Nolletia chrysocomoides

Herman, Paul P. J., 2013, Cypsela morphology in the genus Nolletia (Asteraceae, Astereae) and a revision of the genus, Phytotaxa 122 (1), pp. 1-44 : 32-34

11. Nolletia chrysocomoides (Desfontaines 1799: 269, t. 232) Cassini ex Lessing (1832: 187); Cosson & Kralik

(1857: 180); Willkomm & Lange (1870: 35); Battandier & Trabut (1888: 424; 1902: 172); Bonnet & Barratte

(1896: 206); Durand & Barratte (1910: 121); Tutin (1976: 120); Ozenda (1977: 423); Dobignard & Chatelain

(2011: 327).

Basionym: Conyza chrysocomoides Desf. Type:— TUNISIA. Kerwan [Kairouan], Desfontaines s.n. (holotype P-Desf., e!) .

= Conyza pulicarioides Cosson & Durand ined. (manuscript name)

= Nolletia arenosa O. Hoffmann (1893: 76) ; Merxmüller (1967: 119); Wild (1975: 19); Gibbs Russell et al. (1984: 125; 1987: 215); Herman (1993: 714; 2003: 261; 2006: 232); Retief & Herman (1997: 327); Craven (1999: 162); Klopper et al. (2006: 142). syn. nov. Type:— NAMIBIA. Zwischen Ausis und Khuias (Huib Plateau), (QDS: 2616CB/CD Aus), March 1885, Schenck 209 (lectotype Z!, designated here).

= Chrysocoma dinteri Muschler (1911: 104) . Type :— NAMIBIA. Anabriver, (QDS: 2717BA Chamaites), 5 September 1897, Dinter 1190 (holotype Z!).

Usually ascending to erect, virgate, rarely caespitose, perennial herb or suffrutex, but also small woody shrubs, up to (0.1–)0.2–0.4(–0.5) m high, mostly divided at the base into a few initially simple stems, only branching upwards into the inflorescences, rarely consisting of a main stem with side branches, these branching into inflorescences; basal part with pale, yellowish brown or reddish bark, herbaceous and green upwards, appressed pubescent, densely to scattered leafy. Leaves alternate, scattered to fairly densely set, sessile, linear, lower leaves mostly (10–)15–20(–30) × (0.5–)1.0–1.5(–2.5) mm but sometimes up to 35–40 mm long, decreasing in size upwards; apex bluntly acute; margins entire; appressed pubescent on both surfaces. Capitula heterogamous, disciform, 8–10(–12) mm in diameter, usually arranged in few-headed, loose corymbs, rarely solitary at ends of branches. Involucre campanulate. Involucral bracts imbricate, in 3 or 4 rows, green but sometimes tinged purplish, persistent and recurved in old inflorescences, very variable in shape; outer row linear or narrowly linear-triangular to narrowly ovate or narrowly obovate, 1.0–2.5 × 0.2–1.0 mm, apex acute, appressed pubescent; second row narrowly ovate to narrowly obovate or narrowly oblong to narrowly elliptic, 1.8–3.6 × 0.4–1.0 mm, apex acute to acuminate, with very narrowly membranous margin, appressed pubescent; third row narrowly ovate to narrowly obovate or narrowly elliptic, 2.2–4.5 × 0.5–1.0 mm, apex acute to acuminate, with narrowly membranous, faintly fimbriate margin, appressed pubescent; inner row linear to narrowly elliptic or narrowly obovate, 3–5 × 0.4–1.0 mm, apex acute to acuminate, with membranous, faintly fimbriate margin, central portion sparsely appressed pubescent. Receptacle epaleate, foveolate. Outer female florets ± 13–50, in 1 row, fertile, filiform; tube 1–2 mm long, shorter than style furcation, with numerous glandular hairs, apex with single lobe or toothed, in North African material one lobe more obvious, with glandular hairs; corolla yellow. Style 1.0– 2.5 mm long, bifurcate; style branches elliptic, 0.5–1.0 mm long, apex rounded; stigmatic areas marginal, confluent at apex, sometimes with short penicillate, apical appendage. Pappus and cypsela as in disc florets. Disc florets ± 40–90, regular, bisexual, fertile; tubular below, tube 2.0–4.0 mm long, widening slightly after about 1 mm but still tubular and then widening slightly towards lobes, with glandular hairs around first widening; lobes 5, ± 0.5 mm long, also with some glandular hairs; corolla yellow but sometimes purplish on upper part of tube and lower part of lobes. Anthers 1.2–2.0 mm long; with narrowly ovate apical appendages and slightly calcarate at base, ecaudate; filament collars with thickened cell walls. Style 1.5–3.5 mm long, bifurcate; style branches linear, 0.5–1.0 mm long; stigmatic areas marginal, not confluent at apex, with deltoid-penicillate apical appendages. Cypsela honey-coloured, shiny, with faint marginal line, narrowly obovoid, laterally compressed, often slightly skew to one side, 1.2–2.5 × 0.5–1.0 mm ( Fig. 1J ); surface covered with circular epicarpic cells each with short twin hair ( Figs. 2E , 3C ), apices of twin hairs acute but subequal in length. Pappus of caducous, barbellate bristles, 2 mm long in female florets, 3 mm long in disc florets. Flowering time: in southern Africa throughout the year, but with peaks in April–May, August and December; in North Africa from January to August but with a peak from March–June.

