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Eight new species of Marasmodes (Asteraceae,Anthemideae) from South Africa
SANTIAGO ORTIZ*
Departamento de Botánica, Facultade de Farmacia, Universidade de Santiago, 15782 Santiago deCompostela, Galiza, Spain
Received 30 October 2006; accepted for publication 19 May 2008
Marasmodes DC. (Asteraceae, Anthemideae) waserected by de Candolle (1837) for two species, M.polycephala DC. and M. oligocephala DC. [for whichthe specific epithet should be feminine followingArticle 62.4 of the International Code of BotanicalNomenclature (ICBN)], first collected by Ecklon inWestern Cape Province (South Africa). Subsequently,only two new species have been described in thegenus, both from the same province: M. dummeriBolus ex Hutch. (Hutchinson, 1917) and M. undulataCompton (Compton, 1946). However, M. dummeri isprobably conspecific with M. polycephala, so that onlythree species should be accepted to date. The firsttwo revisions of the genus, by Harvey (1865) andHutchinson (1917), included M. adenosolen Harv.;however, Nordenstam (1976) subsequently removedthis species from Marasmodes to include it in hisgenus Cymbopappus B.Nord. [C. adenosolen (Harv.)B.Nord.], differentiated from Marasmodes by itsradiate capitula and its cup-shaped pappus or apappus composed of only two scales.
This shrubby genus is characterized with respect tothe other genera of the subtribe Matricariinae by its
ericoid, entire to divided leaves; generally small,discoid, homogamous capitula, these sessile to sub-sessile, solitary or in groups; epaleate receptacle;glandular corolla with a swollen tube and thick vas-cular strands and lobes often reflexed at maturity;five-ribbed, glandular, cypselas with myxogenic cellson the ribs and on the abaxial surface; and pappus of7–14(–18) unequal scales, adaxial scales being longer.As noted by Källersjö (1988) and Bremer &Humphries (1993), the most closely related genus isPentzia Thunb., which, unlike Marasmodes, generallyhas capitula on long peduncles, erect corolla lobes anda coroniform to auriculate pappus, or, rarely [P.dentata Kuntze], a pappus of three to five separatescales or no pappus [P. calva S.Moore and P. tortuosaFenzl ex Harv. & Sond.].
During a stay in 2002 at the Kirstenbosch ResearchCentre in Cape Town (South Africa), the authorstudied several groups of Asteraceae, principally ofthe tribes Mutisieae s.l., Astereae and Anthemidae.One of these was the genus Marasmodes. As a resultof this research, eight collections from the WesternCape Province were finally identified as new species.
MATERIAL AND METHODS
This study was based on a morphological analysis ofspecimens from BM, BOL, G-DC, NBG and K*Corresponding author. E-mail: [email protected]
Botanical Journal of the Linnean Society, 2009, 159, 330–342. With 8 figures
(Holmgren, Holmgren & Barnett, 1990). The materialwas studied with the aid of a light microscope.
DESCRIPTIONMARASMODES TRIFIDA S.ORTIZ SP. NOV.
Type: South Africa, Western Cape Province, CapeDistrict, Cape Town, Groenrivier farm, between theeastern slopes of Dassenberg and Kalabaskraalvillage, on heavy brackish clay in association withSalicornia, 350 ft, 2.v.1980, J. P. Rourke 1672 (holo-type NBG, isotype K).
Description: A small diffuse shrub up to 40 cm high.Stems branched, branches terete to polygonate insection, glabrous, with sparse sessile glands. Leavesalternate, sometimes opposite, sessile, erecto-patent,(4–)6–9(–11) ¥ 0.5–3 mm, linear, entire or oftendivided into (2)3(4) short linear segments in theirdistalmost one-fifth to one-seventh, glabrous, con-spicuously glandular-punctate, with a short apiculusin each division. Capitula discoid, homogamous, soli-tary, rarely in groups of two or three, terminal,sessile. Involucre 4(–5) ¥ 3.5–4.5 mm, campanulate;phyllaries 30–40 per involucre, arranged in 4(–5)rows, lanceolate to narrowly oblong, obtuse, glabrous;outer phyllaries 1–1.5 ¥ 0.5–0.7 mm, without scariousmargins; inner phyllaries 3.5–5 ¥ 0.7–1 mm, withpale or colourless or, more rarely, purple narrow scari-ous margins and tip, this 0.5–1 mm long, with yel-lowish glands at base of tip inconspicuous or absent.Receptacle flat, without pits. Florets c. 15 per capitu-lum. Corolla yellowish, with sessile glands; tube cylin-drical, 1.5–1.7(–2) mm long, broader in apical part;lobes triangular, recurved, c. 0.5 mm long. Anthers1.3–1.5 mm long, shortly exserted, with anther baseobtuse, ecaudate; apical appendage c. 0.2 mm long,lanceolate. Style c. 2 mm long, terete; branchesc. 0.3 mm long, truncate, bristly at tip. Cypselas 1.5–1.7 ¥ 0.4–0.5 mm, oblong, slightly obconical, five-ribbed, with sessile glands between ribs. Pappus of10–15 unequal scarious scales 0.1–1 mm long (Fig. 1).
