Hattoria 9: 11–30. 2018

The () in Govind Wildlife Sanctuary, Uttarakhand, India

Dayanidhi GUPTA1, Vinay SAHU1, Krishna Kumar RAWAT1, Neerja PANDE2 & Ashish Kumar ASTHANA1

1 CSIR-National Botanical Research Institute, Rana Pratap Marg, Lucknow 226001, India 2 Department of Botany, Kumaun University, Nainital 263001, India Author for correspondence: Ashish Kumar ASTHANA, [email protected]

Abstract The diversity of the liverwort genus Porella is studied in the Govind Wildlife Sanctuary (GWLS), Uttarakhand. A total of 12 species and infraspecific taxa are described, all being reported for the first time from the area. A working key is provided for identification along with the taxonomic description of each of them. As Porella is represented by 23 species and infraspecific taxa in the Western Himalaya, the number of species and infraspecific taxa found in Govind Wildlife Sanctuary is relatively high.

Introduction Govind Wildlife Sanctuary is located in Purola Tahsil, District Uttarkashi of Uttarakhand, India and lies between 77°47′–78°37′E longitude and 31°17′–35°35′ N latitude. The sanctuary was established in 1955, covering 485.39 km2 area. In year 1990, it was selected for conservation of Snow leopard (Uncia uncia) and 472 km2 more area of upper reaches of the sanctuary was added and notified as Govind Wildlife Sanctuary and National Park. River Tons is the main stream of this area which forms upper catchment of river Yamuna. The sanctuary has different ecological zones ranging from subtemperate to temperate and subalpine to alpine, providing different habitats and niches for the different groups of as well as animals. The bryophyte flora of Govind Wildlife Sanctuary is very little known. Singh & Singh (2002) reported a total of nine species of liverworts from that area. Recently, Asthana & Gupta (2014), Asthana & Sahu (2013, 2014a, 2014b, 2014c, 2015a, 2015b, 2016), Sahu & Asthana (2014a, 2014b, 2016) and Rawat et al. (2016) contributed towards the knowledge of bryophyte flora of the area. Despite these efforts, the bryophyte flora of the area in general and liverworts in particular, are yet to be explored in detail. The present paper deals with the genus Porella in the Govind Wildlife Sanctuary. A total 32 species and infra specific taxa of Porella have been recognized in India, of which 23 are reported from Western Himalayas including 21 from Uttarakhand (Singh et al. 2016). In India, earlier studies on Porella were carried out by Udar & Shaheen (1983a, 1983b, 1983c),

11 Shaheen & Srivastava (1986a, 1986b, 1989), Singh & Singh (2006), and Kumar & Singh (2002). Porella campylophylla var. ligulifera (Taylor) S.Hatt. and P. hattori Udar & Shaheen have been reported by Singh & Singh (2002) from the study area. Both are treated here as doubtful records (see below). In the present paper, an account of 12 species and infraspecific taxa of the genus Porella from the Govind Wildlife Sanctuary is provided, together with brief morphological descriptions, illustrations and an identification key.

Material and Methods Fresh samples collected from different localities during various field trips in the study area (Fig. 1) as well as herbarium samples have been studied and deposited in the Bryophyte Herbarium of CSIR–NBRI Lucknow (LWG).

Key to the species of genus Porella in Govind Wildlife Sanctuary 1. Leaves entire ···························································································· 2 1. Leaves toothed ·························································································· 6 2. Leaf lobe crispate at margin ············································ P. chinensis var. chinensis 2. Leaf lobes flat or recurved at margin ································································ 3 3. Leaf apex usually flat ··························································· P. madagascariensis 3. Leaf apex strongly recurved ·········································································· 4 4. Leaf lobules large, almost equal to leaf width ·······················P. obtusata var. macroloba 4. Leaf lobules small than above ········································································ 5 5. Underleaves with basal appendages ·································P. gracillima var. gracillima 5. Underleaves without basal appendages ·······························P. chinensis var. decurrens 6. Leaf apex acute to acuminate ········································································· 7 6. Leaf apex broad, obtuse to multidentate ···························································10 7. Leaf apex subacute to acute, irregularly 2–5 toothed ···············P. acutifolia var. acutifolia 7. Leaf apex acuminate to apiculate or setose ························································· 8 8. Leaf apex acute to apiculate, with few additional teeth, margin weakly arched ················· ············································································ P. caespitans var. caespitans 8. Leaf apex sharply setose, margin prominently arched ············································ 9 9. Leaf apex without additional teeth ····································· P. caespitans var. setigera 9. Leaf apex with 1–3 additional teeth near apex ····················· P. caespitans var. nipponica 10. Leaf apex with more than 3 teeth ························································P. plumosa 10. Leaf apex obtuse, truncate or acuminate with 2–3 apical teeth ·······························11 11. Underleaves and lobules with dentate apex ··········································· P. japonica 11. Underleaves and lobules with entire apex ·······················P. densifolia subsp. densifolia

