Bull. Natl. Mus. Nat. Sci., Ser. B, 46(2), pp. 55–64, May 22, 2020

Revision and Checklist of Philonotis (Bartramiaceae, Bryophyta)
in Pakistan, with the status of P. trachyphylla Dixon & Badhw.
and note on Mnium riparium Mitt. (Mniaceae)
Timo Koponen1 and Masanobu Higuchi2,*
1Finnish-Chinese Botanical Foundation, Mailantie 109,
FI-08800 Lohja, Finland & Finnish Museum of Natural History, Botany Unit (Bryology),
P.O. Box 7 (Unioninkatu 4), FI-00014 University of Helsinki, Finland2
Department of Botany, National Museum of Nature and Science,
4–1–1 Amakubo, Tsukuba, Ibaraki 305–0005, Japan*E-mail: higuchi@kahaku.go.jp(Received 5 March 2020; accepted 25 March 2020)

Abstract The Philonotis Brid. ﬂora of Pakistan contains four species, P. falcata (Hook.) Mitt., P.
fontana (Hedw.) Brid., P. laii T.J. Kop. and P. trachyphylla Dixon & Badhw. Philonotis calcarea(Bruch & Schimp.) Schimp., P. marchica (Hedw.) Brid., P. nitida Mitt. and P. turneriana(Schwägr.) Mitt. recorded previously from the country are excluded. A key to the species and lists
of previously published representative illustrations are provided, and P. trachyphylla is illustrated.
The taxa are shortly discussed. The record of Mnium riparium Mitt. from Pakistan is M. lycopodioides Schwägr.
Key words : Bryophyta, checklist, mosses, nomenclature, ranges, taxonomy, Pakistan, Philonotis.

Introduction
This paper deals with the genus Philonotis
based mainly the collection by the second author,
a member of the Botanical Expedition to Pakistan organized by the National Museum of
Nature and Science, Tokyo, during his trips from
July to September, 1990. The investigated area
includes Swat Valley (Fig. 1), Kagahn Valley,
Mt. Nanga Parbat area and Murree area (cf.
Higuchi, 1992).
Nishimura & Higuchi (1993) and Higuchi &Nishimura (2003) listed six species of Philonotis
from Pakistan: P. calcarea (Bruch & Schimp.)© 2020 National Museum of Nature and Science

Schimp., P. falcata (Hook.) Mitt., P. fontana(Hedw.) Brid., P. marchica (Hedw.) Brid., P. nitida Mitt. and P. turneriana (Schwägr). Mitt. Of
them, P. calcarea, P. marchica and P. turneriana(syn.: P. nitida) are excluded from the Pakistan
moss ﬂora in this paper (see below). The Higuchi
collection from Pakistan contains P. falcata, P.
fontana, P. laii T.J. Kop. and P. trachyphylla
Dixon & Badhw., the latter two new to Pakistan
moss ﬂora. P. trachyphylla is a new record to
China and Nepal. We do not repeat all bibliographic references of P. falcata and P. fontana,but refer to the latest checklist of Pakistan
mosses by Higuchi and Nishimura (2003).

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Timo Koponen and Masanobu Higuchi

Fig. 1. Swat Valley, North-West Frontier Province, Pakistan 1. Upper part of Swat Valley (M. Higuchi, 1990).
2. Small stream near Lake Mahodand, Swat Valley (M. Higuchi, 1990). 3. Several mosses growing at small
stream valley near Lake Mahodand, Swat Valley (M. Higuchi, 1990). 4. Philonotis fontana growing at small
stream valley near Lake Mahodand, Swat Valley (M. Higuchi, 1990).

