Campanula jacobaea C. Sm. ex Webb

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 : 16-20

1. Campanula jacobaea C. Sm. ex Webb in Hook., Icon. Pl. 8: tab. 762. 1848 ( Fig. 1 , 2 , 3A , 5C ).

Lectotypus (erroneously designated by PORTER, 1986: 85; corrected and designated here): CABO VERDE: sine loco, “sp. r. to be figured, Cap Verd”, s.d. [IV.1822], Forbes s.n. ( G [ G 00426961 ] image!) .

Sub-frutex 15 – 40 cm tall, highly woody in lower part; floriferous stems branched, decumbent to pendulous 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 0.2–1 mm long. Leaves: rosette leaves elliptic to narrowly elliptic, (1.5–)3–4(–5) × (0.5–)1–1.5(–2.5) cm, base cuneiform to slightly attenuate, apex ± obtuse to acute; cauline leaves narrowly ovate to elliptic, (1.5–)2.5–3.5(–5.5) × (0.5–) 0.8–1.5(–2.5) cm, base attenuate sometimes asymmetric, apex ± obtuse to acute; margin weakly revolute, crenulate; adaxial side light green in vivo, weakly or densely covered with hispidulous to hispid indument, consisting of trichomes 0.2–0.5 mm long; abaxial side light green 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 erect rarely pendulous, pedicel 0.5–1.5(– 2) 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 triangular, 10–13 × 4–6 mm, erect, margin weakly revolute; appendages ovate, reflexed, 1.5–2 mm long; calyx-lobes, lobe edges, appendages and median main vein covered with an indument hispidulous to hispid consisting of trichomes 0.3–0.65 mm long. Corolla campanulate with inflexion point in the upper third, purplish-blue sometimes mauve with veins distinctly marked; base wide round c. 8 mm large; tube, 23–30 mm long, gradually widening and reach the maximum diameter of 14 –18 mm in the upper third; throat slightly constricted then widening up to 20 –26 mm at the mouth; lobes spreading to obliquely erect, 2 –4 × 9–12 mm, apex apiculate; primary external veins hispidulous. 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, 15–18 mm long, included in the corolla, stigma trifid and papillose.

Etymology. – The specific epithet “ jacobaea ”, from the Latin Jacobus, refers to Santiago or “Saint James”; the island was given the name because it was discovered on Saint James Day. Jacobaea was initially chosen by Smith in 1816 to name the bellflowers from Santiago , which he wanted to describe as new to the Cabo Verde flora.

Vernacular name. – GOMES (1994) note “Velho-Teso” but in Brava this name is also use for Spermacoce verticillata L. ( Rubiaceae ) (BARBOSA, 1961; DINIZ et al., 2002; MARTINS, 2002) and Campanula bravensis (BOLLE, 1861) .

Distribution and habitat. – Campanula jacobaea , now circumscribed to Santiago , is a montane chasmophyte that can be found from 600 m to 1100 m in humid rupicolous areas: on cliffs frequently covered in dense fog, with Limonium lobinii N. Kilian & Leyens (Plumbaginaceae) , Polycarpaea gayi Webb (Caryophyllaceae) , Kickxia elegans (G. Forst.) D.A. Sutton (Scrophulariaceae) , sometimes Hypodematium crenatum (Dryopteridaceae) and abundant lichens, or on seeping rock faces and near to springs (chupadeiros) in the depths of valleys, near dense ferns, mainly Adiantum capillus-veneris L. ( Pteridaceae ), and sometimes Christella dentata (Forssk.) Brownsey & Jermy (Thelypteridaceae) . Campanula jacobaea is confined to the main mountains: Serra do Pico da Antónia and Serra da Malagueta, but also Monte Afonso and Monte Tagarrinho, the last two localities being chorological novelties.

Notes. – WEBB (1848: tab. 762) based the description of C. jacobaea on collections from different geographical origin and different collectors; only the names of the collectors were indicated. The following year in his Spicilegia Gorgonea WEBB (1849: 148) added information such as localities, dates and sometimes collection numbers. Eight syntypes have been located. Original material is extant in the Hooker Herbarium (now K) and mounted on two sheets ( Fig. 4 , 5 ) with duplicates in CGE and L and a further syntype has been located in G.

