Phylogenetic Relationships and Morphological Evolution in A Major Clade of

Phylogenetic Relationships and Morphological Evolution in a Major Clade of
Leafy Liverworts (Phylum Marchantiophyta, Order Jungermanniales): Suborder

Jungermanniineae
Author(s): Blanka Shaw, Barbara Crandall-Stotler, Jiří Váňa, Raymond E. Stotler, Matt von Konrat, John
J. Engel, E. Christine Davis, David G. Long, Pavel Sova, and A. Jonathan Shaw
Source: Systematic Botany, 40(1):27-45.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364415X686314
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Systematic Botany (2015), 40(1): pp. 27–45
© Copyright 2015 by the American Society of Plant Taxonomists
DOI 10.1600/036364415X686314
Date of publication February 12, 2015
Phylogenetic Relationships and Morphological Evolution in a Major Clade

of Leafy Liverworts (Phylum Marchantiophyta, Order Jungermanniales):
Suborder Jungermanniineae
Blanka Shaw,1,8 Barbara Crandall-Stotler,2 Jiřı́ Váňa,3 Raymond E. Stotler,2† Matt von Konrat,4 John J. Engel,4
E. Christine Davis,5 David G. Long,6 Pavel Sova,7 and A. Jonathan Shaw1
1
Department of Biology, Duke University, Durham, North Carolina 27708, U. S. A.
2
Department of Plant Biology, Southern Illinois University, Carbondale, Illinois 62901-6509 U. S. A
3
Department of Botany, Charles University, Benátská 2, CZ-12801 Praha 2, Czech Republic.
4
Department of Botany, The Field Museum, 1400 South Lake Shore Drive, Chicago, Illinois 60605–2496, U. S. A.
5
Department of Biology, University of Florida, 614 Carr Hall, Gainesville Florida 32611, U. S. A
6
Royal Botanic Garden, Edinburgh EH3 5LR, Scotland.
7
University of South Bohemia, Faculty of Science, Branisovska 31, 370 05 Ceske Budejovice, Czech Republic.
†
Ray Stotler died in December, 2013, before this paper was accepted for publication; he played a major role
in the research, including preparation of this manuscript.
8
Author for correspondence (blanka@duke.edu)
Communicating Editor: Thomas L. P. Couvreur

Abstract—The suborder Jungermanniineae of the Jungermanniales is a major lineage of leafy liverworts, recognized in recent classifica-
tions to include 15 families. Gametophytes within the suborder are morphologically diverse, but commonly anisophyllous to distichous,
usually with succubous, rarely transverse or incubuous, leaf insertions. Sporophytes are frequently, but not universally, enclosed by stem-
derived perigynia or coelocaules, often accompanied by perianth reduction or loss and some level of geocauly or marsupial development.
We herein provide the first comprehensive molecular phylogeny of this geographically widespread suborder, using sequences generated
from one nuclear (rpb2), two mitochondrial (nad1 and rps3), and seven plastid (atpB, psbA, psbT-H, rbcL, rps4, trnG and trnL) loci, sampled
from 279 accessions representing 163 species in 57 genera. Ancestral states were reconstructed for 14 morphological characters generally
considered taxonomically diagnostic for families in the suborder. Our phylogenetic analyses support the return of Leiomylia (=Mylia anomala)
to the Myliaceae, removal of Myliaceae from the Jungermanniineae, and validation of the monogeneric suborder Myliineae subord. nov.
to house it. Eighteen families are recognized within the Jungermanniineae, nine of which are monogeneric; namely, Endogemmataceae,
Harpanthaceae, Gyrothyraceae, Arnelliaceae, Saccogynaceae, Geocalycaceae, Jackiellaceae, Notoscyphaceae stat. nov., and Trichotemnomaceae.
The generic compositions of other families are modified as follows: Saccogynidium is transferred from Geocalycaceae to a newly named
subfamily of Acrobolbaceae, Acrobolbaceae subf. Saccogynidioideae, and one other subfamily of the Acrobolbaceae is validated,
Acrobolbaceae subf. Austrolophozioideae; Hygrobiella is included in Antheliaceae (previously in Cephaloziaceae or its own family);
Jungermanniaceae is broadened to include Mesoptychiaceae and Delavayellaceae; Cryptocoleopsis and Nardia are transferred from
Solenostomataceae to Gymnomitriaceae; Gottschelia, Herzogobryum, and Nothogymnomitrion are excluded from the Jungermanniineae;
Solenostomataceae is recognized to include Solenostoma, Arctoscyphus, Cryptocolea, and Diplocolea. Additional nomenclatural changes
include recognizing Horikawaella as a synonym of Solenostoma and Apomarsupella as a synonym of Gymnomitrion, establishing two new
subgenera of Solenostoma, Solenostoma subg. Metasolenostoma and Solenostoma subg. Eucalyx, and transferring Jungermannia conchata
to Cephalozia. Morphological character state reconstructions identify dioecious inflorescences, gametangia on leading stems, flagelliform or
stoloniferous branches absent, dorsal leaf insertions not overlapping the stem midline, large underleaves, and lack of gemmae as ancestral
within the Jungermanniineae. All morphological characters appear to be moderately to highly homoplasious within the suborder.
Keywords—Chloroplast sequence data, Jungermanniaceae, liverwort phylogeny, mitochondrial sequence data, morphological character
evolution, nuclear sequence data
Fossil records suggest that the early Paleozoic era, some Concepts of the suborder have changed dramatically since
500–350 mya, marked the beginnings of land plant diver- its validation by Crandall-Stotler and Stotler (2000), pri-
sification, with the ancestors of several major clades estab- marily as a consequence of molecular phylogenetic studies
lished by the Devonian epoch (Kenrick and Crane 1997). (Forrest et al. 2006; He-Nygrén et al. 2006; Heinrichs et al.
Megafossils that are morphologically similar to extant simple 2007; Vilnet et al. 2010). When first proposed by Schuster
thalloid liverworts are known from this epoch, including in 1953, the suborder encompassed all families of what
Riccardiothallus Guo et al., Pallaviciniites R. M. Schust., and is currently recognized as the Jungermanniales, but by
Metzgeriothallus Hernick et al., and there is evidence that 2000 several additional suborders had been established,
liverworts may be the earliest-diverging clade among the and the Jungermanniineae were limited to a very broadly
bryophyte lineages (Qiu 2008; but see also Finet et al. 2010; defined Jungermanniaceae, a monotypic Mesoptychiaceae,
Wodniok et al. 2011; Cox et al. 2014). Over the last ten years, the Gymnomitriaceae, and the Scapaniaceae (Crandall-
molecular phylogenetic analyses have resolved the deep Stotler and Stotler 2000). The most recently published clas-
clades of liverworts (Davis 2004; He-Nygrén et al. 2004; sification of liverworts by Crandall-Stotler et al. (2009), which
Forrest et al. 2006), and made progress in resolving relation- incorporated phylogenetic inferences from Davis (2004),
ships among major lineages of the largest clade, the leafy He-Nygrén (2007), He-Nygrén et al. (2004, 2006), Schill
liverworts, i.e. subclass Jungermanniidae Engl. Fine-scale et al. (2004), Yatsentyuk et al. (2004), Heinrichs et al. (2005,
relationships within many speciose lineages, however, 2007), Forrest et al. (2006), Hentschel et al. (2006, 2007), de
are still poorly understood. The suborder Jungermanniineae Roo et al. (2007), Vilnet et al. (2007), and Feldberg et al.
R.M. Schust. ex Stotler & Crand.-Stotl., as circumscribed (2009), recognized 15 families in the Jungermanniineae. Our
by Crandall-Stotler et al. (2009), comprise one such group. study is designed to assess the relationships among genera
27
28 SYSTEMATIC BOTANY [Volume 40
and families included in this circumscription of the sub- the suborder includes analyses of morphological trait evolu-
order. We also address species relationships within one clade tion in the context of phylogenetic relationships inferred
(Solenostomataceae) by sampling multiple accessions, when from molecular data.
available, of the included species. The present phylogenetic analysis is based on nucleotide
Within the suborder Jungermanniineae, circumscriptions sequence variation from 10 genomic regions representing
of the Jungermanniaceae and the genus Jungermannia L. all three genomes, mitochondrial, plastid, and nuclear, and
have been diverse. In early morphology-based classifica- includes 279 accessions representing 57 genera. The specific
tions (e.g. Schuster 1970; Váňa 1973), the family was goals of this work were to (1) resolve relationships among
broadly circumscribed to include Jungermannia, Nardia Gray, species, genera, and families within the Jungermanniineae,
Notoscyphus Mitt., Cryptocolea R.M. Schust., Cryptocoleopsis (2) reconstruct ancestral states for selected morphological
Amak., Diplocolea Amak., Scaphophyllum Inoue, Horikawaella characters, and (3) propose a revised classification for the
Hatt. & Amak., and Hattoria R.M. Schust. in the sub- Jungermanniineae based on results from our analyses. Our
family Jungermannioideae and numerous other genera general goals for the morphological reconstructions were
classified in eight additional subfamilies (Crandall-Stotler to infer ancestral states for the Jungermanniineae as a
and Stotler 2000). Jungermannia was circumscribed to whole and to assess, qualitatively, the degree to which mor-
include four subgenera, namely J. subg. Jungermannia, phological characters commonly used to delineate genera
J. subg. Liochlaena (Nees) S. Arnell, J. subg. Solenostoma and families in this group are or are not homoplasious.
(Mitt.) Amak., and J. subg. Plectocolea (Mitt.) Amak. As a
consequence of their molecular analysis, Hentschel et al.
(2007) proposed to split this broadly interpreted genus into Materials and Methods
three genera corresponding to the subgenera, Jungermannia, Taxon Sampling—Our total data set includes 163 species representing
Liochlaena Nees, and Solenostoma Mitt. (incl. Plectocolea 57 of the 63 genera classified in 15 families of the Jungermanniineae
(Mitt.) Mitt.). This system, including the broad concept (Crandall-Stotler et al. 2009). In the absence of unambiguous informa-
of Solenostoma, has been accepted by most recent authors tion about circumscription of the Jungermanniaceae, our sampling
includes multiple species within those genera included in the family,
(e.g. Crandall-Stotler et al. 2009; Feldberg et al. 2009; Váňa taxa that have been classified in proximity to the Jungermanniaceae,
and Long 2009; Váňa et al. 2010a) although some (e.g. and selected more distant taxa to resolve polarity within a phylogeneti-
Vilnet et al. 2011; Bakalin and Vilnet 2012) continue to cally circumscribed in-group. Voucher information and GenBank acces-
recognize Solenostoma and Plectocolea as distinct genera. sion numbers are provided in Appendix 1.
Genomic Sampling—Sequences were generated for one nuclear,
Hentschel et al. (2007) proposed that Jungermanniaceae
two mitochondrial, and seven plastid loci (Table 1). RNA polymerase II
should include only Jungermannia, Delavayella Steph., (rpb2) represented the nuclear genome, and the mitochondrial genome
Eremonotus Lindb. & Kaal., Leiocolea (Müll. Frib.) H. Buch, was represented by intron sequences in the NADH protein-coding
and Liochlaena, but they did not suggest a family place- subunit 1 (nad1) and ribosomal small subunit protein 3 (rps3). From
ment for the remaining genera usually classified in the the plastid genome we sequenced CF1 ATPase beta-subunit gene
(atpB), photosystem II (PSII) reaction center protein D1 (psbA), photo-
subfamily Jungermannioideae. Crandall-Stotler et al. (2008) system II reaction center protein T (psbT-H), ribulose-bisphosphate
recognized three families for the jungermannioid complex, carboxylase gene (rbcL), ribosomal small protein 4 (rps4), tRNA(Gly)
placing Delavayella in Delavayellaceae, Leiocolea and Liochlaena (UCC) (trnG), and the trnL (UAA) 59 exon trnF (GAA) region (trnL).
in Mesoptychiaceae, and the residue of the Jungermannioideae Extractions, amplifications, and sequencing followed methods and
used primers described by Shaw et al. (2003) except for atpB and rpb2.
in the Jungermanniaceae, excluding other subfamilies from
Primer sequences for all the loci used in this study were provided in
the family. In 2009, these families were redefined, the Cooper et al. (2011).
Mesoptychiaceae was combined with the Jungermanniaceae, Phylogenetic Analyses—Sequences were initially aligned using
and the Solenostomataceae was erected (Crandall-Stotler et al. MUSCLE (vers. 3.7; Edgar 2004) with default options and then
2009). There is, however, still ambiguity as to the relation- adjusted manually with PhyDEÒ 0.995 (Müller et al. 2007). Regions of
ambiguous alignment were identified and excluded from further analy-
ships of these taxa. Addressing the phylogenetic affinities sis. Optimal substitution models were selected to maximize the Akaike
of taxa previously considered subgenera of Jungermannia, information criterion (AIC) statistic for each locus using the software
namely, Liochlaena, Solenostoma, and Plectocolea (Amakawa 1959, MrModeltest (vers. 2.3; Nylander 2004) in conjunction with PAUP* (vers.
1960; Váňa 1973) and related genera of the Jungermanniaceae 4.0b10; Swofford 1998). Incongruence among loci (defined as conflicting
clades supported by > 70 bootstrap (BS) support) were identified using the
and Solenostomataceae is, therefore, an additional focus of
TCT (Tree Congruence Tester) program available at http://www.biology
our study. .duke.edu/bryology/cymon.html#software. No significant conflicts were
Within the Jungermanniineae, leaves are usually succubous, detected so no taxa were removed from the concatenated data set.
rarely transverse or incubous, in arrangement; sporophytes Two data sets were analyzed. One included all 279 accessions and
are often, but not always, protected by stem perigynia or the other was pruned to include just one accession per 169 species. For
the larger unpruned data set, maximum likelihood (ML) trees were
coelocaules; perianths are frequently reduced or lacking; constructed for a series of concatenated data sets to test the effects of
and marsupia may be present or absent (Crandall-Stotler missing data on topology and clade support. Analyses were run with
et al. 2009). In addition, the suborder is heterogeneous only those accessions represented by eight or more loci, six or more,
with regards to leaf form, underleaf presence or absence, five or more, and with all accessions including those represented by
only one locus. We found no topological differences among the
branching types and patterns, and form and position of
restricted data sets. Results described below are based on all acces-
reproductive structures, all characters considered of taxo- sions, even those represented in the combined data set by only one of
nomic importance in the delimitation of genera and families the 10 loci. Single locus bootstrap trees are available upon request.
(He-Nygrén 2007). It has been suggested that the pres- Trees from concatenated data sets with taxa having minimum numbers
ence of perigynia is a synapomorphy for the suborder of successfully sequenced loci (as described above) are not presented,
but are available upon request. Reconstructions were obtained using
(He-Nygrén 2007), but patterns of evolutionary change in GARLI, version 1.0 (Zwickl 2006) not specifying nor fixing model
the key taxonomic characters of the suborder have never parameters. These analyses were run on the Cipres portal (http://www
been reconstructed. Our evaluation of relationships within .phylo.org/portal2/, Miller et al. 2010). Clade support was estimated
2015] SHAW ET AL.: PHYLOGENY OF THE JUNGERMANNIINEAE 29
Table 1. Genomic regions sequenced for phylogenetic analyses of the Jungermanniineae (shown in Figs. 1, 2, and Fig. S1). atpB, psbA, psbT,
rbcL, rps4, trnG, and trnL are from the plastid genome, nad1 and rps3 are from the mitochondrial genome, and rpb2 is from the nuclear genome.
N tree S1 = sample size in the analysis shown in Fig. S1; N Fig. 1 = sample size in the analysis shown in Fig. 1. PS = potentially informative
N tree % Missing data N tree % Missing data Aligned Analyzed Autapo- Invariant
Marker Genome S1 (Fig. S1) Fig. 1 (Fig. 1) length length morphic sites sites PS sites % PS sites Model
atpB cp 70 75% 67 60% 1,181 997 595 68 334 34% GTR+I+G

psbA cp 225 19% 154 9% 1,473 1,206 769 101 336 28% GTR+I+GHKY+I+G
psbT cp 184 34% 131 22% 589 443 199 46 198 45% GTR+I+G
rbcL cp 137 51% 106 37% 1,551 1,346 797 74 475 35% GTR+I+G
rps4 cp 204 27% 148 12% 751 583 200 58 325 56% GTR+I+G
trnG cp 160 43% 113 33% 1,013 656 123 67 466 71% GTR+I+G
trnL cp 229 18% 136 20% 913 518 214 50 254 49% GTR+I+G
nad1 mt 202 28% 140 17% 990 844 590 79 175 21% GTR+I+G
rps3 mt 167 40% 125 26% 1,186 1,104 690 146 268 24% GTR+I+G
rpb2 nc 98 65% 66 61% 765 695 434 21 240 35% GTR+I+G
combined 10 loci 279 40% 169 30% 10,412 8,391 4,611 710 3,071 37% GTR+I+G
using GARLI, with 300 bootstrap replicates. The best substitution model are provided in Table 1. Among the plastid loci, trnG was
for every gene in the unpruned data set was found to be GTR+I+ G so most variable (in terms of potentially synapomorphic substi-
the concatenated data were analyzed using that single model.
