Hattoria 12: 1–7.

2021

Lejeunea masamiana (Lejeuneaceae), a new liverwort species

from Indonesian New Guinea
1, 2 3
Gaik Ee LEE & Tamás PÓCS

1
Faculty of Science and Marine Environment, 21030 Kuala Nerus, Universiti Malaysia Terengganu,
Terengganu, Malaysia
2
Institute of Tropical Biodiversity and Sustainable Development, 21030 Kuala Nerus, Universiti
Malaysia Terengganu, Terengganu, Malaysia
3
Eszterházy University, Institute of Biology, Botany Department, Eger, Pf. 43, H-3301 Hungary
Author for correspondence: Gaik Ee LEE, gaik.ee@umt.edu.my

Abstract A new liverwort species, Lejeunea masamiana G.E.Lee & Pócs, is described and illustrated
from Indonesian New Guinea. It is recognised by the strongly recurved to involuted leaves both in dry
and moist condition, leaf cells with well-developed trigones and with conspicuous intermediate
thickenings, smooth cuticle, fully incurved free margin of the lobule, and large, reniform underleaves
with frequently recurved margins and with lobes up to 1/4 of underleaf length. Morphological variations
of some liverwort species due to environmental effect of the forest in New Guinea are discussed, as a
factor which might have influenced also the morphology of the new species.

Introduction
New Guinea is politically divided into two separate regions, i.e. Papua New Guinea and
Indonesian New Guinea (Fig. 1). The vascular plant flora of New Guinea is widely known
with its remarkable diversity higher than the Madagascar and Borneo, and it also rank among
the most biodiverse island on Earth (Cámara-Leret et al. 2020). As compared to vascular plant,
the inventory has been less detailed and in general extremely fragmentary for bryophytes. It
appears that in New Guinea, bryophyte exploration has been more intensive or bryologically
well known in the eastern (Papua New Guinea) than in the western (Indonesian New Guinea)
part of the region. For example, the liverwort species recorded in Papua New Guinea was 557,
which has 63% more species than Indonesian New Guinea, with 342 species (Grolle & Piippo
1984). Reasons maybe due to the lower collecting density in Indonesian New Guinea, a
scenario similar to vascular plant, and the relatively larger areas and good road infrastructure
on some of the highlands, e.g. road to Mt. Kaindi fide Grolle & Piippo (1984), in Papua New
Guinea. Furthermore, much collecting has been done for the bryoflora of Papua New Guinea
due to the Finnish-American expedition in 1981 between University of Helsinki, Academy of

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Figure 1. Map of New Guinea showing two separate regions; Papua New Guinea and Indonesian New
Guinea. The star indicates the locality where the type specimen of the new species was collected.

Finland and Humboldt State University (California) which yielded enormous amount of
specimens. Since then, numerous papers (a series of about 80 chapters) on the liverwort
species have been published resulting from the joint expedition.
The first comprehensive checklist from Western Melanesia which include New Guinea
and Solomon Islands was made by Grolle & Piippo (1984). In their checklist, 28 species of
Lejeunea were reported with the highest representation from Papua New Guinea, with 22
species, followed by Indonesian New Guinea and Solomon Island with 6 and 2 species,
respectively. Moreover, 18 species of Lejeunea in the checklist were not specified to which
regions and had been reported as Nova Guinea or New Guinea , mostly by Stephani (1914–
1915) and Bischler et al. (1961). Several additions were also added from the Huon Peninsula,
Papua New Guinea with a total of 16 species of Lejeunea which had been enumerated by Pócs
et al. (1994, 1995, 2019). Recently, four new species of Lejeunea viz. L. streimannii Y.M.Wei
& R.L.Zhu, L. heinarii G.E.Lee & Pócs, L. madangensis G.E.Lee & Pócs, and L.
marginedentata G.E.Lee & Pócs, have been published, in which the latter two type specimens
were collected during the expedition in 1981, as well as new combinations [L. giulianettii
(Steph.) G.E.Lee & Heinrichs, L. tjibodensis (Steph.) G.E.Lee & Heinrichs] and new variety
[L. pulchriflora var. nymannii (Steph.) G.E.Lee & Heinrichs] reported from Papau New
Guinea (Lee et al. 2018; Wei et al. 2018; Lee et al. 2020). In this paper, we report another new
species of Lejeunea from Indonesian New Guinea. Even though this species is without mature
perianth, it has most of the distinctive characteristics of the genus Lejeunea: ocelli absent,
ventral merophyte 2 cells wide, stem with 7 epidermal cells (thin-walled and larger than
medullary cells), hyaline papilla at the proximal side of the first tooth, branches of the
Lejeunea-type, and gynoecia with lejeuneoid innovations. Several distinguishing characters of
this species including the strongly recurved to involuted leaves, the small leaf lobules with

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fully incurved free margin, and the reniform, frequently recurved underleaves, however, did
not match with any of the Lejeunea species known so far, and hence we describe it here as
new species, L. masamiana, in memory of the late Dr Masami Mizutani.