On the first sheet of original material in K ( Fig. 4 ), four specimens are mounted of which only those on the bottom half display the names of the collectors ( Fig. 4A–B ). On the bottom right, the Darwin’s specimen [K000865902] collected in Santiago represents C. bravensis ( Fig. 4A ). A small footnote cross leads to a label ( Fig. 4 A’) with a printed reference “ from J.S. Henslow ” and “ Campanula ” in Hooker f.’s handwriting and “ C. Darwin’s Ms. No. 279 (γ.) ” and “ 166 ” in an unidentified handwriting – probably numbering by Hooker f. or Henslow ( PORTER, 1986). Just above Darwin’s specimen on the left is the Forbes’ specimen ( Fig. 4B ) from São Nicolau (“ n. 35 ” according WEBB, 1849; [K001134406], identified here as C. fransinea ) and with the locality and collector handwritten: “ Isle San Nicol. Forbes ”. The other two specimens on the upper half [K000865901] ( Fig. 4C ) and [K001134405] ( Fig. 4D ), are erroneously annotated “ Teneriffe ” in the Canary Islands by Hooker f. LEYENS & LOBIN (1995) were misled by the placement of the label ( Fig. 4 A’) and wrongly attributed them to Darwin. These two specimens represent C. fransinea and most likely collected in São Nicolau. Among the collectors cited in the protologue, Forbes was the only one who visited São Nicolau from March 27 TH to 31ST, 1822 ( OWEN, 1833); Darwin only visited Santiago ( VALA, 2009) . Therefore, the two specimens [K000865901] ( Fig. 4C ) and [K001134405] ( Fig. 4D ) must have been collected by Forbes. [K000865901] also holds the same fragment of Hypodematium crenatum found on [K001134406] that reinforced the collection locality, i.e. São Nicolau ( Fig. 4B–C ). Furthermore, [K000865901] ( Fig. 4C ) holds a small leaf of Diplotaxis gracilis (Webb) O.E. Schulz (Brassicaceae) , a species endemic to the W São Nicolau.

The second sheet in K ( Fig. 5 ) contains three different specimens with original labels: on the upper half, Forbes s.n. [K001134390] identified by Webb as “ C. daltonii ” (see Introduction) from Santo Antão (“ n. 4 ” according WEBB, 1849) and identified here as C. feijoana ( Fig. 5A ); on the bottom right, Vogel 73 [K001134391] from São Vicente, identified here as C. monteverdensis ( Fig. 5B ); and on the bottom right, Hooker 125 [K001134407] from Santiago , identified here as C. jacobaea ( Fig. 5C ).

The original material located in G is a specimen collected by Forbes [G00426961]. The locality is not mentioned but should be from Santiago where Forbes made a stopover in early April 1822 ( OWEN, 1833). On the original determination label on the bottom left is written “ Campanula Daltonii Webb ” and “ sp. [specimen] r. [retained] to be figured, Cap Verd, Forbes ” both in Webb’s handwriting. This specimen has served for the preparation of the illustrations of C. jacobaea ( WEBB, 1848: tab. 762) and is here identified as C. jacobaea .

Among these syntypes, PORTER (1986) chose Darwin 279 from K [K000865902] ( Fig. 4A ) as the lectotype of C. jacobaea with a duplicate CGE [CGE03087] but Darwin 279 is in serious conflict with the protologue. WEBB (1848) described the species as having a campanulate corolla, three times longer than the calyx-lobes, illustrating the throat as slightly constricted and the roof of the ovary as flat ( WEBB, 1848: tab. 762). However, the flower of Darwin 279 has a tubular corolla that is barely longer than twice the length of the calyx-lobes, and the roof of the ovary appears to be conical.

LEYENS & LOBIN (1995) correctly identified Darwin 279 as C. bravensis . They chose Forbes s.n. [K001134390] ( Fig. 5A ), identified here as C. feijoana , as lectotype by adding a printed label indicating “ Campanula jacobaea Webb – Lectotype – det. T. Leyens & W. Lobin 11.1994”. This lectotypification has never been effectively published. We propose that the lectotype designated by PORTER (1986: 85) is rejected according to Art. 9.19(c) of the ICN ( TURLAND et al., 2018) and superseded by Forbes s.n. in G [G00426961]. We designated here this collection as a new lectotype for C. jacobaea because it represents unambiguous original material of C. jacobaea and this collection has served for Webb’s illustrations ( WEBB, 1848: tab. 762).

