The new species Caesia sabulosa Boatwr. and J.C.Manning from deep sands along the West Coast of South Africa and sandy flats in the Cederberg and Bokkeveld Escarpment is described. It is distinguished by its extensively branched rhizome resulting in a robust, clump-forming habit, and unique ‘palisade’ root system of closely packed, hard, vertical roots; mostly larger flowers; erect fruiting pedicels; and details of the seed testa sculpturing

Boatwright, James

Manning, J C

Cape Town University OpenUCT

Available online at www.sciencedirect.comSouth African Journal of Botany 76 (2010) 524–529www.elsevier.com/locate/sajbCaesia sabulosa (Hemerocallidaceae), a new species from the Greater CapeRegion of South AfricaJ.S. Boatwright a,b,⁎, J.C. Manning aa Compton Herbarium, South African National Biodiversity Institute, Private Bag X7, Claremont 7735, Cape Town, South Africab Department of Botany, University of Cape Town, Private Bag, Rondebosch 7700, Cape Town, South AfricaReceived 26 January 2010; received in revised form 29 March 2010; accepted 8 April 2010AbstractThe new species Caesia sabulosa Boatwr. and J.C.Manning from deep sands along the West Coast of South Africa and sandy flats in theCederberg and Bokkeveld Escarpment is described. It is distinguished by its extensively branched rhizome resulting in a robust, clump-forminghabit, and unique ‘palisade’ root system of closely packed, hard, vertical roots; mostly larger flowers; erect fruiting pedicels; and details of the seedtesta sculpturing.© 2010 SAAB. Published by Elsevier B.V. All rights reserved.Keywords: Caesia; Greater Cape Region; Hemerocallidaceae; New species; Scanning electron microscopy; Seed testa; Taxonomy1. IntroductionCaesia R.Br. comprises ±11 species of tufted, grass-likeplants with delicate, short-lived, white, blue or purple flowers inwhich the perianth twists spirally when they fade (Henderson,1987; Obermeyer, 1973). Some of the species have verycolourful, horizontally barred filaments with alternate yellow,white and purple transverse bands in various patterns (Fig. 1).The main centre of diversity for the genus is in Australia, whereeight species are found, seven of them endemic (Henderson,1987). One species occurs in Madagascar and an additional twospecies are found in South Africa, predominantly in the WesternCape (Obermeyer, 1973). The two South African species werelast studied by Obermeyer (1973) who transferred Nanolirioncapense (H.Bolus) Benth. and Hook. to the genus Caesia [asCaesia capensis (H.Bolus.) Oberm.] and also placed severalspecies into the synonymy of Caesia contorta (L.f.) Dur. andSchinz, based on studies of herbarium specimens, concludingthat these represent different forms of one variable species.However, further field studies are crucial to understanding the⁎ Corresponding author. Compton Herbarium, South African NationalBiodiversity Institute, Private Bag X7, Claremont 7735, Cape Town, South Africa.E-mail address: S.Boatwright@sanbi.org.za (J.S. Boatwright).0254-6299/$ - see front matter © 2010 SAAB. Published by Elsevier B.V. All righdoi:10.1016/j.sajb.2010.04.007variation in the C. contorta complex to ascertain whether itindeed represents a single species.The familial placement of the genus has been somewhatvolatile with various authors assigning it to different families, i.e.Asphodelaceae subfamily Anthericoideae (Dahlgren and Clif-ford, 1982), Anthericaceae (Dahlgren et al., 1985), Phormiaceae(Chase et al., 1996), Johnsoniaceae (Clifford and Conran, 1998)and Hemerocallidaceae (Chase et al., 2000; Fay et al., 2000).Recent molecular data place Caesia in the “hemerocallid clade”,which includes members of the families Hemerocallidaceae,Johnsoniaceae