Additional phylogenetic analyses were conducted on a pruned data tutions) and psbA was least variable. The two mitochondrial
set containing one accession per species (supplementary Table S1). The loci (nad1, rps3) were relatively nonvariable (21% and 24%
reconstruction from this more restricted data set is provided as Fig. 1, synapomorphic sites, respectively). The nuclear locus, rpb2,
but the more taxon-inclusive reconstruction (all samples) is provided was the least variable of the non-organellar loci (Table 1). The
as supplementary Fig. S1. The ML reconstruction based on all samples
(including multiple accessions per species) for the Solenostomataceae unpruned data set (279 taxa; Fig. S1) contained 39% missing
s. s. is presented as Fig. 2, since that family is also a focus of the study. data; the pruned data set (169 taxa; Fig. 1) contained 30%
Relationships among the 169 accessions (one per species; Fig. 1) were missing data. Specifically, 18 accessions were represented by
reconstructed under ML using the search methods described above. data from all of the ten genomic regions sequenced, 36 for
In addition to the ML analyses, Bayesian analyses were conducted on
the pruned data set using MrBayes v3.1.2. We ran heterogeneous nine, 40 for eight, 41 for seven, 41 for six, 27 for five, 22 for
Bayesian analysis with site-specific (variable) rates for each partition four, 17 for three, eight for two, and 29 for only one.
with the following settings: unlink tratio=(all) revmat=(all); statefreq= Phylogenetic Relationships: Jungermanniineae—ML boot-
(all) shape=(all) pinvar=(all); lset applyto=(all) ngammacat=4; prset strap values and Bayesian posterior probabilities were
ratepr=variable. Based on results from MrModeltest, all partitions for
this data set, pruned to include one accession per species, were run moderate to high for many of the important nodes that
with the GTR+I+G substitution model with the exception of psbA, for define major clades (Figs. 1, 2). The phylogeny was rooted
which the HKY+I+G model was optimal amog those tested. The Bayesian with Schistochila laminigera Hook. f. & Taylor (suborder
search was run as follows: mcmc ngen=60000000; samplefreq=5000; Perssoniellineae) as the outgroup (Fig. 1). For purposes of
printfreq=10000; nchains=4; nruns=2 temp=0.05. Tracer v1.5 (Rambaut
and Drummond 2007) was used to evaluate the burn-in and to examine describing the reconstruction shown in Fig. 1, clades are
log likelihoods, ensuring that the runs were in the stationary phase and labeled with family names. Some of the families, as defined
that adequate effective sample sizes (ESS) were attained. Construction of in the figure, have modified circumscriptions relative to
the consensus tree and estimation of clade posterior probabilities were current classifications (e.g. Crandall-Stotler et al. 2009). The
based on the trees sampled after the chains converged. Consensus topolo-
gies and support values from the different methodological approaches labels are included to facilitate the description of our
were compiled and drawn using TreeGraph 2.0.42–187 beta (http:// results, while modifications of family circumscriptions are
treegraph.bioinfweb.info/) and Figtree v.1.3.1. (http://tree.bio.ed.ac highlighted in the Discussion section below.
.uk/software/figtree/). The datasets used for phylogenetic analyses The Myliaceae, represented by Mylia taylorii (Hook.) Gray
were deposited in TreeBASE (http://purl.org/phylo/treebase/phylows/
study/TB2:S16128). and Leiomylia anomala (Hook.) J.J. Engel & Braggins in our data
Morphological Character State Reconstruction—Fourteen morphological set, has sometimes been included in the Jungermanniineae,
characters were scored for all species included in the molecular analyses. but is resolved as sister to all other ingroup taxa in the current
Eight of the characters were binary and the other six were multistate. All analyses. Although some taxa in the next two clades (moving
scoring was based on personal study of the vouchers and knowledge of
the taxa, supplemented with published descriptions; none were taken
up the tree in Fig. 1) have similarly been associated with
solely from literature sources. Ancestral state reconstructions were accom- the Jungermanniineae, these clades are best considered rep-
plished using the data set pruned to include one sample per species resentative of suborders Lophocoleineae and Cephaloziineae,
(Table S1). The characters and their states are listed in Table 2; charac- respectively (Fig. 1). Accessions resolved in the Lophocoleineae
ter state scores for each species are listed in supplementary Table S2.
Character state reconstructions were performed with Mesquite vers. 2.75
include representatives of the Lepidoziaceae, Pseudo-
(Maddison and Maddison 2011). Traits were mapped onto the maximum lepicoleaceae, Plagiochilaceae, and Lophocoleaceae, includ-
likelihood tree derived from Garli analyses of the pruned data set (Fig. 1). ing the genus Bragginsella R. M. Schust., whose relationships
Reconstructions were performed using the mk1 model. Ancestral states at have been controversial but which is supported with 100% BS
the base of the Jungermanniineae clade were inferred for those characters
as being nested within Lophocoleaceae. Nothogymnomitrion
where the state was not “ambiguous” in the Mesquite analyses. Levels
of homoplasy for each character were evaluated qualitatively by examina- R. M. Schust. and Herzogobryum Grolle, both formerly
tion of character state reconstructions mapped onto the ML tree. classified in the Gymnomitriaceae of the Jungermanniineae, are
likewise nested among representatives of the Cephaloziineae.
The Jungermanniineae (from the Acrobolbaceae upward in
Results
Fig. 1) is strongly supported (BS = 100%) as monophyletic.
Molecular Variation—Sequence characteristics and amounts Within the Jungermanniineae, species of the genera Lethocolea
of missing data for the 10 loci included in phylogenetic analyses Mitt. nom. cons., Saccogynidium Grolle, Goebelobryum Grolle,
Fig. 1. Maximum likelihood phylogenetic reconstruction of the Jungermanniineae (and outgroups) using GARLI (see Materials and Methods
for analytical details). Multiple accessions per species were pruned from the data set; see Fig. S1 for an ML reconstruction based on the unpruned data
set. Values above branches are ML bootstrap percentages / Bayesian PP. Thickened branches indicate ML bootstrap < 90% and Bayesian PP = 1.0.
Fig. 2. Maximum likelihood phylogenetic reconstruction of the Solenostomataceae using GARLI (see Materials and Methods for analytical details).
Multiple samples per species included where available. Based on the same data set as in Fig. S1. Values above branches are ML bootstrap percentages.
Table 2. Morphological characters and states scored for all species Solenostomataceae). Within the latter group, Endogemma
included in the phylogenetic analyses. caespiticium (Lindenb.) Konstant., Vilnet & A. V. Troitsky is
sister (with strong support; 89% ML bootstrap, 0.99 Bayesian
1. Underleaves: 0 = conspicuous; 1 = reduced to a few cells or papillae; PP) to the Gymnomitriaceae plus Solenostomataceae. The six
2 = absent accessions of E. caespiticium from Eurasia and North America
2. Interlocking merophytes: 0 = absent; 1 = present
3. Leaf form: 0 = undivided or emarginate; 1 = distinctly lobed are on an exceptionally long stem branch (and distinguished
4. Frullania-type branching: 0 = absent; 1 = present as the Endogemmataceae), but are little differentiated from
5. Flagelliform or stoloniferous branches: 0 = absent; 1 = present one another (Fig. S1).
6. Gemmae: 0 = absent; 1 = exogenous, angulate; 2 = exogenous, The Gymnomitriaceae contain nine species of Nardia
spherical to ovoid elliptical; 3 = endogenous
and although several of the species represented by multiple
7. Sexual condition: 0 = dioecious; 1 = monoecious
8. Position of androecia: 0 = on leading axes; 1 = on short specialized accessions are non-monophyletic (Fig. S1), the genus is very
branches strongly supported. Seven species of Marsupella Dumort.
9. Position of gynoecia: 0 = on leading axes; 1 = on short specialized form a monophyletic group sister to Poeltia campylata Grolle
branches within the Gymnomitriaceae (Fig. 1). Prasanthus suecicus
10. Stem perigynium: 0 = absent or very low, < 0.2 length of the perianth;
1 = > 0.3 < 1.0 length of the perianth; 2 = ³ the length of the perianth Lindb. is sister to Cryptocoleopsis imbricata Amakawa with
11. Perianth: 0 = present, well developed; 1 = absent or very reduced strong support. The Gymnomitriaceae also encompass
12. Calyptra type: 0 = with little or no shoot involvement; 1 = at least nine species of Gymnomitrion Corda, including four species
upper third of venter origin (shoot calyptra); 2 = little or no venter often classified as Apomarsupella R. M. Schust. that are
tissue present (coelocaule)
nested within the Gymnomitrion clade (shown in Fig. 1).
13. Geocauly: 0 = absent; 1 = moderate (Nardia or Harpanthus type);
2 = extensive (elongate marsupium) Species Relationships Within the Solenostomaceae—The
14. Capsule shape: 0 = spheroidal, length = width; 1 = ovoid or short Solenostomataceae form a monophyletic group supported
elliptical, < 2x as long as wide; 2 = long elliptic or cylindrical, by 100% BS and 1.00 PP. Our sampling of this clade includes
> 2x as long as wide approximately 51 species of Solenostoma (of an estimated
140 species recognized worldwide), and although the
species fall into multiple subclades that are each well-
Austrolophozia R. M. Schust., Acrobolbus Nees, and Tylimanthus supported, relationships among those subclades are in
Mitt. form a well-supported clade (Acrobolbaceae) that is some cases ambiguous (Fig. 2). We herein (below) describe
resolved as sister to the rest of the Jungermanniineae one new subgenus (Metasolenostoma) and elevate Eucalyx
(PP = 1.0). A small, strongly supported clade con- from sectional to subgeneric rank within Solenostoma to
taining Blepharidophyllum Ångstr., Clandarium (Grolle) formalize two of these monophyletic groups. These two
R. M. Schust., Trichotemnoma R. M. Schust., Acroscyphella groups each contain accessions representing Solenostoma
N. Kitag. & Grolle, Isotachis Mitt., and Balantiopsis Mitt. species from the Northern Hemisphere, including North
(Blepharidophyllaceae, Trichotemnomaceae, Balantiopsaceae) America, Europe, and Asia. Multiple accessions of S. obovatum
is then sister to the remaining taxa. Notoscyphus, Jackiella (Nees) C. Massal. (subg. Eucalyx) from North America and
Schiffn., Metacalypogeia (S. Hatt.) Inoue, Eocalypogeia (R.M. Europe are very similar at the nucleotide sequence level. A
Schust.) R. M. Schust., Mizutania Furuki & Z. Iwats., Mnioloma third clade (with 92% BS support) consisting of 12 Solenostoma
Herzog, Calypogeia Raddi, Geocalyx Nees, Southbya Spruce, species is here assigned to the subgenus Plectocolea. Seven
and Gongylanthus Nees form a clade resolved in the ML of the species in this clade are from Asia and three are
reconstruction that is in turn sister to all remaining taxa from Australia, with S. crenuliforme (Aust.) Steph., S. callithrix
in the study, but without support. However, within that (Lindenb. et Gottsche) Steph. and S. hyalinum (Lyell) Mitt.
clade, the Calypogeiaceae, Geocalycaceae, and Southbyaceae from North America and Europe nested within the
are each strongly supported as monophyletic (Fig. 1). group. A collection referred to S. rubripunctatum (S. Hatt.)
The ML tree shows a clade (Antheliaceae, including R. M. Schust., from China, is nested within S. hyalinum, on
Hygrobiellaceae), supported by 100% BS, that includes a long branch, without support. The relationship between
Hygrobiella Spruce, Pseudolepicolea trollii Grolle & Ando, and these two taxa warrants further study.
Anthelia (Dumort.) Dumort. The sister group relationship Subgenera Plectocolea (92% BS), Metasolenostoma (99% BS),
of this clade to the remaining taxa of Jungermanniineae and Eucalyx (BS=95%) form a paraphyletic grade basal to
is, however, without BS support. The Jungermanniaceae, subgenus Solenostoma (98% BS). The subgenus Solenostoma
including Jungermannia s. s. (with J. atrovirens Dumort., the includes Asian accessions from Bhutan, China, Japan, and
type of Jungermannia), Eremonotus, Liochlaena, and Delavayella, South Korea plus accessions from New Zealand and the
and the Mesoptychiaceae (including Mesoptychia (Lindb.) A. Kerguelen Islands, as well as S. pyriflorum Steph. from
Evans and formerly recognized genera Hattoriella (Inoue) North America (resolved with 100% support as sister
Inoue and Leiocolea form sister clades, and this more inclu- to S. faurieanum (Beauvd.) R. M. Schust. from Japan),
sive, fairly well supported clade that includes both families S. pseudopyriflorum Bakalin & Vilnet from Asiatic Russia,
is resolved sister to the rest of the Jungermanniineae (i.e. S. sphaerocarpum (Hook.) Steph. from Europe, S. confertissimum
above it in Fig. 1), albeit with poor support. (Nees) Schljakov, also from Europe, and S. abyssinicum
Monophyly of the remaining terminal clade, while resolved (Nees) Steph. (reduced to S. confertissimum in this study)
in the ML tree, is not supported by the bootstrap. Saccogyna from Africa. Species formerly segregated as the genus
viticulosa (L.) Dumort. is resolved as an early diverging lineage Horikawaella, namely, H. grosse-verrucosa (Amakawa &
within that group. A well-supported clade with Arnellia S. Hatt. and H. subacuta (Herzog) S. Hatt. & Amakawa,
fennica (Gottsche & Rabenh.) Lindb., two species of Harpanthus from China, are nested within Solenostoma subg. Solenostoma.
Nees, and Gyrothyra underwoodiana M. Howe, is sister Some species are clearly non-monophyletic; e.g. American
to the remaining taxa (Endogemmataceae, Gymnomitriaceae, accessions attributed to S. pyriflorum appear to be only
distantly related to Korean accessions, and one accession (2007), Hentschel et al. (2006, 2007), Wilson et al. (2007)
of S. appressifolium (Mitt.) Váňa & D. G. Long from China and Vilnet et al. (2007) resulted in numerous new insights,
is not resolved with the other three, also from China. most of which were incorporated in the most recent com-
Morphological Character Reconstructions—Ancestral states prehensive classification of liverworts by Crandall-Stotler
for morphological characters are summarized in supple- et al. (2009). Our results extend previous inferences and
mentary Fig. S2. Although character states for some traits provide support for additional realignments of several taxa
are ambiguous, it appears likely that the ancestor of of the Jungermanniineae.
Jungermanniineae had dioecious inflorescences, gametangia The Myliaceae are excluded from the Jungermanniineae
on leading stems, flagelliform or stoloniferous branches and recognized to comprise the new suborder Myliineae
absent, dorsal leaf insertions not overlapping the stem subord. nov. Although the suborder was proposed by Engel
midline, large underleaves, and no gemmae. and Braggins (2005), and referred to by He-Nygrén (2007),
The ancestral sexual condition in the Jungermanniineae it has never been validly published. Although Engel and
was unambiguously dioecious, but monoecy appears to Braggins (2005) indicated in the abstract that Trabacellula
have evolved at least 16 times (Fig. S2). Gemmae evolved Fulford should also be included in the suborder based on
multiple times in distantly relatedly families of the similar cell wall architecture, in the taxonomic section of their
Jungermanniineae. Underleaves were gained multiple times manuscript, that genus is retained in the Cephaloziaceae
in the Jungermanniineae and appear to be present in the subfam. Trabacelluloideae (Fulford) R. M. Schust. Cell wall
ancestor of Mesoptychia, Delavayella, Liochlaena, Eremonotus, architecture aside, Trabacellula is morphologically congruent
and Jungermannia within the Jungermanniaceae (Fig. 1, with the Cephaloziaceae, where it has traditionally been
Fig. S2). Underleaves were also present in the common placed. Because Trabacellula was not included in our molec-
ancestor of the Southbyaceae (Fig. 1, Fig. S2) but absent ular sampling, we cannot assess whether transfer out of
in the common ancestor of the Gymnomitriaceae+ the Cephaloziaceae is justified and therefore we retain it in
Solenostomataceae clade. Underleaves were subsequently that family.
regained in the common ancestor of Nardia species. There Within the Myliaceae, the taxonomic status of Leiomylia
appear to have been at least 4–5 additional losses of under- J. J. Engel & Braggins, erected to include only Mylia anomala
leaves in the Jungermanniineae (Table S2 and Fig. S2). We (Hook.) Gray, is equivocal. Our results mirror those of de
cannot accurately count the numbers of gains and losses Roo et al. (2007) in resolving L. anomala and M. taylorii
for all characters because our sampling was not exhaustive as sister taxa in a strongly supported clade. These results
at the species level and and some additional character state indicate a much closer relationship between L. anomala
transitions occurred within families. A qualitative assessment and M. taylorii than that proposed by Engel and Braggins
of additional transitions can be gained from studying Fig. S2. (2005), who placed them in separate suborders, and clearly
All morphological traits are characterized by extensive support the return of Leiomylia to the Myliaceae. What
homoplasy, including parallel losses and gains. Reversals cannot be determined with certainty is whether Leiomylia
also appear to have occurred repeatedly. This is true of both should be reduced to Mylia, as proposed by de Roo et al.
gametophytic and sporophytic characters; capsule shape (2007). There is disagreement among us as to whether
has clearly transitioned between ovoid and spheroidal Leiomylia should be recognized, but our molecular results
in multiple lineages. Early-diverging lineages within the do not support any particular taxonomic conclusion regard-
Jungermanniineae repeatedly evolved more narrowly elon- ing the two genera. For now, the best decision seems to be
gate to cylindrical capsules but this capsule shape seems to recognize Leiomylia, pending more expansive population-
not to have re-evolved in more derived lineages, which are level studies of the Myliaceae and more exhaustive studies
characterized by shorter and broader capsules (Fig. S2). of the chemical and cellular features of the cell wall grids.
