Bull. Natn. Sci. Mus., Tokyo, Ser. B, 31(3), pp. 109–115, September 22, 2005

Mosses from Pakistan Collected by Botanical Expedition of National
Science Museum, Tokyo in 1990. 3. Orthotrichaceae
Masanobu Higuchi
Department of Botany, National Science Museum,
Amakubo 4–1–1, Tsukuba, 305–0005 Japan
E-mail: higuchi@kahaku.go.jp

Abstract Three genera and ten species in the family Orthotrichaceae are recognized based on thecollections from Pakistan in 1990. Among them Orthotrichum diaphanum is newly recorded from
Pakistan. A key to the species of Orhtotrichaceae in Pakistan is provided.
Key words : bryophytes, mosses, Orthotrichaceae, Pakistan.

This paper deals with the orthotrichaceous
mosses collected in Pakistan from July to September 1990 (cf. Higuchi, 1992). The specimens
examined are deposited in the herbarium of National Science Museum, Tokyo (TNS), and someduplicates in the herbarium of Pakistan Museumof Natural History (PMNH). A key to the speciesof Orhtotrichaceae in Pakistan is provided. For
each species, collection localities, specimen
numbers, distribution and some morphologicaland taxonomical notes are also provided.
Orthotrichaceae
Higuchi & Nishimura (2003) lists three genera
and eight species, Amphidium (1 sp.), Drummondia (1 sp.) and Orthotrichum (6 spp.), in the family Orthotrichaceae as occurring in Pakistan.
Lewinsky (1992) reported that Orthotrichum
affine Brid., O. pallens Brid. and O. striatum
Hedw. occurred on Pakistan, and Lewinsky(1993) noted that O. crenulatum Mitt. was also
recorded from Pakistan. Recently Schäfer-Verwimp & Gruber (2002) added Orthotrichum obtusifolium Brid. to the moss flora of Pakistan.
The Orthotrichaceae presently known from Pakistan includes 13 species in three genera. By theexamination of forty eight specimens collected,
ten species in three genera were recognized.
Among them Orthotrichum diaphanum Schrad.

ex Brid. is reported for the first time from Pakistan.
Key to the genera of Orthotrichaceae in Pakistan
1. Stems creeping, freely branched, with numerous, erect branches; capsules not sulcate,smooth. . . . . . . . . . . . . . . . . . . Drummondia
1. Stems erect or ascending, simple or sparsely
branched; capsules strongly sulcate whendry. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Leaves usually strongly crisped when dry;calyptra cucullate; capsules without peristome. . . . . . . . . . . . . . . . . . . Amphidium
2. Leaves appressed, straight, rarely slightly
contorted when dry; calyptra mitrate or
campanulate; capsules with peristome. . .
. . . . . . . . . . . . . . . . . . . . . . . Orthotrichum
Drummondia thomsonii Mitt., J. Linn. Soc. Bot.
Suppl. 1: 46, 1859.
Specimens examined. N. W. Frontier Prov.:
Swat Valley, Utrot, 2250 m, on trunk of Quercus
dentata, Aug. 15, 1990, Higuchi 19720, 19725;
Kaghan Valley, Shinu, 1450 m, on tree trunk,
Sept. 2, 1990, Higuchi 20230, 20232.
Distribution. Afghanistan, West Pakistan,
Kashmir, northwestern India and Tibet (cf. Vitt,
1972).
Notes. Although Vitt (1972) noted that this110

Masanobu Higuchi

Fig. 1. Calyptrae of Amphidium lapponicum (1–4 taken from Higuchi 19643) and A. cyathicarpum (5–8 taken
form Deguchi 25773 in TNS). 1, 5. Calyptrae when wet. 2, 6. Upper parts of calyptra. 3. Upper margin of calyptra. 4, 8. Basal margins of calyptra. 7. Basal cells of calyptra. Scale bars for 1 & 50.5 mm; 2 & 6
100 m m; 3, 4, 7, 850 m m.

species seems to grow most often on saxicolous
substrates, it grows on tree trunk or rarely on
boulder in Pakistan (cf. Nishimura et al., 1993).
The plants (Higuchi 20230) bear many young
sporophytes which are applicable to the ECIstage in Greene (1960). Sporophytes of thisspecies seem to be matured in late autumn in thenorthern Pakistan.

