Third time lucky? Another substantially revised sectional classification for
Australasian Plagiochila (Plagiochilaceae: Jungermanniopsida)
Author(s): Matt A. M. Renner, Simon D. F. Patzak, Margaret M. Heslewood, Alfons SchäferVerwimp and Jochen Heinrichs
Source: Australian Systematic Botany, 30(1):70-104.
Published By: CSIRO Publishing
URL: http://www.bioone.org/doi/full/10.1071/SB16038
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CSIRO PUBLISHING
Australian Systematic Botany, 2017, 30, 70–104
http://dx.doi.org/10.1071/SB16038
Third time lucky? Another substantially revised sectional
classification for Australasian Plagiochila (Plagiochilaceae:
Jungermanniopsida)
Matt A. M. Renner A,D, Simon D. F. Patzak B, Margaret M. Heslewood A,
Alfons Schäfer-Verwimp C and Jochen Heinrichs B
A
Royal Botanic Gardens and Domain Trust, Mrs Macquaries Road, Sydney, NSW 2000, Australia.
Ludwig Maximilian University, Faculty of Biology, Department of Biology and Geobio-Center,
Menzinger Straße 67, D-80638 Munich, Germany.
C
Mittlere Letten 11, D-88634 Herdwangen-Schönach, Germany.
D
Corresponding author. Email: matt.renner@rbgsyd.nsw.gov.au
B
Abstract. Molecular phylogeny reconstruction has motivated recircumscription of all families and most genera within
the Lophocoleinae, and in Plagiochila, the largest genus of this lineage, has refined the sectional classification as well. Here,
we extend this ongoing revision in the first study focusing on species from Australasia, a region to date underrepresented
in molecular phylogenetic datasets. We reconstructed a phylogeny containing more than 300 individuals from Australasia
and the Pacific, and with this tested sectional circumscriptions within the two largely contradictory classifications recently
proposed for Plagiochila. Neither scheme satisfactorily captures relationships among species and all sections prove
paraphyletic or polyphyletic, with the exception of those defying these properties by virtue of containing only one
species. We propose expanding the circumscription of several sections as the best option for achieving a revised
classification representing monophyla that remains stable over the short to medium term, given current knowledge.
Broader circumscriptions are proposed for section Denticulatae with section Tayloriae as a new synonym; section
Arrectae with section Caducifoliae as a new synonym; a reinstated section Deflexifoliae; and section Plagiochila, to
include P. trapezoidea; section Belangerianae to include sections Annotinae, Mitteniae and Strombifoliae as new
synonyms; and section Durae with section Colensoae as a new synonym. Section Fragmentissimae is applied to the
lineage previously named section Deltoideae nom. inval. or section Hodgsoniae nom. inval., as the Tasmanian
P. ratkowskiana and New Zealand P. fragmentissima are the same, and sister to other species in the lineage containing
P. deltoidea. Morphological characters supporting these groups are identified, but more importantly the proposed
revisions provide a robust framework on which informed re-examination of morphology within this variable and
species-rich genus can proceed, and we introduce some encouraging avenues in this area.
Received 19 September 2016, accepted 8 February 2017, published online 31 May 2017
Introduction
Molecular data have enforced whole-scale revision of the
hypotheses of relationship formalised within the Linnean
hierarchy in the diverse flowering plant lineage (Bremer et al.
2009). Although not as diverse, the upheavals wrought by
molecular data to the classification of the non-vascular plant
lineage Marchantiophyta are at least as drastic. The simple
thalloid Metzgeriales (Schuster 1984) was actually two lineages
(He-Nygrén et al. 2006), one of which, the Metzgeriidae, was
sister to the leafy Pleurozia Dumort. (Davis 2004; He-Nygrén
et al. 2004). The putative ancestor of the complex thalloids,
Monocleales (Schuster 1984), was actually highly derived
(Forrest and Crandall-Stotler 2004; Heinrichs et al. 2005a;
Crandall-Stotler et al. 2009), and the Sphaerocarpales was nested
within the Marchantiales (Forrest et al. 2006). The isolated
Journal compilation CSIRO 2017
groups Calobryales and Treubiales (Schuster 1984) were sister
lineages (Forrest and Crandall-Stotler 2004), although this
sister relationship is rather ancient (Cooper et al. 2012).
Substantial re-arrangement of classifications in all major lineages
has occurred, because molecular data repeatedly identified
malleability in morphological character systems believed to be
rigid, and so enforced whole-scale rearrangement of the
classifications founded on those characters, from species to class
(Heinrichs et al. 2005a; Renzaglia et al. 2007; Söderström et al.
2016). In retrospect, morphological characters supporting many
of these relationships (e.g. Schuster 1966) were under-valued.
Although the framework of relationships within the
Jungermanniopsida is now fairly well resolved, the
recircumscription of suborders and families is ongoing (Shaw
et al. 2015; Patzak et al. 2016a). Recent changes at familial
www.publish.csiro.au/journals/asb
Testing relationships within Plagiochila
level include the synonymisation of the small Chonecoleaceae
R.M.Schust. ex Grolle with Cephaloziaceae Mig. (Patzak et al.
2016b) and the reinstatement of the Lophocoleaceae Vanden
Berghen, which was recovered from synonymy with the
Geocalycaceae H.Klinggr. (Hentschel et al. 2006), and the
newly proposed Solenostomataceae Stotler & Crand.-Stotl.
(Crandall-Stotler et al. 2009).
The Lophocoleaceae was resolved in a monophylum with
the families Brevianthaceae J.J.Engel & R.M.Schust. and
Plagiochilaceae Müll. Frib. Whereas the Brevianthaceae is
likely to have fewer than 10 species confined to Melanesia and
Australasia, the family Plagiochilaceae presents an example of
high species diversity, wide geographic distribution, extensive
morphological variation and broad ecological amplitude
(Heinrichs et al. 2004a, 2004b, 2005b; Rycroft et al. 2004;
Patzak et al. 2016c). Currently, the Plagiochilaceae contains
some 450–700 species, most of which belong to Plagiochila
(Dumort.) Dumort. (Heinrichs 2002; Patzak et al. 2016c;
Söderström et al. 2016), meaning that this genus contains
some 5–9% of global liverwort species diversity (von Konrat
et al. 2010; Söderström et al. 2016). The Plagiochilaceae is
also one of the largest families in the liverwort lineage, being
exceeded only by the Lejeuneaceae Cavers (~1500 species;
Söderström et al. 2016) and possibly the Frullaniaceae Lorch.
Like most other liverwort families, composition and
circumscription of the Plagiochilaceae has been refined by
molecular data. The broad circumscription of Schuster (1980)
included genera now distributed among two suborders, and the
composition of the family continues to change, primarily by
the transfer of morphologically isolated elements from other
families within the Lophocoleineae (Söderström et al. 2013)
and the occasional exclusion of morphologically anomalous
species to other suborders (Patzak et al. 2016a). Within the
Plagiochilaceae, as in other families, the ongoing flux is due
to the capture of smaller genera of difficult acquisition in
molecular studies of ever increasing scope and depth of
sampling. Genera have been transferred between families
within a suborder, for example, Tetracymbaliella Grolle from
Lophocoleaceae to Brevianthaceae and Pseudolophocolea R.M.
Schust. & J.J.Engel from Lophocoleaceae to Plagiochilaceae
(Lophocoleineae, Söderström et al. 2013), Saccogynidium
Grolle from Geocalycaceae to Acrobolbaceae E.A.Hodgs.
(Jungermanniineae, Shaw et al. 2015), Rivulariella D.H.Wagner
from Jungermanniaceae Rchb. to Scapaniaceae Mig. (Patzak
et al. 2016b), and sometimes between different suborders, for
example, Pedinophyllopsis R.M.Schust. & Inoue from
Jungermanniineae to Lophocoleineae (He-Nygrén and Piippo
2003), Syzygiella Spruce and Xenochila R.M.Schust. in the
reverse direction (Groth and Heinrichs 2005; Patzak et al.
2016a) and Chonecolea R.M.Schust. ex Grolle from
Lophocoleaceae, Harpanthaceae Arnell or Plagiochilaceae to
Cephaloziellaceae Douin (Patzak et al. 2016b). Often these
transfers involve morphologically and phylogenetically distinct
elements whose unusual morphology has precipitated
misplacement, and the transferred genera are usually retained.
Unsurprisingly, genera have proven paraphyletic or
polyphyletic within familial limits too. A topical example from
the Lophocoleaceae is Chiloscyphus Corda, which proved
paraphyletic with respect of Leptoscyphus Mitt. (Hentschel
Australian Systematic Botany
71
et al. 2007), prompting segregation of Cryptolophocolea L.
Söderstr., Crand.-Stotl., Stotler & Vá
na (Söderström et al.
2013); however, the exact limits of Chiloscyphus have still
not been resolved (He-Nygrén and Piippo 2003; Patzak et al.
2016c). The large, morphologically distinct genus Plagiochila
was paraphyletic before the segregates Dinckleria Trevis.,
Chiastocaulon Carl and Cryptoplagiochila S.D.F.Patzak,
M.A.M.Renner & Heinrichs were excluded (Groth and
Heinrichs 2003; Heinrichs et al. 2006; Patzak et al. 2016c).
Unfortunately, Chiastocaulon and Acrochila R.M.Schust.
rendered the well known and distinctive (Inoue 1964) genus
Plagiochilion S.Hatt. paraphyletic, and Acrochila, with only two
species, was itself resolved polyphyletic within Chiastocaulon
(Patzak et al. 2016c), so even this minimalist attempt at
monophyly on the basis of morphological evidence failed.
Repeatedly, hypotheses of relationship founded on
homoplastic character systems have been flatly refuted by
molecular data (e.g. Yu et al. 2013; Heinrichs et al. 2014;
Bechteler et al. 2016). Every refutation has reinforced
appreciation of the innovative morphological changes achieved
to meet ecological challenges (Feldberg et al. 2014), and
highlighted the role of natural selection in shaping the observed
distribution of character states among groups of organisms.
In turn, this corroborates to some extent earlier hypotheses of
evolutionary dynamism within this lineage of relatively
uncomplicated, structurally simple, plants (e.g. Schuster 2001).
Well resolved, well sampled, and robust classifications are
integral to formulating and refining hypotheses explaining all
aspects of evolutionary dynamics, including character evolution,
divergence and diversification rates. These explanations are
particularly pertinent in lineages that have experienced higher
diversification or lower extinction rates resulting in high extant
diversity in comparison to sister lineages, because these systems
may provide insight into the interaction between trait origin and
phylogenetic diversity within a range of ecological contexts.
Within the Plagiochilaceae, changes in generic (Groth and
Heinrichs 2003; Heinrichs et al. 2006; Patzak et al. 2016c)
and sectional (Heinrichs et al. 2005c; Engel and Merrill 2013;
Söderström et al. 2015) classifications have been motivated by
a range of factors, including nomenclatural issues, changing
concepts, and shifting emphasis on different character systems.
Concomitant with ongoing refinement to generic limits has
been progressive, and sometimes dramatic modification of
infrageneric groups. The first infrageneric classifications for
Plagiochila were proposed by Lindenberg (1843, see
Söderström et al. 2015, Plagiochilaceae and World Checklist,
cited as 1844), including six sections with 11 infrasectional
groups, all circumscribed by shoot architecture, leaf shape,
arrangement and dentition, and perianth shape in Gottsche
et al. (1845). These morphologically circumscribed sections
were also intercontinentally distributed.
In his treatment of Andean and Amazonian species, Spruce
(1885) was not particularly complementary regarding
Lindenberg’s sections, commenting that he found them
intractable in practice, and so proposed another classification
‘which I hope may be found easier to work with, and more
conformable to nature’ (Spruce 1885, p. 453). Spruce considered
shoot architecture, branching pattern, gynoecium position,
plant colour, perianth form, androecium position and even the
72
Australian Systematic Botany
number of antheridia per bract, and recognised five sections.
He commented that a sixth would be needed for ‘our
P. asplenioides and P. gigantea’ that were distinct from South
American species in their long tubular perianths with an
elongated pedicel. This relationship, on the basis of the same
characters, had earlier been postulated by Lindenberg who ‘soli
manerent tum Plagiochilae genus’ [alone endured the genus
Plagiochila] in Gottsche et al. (1845). Spruce divided
Plagiochila into two broad groups, the Cauliflorae Spruce and
Ramiflorae Spruce. The former was proposed for plants with
erect or cernuous stems arising from a prostrate caudex, and
shoots the branching of which was associated mostly with
gynoecium production, each bearing one or two innovations
that were simple or again fertile. The Ramiflorae contained
plants the shoot systems of which were dichotomous in full or
part, rarely one- to three-pinnate, the gynoecia terminal on
ultimate branches, rarely also in ultimate axils, so the plant
developed a flabelliform frond, surrounded by gynoecia.
Spruce’s (1885) broad groups have not been accepted, and his
five sectional names applied to groups for which names had been
proposed by Lindenberg.
Six sections were described by Schiffner (1900), four of
which are accepted today. A geographical section concept
was introduced by Carl (1931) who used characters of the
gametophyte available from herbarium material, especially leaf
shape, leaf cell pattern and branching to propose a classification
containing 52 new sections and one new subgenus. The
geographical–morphological sectional classification inherited
from Carl (1931) has been progressively modified by
subsequent workers who have considered characters of the
sporophyte in addition to those of the gametophyte (Inoue and
Schuster 1971; Schuster 1980; Inoue 1984; So 2001a,
2001b, 2001c; So and Grolle 2001; Heinrichs 2002; Engel and
Merrill 2013). These revisions have considered an everwidening array of data sources, including both gametophyte
and sporophyte generations. Inoue proposed eight new
sections (Inoue 1958, 1965, 1975, 1984), the new subgenera
Paraplagiochila Inoue and Metaplagiochila Inoue (Inoue 1984),
and the new genus Steereochila Inoue (Inoue 1987). With
Schuster, he also proposed three new sections and four new
subsections for New Zealand and Tasmanian species (Inoue
and Schuster 1971). Hässel de Menéndez (2009) described a
new section from southern South America, based on the unusual
female bracts and the lack of subfloral innovations, among
other morphological features. Engel and Smith-Merrill (2013),
likewise, considered evidence from the sterile and fertile
gametophyte and the sporophyte and described four new
sections from New Zealand.
The first study of Plagiochila including molecular data
appeared this millennium (Heinrichs et al. 2002), since
which time ongoing investigations have refined both generic
and sectional circumscriptions (e.g. Groth and Heinrichs 2003;
Heinrichs et al. 2005a, 2006). Molecular and morphological
evidence from gametophyte and sporophyte have supported
the re-circumscription of several sections, and the description
of the new Plagiochila section Africanae Heinrichs from Africa
(Heinrichs et al. 2005c). Diagnostic morphological characters
have been identified in lieu of molecular phylogenies, often
from poorly studied character systems such as oil bodies,
M. A. M. Renner et al.
sporophyte-valve cell thickenings, and elaters, for example,
the blueish-brown oil bodies of P. section Glaucescentes Carl,
and the unispiral elaters of P. section Arrectae Carl in
comparision to the bispiral elaters of P. section Peculiares
Schiffn. (Heinrichs et al. 2004c). New morphological characters,
such as the leaf surface wax in section Fuscoluteae Carl, have also
been identified (Heinrichs et al. 2000a).
Revised sectional circumscriptions have in most cases
proven robust to increased sampling intensity, as the
taxonomic coverage of molecular datasets has broadened and
deepened (Heinrichs 2002; Groth et al. 2003, 2004; Heinrichs
et al. 2004c, 2005d, 2006; Lindner et al. 2004). The most recent
publication on the infrageneric classification (Söderström
et al. 2015) was a review of the sectional classification of
Plagiochilaceae, which incorporated recent molecular evidence
the implications of which had not yet been fully translated into
the sectional classification.
Unfortunately, the progressive refinement of sectional
circumscription undertaken over the past 80 years has not
implied convergence on a single universally accepted
infrageneric classification. In fact, within some geographic
regions, quite the opposite is true. In New Zealand, 12 sections
were accepted for New Zealand by Engel and Merrill (2013), but,
of these, only seven were accepted by Söderström et al. (2015).
Although some of the differences between these two schemes
were a function of the incorporation of results from recent
molecular phylogenetic investigations in one and not the other,
this incompatibility of sectional concepts and classifications has
highlighted the fact that resolution of the relationships within
Plagiochila is far from complete (Heinrichs 2002; Groth et al.
2004; Patzak et al. 2016b), as does the recent re-instatement of P.
section Abietinae Schiffn. (Jamy et al. 2016).
The recent study of Patzak et al. (2016c) included 15 of an
estimated 71 Australasian species (Renner et al. 2017), and
did not include most Australasian sectional type species, so
was not able to test recent revisions of the infrageneric
classification. Other critical type species from Australasia have
been included on the basis of single accessions only. There
has not yet been a comprehensive test of the relationships
encapsulated by the alternative sectional classification for
Australasia proposed by Engel and Merrill (2013) and Söderström
et al. (2015).
In the present study, we expand species sampling within
Australasia to include 64 of the 71 species currently believed
to occur in this region (Renner et al. 2017), including all
sectional types. We employed likelihood and Bayesian
approaches to infer relationships among species. We aimed to
resolve robust relationships between Australian species as the
basis for refining sectional classifications proposed for species of
this region. Our key questions were how are Australasian species
related, both to each other and to species from other parts of
the world?
Materials and methods
Taxon sampling
Specimens from the herbaria AK, CANB, F, GOET, MEL and
NSW were sampled for DNA to capture the range of
morphological and geographic variation represented in
Testing relationships within Plagiochila
herbarium material. Specimens derived from dedicated
fieldwork in both Australia and New Zealand were included in
this set of sampled herbarium specimens. Voucher details are
given in Appendix 1. Sequences from 259 individuals (49%
of the dataset) were generated during our study of Australasian
Plagiochilaceae. This included 64 of the 71 species (90%)
currently believed to occur in Australasia (Renner et al. 2017),
including all sectional types. Another 110 individuals of
Plagiochila species from other parts of the world were also
included. The remaining individuals comprised representatives
of Chiastocaulon, Dinckleria, the Lophocoleaceae and
Herbertus Gray.