Other material examined: South Africa, Western CapeProvince, Malmesbury District, Kalabaskraal, v.1915,Bolus s.n. (NBG).
Notes: These collections were identified by Goldblatt& Manning (2000) as M. oligocephala, the type ofwhich has entire and shorter leaves. Marasmodestrifida is distinguishable from the other species of the
genus by its rather long and usually apically trifidleaves and solitary capitula with campanulateinvolucre.
MARASMODES BEYERSIANA S.ORTIZ SP. NOV.Type: South Africa, Western Cape Province, Stellen-bosch District, Faure, on clay, 11.v.1945, L. Bolus23198 (holotype BOL).
Diagnosis: Foliis erectis vel tantum leviter erecto-patentibus saepe integris, capitulis parvis aggre-gatis involucro campanulato praeditis, phyllariisinterioribus apice atropurpureo scarioso praeditisdistinguitur.
Description: A small diffuse shrub up to 15 cm high.Stems branched, branches terete to polygonate insection, glabrous, with sparse sessile glands. Leavesalternate, sessile, erect to slightly erect-patent, (3–)4–7 ¥ (0.8–)1–4 mm, linear, entire or sometimes withtwo linear lateral divisions in their distalmost one-third, glabrous, glandular-punctate, with a short api-culus in each division, without leaf fascicles at baseor these scarce. Capitula discoid, homogamous, ingroups of three to ten, terminal, sessile. Involucre3–4 ¥ 2–2.5(–3) mm, campanulate; phyllaries 16–18per involucre, arranged in three to four rows, lan-ceolate to narrowly oblong, obtuse, glabrous; outerphyllaries 0.7–1.2 ¥ 0.7–1 mm, with narrow scariousmargins; inner phyllaries 2.5–3 ¥ 1–1.5 mm, withdark-purple, narrow scarious margins and tip, this0.5–1 mm long, with yellowish glands at base. Recep-tacle flat, without pits. Florets c. eight per capitulum.Corolla yellowish, with sessile glands; tube (0.5–)1–1.3 ¥ 0.7–0.8 mm long, cylindrical, broader in apicalpart; lobes triangular, recurved, c. 0.5–1 mm long.Anthers 1–1.2 mm long, shortly exserted, with antherbase obtuse, ecaudate, apical appendage c. 0.2 mmlong, lanceolate. Style c. 1.7 mm long, terete,branches c. 0.5 mm long, truncate, bristly at tip.Cypselas 1–1.5 ¥ 0.5 mm, oblong, slightly obconical,five-ribbed, with sessile glands between ribs. Pappusof 8–12 unequal scarious scales 0.1–0.9 mm long(Fig. 2).
Etymology: The species name is in memory of Dr J. P.B. Beyers, who provided me with invaluable help andadvice during my stay at the Kirstenbosch ResearchCentre, and who sadly died a few months later.
Other material examined: South Africa, Western CapeProvince, near Faure Station, in clayey ground,4.iii.1929. A.V. Duthie 2012 (BOL). Western CapeProvince, Simonstown, Harmony Flats between Gor-don’s Bay and Strand, sandy clay flats with very short
open vegetation, 50 ft altitude, 3.vi.1972, E.G.H.Oliver 3744 (K). Western Cape Province, Simonstown,Faure, western flats close to national road, sandy,loamy, open ground, 200 ft altitude, 14.v.1975, E.G.H.Oliver 5914 (NBG).
