Research Article |
Grzegorz J. Wolski‡, Aamir Shehzad Khan§, Beata Paszko|
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Corresponding author: Grzegorz J. Wolski ( grzegorz.wolski@biol.uni.lodz.pl ) Academic editor: Matt von Konrat
© 2023 Grzegorz J. Wolski, Aamir Shehzad Khan, Beata Paszko.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Wolski GJ, Khan AS, Paszko B (2023) New taxa of Plagiothecium (Plagiotheciaceae) from Pakistan. PhytoKeys 236: 1-16. https://doi.org/10.3897/phytokeys.236.109519
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Abstract
A revision of specimens of Plagiothecium deposited in the herbarium of Pakistan Museum of Natural History (
Key words
Bryophyta, new taxa, Pakistan, Plagiotheciaceae, South Asia, taxonomy
Introduction
The northern part of Pakistan has a suitable climate and ambience for bryophytes, while the southern part of this country is nearly unexplored because it is far drier, having hot weather and some parts with arid and desert ecosystems. Thereby, due to difficult fieldwork, geographic and climatic conditions, this country is generally still poorly studied bryologically but mainly due to the lack of a resident bryologist. The scattered literature principally covers northern Pakistan and Western Himalaya (
The first checklist of mosses of Pakistan based on previous literature, and herbarium specimens, from Punjab and Khyber Pakhtunkhwa provinces was provided by
Currently, the moss flora of Pakistan is represented by 319 species, with the largest family (Pottiaceae) consisting of 79 species (
The earliest mentions of Plagiothecium Schimp. in Pakistan go back to the 19th century, when
The current initiative is to update the moss flora of Pakistan, by taxonomically revising the dominant families and the addition of new records to the moss flora of Pakistan. This self-funded project was started by Mr. Aamir Shehzad Khan, after his master’s work on the Bryopsida in Pakistan (
However, comparing these data with data from other countries in this region (e.g., Iran and India [nine species], or China [20 taxa]) we can see that the Plagiothecium flora of this country is extremely poor (
Materials and methods
The current study is based exclusively on the Pakistani material, on the specimens deposited at the herbarium of Botanical Sciences Division, Pakistan Museum of Natural History (herbarium
In addition, the investigated herbarium collection was supplemented by field research carried out by the second author from August 2018 to July 2020 in the northern areas of Pakistan, including Murree (Punjab Province), Galiyat-region, Swat Valley, Lower Dir (Khyber Pakhtunkhwa Province), and some selected areas of Azad Jammu and Kashmir (Pakistan).
Measurements were made in accordance with the methodology proposed by
The features of two similar specimens – later named Plagiothecium higuchii var. higuchii and Plagiothecium higuchii var. brevicellum – were statistically compared to see whether the differences are significant. However, due to the lack of normality in the distributions of individual variables, a non-parametric U-Mann-Whitney test was performed.
Results and discussion
The conducted revision shows that the entire analyzed material is not very diverse. Most of the tested specimens belong to the Plagiothecium denticulatum complex. Thus, P. cavifolium, P. latebricola and P. nemorale, previously reported from this area, were not identified in the samples surveyed. The P. denticulatum complex dominated the examined material, within which the largest number of samples was represented by P. denticulatum var. denticulatum, the remaining specimens belonged to a new taxon, not previously recorded from Pakistan, P. denticulatum var. obtusifolium (Turner) Moore (M. Higuchi 20499).
Taxonomic treatment
Plagiothecium denticulatum
Plagiothecium denticulatum (Hedw.) Schimp., Bryologia Europaea 5: 190, 501, Tab. VIII (1851).
