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World checklist of hornworts and liverworts 1
World checklist of hornworts and liverworts
Lars Söderström1,*, Anders Hagborg2,*, Matt von Konrat2,*,
Sharon Bartholomew-Began3, David Bell4, Laura Briscoe2, Elizabeth Brown5 , †,
D.Christine Cargill6, Denise P. Costa7, Barbara J. Crandall-Stotler8,
Endymion D. Cooper9, Gregorio Dauphin10, John J. Engel2, KathrinFeldberg11,
David Glenny12, S. Robbert Gradstein13, Xiaolan He14, Jochen Heinrichs11,
JörnHentschel15, Anna Luiza Ilkiu-Borges16, Tomoyuki Katagiri17,
NadezhdaA.Konstantinova18, Juan Larraín2, David G. Long19, MartinNebel20,
Tamás Pócs21, Felisa Puche22, Elena Reiner-Drehwald23, Matt A.M. Renner5,
Andrea Sass-Gyarmati21, Alfons Schäfer-Verwimp24, José Gabriel Segarra Moragues25,
Raymond E. Stotler8, † , Phiangphak Sukkharak26, Barbara M. iers27,
JaimeUribe28, Jiří Váňa29, Juan Carlos Villarreal30, Martin Wigginton31,
Li Zhang32, Rui-Liang Zhu33
1 Department of Biology, Norwegian University of Science and Technology, N-7491 Trondheim, Norway
2 Department of Science and Education, Field Museum, 1400 South Lake Shore Drive, Chicago, IL 60605–
2496, United States of America 3 Department of Biology, West Chester University, West Chester, PA 19383,
United States of America 4 Department of Botany, University of British Columbia, 6270 University Boulevard,
Vancouver, BC, V6T 1Z4, Canada 5 Royal Botanic Gardens and Domain Trust, Mrs Macquaries Road,
Sydney NSW2000, Australia 6 Centre for Australian National Biodiversity Research, Australian National
Herbarium, GPO Box 1600, Canberra, ACT 2601, Australia 7 Instituto de Pesquisas Jardim Botânico do
Rio de Janeiro, Rua Pacheco Leão 915, 22460-030, Rio de Janeiro, RJ, Brazil 8 Department of Plant Biology,
Southern Illinois University, Carbondale, Illinois 62901-6509, United States of America 9 CMNS-Cell
Biology and Molecular Genetics, 2107 Bioscience Research Building, University of Maryland, College Park,
MD 20742-4451, United States of America 10 Apartado 5-1500, Acosta, Costa Rica 11 Systematic Botany
and Mycology, Ludwig Maximilian University of Munich, Menzinger Str. 67, 80638 Munich, Germany
12 Allan Herbarium, Landcare Research, P O Box 69-040, Lincoln 7608, New Zealand 13 Muséum
National d’Histoire Naturelle, Department Systématique et Evolution, C.P. 39, 57 Rue Cuvier, 75231 Paris
05, France 14 Botany Unit, Finnish Museum of Natural History, University of Helsinki, P.O. Box 7, Helsinki
FI-00014, Finland 15 Department of Systematic Botany with Herbarium Haussknecht and Botanical Garden,
Friedrich Schiller University, Fürstengraben 1, 07737 Jena, Germany 16 Museu Paraense Emilio Goeldi,
Coordenaçao de Botanica, Av. Magalhaes Barata 376, 66040-1 70 Belem, Para, Brazil 17 Department of
Biological Science, Graduate School of Science, Hiroshima University, Kagamiyama 1–3–1, Higashihiroshima-
shi, Hiroshima 739–8526, Japan 18 N.A. Avrorin Polar-Alpine Botanical Garden–Institute of Kola SC RAS,
184236 Kirovsk-6, Russia 19 Royal Botanic Garden, Edinburgh EH3 5LR, United Kingdom 20 Staatliches
PhytoKeys 59: 1–828 (2016)
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
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Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany 21 Botany Department, Institute
of Biology, Eszterházy Károly College, Eger, Pf. 43, H-3301, Hungary 22 Departamento de Botánica,
Facultad de Ciencias Biológicas, Universitat de València. C/ Dr. Moliner 50, E-46100, Burjassot (Valencia),
Spain 23 Albrecht-von-Haller-Institut für Panzenwissenschaften, Department of Systematics, Biodiversity
and Evolution of Plants, Untere Karspüle 2, 37073 Göttingen, Germany 24 Mittlere Letten 11, 88634
Herdwangen-Schönach, Germany 25 Centro de Investigaciones sobre Deserticación (CIDE-CSIC-UV-GV),
C/ Carretera de Moncada-Náquera Km. 4.5, E-46113, Moncada (Valencia), Spain 26 Department of Biology,
Faculty of Science, Burapha University, Mueang, 20131 Chonburi, ailand 27 William and Lynda Steere
Herbarium, e New York, Botanical Garden, Bronx, New York 10458-5126, United States of America
28 Instituto de Ciencias Naturales. Universidad Nacional de Colombia. Apartado 7495, Bogotá D.C.,
Colombia 29 Department of Botany, Charles University, Benátská 2, CZ-128 01 Praha 2, Czech Republic
30 Department of Biology, Ludwig-Maximilians-Universität, Menzinger Str. 67, D-80638, München,
Germany 31 36, Big Green, Warmington, Peterborough PE8 6TU, United Kingdom 32 Shenzhen Key
Laboratory of Southern Subtropical Plant Diversity, Fairylake Botanical Garden, 160 Xianhu Rd., Liantang,
Shenzhen 518004, Guangdong, China 33 Department of Biology, School of Life Sciences, East China Normal
University, 3663 Zhong Shan North Road, Shanghai 200062, China
* Editors
Corresponding author: Lars Söderström (lars.soderstrom@ntnu.no)
Academic editor: Lyubomir Penev|Received 20 June 2015|Accepted 25 September 2015|Published 29 January2016
Citation: Söderström L, Hagborg A, von Konrat M, Bartholomew-Began S, Bell D, Briscoe L, Brown E, Cargill DC,
Costa DP, Crandall-Stotler BJ, Cooper ED, Dauphin G, Engel JJ, Feldberg K, Glenny D, Gradstein SR, He X, Heinrichs
J, Hentschel J, Ilkiu-Borges AL, Katagiri T, Konstantinova NA, Larraín J, Long DG, Nebel M, Pócs T, Felisa Puche F,
Reiner-Drehwald E, Renner MAM, Sass-Gyarmati A, Schäfer-Verwimp A, Moragues JGS, Stotler RE, Sukkharak P,
iers BM, Uribe J, Váňa J, Villarreal JC, Wigginton M, Zhang L, Zhu R-L (2016) World checklist of hornworts and
liverworts. PhytoKeys 59: 1–828. doi: 10.3897/phytokeys.59.6261
is checklist is dedicated to Elizabeth Brown and Ray Stotler, who sadly left us
during our work with it.
“……Charles Darwin wrote of his desire to provide nancial support ‘for the formation of a perfect
M.S. catalogue of all known plants’ (Darwin 1881, letter 13570). It is a personal embarrassment to
me, and should be chastening to us all, that more than 120 years later we still have not delivered on
that commitment.” (Crane 2004).
Abstract
A working checklist of accepted taxa worldwide is vital in achieving the goal of developing an online
ora of all known plants by 2020 as part of the Global Strategy for Plant Conservation. We here present
the rst-ever worldwide checklist for liverworts (Marchantiophyta) and hornworts (Anthocerotophyta)
that includes 7486 species in 398 genera representing 92 families from the two phyla. e checklist
has far reaching implications and applications, including providing a valuable tool for taxonomists and
systematists, analyzing phytogeographic and diversity patterns, aiding in the assessment of oristic and
World checklist of hornworts and liverworts 3
taxonomic knowledge, and identifying geographical gaps in our understanding of the global liverwort
and hornwort ora. e checklist is derived from a working data set centralizing nomenclature, taxo-
nomy and geography on a global scale. Prior to this eort a lack of centralization has been a major
impediment for the study and analysis of species richness, conservation and systematic research at both
regional and global scales. e success of this checklist, initiated in 2008, has been underpinned by
its community approach involving taxonomic specialists working towards a consensus on taxonomy,
nomenclature and distribution.
Keywords
Marchantiophyta, Anthocerophyta, nomenclature, taxonomy
Introduction
e natural world is changing fast (Balmford et al. 2005). e Global Strategy for
Plant Conservation (GSPC) was adopted by the Conference of Parties (COP) of the
Convention on Biological Diversity (CBD) in April 2002 (http://www.cbd.int/deci-
sion/cop/?id=7183; accessed 2014.06.02). e Strategy set out 16 outcome-oriented
targets that were to be achieved by 2010. e GSPC was designed as a framework for
action to halt the loss of plant diversity. Target 1 of the Strategy was to complete “a
widely accessible working list of all known plant species, as a step towards a complete
world Flora” (Lughadha 2004). As early as 1881 Charles Darwin expressed a wish to
have a catalogue of all known plants (Crane 2004). However, over 125 years later, this
wish is not yet fullled. Without a working checklist, many of the other objectives in
the GSPC cannot be met and botanists around the world cannot communicate about
plants on a global basis (Crane 2004). A working list of known plant species is criti-
cal for (I) its underpinning role in the eective implementation of the other targets
through provision of baseline information; (II) increasing the accessibility and use of
accurate botanical name information for research, conservation and sustainable use;
and (III) real-world politics and how taxonomists respond to the decisions of policy
makers (Paton et al. 2008).
Version 1.0 of e Plant List (http://www.theplantlist.org/) was released in De-
cember 2010 aimed to be comprehensive for species of vascular plants (owering
plants, conifers, ferns and their allies) and of bryophytes (mosses, liverworts and horn-
worts), as a response to the 2010 Target 1 of the GSPC and to a clear global need for
such data. e Plant List is a broad collaboration, coordinated by the Royal Botanic
Gardens, Kew, and the Missouri Botanical Garden (MO), involving diverse partner-
ships. Paton (2013) noted e Plant List represents a work in progress with future ver-
sions planned to improve the quality of names and decrease the number of unresolved
names. Target 1 has since been revised and agreed at the 10th COP of the CBD in Na-
goya, Japan, with the goal of developing ‘‘an online ora of all known plants’’ by 2020
(http://www.cbd.int/decision/cop/default.shtml?id=12283; accessed 2014.06.02). Pa-
ton (2013) identied several factors that might assist in the implementation of the
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
4
revised target, stressing that achieving a working list for plant taxa worldwide was a
vital step to an online ora.
Earlier, Paton et al. (2008) assessed the progress made at that time and discussed
prospects for the completion of Target 1. Paton noted that good progress had been
made in bryophytes (mosses, liverworts and hornworts), ferns and gymnosperms with
widely accessible working lists either complete or almost so for those groups. For bryo-
phytes alone, he tabulated 13,370 accepted species noting that the data was largely
derived from Tropicos. However, although Tropicos is an indispensable reference for
anyone dealing with bryophyte names and the database is very strong for mosses, the
nomenclatural and auxiliary data for liverworts is less complete, especially for larger
genera (von Konrat et al. 2008a). e lack of a central source providing a synthesis of
nomenclatural data and global distributional data was the impetus toward developing
the current checklist of liverworts and hornworts (von Konrat et al. 2008a, 2010d,
Söderström et al. 2012e). Checklists are powerful and important tools that can inte-
grate the almost overwhelmingly scattered information concerning taxonomy, syste-
matics, nomenclature, distribution, and even frequency (Söderström et al. 2008).
Ecological and biological significance of liverworts and hornworts
Liverworts and hornworts (Figure 1) are of critical biological, ecological, and phylo-
genetic signicance (e.g. Asakawa 1998, Longton 1992, Hallingbäck and Hodgetts
2000, Gradstein et al. 2001b, Wellman et al. 2003, Qiu et al. 2007). Liverworts are
found on soil, rocks, and trees throughout the world, from coastal Antarctica to the
tundra of the Northern Hemisphere, and from semi-arid areas of Australia to the Ama-
zon rainforests (Hallingbäck and Hodgetts 2000). Although there are xero-tolerant
taxa, the majority of liverworts are found in relatively humid and shaded terrestrial
ecosystems (Gradstein and Costa 2003). Liverworts and hornworts are an important
component of the vegetation in many regions of the world, constituting a major part
of the biodiversity in moist forest, wetlands, mountain, and tundra ecosystems (Hal-
lingbäck and Hodgetts 2000).
Liverworts, hornworts and mosses oer microhabitats that are critical to the survival
of a tremendous diversity of organisms such as single-celled eukaryotes, protozoa, and
numerous groups of invertebrates (Gerson 1982). eir structural contribution to levels
of diversity might be as signicant as that of vascular plants, albeit at a smaller scale (von
Konrat et al. 2008a). ese plants are also important environmental and ecological indica-
tors (Rao 1982, Pitcairn et al. 1995, Gradstein et al. 2001b, Giordano et al. 2004). Liver-
worts, in concert with mosses and hornworts, play a signicant role in the global carbon
budget (O’Neill 2000) and CO2 exchange (De Lucia et al. 2003), plant succession (Cre-
mer and Mount 1965), net production and phytomass (Frahm 1990), nutrient cycling
(Coxson 1991), and water retention (Pócs 1980a, Gradstein et al. 2001b). ese groups
of land plants also have been used as indicators of past climate change, to validate climate
models, and as potential indicators of global warming (Gignac 2001, Tuba et al. 2011).
World checklist of hornworts and liverworts 5
Species numbers
ere are estimated to exist only 215 hornwort species. By comparison estimates of
liverwort species richness have varied considerably, by as much as 50%, with esti-
mates ranging from 4,500 to 9,000 (e.g. Pearson 1995, Forrest et al. 2006). In the
last decade or so estimates in the range of 5,000 to 6,000 liverwort species have been
widely accepted (e.g. Gradstein and Costa 2003, Gradstein and Ilkiu-Borges 2009,
Heinrichs et al. 2007). von Konrat et al. (2010a) provided a mean estimate of 7,500
Figure 1. Habit images of selected lineages of Anthocerotophyta (hornworts) and Marchantiophyta
(liverworts). A Phaeomegaceros squamuliger, photo by J. Hollinger B Haplomitrium mnioides, photo by L.
Zhang C Conocephalum japonicum, photo by D. G. Long D Colura vitiensis, photo by T. Pócs E Pleurozia
gigantea, photo by L. Söderström F Riccardia spegazziniana, photo by Juan Larraín.
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
6
for the number of liverwort species based on estimating rates of synonymy in a sample
of recently monographed and revised taxa. A standardized global worldwide liverwort
checklist with strong community participation, as presented here, coupled with the
critical need for ongoing monographs and revisions, will aid in arriving at clearer es-
timates of liverwort and hornwort diversity. Signicantly, a list of standardized names
reviewed by taxonomic experts will enable more meaningful geographical and biogeo-
graphical species comparative studies. e global checklist of accepted liverwort and
hornwort taxa has vast potential, not only in aiding our understanding of liverwort
diversity, patterns, and processes, but also to the broader biological community (von
Konrat et al. 2008a).
Below we provide context to the worldwide checklist, including a brief histori-
cal account and current informational resources, structure and layout, systematic and
classication concepts, and methodology, including a brief overview of the underlying
data set and nomenclatural elements.
Major historical works
Schuster (1966d) has provided a detailed account on the history of hepaticology, up
to the rst half of the 20th century. Earlier, Verdoorn (1934c) also outlined a valuable
historical review of important gures in hepaticology. In the early 19th century, pos-
sibly the two most inuential persons who provided an early framework for taxonomic
concepts were W.J. Hooker and G. Raddi. During the middle of the 19th century
C.G. Nees von Esenbeck, C.M. Gottsche and J.B.W. Lindenberg provided vastly sig-
nicant contributions towards developing a classication framework. Frahm and Egg-
ers (2001) provided a brief biography of these workers. is was later followed by R.
Spruce and the Austrian-born H. Leitgeb in the second half 19th century. Leitgeb in
particular was instrumental in liverwort plant morphology and anatomy inclu ding a
six part treatise, “Untersuchungen ueber die Lebermoose”, from 1874 to 1881. To-
wards the end of 19th century and early 20th century V. Schiner, F. Stephani, and A.
Evans were prominent hepaticologists. In the rst half of the 20th century A. Evans,
H. Buch, F. Verdoorn, and K. Müller (Frib.) were leading inuential hepaticologists.
In the second half of the 20th century, R. Grolle and R.M. Schuster were of extraordi-
nary inuence in working towards contemporary classication schemes. Many of these
groundbreaking concepts still serve the foundation from which our current classica-
tion scheme is derived. R.M. Schuster’s major contribution to botany and hepaticology
lies in the astounding new diversity of liverworts he added to our knowledge. He de-
scribed a staggering 463 species, 83 genera and 15 families new to science. It is dicult
to name another contemporary botanist who discovered this much new diversity of a
major clade of land plants (Qiu et al. 2013). R.M. Schuster’s contribution to botany
went beyond hepatics. He was one of the rst botanists who recognized the importance
of Wallace’s Line in plant biogeography, separating Australia of Gondwanaland from
Southeast Asia of Laurasia (Cronquist 1988).
World checklist of hornworts and liverworts 7
Figure 2 depicts the number of novel liverwort species, excluding new combina-
tions that have been described over the last 250 years. e rst major peak corresponds
to the works of several early 19th century botanists, including Synopsis Hepaticarum
by Gottsche, Lindenberg, and Nees (1844–1847). e three decades leading into, but
prior to the highest peak, between 1860 and 1890, correspond to publications by a
number of prominent bryologists including W. Mitten, J.D. Hooker, T. Taylor, and
V. Schiner. e second and highest peak of almost 1,200 names, in the early 1900s
corresponds largely to the plethora of taxa described by F. Stephani (1898–1924) in
his monumental work Species Hepaticarum. e periods of highest rates of new spe-
cies described in the 1830s and around 1900 are the same for seed plants (Mutke and
Barthlott 2005). e third peak over the four decades between 1950 and 1980 can
be attributed mainly to the works of R.M. Schuster, H. Inoue, and S. Hattori. H.
Inoue and S. Hattori were inuential on large species-rich genera such as Plagiochila,
Frullania, and Porella. e decline in newly described species since 1970 does not
necessarily translate to the conclusion that taxonomists are closer to discovering all
known species. e almost 200 novel liverwort species that have been described in the
past six years alone still represent a signicant number, considering the relatively few
liverwort taxonomists and monographers. Moreover, bryological exploration has been
very uneven in many parts of the world. For example, many areas of the Neotropics
still remain without a single bryophyte record (Gradstein et al. 2001a). Paradoxically,
scores of new species are still being discovered and described in relatively well-studied
Figure 2. Number of novel liverwort species, excluding new combinations, which have been described
over the last 250 years, with an inset of the number described from 2001–2012.
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
8
areas such as New Zealand, e.g. over 50 new taxa since 2000 alone (e.g. Engel and
Schuster 2001, Engel and Glenny 2008a, Renner and de Lange 2011, von Konrat et
al. 2012b). Recent attention to cryptic speciation in bryophytes is also revealing novel
liverwort species (e.g. Szweykowski et al. 2005). e combination of collecting in yet-
to-be explored areas, the continued discovery of species in well-studied regions, and
an increased understanding of the biology of liverworts (including cryptic speciation),
will lead to a signicant number of newly discovered species in the foreseeable future.
e corollary of this, coupled with increased monographic and revisional work, will be
the increased discovery and the unravelling of synonymy.
Contemporary resources
Data availability and information needs associated with liverworts and hornworts have
been reviewed extensively by von Konrat et al. (2008a, 2010a). A fundamental prob-
lem common to all nomenclatural indexing projects is the dispersed nature of the bio-
logical literature, some of which may date back over 250 years (Lughadha 2004). Our
own data records, from 1990 to the present alone, indicate there are over 190 periodi-
cal titles and non-serials in which liverwort nomenclatural novelties were published.
e most useful and successful web-accessible bryophyte nomenclatural database is
Tropicos (http://www.tropicos.org), which oers name data with references and type
information as well as links to specimen data of its holdings. Tropicos has therefore be-
come an indispensable reference for anyone dealing with bryophyte names. However,
as stated above, the database is particularly strong for mosses and far less comprehen-
sive for liverworts. Moreover, Tropicos does not prescribe taxonomic disposition of
names, i.e., there is no attempt to adopt a single consistent view on the status of any
particular name (Lughadha 2004, von Konrat et al. 2008a).
A major nomenclatural work is the Index Hepaticarum (http://www.ville-ge.ch/
musinfo/bd/cjb/hepatic/index.php), which includes all eectively published liverwort
epithets spanning 12 volumes with the closing date of 1973. e indices were pre-
pared as a purely nomenclatural resource and did not claim to express any particular
taxonomic concept. Recently, Crosby and Engel (2006) provided an equally valuable
nomenclatural resource and catalog of names at all ranks for liverworts and hornworts
published during 1974 to 2000. Subsequently, indices of the citations for names pub-
lished for bryophytes have been compiled for the years 2001-2004 (Crosby and Magill
2005) and 2005 (Crosby and Magill 2006). e Early Land Plants Today initiative has
continued this series (von Konrat et al. 2010c, Söderström et al. 2012b, 2014).
e application of molecular phylogenetics continues to generate new insights into
the evolutionary history of liverworts and hornworts. In recent years, inferences made
from these phylogenies have especially revolutionized liverwort classication (Crandall-
Stotler et al. 2009). Yet, the reconstruction of the phylogenetic history of the Marchan-
tiophyta and Anthocerotophyta remains an ongoing eort and the classi cation is u-
id (von Konrat et al. 2010a). e current checklist largely follows the comprehensive
World checklist of hornworts and liverworts 9
phylogenetic liverwort classication scheme provided by Crandall- Stotler et al. (2009)
and the hornwort classication scheme of Stotler and Crandall-Stotler (2005). us
the checklist largely adheres to their higher classication namely from class to subor-
der, and the majority of family and genera arrangements provided therein. However,
there are some notable exceptions, especially in the systematic treatment of liverworts,
which largely either reect publications subsequent to 2009, or are a slight departure
in taxonomic opinion based on earlier works, e.g. the concepts of Saccogynaceae and
Stephaniellaceae depart from Crandall-Stotler et al. (2009). Since 2010 there have
been a number of realignments and novel hypotheses proposed that were generated
from conclusions based on molecular evidence which have inuenced the systematic
treatment presented here. ese include, for example, Adelanthaceae (e.g. Konstanti-
nova and Vilnet 2009, Feldberg et al. 2010b, 2010a, Söderström et al. 2010b, Vilnet
et al. 2010), Anastrophyllaceae (e.g. Söderström et. al 2010b), Gymnomitriaceae (e.g.
Vilnet et al 2007a, 2010, Váňa et al. 2010b, Shaw et al. 2015), Lepidoziaceae (Cooper
2013), Lophocoleaceae (Söderström et al. 2013b), and Scapaniaceae (e.g. Vilnet et al.
2010, Heinrichs et al. 2012a). In all cases, any such departures, new families, or rea-
lignments are noted and very briey discussed throughout the checklist. As noted by
Crandall-Stotler et al. (2009) many small families still remain to be investigated using
molecular data, and many of the large families and genera that have been sampled ap-
pear to be either polyphyletic or paraphyletic.
e systematic treatment as outlined here also includes the addition of subfamily
and infrageneric classication; ranks not presented by Crandall-Stotler et al. (2008b,
2009). Although there are arguably a number of ambiguous placements, especially at
the infrageneric rank, and there remain many areas of contention, particularly at the
familial and generic level, the systematic treatment itself serves as an excellent synop-
sis of contemporary taxonomic opinion, including infrageneric ranks. e infrageneric
treatments will also aid future eorts in providing a taxonomic framework that will be
particularly useful for extremely large genera. In large part, the infrageneric treatments
only deal with those taxa that represent phylogenetic clades with strong support. Care
has been taken to reduce ambiguity and those taxa where infrageneric placement is weak
or without evidence are simply listed under the genus and not placed systematically.
Structure and format of the checklist
e checklist is presented in a taxonomic framework as outlined above with four main
sections. e rst is arranged systematically with taxa ordered alphabetically within
the nearest higher rank. A brief discussion for selected families is provided in the main
body of the text under the respective families. is does not appear uniformly for all
families, but typically for larger more complicated families. Rather, a brief description
is provided mainly where concepts depart from Crandall-Stotler et al. (2009) or if new
information has come to light since that publication. Subfamilies and infrageneric
ranks have been incorporated where appropriate. However, if the subgeneric place-
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
10
ment is unclear the species are listed under “Incertae sedis”. From the rank of genus and
below, all entries include a three star condence ranking (described below), accepted
taxon name, authority, abbreviated publication title, page number and date, which is
cross- referenced to the full citation in the reference list, as well as the basionym if ap-
propriate. A typical entry appears as follows:
*** Oleolophozia L.Söderstr., De Roo et Hedd., Phytotaxa 3: 50, 2010 (Söderström
et al. 2010b).
*** Oleolophozia perssonii (H.Buch et S.W.Arnell) L.Söderstr., De Roo et Hedd.,
Phytotaxa 3: 51, 2010 (Söderström et al. 2010b). Bas.: Lophozia perssonii
H.Buch et S.W.Arnell, Bot. Not. 97: 382, 1944 (Buch 1944).
ere are comprehensive footnotes throughout the systematic section of the check-
list. ese are applied to selected names representing the rank of genus and below only.
Footnotes without any literature citation represent observations from the authors of
the respective family. Footnotes are wide ranging in their content. Generally, footnotes
can be categorized into the following: i) general statements relating to taxonomic opi-
nion, ii) comments on the type specimen, iii) comments about possible conspecicity,
including citation of conicting opinions, and iv) comment about species complexes.
is section is followed by a list of taxa in genera that we do not recognize. ose
names represent taxa that have not been studied recently and have not been recognized
in any recent treatment. However, many of those names are old and may gain priority
over some recognized taxa once their identity is determined.
e next section is an alphabetical list of taxa of the ranks of species and below. e
alphabetical list includes the condence level, the taxon name and authority as well as
a reference to the page of the corresponding entry in the systematic section. us, the
alphabetical list provides a rapid gateway requiring no prior knowledge of higher taxo-
nomy, but also serves as an index to the corresponding name in the systema tic section
that includes more detailed information.
e last section is a reference list where full bibliographic citations are given to all
references for taxa included in the checklist. Titles and references have all been checked
and veried, except in a few cases where we could not get hold of the publications.
Taxon confidence levels
Signicantly, each accepted taxon is qualied using a three level ranking system that
summarizes our knowledge about a taxon. e coding convention we are adopting
largely follows that described by von Konrat et al. (2010d) using one to three stars,
which has been applied to recent regional checklists produced by the Early Land Plants
Today initiative, e.g. Söderström et al. (2010a, 2011a, 2011b, 2012g, 2013e). e
conventions are briey outlined below coupled with samples illustrating how these
conventions are applied in practice. e application of a condence level to a taxon’s
World checklist of hornworts and liverworts 11
status and whether it represents a genuine “species” that is reached through community
consensus may go towards rening species estimates using an evidence-based approach
(von Konrat et al. 2008a, 2008b). e condence levels coupled with the detailed
annotations in the form of footnotes may also provide a rapid assessment of taxa and
help aid and drive future research into specic taxonomic or nomenclatural problems
and issues.
e coding convention:
* Serious doubts. ere are doubts about the value of the taxon. It can be that there
are conicting views without any substantial evidence in any direction, conicting
views with substantiating evidence supporting one or both positions, or evidence
pointing towards synonymization but it is premature to do it. We have adhered to
the principle that it is better to keep a taxon with one star (and preferably a note)
than to synonymize it too quickly. Example: Bazzania asymmetrica is conspecic
with Bazzania macgregorii in Grolle (1968a), but Kitagawa (1979a) kept them
separate. Example: Nardia kamtschatica may be conspecic with Nardia assamica
(Váňa 1976c), but the type specimen could not be studied.
** Knowledge problem. e taxon is not well known by the person evaluating it. It
may be a newly described species or a species originally not well described and not
restudied recently. Example: Jungermannia erectii Ajit P.Singh et V.Nath was re-
cently described and has not been independently studied by someone with a global
overview. It is therefore dicult to evaluate.
*** Accepted. A good taxon as currently understood based on personal experience or
on taxonomic revisions that have been convincingly performed. Nomenclature
and/or taxonomic position may, however, be questioned. Elements may be ex-
cluded from the taxon, but the taxon with the current type will still be accepted.
Example: H. Bischler-Causse revised the genus Marchantia worldwide in a series
of publications. Although she had a broad species concept, she also recognized
infraspecic taxa. e taxa that she recognized without doubt should be accepted
unless new evidence against it exists. Adopting a narrower species concept so that
many of her subspecies are elevated to species does not change her view of what a
good taxon is.
Methodology
e foundation of the checklist is in the underlying data set from which it is de-
rived. It was briey described by von Konrat et al. (2008a, 2010a). e working
data sets now includes a bibliography of 25,000 publications; approximately 39,000
published liverwort names (including “accepted” taxa, synonyms, invalid and ille-
gitimate names). e data quality and standards were outlined by von Konrat et al.
(2010d). In summary, for authorities and for the citation abbreviations, we follow
the standards set by the on-line version of Authors of Plant Names at the Royal Bota-
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
12
nical Gardens, Kew Website (http://www.ipni.org), with whom we collaborate close-
ly and provide with updated data records. Publications and journal abbreviations
follow Taxonomic Literature: A selective guide to botanical publications and collections
with dates, commentaries and types (Staeu and Cowan 1976–2009) (http://www.
sil.si.edu/digitalcollections/tl- 2/search.cfm) and Botanico Periodicum Huntianum
Lawrence et al. (1968) (http://huntbot.andrew.cmu.edu/HIBD/Departments/Data-
bases.shtml).
ere has been an intense systematic eort focusing on data quality. In all but a
few cases nomenclatural data has been veried against original publications. Söder-
ström and Hagborg have checked and conrmed almost every original publication
(three publications have not been available) for correct author, title and journal/
book citation, date of publication as far as possible, page number for the protologue,
and if the name is validly and legitimately published according to the International
Code of Nomenclature for algae, fungi and plants (Melbourne Code; McNeill et al.
2012). Most signicantly, the checklist has been community driven and collabora-
tive. Broader participation by taxonomic specialists and regional experts has lead to
the checklist containing high quality data (cf. Söderström et al. 2008). Development
of the checklist has included three international meetings (the rst one in 2009) and
generated 74 published notes on taxonomy and nomenclature under the auspices of
the Early Land Plants Today initiative from 2012 to 2015. e series of meetings
were instrumental in providing a framework for direct interaction with taxonomic
experts, workshops reviewing names, and helping identify potential participants who
were taxonomic experts in specic taxonomic groups, whether these were individual
genera or entire families.
e notes, published in the journal Phytotaxa, have included 38 authors from
13 countries. ese are the results of several years of revisions of taxonomic groups
by taxonomic experts. Following these revisions and working with participants iden-
tifying nomenclatural and taxonomic problems that would impact the worldwide
checklist, Söderström, Hagborg and von Konrat led editorial eorts in compiling
manuscripts with baseline information and nomenclatural or taxonomic issues that
required resolving. e work was then coordinated with the taxonomic experts who
drove and led the process and often identied further issues. e series of notes
on taxonomy and nomenclature of liverworts and hornworts provided updated no-
menclature, corrected invalid and illegitimate names and described new taxa based
on studies (mainly molecular) that did not draw nomenclatural conclusions. ose
notes are all open access and any changes were eective immediately as the Mel-
bourne Code of Botanical Nomenclature allows publishing on internet from 1 Jan.
2012 (Knapp et al. 2011).
is checklist builds on all published taxonomic and nomenclatural papers, that
have come to our attention until June 30, 2015.
World checklist of hornworts and liverworts 13
Summary statistics
We here present the rst-ever worldwide checklist for liverworts (Marchantiophyta)
and hornworts (Anthocerotophyta) that includes 8,078 taxa (species and below) in
7,486 species representing 398 genera, 92 families, 20 orders and 7 classes from the
two phyla. e list includes 3,533 species with three stars, 2,988 species with two stars
and 915 species with one star. e checklist also has extra utility in that it contains
3,106 references in the bibliography that serve as a powerful bibliographic resource for
liverwort and hornwort systematic and taxonomic research.
Concluding remarks
e marked-up publication form of the current checklist by PhytoKeys provides a
virtual instrument with a linked environment both internally (e.g. within an article)
and externally (GBIF, IPNI, Tropicos, Wikispecies, etc.) that will undoubtedly help
accelerate taxonomic research. e published checklist was the rst phase in providing
a worldwide list of accepted names. e next phase is to establish a generally recog-
nized online repository to augment the huge underlying informational auxiliary data
of over 25,000 publications, almost 39,000 published names, and the over 700,000
geographical observations. Several features of Web-based technology make it an essen-
tial tool, including 1) a vehicle to facilitate data access, 2) to unify the vastly scattered
data on distributional information and nomenclature, 3) oering dynamic rather than
static data in a searchable forum. e broader accessibility to the wealth of auxiliary
data will help augment monographic and revisional work for many taxonomic groups,
aid in identifying the need for increased oristic and survey work in many regions
throughout the world, and have broad implications and applications beyond taxono-
mic research such as conservation science. e current Early Land Plants Today model
has strong participation from taxonomic experts and an online resource will provide an
opportunity to expand stakeholders to include ecologists, conservationists, scientists
from other disciplines and general interest groups. However, such an eort can only be
successful if it comes with sustained funding and infrastructure rather than depending
on an ad hoc commitment by a few individuals, however dedicated.
e project will help augment monographic and revisional work for many taxo-
nomic groups and aid in identifying the need for increased oristic and survey work
in many regions throughout the world. Although there are many challenges ahead to
obtain high quality data, quantifying global liverwort diversity is a tractable, multi-
faceted and scientically important goal, and everyone stands to gain by fostering this
endeavour. e success of the project will lie on strong collaboration between institu-
tions and the bryological community in general.