Notes: This species is characterized by its erect toslightly erect-patent leaves, often entire, and its smallcapitula arranged in groups, with campanulate involu-cre and phyllaries with dark-purple scarious tip. Thisnew taxon is morphologically similar to M. oligo-
Figure 1. Marasmodes trifida S.Ortiz: A, habit; B, leaf; C, capitula; D, phyllaries; E, floret without cypsela; F, cypsela.Based on holotype. J. P. Rourke 1672 (NBG).
Description: A small diffuse shrub up to 30 cm high.Stems branched, branches ± quadrangular, glabrous.Leaves alternate to fasciculate, densely packed alongbranches, especially on sterile shoots (giving planta spiny appearance), rigid, sessile, erecto-patent,2–4(–5) ¥ 1.5–3 mm, pinnately divided into (3–)5–9linear segments, glabrous, sparsely glandular-punctate, each segment of leaf with a short rigidapiculum. Capitula discoid, homogamous, solitary,terminal, sessile. Involucre 4–5.5 ¥ (2.5–)3–3.5 mm,campanulate; phyllaries 30–40 per involucre,arranged in five to six rows, lanceolate to narrowlyoblong, obtuse, glabrous; outer phyllaries 1–1.5 ¥ 0.5–0.7 mm, with narrow scarious margins, light-brown;inner phyllaries 4–5 ¥ 1–1.3 mm, with purple to light-brown scarious margins and tip, this 0.7–1 mm long,with yellowish glands at base of tip inconspicuous orabsent. Receptacle flat to slightly convex, withoutpits. Florets 13–17 per capitulum. Corolla yellowish,with sessile glands; tube 2–2.5 mm long, cylindrical,gradually broader in apical part; lobes triangular,recurved, c. 0.5 mm long. Anthers 1.2–1.4 mm long,shortly exserted, with anther base obtuse, ecaudate;apical appendage c. 0.2 mm long, lanceolate. Style c.2 mm long, terete, branches c. 0.2 mm long, truncate,bristly at tip. Cypselas 1–1.2 ¥ 0.3–0.4 mm, oblong,slightly obconical, nearly terete to obscurely five-ribbed, with sessile glands. Pappus of 12–14 unequalscarious scales, 0.2–0.5 mm long (Fig. 3).
Other material examined: South Africa, Western CapeProvince, Worcester District, Worcester, flats south ofVoëlvlei, Elandsberg Private Natural Reserve, Malm-esbury shale, heavily grazed fynbos, 80 m altitude,13.vii.1978, C. Boucher 3842 (NBG).
Notes: This species is distinguishable from the otherspecies of the genus by its divided, rigid leaves, whichare densely packed on the branches, and by its soli-tary capitula with campanulate involucres.
MARASMODES REFLEXA S.ORTIZ SP. NOV.Type: South Africa, Western Cape Province, Bredas-dorp District, Napier, Renosterbos community,15.vi.1946[?], P. G. Jordaan 51 (holotype NBG,isotype NBG). [There are two sheets of this collection,with year of collection given as 1946 for the holotypeand 1964 for the isotype.]
Diagnosis: Foliis reflexis, divisionibus lateralibusduabus longis in parte apicali saepe praeditis, capitu-lis solitariis involucro campanulato vel magisminusve obconico praeditis, phyllariis interioribus
glandulas sessiles luteolas prope apicem scariosumferentibus, flosculis in dimidio distali abrupte latiori-bus distinguitur.
Description: Small shrub. Stems branched, bra-nches ± quadrangular in section, glabrous, with abun-dant sessile glands, principally towards apex offloriferous branches. Leaves alternate or sometimesfasciculate, sessile, often reflexed, 2–7 ¥ 0.5–5 mm,linear, entire or often with two linear lateral divisionsup to 2.5 mm long in their distalmost one-quarter totwo-thirds, glabrous, glandular-punctate, with a shortapiculus in each division. Capitula discoid, homoga-mous, solitary, terminal on foliose branches, sessile.Involucre 3–4 ¥ 3–3.5 mm, campanulate to slightlyobconical; phyllaries 25–40 per involucre, arranged in(4–)5–6 rows, lanceolate to narrowly oblong, obtuse,glabrous; outer phyllaries 0.7–1 ¥ 0.5–0.7 mm, with-out scarious margins; inner phyllaries 3.5–4(–4.5)¥ 0.6–0.7(–1) mm, with pale to colourless narrow scari-ous margins and tip, this 0.5–1 mm long, with aconspicuous group of sessile yellowish glands at thebase of the tip. Receptacle flat, without pits. Florets c.8–12 per capitulum. Corolla yellowish, with sessileglands; tube 1.8–2 ¥ 0.6–0.9 mm long, cylindrical,abruptly broader in distal half; lobes triangular,recurved, c. 0.5 mm long. Anthers c. 1 mm long, shortlyexserted, with anther base obtuse, ecaudate, apicalappendage c. 0.3 mm long, lanceolate. Style c. 2 mmlong, terete, branches c. 0.3 mm long, truncate, bristlyat tip. Cypselas (0.8–)1–1.2 ¥ 0.3–0.5 mm, oblong, notor only slightly obconical, slightly curved, five-ribbed,with sessile glands between ribs. Pappus of 14–18unequal scarious scales, 0.2–0.8 mm long (Fig. 4).