Taxonomic treatment Porella acutifolia (Lehm. & Lindenb.) Trevis. var. acutifolia (Fig. 2) Plants yellow to brown in herbarium, up to 60 mm long and 5 mm wide, irregularly bi to tripinnately branched, branches obliquely spreading. Leaves imbricate, oblong-ovate,

12 Figure 1. Map of the study area showing various localities.

1.47–2.23×1.05–1.80 mm, margin entire, apex rarely subacute to acuminate, sometimes 2–3 toothed, rarely obtuse; marginal cells of leaf subquadrate, 12–20×12–16 µm; median cells polygonal, 20–32×16–24 µm; basal cells elongated polygonal, 28–52×16–28 µm; trigones triangular; lobules lanceolate-ovate, 0.40–1.05×0.42–0.60 mm, margin entire, slightly undulate, apex obtuse; underleaves orbicular, triangular or oblong, ca. 0.84–1.18×0.76– 1.13 mm, margin entire or with 2–3 teeth near base, apex rounded to obtuse. Fertile not found. Specimen examined: INDIA, Uttarakhand, Uttarkashi, Govind Wildlife Sanctuary, on way to Kedarkantha, elev. ca. 2240 m, Sahu (LWG 264894B). This variety is new to Western Himalayan region in India.

Porella caespitans (Steph.) S.Hatt. var. caespitans (Fig. 3) Plants yellow to brown in herbarium, 50–60 mm long, up to 4 mm wide, pinnately or bipinnately branched. Leaves imbricate, ovate, 1.5–2×1–1.3 mm, margin entire, dorsally arched, more or less undulate, incurved, apex acute or apiculate or 2–3 toothed, teeth spinose, 3–4 celled long and 2–3 celled wide at base; marginal leaf cells quadrate to subquadrate, 15.2–19×7.6–11.4 µm; median leaf cells polygonal, 22.8–30.4×15.2–19 µm; basal cells slightly elongated polygonal, 26.4–38×19–30 µm; trigones triangulate; lobules ligulate, 0.7–1×0.3–0.5 mm, apex obtuse, margins recurved; underleaves ligulate, triangulate or auriculate, margin entire, weakly recurved, apex obtuse or truncate. Fertile plant not found. Specimens examined: INDIA, Uttarakhand, Uttarkashi, Govind Wildlife Sanctuary,

13 Figure 2. Porella acutifolia (Lehm. & Lindnb.) Trevis. var. acutifolia. 1: Plant, dorsal view. 2: Plant, ventral view. 3–8: Leaves. 9–18: Leaf lobules. 19–25: Underleaves. 26: Apical cells of leaf. 27: Median cells of leaf. 28: Marginal cells of leaf. 29: Basal cells of leaf.

14 Figure 3. Porella ceaspitans (Steph.) S.Hatt. var. ceaspitans. 1: Plant, ventral view. 2: Plant, dorsal view. 3–10: Leaves. 11–17: Leaf lobules. 18–24: Underleaves. 25: Apical cells of leaf. 26: Median cells of leaves. 27: Marginal cells of leaf. 28: Basal cells of leaf.

15 Kedarkantha, elev. ca. 2658 m, Asthana (LWG 263038A); Sankri, elev. ca. 1920 m, Sahu (LWG 264880A); Jakhol village, elev. ca. 2220 m, Sahu (LWG 265065 E).

