Bull. Natl. Mus. Nat. Sci., Ser. B, 39(4), pp. 165–172, November 22, 2013

A Collection of Hepatics from the Tottori Prefecture, Japan
Vadim A. Bakalin1,*, Tomotsugu Arikawa2 and Masanobu Higuchi3
1

Botanical Garden-Institute, Far Eastern Branch of the Russian Academy of Sciences,
Makovskogo Street 142, Vladivostok 690024, Russia
2
Keio University, Hiyoshi 4–1–1, Kohoku-ku, Yokohama, Kanagawa 223–8521, Japan3
Department of Botany, National Museum of Nature and Science,
Amakubo 4–1–1, Tsukuba, Ibaraki 305–0005, Japan* E-mail: v_bak@list.ru(Received 27 August 2013; accepted 2 October 2013)

Abstract Basing on the collection of hepatics gathered in the middle of March, 2013, in Tottori
Prefecture, Japan, 56 species of liverwort and one hornwort species are recorded. Compiled checklist provides the information on generative structures and vegetative propagules presence, altitudinal diapason of collected specimens, habitat description and accompanying taxa list for eachrecorded species. Nine species (Aneura maxima, Conocephalum salebrosum, Lejeunea aquatica,
Megaceros ﬂagellaris, Plectocolea horikowana, Plectocolea ovalifolia, Riccardia kodamae, Riccardia latifrons and Riccardia tamariscina) are new for Tottori Prefecture.
Key words : bryophytes, hornworts, Japan, liverworts, Tottori Prefecture.

Tottori Prefecture is located in South–Westpart of Honshu, Japan, and covers over than 3500
km2. It stretches out long and thin from east to
west (approximately 120 km east–west and maximum 50 km north–south). It faces the Sea ofJapan in the north, and it is bordered southwardwith a ridgeline of Chugoku Mountains including
Mt. Daisen (1,729 m alt.), the highest peak in a
Western part of Honshu. The vegetation of thelower part under 300 m alt. was originally represented by warm-temperate, evergreen broadleaved forests (e.g. laurel forest), and that of thehigher part over 600 m alt. was by deciduous
broad-leaved forest (e.g. beech forest). The part
between 300 and 600 m alt. is an ecotone. Mostof these forests, however, are now replaced bysecondary forests dominated by Pinus densiﬂora
and the plantation of Cryptomeria japonica. Tottori Prefecture is the least populous prefecture in
Japan (ca. 590,000), because of the mountainous
topography. In spite of the low latitude, heavy
snowfalls are sometimes occurred in winterowing to the Sea of Japan.
The prefecture has a long history of bryological exploration that was summarized by Arikawa
et al. (2011) provided checklist of bryophytes ofthe prefecture. The list counts 312 species ofmosses, 124 species of liverworts and 3 speciesof hornworts recorded in literature. Later Arikawa et al. (2012) published ﬂoristic results ofﬁeld survey for Tottori red list species recognizing. That list is based on ﬁeld researches curriedin 2010. Arikawa et al. (2012) provide data on
distribution of 183 species of mosses and 93 species of hepatics, respectively with 30 and 14 species of bryophytes new for Tottori Prefecture.
Recently Kobayashi (2012a, 2012b) also reported
2 species of hepatics as new record for Tottori
Prefecture.
Our researches were conducted in three days
at March of 2013 as the part of collaboration
project for studies of biota diversity and development around Sea of Japan, concluded between
National Museum of Nature and Science, Tokyo,Vadim A. Bakalin et al.

166

Fig. 1.

