Journal of Bryology
ISSN: 0373-6687 (Print) 1743-2820 (Online) Journal homepage: http://www.tandfonline.com/loi/yjbr20
Description and phylogenetic investigation
of Pogonatum shevockii N.E.Bell & Hyvönen
(Polytrichaceae), a new East Asian species with a
unique leaf morphology
Neil E. Bell, Jaakko Hyvönen, Kuei-Yu Yao & Wen-Zhang Ma
To cite this article: Neil E. Bell, Jaakko Hyvönen, Kuei-Yu Yao & Wen-Zhang Ma (2017):
Description and phylogenetic investigation of Pogonatum shevockii N.E.Bell & Hyvönen
(Polytrichaceae), a new East Asian species with a unique leaf morphology, Journal of Bryology,
DOI: 10.1080/03736687.2017.1312732
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Date: 11 May 2017, At: 04:15
Description and phylogenetic investigation of
Pogonatum shevockii N.E.Bell & Hyvönen
(Polytrichaceae), a new East Asian species
with a unique leaf morphology
Neil E. Bell
1,
Jaakko Hyvönen
2,
Kuei-Yu Yao
3,
Wen-Zhang Ma
4
1
Royal Botanic Garden Edinburgh, Edinburgh, Scotland, UK, 2Botanical Museum & Department of Biosciences,
Viikki Plant Science Centre, University of Helsinki, Helsinki, Finland, 3Taiwan Endemic Species Research
Institute, JiJi, Nantou County, Taiwan, 4Herbarium, Key Laboratory for Plant Diversity and Biogeography of East
Asia, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, Yunnan, China
We describe Pogonatum shevockii from Taiwan, a new species with a number of unusual features. A single
collection from Yunnan is treated as conspecific, although further sampling is required to confirm whether
morphological and molecular differences from the Taiwanese material are sufficiently consistent to be
recognised taxonomically. The plants have a remarkable leaf structure with a 3-stratose lamina including a
central row of large hyaline cells that may be responsible for the abaxial surfaces of leaves appearing
distinctly pale-glaucous when dry. These cells often have globular inclusions that appear variously dark or
speckled yellow-brown/blue-green depending on magnification. In fresh material of Taiwanese specimens
the calyptra hairs are often distinctly green, another unusual feature in the Polytrichaceae. Molecular
phylogenetic analyses including exemplars from 75% of Pogonatum species strongly support the
monophyly of the new taxon and tentatively place it as sister to a clade including five species with
geminate apical lamellar cells, namely the exclusively Central and Southern American P. campylocarpum,
P. neglectum, P. comosum and P. procerum and the Asian P. microstomum.
Keywords: Bryophyte, Glaucous, Leaf structure, Mosses, Phylogenetics
Introduction
Pogonatum P.Beauv. is the most diverse genus within
the Polytrichaceae containing more than 50 species
(Hyvönen, 1989; Hyvönen & Wu, 1993; Koskinen &
Hyvönen, 2004). Morphologically very well circumscribed by characters of the sporophyte, it is either
monophyletic, or else includes a monophyletic group
in which all but a few of the species occur [18S data
suggest that P. urnigerum (Hedw.) P.Beauv., P. perichaetiale (Mont.) A.Jaeger, P. dentatum (Menzies ex.
Brid.) Brid. and P. japonicum Sull. & Lesq. might be
outside of a clade in which Polytrichum Hedw. is
sister to the majority of Pogonatum species, although
this is contradicted by chloroplast and mitochondrial
data; Bell & Hyvönen, 2010]. In November 2012 Jim
Shevock and the third author, accompanied on one
occasion by En-Liang Chu and with colleagues from
the Taiwan Endemic Species Research Institute
(TAIE), collected an unusual Pogonatum at three
locations in Chiayi and Nantou Counties in the
Correspondence to: Neil E. Bell, Royal Botanic Garden Edinburgh, 20a
Inverleith Row, Edinburgh EH3 5LR, Scotland, UK. Email: n.bell@rbge.ac.uk
© British Bryological Society 2017
DOI 10.1080/03736687.2017.1312732
mountains of central Taiwan. The plants were
growing at altitudes of between 2000 and 2300 m on
small, smooth-surfaced boulders on semi-exposed
forested ridges in dense hardwood-conifer forest
(Figure 1) and notably had green, hairy calyptras
and leaf surfaces that when dry were glaucous-white
abaxially and green adaxially. These specimens were
examined by the fourth author in May 2013 during a
visit to the California Academy of Sciences, and
brought to the attention of the first two authors
when they could not be referred to a named species
based on the synopsis of the genus (Hyvönen, 1989).
During subsequent excursions in 2014, 2015 and
2016, further specimens were found at similar sites in
Chiayi, Nantou and Yunlin counties (the former at
2350 m and the latter just below 1800 m).
Meanwhile, just 10 days before the Yunlin collection
was made in October 2015, the fourth author,
accompanied by Jim Shevock, Y. L. Yao and
X. L. Deng, found a very similar looking plant at
2240 m in the mountains of southern Yunnan, nearly
2000 km to the west.
Journal of Bryology
2017
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Bell et al. – Pogonatum shevockii N.E.Bell & Hyvönen (Polytrichaceae), sp. nov. from east Asia
Figure 1 Habitat and living material of Pogonatum shevockii. (A) Type locality, at 2125 m altitude on the Sun-Link-Sea Trail,
Alishan Township, Chiayi County, Taiwan. The species occurs on the boulder immediately to the left of the trail in the centre
of the picture. (B) Gametophores and young sporophytes in situ. Although the calyptrae appear yellowish here, when mature,
fresh and dry they are usually distinctly green in Taiwanese material.
Our herbarium-based morphological studies confirmed that these collections represent at least one
species new to science with a number of unique and
remarkable features, this being corroborated by
sequence level characters obtained from four of the
seven existing specimens. Here we describe Pogonatum
shevockii and typify it with one of the Taiwanese collections. The single Yunnan specimen, while very similar, is
distinct both morphologically and in terms of molecular
characters (see below) and could conceivably represent a
second closely related species. However, this cannot be
determined until more collections become available
and we currently regard it as conspecific.
Taxonomy
Pogonatum shevockii N.E.Bell & Hyvönen, sp. nov.
Type: Taiwan: Chiayi County, Alishan Township.
Along the 10 km contour route section of the
Sun-Link-Sea Trail at the saddle ridge of Lucynshan
toward Shueiyang Forest, 23°35′ 49′′ N, 120°
48′ 04.5′′ E, 8 November 2012, 2125 m, Shevock, Yao
& Chu 41562 (holotype TAIE, isotypes CAS, H [dup.
E], KUN, MO, NY).
