Bull. Natl. Mus. Nat. Sci., Ser. B, 33(3 · 4), pp. 115–121, December 21, 2007

Studies on the bryophyte flora of the Tien Shan Mountains. 1.
Hypnaceae and Hylocomiaceae (Musci)
Masanobu Higuchi
Department of Botany, National Museum of Nature and Science,
Amakubo 4–1–1, Tsukuba 305–0005, Japan
E-mail: higuchi@kahaku.go.jp

Abstract Eleven species in three genera in Hypnaceae and four species in four genera in Hylocomiaceae are reported based on the collections made in the Tien Shan Mountains in 1995. Hypnum
callichroum and H. subimponens subsp. subimponens are new records for the moss flora of Kazakhstan, and Hypnum cupressiforme var. subjulaceum and H. recurvatum are new to Kyrgyzstan. A
key for the Hypnum species in Kazakhstan and Kyrgystan is shown.
Key words : Hypnaceae, Hylocomiaceae, mosses, Tien Shan Mountains, Russia, Kazakhstan,
Kyrgyzstan

In 1995 a botanical survey led by Dr. Yuichi
Kadota of National Science Museum, Tokyo, wascarried out in the western part of the Tien Shan
Mountains in collaboration with the Central
Siberian Botanical Garden, Novosibirsk (cf.
Kadota 1995). In July–August 1995 I collectedbryophyte specimens as a member of the survey.
Recently Ignatov et al. (2006) compiled thechecklist of the mosses of East Europe and North
Asia. The moss flora of Central Asia is insufficiently known.
Study area
The Tien Shan Mountains are a big mountainsystem in Central Asia and are situated along theborder area between the former USSR countries
and China. The botanical survey was made in the
Dzhungarsky Alatau, the Zailisky Alatau, theTereskey Alatau, the Central Tien Shan, theKirghisky, the Alaisky and the Za Alaisky Range(Fig. 1), and locations are described below. The
specimens examined are kept in the herbarium of
the Department of Botany, National Museum of
Nature and Science (TNS), and the duplicates in
the herbarium of the Central Siberian Botanical
Garden (NS).

Kazakhstan
Dzhungarsky Alatau (45°13N, 80°20E)
Zailisky Alatau (43°02N, 77°00E)
Kyrgyzstan
Tereskey Alatau (42°15N, 78°30E)
Kirghisky (42°34N, 73°36E)
Alaisky (39°35N, 71°58E)
Za Alaisky Range (39°25N, 72°08E)
This paper deals with taxa of the families Hypnaceae and Hylocomiaceae occurring in the Tien
Shan Mountains based on the collections madeby Higuchi in 1995. By the examination of thespecimens, eleven species in three genera andfour species in four genera were recognized in
Hypnaceae and Hylocomiaceae, respectively.
Hypnum callichroum and H. subimponens subsp.
subimponens are new additions to the moss floraof Kazakhstan, and Hypnum cupressiforme var.
subjulaceum and H. recurvatum are new to Kyrgyzstan.

Hypnaceae
Although the recent molecular phylogenetic
analyses indicate that the Hypnaceae as well as
its type genus Hypnum itself were polyphyletic

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Masanobu Higuchi

Fig. 1. Map of the surveyed areas showing the route and collecting sites (revised from Kadota 1995). Solid triangles indicate main base camps.

(e.g., Tsubota et al. 2002), the delimitation of thefamily is still controversial. Here I follow thesystem of Hypnaceae by Goffinet and Buck
(2004) with some changes and Hypnum by a series of study by Ando respectively.

1. Callicladium haldanianum (Grev.) H. A.
Crum, Bryologist 74: 167 (1971).
Specimen examined. Kazakhstan, Mts. Dzhungarsky Alatau, Maly Baskan, 2480 m, July 21,
1995 (Higuchi 27952).
Distribution. Europe, Caucasus, Pakistan, N.
and C. Asia, China, Korea, Japan and N. America.

Bryophyte flora of the Tien Shan Mountains. Hypnaceae and Hylocomiaceae

2. Pylaisia polyantha (Hedw.) Schimp. in
B.S.G., Bryol. Eur. 5: 89 (1851).
Specimens examined. Russia, Novosibirsk,
Central Siberian Botanical Garden, 150 m, July12, 1995 (Higuchi 27723, 27725, 27728, 27730);
July 14, 1995 (27736, 27743, 27752).
Distribution. North Europe, Russia and N.
America (Arikawa 2004).