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
14
Taxonomic synopsis
Anthocerotophyta ................................................................................................. 27
Anthocerotopsida de Bary ex Jancz. .................................................................... 27
Anthocerotidae Rosenv. ................................................................................. 27
Anthocerotales Limpr. ............................................................................... 27
Anthocerotaceae Dumort. ...................................................................... 27
Anthoceros L. ................................................................................ 27
Folioceros D.C.Bharadwaj ........................................................... 31
Dendrocerotidae R.J.Du .............................................................................. 32
Dendrocerotales Hässel .............................................................................. 32
Dendrocerotaceae J.Haseg. .................................................................... 32
Dendroceros Nees ......................................................................... 32
Megaceros Campb. ....................................................................... 27
Nothoceros (R.M.Schust.) J.Haseg............................................... 35
Phaeomegaceros R.J.Du ............................................................. 35
Phymatocerotales R.J.Du ......................................................................... 36
Phymatocerotaceae R.J.Du .................................................................. 36
Phymatoceros Stotler .................................................................... 36
Notothylatidae R.J.Du ................................................................................. 36
Notothyladales Hyvönen et Piippo ............................................................ 36
Notothyladaceae Müll.Frib. ex Prosk. .................................................... 36
Notothyladoideae Grolle ................................................................... 37
Notothylas Sull. ex A.Gray ........................................................... 37
Phaeocerotoideae Hässel .................................................................... 38
Mesoceros Piippo .......................................................................... 38
Paraphymatoceros Hässel ............................................................ 38
Phaeoceros Prosk. ........................................................................ 38
Leiosporocerotopsida Stotler et Crand.-Stotl. ................................................. 41
Leiosporocerotales Hässel ........................................................................... 41
Leiosporocerotaceae Hässel ex Ochyra ................................................... 41
Leiosporoceros Hässel ................................................................... 41
Marchantiophyta ........................................................................................... 41
Haplomitriopsida Stotler et Crand.-Stotl. ...................................................... 41
Haplomitriidae Stotler et Crand.-Stotl. ...................................................... 41
Calobryales Hamlin ............................................................................... 41
Haplomitriaceae Dědeček .................................................................. 41
Haplomitrium Nees ..................................................................... 41
Treubiidae Stotler et Crand.-Stotl. ............................................................. 42
Treubiales Schljakov .............................................................................. 42
Treubiaceae Verd. .............................................................................. 42
Apotreubia S.Hatt. et Mizut. ....................................................... 42
Treubia K.I.Goebel ...................................................................... 43
World checklist of hornworts and liverworts 15
Jungermanniopsida Stotler et Crand.-Stotl. ........................................................ 43
Jungermanniidae Engl. ................................................................................... 43
Jungermanniales H.Klinggr........................................................................ 43
Cephaloziineae Schljakov ....................................................................... 43
Adelanthaceae Grolle ......................................................................... 43
Adelanthoideae K.Feldberg ............................................................ 43
Adelanthus Mitt. ...................................................................... 44
Pseudomarsupidium Herzog ................................................... 44
Wettsteinia Schin. ................................................................. 45
Jamesonielloideae Inoue ................................................................ 45
Cuspidatula Steph. .................................................................. 45
Denotarisia Grolle ................................................................... 46
Nothostrepta R.M.Schust. ....................................................... 46
Pisanoa Hässel ......................................................................... 46
Protosyzygiella (Inoue) R.M.Schust. ....................................... 46
Syzygiella Spruce...................................................................... 46
Vanaea (Inoue et Gradst.) Inoue et Gradst. ........................... 49
Anastrophyllaceae L.Söderstr. ............................................................ 49
Anastrepta (Lindb.) Schin. .................................................... 50
Anastrophyllum (Spruce) Steph. .............................................. 50
Barbilophozia Loeske .............................................................. 51
Biantheridion (Grolle) Konstant. et Vilnet ............................ 52
Chandonanthus Mitt. .............................................................. 52
Crossocalyx Meyl. ..................................................................... 52
Gymnocolea (Dumort.) Dumort. ............................................ 52
Hamatostrepta Váňa et D.G.Long .......................................... 52
Hattoria R.M.Schust. .............................................................. 53
Isopaches H.Buch ..................................................................... 53
Neoorthocaulis L.Söderstr. ...................................................... 53
Orthocaulis H.Buch ................................................................. 53
Plicanthus R.M.Schust. ........................................................... 54
Schizophyllopsis Váňa et L.Söderstr. ....................................... 54
Schljakovia Konstant. et Vilnet ............................................... 54
Schljakovianthus Konstant. et Vilnet ...................................... 55
Sphenolobopsis R.M.Schust. et N.Kitag. ................................. 55
Sphenolobus (Lindb.) Berggr. .................................................. 55
Tetralophozia (R.M.Schust.) Schljakov .................................. 55
Zantenia (S.Hatt.) Váňa et J.J.Engel ....................................... 56
Cephaloziaceae Mig. .......................................................................... 56
Alobielloideae R.M.Schust. ............................................................ 56
Alobiella (Spruce) Schin. ....................................................... 56
Alobiellopsis R.M.Schust. ........................................................ 56
Cephalozioideae Müll.Frib. ........................................................... 57
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16
Cephalozia (Dumort.) Dumort. .............................................. 57
Fuscocephaloziopsis Fulford .................................................... 59
Nowellia Mitt. .......................................................................... 61
Odontoschismatoideae H.Buch ex Grolle ...................................... 62
Odontoschisma (Dumort.) Dumort. ....................................... 62
Schinerioideae R.M.Schust. ......................................................... 64
Schineria Steph...................................................................... 64
Trabacelluloideae R.M.Schust. ...................................................... 64
Haesselia Grolle et Gradst. ...................................................... 64
Trabacellula Fulford ............................................................... 64
Cephaloziellaceae Douin .................................................................... 64
Allisoniella E.A.Hodgs. ............................................................ 65
Amphicephalozia R.M.Schust.................................................. 65
Anastrophyllopsis (R.M.Schust.) Váňa et L.Söderstr. ............. 65
Cephalojonesia Grolle .............................................................. 66
Cephalomitrion R.M.Schust. ................................................... 66
Cephaloziella (Spruce) Schin. ............................................... 66
Cephaloziopsis (Spruce) Schin. ............................................. 72
Chaetophyllopsis R.M.Schust. ................................................. 72
Cylindrocolea R.M.Schust. ...................................................... 72
Gottschelia Grolle .................................................................... 74
Gymnocoleopsis (R.M.Schust.) R.M.Schust. ........................... 74
Herzogobryum Grolle .............................................................. 74
Kymatocalyx Herzog ................................................................ 74
Lophonardia R.M.Schust. ....................................................... 75
Nothogymnomitrion R.M.Schust. ............................................ 75
Obtusifolium S.W.Arnell ......................................................... 75
Oleolophozia L.Söderstr. ......................................................... 75
Phycolepidozia R.M.Schust. .................................................... 75
Protolophozia (R.M.Schust.) Schljakov .................................. 75
Lophoziaceae Cavers .......................................................................... 77
Andrewsianthus R.M.Schust. .................................................. 77
Gerhildiella Grolle ................................................................... 78
Heterogemma (Jørg.) Konstant. et Vilnet ............................... 78
Lophozia (Dumort.) Dumort. ................................................. 79
Lophoziopsis Konstant. et Vilnet ............................................. 80
Pseudocephaloziella R.M.Schust. ............................................ 81
Trilophozia (R.M.Schust.) Bakalin ......................................... 81
Tritomaria Schin. ex Loeske ................................................. 82
Scapaniaceae Mig............................................................................... 82
Diplophyllum (Dumort.) Dumort. .......................................... 82
Douinia (C.E.O.Jensen) H.Buch ............................................ 84
Pseudotritomaria Konstant. et Vilnet ..................................... 84
World checklist of hornworts and liverworts 17
Saccobasis H.Buch ................................................................... 85
Scapania (Dumort.) Dumort. ................................................. 85
Schistochilopsis (N.Kitag.) Konstant. ...................................... 92
Jungermanniineae R.M.Schust. ex Stotler et Crand.-Stotl. ..................... 93
Acrobolbaceae E.A.Hodgs.................................................................. 93
Enigmella G.A.M.Scott et K.G.Beckm. ................................... 93
Acrobolboideae R.M.Schust. ex Briscoe ......................................... 93
Acrobolbus Nees ....................................................................... 93
Austrolophozioideae R.M.Schust. ex Crand.-Stotl. ........................ 96
Austrolophozia R.M.Schust. .................................................... 96
Goebelobryum Grolle ............................................................... 97
Lethocoleoideae Grolle .................................................................. 97
Lethocolea Mitt. ....................................................................... 97
Saccogynidioideae Crand.-Stotl. .................................................... 97
Saccogynidium Grolle .............................................................. 97
Antheliaceae R.M.Schust. .................................................................. 98
Anthelia (Dumort.) Dumort. .................................................. 98
Arnelliaceae Nakai ............................................................................. 99
Arnellia Lindb. ........................................................................ 99
Balantiopsidaceae H.Buch ................................................................. 99
Acroscyphella N.Kitag. et Grolle .............................................. 99
Pseudoisotachis Váňa ............................................................... 99
Balantiopsidoideae J.J.Engel et Váňa ............................................. 99
Balantiopsis Mitt. .................................................................... 99
Isotachidoideae Grolle ................................................................. 101
Isotachis Mitt. ........................................................................ 101
Neesioscyphus Grolle .............................................................. 102
Ruizanthoideae R.M.Schust. ex J.J.Engel et G.L.Merr. ................ 103
Ruizanthus R.M.Schust. ........................................................ 103
Blepharidophyllaceae R.M.Schust. ex J.J.Engel ................................ 103
Blepharidophyllum Ångstr. ................................................... 103
Clandarium (Grolle) R.M.Schust. ........................................ 103
Calypogeiaceae Arnell ...................................................................... 103
Calypogeia Raddi ................................................................... 103
Eocalypogeia (R.M.Schust.) R.M.Schust. .............................. 107
Metacalypogeia (S.Hatt.) Inoue ............................................. 107
Mizutania Furuki et Z.Iwats. ................................................ 107
Mnioloma Herzog .................................................................. 107
Endogemmataceae Konstant. ........................................................... 109
Endogemma Konstant. ........................................................... 109
Geocalycaceae H.Klinggr. ................................................................ 109
Geocalyx Nees ........................................................................ 109
Gymnomitriaceae H.Klinggr. .......................................................... 109
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Acrolophozia R.M.Schust. ..................................................... 110
Nanomarsupella R.M.Schust. ex A.Hagborg ........................ 110
Paramomitrion R.M.Schust. ................................................. 110
Gymnomitrioideae T.Jensen ........................................................ 110
Cryptocoleopsis Amakawa ...................................................... 110
Gymnomitrion Corda ............................................................ 110
Marsupella Dumort. .............................................................. 112
Poeltia Grolle ......................................................................... 114
Prasanthus Lindb. ................................................................. 114
Nardioideae Váňa ........................................................................ 114
Nardia Gray ........................................................................... 114
Gyrothyraceae R.M.Schust. ............................................................. 115
Gyrothyra M.Howe................................................................ 115
Harpanthaceae Arnell ...................................................................... 116
Harpanthus Nees ................................................................... 116
Hygrobiellaceae Konstant. et Vilnet ................................................. 116
Hygrobiella Spruce ................................................................ 116
Jackiellaceae R.M.Schust. ................................................................ 116
Jackiella Schin. .................................................................... 116
Jungermanniaceae Rchb. ................................................................. 117
Delavayelloideae Grolle ............................................................... 117
Delavayella Steph. ................................................................. 117
Liochlaena Nees ..................................................................... 117
Jungermannioideae Dumort. ....................................................... 118
Eremonotus Lindb. et Kaal. ex Pearson ................................ 118
Jungermannia L. .................................................................... 118
Mesoptychioideae R.M.Schust. ................................................... 120
Mesoptychia (Lindb.) A.Evans ............................................... 120
Rivulariella D.H.Wagner ...................................................... 122
Notoscyphaceae Crand.-Stotl. ......................................................... 122
Notoscyphus Mitt. .................................................................. 122
Saccogynaceae Heeg ........................................................................ 122
Saccogyna Dumort. ................................................................ 122
Solenostomataceae Stotler et Crand.-Stotl. ...................................... 123
Aponardia (R.M.Schust.) Váňa ............................................. 123
Arctoscyphus Hässel ............................................................... 123
Cryptocolea R.M.Schust. ....................................................... 123
Diplocolea Amakawa ............................................................. 123
Solenostoma Mitt. .................................................................. 123
Southbyaceae Váňa .......................................................................... 135
Gongylanthus Nees ................................................................ 135
Southbya Spruce .................................................................... 135
Stephaniellaceae R.M.Schust. .......................................................... 136
World checklist of hornworts and liverworts 19
Stephaniella J.B.Jack ............................................................. 136
Stephaniellidium S.Winkl. ex Grolle .................................... 136
Trichotemnomataceae R.M.Schust. ................................................. 136
Trichotemnoma R.M.Schust. ................................................. 136
Lophocoleineae Schljakov .................................................................... 137
Blepharostomataceae W.Frey et M.Stech ......................................... 137
Blepharostoma (Dumort.) Dumort. ...................................... 137
Brevianthaceae J.J.Engel et R.M.Schust. .......................................... 137
Brevianthus J.J.Engel et R.M.Schust. ................................... 137
Tetracymbaliella Grolle ......................................................... 137
Chonecoleaceae R.M.Schust. ex Grolle ............................................ 138
Chonecolea Grolle .................................................................. 138
Grolleaceae Solari ex R.M.Schust..................................................... 138
Grollea R.M.Schust. .............................................................. 138
Herbertaceae Müll.Frib. ex Fulford et Hatcher ................................ 138
Herbertus Gray ...................................................................... 138
Triandrophyllum Fulford et Hatcher .................................... 171
Lepicoleaceae R.M.Schust. ............................................................... 141
Lepicolea Dumort. ................................................................. 142
Vetaforma Fulford et J.Taylor ............................................... 142
Lepidoziaceae Limpr. ....................................................................... 142
Meinungeria Frank Müll. ...................................................... 143
Bazzanioideae Rodway ................................................................. 143
Acromastigum A.Evans .......................................................... 143
Bazzania Gray ....................................................................... 146
Mastigopelma Mitt. ................................................................ 163
Drucelloideae R.M.Schust. .......................................................... 163
Drucella E.A.Hodgs. .............................................................. 163
Lembidioideae R.M.Schust. ......................................................... 163
Dendrolembidium Herzog ..................................................... 163
Hygrolembidium R.M.Schust. ............................................... 164
Isolembidium R.M.Schust. .................................................... 164
Kurzia G.Martens .................................................................. 164
Lembidium Mitt. ................................................................... 168
Megalembidium R.M.Schust. ................................................ 168
Pseudocephalozia R.M.Schust. .............................................. 168
Lepidozioideae Müll.Frib. ........................................................... 169
Ceramanus E.D.Cooper ........................................................ 169
Lepidozia (Dumort.) Dumort. .............................................. 169
Neolepidozia Fulford et J.Taylor ........................................... 176
Tricholepidozia (R.M.Schust.) E.D.Cooper ......................... 179
Micropterygioideae Grolle ........................................................... 181
Micropterygium Gottsche ...................................................... 181
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Mytilopsis Spruce ................................................................... 182
Protocephalozioideae R.M.Schust. ............................................... 182
Protocephalozia (Spruce) K.I.Goebel .................................... 182
Zoopsidoideae R.M.Schust. ......................................................... 183
Amazoopsis J.J.Engel et G.L.Merr. ........................................ 183
Hyalolepidozia S.W.Arnell ex Grolle .................................... 183
Monodactylopsis (R.M.Schust.) R.M.Schust. ........................ 183
Neogrollea E.A.Hodgs. ........................................................... 183
Odontoseries Fulford ............................................................. 183
Paracromastigum Fulford et J.Taylor ................................... 183
Psiloclada Mitt. ...................................................................... 185
Pteropsiella Spruce ................................................................ 185
Telaranea Spruce ex Schin. ................................................. 185
Zoopsidella R.M.Schust. ........................................................ 187
Zoopsis Hook.f. ex Gottsche .................................................. 188
Lophocoleaceae Vanden Berghen ..................................................... 189
Bragginsella R.M.Schust........................................................ 189
Chiloscyphus Corda ................................................................ 189
Clasmatocolea Spruce ............................................................ 191
Conoscyphus Mitt. .................................................................. 193
Cryptolophocolea L.Söderstr. ................................................. 193
Deceptifrons J.J.Engel et Váňa ............................................... 196
Evansianthus R.M.Schust. et J.J.Engel ................................. 196
Hepatostolonophora J.J.Engel et R.M.Schust. ...................... 196
Heteroscyphus Schin. ........................................................... 197
Lamellocolea J.J.Engel ........................................................... 204
Leptophyllopsis R.M.Schust. .................................................. 204
Leptoscyphopsis R.M.Schust. ................................................. 204
Leptoscyphus Mitt. ................................................................. 204
Lophocolea (Dumort.) Dumort. ............................................ 208
Otoscyphus J.J.Engel, Bardat et ouvenot ........................... 215
Pachyglossa Herzog et Grolle ................................................ 216
Perdusenia Hässel .................................................................. 216
Pigafettoa C.Massal. .............................................................. 217
Platycaulis R.M.Schust. ......................................................... 217
Stolonivector J.J.Engel ........................................................... 217
Xenocephalozia R.M.Schust. ................................................. 217
Mastigophoraceae R.M.Schust. ........................................................ 217
Dendromastigophora R.M.Schust. ........................................ 217
Mastigophora Nees ................................................................ 217
Plagiochilaceae Müll.Frib................................................................. 218
Acrochila R.M.Schust. ........................................................... 218
Chiastocaulon Carl ................................................................ 219
World checklist of hornworts and liverworts 21
Dinckleria Trevis. .................................................................. 219
Pedinophyllopsis R.M.Schust. et Inoue ................................. 219
Pedinophyllum Lindb. ex Nordst. ......................................... 219
Plagiochila (Dumort.) Dumort. ............................................ 219
Plagiochilidium Herzog ......................................................... 252
Plagiochilion S.Hatt............................................................... 252
Pseudolophocolea R.M.Schust. et J.J.Engel ........................... 253
Xenochila R.M.Schust. .......................................................... 253
Pseudolepicoleaceae Fulford et J.Taylor ........................................... 253
Archeophylla R.M.Schust....................................................... 253
Castanoclobos J.J.Engel et Glenny ........................................ 254
Chaetocolea Spruce ................................................................ 254
Herzogiaria Fulford ex Hässel............................................... 254
Isophyllaria E.A.Hodgs. et Allison ........................................ 254
Pseudolepicolea Fulford et J.Taylor ...................................... 254
Temnoma Mitt. ...................................................................... 255
Trichocoleaceae Nakai ..................................................................... 256
Eotrichocolea R.M.Schust. ..................................................... 256
Leiomitra Lindb. ................................................................... 256
Trichocolea Dumort. ............................................................. 257
Myliineae J.J.Engel et Braggins ex Crand.-Stotl. .................................. 258
Myliaceae Schljakov ......................................................................... 258
Mylia Gray ............................................................................. 258
Perssoniellineae R.M.Schust. ............................................................... 259
Schistochilaceae H.Buch .................................................................. 259
Schistochila Dumort. ............................................................. 259
Porellales Schljakov .................................................................................. 264
Jubulineae Müll.Frib. .......................................................................... 264
Frullaniaceae Lorch.......................................................................... 264
Frullania Raddi ..................................................................... 264
Jubulaceae H.Klinggr....................................................................... 295
Jubula Dumort. ..................................................................... 295
Neohattoria Kamim. .............................................................. 296
Nipponolejeunea S.Hatt. ....................................................... 296
Lejeuneaceae Cavers ........................................................................ 297
Lejeuneoideae ............................................................................. 297
trib. Brachiolejeuneeae ................................................................ 297
subtrib. Brachiolejeuneinae Gradst. ............................................. 297
Acanthocoleus R.M.Schust. .................................................... 297
Blepharolejeunea S.W.Arnell ................................................ 298
Brachiolejeunea (Spruce) Schin. ......................................... 298
Dicranolejeunea (Spruce) Schin. ........................................ 299
Lindigianthus Kruijt et Gradst. ............................................. 299
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22
Odontolejeunea (Spruce) Schin. ......................................... 299
subtrib. Stictolejeuneinae Gradst. ................................................ 299
Neurolejeunea (Spruce) Schin. ............................................ 299
Stictolejeunea (Spruce) Schin. ............................................ 300
trib. Lejeuneeae Dumort. ............................................................ 300
Dactylophorella R.M.Schust. ................................................. 300
Metalejeunea Grolle .............................................................. 300
Pictolejeunea Grolle .............................................................. 300
subtrib. Ceratolejeuneinae Gradst. ............................................... 301
Ceratolejeunea (Spruce) J.B.Jack et Steph. ........................... 301
Luteolejeunea Piippo ............................................................. 304
Otigoniolejeunea (Spruce) Schin. ....................................... 304
subtrib. Cheilolejeuneinae Gradst. ............................................... 304
Aureolejeunea R.M.Schust. ................................................... 304
Cheilolejeunea (Spruce) Steph. ............................................. 305
Cyrtolejeunea A.Evans ........................................................... 316
Cystolejeunea A.Evans ............................................................ 316
Omphalanthus Lindenb. et Nees ........................................... 316
subtrib. Cololejeuneinae Gradst. ................................................. 317
Aphanotropis Herzog ............................................................. 317
Calatholejeunea K.I.Goebel .................................................. 317
Cololejeunea (Spruce) Steph. ................................................ 317
Colura (Dumort.) Dumort. ................................................... 340
Diplasiolejeunea (Spruce) Schin. ........................................ 344
Haplolejeunea Grolle ............................................................. 348
Macrocolura R.M.Schust. ...................................................... 348
Myriocoleopsis Schin. .......................................................... 348
Nephelolejeunea Grolle .......................................................... 349
Schusterolejeunea Grolle ....................................................... 349
Siphonolejeunea Herzog ........................................................ 350
Tuyamaella S.Hatt. ................................................................ 350
subtrib. Cyclolejeuneinae Gradst. ................................................ 350
Bromeliophila R.M.Schust. ................................................... 350
Cyclolejeunea A.Evans ........................................................... 351
Prionolejeunea (Spruce) Schin. ........................................... 351
subtrib. Drepanolejeuneinae Gradst. ........................................... 353
Capillolejeunea S.W.Arnell ................................................... 353
Drepanolejeunea (Spruce) Steph. .......................................... 353
Vitalianthus R.M.Schust. et Giancotti .................................. 361
subtrib. Echinolejeuneinae Gradst. .............................................. 361
Anoplolejeunea (Spruce) Schin. .......................................... 361
Echinolejeunea R.M.Schust. .................................................. 362
Kymatolejeunea Grolle .......................................................... 362
World checklist of hornworts and liverworts 23
subtrib. Leiolejeuneinae Schäf.-Verw. et Heinrichs ...................... 362
Leiolejeunea A.Evans ............................................................. 362
subtrib. Lejeuneinae Gradst. ........................................................ 362
Harpalejeunea (Spruce) Schin. ........................................... 362
Hattoriolejeunea Mizut. ........................................................ 364
Lejeunea Lib. ......................................................................... 364
Microlejeunea (Spruce) Steph. .............................................. 385
Taxilejeunea (Spruce) Steph. ................................................ 388
subtrib. Lepidolejeuneinae Gradst. .............................................. 391
Lepidolejeunea R.M.Schust. .................................................. 391
Otolejeunea Grolle et Tixier .................................................. 392
Rectolejeunea A.Evans ........................................................... 393
subtrib. Leptolejeuneinae Heinrichs et Schäf.-Verw. .................... 394
Leptolejeunea (Spruce) Steph. ............................................... 394
subtrib. Pycnolejeuneinae Heinrichs et Schäf.-Verw. .................... 396
Pycnolejeunea (Spruce) Schifn. ........................................... 396
subtrib. Xylolejeuneinae Henrichs et Schäf.-Verw. ....................... 398
Xylolejeunea Xiao L.He et Grolle .......................................... 398
trib. Symbiezidieae Gradst. .......................................................... 398
Symbiezidium Trevis. ............................................................ 398
Ptychanthoideae Mizut. ............................................................... 399
Acrolejeunea (Spruce) Schin. .............................................. 399
Archilejeunea (Spruce) Steph. ............................................... 401
Bryopteris (Nees) Lindenb. .................................................... 403
Caudalejeunea Schin. .......................................................... 403
Cephalantholejeunea R.M.Schust. ........................................ 405
Cephalolejeunea Mizut. ......................................................... 405
Frullanoides Raddi ................................................................ 405
Fulfordianthus Gradst. .......................................................... 405
Lopholejeunea (Spruce) Steph. .............................................. 406
Marchesinia Gray .................................................................. 408
Mastigolejeunea (Spruce) Steph. ........................................... 409
Phaeolejeunea Mizut. ............................................................ 410
Ptychanthus Nees ................................................................... 411
Schineriolejeunea Verd. ....................................................... 411
Spruceanthus Verd. ................................................................ 412
ysananthus Lindenb. ......................................................... 413
Tuzibeanthus S.Hatt. ............................................................. 414
Verdoornianthus Gradst. ....................................................... 414
Porellineae R.M.Schust. ....................................................................... 415
Goebeliellaceae Verd. ....................................................................... 415
Goebeliella Steph. .................................................................. 415
Lepidolaenaceae Nakai .................................................................... 415
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Gackstroemia Trevis. ............................................................. 415
Lepidogyna R.M.Schust. ........................................................ 416
Lepidolaena Dumort. ............................................................ 416
Porellaceae Cavers nom. conserv. ....................................................... 417
Ascidiota C.Massal. ................................................................ 417
Porella L. ................................................................................ 417
Radulineae R.M.Schust. ...................................................................... 424
Radulaceae Müll.Frib. ..................................................................... 424
Radula Dumort. .................................................................... 425
Ptilidiales Schljakov ................................................................................. 434
Herzogianthaceae Stotler et Crand.-Stotl. ........................................ 434
Herzogianthus R.M.Schust. ................................................... 434
Neotrichocoleaceae Inoue ................................................................ 434
Neotrichocolea S.Hatt. ........................................................... 434
Trichocoleopsis S.Okamura ................................................... 434
Ptilidiaceae H.Klinggr. .................................................................... 434
Ptilidium Nees ....................................................................... 434
Metzgeriidae Barthol.-Began ........................................................................ 435
Metzgeriales Chalaud ............................................................................... 435
Aneuraceae H.Klinggr. .................................................................... 435
Aneura Dumort. .................................................................... 435
Lobatiriccardia (Mizut. et S.Hatt.) Furuki ........................... 437
Riccardia Gray ....................................................................... 438
Verdoornia R.M.Schust. ........................................................ 456
Metzgeriaceae H.Klinggr. ................................................................ 456
Metzgeria Raddi ..................................................................... 457
Steereella Kuwah. .................................................................. 461
Vandiemenia Hewson ........................................................... 461
Pleuroziales Schljakov .............................................................................. 462
Pleuroziaceae Müll.Frib. .................................................................. 462
Pleurozia Dumort. ................................................................ 462
Pelliidae He-Nygrén .................................................................................... 463
Fossombroniales Schljakov ....................................................................... 463
Calyculariineae He-Nygrén .................................................................. 463
Calyculariaceae He-Nygrén ............................................................. 463
Calycularia Mitt. ................................................................... 463
Fossombroniineae R.M.Schust. ex Stotler et Crand.-Stotl. ................... 463
Allisoniaceae Schljakov .................................................................... 463
Allisonia Herzog .................................................................... 463
Fossombroniaceae Hazsl. nom. conserv. ............................................ 463
Fossombronia Raddi .............................................................. 463
Petalophyllaceae Stotler et Crand.-Stotl. .......................................... 468
Petalophyllum Nees et Gottsche ............................................ 468
World checklist of hornworts and liverworts 25
Sewardiella Kashyap ............................................................. 468
Makinoiineae He-Nygrén .................................................................... 468
Makinoaceae Nakai ......................................................................... 468
Makinoa Miyake .................................................................... 468
Pallaviciniales W.Frey et M.Stech ............................................................ 468
Pallaviciniineae R.M.Schust. ................................................................ 468
Hymenophytaceae R.M.Schust. ....................................................... 468
Hymenophyton Dumort. ........................................................ 468
Moerckiaceae K.I.Goebel ex Stotler et Crand.-Stotl. ........................ 469
Hattorianthus R.M.Schust. et Inoue ..................................... 469
Moerckia Gottsche ................................................................. 469
Pallaviciniaceae Mig. ........................................................................ 469
Pallavicinioideae Mig. ex Grolle .................................................. 469
Jensenia Lindb. ...................................................................... 469
Pallavicinia Gray ................................................................... 470
Podomitrium Mitt. ................................................................ 471
Symphyogynoideae R.M.Schust. ex Grolle .................................. 471
Greeneothallus Hässel ............................................................ 471
Seppeltia Grolle ..................................................................... 471
Symphyogyna Nees et Mont. .................................................. 471
Symphyogynopsis Grolle......................................................... 473
Xenothallus R.M.Schust. ....................................................... 473
Sandeothallaceae R.M.Schust. ......................................................... 474
Sandeothallus R.M.Schust. .................................................... 474
Phyllothalliineae R.M.Schust. .............................................................. 474
Phyllothalliaceae E.A.Hodgs. ex T.Katag. ........................................ 474
Phyllothallia E.A.Hodgs. ....................................................... 474
Pelliales He-Nygrén ................................................................................. 474
Noterocladaceae W.Frey et M.Stech ................................................ 474
Noteroclada Taylor ex Hook.f. et Wilson ............................. 474
Pelliaceae H.Klinggr. ....................................................................... 475
Pellia Raddi ........................................................................... 475
Marchantiopsida Cronquist.............................................................................. 476
Blasiidae He-Nygrén .................................................................................... 476
Blasiales Stotler et Crand.-Stotl. ............................................................... 476
Blasiaceae H.Klinggr. ....................................................................... 476
Blasia L. ................................................................................. 476
Cavicularia Steph. ................................................................. 476
Marchantiidae Engl. ..................................................................................... 476
Lunulariales H.Klinggr. ........................................................................... 476
Lunulariaceae H.Klinggr. ................................................................. 476
Lunularia Adans. ................................................................... 476
Marchantiales Limpr. ............................................................................... 476
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Aytoniaceae Cavers .......................................................................... 476
Asterella P.Beauv. .................................................................. 477
Cryptomitrium Austin ex Underw. ....................................... 481
Mannia Corda ....................................................................... 481
Plagiochasma Lehm. .............................................................. 482
Reboulia Raddi ...................................................................... 483
Cleveaceae Cavers ............................................................................ 484
Athalamia Falc. ...................................................................... 484
Clevea Lindb. ......................................................................... 484
Peltolepis Lindb. .................................................................... 484
Sauteria Nees ......................................................................... 485
Conocephalaceae Müll.Frib. ex Grolle ............................................. 485
Conocephalum Hill ................................................................ 485
Corsiniaceae Engl. ........................................................................... 486
Corsinioideae Schin. ...................................................................... 486
Corsinia Raddi ....................................................................... 486
Cronisioideae R.M.Schust. .............................................................. 486
Cronisia Berk. ........................................................................ 486
Cyathodiaceae Stotler et Crand.-Stotl. ............................................. 486
Cyathodium Kunze ................................................................ 486
Dumortieraceae D.G.Long .............................................................. 487
Dumortiera Nees ................................................................... 487
Exormothecaceae Müll.Frib. ex Grolle ............................................. 487
Aitchisoniella Kashyap........................................................... 487
Exormotheca Mitt. ................................................................. 488
Stephensoniella Kashyap ....................................................... 488
Marchantiaceae Lindl. ..................................................................... 488
Bucegioideae R.M.Schust. ........................................................... 488
Bucegia Radian ...................................................................... 488
Marchantioideae Schin. ............................................................. 489
Marchantia L. ........................................................................ 489
Preissia Corda ........................................................................ 492
Monocleaceae A.B.Frank ................................................................. 492
Monoclea Hook. .................................................................... 492
Monosoleniaceae Inoue ................................................................... 492
Monosolenium Gri. .............................................................. 492
Oxymitraceae Müll.Frib. ex Grolle .................................................. 493
Oxymitra Bisch. ex Lindenb. ................................................. 493
Ricciaceae Rchb. .............................................................................. 493
Riccia L. ................................................................................. 493
Ricciocarpos Corda ................................................................ 503
Targioniaceae Dumort. .................................................................... 503
World checklist of hornworts and liverworts 27
Targionia L. ........................................................................... 503
Wiesnerellaceae Inoue ...................................................................... 504
Wiesnerella Schin. ............................................................... 504
Neohodgsoniales D.G.Long ..................................................................... 504
Neohodgsoniaceae D.G.Long .......................................................... 504
Neohodgsonia Perss. ............................................................... 504
Sphaerocarpales Cavers ............................................................................ 504
Monocarpaceae D.J.Carr ex Schelpe ................................................ 504
Monocarpus D.J.Carr ............................................................ 504
Riellaceae Engl. ................................................................................ 504
Austroriella Cargill et J.Milne ............................................... 504
Riella Mont. ........................................................................... 505
Sphaerocarpaceae Heeg .................................................................... 506
Geothallus Campb. ................................................................ 506
Sphaerocarpos Boehm. .......................................................... 506
Taxonomic list
ANTHOCEROTOPHYTA
ANTHOCEROTOPSIDA de Bary ex Jancz.
Anthocerotidae Rosenv.
Anthocerotales Limpr.
*** Anthocerotaceae Dumort.
by J.C. Villarreal and D.C. Cargill
Notes on nomenclature and taxonomy can also be found in Cargill et al. (2013b) and
Villarreal et al. (2015)
*** Anthoceros L., Sp. Pl. 1: 1139, 1753 (Linnaeus 1753). 1
*** Anthoceros adscendens Lehm. et Lindenb., Nov. Stirp. Pug. 4: 24, 1832 (Lehmann
1832).
*** Anthoceros agrestis Paton, J. Bryol. 10 (3): 257, 1979 (Paton 1979a), nom. conserv. 2
*** Anthoceros alpinus Steph., Sp. Hepat. (Stephani) 6: 425, 1923 (Stephani 1923).
1 Anthoceros includes Aspiromitus, but some taxa have neither been transferred nor synonymized. ey
are listed in the “Names in genera not currently accepted” section below.
2 Anthoceros agrestis is conserved against the earlier Anthoceros nagasakiensis Steph. 1916. Anthoceros
cristatus Steph. 1916 is also an earlier synonym and a proposal to conserve Anthoceros agrestis against
it is in progress.
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
28
* Anthoceros angustifolius Gottsche, Lindenb. et Nees, Syn. Hepat. 4: 582, 1846
(Gottsche et al. 1846). 3
*** Anthoceros angustus Steph., Sp. Hepat. (Stephani) 5: 1001, 1916 (Stephani 1916b).
*** Anthoceros bharadwajii Udar et A.K.Asthana, Proc. Indian Natl. Sci. Acad., B 51
(4): 484, 1985 (Udar and Asthana 1985b).
