Harpochilus neesianus Martius ex Nees (1847: 146)

Costa-Lima, James Lucas Da & Chagas, Earl Celestino De Oliveira, 2019, A revision of Harpochilus sheds light on new combinations under Justicia (Acanthaceae), Phytotaxa 393 (2), pp. 119-130 : 122-124

Harpochilus neesianus Martius ex Nees (1847: 146) .

TYPE:— BRAZIL. Bahia: “ in ripa arenosa fl. S. Fran. ad Joazeiro,” s.d., Martius 2275 (lectotype, designated here: M [barcode M0186236 !]; isolectotypes: M [bc M0186237 !], M [bc M0186238 !]). ( Fig. 2A–B )

Suffrutescent, 0.5–1.5(–2.5) m tall; stems cylindrical, usually with inflated internodes at base, striated longitudinally, tomentose. Leaves opposite, decussate; petioles 0.2–2 cm long; blades 1.5–7.6 × 0.7–2.5 cm, elliptic, lanceolate, ovate or obovate, apex obtuse to acute, margin short-ciliate, base obtuse to attenuate, adaxial surface tomentose, abaxial surface tomentose, villous on veins, cystoliths inconspicuous, venation eucamptodromous, secondary veins in 5–8 pairs, alternate or subopposite, veins impressed on adaxial surface, prominent on abaxial surface. Inflorescences in dichasial cymes, with 3–5 sessile flowers per inflorescence unit, apparently secund with usually only 1 flower per node, main rachis and peduncles tomentose, peduncles 0.2–1.8 cm long; first order bracts 1.5–2.5 × 0.8–1.2 cm, decussate on the main axis, pale green to olive, leaf-like, non-ciliate at margin; second and third order bracts 2, lateral, 0.4–0.6 × 0.3–0.5 cm, spatulate to obovate, apex rounded, non-ciliate at margin; bracteoles 2, lateral, 0.2–1 × 0.1–0.2 mm, linear to oblong, apex rounded, non-ciliate at margin. Calyx 5-lobed; lobes 0.6–2 × 0.1–0.4 cm, equal, lanceolate to linear, tomentose to sparsely pubescent on the outside, sparsely pubescent on the inside, mixed with glandular trichomes on both sides, margins sparsely setose. Corolla greenish to greenish yellow, bilabiate, glabrous to sparsely pilose inside the tube and lobes, sparsely pilose mixed with glandular trichomes outside; tube 1.2–2.5 cm long, cylindrical, inflated at base, erect; upper lip 4–7.5 × 0.4–0.8 mm, spatulate, narrow, curved downward, non-ciliate, sinuous at margin, enveloping the style and the filaments for most of their length, 2-lobed at apex, the lobes 1–1.5 mm long; lower lip 3-lobed, the lobes 2.5–6 × 0.3–0.5 mm, spatulate, narrow, irregularly curled downward, non-ciliate, sinuous at margin. Filaments 4.8–8 cm long, sericeous, arcuate; anthers 5–7.6 mm long, with 2 parallel thecae, equal, non-appendiculate, sagittate at base. Ovary 2–3.5 × 2–3 mm; style 4.5–12 cm long, glabrous for most of its length, sparsely setose at base, arcuate; stigma punctiform; nectary disc annular. Capsule 2–3.5 cm long, sterile basal portion 1–1.5 × 0.3–0.5 cm, fertile apical portion 1–1.5 × 0.3–0.6 cm, broadly ellipsoid, irregularly sulcate in sicco, glabrous, glaucous when young, chestnut-brown to reddish when ripe; seeds 4, 8–10 mm diam., suborbicular, glabrous, flat.

Distribution and habitat: — Harpochilus neesianus occurs in the semiarid region of northeastern Brazil (Rio Grande do Norte, Paraíba, Pernambuco, Alagoas, and Bahia) and has been recorded in both the Caatinga domain, in xerophilic vegetation, and transition areas with the Cerrado domain, in south-central Bahia near Minas Gerais ( Fig. 1 ). It is found on sedimentary substrates, such as sandy and limestone outcrops, as well as crystalline substrates, especially rocky outcrops.