117

Nine species and one subspecies of Hypnum
including H. callichroum, H. cupressiforme var.
subjulaceum and H. subimponens subsp. subimponens newly added here are known from Kazakhstan and/or Kyrgyzstan. The following key isrevised from Ando (1973, 1976, 1993, 1994).
The species with asterisk (*) are not present inthis collection.

Key to the speices of Hypnum in Kazakhstan and Kyrgyzstan
1. Perichaetial leaves not plicate; seta twisted to the right throughout the whole length when dry,rarely only slightly turned to the left below the capsule; spores maturing in fall to winter . . . . . 2
1. Perichaetial leaves plicate; seta twisted to the right below, to the left above when dry; spores maturing in winter to spring, spring to summer or in summer to fall . . . . . . . . . . . . . . . . . . . . . . . . 3
2. Leaves oblong-lanceolate; alar parts remarkably excavated and usually brown-colored; median cells (50–)60–70(–80)3–4(–5) m m . . . . . . . . . . . . . . 5. H. cupressiforme var. subjulaceum
2. Leaves ovate-lanceolate; alar parts not or weakly excavated, occasionally yellow- to browncolored; median cells shorter and wider . . . . . . . . . . . . 6. H. cupressiforme var. cupressiforme
3. Epidermal cells of stem always much larger than the associated inner cortical cells, hyaline withthin outer walls (hyalodermous); sometimes phyllodioicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3. Epidermal cells of stem not enlarged, with thick rarely thin outer walls, rarely partly enlarged andhyaline; not phyllodioicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
4. Plants medium- to large-sized; leaves usually broadly acuminate; growing in moist habitats. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. H. lindbergii
4. Plants small- to medium-sized; leaves slenderly acuminate; growing in drier habitats . . . . . 5
5. Stem leaves weakly narrowed and not or only slightly rounded to the insertion; margins usually
recurved below, sometimes so upwards; alar cells only slightly differentiated . . . . . . . . . . . . . . . 6
5. Stem leaves more markedly narrowed and strongly or weakly rounded to the insertion; margins
plane; alar cells well differentiated, forming a distinct group . . . . . . . . . . . . . . . . 4. H. callichroum
6. Plants small-sized, procumbent or erect to ascending in dense tufts; stem leaves ovate-lanceolate, 1.3–1.50.4–0.5 mm; laminal cells 40–60(–70)3–4 m m in lumina . . . . H. hamulosum*
6. Plants medium-sized, usually procumbent; stem leaves ovate- to oblong-lanceolate withlonger acumen, 1.5–2.80.4–0.6 mm; laminal cells 50–80(–90)3–4 m m . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. H. subimponens subsp. subimponens
7. Alar parts of leaves consisting of heterogeneous cells, with enlarged, thin-walled, hyaline cells at
the outer angle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. fertilis*
7. Alar parts of leaves consisting of homogeneous subquadrate to rectangular cells, in most cases
without enlarged, hyaline cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Alar parts of leaf strongly excavate, composed of porose cells that are thick-walled . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. H. bambergeri
8. Alar parts of leaf not or only slightly excavate, composed of cells that are not thick-walled . . .
......................................................................... 99. Plants large-sized; leaves more markedly dimorphic in stem- and branch leaves; stem leaves
cordate-auriculate at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. H. procerrimum
9. Plants small- to medium-sized; leaves slightly dimorphic in stem- and branch leaves; stem leaves