Map of collecting localities in Tottori Prefecture.

and three institutes of the Far East Division of
the Russian Academy of Sciences (Botanical
Garden-Institute, Institute of Biology and Soil
Sciences and Institute of Geology). Due tospringtime we conducted ﬁeld researches in low
altitudes in Tottori Prefecture (see Fig. 1 andTable 1), when over than 200 specimens ofhepatics were collected. All specimens werestudied for oil-bodies features within one week
after the collection, when the plants were in living condition. Despite the time for collecting wasvery limited we have found 57 species of hepatics (56 liverworts and 1 hornwort), 9 of them are
new for the prefecture. Due to the latter fact, as
well as we found in additional localities somerare in Tottori Prefecture taxa we provide the listof our collection below.
Enumeration of Species
The conspectus lists taxa in alphabetical order.
Taxa new for the prefecture are marked with

A Collection of Hepatics from the Tottori Prefecture, Japan
Table 1.

167

Collecting localities in Tottori Prefecture

Locality
numberCoordinates

Geographic description

1
2

35°30′04.7″N 134°14′59.7″E35°28′55.1″N 133°56′09.3″E

3435°28′54.0″N 133°56′07.3″E
35°27′13.8″N 133°56′50.8″E

535°28′41.3″N 133°58′20.7″E

6
7
8
9

35°25′17.3″N 134°05′28.0″E
35°24′09.6″N 134°06′0.9.4″E35°24′48.3″N 134°09′42.6″E35°27′40.8″N 134°22′18.2″E

1035°28′42.5″N 134°24′04.0″E

Tottori-shi, Ue-machi, Ouchi-dani Park
Tohaku-gun, Yurihama-cho, Urushibara,
Fudo Waterfall
Tohaku-gun, Yurihama-cho, Urushibara
Tohaku-gun, Yurihama-cho, Kawakami,
around the summit of Mt. Hachibuse
Tottori-shi, Aoya-cho, Tawaradani, Fudo
Waterfall
Tottori-shi, Kouchi, Kouchi Shrine
Tottori-shi, Iwatsubo, Iwatsubo Shrine
Tottori-shi, Naka-sunami
Tottori-shi, Kokufu-cho, Sugano, Sugano
MarshTottori-shi, Kokufu-cho, Amedaki, Amedaki
Waterfall

asterisk. The nomenclature mostly followed tothe accepted in the “Catalog of the hepatics ofJapan” (Yamada and Iwatsuki, 2006) with the
exception in recognizing of Plectocolea andSolenostoma as the genera different from,
Jungermannia s. str. (cf. Bakalin and Vilnet,
2012). Each species is annotated by the followings.
(1) Locality number.
(2) Presence of generative structures and vegetative propagules (if present) in the following
abbreviations: per. (perianths including ﬂeshy
calyptrae and pseudoperianths of metzgerioid
hepatics), ant. (antheridia), arch. (archegonia),
spor. (sporangia), gemm. (gemmae).
(3) Altitudinal diapason where the species wascollected.
(4) Habitat description.
(5) Accompanying species (if present).
(6) Selected specimens examined herbarium
number (one per each locality, according to theFig. 1 and Table 1).
(7) In some cases some taxonomical or othernotes are added.
The specimens cited here were mainly collected by Bakalin, and they are deposited in theherbarium of the Botanical Garden-Institute, Far
Eastern Branch of the Russian Academy of Sciences (VBGI) and some duplicates in the herbarium of the Department of Botany, National

Altitude,m alt.

