Campanula feijoana Gardère

Gardère, Mathieu L., Florence, Jacques, Muller, Serge, Savriama, Yoland & Dubuisson, Jean-Yves, 2021, Codonographia Gorgonum, or the description of a pleiad of bellflowers (Campanula, Campanulaceae) from the Cabo Verde archipelago, Candollea 76 (1), pp. 13-40 : 26-28

3. Campanula feijoana Gardère in Phytotaxa 197: 105. 2015 ( Fig. 1 , 3C , 5A , 7 ).

Holotypus: CABO VERDE. Santo Antão: Ribeira da Vinha , rocher humide en station ombrophile, 17°08'1"N 25°04'12"W, 400 m, 13.XII.2013, Gardère 460 ( LISC [ LISC118130 , LISC118131 ]!; iso-: CECV!, K!, P [ P00723702 ]!) .

= Campanula hortelensis Gardère in Phytotaxa 197: 109. 2015 , syn. nov. Holotypus: CABO VERDE. Santo Antão: Monte Hortelão, rochers humides, au bord de la piste, 17°06'03"N 25°10'57"W, 1500 m, 14.XII.2013, Gardère 467 (LISC [ LISC118148 ]!; iso-: CECV!).

Sub-frutex 20– 80 cm tall, highly woody in lower part; floriferous stems branched, decumbent to pendulous arising from the base of one or several sterile basal rosettes, sometimes understated, glabrous to glabrescent in the woody basal parts with indument hispidulous toward the extremity, consisting of trichomes 0.15–0.4 mm long. Leaves: pseudorosette leaves or rosette leaves obovate to spatulate, sometimes falciform, (1.4–) 2–6(–9) × (0.4–)0.9– 2(–2.7) cm, base cuneiform to gradually attenuate concave sometimes asymmetric, apex acute to ± obtuse; cauline leaves narrowly obovate to narrowly elliptic rarely elliptic, (0.7–)1.5 – 5(–7.5) × (0.5 –)0.8 –1.8(– 2.5) cm, base attenuate sometimes asymmetric, apex acute to ± obtuse; margin weakly revolute and sometimes obscurely ondulate, crenelate to serrulate, rarely entire; adaxial side pure green to dark green in vivo, weakly or densely covered with hispidulous to hispid indument, consisting of trichomes 0.2–0.6(–0.8) mm long, sometimes glabrescent; abaxial side greenish in vivo, venation whitish, hispidulous to hispid indument on primary and secondary veins consisting of trichomes (0.2–)0.3–0.6(–0.8) mm long and hispidulous indument on tertiary and ultimate veins consisting of trichomes 0.1–0.3 mm long, lamina glabrescent. Inflorescences in monochasial pauciflorous cyme or rarely in pluriflorous thyrse. Flowers erect, pedicel 0.9–4.5 cm long, with the same indument as the stem; axillate by one or two bracts subopposite, ovato-triangular or ovate to narrowly ovate, base semi-amplexicaul, apex acute, with the same indument as the leaves. Calyx: calyx-lobes narrowly triangular, 12–17 × 3–9.5 mm, pressed up against the corolla to recurvate, margin weakly revolute; appendages ovate, reflexed, 1–3 mm long; lobe edges, appendages and median main vein covered with an indument hispidulous to hispid, consisting of trichomes 0.15–0.7 mm long. Corolla infundibuliform, purple-blue to pure white passing through all intermediate tones; base wide round; tube sub-cylindrical c. 20 mm long, constricted in the middle, 9–12 mm large in the larger basal part and 7–10 mm in the constricted part; throat flared, mouth 22–32 mm large; lobes spreading to recurvate, 6–10 × 11–18 mm, apex apiculate; primary external veins micro-hispidulous to hispidous, 0.1–0.12 mm long. Stamens with glabrous filaments; anthers, 2–4 mm long. Ovary with glabrous to glabrescent roof, flat, topped by a yellowish-with nectary disk. Style thick, fleshy, (8–) 9–11 mm long, included in the corolla, stigma trifid and papillose.