(Figures 3, 4)
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Journal of Bryology
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Paratypes: Taiwan: Chiayi County, Alishan National
Forest Recreation Area. Along the lower ridge of the
trail to Tashan, less than 0.5 km above junction of
Mianyue and Jhushan sections of the historic Alishan
Railway. Open secondary Cryptomeria D.Don and
Chamaecyparis Spach forest. On metamorphic boulders
in filtered light, 23°31′ 40′′ N, 120°48′ 53′ E, 20 October
2016, 2350 m, Shevock, Yang, Yao & Schäfer-Verwimp
49629 (CAS, E, TAIE); Nantou County. Jhushan
Township, Sun-Link-Sea Forest Recreation Area.
Along the 4 km trail near the summit ridge of
Jinganshu Mountain, 23°38′ 29′′ N, 120°48′ 50.4′′ E, 7
November 2012, 2090 m, Shevock & Yao 41505 (CAS,
H [dup. E], NY, TAIE); Sun-Link-Sea Trail along
ridge forming boundary between Nantou and Chiayi
counties at summit of Lucynshan, 23°36′ 03′′ N, 120°
47′ 28.5′′ E, 9 November 2012, 2290 m, Shevock & Yao
41614 (CAS, H [dup. E], KUN, MO, TAIE); SunLink-Sea Trail under the saddle ridge of Lucynshan
toward Shueiyang Forest, 23°35′ 52.3′′ N, 120°
47′ 59.6′′ E, 14 November 2014, 2090 m, Yao 6092
(TAIE); Yunlin County, south route trail to Mt.
Shihbi and Jiananyun Peak on divide just below
summit
of
Jiananyun
Peak,
23°36′ 39.5′′ N,
Bell et al. – Pogonatum shevockii N.E.Bell & Hyvönen (Polytrichaceae), sp. nov. from east Asia
Figure 2 Bayesian 50% majority consensus phylogram resulting from the MrBayes analysis of the combined dataset. Numbers
above branches are posterior probabilities, numbers in italics below branches are bootstrap values for corresponding branches
of the maximally likely topology resulting from the RAxML analysis of the same dataset.
Journal of Bryology
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Bell et al. – Pogonatum shevockii N.E.Bell & Hyvönen (Polytrichaceae), sp. nov. from east Asia
Figure 3 Pogonatum shevockii N.E.Bell & Hyvönen. (A, C) Leaves. (B, D) Transverse sections of leaves. (A, B) from Shevock, Yao
& Chu 41562, Taiwan (isotype, H). (C, D) from Shevock & Yao 47925, Yunnan (E).
120°43′ 50′′ E, 10 October 2015, 1795 m, Shevock & Yao
47925 (CAS, E, TAIE); YUNNAN: Jin-Ping County,
Liang-Zi Village, Fen Shui-Ling National Nature
Reserve, Xi-Long Mountain, on granitic rock along
trail in old growth broadleaved forest, 22°40′ 10′′ N,
102°46′ 48′′ E, 30 September 2015, 2240 m, Ma, Yao &
Deng 15-7040 (CAS, E, KUN, PE, TNS).
Etymology: The species is named in honour of
James R. (Jim) Shevock in recognition of his outstanding contribution to knowledge of tropical bryophyte
diversity, particularly through his extensive fieldwork
in East Asia.
Plants small to medium sized with short unbranched
shoots developing from a persistent protenemal mat.
Well-defined polytrichoid central strand lacking.
Dioicous, male plants unknown. Female plants
mostly 4–6 mm high, comose, generally with only
8–10 mature leaves present. Perichaetial leaves
mostly not distinctly differentiated from upper vegetative leaves. Leaves when moist erecto-patent to spreading, moderately soft in appearance. Dry leaves in
different specimens ranging from erect to moderately
incurved with tips slightly curled but not clearly
twisted, to distinctly incurved and somewhat twisted
and contorted (this variation more or less inversely
correlating with lamellar height). Abaxial sides of
leaves when dry appearing distinctly glaucous-white
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Journal of Bryology
2017
to very pale green, with some darker areas (margins
and nerve as well as single longitudinal rows of cells
elsewhere) strongly contrasting. Adaxial surfaces uniformly deep green.
Leaves (2.7)3.5–5(5.5) mm long, (0.7)0.8–1.1(1.3)
mm wide at widest point. Sheathing lamina variable
in relative length, comprising 20–40% of total leaf
length, (0.8)1–1.3 mm wide at base. Nerve as visible
in sheathing lamina from above or below (150)200–
550 μm wide at insertion. Chlorophyllose and sheathing laminae well differentiated, hinge cells not clearly
differentiated. Chlorophyllose lamina obovate-lanceolate, narrowed towards junction with sheath; sheath
ovate-subtriangular to oblong. Adaxial cells of
lamina with thin walls, approximately 15–25 μm in
central part. Margin of chlorophyllose lamina ± prominently toothed from (14)13–12 of distance from base
upwards, teeth multicellular, mostly with enlarged,
pointed end-cells, these becoming spinose and often
brown-orange coloured towards leaf apex. Lamina/
nerve not toothed abaxially (occasionally some very
low projections at extreme apex). Lamellae (30)40–
52, extending over most of lamina surface, lamina
with single-stratose border (3)4–7(8) cells wide.
Lamellae 1–3(4) cells high (not including enlarged
basal cell corresponding to upper surface of multistratose part of lamina), apical cells undifferentiated. In
lateral view lamellae straight, ± smooth (not
Bell et al. – Pogonatum shevockii N.E.Bell & Hyvönen (Polytrichaceae), sp. nov. from east Asia
Figure 4 Transverse sections of leaves of Pogonatum shevockii N.E.Bell & Hyvönen. (A) From Shevock, Yao & Chu 41562,
Taiwan (holotype, TAIE). (B) From Shevock, Yao & Chu 41562, Taiwan (isotype, H); composite image from two separate
photographs. The orange-brown inclusions that can be seen clearly in one of the large hyaline cells in the centre of image (B)
are often much more abundant, and appear distinctly dark-coloured at lower magnifications.
crenate). Leaves in TS showing distinct, thickened
central nerve, relatively narrow compared to some
other ‘polytrichoid’ (Smith, 1971) taxa, this fairly
abruptly transitioning into extensive 3-stratose
lamina underlying the lamellae and extending fully
or almost fully to transition with unistratose border.
Practically only an abaxial stereid band of cells with
firm walls present, only few scattered cells with incrassate walls adaxially. Adaxial layer of 3-stratose lamina
comprising markedly enlarged chlorophyllose cells
forming bases of lamellae, 12–25 μm. Abaxial layer
comprising smaller chlorophyllose cells, 10–17(19) μm,
with ± incrassate cell walls often orange-brown in
defined central part of nerve. Central layer of generally
large (10–25 μm) cells sandwiched between adaxial
and abaxial layers, these being mostly hyaline, but
(in reasonably fresh material at least) often with
numerous agglutinated globular inclusions that
appear black under lower-power light microscopy
and speckled brown-yellow/blue-green under 100 ×
oil-immersion objective. Cells of unistratose border
mostly 8–15 μm wide viewed from above or below,
square to irregularly short-rectangular or oval.