Gynoecia and androecia were borne on the leading axes Monophyly of the Myliineae is strongly supported in
of ancestral Jungermanniineae, but shifts to short lateral our analyses, but its phylogenetic position is ambiguous.
branches occurred multiple times; in some lineages this It is resolved as sister to the large Lophocoleineae/
shift in the position of female and male reproductive struc- Cephaloziineae/Jungermanniineae clade but with poor
tures occurred together, but shifts in gynoecia or androecia, support (Fig. 1), a position also resolved in the parsimony
but not both, occurred in a few lineages. Flagelliform or analysis of He-Nygrén (2007). But in other analyses, e.g.
stoloniferous branches evolved multiple times, but do not those of de Roo et al. (2007), Heinrichs et al. (2007),
currently characterize large clades but rather one or a few Hentschel et al. (2007) and the maximum likelihood analysis
species scattered across the phylogeny. One of the least of He-Nygrén (2007), it was resolved as the earliest diverg-
homoplasious characters pertains to the dorsal leaf inser- ing lineage of the Jungermanniineae. Considering the
tion overlapping (or not) the stem midline, but even that unique blend of morphological characters displayed by
character shows some homoplasy. the group, in combination with our molecular data, rec-
ognition of a separate suborder seems justified.
Circumscription of the Acrobolbaceae is broadened to
Discussion
include Saccogynidium, a genus formerly aligned with the
New Phylogenetic and Taxonomic Inferences—Early molecu- Geocalycaceae (Grolle 1960; Crandall-Stotler et al. 2009),
lar phylogenetic analyses addressed primarily the back- with which it shares many morphological features, and
bone relationships among classes, orders and suborders a new subfamily, Acrobolbaceae subf. Saccogynidioideae
of liverworts (e.g. Davis 2004; Forrest and Crandall-Stotler subf. nov., is established for it. Schuster (2001) recognized
2004, 2005; Heinrichs et al. 2005; He-Nygrén et al. 2006; three subfamilies in the Acrobolbaceae, namely, subf.
Forrest et al. 2006). These and subsequent family level Acrobolboideae R. M. Schust. nom. inval. (without diagnosis)
studies, such as those of de Roo et al. (2007), He-Nygrén comprising Acrobolbus, Marsupidium Mitt., and Tylimanthus,
a monogeneric subf. Austrolophozioideae R. M. Schust. phological and molecular differences between them justify
nom. inval. (without diagnosis), and subf. Lethocoleoideae, their placement in separate families, accommodated by the
as established and delimited by Grolle (1972) to include elevation of Jungermanniaceae subfam. Notoscyphoideae
Lethocolea and Goebelobryum. Although all taxa in the (Schuster 1970, 2002) to the monogeneric Notoscyphaceae.
Acrobolbaceae form well-developed marsupia, in the latter Geocalycaceae, in the sense of Crandall-Stotler et al.
subfamily the marsupia are hollow and derived from a (2009), is polyphyletic and is consequently now recognized
stem perigynium; i.e. Calypogeia type, while in the other two to include only the genus Geocalyx. Additional monogeneric
subfamilies, they are solid and derived from a coelocaule; families are established for Notoscyphus, Saccogyna, and
i.e. Tylimanthus type. In our study, Goebelobryum, which has Harpanthus (see taxonomic section below) and Saccogynidium
a Calypogeia type marsupium and Austrolophozia, with a is transferred to the Acrobolbaceae as already discussed.
Tylimanthus type marsupium, are resolved as sister taxa The characters that have suggested a relationship among
in a strongly supported lineage; therefore, Goebelobryum is these genera; i.e. the restriction of gametangia to abbre-
transferred to the subfamily Austrolophozioideae. Despite viated ventral branches, Calypogeia type marsupia and
these significant differences in marsupial development, the bifid underleaves, have evolved multiple times in the
taxa share the following suite of characters: leaves spinose- Jungermanniineae and are not indicative of a family level
dentate to lobulate, often with reddish to brown pigments, relationship, as previously assumed.
cell walls smooth, oil bodies large and few in number, rhi- Hygrobiella and Anthelia are included in a single family, as
zoids in dense mats that often form decurrent bundles previously postulated by Müller (1948, 1954), who placed
on the ventral surface of the stem, branches only of the them in the Hygrobiellaceae. Much later, Konstantinova
lateral Plagiochila type, and male bracts little modified and Vilnet (2009: 65) proposed to establish Hygrobiellaceae
from the leaves (Schuster 2001; Engel and Glenny 2012). as a new family, but this publication of the name was not
Saccogynidium is resolved as a separate lineage within the valid since the proper basionym was not cited; however,
family (Fig. 1) and is also morphologically distinct in the name was validly published by Konstantinova et al.
having large, bilobed underleaves and gametangia always (2014). Based on the principle of priority, nonetheless, the
borne on very short, determinant ventral branches. Although family name that must be applied to this lineage is
Marsupidium sometimes produces androecia and occasionally Antheliaceae, validly published by Schuster (1963). In more
gynoecia on short to long ventral branches, the androecium- recent treatments, Hygrobiella has been classified in the
bearing branches can continue growth after gametangial Cephaloziaceae based on Schuster’s (1973) transfer of the
maturation (Engel and Grolle 1971; Engel and Glenny genus to that family (e.g. Paton 1999; Damsholt 2002;
2008), in contrast to the very specialized reduced sexual Crandall-Stotler et al. 2009). Schuster (1979), in contrast
branches of Saccogynidium. In Saccogynidium marsupia are to earlier workers, considered Anthelia to be a “highly
of the Calypogeia type, developmentally most like those of isolated taxon” unrelated to Hygrobiella and proposed its
Goebelobryum, and its capsule wall is multistratose, a char- placement in a monogeneric suborder Antheliineae, which
acter shared with all other taxa in the Acrobolbaceae. In was subsequently recognized and validly published by
addition to naming a new subfamily for Saccogynidium, we Crandall-Stotler and Stotler (2000). In 2006 He-Nygrén et al.
also provide a diagnosis (below) to validate subfamily showed that Anthelia belongs in the Jungermanniineae, a
Austrolophozioideae; A. subfam. Acrobolboideae has recently concept accepted by Crandall-Stotler et al. (2009), while
been validated by Briscoe et al. (2015). Vilnet et al. (2012) resolved Hygrobiella as an outlier of
No changes are proposed to the circumscriptions of the Cephaloziineae. Ours is the first study to include both
Blepharidophyllaceae, Trichotemnomataceae, Balantiopsidaceae, genera, and resolve their close relationship.
Jackiellaceae and Gyrothyraceae in Crandall-Stotler et al. The strongly supported resolution of Pseudolepicolea trollii
(2009), and the Calypogeiaceae is the modified only by the ssp. andoi (R. M. Schust.) S. Hatt. & Mizut. within the
inclusion of Mizutania, as already proposed by Masuzaki Antheliaceae lineage is puzzling, considering its very dif-
et al. (2010). Our molecular data show that Blepharidophyllum ferent morphology from its sister taxon, Anthelia. These
and Clandarium are closely related, and Grolle (1965) sug- results also render the genus Pseudolepicolea Fulford &
gested that they be classified in a single genus. Since J. Taylor, as circumscribed by Hässel de Menéndez and
B. densifolium (Hook.) Ångstr. ex C. Massal., the type of Solari (1975), Grolle (1983) and Crandall-Stotler et al.
Blepharidophyllum, was not included in our study, we main- (2009), polyphyletic, with the generitype, P. quadrilaciniata
tain them here as separate genera pending future studies. (Sull.) Fulford & J. Taylor, resolved in the Lophocoleineae.
The Trichotemnomataceae is strongly supported as sister to We have verified that the voucher specimen for the
the Balantiopsidaceae, as also shown by He-Nygrén (2007). P. trollii ssp. andoi sequences is correctly identified. Female
In agreement with the analysis of Hendry et al. (2007), plants with young sporophytes found in the collection
Jackiella is resolved as a monophyletic lineage of the have perianths that are elongate, 6-keeled, and deeply
Jungermanniineae, but with ambiguity as to its precise 12-lobed at the mouth, with the lobes long, acuminate.
relationships (Fig. 1). A weakly supported sister relation- Sporophytes are enclosed in a well-developed shoot calyp-
ship to Notoscyphus is suggested, and there are shared tra and penetrate the stem to the level of the bracts; the
morphological characters that support at least a distant capsule is shortly ellipsoidal, with a 2-layered wall, and
relationship between these taxa; e.g. unlobed leaves, the seta averages six cells in diameter, with 18–22 rows of
restriction of branching to the ventral Bazzania type, leaf epidermal cells that are slightly larger than the internal
cells with large trigones and 1–3 large, granular oil bodies, cells. These features are held in common with P. fryei
a seta comprised of eight rows of large epidermal cells and (Perss.) Grolle & Ando, a species recognized by Schuster
a core of much smaller inner cells, and elongate capsules (1960) as the type of the genus Lophochaete R. M. Schust.,
with 2-stratose walls. On the other hand, substantial mor- and differ from the true calyptra and multistratose capsule
wall of P. quadrilaciniata (Schuster 1966; Hässel de Menéndez and Gymnomitrion, a topology also resolved in Vilnet et al.
and Solari 1975). It is tempting to conclude from our analy- (2010, 2011). The fairly small molecular distance between
sis that Lophochaete should be recognized as a genus, but the two lineages and several shared morphological features
we have no molecular data on P. fryei or other species of (e.g. the development of stem perigynia, reduction in size
Pseudolepicolea. In addition, there seems to be little morpho- of perianths, occurrence of two or three large, granular oil
logical connection between P. trollii ssp. andoi and the other bodies per cell, and isodiametric outer capsule wall cells
genera of the Antheliaceae. Consequently, we are reserving with nodular thickenings on all walls) support our transfer
judgment on the disposition of this taxon, pending addi- of Nardia to the Gymnomitriaceae. The major morphological
tional sampling within the Pseudolepicoleaceae. divergences between the two lineages are the evolution of
Jungermanniaceae is broadened to include five genera, underleaves in Nardia and dorsally overlapping leaf inser-
namely Jungermannia, Eremonotus, Liochlaena, Delavayella, tions (i.e. dorsally interlocking merophytes) in the “Poeltia to
and Mesoptychia, including all species of the former Gymnomitrion clade,” but both of these traits are homoplasious
Leiocolea and Hattoriella (Váňa et al. 2012b). As in earlier (see supplementary Fig. S2). Although nine species of Nardia
analyses by de Roo et al. (2007), Hentschel et al. (2007) were included in our sampling, interspecific relationships
and Vilnet et al. (2010, 2011), three lineages are resolved within the genus are unclear, due in part to the non-
within the Jungermanniaceae clade, each of which has been monophylly of multiple accessions of N. geoscyphus (De Not.)
previously recognized as a family; i.e. Jungermanniaceae, Lindb. and N. compressa (Hook.) Gray. Nardia succulenta
Delavayellaceae, and Mesoptychiaceae (e.g. He-Nygrén (Rich. ex Lehm. & Lindenb.) Spreng. and N. assamica (Mitt.)
et al. 2006). Our analyses suggest that these lineages are Amakawa, which were recognized as related taxa by Váňa
closely related and we therefore agree with Hentschel et al. (1976) and Engel (1988), are strongly supported as sister
(2007) and define the Jungermanniaceae to include all taxa. However, N. insecta Lindb., a species that is mor-
three. Morphological characters that also support recogni- phologically unlike them and quite like N. geoscyphus and
tion of a single family include the following: branching N. lescurii (Austin) Underw. (Schuster 1969; Váňa 1976) is
only from lateral merophytes, gynoecia and androecia only sister to them, forming a clade that is sister to the rest of
on leading stems, long emergent perianths that are the genus. Although our sampling within Nardia is more
contracted at the mouth, bistratose capsule walls, and in extensive than any previously published treatment, expanded
all taxa except M. sahlbergii (Lindb. & Arnell) A. Evans, population level studies are needed to resolve species level
calyptral development with little or no shoot involvement, relationships and evolutionary groups within the genus.
no perigynial development, and no geocauly. Since its erection by Amakawa (1959), Cryptocoleopsis has
The Arnelliaceae are resolved as polyphyletic, with Arnellia been aligned with taxa of the Jungermanniaceae, although
sister to a Gyrothyra/Harpanthus clade and the Southbya/ some resemblance to Prasanthus was noted by Amakawa
Gongylanthus lineage sister to the Geocalycaeae. These results (1959). Inoue (1983) reduced Gymnomitrion integerrimum
support the transfer of Southbya and Gongylanthus to the N. Kitag., named by Kitagawa (1959), to synonymy under
recently named Southbyaceae as discussed by Váňa et al. Cryptocoleopsis imbricata, with an indication that the taxon
(2012a), and the circumscription of Arnelliaceae as a should remain in Jungermanniaceae. It is noteworthy that
monogeneric family. A close phylogenetic relationship among Cryptocoleopsis is not only resolved within the Gymnomitriaceae,
Arnellia, Gyrothyra, and Harpanthus lineages is moderately but sister to Prasanthus, which it resembles in lacking a perianth
supported, but substantial levels of molecular and mor- and in expressing moderate geocauly.
phological divergence among them lead us to recognize Also within the Gymnomitriaceae, Poeltia, considered a
three separate families, as labeled in Fig. 1. The transfer subgenus of Marsupella by Schuster (1996, 2002), is resolved
of Stephaniella and Stephaniellidium Winkler ex Grolle to as sister to the Marsupella clade, and is herein recognized
Arnelliaceae by Crandall-Stotler et al. (2009) was based on as a monotypic genus. Although Poeltia is related to
the nested position of Stephaniella between Southbya and Marsupella, it is unique in the family in having strongly
Gongylanthus in de Roo et al. (2007). Personal study of the patent, squarrose leaves and gynoecia borne on short,
voucher specimen for the Stephaniella sequence confirmed, endogenous branches (Grolle 1966). Apomarsupella is nested
however, that it had been misidentified, and was actually within Gymnomitrion and is reduced to it. This reduction is
Gongylanthus liebmannianus (Lindenb. et Gottsche) Steph. morphologically supported by the absence of perianth and
(labeled FATOL774 in Table S1 and Fig. 1). In the absence perigynial development in Apomarsupella, as is also diag-
of molecular data, the phylogenetic affinities of Stephaniella nostic of Gymnomitrion (Schuster 1996; Vilnet et al. 2010).
and Stephaniellidium remain equivocal, but there is no evi- The inclusion of Herzogobryum in the Gymnomitriaceae has
dence to support their placement in either the Arnelliaceae been equivocal since H. teres (Carrington & Pearson) Grolle
or Southbyaceae. Morphologically they have more characters was resolved in the Cephaloziineae (Clade B) by Davis
in common with the Gymnomitriaceae, in which they were (2004) and Forrest et al. (2006). Although this species has
placed by Schuster (1984), than in other families represented recently been transferred to Syzygiella (Váňa et al. 2013)
in our dataset. In 2002 Schuster elevated his subfamily because of its morphological similarities to that genus, the
Stephanielloideae of the Gymnomitriaceae to family status, phylogenetic affinities of the generitype have not previ-
still noting, however, a close relationship to that family. ously been tested. In our study Herzogobryum vermiculare
The Gymnomitriaceae is modified to include Nardia and (Schiffn.) Grolle (= H. cucullatum (Herzog) Grolle, the type
Cryptocoleopsis, previously classified in the Solenostomataceae of Herzogobryum) and H. atrocapillum (Hook. f. & Taylor)
(Crandall-Stotler et al. 2009), and to exclude Herzogobryum Grolle were resolved in the Cephaloziineae, near the
and Nothogymnomitrion. In our study, the 19 accessions of Cephaloziellaceae, and the genus is, therefore, removed
Nardia form a lineage that is sister to a group that includes from Gymnomitriaceae; H. teres is nested in Syzygiella, con-
five genera: Poeltia, Marsupella, Cryptocoleopsis, Prasanthus, firming its transfer to that genus. The monospecific genus
Nothogymnomitrion was also resolved in the Cephaloziineae, that are bordered by a row of larger cells, whereas those
in the vicinity of the Anastrophyllaceae and Cephaloziellaceae, resolved in S. subg. Solenostoma do not express this char-
supporting its removal from the Gymnomitriaceae as well. acter, but whether this character is constantly diagnostic
Whether Acrolophozia R. M. Schust., Aponardia (R. M. Schust.) is equivocal since not all species with solenostomatoid
Váňa, Nanomarsupella R. M. Schust. ex A.Hagborg, L. perianths and bordered leaves have been tested. No
Söderstr. & von Konrat and Paramomitrion R. M. Schust. other morphological characters, including those used by
should remain in the family remains to be tested. Schuster Amakawa (1960, 1966); e.g. plant size, growth form, or
(2002) and Engel and Glenny (2008) placed Paramomitrion rhizoid distribution, separate infrageneric groups, nor
in the Jungermanniaceae near Eremonotus, but it is still do our results support the sectional classifications pro-
recognized in Gymnomitriaceae by Váňa et al. (2010b). posed by Amakawa (1960, 1966). In this regard, note that
Solenostomataceae was erected by Crandall-Stotler et al. S. hasskarlianum and S. rosulans, the only species of S. subg.
