A taxonomic revision of the genus Lagenophora Cass. (Asteraceae: Astereae) in New Guinea Jian Wang & A.R. Bean Summary Wang, J. & Bean, A.R. (2020). A taxonomic revision of the genus Lagenophora Cass. (Asteraceae: Astereae) in New Guinea. Austrobaileya 10(4): 576-582. The genus Lagenophora Cass, is taxonomically revised for New Guinea with two species recognised. One (Lagenophora sporadica Jian Wang ter & A.R.Bean sp. nov.) is endemic to New Guinea and the other (L. sublyrata (Cass.) A.R.Bean & Jian Wang ter) is widespread in Asia, Australia and Malesia. The species are described and illustrated. Notes are provided on the distribution (including a map), habitat and phenology. An identification key is provided. Key Words: Asteraceae; Astereae., Lagenophora sporadica; Lagenophora sublyrata; New Guinea flora; new species; identification key; distribution map Jian Wang & A.R. Bean, Queensland Herbarium, Department of Environment and Science, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: jian.wang@des. qld.gov. au; [email protected]. au Introduction Lagenophora Cass, is a small genus of daisies The present authors concur with Koster in the Tribe Astereae and found mainly in the (1966) that two species are present - one is L. temperate parts of the southern hemisphere. sublyrata (Cass.) A.R.Bean & Jian Wang ter, Twelve species are found in Australia (Wang of which L. lanata is a synonym and the other & Bean 2019), nine species occur in New is L. sporadica Jian Wang ter & A.R.Bean, a Zealand (Breitwieser et al. 2012), three new species closely related to L. stipitata. species in southern South America (Cabrera Materials and methods 1966) and two in New Guinea. The genus also extends to New Caledonia and Indonesia, This revision is based on morphological and as far as Sri Lanka and southern Japan examination of Lagenophora material from (Koidzumi 1928; Wang & Bean 2019). the following herbaria: A, AK, BRI, CANB, GH, HO, L, MEL and NSW. Images of type The genus Lagenophora was first recorded specimens held at FI, G, HAL, K, M, NY, for New Guinea by Mueller (1889), based on P and W have also been examined and are a specimen collected by William MacGregor indicated as i.d.v. (imago digitalis visa). Most from the summit of the Owen Stanley Range, measurements are based on dried material, earlier that same year. Cabrera (1966) cited but the dimensions of florets are based on New Guinea specimens for three species, material reconstituted with boiling water. L. gracilis Steetz, L. lanata A.Cunn. and Dimensions in the descriptions are inclusive, L. stipitata (Labill.) Druce. Koster (1966) viz. 1.0-17 is given as 1—1.7. reduced this to two species, L. stipitata and L. lanata, and stated that L. gracilis and L. lanata could not be maintained as separate species, as the former differs only by the sparser tomentum. Accepted for publication 24 February 2020 Wang & Bean, Lagenophora in New Guinea 577 Taxonomy Key to the New Guinea species of Lagenophora Roots tuberous and fleshy, bunched; rhizomes short; stem very short (leaves in basal rosette); scape hairs appressed to antrorse; achene glands confined to the dorsal side of beak and adjacent upper dorsal edge; achenes usually with 1-3 hairs at base; achene 2.2-2.4 mm long excluding beak. L. sublyrata Roots fibrous and wiry, not bunched, rhizomes spreading; stem usually elongated (leaves alternate along stem), scape hirsute, hairs retrorse to patent; achene papillae extending all along ventral and dorsal edges from beak to carpopodium, and on the basal and distal portions of both faces; achenes without basal hairs; achene 2.9-3.4 mm long excluding beak. L. sporadica 1. Lagenophora sublyrata (Cass.) A.R.Bean (c. 2.5 x longer than wide), sessile or with a & Jian Wang ter, Austrobaileya 10: 