** Anthoceros buettneri Steph., Sp. Hepat. (Stephani) 5: 997, 1916 (Stephani 1916b).
*** Anthoceros capricornii Cargill et G.A.M.Scott, J. Hattori Bot. Lab. 82: 55, 1997
(Cargill and Scott 1997).
*** Anthoceros caucasicus Steph., Izv. Kavkazsk. Muz. 8: 87, 1914 (Voronov’ 1914).
** Anthoceros cavernosus Steph., Sp. Hepat. (Stephani) 5: 998, 1916 (Stephani
1916b).
* Anthoceros chambensis Kashyap, J. Bombay Nat. Hist. Soc. 25 (2): 281, 1917
(Kashyap 1917).
** Anthoceros chungii Khanna, J. Indian Bot. Soc. 17 (5/6): 316, 1938 (Khanna 1938).
** Anthoceros crispatus Gri., Not. pl. asiat. 2: 349, 1849 (Grith 1849).
** Anthoceros dimorphus Sim, Trans. Roy. Soc. South Africa 15 (1): 114, 1926 (Sim
1926).
*** Anthoceros erectus Kashyap, New Phytol. 14 (1): 9, 1915 (Kashyap 1915).
** Anthoceros expansus (Steph.) J.C.Villarreal et Cargill, Phytotaxa 208 (1): 92, 2015
(Villarreal et al. 2015). Bas.: Aspiromitus expansus Steph., Sp. Hepat. (Stephani) 5:
961, 1916 (Stephani 1916b).
* Anthoceros ferdinandi-muelleri Steph., Sp. Hepat. (Stephani) 5: 1007, 1916
(Stephani 1916b). 4
*** Anthoceros fragilis Steph., Sp. Hepat. (Stephani) 5: 1006, 1916 (Stephani 1916b).
*** Anthoceros fusiformis Austin, Bull. Torrey Bot. Club 6 (4): 28, 1875 [1876] (Austin
1875a).
*** Anthoceros fusiformis var. taiwanensis J.Haseg., Acta Phytotax. Geobot. 44 (2): 100,
1993 (Hasegawa 1993b).
** Anthoceros gasongorii Gola, Mem. Reale Accad. Sci. Torino (ser. 2) 65 (1): 10,
1916 (Gola 1916).
** Anthoceros granulatus Gottsche, Mexik. Leverm.: 275, 1863 (Gottsche 1863).
** Anthoceros harrisanus (Steph.) Parihar, Univ. Allahabad Stud., Bot. 1961-2: 31,
1962 (Parihar 1962). Bas.: Aspiromitus harrisanus Steph., Sp. Hepat. (Stephani) 5:
965, 1916 (Stephani 1916b).
* Anthoceros helmsii Steph., Hedwigia 32 (3): 142, 1893 (Stephani 1893b). 5
** Anthoceros jamesonii Taylor ex Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 171,
1855 (Mitten 1855).
3 Anthoceros angustifolius (type from Europe) has neither been recognized in any recent treatment nor
synonymized. e name may have priority once its identity is determined.
4 Anthoceros ferdinandi-muelleri is similar to Anthoceros punctatus, but Anthoceros still needs to be re-
vised for Australia.
5 Anthoceros helmsii was treated as conspecic with Anthoceros muscoides by Campbell (1982), but it is
in need of a revision.
World checklist of hornworts and liverworts 29
* Anthoceros javanicoides H.A.Mill., Phytologia 47 (4): 319, 1981 (Miller 1981).
Nom. nov. pro Anthoceros javanicus Steph., Sp. Hepat. (Stephani) 5: 988, 1916
(Stephani 1916b), nom. illeg. 6
** Anthoceros jungermannioides Schwein., Spec. Fl. Amer. Crypt.: 25, 1821 (Schweinitz
1821).
* Anthoceros kajumas (K.I.Goebel) Prosk., Bull. Torrey Bot. Club 78 (4): 347, 1951
(Proskauer 1951a). Bas.: Aspiromitus kajumas K.I.Goebel, Ann. Jard. Bot. Buiten-
zorg 39: 69, 1928 (Goebel 1928).
* Anthoceros koshii Khanna, J. Indian Bot. Soc. 15 (4): 237, 1936 (Khanna 1936).
*** Anthoceros lamellatus Steph., Sp. Hepat. (Stephani) 5: 1000, 1916 (Stephani
1916b).
*** Anthoceros laminifer Steph., J. Linn. Soc., Bot. 29 (201): 266, 1892 (Stephani 1892b).
*** Anthoceros macounii M.Howe, Bull. Torrey Bot. Club 25 (1): 19, 1898 (Howe
1898a).
*** Anthoceros macrosporus Steph., Sp. Hepat. (Stephani) 5: 1005, 1916 (Stephani 1916b).
** Anthoceros maritimus Steph., Sp. Hepat. (Stephani) 5: 984, 1916 (Stephani 1916b).
* Anthoceros megasporus Meijer, J. Hattori Bot. Lab. 18: 6, 1957 (Meijer 1957).
** Anthoceros muscoides Colenso, Trans. & Proc. New Zealand Inst. 16: 361, 1884
(Colenso 1884).
*** Anthoceros myriandroecius Steph., Wiss. Ergebn. Deut. Zentr.-Afr. Exped. (1907-
08), Bot. 2: 134, 1911 (Stephani 1911a).
** Anthoceros natalensis Steph., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr.
88: 732, 1913 (Stephani 1913b).
*** Anthoceros neesii Prosk., Leberm. Eur. 2 (9): 1312, 1958 (Proskauer 1958). Nom.
nov. pro Anthoceros punctatus α* monocarpus Nees, Naturgesch. Eur. Leberm. 4:
339, 1838 (Nees 1838a).
** Anthoceros niger Steph., Sp. Hepat. (Stephani) 5: 1005, 1916 (Stephani 1916b).
** Anthoceros orizabensis (Steph.) Hässel, Candollea 45 (1): 211, 1990 (Hässel
1990b). Bas.: Aspiromitus orizabensis Steph., Sp. Hepat. (Stephani) 5: 965, 1916
(Stephani 1916b).
*** Anthoceros pandei Udar et A.K.Asthana, J. Indian Bot. Soc. 64: 305, 1985 (Udar
and Asthana 1985a).
*** Anthoceros patagonicus Hässel, Candollea 45 (1): 207, 1990 (Hässel 1990b).
*** Anthoceros patagonicus subsp. gremmenii J.C.Villarreal, J.J.Engel et Váňa, Mem.
New York Bot. Gard. 105: 32, 2013 (Váňa and Engel 2013).
** Anthoceros peruvianus Steph., Sp. Hepat. (Stephani) 5: 999, 1916 (Stephani 1916b).
** Anthoceros pinnatus Steph., Bol. Soc. Brot. 4: 154 [182], 1885 [1886] (Stephani
1885c).
*** Anthoceros punctatus L., Sp. Pl. 1: 1139, 1753 (Linnaeus 1753). 7
6 Anthoceros javanicoides belongs to the Anthoceros punctatus species complex (Söderström et al. 2010a).
7 Anthoceros punctatus is a species complex in need of a revision.
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
30
* Anthoceros pusillus Colenso, Trans. & Proc. New Zealand Inst. 18: 255, 1886
(Colenso 1886b).
*** Anthoceros rosulans J.Haseg., J. Hattori Bot. Lab. 60: 379, 1986 (Hasegawa 1986b).
*** Anthoceros sambesianus Steph., Sp. Hepat. (Stephani) 5: 996, 1916 (Stephani 1916b).
*** Anthoceros scariosus Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 230, 1869 (Aus-
tin 1869).
** Anthoceros schroederi Steph., 52 (5): 306, 1912 (Stephani 1912a).
** Anthoceros serratus Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9):
90, 1911 (Stephani 1911b).
** Anthoceros simulans M.Howe, Proc. Calif. Acad. Sci. (ser. 4) 21 (17): 204, 1934
(Howe 1934).
** Anthoceros spongiosus Steph., Sp. Hepat. (Stephani) 5: 1003, 1916 (Stephani
1916b).
*** Anthoceros subtilis Steph., Sp. Hepat. (Stephani) 5: 1003, 1916 (Stephani 1916b).
*** Anthoceros telaganus Steph., Sp. Hepat. (Stephani) 5: 1005, 1916 (Stephani 1916b).
*** Anthoceros tristanianus J.C.Villarreal, J.J.Engel et Váňa, Mem. New York Bot.
Gard. 105: 33, 2013 (Váňa and Engel 2013).
*** Anthoceros tuberculatus Lehm. et Lindenb., Nov. Stirp. Pug. 4: 25, 1832 (Lehmann
1832).
** Anthoceros venosus Lindenb. et Gottsche, Syn. Hepat. 4: 584, 1846 (Gottsche et al. 1846).
Excluded from the genus
* Anthoceros aethyopicus Gola, Ann. Bot. (Rome) 13 (1): 73, 1914 (Gola 1914a). 8
* Anthoceros brunnthaleri Steph., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denk-
schr. 88: 732, 1913 (Stephani 1913b). 9
* Anthoceros oribundus Steph., Sp. Hepat. (Stephani) 5: 977, 1916 (Stephani
1916b). 10
* Anthoceros mildbraedii Steph., Sp. Hepat. (Stephani) 6: 428, 1923 (Stephani 1923). 11
* Anthoceros parvifrons Steph., 52 (5): 307, 1912 (Stephani 1912a). 12
* Anthoceros pseudocostus Steph., Sp. Hepat. (Stephani) 5: 977, 1916 (Stephani 1916b). 13
* Anthoceros rossoi Gola, Mem. Reale Accad. Sci. Torino (ser. 2) 65 (1): 10, 1916
(Gola 1916). 14
8 Anthoceros aethyopicus is probably a Phaeoceros species (Wigginton 2009).
9 Anthoceros brunnthaleri was annotated by Hasegawa in 1993 as probably conspecic with Phymatoceros
bulbiculosus but the synonymy seems not to have been published.
10 Anthoceros oribundus type specimen in G was annotated by Hasegawa in 1993 as conspecic with
Anthoceros parvifrons which is a Phaeoceros species.
11 Anthoceros mildbraedii is probably conspecic with Phaeoceros carolinianus (Wigginton and Grolle 1996).
12 Anthoceros parvifrons is annotated as a Phaeoceros species by Hasegawa in 1993 but the new combination
has never been published.
13 Anthoceros pseudocostus is a Phaeoceros species of uncertain status (Grolle 1995).
14 Anthoceros rossoi is a Phaeoceros species of uncertain status (Wigginton 2009).
World checklist of hornworts and liverworts 31
*** Folioceros D.C.Bharadwaj, Geophytology 1 (1): 9, 1971 (Bharadwaj 1971).
*** Folioceros amboinensis (Schin.) Piippo, Acta Bot. Fenn. 148: 36, 1993 (Piippo
1993b). Bas.: Anthoceros amboinensis Schin., Leberm., Forschungsr. Gazelle 4
(4): 45, 1890 (Schiner 1890).
** Folioceros apiahynus (Steph.) Hässel, Candollea 45 (1): 215, 1990 (Hässel 1990b).
Bas.: Anthoceros apiahynus Steph., Sp. Hepat. (Stephani) 5: 999, 1916 (Stephani
1916b).
** Folioceros argillaceus (Steph.) J.C.Villarreal et Cargill, Phytotaxa 208 (1): 93, 2015
(Villarreal et al. 2015). Bas.: Aspiromitus argillaceus Steph., Sp. Hepat. (Stephani)
5: 970, 1916 (Stephani 1916b).
*** Folioceros assamicus D.C.Bharadwaj, Geophytology 1 (1): 10, 1971 (Bharadwaj 1971).
** Folioceros dilatatus (Steph.) J.C.Villarreal et Cargill, Phytotaxa 208 (1): 93, 2015
(Villarreal et al. 2015). Bas.: Anthoceros dilatatus Steph., Bot. Jahrb. Syst. 8 (2): 95,
1886 (Stephani 1886d).
*** Folioceros dixitianus (Mahab.) D.C.Bharadwaj, Geophytology 5 (2): 227, 1975
(Bharadwaj 1975). Bas.: Aspiromitus dixitianus Mahab., Curr. Sci. 10 (12): 532,
1941 (Mahabale 1941).
*** Folioceros fuciformis (Mont.) D.C.Bharadwaj, Geophytology 5 (2): 227, 1975
(Bharadwaj 1975). Bas.: Anthoceros fuciformis Mont., Ann. Sci. Nat. Bot. (sér. 2)
20: 296, 1844 (Montagne 1844b).
*** Folioceros glandulosus (Lehm. et Lindenb.) D.C.Bharadwaj, Geophytology 5 (2):
227, 1975 (Bharadwaj 1975). Bas.: Anthoceros glandulosus Lehm. et Lindenb.,
Nov. Stirp. Pug. 4: 26, 1832 (Lehmann 1832).
*** Folioceros incurvus (Steph.) D.C.Bharadwaj, Geophytology 5 (2): 227, 1975
(Bharadwaj 1975). Bas.: Anthoceros incurvus Steph., Hedwigia 32 (3): 143, 1893
(Stephani 1893b).
*** Folioceros indicus D.C.Bharadwaj, Geophytology 8 (1): 114, 1978 (Bharadwaj 1978).
*** Folioceros kashyapii S.C.Srivast. et A.K.Asthana, Bryologist 92 (2): 219, 1989
(Srivastava and Asthana 1989).
*** Folioceros mangaloreus (Steph.) D.C.Bharadwaj, Geophytology 5 (2): 227, 1975
(Bharadwaj 1975). Bas.: Aspiromitus mangaloreus Steph., Sp. Hepat. (Stephani) 5:
967, 1916 (Stephani 1916b).
*** Folioceros paliformis D.K.Singh, Bull. Bot. Surv. India 29: 176, 1987 [1989] (Singh
1987b).
*** Folioceros physocladus D.C.Bharadwaj, Geophytology 8 (1): 115, 1978 (Bharadwaj
1978). Based on: Anthoceros physocladus Schin. ex Pandé, Proc. Indian Sci. Congr.
Assoc. 47 (2): 98, 1960 (Pandé 1960), nom. inval.
*** Folioceros pinnilobus (Steph.) D.C.Bharadwaj, Geophytology 5 (2): 227, 1975
(Bharadwaj 1975). Bas.: Anthoceros pinnilobus Steph., Akad. Wiss. Wien, Math.-
Naturwiss. Kl., Denkschr. 81: 299, 1907 (Stephani 1907a).
*** Folioceros satpurensis D.C.Bharadwaj et K.P.Srivast., Geophytology 8 (1): 112,
1978 (Bharadwaj 1978). Based on: Anthoceros satpurensis K.P.Srivast., Proc. In-
dian Sci. Congr. Assoc. 47 (3): 337, 1960 (Srivastava 1960), nom. inval.
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
32
*** Folioceros udarii A.K.Asthana et S.C.Srivast., Cryptog. Bryol. Lichénol. 7 (2): 151,
1986 (Asthana and Srivastava 1986).
*** Folioceros verruculosus (J.Haseg.) R.L.Zhu et M.J.Lai, Ann. Bot. Fenn. 48 (5): 383,
2011 (Wang et al. 2011). Bas.: Anthoceros verruculosus J.Haseg., Acta Phytotax.
Geobot. 44 (2): 103, 1993 (Hasegawa 1993b).
Dendrocerotidae R.J.Duff, J.C.Villarreal, Cargill et Renzaglia
Dendrocerotales Hässel
*** Dendrocerotaceae J.Haseg.
by J.C. Villarreal and D.C. Cargill
*** Dendroceros Nees, Syn. Hepat. 4: 579, 1846 (Gottsche et al. 1846).
** subg. Apoceros R.M.Schust., Phytologia 63 (3): 200, 1987 (Schuster 1987b).
*** Dendroceros cavernosus J.Haseg., J. Hattori Bot. Lab. 47: 306, 1980 (Hasegawa 1980).
*** Dendroceros dicilis Steph., Sp. Hepat. (Stephani) 5: 1009, 1917 (Stephani
1917b).
*** Dendroceros muelleri Steph., Hedwigia 28 (2): 133, 1889 (Stephani 1889a).
*** Dendroceros ogeramnangus Piippo, Acta Bot. Fenn. 148: 40, 1993 (Piippo 1993b).
*** Dendroceros pedunculatus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 14, 1909
(Stephani 1909c).
** Dendroceros subdicilis S.Hatt., Bot. Mag. (Tokyo) 64 (755/756): 119, 1951
(Hattori 1951c).
** subg. Dendroceros
*** Dendroceros acutilobus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 18, 1909
(Stephani 1909c).
*** Dendroceros borbonicus Steph., Bull. Soc. Roy. Bot. Belgique, Compt. Rend. 32
(2): 31, 1893 [1894] (Stephani 1893e).
*** Dendroceros crispus (Sw.) Nees, Syn. Hepat. 4: 581, 1846 (Gottsche et al. 1846).
Bas.: Anthoceros crispus Sw., Prodr. (Swartz): 146, 1788 (Swartz 1788).
*** Dendroceros foliicola J.Haseg., J. Hattori Bot. Lab. 47: 296, 1980 (Hasegawa 1980).
*** Dendroceros japonicus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 15, 1909
(Stephani 1909c).
*** Dendroceros javanicus (Nees) Nees, Syn. Hepat. 4: 582, 1846 (Gottsche et al. 1846).
Bas.: Anthoceros javanicus Nees, Enum. Pl. Crypt. Javae: 1, 1830 (Nees 1830).
*** Dendroceros subplanus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 20, 1909
(Stephani 1909c).
*** Dendroceros tubercularis S.Hatt., Bot. Mag. (Tokyo) 58 (685): 6, 1944 (Hattori 1944b).
World checklist of hornworts and liverworts 33
*** Dendroceros validus Steph., Sp. Hepat. (Stephani) 5: 1016, 1917 (Stephani 1917b).
** Dendroceros vesconianus Gottsche, J. Bot. (Morot) 12: 150, 1898 (Bescherelle 1898).
** Dendroceros wattsianus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 17, 1909
(Stephani 1909c).
Incertae sedis
* Dendroceros adglutinatus (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn.
Hepat. 4: 580, 1846 (Gottsche et al. 1846). Bas.: Monoclea adglutinata Hook.f. et
Taylor, London J. Bot. 4: 96, 1845 (Hooker and Taylor 1845). 15
*** Dendroceros africanus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 18, 1909
(Stephani 1909c).
** Dendroceros allionii Steph., Sp. Hepat. (Stephani) 5: 1014, 1917 (Stephani 1917b).
** Dendroceros australis Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 17, 1909 (Ste-
phani 1909c).
** Dendroceros breutelii Nees, Syn. Hepat. 4: 581, 1846 (Gottsche et al. 1846).
** Dendroceros breutelii var. surinamensis Lindenb. et Gottsche, Linnaea 24 (6): 639,
1851 [1852] (Lindenberg and Gottsche 1851a).
*** Dendroceros cichoraceus (Mont.) Gottsche, Bot. Zeitung (Berlin) Beil. 16: 16, 1858
(Gottsche 1858). Bas.: Anthoceros cichoraceus Mont., Ann. Sci. Nat. Bot. (sér. 3) 4:
355, 1845 (Montagne 1845b).
* Dendroceros crassicostatus Steph., Sp. Hepat. (Stephani) 5: 1015, 1917 (Stephani 1917b).
** Dendroceros crassinervis (Nees) Gottsche, Bot. Zeitung (Berlin) Beil. 16: 18, 1858
(Gottsche 1858). Bas.: Anthoceros crassinervis Nees, Syn. Hepat. 4: 589, 1846
(Gottsche et al. 1846).
*** Dendroceros crispatus (Hook.) Nees, Syn. Hepat. 4: 579, 1846 (Gottsche et al.
1846). Bas.: Monoclea crispata Hook., Bot. Misc. 1: 117, 1830 (Hooker 1830).
** Dendroceros crispatus var. simplicior Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 574, 1885 (Spruce 1885).
** Dendroceros cucullatus Steph., Sp. Hepat. (Stephani) 6: 429, 1923 (Stephani 1923). 16
* Dendroceros exalatus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 14, 1909
(Stephani 1909c). 17
* Dendroceros gracilis Steph., Sp. Hepat. (Stephani) 5: 1015, 1917 (Stephani 1917b).
*** Dendroceros granulatus Mitt., Fl. vit.: 419, 1871 [1873] (Mitten 1871).
** Dendroceros herasii M.Infante, J. Bryol. 32 (4): 285, 2010 (Infante 2010).
** Dendroceros humboldtensis Hürl., Bauhinia 1 (3): 254, 1960 (Hürlimann 1960).
** Dendroceros paivae C.A.Garcia, Sérgio et J.C.Villarreal, Cryptog. Bryol. 33 (1): 5,
2012 (Garcia et al. 2012).
15 Dendroceros adglutinatus is conspecic with Dendroceros crispus in Proskauer (1960), but later authors
accept it.
16 Dendroceros cucullatus was treated as conspecic with Dendroceros dicilis by Hasegawa (1980), but
re-instated by Chantanaorrapint et al. (2014).
17 Dendroceros exalatus was reported from the Moluccas in Stephani (1909c), but Hasegawa (1980)
states that this is a Brazilian species. Dendroceros has not been recently revised for Brazil.
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
34
* Dendroceros rarus Steph., Sp. Hepat. (Stephani) 5: 1014, 1917 (Stephani 1917b).
* Dendroceros reticulus Herzog, Memoranda Soc. Fauna Fl. Fennica 26: 37, 1950
[1951] (Herzog 1950b). 18
** Dendroceros rigidus Steph., Sp. Hepat. (Stephani) 5: 1017, 1917 (Stephani 1917b).
*** Dendroceros seramensis J.Haseg., Acta Phytotax. Geobot. 37 (1/3): 10, 1986
(Hasegawa 1986a).
** Dendroceros subtropicus C.J.Wild, Trans. Nat. Hist. Soc. Queensland 1: 49, 1893
(Wild 1893).
** Dendroceros tahitensis Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 30 (5): 138,
1873 (Ångström 1873).
*** Megaceros Campb., Ann. Bot. (Oxford) 21 (4): 484, 1907 (Campbell 1907).
** Megaceros aneuriformis Steph., Sp. Hepat. (Stephani) 5: 949, 1916 (Stephani 1916b).
*** Megaceros austronesophilus Cargill et Seppelt, Austral. Syst. Bot. 26 (5): 372, 2013
(Cargill et al. 2013a).
** Megaceros ciliatus K.I.Goebel, Ann. Jard. Bot. Buitenzorg 39: 74, 1928 (Goebel 1928).
*** Megaceros denticulatus (Lehm.) Steph., Sp. Hepat. (Stephani) 5: 956, 1916
(Stephani 1916b). Bas.: Anthoceros denticulatus Lehm., Nov. Stirp. Pug. 10: 25,
1857 (Lehmann 1857).
*** Megaceros agellaris (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 951, 1916 (Stephani
1916b). Bas.: Anthoceros agellaris Mitt., Fl. vit.: 419, 1871 [1873] (Mitten 1871).
*** Megaceros gracilis (Reichardt) Steph., Sp. Hepat. (Stephani) 5: 955, 1916 (Stepha-
ni 1916b). Bas.: Anthoceros gracilis Reichardt, Verh. K.K. Zool.-Bot. Ges. Wien
16: 957, 1866 (Reichardt 1866).
** Megaceros leptohymenius (Hook.f. et Taylor) Steph., Sp. Hepat. (Stephani) 5: 955,
1916 (Stephani 1916b). Bas.: Monoclea leptohymenia Hook.f. et Taylor, London J.
Bot. 3: 575, 1844 (Hooker and Taylor 1844d).
*** Megaceros pellucidus (Colenso) E.A.Hodgs., J. Roy. Soc. New Zealand 2 (1): 115,
1972 (Hodgson 1972). Bas.: Anthoceros pellucidus Colenso, Trans. & Proc. New
Zealand Inst. 17: 263, 1885 (Colenso 1885).
** Megaceros tjibodensis Campb., Ann. Bot. (Oxford) 21 (4): 484, 1907 (Campbell 1907).
Excluded from the genus 19
* Megaceros avens (Spruce) Campb., Ann. Bot. (Oxford) 21 (4): 483, 1907 (Camp-
bell 1907). Bas.: Anthoceros avens Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 575, 1885 (Spruce 1885).
* Megaceros jamesonii (Taylor) Steph., Biblioth. Bot. 87 (2): 268, 1916 (Stephani 1916a).
Bas.: Dendroceros jamesonii Taylor, London J. Bot. 7: 285, 1848 (Taylor 1848b).
18 Dendroceros reticulus was mentioned by Hasegawa (1980). He did not synonymise it with Dendro ceros
javanicus, he just stated that Herzog thought it similar to Dendroceros elegans which is conspecic with
Dendroceros javanicus.
19 Megaceros avens and Megaceros jamesonii are probably conspecic with Nothoceros vincentianus
(Villarreal et al. 2010a).
World checklist of hornworts and liverworts 35
*** Nothoceros (R.M.Schust.) J.Haseg., J. Hattori Bot. Lab. 76: 32, 1994 (Hasegawa
1994b). Bas.: Megaceros subg. Nothoceros R.M.Schust., Phytologia 63 (3): 200,
1987 (Schuster 1987b).
*** Nothoceros aenigmaticus J.C.Villarreal et K.D.McFarland, Bryologist 113 (1): 109,
2010 (Villarreal et al. 2010b). Based on: Megaceros aenigmaticus R.M.Schust.,
Hepat. Anthocerotae N. Amer. 6: 830, 1992 (Schuster 1992d), nom. inval.
*** Nothoceros canaliculatus (Pagán) J.C.Villarreal, Hässel et N.Salazar, Bryologist 110
(2): 283, 2007 (Villarreal et al. 2007). Bas.: Dendroceros canaliculatus Pagán, Bry-
ologist 45 (4): 111, 1942 (Pagán 1942a).
*** Nothoceros endiviifolius (Mont.) J.Haseg. ex J.C.Villarreal, Hässel et N.Salazar,
Bryologist 110 (2): 283, 2007 (Villarreal et al. 2007). Bas.: Anthoceros endiviifolius
Mont., Voy. Pole Sud, Bot. 1: 211, 1845 (Montagne 1845c).
** Nothoceros fuegiensis (Steph.) J.C.Villarreal, Bryologist 113 (1): 109, 2010 (Villar-
real et al. 2010b). Bas.: Megaceros fuegiensis Steph., Kungl. Svenska Vetensk.-Akad.
Handl. (n.ser.) 46 (9): 91, 1911 (Stephani 1911b).
*** Nothoceros giganteus (Lehm. et Lindenb.) J.Haseg. ex J.C.Villarreal, Hässel et
N.Salazar, Bryologist 110 (2): 283, 2007 (Villarreal et al. 2007). Bas.: Anthoceros
giganteus Lehm. et Lindenb., Nov. Stirp. Pug. 4: 25, 1832 (Lehmann 1832).
*** Nothoceros minarum (Nees) J.C.Villarreal, Molec. Phylogen. Evol. 78: 34, 2014
(Villarreal and Renner 2014). Bas.: Anthoceros minarum Nees, Naturgesch. Eur.
Leberm. 4: 340, 1838 (Nees 1838a).
** Nothoceros renzagliensis J.C.Villarreal, L.V.Campos et Uribe, Syst. Bot. 37 (1): 32,
2012 (Villarreal et al. 2012).
*** Nothoceros schizophyllus (Steph.) J.C.Villarreal, Molec. Phylogen. Evol. 78: 34,
2014 (Villarreal and Renner 2014). Bas.: Megaceros schizophyllus Steph., Sp.
Hepat. (Stephani) 5: 949, 1916 (Stephani 1916b).
*** Nothoceros superbus J.C.Villarreal, Hässel et N.Salazar, Bryologist 110 (2): 280,
2007 (Villarreal et al. 2007).
*** Nothoceros vincentianus (Lehm. et Lindenb.) J.C.Villarreal, Bryologist 113 (1):
111, 2010 (Villarreal et al. 2010b). Bas.: Anthoceros vincentianus Lehm. et Lin-
denb., Nov. Stirp. Pug. 6: 16, 1834 (Lehmann 1834).
** Phaeomegaceros R.J.Du, J.C.Villarreal, Cargill et Renzaglia, Bryologist 110
(2): 241, 2007 (Du et al. 2007).
*** Phaeomegaceros coriaceus (Steph.) R.J.Du, J.C.Villarreal, Cargill et Renzaglia,
Bryologist 110 (2): 241, 2007 (Du et al. 2007). Bas.: Anthoceros coriaceus Steph.,
Sp. Hepat. (Stephani) 5: 991, 1916 (Stephani 1916b).
*** Phaeomegaceros mbriatus (Gottsche) R.J.Du, J.C.Villarreal, Cargill et Renzag-
lia, Bryologist 110 (2): 241, 2007 (Du et al. 2007). Bas.: Anthoceros mbriatus
Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 187, 1864 (Gottsche 1864).
*** Phaeomegaceros foveatus (J.Haseg.) J.C.Villarreal, Nova Hedwigia 91 (3/4): 352,
2010 (Villarreal et al. 2010a). Bas.: Phaeoceros foveatus J.Haseg., Bryol. Res. 7 (12):
374, 2001 (Hasegawa 2001).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
36
*** Phaeomegaceros hirticalyx (Steph.) R.J.Du, J.C.Villarreal, Cargill et Renzaglia,
Bryologist 110 (2): 241, 2007 (Du et al. 2007). Bas.: Aspiromitus hirticalyx Ste-
ph., Sp. Hepat. (Stephani) 5: 966, 1916 (Stephani 1916b).
*** Phaeomegaceros plicatus (Mitt.) J.C.Villarreal, J.J.Engel et Váňa, Mem. New York
Bot. Gard. 105: 85, 2013 (Váňa and Engel 2013). Bas.: Anthoceros plicatus Mitt.,
Rep. Challenger, Bot. 1 (2): 178, 1884 (Mitten 1884a).
*** Phaeomegaceros skottsbergii (Steph.) R.J.Du, J.C.Villarreal, Cargill et Renzaglia,
Bryologist 110 (2): 241, 2007 (Du et al. 2007). Bas.: Anthoceros skottsbergii Steph.,
Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 90, 1911 (Stephani 1911b).
*** Phaeomegaceros squamuliger (Spruce) J.C.Villarreal, Nova Hedwigia 91 (3/4): 351,
2010 (Villarreal et al. 2010a). Bas.: Anthoceros squamuliger Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 576, 1885 (Spruce 1885).
** Phaeomegaceros squamuliger subsp. hasselii J.C.Villarreal, Cargill et Gonet, Nova
Hedwigia 91 (3/4): 352, 2010 (Villarreal et al. 2010a).
Phymatocerotales R.J.Duff, J.C.Villarreal, Cargill et Renzaglia
*** Phymatocerotaceae R.J.Duff, J.C.Villarreal, Cargill et Renzaglia
by J.C. Villarreal and D.C. Cargill
*** Phymatoceros Stotler, W.T.Doyle et Crand.-Stotl., Phytologia 87 (2): 113,
2005 (Stotler et al. 2005).
*** Phymatoceros bulbiculosus (Brot.) Stotler, W.T.Doyle et Crand.-Stotl., Phytologia
87 (2): 114, 2005 (Stotler et al. 2005). Bas.: Anthoceros bulbiculosus Brot., Fl. lusit.
2: 430, 1804 [1805] (Brotero 1804).
*** Phymatoceros phymatodes (M.Howe) R.J.Du, J.C.Villarreal, Cargill et Renzaglia,
Bryologist 110 (2): 240, 2007 (Du et al. 2007). Bas.: Anthoceros phymatodes
M.Howe, Bull. Torrey Bot. Club 25 (1): 12, 1898 (Howe 1898a).
Notothylatidae R.J.Duff, J.C.Villarreal, Cargill et Renzaglia
Notothyladales Hyvönen et Piippo
*** Notothyladaceae Müll.Frib. ex Prosk.
by J.C. Villarreal and D.C. Cargill
World checklist of hornworts and liverworts 37
** Notothyladoideae Grolle
*** Notothylas Sull. ex A.Gray, Amer. J. Sci. Arts (ser. 2) 1 (1): 74, 1846 (Gray 1846).
** subg. Notothylas
*** Notothylas anaporata Udar et D.K.Singh, Rev. Bryol. Lichénol. 45 (2): 202, 1979
(Udar and Singh 1979b).
*** Notothylas breutelii (Gottsche) Gottsche, Bot. Zeitung (Berlin) Beil. 16: 21, 1858
(Gottsche 1858). Bas.: Anthoceros breutelii Gottsche, Syn. Hepat. 4: 583, 1846
(Gottsche et al. 1846).
*** Notothylas depressispora J.Haseg., Acta Phytotax. Geobot. 30 (1/3): 26, 1979
(Hasegawa 1979).
*** Notothylas dissecta Steph., Sp. Hepat. (Stephani) 5: 1020, 1917 (Stephani 1917b).
*** Notothylas himalayensis Udar et D.K.Singh, J. Bryol. 11 (3): 451, 1981 (Udar and
Singh 1981a).
*** Notothylas indica Kashyap, Proc. Lahore Philos. Soc. 4: 54, 1925 (Kashyap and
Dutt 1925).
*** Notothylas javanica (Sande Lac.) Gottsche, Bot. Zeitung (Berlin) Beil. 16: 20,
1858 (Gottsche 1858). Bas.: Blasia javanica Sande Lac., Syn. hepat. jav.: 94, 1856
[1857] (Sande Lacoste 1856b).
*** Notothylas orbicularis (Schwein.) Sull., Amer. J. Sci. Arts (ser. 2) 1 (1): 75, 1846
(Gray 1846). Bas.: Targionia orbicularis Schwein., Spec. Fl. Amer. Crypt.: 23,
1821 (Schweinitz 1821).
*** Notothylas pandei Udar et V.Chandra, Geophytology 7 (2): 142, 1977 (Udar and
Chandra 1977).
** subg. Notothyloides A.K.Asthana et S.C.Srivast., Bryophyt. Biblioth. 42: 106,
1991 (Asthana and Srivastava 1991).
*** Notothylas khasiana Udar et D.K.Singh, J. Indian Bot. Soc. 60: 112, 1981 (Udar
and Singh 1981b).