Sheath cells long-rectangular or long-hexagonal at
Journal of Bryology
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Bell et al. – Pogonatum shevockii N.E.Bell & Hyvönen (Polytrichaceae), sp. nov. from east Asia
insertion, 40–100 μm long, becoming progressively
shorter (to 16–30 μm) towards contracted junction
with chloroplyllose lamina.
Calyptrae densely and robustly hairy, with hairs
longer than mature capsule and forming a thick
mat considerably wider than capsule itself; often distinctly green when fresh, gradually becoming goldenyellow in herbarium specimens. Sporophytes observed
in all specimens but (±) fully mature only in three,
including the holotype (Shevock, Yao & Chu 41562,
Shevock & Yao 41505, Shevock & Yao 41614). Seta
10–14 mm, pale orange-brown when mature, straight
to sinuose. Capsules ±2 mm, short-cylindrical, erect
to inclined, terete (occasionally with ±5 slight
angles), usually slightly contracted immediately
below wide mouth. Exothecium slightly scabrous
but not distinctly papillose or mamillose, outer cells
isodiametric square-hexagonal to short-rectangular
or short-hexagonal, only moderately incrassate,
22–65 × 18–40 μm. Peristome 150–215 μm high,
individual teeth 100–200 μm. Teeth ±32, double,
red-orange with yellow-brown margins when dry.
Operculum flat to slightly convex, rostrate, with
rostrum slightly curved. Spores (7.5) 9–10 μm in
diameter.
Phylogeny
Materials and methods
We assembled a matrix of 51 exemplars of Pogonatum
from 41 species (approximately 75% of currently
accepted species), comprising partial sequences of
the nuclear 18S and ITS2 regions, the chloroplast
rbcL and rps4 genes, the chloroplast rps4-trnS intergenic spacer, the chloroplast trnL-F region (including
the trnL group I intron, the trnL-F intergenic spacer
and parts of the trnL and trnF exons) and the mitochondrial nad5 gene (including the group I intron).
Most of the sequences were generated for previous
studies by the authors (e.g., Koskinen & Hyvönen,
2004; Bell & Hyvönen, 2010; Daniels et al., 2016),
with a small number being previously unpublished
(see Appendix 1). A further 14 species of
Polytrichaceae were included as outgroups along
with four exemplars of P. shevockii, resulting in a
matrix of 69 terminals in total. One or several of the
regions sampled were unavailable for most terminals
(Appendix 1) and this is likely to have negatively
affected resolution and support values. The precise
manner in which missing data affect the results of
different types of phylogenetic analyses is still
debated (e.g., Wolsan & Sato, 2010; Wiens &
Morrill, 2011; Simmons, 2012; Simmons &
Goloboff, 2014), although it is clear that this is not a
simple problem with a single solution but has many
aspects, both practical and theoretical (Wheeler,
2012). Our own experience suggests that missing data
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Journal of Bryology
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are unlikely to result in high support values for erroneous phylogenetic reconstructions using modelbased methods where sufficient informative characters
have been sampled and the data are appropriately
partitioned.
For P. shevockii all seven sampled regions were
obtained for all four exemplars, except that the 18S
region was only sequenced for a single Taiwanese
specimen. Genomic DNA was isolated using the
Invisorb spin plant mini kit (Invitek, Berlin,
Germany) and eluted DNA stored in the supplied
buffer. PCR amplifications were performed either in
50 μl reactions with 1.25 U Fermentas DreamTaq
polymerase and DreamTaq buffer (Fermentas,
Burlington, Ontario, Canada), 200 μM dNTPs and
0.3 μM of each primer, or else in 20 μl reactions with
0.15 μl of Biotaq DNA polymerase (Bioline,
London, UK), 2 μl of the supplied 10× reaction
buffer, 1 μl of 50 mM MgCl2, 0.75 μl of each primer
at 10 μM and 2 μl of dNTPs at 200 μM. Protocols
for PCR and primer sequences for all regions other
than ITS2 were as detailed in Bell & Hyvönen
(2010). For ITS2, the protocol consisted of an initial
step of 95° for 4 min, followed by 35 cycles of 94°
for 1 min, 55° for 1 min, 72° for 1 min 30 s and then
a final extension step of 72° for 7 min. Primers were
seqITS2 (Olsson et al., 2009) and ITS4bryo (Stech
et al., 2003). Sequencing of PCR products was
carried out either by Macrogen Europe (Amsterdam,
The Netherlands) or by the Genepool facility at the
University of Edinburgh using the sequencing
primers detailed in Bell and Hyvönen (2010), and
the above PCR primers in the case of ITS2.
Sequences of 18S, rbcL, rps4, the rps4-trnS intergenic
spacer, the trnL-F region and nad5 were manually
aligned following the methods described for these
same regions in Bell et al. (2015). There was little alignment ambiguity for these regions although a small
number of short areas were excluded from 18S, the
rps4-trnS spacer, the nad5 intron and the trnL-F
region as described in detail in Bell et al. (2015). The
ITS2 region was considerably more variable and alignment was accomplished using MUSCLE (Edgar, 2004),
with areas of significant alignment ambiguity subsequently excluded. Phylogenetic analysis of the concatenated matrix was conducted using maximum
likelihood (ML) and Bayesian model-based methods
in addition to parsimony as an optimality criterion.
Further separate ML analyses of the 18S region, the
ITS2 region and the concatenated chloroplast and
mitochondrial data were undertaken to screen for possible supported incongruence between either of the two
nuclear regions and the exclusively maternally inherited
chloroplast and mitochondrial data (which itself is
invariably congruent in studies of the family, e.g., Bell
& Hyvönen, 2010). Seven partitions were defined
Bell et al. – Pogonatum shevockii N.E.Bell & Hyvönen (Polytrichaceae), sp. nov. from east Asia
corresponding to each of the aforementioned regions.
MrModeltest v.2.2 (Nylander, 2004) was used to estimate the best-fitting model for each partition using
the Akaike information criterion (AIC; Akaike,
1974), although we chose not to implement models specifying both a gamma parameter (G) for rate variation
between sites and a proportion of invariant sites (I) due
to concerns that these parameters may influence each
other’s correct estimation (e.g., Mayrose et al., 2005),
or at least make a stable estimation (and thus run convergence) time consuming under Bayesian analysis.
For analysis of the concatenated datasets under both
ML and Bayesian methods the same model was
applied to all compartments, this being the one with
the highest AIC score for the majority of the regions
after I + G models were excluded.
ML analyses were conducted using RAxML v.7.4.2.