DNA extraction, polymerase chain reaction
(PCR), sequencing
Clean shoot tips comprising two or three leaves were excised
from dried herbarium specimens and used to extract genomic
DNA with the Qiagen DNeasy 96 Plant Kit (QIAGEN, Valencia,
CA, USA). Sequences for the nuclear ribosomal internal
transcribed spacer region (nrITS1–5.8S-ITS2, hereafter ITS)
and the chloroplast regions rps4 plus the rps4–trnS intergenic
spacer (hereafter rps4) and rbcL were obtained by PCR, with
the following parameters. All reaction volumes were 25 mL and
contained 17 mL of MilliQ water, 2.5 mL of 10 Immolase buffer
(Bioline, London), 2 mL of 2.5 mM dNTPs, 1.25 mL of 50 mM
MgCl2, 0.25 mL of 0.4% bovine serum albumin (BSA), 0.5 mL of
10-mM forward primer, 0.5 mL of 10-mM reverse primer, 0.2 mL of
immolase Taq and 1 mL of DNA. For amplification of rbcL
and ITS, nested PCRs were performed. Temperature profile for
PCR 1 was 95C for 10 min, followed by 25 cycles of 95C for
60 s, 51C for 50 s and 72C for 90 s. A final extension step of 72C
for 10 min completed the thermal cycle. PCR 2 included an extra
five cycles. The primer pairs used for the nested rbcL PCR were
rbcL1-Pl-F (ATGTCACCACAAACGGAGACTAAARCAGGT)
and rbcL-M1390-R (CTTTCCAWAYTTCRCAAGCAGCRG;
Wilson et al. 2004) for the first step using genomic DNA, and
rbcL-38-F (GGTGTTGGATTTAAAGCTGGTG; Wilson et al.
2004) and rbcL-1379-Pl-R (TCACAAGCAGCAGCTAGTTCA
GGACTC; Groth and Heinrichs 2003) in the second PCR seeded
with 1 mL of the product from PCR 1. For nrITS, the primers
Hep2-F (GAGTCATCAGCTCGCGTTGAC) and HepC-R
(TCTCCAGACTACAATTCGCAC) were used in the first step
using genomic DNA, and Hep3-F (CGGTTCGCCGCCGGT
GACG) and HepA-R (CGCCGCTACTAGGGAAATCCTA;
Groth and Heinrichs 2003) were used in the second PCR,
which was seeded with 1 mL of the product from PCR 1.
Amplification of rps4 used the primers rps5-F (ATGTCCCG
TTATCGAGGACCT) and trnS-R (TACCGAGGGTTCG
AATC; Groth and Heinrichs 2003). Temperature profile for
rps4 was 95C for 10 min, followed by 30 cycles of 95C for
60 s, 52C for 60 s and 72C for 150 s. A final extension step of
72C for 10 min completed the thermal cycle. Some samples
required a second semi-nested PCR for rps4, using the alternate
reverse primer rps4-R (TTARRCTTGRCGAGAATAATATTC;
Groth and Heinrichs 2003) and identical cycling conditions.
PCR cleanup, sequencing reactions and reads were completed
by Macrogen Inc. (Seoul, South Korea; http://dna.macrogen.
com/eng/, accessed 21 February 2017). Sequences were
Australian Systematic Botany
73
assembled, checked and edited in Geneious, ver. 6 (Biomatters
Ltd, Auckland, New Zealand, see http://www.geneious.com,
accessed 21 February 2017). Specimen and GenBank data are
provided in Appendix 1.
Phylogeny reconstruction
Sequence editing followed Patzak et al. (2016c) and Renner
et al. (2016). Sequences were aligned manually in BioEdit
version (Hall 1999). Only unambiguously alignable regions
were included in the final matrix. The best-fit model of
evolution for each partition was identified by jModelTest 2
(Guindon and Gascuel 2003; Darriba et al. 2012) with the
Akaike information criterion (AIC). Among models specifiable
within MrBayes, GTR+I+G was selected by AIC for all three
partitions. Each partition was assessed for topologically
conflicting nodes supported by >70% bootstrap support in
maximum-likelihood trees estimated by RAxML, ver. 8
(Stamatakis 2014).
We inferred relationships among individuals on the basis of
the three-marker concatenated dataset by using likelihood and
Bayesian inference methods. The most likely tree topology and
support values were estimated using RAxML, ver. 8, with a
GTRGamma substitution model and 1000 bootstrap replicates.
Trees with high posterior probability were sampled by MrBayes,
ver. 3.2 (Ronquist and Huelsenbeck 2003; Ronquist et al.
2012), with the matrix partitioned by marker, partitions
unlinked, and a GTR+I+G model applied to each. Two runs
each with four chains were executed simultaneously, and
completed 15.5 million generations in 150 h of computational
time with eight cores on the CIPRES Portal (Miller et al. 2010).
Runs were sampled every 10 000 generations. Convergence and
mixing of the runs were confirmed with Tracer, ver. 1.6, available
with BEAST (Drummond et al. 2012). Majority-rule consensus
trees with median node heights were calculated after combining
the runs, minus burnin of 10%, in TreeAnnotator, ver. 1.8.4, to
represent our best estimate of phylogeny. Herbertus sendtneri
(Nees) A.Evans was chosen as outgroup, following Patzak
et al. (2016c).
Results
The concatenated alignment comprised 533 terminals and
2453 bases, nrITS contributing 686 bases (541 parsimony
informative), rps4 573 bases (362 parsimony informative), and
rbcL 1194 bases (1005 parsimony informative). Missing data
comprised 5% of nrITS (24 sequences) 6% of rps4 (30
sequences) and 14% of rbcL (72 sequences). Effective sample
sizes from the Bayesian analysis were LnL 1173, LnPr 13890
and TL[all] 13 911 from a combined 28 million generations
after burnin was excluded.
Within Plagiochilaceae, the tree topology recovered by
maximum likelihood analysis was the same as the majorityrule tree in Renner et al. (2017). Plagiochila was recovered
monophyletic with full support (Figs 1–4). A fully supported
lineage corresponding to Plagiochila section Denticulatae
Schiffn. sensu Patzak et al. (2016c) was placed sister to the
remainder of the genus (Fig. 2). This lineage contained several
predominantly dendroid species, including P. gigantea Lindenb.
and P. fuscella (Hook.f. & Taylor) Gottsche, Lindenb. & Nees,
74
Australian Systematic Botany
1
M. A. M. Renner et al.
1. Dinckleria
2
2. Pseudolophocolea
3
3. Chiastocaulon lineage
4. Plagiochila
Fig. 2
4
equally sized lineages, and, at least on the basis of our
sampling, both have strong representation within Australasia
(Fig. 3). The Australasian representatives of the first lineage are
predominantly tropical species, being regional representatives of
the species rich and mostly tropical sections Cucullatae Schiffn.
and Vagae Lindenb., including P. bantamensis Dumort. and
P. chauviniana Mont. as examples of the former and P. obtusa
Lindenb. and P. metcalfii Steph. as examples of the latter. This
lineage also contains Australasian representatives in other smaller
sections (Fig. 3). The second lineage contains a strong
representation of mostly southern temperate Australasian
species in several sublineages (Fig. 4). Many of these
lineages have been recognised as distinct and formalised as
sections, including section Annotinae Carl, section
Strombifoliae Inoue & R.M.Schust. and section Mitteniae
J.J.Engel & G.L.Merr., see Söderström et al. (2015, p. 83).
However, the relationships between different sections and
their constituent species warrants scrutiny in lieu of the
robust topology resolved by the present analysis.
Fig. 3
Discussion
Summary of sectional groups
Fig. 4
0.03
Fig. 1. Condensed maximum-likelihood phylogeny of the Plagiochilaceae,
showing relationships among major lineages. The Chiastocaulon lineage
includes representatives of the genera Chiastocaulon, Cryptoplagiochila
and Pedinophyllum in our sampling. Pruned tree segments (Figs 2–4) are
indicated.
but also species with more usual growth form, and one species,
P. banksiana Gottsche, whose shoots are prostrate. The next
node separated a fully supported lineage containing
P. fragmentissima Inoue & R.M.Schust. (= P. ratkowskiana
Inoue) and the P. deltoidea Lindenb. species complex, along
with two species tentatively attributed to P. gymnoclada Sande
Lac. from Papua New Guinea and Fiji. The monophyly of the
remainder of Plagiochila was strongly supported. The next
node separated the remainder of Plagiochila into two nearly
Neither sectional classification for Australasian Plagiochila is
fully reflective of the relationships between species recovered
by this study (Fig. 5). The major discrepancies are detailed
below.
Section Annotinae sensu Engel and Merrill (2013) was
recognised for five species sharing a distinctive ventral aspect,
and included section Radiculosae Inoue & R.M.Schust. as a
synonym. None of the species attributed to this section are
closely related to P. annotina Lindenb., although P. baylisii
Inoue & R.M.Schust. was resolved within the broader
Belangerianae lineage. Of the remaining species, P. radiculosa
Mitt. has been transferred to its own genus on the basis of its sister
relationship with Pedinophyllopsis and Pedinophyllum Lindenb.,
and P. bazzanioides J.J.Engel & G.L.Merr. and P. circumdentata
Steph. have been placed in a lineage with P. circinalis (Lehm. &
Lindenb.) Lindenb. and sister to the broader section Belangerianae
Carl. Sections Annotinae and Radiculosae were placed into
synonymy of section Tayloriae Carl by Söderström et al.
(2015); however, the type of section Tayloriae is P. fuscella, a
member of section Denticulatae as discussed further below.
Section Belangerianae sensu Engel & Merrill (2013) was
recognised for five species sharing terminal Frullania-type
vegetative branching, including P. subflabellata Colenso,
P. aculeata (Hook.f. & Taylor) Gottsche, Lindenb. & Nees,
P. fasciculata Lindenb., P. pacifica Mitt. and P. arbuscula
(Brid. ex Lehm. & Lindenb.) Lindenb., the sectional type.
A monophylum including these species was recovered by the
analysis, but this section was rendered paraphyletic by
the monotypic sections Mitteniae and Strombifoliae, by the type
of section Annotinae, by a specimen identified as P. fusca
Sande Lac. and another lineage corresponding to an unidentified
species from Queensland.
Section Banksianae Carl included two species, P. banksiana
and P. intertexta Hook.f. & Taylor, in the sectional scheme of
Engel and Merrill (2013). These two species both belong to
section Denticulatae but were not sister species. Section
Testing relationships within Plagiochila
Australian Systematic Botany
Fig.3
Fig. 2. Maximum-likelihood phylogeny for pruned Segment 1, showing sections accepted by the present study, and voucher identifications as
presented on herbarium specimens or determined using currently available literature and, for Plagiochila aff. peculiaris, comparison with type
material. Numbers above branches are maximum-likelihood bootstrap-support values and Bayesian posterior probabilities respectively, with
asterisk indicating full support. Note that P. andina has not yet been assigned to a section.
75
76
Australian Systematic Botany
M. A. M. Renner et al.
Fig. 2
Fig. 4
Fig. 3. Maximum-likelihood phylogeny for pruned Segment 2, showing sections accepted by the present study, and voucher
identifications as presented on herbarium specimens or determined using currently available literature. Numbers above branches
are maximum-likelihood bootstrap-support values and Bayesian posterior probabilities respectively, with asterisk indicating
full support.
Testing relationships within Plagiochila
Australian Systematic Botany
Fig. 3
Fig. 4. Maximum-likelihood phylogeny for pruned Segment 3, showing sections accepted by the present study, and
voucher identifications as presented on herbarium specimens or determined using currently available literature.
Numbers above branches are maximum-likelihood bootstrap-support values and Bayesian posterior probabilities
respectively, with asterisk indicating full support.
77
M. A. M. Renner et al.
al
20
Giganteae
Banksianae
Giganteae
Hodgsonia
Banksianae
Tayloriae
Hodgsonia
st
ud
y
rs
trö
m
Th
is
de
Sö
ge
l&
Chiastocaulon
et
M
er
ril
l
Pseudolophocolea
Cryptoplagiochila
P. gigantea
P. baileyana
P. intertexta
P. rutlandii
P. sp. nov.
P. gregaria
P. banksiana
P. retrospectans
P. fuscella
P. ratkowskiana
P. sp. nov.
P. spenceriana
P. deltoidea
.2
13
Dinckleria
01
5
Australian Systematic Botany
En
78
Denticulatae
Denticulatae
Tayloriae
Denticulatae
iss
ent
gm
Fra
P. sp. indet. 1
P. abietina
P. aff. pecularis
Peculiares
Peculiares
P. aff. trapezoidea
Plagiochila
Plagiochila
Cucullatae
Cucullatae
P. chauviniana
P. vitiensis
P. bantamensis
P. daviesiana
P. trigona
P. parvifolia
P. sp. nov.
P. teysmannii
P. metcalfii
P. obtusa
P. queenslandica
P. sp. nov.
P. ramosissima
P. colensoi
P. sp. nov.
P. sp. nov.
P. caducifolia
P. bazzanioides
P. circumdentata
P. circinalis
P. incurvicolla
P. conturbata
P. hartziana
P. subflabellata
P. baylisii
P. fasciculata
P. annotina
P. strombifolia
P. sp. nov.
P. stephensoniana
P. pacifica
P. trispicata
P. heterospina
P. arbuscula
Vagae
Cucullatae
Tayloriae
Colensoa
Caducifoliae
Annotinae
Deflexifoliae
Colensoa
e
ima
Durae
Abietinae
Vagae
Vagae
Durae
Tayloriae
Durae
Arrectae
Caducifoliae
Arrectae
Tayloriae
Deflexifoliae
Tayloriae
Belangerianae
Annotinae
Belangerianae
Annotinae
Strombifoliae
Tayloriae
Strombifoliae
Mittenia
Tayloriae
Belangerianae
Belangerianae
Vagae
Fig. 5. Phylogeny of Plagiochila, showing sectional circumscription of two previously published schemes, on the basis of explicit statements
regarding sectional membership and section synonymy, or sectional synonymy and attribution of sectional types alone, and the sectional
circumscription adopted by this study. Species names were resolved on the basis of comparison with type material in lieu of phylogeny
reconstruction and revision of specimen morphology. Blank areas indicate sections not included, or species explicitly assigned to section, by
respective treatments.
Testing relationships within Plagiochila
Banksianae was treated as a synonym of section Denticulatae
by Söderström et al. (2015). Section Belangerianae was not
recognised by Söderström et al. (2015), who regarded it as a
synonym of section Vagae on the basis of a single individual of
P. arbuscula included in Groth et al. (2004). This synonymy
created a polyphyletic section Vagae. Section Caducifoliae
J.J.Engel & G.L.Merr. was proposed by Engel and Merrill
(2013) for the regionally anomalous P. caducifolia Inoue &
R.M.Schust., and was accepted by Söderström et al. (2015);
however, P. caducifolia is nested within section Arrectae.
Section Colensoae J.J.Engel & G.L.Merr. sensu Engel &
Merrill (Engel and Merrill 2013) was proposed for the
outwardly similar species P. colensoi Hook.f. & Taylor and
P. incurvicolla (Hook.f. & Taylor) Gottsche, Lindenb. &
Nees; however, these are not each other’s closest relatives and
this section of two species is polyphyletic. Section Colensoae
was not recognised by Söderström et al. (2015), who regarded
it as a synonym of section Tayloriae. Section Cucullatae
Schiffn. was recognised by both Engel and Merrill (2013) and
Söderström et al. (2015), and was resolved monophyletic in the
present study. Section Deflexifoliae Carl was accepted with
two members, P. circinalis (Lehm. & Lindenb.) Lehm. and
P. microdictyon Mitt. by Engel and Merrill (2013), but was
regarded as a synonym of section Tayloriae by Söderström
et al. (2015). Plagiochila circinalis was resolved sister to
P. bazzanioides and P. circumdentata in a well supported
lineage sister to the broader Belangerianae Carl. Section
Giganteae was accepted with two species, P. gigantea
Lindenb. and P. rutlandii Steph. by Engel and Merrill (2013),
but was regarded as a synonym of section Denticulatae by
Söderström et al. (2015). In our phylogeny, Plagiochila
gigantea and P. rutlandii are not sister species. Section
Hodgsoniae J.J.Engel & G.L.Merr. was proposed for
P. deltoidea, P. fragmentissima and P. gregaria (Hook.f. &
Taylor) Gottsche, Lindenb. & Nees by Engel and Merrill
(2013), but these species were distributed among sections
Durae Carl and Tayloriae by Söderström et al. (2015).
Plagiochila deltoidea and P. fragmentissima belong to the
same lineage, but neither is closely related to P. gregaria
(Hook.f. & Taylor) Gottsche, Lindenb. & Nees, which belongs
within section Denticulatae, making this section polyphyletic.
The synonymisation of section Hodgsoniae with section
Durae renders the latter polyphyletic under the scheme of
Söderström et al. (2015). Section Mitteniae was proposed for
P. stephensoniana Mitt. by Engel and Merrill (2013) but was
placed in synonymy of section Tayloriae by Söderström et al.
(2013). Plagiochila stephensoniana is one of four elements
rendering section Belangerianae sensu Engel & Merrill
paraphyletic. Section Strombifoliae was proposed for
P. strombifolia (Taylor) Taylor, which is the only species
attributed to it. Both Engel and Merrill (2013) and Söderström
et al. (2015) accepted this section, which was resolved nested
within the broader section Belangerianae sensu Engel &
Merrill. In our phylogeny the four species of section Tayloriae
accepted by Engel and Merrill (2013) were resolved in different
lineages, rendering this section polyphyletic. Many sections
were synonymised with section Tayloriae by Söderström et al.
(2015), and these were resolved throughout the phylogeny,
rendering the section polyphyletic under this scheme as well.
Australian Systematic Botany
79
The type species Plagiochila taylorii Steph. was recently made
a synonym of P. fuscella (Engel and Merrill 2009), which is a
member of section Denticulatae. Section Peculiares was
recognised by Söderström et al. (2015), and was resolved
monophyletic in the present study, including one Australian
species, namely P. aff. peculiares. Plagiochila trapezoidea
Lindenb. was placed in a supported sister relationship with
section Plagiochila.
Implications
The topology of the molecular phylogeny resolved from our
comprehensive sampling of species indicated that several
refinements to sectional circumscriptions within Plagiochila
are necessary to achieve a natural classification. These
refinements vary in their complexity and the severity of their
taxonomic implications.
At the simplest end of the spectrum, the placement of the
monotypic section Caducifoliae within section Arrectae is, in
hindsight, compatible with morphological characters shared
by P. caducifolia and other species of section Arrectae, in
particular P. spinulosa (Dicks.) Dumort. and P. lunata S.W.