*** Notothylas peidereri Udar et D.K.Singh, Lindbergia 5 (1): 28, 1979 (Udar and
Singh 1979a).
Incertae sedis
*** Notothylas decurva (Mitt.) Steph., Cat. Afr. Pl. (Hiern) 2 (2): 320, 1901 (Stephani
1901d). Bas.: Anthoceros decurvus Mitt., Trans. Linn. Soc. London 23 (1): 58,
1860 (Mitten 1860a).
*** Notothylas abellata Steph., Cat. Afr. Pl. (Hiern) 2 (2): 320, 1901 (Stephani 1901d).
*** Notothylas galapagensis M.Howe, Proc. Calif. Acad. Sci. (ser. 4) 21 (17): 203, 1934
(Howe 1934).
** Notothylas irregularis Chantanaorr., Acta Bot. Hung. 56 (3/4): 270, 2014 (Chan-
tanaorrapint 2014).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
38
*** Notothylas kashyapii D.K.Singh, Indian J. Forest. 23 (4): 386, 2000 (Singh and
Semwal 2000).
*** Notothylas nepalensis D.K.Singh, J. Bombay Nat. Hist. Soc. 84 (3): 650, 1987
[1988] (Singh 1987a).
*** Notothylas temperata J.Haseg., Acta Phytotax. Geobot. 30 (1/3): 20, 1979 (Hasega-
wa 1979).
*** Notothylas udarii D.K.Singh et Semwal, Phytotaxonomy 1: 35, 2001 (Singh and
Semwal 2001).
* Notothylas verdoornii Khanna, Rev. Bryol. Lichénol. 6: 118, 1933 (Khanna 1933).
*** Notothylas vitalii Udar et D.K.Singh, Misc. Bryol. Lichenol. 8 (9): 173, 1980
(Udar and Singh 1980).
** Notothylas yunnanensis T.Peng et R.L.Zhu, Phytotaxa 156 (3): 157, 2014 (Peng
and Zhu 2014).
** Phaeocerotoideae Hässel
* Mesoceros Piippo, Acta Bot. Fenn. 148: 30, 1993 (Piippo 1993b). 20
* Mesoceros mesophoros Piippo, Acta Bot. Fenn. 148: 30, 1993 (Piippo 1993b).
* Mesoceros porcatus Piippo, Haussknechtia, Beih. 9: 279, 1999 (Piippo 1999).
** Paraphymatoceros Hässel, Phytologia 88 (2): 208, 2006 (Hässel 2006a).
*** Paraphymatoceros diadematus Hässel, Phytologia 88 (2): 209, 2006 (Hässel 2006a).
*** Paraphymatoceros hallii (Austin) Hässel, Phytologia 88 (2): 209, 2006 (Hässel
2006a). Bas.: Anthoceros hallii Austin, Bull. Torrey Bot. Club 6 (4): 26, 1875
[1876] (Austin 1875a).
*** Paraphymatoceros pearsonii (M.Howe) J.C.Villarreal et Cargill, Phytotaxa 208 (1):
94, 2015 (Villarreal et al. 2015). Bas.: Anthoceros pearsonii M.Howe, Bull. Torrey
Bot. Club 25 (1): 8, 1898 (Howe 1898a).
*** Paraphymatoceros proskaueri (Stotler, Crand.-Stotl. et W.T.Doyle) J.C.Villarreal
et Cargill, Phytotaxa 208 (1): 94, 2015 (Villarreal et al. 2015). Bas.: Phaeoceros
proskaueri Stotler, Crand.-Stotl. et W.T.Doyle, Fieldiana, Bot. (n.ser.) 47: 232,
2008 (Crandall-Stotler et al. 2008a).
*** Phaeoceros Prosk., Bull. Torrey Bot. Club 78 (4): 346, 1951 (Proskauer 1951a).
** Phaeoceros austroandinus Hässel, Candollea 44 (2): 721, 1989 (Hässel 1989a).
** Phaeoceros bolusii (Sim) S.W.Arnell, Hepat. South Africa: 401, 1963 (Arnell
1963b). Bas.: Anthoceros bolusii Sim, Trans. Roy. Soc. South Africa 15 (1): 114,
1926 (Sim 1926).
20 Mesoceros was described from a collection that contains a mixture of both Anthoceros and Phaeoceros
material.
World checklist of hornworts and liverworts 39
** Phaeoceros brevicapsulus (Steph.) Hässel, Candollea 44 (2): 725, 1989 (Hässel
1989a). Bas.: Anthoceros brevicapsulus Steph., Sp. Hepat. (Stephani) 5: 981, 1916
(Stephani 1916b).
*** Phaeoceros carolinianus (Michx.) Prosk., Bull. Torrey Bot. Club 78 (4): 347, 1951
(Proskauer 1951a). Bas.: Anthoceros carolinianus Michx., Fl. bor.-amer. (Michaux)
2: 280, 1803 (Michaux 1803).
*** Phaeoceros delicatus E.O.Campb. et Outred, New Zealand J. Bot. 33 (3): 285,
1995 (Campbell and Outred 1995).
*** Phaeoceros dendroceroides (Steph.) Hässel, Beih. Nova Hedwigia 134: 487, 2009
(Hässel and Rubies 2009). Bas.: Anthoceros dendroceroides Steph., Sp. Hepat. (Ste-
phani) 5: 984, 1916 (Stephani 1916b). 21
*** Phaeoceros engelii Cargill et Fuhrer, Fieldiana, Bot. (n.ser.) 47: 248, 2008 (Cargill
and Fuhrer 2008).
** Phaeoceros erectus Udar et D.K.Singh, Geophytology 11 (2): 257, 1981 (Udar and
Singh 1981c). Nom. nov. pro Anthoceros erectus Steph., Sp. Hepat. (Stephani) 5:
984, 1916 (Stephani 1916b), nom. illeg.
*** Phaeoceros evanidus (Steph.) Cargill et Fuhrer, Fieldiana, Bot. (n.ser.) 47: 245,
2008 (Cargill and Fuhrer 2008). Bas.: Anthoceros evanidus Steph., Sp. Hepat.
(Stephani) 5: 990, 1916 (Stephani 1916b).
*** Phaeoceros exiguus (Steph.) J.Haseg., J. Hattori Bot. Lab. 60: 387, 1986 (Hasega-
wa 1986b). Bas.: Anthoceros exiguus Steph., Sp. Hepat. (Stephani) 5: 988, 1916
(Stephani 1916b).
*** Phaeoceros exivalvis (Gottsche et Nees) Hässel, Candollea 44 (2): 728, 1989 (Häs-
sel 1989a). Bas.: Anthoceros exivalvis Gottsche et Nees, Syn. Hepat. 4: 586, 1846
(Gottsche et al. 1846).
*** Phaeoceros fulvisporus (Steph.) J.Haseg., Trop. Bryol. 8: 52, 1993 (Hasegawa
1993a). Bas.: Anthoceros fulvisporus Steph., 52 (5): 306, 1912 (Stephani 1912a).
*** Phaeoceros gemmifer (Horik.) J.Haseg., J. Hattori Bot. Lab. 57: 252, 1984 (Hasega-
wa 1984). Bas.: Anthoceros gemmifer Horik., Sci. Rep. Tôhoku Imp. Univ., Ser. 4,
Biol. 4 (2): 426, 1929 (Horikawa 1929a).
** Phaeoceros gualaquizanus (Steph.) Gradst., J. Hattori Bot. Lab. 45: 123, 1979
(Gradstein and Hekking 1979). Bas.: Anthoceros gualaquizanus Steph., Sp. Hepat.
(Stephani) 5: 979, 1916 (Stephani 1916b).
*** Phaeoceros himalayensis (Kashyap) Prosk. ex Bapna et G.G.Vyas, J. Hattori Bot.
Lab. 25: 88, 1962 (Bapna and Vyas 1962). Bas.: Anthoceros himalayensis Kashyap,
New Phytol. 14 (1): 8, 1915 (Kashyap 1915).
*** Phaeoceros huebschmannii Hässel, Verö. Geobot. Inst. ETH Stiftung Rübel
Zürich 91: 301, 1986 (Hässel 1986b).
*** Phaeoceros inatus (Steph.) Cargill et Fuhrer, Fieldiana, Bot. (n.ser.) 47: 246, 2008
(Cargill and Fuhrer 2008). Bas.: Anthoceros inatus Steph., Sp. Hepat. (Stephani)
5: 990, 1916 (Stephani 1916b).
21 Phaeoceros dendroceroides may belong to Phaeomegaceros
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
40
*** Phaeoceros kashyapii A.K.Asthana et S.C.Srivast., Bryophyt. Biblioth. 42: 129,
1991 (Asthana and Srivastava 1991).
*** Phaeoceros laevis (L.) Prosk., Bull. Torrey Bot. Club 78 (4): 347, 1951 (Proskauer
1951a). Bas.: Anthoceros laevis L., Sp. Pl. 1: 1139, 1753 (Linnaeus 1753).
** Phaeoceros maranguensis (Steph.) Bapna, Univ. Udaipur Res. Stud. 3: 138, 1966
(Bapna 1966). Bas.: Aspiromitus maranguensis Steph., Sp. Hepat. (Stephani) 5:
960, 1916 (Stephani 1916b).
* Phaeoceros microsporus (Steph.) Hässel, Candollea 44 (2): 721, 1989 (Hässel
1989a). Bas.: Aspiromitus microsporus Steph., Sp. Hepat. (Stephani) 5: 963, 1916
(Stephani 1916b). 22
*** Phaeoceros minutus (Mitt.) S.W.Arnell, Hepat. South Africa: 403, 1963 (Arnell 1963b).
Bas.: Anthoceros minutus Mitt., J. Linn. Soc., Bot. 16 (91): 195, 1877 (Mitten 1877).
*** Phaeoceros mohrii (Austin) Hässel, Candollea 44 (2): 721, 1989 (Hässel 1989a). Bas.:
Anthoceros mohrii Austin, Bull. Torrey Bot. Club 6 (52): 304, 1879 (Austin 1879).
*** Phaeoceros oreganus (Austin) Hässel, Candollea 44 (2): 718, 1989 (Hässel 1989a).
Bas.: Anthoceros oreganus Austin, Bull. Torrey Bot. Club 6 (4): 26, 1875 [1876]
(Austin 1875a).
*** Phaeoceros parvulus (Schin.) J.Haseg., J. Hattori Bot. Lab. 57: 248, 1984 (Hasega-
wa 1984). Bas.: Anthoceros parvulus Schin., Österr. Bot. Z. 49 (11): 391, 1899
(Schiner 1899c).
*** Phaeoceros perpusillus Chantanaorr., Acta Bot. Hung. 51 (1/2): 30, 2009 (Chanta-
naorrapint 2009).
** Phaeoceros pichinchensis (Spruce) Hässel, Candollea 44 (2): 725, 1989 (Hässel
1989a). Bas.: Anthoceros pichinchensis Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 577, 1885 (Spruce 1885).
** Phaeoceros propagulifer (Steph.) Prosk., Bull. Torrey Bot. Club 78 (4): 347, 1951
(Proskauer 1951a). Bas.: Anthoceros propagulifer Steph., Sp. Hepat. (Stephani) 5:
1001, 1916 (Stephani 1916b).
* Phaeoceros striatisporus J.Haseg., J. Hattori Bot. Lab. 75, 268 (Hasegawa 1994a).23
*** Phaeoceros tenuis (Spruce) Hässel, Verö. Geobot. Inst. ETH Stiftung Rübel
Zürich 91: 303, 1986 (Hässel 1986b). Bas.: Anthoceros tenuis Spruce, Bull. Soc.
Bot. France (Congr. Bot.) 36: cxcvi, 1889 [1890] (Spruce 1889).
** Phaeoceros tigrinus (Gola) J.C.Villarreal, Phytotaxa 208 (1): 95, 2015 (Villarreal et al.
2015). Bas.: Anthoceros tigrinus Gola, Ann. Bot. (Rome) 13 (1): 72, 1914 (Gola 1914a).
*** Phaeoceros tuberosus (Taylor) Prosk., J. Indian Bot. Soc. 42A: 185, 1964 (Proskauer
1964). Bas.: Anthoceros tuberosus Taylor, London J. Bot. 5: 412, 1846 (Taylor 1846b).
*** Phaeoceros udarii A.K.Asthana et V.Nath, Proc. Natl. Acad. Sci. India, B 63 (4):
461, 1993 (Asthana and Nath 1993).
** Phaeoceros wrightii (Steph.) Hässel, Candollea 44 (2): 722, 1989 (Hässel 1989a). Bas.:
Anthoceros wrightii Steph., Sp. Hepat. (Stephani) 5: 999, 1916 (Stephani 1916b).
22 Phaeoceros microsporus may be conspecic with Phaeoceros mohrii in Schuster (1992d), but he did not
study any material.
23 Phaeoceros striatisporus was nested within Phaeoceros in Li et al. (2011). Morphologically it was supposed
to dier by its unique spores which, however, turns out to be fungal infections (Villarreal & Renner 2012).
World checklist of hornworts and liverworts 41
LEIOSPOROCEROTOPSIDA Stotler et Crand.-Stotl.
Leiosporocerotales Hässel
*** Leiosporocerotaceae Hässel ex Ochyra
by J.C. Villarreal and D.C. Cargill
*** Leiosporoceros Hässel, J. Bryol. 14 (2): 255, 1986 (Hässel 1986a).
*** Leiosporoceros dussii (Steph.) Hässel, J. Bryol. 14 (2): 255, 1986 (Hässel 1986a).
Bas.: Anthoceros dussii Steph., Hedwigia 32 (3): 142, 1893 (Stephani 1893b).
MARCHANTIOPHYTA
HAPLOMITRIOPSIDA Stotler et Crand.-Stotl.
Haplomitriidae Stotler et Crand.-Stotl.
Calobryales Hamlin
*** Haplomitriaceae Dědeček
by S. Bartholomew-Began
e treatment of Haplomitriaceae follows Bartholomew-Began (1991). Morphoge-
netic and molecular studies support the infrageneric ranks of Haplomitrium (Crandall-
Stotler et al. 2009, Forrest et al. 2006).
*** Haplomitrium Nees, Naturgesch. Eur. Leberm. 1: 109, 1833 (Nees 1833c) nom.
conserv.
*** subg. Calobryum (Nees) R.M.Schust., Nova Hedwigia 13 (1/2): 40, 1967
(Schuster 1967c). Bas.: Calobryum Nees, Syn. Hepat. 4: 507, 1846 (Gottsche et
al. 1846).
*** Haplomitrium blumei (Nees) R.M.Schust., J. Hattori Bot. Lab. 26: 225, 1963
(Schuster 1963b). Bas.: Monoclea blumei Nees, Enum. Pl. Crypt. Javae: 2, 1830
(Nees 1830).
*** Haplomitrium mnioides (Lindb.) R.M.Schust., J. Hattori Bot. Lab. 26: 225, 1963
(Schuster 1963b). Bas.: Rhopalanthus mnioides Lindb., Not. Sällsk. Fauna Fl.
Fenn. Förh. 13: 391, 1874 (Lindberg 1874a).
*** subg. Haplomitrium
** sect. Archibryum (R.M.Schust.) J.J.Engel, Ann. Missouri Bot. Gard. 68 (4):
675, 1981 (Engel 1981). Bas.: Haplomitrium subg. Archibryum R.M.Schust.,
Nova Hedwigia 13 (1/2): 28, 1967 (Schuster 1967c).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
42
*** Haplomitrium gibbsiae (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 225, 1963
(Schuster 1963b). Bas.: Calobryum gibbsiae Steph., Sp. Hepat. (Stephani) 6: 76,
1917 (Stephani 1917a).
*** Haplomitrium intermedium Berrie, Proc. Linn. Soc. New South Wales (ser. 2) 87
(399): 191, 1963 (Berrie 1963).
** sect. Haplomitrium
*** Haplomitrium hookeri (Lyell ex Sm.) Nees, Naturgesch. Eur. Leberm. 1: 111, 1833
(Nees 1833c). Bas.: Jungermannia hookeri Lyell ex Sm., Engl. Bot. 35: tab. 2555,
1814 (Smith and Sowerby 1814).
*** Haplomitrium hookeri var. minutum (E.O.Campb.) Barthol.-Began, Bryophyt.
Biblioth. 41: 230, 1991 (Bartholomew-Began 1991). Bas.: Steereomitrium
minutum E.O.Campb., Mem. New York Bot. Gard. 45: 569, 1987 (Campbell
1987).
*** Haplomitrium monoicum J.J.Engel, Ann. Missouri Bot. Gard. 68 (4): 668, 1981
[1982] (Engel 1981).
*** Haplomitrium ovalifolium R.M.Schust., Bryologist 74 (2): 136, 1971 (Schuster 1971d).
Treubiidae Stotler et Crand.-Stotl.
Treubiales Schljakov
*** Treubiaceae Verd.
by R. Stotler and B.J. Crandall-Stotler
Stech et al. (2002) showed a clear molecular distinction between Treubia and Apotreu-
bia, which is in accordance with their morphological dierences and supports their
recognition as separate genera.
*** Apotreubia S.Hatt. et Mizut., Bryologist 69 (4): 491, 1966 [1967] (Hattori et al.
1966).
*** Apotreubia hortoniae Konstant., Phytotaxa 76 (3): 33, 2013 (Konstantinova et al.
2013b). Based on: Apotreubia hortoniae R.M.Schust. et Konstant., J. Hattori Bot.
Lab. 78: 55, 1995 (Schuster and Konstantinova 1995), nom. inval.
*** Apotreubia nana (S.Hatt. et Inoue) S.Hatt. et Mizut., Bryologist 69 (4): 492, 1966
[1967] (Hattori et al. 1966). Bas.: Treubia nana S.Hatt. et Inoue, J. Hattori Bot.
Lab. 11: 99, 1954 (Hattori and Inoue 1954).
** Apotreubia pusilla (R.M.Schust.) Grolle, Acta Bot. Fenn. 125: 63, 1984 (Grolle
and Piippo 1984). Bas.: Treubia pusilla R.M.Schust., Nova Hedwigia 15: 515,
1968 (Schuster 1968b).
World checklist of hornworts and liverworts 43
*** Apotreubia yunnanensis Higuchi, Cryptog. Bryol. Lichénol. 19 (4): 321, 1998 (Hi-
guchi 1998).
*** Treubia K.I.Goebel, Ann. Jard. Bot. Buitenzorg 9 (1): 1, 1890 [1891] (Goebel
1890) nom. conserv.
*** Treubia insignis K.I.Goebel, Ann. Jard. Bot. Buitenzorg 9 (1): 1, 1890 [1891]
(Goebel 1890).
** Treubia insignis subsp. bracteata (Steph.) R.M.Schust. et G.A.M.Scott, J. Hattori
Bot. Lab. 32: 241, 1969 (Schuster and Scott 1969). Bas.: Treubia bracteata Steph.,
Bot. Jahrb. Syst. 23 (1/2, 3): 302, 1896 (Stephani 1896a).
** Treubia insignis subsp. caledonica R.M.Schust. et G.A.M.Scott, J. Hattori Bot.
Lab. 32: 243, 1969 (Schuster and Scott 1969).
** Treubia insignis subsp. vitiensis R.M.Schust. et G.A.M.Scott, J. Hattori Bot. Lab.
32: 242, 1969 (Schuster and Scott 1969).
*** Treubia lacunosa (Colenso) Prosk., Bryologist 58 (3): 199, 1955 (Proskauer 1955).
Bas.: Noteroclada lacunosa Colenso, Trans. & Proc. New Zealand Inst. 18: 248,
1886 (Colenso 1886b).
** Treubia lacunosoides T.Pfei., W.Frey et M.Stech, Nova Hedwigia 75 (1/2): 249,
2002 (Pfeier et al. 2002).
*** Treubia pygmaea R.M.Schust., Phytologia 56 (7): 460, 1985 (Schuster 1985c).
*** Treubia scapanioides R.M.Schust., J. Hattori Bot. Lab. 32: 246, 1969 (Schuster
and Scott 1969).
*** Treubia tahitensis (Nadeaud) Besch., J. Bot. (Morot) 12: 147, 1898 (Bescherelle
1898). Bas.: Gottschea tahitensis Nadeaud, Énum. Pl. Tahiti: 7, 1873 (Nadeaud
1873).
*** Treubia tasmanica R.M.Schust. et G.A.M.Scott, J. Hattori Bot. Lab. 32: 248,
1969 (Schuster and Scott 1969).
JUNGERMANNIOPSIDA Stotler et Crand.-Stotl.
Jungermanniidae Engl.
Jungermanniales H.Klinggr.
Cephaloziineae Schljakov
*** Adelanthaceae Grolle
by K. Feldberg, J. Váňa and J. Heinrichs
e subfamily Jamesonielloideae was excluded from Lophoziaceae/Scapaniaceae by De
Roo et al. (2007) and this status was conrmed by e.g. Vilnet et al. (2010). e family
was described and dened by Feldberg et al. (2010a). Some taxonomic and nomen-
clatural notes can also be found in Feldberg et al. (2010b, 2011), Váňa et al. (2014d).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
44
*** Adelanthoideae K.Feldberg, Heinrichs et Váňa
*** Adelanthus Mitt., J. Proc. Linn. Soc., Bot. 7 (28): 243, 1864 (Mitten 1864b)
nom. conserv.
** sect. Adelanthus
*** Adelanthus falcatus (Hook.) Mitt., J. Proc. Linn. Soc., Bot. 7 (28): 243, 1864
(Mitten 1864b). Bas.: Jungermannia falcata Hook., Musci Exot. 1: tab. 89, 1818
(Hooker 1818).
*** Adelanthus occlusus (Hook.f. et Taylor) Carrington, Trans. Bot. Soc. Edinburgh
10: 381, 1870 (Carrington 1870). Bas.: Jungermannia occlusa Hook.f. et Taylor,
London J. Bot. 3: 369, 1844 (Hooker and Taylor 1844a).
** sect. Calyptrocolea (R.M.Schust.) Grolle, J. Hattori Bot. Lab. 35: 331, 1972
(Grolle 1972c). Bas.: Calyptrocolea R.M.Schust., Rev. Bryol. Lichénol. 34 (3/4):
685, 1966 (Schuster 1966a).
*** Adelanthus aureomarginatus R.M.Schust., Phytologia 39 (4): 250, 1978 (Schuster
1978a).
*** Adelanthus gemmiparus (R.M.Schust.) E.A.Hodgs., Trans. Roy. Soc. New Zea-
land, Biol. Sci. 11 (18): 241, 1970 (Hodgson 1970). Bas.: Calyptrocolea gem-
mipara R.M.Schust., Rev. Bryol. Lichénol. 34 (3/4): 695, 1966 [1967] (Schuster
1966a).
** Adelanthus lingulatus J.J.Engel et Váňa, Mem. New York Bot. Gard. 105: 26,
2013 (Váňa and Engel 2013).
*** Adelanthus tenuis J.J.Engel et Grolle, J. Hattori Bot. Lab. 35: 333, 1972 (Grolle
1972c).
** sect. Lindenbergiani Grolle, J. Hattori Bot. Lab. 35: 331, 1972 (Grolle
1972c).
*** Adelanthus carabayensis (Mont.) Grolle, J. Hattori Bot. Lab. 35: 348, 1972 (Grolle
1972c). Bas.: Plagiochila carabayensis Mont., Ann. Sci. Nat. Bot. (sér. 4) 5: 348,
1856 (Montagne 1856c).
*** Adelanthus integerrimus Grolle, J. Hattori Bot. Lab. 35: 340, 1972 (Grolle
1972c).
*** Adelanthus lindenbergianus (Lehm.) Mitt., J. Proc. Linn. Soc., Bot. 7 (28): 244,
1864 (Mitten 1864b). Bas.: Jungermannia lindenbergiana Lehm., Linnaea 4: 367,
1829 (Lehmann 1829).
** sect. Pittieri Grolle, J. Hattori Bot. Lab. 35: 331, 1972 (Grolle 1972c).
*** Adelanthus pittieri (Steph.) Grolle, J. Hattori Bot. Lab. 35: 337, 1972 (Grolle
1972c). Bas.: Tylimanthus pittieri Steph., Sp. Hepat. (Stephani) 6: 250, 1922
(Stephani 1922).
*** Adelanthus squarrosus Grolle, J. Hattori Bot. Lab. 67: 243, 1989 (Grolle 1989d).
World checklist of hornworts and liverworts 45
*** Pseudomarsupidium Herzog, Svensk Bot. Tidskr. 47 (1): 42, 1953 (Herzog
1953b).
*** Pseudomarsupidium aureocinctum (R.M.Schust.) J.J.Engel, Novon 17 (3): 312,
2007 (Engel 2007). Bas.: Adelanthus decipiens subsp. aureocinctus R.M.Schust.,
Phytologia 39 (4): 250, 1978 (Schuster 1978a).
*** Pseudomarsupidium borneensis (Grolle) Váňa, L.Söderstr., A.Hagborg et von Kon-
rat, Phytotaxa 65: 60, 2012 (Váňa et al. 2012f). Bas.: Adelanthus borneensis Grolle,
J. Hattori Bot. Lab. 35: 362, 1972 (Grolle 1972c).
*** Pseudomarsupidium decipiens (Hook.) Grolle, Trans. Brit. Bryol. Soc. 4 (3): 443,
1963 (Grolle 1963a). Bas.: Jungermannia decipiens Hook., Brit. Jungermann.: tab.
50, 1813 (Hooker 1813).
*** Pseudomarsupidium piliferum (Steph.) Herzog ex Grolle, Trans. Brit. Bryol. Soc.
4 (3): 443, 1963 (Grolle 1963a). Bas.: Marsupidium piliferum Steph., Bull. Herb.
Boissier (sér. 2) 8 (8): 602 (386), 1908 (Stephani 1908e).
*** Wettsteinia Schin., Ann. Jard. Bot. Buitenzorg, suppl. 2: 44, 1898 (Schiner
1898c).
*** Wettsteinia densiretis (Herzog) Grolle, J. Hattori Bot. Lab. 28: 99, 1965 (Grolle
1965f). Bas.: Tylimanthus densiretis Herzog, Nat. Hist. Juan Fernandez (Botany) 2
(5): 712, 1942 (Herzog 1942a).
*** Wettsteinia inversa (Sande Lac.) Schin., Ann. Jard. Bot. Buitenzorg, suppl. 2:
45, 1898 (Schiner 1898c). Bas.: Plagiochila inversa Sande Lac., Ann. Mus. Bot.
Lugduno-Batavi 1: 289, 1864 (Sande Lacoste 1864).
*** Wettsteinia rotundifolia (Horik.) Grolle, J. Hattori Bot. Lab. 28: 100, 1965 (Grolle
1965f). Bas.: Adelanthus rotundifolius Horik., J. Sci. Hiroshima Univ., Ser. B, Div.
2, Bot. 2: 181, 1934 (Horikawa 1934).
*** Wettsteinia schusteriana Grolle, J. Hattori Bot. Lab. 28: 99, 1965 (Grolle 1965f).
** Jamesonielloideae Inoue
*** Cuspidatula Steph., Bull. Herb. Boissier (sér. 2) 1 (10): 1141 (124), 1901
(Stephani 1901c).
*** Cuspidatula contracta (Reinw., Blume et Nees) Steph., Bull. Herb. Boissier (sér. 2)
1 (11): 1141 (124), 1901 (Stephani 1901c). Bas.: Jungermannia contracta Reinw.,
Blume et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1):
233, 1824 [1825] (Reinwardt et al. 1824a).
*** Cuspidatula accida (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 142, 2010 (Feldberg et al. 2010b). Bas.: Anastrophyllum accidum
Steph., Sp. Hepat. (Stephani) 6: 105, 1917 (Stephani 1917a).
*** Cuspidatula exicaulis (Nees) Váňa et L.Söderstr., Phytotaxa 76 (3): 35, 2013
(Váňa et al. 2013h). Bas.: Jungermannia exicaulis Nees, Linnaea 6 (4): 604, 1831
(Nees 1831).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
46
*** Cuspidatula kirkii (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 142, 2010 (Feldberg et al. 2010b). Bas.: Jamesoniella kirkii Steph.,
Hedwigia 34 (2): 47, 1895 (Stephani 1895c).
*** Cuspidatula monodon (Taylor) Steph., Bull. Herb. Boissier (sér. 2) 1 (11): 1143
(126), 1901 (Stephani 1901c). Bas.: Jungermannia monodon Taylor, Nov. Stirp.
Pug. 8: 7, 1844 (Lehmann 1844).
*** Cuspidatula orbicularis (Grolle) Váňa et L.Söderstr., Phytotaxa 76 (3): 36, 2013
(Váňa et al. 2013h). Bas.: Jamesoniella orbicularis Grolle, Feddes Repert. 82 (1):
42, 1971 (Grolle 1971b).
*** Cuspidatula robusta (Austin) Váňa et L.Söderstr., Phytotaxa 76 (3): 36, 2013 (Váňa
et al. 2013h). Bas.: Jungermannia robusta Austin, Proc. Acad. Nat. Sci. Philadel-
phia 21: 219, 1869 (Austin 1869).
** Denotarisia Grolle, Feddes Repert. 82 (1): 6, 1971 (Grolle 1971b).
*** Denotarisia linguifolia (De Not.) Grolle, Feddes Repert. 82 (1): 6, 1971 (Grolle 1971b).
Bas.: Plagiochila linguifolia De Not., Epat. Borneo: 13, 1874 (De Notaris 1874).
** Nothostrepta R.M.Schust., Phytologia 45 (5): 420, 1980 (Schuster 1980b).
*** Nothostrepta bida (Steph.) R.M.Schust., Phytologia 45 (5): 420, 1980 (Schuster
1980b). Bas.: Plagiochila bida Steph., Annuario Reale Ist. Bot. Roma 2: 86, 1885
[1886] (Stephani 1885d).
*** Nothostrepta longissima (Steph.) R.M.Schust., Phytologia 45 (5): 420, 1980
(Schuster 1980b). Bas.: Anastrophyllum longissimum Steph., Bih. Kongl. Svenska
Vetensk.-Akad. Handl. 26 (III, 17): 13, 1901 (Stephani 1901b).
** Pisanoa Hässel, Lindbergia 14 (3): 179, 1988 [1989] (Hässel 1988).
*** Pisanoa chilensis Hässel, Lindbergia 14 (3): 179, 1988 [1989] (Hässel 1988).
** Protosyzygiella (Inoue) R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 334,
1980 (Schuster 1980c). Bas.: Syzygiella subg. Protosyzygiella Inoue, J. Hattori Bot.
Lab. 29: 180, 1966 (Inoue 1966c).
*** Protosyzygiella pseudoconnexa (Herzog) R.M.Schust., Hepat. Anthocerotae N.
Amer. 4: 334, 1980 (Schuster 1980c). Bas.: Plagiochila pseudoconnexa Herzog,
Rev. Bryol. Lichénol. 21 (3/4): 259, 1952 [1953] (Herzog 1952d).
*** Syzygiella Spruce, J. Bot. 14: 234, 1876 (Spruce 1876a). 24
*** subg. Anomalae (Inoue) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Syzygiella sect. Anomalae
Inoue, J. Hattori Bot. Lab. 29: 183, 1966 (Inoue 1966c).
24 Syzygiella includes Jamesoniella (cf. Feldberg et al. 2010a), but one taxon has neither been transferred
nor synonymized. It is listed in the “Names in genera not currently accepted” section below.
World checklist of hornworts and liverworts 47
*** Syzygiella anomala (Lindenb. et Gottsche) Steph., Bull. Herb. Boissier (sér. 2)
2 (5): 471 (190), 1902 (Stephani 1902f). Bas.: Plagiochila anomala Lindenb. et
Gottsche, Syn. Hepat. 5: 646, 1847 (Gottsche et al. 1847).
*** Syzygiella bilobata Inoue, J. Hattori Bot. Lab. 29: 186, 1966 (Inoue 1966c).
** Syzygiella ciliata Gradst. et A.R.Benitez, Nova Hedwigia 99 (1/2): 115, 2014
(Gradstein and Benitez 2014).
*** Syzygiella concreta (Gottsche) Spruce, J. Bot. 14: 234, 1876 (Spruce 1876a). Bas.:
Jungermannia concreta Gottsche, Mexik. Leverm.: 82, 1863 (Gottsche 1863).
*** Syzygiella manca (Mont.) Steph., Hedwigia 31 (1): 14, 1892 (Jack and Stephani
1892). Bas.: Chiloscyphus mancus Mont., Syll. Gen. Sp. Crypt.: 63, 1856 (Mon-
tagne 1856b).
*** Syzygiella pectiniformis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 501, 1885
(Spruce 1885).
*** Syzygiella tonduzana Steph., Sp. Hepat. (Stephani) 6: 118, 1917 (Stephani 1917a).
*** Syzygiella trigonifolia (Steph.) Herzog, Hedwigia 74 (2): 87, 1934 (Herzog 1934a).
Bas.: Jamesoniella trigonifolia Steph., Biblioth. Bot. 87 (2): 185, 1916 (Stephani 1916a).
*** subg. Cryptochila (R.M.Schust.) K.Feldberg, Váňa, Hentschel et Heinrichs,
Cryptog. Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Cryptochila
R.M.Schust., J. Hattori Bot. Lab. 26: 284, 1963 (Schuster 1963b).
*** Syzygiella acinacifolia (Hook.f. et Taylor) K.Feldberg, Váňa, Hentschel et Hein-
richs, Cryptog. Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Junger-
mannia acinacifolia Hook.f. et Taylor, London J. Bot. 3: 367, 1844 (Hooker and
Taylor 1844a).
*** Syzygiella nigrescens (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryp-
tog. Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Jamesoniella nigrescens
Steph., Hedwigia 34 (2): 48, 1895 (Stephani 1895c).
*** Syzygiella paludosa (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Jamesoniella paludosa Steph.,
Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 (III, 17): 11, 1901 (Stephani 1901b).
*** Syzygiella pseudocclusa (E.A.Hodgs.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryp-
tog. Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Jamesoniella pseudocclusa
E.A.Hodgs., Trans. Roy. Soc. New Zealand 85 (4): 583, 1958 (Hodgson 1958).
*** Syzygiella sonderi (Gottsche) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryp-
tog. Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia sonderi
Gottsche, Linnaea 28 (5): 550, 1856 [1857] (Gottsche 1856).
*** Syzygiella spegazziniana (Spruce ex C.Massal.) K.Feldberg, Váňa, Hentschel et
Heinrichs, Cryptog. Bryol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Jung-
ermannia spegazziniana Spruce ex C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 216,
1885 (Massalongo 1885).
