The current state of moss systematics
is discussed with respect to the impact of this approach as well as recently published
molecular phylogenies and phylogenomic inferences. Finally, we propose a new
synopsis of the suprageneric classification of the Bryophyta, which updates that of
Goffinet & Buck (2004). A detailed description of the taxa down to genus level will
be provided by a forthcoming new edition of Engler's Syllabus of Plant Families,
for which additional taxonomic changes are made here.

Material and methods

TAXON SAMPLING. A molecular data set of 85 species of Bryopsida was assembled by compiling
sequences from own previous analyses and by sequencing of herbarium material. Voucher information
and GenBank accession numbers are summarised in Table 1, together with indications of the newly
sequenced accessions. From 21 species, all three non-coding markers employed here were newly
sequenced. In few cases sequences available in GenBank were used to complete the data set. Andreaea
rupestris Hedw. (Andreaeopsida) and Pogonatum subulatum (Menzies ex Brid.) Brid. (Poly-
trichopsida) were used as outgroup representatives.
DNA EXTRACTION, PCR AND SEQUENCING. DNA was extracted using the mixer mill MM200 (Retsch)
and following the method of Doyle & Doyle (1990). PCR reactions for trnL intron and atpB-rbcL
spacer followed the protocols given in Frey et al. (1999) and Stech (2004), and were performed with

2
 Table 1. Voucher information and GenBank accession numbers of the species used for the three-marker analysis of Bryopsida. * sequences taken from
GenBank; ** species of which all three markers were newly sequenced for the present study.

Family Species Voucher Herb. Acc. no. Acc. no. Acc. no.
trnL atpB-rbcL psbA-trnH

Andreaeaceae Andreaea rupestris Hedw. Stech B970827.3 L AY050278 AY864297 EU163506

Polytrichaceae Pogonatum subulatum (Menzies ex Brid.) Brid. Pfeiffer 98-T207A CHR, Frey AY050277 EU124442 EU163505
Diphysciaceae Diphyscium foliosum (Hedw.) D.Mohr Frey 1-4982 (trnL) Frey EU186537 *DQ397161 *AY312899
Timmiaceae Timmia megapolitana Hedw. (GenBank) - *AF478320 *DQ397137 *AY312923
Timmia austriaca Hedw. ** Stech B970831.4 L EU186543 EU186584 EU163513
Encalyptaceae Encalypta streptocarpa Hedw.** Stech B060412.2 L EU186541 EU186582 EU163511
Encalypta vulgaris Hedw. Stech B950328.3 L EU128000 EU124423 EU163510
Funariaceae Funaria hygrometrica Hedw. Stech B981130.1 L EU186538 EU186579 EU163507
Physcomitrella patens (Hedw.) (sterile culture) - EU186539 EU186580 EU163508
Bruch & Schimp.
Physcomitrium pyriforme (Hedw.) Hampe** Stech B040407.1 L EU186540 EU186581 EU163509
Catoscopiaceae Catoscopium nigritum (Hedw.) Brid. ** Stech B970828.13 L EU186545 EU186592 EU163524
Bryoxiphiaceae Bryoxiphium norvegicum (Brid.) Mitt. Koponen 36664 B AF135101 EU186590 EU163521

3
Grimmiaceae Racomitrium microcarpum (Hedw.) Brid. Stech B970818.1 L AF127186 EU186589 EU163519
Schistidium apocarpum (Hedw.) Stech B970226.2 L AF127185 EU186588 EU163518
Bruch & Schimp.
Ptychomitriaceae Ptychomitrium polyphyllum (Sw.) Stech et al. 04-40 L EU186542 EU186583 EU163512
Bruch & Schimp. **
Archidiaceae Archidium alternifolium (Dicks. ex Hedw.) Mitt. Frahm s.n. BONN AF135114 EU186597 EU163529
Amphidiaceae Amphidium mougeotii (Bruch & Schimp.) Frahm s.n. BONN AF127187 AY159894 EU163520
Schimp.
Dicranaceae Dicranum polysetum Sw. Stech B970518.1 L AF129587 AY159895 EU163523
Dicranellaceae Dicranella cerviculata (Hedw.) Schimp. Stech B970824.1 L AF129597 EU186591 EU163522
Ditrichaceae Pleuridium acuminatum Lindb. Frey 1-4991 Frey EU186546 EU186596 EU163528
Fissidentaceae Fissidens bryoides Hedw. Darmer 13017 BSB AF135105 EU186586 EU163515
Octodiceras fontanum (Bach. Pyl.) Lindb. Haapasaari s.n. L AF135107 EU186585 EU163514
Leucobryaceae Campylopus flexuosus (Hedw.) Brid. Stech B960905.2 L AF129593 AY159919 EU163530
Leucobryum javense (Brid.) Mitt. Menzel et al. 3841 Frey AF442659 AY159900 EU163531
Oncophoraceae Cynodontium polycarpum (Hedw.) Schimp. Stech B 930721.2 L AF129599 EU186595 EU163527
Oncophorus virens (Hedw.) Brid. Stech B960801.1 L AF129598 EU186593 EU163525
Rhabdoweisia crenulata (Mitt.) H.Jameson Frahm s.n. 18.10.97 BONN AF127181 EU186594 EU163526
 Table 1 continued.
Pottiaceae Cinclidotus nigricans (Brid.) Wijk & Margadant Stech B920517.4 L EU186544 EU186587 EU163517
Tortula muralis Hedw. Stech B970226.3 L AF135108 AY159892 EU163516
Splachnaceae Splachnum luteum Hedw. ** Stech B970828.12 L EU186555 EU186613 EU163548
Tayloria mirabilis (Cardot) Broth. ** Frey & Schaumann F, Frey EU186554 EU186612 EU163547
01-36
Tetraplodon mnioides (Sw. ex Hedw.) Stech B910615.4 L EU186556 EU186614 EU163549
Bruch & Schimp. **
Meesiaceae Meesia uliginosa Hedw. ** Stech B970828.5 L EU186552 EU186610 EU163545
Paludella squarrosa (Hedw.) Brid. ** Stech B970820.1 L EU186553 EU186611 EU163546
Pulchrinodaceae Pulchrinodus inflatus (Hook.f. & Wilson) Pfeiffer & Lindlar CHR AF135065 EU186603 EU163538
B.H.Allen 98-Z215D
Leptostomataceae Leptostomum menziesii R.Br. Frey & Schaumann F, Frey EU186550 EU186608 EU163543
01-388
Bryaceae Bryum billardierei Schwägr. Frey & Pfeiffer 98-Mo17 Frey EU186549 EU186607 EU163542
Rhodobryum giganteum (Schwägr.) Paris (GenBank) - *AF023737 *AF546860 *AY312918
Mniaceae Mielichhoferia mielichhoferiana (Funck) Loeske Stech B910615.3 L EU186548 EU186605 EU163540
Mnium hornum Hedw. Frey 1-4985 (trnL) Frey AF363279 *AF546857 *AY312908
Plagiomnium undulatum (Hedw.) T.J.Kop. Stech et al. 04-607 L DQ108967 EU186604 EU163539
Rhizomnium magnifolium (Horik.) T.J.Kop. Stech B970829.6 L DQ108959 EU186606 EU163541