Materials and Methods

The new species of Lejeunea masamiana was described and illustrated based on the
specimen kept in Herbarium Haussknecht (JE, Zanten 716a). This plant was first examined
under an Olympus SZ51 stereomicroscope both in dry and wet condition (moistened with tap
water), and then detailed morphological characteristics of the species were examined using
Leica DM1000 microscope. The illustration of the type specimen was produced using the
same compound microscope equipped with a drawing tube.

Taxonomy
Lejeunea masamiana G.E.Lee & Pócs, sp. nov. Fig. 2

Diagnosis: The species is similar to the Papua New Guinean Lejeunea giulianettii in the
relatively large size, the strongly recurved leaves, the broadly rounded leaf apices, the absence
of a large disc cell of lobule, and the large, distantly arranged underleaves. The latter species,
however, differs in the presence of two intermediate thickenings in basal leaf cells between
adjacent trigones, the flat free margin of leaf lobules, and the finely punctate-papillose cuticle.

Type: INDONESIA. Western New Guinea, Antares Mts., Camp 39a, 1,500 m elev., 1959,
Zanten 716a (holotype: JE).

Description: Plants dioicous?, 1.8–2.5 mm wide, irregularly and slightly, seldom

bipinnately branched, branches erect-spreading to spreading, with three rather large basal
collar lobes. Stem 0.15–0.20 mm in diameter, about 7 cells high in cross-section; epidermal
cells 38–63 µm wide, medullary cells 15–17, 13–25 µm wide, ventral merophyte 2 cells wide.
Leaf lobes ovate-orbicular, 1.0–1.3 mm long, 0.8–1.0 mm wide (when flattened), strongly
imbricate; leaf apex broadly rounded, strongly recurved to frequently involute when in dry or
moist condition; leaf margins entire; the ventral margin forming an angle of 100°–130° with
the keel when flattened; insertion 12 lobe cells long. Leaf cells rather uniform, gradually
becoming smaller towards the leaf margin, usually round, irregularly quadrate to rectangular
towards the leaf margin; apical cells 13–25×13–25 µm, median cells 25–38×13–25 µm,
basal cells 38–50×25–33 µm; cell walls hyaline, with well-developed (relatively large)
trigones and infrequently with intermediate thickenings, 1–2 per cell, 0–1 between 2 adjacent
trigones. Cuticle smooth. Oil bodies not seen. Leaf lobules relatively small, sometimes
reduced, 0.15–0.20 mm long, 0.10 mm wide, to 1/5 the length of the lobe, at an angle of
60°–70° to the stem, ovate to oblong, inflated along the keel; apex obliquely truncate; keel
somewhat straight to curved; free margin incurved fully; first tooth 38–45 µm long, oblong,
somewhat erect, apex obtuse; margin between tooth and sinus 3 cells long, large disc cell (cell

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Figure 2. Lejeunea masamiana G.E.Lee & Pócs. A: Part of plant in ventral view. B–C: Leaves. D–F:
Underleaves (flattened in F). G: Cross-section of stem. H: Apical leaf cells of leaf lobe. I: Stem portion
and leaf lobule. J: Upper part of leaf lobule when flattened (hyaline papilla shown in gray). K–M: Cross-
section of leaf lobe. N: Marginal leaf cells. O: Female bracts and bracteole. P–Q: Female bracts. R:
Median leaf cells. All figures drawn from the holotype, Zanten 716a (JE).

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below the first tooth) lacking, 25–33 µm long, 20–25 µm wide. Underleaves reniform (wider
than long), (0.3) 0.40–0.45 mm long, (0.5) 0.6–0.8 mm wide, to 3–4 times wider than the
stem, contiguous to distant; covering the leaf lobules; bilobed, lobes to 1/5–1/4 of underleaf
length, about 9–12 cells wide, triangular, sometimes connivent to overlapping; sinus narrow,
acute, V-shaped; tips acute to obtuse; underleaf margin entire, recurved; two large basal
underleaf cells differentiated; base ± cuneate, insertion line curved. Gynoecia on short or long
lateral branches, female bracts somewhat crowded, with one innovation, with 1–2 gynoecia in
a lateral position. Female bracts smaller than the leaf, erect-spreading when moist. Lobes
ovate to somewhat oblong, 0.75–0.80 mm long, 0.5–0.7 mm wide, apex broadly rounded,
margin entire. Lobules oblong, 0.3–0.4 mm long, 0.05–0.08 mm wide, rarely reduced, 1/10 the
width and 1/3–1/2 the length of the lobe, apex obtuse; keels straight, smooth, 0.10–0.25 mm
long. Female bracteoles ovate, 0.6 mm long, 0.45 mm wide; bilobed, lobes to 1/4–1/3 of
female bracteole length, overlapping, sinus narrow, acute; margin entire. Androecia, perianth,
sporophyte and vegetative propagation not seen.