Notes: This species is characterized by its reflexedleaves, often with two long lateral divisions in theapical part, solitary capitula with campanulate toslightly obconical involucres with inner phyllariesshowing a conspicuous group of sessile yellowishglands at the base of the scarious tip and floretsabruptly broader in the distal half. From a morpho-logical point of view, the most similar species seems tobe M. oubinae S.Ortiz sp. nov.
MARASMODES FASCICULATA S.ORTIZ SP. NOV.Type: South Africa, Western Cape Province, Malmes-bury, east side of Cape Town to Malmesbury road,near the turning to road to Malmesbury, 20.v.1956,N. S. Pillans 10114 (holotype BOL, isotype K).
Description: Small diffuse shrub. Stems branched,branches polygonate to terete in section, glabrous,with sessile glands. Leaves alternate, sessile, gener-ally patent, sometimes somewhat recurved, 2.5–5(–8) ¥ (0.5–)0.7–3 mm, linear, entire or sometimes withtwo linear lateral divisions in distalmost one-third,more rarely with four lateral divisions, glabrous,glandular-punctate, with a short apiculus in each
division, often with an axillary fascicle of short linearleaves. Capitula discoid, homogamous, in groups of(2–)4–20, sessile. Involucre (2.5–)3–4(–4.5) ¥ (2–)2.3–3(–3.5) mm, campanulate; phyllaries 20–28 perinvolucre, arranged in (4–)5(–6) rows, lanceolate tonarrowly oblong, obtuse, glabrous; outer phyllaries0.5–1 ¥ 0.5–1 mm, without scarious margins; innerphyllaries 2.5–3.5 ¥ 0.7–1.2 mm, with pale to colour-less narrow scarious margins and tip, this 0.5–1 mmlong, with yellowish glands at base of tip inconspicu-ous or absent. Receptacle flat, without pits, with
Figure 3. Marasmodes spinosa S.Ortiz: A, habit; B, leaf; C, capitulum; D, phyllaries; E, floret. Based on holotype.M. E. de Villiers 18 (NBG).
sessile yellowish glands. Florets c. 8–16 per capitu-lum. Corolla yellowish, with sessile glands; tube 1.2–1.5 ¥ 0.5–0.7 mm long, cylindrical, abruptly broaderin apical part; lobes triangular, recurved, c. 0.5 mmlong. Anthers c. 1 mm long, shortly exserted, withanther base obtuse, ecaudate; apical appendage 0.2–0.3 mm long, lanceolate. Style 1.7–2 mm long, terete,branches c. 0.5 mm long, truncate, bristly at tip.Cypselas 1–1.3 ¥ 0.4–0.5 mm, oblong, slightly obconi-
cal, five-ribbed, with sessile glands between ribs.Pappus of 12–14 unequal scarious scales, 0.05–0.7 mm long (Fig. 5).
Other material examined: South Africa, WesternCape Province, Cape District, near to the east sideon the main road Cape Town to Malmesbury, about15 m out; abundant in a small area on clay,7.vi.1942, N.S. Pillans 9778 (BOL, K). Western Cape
Figure 4. Marasmodes reflexa S.Ortiz: A, habit; B, leaf; C, capitulum; D, phyllaries; E, floret. Based on holotype. P. G.Jordaan 51 (NBG).
Province, Cape, Schoongezicht, 22.iv.1992, A. DainesAD932 (BOL). Western Cape Province, MalmesburyDistrict, along Malmesbury road, c. 15–20 milesbeyond the turn-off from the N, in zone betweensand and clay, 19.iii.1971, E. Esterhuysen 32584(BOL, K).