4
Hedwigiaceae Hedwigia ciliata (Hedw.) P.Beauv. (GenBank) - *AF478336 *AJ249038 *AY312905
Rhacocarpaceae Rhacocarpus purpurascens (Brid.) Paris (sterile culture) - AF023724 EU186598 EU163532
Helicophyllaceae Helicophyllum torquatum (Hook.) Brid. Frahm 1507 Frey EU186547 EU186602 EU163537
Bartramiaceae Anacolia laevisphaera (Taylor) Flowers Shevock 17853 MO EU 301607 *AF413557 EU163533
Bartramia stricta Brid. Müller C1552 Müller AF023756 EU186599 EU163534
Conostomum pentastichum (Brid.) Lindb. Frahm s.n. BONN EU 301606 EU186600 EU163535
Philonotis fontana (Hedw.) Brid. Garton 20748 BONN EF107540 EU186601 EU163536
Orthotrichaceae Orthotrichum anomalum Hedw. Stech B970226.1 L AF130314 EU186616 EU163551
Orthotrichum stramineum Hornsch. (sterile culture) - AF127183 EU186618 EU163553
Ulota crispa (Hedw.) Brid. ** Stech B050324.3 L EU186557 EU186617 EU163552
Zygodon baumgartneri Malta Frahm s.n. 4.5.1985 BSB AF127184 EU186615 EU163550
Aulacomniaceae Aulacomnium palustre (Hedw.) Schwägr. ** Stech B030529.4 L EU186551 EU186609 EU163544
Aulacomnium turgidum (Wahlenb.) Schwägr. (GenBank) - *AF023728 *AF413532 *AY312894
Hymenodontopsis mnioides (Hook.) Frey & Schaumann F, Frey EU186563 EU186625 EU163560
N.E.Bell, A.E.Newton & D.Quandt** 01-186a
Orthodontiaceae Hymenodon pilifer Hook.f. & Wilson Frahm 1-14 Frey EU186559 EU186620 EU163555
Orthodontium lineare Schwägr. ** Stech B950616.2 L EU186558 EU186619 EU163554
Leptotheca gaudichaudii Schwägr. Frey & Schaumann F, Frey *AF023750 EU186624 EU163559
01-09
 Table 1 continued.
Rhizogoniaceae Calomnion complanatum Frahm 20-4 Frey EU186560 EU186621 EU163556
(Hook.f. & Wilson) Lindb.
Cryptopodium bartramioides (Hook.) Brid. ** Frey & Schaumann 01-40 F, Frey EU186561 EU186622 EU163557

Rhizogonium distichum (Sw.) Brid. Frahm 2-11 Frey EU186562 EU186623 EU163558
Hypnodendraceae Cyrtopus setosus (Hedw.) Hook.f. Frahm X16-1 Frey EU186569 EU186631 EU163566
Hypnodendron spininervium (Hook.) Pfeiffer 98-T162 CHR, Frey EU186567 EU186629 EU163564
A.Jaeger & Sauerb.
Hypnodendron vitiense Mitt. Streimann B EU186568 EU186630 EU163565
Sciadocladus kerrii (Mitt.) A.Jaeger ex Broth. Pfeiffer T210 CHR, Frey EU186565 EU186627 EU163562
Sciadocladus menziesii (Hook.) Frey & Pfeiffer CHR, Frey EU186566 EU186628 EU163563
Lindb. ex Broth. 98-Mo5
Racopilaceae Racopilum convolutaceum (Müll.Hal.) Zank s.n. Frey EU186564 EU186626 EU163561
Reichardt
Ptychomniaceae Ptychomnion ptychocarpon (Schwägr.) Frey & Schaumann F, Frey EU186570 EU186632 EU163567
Mitt. ** 01-266a
Hookeriaceae Hookeria lucens (Hedw.) Sm. Stech B880404.8 L AF152380 EU186635 EU163570
Lepidopilum virens Cardot** Stech et al. 04-109 L EU186571 EU186633 EU163568
Tetrastichium fontanum (Mitt.) Cardot** Stech et al. 04-124 L EU186572 EU186634 EU163569