Date

60
40

10 March 2013
11 March 2013

40
500

11 March 2013
11 March 2013115

11 March 2013

170
240
210
400

12 March 201312 March 201312 March 201312 March 2013390

12 March 2013

Museum of Nature and Science (TNS).
* Aneura maxima (Schiffn.) Steph. L10, 390
m. Wet cliff in open place in the spray zone ofwaterfall. Only one collection. With Conocephalum salebrosum, Heteroscyphus coalitus, Pallavicinia subciliata, Pellia neesiana and Riccardia chamaedryfolia. J-10-9-13.
Bazzania tridens (Reinw., Blume & Nees)
Trevis. L1, 60 m. Wet clay road cuts and tree
trunks in part shade in broadleaved-coniferous
forest. In pure mats or with Blepharostoma
minus and Kurzia makinoana. J-1-45-13.
Blepharostoma minus Horik. L1, 60 m. Wet
clay road cuts and stone walls in the valley inpart shade in the secondary forests. In pure matsor with Bazzania tridens, Diplophyllum taxifolium, Kurzia makinoana and Scapania parvitexta. J-1-44-13.
Calypogeia japonica Steph. L1, 60 m. Wet
stone walls in the valley in the seconcary forests.
In pure mats or with Conocephalum salebrosum,
Pallavicinia subciliata and Riccardia tamariscina. J-1-15-13.
Calypogeia tosana (Steph.) Steph. L1, 2, 7,
40–240 m. Wet sandy and clayish road cuts inpart shade, clay on slope to stream, moist cliffs
along stream in the secondary forests. In puremats or with Lejeunea japonica, Pallavicinia
subciliata, Plagiochila ovalifolia and Plectocolea truncata. J-1-26-13, J-2-9-13, J-7-18-13.

168

Vadim A. Bakalin et al.

Cavicularia densa Steph. L7, 240 m. Very
wet cliff along stream in the secondary forest.
Only one collection. With Conocephalum salebrosum and Plectocolea unispiris. J-7-13-13.
Cephalozia otaruensis Steph. L1,5, per.
60–110 m. Moist rotten log in part shade in thesecondary forests. In pure mats or with Heteroscyphus coalitus, Odontoschisma denudatum
and Riccardia latifrons var. miyakeana. J-1-4-13,
J-5-30-13.
Chiloscyphus polyanthos (L.) Corda L2, 5, 7,
ant., per., spor. 40–240 m. Very wet cliffs inspray zone of waterfall in part shade, wet boulders and cliffs in the stream valley in open place
or in part shade in the secondary forests. In puremats or with Conocephalum salebrosum. J-2-1513, J-5-12-13, J-7-16-13.
Cololejeunea japonica (Schiffn.) S.Hatt. ex
Mizut. gemm. L1, 60 m. Tree trunk in the secondary forest. Only one collection. With Radulaconstricta. J-1-19-13.
Cololejeunea longifolia (Mitt.) Benedix in
Mizut. L6, 170 m. Boulders in part shade in thesecondary forests. Only one collection. In puremats. J-6-16-13.
Conocephalum japonicum (Thunb.) Grolle
L5, 110 m. Clay covering rocks in slope tostream in mesic conditions in part shade in thesecondary forests. Only one collection. With
Dumortiera hirsuta and Plectocolea horikawana.
J-5-2-13.
* Conocephalum salebrosum Szweyk., Buczkowska & Odrzykoski spor. L1, 5, 7, 10, 60–390
m. Wet clay and sandy road cuts, moist rotten
log, wet cliffs along stream and in the spray zone
of waterfall, clay covering rocks in slope tostream in mesic conditions, in part shade to open
places mostly in the secondary forests. Verycommon species in pure mats or with Aneura
maxima, Calypogeia japonica, Cavicularia
densa, Chiloscyphus polyanthos, Dumortiera hirsuta, Heteroscyphus coalitus, H. planus, Makinoa crispata, Pallavicinia subciliata, Pellia neesiana, Plectocolea unispiris, Radula kojana,
Riccardia chamaedryfolia, R. tamariscina,
Trichocolea tomentella and Wiesnerella denu-

