Corresponding author: Vadim A. Bakalin (
Academic editor: Peter de Lange
The eastern part of the southern macroslope of the Himalayan Range, Hengduan Mountains and the complex of smaller ranges from Hengduan southward to northern Indochina is one of the taxonomic hotspots of
Bakalin VA, Klimova KG, Nguyen VS (2020) A review of
The basic and inevitable tasks for progress in the knowledge of
The vast majority of
1) Taxa undoubtedly occurring in the study area (e.g., if the type specimen is from there).
2) Doubtful records of taxa that have a very low probability of being observed there.
3) Taxa that are not recorded in the study area but may be expected. We treat the latter definition very broadly, as involving some taxa from as far as India and Japan that look reasonable taking into account, e.g., the undoubted occurrence of
4) A dichotomous key to the taxa observed (also including doubtful records) and expected in the study area.
Each taxon in the taxonomic section is annotated as usual, with data on studied type specimens, morphological description based on the type and other comments on morphology or ecology. When providing the distribution of taxa, we do not limit the data to the study area only, but largely also include data from other, nearby regions or areas that have distinct relations in mountain flora with the Sino-Himalaya, e.g., the mountain flora of Taiwan.
The valuable problem of the present work is the inability to evaluate the morphological variation parameters clearly for many species; since there are only a few specimens known (e.g.
It is quite difficult to describe the ‘Sino-Himalaya’ using definite terminology. In very general terms, it is a large territory including the Himalaya Range with some spurs as well as mountain ranges in Southwest China, where it generally includes the Hengduan Shan – a very unclear term for the large mountain massif stretching from the Tibetan Plateau to the southeast until intersection with the mountainous northern end in Indochina. Despite the unclear definition, “the biogeographic unit informally known as the ‘Sino-Himalayan region’” (
The eastern Sino-Himalaya is identified here as the land included in the Sino-Himalaya eastward of eastern Nepal. The Meta-Himalaya is identified as an area surrounding the southeastern part of the Sino-Himalaya, although not belonging to the Sino-Himalaya in its common sense. It includes eastern Sichuan, western Guizhou, eastern Yunnan and the mountains of northern Indochina. This is an area where Sino-Himalayan species deeply penetrate, although sometimes represented by transformed races or the speciation derivates of species status (liverwort examples are in
The area treated in this work covers four floristic provinces in the sense of
1) Sikang-Yunnan floristic province that covers western Sichuan, western Yunnan, northeastern Myanmar, northern Laos and northwestern Vietnam (including the Hoang Lien Range).
2) North Burma (= Myanmar) floristic province.
3) East Himalayan province, including eastern Nepal (excluding low elevations with tropical vegetation), Darjeeling, Sikkim, Bhutan, the Assam Himalaya and the southern and southeastern flanks of Xizang (Tibet), where the monsoon climate is still pronounced.
4) Eastern part of the central Chinese province (western Guizhou, eastern Sichuan and eastern Yunnan).
We do not include Chinese Tibet (Xizang) to the eastern Sino-Himalaya as was done, e.g., by
To identify the general character of the vegetation in the study area, this is alpine vegetation in forest-free landscapes at high elevations extending down to the vegetation developed above tropical communities, starting from the mountain subtropics. The Sino-Himalaya is dominated by a strong monsoon climate and has a distinct cool season with at least occasional snowfall, even at the southern extremes, such as peaks of the Hoang Lien Range. The area under consideration is depicted in Fig.
The area considered in the present paper.
India. Sikkim: 12000 ped. alt. (4000 m a.s.l.), J.D. Hooker, no. 1319 (isotype: G [G00064244/5286!]).
This taxon is broadly Sino-Himalayan-Taiwan-Japanese endemic, distributed within this large area quite disjunctively (although this may reflect the data deficiency). It was described from Sikkim (
Japan. Ozoresan: 11 October 1902, U. Faurie, 1181 (Lectotype (designated here): G [G00067716!]).
The species was described in Japan (
Java. Prof. Stahl (Lectotype (designated here): G [G00061103!]).
The species was originally described from Java (
The description based on the lectotype of
Montagne (holotype: STR [(n. 163)!]).
The species is described in “südlichen Frankreich, auf der Erde” (
Within East Asia,
China. Yunnan: Hokin Delavay, no 1623 (Lectotype (designated here): G [G00061105/10811!]).
