Utricularia sect. Nigrescentes (Oliv.) Komiya, J. Jap. Bot.

Utricularia sect. Nigrescentes (Oliv.) Komiya, J. Jap. Bot. 48(5): 151. 1973; P.Taylor, Kew Bull. 41(1): 6. 1986, Gen. Utricularia 186. 1989. Type: Utricularia caerulea L.

Terrestrial, semi aquatic herbs. Rhizoids a few, simple or branched. Stolons many, branched. Leaves obovate to spathulate, 1-nerved. Traps numerous on stolons and leaf petiole, ovoid; mouth lateral, extends to an oblique funnel-shaped rim, glandular hairy towards margin; internal glands 2–4-armed. Scales and bracts basisolute; bracteoles basifixed or basisolute. Inflorescence simple or branched. Calyx lobes sub-equal, minutely papillate. Corolla blue– purple, white or yellow. Pollen 3 or 4-colporate, oblate, spherical or prolate. Capsule wall thickened, dehisce by a ventral slit. Seeds obovoid–oblongoid; testa cells slightly elongate, anticlinal wall concave, periclinal wall convex, surface verrucose or rugulose.

Distribution: Tropical Africa, Madagascar, Asia to

Japan and Australia.

Notes: The name ‘ Nigrescentes ’ was originally given by Oliver (1859) to a group of Utricularia species with shorter pedicels sub-equal to the length of bracts; medifixed scales and bracts with acute apices, minutely papillate calyx lobes and spur shorter or longer than the lower lip of the corolla. He segregated species of this group under two subsections: 1. Corollae calcar labio inferior non longius (short spurred: U. rosea Edgew. ) 2. Corollae calcar labium inferius excedens (long spurred: U. racemosa Wall. ex Walp. , U. nivea Vahl and U. filicaulis Wall. ). Komiya (1973) raised ‘Nigrescentes’ into a formal section and included the species treated by Oliver under it. The section currently includes four species: U. bracteata , restricted to South Tropical Africa, U. warburgii , distributed in Southeast China, U. caerulea and U. roseopurpurea ( U. rosea , U. racemosa , U. nivea , and U. filicaulis are presently considered as synonyms of U. caerulea ).

Utricularia roseopurpurea Stapf ex Gamble, Fl. Madras View in CoL 2: 983. 1924; Fyson, Fl. S. Ind. Hill Sta. 1: 438. 1932, as “ Utricularea rosa-purpurea Staff ”; Subr. & L.K.Banerjee, Bull. Bot. Surv. India 10(1): 103. 1968; V.Chandras. in A.N.Henry et al. (eds.), Fl. Tamil Nadu Ind., Ser. 1: Analysis 131. 1987; Janarth. & A.N.Henry, Taxon 38: 141. 1989, Bladderworts India 95. f. 28. 1992; Anil Kumar et al., Fl. Pathanamthitta 368. 2005; Nayar et al., Fl. Pl. W. Ghats India 587. 2014. Lectotype (designated by Janarthanam & Henry, 1989): INDIA, Anamalais, Paralai, 07.11.1901, C.A. Barber 3982 (MH [ MH00002434 !]).

Utricularia racemosa sensu Wight, Icon. Pl. Ind. Orient. f. View in CoL 1. 4: 1584. 1850, non Wall. ex Walp., 1843.

Utricularia rosea sensu Oliv., J. Proc. Linn. Soc., Bot. View in CoL 3: 184. 1858, non Edgew. 1847; C.B.Clarke in Hook.f., Fl. Brit. India 4: 333. 1884. Figs. 1 & 2 (a–d)