(2009) to accommodate Solenostoma and the presumably Plectocolea sect. Plectocolea included in our dataset are
related Arctoscyphus, Bragginsella, Cryptocolea, Cryptocoleopsis, resolved in separate clades of S. subg. Plectocolea, the
Cryptostipula, Diplocolea, Horikawaella, Nardia, and Scaphophyllum. former in S. subg. Plectocolea II and the latter in S. subg.
Because the family included Nardia, the type of the earlier Plectocolea I (Fig. 2). Since our current dataset does not
named Mesophyllaceae Heeg (= the superfluous Aliculariaceae include S. radicellosum Mitt., the type of S. subg. Plectocolea,
Warnst., published 11 yr later), Solenostomataceae was incor- we prefer to recognize both of these clades in S. subg.
rect at the time of its naming. With the transfer of Nardia to Plectocolea. Although Bakalin and Vilnet (2012) recover
the Gymnomitriaceae, however, it becomes a correct family clades that are comparable to our four subgeneric lineages,
name, the type of which is Solenostoma. Our results support their analysis differs in several regards, the most significant
the placement of S. caespiticium (Lindenb.) Steph. in the mono- of which is the resolution of S. hyalinum sister to the
typic Endogemmataceae by Vilnet et al. (2011), the transfer Solenostoma clade. In all of our analyses, accessions of
of Gottschelia patoniae Grolle, D. B. Schill & D. G. Long and S. hyalinum, including one from Russia, form a clade that
Scaphophyllum speciosum (Horik.) Inoue to Solenostoma by is nested in S. subg. Plectocolea, with which it shares peri-
Feldberg et al. (2009), and the removal of the genus anth and perigynial characters. Since there is little sup-
Gottschelia, based on G. schizopleura (Spruce) Grolle, from port for the clade designated by Bakalin and Vilnet (2012)
the Jungermanniineae to the Cephaloziineae. In our analy- as Solenostoma, but strong support for the clade that
ses, both species of Horikawaella are nested in Solenostoma includes both the Solenostoma and Plectocolea lineages, it
subg. Solenostoma, supporting the synonymy of that seems more reasonable to recognize Plectocolea as one
genus under Solenostoma. Cryptocoleopsis is transferred to of four subgenera of Solenostoma, as we propose, than
Gymnomitriaceae (as already discussed), Bragginsella has to modify its circumscription as a genus (Bakalin and
been placed in Lophocoleaceae (Glenny and Malcolm Vilnet 2012).
2005; Söderström et al. 2013) and Cryptostipula has been In addition to the transfer of Horikawaella subacuta and
synonymized with Hepatostolonophora J. J. Engel & R. M. H. grosse-verrucosa to S. subg. Solenostoma, our analyses sug-
Schust. in Lophocoleaceae (Engel 2011). The phylogenetic gest that S. abyssinicum is nested within S. confertissimum,
affinities of Arctoscyphus, Cryptocolea and Diplocolea remain and S. subellipticum (Lindb. ex Heeg) R. M. Schust. is nested
to be tested. within S. obovatum. We propose (below) that the two nested
Our analyses resolved four major clades within a broadly taxa be synonymized with their respective inclusive species.
defined, strongly supported Solenostoma (Solenostomataceae) Schuster (1988) postulated that Jungermannia subellipticum
clade (Fig. 2). Subgeneric status is proposed for each of these (Lindb. ex Heeg) Levier was likely an ecological variant
lineages as indicated in Fig. 2, namely, S. subg. Solenostoma, of J. obovata Nees, and Damsholt (2002) reduced it to
S. subg. Eucalyx, S. subg. Metasolenostoma, and S. subg. infrageneric status as J. obovata subsp. minor (Carrington)
Plectocolea. Relationships among these subgenera are not Damsh. (Basionym: Nardia obovata Lindb. var. minor
unambiguously resolved, and there are clear morphological Carrington. Brit. Hep.: 33. 1876). Since S. subellipticum
overlaps among them, especially as regards perigynial is nested within S. obovatum and not recovered as a
and perianth characters. Taxa in S. subg. Eucalyx and monophyletic group (see also Bakalin and Vilnet 2012),
S. subg. Plectocolea have plectocoleoid perianths that are this subspecies is not recognized. Váňa (1974a) noted
fusiform, comprised of elongate cells, and contracted but that S. abyssinicum is morphologically very similar to
not beaked at the mouth. Of species sampled, all resolved S. confertissimum and likely should be reduced to it, as
in S. subg. Eucalyx have well-developed perigynia that are supported by its nested position in that clade.
1–2 times the length of the perianths, which are barely That Solenostoma rufiflorum (Colenso) J. J. Engel is not
emergent from the bracts, whereas all resolved in S. subg. resolved as distinct from S. inundatum (Hook. f. & Taylor)
Plectocolea have perigynia that are shorter than the peri- Mitt. ex Steph. supports the reduction of this species to
anths, which generally extend well beyond the bracts. S. inundatum by Váňa (1975) based on morphology alone.
Whether species that were not included in our sampling Our evidence does not refute this reduction. At the same
will also sort into these two subgenera based on this dif- time, the two samples identified as S. totipapillosum (E. A.
ference in perigynial development remains to be tested. Hodgs.) R. M. Schust. are resolved as sister to the
Solenostomatoid perianths that are pyriform, comprised S. inundatum/S. rufiflorum clade. Although these results
of isodiametric cells in the upper and middle part, and indicate that S. inundatum and S. totipapillosum are closely
contracted into a distinct beak at the mouth, and low related, they do not necessarily contradict the species-level
to scarcely developed perigynia are characteristic of both recognition of S. totipapillosum. Cell walls are always
S. subg. Solenostoma and S. subg. Metastolenostoma. All coarsely papillose in S. totipapillosum, a relatively small
species resolved in S. subg. Metasolenostoma have leaves plant, as compared to usually smooth to finely striolate,
or with delicate, low papillae, in S. inundatum, which is Taxonomic Treatment

typically larger (Schuster 2002; Engel and Glenny 2008).
New Taxa—We here name and describe a new suborder,
However, the two taxa are quite similar in other aspects of
two new subfamilies, a new family, and two new subgenera.
their morphology, including leaf shape and stance, perianth/
perigynial structure, subglobose capsule shape and color, Myliineae J. J. Engel & Braggins ex Crand.-Stotl., Váňa,
capsule wall anatomy and spore wall architecture. Resolu- Stotler & J. J. Engel, subord. nov. —TYPE: Mylia Gray,
tion of their taxonomic status requires additional sampling of Nat. Arr. Brit. Pl. I: 693. 1821.
both populations and genomic regions. Likewise, determin-
Plants sparsely branched, with branches of the Frullania-
ing the significance of the apparent polyphyly of S. pyriflorum
type; leaves succubous, undivided, orbicular to oblong-
and S. appressifolium requires expanded population level
ovate, usually with cell wall grids and perforations, margins
studies within Solenostoma.
entire; underleaves lanceolate to subulate, entire, smaller
Solenostoma pusillum (C. E. O. Jensen) Steph. and
than the leaves; gyneocia and androecia on leading axes;
S. sphaerocarpum are not resolved as distinct with our
antheridial stalks uniseriate; sporophytes enclosed by a
data. According to Schuster (1969), Váňa (1974b) and
shoot calyptra and perianth; perianths laterally compressed
Damsholt (2002), there are only minor morphological dif-
above, with the mouth broad, bilabiate; capsules ovoid,
ferences between these species, the most notable being
with the wall 3- to 5-stratose; gemmae 1- or 2-celled, on
the frequent presence of small-leaved shoots, reddish brown
leaf tips.
pigmentation and a tendency for leaves to be reniform in
This suborder includes only the Myliaceae Schljakov,
the former, as compared to an absence of these features in
which was formerly included in the Jungermanniineae
the latter. However, these differences are not consistently
(Crandall-Stotler et al. 2009). Engel and Braggins (2005)
expressed, making separation of the two taxa also ques-
suggested that this suborder should be erected, but did
tionable on morphological grounds. Based on their similar
not validly publish it. They proposed the suborder to
morphologies and our molecular evidence, we propose the
include Mylia and Trabacellula Fulford, having segregated
reduction of S. pusillum to S. sphaerocarpum.
Mylia anomala (Hook.) Gray to the monotypic genus
Morphological Evolution—Despite ambiguities for some
Leiomylia J. J. Engel & Braggins, which they retained in the
characters at the base of the Jungermanniineae tree, we
Jungermanniaceae Rchb. of the suborder Jungermanniineae
were able to reconstruct ancestral conditions for a number
R. M. Schust. ex Stotler & Crand.-Stotl. As discussed pre-
of characters. Molecular analyses have repeatedly shown
viously, we retain Trabacellula in the Cephaloziaceae Mig.
that traditional hypotheses of morphological evolution in
(suborder Cephaloziineae Schljakov), and return Leiomylia
hepatics (e.g. Evans 1939; Schuster 1966) are incongruent
to the Myliaceae, suborder Myliineae. Even though our cir-
with phylogenetic relationships that have been resolved
cumscriptions of the Myliineae differ, we ascribe the name
across the phylum (Davis 2004; He-Nygrén et al. 2004,
to Engel and Braggins (2005).
2006; Heinrichs et al. 2005; Forrest et al. 2006). Maxi-
mum parsimony reconstructions of morphological char- Acrobolbaceae E. A. Hodgs. subfam. Saccogynidioideae
acter state evolution have further verified that many Crand.-Stotl., Váňa & Stotler, subfam. nov. —TYPE:
characters once considered taxonomically diagnostic are Saccogynidium Grolle, J. Hattori Bot. Lab. 23: 43.
homoplasious (Crandall-Stotler et al. 2005; Yu et al. 2013), 1960 [1961].
therein challenging the utility of morphological data in
Plants green to brown, irrregularly branched, with all
liverwort systematics. Our results support that view; we
branches of the Bazzania-type; leaves succubous, unlobed,
were not able to identify a single one of the 14 charac-
with the apices rounded or emarginate; leaf cells papillose,
ters we scored as having evolved without homoplasy.
with the oil bodies small, finely granulate, ellipsoidal to
It is impossible to count the precise number of state
fusiform, eight to 20 per cell; underleaves large, deeply
transitions for any character without exhaustive sam-
bilobed, with margins entire; androecia on very short
pling at the species level, but even with our incomplete
catkin-like ventral branches; gynoecia also on very short
sampling we demonstrated that most character states
ventral branches; perianths lacking; marsupia hollow, of the
evolved 4–10 times across the Jungermanniiineae tree as
Calypogeia-type; capsule ellipsoidal, beaked, with the wall
we reconstructed it.
4- to 6-layered; asexual reproductive structures lacking.
We did not quantify correlations among state transi-
Saccogynidium, the only genus recognized in this sub-
tions in this study, but examination of our reconstruc-
family, was previously classified in the Geocalycaceae
tions indicates largely independent patterns of homoplasy
H. Klinggr. (Crandall-Stotler et al. 2009). Its transfer to the
across characters. The correlation between the position
Acrobolbaceae is supported not only by our analysis, but
of gynoecia and androecia is not surprising, and pos-
also by its possession of a thick multistratose capsule
sibly adaptive. When gametangia are produced on short
wall that is structurally different from the bistratose wall
branches, they are ventrally exposed, being overlaid by
characteristic of Geocalycaceae, and by its highly papillose
the stem and/or leaves of the parent plant, while those
leaf cell walls. The subfamily differs from other sub-
on leading stems are dorsally exposed. Having archegonia
families of Acrobolbaceae by the presence of large, bilobed
and antheridia in the same plane would presumably
underleaves and androecia and gynoecia borne on very
enhance fertilization. Future work on morphological evo-
short ventral branches.
lution in this group should also include assessments
of habitat shifts in order to assess possible adaptive pro- Acrobolbaceae E. A. Hodgs. subfam. Austrolophozioideae
cesses underlying trait homoplasy. Transitions from dioecy R. M. Schust. ex Crand.-Stotl., Váňa & Stotler, subfam.
to monoecy might involve polyploidization, but too few nov. —TYPE: Austrolophozia R. M. Schust., J. Hattori
data are available at present to test that hypothesis. Bot. Lab. 26: 282. 1963.
Plants with branches only of the lateral Plagiochila type; these subgenera, two new subgenera are established to
leaves succubous, spinose-dentate to lobulate, often with accomodate these lineages. Of the species treated by us,
reddish to brown pigments, with the cell walls smooth; oil Solenostoma subg. Metasolenostoma includes S. gracillimum,
bodies large and few in number; rhizoids in dense mats S. fusiforme (Steph.), S. rubrum (Gottsche ex Underw.)
that often form decurrent bundles on the ventral surface of R. M. Schust., and S. handelii (Schiffn.) Müll. Frib., species
the stem; androecia on leading stems, with male bracts previously classified in S. subgenus Solenostoma.
little modified from the leaves; gynoecia terminal on lead-
Solenostoma Mitt. subg. Eucalyx (Lindb.) Váňa, Crand.-
ing stems; perianths vestigial or lacking; marsupium of
Stotl. & Stotler, stat. et comb. nov.; Basionym: Nardia
either the solid Tylimanthus-type (Austrolophozia) or hollow
sect. Eucalyx Lindb., Not. Sällsk. Fauna Fl. Fennica
Calypogeia-type (Goebelobryum); capsule ellipsoidal, not
Förh. 13: 369. 1874. Nardia subg. Eucalyx (Lindb.)
beaked,with the wall 4- or 5-layered; gemmae, if present,
Lindb., Musci Scand.: 8. 1879. —TYPE: Solenostoma
2-celled from reduced leaves.
obovatum (Nees) C. Massal., Epat. Erb. Critt. Ital.: 17
This subfamily includes Austrolophozia R. M. Schust. and
[=Accad. Sci. Med. Nat. Ferrara 1903: 199]. 1903.
Goebelobryum Grolle. Although Schuster (1980: 543; 2001:
Basionym: Jungermannia obovata Nees, Naturgesch. Eur.
107) included “subf. Austrolophozioideae Schust.” in treat-
Leberm. 1: 279, 332. 1833.
ments of the Acrobolbaceae, he failed to provide a Latin
description or diagnosis for the subfamily, rendering it Plants often with reddish-purple pigmentation on stems
invalid (ICN Art. 39.1). We here validate this name with and leaf bases; stoloniferous, flagelliform branches common,
an English description (ICN Art. 39.2), and circumscribe of the Plagiochila-type; rhizoids purple; perianth of the
it to include Austrolophozia and Goebelobryum. Plectocolea-type, 4- to 6- plicate, with the cells elongated and
the mouth lobulate, scarcely emergent from the bracts;
Notoscyphaceae Crand.-Stotl., Váňa & Stotler, fam. et stat.
perigynium terete, bearing the bracts on its surface, exceed-
nov. —TYPE: Notoscyphus Mitt. in B. C. Seemann, Fl.
ing the perianth in length.
Vitiensis: 407. 1873. [The date of this publication is
The name Eucalyx was first proposed as a subgenus of
often cited as 1871 and on p. 405 of our copy is printed
Nardia Gray nom. cons. by Lindberg (Bot. Not. 1872: 167.
“published June 1, 1871,” but Grolle (1983) pointed out
1872) but was not validly published because it was not
that it was not actually issued until Feb 1873.]
described. In 1874 it was described as Nardia sect. Eucalyx
This monogeneric family is here validated by the diagno- Lindb. in the publication “Manipulus Muscorum Secundus”
sis of Jungermanniaceae Rchb. subfamily Notoscyphoideae (Lindberg 1874), which validated the name. The Lindberg
R. M. Schust., Trans. Brit. Bryol. Soc. 6: 94–95. 1970. Note use of this epithet as a subgenus of Nardia (stat. nov.) is
that in accordance with Article 49.2 (McNeill et al. 2012) therefore valid in any of the Lindberg publications after
parenthetical authors are not cited for suprageneric names, that date, the first of which is Nardia A. Eucalyx (Lindb.)
even if validly published by reference. Lindb., Musci Scand.: 8. (8 Nov.- 30 Dec. 1879) according
Schuster (2002: 447) remarked that following intensive to Isoviita (1980: 72). Of the species resolved in the S. subg.
restudy of Notoscyphus, he thought it ought to be placed Eucalyx clade, namely, S. obovatum, S. flagellatum (Hatt.)
in its own family although he did not do so. He stated that Váňa & Long, and S. schusterianum (J. D. Godfrey & G.
the affinities of the genus were probably with Geocalyx Godfrey) Váňa, Hentschel & Heinrichs, only S. obovata
Nees and that it should go into the Geocalycaceae or else ( Jungermannia obovata Nees Nardia obovata (Nees)
be a family adjoining the Geocalycaceae. Our molecular Lindb.) was included in Lindberg’s original treatment
data support his view that it should be placed in its own (1872) and in the validly published section (1874); we,
family, but do not support a close relationship with Geocalyx. therefore, designate this as the type of the subgenus. Other
species placed by Lindberg in subgenus Eucalyx have been
Solenostoma Mitt. subg. Metasolenostoma Váňa, Crand.-
resolved in S. subg. Plectocolea. Although both S. subg.