(Stamatakis, 2006) with the raxmlGUI v.1.3 front end
(Silvestro & Michalak, 2012). The ‘ML + thorough
bootstrap’ option within the raxmlGUI (RAxML
option ‘-b’ followed by an ML search) was used on
the partitioned dataset, with 50 runs and 500 replications. Bayesian phylogenetic analyses were conducted
using MrBayes v.3.1.2 x64 (Ronquist &
Huelsenbeck, 2003). Partitions were unlinked to
allow parameters other than topology to vary independently. In each analysis, three independent runs using
the default prior settings, each with five chains
(‘temp’ parameter = 0.1), were run simultaneously
for 2.5 × 107 generations with trees sampled every
1000 generations. Adequate sampling from the cold
chain at stationarity and convergence of independent
runs was assessed by checking that the average standard deviation of split frequencies was <0.01, potential scale reduction factor values were near 1.00,
effective sample sizes for each parameter were meaningful, and sampling from the posterior probability
(PP) distribution was accurate as assessed by examination of log-likelihood trace files in Tracer v.1.6.0.
The first 60% of trees (including those from the
burn-in phase) were discarded and a majority-rule
consensus tree constructed using the remaining
sample of 10,000 trees.
Parsimony analyses were performed using Nona
(Goloboff, 1994) within the WinClada shell
(Nixon, 2002), and TNT (Goloboff et al., 2003,
2008). The molecular matrix used for the modelbased analyses was supplemented with 49 morphological characters (Appendix S1) following
Koskinen & Hyvönen (2004), but with the addition
of two further characters and two character states
as follows: character 9 was supplemented with a
state (2) to describe apices typical for some species
of Polytrichum (costa excurrent, margins folded
over lamellae) and character 39 with a state (3) to
describe capsules with 4(–5) angles, while a new
character 32 described the walls of lamellar marginal
cells as either thin to firm (0), outer walls incrassate
(1) or all walls incrassate (2), and a new character 44
the position of stomata as either scattered (0) or
restricted to the apophysis/basal part of the
capsule (1). Note that these additions change the
numbering sequence of characters 32 onwards as
compared to Koskinen & Hyvönen (2004). Prior to
these analyses we used the WinClada command
‘Mop uninformative characters’. This resulted in a
matrix of 741 characters. Nona analyses were performed using processor time as a seed to randomise
the order of the terminals with the following settings:
hold 30,000 (holding a defined number of trees), 100
replications (search performed with multiple tree
bisection–reconnection algorithm mult*max*), and
hold/3 (to define the starting trees for each replication). This was supplemented by another search
with 1000 replications and the number of starting
trees defined as 20. Jackknife (Farris et al., 1997)
support values were obtained with 100 replications
of ten search replications (mult*10) and with one
starting tree per replication (hold/1). As with the
search for most parsimonious trees these analyses
used processor time as a seed for randomisation
and the tree bisection–reconnection algorithm
(max*).
Results
After exclusion of areas of significant alignment ambiguity the concatenated matrix comprised 5850 characters, distributed among the sampled regions as follows:
18S—1693, ITS2—338, nad5—1780, rbcL—619, rps4
—536, rps4-trnS spacer—356, trnL—528. The
optimal models of nucleotide substitution for each
compartment selected using the Akaike criterion
within MrModeltest (excluding I + G models, as
above) were GTR + I, HKY + G, GTR + I, GTR +
I, GTR + G, GTR + G and GTR + G respectively,
thus a GTR + G model was applied to all compartments in the combined analyses. For independent
analysis of the 18S, ITS2 and combined chloroplast
/ mitochondrial datasets in RAxML the models
applied were GTR + I, GTR + G and GTR + G
(the HKY model is not implemented in RAxML).
The only supported topological incongruence
between the results of the independent RAxML analyses (incompatible clades supported with 70% BS
values or above in both topologies) was the expected
paraphyly of Pogonatum in the 18S analysis compared
with its monophyly in the analysis of the combined
organellar data (see Bell & Hyvönen, 2010), and the
grouping of P. neesii (Müll.Hal.) Dozy with P. shevockii (84% BS) in the 18S analysis and with P.
inflexum Sande Lac. (99% BS) in the chloroplast and
mitochondrial analysis. However, as P. neesii has an
Journal of Bryology
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Bell et al. – Pogonatum shevockii N.E.Bell & Hyvönen (Polytrichaceae), sp. nov. from east Asia
extraordinarily large number of autapomorphies in the
18S region (confirmed by sequencing of multiple
exemplars), and a correspondingly long branch
(more than six times longer than any other branch in
the 18S topology and nearly 10 times longer than the
next longest terminal branch), this result can fairly
confidently be attributed to long branch attraction
(Felsenstein, 1978) in the 18S dataset. We thus
repeated the RAxML analysis of the 18S data with
P. neesii excluded, resulting in an otherwise identical
topology. As the incongruence of 18S with the other
data regarding the monophyly or otherwise of
Pogonatum only affects single nodes in each topology
and does not extend to relationships within the large
clade in which the majority of species occur (as also
found previously, see Bell & Hyvönen, 2010), we considered it appropriate to perform the final analyses on
the total combined data.
Topologies resulting from the ML and Bayesian analyses of the concatenated data were completely consistent, other than that Pogonatum usambaricum (Broth.)
Paris grouped with P. gracilifolium Besch. in the
Bayesian analysis (74% PP) and with P. belangeri
(Müll.Hal.) A.Jaeger using ML (BS <50%), while P.
contortum (Menzies ex Brid.) Lesq. was sister to the
P. neesii / P. inflexum clade under Bayesian (71% BS)
and to the P. aloides (Hedw.) P.Beauv. / P. nanum
(Hedw.) P.Beauv. / P. spinulosum Mitt. / P. pensilvanicum (Bartram ex Hedw.) P.Beauv. / P. brachyphyllum
(Michx.) P.Beauv. / P. tahitense Schimp. / P. semipellucidum (Hampe) Mitt. / P. tortile (Sw.) Brid. clade under
ML (BS <50%). Figure 2 shows the Bayesian majority
consensus tree with PP values as well as BS values from
the ML analysis on corresponding nodes.
Pogonatum shevockii is resolved as monophyletic
(100% PP & BS) with the three Taiwanese specimens
forming a clade (100% PP & BS) distinct from the
single Yunnan exemplar. It is tentatively placed
(98% PP, 62% BS) as sister to a clade (100% PP &
BS) of five species with geminate apical lamellar
cells [P. campylocarpum (Müll.Hal.) Mitt., P.
neglectum (Hampe) A.Jaeger, P. microstomum
(R.Br. ex Schwäegr.) Brid., P. comosum (Müll.Hal.)
Mitt. and P. procerum (Lindb.) Schimp.], four of
which are exclusively Central and Southern
American and one of which (P. microstomum) is
widespread in South and South East Asia. This
larger clade in turn is very tentatively supported
(88% PP, BS <50%) as sister to a well-supported
(100% PP, 83% BS) clade of three other SE Asian
/ Sino-Himalayan species [P. sinense (Broth.)