Key to Philonotis in Pakistan
1. Mammillae/papillae in mid-leaf either central on cell or/and at proximal cell end ..... P. trachyphylla
1. Mammillae/papillae in mid-leaf cells either at proximal or distal cell end, cells with central papillae
absent................................................................................................................................................... 2
2. Mammillae/papillae in mid-leaf at the proximal cell end .....................................................P. fontana
2. Mammillae/papillae in mid-leaf at distal cell end ............................................................................... 3
3. Leaves on stem in ﬁve ranks, carinate and falcate; basal part of leaf±triangular.................P. falcata
3. Leaves not ranked, plane or slightly concave; basal part of leaf ovate ............................................... 4[4. Plants lustrous; leaf apex piliferous (1/5 of leaf or more); basal leaf cells subquadrate to narrowrectangular; areolation opaque........................................................................................P. turneriana]
4. Plants green to brownish; leaf acuminate with atmost a short uniseriate apex; basal leaf cells elongate rectangular; areolation translucent .......................................................................................P. laii

Taxa and Specimens
The collector numbers are by M. Higuchi. The
voucher specimens are deposited in the herbariaof Finnish Museum of Natural History, Botany

Unit (H) and Department of Botany, National
Museum of Nature and Science (TNS). Abbreviation: “c.fr.”＝with sporophytes.

Revision and checklist of Philonotis (Bartramiaceae) in Pakistan

Philonotis falcata (Hook.) Mitt.
J. Proc. Proc. Linn. Soc., Bot., Suppl. 1: 62.
1859. ―Bartramia falcata Hook., Trans. Linn.
Soc. London 9: 317, 27 ﬁg. 4. 1808. ―Philonotis
fontana var. falcata (Hook.) Brid., Bryol. Univ.
2: 21. 1827. ―Lectotype (designated by Koponen 2012): Nepal, coll. Dr. Buchanan, ex herb.
Hooker, ex herb. Kew, ʻDr. Smith 1808, No.
H/2359ʼ (BM); isolectotypes: ex herb. Dawson
Turner, ex herb. Hooker (BM); ex herb. Mitten,
ex herb. Hooker (FH).
P. carinata Mitt., Trans. Linn. Soc. London,
Bot., ser. 2, 3: 164. 1891. ―Isotype: Japan. Kiga,
Miyanoshita, V.1887 J. Bisset 67 (H-BR). ―Synonymized by Dismier (1912).
Illustrations. Ochi 1962: pl. 4 (ﬁgs. A–D,
drawn from the type of P. falcata); pl. 5 (ﬁgs.
A–F, drawn from the type of P. carinata); Noguchi 1989: 569 (ﬁg. 252, A, B); Koponen 1996:
115 (ﬁg. 1); 2012: 154 (ﬁg. 1); Kürschner andOchyra 1999: 272 (ﬁg. 2); Eddy 1996: 234 (ﬁg.
489); 235 (ﬁg. 490, as P. turneriana); Koponen etal. 2012: 29 (ﬁg. 3); Koponen et al. 2019a: 79:(ﬁg. 3, J–N); 97 (ﬁg. 13 H).
Specimens examined, leg. M. Higuchi. Pakistan. North-West Frontier Prov. Besham –Dassu, 20324, 20327, on wet rock-cliff at
1000 m; Kaghan Valley, Shogran, 20025, on wet
soil at 2270 m; Sharan 20263, on wet humus at
2370 m; Swat Valley, Matiltan, 19404, on wet
boulder at 2100 m, 19410, on wet soil at 2240 m;
Lake Mahodand (N of Matiltan), 19548, on wet
soil at 2850 m, 19561, on rock-cliff at 2900 m;
Mt. Nanga Parbat. Tarashing – Rupal, 20329
c.fr., 20333, on soil at 2960 m.
Other specimens. Hindukush Expedition. NW
Indien, Rawalpindi, kleines Tälchen in d. Ebene,
XI.1935 Karsten 63 (JE, as P. nitida). Baltistan.
Karakorum Range, Hushe Valley, Hushe Village,
wet ground near springs at 10 000 ft., 4.VII.1955
G.L. Webster & E. Nasir 5977, 5983 (US, as P.
turneriana), Hushe Valley, Ghondakoro Glacier,along lateral moraine at Ghondagoro at c. 13 500
ft., 14.VII.1955 G.L. Webster & E. Nasir 6126
(US, as P. turneriana), upper Hushe Valley, Atosar Valley, mesophytic juniper woods at c. 12

