Bull. Natn. Sc i. Mus., Tokyo, Ser. B, 24(4), pp. 137- 146, December 22, 1998

Neodolichomitra y unnanensis (Musci) New to Japan
Masanobu Higuchi 1 and Naoki Nishimura 2
1
Departme nt of Botany, Nationa l Science Museum ,4- 1- 1 Amakubo, Tsukuba, lbaraki, 305- 0005 Japan
2
Hiruzen Branch, Research Institute of atura l Sciences, Okayama Universi ty of Science,
Kawaka mi-son, Maniwa-gun , Okayama, 7 17- 0602 Japan

Abstract Neodolichomitra y unnanensis (Besch.) Kop . is reported as new to Japan.
The description and illustrat ion based on Japanese plants a nd the type specimens
are given. Some characters, including overlooked or neg lected ones, are noted.
Neodolichomitra is co nsidered monospecific and closely related to Rhy tidiadelphus.

Key words:

Neodo /ichomilra yunnanensis , Musci , Japan, Taxono my.

In 1996, during the post foray of the 25th Annual Meeting of the BryologicalSociety of Japan, some of the participants, Mr. Y. Tateishi, Mrs. T. Hiraoka and Mr.
Y. Ashida, collected a robu st moss in central Honshu, Japan. In appearance it is reminescent of some species of Rhytidiadelphus . However, a microscopic examination
confirmed that the pl ants are not any species of that genus, but rather are
Neodolichomitra yunnanensis (Besch.) Kop . This species, previously known from
Taiwan and southern provinces in mainland China (cf. Redfearn et al., 1996), is new
to Japan.
The genus Neodo lichomitra was described by Noguchi ( 1937) based on the new
species, Neodolichomitra gigantea Nog., from Taiwan, and subsequently he (Noguchi1966) made the new combination, Neodo lichomitra robusta (Broth.) Nog. based on
Penzigiella robusta Broth . from Hunan, China. Later Koponen ( 1971) synonymized
N. robusta (Broth.) Nog. with Hylocomium yunnanense Besch. from Yunnan , China,
and he made the new combination, Neodolichomitra y unnanensis (Besch.) Kop . We
have examined the type specimens of these three species, and agree with Koponen's
treatment. We consider the genus Neodolichomitra monotypic and closely related to
Rhytidiadelphus.
Description

Neodo lichomitra y unnanensis (Besch .) Kop ., Hikobia 6(1 - 2): 53, 1971.
Hy locomium y unnanense Besch. , Ann. Sci. Nat. Bot. Ser. 7, 15 : 93, 1892. Type :Yunnan, boi s de Koutoui , au-dessus de Mo-so-yu , (I )7 j anvier 1890, Delavay s. no.
(= no. 4636). Lectotype (fid. Koponen, 1971) in BM! ; isolectotypes in H-BR!, NY!,

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Masanobu Higuchi and Naoki Nishimura