5
Hypopterygiaceae Arbusculohypopterygium arbuscula (Brid.) Frey 95-17 CHR, Frey AF134637 EU186637 EU163573
M.Stech, T.Pfeiff. & W.Frey
Catharomnion ciliatum (Hedw.) Wilson Beever 97-69 CHR, Frey EU186573 EU186636 EU163572
Hypopterygium discolor Mitt. Streimann 52819 Frey ex L AF363282 AY864293 EU163575
Hypopterygium tamarisci (Sw.) Pfeiffer & Sipman B DQ077169 AY864294 EU163574
Brid. ex Müll.Hal. 99-69
Lopidium concinnum (Hook.) Wilson Frey 95-62 Frey AF033231 AY864295 EU163571
Fontinalaceae Fontinalis antipyretica Hedw. ** Stech et al. 04-14 L EU186574 EU186638 EU163576
Amblystegiaceae Cratoneuron filicinum (Hedw.) Spruce Stech B900615.1 L AF129586 EU186645 EU163583
Brachytheciaceae Platyhypnidium riparioides (Hedw.) Dixon** Stech et al. 04-237 L EU186577 EU186644 EU163582
Hypnaceae Hypnum andoi A.J.E.Sm. ** Stech et al. 04-454 L EU186578 EU186647 EU163585
Rhytidiaceae Rhytidium rugosum (Ehrh. ex Hedw.) Kindb. Stech B930910.1 L AF264046 EU186639 EU163577
Cryphaeaceae Cryphaea tasmanica Mitt. Frahm 31-1 Frey EU186576 EU186643 EU163581
Leucodontaceae Antitrichia curtipendula (Timm ex Hedw.) Brid. Stech B940717.3 L AY010286 EU186641 EU163579
Neckeraceae Thamnobryum pandum (Hook.f. & Wilson) Frey & Frey 94-117 Frey AY010297 EU186642 EU163580
I.G.Stone & G.A.M.Scott
Plagiotheciaceae Plagiothecium undulatum (Hedw.) Schimp. Stech B881003.2 L AF264045 EU186646 EU163584
Catagoniaceae Catagonium nitens (Brid.) Cardot Frey & Schaumann F, Frey EU186575 EU186640 EU163578
01-187a
primers CM / DM (slightly modified after Taberlet et al. 1991, see Frey et al. 1999) and atpB-1/rbcL-1
(Chiang et al. 1998), respectively. For amplification of the psbA-trnH spacer, primers psbAF
(5'-GTT ATG CAT GAA CGT AAT GCT C-3', Sang et al. 1997) and trnHR-2 (5'-CGC GCA TGG
TGG ATT CAC AAT CC-3', slightly modified after Sang et al. 1997) were used with a PCR
protocol of 5 min 94°C, 35 cycles (30 sec 94°C, 1 min 48°C, 1 min 40 sec 72°C), 7 min 72°C. PCR
products were purified using DNA purification kits (Qiagen, Macherey-Nagel). Sequencing was
performed as described in Stech (2004), using the amplification primers.
ALIGNMENT AND PHYLOGENETIC ANALYSES. DNA sequences were manually aligned in PhyDE v0.992
(Müller et al. 2005). Following Quandt et al. (2003) and Quandt & Stech (2005), inversions in
putative hairpin secondary structures were positionally separated in the alignment; their presence or
absence was not coded for the phylogenetic analyses. Phylogenetic reconstructions under the maximum
parsimony criterion were performed using winPAUP 4.0b10 (Swofford 2002) in combination with
PRAP (Müller 2004), as implemented in SeqState (Müller 2005). PRAP generates command files
for PAUP that allow parsimony ratchet searches as designed by Nixon (1999). Heuristic searches
under parsimony were performed with the following options: all characters equally weighted and
unordered, TBR branch swapping, collapse branches if maximum branch length is zero, MulTrees
option in effect, gaps treated as missing data. Initially, a heuristic search with 1000 addition sequence
replicates was performed. For the parsimony ratchet, ten random addition cycles of 200 ratchet
iterations each were used, with TBR branch swapping on a randomly re-weighted data set (25% of
the positions). Heuristic bootstrap searches were performed with 1,000 replicates with 10 addition
sequence replicates per bootstrap replicate.

Results
In the present data set of non-coding plastid markers, sequence lengths ranged from
243 to 368 nucleotides (nt) in the trnL intron and from 445 to 615 nt in the atpB-
rbcL spacer. The psbA-trnH spacer was considerably shorter than the other two
markers and ranged from 103 to 186 nt. The combined alignment of sequences from
85 ingroup and two outgroup taxa comprised 2430 positions (trnL intron 668, atpB-
rbcL spacer 1366, psbA-trnH spacer 396 positions). The most variable regions with
ambiguous alignment (781 positions) were excluded from phylogenetic analysis. Of
the 1649 included positions, 628 (38.1%) were variable, and 455 of the variable
positions (72.5%, or 27.6% of the total number of positions) were parsimony-
informative. The trnL intron comprised 124, the atpB-rbcL spacer 286 and the psbA-
trnH spacer 45 of these included parsimony-informative positions.
The shortest trees found in the heuristic maximum parsimony analysis of the combined
data set had a length of 2433 steps. By applying the parsimony ratchet, 129 trees
were retained that were one step shorter (length 2432, CI = 0.419, RI = 0.624). The
strict consensus tree of the ratchet analysis is shown in Fig. 1. The ingroup (Bryopsida)
is supported with 100% bootstrap support (BS), Diphysciidae are sister to the
remaining Bryopsida with 54% BS. Encalyptidae, Funariidae, Timmiidae and
Dicranidae all receive > 95% BS, while Bryidae are slightly less supported (89%
BS). Relationships between these major lineages are resolved but unsupported. At
ordinal level within the Bryidae, a lower resolution is observed. Splachnales,
Orthotrichales, Aulacomniales, Hypnodendrales, Hookeriales and Hypnales are
resolved as monophyletic with 67-100% BS, whereas Hedwigiales, Bartramiales,
Bryales, Orthodontiales and Rhizogoniales are not. A closer relationship of
Orthotrichales with the pleurocarpous mosses s.l. than with the more basal orders
Hedwigiales, Bartramiales, Splachnales and Bryales is indicated.