57

500 ft., 17.VII.1955 G.L. Webster & E. Nasir
6161 (US, as P. turneriana). Barum Gol. Shokor
Shal, on wet ground, associated with Poa supina
Schrad., and on a slope above the camp, bothsterile, at c. 3500 m, 15. and 16.VII.1950 P. Wendelbo, s.n. (OSLO, as P. marchica). Oghi – Battagram road. N side of hill, vertical earth cliff ofmicaceous soil, dripping rock, 7.VI.1958 B.L.
Burtt B 752 (E, as P. turneriana). Swat State.
VIII.1952 Sultan Ahmed 22 (NICH-308006,
herb. Dr. A. Noguchi 35921 as P. turneriana).
The following specimens listed from Pakistan as
Philonotis calcarea in Brotherus (1898) are P.
falcata: Kashmir. Kaylee-ban, Gurais Valley, 10500 p., 1.VII.1901 Inayal Khan 2933, Bryotheca
E. Levier. Musci Indiae orientalis, curante cl. W.
Gollan lecti, det. V. F. B. (H-BR 3125 007, as P.
calcarea); Vallis Surú, Surú – Purkutte, 3200–
3400 m, 11. VI.1913 L. Borelli (H-BR 3125 004,
as P. calcarea); Vallis Brahma in jugo Nun –Kun, c. 3800 m, VII.1913 L. Borelli (H-BR 3125
001, as P. calcarea).
The taxonomy and range of Philonotis falcata
has been discussed in several papers by Koponen(1996, 2003, 2012, 2019a, Koponen et al. 2012,
Koponen et al. 2019b). It is widely distributed
from Africa through SW Asia, the Himalayas,
China and Japan to Russian Far East and to Java
Island in south. Range maps are in Koponen(1996, 2003). The range in Asia is mainly in temperate and warm temperate (meridional) vegetation zones. Records from the Paciﬁc area need tobe conﬁrmed (Koponen, in prep.). P. falcata iseasily distinguished from the other Asiatic species of Philonotis by the±triangular basal part ofthe leaf, the foliation in ﬁve ranks on the stem,
the leaf serrate to serrulate in lower half of theleaf and faintly biseriate in the apical leaf. The
basal leaf cells are more or less translucent, subquadrate to short rectangular with a papilla at the
distal cell end.
Philonotis fontana (Hedw.) Brid.
Bryol. Univ. 2: 18. 1827. ―Mnium fontanum
Hedw., Spec. Musc. 195. 1801. ―Type: Europe(not seen).

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Timo Koponen and Masanobu Higuchi

Illustrations. In many American, European and
Asiatic ﬂora works. A selection: Brotherus 1923:
358 (ﬁg. 63, F, G, perigonial leaves); Lawton
1971: pl. 116 (1–6, as P. fontana var. fontana);
Gangulee 1974: 1115 (ﬁg. 541); Smith 1978: 463(ﬁg. 223, 1–6); 2004: 652 (ﬁg. 215, 1–6); Crum
& Anderson 1981: 654 (ﬁg. 310); Noguchi 1989:
571 (ﬁg, 253 A, as P. fontana var. fontana);
Nyholm 1998: 263 (ﬁg. 222); Ignatov &Ignatova 2003: 584 (ﬁg. 418); Hallingbäck 2008:
295, 296; Ederra 2010: 268 (ﬁg. 95a–f);
Koponen et al. 2012: 38 (ﬁg. 11); 39 (ﬁg. 12); 40(ﬁg. 13); Grifﬁn 2014: 110 (as P. fontana var.
fontana).
Specimens examined, leg. M. Higuchi. Pakistan. North-West Frontier Prov. Swat Valley,
Gabral, 19650, on sandy soil at river at 2300 m;
Kaghan Valley, Batakundi, 19854, on soil at
2800 m; Lake Mahodand (N of Matiltan), 19520,
19545 c.fr., on soil and on wet soil at 2850,
Higuchi no. 19557 (coll. T. Nakaike), on rockcliff at 2900 m. Mt. Nanga Barbat. Rupal –Nanga Parbat Base Camp, 20392, 20396, on wet
humus at 3400 m; Mazeno Base Camp, 20443,
on soil at 3900 m, 20456, 20458, on soil at
3950 m.
Philonotis fontana is the most common species of Philonotis in the northern hemisphereoccurring from the arctic to cool temperate(nemoral) areas (see e.g. Koponen et al., 2012).
The southern stations are at the corresponding
vegetation zones on high mountains. The present
collections suit to this pattern, as well as thepresent substrate observations (Fig. 1: 4). It differs from most of SE Asiatic species of Philonotis in having the mammillae/papillae on leaves atthe proximal cell end and by double-crenulate
serration of the leaf margin. As to the taxonomy
and Asiatic ranges of P. seriata Mitt. and P.
tomentella Molendo having leaf cell and leaf
margin characters similar to P. fontana, see
Koponen et al. (2012).
Philonotis laii T.J. Kop.
Acta Bryolich. Asiat. 3: 137. 2010a. ―Holotype:
China. Hunan Prov.: Wulingyuan World Heri-