Terrestrial or lithophytic annual herbs. Rhizoids up to 35 mm long, c. 0.2 mm thick, capillary. Stolons c. 75 mm long, terete, glabrous, branches up to 40 mm long. Leaves usually 4–6 on peduncle base, rosulate, numerous on stolons; petioles up to 8 mm long; lamina oblanceolate to oblance-ovate, 6.35–8.8 × 1–1.5 mm, rounded at apex, 1-nerved, nerves rarely branched. Traps more on stolons, few on petiole, oblique, globose, 1.08–1.3 mm across; stalk 0.3–1 mm long; mouth latero-terminal, body expanded into a circular, coiled or reflexed structure, with glandular hairs. Racemes erect, simple or rarely branched; peduncles terete, 45–180 mm long, 0.3–1.3 mm thick, purplish green, glabrous, 1–8-flowered. Scales attached obliquely below the middle, oblong, 1.6–2.3 × 0.6–0.85 mm, 1-nerved, acute at apex, bi-tridentate at lower part, minutely papillate. Bracts basisolute, attached just above the middle, rhombic, 1.25–2.4 × 0.4–0.9 mm, upper acute at apex, lower acute to bi-dentate at apex, minutely papillate; bracteoles basisolute, attached just below the middle, ovate to rhombic, 1.3–1.45 × 0.4–0.6 mm long, 1-nerved, acute at both ends, minutely papillate. Flowers 6–13 mm long; pedicels 0.8 – 1.2 mm long, erect, terete, slightly deflexed in fruit, papillate. Calyx lobes unequal, greenish-violet, or rarely yellowish green, convex, glabrous within, outer papillate; upper lip widely ovate, 2.7–4 × 3–4 mm, inflated at middle, acute-obtuse at apex; lower lip sub-orbicular, 1.5–2.6 × 2.2–2.7 mm, margins involute, obtuse-rounded at apex. Corolla pink to purple, or rarely greenish white to white; upper lip oblong, galeate, constricted at middle, 3–10 mm long, apical margin undulate, lower margins hairy, truncate-emarginate or rounded at apex; lower lip oblong, moderately flared towards tip, purple with a yellow patch at the palate, crested with two hooklike process at margins of throat with glandular hairs, palate extends into a prominent 4-lobed swelling with white patches, expands gradually into a sub-orbicular structure, margin undulate, 5.5–13 mm long, emarginate-rounded at apex; spur subulate, 5–8.5 mm long, shorter than or slightly equalling the lower lip of corolla, inner surface glandular, outer surface glabrous, obtuse-rounded at apex. Stamens 1.5–2 mm long; filaments flattened, strap-shaped, 1.2–1.6 × 0.3–0.55 mm, twisted at base; anther thecae distinct. Pistil c. 2.5 mm long; ovary ovoid-globose, 1.7–2.2 mm long, greenish purple; style c. 0.4 mm long, distinct, glandular hairy; stigma 2-lipped, lower lip sub-orbicular, curved inwards, upper lip tri-fid, obsolete, subulate, glandular hairy. Capsules ovoid, 2–3 mm long, wall prominently thickened, dehiscing by both dorsal and ventral longitudinal slits; placenta ovoid, 1–1.5 mm long. Seeds numerous, obovoid–oblongoid, 0.23–0.35 × 0.17–0.23 mm; hilum sub-terminal; testa cells irregularly shaped; surface rough, finely verrucate.

Flowering & fruiting: Flowering from early September till mid of January and fruiting from late September till the end of January.

Habitat: As small patches on mossy wet rocks, stream margins and marshy areas exposed to bright light, between 1000–2200 m elevation, along with Utricularia nayarii Janarth. & A.N.Henry , Burmannia munnarensis Dani & Nampy ( Burmanniaceae ), Apocopsis mangalorensis (Hochst. ex Steud.) Henrad , Arundinella sp. , Dimeria sp. , Jansenella griffithiana (Mull.Hal.) Bor (all Poaceae ), Cyanotis sp. ( Commelinaceae ), Drosera indica L. ( Droseraceae ), Eriocaulon brownianum Mart. , E. nairii Chandrab. & V.Chandras. , E. odoratum Dalzell , E. thwaitesii Korn. (all Eriocaulaceae ), Fimbristylis sp. ( Cyperaceae ), Impatiens rakthakesara M.Vishnu & Nampy ( Balsaminaceae ), Laurembergia coccinea Kanitz. ( Haloragaceae ), Smithia racemosa B.Heyne ( Fabaceae ) and some other grasses ( Fig. 3a–d).

Distribution: Western Ghats and Sri Lanka ( Fig. 4).