Hyvönen & P.C.Wu, P. fastigiatum Mitt. and P. subtortile (Müll.Hal.) A.Jaeger]. As in the 18S topology,
but contrary to the signal from the organellar data,
Pogonatum itself appears as paraphyletic with the
majority of species forming a well-supported (100%
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Journal of Bryology
2017
BS & PP) clade with a relatively long branch that
is tentatively supported (88% PP, 60% BS) as sister
to Polytrichum but excludes Pogonatum urnigerum,
P. dentatum, P. perichaetiale and P. japonicum.
The topology obtained from parsimony analysis
based exclusively on sequence level data (not
shown) was to a large extent unresolved. Several
clades are shared with the topology found in the
model-based analyses but resolution is lacking for
deeper nodes. Nonetheless, the new species was
found to be monophyletic. When the matrix was
supplemented with morphological characters 36
equally parsimonious trees of length 1857 steps
were found with a CI (consistency index) of 0.41
(Kluge & Farris, 1969) and RI (retention index) of
0.73 (Farris, 1989). The (strict) consensus of these
trees (see Supplementary Figure S1) deviates in
many respects from the one obtained in the modelbased analyses, although in all cases of incongruence
within the ingroup jackknife support values are low
or below 50%. Pogonatum shevockii is resolved in a
clade together with four diminutive Asian species
with persistent protonema [P. piliferum (Dozy &
Molk.) Touw, P. camusii (Thér.) Touw, P. neo-caledonicum Besch. and P. marginatum Mitt.], while
these together form the sister clade to the group of
five species with geminate apical cells (see above).
Discussion
The general habit of Pogonatum shevockii is reminiscent of P. microstomum, a common (and normally distinctly larger) plant of the same region. However, it is
quite distinct from all other known species of the genus
in combining lamellae that are smooth (not crenate)
when viewed laterally together with a number of
other features, including low lamellae, a narrow
nerve, undifferentiated and non-geminate lamellar
apical cells, lack of abaxial teeth and a lack of
heavily incrassate cells with rounded lumens on the
back of the lamina (as are found in P. cirratum (Sw.)
Brid. s.l., and P. convolutum (Hedw.) P.Beauv., other
species with low, smooth, non-geminate lamellae).
Furthermore the 3-stratose structure of the leaf in TS
with the abaxial surface glaucous when dry is
unknown in any other species. The green calyptrae
(when fresh) of the Taiwanese collections are also
unusual and distinctive. This is corroborated by the
molecular analyses, which strongly support monophyly and distinctness of the new taxon in a dataset
that includes sampling from around 75% of all
known species of Pogonatum (Figure 2). In parsimony
analyses including morphological characters P. shevockii is placed as sister to a group of diminutive
plants from SE Asia [recognised as section Racelopus
(Dozy & Molk.) Touw by Touw (1986)]. Features
shared with these species include persistent
Bell et al. – Pogonatum shevockii N.E.Bell & Hyvönen (Polytrichaceae), sp. nov. from east Asia
protonema, small size, absence of the well-defined
( polytrichoid) central strand, dorsal stereid band
present only in costa in the central part of the leaf
and low adaxial leaf lamellae. Whether P. shevockii
also possesses the echinoid spore surface structure
that this group shares with the genus Polytrichum is,
as yet, unknown. However, this placement is at odds
with the results of the model-based analyses of the
molecular dataset, and thus it is possible that these
shared features represent convergence of characters
associated with small plant size.
The single collection from Yunnan differs from all
of the Taiwanese specimens most notably in its lower
lamellae; excluding the enlarged basal cell in the
lamina these are mostly only one cell high (with a
few being two cells high), as opposed to 2–4 (mostly
3) cells high in the Taiwanese material). Other differences are leaves that are more strongly incurved and
twisted when dry, a generally longer leaf sheath that
is oblong and less sharply differentiated from the
chlorophyllose lamina, and slightly larger cells in
the unistratose border (12–20 μm) and the lamellae
(Figure 3). This is consistent with the strongly supported monophyly of the Taiwanese exemplars in
the molecular phylogeny (Figure 2) and the relatively
long branch separating these samples from the
Yunnanese plant, which is comparable to the degree
of molecular difference separating many other
species within the genus. However, with only a
single specimen from Yunnan available it is impossible to say at this stage whether any population level
structure within the taxon has a geographical basis,
and also whether the morphological differences
observed are consistent. The distinct morphology of
the Yunnanese plant is similar to infraspecific morphotypes that occur in other species and are unsupported by molecular data (e.g., the laxer forms of
the recently recombined Delongia cavallii (G.Negri)
N.E.Bell, Kariyawasam, Hedd. & Hyvönen; Bell
et al., 2015). Nonetheless, if increased future
sampling continued to support reciprocal monophyly
of the Yunnanese and Taiwanese populations as well
as consistent morphological differences between
them, recognition of a second species in Yunnan
would be justified.
The most distinctive macro-morphological feature
of Pogonatum shevockii is the abaxial (lower) surfaces
of the leaves appearing dramatically pale-glaucous
when dry, this contrasting with the deep green
adaxial surfaces. This is presumably related to the
unique leaf structure as seen in TS, in which an
‘extra’ layer of large, hyaline cells is sandwiched
between the layer of similar-sized adaxial chlorophyllose cells underlying the lamellae and the smaller
chlorophyllose cells forming the abaxial surface
(Figures 3–4). In other Polytrichaceae with extensive
lamellae and/or wide nerves, there may be one or
two rows of large empty cells (deuters) together with
small, heavily incrassate cells (stereids) in the multistratose central part of the nerve, but to the extent
that these can be said to extend into the rest of the
lamina (outside the multistratose zone containing stereids) they could only be equated with the large cells
immediately underlying the lamellae. Thus either
there are only two layers of cells in the extended
lamina, or else (in more robust species such as many
Polytrichum spp.) a fully differentiated nerve including
stereids may extend to most of the width of the leaf. In
P. shevockii the great majority of the lamina is uniformly 3-stratose, lacking stereids but with the
central layer composed of large hyaline cells. We
hypothesise that transmission of reflected light
through these hyaline cells is responsible for the glaucous appearance of the leaves abaxially (where they are
only obscured by a single layer of small chlorophylose
cells, whereas adaxially they are comprehensively
obscured by the lamellae). Of further note are the
agglutinated, dark, globular inclusions that can often
be seen inside these enlarged hyaline cells under lowand medium-power light microscopy and appear as
brown-yellow masses (Figure 4) with blue-green speckles under a high-power oil-immersion objective. While
we speculated that these might represent an algal or
fungal endosymbiont, in the lack of further evidence
it seems most likely that they are simply cellular
debris, a byproduct of the senescence process that produces the hyaline cells. They rather resemble dead
chloroplasts (J. Duckett, pers. comm.). It seems that
these inclusions may disappear or become more difficult to observe in some older specimens, although
this has yet to be confirmed. They are certainly deserving of further investigation.