Notes: This species is distinguishable from the otherspecies of the genus by its patent, often recurved,leaves, generally with an axillary fascicle of shortleaves, and by its small capitula arranged in groupswith campanulate involucres and phyllaries withlight-coloured scarious tip.
Figure 5. Marasmodes fasciculata S.Ortiz: A, habit; B, leaf with axillary fascicle; C, capitulum; D, phyllaries; E, floret.Based on holotype. N. S. Pillans 10114 (BOL).
Description: A small diffuse shrub. Stems branched,branches quadrangular in section, glabrous, withsparse sessile glands. Leaves alternate, sessile, patent,slightly recurved, 2–3(–5) ¥ 0.5–2(–5) mm, linear,entire or often with 2(–4) linear lateral divisions1–1.5(–2) mm long in distal one-half to one-quarter,glabrous, glandular-punctate, with a short apiculus ineach division, sometimes with a fascicle of short axil-lary, linear leaves. Capitula discoid, homogamous,solitary, rarely in groups of two or three, terminal onshort foliose shoots, sessile. Involucre 5–6 ¥ 4–5 mm,obconical; phyllaries 30–40 per involucre, arranged infive to seven rows, lanceolate to narrowly oblong,obtuse, glabrous; outer phyllaries 0.7–1 ¥ 0.5–0.7 mm,without scarious margins; inner phyllaries 3.5–4(–5) ¥ 0.8–1(–1.2) mm, with pale to colourless or some-times purple narrow scarious margins and tip, this0.5–1 mm long, with yellowish glands at base of tipinconspicuous or absent. Receptacle flat to slightlyconcave, without pits. Florets c. 10–15 per capitulum.Corolla yellowish, with sessile glands; tube 1.5–2 ¥ 0.5–0.7 mm long, cylindrical, gradually broadeningin its apical half; lobes triangular, recurved, 0.3–0.5 mm long. Anthers 1.2–1.3 mm long, shortlyexserted, with anther base obtuse, ecaudate, apicalappendage c. 0.3 mm long, lanceolate. Style 2–2.5 mmlong, terete, branches 0.3(–0.5) mm long, truncate,bristly at tip. Cypselas 1.3–1.5(–1.7) ¥ 0.5–0.7 mm,oblong, slightly obconical, five-ribbed, with sessileglands between ribs. Pappus of 12–14 unequal scariousscales (0.5–)0.7–1(–1.5) mm long (Fig. 6).
Etymology: The species name is in reference to mycolleague and friend, Juan Rodríguez Oubiña.
Other material examined: South Africa, Western CapeProvince, Paarl Division, low flats at east base of thePaadeberg, vi.1937, N. S. Pillans 8367 (BOL).
Notes: This species is characterized by its small,patent leaves, entire or divided, and its small capitulawith conspicuously obconical involucres. It can beeasily distinguished from M. macrocephala, whichalso has a conspicuously obconical involucre (see nextsection), by its much smaller leaves, often divided,and its much smaller capitula.
The specimen Western Cape Province, MalmesburyDistrict, Selicfontein, prope Hopefield, ix.1889, Bach-mann 5969 (K) is incomplete and damaged, and thereare some doubts about its correct identification.
MARASMODES MACROCEPHALA S.ORTIZ SP. NOV.Type: South Africa, in convalle Hex River, prope DeDoorns, iv.1907, H. Bolus 13126 (holotype BOL, iso-types BM, K).
Description: Shrub. Stems branched, branchespolygonate to nearly terete in section, glabrous, withsparse sessile glands. Leaves alternate, sessile,erecto-patent, (4–)10–18 ¥ 0.5–0.8 mm, linear, entire,glabrous, glandular-punctate, with a short apiculus.Capitula discoid, homogamous, solitary, terminal, onfoliose shoots 0.5–5 cm long, sessile. Involucre6–7 ¥ (5–)6(–7) mm, obconical; phyllaries 30–35 perinvolucre, arranged in five to six rows, lanceolate tonarrowly oblong, obtuse, glabrous; outer phyllaries(1.5–)2–2.5 ¥ 1(–1.5) mm, without or with narrowscarious margins; inner phyllaries 4–5 ¥ 1.3–1.5 mmwith pale to colourless scarious margins and tip, this1.5(–2) mm long, without yellowish glands at base.Receptacle flat, with sessile, yellowish glands, andwith pits absent or faintly marked. Florets c. 18 percapitulum. Corolla yellowish, with sessile glands;tube 2–2.5(–3) ¥ 0.7–1 mm long, cylindrical, graduallybroadening in its apical part; lobes triangular,recurved, 0.5–0.7 mm long. Anthers 1.5–1.8 mm long,shortly exserted, with anther base obtuse, ecaudate;apical appendage c. 0.4 mm long, lanceolate. Style c.2.5–3 mm long, terete, branches c. 0.5 mm long, trun-cate, bristly at tip. Cypselas (1–)1.5(–1.7) ¥ 0.3–0.7 mm, oblong, slightly obconical, five-ribbed, withsessile glands between ribs. Pappus of seven to tenunequal scarious scales, 0.2–1 mm long, somewhatacute, slightly joined at base (Fig. 7).