Arnell. The synonymisation of section Caducifoliae with
section Arrectae is uncontroversial, and means that section
Arrectae is newly recorded for southern temperate Australasia.
Another simple issue concerns section Deflexifoliae; this
section is reinstated from synonymy of section Tayloriae for
P. circinalis, P. bazzanioides and P. circumdentata, all of
which share closely imbricate leaves with deep U-shaped
insertion lines, granular oil bodies, exclusively lateral–
intercalary vegetative branching, and long, ligulate perianths
with parallel dorsal and ventral keels. This section is endemic
to southern temperate Australasia, and all species occur in New
Zealand where two are endemic. The type species, P. circinalis,
also occurs in Tasmania, Victoria, and south-eastern New South
Wales.
What happens with section Hodgsoniae? Plagiochila
deltoidea and P. fragmentissima, two of the three species
placed in section Hodgsoniae by Engel and Merrill (2013), are
related, and belong to a newly identified lineage within
Plagiochila with species from Papua New Guinea and Fiji.
This lineage belongs between section Denticulatae and the
monophylum containing all other species. The third species,
P. gregaria, was nested within section Denticulatae. There are
defining morphological characters for this lineage in the
ventral–intercalary stolon origin and lack of vegetative
Frullania-type branching. As pointed out by Söderström et al.
(2015) section Hodgsoniae is invalid because it was published
without description. An earlier name, section Denticulatae, was
also published without description. The earliest valid name
available for this lineage is section Fragmentissimae (Inoue &
R.M.Schust.) R.M.Schust., which we reinstate below. Section
Deltoideae was synonymised with section Durae by Groth et al.
(2004) who recovered P. deltoidea in a sister relationship with
P. dura. However, their P. deltoidea sequence actually comes
from P. ramosissima (Hook.) Lindenb.
For more than a decade, section Durae has been accepted
as having a South-American–Australasian distribution
(Heinrichs et al. 2005c), on the basis of the sister relationship
between Plagiochila dura De Not. from South America and a
80
Australian Systematic Botany
New Zealand individual identified as P. ramosissima, as
discussed above. An additional member of this lineage
resolved by the present study is P. colensoi Hook.f. & Taylor,
which carries section Colensoae with it into synonymy of
section Durae. The Australasian–South American distribution
of this section holds, and its range extends into the Wet Tropics
Bioregion of north-eastern Queensland, with the resolution of
relationships to a lineage of small epiphytes.
Section Denticulatae was resolved sister to the remainder
of Plagiochila by Patzak et al. (2016c), who accepted a
circumscription encompassing all species within the lineage,
including P. chonotica Taylor, P. banksiana, and two lineages
of P. gigantea. Of these, P. gigantea II is as labelled, whereas
P. gigantea I and P. sp. are both P. fuscella, the type species of
the grossly polyphyletic section Tayloriae, and which rendered
section Denticulatae paraphyletic under the scheme of
Söderström et al. (2015). There are two options for resolving
this paraphyly (the polyphyly of Tayloriae being a separate
issue). Under the first option, section Denticulatae could be
restricted to the type species, P. nobilis Gottsche, and all other
species removed to some or all of the Alternantes Carl,
Banksianae, Giganteae and Baileyanae Inoue & R.M.Schust.,
and a new section for P. rutlandii. There are two difficulties
inherent to this approach. First, there are no clear morphological
grounds for identifying limits to sectional circumscription
compatible with the phylogeny, as it is currently known.
Second, the fragmentation will result in a sectional
classification potentially incapable of accommodating new
species added to the phylogeny, and it may be necessary to
propose more new sections if these are resolved on the
backbone of the clade. Third, although not problematic per se,
the monotypic sections necessary to maintain the Tayloriae
themselves communicate little about the relationships among
species. Under the second option, section Denticulatae could
be maintained in its current broad circumscription by the
formal synonymisation of section Tayloriae with section
Denticulatae, and the maintenance in synonymy of other
sections, following the proposals of Söderström et al. (2015).
This approach would be robust to the addition of previously
unplaced species, avoids difficulties associated with defining
subgroups within this lineage, and formalises a robust
hypothesis of relationship among a distinct group of species.
This is the approach we favour, and we formalise the synonymy
of section Tayloriae below. This also effectively deals with
section Tayloriae, the most polyphyletic section under the
classification proposed by Söderström et al. (2015), by
eliminating it.
Where does Plagiochila trapezoidea belong? This species
could well fit into section Plagiochila, given its perianth
morphology and branching types, and spores having low
surface structure that seems typical of section Plagiochila.
However, according to Inoue (1984), P. trapezoidea has a
thin-walled capsule epidermis, a feature otherwise occurring
only in sections Hylacoetes and Cucullatae (Inoue 1984;
Heinrichs 2002). Here we adopt a conservative position
regarding the sectional placement of this species, and assign
it to section Plagiochila. The alternative, proposal of a new
section, would create a monotypic group of unknown
durability.
M. A. M. Renner et al.
The following five species of the widespread
paleotropical section Belangerianae were accepted for New
Zealand by Engel and Merrill (2013): P. aculeata (Hook.f. &
Taylor) Gottsche, Lindenb. & Nees, P. arbuscula, P. pacifica,
P. subflabellata and P. fasciculata. However, section
Belangerianae was regarded as a synonym of section Vagae
(Söderström et al. 2015) on the basis of a specimen from
Japan included in the study of Groth et al. (2004). In our
study, specimens corresponding with the type of P. arbuscula
were resolved in a lineage sister to the morphologically similar
P. stephensoniana in a rather different part of the phylogeny,
suggesting that the Japanese specimen was misidentified.
This topology carries the section Belangerianae out of
synonymy with section Vagae, but may not necessarily
mandate its re-instatement. The section as circumscribed by
Engel and Merrill (2013) was rendered paraphyletic by section
Strombifoliae, the type species of section Annotinae and
P. baylisii the latter also being attributed to section Annotinae
by Engel and Merrill (2013), one species attributed to section
Colensoae, one species identified as P. fusca, and an
unidentified species from north-eastern Queensland. The
paraphyly of section Belangerianae sensu Engel & Merrill
(2013) is probably the biggest issue of the present study
because it involves making a decision about the status of three
or possibly four sections that are resolved nested within the
broadly circumscribed section Belangerianae accepted by
Engel and Merrill (2013). There are two options. The first is to
retain these embedded sections and recognise additional
sections to achieve monophyletic groups across the phylogeny.
This could be achieved by proposing a new section
for Plagiochila incurvicolla and the P. fasciculata complex.
The challenge is achieving a circumscription for the new
section that successfully differentiates the species belonging to
it from those belonging to section Belangerianae in a narrower
sense. As the circumscription accepted by Engel and Merrill
(2013) suggests, this is not an easy task. All share pinnately
branched, dimorphic leafy shoot systems with Frullania-type
branching, a five- to six-stratose capsule with thickenings in all
layers, and colourless, homogenous or at most few-segmented
oil bodies with a smooth surface. The leaves of all species bear
triangular teeth. The sections nested within the broader
Belangerianae also share these features, with the exception of
P. annotina and P. strombifolia, the vegetative branching of
which is predominantly or exclusively lateral–intercalary.
Plagiochila stephensoniana differs in its multicellular spores
and unispiral elaters adherent to the capsule valves, but shares
the same shoot architecture, and branching-, oil body- and
capsule-anatomy characters as P. arbuscula and its allies.
The nestedness of section Fuscae Carl within the broader
section
Belangerianae
requires
confirmation
against
additional specimens, particularly as species circumscription and
identification are challenging issues within this large genus.
The relationships among the Queensland species, the P. fusca
specimen, P. strombifolia and P. annotina, are all poorly
resolved and subject to fluctuation among inferential approaches.
This lack of signal surrounding nodes critical to inference of
robust sectional limits is a considerable impediment to attempts
to subdivide the broader Belangerianae that strive for stability
in the medium to long term. The broad morphological similarity
Testing relationships within Plagiochila
of species across the lineage, and the lack of clear internal
phylogenetic subdivision correlated with defining morphological
characters are strong arguments in favour of accepting a broad
sectional circumscription. We believe this has two significant
advantages. First, on the basis of current sampling, the
monophylum is well supported. We believe that this is unlikely
to change in response to the inclusion of additional species. Part of
our basis for this assertion is that the monophylum can be
defined using morphological characters of known utility for
sectional circumscription in other parts of the phylogeny, and
these characters encompass a range of morphological features,
including habit, branching types, Frullania-type branching oil
bodies and capsule anatomy. In these features, the broader
Belangerianae consistently differ from its sister lineage, the
Deflexifoliae, which lacks terminal vegetative branching entirely,
has granular oil bodies and ligulate perianths. It is likely that
species could be attributed to a broad section Belangerianae on
the basis of these morphological characters alone, and this is the
second significant advantage.
The alternative, splitting the Belangerianae, would have
the opposing disadvantages, in that further refinement to sectional
circumscription may be necessary in response to species being added
to phylogenetic datasets, and morphological circumscription of
sections may have little predictive power, and may need to
change in response to the inclusion of each additional species. As
evidence for the utility of broad sectional circumscriptions within
Plagiochila, we note that section Vagae encompasses a large
clade of equivalent phylogenetic distinctiveness to our proposed
Belangerianae, and that this group is both morphologically distinct,
but also internally variable. In particular, P. obtusa possesses unusual
morphological characters, but shares with all other species the
characteristic alternating Frullania-type vegetative branching and
production of vegetative propagules from leaf surfaces.
With regard to sectional name, there is high potential here
for us to contribute to a legacy of inadequately founded
sectional synonymies. The identity, and monophyly of
Plagiochila fusca, both need to be corroborated before a
formal synonymy between section Belangerianae and section
Fuscae is proposed. As we have resolved with certainty only the
type species of section Belangerianae, we apply this name to
the lineage, and abstain from further comment on the status of
section Fuscae.
Morphological characters
Relationships currently inferred among species of Plagiochila
mostly prove unreliable, and both sectional classifications
recently applied to Australasian Plagiochila require revision
to achieve monophyly of recognised groups. The severity of
para- and polyphyly varies within both schemes. The most
extreme case of polyphyly is presented by section Tayloriae
as circumscribed by Söderström et al. (2015), which has, on
the basis of Australasian sectional type species alone, six
independent origins and encompasses nearly all major lineages
within Plagiochila, and the segregate genus Cryptoplagiochila.
The only sections resolved monophyletic are those that defy
polyphyly by virtue of being monotypic, such as section
Strombifoliae and section Mitteniae and those proposed by
Inoue (1984) as subgenera. Section Deflexifoliae sensu Engel
Australian Systematic Botany
81
& Merrill (2013) may be monophyletic, but P. microdictyon
was not included in our dataset, so its monophyly has not
been tested.
Morphology, or at least the morphological characters that
have been employed, have proven positively misleading in
establishing higher-order relationships. Some examples below
highlight the extent, both structurally and phylogenetically, of
the challenge faced by taxonomists inferring relationships
within Plagiochila using morphological data. The first
significant difficulty is that shared possession of unusual
features does not necessarily imply a close relationship. Several
unusual features have motivated the placement of unrelated
species in section Annotinae, including the following: presence
of teeth on the dorsal leaf margin in P. baylisii, P. bazzanioides
and P. circumdentata; paraphyllia on the stems of P. bazzanioides
and P. circumdentata; and lamelliform appendages on
perianths of P. annotina, and P. baylisii. Of these species, only
two, P. bazzanioides and P. circumdentata, are related and
they are related to another species with none of these
characters, namely P. circinalis. A ventral lobe on the perianth
is an unusual feature within Plagiochila, and occurs in
P. banksiana and P. intertexta. Although both species are
members of section Denticulatae, they are not each other’s
closest relatives, even among New Zealand taxa. The unusual
monospiral elaters of section Mitteniae are shared with species
of the unrelated section Arrectae. Tightly spicate androecia
with entire male bracts united with the opposing bract occur
in both P. deltoidea and P. gregaria, which belong in different
major lineages. Within P. deltoidea, expression of this character
is inconsistent. Shared possession of this combination of
distinctive reproductive features does not necessarily indicate a
close relationship. These two species also share spores that are
two- to four-celled while still in the capsule (Engel and Merrill
2013), as do species of the Neotropical section Glaucescentes
and species of the pantropical section Vagae (Heinrichs et al.
2000b).
Branch types were repeatedly emphasised as of critical
importance to all levels of leafy liverwort classification
(Schuster 1984, 2001) and, within the Plagiochilaceae, have
been employed as the basis for distinctions among genera,
sections and species (Carl 1931; Hattori 1947; Inoue and
Schuster 1971; Heinrichs et al. 2006). Generally, the presence
or absence of a particular branch type, whether vegetative or
associated with the production of reproductive structures, has
been interpreted as significant. However, in some families such
as the Lophocoleaceae, the shared possession of branching
types did not always serve to delimit natural groups (Engel
1991, 2009; Schuster 2001; Engel and He 2010; Engel et al.
2010). Branch types exhibit homoplasy within Plagiochila
as well. For instance, three lineages within section
Belangerianae, as accepted here, lack terminal branching and
other unrelated lineages such as section Vagae have Frullaniatype vegetative branches. The occurrence of Frullania-type
branches and lateral–intercalary branches seems to be highly
homoplastic in Plagiochila, and several sections consistently
have both branching types, for example, sections
Glaucescentes and Vagae.
Branch types have been employed to circumscribe
generic segregates or re-instatements, Chiastocaulon differs
82
Australian Systematic Botany
from Plagiochila in the widespread occurrence of ventral–
intercalary vegetative branches (Groth and Heinrichs 2003,
Heinrichs et al. 2006, Patzak et al. 2016c), and Dinckleria
differs from most Plagiochila species by stolon production by
ventral–intercalary, rather than lateral–intercalary branching
(Renner et al. 2016).
The second significant difficulty is that overall similarity in a
wide range of features, including habit, branching, leaf-shape
and dentition, does not necessarily imply a close relationship.
Two species, P. fragmentissima and P. gregaria, are so similar
that they have been regarded, but not formalised, as synonyms
(Söderström et al. 2015). The New Zealand P. fragmentissima is
the same as the Tasmanian P. ratkowskiana, which is unrelated
to P. gregaria, despite their overall morphological similarity.
In another example, a relationship between Cryptoplagiochila
radiculosa and section Annotinae was hypothesised by
Engel and Merrill (2013, p. 405), who noted that ‘the
relationship between P. radiculosa and P. annotina, however,
M. A. M. Renner et al.
is underscored by the striking similarities in habit, leaf shape
and arrangement, androecial position, and spores, despite the
total absence of leaf marginal teeth in P. radiculosa’. That
similarity can be an unreliable indicator of relationships has
been emphasised by molecular phylogenetic studies in many
other lineages of bryophytes (Ramaiya et al. 2010; Renner
et al. 2011; Dong et al. 2012).
Instances such as those described above beg the question
whether any relationships within Plagiochila can be confidently
inferred independent of molecular data. The difficulties they
reflect are unlikely to be endemic to the Australasian region,
given that representatives of nearly every major lineage occur
here, these issues show no inclination towards phylogenetic
restriction, and the manifestation of homoplastic characters
in extraterritorial species suggest that they show no inclination
towards geographic restriction either. More than anything
else, the parlous state of the current infrageneric classification
reflects the difficulties involved when inferring relationships
A
B
C
D
Fig. 6. A, B. Ventral intercalary stolons from Plagiochila deltoidea F1141750, showing origin from ventral stem surface between leaf insertion lines (A),
and a newly forming ventral intercalary branch showing the proliferating, elongated mass of dividing cells, which separates the rows of matured ventral
cortical cells (B). C. Lateral intercalary stolons from P. abietina (NSW848776). D. Lateral intercalary stolons from P. gregaria (AK322588), showing origin
from within leaf insertion lines.
Testing relationships within Plagiochila
among species within a lineage expressing considerable
morphological variation and homoplasy.
This is nothing new, relationships resolved on the basis of
molecular data have frequently contradicted relationships
formalised within the Linnean hierarchy, every instance of which
indicates complexities in morphological evolution involving
symplesiomorphy, convergence and multiple independent origins
(Long et al. 2000; Shaw 2000; Vanderpoorten et al. 2002). These
complexities have the downside of confounding attempts to
derive natural classifications from morphological data alone
(Weigend et al. 2013; Clements et al. 2015), but they also have
the upside of informing our appreciation for how dynamic plants
actually are.
Within the Plagiochilaceae, evolutionary lability and
associated homoplasy in morphological character systems
regarded as having high taxonomic utility, particularly from
the vegetative and reproductive gametophyte, has not yet been
quantified. Homoplasy is a known issue within lineages where
high species richness contrasts with low morphological
complexity and variation (Heinrichs et al. 2004d; Devos et al.
2011). Plagiochila, in contrast to many leafy liverwort genera,
expresses both high species richness and high morphological
complexity and variation.
Plagiochilaceae species have, with a small number of
exceptions, shoot systems divided into erect leafy shoots and
creeping stolons. The combination of shoot systems
differentiated into stolons and leafy shoots interacts with
varying modes of stolon and leafy-shoot production to produce
a wide diversity of shoot architectures, possibly the widest of
any liverwort lineage, which may contribute to the high species
diversity and broad ecological range of the lineage. For example,
within section Denticulatae, shoot systems grow by monopodial
replication of an integrated stolon plus leafy-shoot module,
whereas in all other lineages shoot systems grow by sympodial
replication of a module with a differentiated creeping basal
stolon, and an erect leafy shoot. Plant architecture can
contribute to circumscribing monophyletic groups in addition to
branch types, which have been the focus of so much attention
both across leafy liverworts, and within Plagiochila.
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Australian Systematic Botany
83
Branch types exhibit informative patterns of tissue-specific
production. This is particularly valuable as branch types
are present in most individuals, and captured within nearly all
herbarium specimens regardless, and can be used as a universally
applicable framework for placing specimens into a section.
The genera Dinckleria and Chiastocaulon both produce
stolons by ventral–intercalary branching, whereas most Plagiochila
species have lateral–intercalary stolons. The exception is section
Fragmentissimae, which has ventral–intercalary stolons (Fig. 6).
