Justicia quinqueangularis J.Koenig ex Roxb., Fl. Ind.

P., Soumya, Y. F., Deng, S. S., Dash & Sunojkumar, P., 2023, A taxonomic revision of the genus JUStICIa (Acanthaceae) in Peninsular India, Rheedea 33 (4), pp. 270-331 : 305-310

Justicia quinqueangularis J.Koenig ex Roxb., Fl. Ind. 1: 134. 1820; C.B.Clarke in Hook.f., Fl. Brit. India 4: 536. 1885; Gamble, Fl. Madras 2: 1081. 1921; V.A.W.Graham, Kew Bull. 43: 597. 1988. Rostellaria quinquangularis (J.Koenig ex Roxb.) Nees in Wall ., Pl. Asiat. Rar. 3: 101. 1832. Rostellularia quinquangularis (J.Koenig ex Roxb.) Nees in DC., Prodr. 11: 375. 1847. Type: INDIA, s.loc., s.d., Banks s.n. (C [C10004992 digital image!]).

Rostellaria peploides Nees in Wall ., Pl. Asiat.Rar.3:101. 1832. Rostellularia peploides Nees in DC., Prodr. 11: 375.1847. Justicia peploides (Nees) T.Anderson J. Linn. Soc., Bot. 9: 511. 1867; Malik & Ghafoor in E.Nasir & S.I.Ali (eds.), Fl. Pakistan 188: 33. 1988; Y.F.Deng, Fl. Pan Himalaya 46: 407. 2020; V.A.W.Graham, Kew Bull. 43: 597. 1988. Justicia quinquangularis J.Koenig ex Roxb. var. peploides C.B.Clarke in Hook.f., Fl. Brit. India 4: 536. 1885. Lectotype (designated by Deng, 2020): INDIA, Ripa Gangis and Bhagulpur, 1820, Wall . Cat . 2441B (K-W [K001115997 digital image!]).

Justicia quinqueangularis J.Koenig ex Roxb. var. kawalii Annamma & Venu , in, Indian J. Forest. 39(2): 179. 2016, syn. nov. Type: INDIA, Telangana , Adilabad district , Kawal Tiger Reserve , Jannaram Forest Division , Indhanpalli range, 15.09.2014, Annamma P.S. 4044 a (holo CAL [ CAL0000024888 !]; iso BSID [ BSID0017169 !]). FigS. 22 & 23

Perennial herbs, up to 70 cm tall., Stems decumbent to erect, usually rooting at lower nodes, angular, dichotomously branched; branches slender, greenish, scabrous, pubescent with strigose hairs; internodes 2–9 cm long. Leaves sub-sessile or shortly petiolate; laminae linear to lanceolate or elliptic to narrowly oblong, 1.5–8 × 0.4–0.8 cm, base attenuate, scabrous, margins entire to distantly crenate with minute hairs, apex acute or obtuse, chartaceous, puberulous with dense hairs on ribs; veins inconspicuous, secondary veins 4–7 pairs, venation weak brochidodromous; petioles up to 5 mm long, densely pubescent. Inflorescence terminal spike, up to 9 cm long; peduncles 0.5–1.5 cm long, densely pubescent. Flower sessile, two per inflorescence node, 4–9 × 2–4 mm, bracts and bracteoles almost equal to calyx; bracts linear to narrowly oblong or ovate to lanceolate, 3.5–4.5 × 1–1.3 mm, margin scarious, ciliate, apex acute to obtuse or cuspidate; bracteoles similar to bracts but narrower, 2.5–3.5 × c. 1 mm. Calyx deeply 5-lobed, 3.5–5 × 1–1.25 mm at anthesis, accrescent, greenish; lobes unequal, lanceolate, the fifth lobe much reduced, margins scarious, ciliate, apex acute or obtuse. Corolla purplish white with white striations on lower lip, pubescent outside and glabrous inside; tube cylindrical, 2–2.5 × 1–1.3 mm, slightly enlarged at throat, glabrous outside and pubescent inside on throat; upper lip erect, triangular to ovate, 2.5–3.5 × 2–2.2 mm, apex obcordate; lower lip orbicular to obovate, 4.5–5 × 5–5.5 mm, 3-lobed at apex; lobes imbricate, apex rounded or obtuse. Stamens exserted; filaments 2 mm long, declinate, white, basally hirsute; anther thecae slightly superposed, equal, elliptic, c. 0.8 mm long, purplish with pale brown shades, upper theca dorsally hirsute, lower theca glabrous; appendage white, c. 0.5 mm long, apex acute or rounded. Ovary ellipsoid, 1–1.2 × 0.4– 0.6 mm, pale green, densely pubescent throughout; style slender, 3.5–4 mm long, whitish, hairy in lower half; stigma slightly bifid; nectariferous disc c. 0.5 mm long, pale green, margins undulate. Capsule oblong, 3.5–4 × 1–1.2 mm, apex acute, pale brown, pubescent at apex and along margins. Seeds 4, reniform to orbicular, compressed, 1–1.2 × 1 mm, brownish; testa rugulose.