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not cordate-auriculate at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Leaves distinctly serrate; operculum rostrate; growing chiefly on tree-trunks . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. pallescens*
10. Leaves almost entire to serrulate; operculum shorter-apiculate; growing chiefly on rocks orearth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Autoicous; plants smaller (branch leaves 0.2–0.4 mm wide); pseudoparaphyllia numerous, narrowlanceolate to filamentous; subquadrate alar cells 4–10 in the marginal row . . . . . 8. H. recurvatum
11. Dioicous; plants usually larger (branch leaves 0.3–0.6 mm wide or more); pseudoparaphyllia
fewer, wider-foliose; subquadrate alar cells 8–16(–20) in the marginal row . . . . . . . . . . . . . . . . . 12
12. Pseudoparaphyllia semicircular or round-rectangular; leaves broader, more strongly concave,smooth, with plane margins; subquadrate alar cells numerous, 10–16(–20) in the marginal row. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. H. vaucheri
12. Pseudoparaphyllia ovate to lanceolate; leaves narrower, slightly concave, usually plicate, withrevolute, occasionally plane margins; subquadrate alar cells fewer, 8–15 in the marginal row. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. H. revolutum
3. Hypnum bambergeri Schimp., Syn. Musc.
Eur., ed. 1, Addend. 698 (1860).
Specimens examined. Kazakhstan, Mts. Dzhungarsky Alatau, Maly Baskan, 2000 m, July 21,
1995 (Higuchi 27957); 2100 m, July 22, 1995
(Higuchi 28023).
Distribution. Spitsbergen, Iceland, Europe,
Russia, Canada, Alaska and Greenland (Ando
1996).
Notes. This species is characterized by having falcate, smooth leaves with entire channelled
acumen, incrassate, porose laminal cells andbrown-colored alar parts of leaves. According toAndo (1996), the distribution of the species in
Asia is limited to Siberia north of about 60°N
and the Altai Mountains. Ignatov et al. (2006) reported this species from Kazakhstan and Kyrgyzstan as Stereodon bambergeri (Schimp.)
Lindb.
4. Hypnum callichroum Brid., Bryol. Univ. 2:
631 (1827).
Specimen examined. Kazakhstan, North of
Semipalatinsk, 340 m, July 17, 1995 (Higuchi
27762).
Distribution. Europe, Siberia and northern
N. America (Ando 1997, as H. callichroum
subsp. callichroum). New to Kazakhstan.
Notes. This species is characterized by thehyalodermatous stems and inflated hyaline alarcells of leaves. Ando (1997) described a newsubspecies of the species, H. callichroum subsp.
japonicum, from alpine region of Japan, which isdistinguished from subsp. callichroum by thesmaller-sized plants usually showing more closebranching and foliation, and the somewhat shorter laminal cells and setae.
5. Hypnum cupressiforme Hedw. var. cupressiforme, Spec. Musc. 291 (1801).
Specimens examined. Kazakhstan, Sarkand,
900 m, July 20, 1995 (Higuchi 27814Hypnum
revolutum); Mts. Dzhungarsky Alatau, Maly
Baskan, 1900 m, July 22, 1995 (Higuchi 27985,
28085, 28088, 28089, 28096, 28106); 2100 m,
July 22, 1995 (Higuchi 28029); Mts. Zailisky
Alatau, north of Almaty, 1750 m, July 25, 1995
(Higuchi 28131). Kyrgyzstan, Tien Shan, Mts.
Tereskey Alatau, 2800 m, Aug. 1, 1995 (Higuchi
28411); 2900 m, July 31, 1995 (Higuchi 28355);
2920 m, Aug. 1, 1995 (Higuchi 28428).
Distribution. Widely distributed in the world.
6. Hypnum cupressiforme Hedw. var. subjulaceum Molendo, Ber. Naturh. Ver. Augsburg.
18: 183 (1865).
Specimens examined. Kazakhstan, Sarkand,
900 m, July 20, 1995 (Higuchi 27785); Mts.
Dzhungarsky Alatau, Maly Baskan, 1900 m, July22, 1995 (Higuchi 28083, 28098, 28100). Kyr-

Bryophyte flora of the Tien Shan Mountains. Hypnaceae and Hylocomiaceae

gyzstan, Tien Shan, Mts. Tereskey Alatau, 3600
m, July 31, 1995 (Higuchi 28375); near glacier,3600 m, Aug. 2, 1995 (Higuchi 28459, 28466);
Mts. Alaisky, south of Gulcha, 2000 m, Aug. 9,
1995 (Higuchi 28643, 28652, 28663, 28668).
Distribution. Widely distributed in the
Northern Hemisphere and New Zealand. New to
Kyrgyzstan.
Notes. This variety is characterized by having subjulaceous branches and the not or weakly
falcate leaves with excavated and usually browncolored alar areas. Although Ignatov et al. (2006)
doesn’t list the variety from Kazakhstan, Ando
(1990) cited the specimen of the variey from
“Kasachstania, Montes Kungej-Alatau, 2000 m,
Lisowski (NICH 208140).”
7. Hypnum lindbergii Mitt., J. Bot. 2: 123(1864).
Specimen examined. Kazakhstan, Mts. Dzhungarsky Alatau, Maly Baskan, 2400 m, July 21,
1995 (Higuchi 27890).
Distribution. Widely distributed in the
Northern Hemisphere and Brasil.
Notes. Hedenäs (1990) transferred thisspecies to the genus Calliergonella based onsome characteristics, such as appendiculate cilia
and large groups of strongly inflated alar cells ofstem leaves.
8. Hypnum recurvatum (Lindb. & Arn.)
Kindb., Enum. Bryin. Exot. 100 (1891).
Specimens examined. Kazakhstan, Sarkand,
900 m, July 20, 1995 (Higuchi 27789); Mts.
Zailisky Alatau, north of Almaty, 1750 m, July25, 1995 (Higuchi 28126). Kyrgyzstan, Tien
Shan, Mts. Tereskey Alatau, 2500 m, Aug. 2,
1995 (Higuchi 28502, 28503); 2920 m, Aug. 1,
1995 (Higuchi 28432, 28434).
Distribution. Urals, Fennoscandia, Tatra,
Carpathians, Alps, Jura, Pyrenees, Caucasus,
Siberia, Mongolia, Canada, Montana and Greenland (Ando 1973). New to Kyrgyzstan.
Notes. This species is closely related to Hypnum revolutum, but is distinguished from itby having narrow-lanceolate to filamentous