The description based on the lectotype is as follows: plants greenish brownish to grayish brown, 1.5–2.1 mm wide, translucent, slightly glistening; stem ~2500 µm wide, branching not seen; rhizoids common, in brownish fascicles erect to upward obliquely spreading; leaves obliquely inserted, subhorizontally oriented, overlapping 1/2 of the next leaf in the base, loosely concave to almost planar, with apex slightly turned to dorsal side, not or for 1/3 of stem width decurrent, 800–1200 × 800–1200 µm, obliquely widely ovate-triangular, apex acute, never divided; underleaves appressed to the stem to obliquely spreading, 1.8–2.5 as wide as stem, decurrent for 1/2–2/3 of stem width, divided by U- to V-shaped sinus into two lobes without additional teeth, lobes obtuse, undivided portion 3–5 cells high; midleaf cells 40–50 × 40–68 µm, thin-walled, trigones small to very small, concave, cuticle virtually smooth.
Japan. Saitama Prefecture: Kuroyama, 500 m a.s.l., 24 June 1968 H. Inoue 18004 (holotype: TNS [174361!]; isotype: G [G00114896!]).
Morphologically, the taxon is similar to
The description based on type specimens is as follows: plants green, strongly glistening, translucent, 1.5–2.1 mm wide, 1–3 cm long; stem greenish, soft, 220–320 µm wide, sparsely ventrally branched; rhizoids sparse to common, in unclear loose fascicles, obliquely spreading, grayish; leaves contiguous to subimbricate (overlap 1/3 of above situated leaf), very obliquely to subhorizontally inserted, slightly convex, apical third turned to ventral side, not or shortly decurrent, when flattened -– obliquely triangular-ovate, 900–1000 × 900–1000 µm, very shortly incised or apex apiculate; underleaves obliquely spreading, decurrent for 1/3–2/3 of leaf length, commonly bisbifid, divided by U-shaped sinus, undivided area 2–3 cell high, 250–300 × 550 µm, 1.1–1.6 as wide as stem; cells in the midleaf thin-walled, with vestigial trigones, 32.5–52.5 × 30.0–37.5 µm, cuticle smooth.
Japan. “Japonia, Uematsu” (neotype by
For a long time regarded as a Japanese endemic species, it was later reported from Fujian (
The description based on the neotype is as follows: plants 1.8–2.0 mm wide, yellowish brownish, merely soft, loosely translucent; stem 220–280 µm wide, sparsely ventrally branched rhizoids common, but not numerous, in loose, obliquely to erect spreading fascicles or separated; leaves obliquely inserted and oriented, slightly concave-canaliculate, contiguous, to slightly overlapping above situated leaves, somewhat loosely crispate along margin, widely ovate-triangular, apex obtuse to narrowly rounded, 1000–1130 × 1000–1250 µm; underleaves obliquely spreading, decurrent for 1/3–2/3 of stem width, divided by V- to U-shaped sinus into two lobes without additional lateral teeth, undivided zone 4–6 cells high, 2.5–3.0 as wide as stem; midleaf cells 25–55 × 25–35 µm, thin-walled, trigones small to very small, cuticle smooth.
India. Assam: Griffith (syntype: G [G00064229/5288!]).
This is a broadly Sino-Himalayan endemic species that seems locally abundant in the eastern Sino-Himalaya.
The description based on the isotype is as follows: plants brownish to blackish brownish in the herbarium, translucent, glistening, 1.5–2.2 mm width; stem 120–220 µm wide, branching not seen; rhizoids sparse to common in brownish grayish, erect to obliquely spreading loose fascicles; leaves overlapping ~1/4–1/3 of next leaf basal part, slightly convex, with apices somewhat turned to ventral side, obliquely inserted and oriented, ventrally clearly decurrent to 1.0 of stem width or less, widely triangular-ovate, apex acute to obtuse (very rarely bilobed), 850–1200 × 770–1200 µm, margin entire to somewhat crispate; underleaves decurrent for (0.3–)0.5–1.0 of stem width, 1.5–3.5 as wide as stem, bisbifid or with each main lobe divided into three small lobes, or bisbifid with additional lateral tooth on each side; midleaf cells thin-walled, trigones vestigial, cuticle smooth, 30–38 × 20–33 µm (the cell measurements may be incorrect because of collapsed leaf cells).