Vegetative branching within leafy-shoot sectors of
Chiastocaulon is ventral–intercalary or Frullania-type. In
contrast, within leafy-shoot sectors of Dinckleria branches arise
by either lateral–intercalary or Frullania-type branching, and
subfloral innovations are exclusively lateral–intercalary in
origin. Vegetative branching within leafy-shoot sectors of
Plagiochila is almost exclusively lateral, either intercalary or
terminal Frullania-type, except when the shoot tip is damaged,
in which case lateral– and ventral–intercalary branches arise
below the damaged tip.
Species in the lineages Vagae and Belangerianae produce
vegetative branches predominantly of the Frullania-type.
Species in other lineages, the vegetative branching of which
is exclusively lateral–intercalary in origin may produce
Frullania-type male branches, the only location within the
plant where terminal branching occurs (Engel and Merrill
2013). Plagiochila incurvicolla occasionally produces ventral–
intercalary vegetative branches from undamaged shoots.
Subfloral innovations in Dinckleria and most Plagiochila
species are lateral–intercalary in origin. In Chiastocaulon,
subfloral innovations are ventral–intercalary when a single
subfloral innovation is produced, and lateral when two or
more are initiated. One lineage of Plagiochila, section
Denticulatae, also produces ventral–intercalary subfloral
innovations.
When aggregations of male branches occur, these are
usually produced by Frullania-type branching. In section
Fragmentissimae, at least one species produces ventral–
intercalary branches at the base or from within male branches
and these may be again fertile.
B
Fig. 7. Teeth. A. Plagiochila baileyana NSW880459. B. P. fuscella var. fuscella MEL2235519.
84
Australian Systematic Botany
M. A. M. Renner et al.
A
B
C
D
Fig. 8. Oil bodies. A. Plagiochila baileyana (NSW880459). B. Plagiochila sp. indet. of section Vagae (NSW Renner 6806). C. Plagiochila fasciculata
(NSW, Renner 6800). D. Plagiochila fasciculata (NSW, Renner 6801). Scale bars: 25 mm.
A distinction between sharply spinose-aciculate teeth, and
teeth of other form was utilised by Inoue and Schuster (1971) in
their artificial key to species. In our phylogeny, species with
spinose-aciculate teeth group within section Denticulatae, and,
thus, the character can contribute to circumscription of this
group. In addition to the spinose teeth, many species in this
section have cell walls around the leaf margin thickened and
intensely pigmented, being often golden or golden-brown
(Fig. 7).
Oil bodies again prove phylogenetically informative. All
regional species of the section Belangerianae share colourless,
homogeneous (or nearly so), smooth oil bodies, which is a
convenient differentiating character from outwardly similar
species belonging to section Vagae that have granular oil bodies
(Fig. 8). This character, in addition to vegetative branching,
distinguishes section Belangerianae from section Deflexifoliae.
Conclusions
Substantial modification of the sectional classification is
required to achieve natural groups corresponding to
monophyletic lineages. A combination of broadening the
circumscription of some recognised entities, and synonymising
smaller embedded sections will achieve a natural classification.
Review and revision of the morphological circumscription of
each section is necessary in lieu of the relationships recovered
by reconstruction of molecular phylogeny. The current
classification suggests that new defining characters, if they
exist, must be identified in lieu of phylogeny, following a
comprehensive re-appraisal of the full range of Australasian
species. Morphological character systems employed have
failed to accurately predict relationships. This task will be
facilitated by full resolution of the substantial nomenclatural
and taxonomic confusion surrounding Australasian species. In
Australasia, as in other parts of the world, informative characters
will likely be derived from facets of morphology so far
neglected and under-utilised, including oil body-, spore-, and
sporophyte-derived characters. Acknowledgement that an
architectural component is key to understanding patterns of
branch-type diversity and variation within Plagiochilaceae
may also increase the utility of this character system.
Testing relationships within Plagiochila
Taxonomic treatment
Here, we present a sectional classification and arrangement for
Australasian species. Only sections with Australasian types are
included, and species lists within each section include species
currently recognised on the basis of published literature, or
known to occur on the basis of published data (Renner et al. 2017).
Plagiochila sect. Denticulatae Schiffn., Hepat. Fl. Buitenzorg
106 (1900)
Type: Plagiochila nobilis Gottsche.
Plagiochila sect. Banksianae Carl, Ann. Bryol. Suppl. 2: 138 (1931).
Type: Plagiochila banksiana Gottsche.
Plagiochila sect. Giganteae Carl, Ann. Bryol. Suppl 2: 143 (1931).
Type: Plagiochila gigantea Lindenb.
Plagiochila sect. Tayloriae (as ‘Taylori ‘) Carl, Ann. Bryol. Suppl 2:
140 (1931), syn. nov.
Type: Plagiochila taylorii Steph. [=P. fuscella (Hook.f. &
Taylor) Gottsche, Lindenb. & Nees].
Plagiochila subsect. Baileyanae Inoue & R.M.Schust., J. Hattori Bot.
Lab. 34: 172 (1971)
Type: Plagiochila baileyana Steph.
Shoot systems in species of section Denticulatae grow by
monopodial replication of a unit having a short erect basal
stolon-like sector and a leafy shoot section on top. The stolonlike section in these species does not creep. However, a few of
species, such as P. gregaria and P. rutlandii, have a shortcreeping basal stolon, and P. banksiana hardly has any stolons
at all. Another feature shared by many species is that the cell
walls of marginal leaf cells are evenly and continuously
thickened, and may be distinctly gold or yellow–brown
pigmented. This is particularly prominent in P. baileyana,
P. gigantea, P. fuscella and P. retrospectans. Third, teeth are
often spinose to acicular, triangular teeth are not a conspicuous
feature of leaves in species of this section, but P. gregaria
and an undescribed species from Tasmania and New Zealand
are again exceptions. Fourth, subfloral innovations are ventral–
intercalary in origin when one is present, but lateral–intercalary
when two or more are present. Species in other sections of
Plagiochila produce lateral–intercalary subfloral innovations
alone, regardless of the number present below the gynoecium.
Australasian species
• Plagiochila baileyana Steph.
• Plagiochila banksiana Gottsche.
• Plagiochila fuscella (Hook.f. & Taylor) Gottsche, Lindenb.
& Nees.
• Plagiochila gigantea Lindenb.
• Plagiochila gregaria (Hook.f. & Taylor) Gottsche,
Lindenb. & Nees.
• Plagiochila intertexta Hook.f. & Taylor.
• Plagiochila retrospectans Lindenb.
• Plagiochila rutlandii Steph.
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Australian Systematic Botany
85
Plagiochila sect. Fragmentissimae (Inoue & R.M.Schust.)
R.M.Schust., Hepaticae and Anthocerotae of North America
4: 449 (1980)
Plagiochila subsect. Fragmentissimae Inoue & R.M.Schust., J. Hattori
Bot. Lab. 34: 154 (1971).
Type: Plagiochila fragmentissima Inoue & R.M.Schust.
Plagiochila sect. Deltoideae E.A.Hodgs., Trans. Roy. Soc. New Zealand
73: 271 (1944), nom. inval. [Art. 38.1a: no description].
Plagiochila sect. Hodgsoniae J.J.Engel & Merr., Nova Hedwigia 96:
403 (2013), nom. inval. [Art.38.1a: no description].
In species belonging to section Fragmentissimae, stolons
initiate by ventral–intercalary branching, either exclusively or in
combination with lateral–intercalary branching from the ventral
part of the lateral merophyte, as in one species within the
P. gymnoclada aggregate. In all other sections of Plagiochila,
stolons originate by lateral–intercalary branches only. Terminal
vegetative branching is absent from species in this section. The
outer surfaces of the capsules of at least one species are water
repellent and covered with a distinctive low ornamentation that
can be removed by chloroform. Whether this ornamentation is
wax and whether it is produced by the sporophyte itself, or is
deposited onto the capsule by surrounding gametophyte tissue
during sporophyte maturation, should be investigated.
Australasian species
• Plagiochila deltoidea Lindenb.
• Plagiochila axillaris Colenso.
• Plagiochila fragmentissima Inoue & R.M.Schust.,
J. Hattori Bot. Lab. 34: 155 (1971).
Type: New Zealand, South Island: Deep Cove, Doubtful Sd.,
Fiordland Natl. Park, R.M.Schuster 53315 (MASS).
Plagiochila ratkowskiana Inoue, Brunonia 3: 141. 1980. syn. nov.A
Type: Tasmania, Mt Wedge, 1000 m, leg. A.V.Ratkowsky n.
78/59 (holotype: TNS, isotype: HO!).
Plagiochila sect. Belangerianae Carl, Ann. Bryol. Suppl. 2:
117 (1931)
Type: Plagiochila belangeriana Lindenb. [= P. arbuscula
(Brid. ex Lehm.) Lindenb.]
Plagiochila sect. Annotinae Carl, Ann. Bryol. Suppl. 2: 143 (1931),
syn. nov.
Type: Plagiochila annotina Lindenb.
Plagiochila sect. Mitteniae J.J.Engel & Merr., Nova Hedwigia 96:
401 (2013), syn. nov.
Type: Plagiochila stephensoniana Mitt.
Plagiochila subsect. Obscurae Inoue & R.M.Schust., J. Hattori Bot.
Laboratory 34: 160 (1971), syn. nov.
Type: Plagiochila obscura Colenso (= P. stephensoniana
Mitt.).
NOTE ADDED IN PROOF. Ongoing revision of Australasian specimens suggests that Plagiochila fragmentissima is likely the same as our
Plagiochila aff. gregaria in Fig. 2. If this is indeed the case the lineage referred to in this paper as sect. Fragmentissimae will require a new name.
86
Australian Systematic Botany
Plagiochila sect. Strombifoliae Inoue & R.M.Schust., J. Hattori Bot.
Lab. 34: 130 (1971), syn. nov.
Type: Plagiochila strombifolia Taylor.
Species of section Belangerianae bear Frullania-type
vegetative branches, at least occasionally, and most species
have pinnate di- or tri-morphic shoot systems, wherein stature
is negatively correlated with branch order. The plants do not
often produce vegetative propagules. Plagiochila annotina is
anomalous in its lack of Frullania-type branches. Teeth on
leaves tend to increase in stature towards the leaf base.
Australasian species
• Plagiochila annotina Lindenb.
• Plagiochila arbuscula (Brid. ex Lehm. & Lindenb.)
Lindenb.
• Plagiochila baylisii Inoue & R.M.Schust.
• Plagiochila conturbata Steph.
• Plagiochila fasciculata Lindenb.
• Plagiochila hartziana Pearson.
• Plagiochila incurvicolla (Hook.f. & Taylor) Gottsche,
Lindenb. & Nees.
• Plagiochila pacifica Mitt.
• Plagiochila stephensoniana Mitt.
• Plagiochila strombifolia Taylor.
• Plagiochila subflabellata Colenso.
• Plagiochila trispicata Colenso.
M. A. M. Renner et al.
not differentially thickened and pigmented, and the teeth on the
leaves are triangular, with a broad base.
Australasian species
• Plagiochila colensoi Hook.f. & Taylor.
• Plagiochila ramosissima (Hook.) Lindenb.
• Plagiochila sp. indet.
Plagiochila sect. Arrectae Carl, Ann. Bryol. Suppl. 2:
52 (1931)
Type: Plagiochila arrecta Gottsche.
Plagiochila sect. Caducifoliae J.J.Engel & Merr., Nova Hedwigia 96:
407 (2013), syn. nov.
Type: Plagiochila caducifolia Inoue & R.M.Schust.
Members of section Arrectae produce exclusively lateral–
intercalary vegetative branches, have simple, intercalary
androecia; homogeneous or indistinctly coarse-segmented
oil bodies, and unispiral elaters (Heinrichs et al. 2004c).
Plagiochila caducifolia shares with some other species of
section Arrectae the striolate ornamentation on the leaf-cell
surfaces, particularly on the cells towards the leaf base.
Australasian species
• Plagiochila caducifolia Inoue & R.M.Schust.
Plagiochila sect. Deflexifoliae Carl, Ann. Bryol. Suppl. 2:
139 (1931)
Plagiochila sect. Vagae Lindenb., Monogr. Hep. Gen.
Plagiochilae xv (1844)
Type: Plagiochila deflexifolia Steph. [= P. circinalis (Lehm. &
Lindenb.) Lehm.]
The species of section Deflexifoliae have irregularly
branched, monomorphic leafy shoots arising by lateral–
intercalary branches only. The leaves are transversely inserted
and orientated, and toothed to varying degrees. Two of the
species bear paraphyllia on the dorsal stem surface. The
perianths are oblong with parallel dorsal and ventral keels.
Type: Plagiochila patula (Sw.) Lindenb.
Species of section Vagae share Frullania-type vegetative
branching, which is often partially or completely
pseudodichotomous, in that the branch displaces to some
degree the parent shoot. The species frequently produce
propagules, mostly from the ventral leaf surface, and often
have endosporous protonemata, perianths often with one or
two wings, all layers of capsule walls with thickenings, and
colorless, granular oil bodies.
Australasian species
• Plagiochila bazzanioides J.J.Engel & G.L.Merr.
• Plagiochila circinalis (Lehm. & Lindenb.) Lehm.
• Plagiochila circumdentata Steph.
Plagiochila sect. Durae Carl, Ann. Bryol. Suppl. 2: 123 (1931)
Type: Plagiochila dura De Not.
Plagiochila sect. Ramosissimae Inoue & R.M.Schust., J. Hattori Bot.
Lab. 34: 202 (1971).
Type: Plagiochila ramosissima (Hook.) Lindenb.
Plagiochila sect. Colensoae [as ‘Colensoa’] J.J.Engel & Merr., Nova
Hedwigia 96: 406 (2013), syn. nov.
Type: Plagiochila colensoi Hook.f. & Taylor.
Both P. colensoi and P. ramosissima grow by monopodial
replication of a shoot unit, having a short erect basal stolon-like
sector and a leafy shoot section on top, a feature they share
with species of section Denticulatae. However, unlike that
section, the leaf marginal cell walls in these two species are
Australasian species
•
•
•
•
•
•
•
•
•
Plagiochila daviesiana Steph.
Plagiochila metcalfii Steph.
Plagiochila norfolkiensis Steph.
Plagiochila obtusa Lindenb.
Plagiochila parvifolia Lindenb.
Plagiochila queenslandica Steph.
Plagiochila teysmannii Sande Lac.
Plagiochila trigona Steph.
Plagiochila wattsiana J.J.Engel & G.L.Merr.
Plagiochila sect. Peculiares Schiffn. Hepat. Fl. Buitenzorg
107 (1900)
Type: Plagiochila peculiaris Schiffn.
Many species of section Peculiares have flexuose leaf cells
and rhizoids continuously distributed along the stem within
leafy shoot sectors. They are also conspicuously nitid. Stolons
and vegetative branches are exclusively lateral–intercalary.
Testing relationships within Plagiochila
Australasian species
• Plagiochila aff. peculiaris (NSW 880483).
Plagiochila sect. Abietinae Schiffn. Hepat. Fl. Buitenzorg:
106 (1900)
Type: Plagiochila abietina (Nees) Mont.
Species in section Abietinae share regularly dimorphic,
pinnately branched shoot systems with Frullania-type
vegetative branches, paraphyllia on the stems, and no vegetative
reproduction.
Australasian species
• Plagiochila abietina (Nees) Nees & Mont.
Plagiochila Dumort. sect. Plagiochila
Type: Plagiochila asplenioides (L.) Dumort.
Species of Plagiochila section Plagiochila have lateral–
intercalary stolons and vegetative branches, granular oil
bodies, long cylindrical perianths and an ellipsoidal capsule
with thickenings in all cell layers, and bispiral elaters
(Heinrichs et al. 2004c); however, P. trapezoidea has a
capsule with a thin-walled outer layer (Inoue 1984).
Australasian species
• Plagiochila aff. trapezoidea (NSW 855092).
Plagiochila sect. Cucullatae Schiffn., Denkschr. Kaiserl.
Akad. Wiss., Wien. Math.-Naturwiss. Kl. 70: 160 (1900)
Type: Plagiochila sandei Dozy ex Sande Lac.
Species in section Cucullatae share a folded or pouchlike elaboration of the antical leaf base, and male branches
with relatively loosely spaced, contiguous male bracts with
rounded, leaf-like lobes. Leaves are often spinose-ciliate.
Small underleaves are usually present on leafy shoots.
Australasian species
Plagiochila bantamensis (Reinw., Blume & Nees) Mont.
Plagiochila blepharophora (Nees) Lindenb.
Plagiochila chauviniana Mont.
Plagiochila reischeckiana Steph. fide Engel and Merrill
(2013).
• Plagiochila sandei Dozy ex Sande Lac.
• Plagiochila sciophila Nees.
• Plagiochila vitiensis Mitt.
•
•
•
•
Incertae sedis
Plagiochila microdictyon Mitt.
Acknowledgements
This study was funded by the Australian Government through the
Australian Biological Resources Study (ABRS) by a Research Fellowship
(grant RFL213–14) awarded to M. A. M. Renner and the Royal Botanic
Gardens & Domain Trust; and an a Ludwig Maximilian University student
research grant to S. D. F. Patzak. The Foundation and Friends of the Royal
Botanic Gardens provided funding for M. A. M. Renner to visit herbaria in
Chicago, Geneva, London, and Munich, which was critical for establishing
Australian Systematic Botany
87
the names for entities resolved by this study. We thank Peter de Lange and
David Meagher for providing material for study; Ewen Cameron (AK),
Brendan Lepschi (CANB), and the curators at F, FH, GOET, HO, JE, M,
MEL, S for loans, access to specimens and permission for destructive
sampling; and John Engel, Matt von Konrat, and David Glenny for their
support of the project.
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Testing relationships within Plagiochila
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Appendix 1. Specimens used in the present study, including information about the origin of the studied material, voucher information, as well as
GenBank accession numbers
Accession numbers in bold were generated during this study and first published in Renner et al. (2017), accession numbers without asterisks were first
published in Patzak et al. (2016c) and those with asterisks were sourced from GenBank
Species
Voucher
nrITS
rps4
rbcL
Brevianthus flavus (Grolle) R.M.Schust. &
J.J.Engel
Chiastocaulon aff. dendroides
Chiastocaulon aff. dendroides
Chiastocaulon biserialis (Lehm. & Lindenb.)