6
 Discussion

Classification of mosses - recent progress and the current state

For a long time, mosses were subdivided into three main lineages, the subclasses
Sphagnidae, Andreaeidae and Bryidae (e.g. Fleischer 1920, Vitt 1984, Frahm &
Frey 1987), the latter with a number of (super-)orders (e.g. Frey 1977). However,
molecular data confirmed that mosses comprise a number of isolated basal lineages,
represented by morphologically and ecologically distinct extant taxa such as Takakia,
Sphagnum, Andreaea, Andreaeobryum, Oedipodium, Tetraphis, the Polytrichaceae
and Buxbaumia, which probably have long independent phylogenetic histories (cf.
Newton et al. 2007). As a result of molecular phylogenies and a re-evaluation of
morphological-anatomical characters, the classification of mosses has changed
considerably in the last few years, with the main lineages nowadays treated at
subdivision, class and subclass levels within division Bryophyta (e.g. Doweld 2001,
Ochyra et al. 2003, Goffinet & Buck 2004, this study).
Takakia and Sphagnum are the earliest diverging moss taxa in molecular phylogenies
(e.g. Newton et al. 2000, Beckert et al. 2001, Cox et al. 2004, Qiu et al. 2006,
Newton et al. 2007). That Takakia is a moss and not a liverwort was confirmed by
analysis of DNA sequences and phylogenomic characters such as the distribution of
mitochondrial introns (e.g. Beckert et al. 2001). Its taxonomic status, however, still
remains to be clarified. On the one hand, most molecular studies resolve Takakia
and Sphagnum as sister genera, although only partly with high statistical support.
These molecular topologies do not support the recognition of division Takakiophyta
as proposed by Crandall-Stotler (1986). One the other hand, no morphological
synapomorphies unite Takakia with Sphagnum, and significant molecular differences
separating Takakia from all other mosses were also observed. For example, a region
in the plastid ITS3 is similar to that of other land plants but absent in mosses
(Samigullin et al. 2002), and a transition in the trnLUAA gene changing the anticodon
to CAA was otherwise only found in leptosporangiate ferns and Ginkgo (Quandt et
al. 2004). Thus, “resolution of Sphagnum - Takakia clade may be an artifact” (Shaw
& Renzaglia 2004). Also Goffinet & Buck (2004) did not unite Takakia and Sphagnum
under a formal taxonomic rank but placed them into different informal superclasses.
In our opinion Takakia should be treated at the same level as Sphagnophytina and
Bryophytina (cf. Doweld 2001), i.e., as a subdivision of Bryophyta. However, more
fundamental nomenclatural changes are necessary, as the family name Takakiaceae
was not validated in the original publication by Hattori & Inoue (1958) (article
41.1. of ICBN, McNeill et al. 2006), and consequently, all higher taxa erected later
are invalid. The corrected taxonomic treatment of Takakia is provided here, with
sporophytic characters included in the latin diagnosis of Takakiaceae (see taxonomy).
Within Bryophytina, we recognise four morphologically and molecularly isolated
basal lineages as separated from the arthrodontous mosses (Bryopsida): Andreaeopsida
(with linear capsule dehiscence), the operculate but eperistomate Oedipodiopsida as
well as Polytrichopsida and Tetraphidopsida with nematodontous peristomes. A fifth
class, Andreaeobryopsida, was erected by Buck & Goffinet (2000) but re-included
in Andreaeopsida as a subclass by Ochyra et al. (2003). The systematic status of

7
Andreaeobryum macrosporum Steere & B.M.Murray and possible relationships with
Andreaea and Takakia were discussed by Murray (1988) and summarised, including
molecular results, by Shaw & Renzaglia (2004).
On the next lower level, a number of morphologically and molecularly divergent
lineages are nowadays treated as subclasses of Bryopsida: Buxbaumiidae and
Diphysciidae are generally branching off first in molecular topologies (e.g. Beckert
et al. 2001, Magombo 2003, Cox et al. 2004), followed by Timmiidae, Encalyptidae
and Funariidae, as well as Dicranidae (haplolepideous mosses) sister to Bryidae s.l.
These subclasses (except Buxbaumiidae, not included) are also well recognizable in
the present analysis of non-coding plastid sequences (Fig. 1).
The morphologically very different Encalyptidae and Funariidae are revealed as
sister groups, although mostly without high support, in several molecular phylogenies
(e.g. Goffinet et al. 2001, Cox et al. 2004, Hedderson et al. 2004, Tsubota et al.
2003, 2004), and also in the present study (Fig. 1). Goffinet et al. (2007) confirmed
this relationship by phylogenomic evidence, namely a large (71 kb) inversion in the
chloroplast genome of Funariaceae, Disceliaceae and Encalyptaceae, but not in
Gigaspermaceae. Consequently, the latter family was separated at ordinal level from
the two remaining families of Funariales by Goffinet et al. (2007). Gigaspermales,
Funariales and Encalyptales may share a common ancestor according to some, but
not all, molecular phylogenies (e.g. Goffinet et al. 2001, 2007, Hedderson et al.
2004, but not in Tsubota et al. 2004). Although ontogenetic data from peristome
development may support their common ancestry (Goffinet et al. 2007), unambiguous
morphological synapomorphies are lacking. We therefore propose to treat
Gigaspermaceae at subclass level within the Bryopsida (see taxonomy).
The haplolepideous mosses (Dicranidae) are clearly supported as a monophyletic
lineage, which obviously evolved from a diplolepideous ancestor according to their
position in the molecular trees (e.g. Newton et al. 2000, Tsubota et al. 2003, Cox et
al. 2004, Goffinet & Buck 2004). Molecular studies suitable for inferring ordinal
and family level relationships within Dicranidae either focused on the haplolepideous
mosses (e.g. Stech 1999a,b, 2004, LaFarge et al. 2000, 2002, Tsubota et al. 2003,
Hedderson et al. 2004) or included more than 15 haplolepideous taxa in analyses of
a broader range of mosses (e.g. Goffinet et al. 2001, Tsubota et al. 2004, this study).
Major lineages supported by these analyses are, for example, Grimmiales
(Grimmiaceae, Ptychomitriaceae and Seligeriaceae), Leucobryaceae (incl. former
Dicranaceae-Campylopoideae and Paraleucobryoideae p.p.), Dicranaceae s.str.
(Dicranoideae plus former Dicnemonaceae and Paraleucobryoideae p.p.), Pottiaceae
(incl. Cinclidotaceae and Splachnobryum) and Calymperaceae (probably excl.
Octoblepharum). Furthermore, several genera and families belong to Dicranidae
whose systematic positions have been controversial (e.g. Stech et al. 1999b, La
Farge et al. 2002, Hedderson et al. 2004, O'Brien 2007, Quandt et al. 2007, this
study): Amphidium, Archidium, Catoscopium, Drummondia, Ephemeraceae,
Erpodiaceae, Mittenia (cf. O'Brien 2007, sequences in some earlier analyses were
based on misdentified specimens), Rhachitheciaceae, Schistostega, Splachnobryum
and Wardia. On the contrary, several problems remain, such as the circumscription
of Ditrichaceae and relationships of the major haplolepideous lineages in general.