and Phormiaceae and which is sister to Asphode-laceae and Xanthorrhoeaceae (Devey et al., 2006). The mostrecent classification system of the Angiosperm Phylogeny Group(APG III, 2009), proposes a greatly expanded concept of thepreviously monotypic family Xanthorrhoeaceae to include boththe Asphodelaceae and the “hemerocallid clade” but we followthe more conservative classification of APG II (2003), whichretains Hemerocallidaceae in an expanded circumscription toinclude Johnsoniaceae and Phormiaceae rather than including itwithin a broad concept of Xanthorrhoeaceae.Here we describe a new species ofCaesia that was previouslyrecognized informally as Caesia sp. 1 in Goldblatt and Manning(2000), raising to the 12 number of species in the genus. Thiswork forms part of a broader systematic and taxonomic study ofts reserved.Fig. 1. Habit and flowers of Caesia sabulosa (a–d) and C. contorta (e–f). Note the transverse bands on the filaments and styles (a–c, f). (For interpretation of thereferences to colour in this figure legend, the reader is referred to the web version of this article.)525J.S. Boatwright, J.C. Manning / South African Journal of Botany 76 (2010) 524–529the South African species and their relationship to theirAustralian counterparts, including field work to unravel thevariation within the species and their circumscription.2. Materials and methodsMorphological data on the new species were gained throughfield studies, as well as examination of herbarium material fromBOL, K, NBG and PRE (abbreviations according to Holmgrenet al., 1990). Seed surface sculpturing of the three South AfricanCaesia species was examined with an LEO S440 fullyanalytical Scanning Electron Microscope (SEM) after beingcoated with gold. This was done at the Electron MicroscopeUnit, University of Cape Town. Voucher information of thespecies studied is listed in Table 1. Drawings were done using astereoscope (WILD M4A) with a camera lucida attachment.3. Species treatmentCaesia sabulosa Boatwr. and J.C.Manning sp. nov., Caesiaecontortae (L.f.) Dur. and Schinz foliis longis distichisconduplicatis, panicula divaricate ramosa et capsulis tripartibusTable 1Voucher data of Caesia species examined for seed testa sculpturing.Species LocalityC. capensis Western Cape: Groot Winterhoek MounC. sabulosa Western Cape: Bokbaai farm, south-wesC. contorta Western Cape: Nature's Valley ReserveC. contorta Western Cape: Jonker's Hoekglobosis ad obovoideas similis, sed rhizomate extensi horizon-tali cum systemate radicum verticali, habitu robusto formanticaespites, pedicellis erectis fructiferis differt.Type: Western Cape, Bokbaai Farm, south-west of Darling[3318 AC], 1 January 1993, Goldblatt & Manning 9496 (NBG,holo.; K, MO, iso.).Rhizomatous perennial to 0.6 m, grass-like and growing inclumps, forming extensive patches up to 2 m diam. Rhizomewoody, extensively branched; roots stiff and tapering, extend-ing vertically downwards, closely packed and palisade-like(horizontal rhizome with many long, vertical roots creating theimpression of a palisade fence). Leaves distichous, erect,conduplicate, linear-lanceolate, 200–600×7–10 mm, acute,glabrous, bases sheathing and pink. Inflorescence a divaricatelybranched panicle; peduncle erect, terete, 3–5 mm diam. at base;bracts ovate, 2–4×2–3 mm, membranous, white or pink;pedicels short, buds subsessile, ±3–5 mm long at anthesis,ultimately ±5–8 mm long in fruit. Flowers 1–2 in each axil,nodding, pale pink or mauve to blue with purplish midribs, openall day and fading in the afternoon, apparently unscented,caducous with perianth twisting spirally when faded; tepalsrecurved, connate in basal ±1 mm, outer elliptical, mucronateVoucher specimentains