119

pseudoparaphyllia (ovate to lanceolate in H. revolutum), leaves with margins plane or recurved
below (usually margins revolute from the base up
to near the apex in H. revolutum) and subquadrate alar cells 4–10 in the marginal row(8–15 in the marginal row in H. revolutum).
9. Hypnum revolutum (Mitt.) Lindb., Oefv. K.
Vet. Ak. Foerh. 23: 542 (1867).
Specimens examined. Kazakhstan, Sarkand,
900 m, July 20, 1995 (Higuchi 27814Hypnum
cupressiforme var. cupressiforme); Mts. Dzhungarsky Alatau, Maly Baskan, 900 m, July 21,
1995 (Higuchi 27820, 27825); 1900 m, July 22,
1995 (Higuchi 27979, 27980, 27984, 28001,
28067, 28073, 28079); 2100 m, July 22, 1995
(Higuchi 27834); 2270 m (Higuchi 28019); 2400m, July 21, 1995 (Higuchi 27851, 27856, 27868,
27881, 27899); 2550 m (Higuchi 27907, 27931,
27939); Mts. Zailisky Alatau, north of Almaty,
1750 m, July 25, 1995 (Higuchi 28136, 28139);
near reservoir, 2500 m (Higuchi 28142, 28153,
28159); 2560 m, July 27, 1995 (Higuchi 28259,
28265); 2600 m (Higuchi 28287); 2680 m, July
26, 1995 (Higuchi 28254); 2800 m (Higuchi
28177): 2850 m (Higuchi 28182, 28183); 3100 m(Higuchi 28205, 28208); 3210 m (Higuchi
28219, 28233, 28243, 28246). Kyrgyzstan, Tien
Shan, Mts. Tereskey Alatau, 2480 m, July 30,
1995 (Higuchi 28317); 2700 m (Higuchi 28339);
2920 m, Aug. 1, 1995 (Higuchi 28427); 3370 m,
July 31, 1995 (Higuchi 28398); 3600 m, Aug. 2,
1995 (Higuchi 28454, 28468, 28469, 28479);
3640 m (Higuchi 28386); 3950 m, (Higuchi
28381); Mts. Kirghisky, 3400 m, Aug. 6, 1995
(Higuchi 28637); 3550 m (Higuchi 28561,
28563, 28575, 28576); near glacier, 3600 m(Higuchi 28583, 28586, 28604, 28606, 28608);
Mts. Alaisky, south of Gulcha, 3550 m, Aug. 9,
1995 (Higuchi 28658); 3600 m (Higuchi 28649);
Mts. Za Alaisky, 2700 m, Aug. 11, 1995 (Higuchi
28694, 28698).
Distribution. Widely distributed in the
Northern Hemisphere and Antarctic Peninsula(Ando 1973).
Notes. This species is the commonest

