Arctoa (2007) 16: 127-132

THE GENUS PYLAISIADELPHA (PYLAISIADELPHACEAE, MUSCI) IN RUSSIAРОД PYLAISIADELPHA (PYLAISIADELPHACEAE, MUSCI) ВО ФЛОРЕ РОССИИOLGA M. AFONINA1, HIROMI TSUBOTA2 & ELENA A. IGNATOVA3
ОЛЬГА М. АФОНИНА1, ХИРОМИ ЦУБОТА2, ЕЛЕНА А. ИГНАТОВА3Abstract
The genus Pylaisiadelpha in Russia includes two species, Pylaisiadelphatenuirostris (Bruch & Schimp. ex Sull.) W.R.Buck and P. tristoviridis (Broth.) Afonina,
Tsubota & Ignatova, comb. nov. Descriptions, illustrations, comparisons with similar species and distribution maps in Russia are provided.
РезюмеРод Pylaisiadelpha в России включает два вида, Pylaisiadelpha tenuirostris(Bruch & Schimp. ex Sull.) W.R.Buck и P. tristoviridis (Broth.) Afonina, Tsubota &Ignatova, comb. nov. Даны описания, иллюстрации, сравнение с близкимивидами, карты распространения видов в России.

INTRODUCTIONThe genus Pylaisiadelpha was established byCardot (1912) for two Mexican species, and for along time was treated as an American endemic,
until Buck (1984) found that species classified inBrotherella fit well the concept of this genus. Buckincluded all the species described in Brotherella
into Pylaisiadelpha, which did not meet full acceptance, and Ando et al. (1989) and subsequentauthors (e.g. Tan & Jia Yu, 1999) demonstrated
that some species of Brotherella are distinct andthat the genus has to be retained. All the authors,
however, agreed that Buck was correct in placing
one of the most common species, Brotherellayokohamae in the genus Pylaisiadelpha.
Ando et al. (l.c.) concluded also that the North
American P. tenuirostris (Bruch & Schimp. exSull.) W.R.Buck and the East Asian P. yokohamae (Broth.) W.R.Buck can not be satisfactory distinguished and thus the former name should
be applied to the widespread East Asian species.
The idea that both species are probably closelyrelated was pointed out already by Horikawa &Seki (1960) and Crum (1965).

Recently, Tan & Jia Yu (1999) found in thecourse of revision of the family Sematophyllaceae
in China that P. tenuirostris and P. yokohamae are
not totally identical and accepted both of them. The
latter species however is out of scope of the presentpaper, because it was not found in Russia. In China P. yokohamae, as circumscribed by Tan & Jia
Yu (l.c.), is a more southern species, whereas P.
tenuirostris has broader range and it is the onlyspecies of the genus present in Heilongjiang Province that is the northernmost in China, situating
at the border with the Russian Far East.
From the other side, in the course of revisionof the genus Stereodon for Russia, Afonina (2007)
came to conclusion, that one East Asian species,
Hypnum (Stereodon) tristoviride, traditionallyclassified within Hypnum sensu lato (cf. Ando,1972, 1995), can not be placed comfortably in neither Hypnum, nor Stereodon, but in some characters, especially alar cell pattern, strong margin serration, and suberect capsule is similar to Pylaisiadelpha tenuirostris. The same idea was firstoutlined by Ando (1956, also 1995), who however retained H. trisoviride in the genus Hypnum.

1 – V. L. Komarov Botanical Institute Rus. Acad. Sci., Prof. Popov Str., 2, St. Petersburg, 197376 Russia – Россия

197376, Санкт-Петербург, ул. Проф. Попова, 2, Ботанический институт им. В. Л. Комарова РАН.
Hiroshima University, Mitsumarukoyama 1156-2,Miyajima-cho, Hatsukaichi-shi, Hiroshima 739-0543, Japan
3 – Biological Faculty, Moscow State University, Moscow 119991 Russia – Россия 119991 Москва, Московскийгосударственный университ, Биологический факультет, каф. геоботаники.
2 – Miyajima Natural Botanical Garden, Graduate School of Science,