data. J-1-38-13, J-5-28-13, J-7-17-13, J-10-1213.
This species was recorded for Japan (Kyushu,
Miyazaki-ken) relatively recently (Szweykovsky
et al., 2005). In the same paper (l.c.) it was
shown that the distribution of C. conicum (L.)
Underw. mainly restricted to Europe, with thewesternmost known locality in Turkey. Further
researches by Borovichev et al. (2009) showed
the similar tendency (the westernmost locality of
C. conicum in Eurasia is in Altai Mts.). In our
researches we did not recognize this species in
the Russian Far East, Korean Peninsula andChina (unpublished), where all population of thespecies previously named as C. conicum arereally belonging to C. salebrosum. Akiyama(2006) suggested that Conocephalum conicum
doesnʼt occur in Japan. The present researchrather conﬁrms this point of view.
Diplophyllum taxifolium (Wahlenb.) Dumort.
L1, 7, ant., per. 60–240 m. Moist to wet cliffs
along streams and wet stone walls in the valley
in the secondary forests. In pure mats or with
Blepharostoma minus and Scapania parvitexta.
J-1-10-13, J-7-14-13.
Dumortiera hirsuta (Sw.) Nees in Reinw. L1,
2, 5, 7, 10, arch. 40–390 m. Moist to wet clayish
road cuts, cliffs along stream, boulders and clay
covering rocks on slopes to valleys, cliff in sprayzone of waterfall, mostly in part shade in the secondary forests. Very common species growing inpure mats or with Conocephalum japonicum,
C. salebrosum, Heteroscyphus coalitus, Jubula
hutchinsiae, Pallavicinia subciliata, Pellia neesiana, Plectocolea sp. and Wiesnerella denudata.
J-1-41-13, J-2-10-13, J-5-2-13, J-7-21-13, J-104-13.
Frullania appendiculata Steph. L1, 60 m.
Tree trunks in the secondary forest. In pure mats.
J-1-22-13.
Frullania appendiculata Steph. was considered by Hattori (1972) as one of synonyms of
F. tamarisci subsp. obscura. Recent molecularresearches, however, conﬁrmed the species rank
for F. appendiculata (cf. Heinrichs et al., 2010).
Frullania davurica Hampe L1, 60 m. TreeA Collection of Hepatics from the Tottori Prefecture, Japan