Specimens examined: INDIA, Kerala, Idukki district, Elaveezhapunchira, 29.10.2020, Santhosh Nampy & M.P. Krishnapriya 164389; Eravikulam National Park, 2100 m, 29.11.1979, Nambiar 1124 (KFRI); Ibid., grassy slope, ± 1722 m, 23.12.2019, M.P. Krishnapriya & Santhosh Nampy 164338; Kattadikadavu, 07.12.2018, Santhosh Nampy & M.P. Krishnapriya 168045 (CALI); Kulamavu, 03.10.1983, C.N. Mohanan 79967 (MH00126665); about 15 km from Idukki on the road to Kulamavu, 31.08.1973, Cook, Rix & Schneller 67 (P [P04425315 digital image]); Lockhart gap, Devikulam, 1675 m, 12.10.1963, K.M. Sebastine 17542 (MH [MH00126663, MH00126966]); Kumali road on dripping rocks, 13.12.1985, M.K. Janarthanam 82964 (MH [MH00126666, MH00126667]; BSID [BSID0012642]); Mathiketan Shola National Park, Vattachola grassland, 08.11.2018, Syam Radh & M.P. Krishnapriya 168033; Ibid., 07.12.2018, Syam Radh & M.P. Krishnapriya 168044 (CALI); Munnar, 15.11.1961, C. Saldanha CS 8045 (BLAT); Pettimudi grassland, ± 1800 m, 10.12.2019, M.P. Krishnapriya & Santhosh Nampy 164335; Ibid., ± 1903 m, 29.12.2019, M.P. Krishnapriya & Dani Francis 164341; Pettimudi Hilltop,waytoSubranshed,± 2089m, 30.12.2019 M,. P. Krishnapriya & Santhosh Nampy 164344; Subran shed, near stream, 30.12.2019, M.P. Krishnapriya 164348; Upputhara, Vakavanam, 14.09.2019, Dani Francis & M.P. Krishnapriya 164308 (CALI); Palakkad district, Nelliyampathy, Kaarasoori, ± 1130 m, 23.11.2022, M.P. Krishnapriya 186026; Nelliyampathy Hill top, ± 1175 m, 24.11.2022, M.P. Krishnapriya 186028, 186029; Silent Valley National Park, Aruvampara, 12.11.2020, M.P. Krishnapriya & Santhosh Nampy 164396 (CALI); Pathanamthitta district, Kokkathode, 08.09.1989, N. Anilkumar 2024 (CAL); Thiruvananthapuram district, Agasthyamala, 21.12.2018, M.P. Krishnapriya 168053 (CALI); Western slopes of Agasthymala, 1400 m, 06.10.1973 J. Joseph 44629 (MH00126954); Ponmudi, opposite Indian Institute of Space Science complex, 02.11.2021, M.P. Krishnapriya 177227 (CALI). Tamil Nadu, Coimbatore district, Anamalai, around Attakatti, 25.01.1962, J. Joseph 13565 (MH [MH00126674]); Ibid., Konalar, ± 1300 m, 13.12.2018, Resmi S. & M.P. Krishnapriya 168050; Ibid., December 2019, S. Resmi 164329 (CALI); Grass hills above Iyerpadi, 10.01.1901, C.A. Barber 4025 (MH [MH00126676, MH00126677]); Poonachi, 10.10.1901, C.A. Barber 3716 (MH [MH00127006, MH00127007]); Dindigul district, Kodaikanal, new reservoir, 04.08.1985, K.M. Matthew 41687 (RHT [RHT044648]); swampy high ground near lake outflow, 20.05.1987, Bourne 16 (K [K000779503, K000779504 digital images]); Blackburn slopes facing Palamalai (Kavalamalai), 1400 m, 11.11.1987, K.M. Matthew 48220 (RHT [RHT044605]); Palni Hills, lake marsh by south, 08.09.1986, K.M. Matthew & M. Charles 46762 (RHT [RHT044602]); Peak north of Perumal Peak, 2100 m, 12.12.1986, K.M. Matthew 47915 (RHT [RHT044603]); Madurai district, Lake marsh, 10.05.1986, s.col., 8675 (RHT [RHT044594]); Tirunelveli district, Agasthiar malai, 22.05.1901, C.A. Barber 2936 (MH [MH00126670]); way to Mahendragiri beyond Sengamal estate, 1025 m, 05.12.1969, B.V. Shetty 33050 (MH [MH00126671, MH00126672]); Muthukuzhivayal, ± 1400 m, 29.09.1980, A.N. Henry 68866 (MH [MH00126668, MH00126669]); SRI LANKA, s.loc., s.d., s.col. CV277 (MH [MH00274881]); s.loc., 1873, R.H. Beddome s.n. (MH [MH00126417]); s.loc., 1847, Gardner 507 (K [K000779493, K000779502 digital images]); Central Province, Kandy district, 4 mi. directly SW of Maskeliya fishing hut area on the margin of the Moray Occurrence group tea estate, at the SE base of Adams Peak, 1400 m, 21.11.1974, Gerrit Davidse & D.B. Sumithraarachchi 8705 ( US [ US 03216814 digital image]); Nuwara Eliya district, boggy ground along right side of route A-5 near pass going from Hakgala to Nuwara Eliya, 13.09.1969, Robert W. Read & P.E. Desautels 2276 ( US [ US 03216859 digital image]).

Conservation status: It grows in a few localities in southern Western Ghats (Kerala and Tamil Nadu) and Sri Lanka in protected areas with an estimated Area of Occupancy (AOO) of 108 km 2 and an Extent of Occurrence (EOO) of 60011.492 km 2. It is prone to habitat alterations due to landslides and climate change. With the available data and considering the threats, the status is provisionally assessed as Endangered (EN) B2ab(ii,iii,iv,v)D) in the Indian context ( IUCN 2012, 2022).

Notes: The name Utricularia roseopurpurea was originally based on a specimen by Stapf in the Kew herbarium (K), but we could not locate this specimen even after thorough search. Gamble (1924) validated this name in his Flora of the Presidency of Madras, and cited the locality as “ W. Ghats, Anamalais, Pulneys and hills of Travancore, to 7000 ft., in swamps ”. Janarthanam and Henry (1989) designated a lectotype. Bhattacharyya (1976) treated U. caerulea ( U. racemosa wall ex Walp.), U. roseopurpurea , and U. nivea under the U. caerulea complex, in which U. nivea was the white morphotype of the latter (see Abraham et al., 1974). Taylor (1977, 1989) synonymized U. roseopurpurea under U. caerulea , and this was followed by others ( Li, 2007; Janarthanam et al., 2020; Narasimhan & Sheeba, 2021; POWO, 2023), except Janarthanam and Henry (1992), who considered U. roseopurpurea distinct. Poorly preserved or incomplete herbarium specimens, that lack sufficient floral detail, often result in misidentification of specimens ( Baleeiro et al., 2022) and this might be applicable to U. roseopurpurea (for further notes, see under discussion).