≡Hypnum denticulatum Hedw., Species Muscorum Frondosorum 237 (1801). Lectotype (fide
Description
Plants green to yellowish green, with metallic luster; stem in cross section rounded, 200–250 μm in diameter; leaves not folded, julaceous, elliptical-ovate, asymmetrical; leaves 2.2–3.0 × 1.0–1.3 (M 2.6 × 1.1) mm, shortly tapering to acute, denticulate apex; costae two, extending usually to ½ of the leaf length, ranging 440–810 (M 625) μm; cell areolation loose; length and width of cells very variable, but dependent on location: 85–140 × 15–28 (M 113 × 22) μm at the apex; 120–210 × 15–30 (M 165 × 23) μm at midleaf; 116–205 × 18–37 (M 160 × 28) μm toward insertion; broad decurrency 400–700 (M 555) μm; alar cells rounded, clearly inflated, 45–90 × 20–50 (M 68 × 35) μm; sporophytes not found in this material (Fig.
Plagiothecium denticulatum var. obtusifolium
Plagiothecium denticulatum var. obtusifolium (Turner) Moore, Proceedings of the Royal Irish Academy 1: 424 (1873).
≡Hypnum denticulatum var. obtusifolium Turner, Muscologiae Hibernicae Spicilegium 146, T. 12, f. 2 (1804).
≡Plagiothecium obtusifolium (Turner) J.J.Amann, Mémoire de la Société Vaudoise des Sciences Naturelles 3: 61 (1928). Holotype: fig. 2, tabela 12 “T. 12, f. 2” (Turner 1804: 237) (fig. 3). Epitype (selected by
=Plagiothecium sandbergii Renauld & Cardot, Contributions from the United States National Herbarium, 3: 274 (1895), Lectotype (selected by
=Plagiothecium denticulatum var. auritum Kern, Jahresbericht der Schlesischen Gesellschaft für Vaterländische Cultur 91(Abt. 2b): 97 (1914). Lectotype (selected by
Description
Plants light green, with metallic luster; stem in cross section rounded, 150–200 μm in diameter; the central strand well developed, epidermal cells 12–20 × 15–31 (M 16 × 23) μm, the parenchyma thin-walled, 25–43 × 21–39 (M 34 × 30) μm; leaves very concave, not folded, julaceous, elliptical-ovate, very slightly asymmetrical, 1.6–1.7 × 0.8–0.95 (M 1.65 × 0.87) mm; the apex obtusely-apiculate; costae two, extending usually up to 1/3 or ½ of the leaf length, ranging 250–530 (M 390) μm; cell areolation loose; length and width of cells very variable, but dependent on location: 85–140 × 15–18 (M 113 × 17) μm at the apex; 95–170 × 12–18 (M 133 × 15) μm at midleaf; 98–200 × 15–25 (M 150 × 20) μm toward insertion; broad decurrency 260–400 (M 330) μm; alar cells rounded-rectangular, 51–98 × 21–31 (M 75 × 26) μm; sporophytes not found in this material (Fig.
The next two samples (M. Higuchi 20460 and M. Higuchi 20479) were characterized by a unique combination of qualitative and quantitative features of their gametophytes. They did not match any of the taxa of the genus currently known in the Northern and Southern Hemispheres.
The specimen collected by M. Higuchi (M. Higuchi 20460) represents two morphotypes that are microscopically very different from each other. This material differs in the length and width of the leaf, the shape of the apex, the length and width of the cells of the top, middle and basal part of the leaf – which are one of the most important taxonomic features for the whole genus. In addition, the result of the U Mann-Whitney test showed that the length and width of the cells of individual parts of the leaf of these morphotypes differ statistically significantly (p<0.001) from each other. Thus, it was proposed to recognise two varieties – Plagiothecium higuchii var. higuchii and P. higuchii var. brevicellum.
Plagiothecium higuchii , sp. nov.