Porella caespitans (Steph.) S.Hatt. var. nipponica S.Hatt. (Fig. 4) Plants yellow to brown in herbarium, upto 80 mm long and 2–2.5 mm wide, bipinnately branched. Leaves imbricate, leaf lobe ovate, 1.60–2.45×0.84–1.70 mm, dorsal margin arched, entire, apex acute to apiculate with 1–3 teeth; marginal leaf cells quadrate to subquadrate, 16–25×16–25 µm; median cells polygonal, 16–28×16–25 µm; basal cells elongated polygonal, 28–44×16–24 µm; trigones nodulose; lobule ligulate-lanceolate, 0.67–1.10× 0.46–0.60 mm, obliquely arranged, sinuately inserted, margin entire, apex obtuse rarely 1–2 dentations, base laterally broad; underleaves auriculate, 0.51–20×0.51–0.93 mm, margin entire; apex recurved, truncate to bispinose, rarely unispinose, base long decurrent, rarely toothed. Fertile plant not found. Specimens examined: INDIA, Uttarakhand, Uttarkashi, Govind Wildlife Sanctuary, on the way from Taluka to Osla, elev. ca. 2097 m, Asthana (LWG 263001A).

Porella caespitans (Steph.) S.Hatt. var setigera (Steph.) S.Hatt. (Fig. 5) Plants yellow to brown in herbarium, up to 60 mm long and 1.5–2 mm wide, bipinnately branched. Leaves ovate, imbricate, 1.14–2.0×0.97–1.34 mm, margin arched, apex setiferous or apiculate; marginal cells of leaf quadrate to subquadrate, 8–16×8–16 µm, thick walled; median cells polygonal, 12–24×12–16 µm; basal cells elongated polygonal, 24–36× 16–20 µm; trigones nodulose; lobule obliquely spread, lanceolate, margin entire, apex obtuse to subacute, long decurrent base; underleaves ovate-triangular, 0.88–1.34×0.67–1.05 mm, margin entire, apex bispinose (teeth acute to setose) or obtuse or subtruncate. Fertile plants not found. Specimens examined: INDIA, Uttarakhand, Uttarkashi, Govind Wildlife Sanctuary, on way Taluka to Osla, elev. ca. 2720 m, Nath (LWG 204844A); Har Ki Doon, elev. ca. 3000 m, Nath (LWG 204915A); Har Ki Doon to Osla, elev. ca. 3000 m, Nath (LWG 204929A); Taluka to Osla, elev. ca. 2097 m, Asthana (LWG 252488A); Taluka to Osla, elev. ca. 2229 m, Rawat (LWG 264847A); Gangar village, elev. ca. 2483 m, Rawat (LWG 265084B). This variety is new to Western Himalayan region in India.

Porella chinensis (Steph.) S.Hatt. var. chinensis (Fig. 6) Plants yellow to brown in herbarium, up to 30 mm long and 1.5–2 mm wide. Leaves incubous, imbricate, oblong, 1.05–1.47×0.75–1.40 mm, margin entire, crispate, apex rounded; marginal cells thick walled, quadrate to subquadrate, 12–20×16–20 µm; median cells hexagonal to polygonal, 16–20×20–28 µm; basal cells elongated polygonal, 24–40× 20–28 µm; trigones nodulose; lobules lanceolate; 0.35–0.80×0.17–0.47 mm; margin entire, apex rounded; underleaves ovate or oblong, ca. 0.50–0.63×0.42–0.60 mm, margin entire, crispate, apex rounded, recurved. Fertile plants not found. Specimen examined: INDIA, Uttarakhand, Uttarkashi, Govind Wildlife Sanctuary, Sankri, elev. ca. 1984 m, Rawat (LWG 265087A).

16 Figure 4. Porella ceaspitans (Steph.) S.Hatt. var. nipponica S.Hatt. 1: Plant, dorsal view. 2: Plant, ventral view. 3–7: Leaves. 8–13: Leaf lobules. 14–18: Underleaves. 19: Apical cells of leaf. 20: Median cells of leaf. 21: Marginal cells of leaf. 22: Basal cells of leaf.

17 Figure 5. Porella ceaspitans (Steph.) S.Hatt. var. setigera (Steph.) S.Hatt. 1: Plant, dorsal view. 2: Plant, ventral view. 3–9: Leaves. 10–13: Leaf lobules. 14–16: Underleaves. 17: Apical cells of leaf. 18: Median cells of leaf. 19: Marginal cells of leaf. 20: Basal cells of leaf.