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Masanobu Higuchi

species of the genus in the area investigated.
Hypnum revolutum is variable in size, but is mostdistinct in having more or less plicate leaves withstrongly revolute margins and alar parts of leaves
consisting of homogeneous subquadrate cells.
10. Hypnum subimponens Lesq. subsp. subimponens, Trans. Am. Phil. Soc., n. ser. 13: 14(1880).
Specimens examined. Kazakhstan, Sarkand,
900 m, July 20, 1995 (Higuchi 27790, 27809).
Distribution. Siberia and northwestern N.
America (Ando 1997). New to Kazakhstan.
Notes. This subspecies differs from H.
subimponens subsp. ulophyllum (Müll.Hal.)
Ando distributed in Himalayas, China, Taiwan,
Korea and Japan by its larger plants, shorter
costae of leaves and longer, arched capsules.
11. Hypnum vaucheri Lesq., Mem. Soc. Sc.
Nat. Neuchatel 3(3): 48 (1846).
Specimen
examined. Kazakhstan,
Mts.
Dzhungarsky Alatau, Maly Baskan, 1900 m, July21, 1995 (Higuchi 28101).
Distribution. Widely distributed in the
Northern Hemisphere.
12. Hypnum procerrimum Molendo, Flora
49: 458 (1866).
Specimens examined. Kazakhstan, Mts. Dzhungarsky Alatau, Maly Baskan, 2550 m, July 21,
1995 (Higuchi 27932, 27941); Mts. Zailisky
Alatau, north of Almaty, 3210 m (Higuchi
28222). Kyrgyzstan, Tien Shan, Mts. Tereskey
Alatau, 2920 m, Aug. 1, 1995 (Higuchi 28412,
28414, 28438, 28439); near glacier, 3600 m,
Aug. 2, 1995 (Higuchi 28476).
Distribution. Widely distributed in the
Northern Hemisphere.
Notes. This species is characterized by having smooth or slightly plicate only in upper partsof leaves, plane margins of branch leaves and alarparts of leaves consisting of numerous homogeneous subquadrate cells.

Hylocomiaceae
13. Hylocomium splendens (Hedw.) Schimp.,
Bryol. Eur. 5: 173 (1852).
Specimens examined. Russia, Novosibirsk,
Central Siberian Botanical Garden, 150 m, July
14, 1995 (Higuchi 27747). Kazakhstan, Mts.
Dzhungarsky Alatau, Maly Baskan, 1900 m, July22, 1995 (Higuchi 27993); 2100 m, July 21, 1995
(Higuchi 27836, 27837); 2500 m (Higuchi
27901). Kyrgyzstan, Tien Shan, Mts. Tereskey
Alatau, 2430 m, July 30, 1995 (Higuchi 28331,
28336).
Distribution. Widely distributed in the
Northern Hemisphere.
14. Pleurozium schreberi (Brid.) Mitt., J.
Linn. Soc. Bot. 12: 537 (1869).
Specimens examined. Kyrgyzstan, Tien Shan,
Mts. Tereskey Alatau, 2920 m, Aug. 1, 1995
(Higuchi 28426); 3640 m, July 31, 1995 (Higuchi
28389).
Distribution. Widely distributed in the
Northern Hemisphere.
15. Rhytidiadelphus
triquetrus(Hedw.)
Warnst., Krypt. Fl. Brandenburg 920: 996(1906).
Specimens examined. Kyrgyzstan, Tien Shan,
Mts. Tereskey Alatau, 2430 m, July 30, 1995
(Higuchi 28318, 28320, 28326).
Distribution. Widely distributed in the
Northern Hemisphere.
16. Rhytidium rugosum (Hedw.) Kindb., Bih.
K. Svensk. Vet. Ak. Handl. 7(9): 15 (1883).
Specimens examined. Kazakhstan, Mts.
Dzhungarsky Alatau, Maly Baskan, 2400 m, July21, 1995 (Higuchi 27891); 2480 m (Higuchi
27953); Zailisky Alatau, north of Almaty, 3000m, July 26, 1995 (Higuchi 28202). Kyrgyzstan,
Tien Shan, Mts. Tereskey Alatau, 2920 m, Aug.
1, 1995 (Higuchi 28413).
Distribution. Morocco, Europe, Caucasus,
Himalayas, N. and C. Asia, China, Japan, N.
America, Mexico, Guatemala and Greenland.

Bryophyte flora of the Tien Shan Mountains. Hypnaceae and Hylocomiaceae

Acknowledgements
I wish to thank Dr. Yuichi Kadota, the leaderof the expedition, for his encouraging studying
bryophytes of the Tien Shan Mountains. I am
grateful to Dr. Ivan M. Krasnoborov, Dr. Elena
Wiebe and Dr. Alexandre A. Krasnikov, Central
Siberian Botanical Garden, for their kind help in
the field research in 1995. This study was supported in part by Grant-in Aid (no. 18570097 to
M. Higuchi) from the Ministry of Education,
Culture, Sports, Science and Technology, Japan,
and collection of plants was no. 07041152 to Y.
Kadota.
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