Etymology. – The species is dedicated to the Portuguese naturalist João da Silva Feijó (GARDÈRE, 2015) who undertook the first scientific expedition entirely dedicated to the study of the natural history of the Cabo Verde islands between 1783 and 1796 during the “Philosophical Journeys” ( GARDÈRE et al., 2019a) and who made the first collections of CVB between 1783 and 1789.

Vernacular names and uses. – “Guinchino” (CARDOSO JÚNIOR, 1905; CHEVALIER, 1935), “Mataquim” (in Água das Caldeiras, Hiemstra H236; CARDOSO JÚNIOR, 1905), “Contra-Bruxas” and “Dedal” (CHEVALIER, 1935; BARBOSA, 1961; LEYENS & LOBIN, 1995; FIGUEIREDO, 1995); the colour adjectives “branco” (white) or “azul” (blue) are sometimes added to the name; “Hortelãoda-Rocha” in Moroços areas (Gardère 1560). However, in the archipelago according CHEVALIER (1935) “Mataquim” can also designate Corchorus trilocularis L. ( Malvaceae ) and on the island of Santo Antão, BARBOSA (1961) records this name for Antirrhinum orontium L. ( Scrophulariaceae ).

Campanula feijoana was once used for its magical and medical properties. Healers [curandeiros] used the leaves and flowers to make a tea to treat flu (LEYENS & LOBIN, 1995) and used it as a fetish plant against curses: “Contra-Bruxas” meaning “anti-witch”; the practice was to wash the body with the infused water and to put a few drops on the tongue. This tradition was still very much alive in the beginning of the last century in the Alto Mira region and in the Ribeira Corvo (Gardère, unpubl. data). CARDOSO JÚNIOR (1905) also mentioned, without going into detail, the use of this plant for medical purposes by the islanders.

Distribution and habitat. – Campanula feijoana , endemic to Santo Antão, is a rupicolous species confined to steep, moist rocks. It is found from 150 m to 1700 m, in deep and shadowy valleys [ribeiras] ( Fig. 7A ), surrounding waterfalls, on seeping rock faces or near to springs [chupadeiros], or rarely on river banks; and up to the highest mountainous areas, on rock faces ( Fig. 7F ) regularly submitted to dense fog. Campanula feijoana grows with ferns, notably Adiantum capillus-veneris (Pteridaceae) and Pteris vittata (Pteridaceae) and with other species that are characteristic components of these rupicolous environments, like Kickxia elegans (G. Forst.) D.A. Sutton (Scrophulariaceae) and Blumea axillaris (Lam.) DC. (Asteraceae) .

Notes. – The bellflowers from Santo Antão have long been identified as C. jacobaea (COUTINHO, 1914; CHEVALIER, 1935; SUNDING, 1973, 1982; ERIKSSON et al., 1974, 1979; NOGUEIRA, 1976; HANSEN & SUNDING, 1985, 1993; RUSTAN & BROCHMANN, 1993; LEYENS & LOBIN, 1995; FIGUEIREDO, 1995; SÁNCHEZ-PINTO et al., 2005), moreover material from this island (Forbes s.n. [K001134390]) was chosen by WEBB (1848: tab. 762) to be part of the syntypes of this species (see under C. jacobaea ). In his infraspecific division of C. jacobaea, BOLLE (1861) considered the bellflowers from Santo Antão and those from the upper humid areas of São Nicolau (see under C. fransinea ), under the type variety “ genuina ”. Then, CARDOSO JÚNIOR (1902) designated the white-flowered bellflowers from Santo Antão under the variety “ albiflora ”. Without description this variety remains a nomen nudum.