Flowering & fruiting: Flowering from July to December; fruiting in January.

Habitat: Grows in humus-rich soil, on moist rocks, along the margins of paddy fields and streams at an elevation of 450 to1100 m.

Distribution: Afghanistan, Bangladesh, China, India, Nepal, Pakistan, Sri Lanka and Taiwan.

Specimens examined: INDIA, Andhra Pradesh, East Godavari district, Samalkota, 29.04.1913, s.coll. s.n.; Ibid., 07.01.1917, s.coll. 14024; Ibid., 19.12.1924, G.V. Narayanan 81943 (MH); Prakasam district, Diguvametta, Nallamalais, 18.11.1969, J.L. Ellis 38489 (CAL); Visakhapatnam district, Araku Valley near stream, 23.08.1960, N.P. Balakrishnan 10748; Srungavarpukota, 22.10.1964, G. V. Subba Rao 21824 (MH); Warangal district, Western side of Pakhal Lake, 08.08.1961, K.M. Sebastine 13131 (CAL). Karnataka, Kodagu district, Madhapur- Mercara road, 08.10.1961, s.coll. 74985 (CAL); Mysore district, Siligiri Rangan hills, 07.09.1978, S.R. Ramesh 2605 (CAL). Kerala, Kollam district, Mannarappara, 15.11.1976, M. Chandrabose 49069 (MH); Ibid., 15.11.1970, M. Chandrabose 49069 (CAL); Palaruvi, 28.11.1961, K.N. Subramanian 77199 (FRC); Kottayam district, Muthuvara, s.d. Robi A.J. 06589 (CMPR); Palakkad district, Kunthipuzha bank, 16.01.1980, P. Bhargavan 65504 (CAL); Silent valley R.F., 11.12.1980, N.C. Nair 69549 (CAL); Thiruvananthapuram district, Bonacaud, 09.10.1973, J.Joseph 44644 (MH); Ibid., 22.12.1987, N. Mohanan 9096 (CALI); Ibid., 17.05.1988, N. Mohanan 9894 (CALI); Kulathupuzha range, Velayankala coup 10, 14.09.1984, K.R. Sasidhararn 10482 (FRC); Ponmudi, 18.05.1973, M. Mohanan 54873 (MH); Ibid., 09.11.2018, P. Soumya, Resmi & P. Sunojkumar 138573 (CALI); Wayanad district, Baveli, Begur R.F., 22.06.1979, V.S Ramachandran 62723 (CAL); Nadavayal, s.d., P. Soumya, Drisya, A.K. Pradeep & P. Sunojkumar 145302 (CALI).

Conservation status: Due to its wide distribution, this species is assessed here as Least Concern (LC).

Notes: This species is often mistaken with another annual species Justicia vahliana Schult. , based on the similarities in the leaf shape and size. However, J. quinqueangularis J.Koenig ex Roxb. can be distinguished from it by its dense cylindrical spikes (vs. less congested spike), a 5-lobed calyx with the fifth lobe much reduced (vs. four lobed), linear to narrowly lanceolate bracts and bracteoles (vs. ovate) with widely scarious margins (vs. narrow scarious). The species exhibits considerable variability in its vegetative characters, with plants in wet and dry habitats often looking different. Its habit varies from small decumbent herb to erect subshrub and the leaves are linear to lanceolate, occasionally elliptic to narrowly oblong.

Annamma et al. (2016) described Justicia quinqueangularis var. kawalii Annamma & Venu from the Kawal Tiger Reserve in Telangana, differentiating it from other varieties based on characteristics such as cylindrical spikes with flowers of the same size on successive nodes. However, upon further investigation and comparison with different populations from Peninsular India, it was observed that the characteristics used to describe the new variety were inconsistent and varied within individuals of the same population and between populations. Consequently, this variety is treated as a new synonym of J. quinqueangularis in the present study.