Type
Pakistan, Mt. Nanga Parbat, Mazeno Base Camp, 4000 m alt, on soil, 15 September 1990, M. Higuchi 20460, holotype LOD15016, isotype
Description
Plants green-yellow to golden-gold, without metallic luster; stems complanate-foliate, 3.0–4.0 cm long, in cross-section rounded, with a diameter of 290–360 (M 325) μm, the central strand well developed, epidermal cells 11–22 × 13–32 (M 16 × 22) μm, the parenchyma thin-walled, 26–50 × 23–45 (M 38 × 34) μm; leaves quite loosely arranged on the stem, concave, symmetrical to asymmetrical, ovate, those leaves from the middle of the stem 3.4–4.3 × 1.2–1.9 (M 3.8 × 1.5) mm; the apex acuminate and denticulate; costae two, thick and strong, usually to 1/2 of the leaf length, reaching 0.5–1.2 mm; laminal cells symmetrical, in unregulated transverse rows, the length and width very variable, but dependent on location: 107–250 × 16–24 (M 178 × 20) μm at apex, 139–266 × 21–33 (M 203 × 27) μm at mid-leaf and 139–266 × 21–33 (M 203 × 27) μm towards insertion, cell areolation very loose; decurrency long, 300–500 (M 400) μm, composed of 4–6 rows of rectangular and spherically inflated cells, 48–144 × 26–70 (M 96 × 48) μm. Sporophytes so far unknown (Fig.
Etymology
The present species is named in honor of Professor Masanobu Higuchi, who participated in the Studies on Cryptogams in the Western Himalayas in Pakistan project, and who collected the specimen (Higuchi 20460) chosen here as the holotype of Plagiothecium higuchii.
Plagiothecium higuchii var. brevicellum , var. nov.
Type
Pakistan, Mt. Nanga Parbat, Mazeno Base Camp, 4000 m alt, on soil, 15 September 1990, M. Higuchi 20460pp, holotype LOD 15017, isotype
Description
Plants green-yellow to golden-gold, without metallic luster. Stems complanate-foliate, 3.0–4.0 cm long, in cross-section rounded, with a diameter of 310–388 μm, epidermal cells 10–16 × 21–33 (M 13 × 27) μm, the parenchyma thin-walled, 14–58 × 18–51 (M 34 × 35) μm; leaves quite loosely arranged on the stem, concave, symmetrical to gently asymmetrical, ovate, those leaves from the middle of the stem 2.9–3.2 × 1.0–1.2 (M 3.1 × 1.1) mm; the apex acute and apiculate, denticulate; costae two, thick and strong, usually to 1/2 of the leaf length, reaching 0.5–1.5 (M 1.0) mm; laminal cells symmetrical to slightly asymmetrical, the length and width very variable, but dependent on location: 70–170 × 10–19 (M 120 × 15) μm at apex, 100–200 × 14–26 (M 150 × 20) μm at mid-leaf and 100–200 × 19–36 (M 150 × 28) μm towards insertion, cell areolation loose; decurrency long, 390–650 (M 520) μm, composed of 4–6 rows of rectangular and spherically inflated cells, 36–116 × 18–67 (M 76 × 43) μm. Sporophytes unknown so far (Fig.
Etymology
The variety name brevicellum refers to breve- [Lat.] short; -cellus [Lat.] cells and indicates the most distinctive features between varieties – leaves with short cells.
Plagiothecium filifolium , sp. nov.
Type
Pakistan, Mt. Nanga Parbat, Mazeno Base Camp, 4000 m alt, on boulder, 15 September 1990, M. Higuchi 20479, holotype LOD 15018, isotype
Description
Plants green-yellow, without metallic luster; stems complanate-foliate, 1.0–1.5 cm long; cross-section rounded, with a diameter of 200–300 μm, the central strand well developed, epidermal cells 9–13 × 11–15 (M 11 × 13) μm, the parenchyma thin-walled, 15–21 × 16–28 (M 18 × 22) μm; leaves folded, most strongly in the upper part of leaves, concave, asymmetrical, those leaves from the middle of the stem 1.6–2.2 × 0.5–0.8 (M 1.9 × 0.66) mm; leaves gradually tapering to long, acuminate, not denticulate apex; costae two, very thin and delicate, extending up to 1/2 of the leaf length, reaching 140–423 (M 282) μm; laminal cells rather symmetrical, the length and width very variable, but dependent on location: 97–160 × 7–10 (M 127 × 8) μm at apex, 105–180 × 7–9 (M 150 × 8) μm at mid-leaf and 70–165 × 8–11 (M 118 × 9) μm towards insertion, cell areolation tight; decurrency short, 220–530 μm, composed of 3 rows of gently inflated, rectangular cells, 47–147 × 12–21 (M 97 × 16) μm. Sporophytes unknown so far (Fig.