The species is most morphologically similar to
One more observation should be made on the type specimen identification. The specimen in JE marked as the possible type (JE-H2316 = JE04005930!) is actually not the type. The label means that the specimen was collected in “Khasia, Churra”, not in Upper Assam, as in the original description by
China. Guizhou Province: Duyun Municipality (
The species was described in Guizhou Province, China, and confirmed in northern Vietnam (
Japan: Tosa Makino (LECTOTYPE (designated here): G [G00047274/26013, packet b!] The holotype should be in ‘herb. Polytechnicum Zurich’, but such specimen is absent in Zurich herbaria (
This is a widely amphi-Pacific East Asian species whose area stretches from the southern Kurils and East Manchurian mountains in Russia via the Korean Peninsula and Japanese Archipelago to southeastern China, namely, Taiwan (
There is a problem with the type of plants in the type specimen due to mixture within. The type specimen (Makino 25, G), as correctly noted by T. Furuki
The brief description based on the plants belonging to
The plants in
Vietnam. Lao Cai Province: SaPa District, San Sa Ho Commune, Hoang Lien Range, Phan Xi Pang Peak area,
This taxon was recently described in moist rocky outcrops at the highest elevation in Indochina – Phan Xi Pang Mt. – in the somewhat unique formation of a ‘mossy’
Not seen.
There are several records of this taxon in East and Southeast Asia.
Nevertheless,
Java. K. Goebel (syntype: G [G00115804!]).
The species was described from Java based on K. Goebel specimen (
The description based on the isotype is as follows: plants brownish, pellucid, glistening, 1.5–2.5 mm wide, 5–8 cm long; stem 150–200 µm wide, sparsely ventrally branched; rhizoids brownish, common to numerous, obliquely to erect spreading fascicles; leaves contiguous to somewhat distant, slightly convex, decurrent for 1–2 stem widths, 750–1250 × 575–1050 µm, divided by U-shaped sinus into two acute lobes; underleaves, obliquely spreading, 1.5–2.5 as wide as stem, bisbifid, the undivided portion in the underleaf middle 2 cells high, arcuately inserted, not or barely decurrent; midleaf cells thin-walled, trigones very small, concave, 37.5–55.0 × 25.0–37.5 µm.
Czech Republic, Bohemia, Schiffner; not seen.
This boreal species was originally described from the border between the Czech Republic and German Bavaria (Bohemian Forest) and was found to have circumpolar distribution in the hemiarctic and boreal zones of the Northern Hemisphere. However, even in the hemiboreal zone of East Asia (e.g., in the southern Russian Far East at 43–48°N), the species rarely occurs.
Italia, Rive Valsesia; not seen.
The species was described from the Italian Alps (
Sweden, Dalarne; not seen.
There are several records of this rejected name.
Japan. Tokyo: Asakawa Experimental forest, 12 June 1954, U. Mizushima, no. 5 (holotype: TNS [TNS-174359!]; isotype NICH [NICH-55582!]).
The species is regarded as Japanese endemic (known in Honshu only, cf.
The description based on the holotype is as follows: plants pale brownish in herbarium, prostrate, translucent, slightly glistening, 1.1–1.5 mm wide and 8–20 mm long, forming loose mats; rhizoids rather numerous, originating as several unclear fascicles near underleaf bases and obliquely to erect spreading, attaching plants to the substratum; stem brownish (in the herbarium), 100–160 µm in diameter, branching not seen; leaves subhorizontally inserted, dorsally insertion line subtransverse to loosely arcuate, ventrally decurrent for 1/2–2/3 of stem width, obliquely lingulate, to obliquely ovate-lingulate, slightly convex to planar, slightly undulate along margin, laterally spreading, 650–700 × 400–550 µm, leaf apex rounded; underleaves loosely sinuately to transversely inserted, shortly decurrent (up 1/3 of stem width), 120–170 × 200–220 µm, bilobed by U-shaped sinus descending to 2/3 of leaf length, undivided zone 1–2 cells, lobes in the base 3–5 cells wide, midleaf cells oblong, thin-walled, 32–62 × 20–33 µm, trigones vestigial, cuticle virtually smooth; cells along leaf margin subquadrate to oblong, 20–38 µm, thin-walled, with very small concave trigones, cuticle smooth.
Sri Lanka. Central Province: Nuwara-Eliya, 1950 m a.s.l., 24–27 February 1954, F. Schmid 10334 (isotype: G [G00064248!]).