Amphidium lapponicum (Hedw.) Schimp., Coroll.
39, 1856.
Specimens examined. N. W. Frontier Prov.:
Swat Valley, Gabral, 2300 m, on rock-cliff, Aug.
13, 1990, Higuchi 19643; Matiltan, 2450 m, onrock crevice, Aug. 11, 1990, Higuchi 19591;
Lake Mahodand (north of Matiltan), 2900 m, onrock crevice, Aug. 10, 1990, Higuchi 19554.
Distribution. Widely distributed in the Northern Hemisphere.

Mosses of Pakistan

Notes. Lewinsky (1976) firstly observed papillae on the surface of the calyptra of Amphidium
cyathicarpum, and pointed out that the occurrence of papillose calyptra in Amphidium indicates relationship to the Orthotrichaceae rather
than to the Dicranaceae (Rhabdoweisiaceae). I
also observed papillose calyptra in the plants(Higuchi 19643) of A. lapponicum. In Amphidium lapponicum the papillae are restricted on theupper part of the calyptra (Fig. 1: 3, 4), while inA. cyathicarpum they are on the lower part(Fig. 1: 7, 8).
Twelve species of Orthotrichum including O.
diaphanum newly added here are known from
Pakistan. The following key is revised from
Lewinsky (1992, 1993) and Schäfer-Verwimp &Gruber (2002). The species with asterisk (*) are
not present in this collection.
Key to the speices of Orthotrichum in Pakistan
1. Leaves with a hair-point of hyaline cells. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . O. diaphanum
1. Leaves without hyaline hair-points. . . . . 2
2. Stomata immersed. . . . . . . . . . . . . . . . . . 3
2. Stomata superficial. . . . . . . . . . . . . . . . . . 8
3. Exostomes erect-spreading when dry; preperistomes present; most commonly growing on
rocks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3. Exostomes reflexed, recurved or revolute
when dry; preperistomes present or absent;
most commonly growing on tree trunks. . . . 5
4. Capsules exserted on a long seta. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . O. anomalum
4. Capsules immersed to emergent; setae
short. . . . . . . . . . . . . . . . . . . . O. cupulatum
5. Calyptra with papillose hairs reaching to thetop. . . . . . . . . . . . . . . . . . . . . . . . . . . O. alpestre
5. Calyptra naked. . . . . . . . . . . . . . . . . . . . . . . . 6
6. Leaves broadly obtuse, rounded acute or
acute; margins crenulate near the apex. . . .
. . . . . . . . . . . . . . . . . . . . . . . O. crenulatum
6. Leaves acute or rouned acute; margins entire or very rarely sharply dentate near theapex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Segments 16, well developed; leaves lanceo-

111

late to oblong-lanceolate; stomata almost uncovered by subsidiary cells. . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . O. pallens*
7. Segments 8, or rarely with 8 rudimentary segments in between; leaves ovate-lanceolate;stomata half to almost completely covered bysubsidiary cells. . . . . . . . . . . . . . O. pumilum*
8. Plants dioicous; leaf margins erect-incurved to incurved; leaf apices broadly obtuse. . . . . . . . . . . . . . . . . . O. obtusifolium*
8. Plants not with the combination of characters mentioned above. . . . . . . . . . . . . . . . 9
9. Exostomes erect to spreading when dry. . . 10
9. Exostomes reflexed, recurved or revolute
when dry. . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10. Capsules immersed to emergent, oblong,often furrowed when dry; endostomeoften missing; calyptra with hairs reaching over the top. . . . . . . . . . . . O. rupestre
10. Capsules long exserted, cylindric, smoothwhen when dry; endostome well developed; calyptra with hairs reaching over the
top . . . . . . O. laevigatum var. japonicum
11. Endostome of 16 lanceolate segments. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . O. striatum
11. Endostome of 8 triangular segments. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . O. affine*Orthotrichum alpestre Hronsch. ex B. S. G.,
Bryol. Eur. 3: 75, 1849.
Specimens examined. N. W. Frontier Prov.:
Swat Valley, Gabral, 2300 m, on boulder, Aug.
13, 1990, Higuchi 19631 (Orthotrichum laevigatum); Matiltan, 2400 m, on tree trunk, Aug. 7,
1990, Higuchi 19431, 19434; 2450 m, Aug. 11,
1990, Higuchi 19586; Kaghan Valley, Kamal Ban
Forest (southwest of Kaghan), 2050 m, on tree
trunk, Sept. 2, 1990, Higuchi 20176; Naran, 2430
m, on trunk of Juglans sp., Aug. 26, 1990,
Higuchi 19828 (Orthotrichum crenulatum);
Batakundi, 2600 m, on rock-cliff, Aug. 26, 1990,
Higuchi 19864 (Orthotrichum cupulatum). Mt.
Nanga Parbat, Rupal–Nanga Parbat Base Camp,
3330 m, on tree trunk, Sept. 13, 1990, Higuchi
20369; 3400 m, on boulder, Higuchi 20406.
Distribution. Western North America, Green-