8
Fig. 1. Strict consensus tree of 129 most parsimonious trees inferred from combined trnLUAA intron,
atpB-rbcL spacer, and psbA-trnH spacer sequences of 85 taxa of Bryopsida as well as Andreaea
rupestris (Andreaeopsida) and Pogonatum subulatum (Polytrichopsida) as outgroup representatives.
Bootstrap support values >50% are depicted above the branches.

9
Consequently, resolving the phylogeny of Dicranidae with confidence is one of the
major future challenges in moss systematics.
As mentioned above, Dicranaceae are monophyletic in a narrow circumscription
according to molecular data. In contrast to former Campylopoideae and
Paraleucobryoideae, the status of Rhabdoweisioideae (= Oncophoroideae, Ochyra
2002) and Dicranelloideae needs further consideration. Stech (1999a,b) suggested
to recognise Rhabdoweisioideae/Rhabdoweisiaceae in a broad sense, including
Dicranoweisia, Amphidium and Aongstroemia. Subsequent molecular analyses
confirmed a close relationship of the genera Arctoa, Cynodontium, Dicranoweisia,
Kiaeria, Oncophorus, Oreoweisia, Oreas, Rhabdoweisia and Symblepharis, although
without significant support (La Farge et al. 2002, Tsubota et al. 2003, Hedderson et
al. 2004). With respect to the molecular evidence available so far, we propose to
elevate Oncophoroideae to family level to accomodate these genera and perhaps
Glyphomitrium (cf. Tsubota et al. 2003), whereas we consider Aongstroemia,
Dichodontium and Diobelonella to belong to Aongstroemiaceae (cf. Stech 1999a).
Amphidium is neither closely related to Dicranaceae s.str. nor to Oncophoroideae
(La Farge et al. 2002, Tsubota et al. 2003, Hedderson et al. 2004, Stech 2004, this
study), and should be treated at family level rather than placed in Dicranaceae as
subfam. Amphidioideae (Ochyra et al. 2003). The circumscription of Dicranelloideae
is more difficult to evaluate, as few representatives have been included in molecular
analyses. However, Dicranella heteromalla (Hedw.) Schimp. as well as the genera
Campylopodium and Microcampylopus are clearly separated from Dicranaceae s.str.
(Stech 1999a, 2004, La Farge et al. 2002, Tsubota et al. 2003, Hedderson et al.
2004), and consequently, the subfamily deserves family status as well. Finally,
molecular data showed that the genus Hypodontium does not belong to Pottiaceae
(Hedderson et al. 2004, Tsubota et al. 2004). Taxonomic conclusions have not been
drawn from these results, which is made up here (see taxonomy).
All mosses with diplolepideous-alternate peristomes together with Splachnales with
a diplolepideous-opposite (Funaria-type) peristome form a monophyletic group in
molecular analyses (e.g. Newton et al. 2000, Beckert et al. 2001, Goffinet et al.
2001, Cox et al. 2004, Tsubota et al. 2004, references in Table 2). Different
classifications of this by far most species-rich group of mosses were proposed (e.g.
Vitt et al. 1998, Ochyra et al. 2003, Goffinet & Buck 2004). However, at least some
of the recognised subclasses and superorders are paraphyletic according to the
molecular topologies, and assigning formal ranks to all evolutionarily significant
monophyletic entities is problematic if the main taxonomic framework of Bryophyta
and Bryopsida should be maintained (cf. discussion in Bell et al. 2007). Therefore,
in the present synopsis only one subclass, Bryidae, is recognised for all diplolepideous-
alternate mosses and Splachnales. This broadly defined Bryidae is subdivided into
13 orders, most of them resolved as monophyletic in the majority of the phylogenetic
analyses summarised in Table 2, although with different levels of statistical support
in maximum parsimony analyses. To define monophyletic lineages at supraordinal
level, informal node-based names may be chosen according to Bell et al. (2007). For
example, the pleurocarpous mosses (pleurocarpy sensu Bell & Newton 2007) and
their closest acrocarpous relatives (Orthodontiales to Hypnales) were already

10
Table 2. Taxon sampling and bootstrap support (BS) for Dicranidae, Bryidae, and orders within the
Bryidae in recent phylogenetic analyses (MP only) based on combined molecular markers. Markers
are given in square brackets (non-coding regions in bold), numbers of included taxa in brackets. ++
respective clade present with ≥ 80% BS, + clade present with < 80% BS, - clade not present, * no or
only one representative included.

Bell & Newton O'Brien Quandt et al. Bell et al. this study
(2004) (2007) (2007) (2007)
[rbcL, rps4, [atpB, atpB- [rbcL, rps4, [rbcL, rps4, [atpB-rbcL,
nad5] rbcL, rbcL, trnL-F, nad5] trnL-F, nad5] psbA-trnH,
rps4, trnL-F] trnL]

Dicranidae + (2) + (4) ++ (3) * (1) ++ (19)

Bryidae ++ (57) + (53) ++ (52) ++ (98) ++ (58)
Hedwigiales ++ (2) * (1) ++ (2) ++ (2) - (3)
Bartramiales ++ (2) ++ (4) ++ (4) ++ (3) - (4)
Splachnales * (0) ++ (2) ++ (3) ++ (2) ++ (5)
Bryales - (4) - (5) ++ (18) + (17) - (8)
Orthotrichales * (1) ++ (3) ++ (3) ++ (2) ++ (4)
Orthodontiales + (4) + (4) + (3) + (6) - (3)
Aulacomniales + (6) + (7) ++ (2) + (7) + (3)
Rhizogoniales ++ (13) + (8) ++ (5) ++ (14) - (3)
Hypnodendrales ++ (13) + (7) ++ (4) ++ (29) ++ (6)
Ptychomniales ++ (3) ++ (3) * (1) ++ (4) * (1)
Hookeriales - (4) + (2) * (1) + (2) + (8)
Hypnales - (5) - (7) ++ (5) ++ (10) + (10)