*** subg. Pseudoplagiochila Inoue, J. Hattori Bot. Lab. 29: 182, 1966 (Inoue
1966c).
*** Syzygiella ovalifolia Inoue, J. Hattori Bot. Lab. 29: 191, 1966 (Inoue 1966c).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
48
* Syzygiella securifolia (Nees) Inoue, J. Hattori Bot. Lab. 46: 232, 1979 (Inoue
1979a). Bas.: Plagiochila securifolia Nees, Sp. Hepat. (Lindenberg) 2-4: 58, 1840
(Lindenberg 1840). 25
*** Syzygiella subintegerrima (Reinw., Blume et Nees) Spruce, J. Linn. Soc., Bot.
30 (210): 362, 1895 (Gepp 1895b). Bas.: Jungermannia subintegerrima Reinw.,
Blume et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1):
238, 1824 [1825] (Reinwardt et al. 1824a).
*** Syzygiella tasmanica (Hook.f. et Taylor) K.Feldberg, Váňa, Hentschel et Hein-
richs, Cryptog. Bryol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Junger-
mannia tasmanica Hook.f. et Taylor, London J. Bot. 5: 274, 1846 (Taylor 1846a).
*** subg. Roivainenia (Perss.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryp-
tog. Bryol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Roivainenia Perss.,
Nova Hedwigia 3 (1): 43, 1961 (Persson and Grolle 1961).
*** Syzygiella jacquinotii (Mont.) Hentschel, K.Feldberg, Váňa et Heinrichs, Cryptog.
Bryol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia jacquinotii
Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 250, 1843 (Montagne 1843).
*** subg. Syzygiella
*** Syzygiella autumnalis (DC.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog. Bry-
ol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia autumnalis DC., Fl.
Franç. (DC. & Lamarck), 5 (6): 202, 1815 (De Candolle and Lamarck 1815).
*** Syzygiella campanulata Herzog, Rev. Bryol. Lichénol. 11 (1): 9, 1938 [1939] (Herzog
1938a).
*** Syzygiella colorata (Lehm.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia colorata
Lehm., Linnaea 4: 366, 1829 (Lehmann 1829).
** Syzygiella colorata var. collenchymata J.J.Engel et Váňa, Mem. New York Bot.
Gard. 105: 102, 2013 (Váňa and Engel 2013).
*** Syzygiella contigua Steph., Bull. Herb. Boissier (sér. 2) 2 (5): 470 (189), 1902
(Stephani 1902f). Nom. nov. pro Jungermannia contigua Gottsche, Ann. Sci. Nat.
Bot. (sér. 5) 1: 118, 1864 (Gottsche 1864), nom. illeg.
*** Syzygiella elongella (Taylor) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia elongella
Taylor, London J. Bot. 5: 274, 1846 (Taylor 1846a).
*** Syzygiella macrocalyx (Mont.) Spruce, J. Bot. 14: 234, 1876 (Spruce 1876a). Bas.:
Jungermannia macrocalyx Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 248, 1843 (Mon-
tagne 1843).
*** Syzygiella nipponica (S.Hatt.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryp-
tog. Bryol. 31 (2): 145, 2010 (Feldberg et al. 2010b). Bas.: Jamesoniella nipponica
S.Hatt., J. Jap. Bot. 19 (11): 350, 1943 (Hattori 1943b).
25 Syzygiella securifolia is a sister taxon to Syzygiella subintegerrima (Feldberg et al. 2010b) and it is debat-
able if they are dierent enough to warrant separation on the species level. Morphologically they are
very similar (Söderström et al. 2010a).
World checklist of hornworts and liverworts 49
** Syzygiella oenops (Lindenb. et Gottsche) K.Feldberg, Váňa, Hentschel et Heinrichs,
Cryptog. Bryol. 31 (2): 145, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia
oenops Lindenb. et Gottsche, Syn. Hepat. 5: 673, 1847 (Gottsche et al. 1847).
*** Syzygiella perfoliata (Sw.) Spruce, J. Bot. 14: 234, 1876 (Spruce 1876a). Bas.: Jung-
ermannia perfoliata Sw., Prodr. (Swartz): 143, 1788 (Swartz 1788).
*** Syzygiella purpurascens (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryp-
tog. Bryol. 31 (2): 145, 2010 (Feldberg et al. 2010b). Bas.: Jamesoniella purpu-
rascens Steph., Bull. Soc. Roy. Bot. Belgique, Compt. Rend. 30 (2): 200, 1891
[1892] (Stephani 1891b).
*** Syzygiella rubricaulis (Nees) Steph., Bull. Herb. Boissier (sér. 2) 2 (5): 468 (187),
1902 (Stephani 1902f). Bas.: Jungermannia rubricaulis Nees, Fl. Bras. (Martius) 1
(1): 344, 1833 (Nees 1833a).
*** Syzygiella setulosa Steph., Bull. Herb. Boissier (sér. 2) 2 (5): 469 (188), 1902
(Stephani 1902f).
*** Syzygiella teres (Carrington et Pearson) Váňa, Phytotaxa 76 (3): 35, 2013 (Váňa et
al. 2013h). Bas.: Jungermannia teres Carrington et Pearson, Pap. & Proc. Roy. Soc.
Tasmania 1887: 9, 1888 (Carrington and Pearson 1888b).
*** Syzygiella undata (Mont.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 145, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia undata
Mont., Ann. Sci. Nat. Bot. (sér. 4) 14: 183, 1860 (Montagne 1860).
Incertae sedis
*** Syzygiella eatonii (Austin) Inoue, J. Jap. Bot. 37 (12): 359, 1962 (Inoue 1962a).
Bas.: Plagiochila eatonii Austin, Trans. Connecticut Acad. Arts 8 (15): 257, 1891
(Evans 1891).
*** Syzygiella uleana Steph., Hedwigia 44 (4): 224, 1905 (Stephani 1905a).
** Vanaea (Inoue et Gradst.) Inoue et Gradst., Trop. Bryol. 1: 33, 1989 (Gradstein
and Florschütz-de Waard 1989). Bas.: Anastrophyllum subg. Vanaea Inoue et Gradst.,
Bull. Natl. Sci. Mus. Tokyo, B 14 (3): 88, 1988 (Inoue and Gradstein 1988).
*** Vanaea plagiochiloides (Inoue et Gradst.) Inoue et Gradst., Trop. Bryol. 1: 33,
1989 (Gradstein and Florschütz-de Waard 1989). Bas.: Anastrophyllum plagi-
ochiloides Inoue et Gradst., Bull. Natl. Sci. Mus. Tokyo, B 14 (3): 88, 1988 (Inoue
and Gradstein 1988).
*** Anastrophyllaceae L.Söderstr., De Roo et Hedd.
by J. Váňa and L. Söderström
Anastrophyllaceae was described by Söderström et. al (2010b) from elements usually
included in Lophoziaceae. Further taxonomic and nomenclatural notes can be found
in Váňa et al. (2013k, 2013a). e complex of Chandonanthus/Plicanthus/Tetralophozia
should be checked using molecular methods before generic and specic status of most
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
50
of the species can be conrmed. e placement of Hattoria and Zantenia in the family is
still provisional. e placement of Isopaches is also unclear, de Roo et al. (2007) placed it
in the family, but the study of Vilnet et al. (2010) showed that it can not be placed there.
*** Anastrepta (Lindb.) Schin., Hepat. (Engl.-Prantl): 85, 1893 (Schiner 1893b).
Bas.: Jungermannia sect. Anastrepta Lindb., Kongl. Svenska Vetensk.-Akad. Handl.
(n.ser.) 23 (5): 40, 1889 (Lindberg and Arnell 1889).
*** Anastrepta orcadensis (Hook.) Schin., Hepat. (Engl.-Prantl): 85, 1893 (Schiner
1893b). Bas.: Jungermannia orcadensis Hook., Brit. Jungermann.: tab. 71, 1815
(Hooker 1815).
*** Anastrophyllum (Spruce) Steph., Hedwigia 32 (3): 139, 1893 (Stephani 1893b).
Bas.: Jungermannia subg. Anastrophyllum Spruce, J. Bot. 14: 235, 1876 (Spruce
1876a).
*** Anastrophyllum alpinum Steph., Sp. Hepat. (Stephani) 6: 103, 1917 (Stephani 1917a).
*** Anastrophyllum assimile (Mitt.) Steph., Hedwigia 32 (3): 140, 1893 (Stephani
1893b). Bas.: Jungermannia assimilis Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 93,
1860 [1861] (Mitten 1860c).
*** Anastrophyllum auritum (Lehm.) Steph., Bull. Herb. Boissier (sér. 2) 1 (11): 1137
(120), 1901 (Stephani 1901c). Bas.: Jungermannia aurita Lehm., Linnaea 4: 368,
1829 (Lehmann 1829).
*** Anastrophyllum ciliatum Steph., Hedwigia 32 (3): 139, 1893 (Stephani 1893b).
*** Anastrophyllum donnianum (Hook.) Steph., Hedwigia 32 (3): 140, 1893 (Stephani
1893b). Bas.: Jungermannia donniana Hook., Brit. Jungermann.: tab. 39, 1813
(Hooker 1813).
*** Anastrophyllum ellipticum Inoue, Bull. Natl. Sci. Mus. Tokyo, B 4 (1): 13, 1978
(Inoue 1978b). 26
** Anastrophyllum esenbeckii (Mont.) Steph., Hedwigia 32 (3): 140, 1893 (Stephani
1893b). Bas.: Jungermannia esenbeckii Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 247,
1843 (Montagne 1843).
*** Anastrophyllum ssum Steph., Bull. Herb. Boissier 5 (10): 845, 1897 (Stephani 1897c).
*** Anastrophyllum joergensenii Schin., Hedwigia 49 (4): 396, 1910 (Schiner 1910b).
* Anastrophyllum lignicola D.B.Schill et D.G.Long, Ann. Bot. Fenn. 39 (2): 130,
2002 (Schill and Long 2002). 27
*** Anastrophyllum michauxii (F.Weber) H.Buch, Memoranda Soc. Fauna Fl. Fennica
8: 289, 1932 [1933] (Buch 1932). Bas.: Jungermannia michauxii F.Weber, Hist.
Musc. Hepat. Prodr.: 76, 1815 (Weber 1815).
26 Anastrophyllum ellipticum is only tentatively placed in the genus following Mamontov & Vilnet
(2013).
27 Anastrophyllum lignicola is possibly conspecic with Anastrophyllum ellipticum and its generic place-
ment is uncertain.
World checklist of hornworts and liverworts 51
*** Anastrophyllum nigrescens (Mitt.) Steph., Hedwigia 32 (3): 140, 1893 (Stephani
1893b). Bas.: Jungermannia nigrescens Mitt., Hooker’s J. Bot. Kew Gard. Misc. 3:
358, 1851 (Mitten 1851).
*** Anastrophyllum obtusum Herzog, Trans. Brit. Bryol. Soc. 1 (4): 285, 1950 (Herzog
1950a).
*** Anastrophyllum piligerum (Nees) Steph., Hedwigia 32 (3): 140, 1893 (Stephani
1893b). Bas.: Jungermannia piligera Nees, Nova Acta Phys.-Med. Acad. Caes.
Leop.-Carol. Nat. Cur. 12 (1): 414, 1824 [1825] (Reinwardt et al. 1824b).
*** Anastrophyllum squarrosum Herzog, Ann. Bryol. 5: 72, 1932 (Herzog 1932b).
*** Anastrophyllum stellatum R.M.Schust., Phytologia 39 (4): 243, 1978 (Schuster
1978a).
*** Anastrophyllum tubulosum (Nees) Grolle, J. Hattori Bot. Lab. 28: 101, 1965
(Grolle 1965e). Bas.: Jungermannia tubulosa Nees, Enum. Pl. Crypt. Javae: 32,
1830 (Nees 1830).
*** Barbilophozia Loeske, Verh. Bot. Vereins Prov. Brandenburg 49 (1): 37, 1908
(Loeske 1908).
** subg. Barbilophozia
*** Barbilophozia barbata (Schmidel ex Schreb.) Loeske, Verh. Bot. Vereins Prov.
Brandenburg 49 (1): 37, 1908 (Loeske 1908). Bas.: Jungermannia barbata Schmi-
del ex Schreb., Spic. Fl. Lips.: 107, 1771 (Schreber 1771).
*** Barbilophozia hatcheri (A.Evans) Loeske, Verh. Bot. Vereins Prov. Brandenburg
49 (1): 37, 1908 (Loeske 1908). Bas.: Jungermannia hatcheri A.Evans, Bull. Torrey
Bot. Club 25 (8): 417, 1898 (Evans 1898).
*** Barbilophozia lycopodioides (Wallr.) Loeske, Verh. Bot. Vereins Prov. Brandenburg
49 (1): 37, 1908 (Loeske 1908). Bas.: Jungermannia lycopodioides Wallr., Comp. .
Germ. 2 (III): 76, 1831 (Blu and Fingerhuth 1831).
*** Barbilophozia rubescens (R.M.Schust. et Damsh.) Kartt. et L.Söderstr., Ann. Bot. Fenn.
29 (2): 120, 1992 (Söderström et al. 1992). Bas.: Lophozia rubescens R.M.Schust. et
Damsh., Phytologia 63 (5): 325, 1987 (Schuster and Damsholt 1987).
** subg. Sudeticae (Schljakov) L.Söderstr., De Roo et Hedd., Phytotaxa 3: 50,
2010 (Söderström et al. 2010b). Bas.: Lophozia sect. Sudeticae Schljakov, Pečen.
Mchi Sev. SSSR 3: 113, 1980 (Shliakov 1980a).
*** Barbilophozia sudetica (Nees ex Huebener) L.Söderstr., De Roo et Hedd., Phyto-
taxa 3: 50, 2010 (Söderström et al. 2010b). Bas.: Jungermannia sudetica Nees ex
Huebener, Hepaticol. germ.: 142, 1834 (Hübener 1834). 28
28 Barbilophozia sudetica was transferred from Lophozia by Söderström et al. (2010a), but transferred to
the new genus Pseudolophozia Konstant. and Vilnet (sister to Barbilophozia) by Konstantinova and
Vilnet (2009).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
52
*** Biantheridion (Grolle) Konstant. et Vilnet, Arctoa 18: 67, 2009 [2010] (Kons-
tantinova and Vilnet 2009). Bas.: Jamesoniella sect. Biantheridion Grolle, Trans.
Brit. Bryol. Soc. 4 (4): 662, 1964 (Grolle 1964i).
*** Biantheridion undulifolium (Nees) Konstant. et Vilnet, Arctoa 18: 67, 2009 [2010]
(Konstantinova and Vilnet 2009). Bas.: Jungermannia schraderi β undulifolia Nees,
Naturgesch. Eur. Leberm. 1: 306, 1833 (Nees 1833c).
*** Chandonanthus Mitt., Handb. N. Zeal. . 2: 753, 1867 (Hooker 1867).
*** Chandonanthus squarrosus (Menzies) Mitt., Handb. N. Zeal. . 2: 753, 1867
(Hooker 1867). Bas.: Jungermannia squarrosa Menzies, Musci Exot. 1: tab. 78,
1818 (Hooker 1818).
*** Crossocalyx Meyl., Bull. Soc. Vaud. Sci. Nat. 60 (249): 266, 1939 (Meylan 1939).
*** Crossocalyx hellerianus (Nees ex Lindenb.) Meyl., Bull. Soc. Vaud. Sci. Nat. 60
(249): 266, 1939 (Meylan 1939). Bas.: Jungermannia helleriana Nees ex Lindenb.,
Syn. hepat. eur: 64, 1829 (Lindenberg 1829).
*** Crossocalyx tenuis (Harry Williams) Schljakov, Novosti Sist. Nizš. Rast. 15: 246,
1978 (Shliakov 1978). Bas.: Anastrophyllum tenue Harry Williams, Bryologist 71
(1): 34, 1968 (Williams 1968).
*** Gymnocolea (Dumort.) Dumort., Recueil Observ. Jungerm.: 17, 1835 (Dumor-
tier 1835). Bas.: Jungermannia sect. Gymnocolea Dumort., Syll. Jungerm. Europ.:
52, 1831 (Dumortier 1831).
*** Gymnocolea borealis (Frisvoll et Moen) R.M.Schust., Lindbergia 12 (1): 7, 1986
(Schuster 1986). Bas.: Lophozia borealis Frisvoll et Moen, Lindbergia 6 (2): 138,
1980 [1981] (Frisvoll and Moen 1980).
** Gymnocolea fascinifera Potemkin, Arctoa 2: 76, 1993 (Potemkin 1993).
*** Gymnocolea inata (Huds.) Dumort., Recueil Observ. Jungerm.: 17, 1835 (Du-
mortier 1835). Bas.: Jungermannia inata Huds., Fl. Angl. (Hudson), ed. 2: 511,
1778 (Hudson 1778).
* Gymnocolea inata subsp. acutiloba (Schin.) R.M.Schust. et Damsh. ex L.Söder-
str. et Váňa, Lindbergia 27 (1): 43, 2002 (Söderström et al. 2002). Bas.: Lophozia
acutiloba Schin., Hedwigia 48 (3): 187, 1909 (Schiner 1909a). 29
*** Hamatostrepta Váňa et D.G.Long, Fieldiana, Bot. (n.ser.) 47: 134, 2008 (Váňa
and Long 2008).
*** Hamatostrepta concinna Váňa et D.G.Long, Fieldiana, Bot. (n.ser.) 47: 134, 2008
(Váňa and Long 2008).
29 Gymnocolea inata subsp. acutiloba is a problematic taxon that sometimes has been treated as conspecic
with Gymnocolea inata, sometimes as a separate species.
World checklist of hornworts and liverworts 53
** Hattoria R.M.Schust., Rev. Bryol. Lichénol. 30 (1/2): 69, 1961 (Schuster 1961a).
*** Hattoria yakushimensis (Horik.) R.M.Schust., Rev. Bryol. Lichénol. 30 (1/2): 70,
1961 (Schuster 1961a). Bas.: Anastrophyllum yakushimense Horik., J. Sci. Hiroshi-
ma Univ., Ser. B, Div. 2, Bot. 2: 149, 1934 (Horikawa 1934).
*** Isopaches H.Buch, Memoranda Soc. Fauna Fl. Fennica 8: 287, 1932 [1933]
(Buch 1932).
*** Isopaches alboviridis (R.M.Schust.) Schljakov, Novosti Sist. Nizš. Rast. 16: 205,
1979 (Shliakov 1979). Bas.: Lophozia alboviridis R.M.Schust., Hepat. Anthocero-
tae N. Amer. 2: 487, 1969 (Schuster 1969b).
*** Isopaches bicrenatus (Schmidel ex Hom.) H.Buch, Memoranda Soc. Fauna Fl.
Fennica 8: 288, 1932 [1933] (Buch 1932). Bas.: Jungermannia bicrenata Schmidel
ex Hom., Deutschl. Fl., eil 2 (Hom.): 11 (addenda), 1795 [1796] (Ho-
mann 1795).
*** Isopaches decolorans (Limpr.) H.Buch, Memoranda Soc. Fauna Fl. Fennica 8:
288, 1932 [1933] (Buch 1932). Bas.: Jungermannia decolorans Limpr., Jahresber.
Schles. Ges. Vaterl. Cult. 57: 316, 1879 [1880] (Limpricht 1879).
*** Isopaches pumicicola (Berggr.) Bakalin, Arctoa 17: 162, 2008 [2009] (Bakalin
2008b). Bas.: Lophozia pumicicola Berggr., New Zealand Hepat.: 21, 1898
(Berggren 1898).
*** Neoorthocaulis L.Söderstr., De Roo et Hedd., Phytotaxa 3: 49, 2010 (Söder-
ström et al. 2010b).
*** Neoorthocaulis attenuatus (Mart.) L.Söderstr., De Roo et Hedd., Phytotaxa 3: 49,
2010 (Söderström et al. 2010b). Bas.: Jungermannia quinquedentata δ attenuata
Mart., Fl. crypt. erlang.: 177, 1817 (Martius 1817).
*** Neoorthocaulis binsteadii (Kaal.) L.Söderstr., De Roo et Hedd., Phytotaxa 3: 49,
2010 (Söderström et al. 2010b). Bas.: Jungermannia binsteadii Kaal., Skr. Vidensk.-
Selsk. Christiana, Math.-Naturvidensk. Kl. 1898 (9): 9, 1898 (Kaalaas 1898).
*** Neoorthocaulis oerkei (F.Weber et D.Mohr) L.Söderstr., De Roo et Hedd., Phy-
totaxa 3: 50, 2010 (Söderström et al. 2010b). Bas.: Jungermannia oerkei F.Weber
et D.Mohr, Bot. Taschenb. (Weber): 410, 1807 (Weber and Mohr 1807).
** Neoorthocaulis hyperboreus (R.M.Schust.) L.Söderstr., De Roo et Hedd., Phyto-
taxa 3: 50, 2010 (Söderström et al. 2010b). Bas.: Lophozia oerkei var. hyperborea
R.M.Schust., Bull. Natl. Mus. Canada 164: 21, 1959 (Schuster et al. 1959).
*** Orthocaulis H.Buch, Memoranda Soc. Fauna Fl. Fennica 8: 293, 1932 [1933]
(Buch 1932).
*** Orthocaulis atlanticus (Kaal.) H.Buch, Memoranda Soc. Fauna Fl. Fennica 8: 294,
1932 [1933] (Buch 1932). Bas.: Jungermannia atlantica Kaal., Skr. Vidensk.-Selsk.
Christiana, Math.-Naturvidensk. Kl. 1898 (9): 11, 1898 (Kaalaas 1898).
* Orthocaulis cavifolius H.Buch et S.W.Arnell, Memoranda Soc. Fauna Fl. Fennica
26: 71, 1951 (Buch 1951).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
54
** Plicanthus R.M.Schust., Nova Hedwigia 74 (3/4): 484, 2002 (Schuster 2002a).
*** Plicanthus birmensis (Steph.) R.M.Schust., Beih. Nova Hedwigia 119: 223, 2002
(Schuster 2002b). Bas.: Chandonanthus birmensis Steph., Bull. Soc. Roy. Bot. Bel-
gique 38 (1): 43, 1899 (Stephani 1899h). 30
** Plicanthus dicilis (Steph.) L.Söderstr. et Váňa, Phytotaxa 81 (1): 30, 2013 (Váňa
et al. 2013a). Bas.: Chandonanthus dicilis Steph., J. & Proc. Roy. Soc. New
South Wales 48 (1/2): 101, 1914 (Stephani and Watts 1914).
** Plicanthus giganteus (Steph.) R.M.Schust., Nova Hedwigia 74 (3/4): 485, 2002
(Schuster 2002a). Bas.: Chandonanthus giganteus Steph., Wiss. Ergebn. Deut.
Zentr.-Afr. Exped. (1907-08), Bot. 2: 124, 1911 (Stephani 1911a). 31
*** Plicanthus hirtellus (F.Weber) R.M.Schust., Nova Hedwigia 74 (3/4): 492, 2002
(Schuster 2002a). Bas.: Jungermannia hirtella F.Weber, Hist. Musc. Hepat. Prodr.:
50, 1815 (Weber 1815).
*** Schizophyllopsis Váňa et L.Söderstr., Phytotaxa 152 (1): 48, 2013 (Váňa et al.
2013f). Nom. nov. pro Anastrophyllum subg. Schizophyllum R.M.Schust., Hepat.
Anthocerotae N. Amer. 2: 739, 1969 (Schuster 1969b). 32
** Schizophyllopsis aristata (Herzog ex N.Kitag.) Váňa et L.Söderstr., Phytotaxa 152
(1): 48, 2013 (Váňa et al. 2013f). Bas.: Anastrophyllum bidens var. aristatum Her-
zog ex N.Kitag., J. Hattori Bot. Lab. 33: 216, 1970 (Kitagawa 1970).
*** Schizophyllopsis bidens (Reinw., Blume et Nees) Váňa et L.Söderstr., Phytotaxa
152 (1): 48, 2013 (Váňa et al. 2013f). Bas.: Jungermannia bidens Reinw., Blume
et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 208,
1824 [1825] (Reinwardt et al. 1824a).
** Schizophyllopsis lanciloba (Steph.) Váňa et L.Söderstr., Phytotaxa 152 (1): 48, 2013
(Váňa et al. 2013f). Bas.: Anastrophyllum lancilobum Steph., Sp. Hepat. (Stephani)
6: 107, 1917 (Stephani 1917a).
*** Schizophyllopsis papillosa (J.J.Engel et Braggins) Váňa et L.Söderstr., Phytotaxa 152
(1): 48, 2013 (Váňa et al. 2013f). Bas.: Anastrophyllum papillosum J.J.Engel et
Braggins, J. Bryol. 20 (2): 381, 1998 (Engel and Braggins 1998).
*** Schizophyllopsis sphenoloboides (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 152
(1): 49, 2013 (Váňa et al. 2013f). Bas.: Anastrophyllum sphenoloboides R.M.Schust.,
Hepat. Anthocerotae N. Amer. 2: 741, 1969 (Schuster 1969b).
*** Schljakovia Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010] (Konstantinova and
Vilnet 2009).
30 Plicanthus birmensis seems to be closely related to Tetralophozia (Vilnet et al. 2010), but Grolle (1995)
considered it a possibly depauperate form of Plicanthus hirtellus. However, it may be a good species,
possibly of the genus Tetralophozia.
31 Plicanthus giganteus is often treated as a subspecies of Plicanthus hirtellus (e.g. Pócs and Lye 1999).
32 Schizophyllopsis was originally published as Schizophyllum (R.M.Schust.) Váňa et L.Söderstr. (Váňa et al.
2013k), but that is a later homonym of Schizophyllum Fr. (1815; Fungus) and Nutt. (1841; Asteraceae).
World checklist of hornworts and liverworts 55
*** Schljakovia kunzeana (Huebener) Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010]
(Konstantinova and Vilnet 2009). Bas.: Jungermannia kunzeana Huebener, Hepa-
ticol. germ.: 115, 1834 (Hübener 1834).
*** Schljakovianthus Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010] (Konstanti-
nova and Vilnet 2009).
*** Schljakovianthus quadrilobus (Lindb.) Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010]
(Konstantinova and Vilnet 2009). Bas.: Jungermannia quadriloba Lindb., Kongl.
Svens ka Vetensk.-Akad. Handl. (n.ser.) 23 (5): 55, 1889 (Lindberg and Arnell 1889).
*** Sphenolobopsis R.M.Schust. et N.Kitag., Nova Hedwigia 22: 152, 1971 [1972]
(Schuster 1971b).
*** Sphenolobopsis pearsonii (Spruce) R.M.Schust., Nova Hedwigia 22: 153, 1971
[1972] (Schuster 1971b). Bas.: Jungermannia pearsonii Spruce, J. Bot. 19: 33,
1881 (Spruce 1881b).
*** Sphenolobus (Lindb.) Berggr., New Zealand Hepat.: 22, 1898 (Berggren 1898).
Bas.: Jungermannia sect. Sphenolobus Lindb., Not. Sällsk. Fauna Fl. Fenn. Förh.
13: 369, 1874 (Lindberg 1874a).
*** Sphenolobus austroamericanus (Váňa) Váňa, Phytotaxa 81 (1): 30, 2013 (Váňa et
al. 2013a). Bas.: Anastrophyllum austroamericanum Váňa, J. Hattori Bot. Lab. 48:
225, 1980 (Váňa 1980).
*** Sphenolobus minutus (Schreb. ex D.Crantz) Berggr., New Zealand Hepat.: 22,
1898 (Berggren 1898). Bas.: Jungermannia minuta Schreb. ex D.Crantz, Forts.
Hist. Grönland: 285, 1770 (Crantz 1770; non vidi).
*** Sphenolobus saxicola (Schrad.) Steph., Bull. Herb. Boissier (sér. 2) 2 (2): 168 (160),
1902 (Stephani 1902d). Bas.: Jungermannia saxicola Schrad., Syst. Samml. Crypt.
Gew. 2: 4, 1797 (Schrader 1797).
** Tetralophozia (R.M.Schust.) Schljakov, Novosti Sist. Nizš. Rast. 13: 227, 1976
(Shliakov 1976). Bas.: Chandonanthus subg. Tetralophozia R.M.Schust., J. Hattori
Bot. Lab. 23: 206, 1960 [1961] (Schuster 1960a).
*** Tetralophozia cavallii (Gola) Váňa, Trop. Bryol. 8: 102, 1993 (Váňa 1993). Bas.:
Blepharostomum cavallii Gola, Ann. Bot. (Rome) 6 (2): 274, 1907 (Gola 1907).
*** Tetralophozia liformis (Steph.) Urmi, J. Bryol. 12 (3): 394, 1983 (Urmi 1983).
Bas.: Chandonanthus liformis Steph., Sp. Hepat. (Stephani) 3: 644, 1909 (Stephani
1909a).
** Tetralophozia pilifera (Steph.) R.M.Schust., Nova Hedwigia 74 (3/4): 482, 2002
(Schuster 2002a). Bas.: Chandonanthus pilifer Steph., Sp. Hepat. (Stephani) 3:
644, 1909 (Stephani 1909a).
*** Tetralophozia setiformis (Ehrh.) Schljakov, Novosti Sist. Nizš. Rast. 13: 228, 1976
(Shliakov 1976). Bas.: Jungermannia setiformis Ehrh., Hannover. Mag. 22 (8):
142, 1784 (Ehrhart 1784).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
56
*** Zantenia (S.Hatt.) Váňa et J.J.Engel, Mem. New York Bot. Gard. 105: 29, 2013
(Váňa and Engel 2013). Bas.: Anastrophyllum subg. Zantenia S.Hatt., Bot. Mag.
(Tokyo) 79 (937): 342, 1966 (Hattori 1966a).
*** Zantenia borneensis (Herzog) Váňa et J.J.Engel, Mem. New York Bot. Gard. 105:
29, 2013 (Váňa and Engel 2013). Bas.: Anastrophyllum borneense Herzog, Trans.
Brit. Bryol. Soc. 1 (4): 282, 1950 (Herzog 1950a).
*** Zantenia denticulata (Grolle) Váňa et J.J.Engel, Mem. New York Bot. Gard. 105:
30, 2013 (Váňa and Engel 2013). Bas.: Anastrophyllum denticulatum Grolle, Nova
Hedwigia 16: 148, 1968 (Grolle 1968d).
*** Zantenia karstenii (Schin.) Váňa et J.J.Engel, Mem. New York Bot. Gard. 105:
30, 2013 (Váňa and Engel 2013). Bas.: Anastrophyllum karstenii Schin., Nova Acta
Acad. Caes. Leop.-Carol. German. Nat. Cur. 60 (2): 268, 1893 (Schiner 1893a).
*** Zantenia prionophylla (S.Hatt.) Váňa et J.J.Engel, Mem. New York Bot. Gard.
105: 30, 2013 (Váňa and Engel 2013). Bas.: Anastrophyllum prionophyllum S.Hatt.,
Bot. Mag. (Tokyo) 79 (937): 342, 1966 (Hattori 1966a).
*** Cephaloziaceae Mig.
by J. Váňa with contributions by S.R. Gradstein (Odontoschisma)
Cephaloziaceae was recently studied by Vilnet et al. (2012) and Feldberg et al. (2013).
Some taxonomic and nomenclatural notes were published by Váňa et al. (2013g,
2013i) and Gradstein et al. (2014b).
** Alobielloideae R.M.Schust.
** Alobiella (Spruce) Schin., Hepat. (Engl.-Prantl): 98, 1893 (Schiner 1893b).
Bas.: Cephalozia subg. Alobiella Spruce, Cephalozia: 28, 1882 (Spruce 1882).
*** Alobiella husnotii (Spruce) Schin., Hepat. (Engl.-Prantl): 98, 1893 (Schiner
1893b). Bas.: Cephalozia husnotii Spruce, Cephalozia: 30, 1882 (Spruce 1882).
** Alobiellopsis R.M.Schust., Nova Hedwigia 10 (1/2): 25, 1965 (Schuster 1965b).
*** Alobiellopsis acroscypha (Spruce) R.M.Schust., Nova Hedwigia 10 (1/2): 25, 1965
(Schuster 1965b). Bas.: Cephalozia acroscypha Spruce, Cephalozia: 30, 1882
(Spruce 1882).
*** Alobiellopsis dominicensis (Spruce) Fulford, Mem. New York Bot. Gard. 11 (3):
350, 1968 (Fulford 1968). Bas.: Alobiella dominicensis Spruce, J. Linn. Soc., Bot.
30 (210): 355, 1895 (Gepp 1895b).
*** Alobiellopsis heteromorpha (Lehm.) R.M.Schust., Bull. Natl. Sci. Mus. Tokyo
(n.ser.) 12 (3): 682, 1969 (Schuster 1969a). Bas.: Jungermannia heteromorpha
Lehm., Linnaea 4: 362, 1829 (Lehmann 1829).
World checklist of hornworts and liverworts 57
*** Alobiellopsis parvifolia (Steph.) R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.)
12 (3): 679, 1969 (Schuster 1969a). Bas.: Alobiella parvifolia Steph., Bull. Herb.
Boissier (sér. 2) 8 (8): 568 (352), 1908 (Stephani 1908e).
*** Alobiellopsis pillansii (Sim) R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 12
(3): 683, 1969 (Schuster 1969a). Bas.: Cephalozia pillansii Sim, Trans. Roy. Soc.
South Africa 15 (1): 87, 1926 (Sim 1926).
** Cephalozioideae Müll.Frib.
*** Cephalozia (Dumort.) Dumort., Recueil Observ. Jungerm.: 18, 1835 (Dumor-
tier 1835). Bas.: Jungermannia sect. Cephalozia Dumort., Syll. Jungerm. Europ.:
60, 1831 (Dumortier 1831).
*** Cephalozia acuminata (Herzog) Váňa, Phytotaxa 112 (1): 11, 2013 (Váňa et al.
2013g). Bas.: Hygrobiella acuminata Herzog, Trans. Brit. Bryol. Soc. 1 (4): 292,
1950 (Herzog 1950a).
** Cephalozia acutiloba (Inoue) Váňa, Phytotaxa 112 (1): 11, 2013 (Váňa et al.
2013g). Bas.: Metahygrobiella acutiloba Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.)
10 (2): 155, 1967 (Inoue 1967c).