435-437 winged petiole-like base to 1 cm long; leaf (2019); Ixauchenussublyratus Cass., Diet. Sci. apex obtuse or rounded; leaf margins toothed, Nat. ed. 2. [F. Cuvier] 56: 176 (1828). Type: crenate to sinuate, with 5-7(-9) teeth, each Australia. [New South Wales.] Port Jackson, tooth 0.2-0.4 mm long; upper leaf surface November-December 1819, C. Gaudichaud green, with 2-7 hairs per mm2, each 0.1-0.3 s.n. (lecto: image of P 00742955^ i.d.v.), fide mm long; lower leaf surface pale green, with Bean & Wang (2017: 168). Epitype: New 3-7 hairs per mm2, each 0.1-0.4 mm long; South Wales. Hornsby, April 1914, W.F. leaf margins with 6-12 hairs per mm2, each Blakely s.n. (NSW 10275), fide Bean & Wang 0.1-0.3 mm long; net veins usually obscure (2017: 168). on dried material on both surfaces. Scapes channelled, 1-6 per tuft, each (2.5-)4-8 cm Ixauchenus lyratus Less., Syn. Gen. Compos. long at anthesis, 4-11 cm long at fruiting 193 (1832), nomen nudum. stage, 0.3-0.4 mm diameter, 0.3-0.5 mm thick Lagenophora billardierei var. media DC., from lowest to upmost section; scape bracts Prodr. [A. P. de Candolle] 5: 307 (1836). 2-4, upper ones c. 0.6 x 0.2 mm, lower ones Type: Nova Hollandia, [in 1823], F.W. Sieber c. 1.4 x 0.4 mm; scape indumentum c. 0.1 mm 505 (syn: G 00454010 i.d.v:, HAL i.d.v:, NY long, antrorse, more or less appressed; 2-6 00180436 i.d.v). hairs per mm at midpoint of scape, slightly denser towards apex. Capitula 2.5-3.5 mm Lagenophora billardierei var. glabrata DC., long, 2.5-5 mm diameter; involucral bracts Prodr. [A. P. de Candolle] 5: 307 (1836). Type: (20-)25 in 3 or 4 rows, lanceolate, oblong to Nouvelle Holland, in 1816, from Lambert’s obovate, apex obtuse, acute, ciliate or with herbarium (syn: G 00454009 i.d.v). fringed margin on distal part, glabrous, outer Lagenophora lanata A.Cunn., Ann. Nat. Hist. bracts 1.2-1.5 x 0.3-0.4 mm, inner bracts 2: 126 (1839). Type: New Zealand. Between 2.2-2.8 x 0.5-0.6 mm. Receptacle convex, the Waitangy and Keri-Keri Rivers, in 1834, 0.6-0.8 mm diameter and 0.5-0.8 mm high. R. Cunningham s.n. (lecto: K 000890104 Ray florets 20-30 in 2-4 rows; tube c. 0.4 mm i.d.v), fide Allen (1961: 606). long, 0.1-0.2 mm wide, glandular hairy; style branches c. 0.4 mm long; ligules 1.8-2 mm Perennial rhizomatous herb; roots fleshy, long, 0.15-0.35 mm wide, with longitudinal bunched, 0.2-1 mm diameter; no stem or short veins obscure, creamy, yellow or purple, apex stem to 10 mm long; leaves and scapes firmly obtuse. Disc florets 10—15, corolla tubular, attached to stem and/or rootstock. Leaves 1.5-1.8 mm long, light yellow, outer surface 4—9(—11), obovate, oblanceolate, elliptical, with sparse glandular hairs; corolla lobes spathulate, 1-2 cm long by 0.4-0.8 cm wide 5, deflate, 0.2-0.3 mm long x 0.3-0.35 mm 578 Austrobaileya 10(4): 576-582 (2020) wide; stamens 5, anthers c. 0.6 mm long; style sandy or rocky flats. branches c. 0.2 mm long; sterile ovary 0.6-0.7 Phenology: Flowers and fruits have been mm long; pappus scales 1 or 2, c. 0.1 mm long. recorded throughout the year except February Achenes compressed, lanceolate, obliquely and March. oblanceolate, 2.2-2.4 x 0.4-0.6 mm excluding beak, light brown to dark brown at maturity; Note: Lagenophora sublyrata is a widespread achene edges more or less thickened; with species with variable leaf shape, indumentum 1-3 eglandular hairs present usually at base of and plant size. New Guinea specimens are achene; achene glands confined to dorsal side usually quite hairy and small in stature, as are of beak and adjacent area of achene; achene specimens from New Zealand, but features beak 0.4-0.5 mm long, with a thickened of the roots, the achene, scape and involucral white annular collar at its apex, 0.15-0.2 mm bracts are consistent with typical plants from diameter. Fig. 1. eastern Australia. Additional specimens examined: Indonesia. Papua. 