The description based on isotype plants is as follows: plants yellowish brownish in herbarium, glistening, translucent, 2.2–3.5 mm wide 3–5 cm long; stem 370–450 µm wide, branching not seen; rhizoids in loose colorless to brownish fascicles, sparse to numerous; leaves obliquely inserted, slightly concave or convex, somewhat turned to ventral side, not or barely decurrent ventrally, obliquely ovate, well developed 560–670 × 450–550 µm, apex acute to (rarely) unclearly and very shortly bidentate; underleaves obliquely spreading, 1.1–1.3 as wide as stem, decurrent for ¼–1/3 of stem width, divided by V- to U-shaped sinus into two lobes, lateral teeth absent or present and unclear, undivided portion 2–3 cells high; midleaf cells thin-walled, trigones very small to vestigial, 35–80 × 35–58 µm, cuticle smooth.
Hawaii, Heller 2308 (LECTOTYPE (designated here): G [G00069713!] there are no other known authentic materials for this taxon in G).
The species was described from Hawaii and is somewhat morphologically similar to Indochinese-Malesian
The status of
Due to plant features in the type specimen
Another possible synonym of
The species status is seriously doubted by
Taiwan (Formosa). Mt. Morrison, August 1932, Y. Horikawa, no. 9124; not seen.
This is a Taiwan endemic species (
Thailand. Nakawn Sritamarat: Mt. Khao Luang, M. Tagawa & N. Kitagawa (holotype: KYO [T4737]); not seen.
The taxon is known only from the type that is from southern Thailand (
Germany. Saxonia: July 1888, F. Stephani (lectotype, designated by
This is a generally boreal circumpolar species widely spreading to hemiarctic and hemiboreal zones and southward in corresponding belts in the mountains (especially in Japan, although surprisingly not known in China and the Korean Peninsula). The description based on the lectotype is as follows: plants 2.2–3.0 mm wide, soft, greenish to yellowish greenish, loosely translucent; stem 200–300 µm wide, freely ventrally branched with 1–2 branches from one underleaf sinus; rhizoids common, in obliquely spreading brownish fascicles; leaves very obliquely inserted, not or barely decurrent ventrally, contiguous to overlapping 1/4 of above situated leaf in the leaf base, slightly convex to nearly planar, ovate to obliquely ovate, 1200–1900 × 1000–1500 µm, with rounded apex; underleaves appressed to the stem, retuse to emarginate at apex, 1.7–2.5 as wide as stem; midleaf cells thin-walled, trigones vestigial, cuticle smooth to very finely verruculose, 37.5–70.5 × 32.5–55.0 µm.
India. Meghalaya: East Khasi Hills, Langkyrdum-Dawki Road, 07 Nov 1998, V. Nath et al. (holotype: LWG [206109-A]; not seen).
Philippines. Luzon, Merril (LECTOTYPE (designated here): G [G00061102!]).
The species was described from the Philippines (“Luzon”) and is very similar to the Meta-Himalayan
There are two original specimens of the species in G. Both represent the parts of one original specimen (one was probably scheduled to be preserved in the Stephani herbarium, and the other should be returned to the collector) of which we prefer to select G00061102 as the lectotype because the second one (G00061101) has no original label (and is probable a duplicate).
The description from the lectotype is as follows: plants pale brownish (perhaps were bluish green when fresh), glistening and translucent, 1.5–2.0 mm wide; stem 180–230 µm wide, branching not seen; rhizoids common to numerous in brownish, loose, obliquely spreading fascicles; leaves contiguous or overlapping to 1/2 of leaf width, obliquely inserted and oriented, slightly convex, with apex commonly turned to ventral side, not decurrent, 800–1000 × 800–1000 µm, widely triangular-ovate, shortly bifid at the apex; underleaves obliquely spreading, decurrent for 1/3–1/2 of stem width, 1.2–2.2 as wide as stem, mostly bisbifid or with lateral tooth on one or on both sides, rarely bifid; cells in the midleaf 30–40 × 25–35 µm; nearly thin-walled, trigones small, cuticle nearly smooth in the leaf middle to very finely verruculose near leaf apices.
India. Khasia 1849 Hooker, no. 1339 (syntype: JE [JE-04005904, =JE-H4084!]; syntype: G [G00113555/5289!]).