summarised under the informal node based name 'pleurcarpids' (Bell et al. 2007),
and can be distinguished from a basal grade of acrocarpous or cladocarpous lineages
(Hedwigiales to Orthotrichales). Within the latter, monophyly of Bartramiales,
Splachnales and Orthotrichales is well-supported in all analyses including two or
more taxa, except Bartramiales in the present data set (cf. Fig. 1). In contrast, Bryales
receive significant support less frequently (but see Bell et al. 2007, Quandt et al.
2007). Hedwigiales are well supported if only Hedwigia (Hedwigiaceae) and
Rhacocarpus (Rhacocarpaceae) are included, whereas the monospecific Helicophyllum
(Helicophyllaceae) is not closely related to these taxa but shows affinities with
Bartramiales (Goffinet et al. 2001, this study). With respect to the different
morphologies of Helicophyllaceae and Bartramiaceae, however, the former should
be separated at ordinal level (see taxonomy).
Clarifying relationships of the Orthotrichales is still a difficult issue. Analyses of
single chloroplast markers or parsimony analyses of combined markers from different
genomes mostly indicated a close relationship either with Splachnales (e.g. Cox et
al. 2000, Newton et al. 2000, Tsubota et al. 2004, Quandt et al. 2007) or with the
pleurocarpids (Goffinet et al. 2001, this study). Although none of these alternatives
is well-supported, the latter seems more likely with regard to the moderately to
maximally supported Orthotrichales - pleurocarpid sister group relationship in all
analyses of mitochondrial data and in likelihood analyses of combined plastid/
mitochondrial markers (Beckert et al. 2001, Quandt et al. 2007).

11
The molecular polyphyly of Rhizogoniaceae (e.g. Bell & Newton 2004) was solved
by the transfer of rhizogonioid genera into broadly defined Aulacomniaceae and
Orthodontiaceae (Bell et al. 2007), which now include both acrocarpous and
pleurocarpous taxa. These systematic changes are also comprehensible by non-
molecular characters, such as morphological traits shared by Mesochaete and
Aulacomnium heterostichum (Hedw.) Bruch & Schimp., or the variously reduced
peristomes of Orthodontiaceae p.p. in contrast to the generally fully developed Bryum-
type peristomes of rhizogonioid and hypnodendroid mosses (Bell et al. 2007). The
position of the solely pleurocarpous Hypnodendrales sister to the former Hypnidae
led to the recognition of the informal group 'core pleurocarps' (Bell et al. 2007).
The former Hypnidae (Ptychomniales, Hookeriales and Hypnales, summarised as
'homocostate pleurocarps') are monophyletic in most molecular analyses (e.g. Goffinet
et al. 2001, Tsubota et al. 2004, Bell et al. 2007), with Ptychomniales being well-
supported (Pedersen & Newton 2007, Table 2). Clarifying relationships between
Hookeriales and Hypnales and within the latter is still a challenge, but recent molecular
studies seem to support a sister group relationship of both orders (Bell et al. 2007,
present study).

Phylogenetic utility of non-coding plastid markers

Non-coding plastid markers that comprise both variable and conserved regions,
such as the trnLUAA intron, have repeatedly been shown to possess potential for
resolving relationships at high taxonomic levels (e.g., Borsch et al. 2003, Quandt
et al. 2004). Both the trnL intron and atpB-rbcL spacer were characterised in
detail in mosses with respect to their structure and molecular evolution (Quandt &
Stech 2005, Stech & Quandt 2006). They include conservative regions that can be
unambiguosly aligned, and even parts of the more variable regions can be used if
repeats, hairpin loops etc. are correctly identified and adequately treated in the
phylogenetic analyses (e.g. Quandt et al. 2003, Quandt & Stech 2005). Using the
atpB-rbcL spacer alone (Stech & Quandt 2006), relationships of the major moss
lineages were largely consistent with the “total evidence” molecular tree of Goffinet
& Buck (2004). However, in addition to general limitations that result from single
marker analyses, the use of non-coding markers may pose further problems, such
as a high amount of homoplasy resulting in low resolution or low statistical support
for many clades. Therefore, several recent molecular studies of relationships within
Bryidae have employed both coding and non-coding markers (Table 2).
Nevertheless, the usefulness of analyses of combined non-coding plastid markers
cannot be ruled out. In fact, Fig. 1 and Table 2 indicate a phylogenetic signal in
the present data set, as many clades are recovered that are usually also resolved
in analyses of coding or combined coding and non-coding markers. Especially
with regard to statistical support, however, the present analysis is inferior to
those of other recent data sets, especially Bell et al. (2007) and Quandt et al.
(2007). In addition, the non-coding markers used here are of different suitability,
and even the two intergenic spacers differ considerably from each other. The
psbA-trnH spacer contributes much less to the tree's resolution than the atpB-
rbcL spacer, first because it is much shorter, and second because relatively large
parts are attributed to hairpin loops or otherwise hypervariable regions. These

12
results indicate that future studies should evaluate the molecular evolution and
phylogenetic potential of new markers, both coding and con-coding, and choose
the most suitable ones in a combined analysis to resolve the remaining uncertainties
in moss phylogeny.

Taxonomy

Takakiaceae Stech & W.Frey, stat. nov.

Plantae virides, nusquam rhizophorae, sine protonemate, e caudicis rhizomatis erectae. Caudices
rhizomati et axes geotropi radiformes frequenter intercalariter ramificantes, sine phyllidiis, cum
papillis mucilaginis rostratis. Caules phyllidiiferi saepe furcati. Phyllidia sine lamina et costa
distincta, profunde 2-4(-5)-lobata, pluristratosa, tristicha sed torsione spiraliter disposita. Filum
centralium in ambabus generationibus (axibus gametophyti et setis) adest. Plantae dioicae. Antheridia
et archegonia nuda. Sporophytum terminale, solitarium. Seta persistens. Capsula tortilis ad
maturitatem, schizocarpa, secus unam rimam spiralem longitudinalem dehiscens. Stomata et
operculum absunt, peristomium nullum. Calyptra mitriformis.
TYPUS: Takakia S.Hatt. & Inoue, J. Hattori Bot. Lab. 19: 137. 1958.
Takakiaceae S.Hatt. & Inoue, J. Hattori Bot. Lab. 19: 137. 1958, nom. inval. (Art. 41.1.).

Takakiales Stech & W.Frey, stat. nov.