S.D.F.Patzak, M.A.M.Renner, Schäf.Verw. & Heinrichs
Chiastocaulon biserialis
Chiastocaulon biserialis
Australia, Tasmania, Renner 5927
(NSW895251)
Japan, Kurita 84 (HIRO)
Japan, Ohnishi 5570 (HIRO)
Australia, Pócs 0060B (JE)
KT992494
KT992643
KT992570
AY438232
AY438233
KT992526
AY438212
AY438213
–
AY699991
DQ194105
KT992601
New Zealand, Renner 6778 (NSW870823)
Australia, Tasmania, Klazenga N5793
(MEL2111031)
Indonesia, Sumatra, Schäfer-Verwimp 24847
(M)
KT992527
KX828513
KT992671
KX828544
KT992602
KX828528
KT992487
KT992636
–
Indonesia, Sumatra, Schäfer-Verwimp &
Verwimp 25009 (M)
Malaysia, Schäfer-Verwimp & Verwimp
18728 (M)
Australia, Queensland, Renner 6925
(NSW855115)
New Caledonia, Mueller NC344 (JE)
KT992483
KT992632
KT992560
KT992482
KT992631
KT992559
KX828512
KX828545
KX828529
KT992528
KT992672
KT992603
New Zealand, South Island, Schäfer-Verwimp
& Verwimp 14080 (M)
KT992490
KT992639
KT992566
New Zealand, Renner Hf/T18 (AK282526)
Australia, Queensland, Renner 6920b
(NSW855500)
Australia, Queensland, Renner 6922
(NSW855484)
Fiji, Renner 5547 (NSW894781)
Indonesia, Sumatra, Schäfer-Verwimp &
Verwimp 25041 (JE)
Malaysia, Duerhammer 26 (GOET)
Malaysia, Gradstein 10391 (GOET)
Australia, Queensland, Renner 6426
(NSW896965)
New Zealand, Engel 22421 (F1141172)
New Zealand, Renner 6783 (NSW898898)
New Zealand, Braggins 13/240C (NSW)
Australia, Queensland, Renner 2218
(NSW980503)
Australia, Queensland, Renner 6424
(NSW896963)
Australia, Queensland, Renner 6921
(NSW870359)
Australia, Queensland, Renner 7324
(NSW880489)
Fiji, Renner 5815, (NSW980518)
Australia, Queensland, Brown 12/346
(NSW879207)
Japan, Onishi 5588 (HIRO)
KT992540
KX828516
KT992684
KX828548
KT992614
KX828532
KX828515
KX828547
KX828531
KT992529
KT992491
KT992673
KT992640
–
KT992567
KT992492
KT992480
KX828514
KT992641
KT992629
KX828546
KT992568
KT992557
KX828530
KX828518
KX828519
KX828517
KX828522
KX828550
KX828551
KX828549
KX828554
KX828534
KX828535
KX828533
KX828538
KT992533
KT992677
KT992607
KX828521
KX828553
KX828537
KX828520
KX828552
KX828536
KT992534
KX828523
KT992678
KX828555
KT992608
KX828539
AY438238*
AY438220*
AY699999*
KX828524
KX828556
KX828540
Chiastocaulon braunianum (Nees)
S.D.F.Patzak, M.A.M.Renner, Schäf.Verw. & Heinrichs
Chiastocaulon braunianum
Chiastocaulon braunianum
Chiastocaulon braunianum
Chiastocaulon caledonicum (Steph.)
S.D.F.Patzak, M.A.M.Renner, Schäf.Verw. & Heinrichs
Chiastocaulon conjugatum (Hook.)
S.D.F.Patzak, M.A.M.Renner, Schäf.Verw. & Heinrichs
Chiastocaulon conjugatum
Chiastocaulon dendroides (Nees) Carl
Chiastocaulon dendroides
Chiastocaulon dendroides
Chiastocaulon dendroides
Chiastocaulon dendroides
Chiastocaulon dendroides
Chiastocaulon dendroides
Chiastocaulon flamabilis M.A.M.Renner
Chiastocaulon flamabilis
Chiastocaulon flamabilis
Chiastocaulon geminifolium (Mitt.)
M.A.M.Renner
Chiastocaulon geminifolium
Chiastocaulon geminifolium
Chiastocaulon geminifolium
Chiastocaulon geminifolium
Chiastocaulon geminifolium
Chiastocaulon maybarae (S.Hatt.)
S.D.F.Patzak, M.A.M.Renner, Schäf.Verw. & Heinrichs
Chiastocaulon oppositum (Reinw., Blume &
Nees) S.D.F.Patzak, M.A.M.Renner,
Schäf.-Verw. & Heinrichs
Australia, Queensland, Renner 6886
(NSW870834)
(continued next page)
92
Australian Systematic Botany
M. A. M. Renner et al.
Appendix 1. (continued )
Species
Voucher
nrITS
rps4
rbcL
Chiastocaulon oppositum
Australia, Queensland, Renner 7239
(NSW880626)
Australia, Streimann 57083 (CANB9519226)
Indonesia (V), Bali, Schäfer-Verwimp &
Verwimp 16774 (M)
Indonesia, Bali, Schäfer-Verwimp & Verwimp
20711 (M)
Indonesia, Java, Gradstein 11062 (BIOT)
Indonesia, Malaysia, Gradstein 10347
(GOET)
Indonesia, Malaysia, Gradstein 10392
(GOET)
Indonesia, Sulawesi, Gradstein 12005
(GOET)
Indonesia, Sumatra, Schäfer-Verwimp &
Verwimp 24959 (M)
Indonesia, Sumatra, Schäfer-Verwimp &
Verwimp 25015 (M)
Indonesia, Sumatra, Schäfer-Verwimp &
Verwimp 25158 (M)
Australia, Queensland, Renner 6459
(NSW897000)
New Zealand, Braggins 98/237 (AK254223)
KX828525
KX828557
KX828541
KX828526
KT992489
KX828558
KT992638
KX828542
KT992565
KT992488
KT992637
KT992564
KT992479
KT992477
KT992628
KT992626
KT992556
KT992554
KT992478
KT992627
KT992555
KT992476
KT992625
KT992553
KT992486
KT992635
KT992563
KT992485
KT992634
KT992562
KT992484
KT992633
KT992561
KT992535
KT992679
KT992609
KT992538
KT992682
KT992612
New Zealand, Braggins 99/313 (AK254563)
New Zealand, Renner 6776c (NSW870827)
New Zealand, Braggins 00/296D
(AK254836)
Malaysia, Schäfer-Verwimp & Verwimp
18828 (M)
KT992536
KT992537
KT992539
KT992680
KT992681
KT992683
KT992610
KT992611
KT992613
KT992481
KT992630
KT992558
Jamaica, Schäfer-Verwimp 35035 (JE)
KT992510
KT992657
KT992585
Costa Rica, Gradstein 9404 (GOET)
AM282806*
–
DQ312483*
Indonesia, Sumatra, Schäfer-Verwimp &
Verwimp 24801 (JE)
Chile, Drehwald 3206 (GOET)
São Tomé Island, Shevock 39727 (JE)
KT992522
–
KT992597
KT992503
KT992508
KT992651
KT992655
KT992578
KT992583
South Africa, Arts RSA 129/38 (JE)
Portugal, Azores, Schwab 113 (JE)
KT992504
AM282810*
KT992652
–
KT992579
DQ312484*
Chile, Franzaring HF-00-44B (GOET)
AM491290*
KT992659
KT992587
India, Schäfer-Verwimp & Verwimp 28105
(JE)
Ecuador, Gradstein 10119 (GOET)
KT992511
KT992658
KT992586
AM282817*
–
DQ312485*
São Tomé Island, Shevock 39856 (JE)
Australia, Streimann 51629 (JE)
KT992509
AM282819*
KT992656
–
KT992584
DQ312486*
New Zealand, Schäfer-Verwimp & Verwimp
14318 (JE)
Chile, Busch et al. Bryo 01396 (JE)
Australia, Streimann 58464 (GOET)
KT992523
–
KT992598
AM282835*
AM282836*
–
–
DQ312488*
DQ312489*
KT992512
–
KT992588
Chiastocaulon oppositum
Chiastocaulon oppositum
Chiastocaulon oppositum
Chiastocaulon oppositum
Chiastocaulon oppositum
Chiastocaulon oppositum
Chiastocaulon oppositum
Chiastocaulon oppositum
Chiastocaulon oppositum
Chiastocaulon oppositum
Chiastocaulon oppositum
Chiastocaulon proliferum (Mitt.)
S.D.F.Patzak, M.A.M.Renner, Schäf.Verw. & Heinrichs
Chiastocaulon proliferum
Chiastocaulon proliferum
Chiastocaulon proliferum
Chiastocaulon theriotanum (R.M.Schust.)
S.D.F.Patzak, M.A.M.Renner, Schäf.Verw. & Heinrichs
Chiloscyphus coadunatus (Sw.) R.M.Schust.
& J.J.Engel
Chiloscyphus connatus (Sw.) J.J.Engel &
R.M.Schust.
Chiloscyphus costatus (Nees) J.J.Engel &
R.M.Schust.
Chiloscyphus cucullistipulus (Steph.) Hässel
Chiloscyphus difformis (Nees) J.J.Engel &
R.M.Schust.
Chiloscyphus fasciculatus (Nees) Nees
Chiloscyphus fragrans (Moris & De Not.)
J.J.Engel & R.M.Schust.
Chiloscyphus humilis (Hook. f. & Taylor)
Hässel
Chiloscyphus latifolius (Nees) J.J.Engel &
R.M.Schust. agg.
Chiloscyphus martianus (Nees) J.J.Engel &
R.M.Schust.
Chiloscyphus martianus
Chiloscyphus muricatus (Lehm.) J.J.Engel &
R.M.Schust.
Chiloscyphus novae-zeelandiae (Lehm. &
Lindenb.) J.J.Engel & R.M.Schust.
Chiloscyphus sabuletorum
Chiloscyphus semiteres (Lehm.) Lehm. &
Lindenb.
Chiloscyphus vermicularis (Lehm.) Hässel de
Menéndez
Dominican Republic, Schäfer-Verwimp &
Verwimp 26738 (JE)
(continued next page)
Testing relationships within Plagiochila
Australian Systematic Botany
93
Appendix 1. (continued )
Species
Voucher
nrITS
rps4
rbcL
Cryptolophocolea costata (Nees) L.Söderstr.
Indonesia, Sumatra, Schäfer-Verwimp &
Verwimp 24801 (JE)
New Zealand (I), Engel 23337 (GOET)
KT992522
–
KT992597
DQ194059*
DQ194017*
DQ194182*
KT992541
KU888882
KT992685
KU888859
KT992615
KU888871
KT992532
DQ194051
AM180613
KT992676
DQ193991
AM180506
KT992606
DQ194073
KU888883
KT992531
KU888860
KU888858
KT992675
KU888872
KU888870
KT992605
KU888886
KU888863
KU888875
KU888885
KU888862
KU888874
AM180614*
AM180507*
–
KU888884
KU888888
KT992530
DQ194052
KU888861
KU888864
KT992674
DQ194015
KU888887
KU888865
KU888873
KU888876
KT992604
DQ194074 &
DQ194088
KU888877
–
KU888867
KU888879
KU888889
KU888866
KU888878
AJ413177*
AM491300*
AY438203*
–
AY699993*
DQ312494*
KT992513
KT992660
KT992589
AM180588*
–
DQ312493*
Cryptoplagiochila radiculosa (Mitt.)
S.D.F.Patzak, M.A.M.Renner & Heinrichs
Cryptoplagiochila radiculosa
Dinckleria fruticella (Hook.f. & Taylor)
J.J.Engel & Heinrichs
Dinckleria fruticella
Dinckleria fruticella
Dinckleria fruticella
Dinckleria fruticella
Dinckleria fruticella
Dinckleria pleurata (Hook.f. & Taylor)
Trevis.
Dinckleria pleurata
Dinckleria pleurata
Dinckleria pleurata
Dinckleria pleurata
Dinckleria pleurata
Dinckleria pleurata
Dinckleria pleurata
Dinckleria pleurata
Dinckleria singularis (Schiffn.)
M.A.M.Renner, Schäf.-Verw. & Heinrichs
Dinckleria singularis
Herbertus sendtneri (Nees) A.Evans
Heteroscyphus allodontus (Hook.f. & Taylor)
J.J.Engel & R.M.Schust.
Heteroscyphus argutus (Nees) Schiffn.
Heteroscyphus aselliformis (Reinw., Blume &
Nees) Schiffn.
Heteroscyphus ciliatus (Steph.) Schiffn.
Heteroscyphus dubius (Gottsche) Schiffn.
Heteroscyphus fissistipus (Hook.f. & Taylor)
Schiffn.
Heteroscyphus integrifolius (Lehm. &
Lindenb.) Fulford
Heteroscyphus integrifolius
Heteroscyphus knightii (Steph.) Grolle
Heteroscyphus polyblepharis (Spruce)
Schiffn.
Heteroscyphus polychaetus (Spruce)
Hentschel & Heinrichs
Heteroscyphus spectabilis (Steph.) Schiffn.
Heteroscyphus succulentus (Gottsche)
Schiffn.
Heteroscyphus wettsteinii (Schiffn.) Schiffn.
Leptoscyphus amphibolius (Nees) Grolle
New Zealand, Renner 6787 (NSW880455)
New Zealand, Braggins 04/140D
(AK288985)
New Zealand, de Lange 11766 (NSW986163)
New Zealand, Engel 20426 (GOET)
New Zealand, Engel 23943 & von Konrat
(GOET)
New Zealand, Glenny I34 (NSW986162)
New Zealand, Braggins 02/171 (AK287362)
Australia, Queensland, Renner 6822a
(NSW870364)
Australia, Queensland, Renner 6827
(NSW870365)
Australia, Victoria, Meagher s.n.
(MEL2326625)
New Zealand (II), Schäfer-Verwimp &
Verwimp 14071 (GOET)
New Zealand, Braggins 06/004 (AK294976)
New Zealand, Renner 6789 (NSW870825)
New Zealand, Renner 6790 (NSW870826)
New Zealand, Schäfer-Verwimp & Verwimp
13777 (GOET)
Australia, New South Wales, Renner 6756
(NSW978715)
Australia, Queensland, Renner 6877
(NSW859316)
Australia, Queensland, Renner 6750
(NSW970423)
Bolivia, Groth s.n. (GOET)
New Zealand, Frahm 20-6 (GOET)
Thailand, Schäfer-Verwimp & Verwimp
23900 (JE)
Indonesia, Gradstein 10240 (GOET)
New Zealand, Schäfer-Verwimp & Verwimp
13770 (JE)
Principe Island, Shevock 40146 (JE)
Ireland, Long H4064 (JE)
KT992514
KT992661
KT992590
KT992515
AM282841*
KT992662
–
KT992591
DQ312496*
Chile (I), Franzaring CH-00-132 (JE)
KT992516
KT992663
KT992592
Chile (II), Zündorf 21260 (JE)
New Zealand, South Island, Renner 6136
(NSW)
Ecuador, Schäfer-Verwimp & Nebel 31844
(JE)
Ecuador, Gradstein & Mandl 10139 (GOET)
KT992517
KT992493
KT992664
KT992642
KT992593
KT992569
KT992518
KT992665
KT992594
AM491296*
-
DQ312487*
Principe Island, Shevock 40257 (JE)
Malaysia, Schäfer-Verwimp & Verwimp
18640 (JE)
Malaysia, Schäfer-Verwimp & Verwimp
19023 (JE)
Brazil, Schäfer-Verwimp & Verwimp 14748
(JE)
KT992505
KT992519
KT992653
KT992666
KT992580
KT992595
KT992506
KT992654
KT992581
EU350435*
EU661830*
–
(continued next page)
94
Australian Systematic Botany
M. A. M. Renner et al.
Appendix 1. (continued )
Species
Voucher
nrITS
rps4
rbcL
Leptoscyphus amphibolius
Leptoscyphus amphibolius
Leptoscyphus amphibolius
Leptoscyphus amphibolius
Leptoscyphus amphibolius
Leptoscyphus australis (Gottsche, Lindendb.
& Nees) R.M.Schust.
Leptoscyphus cleefii Fulford
Leptoscyphus cuneifolius (Hook.) Mitt.
Mexico (I), Long 29617a (RBGE)
Mexico (II), Long 29617b (RBGE)
Mexico (III), Burghardt 4469 (GOET)
Panama, Dauphin et al. 1578 (GOET)
Ecuador, Kottke & Preussing s.n. (JE)
Australia, Glenny 9431 (LG)
EU350433*
EU350434*
EU350432*
AM491302*
EU350431*
EU350455*
EU661828*
EU661829*
EU661827*
–
EU661826*
EU661825*
–
–
–
DQ312502*
–
–
Venezuela, Long 33049 (RBGE)
Ecuador (I), Schäfer-Verwimp & Preussing
23299/b (JE)
Great Britain (I), Schumacker SBCO 768 (hb.
R. Schumacker)
Great Britain (II), Vanderpoorten 3101 (LG)
Great Britain (III), Long 29812 (RBGE)
Ireland, Blockeel TB 31/1325 (hb. T.
L. Blockeel)
Portugal, Madeira, Schäfer-Verwimp &
Verwimp 26050 (JE)
Ecuador (II), Schäfer-Verwimp & Preussing
23312/c (JE)
South Africa, Prince Edward Islands,
Gremmen, 98-124 (JE)
Costa Rica, Schäfer-Verwimp & Holz SV/H0364 (JE)
Dominica, Schäfer-Verwimp & Verwimp
17647 (JE)
Costa Rica, Gradstein 9694 (GOET)
EU350449*
EU350458*
EU661835*
EU661836*
–
–
EU350461*
EU661857*
–
EU350462*
EU350459*
EU350463*
EU661839*
EU661838*
EU661837*
–
–
–
EU350460*
EU661854*
–
EU350464*
EU661853*
–
AM491303*
-
DQ312503*
EU350457*
EU661855*
–
EU350456*
EU661840*
–
EU350441*
EU661852*
–
KT992497
KT992645
KT992572
EU350446*
KT992520
EU661850*
KT992667
–
KT992596
EU350450*
KT992521
KT992507
EU350453*
EU350454*
EU661841*
KT992668
–
EU661843*
EU661844*
–
–
KT992582
–
–
AM491304*
EU661846*
DQ312504*
EU350447*
EU661845*
–
KT992496
KT992644
KT992571
EU350440*
EU661851*
–
EU350451*
EU661847*
–
EU350442*
EU661848*
–
EU350444*
EU661856*
–
EU350443*
EU661849*
–
EU350437*
EU661832*
–
EU350438*
EU661833*
–
Leptoscyphus cuneifolius
Leptoscyphus cuneifolius
Leptoscyphus cuneifolius
Leptoscyphus cuneifolius
Leptoscyphus cuneifolius
Leptoscyphus cuneifolius
Leptoscyphus expansus (Lehm.) Grolle
Leptoscyphus gibbosus (Taylor) Mitt.