trunks in the secondary forests. In pure mats. J-120-13.
Frullania muscicola Steph. L1, 3, per., spor.
40–60 m. Tree trunk in open places near theroads or trunk in the secondary forests in partshade. In pure mats. J-1-23a-13, J-3-4-13.
Haplomitrium mnioides (Lindb.) R.M.
Schust. L1, 7, ant., per. 60–240 m. Moist to wet
cliffs along streams and wet sandy to clayish
road cuts in part shade in the secondary forests.
In pure mats. J-1-27-13, J-7-30-13.
Heteroscyphus coalitus (Hook.) Schiffn. L1,
2, 5, 7, 10, 40–390 m. Moist to wet clay to sandyroad cuts in part shade, moist to wet cliffs along
streams, near waterfalls, mostly in part to fullshade in the secondary forests. Very common
species. In pure mats or with Aneura maxima,
Cephalozia otaruensis, Conocephalum salebrosum, Dumortiera hirsuta, Heteroscyphus planus,
Makinoa crispata, Pallavicinia subciliata, Plagiochila ovalifolia, P. sciophila, Radula kojana,
Riccardia chamaedryfolia, Pellia neesiana,
Trichocolea tomentella and Wiesnerella denudata. J-1-57-13, J-2-22-13, J-5-22-13, J-7-23-13,
J-10-12-13.
Heteroscyphus planus (Mitt.) Schiffn. L1, 5,
6, 7, 60–240 m. Moist to wet clay to sandy roadcuts in part shade, moist cliffs and tree trunks inpart shade in stream valleys in the secondary forests. In pure mats or with Conocephalum salebrosum, Heteroscyphus coalitus, Plagiochila sciophila and Trichocolea tomentella. J-1-39-13,
J-5-19-13, J-6-4-13, J-7-25-13.
Jubula hutchinsiae (Hook.) Dumort. ssp.
javanica (Steph.) Verd. L2, 5, 6, 7, 10, ant.,arch., per. 40–390 m. Moist boulders and humus
on slopes to and along of the streams in part tofull shade in the secondary forests. In pure matsor with Dumortiera hirsuta, Megaceros ﬂagellaris, Makinoa crispata, Plagiochila ovalifolia,
Riccardia chamaedryfolia and Wiesnerella denudata. J-2-5-13, J-5-33-13, J-6-13-13, J-7-1-13,
J-10-5-13.
Kurzia makinoana (Steph.) Grolle L1, per.
60 m. Rotten logs and wet clay road cuts in partshade in the secondary forests. In pure mats or169with Bazzania tridens, Blepharostoma minus,
Plagiochila ovalifolia and Plectocolea ovalifolia.
J-1-2-13.
* Lejeunea aquatica Horik. L2, 5, 40–110 m.
Very wet to moist cliffs in spray zone of waterfall
or along steam valley, in part shade in the secondary forests. In pure mats or with Riccardia
chamaedryfolia. J-2-18-13, J-5-21-13.
Lejeunea japonica Mitt. L5, 7, ant., per. 110–240 m. Wet to moist stone walls and cliffs, moist
humus slope to valleys and along streams, tree
trunks in part shade in the secondary forests. In
pure mats or with Calypogeia tosana, Plagiochila ovalifolia, P. sciophila and Wiesnerella denudata. J-5-15-13, J-7-18-13.
Lophocolea minor Nees L1, 4, 5, 6, gemm.,
per. 60–500 m. Tree trunks in open places andmoist rotten log in part shade in the secondary
forests. In pure mats or with Metzgeria lindbergii. J-1-1-13, J-4-1-13, J-5-8-13, J-6-3-13 .
This species is listed as Chiloscyphus minor(Nees) J.J.Engel & R.M.Schuster in Katagiri and
Furuki (2012).
Macvicaria ulophylla (Steph.) S.Hatt L3, 40
m. Tree trunk in open place near the road acrossrice ﬁelds. In pure mats or with Radula constricta. J-3-2-13.
Makinoa crispata (Steph.) Miyake L1, 5, 6,
60–170 m. Wet stone walls, boulders and moist
cliffs in part shade along streams in the secondary forests. In pure mats or with Conocephalum
salebrosum, Heteroscyphus coalitus, Jubula
hutchinsiae, Megaceros ﬂagellaris and Riccardia
chamaedryfolia. J-1-12-13, J-5-22-13, J-6-1113; Higuchi 51330.
*Megaceros ﬂagellaris (Mitt.) Steph. L2, 6, 7,
spor. 40–240 m. Very wet cliffs in spray zone ofwaterfalls, wet to moist boulders in the stream
valleys in part shade in the secondary forests. In
pure mats or with Jubula hutchinsiae, Makinoa
crispata and Riccardia chamaedryfolia. J-2-2113, J-6-11-13, J-7-1-13; Higuchi 51349.
Metzgeria furcata (L.) Dumort. L6, 170 m.
Moist boulders along stream in part shade in thesecondary forests. Only one collection. In puremat. J-6-10-13.

170

Vadim A. Bakalin et al.

Metzgeria lindbergii Schiffn. L5, ant., arch.
110 m. Tree trunk in open place in the secondary
forests. Only one collection. With Lophocolea
minor. J-5-7-13.
Nardia assamica (Mitt.) Amakawa L3, 6, per.,spor. 40–170 m. Clayish roadside in mesic condition in open to partly shaded places along roadacross rice ﬁelds or in the secondary forests. In
pure mats or with Plectocolea infusca var. ovicalyx. J-3-1-13, J-6-5-13.
Odontoschisma denudatum (Mart.) Dumort.
L1, gemm. 60 m. Rotten log in part shade in thesecondary forests. In pure mats or with Cephalozia otaruensis and Riccardia latifrons var.
miyakeana. J-1-5-13.
Pallavicinia subciliata (Austin) Stephani L1,
5, 6, 7, 10, ant., arch., per., spor. 60–390 m. Wet
to moist clay, humus, stones on slopes to andalong streams, near waterfalls, along roads or inforest ﬂoor, mostly in part shade in the secondary
forests. Very common species. In pure mats orwith Aneura maxima, Calypogeia japonica, C.
tosana, Conocephalum salebrosum, Dumortiera
hirsuta, Heteroscyphus coalitus, Pellia neesiana,
Plectocolea truncata, Radula kojana, Riccardia
chamaedryfolia and Trichocolea tomentella. J-117-13, J-5-25-13, J-6-6-13, J-7-31-13, J-10-1213; Higuchi 51313.
Pellia neesiana (Gott.) Limpr. L1, 7, 10, arch.,
spor. 60–390 m. Wet clay road cuts, wet to moist
cliffs along streams or in the spray zone of waterfalls in part shade in the secondary forests. In
pure mats or with Aneura maxima, Conocephalum salebrosum, Dumortiera hirsuta, Heteroscyphus coalitus, Pallavicinia subciliata andRiccardia chamaedryfolia. J-1-37-13, J-7-31-13,
J-10-9-13.
Plagiochila ovalifolia Mitt. L1, 2, 7, 10,
40–390 m. Wet to moist sandy road cuts, boulders along streams, cliffs in the spray zone ofwaterfalls, mostly in part shade in the secondary
forests. In pure mats or with Heteroscyphus
coalitus, Kurzia makinoana, Plagiochila sciophila and Wiesnerella denudata. J-1-31-13, J-2-2213, J-7-2-13, J-10-6-13.
Plagiochila sciophila Neew in Lindenb. L5, 7,