We also treat Justicia peploides (Nees) T.Anderson as a synonym of J. quinqueangularis . It was originally described as Rostellaria peploides Nees ( Nees von Esenbeck, 1832) . Anderson (1867) transferred it to Justicia as J. peploides . Subsequently, Clarke (1885) noticed that J. peploides is morphologically similar to J. quinqueangularis and reduced it as a variety under the latter based on its diffuse branched stem with oblong or ovate leaves and shorter spikes. In the present study we confirmed that the oblong to ovate leaf with short spike falls within the variation range represented by Justicia quinqueangularis . However, taxon is still treated as a distinct species in some major floras ( Stewart, 1972; Malik & Ghafoor, 1988; Graham, 1988; Deng, 2020). The morphological variation in J. quinqueangularis primarily pertains to its habit, leaves and inflorescences, with plant size and inflorescences ranging from small to large and leaf shape from linear to lanceolate or elliptic to narrowly oblong. Occasionally, bracts and bracteoles may also exhibit slight variations in shape from linear or narrowly oblong to lanceolate.

Justicia santapaui Bennet, J. Bombay Nat. Hist. Soc. 67: 358. 1970. Hemichoriste montana Nees in Wall ., Pl. Asiat. Rar. 3: 102. 1832; in DC., Prodr. 11: 367. 1847; Wight Ic. t. 1538. 1850. Justicia montana (Nees) T.Anderson , in J. Linn. Soc. Bot. 9: 509. 1867 nom. illeg., non Roxb. 1805; C.B.Clarke in Hook.f., Fl. Brit. India 4: 525. 1885. Gamble, Fl. Madras 2: 1078. 1921; V.A.W.Graham, Kew Bull. 43: 589. 1988. Justicia andersonii Ramamoorthy in C.J.Saldanha & Nicolson, Fl. Hassan district, 551. 1976, nom. superfl. Lectotype (designated by Graham, 1988): INDIA, Wall . Cat . 2471a (K [K001116149, digital image!]). FigS. 24 & 25

Perennial shrubs, up to 2 m tall. Stems erect, angular, dichotomously branched; branches stout, green to purplish green, glabrous or puberulous; internodes 2–13 cm long. Leaves petiolate; laminae elliptic or oblanceolate, 11–28 × 3.5–8 cm, base cuneate, margins entire, slightly undulate, apex acuminate or acute, sub-coriaceous, glabrous; veins prominent, secondary veins 6–11 pairs, venation simple craspedodromous; petioles 2–5 cm long, glabrous. Inflorescence terminal, simple or compound spike, up to 35 cm long, many-flowered, dense towards base and lax at apex; peduncles 1.5–5 cm long, glabrous. Flowers sessile or sub-sessile, fascicled, 1–3 flowers on each side, 2.75–3.5 × 1.5–2 cm; bracts ovate to widely lanceolate, 0.7–1.5 × 0.4–0.7 cm, margins entire, apex acuminate or acute, green or purplish green, pubescent. Bracteoles oblong to elliptic, 0.5–0.7 × 0.25–0.4 cm, apex acute, green, pubescent. Calyx deeply 5-lobed, 0.7–1.3 × 0.4–0.6 cm at anthesis, accrescent, green or purplish green; lobes lanceolate to subulate, equal, densely pubescent, apex acuminate, margins entire, finely ciliate. Corolla whitish with purple striations on lips, pubescent outside, glabrous inside except in throat; tube cylindrical, 4–8 × 4–5.5 mm, slightly enlarged in throat; upper lip erect, elliptic to narrowly oblong, 2–2.3 × 0.6–0.75 cm, apex minutely two lobed, hooded; lower lip obovate to sub-orbicular, 2–2.25 × 1.5–1.75 cm, 3-lobed; lobes sub-equal, middle lobe broadly oblong to ovate, apex rounded; lateral lobes narrowly oblong, apex acute or rounded. Stamens exserted; filaments c. 1.5 cm long, declinate or sometimes curled, white, basally hirsute; anther thecae superposed, equal, elliptic, 3–4 mm long, greenish with yellow margin, upper theca dorsally hirsute, lower theca glabrous; appendage white, c. 1 mm long, apex acute, pointed. Ovary conical, c. 3 × 2 mm, pale green, densely tomentose; ovules 4; style filiform, 2.1–2.4 cm long, whitish recurved, hirsute in lower half; stigma simple; nectariferous disc c. 1.5 × 3 mm, greenish yellow. Capsule clavate, 2.5–3 × 0.5–1 cm, apex acute or shortly pointed, green to brown, densely pubescent; base sterile, 1–1.3 cm long, flattened. Seeds 4, orbicular or sub-orbicular, 2.7–2.9 × 3–3.2 cm, glabrous; testa tuberculate.