Stotl. & Stotler, subg. nov. —TYPE: Solenostoma
Eucalyx and S. subg. Plectocolea have perigynia and
gracillimum (Sm.) R. M. Schust., Hepat. Anthocerotae
Plectocolea-type perianths, only those species resolved in
N. Amer. 2: 972. 1969. Basionym: Jungermannia gracillima
the S. subg. Eucalyx lineage have frequent flagelliform branches
Sm. in Sowerby, Engl. Bot. 32. pl. 2238. 1811.
and very long perigynia, with scarcely emergent perianths.
Plants with leaves bordered by one row of larger cells New Combinations—Six new combinations at the spe-
(only in well-developed plants in S. gracillimum and cies level are provided based on results of this study.
S. fusiforme (Steph.) R. M. Schust.); perianth of the Solenostoma- Grolle and Váňa (1992) named Jungermannia conchata
type, pyriform, suddenly contracted to a beaked mouth Grolle & Váňa based on a collection of small, entire-leafed
and made up of isodiametric (not elongated) cells; peri- plants from 4100 m elevation in Nepal. At the time of
gynium scarcely developed or less than 0.2 times the length its naming, neither males nor sporophytes were known.
of the perianth. In our analyses this taxon is resolved in Cephalozia, as
Previous classifications of Solenostoma Mitt. often recog- also supported by new findings of male plants (Váňa,
nized two subgenera, S. subg. Solenostoma and S. subg. unpubl. data).
Plectocolea Mitt. (Schuster 2002, Hentschel et al. 2007),
Cephalozia conchata (Grolle & Váňa) Váňa, comb. nov.;
based primarily on the form of the perianth cells and
Basionym: Jungermannia conchata Grolle & Váňa, Fragm.
absence or presence of a well-developed perigynium.
Flor. Geobot. 37: 3. 1992.
However, in our analyses four well-supported, mono-
phyletic lineages are resolved within Solenostoma, with The genus Apomarsupella R.M. Schust. was established
S. subg. Solenostoma and S. subg. Plectocolea, as previously by Schuster in 1996 to include four taxa that resemble
circumscribed, paraphyletic. In addition to recognizing Marsupella in vegetative characters, but Gymnomitrion in
gynoecial characters, namely, A. revoluta (Nees) R.M. Schust. Solenostoma pusillum (C. E. O. Jensen) Steph., Sp. Hepat. 6:
(the generitype), with two subspecies, A. africana (Steph. ex 83, 1917. Basionym: Aplozia pusilla C. E. O. Jensen, Rev.
Bonner) R.M. Schust., and A. rubida (Mitt.) R.M. Schust. Bryol. 39: 92, 1912, syn. nov. = SOLENOSTOMA SPHAEROCARPUM
In 1999, Váňa transferred two additional species from (Hook.) Steph., Bull. Herb. Boissier, sér. 2, 1(5): 499 [Sp.
Marsupella, A. crystallocaulon (Grolle) Váňa and A. verrucosa Hepat. 2: 61], 1901. Basionym: Jungermannia sphaerocarpa
(Nichols.) Váňa. Since all species of Apomarsupella, includ- Hook., Brit. Jungerm. Table 74, 1815.
ing the generitype, are nested in Gymnomitrion, we place
Solenostoma subellipticum (Lindb. ex Heeg) R. M. Schust.,
Apomarsupella in synonymy of Gymnomitrion (see below),
Hepat. Anthocerotae N. Amer. 2: 1021. 1969. Basionym:
necessitating the following new combinations:
Nardia subelliptica Lindb. ex Heeg, Verh. K. K. Zool.-
Gymnomitrion crystallocaulon (Grolle) Váňa, Crand.-Stotl. & Bot. Ges. Wien 43: 69. 1893, syn. nov. = SOLENOSTOMA
Stotler, comb. nov.; Basionym: Marsupella crystallocaulon OBOVATUM (Nees) C. Massal., Epat. Erb. Critt. Ital.: 17.
Grolle, Khumbu Himal: 281. 1966. [= Accad. Sci. Med. Nat. Ferrara 1903: 199.] 1903.
Basionym: Jungermannia obovata Nees, Naturgesch. Eur.
Gymnomitrion revolutum (Nees) H. Philib. subsp.
Leberm. 1: 332. 1833. This agrees with the concept
novoguineanensis (R. M. Schust.) Váňa, Crand.-Stotl. &
of Damsholt (2002: 230) who reduced J. subelliptica to
Stotler, comb. nov.; Basionym: Apomarsupella revoluta
J. obovata subsp. minor Carrington.
(Nees) R. M. Schust. subsp. novoguineanensis R. M.
Schust., J. Hattori Bot. Lab. 80: 90. 1996. New Circumscriptions—New circumscriptions for seven
families are required based on results of this study.
Gymnomitrion rubidum (Mitt.) Váňa, Crand.-Stotl. &
The following three families were all validly published
Stotler, comb. nov.; Basionym: Jungermannia rubida Mitt.,
but were incorrect when named because they included
J. Proc. Linn. Soc., Bot. 5: 90. 1860.
the type genus of an earlier published family name. How-
Both of the named species of Horikawaella S. Hatt. & ever, by exclusion of that genus from each of these families,
Amakawa are nested with strong support within Solenostoma they become the correct family name with new circum-
subg. Solenostoma. Placement of the two genera in synonymy scriptions. In accordance with Article 49.2 (McNeill et al.
(see below) is supported, not only by our analysis, but 2012), parenthetical authors are not cited for suprageneric
also by their similar morphologies, with acute, rather than names. HARPANTHACEAE Arnell in Holmberg, Skand. Fl. 2(a):
rounded, leaf apices, being the major distinguishing charac- 147. 1928. Type: Harpanthus Nees, Naturg. Eur. Leberm.
ter of the Horikawaella species, which are herein transferred 2: 351. 1836. We here recognize this family as mono-
to Solenostoma. generic. When published, this family name was incorrect
because it included Geocalyx, the type genus of the older
Solenostoma subacutum (Herzog) Váňa, Crand.-Stotl. &
Geocalycaceae. With the removal here of Geocalyx from
Stotler, comb. nov.; Basionym: Anastrophyllum subacutum
Harpanthaceae and its retention in the Geocalycaceae, this
Herzog, Ann. Bryol. 12: 75. 1939.
family becomes correct under Article 52.3 (McNeill et al.
Solenostoma grosse-verrucosum (Amakawa & S. Hatt.) 2012; ex. 18). SACCOGYNACEAE Heeg, Verh. Zool.-Bot. Ges.
Váňa, Crand.-Stotl. & Stotler, comb. nov.; Basionym: Wien 41: 571. 1891. Type: Saccogyna Dumort., Comment.
Horikawaella grosse-verrucosa Amakawa & S. Hatt. in Bot.: 113. 1822. We here recognize this family as being
Hara, Bull. Univ. Mus. Univ. Tokyo 8 [Fl. East Himalaya, monogeneric. When published, this family name was incor-
third Rep.]: 216. 1975. rect because it included Geocalyx, the type genus of the
older Geocalycaceae. As with the Harpanthaceae, the
New Synonyms—The following new synonyms are pro-
removal here of Geocalyx from Saccogynaceae and its
vided based on this study:
retention in the Geocalycaceae, makes this family name
Delavayellaceae R. M. Schust., Bryologist 64: 202. 1961, syn. correct under Article 52.3 (McNeill et al. 2012; ex. 18).
nov. = JUNGERMANNIACEAE Rchb., Bot. Damen: 256. 1828. SOLENOSTOMATACEAE Stotler & Crand.-Stotl., Edinburgh
J. Bot. 66: 190. 2009. Type: Solenostoma Mitt., J. Proc. Linn.
Hygrobiellaceae Konstant. & Vilnet, Phytotaxa 167(2): 217.
Soc., Bot. 8: 51. 1865. The genera that now remain in
2014 (Arctoa 18: 65. 2009[2010], nom. inval.), syn.
Solenostomataceae are Solenostoma Mitt. [including Plectocolea
nov. = ANTHELIACEAE R. M. Schust., J. Hattori Bot. Lab.
(Mitt.) Mitt.], Arctoscyphus Hässel, Cryptocolea R. M.Schust.,
26: 236. 1963.
and Diplocolea Amak. When published, this family name
Apomarsupella R. M. Schust., J. Hattori Bot. Lab. 80: 79. 1996, was incorrect because it included Nardia, which is the type
syn. nov. = GYMNOMITRION Corda in Opiz, Naturalientausch genus of the earlier Mesophyllaceae Heeg (Mesophylla
12: 651. 1829. Nardia). With transfer of Nardia to the older Gymnomitriaceae,
this family becomes correct under Article 52.3 (McNeill et al.
Horikawaella S. Hatt. & Amakawa, Misc. Bryol. Licheol. 5:
2012; ex. 18). In addition to the removal of Nardia here, the
164. 1971, syn. nov. = SOLENOSTOMA Mitt., J. Proc. Linn.
following genera formerly placed here are transferred or
Soc., Bot. 8: 51. 1865.
reduced as follows: Bragginsella R. M.Schust. transferred to
Solenostoma abyssinicum (Nees) Steph., Bull. Herb. Boissier, Lophocoleaceae (Söderström et al. 2013); Cryptocoleopsis
sér. 2, 1(5): 491 [=Sp. Hepat. 2: 53]. 1901. Basionym: Amak. transferred to Gymnomitriaceae (done here) ;
Jungermannia abyssinica Nees in Gottsche, Lindenberg & Cryptostipula R. M. Schust. reduced to Hepatostolonophora
Nees, Syn. Hepat.: 93. 1844, syn. nov. = SOLENOSTOMA J. J. Engel & R. M. Schust. (Lophocoleaceae) (Engel 2011:
CONFERTISSIMUM (Nees) Schljakov, Pečen. Mhi Severa 402); Horikawaella S. Hatt. & Amakawa reduced to
SSSR 4: 51. 1981. Basionym: Jungermannia confertissima Solenostoma Mitt. (done here), and Scaphophyllum Inoue
Nees, Naturgesch. Eur. Leberm. 1: 277, 291. 1833. reduced to Solenostoma Mitt. (Feldberg et al. 2009).
Circumscriptions of the following four families have also Engel, J. J. and D. Glenny. 2008. A flora of the liverworts and hornworts
been modified by having genera added or genera trans- of New Zealand, Vol. 1. Monographs in Systematic Botany from the
Missouri Botanical Garden 110: 1–897.
ferred to other families without nomenclatural conse- Engel, J. J. and D. Glenny. 2012. Austral Hepaticae 48. Goebelobryum
quences: ANTHELIACEAE, ARNELLIACEAE, GYMNOMITRIACEAE and Grolle (Acrobolbaceae). Nova Hedwigia 95: 319–336.
JUNGERMANNIACEAE. Full details of these changes and their Engel, J. J. and R. Grolle. 1971. Marsupidium in South America. The
justifications are provided in the Discussion section. Journal of the Hattori Botanical Laboratory 34: 437–444.
Evans, A. W. 1939. The classification of the Hepaticae. Botanical Review
5: 49–96.
Acknowledgments. This research was supported by a multi- Feldberg, K., J. Hentschel, A. Bombosch, D. G. Long, J. Váňa, and J.
institutional, collaborative grant from the Assembling the Tree of Heinrichs. 2009. Transfer of Gottschelia grollei, G. patoniae and
Life program at NSF. The following NSF grants supported this work: Scaphophyllum speciosum to Solenostoma based on chloroplast DNA
EF-0531730-002 (to AJS) EF-0531680 (to MvK and JJE), EF-0531750 rbcL sequences. Plant Systematics and Evolution 280: 243–250.
(to BC-S & RES). Numerous collectors supplied collections for the Finet, C., R. E. Timme, C. F. Delwiche, and F. Marlétaz. 2010. Multi-
phylogenetic analyses and all such collaborators are gratefully acknowl- gene phylogeny of the green lineage reveals the origin and diver-
edged. This work would be impossible without assistance from many sification of land plants. Current Biology 20: 2217–2222.
talented collectors. We thank Sandra Boles, Lisa Karst, Nicolas Devos, Forrest, L. L. and B. J. Crandall-Stotler. 2004. A phylogeny of the
and Laura Briscoe for technical assistance. simple thalloid liverworts (Jungermanniopsida, Metzgeriidae) as
inferred from five chloroplast genes. Pp. 119–140 in Molecular
systematics of bryophytes: progress, problems & perspectives eds.
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to the genera of antipodal Hepaticae with special reference to cal rearrangements of Solenostomataceae (Marchantiophyta) with
description of new family Endogemmataceae based on trnL-F DNA KF943135, KF942720,—; Gottschelia schizopleura: L1206, Malaysia:
analysis. Folia Cryptogamica Estonica 48: 125–133. Pahang, D.G. Long 36922 (E),—, KF851920,—, KF852362, KF851453,—,
Vilnet, A. A., N. A. Konstantinova, and A. V. Troitsky. 2012. Molecular KJ802085, KF852521, KF851624, KJ802021; Gymnocolea inflata: FATOL535,
phylogeny and systematics of the suborder Cephaloziineae with Svalbard, N.A. Konstantinova & A.N. Savchenko 118-1-04 (F), KF852019,
special attention to the family Cephaloziaceae s. l. (Jungermanniales, KF851861, KF852141, KF852306, KF851412, KJ802048, KJ802076, KF852452,
Marchantiophyta). Arctoa 21: 113–132. KF851553, KJ802022. Antheliaceae: “Pseudolepicolea” trollii subsp. andoi:
Vilnet, A. A., I. A. Milyutina, N. A. Konstantinova, M. S. Ignatov, and E53, China: Yunnan, D.G.Long 33719 (E),—,—, KF943263, KF943547,
A. V. Troitsky. 2007. Phylogeny of the genus Lophozia (Dumort.) KF943399,—,—, KF943117, KF942703,—; Anthelia julacea: E67, UK:
Dumort. s. s. inferred from nuclear and chloroplast sequences Scotland, D.G. Long 31292 (E),—, KF851830, KF852121, KF852280,
ITS1–2 and TRNL-F. Russian Journal of Genetics 43: 1306–1313. KF851384, KJ802043, KJ802070, KF852428, KF851527, KJ802014;
Wilson, R., S. R. Gradstein, H. Schneider, and J. Heinrichs. 2007. L1404, USA: Alaska, B. Shaw F952/5 (DUKE),—, KF942570, KF943340,
Unraveling the phylogeny of Lejeuneaceae (Jungermaniopsida): KF943594, KF943488, KF942894, KF943055, KF943212, KF942772,
evidence for four main lineages. Molecular Phylogenetics and Evolu- KF942677; Hygrobiella laxifolia: FATOL512, Russia, N.A. Konstantinova
tion 453: 270–282. 31-2-03 (F),—, KF942489, KF943285,—, KF943420,—,—,—,—,—; FATOL683,
Wodniok, S., H. Brinkmann, G. Glöckner, A. J. Heidel, H. Philippe, M. Russia, V. A. Bakalin s.n. (F),—,—, KF852154, KF852320, KF851422,—,—,
Melkonian, and B. Becker. 2011. Origin of land plants: do conju- KF852462, KF851560,—. Arnelliaceae: Arnellia fennica: L1498, Finland,
gating green algae hold the key? BMC Evolutionary Biology 11: 104. B. Shaw 1250 (DUKE), KF852098, KF851969, KF852251, KF852402,
Yatsentyuk, S. P., N. A. Konstantinova, M. S. Ignatov, J. Hyvönen, and KF851493, KJ802058, KJ802096,—, KF851663,—. Balantiopsidaceae:
A. V. Troitsky. 2004. On phylogeny of Lophoziaceae and related Acroscyphella phoenicorhiza: FATOL865, New Zealand, J.E. Braggins &
families (Hepaticae, Jungermanniales) based on TrnL-TrnF intron- M.A.M. Renner 4174 (F), KF852039, KF851900, KF852165, KF852339,
spacer sequences of chloroplast DNA. Pp. 151–167 in Molecular —,—,—,—,—,—; Balantiopsis diplophylla: IBC7, Australia, H. Streimann
systematics of bryophytes: progress, problems & perspectives eds. 59554 (NY),—,, KF852201, KF852348,—,—,—,—, KF851610,—; B. splendens:
B. Goffinet, V. C. Hollowell, and R. E. Magill. Monographs in FATOL783, Chile, J.J. Engel 25363 (F),—, KF851887,—, KF852330,—,—,—,
Systematic Botany from the Missouri Botanical Garden. Vol. 98. —,—,—; Isotachis lyallii: IBC30, New Zealand, J.J. Engel 21825 (F),
St. Louis: Missouri Botanical Garden. AY607905, AY607952, AY608010, KF852343, AY608073, AY608178,
Yu, Y., T. Pócs, A. Schäfer-Verwimp, J. Heinrichs, R.-L. Zhu, and H. AY608130, KF852497, KF851600,—. Blepharidophyllaceae: Blepharidophyllum
Schneider. 2013. Evidence for rampant homoplasy in the phy- vertebrale: L1209, New Zealand, J.J. Engel & M. von Konrat s.n. (F),
logeny of the epiphyllous liverwort genus Cololejeuenea (Lejeuneaceae). KF852073, KF851921, KF852217, KF852363, KF851454,—, KJ802086,
Systematic Botany 38: 553–563. KF852522, KF851625,—; Clandarium gottscheanum: FATOL777, Chile, J.J.