Pogonatum shevockii represents a further addition
to the rich polytrichaceous flora of China and
Taiwan (Hyvönen & Lai, 1991), demonstrating that
even in regions that are relatively well known, with a
long history of exploration (e.g., Cardot, 1906;
Noguchi, 1934; Ihsiba, 1935), species new to science
can still be found. Pogonatum shevockii clearly
belongs within Pogonatum s. str., and based on the
results of our model-based analyses it is sister to a
clade containing P. microstomum and a number of
American species that were already considered to be
closely allied by Hyvönen (1989). The general appearance of the new species hints at this affinity, but its
undifferentiated lamellae readily distinguish it from
all other species of the group. Although currently analyses including morphological characters and using
the parsimony optimality criterion disagree with this
phylogenetic placement, we are confident that with
further sampling of both terminals and characters
this uncertainty will be resolved. While as mentioned
Journal of Bryology
2017
9
Bell et al. – Pogonatum shevockii N.E.Bell & Hyvönen (Polytrichaceae), sp. nov. from east Asia
above the overall habit of the plant closely resembles
much larger species such P. microstomum, many morphological details are shared with the group of
diminutive species previously placed in Racelopus
and allied taxa.
Biogeographically P. shevockii represents a further
link between Taiwan and the eastern Himalayas.
This type of disjunction is already known in a
number of bryophyte species as well as in other plant
groups such as conifers (Lu et al., 2001; Chung
et al., 2004; Wei et al., 2010), while recent findings in
angiosperms further strengthen the link, as exemplified by new records of Begonia grandis Dry. from
Taiwan (Nakamura et al., 2015). That these two
regions should share some biotic elements is not surprising given the high elevation of some of the
Taiwanese mountains, the great diversity of habitats
available, and the relatively recent isolation (5 Ma
approx.) of the island from the mainland (e.g.,
Sibuet & Hsu, 2004). Furthermore it seems that for
some plants at least, genetic interchange between
Taiwanese and mainland populations has continued
in spite of the considerable geographical distances
involved (Chung et al., 2004).
Online Supplementary Material
Supplementary material is available at [10.1080/
03736687.2017.1312732].
Acknowledgements
We thank Jim Shevock for kindly sending us material
of the new species as well as staff at E and H for
proving facilities and support. The Royal Botanic
Garden Edinburgh is supported by the Scottish
Government’s Rural and Environment Science and
Analytical Services Division. During 2017 we are
also grateful for the support of players of People’s
Postcode Lottery towards our scientific research.
Field work in Yunnan by the fourth author was supported by the National Geographic Society (Grant
Number 9697-15).
Taxonomic Additions and Changes: Pogonatum shevockii N.E.Bell & Hyvönen, sp. nov.
ORCID
Neil E. Bell http://orcid.org/0000-0002-2401-2968
Jaakko Hyvönen http://orcid.org/0000-0001-75598295
Kuei-Yu Yao http://orcid.org/0000-0002-3737-6832
http://orcid.org/0000-0003-3144Wen-Zhang Ma
001X
References
Akaike, H. 1974. A new look at the statistical model identification.
IEEE Transactions on Automatic Control, 19: 716–23.
10
Journal of Bryology
2017
Bell, N.E. & Hyvönen, J. 2010. Phylogeny of the moss class
Polytrichopsida (BRYOPHYTA): generic-level structure and
incongruent gene trees. Molecular Phylogenetics & Evolution,
55: 381–98.
Bell, N.E., Kariyawasam, I.U., Hedderson, T.A.J. & Hyvönen, J.
2015. Delongia gen. nov., a new genus of Polytrichaceae
(Bryophyta) with two disjunct species in East Africa and the
Himalaya. Taxon, 64: 893–910.
Cardot, J. 1906. Mousses de ĺile Formose. Beiheft zum Botanischen
Centralblatt, 19: 85–148.
Chung, J.D., Lin, T.P., Tan, Y.C., Lin, M.Y. & Hwang, S.Y.
2004. Genetic diversity and biogeography of Cunninghamia
konishii (Cupressaceae), an island species in Taiwan: a
comparison with Cunninghamia lanceolata, a mainland
species in China. Molecular Phylogenetics & Evolution, 33:
791–801.
Daniels, A.E.D., Kariyappa, K.C., Hyvönen, J. & Bell, N. 2016. The
first Indian record of Pogonatum marginatum Mitt.
(Polytrichaceae) from the Western Ghats. Bryophyte Diversity
& Evolution, 38: 41–46.
Edgar, R.C. 2004. MUSCLE: multiple sequence alignment with
high accuracy and high throughput. Nucleic Acids Research,
32: 1792–7.
Farris, J.S. 1989. The retention index and the rescaled consistency
index. Cladistics, 5: 417–9.
Farris, J.S., Albert, V.A., Kallersjo, M., Lipscomb, D. & Kluge, A.G.
1997. Parsimony jackknifing outperforms neighbor-joining.
Cladistics, 12: 99–124.
Felsenstein, J. 1978. Cases in which parsimony or compatibility
methods will be positively misleading. Systematic Zoology, 4:
401–10.
Goloboff, P.A. 1994. Nona: a Tree-searching Program. Available at:
www.cladistics.com/aboutNona.htm.
Goloboff, P.A., Farris, J.S. & Nixon, K.C. 2003. TNT: Tree Analysis
Using New Technology. Available at: www.lillo.org.ar/phylo
geny/tnt/.
Goloboff, P.A., Farris, J.S. & Nixon, K.C. 2008. TNT, a free
program for phylogenetic analysis. Cladistics, 24: 774–86.
Hyvönen, J. 1989. A synopsis of genus Pogonatum (Polytrichaceae,
Musci). Acta Botanica Fennica, 138: 1–87.
Hyvönen, J. & Lai, M.-J. 1991. A synopsis of family Polytrichaceae
(Musci) in Taiwan (China). Journal of the Hattori Botanical
Laboratory, 70: 119–41.
Hyvönen, J. & Wu, P.-C. 1993. The identity of Microdendron sinense
(Polytrichaceae). Bryologist, 96: 631–4.
Ihsiba, E. 1935. Index Muscorum Formosarum. Transactions of
Tropical Agricultural Society Taihoku Imperial University, 7:
197–204.
Kluge, A.G. & Farris, J.S. 1969. Quantitative phyletics and the evolution of Anurans. Systematic Zoology, 18: 1–32.
Koskinen, S. & Hyvönen, J. 2004. Pogonatum (Polytrichales,
Bryophyta) revisited. In: B. Goffinet, V. Hollowell, & R.
Magill, eds. Molecular systematics of bryophytes.