Leptoscyphus gibbosus
Leptoscyphus gradsteinii Vanderp., Schäf.Verw. & D.G.Long
Leptoscyphus gradsteinii
Leptoscyphus gradsteinii
Leptoscyphus hexagonus (Nees) Grolle
Leptoscyphus hexagonus
Leptoscyphus infuscatus (Mitt.) E.W.Jones
Leptoscyphus infuscatus
Leptoscyphus infuscatus
Leptoscyphus intermedius Grolle
Leptoscyphus jackii (Steph.) Grolle
Leptoscyphus jackii
Leptoscyphus jackii
Leptoscyphus lambinonii Vanderp., Schäf.Verw. & D.G.Long
Leptoscyphus physocalyx (Hampe &
Gottsche) Gottsche
Leptoscyphus porphyrius (Nees) Grolle
Leptoscyphus porphyrius
Leptoscyphus porphyrius
Leptoscyphus porphyrius subsp. azoricus
(H.Buch & Perss.) Vanderp. & Heinrichs
Leptoscyphus porphyrius subsp. azoricus
Ecuador (II), Schäfer-Verwimp & Nebel 32205
(M)
Ecuador (I), Schäfer-Verwimp et al. 24282 (JE)
Ecuador, Schäfer-Verwimp & Nebel 31883
(JE)
Venezuela, Long 33049-b (RBGE)
Equatorial Guinea, Bioko, Mueller B560 (JE)
Réunion, Arts Reu 59/3/ (JE)
Malawi, Wiggington M1696b (hb. Hodgetts)
Guadeloupe, Schäfer-Verwimp & Verwimp
22530 (JE)
Ecuador (I), Schäfer-Verwimp et al. 24296
(GOET)
Ecuador (III), Schäfer-Verwimp et al. 24429
(M)
Ecuador (II), Schäfer-Verwimp & Nebel 32769
(M)
Costa Rica, Schäfer-Verwimp & Holz SV/H0342 (JE)
Venezuela, Pócs 05034 H (EGR)
Ecuador (I), Schäfer-Verwimp & Preussing
23229/a (JE)
Ecuador (II), Schäfer-Verwimp et al. 24214/a
(JE)
Ecuador (IV), Schäfer-Verwimp & Preussing
23448 (JE)
Portugal, Azores (II), Terceira, Schumacker
20030611 (hb. Schumacker)
Portugal, Azores (III), Flores, Schumacker
2000 0818/5 (hb. Schumacker)
(continued next page)
Testing relationships within Plagiochila
Australian Systematic Botany
95
Appendix 1. (continued )
Species
Voucher
nrITS
rps4
rbcL
Leptoscyphus porphyrius subsp. azoricus
Portugal, Azores (IV), Flores, Schumacker
2000 0824 (hb. Schumacker)
Portugal, Azores (I), Schumacker 20020812
(hb. Schumacker)
Costa Rica, Schäfer-Verwimp & Holz SV/H
0345 (JE)
Ecuador, Schäfer-Verwimp et al. 24148 (JE)
Germany, Schäfer-Verwimp 35485 (M)
Great Britain, Rycroft 020907 (GOET)
EU350439*
EU661834*
–
EU350436*
EU661831*
–
KT992525
KT992670
KT992600
KT992524
KT992498
AY438234*
KT992669
KT992646
AY438216*
KX090153
KX090171
KT992599
KT992573
DQ194085 &
DQ194071
KX090165
KX090154
KX090172
KX090166
KY051312
KY051060
KY050813
AM180591*
DQ194046*
AJ422027*
AJ422028*
KX090150
KX090151
KX090152
KY051396
KY051417
KY051404
KY051406
KY051468
KY051387
KY051316
KY051317
AM180487*
DQ193996*
AY438204*
AY438205*
–
KX090170
–
KY051149
KY051172
KY051159
KY051161
KY051221
KY051141
KY051064
KY051065
–
DQ194152*
DQ194108*
DQ194109*
KX090162
KX090163
KX090164
KY050901
KY050922
KY050911
KY050913
KY050969
KY050892
–
KY050817
KY051318
KY051314
KY051066
KY051062
KY050818
KY050815
KY051315
KY051063
KY050816
–
KY051278
KY051023
KY051545
KY051307
KY051055
KY051542
KY051304
KY051052
KY051544
KY051306
KY051054
KY051543
KY051305
KY051053
KY051541
KY051303
KY051051
KY051546
AY550130*
DQ194028*
AM180592*
KY051319
KY051308
AY547691*
DQ193974*
AM180488*
KY051067
KY051056
AY699995*
DQ194111*
–
KY050819
KY051320
KY051321
KY051322
AM180593*
KY051068
KY051069
KY051070
AM180489*
KY050820
–
KY050821
–
Leptoscyphus porphyrius subsp. azoricus
Leptoscyphus trapezoides (Mont.) L.Söderstr.
Leptoscyphus trapezoides
Pedinophyllum interruptum (Nees) Kaal.
Pedinophyllum interruptum
Plagiochila abietina (Nees) Nees & Mont.
Plagiochila abietina
Plagiochila abietina
Plagiochila acanthocaulis Sull.
Plagiochila acanthocaulis (Sw.) Lindenb.
Plagiochila adianthoides (Sw.) Lindenb.
Plagiochila aerea Taylor
Plagiochila aff. abietina
Plagiochila aff. abietina
Plagiochila aff. abietina
Plagiochila aff. deltoidea
Plagiochila aff. deltoidea
Plagiochila aff. deltoidea
Plagiochila aff. deltoidea
Plagiochila aff. gregaria
Plagiochila aff. gregaria
Plagiochila aff. peculiaris
Plagiochila aff. peculiaris
Plagiochila aff. peculiaris
Plagiochila aff. peculiaris
Plagiochila aff. peculiaris
Plagiochila aff. peculiaris
Plagiochila aff. trapezoidea
Plagiochila aff. trapezoidea
Plagiochila aff. trapezoidea
Plagiochila aff. trapezoidea
Plagiochila aff. trapezoidea
Plagiochila aff. trapezoidea
Plagiochila alternans Lindenb. & Gottsche
Plagiochila andina Steph.
Plagiochila andina
Plagiochila annotina Lindenb.
Plagiochila annotina
Plagiochila annotina
Plagiochila annotina
Plagiochila ansata (Hook.f. & Taylor)
Gottsche, Lindenb. & Nees
Australia, Queensland, Renner 6977
(NSW848775)
Australia, Queensland, Renner 7274
(NSW848776)
Australia, Queensland, Renner 7275
(NSW849299)
Chile, Holz 252 (GOET)
Argentina, Long 31680 (GOET)
Costa Rica, Heinrichs et al. 4314 (GOET)
Costa Rica, Heinrichs et al. 4321 (GOET)
Fiji, Renner 5464 (NSW890131)
Fiji, Renner 5486 (NSW890173)
Fiji, Renner 5778 (NSW895657)
New Zealand, Beever 101-14e (AK298125)
New Zealand, de Lange 9989 (AK327793)
New Zealand, Engel 20837 (F1141749)
New Zealand, Engel 20913 (F1141788)
New Zealand, Engel 22583 (F1141352)
New Zealand, Renner 6782 (NSW899353)
Australia, Queensland, Renner (NSW852983)
Australia, Queensland, Renner 2230
(NSW980513)
Australia, Queensland, Renner 6449 (NSW)
Australia, Queensland, Renner 7273
(NSW849269)
Australia, Queensland, Renner 7312
(NSW880483.1)
Australia, Queensland, Renner 7312
(NSW880483.2)
Australia, Queensland, Renner 6480
(NSW897031)
Australia, Queensland, Renner 6928
(NSW855092)
Australia, Queensland, Renner 6976
(NSW858927)
Australia, Queensland, Renner 6982
(NSW858922)
Australia, Queensland, Renner 7273
(NSW849294)
Australia, Queensland, Renner 7278 (NSW)
Bolivia, Heinrichs et al. 4178 (GOET)
Bolivia (II), Heinrichs & Müller 4046 (GOET)
Bolivia (I), Heinrichs & Müller 4100 (GOET)
New Zealand, Braggins 04/055D
(AK290979)
New Zealand, Braggins 06/089 (AK303109)
New Zealand, Engel 20962 (F1141203)
New Zealand, Engel 21731 (F1141204)
Chile, Holz 570A (GOET)
(continued next page)
96
Australian Systematic Botany
M. A. M. Renner et al.
Appendix 1. (continued )
Species
Voucher
nrITS
rps4
rbcL
Plagiochila arbuscula (Brid.) Lindenb.
Australia, Queensland, Renner 6328a
(NSW896723)
Australia, Queensland, Renner 6550
(NSW849132)
Australia, Queensland, Renner 6575
(NSW970430)
Australia, Queensland, Renner 6873
(NSW859315)
Australia, Queensland, Renner 6876
(NSW870537)
Australia, Queensland, Renner 6879
(NSW870542)
Australia, Queensland, Renner 6883
(NSW870547)
Australia, Queensland, Renner 6887
(NSW870551)
Australia, Queensland, Renner 6892
(NSW870653)
Australia, Queensland, Renner 6895
(NSW859116)
Australia, Queensland, Renner 7070
(NSW870688)
Australia, Queensland, Renner 7277
(NSW849263)
Australia, Queensland, Renner 7335
(NSW880510)
New Zealand, Shevock 44130 (JE)
Sumatra, Hoffmann 89-209bis
(CANB9310875)
Australia, Queensland, Renner 2123
(NSW980500)
Japan, Yamaguchi 18280 (HIRO)
Italy, Schäfer-Verwimp & Verwimp 35859 (M)
USA (II), Kentucky, Risk 10849 (DUKE)
Australia, Queensland, Renner 7311
(NSW880456)
Australia, Queensland, Renner 7318
(NSW880459)
New Zealand (II), Frey & Pfeiffer 98-T121
(GOET)
New Zealand, Curnow 3480 (CANB9406344)
New Zealand, de Lange 9934 (AK323586)
New Zealand, Shevock 44032 (JE)
Australia, Queensland, Renner 6287
(NSW896671)
Australia, Queensland, Renner 6293
(NSW849176)
Australia, Queensland, Renner 6947
(NSW870748)
Australia, Queensland, Renner 6948
(NSW870754)
Australia, Queensland, Renner 7117
(NSW853005)
Fiji, Renner 5344 (NSW889357)
Fiji, Renner 5444 (NSW889486)
Japan, Yamaguchi 16890 (HIRO)
KY051349
KY051097
KY050848
KY051334
KY051082
KY050833
KY051350
KY051098
KY050849
KY051337
KY051085
KY050836
KY051338
KY051086
KY050837
KY051339
KY051087
KY050838
KY051340
KY051088
KY050839
KY051341
KY051089
KY050840
KY051342
KY051090
KY050841
KY051336
KY051084
KY050835
KY051343
KY051091
KY050842
KY051335
KY051083
KY050834
KY051344
KY051092
KY050843
KT992499
KY051330
KT992647
KY051078
KT992574
KY050829
KY051351
KY051099
KY050850
AY550131*
KT992544
AJ748130*
KY051464
AY547692*
KT992688
AY608099*
KY051217
DQ194112*
KT992617
DQ439699*
KY050965
KY051465
KY051218
KY050966
AY550132*
AY547694*
DQ194114*
KY051360
KY051359
KT992500
KY051367
KY051109
KY051108
KT992648
KY051117
KY050860
KY050859
KT992575
KY050868
KY051361
KY051110
KY050861
KY051363
KY051112
KY050863
–
KY051113
KY050864
KY051362
KY051111
KY050862
KY051364
KY051365
AY275160*
KY051114
KY051115
AY547695*
KY051366
KY051116
KY050865
KY050866
DQ194084* &
DQ194070*
KY050867
AJ866749*
AJ866764*
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila arbuscula
Plagiochila cf arbuscula
Plagiochila asplenioides (L.) Dumort.
Plagiochila austinii A.Evans
Plagiochila baileyana Steph.
Plagiochila baileyana
Plagiochila banksiana Gottsche
Plagiochila banksiana
Plagiochila banksiana
Plagiochila banksiana
Plagiochila bantamensis (Reinw., Blume &
Nees) Mont.
Plagiochila bantamensis
Plagiochila bantamensis
Plagiochila bantamensis
Plagiochila bantamensis
Plagiochila bantamensis
Plagiochila bantamensis
Plagiochila bantamensis
Plagiochila bantamensis
Plagiochila barteri Mitt.
Australia, Queensland, Renner 6286
(NSW896670)
Malawi, O'Shea M7062a (GOET)
(continued next page)
Testing relationships within Plagiochila
Australian Systematic Botany
97
Appendix 1. (continued )
Species
Voucher
nrITS
rps4
rbcL
Plagiochila baylisii Inoue & R.M.Schust.
Plagiochila bazzanioides J.J.Engel &
G.L.Merr.
Plagiochila bazzanioides
Plagiochila bazzanioides
Plagiochila bicuspidata Gottsche
Plagiochila bifaria (Sw.) Lindenb.
Plagiochila bifaria
Plagiochila boryana Gottsche
Plagiochila britannica Paton
Plagiochila buchtiniana Steph.
Plagiochila caducifolia Inoue & R.M.Schust.
Plagiochila caducifolia
Plagiochila caducifolia
Plagiochila caducifolia
Plagiochila carringtonii ssp. lobuchensis
Grolle
Plagiochila chauviniana Mont.
New Zealand, Renner 01/147 (AK255189)
New Zealand, de Lange 8301 (AK308901)
KY051368
KY051370
KY051118
KY051120
KY050869
KY050871
New Zealand, de Lange 9696 (AK322361)
New Zealand, Braggins s.n. (AK257177)
Chile, Drehwald 3201 (GOET)
Ecuador (II), Holz EC-01-416 (GOET)
Spain, Drehwald 3922 (GOET)
Bolivia, Heinrichs et al. 4182 (GOET)
Great Britain, Rycroft 00015 (GOET)
Bolivia, Groth s.n. (GOET)
New Zealand, Engel 22932 (F1141635)
New Zealand, Engel 27071B (C0311977F)
New Zealand, Engel 26697 (C0311975F)
New Zealand, Engel 22807 (F1141636)
Bhutan, Long 28857 (GOET)
KY051371
KY051369
AY390532*
AJ422010*
AJ413173*
AJ866753*
AY275162*
AJ413306*
KY051373
–
KY051372
–
AJ414631*
KY051121
KY051119
DQ194019*
AY438206*
–
AJ866768*
AY438207*
AY438208*
KY051124
KY051123
KY051122
KY051125
AY438209*
KY050872
KY050870
DQ194188*
–
DQ194117*
–
DQ194119*
DQ194120*
KY050875
KY050874
KY050873
KY050876
DQ194121*
Australia, Queensland, Renner 6549
(NSW880463)
Fiji, Renner 5443 (NSW889647)
Fiji, Renner 5449 (NSW889658)
Fiji, Renner 5451 (NSW889660)
Papua New Guinea, Streimann 40603
(CANB8911426)
Papua New Guinea, Streimann 41062
(CANB8911877)
Chile (I), Holz 631 (GOET)
Chile (III), Rycroft 020720-24 (GOET)
Chile (II), Holz & Franzaring CH-00-108
(GOET)
Australia, Victoria, Renner 5199
(NSW893123)
New Zealand, Braggins 04/080B (AK290529)
New Zealand, Braggins s.n. (AK286464)
New Zealand, Braggins s.n. (AK287614)
New Zealand, Engel 20651 (F1141269)
New Zealand, Engel 21406 (F1141268)
New Zealand, Engel 22810 (F1141095)
New Zealand, Pfeiffer 98-T166 (GOET)
New Zealand, Renner 6774 (NSW903565)
New Zealand, Renner 6781 (NSW903564)
New Zealand, Streimann 51055
(CANB9304386)
Australia, Tasmania, Jobson 3415
(MEL2293031)
New Zealand, Braggins 95/121 (AK255132)
New Zealand, Engel 21192 (F1141079)
New Zealand, Braggins 01/691 (AK309826)
New Zealand, Renner 6793 (NSW900040)
New Zealand, Renner 6777a (NSW900148)
New Zealand, Renner 2572a (AK298571)
Rwanda (I), Buchbender & Fischer 1115
(GOET)
Australia, Queensland, Renner 2265
(NSW980505)
Australia, Queensland, Renner 6425
(NSW896964)
Australia, Queensland, Renner 6984
(NSW880559)
KY051376
KY051128
KY050879
KY051377
KY051378
KY051379
KY051374
KY051129
KY051130
KY051131
KY051126
KY050880
KY050881
KY050882
KY050877
KY051375
KY051127
KY050878
AM180594*
DQ194050*
AY550134*
AM180490*
DQ194002*
AY547697*
–
DQ194157*
DQ194122*
KY051386
KY051140
KY050891
KY051381
KY051380
–
KY051384
–
KY051383
DQ194032*
KY051389
KY051388
KY051382
KY051134
KY051132
KY051133
KY051138
KY051137
KY051136
DQ193977*
KY051143
KY051142
KY051135
KY050885
KY050883
KY050884
KY050889
KY050888
KY050887
–
KY050894
KY050893
KY050886
KY051385
KY051139
KY050890
KY051390
KY051392
KY051391
KY051412
KY051476
KY051416
AJ866751*
–
KY051145
KY051144
KY051167
KY051229
KY051171
AJ866765*
KY050895
KY050897
KY050896
–
–
–
DQ194123*
KY051454
KY051207
KY050959
KY051450
KY051203
KY050955
KY051449
KY051202
KY050954
Plagiochila chauviniana
Plagiochila chauviniana
Plagiochila chauviniana
Plagiochila chauviniana
Plagiochila chauviniana
Plagiochila chonotica Taylor
Plagiochila chonotica
Plagiochila chonotica
Plagiochila circinalis (Lehm. & Lindenb.)
Lindenb.
Plagiochila circinalis
Plagiochila circinalis
Plagiochila circinalis
Plagiochila circinalis
Plagiochila circinalis
Plagiochila circinalis
Plagiochila circinalis
Plagiochila circinalis
Plagiochila circinalis
Plagiochila circinalis
Plagiochila circinalis
Plagiochila circumdentata Steph.