Marloth 1643 (PRE)t of Darling Goldblatt & Manning 9496 (NBG)Compton 15333 (NBG)Lubke et al. 66 (PRE)526 J.S. Boatwright, J.C. Manning / South African Journal of Botany 76 (2010) 524–529with conspicuous papillose apical tuft, 10–12×2.0–2.5 mm, in-ner narrowly obovate, emarginate, 10–12×2.5–3.0 mm. Stamens6, erect but often with sigmoid bend near base; filaments filiform,retrorsely scabridulous, basal two-thirds yellow and upper thirdwhite separated by dark purple band or upper third purple withwhite tip or yellow with purple bands, inserted at top of perianthtube, outer ±8 mm long, inner ±10 mm long; anthers yellow,1.0–1.5 mm long. Ovary globose, ±1.0–1.5 mm long, green,Fig. 2. Morphology of Caesia sabulosa (a–i) and C. contorta (j–o). (a) whole plant;(e2) seed in lateral view; (f) gynoecium; (g1) outer tepal; (g2) inner tepal; (h) ripe dehstamen; (l2) inner stamen; (m) tufted node on the peduncle; (n) capsule; (o) whole(c) Boatwright & Manning 258 (NBG); (e, i) Goldblatt & Manning 9496 (NBG); (j–l(a, c, m, o) 1 cm; (b, d–l, n) 1 mm.with 2 or 3 ovules per locule; style straight, filiform, scabridulous,basal two-thirds yellow and upper third white separated by darkpurple band or upper third purple with white tip or yellowwith purple bands, ±10 mm long. Capsule turbinate, tripartite,3–5×3–5 mm, abscising when ripe and tardily dehiscent, palebrown. Seeds globose to subreniform, ±2.0–2.5×2×2 mm, black,surface striate-verrucose; strophiole white, fleshy.Flowering time:November–January (Figs. 1a–c; 2).(b) flower; (c) extensive horizontal rhizome; (d) bract; (e1) seed in ventral view;isced capsule; (i) capsule on upright pedicel; (j) flower; (k) gynoecium; (l1) outerplant. Voucher specimens: (a–b, d, f–h) Boatwright & Manning 362 (NBG);, n) Boatwright & Manning 257 (NBG); (m, o)Manning s.n. (NBG). Scale bars:527J.S. Boatwright, J.C. Manning / South African Journal of Botany 76 (2010) 524–5294. Diagnostic charactersCaesia sabulosa resembles the widespread C. contorta in itslong, distichous, conduplicate leaves, divaricately branchedpanicle, and deciduous, tripartite, turbinate capsules in whichmostly one or sometimes two locules ripen, but the two speciesdiffer considerably in habit and root structure, flower colour insome forms, and details of the fruit and seeds. The rhizome in C.sabulosa is extensively branched, with the result that the speciesdevelops a distinctive, clumped growth habit (Fig. 1d), with thelarger clumps reaching up to 2 m in diameter. The species growsin deep, sometimes coastal sands often in association with restioidtaxa, and the root system is well-adapted to the loose substrate,producing numerous hard, tapering roots that are closely packedalong the rhizomes and that grow vertically downwards into thesoil, forming a dense, palisade-like structure serving to anchor theplants securely. The pink to deep blue flowers are borne in stifflyerect panicles that project above the foliage (Fig. 1d), and anthesistakes place during themorning, with the flowers fading during theafternoon. The fruiting pedicels remain erect, and the capsules areshed as soon as they ripen, collecting around the base of theplant. The rhizome in C. contorta is far less extensive, with theplants typically solitary (Fig. 1e), and the roots are wiry andspreading. The pale blue to purple flowers are borne in sprawlingFig. 