PC!, S-PA not seen. - Rhytidiadelphus yunnanensis (Besch.) Broth., Nat. Pfl . 1(3):1057, 1907.
Penzigiella (?) robusta Broth., Sitzungsber. Akad. Wiss. Wien , Abt. I, 131 : 216,
1922. Type: China, Hunan, Wukang, Yun-schan, 1200- 1300 m, Handel-Mazzetii
11178. Holotype in H-BR!, isotype in PC! - Neodolichomitra robusta (Broth.) Nog. ,Bryologist 69: 232, 1966.
Neodolichomitra gigantea Nog. , J. Jpn. Bot. 13: 92, 1937. Type: Formosa, Prov.
Taihoku, Mt. Taihei, 2200m, on humus in forest, Noguchi 6561. Holotype in NICH! ,isotypes in BM! , PC!
The following description is based on the Japanese plants.
Plants robust in loose green mats. Stems reddish-brown , mostly arcuate, irregularly or regularly pinnate-branched, sometimes bipinnate-branched, more or less frondose from a stipitate base, usually stepwise layered because of repeated annual
growth from an arched-ascending stoloniform innovation, to 6 cm long (the length isthat of the innovation shoot from a single growing season), with a slightly developed
central strand; branches tapered, occasionally radiculose at the tips; paraphyllia absent. Pseudoparaphyllia foliose , wide-triangular, a few. Stem leaves spreading when
wet and dry, deeply concave, smooth, slightly decurrent, broadly ovate to oblonglanceolate, cordate at the base, abruptly or gradually narrowed to a short acumen ,usually twisted at the apex in upper leaves, 2.4- 3.2 mm long, 1.5- 2.4 mm wide at median parts of stem; margings plane or erect, usually slightly recurved at the base,sometimes incurved above, serrulate or serrate near the apex , minutely serrulate
throughout; costa variable, usually single and branched, long (1 /2- 3/4 the leaf
length), sometimes double and shorter, rarely triple or lacking; median cells smooth,linear; basal cells shorter and wider, incrassate, porose, colored; alar cells scarcely
differentiated, porose, colored. Branch leaves similar to stem leaves, smaller, 1.42.0mm long, 0.9- 1.3 mm wide at median parts of branch.
Dioicous. Perichaetia on the main stem (unfertilized perichaetia seen).
Sporophytes not seen.
Specimens examined. Japan. Central Honshu, Nagano-ken, Shimoina-gun,
Oshika-mura, Shiokawa, 1330- 1420 m alt., on humus in deciduous forest at riverside, 26 July 1996, coli. Y Tateishi 8700 (HIRU, TNS) ; Y Ashida 2891 (HIRU); T.
Hiraoka 10030 (HIRU).
Habitat. The site, "Oshika-mura, Shiokawa", is one of the famous areas forcollecting the calcicolous bryophytes in central Japan. According to the collectors,
however, the place where they collected the specimens was somewhat apart from thecalcareous area. They found the species separately at or around the same place before
reaching the calcareous area. The plants were growing on humus-covered, non-calcareous boulders, in a soft mat like Rhy tidiadelphus japonicus, in the deciduous forest of Euptelea poly andra, Pterocarya rhoifolia and Cercidiphy llum japonicum along
the edge of the Shiokawa River.

Neodolichomitra yunnanensis (Musci) New to Japan

139

Fig. I. Neodo/ichomitra y unnanensis (Besch.) Kop. A. A portion of branch. B. Innovation. C- M.
Stem leaves. C- E. Whole leaves. F, G. Apical parts. H. Marginal cells at upper portion. I. Marginal cells at median portion. J. Ma rginal cells at basal portion. K. Median cells. L. Basal parton the stem. M. Basal angle . N, 0. Pseudoparaphyllia. P. Unfertilized peirchaetium. Q. Axillary hairs. R- T. Branch leaves. U. Portion of a cross section of stem. (A, C- U drawn from

Tateishi 8700; B from iso lectotype of Hylocomium y unnanense in PC.)

Masanobu Higuchi and Naoki Nishimura

140

E
E
L____..__

I mm ___.J

~

5

I

N

I

E

0

E

0

oo j
J

F

c

E
E

d

j
D
Fig. 2. Neodolichomitra y unnanensis (Besch.) Kop. A. A plant. B. Peri chaet ium . C. Inner
perichaetial leaf. D. Median cells of inner perichaetia l leaf. E. Ca lyptra. F- H. Capsules (left:when dry, central & right: when wet). I. Operculum. J. Spores. K. Outside view of peristom ewith upper exothecial cells. L. Inside view of peristome. M- P. Exothecial cells. M. Upper portion. N. Median portion. 0. Lower portion . P Basal portion with stom a. (All drawn fromisolectotype of Hy/ocomium y unnanense in PC.)

Neodolichomitra yunnanensis (Musci) New to Japan

141

D

.
0

0

c

J
E

Fig. 3. Neodo lichomitra yunnanensis (Besch.) Kop. A. Stem leaves (left: lower ones, right: upperones). B. Branch leaves (left: lower ones, right: upper ones). C. Apical parts of stem leaves(be low: lower ones, upper: upper ones). D. Variation of costae of stem leaves. E. Portions ofserious cross sections of costa in stem leaves (below: basal portion, upper: upper portion). (A lldrawn from isolectotype of Hvlocomium vunnanense in PC.)

Masanobu Hi guchi and Naoki Nishimura

142

Distribution.