Etymology
Filum- [Lat.] thread; -folium [Lat.] leaf. The present species is named in reference to the most distinctive feature – its threadlike leaf apex.
Discussion
Previously, only four species had been reported from Pakistan: Plagiothecium cavifolium, P. denticulatum, P. latebricola and P. nemorale (
Compared to neighboring countries, the four species previously reported from Pakistan (Plagiothecium cavifolium, P. denticulatum, P. latebricola and P. nemorale) were considered a very low number (
As indicated by
This research showed that some of the specimens have unique combinations of qualitative and quantitative gametophytic features. A specimen with complanate-foliate stems; folded, concave, asymmetrical leaves gradually tapering to long, acuminate, not denticulate apex, with a decurrency composed of gently inflated, rectangular cells has been named Plagiothecium filifolium. There is no species in the Northern Hemisphere with this set of features. The most similar to this taxon is P. latebricola, which, however, is characterized by a symmetrical leaf, narrow decurrency with non-inflated cells (e.g.
Specimens with leaves quite loosely arranged on the stem, concave, symmetrical to asymmetrical, ovate, with a denticulate apex, loose cell areolation and wide decurrency, composed of rectangular and spherically inflated cells have been named Plagiothecium higuchii. These specimens also represent a unique set of features hitherto unknown in the Northern or Southern Hemispheres (
The taxa from the Northern Hemisphere most similar to P. higuchii would be: P. denticulatum, P. platyphyllum Mönk. or P. ruthei Limpr., however, they differ in the arrangement of the leaves on the stem, in the shape and dimensions of the leaf, and the dimensions of the leaf cells, which shows the striking distinctiveness of the examined material against the previously described taxa (
As studies of Plagiothecium indicate (e.g.
The present study describes three taxa, Plagiothecium higuchii var. higuchii, P. higuchii var. brevicellum, and P. filifolium, as new to science and reports another taxon, P. denticulatum var. obtusifolium, as new to Pakistan. With the four species reported earlier this brings the number of recognized species and infraspecific taxa of Plagiothecium in Pakistan to eight.
On the basis of herbarium specimens and literature analysis, it was possible to determine the range of the described species of the genus in Pakistan:
-
Plagiothecium cavifolium
has been recorded so far from: Khyber Pakhtunkhwa, Kaghan Valley, Lake Saif Ul Maluk, 3150 m (Higuchi 19888,
Higuchi 19891,
Higuchi 1992 ) (Fig.6 ). -
Plagiothecium denticulatum
has been recorded so far from: Khyber Pakhtunkhwa, Chitral, Ziarat, 7400 m and Gilgit-Baltistan, Minimarg, Burzil Pass, 10 000 to 11 000 m (
Brotherus 1898a ,b ); Gilgit-Baltistan, Nanga Parbat Base camp 3430 m, and Mazeno Base camp, 4000 m (Higuchi 20462, 20475,Higuchi 1992 ); Khyber Pakhtunkhwa, Mansehra, Sikyan near Nadi, 2020 m (Nishimura et al. 1993b ) and Nanga Parbat Base Camp and Nanga Parbat Mazeno Base Camp, at 3430–4000 m (M. Higuchi 20462; M. Higuchi 20464; M. Higuchi 20479; M. Higuchi 20510) (Fig.6 ). -
Plagiothecium denticulatum
var.
obtusifolium has been recorded so far from: Gilgit-Baltistan, Nanga Parbat, Base Camp, at 3430 m (M. Higuchi 20499) (Fig.