3. Scanning electron micrographs of the seeds surfaces of Caesia capensis (a–b1643 (PRE); (c–d) Goldblatt & Manning 9496 (NBG); (e–f) Compton 15333 (NBGor trailing panicles that develop characteristic tufts of leavesfrom the lower leaf axils (Fig. 1e), and anthesis takes placeonly in the early afternoon, with the flowers fading in the earlyevening. The fruiting pedicels are recurved, with the capsulesnodding.The remaining South African species, C. capensis is a smallcaespitose and mat-forming plant less than 80 mm high withsubumbellate inflorescences or solitary flowers with smoothfilaments.Surface sculpturing of the seeds is taxonomically informa-tive in the three South African species and has also been usedas a key character in the Australian species (Henderson, 1987;Fig. 3). The seeds of the three South African species are blackwith fleshy white strophioles, reniform in C. capensis andC. contorta but globose in C. sabulosa. The fleshy strophioles(generally referred to as elaiosomes) attract ants which carry theseeds to their nest thus distributing the seeds away from themother plant and providing protection from predation or unfa-vourable conditions (myrmecochory; Lengyel et al., 2010).Surface sculpturing in all three species is striate-verrucose,with a mix of large and small tubercles. In C. sabulosa andC. capensis the large tubercles are surrounded by valleys ofmuch smaller tubercles, with the larger tubercles arranged inirregular clusters in C. capensis and in short, narrow rows in), C. sabulosa (c–d) and C. contorta (e–f). Voucher specimens: (a–b) Marloth). Scale bars: (a, c) 20 µm; (b, d, and f) 100 µm; (e) 10 µm.528 J.S. Boatwright, J.C. Manning / South African Journal of Botany 76 (2010) 524–529C. sabulosa (Fig. 3a–d). In C. contorta the tubercles in thevalleys are diagnostically only somewhat smaller than the largetubercles, which are distributed randomly (Fig. 3e–f). InC. capensis and C. contorta the tubercles are conical, ter-minating in a small conspicuous point, while in C. sabulosa thetubercles are spherical and smooth.5. Distribution and habitatCaesia sabulosa grows in deep sands along the WesternCape coast, from Silverstroomstrand to Redelinghuys (inAtlantis Sand Fynbos, Hopefield Sand Fynbos and LeipoldvilleSand Fynbos plant communities; Rebelo et al., 2006) and as farFig. 4. Known geographical distnorth as Kotzesrus in Namaqualand (in Namaqualand Strand-veld; Rebelo et al., 2006), and inland along the Cederberg andBokkebeld Mountains (in Cederberg Sandstone Fynbos andBokkeveld Sandstone Fynbos; Rebelo et al., 2006; Fig. 4).A similar distribution pattern is evident in another psammo-philous species, Babiana ringens (L.) Ker Gawl. (Goldblatt andManning, 2007). Typical populations of C. sabulosa along theWest Coast south of Redelinghuys have distinctive, pinkflowers but populations from the Bokkeveld Escarpment andCederberg have a blue or purple perianth. They show thecharacteristic rather rigid panicles with large flowers and erectcapsules of the coastal forms and we include them here pendingfurther field work to understand this floral variation.ribution of Caesia sabulosa.529J.S. Boatwright, J.C. Manning / South African Journal of Botany 76 (2010) 524–529C. contorta has a widespread, mainly more inland distribu-tion from the Cederberg and Clanwilliam east as far as Port St.Johns, usually in stony clay or gravelly soils, while C. capensisis a high altitude species restricted to mountain peaks aroundWorcester (Du Toit's Peak, Groot Winterhoek Mountains, HexRiver Mountains, Matroosberg, Wemmershoek Mountains) andon the Seweweekspoort Mountains (Goldblatt and Manning,2000).5.1. Additional specimens examined- 3017 (Hondeklipbaai): 4.8 mls [7.72 km] north–north-westof Kotzerus (–DD), 11 January 1963, Acocks 23400 (BOL,PRE).- 3119 (Calvinia): Nieuwoudtville, Oorlogskloof NatureReserve, M21, 31°30′26″S 19°7′1″E (–CA), 7 November2000, Pretorius 636 (NBG); near the top of Botterkloof Pass(–CD), 8 November 1961, Barker 9633 (NBG 3 sheets).