18 Figure 6. Porella chinensis (Steph.) S.Hatt. var. chinensis. 1: Plant, ventral view. 2–4: Leaves with lobules. 5–8: Underleaves. 9: Marginal cells of leaf. 10: Basal cells of leaf. 11: Median cells of leaf.

19 Figure 7. Porella chinensis (Steph.) S.Hatt. var. decurrens (Steph.) S.Hatt. 1: A portion of male plant. 2: Plant, ventral view. 3–6: Leaves with lobule. 7–10: Underleaves. 11: Marginal cells of leaf. 12: Basal cells of leaf. 13: Bracts. 14: Bracteole. 15: Antheridium.

20 Porella chinensis (Steph.) S.Hatt. var. decurrens (Steph.) S.Hatt. (Fig. 7) Plants light green, up to 39 mm long and 2.5–3.0 mm wide, bipinnately branched. Leaves imbricate, ovate to orbicular-ovate, 1.25–1.53×0.89–1.46 mm, margin entire, ventrally concave, apex rounded, incurved, base decurrent; marginal cells of leaf quadrate to subquadrate, 16–24×12–20 mm; basal cells slightly elongated, polygonal, 24–32× 24–32 mm; trigones nodulose; lobules ovate-triangular, 0.63–0.65×0.43–0.63 mm, margin recurved, apex obtuse or rounded, entire, base decurrent; underleaves suborbicular, 0.32–0.66 ×0.43–0.86 mm wide, margins entire, recurved at apex, base decurrent. Plants dioecious, androecia usually on primary branch, bracts in 3–5 pairs, ovate-triangulate, 0.61–0.95×0.35– 0.55 mm; bracteole ovate, 0.27×0.16 mm; antheridia 1 in each bract, globose, 0.56× 0.36 mm; Gynoecia not found. Specimen examined: INDIA, Uttarakhand, Uttarkashi, Govind Wildlife Sanctuary, Bithri, elev. ca. 2401 m, Rawat (LWG 300443A).

Porella densifolia (Steph.) S.Hatt. subsp. densifolia (Fig. 8) Plants yellow to green to brown in herbarium, up to 70 mm long and 2.5–3 mm wide, irregularly branched. Leaves densely imbricate, ovate to oblong, 1.40–2.10×1.38–1.60 mm, margin entire, apex almost round to spinose-toothed; marginal cells of leaf quadrate to subquadrate, 12–20×12–16 µm; median cells polygonal, 16–32×12–24 µm; basal cells elongated polygonal, 20–36×12–24 µm; trigones nodulose; lobules ovate, entire margin with rounded apex, 0.63–0.71×0.34–0.50 mm; underleaves ovate or oblong, 0.67–0.84×0.33– 0.46, margin entire, apex recurved, rounded, base appendiculate. Fertile plants not found. Specimen examined: INDIA, Uttarakhand, Uttarkashi, Govind Wildlife Sanctuary, Juda Tal, elev. ca. 2209 m, Sahu (LWG 254803B).

Porella gracillima Mitt. var. gracillima (Fig. 9) Plants yellow to green to brown in herbarium, up to 48 mm long, 1–1.5 mm wide, irregularly bipinnately branched. Leaves imbricate, oblong to ovate, 1.21–1.50×1–1.26 mm, margin entire, apex rounded, incurved, slightly decurrent; marginal cells 8–20×8–16 µm, quadrate to subquadrate; median cells polygonal, 12–24×12–20 µm; basal cells elongated polygonal, 20–36×16–24 µm; leaf lobules lanceolate to ligulate, 0.55–0.71×0.30–0.40 mm, margin entire, apex obtuse, recurved, outer base appendaged; underleaves oblong-quadrate, 0.50–0.7×0.42–0.60 mm, margin obtuse, entire or with few teeth on basal margin, apex entire, recurved, base decurrent at both side. Fertile plants not found. Specimens examined: INDIA, Uttarakhand, Uttarkashi, Govind Wildlife Sanctuary, between Taluka and Osla, elev. ca. 2720 m, Nath (LWG 204846A, 204847A).