Distribution and habitat:— In southern Africa: Namibia, Free State, Northern Cape; in North Africa: Mauritania, Morocco, Algeria, Tunisia and Libya and in Europe: Spain ( Fig. 13 ), growing in sand under desert or semi-desert conditions. According to Wild (1975) this species also occurs in Angola but Figueiredo & Beentje (2008) did not list any Nolletia taxa for that country. Wild (1975) also recorded this species for Botswana, but a duplicate of the specimen he quoted (Wild 5157), housed in PRE, was identified as the new species N. annetjieae . The red list status of this species in South Africa is LC (Least Concern) ( Raimondo et al. 2009).

Discussion:—A very variable species as far as growth form and involucral bracts are concerned. I could not find any reason to keep the northern African and southern African specimens separate, although there is some degree of variation in the growth form (the North African specimens tend to be more dwarf shrubs), the degree of hairiness (often more densely hairy in North African specimens), the slightly longer lobe of the female florets in the North African specimens and the short penicillate, apical appendages more often present on the apices of the style branches of the female florets in the North African specimens. Abnormal inflorescences were observed in some specimens from Morocco, Algeria and Tunisia (North Africa) (Mairé s.n. (P), Cosson s.n. (P), Cosson et al. (P) and Chevallier s.n. (P)). The individual florets are borne on elongated pedicels or ovaries, crowned with abnormal pappus hairs, so that the individual florets rise above the involucral bracts. Some inflorescences appear spicately arranged: the capitula arranged one above the other.

Typification:— Hoffmann (1893), in his original description of N. arenosa , listed the following specimens: Schinz, with no number(-s), collected at Aus, Byzondermeid [Besondermaid] and Schenck 209, collected between Ausis and Khuias (Huib-Plateau). Merxmüller (1967) listed the following specimens as Syntypes: Namibia, Aus, Schinz 699 and Schinz 700; Byzondermeid, Schinz 701; zwischen Ausis und Khuias, Schenck 209. The following specimens received from Z were studied: Schenck 209 collected March 1885 zwischen Ausis und Khuias (between Ausis and Khuias); Schinz 699, collected at Aus, 20 January. On the label the year was altered and it is unclear whether it is 1875 or 1895. According to Glen & Germishuizen (2010) the date should be 1885. Schinz 700 has no indication of an exact locality except for the region: Gross- Namaland. The date of collection is given as 5 December but again it is unclear from the label whether it was in 1874 or 1894. According to Glen & Germishuizen (2010) the date should be 1884. Of the two Schinz 701 specimens, the label on one sheet indicated only a year (1885), no month, and that it was collected at Byzondermeid [Besondermaid]. The label on the second sheet indicated that it was collected at the same locality in January, in the year 1885 according to Glen & Germishuizen (2010). On the JSTOR website (2011) Schinz 707 is listed as the type specimen but the label on the scanned specimen clearly shows 701. As it is unclear whether Hoffmann (1893) saw all the Schinz specimens, I decided to choose the Schenck specimen as lectotype, in order to eliminate all uncertainties.