*** Cephalozia albula Steph., Hedwigia 32 (5): 318, 1893 (Stephani 1893d). Nom.
nov. pro Jungermannia albula Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 93, 1860
[1861] (Mitten 1860c), nom. illeg.
*** Cephalozia ambigua C.Massal., Malpighia 21 (7/8): 310, 1907 (Massalongo 1907).
*** Cephalozia austrigena R.M.Schust. ex J.J.Engel, Novon 17 (3): 312, 2007 (Engel
2007). Based on: Cephalozia bicuspidata subsp. austrigena R.M.Schust., Hepat.
Anthocerotae N. Amer. 3: 712, 1974 (Schuster 1974), nom. inval.
*** Cephalozia badia (Gottsche) Steph., Bull. Herb. Boissier (sér. 2) 8 (7): 483 (313),
1908 (Stephani 1908f). Bas.: Jungermannia badia Gottsche, Int. Polarforsch.,
Deutsch. Exped. 2: 452, 1890 (Gottsche 1890).
*** Cephalozia bicuspidata (L.) Dumort., Recueil Observ. Jungerm.: 18, 1835 (Dumor-
tier 1835). Bas.: Jungermannia bicuspidata L., Sp. Pl. 1: 1132, 1753 (Linnaeus 1753).
* Cephalozia bicuspidata subsp. lammersiana (Huebener) R.M.Schust., Hepat. An-
thocerotae N. Amer. 3: 730, 1974 (Schuster 1974). Bas.: Jungermannia lammersi-
ana Huebener, Flora 15 (20): 306, 1832 (Hübener 1832).
*** Cephalozia chilensis (J.J.Engel et R.M.Schust.) R.M.Schust., Beih. Nova Hedwi-
gia 119: 29, 2002 (Schuster 2002b). Bas.: Metahygrobiella chilensis J.J.Engel et
R.M.Schust., Brittonia 40 (2): 203, 1988 (Engel and Schuster 1988).
*** Cephalozia conchata (Grolle et Váňa) Váňa, Syst. Bot. 40 (1): 38, 2015 (Shaw et
al. 2015). Bas.: Jungermannia conchata Grolle et Váňa, Fragm. Florist. Geobot. 37
(1): 3, 1992 (Grolle and Váňa 1992).
*** Cephalozia crossii Spruce, Cephalozia: 46, 1882 (Spruce 1882).
*** Cephalozia darjeelingensis Udar et D.Kumar, Geophytology 6 (1): 36, 1976 (Udar
and Kumar 1976).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
58
*** Cephalozia drucei (R.M.Schust.) Váňa, Phytotaxa 112 (1): 11, 2013 (Váňa et al.
2013g). Bas.: Metahygrobiella drucei R.M.Schust., J. Hattori Bot. Lab. 26: 273,
1963 (Schuster 1963b).
*** Cephalozia fuegiensis Váňa, Phytotaxa 112 (1): 12, 2013 (Váňa et al. 2013g). Nom. nov.
pro Hygrobiella dusenii Steph., Sp. Hepat. (Stephani) 6: 444, 1924 (Stephani 1924).
*** Cephalozia hamatiloba Steph., Bull. Herb. Boissier (sér. 2) 8 (6): 427 (303), 1908
(Stephani 1908g).
*** Cephalozia hamatiloba subsp. siamensis (N.Kitag.) Váňa, Acta Bot. Fenn. 177: 16,
2004 (Koponen et al. 2004). Bas.: Cephalozia siamensis N.Kitag., J. Hattori Bot.
Lab. 32: 293, 1969 (Kitagawa 1969c).
*** Cephalozia lacinulata (J.B.Jack ex Gottsche et Rabenh.) Spruce, Cephalozia: 45,
1882 (Spruce 1882). Bas.: Jungermannia lacinulata J.B.Jack ex Gottsche et Ra-
benh., Hepat. Eur., Leberm. 62-64: no. 624, 1877 (Gottsche and Rabenhorst
1877).
*** Cephalozia lucens (A.Evans) Steph., Bull. Herb. Boissier (sér. 2) 8 (7): 496 (326),
1908 (Stephani 1908f). Bas.: Jungermannia lucens A.Evans, Trans. Connecticut
Acad. Arts 8 (15): 258, 1891 (Evans 1891).
*** Cephalozia macgregorii (Steph.) Váňa, Phytotaxa 112 (1): 12, 2013 (Váňa et al. 2013g).
Bas.: Hygrobiella macgregorii Steph., Hedwigia 34 (2): 45, 1895 (Stephani 1895c).
*** Cephalozia macounii (Austin) Austin, Hepat. bor.-amer.: 14, 1873 (Austin 1873).
Bas.: Jungermannia macounii Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 222,
1869 (Austin 1869).
*** Cephalozia maxima Steph., Sp. Hepat. (Stephani) 6: 441, 1924 (Stephani 1924).
*** Cephalozia mollusca (De Not.) Váňa, Phytotaxa 112 (1): 12, 2013 (Váňa et al. 2013g).
Bas.: Jungermannia mollusca De Not., Epat. Borneo: 16, 1874 (De Notaris 1874).
* Cephalozia neesiana Steph., Bull. Herb. Boissier (sér. 2) 8 (6): 429 (305), 1908
(Stephani 1908g). 33
*** Cephalozia nishimurae (N.Kitag.) Váňa, Phytotaxa 112 (1): 12, 2013 (Váňa et al.
2013g). Bas.: Hygrobiella nishimurae N.Kitag., Misc. Bryol. Lichenol. 9 (4): 69,
1982 (Kitagawa 1982).
** Cephalozia pachygyna R.M.Schust. ex J.J.Engel, Novon 17 (3): 313, 2007 (Engel 2007).
** Cephalozia physocaula (Hook.f. et Taylor) Steph., Bull. Herb. Boissier (sér. 2) 8
(7): 485 (315), 1908 (Stephani 1908f). Bas.: Jungermannia physocaula Hook.f. et
Taylor, London J. Bot. 3: 455, 1844 (Hooker and Taylor 1844b).
** Cephalozia schusteriana J.J.Engel, Novon 17 (3): 313, 2007 (Engel 2007).
*** Cephalozia stolonacea (Herzog) Váňa, Phytotaxa 112 (1): 12, 2013 (Váňa et al.
2013g). Bas.: Hygrobiella stolonacea Herzog, Trans. Brit. Bryol. Soc. 1 (4): 293,
1950 (Herzog 1950a).
*** Cephalozia tubulata (Hook.f. et Taylor) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 417, 1877 (Trevisan 1877). Bas.: Jungermannia tubu-
lata Hook.f. et Taylor, London J. Bot. 3: 463, 1844 (Hooker and Taylor 1844b).
33 Cephalozia neesiana may be conspecic with Cephalozia hamatiloba (Váňa 1988).
World checklist of hornworts and liverworts 59
Excluded from the genus
* Cephalozia hians Steph., Sp. Hepat. (Stephani) 6: 441, 1924 (Stephani 1924). 34
* Cephalozia indica Udar et D.Kumar, Geophytology 6 (1): 42, 1976 (Udar and
Kumar 1976). 35
* Cephalozia kodaikanalensis G.Asthana et Saumya Srivast., Geophytology 43 (1):
63, 2013 (Asthana and Srivastava 2013). 36
* Cephalozia parvifolia Arnell, Rev. Bryol. 25 (1): 1, 1898 (Arnell 1898). 37
* Cephalozia tricuspidata (Nees) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 417, 1877 (Trevisan 1877). Bas.: Jungermannia tricuspidata
Nees, Enum. Pl. Crypt. Javae: 31, 1830 (Nees 1830). 38
*** Fuscocephaloziopsis Fulford, Mem. New York Bot. Gard. 11 (3): 353, 1968
(Fulford 1968).
** Fuscocephaloziopsis anis (Lindb. ex Steph.) Váňa et L.Söderstr., Phytotaxa 112
(1): 9, 2013 (Váňa et al. 2013g). Bas.: Cephalozia anis Lindb. ex Steph., Bull.
Herb. Boissier (sér. 2) 8 (4): 277 (291), 1908 (Stephani 1908j).
*** Fuscocephaloziopsis africana (Váňa) Váňa et L.Söderstr., Phytotaxa 112 (1): 9,
2013 (Váňa et al. 2013g). Bas.: Cephalozia africana Váňa, Beih. Nova Hedwigia
90: 187, 1988 (Váňa 1988).
*** Fuscocephaloziopsis albescens (Hook.) Váňa et L.Söderstr., Phytotaxa 112 (1): 9,
2013 (Váňa et al. 2013g). Bas.: Jungermannia albescens Hook., Brit. Jungermann.:
tab. 72, 1815 (Hooker 1815).
* Fuscocephaloziopsis albescens var. islandica (Nees) Váňa et L.Söderstr., Phytotaxa
112 (1): 9, 2013 (Váňa et al. 2013g). Bas.: Jungermannia islandica Nees, Natur-
gesch. Eur. Leberm. 2: 29, 1836 (Nees 1836).
*** Fuscocephaloziopsis baldwinii (C.M.Cooke) Váňa et L.Söderstr., Phytotaxa 112
(1): 9, 2013 (Váňa et al. 2013g). Bas.: Cephalozia baldwinii C.M.Cooke, Trans.
Connecticut Acad. Arts 12 (1): 35, 1904 (Cooke 1904).
** Fuscocephaloziopsis biloba (Herzog) Fulford, Mem. New York Bot. Gard. 11 (3):
355, 1968 (Fulford 1968). Bas.: Alobiella biloba Herzog, Feddes Repert. Spec.
Nov. Regni Veg. 57 (1/2): 166, 1955 (Herzog 1955).
*** Fuscocephaloziopsis catenulata (Huebener) Váňa et L.Söderstr., Phytotaxa 112 (1):
9, 2013 (Váňa et al. 2013g). Bas.: Jungermannia catenulata Huebener, Hepaticol.
germ.: 169, 1834 (Hübener 1834).
*** Fuscocephaloziopsis catenulata subsp. nipponica (S.Hatt.) Váňa et L.Söderstr., Phy-
totaxa 112 (1): 9, 2013 (Váňa et al. 2013g). Bas.: Cephalozia nipponica S.Hatt.,
Bull. Tokyo Sci. Mus. 11: 74, 1944 (Hattori 1944d).
34 Cephalozia hians is probably an Isotachis or Lophocolea species.
35 Cephalozia indica is a Lophocolea species and probably conspecic with “Cephalozia” kodaikanalensis.
36 Cephalozia kodaikanalensis is a Lophocolea species, possibly conspecic with “Cephalozia” indica.
37 Cephalozia parvifolia is a Cephaloziella species although the type specimen has not been found.
38 Cephalozia tricuspidata is a Lepidozia species (Schiner 1898b).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
60
*** Fuscocephaloziopsis connivens (Dicks.) Váňa et L.Söderstr., Phytotaxa 112 (1): 9,
2013 (Váňa et al. 2013g). Bas.: Jungermannia connivens Dicks., Fasc. Pl. Crypt.
Brit. 4: 19, 1801 (Dickson 1801).
*** Fuscocephaloziopsis connivens subsp. ssa (Steph.) Váňa et L.Söderstr., Phytotaxa
112 (1): 9, 2013 (Váňa et al. 2013g). Bas.: Cephalozia ssa Steph., Hedwigia 30
(5): 204, 1891 (Stephani 1891a).
*** Fuscocephaloziopsis connivens subsp. sandwicensis (Mont.) Váňa et L.Söderstr.,
Phytotaxa 112 (1): 9, 2013 (Váňa et al. 2013g). Bas.: Jungermannia sandwicensis
Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 249, 1843 (Montagne 1843).
*** Fuscocephaloziopsis crassifolia (Lindenb. et Gottsche) Váňa et L.Söderstr., Phyto-
taxa 112 (1): 10, 2013 (Váňa et al. 2013g). Bas.: Jungermannia crassifolia Lindenb.
et Gottsche, Syn. Hepat. 5: 685, 1847 (Gottsche et al. 1847).
*** Fuscocephaloziopsis gollanii (Steph.) Váňa et L.Söderstr., Phytotaxa 112 (1): 10,
2013 (Váňa et al. 2013g). Bas.: Cephalozia gollanii Steph., Bull. Herb. Boissier
(sér. 2) 8 (6): 428 (304), 1908 (Stephani 1908g).
*** Fuscocephaloziopsis leucantha (Spruce) Váňa et L.Söderstr., Phytotaxa 112 (1): 10,
2013 (Váňa et al. 2013g). Bas.: Cephalozia leucantha Spruce, Cephalozia: 68, 1882
(Spruce 1882).
*** Fuscocephaloziopsis loitlesbergeri (Schin.) Váňa et L.Söderstr., Phytotaxa 112 (1):
10, 2013 (Váňa et al. 2013g). Bas.: Cephalozia loitlesbergeri Schin., Österr. Bot.
Z. 62 (1): 10, 1912 (Schiner 1912b).
*** Fuscocephaloziopsis lunulifolia (Dumort.) Váňa et L.Söderstr., Phytotaxa 112 (1):
10, 2013 (Váňa et al. 2013g). Bas.: Jungermannia lunulifolia Dumort., Syll. Jung-
erm. Europ.: 61, 1831 (Dumortier 1831). 39
*** Fuscocephaloziopsis macrostachya (Kaal.) Váňa et L.Söderstr., Phytotaxa 112 (1):
10, 2013 (Váňa et al. 2013g). Bas.: Cephalozia macrostachya Kaal., Rev. Bryol. 29
(1): 8, 1902 (Kaalaas 1902).
*** Fuscocephaloziopsis macrostachya subsp. australis (R.M.Schust.) Váňa et L.Söder-
str., Phytotaxa 112 (1): 10, 2013 (Váňa et al. 2013g). Bas.: Cephalozia macrosta-
chya subsp. australis R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 754, 1974
(Schuster 1974).
*** Fuscocephaloziopsis macrostachya subsp. macrostachya var. spiniora (Schin.) Váňa
et L.Söderstr., Phytotaxa 112 (1): 10, 2013 (Váňa et al. 2013g). Bas.: Cephalozia
spiniora Schin., Hedwigia 54 (6): 323, 1914 (Schiner 1914a).
*** Fuscocephaloziopsis monticola (J.D.Godfrey) Váňa et L.Söderstr., Phytotaxa 112
(1): 11, 2013 (Váňa et al. 2013g). Bas.: Schoeldia monticola J.D.Godfrey, Bryolo-
gist 79 (3): 315, 1976 (Godfrey 1976).
*** Fuscocephaloziopsis pachycaulis (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 112
(1): 11, 2013 (Váňa et al. 2013g). Bas.: Cephalozia pachycaulis R.M.Schust., Bryo-
logist 96 (4): 623, 1993 (Schuster 1993b).
39 Fuscocephaloziopsis lunulifolia is a species complex also including Fuscocephaloziopsis anis and Fusco-
cephaloziopsis schusteri.
World checklist of hornworts and liverworts 61
*** Fuscocephaloziopsis pleniceps (Austin) Váňa et L.Söderstr., Phytotaxa 112 (1): 11,
2013 (Váňa et al. 2013g). Bas.: Jungermannia pleniceps Austin, Proc. Acad. Nat.
Sci. Philadelphia 21: 222, 1869 (Austin 1869).
** Fuscocephaloziopsis pleniceps var. caroliniana (R.M.Schust.) Váňa et L.Söderstr., Phy-
totaxa 112 (1): 11, 2013 (Váňa et al. 2013g). Bas.: Cephalozia pleniceps var. carolini-
ana R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 780, 1974 (Schuster 1974).
*** Fuscocephaloziopsis pulvinata (Steph.) Fulford, Mem. New York Bot. Gard. 11 (3):
355, 1968 (Fulford 1968). Bas.: Alobiella pulvinata Steph., Bull. Herb. Boissier
(sér. 2) 8 (8): 572 (356), 1908 (Stephani 1908e).
* Fuscocephaloziopsis schusteri (Sushil K.Singh et D.K.Singh) Váňa, Phytotaxa 183
(4): 291, 2014 (Váňa et al. 2014a). Bas.: Cephalozia schusteri Sushil K.Singh et
D.K.Singh, Lindbergia 32 (1): 1, 2007 (Singh and Singh 2007a).
*** Fuscocephaloziopsis subintegra Gradst. et Váňa, Cryptog. Bryol. 25 (3): 274, 2004
(Parolly et al. 2004).
*** Fuscocephaloziopsis zoopsioides (Horik.) Váňa et L.Söderstr., Phytotaxa 112 (1): 11,
2013 (Váňa et al. 2013g). Bas.: Cephalozia zoopsioides Horik., J. Sci. Hiroshima
Univ., Ser. B, Div. 2, Bot. 2: 178, 1934 (Horikawa 1934).
*** Nowellia Mitt., Nat. hist. Azores: 321, 1870 (Mitten 1870).
** sect. Acronowellia R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 816, 1974
(Schuster 1974).
*** Nowellia reedii H.Rob., Bryologist 73 (1): 150, 1970 (Robinson 1970).
*** Nowellia yunckeri Fulford, Mem. New York Bot. Gard. 11 (3): 329, 1968 (Fulford
1968).
** sect. Metanowellia (Grolle) R.M.Schust., Hepat. Anthocerotae N. Amer. 3:
816, 1974 (Schuster 1974). Bas.: Nowellia subg. Metanowellia Grolle, J. Hattori
Bot. Lab. 31: 33, 1968 (Grolle 1968b).
*** Nowellia borneensis (De Not.) Schin., Hepat. (Engl.-Prantl): 98, 1893 (Schiner
1893b). Bas.: Jungermannia curvifolia var. borneensis De Not., Epat. Borneo: 19,
1874 (De Notaris 1874).
*** Nowellia dominicensis Steph., Sp. Hepat. (Stephani) 6: 443, 1924 (Stephani 1924).
*** Nowellia evansii Grolle, J. Hattori Bot. Lab. 31: 33, 1968 (Grolle 1968b).
*** Nowellia langii Pearson, J. Linn. Soc., Bot. 46 (305): 25, 1922 (Pearson 1922b).
*** Nowellia pusilla Grolle, J. Hattori Bot. Lab. 31: 37, 1968 (Grolle 1968b).
*** Nowellia wrightii (Gottsche ex Spruce) Steph. ex Duss, Enum. musc. Antilles
franç., Hép.: 21, 1903 (Duss 1903). Bas.: Cephalozia wrightii Gottsche ex Spruce,
J. Linn. Soc., Bot. 30 (210): 354, 1895 (Gepp 1895b).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
62
** sect. Nowellia
*** Nowellia aciliata (P.C.Chen et P.C.Wu) Mizut., Hikobia 11: 469, 1994 (Mizu-
tani 1994). Bas.: Nowellia curvifolia var. aciliata P.C.Chen et P.C.Wu, Obs. .
Hwangs.: 6, 1965 (Chen and Wu 1965).
*** Nowellia curvifolia (Dicks.) Mitt., Nat. hist. Azores: 321, 1870 (Mitten 1870). Bas.:
Jungermannia curvifolia Dicks., Fasc. Pl. Crypt. Brit. 2: 15, 1790 (Dickson 1790).
** Odontoschismatoideae H.Buch ex Grolle
*** Odontoschisma (Dumort.) Dumort., Recueil Observ. Jungerm.: 19, 1835 (Du-
mortier 1835). Bas.: Pleuroschisma sect. Odontoschisma Dumort., Syll. Jungerm.
Europ.: 68, 1831 (Dumortier 1831).
** sect. Cladopodiella (H.Buch) Gradst., S.C.Aranda et Vanderp., Taxon 63 (5):
1017, 2014 (Aranda et al. 2014). Bas.: Cladopodiella H.Buch, Memoranda Soc.
Fauna Fl. Fennica 1: 89, 1927 (Buch 1927).
*** Odontoschisma francisci (Hook.) L.Söderstr. et Váňa, Phytotaxa 112 (1): 12, 2013
(Váňa et al. 2013g). Bas.: Jungermannia francisci Hook., Brit. Jungermann.: tab.
49, 1813 (Hooker 1813).
** sect. Denudata R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 833, 1974
(Schuster 1974).
*** Odontoschisma brasiliense Steph., Bull. Herb. Boissier (sér. 2) 8 (8): 585 (369),
1908 (Stephani 1908e).
** Odontoschisma cleei Gradst., S.C.Aranda et Vanderp., Taxon 63 (5): 1017, 2014
(Aranda et al. 2014).
*** Odontoschisma denudatum (Mart.) Dumort., Recueil Observ. Jungerm.: 19, 1835
(Dumortier 1835). Bas.: Jungermannia scalaris var. denudata Mart., Fl. crypt. er-
lang.: 183, 1817 (Martius 1817).
** Odontoschisma denudatum subsp. naviculare (Steph.) Gradst., S.C.Aranda et Van-
derp., Taxon 63 (5): 1019, 2014 (Aranda et al. 2014). Bas.: Jamesoniella navicula-
ris Steph., Sp. Hepat. (Stephani) 6: 101, 1917 (Stephani 1917a).
** Odontoschisma denudatum subsp. sandvicense (Ångstr.) Gradst., S.C.Aranda et
Vanderp., Taxon 63 (5): 1019, 2014 (Aranda et al. 2014). Bas.: Sphagnoecetis
sandvicensis Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 29 (4): 22, 1872 (Ång-
ström 1872).
*** Odontoschisma elongatum (Lindb.) A.Evans, Rhodora 14 (157): 13, 1912 (Evans
1912b). Bas.: Odontoschisma denudatum f. elongatum Lindb., Helsingf. Dagbl.
1874 (45, 16 Feb): 2, 1874 (Lindberg 1874b).
*** Odontoschisma engelii Gradst. et Burghardt, Fieldiana, Bot. (n.ser.) 47: 194, 2008
(Gradstein and Burghardt 2008).
World checklist of hornworts and liverworts 63
*** Odontoschisma longiorum (Taylor) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 419, 1877 (Trevisan 1877). Bas.: Sphagnoecetis longiora
Taylor, London J. Bot. 5: 281, 1846 (Taylor 1846a).
*** Odontoschisma macounii (Austin) Underw., Bull. Illinois State Lab. Nat. Hist. 2
(1): 92, 1884 (Underwood 1884). Bas.: Sphagnoecetis macounii Austin, Bull. Tor-
rey Bot. Club 3 (3): 13, 1872 (Austin 1872).
*** Odontoschisma portoricense (Hampe et Gottsche) Steph., Hedwigia 27 (11/12):
296, 1888 (Stephani 1888c). Bas.: Sphagnoecetis portoricensis Hampe et Gottsche,
Linnaea 25 (3): 343, 1852 [1853] (Hampe and Gottsche 1852).
** Odontoschisma pseudogrosseverrucosum Gradst., S.C.Aranda et Vanderp., Taxon 63
(5): 1019, 2014 (Aranda et al. 2014).
*** Odontoschisma purpuratum Herzog, Trans. Brit. Bryol. Soc. 1 (4): 297, 1950
(Herzog 1950a).
*** Odontoschisma soratamum Fulford, Mem. New York Bot. Gard. 11 (3): 338, 1968
(Fulford 1968).
*** Odontoschisma variabile (Lindenb. et Gottsche) Trevis., Mem. Reale Ist. Lombar-
do Sci. (Ser. 3), C. Sci. Mat. 4 (13): 419, 1877 (Trevisan 1877). Bas.: Sphagnoecetis
variabilis Lindenb. et Gottsche, Syn. Hepat. 5: 688, 1847 (Gottsche et al. 1847).
** Odontoschisma zhui Gradst., S.C.Aranda et Vanderp., Taxon 63 (5): 1020, 2014
(Aranda et al. 2014).
** sect. Iwatsukia (N.Kitag.) Gradst., S.C.Aranda et Vanderp., Phytotaxa 162 (4):
232, 2014 (Gradstein et al. 2014b). Bas.: Iwatsukia N.Kitag., J. Hattori Bot. Lab.
27: 178, 1964 (Kitagawa 1964).
*** Odontoschisma bidum (Fulford) Gradst., S.C.Aranda et Vanderp., Phytotaxa 162
(4): 232, 2014 (Gradstein et al. 2014b). Bas.: Cladomastigum bidum Fulford,
Acta Bot. Venez. 2 (5/8): 80, 1967 (Fulford 1967).
*** Odontoschisma jishibae (Steph.) L.Söderstr. et Váňa, Phytotaxa 112 (1): 13, 2013
(Váňa et al. 2013g). Bas.: Cephalozia jishibae Steph., Sp. Hepat. (Stephani) 6: 437,
1924 (Stephani 1924).
*** Odontoschisma spinosum (Fulford) Gradst., S.C.Aranda et Vanderp., Phytotaxa
162 (4): 232, 2014 (Gradstein et al. 2014b). Bas.: Cladomastigum spinosum Ful-
ford, Mem. New York Bot. Gard. 23: 840, 1972 (Fulford 1972).
** sect. Neesia Gradst., S.C.Aranda et Vanderp., Taxon 63 (5): 1017, 2014 (Aran-
da et al. 2014).
*** Odontoschisma uitans (Nees) L.Söderstr. et Váňa, Phytotaxa 112 (1): 12, 2013 (Váňa
et al. 2013g). Bas.: Jungermannia uitans Nees, Flora 6 (2): 30, 1823 (Link 1823).
** sect. Odontoschisma
*** Odontoschisma grosseverrucosum Steph., Bull. Herb. Boissier (sér. 2) 8 (8): 593
(377), 1908 (Stephani 1908e).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
64
*** Odontoschisma sphagni (Dicks.) Dumort., Recueil Observ. Jungerm.: 19, 1835
(Dumortier 1835). Bas.: Jungermannia sphagni Dicks., Fasc. Pl. Crypt. Brit. 1: 6,
1785 (Dickson 1785).
** Odontoschisma steyermarkii Gradst. et Ilk.-Borg., Nova Hedwigia 100 (1/2): 38,
2015 [2014 online] (Gradstein and Ilkiu-Borges 2015).
Incertae sedis
* Odontoschisma obcordatum (Spruce) Steph., Bull. Herb. Boissier (sér. 2) 8 (8): 586
(370), 1908 (Stephani 1908e). Bas.: Cephalozia obcordata Spruce, Cephalozia: 61,
1882 (Spruce 1882). 40
** Schiffnerioideae R.M.Schust.
** Schineria Steph., Österr. Bot. Z. 44 (1): 1, 1894 (Stephani 1894c).
*** Schineria hyalina Steph., Österr. Bot. Z. 44 (1): 1, 1894 (Stephani 1894c).
** Trabacelluloideae R.M.Schust.
** Haesselia Grolle et Gradst., J. Hattori Bot. Lab. 64: 327, 1988 (Grolle and Grad-
stein 1988).
*** Haesselia acuminata Gradst., Trop. Bryol. 1: 30, 1989 (Gradstein and Florschütz-
de Waard 1989).
*** Haesselia roraimensis Grolle et Gradst., J. Hattori Bot. Lab. 64: 327, 1988 (Grolle
and Gradstein 1988).
** Trabacellula Fulford, Acta Bot. Venez. 2 (5/8): 86, 1967 (Fulford 1967).
*** Trabacellula tumidula Fulford, Acta Bot. Venez. 2 (5/8): 86, 1967 (Fulford 1967).
*** Cephaloziellaceae Douin
by J. Váňa
e treatment of Cephaloziellaceae follows what was outlined in Váňa et al. (2013e).
Some nomenclatural and taxonomic notes can also be found in Söderström et al. (2013a)
and Váňa et al. (2013b, 2013l). e placement of Phycolepidozia follows Gradstein et al.
(2014a).
40 Odontoschisma obcordatum is to be excluded from Odontoschisma but its identity remains unclear
as the type is not found. Based on the original description this species might belong to Alobiellopsis
(Gradstein & Ilkiu-Borges 2014: 81).
World checklist of hornworts and liverworts 65
*** Allisoniella E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 3 (4): 80, 1965
(Hodgson 1965).
** sect. Allisoniella
*** Allisoniella nigra (Rodway) R.M.Schust., Nova Hedwigia 22: 137, 1971 [1972]
(Schuster 1971b). Bas.: Sphenolobus niger Rodway, Tasm. Bryoph.: 33, 1917
(Rodway 1917b).
** Allisoniella nigra var. acutiloba J.J.Engel, Novon 17 (3): 313, 2007 (Engel 2007).
** Allisoniella nigra subsp. novaezelandiae R.M.Schust., Nova Hedwigia 22: 143,
1971 [1972] (Schuster 1971b).
*** Allisoniella recurva R.M.Schust., Nova Hedwigia 22: 146, 1971 [1972] (Schuster
1971b).
*** Allisoniella subbipartita (C.Massal.) R.M.Schust. et J.J.Engel, Nova Hedwigia
22: 147, 1971 [1972] (Schuster 1971b). Bas.: Cephalozia subbipartita C.Massal.,
Nuovo Giorn. Bot. Ital. 17 (3): 235, 1885 (Massalongo 1885).
*** Allisoniella tasmanica R.M.Schust., Nova Hedwigia 22: 145, 1971 [1972] (Schuster
1971b).
** sect. Protomarsupella (R.M.Schust.) R.M.Schust., Nova Hedwigia 22: 150,
1971 (Schuster 1971b). Bas.: Protomarsupella R.M.Schust., Rev. Bryol. Lichénol.
34 (1/2): 264, 1966 (Schuster 1966b).
*** Allisoniella scottii (R.M.Schust.) R.M.Schust., Nova Hedwigia 22: 151, 1971
[1972] (Schuster 1971b). Bas.: Protomarsupella scottii R.M.Schust., Rev. Bryol.
Lichénol. 34 (1/2): 267, 1966 (Schuster 1966b).
*** Amphicephalozia R.M.Schust., Nova Hedwigia 22: 131, 1971 [1972] (Schuster
1971b).
*** Amphicephalozia africana Váňa et M.Wigginton, J. Bryol. 30 (1): 55, 2008 (Váňa
and Wigginton 2008).
*** Amphicephalozia amplexicaulis R.M.Schust., Nova Hedwigia 22: 133, 1971 [1972]
(Schuster 1971b).
*** Amphicephalozia geisslerae Pócs et Váňa, Polish Bot. J. 46 (2): 145, 2001 (Pócs and
Váňa 2001).
*** Anastrophyllopsis (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 81 (1): 15,
2013 (Váňa et al. 2013k). Bas.: Anastrophyllum sect. Anastrophyllopsis R.M.Schust.,
Beih. Nova Hedwigia 119: 310, 2002 (Schuster 2002b).
*** Anastrophyllopsis involutifolia (Mont. ex Gottsche, Lindenb. et Nees) Váňa et L.Söderstr.,
Phytotaxa 81 (1): 15, 2013 (Váňa et al. 2013k). Bas.: Jungermannia involutifolia Mont.
ex Gottsche, Lindenb. et Nees, Syn. Hepat. 1: 81, 1844 (Gottsche et al. 1844).
*** Anastrophyllopsis revoluta (Steph.) Váňa et L.Söderstr., Phytotaxa 81 (1): 16, 2013
(Váňa et al. 2013k). Bas.: Anastrophyllum revolutum Steph., Hedwigia 32 (3): 139,
1893 (Stephani 1893b).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
66
*** Anastrophyllopsis subcomplicata (Lehm. et Lindenb.) Váňa et L.Söderstr., Phyto-
taxa 81 (1): 16, 2013 (Váňa et al. 2013k). Bas.: Jungermannia subcomplicata Lehm.
et Lindenb., Nov. Stirp. Pug. 7: 4, 1838 (Lehmann 1838).
** Cephalojonesia Grolle, Rev. Bryol. Lichénol. 37 (4): 763, 1970 [1971] (Grolle
and Vanden Berghen 1970).
*** Cephalojonesia incuba Grolle et Vanden Berghen, Rev. Bryol. Lichénol. 37 (4):
764, 1970 [1971] (Grolle and Vanden Berghen 1970).
*** Cephalojonesia incuba subsp. mexicana Burghardt, Gradst. et Váňa, J. Hattori Bot.
Lab. 100: 35, 2006 (Burghardt et al. 2006).
** Cephalomitrion R.M.Schust., Nova Hedwigia 61 (3/4): 550, 1995 (Schuster 1995b).
*** Cephalomitrion aterrimum (Steph.) R.M.Schust., Nova Hedwigia 61 (3/4): 554,
1995 (Schuster 1995b). Bas.: Cephalozia aterrima Steph., Bull. Herb. Boissier (sér.
2) 8 (7): 501 (331), 1908 (Stephani 1908f).
*** Cephaloziella (Spruce) Schin., Hepat. (Engl.-Prantl): 98, 1893 (Schiner
1893b) nom. conserv. Bas.: Cephalozia subg. Cephaloziella Spruce, Cephalozia:
62, 1882 (Spruce 1882). 41
** subg. Cephaloziella
*** Cephaloziella aenigmatica R.M.Schust., Nova Hedwigia 63 (1/2): 20, 1996 (Schuster
1996d).
*** Cephaloziella anthelioides S.W.Arnell, Bot. Not. 105: 322, 1952 (Arnell 1952a).
*** Cephaloziella arctogena (R.M.Schust.) Konstant., Arctoa 3: 126, 1994 (Konstan-
tinova and Vasil’ev 1994). Bas.: Cephaloziella rubella var. arctogena R.M.Schust.,
Hepat. Anthocerotae N. Amer. 4: 125, 1980 (Schuster 1980c).
** Cephaloziella arenaria (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 280, 1963
(Schuster 1963b). Bas.: Cephalozia arenaria Steph., Bull. Herb. Boissier (sér. 2) 8
(7): 504 (334), 1908 (Stephani 1908f).
*** Cephaloziella aspericaulis Jørg., Bergens Mus. Skr. (n.ser.) 16: 197, 1934 (Jørgensen
1934).
*** Cephaloziella baumgartneri Schin., Verh. K.K. Zool.-Bot. Ges. Wien 56 (3): 273,
1906 (Schiner 1906c).
** Cephaloziella breviperianthia C.Gao, Fl. Hepat. Chin. Boreali-Orient.: 208, 1981
(Gao and Chang 1981).
** Cephaloziella brinkmannii Douin, Mém. Soc. Bot. France 29: 75, 1920 (Douin 1920).
*** Cephaloziella capensis (Sim) S.W.Arnell, Bot. Not. 105: 326, 1952 (Arnell 1952a). Bas.:
Cephalozia capensis Sim, Trans. Roy. Soc. South Africa 15 (1): 87, 1926 (Sim 1926).
41 Cephaloziella is a large genus and the subgeneric division is unclear. e traditional subdivisions used
here do not necessary agree with recent phylogenetic studies.