Plagiochila circumdentata
Plagiochila circumdentata
Plagiochila colensoi Hook.f. & Taylor
Plagiochila colensoi
Plagiochila colensoi
Plagiochila colorans Steph.
Plagiochila conturbata Steph.
Plagiochila conturbata
Plagiochila conturbata
(continued next page)
98
Australian Systematic Botany
M. A. M. Renner et al.
Appendix 1. (continued )
Species
Voucher
nrITS
rps4
rbcL
Plagiochila conturbata
Australia, Queensland, Renner 7263
(NSW849236)
Australia, Queensland, Renner 7298
(NSW880471)
Australia, Queensland, Renner 7310
(NSW880457)
Australia, Queensland, Renner 1915
(NSW980506)
Costa Rica, Heinrichs et al. 4192 (GOET)
South Africa, Marion-Island, Gremmen
94-007 (GOET)
Ecuador, Holz 558 E/5-01 (GOET)
KY051443
KY051196
KY050948
KY051448
KY051201
KY050953
KY051447
KY051200
KY050952
KY051455
KY051208
KY050960
AJ422015*
DQ194033*
AY438210*
DQ193979*
–
–
AJ422012*
AY438211*
DQ194126*
KY051393
KY051146
KY050898
AJ416083*
AY550135*
–
KY051403
KY051400
–
KY051399
KY051402
AY550136*
KY051410
KY051409
KY051394
AM180597
AJ422014*
AY569439*
AY547698*
KY051153
KY051158
KY051155
KY051152
KY051154
KY051157
AY547699*
KY051165
KY051164
KY051147
AM180493
AY438214*
–
DQ194128*
KY050905
KY050910
KY050907
KY050904
KY050906
KY050909
DQ194129*
KY050917
KY050916
KY050899
–
DQ194130*
DQ194027*
AJ413308*
AY550137*
DQ194037*
AM180599*
AY275163*
DQ193981*
AY438215*
AY547700*
DQ193984*
AM180495*
AY547701*
DQ194102*
DQ194131*
DQ194133*
DQ194135*
–
–
AM180600*
KY051453
AM180496*
KY051206
DQ194127*
KY050958
KY051451
KY051204
KY050956
KY051356
KY051105
KY050856
KY051357
KY051106
KY050857
KY051452
KY051205
KY050957
KY051433
KY051188
KY050938
KY051437
KY051192
KY050942
KY051425
KY051180
KY050930
KY051427
KY051182
KY050932
KY051428
KY051183
KY050933
Plagiochila conturbata
Plagiochila conturbata
Plagiochila conturbata
Plagiochila cristata (Sw.) Lindenb.
Plagiochila crozetensis Kaal.
Plagiochila cucullifolia var. anomala
Heinrichs & Gradst.
Plagiochila daviesiana Steph.
Plagiochila deflexa Mont. & Gottsche
Plagiochila deflexirama Taylor
Plagiochila deltoidea Lindenb.
Plagiochila deltoidea
Plagiochila deltoidea
Plagiochila deltoidea
Plagiochila deltoidea
Plagiochila deltoidea
Plagiochila ramosissima (as P. deltoidea)
Plagiochila deltoidea
Plagiochila deltoidea
Plagiochila deltoidea
Plagiochila dependula Taylor
Plagiochila disticha (Lehm. & Lindenb.)
Lindenb.
Plagiochila divergens Steph.
Plagiochila diversifolia Lindenb. & Gottsche
Plagiochila dura De Not.
Plagiochila dusenii Steph.
Plagiochila elegans Mitt.
Plagiochila ensiformis Taylor
Plagiochila equitans Gottsche
Plagiochila fasciculata Lindenb.
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Australia, Queensland, Renner 6918
(NSW858917)
Costa Rica (II), Heinrichs et al. 4160 (GOET)
Costa Rica, Heinrichs 11 (GOET)
New Zealand, Engel 18409 (F1141063)
New Zealand, Engel 20652 (F1141748)
New Zealand, Engel 22727 (F1141080)
New Zealand, Engel 22920 (F1140885)
New Zealand, Engel 23325 (F1141064)
New Zealand, Engel 23428 (F1141729)
New Zealand, Frey 98-T116 (GOET)
New Zealand, Renner 6776b (NSW899342)
New Zealand, Renner 6780 (NSW899341)
New Zealand, Braggins s.n. (AK287128)
Ecuador, Heinrichs et al. 4442 (GOET)
Ecuador, Holz 436 E/5-01 (GOET)
Kenia, Solga s.n. (GOET)
Ecuador, Holz 017 E/5-01 (GOET)
Chile, Drehwald 3190 (GOET)
Argentina, Holz 146 (GOET)
China, Long 33675 (GOET)
Ecuador, Sauer & Gradstein MS-E 115, BEGP
149 (GOET)
Chile, Holz 625 (GOET)
Australia, New South Wales, Renner 6771
(NSW978730)
Australia, New South Wales, Renner 6801
(NSW900149)
Australia, New South Wales, Renner 6813
(NSW850854)
Australia, New South Wales, Renner 6842
(NSW859320)
Australia, New South Wales, Renner 6848
(NSW900152)
Australia, New South Wales, Steimann 44457
(CANB9008881)
Australia, New South Wales, Steimann 61245
(CANB9910863.1)
Australia, New South Wales, Streimann 38243
(CANB9006252)
Australia, New South Wales, Streimann 47674
(CANB9107719)
Australia, New South Wales, Streimann 47856
(CANB9107902)
(continued next page)
Testing relationships within Plagiochila
Australian Systematic Botany
99
Appendix 1. (continued )
Species
Voucher
nrITS
rps4
rbcL
Plagiochila fasciculata
Australia, New South Wales, Streimann 49561
(CANB9216583)
Australia, Queensland, Renner 6809
(NSW850851)
Australia, Queensland, Renner 6811
(NSW850853)
Australia, Queensland, Renner 6824
(NSW870747)
Australia, Renker A18.2 (GOET)
Australia, Tasmania, Curnow 2141
(CANB8807469)
Australia, Tasmania, Moscal 14255
(CANB781957)
Australia, Tasmania, Whinray 1291
(MEL2238068)
Australia, Tasmania, Whinray B1877
(MEL2209256)
Australia, Victoria, Curnow 4176
(CANB9501564)
Australia, Victoria, Streimann 42679
(CANB8915211)
Australia, Victoria, Streimann 43785
(CANB9007770)
Australia, Victoria, Streimann 58768
(CANB9802439)
Australia, Victoria, Streimann 58820
(CANB9802491)
Australia, Victoria, Streimann 58825
(CANB9802496)
Australia, New South Wales, Renner 6758
(NSW978717)
Mexico, Gradstein 8274 (GOET)
Japan, Kurita 147 (HIRO)
Australia, Tasmania, Ratkowsky H1571
(CANB8409926)
Australia, Tasmania, Whinray B2227
(MEL2354417)
Indonesia, Bali, Schäfer-Verwimp & Verwimp
20704 (GOET)
India, Long 23002 (GOET)
Indonesia, Gradstein 10259 (GOET)
New Zealand, de Lange 7355 (AK303495)
New Zealand, Streimann 51026
(CANB9304357)
New Zealand, Streimann 51191
(CANB9306641)
New Zealand, Streimann 51342
(CANB9306792)
Australia, Victoria, Streimann 50924
(CANB9304254)
New Zealand, Frey & Pfeiffer 98-T88 (GOET)
Costa Rica, Heinrichs et al. 4400 (GOET)
Seychelles, Pócs 9342/B (GOET)
New Zealand (II), Engel & von Konrat 23911
(F)
New Zealand, Shevock 43981 (JE)
New Zealand, Streimann 51236
(CANB9306686)
New Zealand, Glenny I18 (NSW)
KY051429
KY051184
KY050934
KY051355
KY051104
KY050855
KY051444
KY051197
KY050949
KY051446
KY051199
KY050951
DQ194040*
KY051421
DQ193988*
KY051176
DQ194138*
KY050926
KY051419
KY051174
KY050924
KY051441
–
KY050946
KY051440
KY051195
KY050945
KY051430
KY051185
KY050935
KY051424
KY051179
KY050929
KY051426
KY051181
KY050931
KY051434
KY051189
KY050939
KY051435
KY051190
KY050940
KY051436
KY051191
KY050941
KY051358
KY051107
KY050858
AJ744790*
AY550138*
KY051496
DQ193989*
AY547703*
KY051257
DQ194139*
DQ194140*
KY051003
KY051497
KY051258
KY051004
AY438237*
AY438219*
DQ194141*
AY438235*
DQ194041*
KY051456
KY051458
AY438217*
DQ193992*
KY051209
KY051211
–
DQ194143*
–
–
KY051459
KY051212
KY050961
KY051460
KY051213
–
KY051457
KY051210
–
AY550143*
AJ416086*
AJ866746*
AM180604*
AY547721*
AY547704*
AJ866760*
AM180499*
DQ194186*
DQ194144*
DQ194145*
DQ194146*
KT992501
KY051466
KT992649
–
KT992576
KY050962
KY051462
KY051215
KY050963
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fasciculata
Plagiochila fastigiata Lindenb. & Gottsche
Plagiochila flexuosa Mitt.
Plagiochila fragmentissima Inoue &
R.M.Schust.
Plagiochila fragmentissima
Plagiochila frondescens (Nees) Lindenb.
Plagiochila fruticosa Mitt.
Plagiochila fusca Sande Lac.
Plagiochila fuscella Taylor
Plagiochila fuscella
Plagiochila fuscella
Plagiochila fuscella
Plagiochila fuscella
Plagiochila fuscella
Plagiochila fuscolutea Taylor
Plagiochila fusifera Taylor
Plagiochila gigantea Lindenb.
Plagiochila gigantea
Plagiochila gigantea
Plagiochila gigantea
(continued next page)
100
Australian Systematic Botany
M. A. M. Renner et al.
Appendix 1. (continued )
Species
Voucher
nrITS
rps4
rbcL
Plagiochila gregaria (Hook.f. & Taylor)
Gottsche, Lindenb. & Nees
Plagiochila gregaria
Plagiochila gregaria
Plagiochila gymnocalycina (Lehm. &
Lindenb.) Mont.
Plagiochila gymnoclada Sande Lac.
Plagiochila gymnoclada
New Zealand, de Lange 9936 (AK323588)
KY051467
KY051220
KY050968
New Zealand, Engel 21089 (F1141554)
New Zealand, de Lange 8302 (AK308902)
Bolivia, Drehwald 4729 (GOET)
KY051469
–
AY390531*
KY051222
KY051219
AY438218*
KY050970
KY050967
DQ194147*
Fiji, Renner 5538 (NSW890228)
Papua New Guinea, De Sloover 42776
(CANB9312443)
Japan, Yamaguchi 12271 (HIRO)
Indonesia, Gradstein 11036 (GOET)
KY051471
KY051470
KY051224
KY051223
KY050972
KY050971
AY275164*
AJ866735
AY547705*
AJ866757
KY051423
KY051178
DQ194149*
DQ194080 &
DQ194066
KY050928
KY051420
KY051175
KY050925
KY051422
KY051177
KY050927
DQ194045*
–
DQ194150*
Plagiochila hakkodensis Steph.
Plagiochila hampeana Gottsche
Plagiochila hartziana Pearson
Plagiochila hartziana
Plagiochila hartziana
Plagiochila heterodonta (Hook.f. & Taylor)
Gottsche, Lindenb. & Nees
Plagiochila heterophylla Lindenb. ex Lehm.
Plagiochila heterophylla
Plagiochila heterospina Steph.
Plagiochila heterospina
Plagiochila heterospina
Plagiochila heterospina
Plagiochila heterostipa Steph.
Plagiochila hookeriana Lindenb.
Plagiochila incerta Gottsche
Plagiochila incurvicolla (Hook.f. & Taylor)
Gottsche, Lindenb. & Nees
Plagiochila incurvicolla
Plagiochila incurvicolla
Plagiochila incurvicolla
Plagiochila integerrima Steph.
Plagiochila intertexta Mitt.
Plagiochila intertexta
Plagiochila intertexta
Plagiochila intertexta
Plagiochila javanica (Sw.) Nees & Mont.
Plagiochila kiaerii Gottsche
Plagiochila korthalsiana Molk.
Plagiochila lastii Mitt.
Plagiochila longiramea Steph.
Plagiochila lunata S.W.Arnell
Plagiochila macrostachya Lindenb.
Plagiochila maderensis Gottsche ex Steph.
Plagiochila magna Inoue
Plagiochila metcalfii Steph.
Plagiochila metcalfii
Plagiochila metcalfii
Plagiochila minutula (Hook.f. & Taylor)
Gottsche, Lindenb. & Nees
Plagiochila nobilis Gottsche
Australia, Tasmania, Curnow 2660
(CANB8808028)
Australia, Tasmania, Ratkowsky H1553
(CANB8408762)
Australia, Tasmania, Curnow 2606
(CANB8807975)
South Africa, Marion Island, Ochyra 1317/03
(GOET)
Great Britain, Rycroft 97006, B.E.G.P. 80
(GOET)
Jamaica, Schäfer-Verwimp 35404 (JE)
Fiji, Renner 5333 (NSW889332)
Fiji, Renner 5417 (NSW889522)
Fiji, Renner 5476 (NSW890152)
Fiji, Renner 5323 (NSW889302)
Malawi, O'Shea M7070a (GOET)
AY275165*
AY547706*
DQ194151*
KT992542
KY051346
KY051347
KY051348
KY051345
AJ866735*
KT992686
KY051094
KY051095
KY051096
KY051093
AJ866757*
Chile, Holz 560 (GOET)
Madagascar, Pócs 9447/L (GOET)
New Zealand, Engel 23137 (F1132133)
AM180605*
AJ866737*
KY051473
AM180500*
AJ866761*
KY051226
KY050845
KY050846
KY050847
KY050844
DQ194080* &
DQ194066*
–
DQ194101*
KY050974
New Zealand, Renner 01/136 (AK255217)
New Zealand, Renner 6796 (NSW899787)
New Zealand, Braggins 13/031A (NSW)
Malawi, O'Shea M7552a (GOET)
New Zealand, Engel 21764A (F1141311)
New Zealand, Engel 23347 (F1182057)
New Zealand, Renner 01/253B (AK282335)
New Zealand, Engel 21090 (F1141683)
Indonesia, Java, Gradstein 10209 (GOET)
Madagascar, Pócs 9477/A6 (GOET)
Indonesia, Gradstein 10258 (GOET)
Comoros, Pócs et al. 9273/U (GOET)
Bolivia, Heinrichs et al. 4071 (GOET)
KY051413
KY051474
KY051472
AY275166*
KY051502
KY051503
KY051501
–
AJ744791*
AJ866747*
DQ194049*
AY550139*
AF539463*
KY051168
KY051227
KY051225
AY547707*
KY051263
KY051265
KY051262
KY051264
DQ193998*
–
DQ194000*
AY547708*
AY547709*
Lesotho, Duckett 34026 (GOET)
Ecuador, Holz 080 E/5-01 (GOET)
Portugal, Madeira (II), Rycroft 99030 (GOET)
Japan, Kurita 258 (HIRO)
Norfolk Island, Streimann (CANB9513847)
Norfolk Island, Ward 94-1A (CANB9507854)
Norfolk Island, Curnow 4867
(CANB9513836)
South Africa, Prince Edward Islands, Ochyra
434/01 (GOET)
Indonesia, Gradstein 10253 (GOET)
DQ194026*
AJ422016*
AY462143*
AY275167*
–
–
–
DQ194003*
AY547710*
DQ194005*
AY438221*
KY051232
KY051230
KY051231
KY050919
KY050975
KY050973
DQ194095*
KY051008
KY051010
KY051007
KY051009
DQ194154*
DQ194094*
DQ194156*
DQ194097*
DQ194089* &
DQ194075*
DQ194100*
DQ194160*
DQ194161*
DQ194162*
KY050979
KY050977
KY050978
DQ194054*
DQ194007*
–
DQ194055*
DQ194010*
DQ194166*
(continued next page)
Testing relationships within Plagiochila
Australian Systematic Botany
101
Appendix 1. (continued )
Species
Voucher
nrITS
rps4
rbcL
Plagiochila obovata Steph.
Plagiochila obovata
Plagiochila obtusa Lindenb.