Selected material seen. – CABO VERDE. Santiago: Alto da Serra da Malagueta, 1000 m, 8.II.1986, Cardoso de Matos & Matos 6023 ( LISC); Drago [Dragoeira], III.1998, Santos s.n. ( ORT); João Teves , 550 m, 17.XII.1981, Rustan & Brochmann ØHR -1129 ( O); Monte Afonso , 630 m, 11.VIII.2016, Gardère 1301 ( P), 1303 ( P); Monte Chota , c. 1000 m, 16.XII.1995, Leyens CV -95-529 ( FR); Monte Ribão de Cana , 650 m, 22.VII.2013, Gardère 110 ( LISC); Monte Tagarrinho , 850 m, 11.VIII.2016, Gardère 1308 ( P); Orgãos Grandes , 200 m, IV– V.1898, Fea s.n. ( GDOR); ibid. loco, c. 250 m, IV– V.1898, Fea s.n. ( GDOR); Os Orgãos , 31. I.1866, Lowe s.n. ( BM); path from Chã da Figueira to Coruja, c. 700 m, 18.XII.1993, Kilian 2779 & Leyens ( B); Ribeira da Aguada , 25. I.1983, Grandvaux Barbosa et al. 14320 ( CECV, LISC); Ribeira Cantada , 580 m, 26.XI.2014, Gardère 934 ( P); Ribeira Gon Gon , 710 m, 11.VIII.2016, Gardère 1345 ( P); ibid. loco, 700 m, 11.VIII.2016, Gardère 1347 ( CECV, LISC, P); ibid. loco, 700 m, 27.XII.2017, Gardère 1619 ( P); Ribeira da Janela , 770 m, 17. I.1980, Borgen 3390 ( O); Ribeira Longueira , 800 m, 25.XI.2014, Gardère 928 ( P); Ribeira Xáxá , 610 m, 27.XII.2017, Gardère 1620 ( P); Rui Vaz , 15.VII.1934, Chevalier 44593 ( COI, K, P); ibid. loco, 750 m, 27.XI.2014, Gardère 936 ( P); ibid. loco, 750 m, 17.VII.2016, Gardère 1220 ( CECV, LISC, P); ibid. loco, 700 m, 24.III.1968, de Naurois s.n. ( LISC); São Jorge , Ribeira Matom [Mato Moniz], 400 m, 2.IV.1984, Veiga 19 ( LISC); Serra da Malagueta , 1040 m, 31.XII.2013, Aedo 21160 ( MA); ibid. loco, 850 m, 22.XI.1987, Cardoso de Matos 6333 ( LISC, MA); ibid. loco, c. 800 m, 11.X.1988, Diniz et al. 214 ( LISC); ibid. loco, V.1989, Doutre 19 ( ALF); ibid. loco, 800 m, 28.VIII.2013, Gardère 284 ( LISC), 285 ( LISC); ibid. loco, 800 m, 25.VIII.2013, Gardère 287 ( P); ibid. loco, 800 m, 22.XII.2013, Gardère 603 ( LISC); ibid. loco, 750 m, 22.XII.2013, Gardère 604 ( P); ibid. loco, 800 m, 22.XII.2013, Gardère 605 ( CECV); ibid. loco, 840 m, 15.VII.2016, Gardère 1211 ( P); ibid. loco, 850 m, 27.X.2016, Gardère 1428 ( P); ibid. loco, 840 m, 30.X.2016, Gardère 1435 ( P); ibid. loco, 1310 m, 27.XII.2017, Gardère 1618 ( CECV, MARS, P); ibid. loco, 27.XII.1955, Grandvaux Barbosa 6087 ( CECV, COI, LISC); ibid. loco, c. 800 m, 17.XII.1993, Kilian & Leyens 2751 ( B, FR); ibid. loco, 1000 m, 26.X.1986, Mies 383 ( FR); ibid. loco, c. 800 m, 5.II.1994, Leyens CV -94-079 ( FR); ibid. loco, 1.X.1979, Lobin 759 ( FR); ibid. loco, 920–950 m, 15. I.1980, Rustan 790 ( O); ibid. loco, c. 900 m, 1.XII.1996, Schmidt CV /KS -1996-59 ( FR), CV /KS -1996-60 ( FR); ibid. loco, 800 m, 17.XI.1976, Sunding 3659 ( O); Serra do Pico da Antónia , 800–1200 m, 16.IV.1898, Fea s.n. ( GDOR); ibid. loco, 1135 m, 27.XII.2015, Gardère 1198 ( P); sine loco [ Serra do Pico da Antónia ], 11.IV.1816, Smith 39 ( BM); entre a Trindade e o Curralinho, 720 m, 24.XI.1955, Grandvaux Barbosa 5676 ( CECV, LISC); “a most beautiful sp. […] on a peak in the valley of St Domingo at 2000 ft ”, XI.1839, Hooker 125 ( K p.p.: remaining syntype for C. jacobaea ) .

Figures

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

Fig. 2. – Campanula jacobaea C. Sm. ex Webb. A. Habit; B. Rosette-leaves; C. Inflorescence; D. Flower in lateral view; E. Flower in face view. [A–B: Gardère 1620; C: Gardère 1435; D: Gardère 928; E: Gardère 1211] [Photos: M.L. Gardère]

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. 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]