Campanula bravensis (Bolle) A. Chev., 1935

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 : 20-24

2. Campanula bravensis (Bolle) A. Chev. in Rev. Bot. Appl. Agric. Trop. 15: 889. 1935 ( Fig. 1 , 3B , 4A –A’, 6).

 Campanula jacobaea var. bravensis Bolle in Bonplandia 9: 51. 1861.

Lectotypus (designated by LEYENS & LOBIN, 1995: 222): CABO VERDE. Brava: “in rupestribus ins.: Brava frequens”, XII.1852, Bolle s.n. ( K [ K 001134396 ]!) .

Sub-frutex 20 – 60 cm tall, highly woody in lower part; floriferous stems branched, erect or decumbent to procumbent arising from the base of one or several sterile basal rosettes, glabrous to glabrescent in the woody basal parts with indument hispidulous to hispid toward the extremity, consisting of trichomes 0.2–0.6 mm long. Leaves: rosette leaves narrowly obovate to narrowly elliptic, (2–)3.5– 6(–8) × (0.6–)0.9–1.7 (–2.3) cm, base cuneiform to attenuate, apex obtuse to acute; cauline leaves narrowly obovate to narrowly elliptic, rarely ovate, (1.5–)3–5(–8) × (0.5–)1–1.5(–3) cm, base attenuate sometimes asymmetric, apex acute to ± obtuse; margin weakly revolute, crenulate to serrulate; adaxial side light green to pure green in vivo, weakly or densely covered with hispidulous to hispid indument, consisting of trichomes (0.1–)0.2–0.6(–0.75) mm long sometimes with a slight canescent aspect in vivo; abaxial side greenish in vivo, venation whitish, hispidulous to hispid indument on primary and secondary veins consisting of trichomes 0.4–0.6(–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 generally ± pendulous or erect, pedicel 1.5–4 cm long, with the same indument as the stem; axillate by one or two bracts subopposite, ovato-triangular or ovate to narrowly ovate, base semiamplexicaul, apex acute, with the same indument as the leaves. Calyx : calyx-lobes narrowly triangular, 10–15 × 3–5 mm, erect to recurved rarely pressed up against the corolla tube, margin weakly revolute; appendages ovate, reflexed, 1.5–2 mm long; lobe edges, appendages and median main vein covered with an indument hispidulous rarely hispid, consisting of trichomes 0.3–0.45(–0.6) mm long. Corolla tubulate, generally whitishcream (never pure white) with the veins greenish, lobe edges sometime slightly purplish, rarely corolla entirely purplishblue; base wide round; tube cylindrical, 22–33 × 8 –13 mm, sometimes slightly constricted in the lower quarter giving an aspect bounded at the base; throat straight to slightly constricted occasionally highly constricted giving in extreme cases an urceolate shape to the corolla, mouth c. 20 mm; lobes erect to recurved, 2 –4 × 7.5–11 mm, apex apiculate; primary external veins micro-hispidulous to hispidulous, 0.1 × 0.15 mm long. Stamens with glabrous filaments; anthers 2–4 mm long. Ovary with pubescent roof, conical, topped by a yellowish-with nectary disk. Style thick, fleshy, 17–22 mm long, included in the corolla, stigma trifid and papillose.

Etymology. – The epithet bravensis refers to the type locality, the island of Brava; brava meaning “wild” in Portuguese.

Vernacular names. – Among the CVB species, C. bravensis holds the most of vernacular names: in Brava, “Ortiga-Branca” and “Velho-Teso” (both according BOLLE, 1861); in Fogo, “Frol-Branca” (CHEVALIER, 1935), “João-Copinho” in Campanas de Cima and Ribeira Zória (Gardère 1253), “Pabil” in Bordeira and Chã das Caldeiras (Gardère 1610), “Palha-Barquinho” (in Espigão, Grandvaux Barbosa 6277; FIGUEIREDO, 1995) and “Palha-Caneca” (in Ribeira de São Filipe, Cardoso de Matos 5512; FIGUEIREDO, 1995); in Santiago , “Flôr-Branca” in Pico da Antónia and “Ortiga-Branca” in São Jorge dos Órgãos (both according GOMES, 1994). “Velho-Teso” is also use for Spermacoce verticillata L. ( Rubiaceae ) in Brava (BARBOSA, 1961; DINIZ et al., 2002; MARTINS, 2002).