Conservation status: —Due to Harpochilus neesianus having a relatively wide distribution in the Brazilian semiarid region (EOO = ca. 312,000 km 2), it is not under any threat and falls under the category Least Concern (LC), according to IUCN (2017) criteria.Although the species is widely distributed, it occurs in areas that have suffered from severe deforestation (e.g., extensive cattle farming, removal of trees for firewood, extraction of sand and kaolin for construction, and the creation of wind farms). Despite the abundance of specimens deposited in herbarium collections, evidence suggests that only two subpopulations are within protected areas, Estação Biológica de Canudos (Bahia) and Parque Nacional do Catimbau (Pernambuco).

Specimens examined: — BRAZIL. Alagoas: Mun. Olho D’Água do Casado, Fazenda Morro Grande , 9º31’03.3’’S, 37º48’49.5’’W, 8 August 2000 (fr), D. Moura & R. A. Silva 1327 ( RB) ; ibidem, Xingó , caminhos para os cânios [sic], 9º31’53’’S, 37º50’36’’W, 22 October 2016 (fl), J. W. Alves-Silva et al. 1589 ( MAC) . Bahia: Sine loco accuratio, Rio S. Francisco, Marais d’Ilhabira , 1838–1839, Blanchet 2884 (syntypes: BM, F, K, P, W) ; Mun. Barra, Ibiraba (= Icatu ), em frente à vila de Ibiraba no caminho para os brejos, 10º48’S, 42º50’W, 23 February 1997 (fl), L. P. de Queiroz 4793 ( ESA, HUEFS, MBM) ; ibidem, 26 February 1997 (fl), L. P. de Queiroz 4887 ( HUEFS, MBM) ; ibidem, Serra do Estreito, subindo a partir da localidade de Olho D’Água , 11º1’13’’S, 43º20’15’’W, 409 m, 19 May 2010 (fl), L. P. de Queiroz et al. 14649 ( ASE, HUEFS, UEC) ; ibidem, duna no Distrito de Ibiraba , 10º47’24’’S, 42º49’19’’W, 399 m, 31 July 2013 (fl, fr), D. M. Neves et al. 1518 ( HUEFS – 2 sheets, RB) ; Mun. Caetité , Brejinho das Ametistas, 900 m, 15 April 1983 (fl, fr), A. M. de Carvalho et al. 1777 ( CEPEC, HUEFS) ; ibidem, 18 km da cidade, localidade de Santa Luzia , 14º8’19’’S, 42º27’33’’W, 850 m, 10 March 1994 (fl), V. C. Souza et al. 5413 ( ESA, MBM, MO, RB, UFRN) ; ibidem, 14º8’48’’S, 42º32’29’’W, 960 m, 11 February 1997 (fl), M. L. Guedes et al. PCD 5441 ( CEPEC, HUEFS, RB) ; ibidem, 14º15’22’’S, 42º31’20’’W, 880 m, 13 March 2002 (fl), N. Roque et al. 634 ( HUEFS) ; ibidem, 14º8’22’’S, 42º27’48’’W, 800 m, 14 April 2002 (fl, fr), F. França et al. 3744 ( HST, HUEFS, UB) ; ibidem, Distrito de Brejinho das Ametistas , 14º16’12’’S, 42º31’40’’W, 10 January 2008 (fl), A. L. Côrtes et al. 66 ( HUEFS) ; Mun. Canudos, estrada para Reserva Biológica de Biodiversitas , 10º1’58’’S, 39º9’00’’W, 22 February 2000 (fl), A. M. Giulietti & R. M. Harley 1729 ( ALCB, HUEFS, UFRN) ; ibidem, ca. 10 km de Canudos, na estrada para Uauá , 9º55’15’’S, 38º1’55’’W, 416 m, 26 June 2002 (fl), L. P. de Queiroz et al. 7158 ( HUEFS) ; ibidem, ca. 6 km SE de Canudos na estrada para Estação Biológica de Canudos , 9º55’22’’S, 38º59’13’’W, 391 m, 26 June 2002 (fl, fr), L. P. de Queiroz et al. 7172 ( HUEFS, UFP) ; ibidem, Estação Biológica de Canudos , 10º1’00’’S, 39º9’00’’W, 24 April 2003 (fl, fr), F. H. M. Silva et al. 375 ( HUEFS) ; ibidem, caminho para a Gruta do Minadouro , 9º56’35’’S, 38º59’18’’W, 390 m, 22 May 2003 (fl), A. A. de Oliveira et al. 164 ( HUEFS) ; ibidem, 9º56’44’’S, 38º59’18’’W, 381 m, 17 February 2004 (fl), R. M. Harely et al. 54865 ( HUEFS) ; ibidem, 9º56’41’’S, 39º00’55’’W, 23 March 2007 (fl), M. C. Dórea & R. P. Oliveira 21 ( HUEFS) ; ibidem, 9º57’S, 39º00’W, 27 April 2007 (fl), A. L. Côrtes et al. 01 ( HUEFS – 2 sheets) ; ibidem, 9º57’S, 39º00’W, 27 April 2007 (fl), A. L. Côrtes et al. 02 ( HUEFS) ; ibidem, próximo à cidade, 9º53’19’’S, 38º51’54’’W, 393 m, 5 June 2008 (fl, fr), F. França et al. 5757 ( HUEFS) ; Mun. Casa Nova, próximo a Casa Nova Velha , 17 km do entroncamento na estrada BA 235 para Remanso , 9º24’36’’S, 41º8’58’’W, 27 April 2001 (fl, fr), R. M. Harley et al. 54328 ( ALCB, CEPEC, HUEFS) ; ibidem, ca. de 30 km de Remanso para Casa Nova , 9º23’8’’S, 41º49’4’’W, 481 m, 16 June 2001 (fl), T. S. Nunes et al. 505 ( ALCB, CEPEC, HUEFS) ; ibidem, Fazenda Santarém , 9º32’20’’S, 41º21’31’’W, 440 m, 5 October 2003 (fl, fr), K. R. B. Leite et al. 404 ( HUEFS) ; ibidem, ca. 17 km do entroncamento para Casa Nova Velha com a BR, próximo à entrada do Balneário Dunas do Velho Chico , 9º24’57’’S, 41º8’24’’W, 418 m, 5 July 2003 (fl, fr), L. P. de Queiroz et al. 7899 ( HUEFS) ; ibidem, 9º32’4’’S, 41º21’12’’W, 452 m, 28 November 2003 (st), L. P. de Queiroz et al. 7997 ( HUEFS) ; ibidem, 9º32’4’’S, 41º21’12’’W, 452 m, 28 November 2003 (fl), L. P. de Queiroz et al. 8019 ( CEN, HUEFS) ; ibidem, 9º24’53’’S, 41º9’7’’W, 414 m, 30 November 2003 (fr), L. P. de Queiroz et al. 8082 ( HUEFS) ; ibidem, estrada para Pau a Pique ca. de 11 km da entrada, 9º23’48’’S, 41º39’59’’W, 591 m, 18 April 