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OLGA M. AFONINA, HIROMI TSUBOTA & ELENA A. IGNATOVAThis morphological observations fall in a very
well agreement with the molecular data obtainedby Tsubota et al. (2002, 2004), revealing Hypnum
tristoviride unrelated to any other species of theHypnum in the phylogenetic analysis. The genus
Hypnum was found polyphyletic, but most of its
species formed monophyletic groups (usually corresponding to subgeneric units of the genus), in away similar to the result of Gardiner et al. (2005),
based on other selection of taxa and different generegions.
The analyses of Tsubota et al. (2002, 2004)
clearly indicate the affinity of Hypnum tristoviride with Pylaisiadelpha tenuirostris. In the first
analysis, the single accession of Hypnum tristoviride has been found as a sister to clade of Pylaisiadelpha tenuirostris + Wijkia hornschuchii. In
the second analysis three specimens of Hypnum
tristoviride were studied and they were foundwithin a grade among three specimens of Pylaisiadelpha tenuirostris. The latter grade was a partof clade otherwise composed by species of Brotherella, Wijkia and Heterophyllium nematosum(note that Heterophyllium affine was found inanother group).
An attempt to study Hypnum tristoviride with
ITS was not successful, as PCR product with thestandard primers, both internal and external, was
not obtained (Milyutina, pers. corresp.). The samedifficulty we faced with Pylaisiadelpha tenuirostris and some other Sematophyllaceae and Pylaisiadelphaceae, but never with Hypnaceae andPylaisiaceae. This may be considered as an additional (although weak) evidence of the relationship of these taxa.
Basing on the above evidences, we transfer
Hypnum tristoviride to the genus Pylaisiadelpha.
Pylaisiadelpha Cardot, Rev. Bryol. 39: 57.
1912.
Type species: P. rhaphidostegioides (Cardot)
Cardot, = P. tenuirostris (Bruch & Schimp. exSull.) W.R.Buck
Plants small to medium-sized, in green, yellowish-green or brownish-golden mats. Stems
creeping, brownish or dark red, rather regularly pinnately branched in one plane, teretely foliate;
branches teretely to subcomplanately foliate; hyalodermis well-developed or only partial; central
strand absent. Pseudoparaphyllia lanceolate, some-times slightly branched, distinctly serrate. Stemleaves erect, straight to falcate-secund, lanceolate
to triangular-lanceolate, rather gradually longly andnarrowly acuminate, slightly or abruptly narrowed
to insertion, not plicate, concave; margins sometimes recurved below, strongly serrate above, serrulate to subentire below; costa double, short, sometimes almost lacking; laminal cells linear, withacute ends; basal cells yellow or brownish-orange
with thick, subporose walls; alar cells few, in 1 row
along the base, large and inflated, somewhat thickwalled, supra-alar cells distinct, quadrate. Branchleaves smaller, more strongly falcate to circinate,
more strongly serrate. Autoicous or dioicous. Perichaetial leaves lanceolate, subulate, almost not plicate, serrate above. Seta yellowish-red. Capsuleslightly inclined to suberect, oblong-cylindric;
operculum conic and indisctincly rostrate to distinctly longirostrate; exothecial cells non-collenchimatous, exostome teeth cross-striolate below, yellowish; endostome with rather high basal membrane, with or without cilia. Spores relatively small,
12-20 μm.
Familial placement of Pylaisiadelpha shifted from Entodontaceae (Cardot, 1912) to Sematophyllaceae (Brotherus, 1925; Buck, 1998),
Hypnaceae (Ando et al., 1989), and finally Goffinet & Buck (2004) classified this genus in thefamily Pylaisiadelphaceae.
KEY FOR IDENTIFICATION OF PYLAISIADELPHA
SPECIES IN RUSSIA
1. Leaves slightly narrowed to the insertion;
supra-alar cells (2-)3-7 at margin; endostome
cilia rudimentary or absent; autoicous; plants
usually yellow-brownish .. 1. P. tenuirostris— Leaves rather abruptly narrowed to the insertion; supra-alar cells 1-2(-3) at margin; endostome cilia usually single, occasionally rudimentary; dioicous; plants often dark-green
or yellowish-green ........... 2. P. tristoviridis
1. Pylaisiadelpha tenuirostris (Bruch & Schimp.
ex Sull.) W.R.Buck, Yushamia 1(2): 13.
1984.– Leskea tenuirostris Bruch & Schimp.
ex Sull in Gray, Man. Bot. N.U. States 668.
1848.
Brotherella tenuirostris (Bruch & Schimp. exSull.) Broth. in Engler & Prantl, Nat. Pfl.
(ed. 2) 11: 425.1925.
Figs. 1, 3