10, ant., arch. 110–390 m. Cliffs, tree trunks andbranches mostly in mesic, but rarely in wet conditions, in part shade in the secondary forests. In
pure mats or with Heteroscyphus coalitus, H.
planus, Lejeunea japonica, Plagiochila ovalifolia, Radula kojana and Wiesnerella denudata.
J-5-18-13, J-7-7-13, J-10-1-13; Higuchi 51343.
* Plectocolea horikawana Amakawa L5, ant.,
per., spor. 110 m. Clay covering rocks in slope tostream in mesic conditions in part shade in thesecondary forests. In pure mats or with Conocephalum japonicum. J-5-1-13.
This species is listed as Solenostoma horikawanum (Amakawa) Váňa, Hentschel & J.Heinrichs in Katagiri and Furuki (2012).
Plectocolea infusca var. ovicalyx (Steph.)
Bakalin L1, 40 m. Clayish road cut along roadacross rice ﬁelds in open place. Only one collection. With Nardia assamica. J-3-1-13.
This variety is listed as Solenostoma infuscum(Mitt.) J.Hentschel var. ovicalyx (Steph.) Potemkin & Sofronova in Katagiri and Furuki (2012).
* Plectocolea ovalifolia (Amakawa) Bakalin& Vilnet L1, per., spor. 60 m. Wet clay road cut
in part shade in the secondary forest. Only one
collection. With Diplophyllum taxifolium, Kurzia
makinoana. J-1-56-13.
This species is listed as Solenostoma infuscum(Mitt.) J.Hentschel var. ovalifolium (Amakawa)
Potemkin & Sofronova in Katagiri and Furuki(2012).
Plectocolea truncata (Nees) Bakalin L1, 5,
ant., per., spor. 60–110 m. Mesic to wet clay on
slope to stream or along road cuts in part shadein the secondary forests. In pure mats or with
Calypogeia tosana and Pallavicinia subciliata.
J-1-50-13, J-5-10-13.
This species is listed as Solenostoma truncatum (Nees) R.M.Schust. ex Váňa & D.G.Long in
Katagiri and Furuki (2012).
Plectocolea unispiris Amakawa L5, 7, ant.,
per., spor. 110–240 m. Moist to wet cliffs andboulders on slopes and along streams, in partshade to open places in the secondary forests. In
pure mats or with Cavicularia densa and Conocephalum salebrosum. J-5-23-13, J-7-11-13.