Taiwan, southern provinces in mainland China. New to Japan.
When Noguchi ( 193 7) established the genus Neodolichomitra, the sporophytes
were unknown. Subsequently Koponen ( 1971) and Rohrer ( 1985b) described thesporophytic characters of Neodolichomitra y unnanensis. We examined the sporophytes in the isolectotype specimen of Hy locomium y unnanense Besch. in PC, andfound some previously undescribed characters, such as perichaetial leaves, calyptra,seta twisting and detailed surface ornamentation of the peristome.
The following description is based on the isolectotype of Hylocomium y unnanense Besch. in PC.
Inner perichaetial leaves oblong-lanceolate, filiform-acuminate , reflexed and serrulate above; median cells thick-walled, porose. Calyptra cuculate .
Setae I 0- 16 mm long, reddish-brown , smooth, sinistrorsely twisted in lower halfand dextrorsely twisted at upper part when dry. Capsules inclined to horizontal , reddish-brown , ovoid, 1.5- 2.2 mm long, 1.1 - 1.3 mm thick; exothecial cells oblong toelongate-rectangular, unevenly but distinctly thick-walled; stomata few, confined to
the neck ; annuli present; opercula short-rostrate. Peristome perfect; exostome teeth
lance-subulate, yellow, bordered, cross-striolate or irregularly cross-striolate below,
pale and papillose above and with a weak zig-zag median line on the external surface,smooth below, roughly papillose above and with projecting trabeculae on the internalsurface; endostome pale or hyaline, papillose on the internal surface; segments subulate, keeled, spiculose-papillose, usually narrowly, but sometimes broadly perforated;cilia 2- 3, finely papillose; basal membrane about 1/2 as high as exostome teeth,
densely papillose. Spores spherical , finely papillose, 15- 19 mm in diameter.

Discussion
Japanese plants agree well with the type specimens of the above three names intheir essential characters, but differ slightly in having stem leaves with less serration(Fig. I : H-J). To clarify the relationship between Neodolichomitra and its allied genera, some selected characters, including overlooked or neglected ones, are discussed
as follows.
Our examination on the peristome by SEM showed that the lower part of theouter exostome is cross-striolate (Fig. 4 : E) or irregularly cross-striolate (Fig. 4: F),confirming Rohrer's ( 1985b) observation that it is cross-striolate. As we examined thesporophytes from only the isolectotype of Hylocomium y unnanense Besch. in PC, we
could not judge whether this feature is stable or not. In Hylo comium (s. lat) and Rhytidiadelphus species the ornamentation on the outer surface of the exostome teeth
varies from cross-striolate to irregularly cross-striolate or reticulate (cf. Buck, 1980;
Rohrer, 1985b ).
Rohrer ( 1985b) pointed out that the branching pattern of specimens of Taiwan
were often monopodia!, whereas those of the mainland China were nearly always