Zwickl, D. J. 2006. Genetic algorithm approaches for the phylogenetic Engel 26229 (F), KF852032, KF851885,, KF852329, KF851425,—,—,
analysis of large biological sequence datasets under the maxi- KF852472, KF851564,—; C. xiphophyllum: L1373, Australia: Tasmania,
mum likelihood criterion. Ph.D. dissertation. Austin: University B. Shaw 6433 (DUKE), KF942448,—, KF943334, KF943590, KF943482,
of Texas. —,—, KF943206, KF942767,—. Calypogeiaceae: Eocalypogeia quelpartensis:
gb242, Japan, T. Furuki 20233 (HIRO),—,—,—, AB476566, AB476605,
—,—,—,—,—; Calypogeia muelleriana: E77, UK: Scotland, D.G. Long
Appendix 1. List of sampled taxa by family, voucher information, 31227 (E),—, KF942466, KF943266,—,—, KF942826, KF942955,
and GenBank Accession numbers (atpB, psbA, psbT, rbcL, rps4, trnG, trnL, KF943121, KF942707, KF942633; IBC11, USA, E.C. Davis 130 (DUKE),
nad1, rps3, rpb2, respectively).—: missing data. Voucher numbers begin- JF315941, AY6079311, JF316196, JF316291, AY6080521, AY608169,
ning with “gb” designate sequences downloaded from GenBank. AY608121, AY607870, AY608169, KJ802016; Metacalypogeia alternifolia:
E30, Bhutan, D.G. Long 28712 (E),—, JF513396, JF513412, JF513455,
Acrobolbaceae: Acrobolbus ochrophyllus: FATOL118, Chile, J.J. Engel JF513474, KJ802040, KJ802068, KF852423, KF851522,—; Mizutania
26633 (F), KF852005, KF851834, KJ451735, KF852285, KF851387, riccardioides: L1510, Malaysia: Pahang, D.G. Long, D.S. Rycroft &
KJ802044, KJ802071, KF852430, KF851532,—; Austrolophozia paradoxa: B. O’Shea 36926 (E),—, KF851971, KF852254,—,—,—,—, KF852568,
FATOL785, New Zealand, J.J. Engel & M. von Konrat 28538 (F), KF851664,—; Mnioloma fuscum: L1516, Fiji, J.E. Braggins et al. 16. IV. 2008
KF852035, KF851889,—,—, KF851427,—,—, KF852475, KF851567,—; (NSW),—,—, KF852256, KF852404, KF851498,—, KJ802100, KF852569,
Goebelobryum unguiculatum: L1274, New Zealand: North Island, Y. Qiu, KF851665, KJ802029. Cephaloziaceae: Alobiellopsis parvifolius: L1532,
M.A.M. Renner & R.M. Schuster NZ-03054 (AK), KF852077, JF513406,—, Japan: Honshu, B. Shaw 8240 (DUKE), KF852105, KF851979, KF852262,
JF513466, JF513487,—,—, JF513386, KF851638,—; Lethocolea glossophylla: KF852411, KF851505, KJ802064, KJ802104, KF852576, KF851672,—;
IBC41, Ecuador, E.C. Davis 259 (DUKE), KF852059, AY607963, KF852195, Cephalozia (Jungermannia) conchata: L1198, China: Yunnan, D.G. Long
DQ439692, AY608084, KJ802051, KJ802080, KF852498, KF851602, KJ802025; 34888 (E),—, KF942541, KF943323, KF943576, KF943462, KF942883,
Saccogynidium australe: FATOL135, New Zealand, J.J. Engel, M. von KF943028, KF943183, KF942748,—; L1526, China: Yunnan, D. G. Long
Konrat & J.E. Braggins 24166 (F),—, KF942473, KF943272,—, KF943404, & J. Shevock 37157 (DUKE),—, KF942623, KF943389,—,—, KF942942,
KF942832, KF942960,—, KF942712,—; S. decurvum: L1368, Australia: KF943107,—, KF942814,—; Cephalozia bicuspidata: FATOL587, Russia,
Tasmania, B. Shaw 6410 (DUKE), KF852088,—, KF852238, KF852389, N.A. Konstantinova s.n. (F),—, KF942504, KF943295, KF943563,
KF851479,—,—, KF852555, KF851652,—; Tylimanthus laxus: IBC44, Costa KF943435, KF942855, KF942986, KF943151,—,—; L1240, Ireland, R.E.
Rica, G. Dauphin 2920 (NY),—,—,—,—,—,—,—, KF852499, KF851603, Stotler & B.J. Crandall.-Stotler s.n. (ABSH),—, JF513405, KF852220,
KJ802035. Adelanthaceae: Cuspidatula monodon: L1210, New Zealand, J.J. JF513465, JF513486,—, KJ802090, JF513385, KF851633,—; Odontoschisma
Engel & M. von Konrat 28380 (F),—, KF851922,—, KF852364, KF851455,—, sphagni: L1494, Canada: Newfoundland, B. Shaw 6684 (DUKE),—,
KJ802087, KF852523, KF851626,—; Syzygiella autumnalis: L1300, USA: KF942613, KF943377,—, KF943531, KF942930, KF943097,—,—,—.
Vermont, B. Shaw 6969 (DUKE), KF852084, KF851944, KF852226, Cephaloziellaceae: Cephaloziella divaricata: L1426, Czech Republic,
KF852382, KF851473, KJ802054, KJ802092, KF852544, KF851646, KJ802034; P. Sova s.n. (DUKE), KF852095, KF851965, KF852248, KF852399,
S. colorata: L1371, Australia: Tasmania, B. Shaw 6533 (DUKE), KF942447, KF851489, KJ802056, KJ802094, KF852565, KF851660, KJ802017; C. hirta:
KF942564, KF943333, KF943589, KF943481, KF942888, KF943049, IBC14, Australia, H. Streimann 59793 (NY), KF942433,—, JF513433,—,—,
KF943205, KF942766, KF942671; S. nipponica: E22, Nepal, D.G. Long KF942864, KJ802079,—, KF942739, KF942650; Herzogobryum atrocapillum:
30702 (E),—, KF942461, KF943261,—, KF943397, KF942822, KF942950, L1497, Marion Island, R. Ochyra 1402/01 (DUKE),—,—, KF943380,—,
KF943116,—, KF942631; S. sonderi: FATOL108, Gough Island, N.J.M. KF943534, KF942933,—,—,—,—; H. vermiculare: L1351, Iles Kerguelen,
Gremmen 2000-0374 (F),—, KF942472, KF943270,—, KF943402, R. Ochyra 1011/06 (DUKE),—, KF942562, KF943332, KF943587, KF943479,
KF942831,—, KF943124, KF942710,—; S. tasmanica: FATOL02, New KF942886, KF943047,—, KF942764, KF942669; Nothogymnomitrion erosum:
Zealand: South Auckland Province, J.J. Engel, M. von Konrat, & J.E. FATOL177, New Zealand, J.J. Engel, M. von Konrat & J.E. Braggins 24569
Braggins 23900 (F),—, KF942471, KF943269,—,—,—,—,—,—,—; S. teres: (F),—, KF942474,—, KF943548, KF943395,—,—,—, KF942713,—; gb046,
FATOL39, New Zealand, J.J. Engel, M. von Konrat & J.E. Braggins 24539 Australia, H. Streimann 53475 (JE),—, GQ900110,—, GQ900318,—,—,
(F), KF852010, KF851847, KF852129, KF852296, KF851398,—,—, GQ900216,—,—,—; L1235, New Zealand, J.J. Engel & M. von Konrat s.n.
KF852444,—,—; L1225, New Zealand, J.J. Engel & M. von Konrat s.n. (F),—, KF851928,—, KF852369, KF851461,—, KJ802089, KF852528,
(F),—, KF942549,—, KF943582, KF943469,—, KF943035, KF943191, KF851632,—. Endogemmataceae: Endogemma caespiticium: FATOL505,
KF942753, KF942665. Anastrophyllaceae: Anastrophyllopsis subcomplicata: Russia, N.A. Konstantinova 213-02 (F),—, KF942486,—,—, KF943417,—,
FATOL483, Australia, D. A. Meagher & N. Klazenga 07-142 (F), KF942423, KF942970,—,—,—; gb322, Russia: Buryatiya, N.A. Konstantinova 101-1-01
KF942484, KF943280, KF943550, KF943414, KF942838, KF942968, (KPABG),—,—,—,—,—,—, GU220585,—,—,—; L1413, USA: Alaska,
B. Shaw 7211 (DUKE), KF942450, KF942578, KF943346,—, KF943496,—, KF942742, KF942652; L1075, USA: Alaska, B. Shaw 7210 (DUKE),
KF943062, KF943220,—,—; L1443, Czech Republic, P. Sova s.n. (DUKE),—, KF942437, KF942520, KF943312,—, KF943449, KF942874, KF943012,
KF942603, KF943368,—, KF943521, KF942924, KF943087,—,—, KF942699; KF943167,—, KF942653; L1418, Norway, P. Sova s.n. (DUKE),—, KF942582,
L1444, Austria, P. Sova s.n. (DUKE),—, KF942604, KF943369,—, KF943522, KF943349,—, KF943500, KF942905, KF943066, KF943224, KF942780,
KF942925, KF943088, KF943245,—, KF942700; L1445, Norway, P. Sova KF942683; L1419, Czech Republic, P. Sova s.n. (DUKE),—, KF942583,—,
s.n. (DUKE),—, KF942605, KF943370,—, KF943523,—, KF943089, KF943246, KF943602, KF943501, KF942906, KF943067, KF943225, KF942781, KF942684;
—,—. Geocalycaceae: Geocalyx caledonicus: FATOL132, New Zealand, N. insecta: L1284, Russia: Murmanskaya Oblast, J. Kučera 11367 (CBFS),—,
J.J. Engel & M. von Konrat 23659 (F),—,—, KF943271,—, KF943403,—,—, KF942558, KF943328,—, KF943475,—, KF943043, KF943200, KF942761,—;
KF943125, KF942711,—; L1233, New Zealand, J.J. Engel & M. von N. japonica: L1509, USA: Alaska, W.B. Schofield (DUKE),—, KF942620,
Konrat 28342 (F),—, KF942551,—, KF943584, KF943470,—,—, KF943193, KF943386,—, KF943539, KF942939, KF943104,—,—,—; N. lescurii: L1077,
KF942755,—; G. graveolens: E13, China: Yunnan, D. G. Long 34828 USA: North Carolina, B. Shaw 12987 (DUKE),—, KF942522,—,—,
(E), KF851995, KF851821, KF852113, KF852271, KF851378, KJ802038, KF943451, KF942875, KF943013,—,—, KF942654; L1363, USA: North
KJ802067, KF852417, KF851516, KJ802019. Gymnomitriaceae: Cryptocoleopsis Carolina, B. Shaw 7209 (DUKE),—, KF942563,—, KF943588, KF943480,
imbricata: L1503, Russia, V.A. Bakalin s.n. (DUKE),—,—,—,—, KF851495, KF942887, KF943048, KF943204, KF942765, KF942670; N. scalaris: E101,
KJ802059, KJ802098,—,—,—; Gymnomitrion (Apomarsupella) africanum: Ireland, D.G. Long 35628 (E),—, KF942458, KF943259,—,—, KF942819,
L1502, Rwanda, T. Pócs 8210 (F),—, KF942616, KF943382,—,—, KF942935, KF942948, KF943113,—, KF942629; FATOL540, Russia, N.A. Konstantinova &
KF943101,—,—,—; Gymnomitrion (Apomarsupella) revolutum: L1135, China: A.N. Savchenko 1/3/2002 (F), KF852020, JF513400, JF513419, JF513462,
Yunnan, B. Shaw 5764 (DUKE), KF942441, KF942537, KF943320,—, JF513481, KJ802049, KJ802077, JF513354, KF851554,—; L1428, Austria,
KF943457,—, KF943024, KF943178, KF942743, KF942659; Gymnomitrion P. Sova s.n. (DUKE),—, KF942590, KF943355, KF943606, KF943508,
(Apomarsupella) rubidum: L1508, China, D.G. Long 34462 (DUKE), KF942911, KF943074, KF943232, KF942786, KF942690; N. succulenta:
KF942453, KF942619, KF943385,—, KF943538, KF942938, KF943103,—, L1491, Colombia, J.C. Benavides s.n. (SIU),—, KF942610,—, KF943622,
KF942811,—; Gymnomitrion (Apomarsupella) verrucosum: L1524, China: KF943528, KF942927, KF943094, KF943251, KF942803, KF942628; Poeltia
Yunnan, D.G. Long & J. Shevock 37182 (DUKE), KF852103, KF851977, campylata: L1499, Bhutan, G. Miehe & S. Miehe 00-356-12 (DUKE),—,—,
KF852260, KF852409, KF851503, KJ802062, KJ802102, KF852574, KF852252,—, KF851494, KJ802036, KJ802097,—,—,—; Prasanthus suecicus:
KF851670,—; Gymnomitrion alpinum: L1519, UK: Scotland, D.G. Long FATOL526, Svalbard, N.A. Konstantinova & A.N. Savchenko K106/1-07
38270 (DUKE), KF852100, KF851974, KF852258, KF852406, KF851500, (F), KF852017, KF851857, KF852137, KF852304, KF851408, KJ802046,
KJ802061, KJ802101, KF852571, KF851667,—; G. commutatum: L1525, China: KJ802074, KF852451, KF851549, KJ802031; FATOL842, Sweden, J. Váňa
Yunnan, D.G. Long 34684 (DUKE), KF942454, KF942622, KF943388, 3.8.2005 (F),—, KF942512,—,—, KF943443,—,—,—, KF942735,—.
KF943624, KF943541, KF942941, KF943106, KF943255, KF942813,—; Gyrothyraceae: Gyrothyra underwoodiana: IBC123, Canada, L.L. Forrest
G. concinnatum: FATOL542, Svalbard, N.A. Konstantinova & A.N. 593 (ABSH),—,—,—,—,—, KF942862, KF942997, JF513364, KF942736,
Savchenko K158/8-07 (F), KF942430, KF942497, KF943290, KF943557, KF942647; L1531, USA: Oregon, B. Shaw F886 (DUKE), KF852104,
KF943428, KF942849, KF942980, KF943145, KF942729, KF942642; KF851978, KF852261, KF852410, KF851504, KJ802063, KJ802103, KF852575,
G. corallioides: FATOL536, Svalbard, N.A. Konstantinova & A.N. Savchenko KF851671,—. Harpanthaceae: Harpanthus flotovianus: E16, Norway, D.G.
K162/2-07 (F), KF942428, KF942494, KF943288, KF943555, KF943425, Long & D.B. Schill 31355 (E),—, KF851822, KF852114, KF852272, KF851379,
KF942846, KF942977, KF943142, KF942726, KF942640; FATOL577, KJ802039, KJ802065, KF852418, KF851517, KJ802023; H. scutatus: IBC26,
Russia, N. Konstantinova s.n. (F),—, KF942500, KF943292, KF943559, USA, A.C. Risk & B. Ratliff 10341 (DUKE),—,—, KF943305,—,—, KF942867,
KF943431, KF942851, KF942983, KF943147,—, KF942644; G. obtusum: KF943002,—,—,—. Jackiellaceae: Jackiella curvata: L1276, New Zealand:
IBC22, Canada, E.C. Davis 424 (DUKE), KF942434,—, KF943304,—,—, North Island, Y. Qiu, M.A.M. Renner & R.M. Schuster NZ-03064 (AK),
KF942866, KF943001, KF943162, KF942741, KF942651; G. pacificum: KF852079, KF851938,, KF852377, KF851467,—,—, KF852538, KF851640,—;
FATOL579, Russia, V. A. Bakalin s.n. (F),—, KF942501,—, KF943560, J. javanica: FATOL254, Fiji, M. von Konrat, J.J. Braggins, & Alivereti
KF943432, KF942852,—, KF943148,—, KF942645; L1381, USA: Naikatini 6/20-9 (F),—,—,—, KF943549,—,—,—, KF943128, KF942715,—.