Distribution and habitat. – Campanula bravensis has the broadest geographical range: it occurs on the three mountainous southern islands, i.e. Santiago, Fogo and Brava (HANSEN & SUNDING, 1993; LEYENS & LOBIN, 1995; BROCHMANN et al., 1997; SÁNCHEZ-PINTO et al., 2005), and is found in the widest range of elevations but in different plant communities according to habitats and islands. In Brava, C. bravensis is found from around 500 m elevation to the highest summits, on rocks regularly submitted to fog with Launaea thalassica N. Kilian et al. (Asteraceae) , Tolpis farinulosa Walp. (Asteraceae) , Daucus sp. (Apiaceae) and sometimes with Nephrolepis undulata J. Sm. (Nephrolepidaceae) or Pteris vittata L. ( Pteridaceae ). In Fogo, it can be found in low-elevation valleys from around 70 m (Brochmann & Rustan CB-916/82) up to the highest point of the island (and the entire archipelago), at around 2850 m (Gardère 1405). In this way, it occupies a diverse range of habitats such as the depths of wet lowland valleys (ribeiras), grassy slopes around Euphorbia tuckeyana Webb (Euphorbiaceae) shrubland, isolated on volcanic ash slopes, on wet rocks, near springs (chupadeiros), or at the entrance of caves with Adiantum capillus-veneris (Pteridaceae) , Pteris vittata L. ( Pteridaceae ) and sometimes with Asplenium adiantum-nigrum L. ( Aspleniaceae ) for the highest elevation locations. Campanula bravensis is very common across these two islands. On the other hand, in Santiago , it is quite rare and is only known from some field stations in the two main mountain ranges, Serra do Pico da Antónia and Serra da Malagueta, where it generally occurs around wet rocks with Pteris vittata (Pteridaceae) or more rarely along riverbeds, and often in sympatry with C. jacobaea .

Notes. – For BOLLE (1861), the concept of his variety bravensis is limited to its type locality, i.e. Brava, because in the protologue he only cited his own collections from this island. Indeed, Bolle made two expeditions to the archipelago in 1851 and 1853 (SALINGER & STREHLOW, 1991), and collected only in Santo Antão, São Vicente, São Nicolau and Brava (BARBOSA, 1962). Later, ANDRADE (1908) extended the concept of the variety bravensis to Fogo. Then, CHEVALIER (1935) raised the variety to the rank of species, adopting a broader concept than currently accepted and which included the islands of Brava, Fogo and Santiago . However, some authors have also extended the distribution of C. bravensis ( PETTERSSON, 1960; SUNDING, 1973; ERIKSSON et al., 1974, 1979) to W São Nicolau probably owing to the presence in this island of plants with white narrow infundibuliform corollas ( C. fransinea ), which slightly resemble C. bravensis .

For C. bravensis , the tubular corolla shape is the most noteworthy and dependable diagnostic feature to distinguish it from the other species. However, this feature is absent from the original description of the basionym. BOLLE (1861) described the variety bravensis using features of the calyx-lobes, a calyx/ corolla length ratio, and on the indument. That said, he did accurately describe the colour of the corolla: yellowish-white with green veins and with the edge of lamina (i.e. edges of corolla-lobes) slightly purplish. CHEVALIER (1935) added depth to the description of BOLLE (1861) by describing the leaf shape which he used, along with the colour of the flower as a diagnostic feature. However, he did not make any descriptions of the shape of the corolla, even if considered unique in the genus, the character being described much later by LEYENS & LOBIN (1995).