The description based on the syntype G00113555/5289 is as follows: plants brownish to greenish brownish, slightly translucent and glistening, 2.0–2.5 mm wide; stem 140–200 µm wide, sparsely ventrally branched; rhizoids virtually absent or solitary, obliquely spreading; leaves contiguous to overlapping 1/3–1/2 of the leaf base of the next leaf, nearly planar to slightly convex, subhorizontally inserted and oriented, shortly or up to 1/2 of stem width decurrent, widely obliquely ovate to rounded-lingulate, with rounded apex, 1100–1300 × 1000–1300 µm; underleaves appressed to the stem, decurrent for 1/2–2/3 of stem width, divided mostly by very narrow V-shaped sinus into two lobes without additional lateral teeth or shortly bisbifid, with rounded to obtuse lobes, undivided portion 3–6 cell high; midleaf cells 30–65 × 17–37 µm, thin-walled, trigones small, cuticle smooth; marginal cells considerable larger and elongate along leaf margin, 70–80 µm long, with thickened external wall.
Figure
Japan. Tochigi Prefecture: Nasu, 700 m a.s.l., August 1968, Empress Nagako (holotype: TNS: TNS-174632!] ; isotype: G [G00064238!]) .
The taxon is currently known from Japan only (Honshu). It was recently synonymized with
The description based on the holotype is as follows: plants merely soft, glistening and translucent, greenish, 1.2–1.6 mm wide; stem 150–200 µm wide, sparsely ventrally branched; rhizoids numerous, in short, divaricate, grayish, erect spreading fascicles; leaves obliquely inserted and oriented, slightly concave-canaliculate or slightly convex (then apex somewhat turned to ventral side), ventrally decurrent for 1/2 of stem width or farther, 750–900 × 750–800 µm, with rounded or truncate apex; underleaves obliquely spreading, 1.0–1.5 as wide as stem, decurrent for 1/3–1/2 of stem width, deeply divided by U-shaped sinus into two lobes, entire at margin or with blunt tooth on one or each lateral side or very shortly bisbifid cells in the midleaf thin-walled, trigones vestigial to nearly absent, 35–52 × 25–40 µm, cuticle finely but distinctly papillose.
Japan, Toyama Prefecture, Tateyama Mt., between Shishindake and Ryodake, 2600–2800 m a.s.l., on humus beneath
Unlike
The description based on the holotype is as follows: plants prostrate to loosely ascending, pale brownish in herbarium, forming loose mats, 1.1–2.0 mm wide and 5–10 mm long; rhizoids sparse to virtually absent, in several bundles obliquely to erect spreading or spreading up by the underleaf surface, from each underleaf base (if rhizoids developed), brownish to nearly colorless; stem brownish, 140–200 µm in diameter; leaves obliquely to subhorizontally inserted, dorsally insertion line transverse to arcuate, ventrally shortly decurrent, contiguous to subimbricate, ovate to obliquely ovate, 925–1075 × 625–875 µm; underleaves appressed to the stem, hyaline, 450–550 × 550–650 µm, nearly orbicular; midleaf cells subisodiametric to slightly oblong, ~25–40 µm in diameter, cells 5–6-gonal, thin-walled, with small and concave but distinct trigones, cuticle loosely verruculose; along margin 37–75 µm, with walls slightly thickened, trigones moderate in size, sometimes confluent on tangential side, concave; cells in underleaf middle mostly thin-walled with small to vestigial, concave trigones, along margin thin-walled, with small concave trigones.
India. Eastern Himalaya: Arunachal Pradesh, Lower Dibang Valley district, Mehao Wildlife Sanctuary, Mayodia top, ~2850 m, 18 Nov 2000, D.K. Singh 98225 (holotype: BSD); not seen.
The species is known only from the type locality cited by
Japan. Mt. Yokogura, May 1901, T. Yoshinaga no. 38 (LECTOTYPE (designated here): G [G00067733!], another, poor specimen [G00282608!] is lectotype duplicate. This species was founded on the gatherings by T. Yoshinaga and U. Faurie, however only Yoshinaga’s collections are now present in G. Both reviewed specimens contain plants fully corresponding to the original description).
The description based on the lectotype is as follows: plants greenish brownish to dirty greenish, 1.8–2.2 mm wide, 2–3 cm long, relatively rigid; stem 200–250 µm wide, branching not seen; rhizoids sparse to common, in brownish, erect spreading fascicles; leaves obliquely inserted and oriented, slightly concave-canaliculate, leaves not decurrent ventrally, triangular-ovate, with acute or rarely obtuse or bidentate apices, 900–1100 × 1000–1200 µm; underleaves 1.5–2.5 as wide as stem, decurrent for 1/3–2/3 of stem width, clearly bisbifid, undivided area 2(–3) cells high; midleaf cells subisodiametric 30–50 × 27–40 µm, thin-walled, trigones small to very small, cuticle smooth.