In Muschler’s (1911) description of Chrysocoma dinteri , he stated that the plant flowered and had fruit in December 1897, but the label on the Dinter s pecimen received from Z (Dinter 1190), collected at Anabriver, clearly shows 5 September 1897.

Figures

FIGURE 1. Cypselae of Nolletia species as seen under the stereomicroscope: (A) N. gariepina (Warren 78, PRE); (B) N. tenuifolia (Giess 10863, PRE); (C) N. ciliaris (Jacot Guillarmod 377, PRE); (D) N. jeanettae (Bester 8534, PRE); (E) N. welmaniae (Winter 7649, PRE); (F) N. rarifolia (Van Hoepen 1825, PRE); (G) N. ruderalis (Compton 27385, PRE); (H) N. vanhoepeniae (Koekemoer 3899, PRE); (I) N. zambesica (Maguire 1693, PRE); (J) N. chrysocomoides (Merxmüller & Giess 3579, PRE); (K) N. annetjieae (Van Rooyen 3714, PRE). Photographs by P.P.J. Herman.

FIGURE 2. Scanning electron micrographs of cypsela features in Nolletia: (A) obovoid cypsela of N. gariepina (Dinter 3577, PRE); (B) oblong epicarpic cells arranged in regular rows and twin hairs on the cypsela of N. tenuifolia (Dinter 7889, PRE); (C) oblong epicarpic cells arranged in regular rows and twin hairs with acute apices on the cypsela of N. gariepina (Zietsman & Zietsman 794, PRE); (D) oblong epicarpic cells arranged in regular rows and twin hairs with broadened apices in cypsela of N. annemarieae (Moss & Jacobsen K180, PRE); (E) obovoid cypsela in the dried state of N. chrysocomoides showing the ‘pocked’ surface (Germishuizen 7763, PRE); (F) circular epicarpic cells each with a twin hair on the cypsela of N. vanhoepeniae in the dried state (De Beer B5, PRE); (G) circular epicarpic cells each with a twin hair, in N. vanhoepeniae in the rehydrated state (De Beer B5, PRE); (H) paired circular to elliptic epicarpic cells with one twin hair between each pair, scattered over the cypsela surface in N. annetjieae in the dried state (Leistner 1581, PRE). Photographs by P.P.J. Herman.

FIGURE 3. Light micrographs of cypsela surfaces of: (A) N. jeanettae illustrating the oblong epicarpic cells arranged in parallel rows after rehydration (Bester 8494, PRE); and (B) N. ruderalis (Vander Schijff 2041, PRE), higher magnification; (C) N. chrysocomoides showing the circular epicarpic cells after rehydration (Wiss 2553, PRE); (D) N. zambesica (Merxmüller 1966, PRE) showing the twin hairs arising from the apical part of the circular epicarpic cells after rehydration; (E) N. annetjieae after rehydration showing the paired circular to elliptic epicarpic cells scattered over the surface, each pair with one twin hair between them (Koekemoer 3901, PRE); (F) N. annetjieae showing the twin hairs arising from two (paired) circular epicarpic cells (Van Rooyen 3714, PRE). Scanning electron micrographs illustrating the different outer female florets of Nolletia species: (G) filiform female floret in N. ciliaris (Koekemoer 1062, PRE); (H) short ray in the outer female floret of N. gariepina (Jensen 208, PRE). Photographs by P.P.J. Herman.

FIGURE 13. Known geographical distribution of N. chrysocomoides in Africa.