tage Area, Zhangjiajie, Huangshizhai. On cliffs
and in low evergreen forest along path, warm
temperate zone, alt. 910–985 m, 29°20′N,
110°25′E, on moist partially shaded cliff, collecting locality 19d, 8.X.1997 T. Koponen, S.
Huttunen & P.-C. Rao 51641 (H).
Illustations and descriptions. Tan and Ho
2008: (ﬁg. 9 and ﬁgs. on p. 78 and 79, as Philonotis hastata); Koponen 2010a: 138 (ﬁg. 1);
Koponen et al. 2019a: 81 (ﬁg. 4).
Specimens examined. Pakistan. Rawalpindi
Distr. Karot, c. 5 km W of Jhelum Valley on roadto Rawalpindi, on wet sand in an almost dry
stream bed in deep ravine with thorn scrub, 4.
IV.1973 C.C. Townsend 73/1624 (E, as P. turneriana). North-West Frontier Prov. Kaghan Valley,
Shinu, Higuchi 20223 c.fr., on rock-cliff at
1450 m.
After its description (Koponen, 2010a), thetaxonomy of Philonotis laii has been discussed
by Koponen (2019a, b, and Koponen et al.,2019a). The range map (Koponen, 2010a) showsa rather large distribution in SE Asia, and hassince then been reported from the Philippines
and Japan (Koponen, 2018), New Guinea(Koponen, 2019b) and Vietnam (Koponen et al.,2019a). The sporophytes of P. laii were previously reported from Yunnan in China andVietnam (Koponen et al., 2019a).
The diagnostic characters of Philonotis laii are
the upright stems with propagules on innovations, which later grow in length to slender
creeping shoots with miniature leaves andnumerous propagules. The young plants, inwhich the propagules have not yet developed,
can be separated from P. hastata (Duby) Wijk &Margad. having wide leaf cells in all parts ofleaves, and from P. mollis (Dozy & Molk.) Mitt.
having spreading leaves with piliferous apex andelongated, rectangular to rhomboidal and wide
cells at the basal leaf. The differences from P.
runcinata Müll.Hal. ex Ångstr. were discussed
by Koponen et al. (2019b). As to the differencewith P. turneriana, see the key and below.
Philonotis trachyphylla Dixon & Badwh. (Fig.

Revision and checklist of Philonotis (Bartramiaceae) in Pakistan

59

Fig. 2. Philonotis trachyphylla Dixon & Badwh. (Higuchi 19487). 1, 3. Leaves. Scale bar＝1 mm. 2. Ventral surface of cells at upper part of leaf. Scale bar＝50 μm. 4. Dorsal surface of cells at upper part of leaf. Scalebar＝50 μm.

2)Rec. Bot. Surv. India 12: 175. 1938. ―Holotype:
India. Kashmir, Zanskar (below Shingo-la), 15
000 ft., 28th Aug. 1928 R.L. Badhwar 931 (BM).
―Paratype: Lahul-Bhaga valley (Ramzak to
Rampozampa), 12 500 ft., R.L. Badhwar 965(BM).
At ﬁrst, the Higuchi specimens from Pakistan
were named as Philonotis yezoana Besch. & Cardot on the basis of the leaf cells with central
papilla. However, the plants looked different

from the Japanese specimens of P. yezoana inhaving, in addition to leaf cells with central
papilla, also cells with papilla at the proximal
cell end. Other differences with P. yezoana s.str.
were discovered, as well.
The Higuchi specimens and a number of similar unnamed specimens from Himalayan area
were studied and compared with 11 JapanesePhilonotis yezoana specimens (listed in Koponen
et al., 2012).
The plants in both groups have similar habit,