Notes: This species is characterized by its copiouslong, linear leaves, always entire, and its compara-tively large, solitary capitula with obconical involu-cres of 30–35 phyllaries arranged in five to six rowsand with a light-coloured scarious apex. These char-acters allow it to be readily separated from M. defo-liata, probably the closest species.
Description: Shrub. Stems branched, branches rami-fied, polygonate to nearly terete in section, glabrous.Leaves deciduous, alternate, rarely opposite, sessile,erecto-patent, 9–23 ¥ 0.7–1 mm, linear, entire, gla-brous, blackish, glandular-punctate, with a shortapiculus. Capitula discoid, homogamous, solitary,rarely in groups of three, terminal or sometimesaxillary, sessile. Involucre (5–)6(–7) ¥ (5–)6–7(–8) mm, broadly campanulate; 20–35(–40) phyllariesper involucre, arranged in four to five rows, lan-ceolate to narrowly oblong, obtuse, glabrous;outer phyllaries 1–1.5(–2) ¥ 0.5–1 mm, with scarious
Figure 6. Marasmodes oubinae S.Ortiz: A, habit; B, leaf; C, capitulum; D, phyllaries; E, floret. Based on holotype.T. M. Salter 4408 (NBG).
margins absent or very narrow; inner phyllar-ies 4–5(–5.5) ¥ 1.2–1.7(–2) mm, with conspicuouslypurple narrow scarious margins and tip, this1–2 mm long, without yellowish glands at base.Receptacle flat, with pits absent or faintly marked.Florets c. 20 per capitulum. Corolla yellowish, withsessile glands; tube (1.8–)2 ¥ 0.7–1(–1.2) mm long,cylindrical, gradually broadening in its apical part;lobes triangular, recurved, c. 0.5 mm long. Anthers1.2–1.3 mm long, shortly exserted, with anther baseobtuse, ecaudate; apical appendage c. 0.3 mm long,
lanceolate. Style c. 2–2.5(–3) mm long, terete,branches c. 0.5 mm long, truncate, bristly at tip.Cypselas 1.5–2 ¥ 0.5–0.7(–1) mm, oblong, slightlyobconical, five-ribbed, with sessile glands betweenribs. Pappus of 10–12 unequal scarious scales, 0.2–1mm long, rounded at apex (Fig. 8).
Other material examined: South Africa, betweenMalmesbury and Darling, 4.iv.1932, T. M. Salter 2077(BM, BOL, K). In the Kew herbarium, there are twoadditional sheets with incomplete and damaged speci-
Figure 7. Marasmodes macrocephala S.Ortiz: A, habit; B, leaf; C, capitulum; D, phyllaries; E, floret. Based onholotype. H. Bolus 13126 (BOL).
mens and incomplete label information, collected byZeyher; these probably belong to M. defoliata.
Notes: This species is characterized by its long, linearcaducous leaves, always entire, and its comparativelylarge, solitary capitula with campanulate involucresof 20–35(–40) phyllaries arranged in four to five rowsand with a purple scarious apex. These characters
allow it to be readily separated from M. macro-cephala, probably the closest species.
ACKNOWLEDGEMENTS
My thanks go to Luis G. Orellana for the illustrations,to M. Laínz and Philip Oswald for the Latin diagnosesand nomenclatural advice, to G. Norman for revisionof the English text, to the staff of the Kirstenbosch
Figure 8. Marasmodes defoliata S.Ortiz: A, habit; B, leaf; C, capitulum; D, phyllaries; E, floret. Based on holotype.E. Esterhuysen 34925 (BOL).
Research Centre, Cape Town and to the keepers of theherbaria mentioned for the loan of study material. Ialso thank an anonymous reviewer for comments thatimproved the final version.
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