2004 (fl), T. S. Nunes et al. 1086 ( HUEFS) ; ibidem, BR-235, Dunas do São Francisco , cerca de 10 km a oeste do entroncamento, 9º22’22’’S, 41º10’49’’W, 380 m, 4 July 2004 (fl, fr), G. Pereira-Silva et al. 9156 ( CEN, HUEFS, RB) ; ibidem, 9º21’32’’S, 41º22’29’’W, 430 m, 9 October 2004 (fl), L. P. de Queiroz et al. 9608 ( ESA, HUEFS) ; [Mun. Gentio do Ouro], 62 km N of Gentio do Ouro on road to São [sic] Inácio , 11º08’S, 42º45’W, 500 m, 23 February 1977 (fl, fr), R. M. Harley et al. 18975 ( CEPEC, K, NY) ; ibidem, ca. 24 km S de Xique-Xique, na estrada para Santo Inácio, 16 July 1994 (fl, fr), L. P. de Queiroz & N. S. Nascimento 3960 ( FLOR, HUEFS) ; ibidem, 27 km de Xique-Xique , 11º3’49’’S, 42º44’2’’W, 400 m, 1 June 1999 (fr), E. Melo et al. 2699 ( HUEFS) ; Mun. Glória, Brejo do Burgo , 1 July 1995 (fr), F. P. Bandeira 168 ( HUEFS) ; ibidem, caminho da Serrota, 3 July 1995 (fl), F. P. Bandeira 243 ( HUEFS, MBM) ; ibidem, APA de Itaparica, Caçimba [sic], 11º7’38’’S, 42º44’23’’W, 5 May 2002 (fl), M. C. Fereira et al. 1265 ( ALCB, HUEFS – 2 sheets) ; ibidem, Baixa de Fontana , 9º20’S, 38º18’W, 9 March 2004 (fl), M. V. Moraes 619 ( HUEFS) ; ibidem, Raso da Catarina , 9º20’S, 38º29’W, 6 June 2004 (fl, fr), M. V. M. Oliveira 702 ( HUEFS) ; ibidem, Baixa do Fontana , 9º20’S, 38º29’W, 6 September 2004 (fl, fr), M. Colaço & T. Moreira 18 ( HUEFS) ; ibidem, 9º20’S, 38º29’W, 419 m, 28 August 2006 (fl), S. Leal & M. Nascimento 01 ( HUEFS, MO) ; Mun. Jaguarari, Morro do Flamengo , 10º6’1’’S, 40º13’43’’W, 635 m, 27 July 2005 (fr), P. D. Carvalho et al. 119 ( HUEFS, MBM) ; Mun. Jeremoabo, a 6 km na estrada de chão sentido Canudos , 10º5’53’’S, 38º25’8’’W, 400 m, s.d. (fl), D. S. Carneiro-Torres et al. 498 ( CEPEC, HUEFS) ; ibidem, Muro , 10º11’45’’S, 38º18’4’’W, 376 m, 18 October 2009 (fl), E. Melo et al. 6783 ( ALCB, HUEFS) ; Mun. Juazeiro, Serra do Mulato , 9º44’40’’S, 40º40’39’’W, 27 March 2000 (fl), G. Cavalcanti et al. 63 ( ALCB, BAH, CEPEC, HUEFS, RB) ; Mun. Paulo Afonso, Raso da Catarina, Baixa do Chico , 9º31’54’’S, 38º38’11’’W, 544 m, 11 August 2005 (fl, fr), J. G. Carvalho-Sobrinho et al. 560 ( HUEFS) ; ibidem, 9º43’53’’S, 38º40’58’’W, 541 m, 29 November 2005 (fl), R. M. Castro et al. 1279 ( HUEFS) ; ibidem, 9º48’32’’S, 38º29’32’’W, 715 m, 1 December 2005 (fr), A. O. Moraes et al. 111 ( HUEFS) ; ibidem, Aldeia Serrote , 9º29’S, 38º5’W, 6 June 2006 (fl), M. Colaço 130 ( HUEFS) ; Mun. Pilão Arcado, localidade de Goiabeira , dunas continentais, 13 August 1998 (fl), A. M. Miranda et al. 3003 ( HST, HUEFS) ; ibidem, Goiabeira do Brejo do Zacarias , 10º1’37’’S, 42º52’4’’W, 427 m, 6 September 2005 (fl, fr), L. P. de Queiroz et al. 10878 ( HST, HUEFS) ; Mun. Ribeira do Pombal , 10º44’49’’S, 38º53’34’’W, 235 m, 24 February 2006 (fl), E. Melo et al. 4265 ( HUEFS) ; Mun. Senhor do Bonfim, Crista de quartzito, 10º2’38’’S, 40º13’3’’W, 710 m, 14April 2005 (fl), E. Melo et al. 3792 ( HUEFS) ; Mun. Sento Sé, estrada Minas do Mimoso para Umburanas , ca. 5, 7 km de Minas do Mimoso , 10º17’2’’S, 41º22’31’’W, 792 m, 8 September 2016 (fl), L. P. de Queiroz et al. 16278 ( HUEFS) ; Mun. Tucano, ca. 23 km na estrada de Tucano para Euclides da Cunha , 22 March 1992 (fr), A. M. de Carvalho et al. 3931 ( CEPEC, HUEFS, NY, CEPEC) ; ibidem, Sítio do Mandacaru, Faz. do Sr. Gildásio , 10º52’29’’S, 38º46’19’’W, 391–411 m, 15 April 2006 (fl), D. Cardoso & M. Guerreiro 1196 ( HUEFS) ; ibidem, BR-116, a N da cidade de Tucano sentido Euclides da Cunha, próximo da Baixa da Areia , 10º57’S, 38º43’W, 6 March 2010 (fl), D. Cardoso 2836 ( HUEFS) ; Mun. Uauá, ca. 51 km N de Monte Santo na estrada para Uauá, 25 August 1998 (fl), L. P. de Queiroz & N. S. Nascimento 4637 ( CEPEC, HUEFS, MBM) ; Mun. Uibaí, Serra Azul , 11º20’10’’S, 42º08’29’’W, 600 m, 17 March 1996 (fl), R. Atkinson et al. PCD 2486 ( CEPEC, HUEFS) ; Mun. Xique-Xique, Dunas do Rio São Francisco , 10º47’16’’S, 42º46’22’’W, 548 m, 23 June 1996 (fl), A. M. Giulietti et al. PCD 2978 ( HUEFS) ; ibidem, Cacimba, Estrada entre Xique-Xique e Barra , próximo à entrada para Gentio do Ouro , 11º7’42’’S, 44º21’8’’W, 412 m, 30 June 2008 (fr), A. A. Conceição et al. 2918 ( HUEFS) ; ibidem, 11º7’32’’S, 42º44’14’’W, 407 m, 16 May 2009 (fl), F. França et al. 5968 ( HUEFS) . Paraíba: Mun. Remígio , regioés secas [sic], proximo a lagêdos [sic], 20 September 1959 (fl), J. C. de Moraes s.n. ( EAN 2242 , NY [barcode NY00484268 ], P [bc P02901556 ]) ; Mun. Serra Branca, Pedra de Ser. Branca , 7º30’51’’S, 36º44’94’’W [sic], 22 February 2002 (fl), M. R. Barbosa et al. 2225 ( HUEFS, JPB) ; ibidem, 7º28’59’’S, 35º39’54’’W, 493 m, 8 March 2002 (fl), M. F. Agra 5711 ( HUEFS – 2 sheets, JPB) ; ibidem, 7º30’29’’S, 35º45’494’’W, 21 May 2002 (fr), M. F. Agra et al. 5949 ( HUEFS, JPB) .