- 3218 (Clanwilliam): 1 km south ofRedelinghuys, 32°29.196′S18°32.045E (–BC), 6 December 2006, Helme 4026 (NBG);5 km south-west of Redelinghuys, mid south slopes ofMierberg, west of Oloff Berghspas, 32°30′52.4″S 18°30′26.8″E (–DA), 20 September 2007, Helme 4853 (NBG).- 3219 (Wuppertal): Heuningvlei (–AA), 6 January 1999,Manning s.n. (NBG, 2 sheets); North Cederberg, PathHeuningvlei to Krakadoupas (–AA), 28 December 1983,Taylor 10863 (NBG); Cederberg, Middelberg Plateau (–AC),December 1939, Esterhuysen 2460 (BOL 2 sheets, NBG 2sheets), 14 December 1941, Esterhuysen 7200 (BOL,NBG); lower eastern slopes below Krakadou near Cederplantation (–AC), 28 December 2004, Gwynn-Evans 2166(BOL); Cederberg Wilderness Area, Groot Koupoort (–AC),19 January 1977, Haynes 1280 (NBG, PRE); Cederberg,Kleine Valey (–AC), 9 November 1956, Taylor 1843 (NBG 2sheets); Matjiesrivier Nature Reserve, 32°29′90″S 19°27′50″E(–AD), 21 November 1999, Low 5841 (NBG, 2 sheets).- 3318 (Cape Town): ca. 6 km south-east of Hopefield onKoperfontein 347 near powerlines, 33°06′48.5″S 18°22′02.0″E (–AB), 26 August 2008, Helme 5604 (NBG);Darling district, Klawervlei Farm (–AD), 15 December1988, Manning 1023 (NBG); just after turn-off to Silwer-stroomstrand on the R27 to Velddrif (–CB), 2 April 2009,Boatwright & Manning 258 (NBG), 26 November 2009,Boatwright & Manning 362; roadside on R27 betweenBokbaai turn-off and Rondeberg (–CB), 18 November 1995,Goldblatt & Manning 10437 (NBG).AcknowledgementsThis research forms part of a Post-Doctoral Fellowship at theCompton Herbarium (South African National BiodiversityInstitute) and the University of Cape Town. The Parker family(Elandsberg Nature Reserve) generously provided funding forthe Fellowship. Dr Gerrit Koorsen (University of Johannesburg)translated the diagnosis into Latin. The curators and staff of Kand PRE are thanked for making their specimens available onloan. Miranda Waldron (Electron Microscope Unit, Universityof Cape Town) kindly assisted with the SEM.ReferencesAPG II, 2003. An update of the angiosperm phylogeny group classification forthe orders and families of flowering plants: APG II. Botanical Journal of theLinnean Society 141, 399–436.APG III, 2009. An update of the angiosperm phylogeny group classification forthe orders and families of flowering plants: APG III. Botanical Journal of theLinnean Society 161, 105–121.Chase, M.W., Rudall, P.J., Conran, J.G., 1996. New circumscriptions and a newfamily of asparagoid lilies: genera formerly included in Anthericaceae. 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(Ed.), Flora of Australia.Canberra, Australian Government Publishing Service, pp. 281–288.Holmgren, P.K., Holmgren, N.H., Barnett, L.C., 1990. Index Herbariorum 1: theHerbaria of the World, 8th ed. Regnum Vegetabile, 120.Lengyel, S., Gove, A.D., Latimer, A.M., Majer, J.D., Dunn, R.R., 2010.Convergent evolution of seed dispersal by ants, and phylogeny andbiogeography in flowering plants: a global survey. Perspectives in PlantEcology, Evolution and Systematics 12, 43–55.Obermeyer, A.A., 1973. Caesia and Nanolirion. Bothalia 11, 122–123.Rebelo, A.G., Boucher, C., Helme, N., Mucina, L., Rutherford, M.C., 2006.Fynbos biome. In: Mucina, L., Rutherford, M.C. (Eds.), Vegetation of SouthAfrica, Lesotho and Swaziland. Strelitzia, 19. South African NationalBiodiversity Institute, Pretoria, pp. 137–139.Edited by B-E Van Wyk

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Caesia sabulosa (Hemerocallidaceae), a new species from the Greater Cape
Region of South Africa

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Caesia sabulosa (Hemerocallidaceae), a new species from the Greater Cape Region of South Africa

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