Monographs in systematic botany from the Missouri
Botanical Garden, vol. 98, pp. 255–69. St. Louis: Missouri
Botanical Garden.
Lu, S.Y., Peng, C.I., Cheng, Y.P., Hong, K.H. & Chiang, T.Y. 2001.
Chloroplast DNA phylogeography of Cunninghamia konishii
(Cupressaceae), an endemic conifer of Taiwan. Genome, 44:
797–807.
Mayrose, I., Friedman, N. & Pupko, T. 2005. A Gamma mixture
model better accounts for among site rate heterogeneity.
Bioinformatics, 21: 151–8.
Nakamura, K., Wang, Y.-F., Ho, M.-J., Chung, K.-F. & Peng, C.-I.
2015. New distribution record of Begonia grandis
(Begoniaceae, section Diploclinium) from Taiwan, with subspecies assignment based on morphology and molecular phylogeny. Taiwania, 60: 49–53.
Nixon, K.C. 2002. WinClada ver. 1.00.08. Available at: www.cladis
tics.com/aboutWinc.htm.
Noguchi, A. 1934. Contributions to the moss flora of Formosa.
Transactions of the Natural History Society of Formosa, 24:
289–97.
Nylander, J.A.A. 2004. MrModeltest, version 2.2. Program distributed by the author. Evolutionary Biology Centre, Uppsala
University.
Available
at:
https://github.com/nylander/
MrModeltest2
Olsson, S., Buchbender, V., Enroth, J., Hedenäs, L., Huttunen, S. &
Quandt, D. 2009. Miyabeaceae, a new family of pleurocarpous
mosses. Bryologist, 112: 447–66.
Bell et al. – Pogonatum shevockii N.E.Bell & Hyvönen (Polytrichaceae), sp. nov. from east Asia
Ronquist, F. & Huelsenbeck, J.P. 2003. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19:
1572–4.
Sibuet, J.C. & Hsu, S.K. 2004. How was Taiwan created?
Tectonophysics, 379: 159–81.
Silvestro, D. & Michalak, I. 2012. raxmlGUI: a graphical
front-end for RAxML. Organisms Diversity and Evolution, 12:
335–7.
Simmons, M.P. 2012. Misleading results of likelihood-based phylogenetic analyses in the presence of missing data. Cladistics,
28: 208–22.
Simmons, M.P. & Goloboff, P.A. 2014. Dubious resolution and
support from published sparse supermatrices: the importance
of thorough tree searches. Molecular Phylogenetics &
Evolution, 78: 334–48.
Smith, G.L. 1971. A conspectus of the genera of Polytrichaceae.
Memoirs of the New York Botanical Garden, 21: 1–83.
Stamatakis, A. 2006. RAxML-VI-HPC: maximum likelihood-based
phylogenetic analyses with thousands of taxa and mixed
models. Bioinformatics, 22: 2688–90.
Stech, M., Quandt, D., Lindlar, A. & Frahm, J.-P. 2003. The
systematic position of Pulchrinodus inflatus (Pterobryaceae,
Bryopsida) based on molecular data. Studies in austral temperate rainforest bryophytes 21. Australian Systematic Botany, 16:
561–8.
Touw, A. 1986. A revision of Pogonatum sect. Racelopus, sect. nov.,
including Racelopus Dozy & Molk., Pseudoracelopus Broth.,
and Racelopodopsis Thér. Journal of the Hattori Botanical
Laboratory, 60: 1–33.
Wei, X.-X., Yang, Z.-Y., Li, Y. & Wang, X.-Q. 2010. Molecular
phylogeny and biogeography of Pseudotsuga (Pinaceae): insights
into the floristic relationship between Taiwan and its adjacent
areas. Molecular Phylogenetics & Evolution, 55: 776–85.
Wheeler, W.C. 2012. Systematics: a course of lectures. New York:
Wiley-Blackwell.
Wiens, J.J. & Morrill, M.C. 2011. Missing data in phylogenetic
analysis: reconciling results from simulations and empirical
data. Systematic Biology, 60: 719–31.
Wolsan, M. & Sato, J.J. 2010. Effects of data incompleteness on
the relative performance of parsimony and Bayesian
approaches in a supermatrix phylogenetic reconstruction of
Mustelidae and Procyonidae (Carnivora). Cladistics, 26:
168–94.
Appendix 1
GenBank accession numbers for terminals used in the molecular
phylogenetic study, including voucher information for newly
sequenced material (asterisked). Voucher specimens are held in the
Botanical Museum of the Finnish Museum of Natural
History, University Helsinki (H) or the Royal Botanic Garden,
Edinburgh (E).
Taxon; (voucher); 18S, rbcL, rps4 / rps4-trnS intergenic spacer, trnL,
nad5, ITS2.
Hebantia rigida (Lorentz) G.L.Merr.; GU569594, GU569416,
GU569770, GU569679, GU569502, —. Oligotrichum falcatum
Steere; JQ639444, JQ639453, JQ639426, JQ639416, JQ639435, —.
Pogonatum aloides (Hedw.) P.Beauv. GU569615, GU569437,
GU569791, GU569703, GU569526, —. Pogonatum belangeri
(Müll.Hal.) A.Jaeger; GU569616, GU569438, GU569792,
GU569704, GU569527, AY396474. Pogonatum brachyphyllum
(Michx.) P.Beauv.; —, —, AY396515, AY396497, —, AY396473.
Pogonatum campylocarpum (Müll.Hal.) Mitt.; GU569617,
GU569439, GU569793, GU569705, GU569528, AY396455.
Pogonatum camusii (Thér.) Touw; —, KU852692, KU852695,
KU852698, —, —. Pogonatum cirratum (Sw.) Brid. (Borneo 1);
GU569618, GU569440, GU569794, GU569706, GU569529, —.
Pogonatum cirratum (Borneo 2); GU569619, GU569441,
GU569795, GU569707, GU569530, —. Pogonatum cirratum
(Taiwan); AY126966, AY118246, AY137691, AF545018,
AY137733, —. Pogonatum comosum (Müll.Hal.) Mitt.; —, —,
AY396503, AY396481, —, AY396446. *Pogonatum congolense
Cardot (1); Shevock 40093, São Tomé and Príncipe, H;
KY793641, KY793619, KY793653, KY793628, KY793646, —.
*Pogonatum congolense (2); Shevock 40161, São Tomé and
Príncipe, H; KY793642, KY793620, KY793654, KY793629,
KY793647, —. Pogonatum contortum (Menzies ex Brid.) Lesq.;
AY126967, AY118247, AF208425, AF545019, AY137734, —.
Pogonatum convolutum (Hedw.) P.Beauv.; GU569620, GU569442,
GU569796, GU569708, GU569531, —. Pogonatum dentatum
(Menzies ex Brid.) Brid.; GU569621, GU569443, GU569797,
GU569709, GU569532, AY396454. Pogonatum fastigiatum Mitt.;
—, —, —, —, —, AY396447. Pogonatum gracilifolium Besch.; —,
—, AY396507, AY396488, —, AY396462. Pogonatum inflexum
(Lindb.) Sande Lac.; —, —, —, AY396486, —, AY396459.