A Collection of Hepatics from the Tottori Prefecture, Japan

This species is listed as Solenostoma unispire(Amakawa) Váňa, Hentschel & J.Heinrichs in
Katagiri and Furuki (2012).
Porella densifolia (Steph.) S.Hatt. L1, 7,
60–240 m. Tree trunks on slope to valleys, inpart shade in the secondary forests. In pure mats.
J-1-21-13, J-7-8-13.
Porella japonica (Sande Lac.) Mitt. L7, per.,spor. 240 m. Tree trunks on slope to valley, inpart shade in the secondary forests. In pure mats.
J-7-5-13.
Porella perrottetiana (Mont.) Trevis. L7,
arch. 240 m. Tree trunks on slope to valley, inpart shade in the secondary forests. In pure mats.
J-7-3-13.
Porella vernicosa Lindb. L6, arch. 170 m.
Tree trunks on slope to valley, in part shade in
the secondary forests. Only one collection. In
pure mat. J-6-2-13.
Radula constricta Steph. L1, 3, gemm. 40–60
m. Tree trunks in open place near road acrossrice ﬁelds and in part shade in the secondary forests. In pure mats or with Cololejeunea japonica
and Macvicaria ulophylla. J-1-24-13, J-3-2-13.
Radula japonica Gottsche ex Steph. L2, per.
40 m. Tree trunk in part shade in the secondary
forests. In pure mat. J-2-6-13.
Radula kojana Steph. L2, 5, 7, 40–240 m.
Moist to wet cliffs in part shade along streams inthe secondary forests. In pure mats or with Conocephalum salebrosum, Heteroscyphus coalitus,
Plagiochila sciophila, Pallavicinia cf. subciliata
and Wiesnerella denudata. J-2-11-13, J-5-20-13,
J-7-26-13.
Reboulia hemisphaerica (L.) Raddi ssp. orientalis R.M.Schust. L5, 6, 7, 10, ant., arch., spor.
110–390 m. Moist to wet and mesic stones andcliffs in part shade to in open places along
streams, on slopes to valleys and in the sprayzone of waterfalls in the secondary forests. In
pure mats or with Wiesnerella denudata. J-5-413, J-6-15-13, J-7-20-13, J-10-13-13.
Riccardia chamedryfolia (With.) Grolle L2,6, 10, gemm. 40–390 m. Moist to wet boulders
and cliffs along streams, in the spray zone ofwaterfall, clayish road cuts in mesic conditions,171

in part shade in the secondary forests. In puremats, but mostly mixed with other hepatics, suchas Aneura maxima, Conocephalum salebrosum,
Heteroscyphus coalitus, Jubula hutchinsiae,
Lejeunea aquatica, Makinoa crispata, Megaceros ﬂagellaris, Pallavicinia subciliata, Pellia
neesiana and Wiesnerella denudata. J-2-7-13,
J-6-14-13, J-10-11-13.
* Riccardia kodamae Mizut. & S.Hatt. L2,
per. spor. 40 m. Clayish road cut in mesic conditions in part shade in the secondary. Only one
collection. In pure mat. J-2-8-13.
* Riccardia latifrons (Lindb.) Lindb. var.
miyakeana (Schiffn.) Furuki L1, arch. 60 m.
Rotten log in part shade in the secondary forests.
Mixed with Cephalozia otaruensis and Odontoschisma denudatum. J-1-3-13.
* Riccardia tamariscina (Steph.) Schiffn. L1,
60 m. Wet stone wall in the valley. Wet clay roadcuts and stone walls along streams, in part shadein the secondary forests. In pure mats or with
Calypogeia japonica and Conocephalum salebrosum. J-1-40-13.
Scapania parvitexta Steph. L1, 2, ant., per.
40–60 m. Wet clay road cuts, boulders in thestream beds, stone walls along streams, in partshade in the secondary forests. In pure mats orwith Blepharostoma minus and Diplophyllum
taxifolium. J-1-48-13, J-2-1-13.
Trichocolea tomentella (Ehrh.) Dumort. L7,
240 m. Moist to wet cliffs along stream in partshade in the secondary forests. In pure mats orwith Conocephalum salebrosum, Heteroscyphus
coalitus, H. planus and Pallavicinia subciliata.
J-7-25-13.
Trocholejeunea sandvicensis (Gottsche)Mizutani L2, per. 40 m. Tree branch in open
place in the secondary forest. Only one collection. In pure mat. J-2-3-13.
Wiesnerella denudata (Mitt.) Steph. L2, 5,
10, ant., arch. 40–390 m. Moist to wet cliffs,
boulders (including clay covered) and humus
along streams, on slopes, in the spray zone ofwaterfalls, mostly in part shade in the secondary
forests. In pure mats or with Conocephalum salebrosum, Dumortiera hirsuta, Heteroscyphus

Vadim A. Bakalin et al.