2. Lagenophora sporadica Jian Wang ter BalimRiver,Dec 1938.Brass II782(BR\,CANB,L);near & A.R.Bean, sp. nov. with affinities to L. Lake Gita, Angi, Arfak Mts, Apr 1940, Kanehira 14115 stipitata (Labill.) Druce and L. adenosa Jian & Hatusima (L); Anggi Gita Lake, Bivouac Noordpool, Jan 1962, Sleumer 14020 & Vink (BRI, CANB, L). Wang ter & A.R.Bean, but differing from Papua New Guinea. Western Highlands Province: L. stipitata by the sinuate to undulate leaf Yobobos grassland area, Laiagam subdistrict, Aug 1960, margins, the two types of scape indumentum, Hoogland 7505 & Schodde (CANB, L). Southern the shorter ligules of the ray florets, and Highlands Province: Between Kendatel and Yombi, c. differing gland and/or papillae distribution 6 miles [10 km] N of Ialibu patrol post, Jul 1961, Pullen 2735 (CANB). Morobe Province: Mannasat, Cromwell on the achene surfaces. It differs from L. Mountains, Huon Peninsula, Jul 1964, Hoogland 9450 adenosa by the smaller sized ligules, and (CANB); ibid, Aug 1964, Hoogland 9614 (BRI, L); the gland distributional pattern on achene Watama, Menyamya subdistrict. May 1968, Streimann NGF35897 & Kairo (L). Central Province: Boridi, surfaces. Typus: Papua New Guinea. Oro Nov 1935, Carr 13441 (CANB, L); Mafulu, Sep 1933, (Northern) Province: Mt Kenive (Nisbet), Brass 5151 (BRI, L); Coastal scarp of Astrolabe Range, Kokoda Subdistrict, 30 July 1974, J.R. Croft SW of Birribi, Aug 1970, Schodde 5637 (L); Coast et al. LAE65116 (holo: BRI [AQ352188]; iso: scarp of Astrolabe Range, Port Moresby subdistrict, A [2 sheets], CANB 559390, L 1815314). Aug 1970, Stevens LAE50342 (BRI, CANB, L). Milne Bay Province: Maneau Peak, Mt Dayman, May 1953, [Lagenophora stipitata, auct. non (Labill). Brass 22255 (CANB, L); E slopes, Goodenough Island, Druce; Roster (1966: 590)]. Oct 1953, Brass 24666 (CANB, L); Mt Wadimana ridge, NE from Mt Simpson, Jul 1969, Pullen 7865 (CANB); Perennial rhizomatous herb; roots and Summit area of Mt Wadimana, eastern Mt Simpson rhizomes fibrous; stem usually elongated Range, Jul 1969, Schodde 5480 (CANB); Mt Suckling Complex, Mayu II, Raba Raba Subdistrict, Jun 1972, (leaves alternate along stem) or occasionally Stevens LAE54980 & Veldkamp (BRI, L); Goropu short (leaves in near basal rosette); leaves Mountains (Mt Suckling), Jun 1972, Veldkamp 5536 & and scapes firmly attached to stem and/or Stevens (L). rootstock. Leaves 5-10, obovate to long- Distribution and habitat: Lagenophora spathulate, 1.5-4 cm long by 1-2.2 cm wide sublyrata is the most widespread species in (1.5-1.8 x longer than wide), with a winged the genus. It has been called by the misapplied petiole-like base 1-4 cm long; leaf apex name L. gracilis or the synonymous name L. obtuse; leaf margins sinuate to undulate, lanata in south Asia (e.g. India, Sri Lanka), usually with 9—11(—17) shallow lobes, each south-east Asia (e.g. Vietnam), China, lobe 0.5-2 mm long; upper leaf surface Malesia (e.g. Java), New Caledonia, Australia grey green; with eglandular hairs 0.3-0.6 and New Zealand. In New Guinea, it has a mm long, 4-10 per mm2; lower leaf surface widespread distribution (Map 1). It mainly pale green; with eglandular hairs 0.2-0.6 inhabits montane grassland near rainforest mm long, 1-6 per mm2; both leaf surfaces margins on sandy or fine gravel deposits in have papillae to 0.01 mm long, more or less wet situations from 670 m to 2750 m above evenly distributed; leaf margins with 6-14 sea level. It has also been recorded on dry eglandular hairs per mm2, each 0.4-0.8 mm Wang & Bean, Lagenophora in New Guinea 579 Fig. 1. Lagenophora sublyrata. A. habit of whole plant with flowering inflorescences >4.5. B. leaf with a section in details x5. C-E. lower, mid and upper-sections of scape x20. F. capitulum with flowers and fruits removed, lateral view x20. G. outer involucral bract x20. H. inner involucral bract x20. I. disc floret x35. J. marginal floret x35. K. achene x35. A from Stevens LAE50342 (BRI), B, F from Brass 5151 (BRI), C-E from Stevens & Veldkamp LAE54980 (BRI), G-J from Hoogland 9614 (BRI), K from Brass 11782 (BRI). Scale bar = 10 mm at xl magnification. Del. N. Crosswell. 