Pogonatum japonicum Sull. & Lesq.; GU569622, GU569444,
GU569798, GU569710, GU569533, AY396463. Pogonatum macrophyllum Dozy & Molk.; GU569623, GU569445, GU569799,
GU569711, GU569534, AY396443. Pogonatum marginatum Mitt.;
—, KU852693, KU852696, KU852699, —, —. Pogonatum microstomum (R.Br. ex Schwägr.) Brid.; GU569624, GU569446,
GU569800, GU569712, GU569535, —. Pogonatum nanum
(Hedw.) P.Beauv.; —, —, AY396506, AY396484, —, AY396456.
Pogonatum neesii (Müll.Hal.) Dozy (1); GU569625, GU569447,
GU569801, GU569713, GU569536, AY396449. *Pogonatum
neesii (2); Zündorf 26860, Georgia, H; KY793643, KY793621,
KY793655, KY793630, —, —. *Pogonatum neesii (3); Zündorf
26054, Georgia, H; KY793644, KY793622, KY793656,
KY793631, —, —. Pogonatum neglectum (Hampe) A.Jaeger; —,
—, AY396510, AY396491, —, AY396466. *Pogonatum neo-caledonicum Besch.; Bell 07.11.08.007, New Caledonia, H; —, KY793623,
KY793657, KY793632, KY793648, AY396471. Pogonatum nipponicum Nog. & Osada; GU569626, GU569448, GU569802,
GU569714, GU569537, AY396460. Pogonatum nudiusculum Mitt.;
—, —, AY396505, —, —, AY396451. Pogonatum pensilvanicum
(Bartram ex Hedw.) P.Beauv.; —, —, —, AY396477, AY137740,
AY396442. Pogonatum pergranulatum P.C.Chen; —, —,
AY396514, AY396496, —, —. Pogonatum perichaetiale (Mont.)
A.Jaeger; GU569627, GU569449, GU569803, GU569715,
GU569538, AY396461. Pogonatum piliferum (Dozy & Molk.)
Touw; —, —, AY396512, —, —, AY396469. Pogonatum procerum
(Lindb.) Schimp.; —, —, AY396508, AY396489, —, AY396464.
Pogonatum proliferum (Griff.) Mitt.; GU569628, GU569450,
GU569804, GU569716, GU569539, AY396465. Pogonatum rufisetum Mitt.; —, —, —, AY396479, —, AY396444. Pogonatum semipellucidum (Hampe) Mitt.; —, —, AY396511, AY396492, —,
AY396467. *Pogonatum shevockii N.E.Bell & Hyvönen (Taiwan 1);
Shevock 41505, Taiwan, H; —, KY793625, KY793659,
KY793634, KY793650, KY793638. *Pogonatum shevockii
(Taiwan 2); Shevock 47925, Taiwan, E; —, KY793627,
KY793661, KY793636, KY793652, KY793640. *Pogonatum shevockii (Taiwan 3); Shevock 41562, Taiwan, H; KY793645,
KY793624, KY793658, KY793633, KY793649, KY793637.
*Pogonatum shevockii (Yunnan); Ma, Yao & Deng 15-7040,
China, E; —, KY793626, KY793660, KY793635, KY793651,
KY793639. Pogonatum sinense (Broth.) Hyvönen & P.C.Wu (1);
KP901278, KP901283, KP901289, KP901295, KP901301, —.
Pogonatum sinense (2); DQ120780, DQ120779, —, —, DQ120778,
AY396445. Pogonatum spinulosum Mitt.; AY126974, AY118254,
AY137698, AF545026, AY137741, AY396457. Pogonatum subtortile
(Müll.Hal.) A.Jaeger; —, —, —, AY396493, —, AY396468.
Pogonatum subulatum (Menzies ex Brid.) Brid. (Australia);
GU569629, GU569451, GU569805, GU569717, GU569540,
AY396453. Pogonatum subulatum (New Zealand); GU569630,
GU569452, GU569806, GU569718, GU569541, —. Pogonatum
tahitense Schimp;. —, —, —, AY396483, —, AY396452.
Pogonatum tortile (Sw.) Brid.; —, —, AY396501, AY396476, —,
AY396441. Pogonatum tubulosum Dixon; —, —, —, AY396485,
—, AY396458. Pogonatum urnigerum (Hedw.) P.Beauv. (Borneo);
GU569631, GU569453, GU569807, GU569719, GU569542, —.
Pogonatum
urnigerum
(China);
GU569632,
GU569454,
GU569808, GU569720, GU569543, —. Pogonatum urnigerum
(Europe); AF208406, AY118256, AF208426, AF545028,
AJ291554, AY396472. Pogonatum urnigerum (Taiwan); AY126970,
AY118250, AY137694, AF545022, AY137737, —. Pogonatum
usambaricum (Broth.) Paris; GU569633, GU569455, GU569809,
GU569721, GU569544, —. Polytrichastrum papillatum G.L.Sm.;
GU569652, GU569474, GU569828, GU569744, GU569567, —.
Polytrichastrum lyallii (Mitt.) G.L.Sm.; EU927331, AY118241,
AF208423, AF545011, AY137726, AY908802, —. Polytrichum
appalachianum L.E.Anderson; GU569644, GU569466, GU569820,
GU569735, GU569558, —. Polytrichum commune Hedw.;
GU569663, GU569485, AF208428, AF545035, GU569578, —.
Polytrichum ericoides Hampe; GU569664, GU569486, GU569839,
GU569755, GU569579, —. Polytrichum hyperboreum R.Br.;
GU569665, GU569487, GU569840, GU569756, GU569580, —.
Polytrichum
juniperinum
Hedw.;
EU927329,
EU927317,
Journal of Bryology
2017
11
Bell et al. – Pogonatum shevockii N.E.Bell & Hyvönen (Polytrichaceae), sp. nov. from east Asia
EU927342, GU569757, GU569581, —. Polytrichum piliferum
Hedw.; AY126981, AY118263, AY137706, AF545037, AY137752,
—. Polytrichum strictum Menzies ex Brid.; GU569666, GU569488,
GU569841, GU569758, GU569582, —. Polytrichum xanthopilum
Wilson ex Mitt.; GU569660, GU569482, GU569836, GU569752,
12
Journal of Bryology
2017
GU569575,
—.
Psilopilum
cavifolium
(Wilson)
I.Hagen; EU927330, EU927318, EU927343, GU569761,
GU569585, —. Steereobryon subulirostrum (Schimp. ex Besch.)
G.L.Sm.; AY126984, AY118265, AY137708, AF545040,
AY137755, —.