Flowering & fruiting: Flowering from November to February; fruiting from March to April.

Habitat: This species is distributed on the margins of evergreen and semi-evergreen forests, along Ghat roadsides and riverbanks at elevations between 550 and 1500 m.

Distribution: South India.

Specimens examined: INDIA, Karnataka, Bellary district, s.d., Law s.n. ( CAL); Chikkamagaluru district, Kemmannugundi , 26.03.1964, R . S . Raghavan 37216 ( CAL); Dakshina Kannada district, Shirady Ghat near border, 27.12.1978, P . Prakesh & S . B . Manohar 5523; Someshwar village , 04.02.1980, N . Rameswara Rao 122 ( CAL); Hassan district, Yettinahala , 27.01.1971, T . P . Ramamoorthy 1346 ( CAL); Kodagu district, Coorg , 04.02.1976, B . C . Banerjee 11553; Ibid., Attic estate, 26.01.1976, B . C . Banerjee 11342 ( CAL); Mysore district , s.loc., 01.02.1906, A . Meebold 8502( CAL); Shimoga district, Agumbe , 05.02.1961, R . S Raghavan s.n. ( CAL); Jog falls , 17.01.2018, P . Soumya, V . S . Hareesh, Adithya & P . Sunojkumar 138503 ( CALI); Kodachadri range, 07.03.1979, B . R . Ramesh & K . R . Kesava murthy 6164 ( CAL) . Kerala, Ernakulam district , Thundathil range, Bhoothathankettu, 12.02.1970, B . V . Shetty 33487 ( MH); Idukki district, Kanchiyar , 17.02.1982, C . N . Mohanan 73328 ( MH); Ibid. , 23.02.1983, C . N . Mohanan 77953 ( CAL, MH); Kulamavu , 14.12.1982, C . N . Mohanan 76054 ( CAL, MH); Ibid. , 07.11.2018, P . Soumya & P . Sunojkumar 138521 ( CALI); Malappara to Thanipara , 15.03.2009, P . S . Udayan et al. 05730 ( CMPR); Manaloda, 22.02.1994, Jomy Augustine 13358 ( CALI); Meenmutty , 18.02.1982, V . S . Raju 71252 ( CAL, MH); Meenmutty to Kulamavu , 25.01.1983, A . G . Pandurangan 76627; Ibid., 25.01.1983, A . G . Pandurangan 76632 ( CAL); Ibid. , 25.02.1984, A . G . Pandurangan 78085 ( CAL, MH); Painavu , 28.04.1984, C . N . Mohanan 81655 ( MH); Valara R . F ., 07.02.1984, K . Ramamurthy 80898 ( MH); Kannur district, Ambayathode , 15.12.1979, V . S . Ramachandran 65227 ( CAL); Chanthanathode , 14.02.1978, V . S . Ramachandran 53957; Ibid., 22.02.1979, V . S . Ramachandran 60100 ( MH); Kollam district, Way to Plappally, 23.02.1979, C . N . Mohanan 61181 ( CAL, MH); Kottayam district, Erumeli , 23.12.1978, C . N . Mohanan 59616 ( CAL, MH); Malappuram district, Nilambur, Thalichola , 03.01.1981, Philip Mathew 25662 ( CALI); Ibid. , 04.02.1982, Philip Mathew 33929 ( CALI); Palakkad district , Dhoni hills, 27.03.1910, C . E .C. Fischer 1838 ( CAL, FRC); Karappara river , 16.04.1914, E . Vajravelu 48969 ( CAL, MH); Ibid. , 01.03.1975, E . Vajravelu 46119 ( MH); Mannarkkad, Koodam , Muthikkulam, 03.02.2009, K . A . Anilkumar 3814; Ibid., Kuruvampadi, 14.01.2007, K . A . Anilkumar 2711 ( CMPR); Olavakkot range, Muthikulam , 24.03.1976, K . N . Subramanian 5810 ( FRC); Silent valley, Cheriya Walakkad , 21.12.1969, E . Vajravelu 33218 ( MH); Ibid. , 08.04.1978, N . C . Nair 56656 ( CAL, MH); Ibid. , 11.01.1979, E . Vajravelu 59112 ( CAL, MH); Ibid. , 04.02.1982, T . Sabu 10044 ( CALI); Pathanamthitta district, Plappally , 27.02.1985, N . Anilkumar 519 ( MH); Thrissur district, Chimmini , 21.03.1988, K . Ramamurthy 66243 ( CAL, MH); Vellarimala , 06.02.1985, N . Sasidharan 3389 ( CALI); Thiruvananthapuram district, Ponmudi , 08.03.1980, M . Mohanan 66611 ( MH); Ibid., Upper sanatorium area, 03.02.1986, K . N . Subramanian, B . Gurudev Singh & N . Venkatasubramanian 11719 ( FRC); Wayanad district, Adivaram , 22.07.2018, P . Soumya & P . Sunojkumar 138526; Puthumala, 08.04.2001, M . K . Ratheesh Narayanan 2548 ( CALI); Wayanad Ghat region, 01.1989, s.coll. s.n.; Ibid. , 21.01.1903, C . A . Barber 5677 ( MH); s.loc., 13.01.1894, T . F . Bourdillon 30 ( CAL) . Tamil Nadu, Coimbatore district , Anamalai, 10.03.1961, J . Joseph 12310; Anamalai waterfall estate, 24.03.1969, R . Maruthan 31752; Ibid., s.d., Beddome s.n.; Karian shola, 15.03.1931, V . Narayanaswami 5343; Ibid., 25.02.1942, S . R . Raju 86373 ( MH); Karian shola top slip, 11.03.1958, K . Subramanyam 5561 ( CAL, MH); Ibid. , 06.03.1978, K . N . Subramanian 7155 ( FRC); Sanjeevi hill, 14.02.1957, K . M . Sebastine 2309 ( CAL, MH); Sholaiyar Dam side, 28.12.1963, K . Ramamurthy 18155; Ibid., 30.3.1965, K . Ramamurthy 22887; Yanaikundhi shola, 29.01.1962, J . Joseph 13818 ( MH); Dindigul district, Thandikudi , 27.04.1965, K . Ramamurthy 23495 ( MH); Nilgiris district, Devala Rockwood forest , 23.11.1972, E . Vajravelu 43457 ( MH); Nadukani Ghat road, s.d., P . Soumya, Shinoj & P . Sunojkumar 145338 ( CALI); Tirunelveli district, 1858, Beddome 25 ( CAL); Vellore district, Kilandur , 21.03.1964, R . S . Raghavan 37044 ( CAL); Virudhunagar district, Sethur hills, 07.03.1981, S . R . Srinasan 68023 ( CAL, MH) .