Based on: Takakiaceae Stech & W.Frey, Nova Hedwigia 86: 13. 2008.
TYPUS: Takakia S.Hatt. & Inoue, J. Hattori Bot. Lab. 19: 137. 1958.
Takakiales S.Hatt. & Inoue, J. Hattori Bot. Lab. 19: 137. 1958, nom. inval.
Takakiales S.Hatt. & Inoue ex R.M.Schust., J. Hattori Bot. Lab. 26: 224: 1963, nom. inval. (cum
descr. latin.).

Takakiopsida Stech & W.Frey, stat.nov.

Based on: Takakiaceae Stech & W.Frey, Nova Hedwigia 86: 13. 2008.
TYPUS: Takakia S.Hatt. & Inoue, J. Hattori Bot. Lab. 19: 137. 1958.
Takakiopsida Goffinet & W.R.Buck, Monogr. Syst. Bot. Missouri Bot. Gard. 98: 232. 2004, nom.
inval.
Takakiopsida Jia et al., Acta Phytotax. Sin. 41: 350, 351. 2003, nom. nud.
Takakiopsida Wang & Wu, Fl. Bryophyt. Sin. 8: 447. 2004, nom. inval. (cum descr. latin.).

Takakiophytina Stech & W.Frey, stat. nov.

Based on: Takakiaceae Stech & W.Frey, Nova Hedwigia 86: 13. 2008.
TYPUS: Takakia S.Hatt. & Inoue, J.Hattori Bot. Lab. 19: 137. 1958.

Gigaspermidae Stech & W.Frey, subclass. nov.

Plantae acrocarpae, terrestres, stoloniferae ramis brevibus erectis. Folia unicostata cellulis laxis
laevibus. Capsula plerumque valde rugosa, gymnostomatica vel cleistocarpa, stomata cum duabus
cellulis sociis. Ab Funariidis et Encalyptidis charactere phylogenomico (71-kb inversio abest)
separatae.

13
TYPUS: Gigaspermum Lindb., Öfversigt af Förhandlingar: Kongl. Svenska Vetenskaps-Akademien
21: 599. 1865.

Amphidiaceae Stech, stat. nov.

Based on: Dicranaceae subfam. Amphidioideae Ochyra in Ochyra, Zarnowiec & Bednarek-Ochyra,
Cens. Cat. Polish Mosses: 110. 2003.
TYPUS: Amphidium Schimp., Coroll. Bryol. Eur.: 39. 1856.
Amphidiaceae A.J.E.Sm., Moss Fl. Britain Ireland, 2nd ed.: 657. 2004, nom. inval. descr. angl.

Dicranellaceae Stech, stat. nov.

Based on: Dicranaceae subfam. Dicranelloideae Lindb., Utkast Eur. Bladmoss.: 33. 1878
[‘Underfamilje Dicranelleae’].
TYPUS: Dicranella (Müll.Hal.) Schimp., Coroll. Bryol. Eur.: 13. 1856.

Oncophoraceae Stech, stat. nov.

Based on: Dicranaceae subfam. Oncophoroideae Lindb., Utkast Eur. Bladmoss.: 34. 1878
[‘Underfamilje Oncophoreae’].
TYPUS: Oncophorus (Brid.) Brid., Bryol. Univ. 1: 389. 1826.

Hypodontiaceae Stech, fam. nov.

Plantae robustae, characteribus morphologicis Pottiaceis et Calymperaceis similes sed
characteribus molecularicis separatae. Bracteae perichaetiales internae e basi vaginantes subulatae
vel aristatae. Capsula aspectu succulenta, cellulae exothecii incrassatae, annulus magnopere
absens. Peristomii dentes breves, triangulati, incurvati, aurantiaci, leves vel infirme verrucosi.
Calyptra cucullata.
TYPUS: Hypodontium Müll.Hal., Hedwigia 38: 96. 1899.

Helicophyllales Stech & W.Frey, ord. nov.

Plantae acrocarpae, prostratae, tomentosae. Folia dimorpha, in duas series laterales et unam seriem
ventralem divisa. Costa percurrens. Cellulae hexagonae ad subquadratae, papillosae. Capsula immersa,
cupulata ad oblonga, infra orificium abrupte incurvata, stomata destituta. Peristomium nullum. Sporae
isomorphae, heteropolares, granulatae.
TYPUS: Helicophyllum Brid., Bryol. Univ. 2: 771. 1827.

Synopsis of the suprageneric classification of Bryophyta

Author citations are in accordance with Art. 49.2. of the Vienna Code (McNeill et al. 2006)

Subdivision Takakiophytina Stech & W.Frey

Class Takakiopsida Stech & W.Frey
Order Takakiales Stech & W.Frey
Takakiaceae Stech & W.Frey
Subdivision Sphagnophytina Doweld
Class Sphagnopsida Schimp.
Order Sphagnales Limpr.

14
 Sphagnaceae Dumort.
Order Ambuchananiales Seppelt & H.A.Crum
Ambuchananiaceae Seppelt & H.A.Crum
Subdivision Bryophytina Engler
Class Andreaeopsida J.H.Schaffn.
Subclass Andreaeidae Engl.
Order Andreaeales Limpr.
Andreaeaceae Dumort.
Subclass Andreaeobryidae Ochyra
Order Andreaeobryales B.M.Murray
Andreaeobryaceae Steere & B.M.Murray
Class Oedipodiopsida Goffinet & W.R.Buck
Order Oedipodiales Goffinet & W.R.Buck
Oedipodiaceae Schimp.
Class Tetraphidopsida Goffinet & W.R.Buck
Order Tetraphidales M.Fleisch.
Tetraphidaceae Schimp.
Class Polytrichopsida Doweld
Order Polytrichales M.Fleisch.
Polytrichaceae Schwägr.
Class Bryopsida Pax
Subclass Buxbaumiidae Doweld
Order Buxbaumiales M.Fleisch.
Buxbaumiaceae Schimp.
Subclass Diphysciidae Ochyra
Order Diphysciales M.Fleisch.
Diphysciaceae M.Fleisch.
Subclass Timmiidae Ochyra
Order Timmiales Ochyra
Timmiaceae Schimp.
Subclass Encalyptidae Ochyra, Żarnowiec & Bednarek-Ochyra
Order Encalyptales Dixon
Encalyptaceae Schimp.
Subclass Funariidae Ochyra
Order Funariales M.Fleisch.
Funariaceae Schwägr., Disceliaceae Schimp.
Subclass Gigaspermidae Stech & W.Frey
Order Gigaspermales Goffinet, Wickett, O.Werner, Ros, A.J.Shaw & C.J.Cox
Gigaspermaceae Lindb.
Subclass Dicranidae Doweld
Order Catoscopiales Ignatov & Ignatova
Catoscopiaceae Broth.
Order Scouleriales Goffinet & W.R.Buck
Drummondiaceae Goffinet, Scouleriaceae S.P.Churchill
Order Bryoxiphiales H.A.Crum & L.E.Anderson
Bryoxiphiaceae Besch.