172

coalitus, Jubula hutchinsiae, Lejeunea japonica,
Plagiochila ovalifolia, P. sciophila, Radulakojana, Reboulia hemisphaerica ssp. orientalis
and Riccardia chamaedryfolia. J-2-19-13, J-5-513, J-10-7-13; Higuchi 51320.
Acknowledgments
The laboratory work of V. Bakalin was partially supported by the grants from RussianFoundation for Basic Researches (12-04-91150,
13-04-00775). The bryophyte collections used inthis study were made under a research program,“Research on the Earthʼs surface processes andbiota in and near the Sea of Japan,” by the
National Museum of Nature and Science, Tokyo.
References
Akiyama, H. 2006. Information of the Japanese Conocephalum conicum complex. Bryological Research 9:
88–91 (in Japanese).
Arikawa, T., Hirooka, R., Kawai, K., Tateishi, Y. andNishimura, N. 2011. A checklist of the bryophytes in
Tottori Prefecture, Japan. Bulletin of the Tottori Prefectural Museum 48: 103–118 (in Japanese).
Arikawa, T., Kawai, K., Tamura, E., Tateishi, Y. andNishimura, N. 2012. A list of the bryophytes, collectedin the ﬁeld survey for red list species of Tottori Prefecture, 2010. Natural History Researches of Sanʼin 8:
17–27 (in Japanese).
Bakalin, V. A. and Vilnet, A. A. 2012. New combinations
and new species of Solenostoma and Plectocolea (Sole-nostomataceae) from the Russian Far East. Bryologist
115: 566–584.
Borovichev, E. A., Kalinauskaite, N. and Konstantinova,
N. A. 2009. On the distribution of Conocephalum conicum and C. salebrosum (Marchantiophyta) in Russia.
Arctoa 18: 115–120.
Hattori, S. 1972. Frullania tamarisci-complex and thespecies concept. Journal of the Hattori Botanical Laboratory 35: 202–251.
Heinrichs, J., Hentschel, J., Bombosch, A., Fiebig, A.,
Reise, J., Edelmann, M., Kreier, H.-P., Schaefer-Verwimp, A., Caspari, S., Schmidt, A., Zhu, R.-L., vonKonrat, M., Shaw, B. and Shaw, A. J. 2010. One species or at least eight? Delimination and distribution of
Frullania tamarisci (L.) Dumort. s.l. (Jungermanniopsida, Porellales) inferred from nuclear and chloroplast
DNA markers. Molecular Phylogenetics and Evolution
56: 1105–1114.
Katagiri, T. and Furuki, T. 2012. Checklist of Japaneseliverworts and hornworts, 2012. Bryological Research
10: 193–210.
Kobayashi, R. 2012a. New record of Syzygiella nipponica(S.Hatt.) K.Feldberg et al. from Tottori Prefecture,
Japan. Bryological Research 10: 266–267 (in Japanese).
Kobayashi, R. 2012b. New record of Diplophyllum serrulatum (Müll.Frib.) Steph. from Tottori Prefecture,
Japan. Bryological Research 10: 267 (in Japanese).
Szweykovski, J., Buczkovska, K. and Odrzykoski, I. J.
2005. Conocephalum salebrosum (Matchantiophyta,
Conocephalaceae) a new Holarctic liverwort species.
Plant Systematics and Evolution 253: 133–158.
Yamada, K. and Iwatsuki, Z. 2006. Catalog of the hepatics of Japan. Journal of the Hattori Botanical Laboratory 99: 1–106.