Alaska, B. Shaw F956/3 (DUKE), KF942449, KF942565, KF943335, Jungermanniaceae: Delavayella serrata: E08, China: Yunnan, D.G. Long
KF943591, KF943483, KF942889, KF943050, KF943207, KF942768, 34809 (E),—, KF851820,—, KF852269, KF851377, KJ802037, KJ802066,
KF942672; Marsupella aquatica: E98, UK: Scotland, D.G .Long 29202 KF852415, KF851514, KJ802018; Eremonotus myriocarpus: FATOL524,
(E),—, KF942470, JF513418, JF513460, JF513478, KF942830, KF942959, Russia, N.A. Konstantinova & A.N. Savchenko K446-6-05 (F),—,—,—,—,
JF513351, KF942709,—; FATOL515, Russia, N.A. Konstantinova 206-1-02 —,—,—, KF943140, KF942723,—; gb308, UK, G. Rothero NH 2116-2410
(F),—, KF942490, KF943286,—, KF943421, KF942842, KF942973, KF943138, (GOET),—,—,—, EF503674,—,—,—,—,—,—; L1138, UK: Scotland,
KF942721,—; M. arctica: FATOL516, Svalbard, N.A. Konstantinova & A.N. B. Buryová 165/93 (DUKE),—, KF851913, KF852210, KF852354, KF851445,
Savchenko k113/2-06 (F),—, KF851853, KF852134, KF852300, KF851404, —, KJ802082, KF852513,—,—; Jungermannia atrovirens: gb309, Russia:
KJ802045, KJ802072, KF852448, KF851545,—; M. emarginata: IBC43, France, Caucasus, N.A.Konstantinova K 421-6-05 (KPABG),—,—,—,—,—,—,
J.-P. Frahm 90/589 (F),—,, KF943307,—,—, KF942868, KF943006,—,—,—; GQ220763,—,—,—; gb310, Russia: Kamchatka, V.A. Bakalin K-74-13a-04
L1425, Czech Republic, P. Sova s.n. (DUKE),—, KF942589, KF943354, (KPABG),—,—,—,—,—,—, GQ220764,—,—,—; L1414, Switzerland,
KF943605, KF943507, KF942910, KF943073, KF943231, KF942785, KF942689; Z. Hradı́lek s.n. (DUKE), KF852094, KF851964, KF852247, KF852398,
M. funckii: FATOL581, Russia, N.A. Konstantinova & A.N. Savchenko s.n. (F), KF851488, KJ802055, KJ802093, KF852564, KF851659, KJ802013;
—, KF942503, KF943294, KF943562, KF943434, KF942854, KF942985, L1415, Switzerland, Z. Hradı́lek s.n. (DUKE),—, KF942579,—,—,
KF943150,—,—; M. sphacelata: FATOL580, Russia, N.A. Konstantinova s.n. KF943497, KF942902, KF943063, KF943221,—,—; J. exsertifolia: gb311,
(F),—, KF942502, KF943293, KF943561, KF943433, KF942853, KF942984, USA: Wyoming, N.A. Konstantinova A69/5-95 (),—,—,—,—,—,—,
KF943149,—,—; M. stoloniformis: L1530, China: Yunnan, D.G. Long 35742 AY327775,—,—,—; J. exsertifolia subsp. cordifolia: E88, UK: Scotland,
(DUKE),—, KF942627, KF943393,—,—, KF942946, KF943111,—,—,—; D.G. Long 29178 (E),—, KF942467, KF943267,—,—, KF942827,
M. yakushimensis: FATOL461, Japan, T. Yamaguchi 24349 (F),—, KF942483, KF942956, KF943122,—, KF942634; E89, UK: Scotland, D.G. Long 31287
KF943279,—, KF943413, KF942837, KF942967, KF943134, KF942719,—; (E),—, KF942468, KF943268,—, KF943401, KF942828, KF942957,
Nardia assamica: L1083, China: Yunnan, B. Shaw 5663 (DUKE), KF942438, KF943123,—, KF942635; FATOL511, Russia, N.A. Konstantinova &
KF942528, KF943316,—,—, KF942878, KF943017, KF943173,—, KF942656; A.N. Savchenko K406/3-07 (F), KF942425, KF942488, KF943284,—,
N. breidleri: FATOL845, Sweden, J. Váňa 15.7.2003 (F),—, KF942513, KF943419, KF942841, KF942972, KF943137,—, KF942638; L1082, Finland,
KF943301,—, KF943444, KF942860, KF942995,—,—,—; L1283, Russia: B. Shaw 1161 (DUKE),—, KF942527,—,—,—,—,—,—,—,—; L1400, USA:
Murmanskaya Oblast, J. Kučera 11398 (CBFS), KF942444, KF942557, Alaska, B. Shaw F951/5 (DUKE),—, KF942566, KF943336, KF943592,
KF943327,—, KF943474,—, KF943042, KF943199, KF942760, KF942668; KF943484, KF942890, KF943051, KF943208,—, KF942673; L1401, USA:
N. compressa: FATOL538, Russia, V.A. Bakalin K-74-9-04 (F), KF942429, Alaska, B. Shaw F951/4 (DUKE),—, KF942567, KF943337,—, KF943485,
KF942495, KF943289, KF943556, KF943426, KF942847, KF942978, KF942891, KF943052, KF943209,—, KF942674; L1421, Norway, P. Sova
KF943143, KF942727, KF942641; L1429, Norway, P. Sova s.n. (DUKE),—, s.n. (DUKE),—, KF942585,,—, KF943503,—, KF943069, KF943227,—,—;
KF942591, KF943356, KF943607, KF943509, KF942912, KF943075, KF943233, J. Konstantinovae: gb313, Russia: Primor’e, V. A. Bakalin P-69-16-08
KF942787, KF942691; L1430, Norway, P. Sova s.n. (DUKE),—, KF942592, (VLA),—,—,—,—,—,—, GU220586,,—,—; J. polaris: L1280, France,
KF943357, KF943608, KF943510, KF942913, KF943076, KF943234, J. Kučera 10710 (CBFS), KF942443, KF942554, KF943324,—, KF943472,
KF942788, KF942692; N. geoscyphus: FATOL528, Russia, N.A. Konstantinova —, KF943039, KF943196,—, KF942666; L1281, Russia: Murmanskaya
148-01 (F), KF942427, KF942493,—, KF943554, KF943424, KF942845, Oblast, J. Kučera 11417 (CBFS),—, KF942555, KF943325,—, KF943473,—,
KF942976, KF943141, KF942725, KF942639; IBC49, USA, E.C. Davis 438 KF943040, KF943197, KF942758, KF942667; L1420, Norway, J. Košnar s.n.
(DUKE), KF942435,—, KF943308,—,—, KF942869, KF943007, KF943163, (DUKE),—, KF942584, KF943350,—, KF943502,—, KF943068, KF943226,—,
KF942685; J. pumila: gb314, Russia: Tuva, V.A. Bakalin 10.VII.1999 (),—,—, KF942705,—; S. nimbosa: L1523, UK: Scotland, D.G. Long & M. Flagmeier
—,—,—,—, AY327771,—,—,—; gb315, Russia: Chita, O.M. Afonina 11606 37028 (DUKE), KF852102, KF851976, KF852259, KF852408, KF851502,—,—,
(KPABG),—,—,—,—,—,—, GU220588,—,—,—; L1422, Norway, P. Sova KF852573, KF851669,—. Schistochilaceae: Schistochila laminigera:
s.n. (DUKE),—, KF942586, KF943351, KF943603, KF943504, KF942907, FATOL78, Chile, J.J. Engel 26644 (F),—, KF942457, KF943297, KF943567,
KF943070, KF943228, KF942782, KF942686; L1423, Czech Republic, KF943439,—, KF942989, KF943155, KF942733,—. Solenostomataceae:
P. Sova s.n. (DUKE),—, KF942587, KF943352,—, KF943505, KF942908, Solenostoma (Horikawaella) grosseverrucosum: L1529, China: Yunnan, D.G.
KF943071, KF943229, KF942783, KF942687; Liochlaena lanceolata: Long 35922 (DUKE), KF942456, KF942626, KF943392,—,—, KF942945,
FATOL548, Russia, unspecified 108000 (F), KF942431, KF942498, KF943110, KF943257, KF942817,—; Solenostoma (Horikawaella) subacutum:
KF943291, KF943558, KF943429, KF942850, KF942981, KF943146, E19, China: Yunnan, D.G. Long 33809 (E),—, KF942460, KF943258,
KF942730, KF942643; IBC136, USA: Illinois, R.E. Stotler & B.J. Crandall- KF943545, KF943396, KF942821, KF942949, KF943115,—, KF942630;
Stotler 107 (ABSH),—,—, JF513431,—,—, KF942863, KF942998,—, L1205, China: Yunnan, D.G. Long 35736 (E),—, KF851919,—, KF852361,
KF942737, KF942648; L1093, USA: Oregon, B. Shaw F578 (DUKE),—, KF851452,—, KJ802084, KF852520, KF851623, KJ802033; Solenostoma
KF942535,—,—,—, KF942882, KF943023,—,—, KF942658; L1290, USA: appressifolium: L1071, China: Yunnan, B. Shaw 5687 (DUKE),—,
Oregon, B. Shaw 6259a (DUKE), KF852082, KF851942, KF852224, KF942517, KF943309,—, KF943446, KF942871, KF943009, KF943164,—,—;
KF852381, KF851471, KJ802053, KJ802091, KF852542, KF851644, KJ802026; L1072, China: Yunnan, B. Shaw 5761 (DUKE),—, KF942518, KF943310,
L1424, Czech Republic, P. Sova s.n. (DUKE),—, KF942588, KF943353, KF943570, KF943447, KF942872, KF943010, KF943165,—,—; L1087, China:
KF943604, KF943506, KF942909, KF943072, KF943230, KF942784, KF942688; Yunnan, B. Shaw 5630 (DUKE),—, KF942532,—, KF943572, KF943455,—,
L. subulata: L1416, Czech Republic, J. Košnar s.n. (DUKE),—, KF942580, KF943021,—,—,—; L1204, China: Yunnan, D.G. Long 35737 (E),—,
KF943347, KF943600, KF943498, KF942903, KF943064, KF943222, KF942547,—, KF943580, KF943467,—, KF943033, KF943189, KF942752,
KF942778, KF942682; L1507, China, D.G. Long 35240 (DUKE), KF942452, KF942663; S. atrorevolutum: L1197, China: Yunnan, D.G. Long 35732
KF942618, KF943384,—, KF943537, KF942937, KF943102, KF943253, (E),—, KF942540,—, KF943575, KF943461,—, KF943027, KF943182,
KF942810,—; Mesoptychia badensis: L1496, Canada: Newfoundland, KF942747, KF942661; S. callithrix: L1308, Portugal, R.E. Stotler & B.J.
B. Shaw F455 (DUKE),—, KF942615, KF943379,—, KF943533, KF942932, Crandall-Stotler 4664 (ABSH),—, KF942561, KF943331,—, KF943478,
KF943099, KF943252, KF942807,—; M. bantriensis: E91, UK: Scotland, KF942885, KF943046, KF943203,—,—; S. chenianum: L1506, China, D.G.
D.G. Long 34172 (E),—, KF942469, JF513416, JF513458, JF513476, Long 35816 (E),—, KF942617, KF943383,—, KF943536, KF942936,—,—,
KF942829, KF942958, JF513349, KF942708, KF942636; M. gillmanii: KF942809,—; S. clavellatum: E23, Bhutan, D.G. Long 28636 (E),—, KF942462,
FATOL532, Russia, N.A. Konstantinova & A.N. Savchenko 117-1-00 (F), KF943262, KF943546, KF943398, KF942823, KF942951, KF943112,
KF852018, KF851859, KF852139, KF852305, KF851410, KJ802047, KJ802075, KF942704,—; S. comatum: FATOL56, Vietnam: Ha Tay Province, T. Pócs &
KJ451736, KF851551, KJ802027; M. heterocolpos: IBC137, USA: Utah, R.E. G. Kosa 98109/D (F),—, KF942499,—,—, KF943430,—, KF942982,—,
Stotler & B.J. Crandall-Stotler 4357 (ABSH), KF942432,—, KF943303,—,—,—, KF942731,—; FATOL835, Japan, M. Higuchi 1146 (F),—,—, KF943298,—,
KF942999, KF943161, KF942738, KF942649; L1406, USA: Alaska, B. Shaw —,—, KF942991, KF943157,—,—; S. confertissimum: FATOL504, Russia,
F965a/3 (DUKE),—, KF942572,—,—, KF943490, KF942896, KF943057, N.A. Konstantinova & A.N. Savchenko K548-1-05 (F),—, KF942485,
KF943214,—, KF942678; M. morrisoncola: gb247, Russia: Buryatiya, V.A. KF943282,—, KF943416, KF942839, KF942969,—,—,—; gb323, Russia:
Bakalin (),—,,—,—,—,—, AY327772,—,—,—; M. polymorpha: L1219, USA: Caucasus, N.A. Konstantinova K 459-8a-05 (KPABG),—,—,—,—,—,—,
California, W.T. Doyle 11541 (ABSH),—, KF942548,—, KF943581, GQ220758,—,—,—; L1099, Austria, B. Shaw F272 (DUKE),—, KF942536,
KF943468,—, KF943034, KF943190,—, KF942664; M. rutheana: L1146, —,—,—,—,—,—,—,—; L1417, Norway, P. Sova s.n. (DUKE),—, KF942581,
Canada: British Columbia, B. Shaw F702 (DUKE), KF942442, KF942538, KF943348, KF943601, KF943499, KF942904, KF943065, KF943223,
KF943322, KF943573, KF943459,—, KF943025, KF943180, KF942745, KF942779,—; S. confertissimum (abyssinicum): FATOL772, Kenya, S. Pócs &
KF942660; M. rutheana: L1487, USA: Alaska, B. Shaw F973/10 T. Pócs, guided by M.S. Chuah-Petiot 04005/AH (F),—, KF942507,—,
(DUKE), KF942451, KF942609, KF943374, KF943621, KF943527,—, KF943566, KF943438, KF942856, KF942988, KF943154, KF942732,—;
KF943093, KF943250, KF942802, KF942701; M. sahlbergii: FATOL592, S. coniflorum: L1412, Iles Kerguelen, R. Ochyra s.n. (DUKE),—, KF942577,
Russia, V. Fedosov 107967 (F), KF852022, KF851864, KF852144, KF852310, —,—, KF943495, KF942901,—, KF943219,—,—; L1495, Kergulelen Islands,
KF851415, KJ802050, KJ802078, KF852455, KF851556, KJ802028. R. Ochyra 1219/06 (DUKE),—, KF942614, KF943378,—, KF943532,
Lepidoziaceae: Bazzania exempta: L1518, New Zealand, P.J. de Lange KF942931, KF943098,—, KF942806,—; S. crenuliforme: IBC33, USA,
7881 (DUKE), KF942420, KF942621, KF943387, KF943623, KF943540, A. Risk 11014 (DUKE),—, KF942515, KF943306,—,—,—, KF943003,—,
KF942940, KF943105, KF943254, KF942812, KF942702; Zoopsis nitida: —,—; L1285, USA: North Carolina, B. Shaw 6187 (DUKE), KF942445,
FATOL826, New Zealand, D. Glenny 10517 (F),—, KF942508,—, KF942559, KF943329,—, KF943476,—, KF943044, KF943201, KF942762,—;
KF943568, KF943440, KF942857, KF942990, KF943156, KF942734, L1440, USA: North Carolina, B. Shaw 7212 (DUKE),—, KF942601,
KF942646. Lophocoleaceae: Bragginsella anomala: L1129, New Zealand, KF943366, KF943617, KF943519, KF942922, KF943085, KF943243,
M. von Konrat & J.J. Engel 51 SW facing (F),—, KF851909,—,—, KF942797,—; S. cryptogynum: FATOL51, New Zealand, J.J. Engel,
KF851441, KJ802052, KJ802081, KF852509, KF851614, KJ802015; M. von Konrat & J.E. Braggins 25036 (F), KF942424, KF942487,
Lophocolea heterophylla: IBC140, USA: Illinois, R.E. Stotler & B.J. Crandall- KF943283,—, KF943418, KF942840, KF942971, KF943136,—,—; S. dulongense:
Stotler s.n. (ABSH),—,—, JF513434,—,—, KF942865,—, JF513369, L1200, China: Yunnan, D.G. Long 33676 (E),—, KF942543,—,—, KF943464,
KF942740,—; Lophozia ventricosa: E96, UK: Scotland, D.G. Long 31226 —,—, KF943185, KF942749,—; S. faurieanum: FATOL460, Japan,
(E),—, KF851831, JF513417, JF513459, JF513477,—,—, JF513350, KF851529, T. Yamaguchi 27075 (F),—, KF942482, KF943278,—, KF943412, KF942836,
—. Myliaceae: Leiomylia anomala: L1220, Camada: Alberta, D.H. Vitt s.n. KF942966,—,—,—; S. flagellatum: FATOL459, Japan, H. Yokoyama 11575
(ABSH), KF852075, KF851925,—, KF852367, KF851458,—, KJ802088, (F),—, KF942481, KF943277,—, KF943411, KF942835, KF942965,
KF852526, KF851629, KJ802024; Mylia taylorii: E32, China: Yunnan, D.G. KF943133, KF942718, KF942637; S. fusiforme: FATOL433, Japan,
Long 34707 (E), KF852000, KF851826, KF852118, KF852277,—, KJ802041,—, M. Higuchi 1147 (F),—, KF942479, KF943275,—, KF943409,—,—,
KF852424, KF851523, KJ802030. Notoscyphaceae: Notoscyphus lutescens: KF943131,—,—; S. gracillimum: L1080, Czech Republic, B. Shaw s.n.