The genus Pylaisiadelpha in Russia
2 mm

1291 mm1100 μm27654

1030.5 mm891112131415

1617Fig. 1. Pylaisiadelpha tenuirostris (Bruch & Schimp. ex Sull.) W.R.Buck (from from Ussurijsky Reserve,
Ignatov & Ignatova #06-2965, MHA): 1-2 – capsules; 3-4 – habit; 5 – upper leaf cells; 6-7 – mid-leaf cells; 8 –stem transverse section; 9 – basal leaf cells; 10 – perichaetial leaf; 11-13 – stem leaves; 14-17 – branch leaves.
Scale bars: 2 mm for 1-3; 1 mm for 4; 0.5 mm for 10-17; 100 μm for 5-9.

Plants small, in soft, green, or more commonly yellow to brownish, slightly glossy mats. Stems
0.5-1.0 cm long, subpinnately branching; branches

to 5 mm long, slightly complanately foliated. Stemleaves 0.7-1.3 x 0.20-0.35 mm, erect and falcatesecund, concave, oblong-lanceolate, gradually130OLGA M. AFONINA, HIROMI TSUBOTA & ELENA A. IGNATOVA

3 100 μm22 mm

1541096111 mm
78121314Fig. 2. Pylaisiadelpha tristoviridis (Cardot) Afonina, Tsubota & Ignatova (from Vladivostok, Ignatov & al.
#06-2017, MHA): 1 – habit; 2 – capsule; 3 – stem transverse section; 4-5 – mid-leaf cells; 6 – upper leaf cells; 7-8 –basal leaf cells; 9-11 – branch leaves; 12-14 – stem leaves. Scale bars: 2 mm for 1-2; 1 mm for 9-14; 100 μm for 3-8.

long-acuminate, yellowish at base, margins ± reflexed below, serrate above; median laminal cells
linear, 50-80 x 4-6 μm, upper laminal cells shorter, 35-40 x 5 μm; alar group triangular, alar cells
oblong, moderately inflated, yellowish, with rather thick walls, in 1(-2) rows, supra-alar cells 3-7
along leaf margin, subquadrate to short rectangular, thick-walled. Branch leaves similar to stem
leaves but smaller. Autoicous. Seta smooth,brown, 10-15 mm. Capsule erect and symmetric,
1.5-2.0 mm long, cylindric or oblong-cylindric,yellowish-brown. Operculum obliquely long-ros-trate, to 0.8 mm. Exostome teeth ca. 300 μm long;endostome rudimentary; basal membrane ca. 80
μm high; cilia none. Spores 12-20 μm. Filamentose multicellular propagulae are characteristic forsome populations of this species, but they were
never found in material from Russia.
SPECIMENS EXAMINED: Amurskaya Province: Zeya Distr., Zeya State Reserve, Teplyj Creek,
7.VIII.1980 L.I. Abramova #5 (MW); same, 21.IX.1979
Petelin s.n. (MW); same, Gilyuj River, 11.VIII.1980
L.I. Abramova #81 (MW). Khabarovsk Territory:
Verkhne-Bureinsky Distr., 5 km to ESE from Tyrma

The genus Pylaisiadelpha in Russia

131Fig. 3. Distribution of Pylaisiadelpha tenuirostris (solidcircles) and P. tristoviridis (open squares) in Russia.