Porella japonica (Sande Lac.) Mitt. (Fig. 10) Plants yellowish green to olive green, up to 60 mm long and 2 mm wide; pinnately branched. Leaves ovate, 1.00–1.47×0.84–1.22 mm in size, margins entire, crispate, apex with 2–4 teeth, marginal cells of leaves quadrate to subquadrate, 12–20×12–20 µm in size; median cells polygonal, 20–32×16–24 µm; basal cells elongated, polygonal, 28–52×20–28 µm; cells slightly thickened; trigones triangulate; lobules ovate to ligulate with broad base, 0.25–0.67×

21 Figure 8. Porella densifolia (Steph.) S.Hatt. subsp. densifolia. 1: Plant, ventral view. 2–4: Leaves. 5–7: Leaf lobules. 8–10: Underleaves. 11: Median cells of leaf. 12: Marginal cells of leaf. 13: Basal cells of leaf.

22 Figure 9. Porella gracillima Mitt. var. gracillima. 1: Plant, ventral view. 2–5: Leaves. 6–8: Leaf lobules. 9–12: Underleaves. 13: Marginal cells of leaf. 14: Basal cells of leaf. 15: Median cells of leaf.

23 Figure 10. Porella japonica (Sande Lac.) Mitt. 1: Plant, ventral view. 2: Plant, dorsal view. 3–7: Leaves. 8–12: Underleaves. 13–18: lobules. 19: Apical cells of leaf. 20: Marginal cells of leaf. 21: Median cells of leaf. 22: Basal cells of leaf.

24 0.17–0.34 mm, apex truncate to spinose, parallel to stem, basal appendages present; underleaves ovate or triangulate, 0.25–0.67×0.17–0.34 mm, margins entire with basal appendages, apex truncate with dentations. Fertile plants not found. Specimens examined: INDIA, Uttarakhand, Uttarkashi, Govind Wildlife Sanctuary, Osla to Taluka, elev. ca. 2097 m, Asthana (LWG 252460A).

Porella madagascariensis (Nees & Mont.) Trevis. (Fig. 11) Plants light green to olive green, old parts brown, up to 38 mm long and 2 mm wide; bipinnately branched. Leaves imbricate, ovate, 0.9–1.2×0.92–0.75 mm, margin entire, apex rounded, base slighty decurrent; marginal cells of leaf quadrate to subquadrate, 12–16× 12–16 µm; median cells hexagonal to polygonal, 16–24×12–20 µm; basal cells slightly elongated, polygonal, 16–32×16–24 µm; trigones nodulose; leaf lobules lanceolate, 0.2–0.32 ×0.08–0.13 mm, broad at base, margin entire or sometimes 1–2 tooth, apex obtuse or rounded; underleaves triangular to subrectangular, 0.38–0.55×0.25–0.38 mm wide, margin entire, acute or shallowly bilobed at apex . Fertile plants not found. Specimen examined: INDIA, Uttarakhand, Uttarkashi, Govind Wildlife Sanctuary, Osla to Taluka, elev. ca. 2712 m, Sahu (LWG 265029A).

Porella obtusata (Taylor) Trevis. var. macroloba (Steph.) S.Hatt. & Zhang (Fig. 12) Plants yellowish brown in herbarium, up to 48 mm long and 1.5–2 mm wide, irregularly pinnately branched. Leaves densely imbricate, oblong-ovate to elliptical, 1.26–1.42×1.38– 1.68 mm, concave, margin entire, incurved, apex rounded; marginal cells of leaf thin walled, quadrate to subquadrate, 16–28×12–20 µm; median cells polygonal, 16–32×16–20 µm; basal cells elongated, polygonal 16–40×16–28 µm; trigones nodulose; lobules large, ovoid-elliptical, 0.50–0.97×0.22–0.84 mm, margin entire, apex round-obtuse, base decurent, minutely appendiculate; underleaves ovate or triangulate, 0.59–0.84×0.76–0.88 mm, margin entire, crispate, apex recurved, base long and appendiculate. Fertile plants not found. Specimens examined: INDIA, Uttarakhand, Uttarkashi, Govind Wildlife Sanctuary, on way of Taluka to Osla, elev. ca. 2097 m, Asthana (LWG 252473B); on way of Sankri to Kedarkantha, elev. ca. 2658 m, Asthana (LWG 263039A); Sankri, elev. ca. 1920 m, Sahu (LWG 264879F).