112

Masanobu Higuchi

land, Svalbard, North & Central Europe, Siberia,
northern Pakistan and northern India (cf. Lewinsky, 1992, 1993).

sky, 1992, 1993).

Orthotrichum anomalum Hedw., Spec. Musc.
162, 1801.
Specimens examined. N. W. Frontier Prov.:
Swat Valley, Matiltan, 2100 m, on boulder, Aug.
9, 1990, Higuchi 19502; 2250 m, on boulder,Aug. 7, 1990, Higuchi 19413; Matiltan – Lake
Mahodand, 2250 m, on rock-cliff, Aug. 8, 1990,
Higuchi 19450; Desan Forest (south of Utrot),
2400 m, on rock-cliff, Aug. 14, 1990, Higuchi
19704; Kaghan Valley, Kawai, 1600 m, on boulder, Sept. 1, 1990, Higuchi 20119, 20137; Sharan, 2370 m, on boulder, Sept. 3, 1990, Higuchi
20286. Mt. Nanga Parbat, Nanga Parbat BaseCamp, 4450 m, on boulder, Sept. 18, 1990,
Higuchi 20522.
Distribution. North & Central America, Haiti,
Europe, North Africa, North & Central Asia,
northeastern Afghanistan, northern Pakistan,
India, Tibet, northern Yunnan, central East China
and Japan (cf. Lewinsky, 1992, 1993).

Orthotrichum cupulatum Hoffm. ex Brid., Musc.
Rec. 2(2): 25, 1801.
Specimens examined. N. W. Frontier Prov.:
Swat Valley, Matiltan, 2250 m, on boulder, Aug.
11, 1990, Higuchi 19575; Kaghan Valley, Naran,
2400 m, on rock-cliff, Aug. 25, 1990, Higuchi
19805; Batakundi, 2600 m, on rock-cliff, Higuchi
19864 (Orthotrichum alpestre).
Distribution. Central and western North America, Mexico, southern South America, Europe,
Caucasus, North Africa, Central Asia, Afghanistan, northern India, Turkestan, Uzbekistan,
New Zealand and southeastern Australia (cf.
Lewinsky, 1992, 1993) and Pakistan (cf. Blatter& Fernandez, 1931; Schäfer-Verwimp & Gruber,
2002).
Notes. Schäfer-Verwimp & Gruber (2002)
cited that the first and single previous record of
Orthotrichum cupulatum from Pakistan is “Blatter & Fernandez (1929).” Probably they confusedDixon (1929) with Blatter & Fernandez (1931),
which were based on the same collection.