Selected material seen. – CABO VERDE. Brava: Cruz Nho Basilo, 22.XI.2014, Gardère 894 ( P); road between Fajã da Água and V.N. Cintra, 500 m, 3.II.1994, Leyens CV -94-065 ( FR); an der Strasse oberhalb Fajã de Água, 26.X.1979, Lobin 1143 ( COI, FR); entre João de Nole et Cruz Nho Basilo, 850 m, 20.XII.2015, Gardère 1155 ( P); Mato Grande, 650 m, 27.X.1984, Cardoso de Matos 5816 ( CECV, LISC); N.S. do Monte, 720 m, 23.XI.2014, Gardère 901 ( CECV, P); am Fussweg zwischen N.S. de Monte und Cova Rodela, c. 650 m, 19.I.1986, Kilian 1186 ( FR); Ribeira Fajã de Água , 580 m, 23.XI.2014, Gardère 906 ( P); de Pedra de Água para V.N. Cintra, 10.X.1956, Grandvaux Barbosa 6611 ( CECV, LISC); Ribeira Tina, 600 m, 17.X.1991, Martins et al. 537 ( LISC); Risco Vermelho , 610 m, 23.XI.2014, Gardère 911 ( P); S of V.N. Cintra, 610 m, 21.II.1982, Rustan & Brochmann ØHR -2400 ( O); ancien chemin de V.N. Sintra à N.S. do Monte, 650 m, 20.VII.2016, Gardère 1234 ( P); ibid. loco , c. 540 m, 30.I.1994, Leyens CV -94-21 ( B, FR); sine loco, 1852, Bolle s.n. ( C, K p.p.: remaining syntype for C. jacobaea var. bravensis ); sine loco, 1853, Bolle s.n. ( Z: remaining syntype for C. jacobaea var. bravensis ); sine loco, “flos sempere flavo albidus”, s.d., Bolle s.n. ( COI: remaining syntype for C. jacobaea var. bravensis ); sine loco, VI.1873, s.c. “Herb. Dr. Sagot” ( P). Fogo: Achada Grande, 70 m, 16.II.1982, Brochmann & Rustan CB -916/82 ( O); Arco, 400 m, 25.I.1994, Kilian 3368 & Leyens ( B, FR); Chã das Caldeiras, 1780 m, 5.I.2014, Aedo 21223 ( MA); ibid. loco , 2800 m, 7.XI.1983, Cardoso de Matos 5611 ( CECV, LISC); ibid. loco , 1800–2000 m, 23–24.VII.1934, Chevalier 44856 ( COI, P); ibid. loco , 1700–1780 m, 14.X.1988, Diniz & Cardoso de Matos 260 ( LISC); ibid. loco , 1800 m, 21.XII.2015, Gardère 1164 ( CECV, P); ibid. loco , 1740 m, 21.X.2016, Gardère 1407 ( CECV, LISC P); ibid. loco , c. 1760 m, 22.I.1994, Kilian 3278 & Leyens ( B, FR); ibid. loco , c. 1600–1750 m, 24.I.1994, Kilian & Leyens 3326 ( B, FR); ibid. loco , 1700 m, 30.X.1979, Lobin 1261 ( FR); ibid. loco , 2.XI.1979, Lobin 1339 ( FR); ibid. loco , 5.XI.1979, Lobin 1413 ( FR); façade nord du cratère et dans le cratère, 4.XII.1985, Peyre de Fabrègues 4216 ( ALF); Chupadeirão , 1350 m, 26.VII.2016, Gardère 1257 ( P); Curral Fundo , 1000 m, 7.VIII.1934, Chevalier 45195 ( P); Domingos Santos , 1920 m, 21.XII.2017, Gardère 1606 ( CECV, MARS, P); Fernão Gomes , 1590 m, 19.XII.2013, Gardère 554 ( LISC); ibid. loco , 1540 m, 13.II.1982, Rustan & Brochmann ØHR -2193 ( O); between Fernão Gomes and Monte Velha, 1500 m, 15.II.1982, Rustan & Brochmann ØHR -2246 ( O); Filho de Palha , 2160 m, 24.XII.2017, Gardère 1615 ( CECV, LISC, P); Fonte de Curral Fumo , 1030 m, 2.VIII.2016, Gardère 1289 ( P); ibid. loco , 1030 m, 3.VIII.2016, Gardère 1291 ( CECV, P); Fonte Djam Djorge , 1710 m, 22.XII.2017, Gardère 1610 ( CECV, P); Fonte Figueirinha , 1660 m, 5.XII.1996, Leyens CV -96-662 ( FR); Furna Defendida, 900 m, 20.X.2016, Gardère 1401 ( P); Espigão , 460 m, 18.X.2016, Gardère 1390 ( CECV, P) ; ibid. loco, 11. I.1956, Grandvaux Barbosa 6277 ( CECV, LISC) ; Monte Duarte , c. 1850 m, 23. I.1994, Kilian 3323 & Leyens ( B, FR) ; entre Monte Cruz e Ponta Alto do Sul, 2100 m, 1.XI.1985, Cardoso de Matos 5992 ( CECV, LISC) ; Monte Sodelho , 880 m, 2.VIII.2016, Gardère 1282 ( P) ; Monte Velha [or M. Velho], 1650 m, 23.X.1985, Cardoso de Matos 5966 ( CECV, LISC) ; ibid. loco, c. 1550 m, 13. I.1986, Kilian 1117 ( B, FR) ; ibid. loco, c. 1400 m, 28.VII.2016, Gardère 1263 ( CECV, P) ; ibid. loco, 1500 m, 17.VII.2004, Marrero & Almeida s.n. ( LPA) ; ibid. loco, 1220 m, 2.II.1980, Rustan 922 ( O) ; Montinho , c. 1800 m, 23.XII.2017, Gardère 1613 ( MARS, P) ; Nhuco , VIII.1934, Chevalier 45178 ( P) ; ibid. loco, 1000 m, 1.VIII.2016, Gardère 1273 ( P) ; ibid. loco, 24.III.1864, Lowe s.n. ( P) ; Penedo Rachado , 1250 m, 20.XI.2016, Gardère 1403 ( CECV, P) ; Pico do Fogo , 2750 m, 21.X.2016, Gardère 1405 ( P) ; Pico Novo , 1750–2800 m, 24. I.1994, Kilian & Leyens 3346 ( B, FR) ; entre Piorno et Monte Cruz , 1540 m, 22.X.2016, Gardère 1416 ( CECV, LISC, P) ; Relva , 360 m, 19.X.2016, Gardère 1394 ( CECV, P) ; Ribeira Campanas , 100 m, 14.X.1991, Martins et al. 492 ( LISC) ; ibid. loco, 490 m, 8.XII.1995, Leyens CV -95-522 ( FR) ; ibid. loco, 5.XI.1979, Lobin 1401 ( FR) ; ibid. loco, 19.X.1982, Lobin 2340 ( FR) ; Ribeira Coxo , c. 1200 m, 28.VII.2016, Gardère 1265 ( P) ; Ribeira Jan Reica , 1200 m, 16.II.1995, Leyens CV -95-421 ( FR) ; Ribeira Monte Preto , 940 m, 17.II.1982, Rustan & Brochmann ØHR -2299 ( O) ; ibid. loco, 1005 m, 25.VII.2016, Gardère 1252 ( P) ; Ribeira São Filipe , 950 m, 1.XI.1983, Cardoso de Matos 5512 ( CECV, LISC) ; Ribeira Zória , 1130 m, 25.VII.2016, Gardère 1253 ( CECV, P) ; Ribeiras im Nordwest-Teil der Insel., c. 250 m, 19.X.1982, Lewejohann CV -82- 165 ( GOET) ; Suspensorio , 1980 m, 24.XII.2017, Gardère 1614 ( P) ; sine loco, 1500 m, X.1898, Newton s.n. ( K) . Santiago: Orgãos Grandes , 300–600 m, IV–V.1898, Fea s.n. ( GDOR) ; Ribeira Cantada , 450 m, 10.VII.1993, Duarte & Gomes 552 ( LISC) ; Ribeira Fundão , 740 m, 14.XI.2016, Gardère 1387 ( CECV, K, LISC, P) ; Ribeira Longueira , 800 m, 25.XI.2014, Gardère 917 ( CECV, P) ; ibid. loco, 15.X.1979, Lobin 1039 ( CECV, COI, FR) ; ibid. loco, 950 m, 18.XI.1976, Sunding 3715 ( O) ; Serra do Pico da Antónia , 25.VIII.1934, Chevalier 44717 ( P) ; ibid. loco, 31.XII.1955, Grandvaux Barbosa 6149 ( CECV, COI, LISC) ; sine loco, s.d. [I–II.1832], Darwin s.n. [279] ( CGE, K p.p.: remaining syntype for C. jacobaea ); sine loco, s.d., s.c. ( Z) . Sine loco: 1783–1789, Feijó V - V -2 ( 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. 4. – First herbarium sheet from K containing the syntypes of Campanula jacobaea C. Sm. ex Webb. A. Darwin’s specimen [K000865902] chosen in 1986 as isolectotype of C. jacobaea by Porter, determined here as C. bravensis (Bolle) A. Chev., black circle showing the small footnote cross; A’. Label of the Darwin’s specimen, black circle showing the small footnote cross; B. Forbes’ specimen [K001134406] determined here as C. fransinea Gardère, white circle showing a fragment of Hypodematium crenatum (Hypodematiaceae); C. Specimen attributed to a Forbes’ collect [K000865901] determined here as C. fransinea, white circle nº 1 showing a small fragment of Hypodematium crenatum, white circle nº 2 showing a fragment of Diplotaxis gracilis (Brassicaceae); D. Specimen attributed to a Forbes’ collect [K001134405] determined here as C. fransinea. [© Royal Botanic Gardens, Kew]