World checklist of hornworts and liverworts 67
*** Cephaloziella capillaris (Steph.) Douin, Mém. Soc. Bot. France 29: 59, 1920
(Douin 1920). Bas.: Cephalozia capillaris Steph., Bull. Herb. Boissier (sér. 2) 8
(7): 496 (326), 1908 (Stephani 1908f).
*** Cephaloziella crassigyna (R.M.Schust.) R.M.Schust., Nova Hedwigia 61 (3/4): 556,
1995 (Schuster 1995b). Bas.: Cephaloziella aterrima var. crassigyna R.M.Schust.,
Nova Hedwigia 22: 211, 1971 [1972] (Schuster 1971b).
*** Cephaloziella crispata N.Kitag., J. Hattori Bot. Lab. 32: 301, 1969 (Kitagawa 1969c).
*** Cephaloziella densifolia R.M.Schust., Nova Hedwigia 22: 199, 1971 [1972]
(Schuster 1971b).
** Cephaloziella densifolia var. dubia R.M.Schust., Nova Hedwigia 22: 202, 1971
[1972] (Schuster 1971b).
*** Cephaloziella divaricata (Sm.) Schin., Hepat. (Engl.-Prantl): 99, 1893 (Schiner
1893b). Bas.: Jungermannia divaricata Sm., Engl. Bot. 10: tab. 719, 1800 (Smith
and Sowerby 1800).
** Cephaloziella divaricata var. scabra (M.Howe) Haynes, Bryologist 12 (4): 68, 1909
(Haynes 1909). Bas.: Cephalozia divaricata var. scabra M.Howe, Mem. Torrey
Bot. Club 7: 129, 1899 (Howe 1899). 42
** Cephaloziella dusenii Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 (III,
6): 49, 1900 (Stephani 1900b).
*** Cephaloziella elachista (J.B.Jack ex Gottsche et Rabenh.) Schin., Sitzungsber.
deutsch. naturwiss.-med. Vereins Böhmen “Lotos” Prag 48: 336, 1900 (Schi-
ner 1900d). Bas.: Jungermannia elachista J.B.Jack ex Gottsche et Rabenh., Hepat.
Eur., Leberm. 58-59: no. 574, 1873 (Gottsche and Rabenhorst 1873a).
** Cephaloziella elachista var. spinophylla (C.Gao) C.Gao, Fl. Bryoph. Sin. 9: 181,
2003 (Gao 2003). Bas.: Cephaloziella spinophylla C.Gao, Fl. Hepat. Chin. Boreali-
Orient.: 208, 1981 (Gao and Chang 1981).
** Cephaloziella elegans (Heeg) Schin., Sitzungsber. deutsch. naturwiss.-med. Vereins
Böhmen “Lotos” Prag 48: 336, 1900 (Schiner 1900d). Bas.: Cephalozia ele gans
Heeg, Rev. Bryol. 20 (5): 82, 1893 (Heeg 1893).
*** Cephaloziella exigua R.M.Schust., Nova Hedwigia 63 (1/2): 49, 1996 (Schuster 1996d).
*** Cephaloziella exiliora (Taylor) Douin, Mém. Soc. Bot. France 29: 72, 1920
(Douin 1920). Bas.: Jungermannia exiliora Taylor, London J. Bot. 5: 279, 1846
(Taylor 1846a).
** Cephaloziella fragillima (Spruce) Fulford, Mem. New York Bot. Gard. 11 (4): 409,
1976 (Fulford 1976). Bas.: Cephalozia fragillima Spruce, Mem. Torrey Bot. Club
1 (3): 131, 1890 (Spruce 1890).
*** Cephaloziella garsidei S.W.Arnell, Rev. Bryol. Lichénol. 23 (1/2): 173, 1954 (Arnell
1954a).
*** Cephaloziella grandiretis (R.M.Schust.) R.M.Schust., Nova Hedwigia 63 (1/2): 35,
1996 (Schuster 1996d). Bas.: Cephaloziella byssacea subsp. grandiretis R.M.Schust.,
Nova Hedwigia 22: 195, 1971 [1972] (Schuster 1971b).
42 e correct name for the variety was discussed by Söderström et al. (2012d).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
68
*** Cephaloziella grimsulana (J.B.Jack ex Gottsche et Rabenh.) Lacout., Hépat. France: 52,
1905 (Lacouture 1905). Bas.: Jungermannia grimsulana J.B.Jack ex Gottsche et Ra-
benh., Hepat. Eur., Leberm. 53–55: no. 526, 1872 (Gottsche and Rabenhorst 1872).
*** Cephaloziella hampeana (Nees) Schin. ex Loeske, Moos. Harz.: 92, 1903 (Loes-
ke 1903). Bas.: Jungermannia hampeana Nees, Naturgesch. Eur. Leberm. 3: 560,
1838 (Nees 1838b).
** Cephaloziella hebridensis Steph., Hedwigia 32 (5): 316, 1893 (Stephani 1893d).
*** Cephaloziella herzogiana (Pandé et K.P.Srivast.) Udar et D.Kumar, Geophytology
6 (1): 45, 1976 (Udar and Kumar 1976). Bas.: Cephalozia herzogiana Pandé et
K.P.Srivast., Feddes Repert. Spec. Nov. Regni Veg. 58: 75, 1955 (Pandé and Sri-
vastava 1955).
** Cephaloziella heteroica (C.M.Cooke) Douin, Mém. Soc. Bot. France 29: 76, 1920
(Douin 1920). Bas.: Cephalozia heteroica C.M.Cooke, Trans. Connecticut Acad.
Arts 12 (1): 38, 1904 (Cooke 1904).
** Cephaloziella hyalina Douin, Mém. Soc. Bot. France 29: 77, 1920 (Douin 1920).
* Cephaloziella hyalina var. rappii (Douin) R.M.Schust., Hepat. Anthocerotae N.
Amer. 4: 56, 1980 (Schuster 1980c). Bas.: Cephaloziella rappii Douin, Mém. Soc.
Bot. France 29: 77, 1920 (Douin 1920).
*** Cephaloziella inaequalis R.M.Schust., Nova Hedwigia 22: 186, 1971 [1972]
(Schuster 1971b).
*** Cephaloziella invisa R.M.Schust., Nova Hedwigia 63 (1/2): 30, 1996 (Schuster
1996d).
** Cephaloziella kilohanensis (C.M.Cooke) Douin, Mém. Soc. Bot. France 29: 85,
1920 (Douin 1920). Bas.: Cephalozia kilohanensis C.M.Cooke, Trans. Connecti-
cut Acad. Arts 12 (1): 37, 1904 (Cooke 1904).
*** Cephaloziella longii Váňa, Folia Geobot. Phytotax. 27 (2): 193, 1992 (Váňa 1992).
*** Cephaloziella lycopodioides (Sim) S.W.Arnell, Bot. Not. 105: 321, 1952 (Arnell
1952a). Bas.: Cephalozia lycopodioides Sim, Trans. Roy. Soc. South Africa 15 (1):
85, 1926 (Sim 1926).
** Cephaloziella mammillifera R.M.Schust. et Damsh., Phytologia 63 (5): 327, 1987
(Schuster and Damsholt 1987).
*** Cephaloziella massalongi (Spruce) Müll.Frib., Lebermoose 2 (17): 191, 1913 (Müller
1913a). Bas.: Cephalozia massalongi Spruce, Cephalozia: 71, 1882 (Spruce 1882).
*** Cephaloziella muelleriana R.M.Schust., Nova Hedwigia 63 (1/2): 24, 1996 (Schuster
1996d).
*** Cephaloziella natalensis (Sim) S.W.Arnell, Hepat. South Africa: 338, 1963 (Arnell
1963b). Bas.: Cephalozia natalensis Sim, Trans. Roy. Soc. South Africa 15 (1): 86,
1926 (Sim 1926).
** Cephaloziella nicholsonii Douin, Rev. Bryol. 40 (6): 81, 1913 (Douin 1913b).
*** Cephaloziella nothogena R.M.Schust., Nova Hedwigia 63 (1/2): 37, 1996 (Schuster
1996d).
** Cephaloziella obtusilobula R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 108,
1980 (Schuster 1980c).
World checklist of hornworts and liverworts 69
** Cephaloziella patulifolia (Steph.) Douin, Mém. Soc. Bot. France 29: 70, 1920
(Douin 1920). Bas.: Cephalozia patulifolia Steph., Bull. Herb. Boissier (sér. 2) 8
(7): 509 (339), 1908 (Stephani 1908f).
*** Cephaloziella phyllacantha (C.Massal. et Carestia) Müll.Frib., Lebermoose 2 (17):
194, 1913 (Müller 1913a). Bas.: Anthelia phyllacantha C.Massal. et Carestia, Nuo-
vo Giorn. Bot. Ital. 12 (4): 340, 1880 (Massalongo and Carestia 1880).
*** Cephaloziella polystratosa (R.M.Schust. et Damsh.) Konstant., Bot. Zhurn. (Mos-
cow & Leningrad) 85 (10): 127, 2000 (Konstantinova 2000). Bas.: Cephaloziella
byssacea var. polystratosa R.M.Schust. et Damsh., Phytologia 63 (5): 327, 1987
(Schuster and Damsholt 1987).
*** Cephaloziella pseudocrassigyna R.M.Schust. ex J.J.Engel, Novon 17 (3): 313, 2007
(Engel 2007).
*** Cephaloziella pulcherrima R.M.Schust., Nova Hedwigia 22: 203, 1971 [1972]
(Schuster 1971b).
** Cephaloziella pulcherrima subsp. sphagnicola R.M.Schust., Nova Hedwigia 22:
205, 1971 [1972] (Schuster 1971b).
** Cephaloziella pungens Steph. ex Fulford, Mem. New York Bot. Gard. 11 (4): 410,
1976 (Fulford 1976).
*** Cephaloziella rubella (Nees) Warnst., Krypt.-Fl. Brandenburg, Leber- & Torf-
moose: 231, 1902 (Warnstorf 1902). Bas.: Jungermannia rubella Nees, Natur-
gesch. Eur. Leberm. 2: 236, 1836 (Nees 1836).
*** Cephaloziella schelpei S.W.Arnell, Bot. Not. 110 (1): 18, 1957 (Arnell 1957a).
*** Cephaloziella spegazziniana (C.Massal.) Douin, Mém. Soc. Bot. France 29: 69,
1920 (Douin 1920). Bas.: Cephalozia spegazziniana C.Massal., Nuovo Giorn. Bot.
Ital. 17 (3): 234, 1885 (Massalongo 1885).
*** Cephaloziella spinicaulis Douin, Mém. Soc. Bot. France 29: 62, 1920 (Douin 1920).
*** Cephaloziella spinigera (Lindb.) Jørg., Bergens Mus. Skr. (n.ser.) 16: 189, 1934
(Jørgensen 1934). Bas.: Cephalozia spinigera Lindb., Musci Scand.: 4, 1879 (Lind-
berg 1879).
*** Cephaloziella stellulifera (Taylor ex Carrington et Pearson) Croz., Rev. Bryol. 30 (2):
31, 1903 (Crozals 1903a). Bas.: Jungermannia stellulifera Taylor ex Carrington et
Pearson, Hepat. Brit. Exsicc. Fasc. I: no. 32, 1878 (Carrington and Pearson 1878).
*** Cephaloziella stephanii Schin. ex Douin, Mém. Soc. Bot. France 29: 85, 1920
(Douin 1920).
** Cephaloziella stolonifera R.M.Schust., Phytologia 39 (6): 426, 1978 (Schuster 1978b).
** Cephaloziella subtilis (Lindenb. et Gottsche) Steph., Hedwigia 32 (5): 318, 1893
(Stephani 1893d). Bas.: Jungermannia subtilis Lindenb. et Gottsche, Syn. Hepat.
5: 685, 1847 (Gottsche et al. 1847).
*** Cephaloziella sumatrana Schin. ex Douin, Mém. Soc. Bot. France 29: 59, 1920
(Douin 1920).
*** Cephaloziella tabularis S.W.Arnell, Bot. Not. 105: 318, 1952 (Arnell 1952a).
*** Cephaloziella transvaalensis S.W.Arnell, Bot. Not. 110 (1): 19, 1957 (Arnell 1957a).
*** Cephaloziella triplicata S.W.Arnell, Bot. Not. 115: 203, 1962 (Arnell 1962a).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
70
*** Cephaloziella umtaliensis S.W.Arnell, Bot. Not. 110 (1): 20, 1957 (Arnell 1957a).
*** Cephaloziella uncinata R.M.Schust., Meddel. Grønland 199 (1): 316, 1974
(Schuster and Damsholt 1974).
** Cephaloziella uncinata var. brevigyna R.M.Schust. et Damsh., Phytologia 63 (5):
327, 1987 (Schuster and Damsholt 1987).
** Cephaloziella uncinata var. sphagnicola R.M.Schust., Meddel. Grønland 199 (1):
323, 1974 (Schuster and Damsholt 1974).
*** Cephaloziella vaginans Steph., Wiss. Ergebn. Deut. Zentr.-Afr. Exped. (1907-08),
Bot. 2: 119, 1911 (Stephani 1911a).
*** Cephaloziella varians (Gottsche) Steph., Wiss. Ergebn. Schwed. Südpolar-Exped.
[1901–1903] 4 (1): 4, 1905 (Stephani 1905e). Bas.: Jungermannia varians
Gottsche, Int. Polarforsch., Deutsch. Exped. 2: 452, 1890 (Gottsche 1890).
*** Cephaloziella verrucosa Steph., Hedwigia 32 (5): 318, 1893 (Stephani 1893d).
* Cephaloziella villaumei (Steph.) Váňa, Phytotaxa 112 (1): 3, 2013 (Váňa et al.
2013e). Bas.: Cephalozia villaumei Steph., Sp. Hepat. (Stephani) 6: 437, 1924
(Stephani 1924). 43
** Cephaloziella violacea Schljakov, Novosti Sist. Nizš. Rast. 15: 242, 1978 (Shliakov 1978).
** Cephaloziella welwitschii (Steph.) Douin, Mém. Soc. Bot. France 29: 58, 1920
(Douin 1920). Bas.: Cephalozia welwitschii Steph., Bull. Herb. Boissier (sér. 2) 8
(6): 432 (308), 1908 (Stephani 1908g).
** subg. Dichiton (Mont.) Müll.Frib., Lebermoose 2 (27): 787, 1916 (Müller
1916). Bas.: Dichiton Mont., Syll. Gen. Sp. Crypt.: 52, 1856 (Montagne 1856b).
*** Cephaloziella calyculata (Durieu et Mont.) Müll.Frib., Lebermoose 2 (27): 787,
1916 (Müller 1916). Bas.: Jungermannia calyculata Durieu et Mont., Ann. Sci.
Nat. Bot. (sér. 3) 11: 34, 1849 (Montagne 1849).
*** Cephaloziella integerrima (Lindb.) Warnst., Krypt.-Fl. Brandenburg, Leber- &
Torfmoose: 232, 1902 (Warnstorf 1902). Bas.: Cephalozia integerrima Lindb.,
Acta Soc. Sci. Fenn. 10: 502, 1875 (Lindberg 1875).
** subg. Evansia (Douin et Schin.) Müll.Frib., Lebermoose 2 (27): 787, 1916
(Müller 1916). Bas.: Evansia Douin et Schin., Rev. Bryol. 40 (5): 66, 1913
(Douin 1913a).
*** Cephaloziella antillana (Besch. et Spruce) Fulford, Mem. New York Bot. Gard. 11
(4): 405, 1976 (Fulford 1976). Bas.: Cephalozia antillana Besch. et Spruce, Bull. Soc.
Bot. France (Congr. Bot.) 36: clxxxiii, 1889 [1890] (Bescherelle and Spruce 1889).
*** Cephaloziella dentata (Raddi) Steph., Bull. Herb. Boissier 5 (2): 78, 1897 (Stephani
1897b). Bas.: Jungermannia dentata Raddi, Jungermanniogr. Etrusca: 21, 1818
(Raddi 1818a).
*** Cephaloziella hirta (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 280, 1963
(Schuster 1963b). Bas.: Cephalozia hirta Steph., Bull. Herb. Boissier (sér. 2) 8 (8):
561 (345), 1908 (Stephani 1908e).
43 Cephaloziella villaumei is possibly conspecic with Cephaloziella anthelioides.
World checklist of hornworts and liverworts 71
** Cephaloziella squarrosula (Trevis.) Steph., Hedwigia 32 (5): 318, 1893 (Stephani
1893d). Bas.: Cephalozia squarrosula Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 417, 1877 (Trevisan 1877).
*** Cephaloziella subspinosa R.M.Schust., Nova Hedwigia 22: 210, 1971 [1972]
(Schuster 1971b).
** subg. Prionolobus (Spruce) Müll.Frib., Lebermoose 2 (16): 110, 1912 (Müller
1912). Bas.: Cephalozia subg. Prionolobus Spruce, Trans. & Proc. Bot. Soc. Edin-
burgh 15: 508, 1885 (Spruce 1885).
*** Cephaloziella acanthophora (S.Hatt.) Horik., Hikobia 1 (2): 79, 1951 (Horikawa
1951c). Bas.: Prionolobus acanthophorus S.Hatt., Bull. Tokyo Sci. Mus. 11: 29,
1944 (Hattori 1944d).
*** Cephaloziella biokoensis Váňa et Frank Müll., Trop. Bryol. 24: 1, 2003 (Váňa and
Müller 2003).
*** Cephaloziella granatensis (J.B.Jack ex Steph.) Fulford, Mem. New York Bot. Gard.
11 (4): 411, 1976 (Fulford 1976). Bas.: Cephalozia granatensis J.B.Jack ex Steph.,
Bull. Herb. Boissier (sér. 2) 8 (7): 500 (330), 1908 (Stephani 1908f).
*** Cephaloziella grisea R.M.Schust., Phytologia 39 (6): 425, 1978 (Schuster 1978b).
* Cephaloziella meghalayensis Udar et Ad.Kumar, Lindbergia 8 (1): 34, 1982 (Udar
and Kumar 1982d). 44
*** Cephaloziella microphylla (Steph.) Douin, Mém. Soc. Bot. France 29: 59, 1920
(Douin 1920). Bas.: Cephalozia microphylla Steph., Bull. Herb. Boissier (sér. 2) 8
(7): 513 (343), 1908 (Stephani 1908f).
*** Cephaloziella tenuissima (Lehm.) Steph., Hedwigia 32 (5): 318, 1893 (Stephani 1893d).
Bas.: Jungermannia tenuissima Lehm., Linnaea 4: 367, 1829 (Lehmann 1829).
*** Cephaloziella turneri (Hook.) Müll.Frib., Lebermoose 2 (17): 202, 1913 (Müller
1913a). Bas.: Jungermannia turneri Hook., Brit. Jungermann.: tab. 29, 1812
(Hooker 1812).
Incertae sedis
** Cephaloziella dentifolia Udar et Ad.Kumar, Lindbergia 8 (1): 30, 1982 (Udar and
Kumar 1982d).
** Cephaloziella lum (Trevis.) Steph., Hedwigia 32 (5): 318, 1893 (Stephani 1893d).
Bas.: Cephalozia lum Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 417, 1877 (Trevisan 1877).
* Cephaloziella exuosa C.Gao et K.C.Chang, Bull. Bot. Res., Harbin 4 (3): 88,
1984 (Chang and Gao 1984). 45
** Cephaloziella intricata Schin. ex Douin, Mém. Soc. Bot. France 29: 59, 1920
(Douin 1920).
44 Cephaloziella meghalayensis is probably conspecic with Cephaloziella acanthophora or Cephaloziella
microphylla.
45 Cephaloziella exuosa may be a form of Plicanthus birmensis or a juvenile form of Tetralophozia liformis.
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
72
** Cephaloziella levieri Schin. ex Douin, Mém. Soc. Bot. France 29: 80, 1920
(Douin 1920).
*** Cephaloziella pellucida R.M.Schust., Nova Hedwigia 63 (1/2): 57, 1996 (Schuster
1996d). 46
* Cephaloziella pygmaea (Spruce) Váňa, Phytotaxa 183 (4): 290, 2014 (Váňa et al.
2014a). Bas.: Cephalozia pygmaea Spruce, Cephalozia: 69, 1882 (Spruce 1882). 47
* Cephaloziella secundifolia Pearson, Ann. Bryol. 4: 106, 1931 (Pearson 1931b). 48
** Cephaloziella sinensis Douin, Rev. Bryol. 41 (1): 8, 1914 (Douin 1914).
** Cephaloziopsis (Spruce) Schin., Hepat. (Engl.-Prantl): 85, 1893 (Schiner
1893b). Bas.: Jungermannia sect. Cephaloziopsis Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 511, 1885 (Spruce 1885).
*** Cephaloziopsis intertexta (Gottsche) R.M.Schust., Nova Hedwigia 22: 183, 1971
[1972] (Schuster 1971b). Bas.: Jungermannia intertexta Gottsche, Syn. Hepat. 1:
107, 1844 (Gottsche et al. 1844).
** Chaetophyllopsis R.M.Schust., J. Hattori Bot. Lab. 23: 69, 1960 [1961] (Schuster
1960b).
*** Chaetophyllopsis whiteleggei (Carrington et Pearson) R.M.Schust. ex Hamlin, Rec.
Domin. Mus. 7: 255, 1972 (Hamlin 1972). Bas.: Jungermannia whiteleggei Car-
rington et Pearson, Proc. Linn. Soc. New South Wales (ser. 2) 2 (4): 1051, 1888
(Carrington and Pearson 1888a).
** Cylindrocolea R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 12 (3): 666,
1969 (Schuster 1969a).
** subg. Cylindrocolea
*** Cylindrocolea kiaeri (Austin) Váňa, Phytotaxa 112 (1): 2, 2013 (Váňa et al. 2013e). Bas.:
Jungermannia kiaeri Austin, Bull. Torrey Bot. Club 6 (3): 18, 1875 (Austin 1875b).
*** Cylindrocolea sanctae-helenae M.Wigginton, Polish Bot. J. 58 (1): 107, 2013 (Wig-
ginton 2013).
** sect. Cylindrocolea
*** Cylindrocolea andersonii R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 33, 1980
(Schuster 1980c).
*** Cylindrocolea brasiliensis D.P.Costa, N.D.Santos et Váňa, Bryologist 111 (4): 667,
2008 (Costa et al. 2008).
*** Cylindrocolea chevalieri (Steph.) R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.)
12 (3): 666, 1969 (Schuster 1969a). Bas.: Alobiella chevalieri Steph., Bull. Herb.
Boissier (sér. 2) 8 (8): 567 (351), 1908 (Stephani 1908e).
46 Cephaloziella pellucida is probably not a Cephaloziella, but the type specimen has not been found.
47 Cephaloziella pygmaea is possibly conspecic with Cephaloziella granatensis (Váňa et al. 2014a).
48 Cephaloziella secundifolia could not be studied by Fulford (1976) and she did not know what it is.
World checklist of hornworts and liverworts 73
*** Cylindrocolea gittinsii (E.W.Jones) R.M.Schust., Nova Hedwigia 22: 172, 1971
[1972] (Schuster 1971b). Bas.: Cephaloziella gittinsii E.W.Jones, Trans. Brit.
Bryol. Soc. 3 (3): 437, 1958 (Jones 1958).
*** Cylindrocolea madagascariensis (Steph.) R.M.Schust., Bull. Natl. Sci. Mus. To-
kyo (n.ser.) 12 (3): 666, 1969 (Schuster 1969a). Bas.: Lophozia madagascariensis
Steph., Sp. Hepat. (Stephani) 6: 112, 1917 (Stephani 1917a).
*** Cylindrocolea nigerica (E.W.Jones) R.M.Schust., Nova Hedwigia 22: 172, 1971
[1972] (Schuster 1971b). Bas.: Cephaloziella nigerica E.W.Jones, Trans. Brit. Bry-
ol. Soc. 3 (3): 435, 1958 (Jones 1958).
*** Cylindrocolea novae-caledoniae (Grolle) R.M.Schust., Nova Hedwigia 22: 161,
1971 [1972] (Schuster 1971b). Bas.: Cephaloziella novae-caledoniae Grolle, Rev.
Bryol. Lichénol. 29 (3/4): 208, 1960 [1961] (Grolle 1960a).
*** Cylindrocolea planifolia (Steph.) R.M.Schust., Nova Hedwigia 22: 164, 1971
[1972] (Schuster 1971b). Bas.: Cephaloziella planifolia Steph., Hedwigia 32 (5):
317, 1893 (Stephani 1893d).
*** Cylindrocolea sprucei R.M.Schust., Nova Hedwigia 22: 163, 1971 [1972] (Schuster
1971b).
*** Cylindrocolea ugandica (E.W.Jones) R.M.Schust., Nova Hedwigia 22: 171, 1971
[1972] (Schuster 1971b). Bas.: Cephaloziella ugandica E.W.Jones, Trans. Brit.
Bryol. Soc. 3 (3): 433, 1958 (Jones 1958).
** sect. Platycaulis R.M.Schust., Nova Hedwigia 22: 173, 1971 (Schuster
1971b).
*** Cylindrocolea recurvifolia (Steph.) Inoue, J. Jap. Bot. 47 (11): 348, 1972 (Inoue
1972a). Bas.: Cephalozia recurvifolia Steph., Bull. Herb. Boissier (sér. 2) 8 (7): 497
(327), 1908 (Stephani 1908f).
*** Cylindrocolea tagawae (N.Kitag.) R.M.Schust., Nova Hedwigia 22: 174, 1971
[1972] (Schuster 1971b). Bas.: Cephaloziella tagawae N.Kitag., J. Hattori Bot.
Lab. 32: 303, 1969 (Kitagawa 1969c).
** subg. Cylindroscyphus (Douin) R.M.Schust., Hepat. Anthocerotae N. Amer.
4: 20, 1980 (Schuster 1980c). Bas.: Cephaloziella subg. Cylindroscyphus Douin,
Mém. Soc. Bot. France 29: 56, 1920 (Douin 1920).
*** Cylindrocolea abyssinica (Gola) Váňa, Phytotaxa 112 (1): 2, 2013 (Váňa et al.
2013e). Bas.: Cephaloziella abyssinica Gola, Ann. Bot. (Rome) 13 (1): 68, 1914
(Gola 1914a).
** Cylindrocolea obtusifolia Fulford, Mem. New York Bot. Gard. 11 (4): 401, 1976
(Fulford 1976).
** Cylindrocolea reticulata Udar et Ad.Kumar, Lindbergia 8 (3): 181, 1982 [1983]
(Udar and Kumar 1982c).
*** Cylindrocolea rhizantha (Mont.) R.M.Schust., Nova Hedwigia 22: 175, 1971
[1972] (Schuster 1971b). Bas.: Jungermannia rhizantha Mont., Hist. Phys. Cuba,
Bot., Pl. Cell.: 454, 1842 (Montagne 1842a).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
74
*** Gottschelia Grolle, J. Hattori Bot. Lab. 31: 13, 1968 (Grolle 1968c).
*** Gottschelia maxima (Steph.) Grolle, J. Bryol. 25 (1): 6, 2003 (Grolle et al. 2003). Bas.:
Tylimanthus maximus Steph., Sp. Hepat. (Stephani) 6: 249, 1922 (Stephani 1922).
*** Gottschelia schizopleura (Spruce) Grolle, J. Hattori Bot. Lab. 31: 16, 1968 (Grolle
1968c). Bas.: Jungermannia schizopleura Spruce, Trans. & Proc. Bot. Soc. Edin-
burgh 15: 517, 1885 (Spruce 1885).
** Gymnocoleopsis (R.M.Schust.) R.M.Schust., Phytologia 39 (4): 243, 1978
(Schuster 1978a). Bas.: Gymnocolea subg. Gymnocoleopsis R.M.Schust., Bryologist
70 (1): 111, 1967 (Schuster 1967b).
*** Gymnocoleopsis capensis (S.W.Arnell) R.M.Schust., J. Hattori Bot. Lab. 78: 123,
1995 (Schuster 1995c). Bas.: Lophozia capensis S.W.Arnell, Svensk Bot. Tidskr. 47
(1): 112, 1953 (Arnell 1953a).
*** Gymnocoleopsis cylindriformis (Mitt.) R.M.Schust., J. Hattori Bot. Lab. 78: 126,
1995 (Schuster 1995c). Bas.: Jungermannia cylindriformis Mitt., J. Linn. Soc., Bot.
15 (84): 196, 1876 (Mitten 1876b).
*** Herzogobryum Grolle, Rev. Bryol. Lichénol. 32 (3/4): 160, 1963 [1964] (Grolle
1963d). Nom. nov. pro Chondrophyllum Herzog, Rev. Bryol. Lichénol. 21 (1/2):
46, 1952 (Herzog 1952f).
*** Herzogobryum atrocapillum (Hook.f. et Taylor) Grolle, Österr. Bot. Z. 113 (2):
228, 1966 (Grolle 1966j). Bas.: Gymnomitrion atrocapillum Hook.f. et Taylor,
London J. Bot. 5: 258, 1846 (Taylor 1846a).
*** Herzogobryum liforme R.M.Schust., Phytologia 45 (5): 422, 1980 (Schuster 1980b).
*** Herzogobryum molle Grolle, Österr. Bot. Z. 113 (2): 226, 1966 (Grolle 1966j).
*** Herzogobryum vermiculare (Schin.) Grolle, J. Hattori Bot. Lab. 28: 103, 1965
(Grolle 1965e). Bas.: Gymnomitrion vermiculare Schin., Leberm., Forschungsr.
Gazelle 4 (4): 2, 1890 (Schiner 1890).
** Kymatocalyx Herzog, Memoranda Soc. Fauna Fl. Fennica 25: 56, 1950 (Herzog
1950c).
*** Kymatocalyx africanus Váňa et M.Wigginton, Haussknechtia, Beih. 9: 158, 1999
(Gradstein and Váňa 1999).
*** Kymatocalyx dominicensis (Spruce) Váňa, Österr. Bot. Z. 118 (5): 575, 1970 (Váňa
1970b). Bas.: Jungermannia dominicensis Spruce, J. Linn. Soc., Bot. 30 (210): 363,
1895 (Gepp 1895b).
*** Kymatocalyx madagascariensis (Steph.) Gradst. et Váňa, Haussknechtia, Beih. 9:
164, 1999 (Gradstein and Váňa 1999). Bas.: Acrobolbus madagascariensis Steph.,
Bull. Herb. Boissier (sér. 2) 2 (5): 460 (179), 1902 (Stephani 1902f).
*** Kymatocalyx rhizomaticus (Herzog) Gradst. et Váňa, Haussknechtia, Beih. 9: 166,
1999 (Gradstein and Váňa 1999). Bas.: Stenorrhipis rhizomatica Herzog, Trans.
Brit. Bryol. Soc. 1 (4): 290, 1950 (Herzog 1950a).
World checklist of hornworts and liverworts 75
*** Lophonardia R.M.Schust., Phytologia 39 (4): 244, 1978 (Schuster 1978a).
*** Lophonardia jamesonii (Mont.) L.Söderstr. et Váňa, Phytotaxa 81 (1): 19, 2013
(Söderström et al. 2013a). Bas.: Jungermannia jamesonii Mont., Syll. Gen. Sp.
Crypt.: 60, 1856 (Montagne 1856b).
*** Lophonardia laxifolia (Mont.) L.Söderstr. et Váňa, Phytotaxa 81 (1): 20, 2013
(Söderström et al. 2013a). Bas.: Sarcocyphos laxifolius Mont., Ann. Sci. Nat. Bot.
(sér. 3) 4: 346, 1845 (Montagne 1845b).
*** Lophonardia tristaniana (S.W.Arnell) L.Söderstr. et Váňa, Phytotaxa 81 (1): 20,
2013 (Söderström et al. 2013a). Bas.: Jungermannia tristaniana S.W.Arnell, Re-
sults Norweg. Sci. Exped. Tristan da Cunha 42: 12, 1958 (Arnell 1958b).
*** Nothogymnomitrion R.M.Schust., J. Hattori Bot. Lab. 80: 43, 1996 (Schuster 1996a).
*** Nothogymnomitrion erosum (Carrington et Pearson) R.M.Schust., J. Hattori Bot.
Lab. 80: 44, 1996 (Schuster 1996a). Bas.: Cesius erosus Carrington et Pearson, Pap.
& Proc. Roy. Soc. Tasmania 1887: 8, 1888 (Carrington and Pearson 1888b).
*** Obtusifolium S.W.Arnell, Ill. Moss Fl. Fennosc. Hep.: 309, 1956 (Arnell 1956b).
*** Obtusifolium obtusum (Lindb.) S.W.Arnell, Ill. Moss Fl. Fennosc. Hep.: 133, 1956
(Arnell 1956b). Bas.: Jungermannia obtusa Lindb., Musci Scand.: 7, 1879 (Lind-
berg 1879).
*** Oleolophozia L.Söderstr., De Roo et Hedd., Phytotaxa 3: 50, 2010 (Söderström
et al. 2010b).
*** Oleolophozia perssonii (H.Buch et S.W.Arnell) L.Söderstr., De Roo et Hedd., Phy-
totaxa 3: 51, 2010 (Söderström et al. 2010b). Bas.: Lophozia perssonii H.Buch et
S.W.Arnell, Bot. Not. 97: 382, 1944 (Buch 1944).
** Phycolepidozia R.M.Schust., Bull. Torrey Bot. Club 93 (6): 438, 1966 [1967]
(Schuster 1966f).
** subg. Metaphycolepidozia Gradst., J.-P.Frahm et U.Schwarz, Taxon 63 (3):
506, 2014 (Gradstein et al. 2014a).
** Phycolepidozia indica Gradst., J.-P.Frahm et U.Schwarz, Taxon 63 (3): 499, 2014
(Gradstein et al. 2014a).
** subg. Phycolepidozia
*** Phycolepidozia exigua R.M.Schust., Bull. Torrey Bot. Club 93 (6): 440, 1966
[1967] (Schuster 1966f).
** Protolophozia (R.M.Schust.) Schljakov, Novosti Sist. Nizš. Rast. 16: 204, 1979
(Shliakov 1979). Bas.: Lophozia subg. Protolophozia R.M.Schust., Nova Hedwigia
15: 472, 1968 (Schuster 1968b).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
76
** Protolophozia androgyna R.M.Schust. ex Váňa et L.Söderstr., Phytotaxa 76 (3): 51,
2013 (Váňa et al. 2013l). Based on: Lophozia androgyna R.M.Schust., Beih. Nova
Hedwigia 119: 266, 2002 (Schuster 2002b), nom. inval.