Orthotrichum crenulatum Mitt., J. Linn. Soc.
Bot. Suppl. 1: 48, 1859.
Specimens examined. N. W. Frontier Prov.:
Swat Valley, Matiltan, 2240 m, on trunk ofJuglans sp., Aug. 7, 1990, Higuchi 19412; onrock-cliff, Higuchi 19420 (Orthotrichum laevigatum); 2250 m, on trunk of Juglans sp., Aug.
11, 1990, Higuchi 19571; Utrot, 2250 m, on
trunk of Quercus dentata, Aug. 15, 1990,
Higuchi 19727; Gabral, 2300 m, on tree trunk,
Aug. 13, 1990, Higuchi 19623; Kaghan Valley,
Shinu, 1450 m, on tree trunk, Sept. 2, 1990,
Higuchi 20231 (Orthotrichum diaphanum);
Shogran, 2200 m, on tree trunk, Aug. 31, 1990,
Higuchi 20083; Naran, 2430 m, on trunk of Populus sp., Aug. 26, 1990, Higuchi 19827; on trunkof Juglans sp., Higuchi 19828 (Orthotrichum
alpestre).
Distribution. Turkestan, Kasachstan, Tadzhikistan, northeastern Afghanistan, northern Pakistan, northern India, western Tibet (cf. Lewin-

Orthotrichum diaphanum Schrad. ex Brid.,Musc. Rec. 2(2): 29, 1801.
(Fig. 2)
Plants in small, dense or sparse turf up to
7 mm tall, brownish below, yellowish green to
dark-green above; stems simple or sometimes
branched, usually growing by innovation. Leaves
erect, appressed, somewhat contorted, reflexed at
the upper part when dry, 2.0–2.80.5–0.6 mm
long, ovate-lanceolate, gradually acuminate to a
serrulate, hyaline hair-point (which is longer inthe upper leaves); costa excurrent into the hairpoint, channeled; margins almost entire, revolutenearly to the base of the hair-point; median laminal cells 14–3410–16 m m, quadrate to rectangular, thin-walled, smooth or weakly papillose;
cells of hair-point linear to vermicular, papillose,
thick-walled; basal cells smooth and pale, rectangular toward the costa, quadrate at the margins.
Gemmae commonly present on both leaf surfaces, filamentous, rarely branched, (3–)5–8(–12)
cells long.

Mosses of Pakistan

113

Fig. 2. Orthotrichum diaphanum. 1. Leaf. 2. Apical part of leaf. 3. Exostome. 4. A part of capsule. Arrows show
stomata. 5. Immersed stoma. 6. Gemmae on leaf. 7. Gemma. All taken from Higuchi 19401. Scale bars for10.5 mm; 2 & 6100 m m; 3, 5 & 750 m m; 4200 m m.

114

Masanobu Higuchi

Autoicous. Perigonia often on separate branches. Perichaetial leaves not differentiated. Setae
seen above vaginula very short, 0.2–0.4 mm long.
Capsules immersed or emergent, 1.4–1.6 mm
long, oblong-cylindric, moderately furrowed
when dry; exothecial cells differentiated into 8
brownish bands and 8 hyaline bands; stomata immersed, below the middle part of the capsule.
Peristome double; exostome 16, lanceolate, erectspreading when young and dry, reflexed when oldand dry, densely papillose on the outer surface,densely papillose or papillose-striolate on the
inner surface; endostome segments 16, narrowlanceolate, consisting of 1 row of cells, finely papillose. Spores 14–20 m m. Calyptra not seen.
Specimens examined. N. W. Frontier Prov.:
Swat Valley, Saidu Sharif, 930 m, on tree trunk,
Aug. 6, 1990, Higuchi 19398, 19399, 19401;
Kaghan Valley, Shinu, 1450 m, on tree trunk,
Sept. 2, 1990, Higuchi 20231 (Orthotrichum
crenulatum).
Distribution. North America, Mexico,
Ecuador, Hawaii, Europe, Canaries, North, East
& South Africa and Western Asia (cf. Lewinsky,1993). New to Pakistan.
Notes. The above description is based on theplants from Pakistan. This species is characterized by having immersed or slightly emergent
capsules, immersed stomata, reflexed exostomes
when dry, linear-lanceolate endostome segments,
leaves with a hairpoint of hyaline cells and filamentous gemmae (Fig. 2). As compared withother species of Orthotrichum in Pakistan, O. diaphanum grows on tree trunk in lowland. This
species is easily recognized in the field by its
hyaline hair-point of leaves, which might cause
to confuse with pottiaceous species such as Torutula when in sterile condition.
Orthotrichum laevigatum J.E.Zetterst. var. japonicum (Z.Iwats.) Lewinsky, J. Hattori Bot. Lab. 72:
42, 1992.
Specimens examined. N. W. Frontier Prov.:
Swat Valley, Matiltan, 2240 m, on rock-cliff,
Aug. 7, 1990, Higuchi 19420 (Orthotrichum
crenulatum); Gabral, 2300 m, on boulder, Aug. 13,