60

Timo Koponen and Masanobu Higuchi

plants are slender, leaves distant on stem, withovate leaf bases and acute to acuminate leaf apices. The shape of basal leaf cells is rather similar,
wide, subquadrate, rectangular or rhomboidal.
The apical leaf cells are narrow and mamillate/papillate at the distal cell end.
The differences are (1) in Philonotis yezoana
the mid-leaf cells are ± quadrate or short rectangular with the papilla at the center of the cell.
Only one specimen with some cells with papillaat the proximal cell end was found. In the Himalayan plants, mid-leaf cells are quadrate rectangular and elongate rectangular. The narrower
central leaf cells are either with a central mammilla or the mammilla/papilla is near the proximal cell end on lumen (Fig. 2: 4). Cells with central papilla are present in any part of the leaf,
also on the narrow elongate cells near leaf border
or costa. The position of the cells with central
papilla on leaves and the number of such cells
varies even on leaves from the same stem. (2)The leaf borders in P. yezoana are plane or only
slightly recurved at leaf base, the leaf border inthe Himalayan plants is strongly recurved frombase to near the apex. (3) The leaf margin is
entire or slightly serrulate or double-crenulate in
P. yezoana. The Himalyan plants have strongly
toothed margin by double-crenulate teeth. (4)The leaves of P. yezoana are plane, the leaves insome Himalayan specimens are plicate. (5) P.
yezoana has thin-walled lamina cells while theplants in the Himalayan populations have thickwalled, ﬁrm cell walls.
In 1996, in an early phase of the study of SE
Asian Philonotis (by T. Koponen) the identity ofthe type specimen of P. trachyphylla Dixon &Badhw. (in BM) remained uncertain. It was hesitatingly named as P. tomentella Molendo,
although the leaf shape is different. In 2004,
Blanka Buryová attached the identiﬁcation labelon the specimen: “Philonotis sp. ― not P. tomentella Jur.” The type specimen of P. trachyphylla
resembles P. tomentella in having the basal marginal cells elongate rectangular and having a tall
papilla at the proximal cell end. Similarly, theleaf margin is recurved and the leaf is plicate.

The leaves are broadly ovate and with a short,acute point, while the leaves in P. tomentella aregradually tapering from triangular base to acuminate apices, the narrow distal part of leaf is 1/4–
1/3 of leaf length and straight or slightly bending
to different directions (Koponen et al., 2012).
In the type of P. trachyphylla most of the cells
have the papilla at the proximal cell end, but several cells with central papilla were found. This,and the other characters such the shape of leaves,strongly recurved leaf margin, serration of theleaf margin and thick-walled lamina cells correspond to the unnamed Himalayan specimens discussed above.
The site of the papilla/mammilla on the lamina
cells and the presence of differently papillate/mammillate cells in different parts of the leaf is
one of the few stable characters by which theidentiﬁcation of Philonotis species can be done.
In the species of Philonotis section Philonotis, inaddition to P. yezoana, cells with central papilla
are occasionally present in the quadrate basal
leaf cells of P. fontana and more regularly in thebasal cells of P. seriata (Koponen et al., 2012).
The presence of cells both with central and proximal papillae/mammillae dispersed in all parts of
the leaves is a unique character of P. trachyphylla. The recurved leaf border with rough dentation and thick-walled lamina cells are otherconstant characters which distinguish it from P.
yezoana and reinforce the speciﬁc status to P.
trachyphylla.
Specimens studied. China. Yunnan. Gaoligongshan region, Gongshan Co.: Eastern slope of theGaoligongshan. Along unimproved trail fromtunnel at Heipu Pass toward lakes, headwaters of
the Pula River, 51.5 km from the city of Gongshan, 27°45′N, 98°27′E. Alpine slopes of bamboo, Salix and Betula, stream from cirque in granitic boulder ﬁeld, wet rocks in stream in sun,
3300 m, 12.IX.2013 J.R. Shevock 43015, with
Ma, W.Z., Yao, Y.L. and Fan, X.Z. (CAS, E, H,
KUN, MO, as P. yezoana in Koponen et al.
(2019b), and: Gaoling Shan Range, Gongshan
Co. at 3270–3715 m, D.G. Long 35756, 35818