Pernambuco: Mun. Buíque, Trilha das Torres, 8º37’S, 37º10’W, 790 m, 17 October 1994 (fl, fr), M. J. N. Rodal 417 ( PEUFR, NY) ; ibidem, Catimbau, 20 September 1997 (fl, fr), A. M. Miranda et al. 2827 ( HST, HUEFS) ; ibidem, Vale do Catimbau , 22 November 2005 (fr), A. Bocage et al. 1114 ( HUEFS) ; ibidem, na estrada em direção à reserva indígena dos Capinauás, 8º35’13’’S, 37º15’40’’W, July 2007 (fr), O. Cano et al. 777 ( HUEFS) ; ibidem, 8º35’42’’S, 37º14’7’’W, 815 m, 21 November 2018 (fl, fr), J. L. Costa-Lima & E. C. O. Chagas 3062 ( HUEFS, MAC, UFRN) ; Mun. Gravatá, margens da BR-232, próximo ao Haras Luís Inácio, 8º12’31’’S, 35º36’39’’W, 16 June 2008 (fl, fr), R. Pereira et al. 2803 ( HUEFS) ; ibidem, 14 December 2013 (fl), M. E. Alencar 2323 ( HUEFS) ; Mun. Ibimirim, estrada para Moxotó, próximo ao sítio Baixa, 8º41’25.54’’S, 37º30’00.35’’W, 516 m, 6 November 2012 (fl, fr), A. C. P. Oliveira et al. 1810 ( HUEFS, HVASF) ; [Mun. Petrolina], arredores de Petrolina, 18 April 1971 (fl), E. P. Heringer et al. 120 ( RB, UB). Rio Grande do Norte : Mun. Equador, Serra das Queimadas, área de extração de caulin [sic] e para instalação do Complexo Eólico Santapape, 6º54’43’’S, 36º42’58’’W, 754 m, 13 August 2015 (fl), E. O. Moura et al. 413 ( UFRN) .