1 | Leaf apex mostly rounded |
|
– | Leaf apex acute to incised or distinctly bilobed |
|
2 | Underleaves shortly bilobed, emarginate or rounded at the apex |
|
– | Underleaves distinctly bilobed, at least for 2/5 of the length |
|
3 | Underleaves as large as leaves or slightly smaller, leaves distinctly curved to ventral side, plants distinctly bluish when fresh due to blue (grading to purple!) oil bodies |
|
– | Underleaves much smaller than leaves, leaves not curved to ventral side, plants greenish to bluish greenish, oil bodies colorless to grayish |
|
4 | Cells along leaf margin elongate and distinctly wider than cells of intramarginal row |
|
– | Cells along leaf margin nearly isodiametric, smaller than in intramarginal row |
|
5 | Cells along leaf margin distinctly swollen |
|
– | Cells along leaf margin not different from intramarginal cell row |
|
6 | Underleaves bifid |
|
– | Underleaves bisbifid or bifid with blunt tooth on each lateral side |
|
7 | Oil bodies biconcentric, with large central eye |
|
– | Oil bodies never biconcentric |
|
8 | Leaves with mostly acute apex, only some leaves on some shoots narrowly rounded |
|
– | Leaves with uniformly rounded apex |
|
9 | Underleaves decurrent for 1/3–1/2 of stem width, blunt teeth on lateral sides commonly present or underleaves bisbifid |
|
– | Underleaves not or shortly decurrent (to 1/3 of stem width), without or rarely with additional blunt tooth on one side |
|
10 | Stem ~1/5–1/6 of shoot width, underleaf lobes 8–10 cells in the base, underleaves 1.5–2.0 of stem width |
|
– | Stem ~1/7–1/8 of shoot width, underleaf lobes 3–5 cells width in the base, underleaves 1.1–1.4 of stem width |
|
11 | Stem relatively narrow, ~1/8 of plant width, leaves nearly planar |
|
– | Stem relatively wide, ~1/4 of plant with, leaves distinctly turned to dorsal side |
|
12 | Leaf apex acute |
|
– | Leaf apex incised (sometimes shortly so) to distinctly bilobed |
|
13 | Leaf cuticle smooth, underleaves mostly distinctly wider than stem |
|
– | Leaf cuticle very finely verruculose, underleaves as wide as stem or slightly wider |
|
14 | Underleaves 1.1–1.5 as wide as the stem |
|
– | Underleaves 1.5–3.5 as wide as the stem |
|
15 | Underleaf lobes 3–5 cells wide in the base, no additional lateral tooth on each side, leaves uniformly acute |
|
– | Underleaf lobes more than 6–8 cells wide in the base, additional lateral teeth commonly present on one or both sides, leaves commonly shortly incised, rarely acute (at least some admixture of incised leaves present) |
|
16 | Underleaves commonly bisbifid, rarely with obtuse lateral teeth on both sides, oil bodies brownish blue to brownish, finely granulate |
|
– | Underleaves commonly bifid with blunt (sometimes very smoothed) teeth on one or both sides, oil bodies not known |
|
17 | Underleaves decurrent for 1/3–1/2 of stem width, Hawaii | |
– | Underleaves decurrent for 1/4–1/3 of stem width, Sri Lanka |
|
18 | Undivided portion of underleaf 2–3 cells high |
|
– | Undivided portion of underleaf more than 4 cells high |
|
19 | Oil bodies colorless to grayish |
|
– | Oil bodies deep blue to blue brown, coarsely granulate |
|
20 | Underleaves commonly bisbifid |
|
– | Underleaves without lateral teeth |
|
21 | Underleaves bisbifid |
|
– | Underleaves bifid |
|
22 | Underleaves decurrent for 0.5–1.0 of stem width, oil bodies blue, leaves sometimes shortly incised |
|
– | Underleaves not or barely decurrent, oil bodies not known, leaves only acute |
|
23 | Underleaf lobes obtuse, underleaves 1.8–2.5 times wider than stem |
|
– | Underleaf lobes prominently acute, with 2–3 celled uniseriate ends, underleaves 3–4 times wider than stem |
|
24 | Underleaves divided by semicrescentic to U-shaped sinus, descending less than 1/7 of underleaf length |
|
– | Underleaves divided by V- to U-shaped sinus descending for 1/3–2/5 of underleaf length (undivided portion of underleaves 3–5 cells high) |
|
25 | Midleaf cell surface finely verruculose |
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– | Midleaf cell surface smooth |