1990, Higuchi 19631 (Orthotrichum alpestre).
Distribution. Northwest India, Nepal, Southeast Tibet, Japan (cf. Lewinsky, 1992) and Pakistan (cf. Schäfer-Verwimp & Gruber, 2002).
Notes. Lewinsky (1992) stated that Orthotrichum laevigatum var. japonicum differsfrom var. laevigatum by having the well developed, strongly ornamented segments and larger
spores (18–22 m m). Although the plants (Higuchi
19420) have the combination of the characterssuch as capsules long exserted, cylindric, smoothwhen dry and calyptra with hairs reaching over
the top, they also have smooth, narrow lanceolate
segments and smaller spores (12–16 m m). As discussed by Schäfer-Verwimp & Gruber (2002),
the plants from Pakistan do not show typicalform of the var. japonicum. Orthotrichum laevigatum was firstly reported from Pakistan byBrotherus (1898 as O. schlotthaueri) and Robinson (1965). Nishimura et al., (1993) also reported O. laevigatum fo. macounii from Pakistan.
Orthotrichum rupestre Schleich. ex Schwaegr.,Spec. Musc. Suppl. 1(2): 27, 1816.
Specimens examined. N. W. Frontier Prov.:
Swat Valley, Matiltan, 2100 m, on boulder, Aug.
9, 1990, Higuchi 19497 (Orthotrichum anomalum); Gabral, 2300 m, on boulder, Aug. 13,
1990, Higuchi 19627; on rock-cliff, Higuchi
19636; Kaghan Valley, Naran, 2400 m, on boulder, Aug. 25, 1990, Higuchi 19817; Batakundi,
2600 m, on rock-cliff, Aug. 26, 1990, Higuchi
19858. Mt. Nanga Parbat, Rupal, 3070 m, on
boulder, Sept. 12, 1990, Higuchi 20352.
Distribution. Western North America, SouthAmerica, South Greenland, Iceland, Europe,
North and East Africa, North and Central Asia,
Northwest India, West China, Hawaii, Southeast
Australia, New Zealand and Antarctica (cf.
Lewinsky, 1992, 1993).
Orthotrichum striatum Hedw., Spec. Musc. 163,
1801.
Specimens examined. N. W. Frontier Prov.:
Swat Valley, Utrot, 2250 m, on trunk of Quecus
dentata, Aug. 15, 1990, Higuchi 19728; Matiltan–

Mosses of Pakistan

Lake Mahodand, 2370 m, on tree trunk, Aug. 8,
1990, Higuchi 19474.
Distribution. Northwestern North America,
Europe, northern Africa, northern Pakistan,
Kashmir and Northeast China (cf. Lewinsky,
1992, 1993).
Notes. This species can be distinguished fromother species in Pakistan by its not distinctly furrowed capsules, reflexed and yellow 16 exostome
teeth and broad 16 endostome segments with irregular margins. The variation of spore size havebeen reported: 17–31 m m in diameter from North
America (Vitt, 1973), 24–28 m m from Fennoscandia (Nyholm, 1974), 31–35 m m from Southeast
Asia (Lewinsky, 1992) and 24–28 m m from Pakistan (Schäfer-Verwimp & Gruber, 2002). The
spores in the plants (Higuchi 19728) are
18–24 m m and smaller than those reported.
Acknowledgements
This study was supported in part by Grant-in
Aid (no. 13640707) from the Ministry of Education, Culture, Sports, Science and Technology,
Japan, and collection of plants was no.
02041091.
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