der species of Stereodon, such as S. pallescens(Hedw.) Mitt., but it is distinguished by erect capsules and inflated alar cells.
2. Pylaisiadelpha tristoviridis (Broth.) Afonina,
Tsubota & Ignatova, comb. nov. – Stereodon
tristoviridis Broth., Hedwigia 38: 234. 1899.
Hypnum tristoviride (Broth.) Paris, Ind. Bryol.
Suppl. 1: 214. 1900.
Figs. 2, 3settlement, 20.IX.1940 Mesherov s.n. (LE); Khekhtsirskij
Reserve, 1.IX.1979 Bondartseva s.n. (LE). Primorsky
Territory: Khasan Distr., “Kedrovaya Pad’’ Reserve,
9.X.1954 Vasil’eva s.n. (LE); Sikhote Alin Reserve, vicinity of Sibichi settlement, 7.IX.1946 M.V. Korchagina& A.A. Korchagin s.n. (LE); Slavyansk Distr., Suputinsky Reserve, IX.1961 Vasil’eva s.n.(LE); Ussurijsky Reserve, Ignatov & Ignatova #06-3456 (MHA); LozovskiReserve, 20.IX.1978, Bardunov & al. s.n. (LE); Shkotovo Distr.: upper Usmuha River, 20.IX.1946 Kolesnikov
s.n. (LE); Litovka (Khualaza) Mt., 15.IX.1995 Konovalova & al s.n. (MHA); Pidan (Livadiyskaya) Mt., Ignatov & Ignatova #06-2146 (MHA); Vicinity of Vladivostok, 27.VIII.1963 Selivanova-Gorodkova s.n. (LE);
Vladivostok Area, Okeansky Ridge, near Shamarsky
Pass, Ignatov & al. #06-2024, 06-2006, 06-2010(MHA); Partizanskij Distr., Olkhovaya Peak, Ignatov& al. #06-2845, 06-2308 (MHA). SakhalinskayaProvince: Kuril Islands, Kunashir Island, Ozernaya
Creek, Ignatov #06-1449 (MHA).

Distribution. Japan (Honshu, Shikoku, Kyushyu), Eastern North America and Central America(Mexico, Guatemala, Panama). Russia: Primorskyand Khabarovsk Territories, Amurskaya and Sakhalinskaya Provinces. It grows in forest mostly on trunkbases, rotten wood, rarely on stones and rocks.
Plants reported from South Siberia, Baikal region (Bardunov, 1969, Kazanovsky, 1991) do notbelong to the genus Pylaisiadelpha and will be
discussed elsewhere.
Differentiation. Pylaisiadelpha tenuirostrisshows some superficial resemblance with slen-Plants rather slender, in yellowish-green or
brownish-green mats. Stems 5-10 cm long, regularly and densely pinnately branching; branches
5-10 mm long, slightly complanately foliate. Stemleaves 0.8-1.5 x 0.3-0.5 mm, falcate-secund, triangular-lanceolate, rather abruptly narrowed to theinsertion, gradually longly and narrowly acuminate;
median leaf cells (50-)70-90 x 4-6 μm. Branchleaves 0.7-1.2 x 0.20-0.35 mm. Dioicous. Seta yellowish-red, to 3.2 cm. Capsule inclined to suberect,
1.2-1.5 mm long, oblong-cylindric. Operculum
short-rostrate. Exostome teeth ca. 0.40-0.43 mm
long. Basal membrane ca. 0.13 mm high; cilia 1-2,
as long as the segments. Spores 14-20 μm.
SPECIMENS EXAMINED: Primorsky Territory: Khasan Distr., “Kedrovaya Pad’’ Reserve,
16.X.1974 Bardunov s.n. (IRK, LE); Vladivostok Area, Okeansky Ridge, near Shamarsky Pass,Ignatov & al. #06-2017 (MHA).
Distribution. Japan (Hokkaido, Honshu,Shikoku, Kyushyu), Korea, Taiwan, China. In Russia this species was collected few times in Primorsky Territory. It grows in forest on rotten logs.
Differentiation. Pylaisiadelpha tristoviridis
shows some superficial resemblance with Stereodon fauriei and S. densirameus, but it is distinguished from both these species by large, inflated, somewhat thick-walled, “bubble-like”-shapedalar cells vs. smaller, thin-walled, isodiametric or
ovate alar cells, central strand absent vs. present,
sexual condition dioicous vs. autoicous.
Alar group indicates the affinity with Pylaisiadelpha tenuirostris, but superficially these species arerather distinct by characters mentioned in the key.
ACKNOWLEDGEMENTS
The work was partly supported by RFBR
grants 05-04-48705 and 07-04-00013, and НШ7063.2007.4.

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