580 Austrobaileya 10(4): 576-582 (2020) long; lateral veins obscure on upper leaf vicinity, Upper Camp, Feb 1939, Clemens 9585 (L surfaces, but obvious on lower leaf surfaces. 1815313, L 1815327); ibid, Feb 1939, Clemens 9910A (A, L); Mt Enggom, Sarawaket Range, Feb 1963, van Scapes channelled, 1—3(—4) per tuft, each 2-7 Royen NGF16204 (BRI, CANB, L). Oro (Northern) cm long at anthesis, 6-14 cm long at fruiting Province: W slopes Mt Kenive (Nisbet), Kokoda stage, c. 0.6 mm diameter but expanding to c. Subdistrict, Aug 1974, Croft LAE65155 (BRI, L). Milne 1.4 mm at apex; scape bracts 2-3, upper ones Bay Province: Mt Ganaina, Aug 1962, Cruttwell 1280 c. 4x0.5 mm, lower ones c. 8 x 1.8 mm; scape (K, L); Goropu Mountains (Mt Suckling), Jun 1972, Veldkamp & Stevens 5678 (CANB, L); S Spur of Goe indumentum including eglandular hairs 0.2- Dendeniwa, Mt Suckling complex, Jun 1972, Stevens 0.5 mm long, appressed, patent or retrorse; & Veldkamp LAE54205 (A, BRI, CANB, K, L); Slopes 1-4 hairs per mm2 at midpoint of scape, 4-8 of Goe Dendeniwa, Mt Suckling complex, Raba Raba hairs per mm2 towards apex; and papillate to Subdistrict, Jun 1972, Stevens & Veldkamp LAE54279 (A, BRI, CANB, K, L); E slope of Manurep, Raba Raba c. 0.01 mm long, 5-15 per mm2 at midpoint Subdistrict, Jul 1972, Stevens & Veldkamp LAE54497 of scape, but very densely distributed towards (A, BRI, CANB, K, L). apex. Capitula 4-5(-6) mm long, 7-12 mm diameter; involucral bracts c. 54 in 3 or 4 rows, Distribution and habitat: Lagenophora linear, lanceolate, apex acute to acuminate, sporadica is endemic to Papua New Guinea where it is restricted to high altitudes with fringed margins on distal half, papillate from 2400 to 3800 m (Map 1). It grows in with eglandular hairs occasionally along conifer dominated submontane rainforest midrib on outer surface; outer bracts c. 2.5 x and relictual forest that can be dominated 0.5 mm, inner bracts 2.5-3.5 x 0.3-0.7 mm. by Prunus, Podocarpus or Papuacedrus. It Receptacle convex to hemispherical, 2.8-4.4 is also recorded from the shade of subalpine mm diameter and 1.4-2.5 mm high. Ray florets shrubberies; subalpine grasslands in rocky c. 83, in 3-4 rows; tube 0.5-0.9 mm long, c. places with low grass; and in the shelter of 0.3 mm wide, with papillae; style branches c. forest borders and logs. It can be a common 0.6 mm long; ligules 1.9-2.2 x 0.3-0.4 mm, species even in burnt forest areas. longitudinal veins obscure, white or pink, apex obtuse and bidentate. Disc florets c. 11; Phenology: Both flowers and fruits were corolla tubular, c 1.8 mm long, greenish or recorded in February, March, May to August, yellow, outer surface covered with papillae; and November. corolla lobes (4-)5, deflate, 0.3-0.4 x c. 0.3 mm; stamens (4—)5, filaments 1.1-1.5 mm Typification: It is indicated on the label of long, anthers 0.6-0.8 mm long; style branches the type specimen of Lagenophora sporadica 0.4-0.7 mm long; sterile ovary 2.2-2.5 mm that duplicates of LAE65116 are also present at BISH, BM, BO, E, K, LAE, M, PNH, SING, long, with a thickened white annular collar at SYD and US, but none of these have been its apex, collar c. 0.3 mm diameter. Achenes seen by the present authors. compressed, oblanceolate, straight or slightly curved, 2.9-3.4 mm long excluding beak, c. Affinities: Lagenophora sporadica is of 1 mm