More recently, the bellflowers from Santo Antão were described as two new species (GARDÈRE, 2015): (1) C. feijoana ( Fig. 7A–E ) was described with pauciflorous inflorescences, calyx-lobes spread-out to obliquely erect prolonged by reflexed appendages and pseudorosette leaves spatulate with papery lamina, whereas (2) C. hortelensis ( Fig. 7F, G ) was described with pluriflorous inflorescences, calyx-lobes appressed against the corolla tube and prolonged by curved appendages and rosette-leaves elliptical to obovate often falciform with subleathery lamina. But the recent discovery of new localities with intermediate forms between C. feijoana and C. hortelensis (e.g. Gardère 1555) obscures the clean morphological and geographical separation of the two species. Campanula feijoana was found mainly in shadowy valleys and C. hortelensis in mountainous areas, hence they seem to represent altitudinal ecotypes belonging to a single and unique species. Given that C. feijoana is more widespread and representative of bellflowers with infundibuliform constricted corollas on the island, we conserve the name C. feijoana and treat C. hortelensis as a synonym of C. feijoana . In this way, the range of C. feijoana now covers almost the entire island of Santo Antão, except for the mountains of the far eastern part (Bordia Perdia) where bellflowers with flared infundibuliform corollas are found (see under C. vicinituba ).