Neodo/ichomitra y unnanensis (Musci) New to Japan143

sympodial. We also examined the specimens from Taiwan and the mainland of China,
but we were unable to observe such a difference. In Neodolichomitra the basal partsof plants show the sympodial branching (Fig. 2: A). If the plants lose the older portions from preceding years, then they appear monopodia! in their branching.
The pseudoparaphyllia, observed in the Japanese material, are foliose , wide-triangular, few, somewhat apart from the branch primordia (Fig. I: N, 0), and also
rarely at the base of young branches. Noguchi ( 1972) illustrated narrow-lanceolate
pseudoparaphyllia which are taller than the branch primordia. However we did notdetect such variation from the specimens examined. The cells of the pseudoparaphyllia usually thick-walled. In the type specimen of Penzigiella robusta Broth. , they are
distinctly thick-walled and porose.
In pleurocarpous mosses axillary hairs are easily observed in the leaf axils of
upper portion of young stems. It has been pointed out that the ax ill ary hairs can be ofsome value in the taxonomy of the hypnaceous mosses (Nishimura, 1985 ; Higuchi,
1985 ; Hedenas, 1989), although the number oftaxa examined are limitted. In N. yunnanensis the axillary hairs consist of two well defined portions. A basal portion consists of two (rarely one), short, yellowish-brown cells, and an upper portion of three(rare ly two), long, hyaline cells (Fig. I: Q). In most of hypnaceous mosses examinedby Nishimura ( 1985), Higuchi ( 1985) and Hedenas ( 1989), a basal portion consists of
one colored cell. The structure of axillary hairs in N. yunnaenensis is similar to thatof Rhytidiadelphus loreus (cf. Nishimura, 1985), R. triquetrus , R. squarrosus and R.
loreus (cf. Hedenas, 1989).
The size and shape of the stem leaves are clearly differentiated by their portion
on the stem (Fig. 3: A, C). This feature is one of the unique characters of the genus
Neodolichomitra . The leaves occurring on the lower I /3 of the stem are differentiatedas stipe leaves. They are widely cochleariform, imbricate, and the smallest in size atthe base of the stem, becoming larger in the upper portion. The leaves from the median portion of the stem are the largest, spreading, broadly ovate, and apiculate at apex .
The leaves on the upper portion of the stem become smaller again , oblong-lanceolate,
and acuminate with apices more di stinctly twisted. Among the hypnaceous or hylocomiaceous mosses the differentiation of the lower stem leaves as stipe leaves is also
found in Pleurozium schreberi (Brid.) Mitt. , but they are not so clearly differentiatedas those of Neodolichomitra yunnanensis. The differentiation of leaves noted above isalso seen in the branch leaves (Fig. 3: B).
The development of the costa is one of the important features circumscribing
genera or families in the Hypnobryales (Fleischer, 1923 ; Brotherus, 1925). As pointed out by the preceding authors examining the Neodolichomitra, the costa is one of
the most notable characters of the genus. Noguchi ( 193 7, 1966) suggested that
Neodo lichomitra is near Rhytidiadelphus, but differs in unicostate leaves. Later, however, Noguchi ( 1972) stated that the singleness or doubleness of costa may not be an
extremely reliable character for separating genera. In N. yunnnanensis the costa is144

Ma sa nob u Hi g uch i and Naoki Nishimura

Fig. 4. S EM micrographs of peristome of Neodolichomitra y unnanensis (Besch .) Kop. A. Outside view of peristo me. B. Inside view of exostome teeth . C- F. Outer surface of exostome
teeth (C: apical part. D: middle pa rt. E, F: lower pa rts). G. In side view of endostome. H.
Lower parts of segme nt a nd ci lium . I. Middl e pa rt of segme nt. J. Cilia. K. Basa l membrane. L.
Spore. (a ll from isolectoty pe of Hy locomium y unnanense in PC.)

very variable even in the leaves of the same stem or branch (Fig. 3: D, E), but is basically singl e. The size and shape of the leaf are differenti ated as stated above, which is
independent from the variation of the costa. A similar variable condition of the costa
is also observed in Hy locomiastrum umbratum (Hedw.) Fleisch. ex Broth. (cf.

Neodo lichomitra vunnanensis (Mu sc i) New to Japan

145

Rohrer, 1985b ). However, our examination indicated that the costa of H. umbratum isdifferent from that of N. y unnanensis by being unbranching (at least in stem leaves),
having spines or protuberances on dorsal sides (smooth in N. y unnanensis), and flaton ventral sides in cross section (convex in N. y unnanensis, Fig. 3: E). Ochyra ( 1990)
pointed out that Streimannia turgida Ochyra ( = Bry ostreimannia turgida Ochyra) has
a similar costa, and Neodolichomitra shared many essential characters with theBrachytheciaceae. Although we did not exam ine this species because we did not haveaccess to specimens, some of characters to which he referred are not restricted only
to the Brachytheciaceae. For example, leaves with twisted apices are observed insome species of Ctenidium as we ll as in Hypnum heseleri Ando & Higuchi .
Neodolichomitra has been placed in the Rhytidiaceae (Noguchi, 1937; Redfearn
et al., 1996), Hypnaceae (Noguchi 1966, 1972), Hylocomiaceae (Rohrer, l985a, b;
Buck & Vitt, 1986) or Brachytheciaceae (Ochyra, 1990). However, it is significant
that they consider Neodo lichomitra to be near Rhy tidiadelphus or Macrothamnium,
except Ochyra ( 1990). The familial position of the genus depends on which family
Rhy tidiadelphus belongs to. Neodo lichomitra is characterized by (I) growth patternwith sympodial branching, (2) differentiated leaves into stipe, median stem and upper
stem , (3) variably branched costa, (4) scarecely differentiated alar cells with porosethick-walls, (5) ovoid capsules, and (6) unevenly distinctly thick-walled exothecial
cells. We do not intend to di sc uss the familial position of Neodo lichomitra here, but
we believe that the genus is allied to Rhy tidiadelphus.