6 ). -
Plagiothecium filifolium
has been recorded so far from: Gilgit-Baltistan, Nanga Parbat, Mazeno Base Camp, at 4000 m (M. Higuchi 20479) (Fig.
6 ). -
Plagiothecium higuchii
var.
higuchii has been recorded so far from: Gilgit-Baltistan, Nanga Parbat, Mazeno Base Camp, at 4000 m (M. Higuchi 20460) (Fig.
6 ). -
Plagiothecium higuchii
var.
brevicellum has been recorded so far from: Gilgit-Baltistan, Nanga Parbat, Mazeno Base Camp, at 4000 m (M. Higuchi 20460) (Fig.
6 ). -
Plagiothecium latebricola
has been recorded so far from: Khyber Pakhtunkhwa, Mansehra, Shogran – Sali Hut, 2710 m (
Nishimura et al. 1993b ) (Fig.6 ). -
Plagiothecium nemorale
has been recorded so far from: Gilgit-Baltistan, Minimarg, Burzil Pass, 9000 to 10 000 m and 10 000 to 11 000 m (
Brotherus 1898a ); Azad Jammu and Kashmir, Shekh Bela, between Shardi and Kel, alt. 2000–2100 m (Noguchi 1954 ) (as P. sylvaticum); Khyber Pakhtunkhwa, Mansehra, Sharan Forest, 2400 m (Nishimura et al. 1993b ) (Fig.6 ).
A key to the species of the genus Plagiothecium from Pakistan
| 1 | Angular cells of decurrency rounded, inflated, forming distinct auricles | 2 |
| – | Angular cells of decurrency rectangular, not inflated, not forming distinct auricles | 5 |
| 2 | Leaves asymmetrical; apex acute and denticulate | P. denticulatum |
| – | Leaves symmetrical or slightly asymmetric; apex denticulate or not | 3 |
| 3 | Plants with acuminate, denticulate apex | 4 |
| – | Plants with obtuse, not denticulate apex | P. denticulatum var. obtusifolium |
| 4 | Cells from the middle part of the leaves long and wide (139–266 × 21–33) | P. higuchii var. higuchii |
| – | Cells from the middle part of the leaves shorter and narrower (100–200 × 14–26 μm) | P. higuchii var. brevicellum |
| 5 | Leaves asymmetric and folded | P. filifolium |
| – | Leaves symmetric and not folded | 6 |
| 6 | Cells from the middle part of the leaves < than 10 μm wide | P. latebricola |
| – | Cells from the middle part of the leaves > than 10 μm wide | 7 |
| 7 | Stem julaceus; leaves strongly concave; apex not denticulate | P. cavifolium |
| – | Stem not julaceus; leaves rather flat; apex denticulate | P. nemorale |
Acknowledgements
The authors thank Dr. Syed Aneel Ahmed Gilani of the Botanical Science Division, Pakistan Museum of Natural History, Islamabad, Pakistan for making the collection available to research and for Professor William “Bill” R. Buck for proofreading the manuscript and for valuable suggestions that contributed to improving the quality of this article.
Additional information
Conflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Funding
No funding was reported.
Author contributions
Data curation: GJW. Investigation: GJW, ASK. Project administration: GJW. Supervision: GJW. Visualization: ASK, GJW. Writing – original draft: GJW, BP, ASK. Writing – review and editing: ASK, BP, GJW.
Author ORCIDs
Grzegorz J. Wolski https://orcid.org/0000-0003-1480-8003
Aamir Shehzad Khan https://orcid.org/0000-0002-9039-2524
Beata Paszko https://orcid.org/0000-0003-3044-1766
Data availability
All of the data that support the findings of this study are available in the main text or Supplementary Information.
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Supplementary material
Raw data of new taxa from Pakistan
Data type: xlsx
Explanation note: The raw data contains two sheets, with morphological data and statistical test results.