Selected material seen. – CABO VERDE. Santo Antão: Água das Caldeiras, 1250 m, 8. I.1987, Hiemstra H236 ( WAG) ; Água dos Velhos , 800 m, X.1986, Hiemstra H669 ( FR) ; Bordeira Norte , 1550 m, 16.XII.2013, Gardère 521 ( LISC) ; Chã da Lagoa , VIII –IX.1893, Cardoso Júnior s.n. ( LISU) ; between Chã de Morte and Tope de Coroa , 1310 m, 14. I.1982, Brochmann & Rustan CB -311/82 ( O) ; ibid. loco, 1430 m, 22. I.1982, Brochmann & Rustan CB -409/82 ( O) ; along the path from Chã de Morte to Tope de Coroa , 1430 m, 22. I.1982, Rustan & Brochmann ØHR -1715 ; Cima Monte Joana , 950 m, 25.X.1972, Sunding 2714 ( O) ; Cova , 1180 m, 4. I.1982, Brochmann & Rustan CB -056/82 ( O) ; ibid. loco, 23–28.IX.1934, Chevalier 45512 ( P) ; ibid. loco, 1250 m, 28.XI.2017, Gardère 1551 ( P) ; ibid. loco, c. 1200 m, 12.XII.1985, Kilian 857 ( FR) ; entre Cova y Lagoa , 1100 m, 18. V.1987, González - Coviella Ulrich 4060 ( CECV, TFC) ; Covão , 16–22.IX.1934, Chevalier 45422 ( P) ; ibid. loco, 800 m, 15.XI.2014, Gardère 797 ( P) ; Delgadinho de Corda , 750–800 m, 15.XII.2013, Gardère 508 ( LISC, P) ; Espanada , 1340 m, 7.XI.2014, Gardère 727 ( P) ; Gudo de Cavaleiro , 1750 m, 7.XI.2014, Gardère 716 ( P) ; Gudo da Fonte , 950 m, 29.XI.2017, Gardère 1555 ( CECV, LISC, P) ; Lombo do Mar , 630 m, 5. I.1982, Rustan & Brochmann ØHR -1341 ( O) ; entre Lombo Pelado e Curral da Ruça , 17.III.1956, Grandvaux Barbosa 6924 ( CECV, LISC) ; Monte Hortelão , 1500 m, 8.XI.2014, Gardère 731 ( P) ; ibid. loco, 1500 m, 29.XI.2017, Gardère 1560 ( CECV, LISC, P) ; Monte Manuel Joelhos , 400 m, 13.XI.2014, Gardère 760 ( P) ; Monte Pingo , 1550 m, 29.XI.2017, Gardère 1562 ( CECV, K, LISC, LPA, MARS, P) ; Moroços , 1460 m, 2.II.1995, Leyens CV -95-262 ( FR) ; Pinhão , 500 m, 16.XI.2014, Gardère 800 ( P) ; Poio , 2.III.1956, Grandvaux Barbosa 6796 ( CECV, LISC) ; Ribeira da Água Amarogosa , c. 1250 m, 8.II.1995, Leyens CV -95-350 ( FR) ; Ribeira de Cabouco Chiqueiro , 1500 m, 9.XI.2014, Gardère 740 ( P) ; Ribeira do Cativo , 28.III.1956, Grandvaux Barbosa 7008 ( CECV) ; Ribeira das Chapas , 970 m, 8.XI.2014, Gardère 734 ( CECV, LISC, P) ; Ribeira do Círio , 1500 m, 16.XII.2013, Gardère 526 ( LISC) ; Ribeira do Corvo , 340 m, 18.XI.2017, Gardère 1513 ( CECV, P) ; Ribeira da Cruz , 1000 m, 6.II.1995, Leyens CV -95-317 ( FR) ; Ribeira Fria , 675 m, 11.XI.2014, Gardère 752 ( P) ; ibid. loco, 580 m, 6.II.1995, Leyens CV -95-314 ( FR) ; Ribeira da Garça , 900 m, 23.XI.2017, Gardère 1531 ( CECV, P) ; ibid. loco, 4.IV.1956, Grandvaux Barbosa 7109 ( CECV, LISC) ; ibid. loco, 9.II.1995, Leyens CV -95-362 ( FR) ; ibid. loco, 15. I.1866, Lowe s.n. ( BM) ; Ribeira da Igreja , 22.XI.1979, Lobin 1654 ( FR) ; Ribeira de Janela , 520 m, 27.XI.2017, Gardère 1544 ( CECV, LISC, P) ; ibid. loco, c. 1050 m, 10.II.1995, Leyens CV -95-375 ( FR) ; Ribeira João Afonso , III.1887, Cardoso Júnior s.n. ( COI) ; Ribeira dos Órgãos , 350 m, 2.XI.2014, Gardère 660 ( P) ; Ribeira das Patas , c. 1550 m, 8.II.1995, Leyens CV -95-346 ( FR) ; Ribeira de Paúl, XII.1852, Bolle s.n. ( FI-W) ; ibid. loco, Chã João Vaz , 400 m, 15.XI.2014, Gardère 794 ( P) ; ibid. loco, 300–400 m, 29.XII.1993, Kilian & Leyens 2989 ( B, FR) ; ibid. loco, Descida de Ribeiraozinho , 675–750 m, 12.VII.2004, Marrero & Almeida s.n. ( LPA) ; ibid. loco, Descida de Santa Isabel , 750–775 m, 12.VII.2004, Marrero & Almeida s.n. ( LPA) ; ibid. loco, 21.XI.1979, Lobin 1607 ( CECV, FR) ; ibid. loco, 6.III.1866, Lowe s.n. ( BM) ; ibid. loco, 820 m, 20. I.1980, Rustan 848 ( O) ; ibid.loco, 400 m, 9. I.1982, Rustan & Brochmann ØHR -1386 ( O) ; ibid. loco, 380 m, 24.X.1972, Sunding 2634 ( O) ; Ribeira das Pedras , 200 m, 6.XI.1976, Sunding 3434 ( O) ; montanhas de Paúl, IX.1892, Cardoso Júnior 55 ( COI, Z) ; Ribeira das Pombas , 250 m, 18.XI.2017, Gardère 1518 ( CECV, LISC, P), 1519 ( MARS); Ribeira da Ponta do Sol , 150 m, 19.XI.2015, Gardère 942 ( P) ; Ponta do Sol , III–IV.1893, Cardoso Júnior s.n. ( LISC) ; ibid. loco, III–IV.1893, Cardoso Júnior s.n. ( LISU) ; between Vila Ribeira Grande and Ponta do Sol , 220 m, 3. I.1982, Rustan & Brochmann ØHR -1273 ( O) ; Ribeira do São Jorge , XI.1893, Cardoso Júnior s.n. ( LISU) ; Ribeira da Torre , 350 m, 20.XI.2015, Gardère 959 ( CECV, P) ; ibid. loco, 180 m, 11.XII.1985, Kilian 808 ( B, FR) ; ibid. loco, 24.XII.1978, Lewejohann CV -78-180 ( GOET) ; ibid. loco, III.1864, Lowe s.n. ( FI-W) ; ibid. loco, 4.III.1864, Lowe s.n. ( K) ; ibid. loco, Xôxô , 400 m, 14.X.1990, Martins & Gomes 320 ( LISC) ; ibid. loco, 15.IX.1986, Mies 50 ( FR) ; ibid. loco, 1460 m, 13.II.1994, Leyens CV -94-124 ( B, FR) ; Ribeira da Vinha , 400 m, 9.VIII.2013, Gardère & da Costa e Silva 196 ( P) ; ibid. loco, 400 m, 21.XI.2017, Gardère 1522 ( P) ; Ribeirãozinho de Cima , 28.III.1956, Grandvaux Barbosa 7014 ( CECV, LISC) ; Selada da Alto Mira , 1127 m, 1.XII.2015, Aedo 23393 ( MA) ; Tope de Coroa , 1700 m, 9.XI.2014, Gardère 742 ( P) ; sine loco, s.d., Cardoso Júnior I ( L) ; sine loco, s.d., Cardoso Júnior II 83 ( L) ; sine loco, s.d., Cardoso Júnior II 122 ( L) ; sine loco, s.d., Cardoso Júnior II 148 ( L) ; sine loco, V.1887, Cardoso Júnior s.n. ( COI) ; sine loco, I.1893, Cardoso Júnior s.n. ( Z) ; sine loco, s.d. [IV.1822], Forbes s.n. [4] ( K p.p.: remaining syntype for C. jacobaea ) ; sine loco, s.d., McWilliam s.n. ( K) ; sine loco, s.d., Missão Técnica de Arborização s.n. ( LISC) ; sine loco, “ in rupestribus ins. S. Antonii ”, III.1851, Schmidt s.n., ( HBG p.p.) .