L1501, Vietnam, T. Pócs 98109/c (DUKE),—,—, KF943381,—, KF943535, (DUKE),—, KF942525, KF943314,—, KF943452, KF942876, KF943015,
KF942934, KF943100,—,—,—; L1505, Australia, T. Pócs 01098/d KF943171,—, KF942655; L1089, USA: North Carolina, B. Shaw 4330
(DUKE),—, KF851970, KF852253,—, KF851496, KJ802060, KJ802099,—, (DUKE),—,—, KF943318,—,—,—,—,—,—,—; L1437, Austria, P. Sova s.n.
—,—. Plagiochilaceae: Pedinophyllum interruptum: FATOL518, Russia, (DUKE),—, KF942598, KF943363, KF943614, KF943516, KF942919,
N.A. Konstantinova & A.N. Savchenko k508/7-07 (F), KF852014, KF943082, KF943240, KF942794, KF942696; L1438, Sweden, P. Sova s.n.
KF851854,—, KF852301, KF851405,—, KJ802073, KF852449, KF851546,—; (DUKE),—, KF942599, KF943364, KF943615, KF943517, KF942920,
Plagiochila porelloides: L1405, USA: Alaska, B. Shaw F955/1 (DUKE),—, KF943083, KF943241, KF942795, KF942697; L1439, Czech Republic,
KF942571, KF943341, KF943595, KF943489, KF942895, KF943056, P. Sova s.n. (DUKE),—, KF942600, KF943365, KF943616, KF943518,
KF943213, KF942773,—. Pseudolepicoleaceae: Pseudolepicolea quadrilaciniata: KF942921, KF943084, KF943242, KF942796, KF942698; S. handelii:
E52, Argentina, D.G. Long 31658 (E),—, KF851829,—, KF852279,—, FATOL210, Japan, M. Itouga s.n. (F),—, KF942475,—,—, KF943405,
KJ802042, KJ802069, KF852426, KF851525, KJ802032. Saccogynaceae: KF942818, KF942961, KF943126, KF942714,—; S. hasskarlianum:
Saccogyna viticulosa: FATOL709, Madeira, L. Söderström s.n. (F),—, FATOL257, Fiji, M. von Konrat, J.J. Braggins & Alivereti Naikatini 6/20-5
KF942506,—, KF943565, KF943437,—,—, KF943153,—,—; L1309, Portugal: (F),—, KF942477, KF943273,—, KF943407, KF942833, KF942963, KF943129,
Sintra, R. E. Stotler & B.J. Crandall-Stotler 4657 (ABSH),—, KF851947, KF942716,—; FATOL838, Australia, T. Pócs & H. Streimann 99111/v (F),—,
KF852229, KF852385, KF851476,—,—, KF852547, KF851649,—. Scapaniaceae: KF942511,—,—,—,—, KF942994,—,—,—; S. hyalinum: FATOL525, Russia,
Scapania nemorea: E56, UK: Scotland, D.G. Long 35418 (E), KF942421, N.A. Konstantinova & A.N. Savchenko 351-2-04 (F), KF942426, KF942492,
KF942463, KF943264,—, KF943400, KF942824, KF942952, KF943118, KF943287, KF943553, KF943423, KF942844, KF942975,—, KF942724,—;
L1076, USA: North Carolina, B. Shaw Fza553 (DUKE),—, KF942521, KF943494, KF942900, KF943061, KF943218, KF942777, KF942681;
KF943313,—, KF943450,—,—, KF943168,,—; L1291, USA: Oregon, B. Shaw S. parvitextum: L1073, China: Yunnan, B. Shaw 5775 (DUKE), KF942436,
6259b (DUKE), KF942446, KF942560, KF943330, KF943586, KF943477, KF942519, KF943311, KF943571, KF943448, KF942873, KF943011,
KF942884, KF943045, KF943202, KF942763,—; L1431, Austria, P. Sova s.n. KF943166,—,—; S. patoniae: E14, China: Yunnan, D.G. Long 34743 (E),—,
(DUKE),—, KF942593, KF943358, KF943609, KF943511, KF942914, KF942459, KF943260,—,—, KF942820, KF942947, KF943114,—,—;
KF943077, KF943235, KF942789,—; L1432, Austria, P. Sova s.n. (DUKE),—, S. polyrhizoides: L1528, China: Yunnan, D.G. Long 36119 (DUKE),—,
KF942594, KF943359, KF943610, KF943512, KF942915, KF943078, KF942625, KF943391,—, KF943543, KF942944, KF943109,—, KF942816,—;
KF943236, KF942790,—; S. infuscum: FATOL226, Japan, H. Deguchi s.n. S. pseudopyriflorum: gb326, Russia: Sakhalin, V.A. Bakalin S-25-1a-06
(221) (F),—, KF942476,—,—, KF943406,—, KF942962, KF943127,—,—; (VLA),—,—,—,—,—,—, GU220592,—,—,—; gb327, Russia: Sakhalin,
L1070, China: Jilin, B. Shaw 5448 (DUKE),—, KF942516,—,—,—, V.A. Bakalin K-12-8-07 (VLA),—,—,—,—,—,—, GU220593,—,—,—;
KF942870, KF943008,—,—,—; L1084, China: Jilin, B. Shaw 5424 (DUKE), gb328, Russia: Primor’e, V.A. Bakalin P-74-79a-05 (VLA),—,—,—,—,—,—,
KF942439, KF942529,—,—, KF943453,—, KF943018, KF943174,—,—; GU220594,—,—,—; gb329, Russia: Primor’e, V.A.Bakalin P-65-1-06
S. inundatum: FATOL37, New Zealand, J.J. Engel & M. von Konrat 23589 (VLA),—,—,—,—,—,—, GU220595,—,—,—; gb330, Russia: Buryatiya,
(F),—, KF942478, KF943274,—, KF943408,—,—, KF943130,—,—; L1483, N.A. Konstantinova 30-2-01 (KPABG),—,—,—,—,—,—, GQ220759,—,—,—;
Australia: Tasmania, B. Shaw 6391 (DUKE),—, KF942606, KF943371, S. pyriflorum: gb324, South Korea: KyongNam, V.A. Bakalin Kor-8-5-09
KF943619, KF943524, KF942926, KF943090, KF943247, KF942799,—; (VLA),—,—,—,—,—,—, GU220590,—,—,—; gb325, South Korea:
L1484, Australia: Tasmania, B. Shaw 6395 (DUKE),—, KF942607, KyongNam, V.A. Bakalin Kor-10-8-09 (VLA),—,—,—,—,—,—, GU220591,
KF943372, KF943620, KF943525,—, KF943091, KF943248, KF942800,—; —,—,—; L1078, USA: North Carolina, B. Shaw 12988 (DUKE),—,
S. inundatum (rufiflorum): FATOL857, New Zealand, D. Glenny 9944 KF942523,—,—,—,—, KF943014, KF943169,—,—; L1079, USA: North
(F),—, KF942514, KF943302, KF943569, KF943445, KF942861, KF942996, Carolina, B. Shaw pSchF (DUKE),—, KF942524,—,—,—,—,—, KF943170,
KF943160,—,—; S. kashyapii: L1199, China: Yunnan, D.G. Long 34448 —,—; L1241, USA: North Carolina, B. Shaw D-4A (DUKE),—, KF942553,
(E),—, KF942542,—, KF943577, KF943463,—, KF943029, KF943184,—, —,—, KF943471,—, KF943038, KF943195, KF942757,—; L1493, USA:
KF942662; S. lanigerum: L1086, China: Yunnan, B. Shaw 5604 (DUKE), North Carolina, B. Shaw SchF 2007 (DUKE),—, KF942612, KF943376,—,
KF942440, KF942531,—,—, KF943454, KF942880, KF943020, KF943176,—, KF943530, KF942929, KF943096,—, KF942805,—; S. rosulans: L1196,
—; S. macrocarpum: L1085, China: Yunnan, B. Shaw 5573 (DUKE),—, China: Yunnan, D.G. Long 36185 (E),—, KF942539,—, KF943574,
KF942530, KF943317,—,—, KF942879, KF943019, KF943175,—,—; L1527, KF943460,—, KF943026, KF943181, KF942746,—; L1201, China: Yunnan,
China: Yunnan, D.G. Long 35702 (DUKE), KF942455, KF942624, D.G. Long 36011 (E),—, KF942544,—, KF943578, KF943465,—, KF943030,
KF943390,—, KF943542, KF942943, KF943108, KF943256, KF942815,—; KF943186, KF942750,—; L1203, China: Yunnan, D.G. Long 33911 (E),—,
S. marginatum: FATOL455, Japan, T. Yamaguchi 29073 (F), KF942422, KF942546,—, KF943579, KF943466,—, KF943032, KF943188, KF942751,—;
KF942480, KF943276,—, KF943410, KF942834, KF942964, KF943132, S. rubripunctatum: L1504, China, D.G. Long 32517 (DUKE),—,—,—,—,—,
KF942717,—; S. montanum: FATOL666, Australia, T. Pócs & H. Streimann —,—,—, KF942808,—; S. rubrum: L1092, USA: Oregon, B. Shaw F595
s.n. (F),—, KF942505, KF943296, KF943564, KF943436,—, KF942987, (DUKE),—, KF942534, KF943319,—, KF943456, KF942881, KF943022,
KF943152,—,—; FATOL836, Australia, T. Pócs & H. Streimann 99105/F KF943177,—, KF942657; L1486, USA: Alaska, B. Shaw 7845 (DUKE),—,
(F),—, KF942509, KF943299,—, KF943441, KF942858, KF942992, KF943158, KF942608, KF943373,—, KF943526,—, KF943092, KF943249, KF942801,—;
—,—; S. obovatum: FATOL519, Russia, N.A. Konstantinova & A.N. S. sanguinolentum: L1202, China: Yunnan, D.G. Long 35755 (E),—,
Savchenko 382-3 -04 (F),—, KF942491,—, KF943552, KF943422, KF942843, KF942545,—,—,—,—, KF943031, KF943187,—,—; S. schusterianum:
KF942974, KF943139, KF942722,—; gb316, Russia: Murmansk, N.A. IBC34, Canada, E.C. Davis 431 (DUKE),—,—, KF943394,—,—,—,
Konstantinova 196-6-02 (KPABG),—,—,—,—,—,—, GQ220754,—,—,—; KF943004,—,—,—; S. sp.: gb320, Russia: Murmansk, N.A. Konstantinova
gb317, Russia: Kemerovo, N.A. Konstantinova 72-2-00 (KPABG),—,—, 30-1-97 (KPABG),—,—,—,—,—,—, GQ220761,—,—,—; gb321, Russia:
—,—,—,—, GQ220753,—,—,—; gb318, Russia: Perm, N.A. Konstantinova Buryatiya, N.A. Konstantinova 70-2-01 (KPABG),—,—,—,—,—,—,
K 324-1-04 (KPABG),—,—,—,—,—,—, GQ220755,—,—,—; L1090, Czech GQ220751,—,—,—; S. speciosum: E57, China: Yunnan, D.G. Long 33745
Republic, B. Shaw 1777 (DUKE),—, KF942533,—,—,—,—,—,—,—,—; (E),—, JF513397, JF513413, JF513456,—, KF942825, KF942953, JF513346,
L1402, USA: Alaska, B. Shaw F960/6 (DUKE),—, KF942568, KF943338,—, KF942706,—; S. sphaerocarpum: L1081, Czech Republic, B. Shaw 3503
KF943486, KF942892, KF943053, KF943210, KF942770, KF942675; (DUKE),—, KF942526, KF943315,—,—, KF942877, KF943016, KF943172,
L1403, USA: Alaska, B. Shaw F961/14 (DUKE),—, KF942569, KF943339, —,—; L1441, Norway, P. Sova s.n. (DUKE),—, KF942602, KF943367,
KF943593, KF943487, KF942893, KF943054, KF943211, KF942771, KF943618, KF943520, KF942923, KF943086, KF943244, KF942798,—;
KF942676; L1433, Norway, P. Sova s.n. (DUKE),—, KF942595, KF943360, L1442, Czech Republic, P. Sova s.n. (DUKE), KF852096, KF851966,
KF943611, KF943513, KF942916, KF943079, KF943237, KF942791, KF852249, KF852400, KF851490, KJ802057, KJ802095, KF852566,
KF942693; L1434, Czech Republic, P. Sova s.n. (DUKE),—, KF942596, KF851661,—; S. sphaerocarpum (pusillum): L1282, Russia: Murmanskaya
KF943361, KF943612, KF943514, KF942917, KF943080, KF943238, Oblast, J. Kučera 11462 (CBFS),, KF942556, KF943326,—,—,—, KF943041,
KF942792, KF942694; L1435, Austria, P. Sova s.n. (DUKE),—, KF942597, KF943198, KF942759,—; S. totipapillosum: FATOL50, New Zealand,
KF943362, KF943613, KF943515, KF942918, KF943081, KF943239, J.J. Engel, M. von Konrat & J.E. Braggins 24933 (F),—,—, KF943281,
KF942793, KF942695; L1492, USA: Alaska, B. Shaw 7755 (DUKE),—, KF943551, KF943415,—,—,—,—,—; L1407, New Zealand, M. Renner
KF942611, KF943375,—, KF943529, KF942928, KF943095,—, KF942804,—; 4201 ? 4021 (DUKE),—, KF942573, KF943342, KF943596, KF943491,
S. obovatum (subellipticum): gb319, Russia: Kamchatka, V.A. Bakalin KF942897, KF943058, KF943215, KF942774,—; S. truncatum: FATOL837,
K-48-13-03 (KPABG),—,—,—,—,—,—, GQ220752,—,—,—; IBC35, USA, Australia, T. Pócs & H. Streimann 9969/J (F),—, KF942510, KF943300,—,
W.B. Schofield 111132 (DUKE),—,—,—,—,—,—, KF943005,—,—,—; KF943442, KF942859, KF942993, KF943159,—,—. Southbyaceae: Gongylanthus
S. orbiculatum: FATOL54, New Zealand: Westland Province, J. J. Engel, ericetorum: L1139, Sicily, B. Buryová 593 (DUKE),—,—, KF943321,—,
M. von Konrat & J.E. Braggins 24788 (F),—, KF942496,—,—, KF943427, KF943458,—,—, KF943179, KF942744,—; Gongylanthus liebmannianus:
KF942848, KF942979, KF943144, KF942728,—; L1232, New Zealand, FATOL774, Venezuela: Merida, L. Söderström, P. Manyanga & R. de Roo
J.J. Engel & M. von Konrat 28158 (F),—, KF942550,—, KF943583,—,—, 2004/030b (F), KF852031, KF851883,—,—,—,—,—, KF852471, KF851563,
KF943036, KF943192, KF942754,—; L1236, New Zealand, J.J. Engel & —; L1479, Colombia, J.C. Benavides s.n. (SIU),—, KF851967,—,—,
M. von Konrat 28166 (F),—, KF942552,—, KF943585,—,—, KF943037, KF851491,—,—,—,—,—; Southbya nigrella: L1140, Italy, B. Buryová 621
KF943194, KF942756,—; L1408, New Zealand, M. Renner 4137 (DUKE),—, KF851914, KF852211, KF852355, KF851446,—, KJ802083,
(DUKE),—, KF942574, KF943343, KF943597, KF943492, KF942898, KF852514, KF851618, KJ802020; Southbya tophacea: E60, Italy, D.G. Long
KF943059, KF943216, KF942775, KF942679; L1410, New Zealand, 35466 (E),—, KF942465, KF943265,—,—,—, KF942954, KF943120,—,
M. Renner s.n. (DUKE),—, KF942575, KF943344, KF943598, KF943493, KF942632. Trichotemnomataceae: Trichotemnoma corrugatum: L1279,
KF942899, KF943060, KF943217, KF942776, KF942680; L1411, New New Zealand: South Island, Y. Qiu & J.E. Braggins NZ-03123 (AK),—,
Zealand, M. Renner s.n. (DUKE),—, KF942576, KF943345, KF943599, KF851940,—, KF852379, KF851469,—,—, KF852540, KF851642,—.

Phylogenetic Relationships and Morphological Evolution in A Major Clade of

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Phylogenetic Relationships and Morphological Evolution in a Major Clade of

Leafy Liverworts (Phylum Marchantiophyta, Order Jungermanniales): Suborder

Phylogenetic Relationships and Morphological Evolution in a Major Clade

Communicating Editor: Thomas L. P. Couvreur

atpB cp 70 75% 67 60% 1,181 997 595 68 334 34% GTR+I+G

or with delicate, low papillae, in S. inundatum, which is Taxonomic Treatment

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