15
 Order Grimmiales M.Fleisch.
Grimmiaceae Arn., Ptychomitriaceae Schimp., Seligeriaceae Schimp.
Order Archidiales Limpr.
Archidiaceae Schimp.
Order Mitteniales Shaw
Mitteniaceae Broth.
Order Dicranales H.Philib. ex M.Fleisch.
Amphidiaceae Stech, Aongstroemiaceae De Not., Bruchiaceae
Schimp., Calymperaceae Kindb., Dicranaceae Schimp., Dicranel-
laceae Stech, Ditrichaceae Limpr., Erpodiaceae Broth., Eustichiaceae
Broth., Fissidentaceae Schimp., Leucobryaceae Schimp., Onco-
phoraceae Stech, Rhachitheciaceae H.Rob., Schistostegaceae Schimp.,
Viridivelleraceae I.G.Stone, Wardiaceae Welch
Order Pottiales M.Fleisch.
Ephemeraceae Schimp., Hypodontiaceae Stech, Pleurophascaceae
Broth., Pottiaceae Schimp., Serpotortellaceae W.D.Reese & R.H.Zander
Subclass Bryidae Engl.
Order Hedwigiales Ochyra
Hedwigiaceae Schimp., Rhacocarpaceae Kindb.
Order Helicophyllales Stech & W.Frey
Helicophyllaceae Broth.
Order Bartramiales D.Quandt, N.E.Bell & Stech
Bartramiaceae Schwägr.
Order Splachnales Ochyra
Meesiaceae Schimp., Splachnaceae Grev. & Arn.
Order Bryales Limpr.
Bryaceae Schwägr., Leptostomataceae Schwägr., Mniaceae Schwägr.,
Phyllodrepaniaceae Crosby, Pseudoditrichaceae Steere & Z.Iwats.,
Pulchrinodaceae D.Quandt, N.E.Bell & Stech
Order Orthotrichales Dixon
Orthotrichaceae Arn.
Order Orthodontiales N.E.Bell, A.E.Newton & D.Quandt
Orthodontiaceae Goffinet
Order Aulacomniales N.E.Bell, A.E.Newton & D.Quandt
Aulacomniaceae Schimp.
Order Rhizogoniales Goffinet & W.R.Buck
Rhizogoniaceae Broth.
Order Hypnodendrales N.E.Bell, A.E.Newton & D.Quandt
Braithwaiteaceae N.E.Bell, A.E.Newton & D.Quandt, Hypnodendra-
ceae Broth., Pterobryellaceae W.R.Buck & Vitt, Racopilaceae Kindb.
Order Ptychomniales W.R.Buck, C.J.Cox, A.J.Shaw & Goffinet
Ptychomniaceae M.Fleisch.
Order Hookeriales M.Fleisch.
Daltoniaceae Schimp., Hookeriaceae Schimp., Hypopterygiaceae
Mitt., Leucomiaceae Broth., Pilotrichaceae Kindb., Saulomataceae
W.R.Buck, C.J.Cox, A.J.Shaw & Goffinet, Schimperobryaceae W.R.

16
 Buck, C.J.Cox, A.J.Shaw & Goffinet
Order Hypnales W.R.Buck & Vitt
Amblystegiaceae G.Roth, Anomodontaceae Kindb., Brachytheciaceae
Schimp., Calliergonaceae Vanderpoorten, Hedenäs, C.J.Cox & A.J.
Shaw, Catagoniaceae W.R.Buck & Ireland, Climaciaceae Kindb.,
Cryphaeaceae Schimp., Echinodiaceae Broth., Entodontaceae Kindb.,
Fabroniaceae Schimp., Fontinalaceae Schimp., Helodiaceae Ochyra,
Heterocladiaceae Ignatov & Ignatova, Hylocomiaceae M.Fleisch.,
Hypnaceae Schimp., Lembophyllaceae Broth., Leptodontaceae
Schimp., Lepyrodontaceae Broth., Leskeaceae Schimp., Leucodon-
taceae Schimp., Meteoriaceae Kindb., Microtheciellaceae H.A.Mill
& A.J.Harr., Myriniaceae Schimp., Myuriaceae M.Fleisch., Neckeraceae
Schimp., Orthorrhynchiaceae S.H.Lin, Phyllogoniaceae Kindb.,
Plagiotheciaceae M.Fleisch., Pleuroziopsidaceae Ireland, Prionodon-
taceae Broth., Pseudoleskeaceae Schimp., Pseudoleskeellaceae
Ignatov & Ignatova, Pterigynandraceae Schimp., Pterobryaceae
Kindb., Pylaisiaceae Schimp., Pylaisiadelphaceae Goffinet & W.R.
Buck, Regmatodontaceae Broth., Rhytidiaceae Broth., Rigodiaceae
H.A.Crum, Rutenbergiaceae M.Fleisch., Scorpidiaceae Ignatov &
Ignatova, Sematophyllaceae Broth., Sorapillaceae M.Fleisch., Ste-
reophyllaceae W.R.Buck & Ireland, Symphyodontaceae M.Fleisch.,
Theliaceae M.Fleisch., Thuidiaceae Schimp., Trachylomataceae
W.R.Buck & Vitt

Acknowledgements

Sincere thanks are due to J.L.Reveal (Ithaca) as well as W.F.Prud'homme van Reine and J.F.Veldkamp
(Leiden) for comments and discussions about suprafamilial taxonomy, to D.Quandt (Dresden) for providing
DNA from Rhacocarpus and Bartramiaceae species, and to B.Giesicke (Berlin) for technical assistance.

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Received 3 September 2007, accepted in revised form 30 October 2007.

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