Discussion: — Harpochilus neesianus was described by Nees (1847) based on the collections Martius 2275 and Blanchet 2884. In accordance with Art. 9.3 of the ICN ( Turland et al. 2018), we designated a lectotype for this species. We chose one of the three specimens of the collection Martius 2275 in the Botanische Staatssammlung München herbarium (M) because it was annotated by C.F.P. von Martius as “ Justicia neesiana .”

Among the three known species of Harpochilus analysed in this study, H. paraibanus and H. neesianus are the most morphologically similar, especially for their complex and peculiar floral morphology. Harpochilus phaeocarpus , an ascending shrub endemic to forests in southern Bahia, is clearly more morphologically similar to Justicia and is here combined under this genus (see comments below).

Figures

FIGURE 1. Distribution map of Harpochilus neesianus (circles), H. paraibanus (squares), Justicia citrina (star), and J. phaeocarpa (triangles), with the Brazilian semiarid region highlighted in orange.

FIGURE 2. Harpochilus neesianus. A. Flowering branch, with a detail of the corolla. B. Capsule and seed, in sicco. Harpochilus paraibanus. C. Flowering branch. D. Capsule and seed, in sicco. Photograph credits:A. Domingos (A), A.S.F. Castro (C), J.L. Costa-Lima (B, D).