Revision and checklist of Philonotis (Bartramiaceae) in Pakistan(E), J.R. Shevock 23306 (E). India. Kashmir.
Lidda Valley, Liddarwat, S-facing mountinside at
10750ʼ, 4.IX.1983 C.A. Chadwell B107 (E). Himachal Pradesh. Udaipur, Chimrat, rocks nearwaterfall at 3650 m, 10.VIII.1990 R.J.D.
McBeath 2438 (E). Sikkim. West Distr.: Dzongri,
27°28′N, 88°09′E. Small marshy valley, in cushions at margin of ﬂush, 3965 m, 16.VII.1992
D.G. Long 22633 (E, as P. yezoana in Koponen
et al., 2019b). Nepal. Sankhuwasabha Distr.
Upper valley of tributary N of Barun Khola, NE
of Pemathang Kharka, open rocky alpine valley,
in boggy ground by stream at 3950 m, 9.X.1991
D.G. Long 21084 (E, as P. yezoana in Koponen
et al. 2019b). Pakistan. North-West Frontier
Prov. Kaghan Valley, Batakundi, Higuchi 19837,
on soil at 2800; Swat Valley, Matildan – Lake
Mahodand, Higuchi 19487, on wet soil at
2800 m; Lake Mahodand (N of Matiltan),
Higuchi 19561 (coll. T. Nakaike), on rock-cliff at
2900 m.
Excluded and Doubtful
Philonotis calcarea (Bruch & Schimp.) Schimp.
Coroll. Bryol. Eur. 86. 1856. ―Bartramia calcarea Bruch & Schimp., Bryol. Eur. 4: 49. 325.
1842. ―Philonotis fontana ssp. calcarea (Bruch& Schimp.) Boulay, Musc. France 214. 1884. ―Type: Europe (not seen).
The specimens reported as Philonotis calcarea
from Pakistan by Brotherus (1898) are P. falcata,
see above. P. calcarea is mostly a European species. Easternmost localities in Asia are in Altai in
Russia (Koponen et al., 2012), Kazakhstan andKyrgyztan (Koponen, in prep.)
Philonotis marchica (Brid.) Hedw.
Bryol. Univ. 2: 23, 735. 1827. ―Mnium marchicum Hedw., Spec Musc. 196. 1801. ―Type:Europe (not seen).
Störmer (1954) recorded Philonotis marchicafrom Pakistan. The voucher specimens (leg. P.
Wendelbo, s.n.) are P. falcata (see above).
Philonotis marchica occurs in Europe, near
East in Asia and North America. Its presence in

61

Korea and Japan was excluded by Ochi (1962,1963), and its presence in China was doubted byKoponen (2010b).
According to Koponen et al. (2012), the leavesof Philonotis marchica are not as carinate as theleaves in P. falcata, have a narrower costa,
always serrate, not partly biseriate leaf margin.
The leaves in some specimens are as rows on
stem, but not in all specimens. The papilla of leaf
cells seem to be taller at the basal leaf than thepapilla in P. falcata, and the cells are narrower,
which makes the cell areolation more dim than
the translucent basal leaf in P. falcata. According
to Ochi (1962, 1963), P. falcata differs from P.
marchica having leaves more triangular, morestrongly carinate, and arranged in regular rows
on stem. Cells of leaf-blade are more conspicuously broader toward base, which is hardly narrowed.
Philonotis turneriana (Schwägr.) Mitt.
J. Proc. Linn. Soc., Bot., Suppl. 1: 62. 1859. ―Bartramia turneriana Schwägr., Spec. Musc.
Suppl. 3(1): 238. 1828 (“Turneriana”). ―Lectotype (designated by Koponen, 2009b): Nepal.
“Bartram Hookeriana Turneriana var Nepal
Hook [illegible] longiss cuspidater fol Schwaegrichen cui Hooker misit (G, herb.
Schwägrichen; also fragment in G); syntype: “cuimisit Hooker” (G, herb. Schwägrichen).
Philonotis nitida Mitt., J. Proc. Linn. Soc.,
Bot., Suppl. 1: 62. 1859. ―Bartramia nitidaWils., Kew J. Bot. 9: 370. 1857, nom. nud. ―Lectotype (designated by Koponen, 1998): NW.
Himalaya, Simla, 7 000 ped., T. Thomson 561(NY); isolectotype (BM); syntypes: N.W. Himalaya, Royle (NY, BM, ex herb. Hooker). ―Synonymized actually by Fleischer (1904) by citing
no. 561 in the list of synonyms, and by Koponen(1998).
We conﬁrmed the identity of the vouchers
cited by Noguchi (1956, Sultan Ahmed 22) andRobinson (1965, G.L. Webster & E. Nasir 5977,
5983, 6161,6126) as Philonotis turneriana from
Pakistan. They are P. falcata (see above). Also
the record by Herzog (1938, Karsten 63) as P.