Etymology: The species is named in honor of the late Dr. Masami Mizutani, an eminent
hepaticologist, in recognition of his significant work on the family Lejeuneaceae.

Discussion
Lejeunea masamiana is easily recognised by 1) the frequently involute leaf lobes when
dry or moist, 2) the broadly rounded leaf apices, 3) the leaf cells with relatively large, well-
developed trigones and conspicuous intermediate thickenings, 4) the smooth cuticle, 5) the
small, ovate to oblong leaf lobules with fully incurved free margin, 6) the absence of large disc
cell (below the first tooth of lobule), and 7) the rather large, reniform, frequently recurved
underleaves with lobes to 1/5–1/4 of underleaf length. The holotype of L. masamiana was
initially determined by Mizutani as Taxilejeunea guilianettii (＝ L. guilianettii), however, the
former species differs from the latter by several distinctive characteristics of which some
already mentioned in the diagnosis. Differences between the two species are summarized in
the Table 1 including the morphology of underleaves and leaf lobules.

Morphological variations due to environmental effect in the forests of New Guinea

Based on the authors observation, many liverwort species in New Guinea were found
growing in a more luxuriant form with sharper dentition, longer cilia, stronger recurvation, and
larger size due to the environmental effect of the perhumid microclimate in some New
Guinean rainforests. For examples in the genus Lejeunea, 1) the male bracts of L. dimorpha
Kodama are up to 30 pairs in New Guinea while only 3–6(10) in other Malesian specimens; 2)
vegetative propagation by specialized caducous branches with numerous marginal rhizoids in
L. dimorpha are up to 18 pairs of leaf lobes with 15 large underleaves in New Guinea while
only 3–5(10) pairs of leaf lobes and 3(7) underleaves in other localities; 3) underleaf bases of
L. kinabalensis Mizut. in New Guinea are prominently auriculate instead of cordate in
comparison to Bornean L. kinabalensis; and 4) the margin of leaf lobes and underleaf bases of
L. lumbricoides (Nees) Nees in New Guinea are strongly crenulate and inconspicuously

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Table 1. Differences between Lejeunea masamiana G.E.Lee & Pócs and L. giulianettii (Steph.)
G.E.Lee & Heinrichs.
Characters L. masamiana L. giulianettii
Leaf cells Smooth Rough, with finely
punctate cuticle
Intermediate thickenings at the basal 0–1, never 2 1–2, frequently 2
cells between adjacent trigones
Shape of underleaves Reniform Ovate
Lobe of underleaves (compared to Connivent to overlapping Divergent (1/2)
underleaf length) (1/5–1/4)
Size of leaf lobules (mm length×mm Small (0.1–0.2×0.1) Large (0.4–0.5×0.2–0.3)
wide)
Free margin of leaf lobules Incurved fully Flat
Position (angle) of the leaf lobules to 60°–70° 30°–45°
the stem

auriculate, respectively while other specimens are entire and cordate-auriculate.

Another example is the case of Jubula hutchinsiae (Hook.) Dumor. The teeth within J.
hutchinsiae are variable in length, subsp. hutchinsiae has none or short, subsp. javanica
(Steph.) Verd., which is widespread from Transcaucasia and Himalaya throughout Southeast
Asia, Indomalesia and Australasia to the Pacific (Guerke 1978), can have already longer spines
on lobe margin. It is widespread also in New Guinea, from where a new species: J.
blepharophylla Grolle was described with very long, uniseriate marginal ciliae (Grolle 1965).
It looks to be a distinct species but we have found specimens from New Guinea in Helsinki
Herbarium, which are transitional between the two taxa concerning the cilia length of lobe
margin. Therefore, we suppose that J. blepharophylla falls within the ecological variation of J.
hutchinsiae and is already isolated as a sympatric taxon. In the latter case probably merits only
subspecies rank.

Acknowledgements
We are very grateful to the curator, Dr Hans-Joachim Zündorf of Herbarium Haussknecht
(JE) for sending, on loans, many specimens and unidentified collections of Asian Lejeunea in
which several new species have been discovered including the new species in this paper. We
are also grateful to the director of Hattori Botanical Laboratory, Dr Tomoyuki Katagiri for the
invitation to contribute to this volume in memory of the late Dr Masami Mizutani. Financial
support by the Ministry of Education (MOE) Malaysia through Fundamental Research Grant
Scheme (FRGS/1/2018/WAB13/UMT/03/1 to G.E.Lee) is gratefully acknowledged.

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manuscript received March 15, 2021; accepted April 14, 2021