Figures

Fig. 1. – Distribution map of the seven Cabo Verdean Campanula L. species illustrated by their flower in lateral view.

Fig. 3. – SEM micrographs of abaxial surface of leaves. A. Campanula jacobaea C. Sm. ex Webb; B. C. bravensis (Bolle) A. Chev.; C. C. feijoana Gardère; D. C.vicinituba Gardère; E. C. monteverdensis Gardère; F. C. fransinea Gardère; G. C. cochleromena Gardère. [A: Gardère 1428; B: Gardère 1615; C: Gardère 959; D: Gardère 1035; E: Gardère 612; F: Gardère 1096; G: Gardère 1120]

Fig. 5. – Second herbarium sheet from K containing the syntypes of Campanula jacobaea C. Sm. ex Webb. A. Forbes’ specimen [K001134390] determined here as C. feijoana Gardère; B. Vogel’s specimen [K001134391] determined here as C. monteverdensis Gardère; C. Hooker’s specimen [K001134407] determined here as C. jacobaea. [© Royal Botanic Gardens,Kew]

Fig. 7. – Campanula feijoana Gardère. Upright form from low-elevation locality: A. Habit; B. Pseudorosette-leaves; C. Inflorescence; D. Flower in lateral view; E. Flower in face view.Form of high-elevation locality in dense rupicolous shrubbery: F. Habit; G. Rosette-leaves. [A: Gardère 1522; B–E: Gardère 1518; F–G: Gardère 1562] [Photos: M.L. Gardère]