Acknowledgments
We wish to appreciate Mr. Y. Tateishi , Mrs. T. Hiraoka , and Mr. Y. Ashida for
their kind offer of the specimens. Thanks are also due to the curators of the herbaria(BM, H, NI CH, NY, PC) for loan of specimens, and Dr. W. R. Buck for reading themanuscript and giving valuable comments. The first author is g rateful to Prof. Dr. A.
Coute for hi s kindness in providing research facilities during his stay in 1998 at theLaboratory ofCryptogamy, the National Museum ofNatural History, Paris, with helpof a Grant-in-Aid for Overseas Research Fellows (no. 9-293) from the Ministry of
Education , Science, Sports and Culture, Japan . The second author is grateful to the
National Science Museum , Tokyo, for inviting him as an inviting researcher of 1998
and providing him research facil ities.

References
Brotherus, V F. , 1925. Hypnob rya les. In Eng ler & Prantl (eds.), Die Natu rli c hen Pftan zenfamilien , ed. 2,
bd. 11 • 278-487. Leipzig.
Fleischer. M., 1923. Die Musci der Flora von Buite nzorg, 4. pp. I- XXX I, 11 05- 1729. Leiden.
Buck, W. R., 1980. A generic revi sion of the Entodontaceae. J Ha11ori Bot. Lab., 48 • 7 1- 159.
Buck. W. R. and D. 1-1. Vitt, 1986. Suggestions for a new fam ili al c lassificat io n of pleurocarpous mosses.

146

Masanobu Hi g uchi and Naok i Nishimura

Taxon, 35(1 ): 21 - 60.
Hedeniis, L., 1989. Axi ll ary hairs in pleurocarpous mosses- a comparative study. Lindbergia, 15: 166180.
Hedeniis, L., 1995 . Higher taxonomic leve l relationships among diplolepidous pleurocarpous mosses a cladi stic overview. J BIJ'O!., 18: 723- 78 1.
Hi guchi , M. , 1985. A taxonomic revision of the gen us Gol/ania Broth. (M usci). J Hattori Bot. Lab. , 59 :1- 77.
Koponen , T. , 197 1. The taxonomic status of Rhy tidiade/phus y unnanensis. Hikobia 6( 1- 2): 53- 56.
Nishimura, N., 1985. A revision of the ge nus Ctenidium (M usc i). J Hallori Bot. Lab., 58: 1- 82.
Nishimura, N. , M. Hi guchi , T. Seki and H. Ando, 1984. Delimitation and subdivisio n of the fam ily Hypnaceae . J Hal/ori Bot. Lab., 55 : 227- 234.
Noguchi , A., 193 7. Contributions to the moss fl ora of Japan and Formosa, 6. J Jpn . Bot., 13 : 86- 95 .
Noguc hi , A. , 1966. Notes on th e moss genus Neodo/ichomitra. Nog. Bryologisr, 69: 232- 234.
Noguchi , A. , 1972. On the delimitation of the ge nera of Hy locomi aceae and Rhytid iaceae. J Hartori
Bot. Lab. , 35: 155- 168.
Oc hyra, R., 1990. Streimannia turgida, a new gen us and spec ies of Brachytheciaceae from Austra li a. J
B1y ol., 16: 223- 229.
Redfearn , P. L. , Jr., B. C. Tan and S. He, 1996. A newly updated and annotated check li st of Chinesemosses. J Hallori Bot. Lab. , 79 : 163- 357.
Rohrer, J. R. , l985a. A phenetic and phy logenetic analys is of the Hy locomiaceae and Rhyt idi aceae. J
Hallori Bot. Lab. , 59: 185- 240.
Rohrer, J. R., 1985 b. A generic rev is io n of th e Hylocomiaceae. J Hallori Bot. Lab., 59 : 241 - 278.