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Timo Koponen and Masanobu Higuchi

nitida from Pakistan is P. falcata (see above).
Townsendʼs (1993, Townsend 73/1624) record
as P. turnerina (see above) is P. laii.
The characters of Philonotis turneriana are its
lustrous, lanceolate, plane or slightly concave, ±straight leaves with piliferous apex, and rather
opaque areolation throughout the leaf. Leaf border may be slightly revolute at basal leaf. Suchplants occur in the Himalayas and Yunnan andSichuan provinces of China and have been identiﬁed from the Philippines and Taiwan (Koponen,in prep).
For understanding the taxonomy and nomenclature of East Asiatic Philonotis, it is necessary
to understand the status of P. turneriana, since itis the second oldest taxon described from theHimalayan area (see Koponen, 1998, 2009b,
2010c). In addition to P. turneriana, P. laii, P.
mollis (Dozy & Molk.) Mitt. and P. secunda(Dozy & Molk.) Bosch & Sande Lac. havelongly acuminate leaves, and P. falcata and P.
speciosa (Griff.) Mitt. may have acuminate
leaves. P. turneriana has a piliferous leaf apex
one ﬁfth of the leaf length. P. mollis has similarly
piliferous leaves but the piliferous part of the leaf
apex is longer. The best microscopic character
separating P. mollis from P. turneriana and fromP. secunda is its lax basal areolation and its low
papillae and, therefore, transparent areolation.
The leaves in P. mollis and P. turneriana aregradually tapering while the leaves in P. secunda
taper more quickly from ovate base, and it hasoften a group of translucent, subquadrate,strongly papillate angular cells. In the forms of P.
speciosa with piliferous leaves, the piliferous
part is fragile and the apex then is truncate.
The plants most commonly named as Philonotis turneriana are forms of P. falcata with longly
acuminate leaves. The long-leaved forms of P.
falcata can be separated from P. turneriana, in
addition to ranked leaves, by carinate and ± triangular leaf base, the more strongly bending
nerve which makes the leaf falcate. The basal
leaf cell areolation in P. falcata is more lax andcells subquadrate to short rectangular, while in P.
turneriana the narrow linear to vermiculate leaf

cells extend to two lowest rows of basal cells. In
P. turneriana the basal leaf cells have prominent
papillae, which are lower in P. falcata.
The leaf shapes of Philonots turneriana and P.
laii are rather similar. P. laii is smaller and has
narrower, long acuminate but not piliferous
leaves. The leaf cell areolation has long, translucent, rectangular and narrow cells, while the cell
areolation in P. turneriana is opaque and cells
shorter. The creeping stem apices or innovations
with propagules and miniature leaves are absent
in P. turneriana.
Mniaceae
Mnium lycopodioides Schwägr.
Spec. Musc. Suppl. 2(2): 24, pl. 160, ﬁgs. 1–9.
1826. ―Lectotype (designated by Koponen2014): Nepal. Wallich (BM, no. 2510 dupl.).
The record of Mnium riparium Mitt. from Pakistan (Störmer 1954) is here corrected to Mnium
lycopodioides. M. riparium represents a dioicous
population of Mnium marginatum (Dicks.) P.
Beauv. It has once been found in Britain(Koponen, 1980).
AcknowledgmentsCollection of plants was supported in part by
Grant-in Aid (no. 02041091) from the Ministryof Education, Culture, Sports, Science and Technology, Japan. The curators of the herbaria (BM,E, JE, NICH, OSLO, US) are thanked for forwarding specimens on loan to our study.
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