wide, uniformly brown at maturity; similar appearance to L. stipitata, but differs achene edges slightly thickened; achene by the sinuate to undulate leaf margins papillae extending all along ventral and dorsal (obtusely serrate for L. stipitata), the two edges from beak to carpopodium, and on the types of scape indumentum (usually one type basal and distal portions of both faces; achene only for L. stipitata), the ligules of the ray beak 0.5-1.1 mm long, densely surrounded by florets 1.9-2.2 mm long, (2.3-3.3 mm long papillae, and with a white annular collar at its for L. stipitata), and different gland and/or apex, 0.2-0.25 mm diameter. Fig. 2. papillae distributional patterns on the achene Additional selected specimens examined: Papua surfaces. It also resembles L. adenosa, but New Guinea. Eastern Highlands Province: Mt differs from that species by the smaller sized Aniata, Mar 1959, Cruttwell 1057 (K, L); Mt Wilhelm, ligules at 1.9-2.2 x 0.3-0.4 mm (3-3.4 x 0.4- Nov 1960, Borgmann 315 (L). Central Province: Mt 0.6 mm for L. adenosa), and the distributional Albert Edward, May/Jul 1933, Brass 4226 (A, BRI, K, patterns of glands and/or papillae on achene L); NW part of Mt Albert Edward, Jun 1974, Craven 2750 (A, CANB, K, L). Morobe Province: Samanzing surfaces. Wang & Bean, Lagenophora in New Guinea 581 Fig. 2. Lagenophora sporadica. A. habit of whole plant with flowering inflorescences x0.5. B. leaf with a section in details *5. C-E. lower, mid and upper-sections of scape xl5. F. capitulum with flowers and fruits removed, lateral view x5. G. outer involucral bract xl5. H. inner involucral bract xl5. I. disc floret with immature achene x20. J. marginal floret with near mature achene x20. K. achene x20. A, C-K from Croft LAE65116 (BRI), B from Craven 2750 (CANB). Scale bar = 10 mm at xi magnification. Del. N. Crosswell. 582 Austrobaileya 10(4): 576-582 (2020) Etymology: From the Latin sporadicus, Breitwieser, I., Brownsey, P.J., Garnock-Jones, P.J., meaning ‘scattered’ or ‘widely dispersed’. Perrie, L.R. & Wilton A.D. (2012). Phylum Tracheophyta - vascular plants. In D P Gordon This refers to the widely dispersed glands (ed.). New Zealand Inventory of Biodiversity. and/or papillae on the faces of the achenes, in Kingdoms Bacteria, Protozoa, Chromista, comparison with Lagenophora adenosa and Plantae, Fungi. 3: 411-459. University of L. stipitata. Canterbury Press: Christchurch. Acknowledgements Cabrera, A.L. (1966). The genus Lagenophora (Compositae). Blumea 14: 285-308. We acknowledge the Directors of A, AK, De Candolle, A.P (1836). Prodromus systematis CANB, GH, HO, L, MEL andNSW for the loan naturalis regni vegetabilis 5: 307-308. Treuttel of herbarium specimens. Nicole Crosswell is & Wurtz: Paris. thanked for providing the illustrations. Peter Koidzumi, G. (1928). Plantae Novae Amami- de Lange is also thanked for comments on an Ohsimensis nee non Insularum Adjacentium. earlier draft of this paper. 1. Phytogeographical notes on the flora of the Loochoo Archipelago; 2. Description of new References species, pp. 3-4. Kagoshima, Japan. Allen, H.H. (1961). Flora of New Zealand, Volume Koster, J.T. (1966). The Compositae of New Guinea I. 1, Indigenous Tracheophyta. R.E. Owen, Nova Guinea Botany 24: 497-614. Government Printer: Wellington. Mueller, F. (1889). Records of observations on Sir Bean, A.R. & Wang, J. (2017). The identity and William MacGregor’s highland plants from typification of Ixauchenus sublyratus Cass. New Guinea. Transactions of the Royal Society (Asteraceae). Adansonia ser. 3, 39: 1-4. of Victoria 1(2): 1-45. Wang, J. & Bean, A.R. (2019). A taxonomic revision of Lagenophora Cass. (Asteraceae) in Australia. Austrobaileya 10: 405-442. Map 1. Distribution of Lagenophora sublyrata • and L. sporadica A in New Guinea.