Porella plumosa (Mitt.) Inoue (Fig. 13) Plants light green to yellowish green, up to 50 mm long and 2.5 mm wide, pinnately branched. Leaves imbricate, oblong-ovate, 1–2.5×1–1.8 mm, margin entire, rarely multi toothed apex, teeth 2–6 celled long and 2–3 celled wide at base, terminal cell obtuse or acute, 22.8–26.6×19–22.8 µm; marginal cells of leaf quadrate to subquadrate, 19–30.4×11.4– 22.8 µm; median cells polygonal, 34.2–38×22.8–30.4 µm; basal cells elongated, polygonal, 49.4–76×26.6–30.4; trigones triangulate; lobule lanceolate-ovate, parallel to stem, 0.35–0.40×0.20–0.30 mm, margin entire, apex obtuse to subacute; underleaves ovate, 0.80–1 ×0.40–0.50 mm, margin entire, apex rounded, 2–5 toothed. Fertile plants not found. Specimens examined: INDIA, Uttarakhand, Uttarkashi, Govind Wildlife Sanctuary, on way from Taluka to Osla, elev. ca. 2097 m, Asthana (LWG 252473C); on way from Taluka to

25 Figure 11. Porella madagascariensis (Nees & Mont.) Trevis. 1: Plant, dorsal view. 2: Plant, ventral view. 3–10: Leaves. 11–14: Leaf lobules. 15–18: Underleaves. 19: Apical cells of leaf. 20: Median cells of leaf. 21: Marginal cells of leaf. 22: Basal cells of leaf.

26 Figure 12. Porella obtusata (Taylor) Trevis. var. macroloba (Steph.) S.Hatt. & Zhang. 1: Plant, ventral view. 2–4: Leaves. 5–7: Underleaves. 8–10: Leaf lobules. 11: Marginal cells of leaf. 12: Median cells of leaf. 13: Basal cells of leaf.

27 Figure 13. Porella plumosa (Mitt.) Inoue. 1: Plant, ventral view. 2–5: Leaves. 6–9: Underleaves. 10–12: Lobules. 13: Marginal cells of leaf. 14: Median cells of leaf. 15: Basal cells of leaf. 16: Apical cells of leaf. 28 Osla, elev. ca. 2360 m, Rawat (LWG 264852A); Kedarkantha, elev. ca. 2871 m, Rawat (LWG 254848B); Near Osla bridge, elev. ca. 2650 m, Rawat (LWG 264998B).

Doubtful species in the area Singh & Singh (2002) have earlier reported Porella campylophylla var. ligulifera (Tayl.) S.Hatt. and P. hattorii Udar & Shaheen from Govind Wildlife Sanctuary. However, the illustrations of lobules and underleaves of P. campylophylla var. ligulifera (Singh & Singh 2002, Fig. II: 4, 11) show entire margins in both, indicating that the plants might belong to P. plumosa. Hence, the report of P. campylophylla var. ligulifera is doubtful. Furthermore, the illustrations of the underleaves of P. hattorii (Singh & Singh 2002, Fig. III: 8, 9) show conspicuously decurrent bases, and the lobule size falls within the size range of the underleaves (lobules 0.41–0.61 mm long, underleaves 0.44–0.89 mm long). However, Udar & Shaheen (1983c) while describing P. hattorii, clearly mentioned that “In P. hattorii the leaf lobules are considerably smaller in relation to the lobe and also smaller than the underleaves. In addition, lobule and underleaves are never decurrent on the stem”. Udar & Shaheen (1983c) while discussing the differences of P. hattorii with P. acutifolia and P. caespitans had pointed out that “they show the equal size of the lobules and underleaves as well as their conspicuous decurrens on the stem”. Hence, it seems that plants identified as P. hattorii closely approach P. acutifolia. The record of P. hattorii is therefore considered doubtful and needs reinvestigation.

Acknowledgements We are thankful to the Director, CSIR–NBRI for kindly providing the facilities. Financial support from CSIR under project BSC–0106 is also acknowledged.

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