Conservation status: In Peninsular India, this species is found in Karnataka, Kerala, and Tamil Nadu with good population in many districts. Currently, the species is classified as Least Concern ( LC), following the IUCN criteria (2022) .

Figures

Fig. 22. Justicia quinqueangularis J.Koenig ex Roxb.: a. Habit; b. Inflorescence; c. Flower; d. Flower bud; e. Bract; f. Bracteoles–adaxial side;g. Calyx; h. Calyx–opened;i. Corolla–opened; j. Upper lip; k. Lower lip; l. Stamens;m. Gynoecium with nectariferous disc; n. Capsule; o. Seeds (from P. Soumya & P.Sunojkumar 145341; photos by P. Soumya).

Fig. 23. Distribution of Justicia quinqueangularis J.Koenig ex Roxb. in Peninsular India.Drawn using R (R Core Team,2023).

Fig. 24. Justicia santapaui Bennet:a. Habit; b. Inflorescence; c. Flower; d. Flower bud; e. Calyx; f. Corolla–opened; g. Upper lip; h. Lower lip; i. Stamens; j. Gynoecium with nectariferous disc;k. Capsule; l. Seed (from P. Soumya & P. Sunojkumar 145338; photos by P.Soumya).

Fig. 25. Distribution of Justicia santapaui Bennet in Peninsular India. Drawn using R (R Core Team,2023).