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26 | Leaves constantly incised, cells in the leaf middle 40–80 × 30–60 µm, underleaves bisbifid |
|
– | Leaves rarely incised, commonly apiculate, cells in the leaf middle 37–58 ×25–35 µm, underleaves bifid |
|
27 | Underleaves bifid or with obscure additional teeth on one or both sides |
|
– | Underleaves constantly bisbifid or with distinct and prominent additional lateral teeth on one or both sides |
|
28 | Underleaves 0.8–1.3 as wide as stem, its undivided portion 1–3 cells high |
|
– | Underleaves 2.8–3.5 times as wide as stem, its undivided portion more than 5 cells high |
|
29 | Leaves apiculate to shortly incised into two strongly unequal or rarely nearly equal (sinus depth 2–3 cells) lobes |
|
– | Leaves shortly bilobed, for two subequal lobes, leaf sinus depth 4–6 cells |
|
30 | Leaf apex commonly obliquely truncate, unequally and very shortly bilobed, underleaves decurrent for 1/3 of stem width or less |
|
– | Leaf apex mostly acute to obtuse, underleaves decurrent for 1/3–1/2 of stem width | |
31 | Undivided portion of underleaves 2–3 cells high, oil bodies blue to grayish, brown and colorless |
|
– | Undivided portion of underleaves 4–5 and more cells high, oil bodies blue |
|
32 | Underleaves 1.1–1.3 as wide as stem, commonly bifid, with obscure additional lateral teeth on each side | |
– | Underleaves commonly more than 1.5 as wide as stem, almost constantly bisbifid |
|
33 | Leaves commonly acute, rarely incised (predominantly acute!) |
|
– | Leaves commonly with incised apex, rarely acute |
|
34 | Oil bodies coarsely granulate, deep blue to blue-brown, plants merely soft, somewhat glistening, commonly wider 2.2 mm wide, leaves somewhat undulate at margins, commonly turned to ventral side |
|
– | Oil bodies not known, plants more or less rigid, not glistening, commonly less than 2.2 mm wide, leaves planar at margins, not turned to ventral side |
|
35 | Oil bodies brownish to brown, blue and blue brown, botryoidal to granulate (in |
|
– | Oil bodies colorless to grayish, botryoidal, leaves with almost constantly shortly divided apex by U-shaped sinus |
|
36 | Oil bodies blue to deep blue botryoidal or not known |
|
– | Oil bodies brownish to brownish blue, finely granulate |
|
37 | Underleaves decurrent for 1/3–1/2 of stem width, oil bodies not known, leaves distinctly bilobed at apex (sinus depth 2–3 cells), leaves subimbricate |
|
– | Underleaves not or barely decurrent, oil bodies presumably deep blue, leaves distinctly bilobed at apex (sinus depth 3–5 or more cells), leaves contiguous to distant |
|
The vertical movements of the Himalaya, Tibetan Plateau and Hengduan Mts. have influenced the speciation of various groups of biota, not only that of liverworts (
The eastern part of the Sino-Himalaya and the eastward adjacent Meta-Himalaya, as identified in this work, are valuable biodiversity hotspots on Earth (
Although 11
In a broader context, taking into account the distribution of
The line art illustrations were kindly provided by Mr. Matvei Bakalin and geographic basis of the map was kindly prepared by Mr. Andreas K. Donadel to whom the authors are sincerely grateful. We are also deeply indebted to the curators of G, JE, NICH, STR, and TNS for the permission to study
– About 80% of type specimens in liverwort herbarium in G have two codes: the first one was provided in pre-computer time, whereas the second one was provided in the digitalization process. The papers before 2000-th cited the only old number, whereas now it is better to cite the new one (linked with the picture in the database). However, we guess, the best choice is to cite both, because only in this case we may identify the numbers widely used before with the recent ones. Therefore, in our citation we use the new number first (sometimes only that, if no old number is available) and then, after slash, the old one.
There are three specimens in G. The largest one is G00113555/5289, and another – G00113557/5291 – is part of the same specimen; the third specimen, G00113556/5290, is the possible syntype of the taxon, but the collection number is absent on the label.