** Protolophozia autoica (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76 (3): 51,
2013 (Váňa et al. 2013l). Bas.: Lophozia autoica R.M.Schust., Nova Hedwigia 15:
479, 1968 (Schuster 1968b).
** Protolophozia crispata (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76 (3): 51,
2013 (Váňa et al. 2013l). Bas.: Lophozia crispata R.M.Schust., Nova Hedwigia 15:
474, 1968 (Schuster 1968b).
*** Protolophozia druceae (Grolle et E.A.Hodgs.) Váňa et L.Söderstr., Phytotaxa 76
(3): 51, 2013 (Váňa et al. 2013l). Bas.: Lophozia druceae Grolle et E.A.Hodgs., J.
Roy. Soc. New Zealand 2 (1): 112, 1972 (Hodgson 1972).
*** Protolophozia elongata (Steph.) Schljakov, Novosti Sist. Nizš. Rast. 16: 204, 1979
(Shliakov 1979). Bas.: Lophozia elongata Steph., Bull. Herb. Boissier (sér. 2) 2 (1):
41 (141), 1902 (Stephani 1902c).
*** Protolophozia herzogiana (E.A.Hodgs. et Grolle) Váňa et L.Söderstr., Phytotaxa 76
(3): 51, 2013 (Váňa et al. 2013l). Bas.: Lophozia herzogiana E.A.Hodgs. et Grolle,
Rev. Bryol. Lichénol. 31 (3/4): 152, 1962 [1963] (Grolle 1962c).
*** Protolophozia lancistipa (Grolle) Váňa et L.Söderstr., Phytotaxa 76 (3): 51, 2013
(Váňa et al. 2013l). Bas.: Andrewsianthus lancistipus Grolle, Marion Prince Edw.
Is: 230, 1971 (Grolle 1971d).
*** Protolophozia leucorhiza (Mitt.) Váňa et L.Söderstr., Phytotaxa 76 (3): 51, 2013
(Váňa et al. 2013l). Bas.: Jungermannia leucorhiza Mitt., J. Linn. Soc., Bot. 15
(82): 68, 1876 (Mitten 1876a).
*** Protolophozia longiora (Herzog) L.Söderstr. et Váňa, Phytotaxa 76 (3): 51, 2013
(Váňa et al. 2013l). Bas.: Orthocaulis longiorus Herzog, Rev. Bryol. Lichénol. 23
(1/2): 32, 1954 (Herzog 1954).
** Protolophozia monoica (E.A.Hodgs.) Váňa et L.Söderstr., Phytotaxa 76 (3): 52,
2013 (Váňa et al. 2013l). Bas.: Metahygrobiella monoica E.A.Hodgs., Trans. Roy.
Soc. New Zealand, Bot. 3 (4): 76, 1965 (Hodgson 1965).
*** Protolophozia multicuspidata (Hook.f. et Taylor) Váňa et L.Söderstr., Phytotaxa
76 (3): 52, 2013 (Váňa et al. 2013l). Bas.: Jungermannia multicuspidata Hook.f. et
Taylor, London J. Bot. 3: 375, 1844 (Hooker and Taylor 1844a).
** Protolophozia nivicola (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76 (3): 52,
2013 (Váňa et al. 2013l). Bas.: Lophozia nivicola R.M.Schust., Nova Hedwigia 15:
477, 1968 (Schuster 1968b).
*** Protolophozia perssoniana (H.A.Mill.) Váňa et L.Söderstr., Phytotaxa 76 (3): 52,
2013 (Váňa et al. 2013l). Bas.: Lophozia perssoniana H.A.Mill., Ark. Bot. (n.ser.)
5 (2): 508, 1963 (Miller 1963).
** Protolophozia subalpina (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76 (3): 52,
2013 (Váňa et al. 2013l). Bas.: Lophozia autoica var. subalpina R.M.Schust., Nova
Hedwigia 15: 482, 1968 (Schuster 1968b).
World checklist of hornworts and liverworts 77
*** Protolophozia tasmanica (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76 (3): 52,
2013 (Váňa et al. 2013l). Bas.: Lophozia tasmanica R.M.Schust., Nova Hedwigia
15: 484, 1968 (Schuster 1968b).
*** Protolophozia verruculosa (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76 (3): 52,
2013 (Váňa et al. 2013l). Bas.: Lophozia verruculosa R.M.Schust., Phytologia 39
(4): 242, 1978 (Schuster 1978a).
*** Lophoziaceae Cavers
by L. Söderström and J. Váňa
Lophoziaceae was broadly dened by de Roo et al. (2007) excluding Anastrophyllaceae and
Jamesonielloideae and also moving several traditionally included genera to other families.
Vilnet et al. (2010) further rened the family. Nomenclatural and taxonomic notes can also
be found in Vilnet et al. (2007b, 2008), Söderström et al. (2013c) and Váňa et al. (2013m).
e placement of Gerhildiella and Pseudocephaloziella in the family is provisional.
*** Andrewsianthus R.M.Schust., Rev. Bryol. Lichénol. 30 (1/2): 66, 1961 (Schuster
1961a).
** subg. Andrewsianthus
*** Andrewsianthus aberrans (Nees et Mont.) Grolle, Trans. Brit. Bryol. Soc. 4 (3):
440, 1963 (Grolle 1963a). Bas.: Jungermannia aberrans Nees et Mont., Ann. Sci.
Nat. Bot. (sér. 2) 19: 250, 1843 (Montagne 1843).
*** Andrewsianthus bidens (Mitt. ex Steph.) R.M.Schust., Nova Hedwigia 15: 492,
1968 (Schuster 1968b). Bas.: Lophozia bidens Mitt. ex Steph., Bull. Herb. Boissier
(sér. 2) 2 (1): 41 (141), 1902 (Stephani 1902c).
*** Andrewsianthus bilobus (Mitt.) Grolle, Trans. Brit. Bryol. Soc. 4 (3): 437, 1963
(Grolle 1963a). Bas.: Gymnanthe biloba Mitt., J. Proc. Linn. Soc., Bot. 7 (27):
166, 1863 (Mitten 1863).
*** Andrewsianthus carinatus Grolle, Marion Prince Edw. Is: 229, 1971 (Grolle 1971d).
*** Andrewsianthus cavifolius Grolle et Váňa, J. Hattori Bot. Lab. 38: 640, 1974 (Váňa
1974b).
*** Andrewsianthus chimbuensis R.M.Schust., Nova Hedwigia 15: 491, 1968 (Schuster
1968b).
*** Andrewsianthus kinabaluensis N.Kitag., J. Hattori Bot. Lab. 33: 205, 1970 (Kitagawa
1970).
*** Andrewsianthus koponenii Váňa et Piippo, Ann. Bot. Fenn. 26 (3): 284, 1989
(Váňa and Piippo 1989).
*** Andrewsianthus mizutanii N.Kitag., J. Hattori Bot. Lab. 32: 307, 1969 (Kitagawa
1969a).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
78
*** Andrewsianthus papillosus N.Kitag., J. Hattori Bot. Lab. 33: 207, 1970 (Kitagawa
1970).
*** Andrewsianthus perigonialis (Hook.f. et Taylor) R.M.Schust., Beih. Nova Hedwi-
gia 119: 336, 2002 (Schuster 2002b). Bas.: Jungermannia perigonialis Hook.f. et
Taylor, London J. Bot. 3: 368, 1844 (Hooker and Taylor 1844a).
*** Andrewsianthus puniceus (Nees) R.M.Schust. ex Grolle, Trans. Brit. Bryol. Soc.
4 (3): 439, 1963 (Grolle 1963a). Bas.: Jungermannia punicea Nees, Enum. Pl.
Crypt. Javae: 32, 1830 (Nees 1830).
*** Andrewsianthus recurvifolius (Nees) R.M.Schust., Hepat. Anthocerotae N. Amer.
2: 710, 1969 (Schuster 1969b). Bas.: Jungermannia recurvifolia Nees, Enum. Pl.
Crypt. Javae: 32, 1830 (Nees 1830).
*** Andrewsianthus sundaicus (Schin.) R.M.Schust., Hepat. Anthocerotae N. Amer.
2: 710, 1969 (Schuster 1969b). Bas.: Anastrophyllum sundaicum Schin., Denk-
schr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 67: 202, 1898 (Schiner 1898a).
*** Andrewsianthus zantenii Váňa, J. Hattori Bot. Lab. 38: 645, 1974 (Váňa 1974b).
** subg. Cephalolobus (R.M.Schust.) R.M.Schust., Beih. Nova Hedwigia 119:
326, 2002 (Schuster 2002b). Bas.: Cephalolobus R.M.Schust., Rev. Bryol. Liché-
nol. 34 (1/2): 244, 1966 (Schuster 1966b).
*** Andrewsianthus hodgsoniae (R.M.Schust.) R.M.Schust. ex J.J.Engel, Novon 17 (3):
311, 2007 (Engel 2007). Bas.: Cephalolobus hodgsoniae R.M.Schust., Rev. Bryol.
Lichénol. 34 (1/2): 254, 1966 (Schuster 1966b).
*** Andrewsianthus marionensis (S.W.Arnell) Grolle, Marion Prince Edw. Is: 232,
1971 (Grolle 1971d). Bas.: Lophozia marionensis S.W.Arnell, Svensk Bot. Tidskr.
47 (3): 421, 1953 (Arnell 1953c).
*** Andrewsianthus scabrellus (C.Massal.) R.M.Schust. ex J.J.Engel, Novon 17 (3):
311, 2007 (Engel 2007). Bas.: Cephalozia scabrella C.Massal., Nuovo Giorn. Bot.
Ital. 17 (3): 233, 1885 (Massalongo 1885).
** Andrewsianthus sphenoloboides (R.M.Schust.) R.M.Schust. ex J.J.Engel, Novon 17
(3): 311, 2007 (Engel 2007). Bas.: Cephalolobus sphenoloboides R.M.Schust., Rev.
Bryol. Lichénol. 34 (1/2): 251, 1966 (Schuster 1966b).
** Gerhildiella Grolle, Rev. Bryol. Lichénol. 34 (1/2): 187, 1966 (Grolle 1966a).
*** Gerhildiella rossneriana Grolle, Rev. Bryol. Lichénol. 34 (1/2): 187, 1966 (Grolle
1966a).
*** Heterogemma (Jørg.) Konstant. et Vilnet, Arctoa 18: 67, 2009 [2010] (Konstan-
tinova and Vilnet 2009). Bas.: Lophozia sect. Heterogemma Jørg., Bergens Mus.
Skr. (n.ser.) 16: 146, 1934 (Jørgensen 1934).
*** Heterogemma capitata (Hook.) Konstant. et Vilnet, Arctoa 18: 67, 2009 [2010]
(Konstantinova and Vilnet 2009). Bas.: Jungermannia capitata Hook., Brit. Jung-
ermann.: tab. 80, 1816 (Hooker 1816a).
World checklist of hornworts and liverworts 79
*** Heterogemma laxa (Lindb.) Konstant. et Vilnet, Arctoa 18: 67, 2009 [2010]
(Kons tantinova and Vilnet 2009). Bas.: Jungermannia laxa Lindb., Acta Soc. Sci.
Fenn. 10: 529, 1875 (Lindberg 1875).
*** Heterogemma patagonica (Herzog et Grolle) L.Söderstr. et Váňa, Phytotaxa 97 (2):
29, 2013 (Söderström et al. 2013c). Bas.: Lophozia patagonica Herzog et Grolle,
Rev. Bryol. Lichénol. 28 (3/4): 343, 1959 [1960] (Grolle 1959a).
*** Lophozia (Dumort.) Dumort., Recueil Observ. Jungerm.: 17, 1835 (Dumortier
1835). Bas.: Jungermannia sect. Lophozia Dumort., Syll. Jungerm. Europ.: 53,
1831 (Dumortier 1831). 49
*** Lophozia ascendens (Warnst.) R.M.Schust., Bryologist 55 (3): 180, 1952 (Schuster
1952). Bas.: Sphenolobus ascendens Warnst., Hedwigia 57 (1/2): 63, 1916 (Warns-
torf 1916).
* Lophozia austrosibirica Bakalin, Ann. Bot. Fenn. 40 (1): 49, 2003 (Bakalin 2003).
** Lophozia ciliata Damsh., L.Söderstr. et H.Weibull, Lindbergia 25 (1): 3, 2000
(Söderström et al. 2000).
*** Lophozia guttulata (Lindb. et Arnell) A.Evans, Proc. Wash. Acad. Sci. 2: 302, 1900
(Evans 1900c). Bas.: Jungermannia guttulata Lindb. et Arnell, Kongl. Svenska Ve-
tensk.-Akad. Handl. (n.ser.) 23 (5): 51, 1889 (Lindberg and Arnell 1889).
* Lophozia jamaicensis (Nees) Steph., Bull. Herb. Boissier (sér. 2) 1 (2): 128 (155),
1901 (Stephani 1901h). Bas.: Jungermannia jamaicensis Nees, Syn. Hepat. 1: 105,
1844 (Gottsche et al. 1844). 50
** Lophozia lacerata N.Kitag., Hikobia 3 (3): 172, 1963 (Kitagawa 1963a).
** Lophozia lantratoviae Bakalin, Ann. Bot. Fenn. 40 (1): 47, 2003 (Bakalin 2003).
** Lophozia murmanica Kaal., Rep. Second Norweg. Arctic Exped. 11: 34, 1906
(Bryhn 1906).
** Lophozia pacica Bakalin, Bryologist 114 (2): 302, 2011 (Bakalin 2011).
* Lophozia pallida (Steph.) Grolle, J. Jap. Bot. 39 (6): 174, 1964 (Grolle 1964a).
Bas.: Anastrophyllum pallidum Steph., Bull. Herb. Boissier (sér. 2) 1 (10): 1131
(114), 1901 (Stephani 1901c). 51
** Lophozia savicziae Schljakov, Novosti Sist. Nizš. Rast. 10: 299, 1973 (Shliakov 1973).
* Lophozia schusterana Schljakov, Novosti Sist. Nizš. Rast. 12: 320, 1975 (Shliakov 1975).
*** Lophozia silvicola H.Buch, Beret. 18 Skand. Naturforskarmøde: 228, 1929 (Buch
1929).
*** Lophozia silvicoloides N.Kitag., J. Hattori Bot. Lab. 28: 276, 1965 (Kitagawa 1965).
49 Lophozia has many nomenclatural and taxonomic problems. is treatment follows Söderström et al.
(2013c).
50 Lophozia jamaicensis is probably not a Lophozia, but the type specimen is missing in Nees’s herbarium
(Söderström et al. 2010a).
51 Lophozia pallida may be conspecic with Lophozia guttulata (Bakalin 2003).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
80
* Lophozia subapiculata R.M.Schust. et Damsh., Meddel. Grønland 199 (1): 104,
1974 (Schuster and Damsholt 1974). 52
** Lophozia udarii S.Srivast., S.C.Srivast. et K.K.Rawat, Nelumbo 55: 130, 2013
(Srivastava et al. 2013). 53
*** Lophozia ventricosa (Dicks.) Dumort., Recueil Observ. Jungerm.: 17, 1835 (Du-
mortier 1835). Bas.: Jungermannia ventricosa Dicks., Fasc. Pl. Crypt. Brit. 2: 14,
1790 (Dickson 1790). 54
* Lophozia wenzelii (Nees) Steph., Bull. Herb. Boissier (sér. 2) 2 (1): 35 (135), 1902
(Stephani 1902c). Bas.: Jungermannia wenzelii Nees, Naturgesch. Eur. Leberm. 2:
58, 1836 (Nees 1836). 55
Excluded from the genus
* Lophozia serpens (Dumort.) Dumort., Recueil Observ. Jungerm.: 17, 1835 (Du-
mortier 1835). Bas.: Jungermannia serpens Dumort., Syll. Jungerm. Europ.: 56,
1831 (Dumortier 1831). 56
** Lophoziopsis Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010] (Konstantinova
and Vilnet 2009).
*** Lophoziopsis excisa (Dicks.) Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010] (Kons-
tantinova and Vilnet 2009). Bas.: Jungermannia excisa Dicks., Fasc. Pl. Crypt.
Brit. 3: 11, 1793 (Dickson 1793).
** Lophoziopsis excisa var. elegans (R.M.Schust.) Konstant. et Vilnet, Arctoa 18: 66,
2009 [2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia excisa var. elegans
R.M.Schust., Hepat. Anthocerotae N. Amer. 2: 522, 1969 (Schuster 1969b).
** Lophoziopsis excisa var. infuscata (R.M.Schust. et Damsh.) Konstant. et Vilnet,
Arctoa 18: 66, 2009 [2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia ex-
cisa var. infuscata R.M.Schust. et Damsh., Meddel. Grønland 199 (1): 94, 1974
(Schuster and Damsholt 1974).
** Lophoziopsis excisa var. succulenta (R.M.Schust. et Damsh.) Konstant. et Vilnet, Arc-
toa 18: 66, 2009 [2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia excisa var.
52 Lophozia subapiculata may be a form of Lophozia guttulata, but Bakalin (2005) treated it as conspe-
cic with Lophozia ventricosa.
53 Lophozia udarii has recently been described using the generic concept of Lophozia in the wide sense
(as e.g. Schuster 1969b). We do not know where to refer the species. It is possibly a Schistochilopsis
species close to Schistochilopsis incisa.
54 Lophozia ventricosa is very variable and many segregates at various subspecic levels have been de-
scribed. e species complex also includes at least Lophozia austrosibirica, Lophozia schusteriana and
Lophozia wenzelii.
55 Lophozia wenzelii is closely related to Lophozia ventricosa according to De Roo et al. (2007) and
Vilnet et al. (2010) based on molecular and morphological evidence. It is variable and many varieties
are described. e variability and relationships to related species needs further study.
56 Lophozia serpens is probably conspecic with Schistochilopsis incisa.
World checklist of hornworts and liverworts 81
succulenta R.M.Schust. et Damsh., Meddel. Grønland 199 (1): 96, 1974 (Schuster
and Damsholt 1974).
*** Lophoziopsis longidens (Lindb.) Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010]
(Konstantinova and Vilnet 2009). Bas.: Jungermannia longidens Lindb., Helsingf.
Dagbl. 1876 (323, 26 Nov.): 2, 1876 (Lindberg 1876a).
** Lophoziopsis longidens subsp. arctica (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa
97 (2): 28, 2013 (Söderström et al. 2013c). Bas.: Lophozia longidens subsp. arctica
R.M.Schust., Hepat. Anthocerotae N. Amer. 2: 539, 1969 (Schuster 1969b). 57
*** Lophoziopsis pellucida (R.M.Schust.) Konstant. et Vilnet, Arctoa 18: 67, 2009
[2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia pellucida R.M.Schust.,
Canad. J. Bot. 39 (4): 978, 1961 (Schuster 1961b).
** Lophoziopsis pellucida var. minor (R.M.Schust.) L.Söderstr. et Váňa, Phytotaxa
97 (2): 28, 2013 (Söderström et al. 2013c). Bas.: Lophozia pellucida var. minor
R.M.Schust., Canad. J. Bot. 39 (4): 984, 1961 (Schuster 1961b).
*** Lophoziopsis polaris (R.M.Schust.) Konstant. et Vilnet, Arctoa 18: 67, 2009
[2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia alpestris subsp. polaris
R.M.Schust., Hepat. Anthocerotae N. Amer. 2: 614, 1969 (Schuster 1969b).
** Lophoziopsis polaris var. sphagnorum (R.M.Schust.) Konstant. et Vilnet, Arctoa
18: 67, 2009 [2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia alpestris f.
sphagnorum R.M.Schust., Hepat. Anthocerotae N. Amer. 2: 619, 1969 (Schuster
1969b).
* Lophoziopsis propagulifera (Gottsche) Konstant. et Vilnet, Arctoa 18: 67, 2009
[2010] (Konstantinova and Vilnet 2009). Bas.: Jungermannia propagulifera
Gottsche, Int. Polarforsch., Deutsch. Exped. 2: 451, 1890 (Gottsche 1890). 58
** Lophoziopsis rubrigemma (R.M.Schust.) Konstant. et Vilnet, Arctoa 18: 67, 2009
[2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia rubrigemma R.M.Schust.,
Hepat. Anthocerotae N. Amer. 2: 621, 1969 (Schuster 1969b).
** Pseudocephaloziella R.M.Schust., Phytologia 39 (4): 243, 1978 (Schuster 1978a).
*** Pseudocephaloziella epiphytica R.M.Schust., Phytologia 39 (4): 243, 1978 (Schuster
1978a).
*** Trilophozia (R.M.Schust.) Bakalin, Monogr. Lophozia: 34, 2005 (Bakalin
2005). Bas.: Tritomaria subg. Trilophozia R.M.Schust., Amer. Midl. Naturalist 49
(2): 382, 1953 (Schuster 1953).
*** Trilophozia quinquedentata (Huds.) Bakalin, Monogr. Lophozia: 34, 2005 (Baka-
lin 2005). Bas.: Jungermannia quinquedentata Huds., Fl. Angl. (Hudson): 433,
1762 (Hudson 1762).
57 Lophoziopsis longidens var. arctica was treated as a form of Lophoziopsis pellucida by Bakalin (2005).
58 Lophoziopsis propagulifera belongs to the Lophoziopsis excisa species complex. It is probably only a
modication (cf. discussion by Váňa and Engel 2013).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
82
** Trilophozia quinquedentata var. asymmetrica (Horik.) L.Söderstr. et Váňa, Phytotaxa
97 (2): 32, 2013 (Söderström et al. 2013c). Bas.: Lophozia asymmetrica Horik., J. Sci.
Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 153, 1934 (Horikawa 1934).
*** Tritomaria Schin. ex Loeske, Hedwigia 49 (1/2): 13, 1909 (Loeske 1909).
Based on: Tritomaria Schin., Ber. Naturwiss.-Med. Vereins Innsbruck 31
[Beilage]: 12, 1908 (Schiner 1908a).
*** Tritomaria exsecta (Schmidel) Schin. ex Loeske, Hedwigia 49 (1/2): 13, 1909
(Loeske 1909). Bas.: Jungermannia exsecta Schmidel, Syst. Samml. Crypt. Gew. 2:
5, 1797 (Schrader 1797), nom. conserv.
** Tritomaria exsecta subsp. novaezelandiae J.J.Engel, Bryologist 109 (1): 61, 2006
(Engel 2006b).
*** Tritomaria exsectiformis (Breidl.) Schin. ex Loeske, Hedwigia 49 (1/2): 13, 1909
(Loeske 1909). Bas.: Jungermannia exsectiformis Breidl., Mitt. Naturwiss. Vereins
Steiermark 30: 321, 1894 (Breidler 1894).
** Tritomaria exsectiformis subsp. arctica R.M.Schust., Hepat. Anthocerotae N.
Amer. 2: 661, 1969 (Schuster 1969b).
** Tritomaria exsectiformis subsp. camerunensis S.W.Arnell ex Váňa, Phytotaxa 167
(2): 216, 2014 (Váňa et al. 2014b). Based on: Tritomaria camerunensis S.W.Arnell,
Svensk Bot. Tidskr. 52 (1): 64, 1958 (Arnell 1958a), nom. inval.
*** Tritomaria ferruginea (Grolle) Váňa, Phytotaxa 81 (1): 24, 2013 (Váňa et al.
2013m). Bas.: Andrewsianthus ferrugineus Grolle, Khumbu Himal 1 (4): 275,
1966 (Grolle 1966k).
* Tritomaria koreana Bakalin, S.S.Choi et B.Y.Sun, Arctoa 18: 163, 2009 [2010]
(Bakalin et al. 2009a).
* Tritomaria mexicana Bakalin, Arctoa 17: 162, 2008 [2009] (Bakalin 2008b).
*** Tritomaria scitula (Taylor) Jørg., Bergens Mus. Aarbok 1919/20 (7): 3, 1922 (Jør-
gensen 1922). Bas.: Jungermannia scitula Taylor, London J. Bot. 5: 274, 1846
(Taylor 1846a).
*** Scapaniaceae Mig.
by J. Váňa
Scapaniaceae, in its classical concept, was recognized as monophyletic and nested
within Lophoziaceae s. lat. (cf. Davis 2004, Schill et al. 2004, Yatsentyuk et al. 2004,
He-Nygrén et al. 2006). More recent molecular studies classied Scapaniaceae as a
sister clade to Lophoziaceae, and the genus Schistochilopsis, a segregate of the genus
Lophozia s. lat., more closely related to Scapaniaceae than to Lophoziaceae (cf. de Roo
et al. 2007, Vilnet et al. 2007). e genus Scapania was recently studied by morpho-
logical methods by Potemkin (2002), by molecular methods by Vilnet et al. (2007)
and especially Heinrichs et al. (2012a). Some taxonomic and nomenclatural notes
World checklist of hornworts and liverworts 83
can be found in Váňa et al. (2012a, 2013j, 2015) and Konstantinova et al. (2013a).
e placement of Pseudotritomaria, Saccobasis and Schistochilopsis in the family follows
Vilnet et al. (2010).
*** Diplophyllum (Dumort.) Dumort., Recueil Observ. Jungerm.: 15, 1835 (Du-
mortier 1835) nom. conserv. Bas.: Jungermannia sect. Diplophyllum Dumort.,
Syll. Jungerm. Europ.: 44, 1831 (Dumortier 1831).
*** subg. Austrodiplophyllum R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 11
(1): 18, 1968 (Schuster 1968a).
** Diplophyllum recurvifolium C.Massal., Atti Reale Ist. Veneto Sci. Lett. Arti 87 (2):
221, 1928 (Massalongo 1928).
*** Diplophyllum squarrosum Steph., Sp. Hepat. (Stephani) 4: 116, 1910 (Stephani
1910b).
*** Diplophyllum verrucosum R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 11 (1):
19, 1968 (Schuster 1968a).
*** subg. Diplophyllum
*** sect. Diplophyllum
*** Diplophyllum albicans (L.) Dumort., Recueil Observ. Jungerm.: 16, 1835 (Dumortier
1835). Bas.: Jungermannia albicans L., Sp. Pl. 1: 1133, 1753 (Linnaeus 1753).
*** sect. Protodiplophyllum (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76
(3): 29, 2013 (Váňa et al. 2013j). Bas.: Diplophyllum subg. Protodiplophyllum
R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 192, 1974 (Schuster 1974).
*** Diplophyllum africanum S.W.Arnell, Ark. Bot. (n.ser.) 3 (16): 527, 1956 (Arnell
1956e).
*** Diplophyllum andicola R.M.Schust., Phytologia 39 (4): 248, 1978 (Schuster
1978a).
*** Diplophyllum andrewsii A.Evans, Bryologist 25 (2): 28, 1922 (Evans 1922).
* Diplophyllum androgynum J.J.Engel et G.L.Merr., J. Hattori Bot. Lab. 84: 277,
1998 (Engel and Smith Merrill 1998).
* Diplophyllum angustifolium J.J.Engel et G.L.Merr., J. Hattori Bot. Lab. 84: 262,
1998 (Engel and Smith Merrill 1998).
*** Diplophyllum apiculatum (A.Evans) Steph., Sp. Hepat. (Stephani) 4: 110, 1910
(Stephani 1910b). Bas.: Diplophylleia apiculata A.Evans, Bot. Gaz. 34 (5): 372,
1902 (Evans 1902b).
** Diplophyllum dioicum R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 11 (1):
20, 1968 (Schuster 1968a).
** Diplophyllum exiguum Steph., Sp. Hepat. (Stephani) 6: 500, 1924 (Stephani 1924).
* Diplophyllum gemmiparum J.J.Engel et G.L.Merr., J. Hattori Bot. Lab. 84: 255,
1998 (Engel and Smith Merrill 1998).
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
84
* Diplophyllum incrassatum J.J.Engel et G.L.Merr., J. Hattori Bot. Lab. 84: 265,
1998 (Engel and Smith Merrill 1998).
*** Diplophyllum nanum Herzog, Memoranda Soc. Fauna Fl. Fennica 26: 48, 1950
[1951] (Herzog 1950b).
* Diplophyllum novum J.J.Engel et G.L.Merr., J. Hattori Bot. Lab. 84: 274, 1998
(Engel and Smith Merrill 1998).
*** Diplophyllum obtusatum (R.M.Schust.) R.M.Schust., Hepat. Anthocerotae N.
Amer. 3: 215, 1974 (Schuster 1974). Bas.: Diplophyllum apiculatum var. obtusa-
tum R.M.Schust., Amer. Midl. Naturalist 49 (2): 432, 1953 (Schuster 1953).
*** Diplophyllum obtusifolium (Hook.) Dumort., Recueil Observ. Jungerm.: 16, 1835
(Dumortier 1835). Bas.: Jungermannia obtusifolia Hook., Brit. Jungermann.: tab.
26, 1812 (Hooker 1812). 59
** Diplophyllum obtusifolium subsp. domesticum (Gottsche) Váňa, Phytotaxa 76 (3):
29, 2013 (Váňa et al. 2013j). Bas.: Jungermannia domestica Gottsche, Linnaea 28
(5): 548, 1856 [1857] (Gottsche 1856).
*** Diplophyllum serrulatum (Müll.Frib.) Steph., Sp. Hepat. (Stephani) 4: 112, 1910
(Stephani 1910b). Bas.: Diplophylleia serrulata Müll.Frib., Bull. Herb. Boissier
(sér. 2) 3 (1): 34, 1903 (Müller 1903).
*** Diplophyllum taxifolium (Wahlenb.) Dumort., Recueil Observ. Jungerm.: 16,
1835 (Dumortier 1835). Bas.: Jungermannia taxifolia Wahlenb., Fl. Lapp. (Wah-
lenberg): 389, 1812 (Wahlenberg 1812).
** Diplophyllum taxifolium var. mucronatum R.M.Schust., Hepat. Anthocerotae N.
Amer. 3: 203, 1974 (Schuster 1974).
*** Diplophyllum trollii Grolle, Khumbu Himal 1 (4): 273, 1966 (Grolle 1966k).
*** Douinia (C.E.O.Jensen) H.Buch, Scapan. N.-Eur. Sib.: 13, 1928 (Buch 1928).
Bas.: Diplophylleia subg. Douinia C.E.O.Jensen, Danmarks mosser: 145, 1915
(Jensen 1915).
*** Douinia imbricata (M.Howe) Konstant. et Vilnet, Phytotaxa 76 (3): 31, 2013
(Konstantinova et al. 2013a). Bas.: Scapania imbricata M.Howe, Bull. New York
Bot. Gard. 2 (6): 104, 1901 (Howe 1901b).
*** Douinia ovata (Dicks.) H.Buch, Scapan. N.-Eur. Sib.: 14, 1928 (Buch 1928). Bas.:
Jungermannia ovata Dicks., Fasc. Pl. Crypt. Brit. 3: 11, 1793 (Dickson 1793).
*** Douinia plicata (Lindb.) Konstant. et Vilnet, Phytotaxa 76 (3): 31, 2013 (Kons-
tantinova et al. 2013a). Bas.: Diplophyllum plicatum Lindb., Acta Soc. Sci. Fenn.
10: 235, 1872 [1873] (Lindberg 1872b).
** Pseudotritomaria Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010] (Konstanti-
nova and Vilnet 2009).
59 Diplophyllum obtusifolium is a species complex also including Diplophyllum androgynum, Diplophyllum
angustifolium, Diplophyllum gemmiparum, Diplophyllum incrassatum and Diplophyllum novum.
World checklist of hornworts and liverworts 85
*** Pseudotritomaria heterophylla (R.M.Schust.) Konstant. et Vilnet, Arctoa 18: 66,
2009 [2010] (Konstantinova and Vilnet 2009). Bas.: Tritomaria heterophylla
R.M.Schust., Canad. J. Bot. 36 (2): 272, 1958 (Schuster 1958b).
** Saccobasis H.Buch, Memoranda Soc. Fauna Fl. Fennica 8: 291, 1932 [1933]
(Buch 1932).
*** Saccobasis polita (Nees) H.Buch, Memoranda Soc. Fauna Fl. Fennica 8: 292, 1932
[1933] (Buch 1932). Bas.: Jungermannia polita Nees, Naturgesch. Eur. Leberm. 2:
145, 1836 (Nees 1836).
** Saccobasis polymorpha (R.M.Schust.) Schljakov, Novosti Sist. Nizš. Rast. 16: 205,
1979 (Shliakov 1979). Bas.: Tritomaria polita subsp. polymorpha R.M.Schust.,
Hepat. Anthocerotae N. Amer. 2: 700, 1969 (Schuster 1969b).
*** Scapania (Dumort.) Dumort., Recueil Observ. Jungerm.: 14, 1835 (Dumortier
1835) nom. conserv. Bas.: Radula sect. Scapania Dumort., Syll. Jungerm. Europ.:
38, 1831 (Dumortier 1831).
** subg. Ascapania Grolle, Khumbu Himal 1 (4): 268, 1966 (Grolle 1966k).
*** Scapania contorta Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 101, 1860 [1861] (Mitten
1860c).
** subg. Gracilidae (H.Buch) Váňa, Hentschel, Joch.Müll. et Heinrichs, Phyto-
Keys 10: 15, 2012 (Váňa et al. 2012a). Bas.: Scapania sect. Gracilidae H.Buch,
Scapan. N.-Eur. Sib.: 106, 1928 (Buch 1928).
*** Scapania ampliata Steph., Bull. Herb. Boissier 5 (2): 106, 1897 (Stephani 1897b).
** Scapania ampliata subsp. queenslandica M.L.Hicks, J. Hattori Bot. Lab. 69: 130,
1991 (Hicks 1991).
*** Scapania bolanderi Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 218, 1869 (Austin
1869).
*** Scapania gracilis Lindb., Morgonbladet (Helsinki) 1873 (286, 9 Dec): 2, 1873
(Lindberg 1873a).
** Scapania macroparaphyllia T.Cao, C.Gao et J.Sun, Acta Phytotax. Sin. 42 (2):
180, 2004 (Cao et al. 2004).
*** Scapania maxima Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 223,
1934 (Horikawa 1934).
*** Scapania nipponica (Amakawa et S.Hatt.) Amakawa, J. Hattori Bot. Lab. 30:
319, 1967 (Amakawa 1967a). Bas.: Scapania bolanderi var. nipponica Amakawa et
S.Hatt., J. Hattori Bot. Lab. 14: 83, 1955 (Amakawa and Hattori 195