Chile (I), Holz 385 (GOET)
Chile (II), Holz 583 (GOET)
Australia, Queensland, Renner 6941
(NSW853117)
Japan, Kurita 132 (HIRO)
Japan, Ohnishi 5723 (HIRO)
Costa Rica, Heinrichs et al. 4158 (GOET)
Ecuador, Anton ECU4 (GOET)
New Zealand, de Lange K212 (AK313957)
New Zealand, de Lange K219 (AK305199)
New Zealand, de Lange K250 (AK305205)
Cook Islands, Rarotonga, de Lange CH196
(NSW)
Australia, Queensland, Renner 6902
(NSW858870)
Thailand, Schäfer-Verwimp & Verwimp
23824 (GOET)
Australia, Queensland, Renner 6332a
(NSW896732)
Brazil, Gradstein 3703 (GOET)
Costa Rica, Holz CR 00-0176 (GOET)
Bhutan, Long 28832 (GOET)
India, Long 22802 (GOET)
Portugal, Schäfer-Verwimp & Verwimp
31246 (M)
Japan, Ohnishi 5771 (HIRO)
Great Britain, Rycroft 01013 (GOET)
Australia, Queensland, Renner 6521
(NSW898761)
Australia, Queensland, Renner 7103
(NSW858831)
Australia, Queensland, Renner 7107
(NSW852980)
Australia, Queensland, Renner 6334
(NSW896736)
Ecuador (II), Holz 045 E/5-01 (GOET)
New Zealand, Engel 22033 (F1141444)
Malaysia, Schäfer-Verwimp & Verwimp
18736/A (GOET)
Costa Rica, Heinrichs et al. 4154 (GOET)
Australia, New South Wales, Streimann 43891
(CANB9007877)
Australia, Tasmania, Jobson (MEL2235519)
Australia, Victoria, Klazenga (MEL2131756)
Australia, Victoria, Streimann 50816
(CANB9300450)
Chile (II), Rycroft 020723-6 (GOET)
AM180610*
AM180611*
–
AM180503*
AM180504*
KY051251
–
–
KY050998
AY275168*
AY275169*
AJ422017*
DQ194057*
KY051493
KY051491
KY051492
KY051494
AY438222*
AY547711*
AY547712*
DQ194013*
KY051254
KY051252
KY051253
KY051255
DQ194167*
–
AY699997*
DQ194171*
KY051001
KY050999
KY051000
KY051002
KY051485
KY051241
KY050988
DQ194058*
DQ194014*
DQ194172*
KY051487
KY051243
KY050990
AY550140*
AY275170*
AY550141*
AY550142*
KT992543
AY547713*
AY547714*
AY547716*
AY547717*
KT992687
DQ194173*
DQ194174*
DQ194176*
DQ194177*
KT992616
AY438239*
AJ413174*
KY051489
AY438223*
AY547719*
KY051245
DQ194179*
DQ194180*
KY050992
KY051480
KY051236
KY050983
KY051477
KY051233
KY050980
KY051488
KY051244
KY050991
AJ422020*
KY051495
AY569441*
AY438225*
KY051256
AY569440*
DQ194181*
–
DQ194184*
AJ422021*
KY051498
AY547720*
KY051259
DQ194185*
KY051005
KY051500
KY051461
KY051499
KY051261
KY051214
KY051260
–
–
KY051006
AJ781752*
AJ866767*
DQ194187*
Bolivia (I), Groth 101 (GOET)
Ecuador, Holz 408 E/5-01 (GOET)
AJ416081*
AJ416080*
AY438226*
DQ194009*
DQ194189*
DQ194165*
New Zealand, de Lange 11767 (NSW848797)
New Zealand, Glenny I20 (NSW)
Indonesia, Gradstein 9970 (GOET)
Japan, Ohnishi 5720 (HIRO)
Japan, Ohnishi 5400 (HIRO)
USA, Smith & Davison s.n. (GOET)
Nepal, Long 21348 (GOET)
KY051524
KY051463
AJ414634*
AY550144*
AY275171*
DQ194039*
AY275172*
KY051289
KY051216
AY438228*
AY547723*
AY547724*
DQ193987*
AY438227*
KY051035
KY050964
DQ194191*
DQ194192*
DQ194193*
DQ194136*
DQ194194*
Plagiochila orbicularis (S.Hatt) S.Hatt.
Plagiochila ovalifolia Mitt.
Plagiochila ovata Lindenb. & Gottsche
Plagiochila pachyloma Taylor
Plagiochila pacifica Mitt.
Plagiochila pacifica
Plagiochila pacifica
Plagiochila pacifica
Plagiochila parvifolia Lindenb.
Plagiochila parvifolia
Plagiochila parvifolia
Plagiochila patentissima Steph.
Plagiochila patriciae J.Heinrichs & H.Anton
Plagiochila peculiaris Schiffn.
Plagiochila poeltii Inoue & Grolle
Plagiochila porelloides (Torr. ex Nees)
Lindenb.
Plagiochila pulcherrima Horik.
Plagiochila punctata (Taylor) Taylor
Plagiochila queenslandica Steph.
Plagiochila queenslandica
Plagiochila queenslandica
Plagiochila queenslandica
Plagiochila raddiana Lindenb.
Plagiochila ramosissima (Hook.) Lindenb.
Plagiochila renitens (Nees) Lindenb.
Plagiochila retrorsa Gottsche
Plagiochila retrospectans Lindenb.
Plagiochila retrospectans
Plagiochila retrospectans
Plagiochila retrospectans
Plagiochila rubescens (Lehm. & Lindenb)
Lindenb.
Plagiochila rutilans Lindenb.var. rutilans
Plagiochila rutilans var. moritziana (Lindenb.
& Gottsche) Heinrichs
Plagiochila rutlandii Steph.
Plagiochila rutlandii
Plagiochila sandei Dozy
Plagiochila satoi S.Hatt.
Plagiochila sciophila Nees
Plagiochila sciophila
Plagiochila semidecurrens (Lehm. &
Lindenb.) Lindenb.
(continued next page)
102
Australian Systematic Botany
M. A. M. Renner et al.
Appendix 1. (continued )
Species
Voucher
nrITS
rps4
rbcL
Plagiochila sp.
Australia, New South Wales, Renner 6753
(NSW978711)
Australia, New South Wales, Renner 6755
(NSW978714)
Australia, New South Wales, Renner 6763
(NSW978721)
Australia, New South Wales, Renner 6769
(NSW978728)
Australia, New South Wales, Renner 6773
(NSW978732)
Australia, New South Wales, Renner 6805
(NSW850847)
Australia, New South Wales, Renner 6807
(NSW850849)
Australia, New South Wales, Streimann 60997
(CANB9910339.1)
Australia, Queensland, Renner 6416
(NSW896955)
Australia, Queensland, Renner 6826
(NSW858805)
Australia, Queensland, Renner 6839
(NSW870815)
Australia, Queensland, Renner 6840
(NSW858815)
Australia, Queensland, Renner 6846
(NSW858826)
Australia, Queensland, Renner 6882
(NSW859317)
Australia, Queensland, Renner 6898
(NSW859318)
Australia, Queensland, Renner 6920a
(NSW855499.1)
Australia, Queensland, Renner 6920a
(NSW855499.2)
Australia, Queensland, Renner 6929
(NSW855503)
Australia, Queensland, Renner 7112
(NSW852994)
Australia, Queensland, Renner 7294
(NSW880570)
Australia, Queensland, Renner 7302
(NSW880473)
Fiji, Renner 5407 (NSW889511)
Fiji, Renner 5523 (NSW890205)
Australia, Queensland, Renner 6406
(NSW900037)
Australia, Queensland, Renner 7331
(NSW880506)
Australia, Queensland, Renner 7334
(NSW880509)
New Zealand, Curnow 3556 (CANB9408642)
New Zealand, Engel 20706 (F1141750)
New Zealand, Engel 22601 (F1141724)
New Zealand, Streimann 51257
(CANB9306707)
New Zealand, Braggins 04/055C (AK290978)
Belgium, Dauphin et al. 3811 (GOET)
Kenya (I), Chuah 0310/AB (GOET)
New Zealand, Braggins 05/084 (AK294765)
New Zealand, Shevock 44043 (JE)
KY051519
KY051282
KY051028
KY051520
KY051283
KY051029
KY051521
KY051284
KY051030
–
KY051285
KY051031
KY051522
KY051286
KY051032
KY051504
KY051266
KY051011
KY051505
KY051267
KY051012
–
KY051287
KY051033
KY051518
KY051281
KY051027
KY051509
KY051271
KY051016
KY051514
KY051276
KY051021
KY051510
KY051272
KY051017
KY051511
KY051273
KY051018
KY051512
KY051274
KY051019
KY051513
KY051275
KY051020
KY051506
KY051268
KY051013
KY051507
KY051269
KY051014
KY051508
KY051270
KY051015
KY051445
KY051198
KY050950
KY051516
KY051279
KY051024
KY051515
KY051277
KY051022
–
KY051517
KY051411
KY051280
–
KY051166
KY051025
KY051026
KY050918
KY051407
KY051162
KY050914
KY051408
KY051163
KY050915
KY051398
KY051405
KY051401
KY051397
KY051151
KY051160
KY051156
KY051150
KY050903
KY050912
KY050908
KY050902
KY051395
AY275173*
AJ744796*
KY051523
KT992502
KY051148
AY547725*
AJ866758*
KY051288
KT992650
KY050900
DQ194195*
DQ194098*
KY051034
KT992577
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp.
Plagiochila sp. nov.
Plagiochila sp. nov.
Plagiochila sp. nov.
Plagiochila spenceriana Colenso
Plagiochila spenceriana
Plagiochila spenceriana
Plagiochila spenceriana
Plagiochila spenceriana
Plagiochila spinulosa (Dicks.) Dumort.
Plagiochila squamulosa Mitt.
Plagiochila stephensoniana Mitt.
Plagiochila stephensoniana
(continued next page)
Testing relationships within Plagiochila
Australian Systematic Botany
103
Appendix 1. (continued )
Species
Voucher
nrITS
rps4
rbcL
Plagiochila stephensoniana
New Zealand (II), South Island, SchäferVerwimp & Verwimp 14091 (GOET)
Indonesia, Gradstein 10309 (GOET)
Madagascar, Pócs 9868/AF (GOET)
Costa Rica, Heinrichs et al. 4401 (GOET)
Malawi, Hodgetts M2490a (GOET)
Australia, New South Wales, Renner 5860
(NSW898641)
Australia, Tasmania, Croft 10377
(CANB8904602)
Australia, Tasmania, Curnow 4310
(CANB9503011)
Australia, Tasmania, Curnow 4476
(CANB9503774)
Australia, Tasmania, Curnow 4492
(CANB9504674)
Australia, Tasmania, Curnow 4512
(CANB9504694)
Australia, Tasmania, Curnow 4551
(CANB9504733)
Australia, Tasmania, Streimann 59563
(CANB9803242)
Australia, Tasmania, Streimann 59690
(CANB9803373)
Australia, Victoria, Curnow 4153
(CANB9501541)
Australia, Victoria, Klazenga N5846
(MEL2114285)
Australia, Victoria, Streimann 43616
(CANB9007597)
Australia, Victoria, Streimann 50471
(CANB9219468)
Australia, Victoria, Streimann 58774
(CANB9802445)
Australia, Victoria, Streimann 58818
(CANB9802489)
Australia, Lord Howe Island, Streimann
55832 (CANB9514159)
Australia, Lord Howe Island, Streimann
55882 (CANB9514209)
New Zealand, Braggins 05/059 (AK293730)
New Zealand, Braggins 05/116c (AK297119)
New Zealand, Braggins 95/695D
(AK286012)
New Zealand, de Lange 9994 (AK327798)
New Zealand, Engel 20862 (F1141137)
New Zealand, Engel 22174 (F1141650)
New Zealand, Engel 22604 (F1141589)
New Zealand, Engel 22661 (F1141192)
New Zealand, Engel 23329 (F1141186)
New Zealand, Engel 23639 (C0311971F)
New Zealand, Salter s.n. (AK322871)
Australia, Lord Howe Island, Brown 00/35
(NSW444725)
French Guiana, Holz FG 32 (GOET)
Nepal, Long 17359 (GOET)
Costa Rica, Heinrichs et al. 4175 (GOET)
Costa Rica, Heinrichs et al. 4146 (GOET)
Australia, Queensland, Renner 6917
(NSW855511.1)
DQ194062*
DQ194021*
DQ194196*
AJ866744*
AJ633128*
AJ416646*
AJ866734*
KY051539
AJ866763*
AJ866766*
AY438229*
AJ866759*
KY051302
DQ194197*
–
DQ194198*
DQ194099*
KY051050
KY051525
KY051290
KY051036
KY051529
KY051294
KY051040
KY051533
–
KY051041
KY051530
KY051295
KY051042
KY051531
KY051296
KY051043
KY051532
KY051297
KY051044
KY051536
KY051299
KY051047
KY051537
KY051300
KY051048
KY051528
KY051293
KY051039
KY051538
KY051301
KY051049
KY051526
KY051291
KY051037
KY051527
KY051292
KY051038
KY051534
–
KY051045
KY051535
KY051298
KY051046
KY051431
KY051186
KY050936
KY051432
KY051187
KY050937
KY051414
KY051415
KY051352
KY051169
KY051170
KY051100
KY050920
KY050921
KY050851
KY051418
KY051475
KY051439
KY051438
KY051354
–
KY051313
KY051353
KY051442
KY051173
KY051228
KY051194
KY051193
KY051103
KY051102
KY051061
KY051101
–
KY050923
KY050976
KY050944
KY050943
KY050854
KY050853
KY050814
KY050852
KY050947
AY275174*
AY550145*
AJ416084*
AJ422026*
KY051540
AY438224*
AY547726*
AY438230*
AY438231*
KY051248
DQ194199*
DQ194200*
DQ194201*
–
KY050995
Plagiochila streimannii Inoue
Plagiochila stricta Lindenb.
Plagiochila stricta
Plagiochila strictifolia Steph.
Plagiochila strombifolia (Taylor) Taylor
Plagiochila strombifolia
Plagiochila strombifolia
Plagiochila strombifolia
Plagiochila strombifolia
Plagiochila strombifolia
Plagiochila strombifolia
Plagiochila strombifolia
Plagiochila strombifolia
Plagiochila strombifolia
Plagiochila strombifolia
Plagiochila strombifolia
Plagiochila strombifolia
Plagiochila strombifolia
Plagiochila strombifolia
Plagiochila subflabellata Colenso
Plagiochila subflabellata
Plagiochila subflabellata
Plagiochila subflabellata
Plagiochila subflabellata
Plagiochila subflabellata
Plagiochila subflabellata
Plagiochila subflabellata
Plagiochila subflabellata
Plagiochila subflabellata
Plagiochila subflabellata
Plagiochila subflabellata
Plagiochila subflabellata
Plagiochila subflabellata
Plagiochila subplana Lindenb.
Plagiochila subtropica Steph.
Plagiochila superba (Nees) Mont. & Nees
Plagiochila tabinensis Steph.
Plagiochila teysmannii Sande Lac.
(continued next page)
104
Australian Systematic Botany
M. A. M. Renner et al.
Appendix 1. (continued )
Species
Voucher
nrITS
rps4
rbcL
Plagiochila teysmannii
Australia, Queensland, Renner 6917
(NSW855511.2)
Australia, Queensland, Renner 6930 (NSW)
Indonesia, Gradstein 10308 (GOET)
Australia, Queensland, Renner 6511
(NSW898713)
Japan, Kurita 257 (HIRO)
Costa Rica, Heinrichs et al. 4324 (GOET)
Australia, Queensland, Renner 6900
(NSW858853)
Australia, Queensland, Renner 6915
(NSW858855)
Australia, Queensland, Renner 6952
(NSW859321)
Australia, Queensland, Renner 6954 (NSW)
Australia, Queensland, Renner 6955
(NSW858653)
Australia, Queensland, Renner 7075
(NSW858867)
Australia, Queensland, Renner 7113
(NSW853008)
Australia, Queensland, Renner 6899
(NSW858850)
New Zealand, Braggins 04/067 (AK290513)
New Zealand, Braggins 13/279 (NSW)
New Zealand, de Lange 10631 (AK330919)
New Zealand, de Lange 10649 (AK330937)
New Zealand, de Lange 9705 (AK322390)
New Zealand, de Lange CH1666 (AK303347)
New Zealand, Engel 20933 (F1141705)
New Zealand, Engel 22686 (F1141703)
New Zealand, Glenny I37 (NSW)
New Zealand, Braggins 04/015A (AK286177)
Ecuador, Holz 070 E/5-01 (GOET)
Bolivia, Heinrichs et al. 4058 (GOET)
Costa Rica, Heinrichs et al. 4331 (GOET)
Australia, Queensland, Renner 6492
(NSW909521)
Australia, Queensland, Renner 6541
(NSW899754)
Fiji, Renner 5448 (NSW889656)
New Zealand, Renner 4189 (F)
–
KY051249
KY050996
–
AJ866745*
–
KY051246
AJ866762*
KY051250
KY050993
DQ194203*
KY050997
AY550146*
AJ416082*
KY051482
AY547727*
AY547728 *
KY051238
DQ194204*
–
KY050985
KY051483
KY051239
KY050986
KY051486
KY051242
KY050989
KY051490
KY051479
KY051247
KY051235
KY050994
KY050982
KY051484
KY051240
KY050987
KY051478
KY051234
KY050981
KY051481
KY051237
KY050984
KY051324
KY051329
KY051327
KY051328
KY051326
KY051325
KY051332
KY051331
KY051333
KY051323
AJ422024*
DQ194064*
AY275175*
KY051549
KY051072
KY051077
KY051075
KY051076
KY051074
KY051073
KY051080
KY051079
KY051081
KY051071
AY547729*
DQ194023*
DQ194024*
KY051311
KY050823
KY050828
KY050826
KY050827
KY050825
KY050824
KY050831
KY050830
KY050832
KY050822
DQ194206*
DQ194207*
DQ194208*
KY051059
KY051548
KY051310
KY051058
KY051547
–
KY051309
KF851433*
KY051057
KF852336*
KT992470
KT992619
KT992547
KT992469
KT992618
KT992546
KT992471
KT992620
KT992548
KT992474
KT992623
KT992551
KT992475
KT992624
KT992552
KT992472
KT992621
KT992549
KT992468
–
KT992545
KT992473
KT992622
KT992550
Plagiochila teysmannii
Plagiochila teysmannii
Plagiochila teysmannii
Plagiochila trabeculata Steph.
Plagiochila trichostoma Gottsche
Plagiochila trigona Steph.
Plagiochila trigona
Plagiochila trigona
Plagiochila trigona
Plagiochila trigona
Plagiochila trigona
Plagiochila trigona
Plagiochila trigona
Plagiochila trispicata Colenso
Plagiochila trispicata
Plagiochila trispicata
Plagiochila trispicata
Plagiochila trispicata
Plagiochila trispicata
Plagiochila trispicata
Plagiochila trispicata
Plagiochila trispicata
Plagiochila trispicata
Plagiochila turgida Herzog
Plagiochila validissima Steph.
Plagiochila vincentina Lindenb.
Plagiochila vitiensis Mitt.
Plagiochila vitiensis
Plagiochila vitiensis
Pseudolophocolea denticulata R.M.Schust. &
J.J.Engel
Tetracymbaliella cymbalifera (Hook.f. &
Taylor) Grolle
Tetracymbaliella cymbalifera
Tetracymbaliella subsimplex (Austin)
J.J.Engel
Tetracymbaliella subsimplex
Tetracymbaliella subsimplex
Tetracymbaliella subsimplex
Tetracymbaliella subsimplex
Tetracymbaliella subsimplex
New Zealand (I), South Island, Renner 6139
(NSW895440)
New Zealand (II), South Island, Renner 6104
(NSW895384)
Australia, Tasmania (I), Renner 5928 & Brown
(NSW895282)
Australia, Tasmania (III), Renner 5960
(NSW880807)
Australia, Tasmania (IV), Renner 5930 &
Brown (NSW895256)
Australia, Tasmania (V), Renner 5896 &
Brown (NSW892110)
Australia, Tasmania (VI), Renner 5946
(NSW880770)
Australia, Tasmania (II), Renner 6009 &
Brown (NSW880772)
www.publish.csiro.au/journals/asb
Alfons Schäfer-Verwimp