Research Article |
Corresponding author: Guillaume Lannuzel ( lannuzel@iac.nc ) Corresponding author: Jian Wang ( jian.wang@des.qld.gov.au ) Academic editor: Peter de Lange
© 2021 Guillaume Lannuzel, Gildas Gâteblé, Anthony R. Bean, Jian Wang.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Lannuzel G, Gâteblé G, Bean AR, Wang J (2021) Lagenophora (Asteraceae, Astereae) in New Caledonia. PhytoKeys 177: 125-138. https://doi.org/10.3897/phytokeys.177.63116
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The genus Lagenophora Cass. is taxonomically revised for New Caledonia with two species recognised. Lagenophora sinuosa Lannuzel, Gâteblé & Jian Wang ter, sp. nov. is endemic to New Caledonia and the other, L. sublyrata (Cass.) A.R.Bean & Jian Wang ter occurs there and in many other countries from the region. Both are fully described and illustrated. An identification key is provided, as are notes on the distribution (including maps), habitat, phenology and conservation status. The generic placement of the new species is also discussed.
Asteroideae, Compositae, identification key, Lagenophora sinuosa, Lagenophora sublyrata, Lagenophorinae, New Caledonia flora, new species
Lagenophora Cass. belongs to subfamily Asteroideae (Cass.) Lindl., tribe Astereae Cass. and subtribe Lagenophorinae G.L.Nesom and is found mainly in the southern hemisphere (
The genus was apparently first recorded in a publication for New Caledonia by
This revision is based on morphological examination of Lagenophora material from the following herbaria: BRI, CANB, L, MEL, NOU, NSW and P (acronyms following
1 | Trichomes on scape patent, 0.2–0.5 mm long. Cypsela with beak 0.1–0.3 mm long; oval in cross section, longitudinal ribs on both surfaces, base with no trichome | L. sinuosa |
– | Trichomes on scape appressed, c. 0.1 mm long. Cypsela with a beak 0.4–0.5 mm long; laterally flattened, smooth on both surfaces, base usually with 1–3 trichomes more or less caducous | L. sublyrata |
New Caledonia. North Prov.: Pouembout, Ouaté, 500 m, 21°9'52.43"S, 165°7'1.5"E, 26 Mar 2019, G. Gâteblé, S. Liede-Schumann, U. Meve &D. Fleurot 1091 (Holotype: P!, isotype: NOU107482!).
Lagenophora sinuosa Lannuzel, Gâteblé & Jian Wang ter differs from all other species in the genus with its usually deeply lobed leaf margins and ribbed cypsela surface. It resembles L. queenslandica Jian Wang ter & A.R.Bean with its very short cypsela beak.
Perennial rhizomatous herb; roots and rhizomes fibrous; stem usually absent (leaves in basal rosette); leaves and scapes firmly attached to stem and/or rootstock. Leaves 5–10, oblanceolate to spathulate, 1.5–4 cm long by 0.5–1.5 cm wide (1.5–3 × longer than wide), winged petiole-like base 0.1–3 cm long; leaf apex obtuse to rounded; leaf margins more or less deeply lobate, sinuate to crenate, usually with 4–10 deep lobes, each lobe 2–5 mm deep; upper leaf surface greyish green; with 4–10 trichomes per mm2, each 0.3–1 mm long; lower leaf surface pale green, with 1–6 trichomes per mm2, each 0.2–0.6 mm long; leaf margins with 6–14 trichomes per mm, each 0.7–1 mm long; secondary veins obscure on upper leaf surface, but sometimes obvious on lower leaf surface. Scapes channelled or not, 1–4 per tuft, 3–25 cm long, c. 0.5 mm diameter; bracts 2–7, upper ones c. 4 × 0.5 mm, lower ones c. 5.0 × 1.5 mm; trichomes 0.2–0.5 mm long, patent or retrorse, erect; 10–15 trichomes per mm2 at midpoint of scape, 15–30 trichomes per mm2 towards apex; papillae to c. 0.01 mm long, 5–15 per mm2 at midpoint of scape, but very densely distributed towards apex. Capitula 3–6 mm long, 2–5 mm diameter; involucral bracts c. 25 in 2–3 rows, with trichomes c. 0.3 mm occasionally along midrib on outer surface, linear to lanceolate, apex purple, acute to acuminate, with fringed margins on distal half, outer bracts 1.3–2.1 × 0.5 mm, inner bracts 2–3.5 × 0.3–0.7 mm. Receptacle convex, 2.8–4.2 mm diameter and 1.4–2.5 mm high. Ray florets 20–30 in 1 or 2 rows; tube 0.1–1 mm long, c. 0.3 mm wide, glandular pilose; style branches c. 0.5 mm long; ligules 1.9–4.2 × 0.4–0.9 mm, with longitudinal veins obscure, white or very occasionally pink, apex obtuse and bidentate. Disc florets 10–30, corolla greenish or yellow, tubular, 1.5–2 mm long, outer surface covered with papillae; corolla lobes 4–5, deltate, 0.4–0.6 × 0.3 mm; stamens 4–5, anthers 0.6–0.8 mm long; style branches 0.4–0.9 mm long; sterile ovary 2.2–2.5 mm long; pappus scales absent. Cypselae oval in cross section, oblanceolate, 2.3–3.2 × 0.5–0.8 mm excluding beak, uniformly brown at maturity; surfaces with 2–4 longitudinal ribs on each side; no trichome at the base; beak 0.1–0.3 mm long, densely covered by glands, without a thickened white annular collar at its apex.
New Caledonia. North Prov.: Haute rivière de Voh, 250 m, 12 Mar 1951, Guillaumin & Baumann-Bodenheim 12112 (P03292561image!†); Haute rivière de Voh, 250 m, 12 Mar 1951, Guillaumin & Baumann-Bodenheim 12154 (P03292559image!†); Vallée de la Moindah (branche nord), 150 m, 3 Oct 1965, MacKee 13515 (P04427664image!†); Mt Paéoua, contrefort nord-est, 600–900 m, 4 Jul 1967, MacKee 17004 (P04427667image!†); Pouembout, 30 m, 26 May 1971, MacKee 23675 (NOU054762!, P04234038image!); Pouembout, 30 m, 16 Feb 1972, MacKee 25003 (P03276832image!); Pouembout, 30 m, 16 Apr 1981, MacKee 38956 (P04427671image!†); Poya, forêt de Nékoro, 2 m, 26 May 1983, MacKee 41502 (CANB718870.1image!, NOU054761!, P04427669image!†); Poya, forêt de Nékoro, 2 m, 16 Aug 1984, MacKee 42136 (NOU072073!, P04295155image!†); Poya nord, entre le creek Hervouet et son affluent nord au dessus de la RT1, 40–50 m, 14 Oct 1998, Veillon 8135 (NOU072074!); Cap Devert, 1861–1867, Vieillard 816 (Deplanche? 109) (P03292531image!†). South Prov.: Mont Dore, 800 ft., 3 Apr 1914, Compton 675 (BM013867015image !); Tontouta, 1 Nov 1924, Däniker 414 (P03292449image!†); Prony, 2 m, 22°19'31.5"S, 166°49'34.44"E, 17 Mar 2020, Gâteblé, Lannuzel & Ititiaty 1184 (NOU107485!, P!); Cap N’Doua, Kô Mwâ Nirê, 20 m, 22°22'33.78"S, 166°56'28.1"E, 17 Mar 2020, Gâteblé, Lannuzel & Ititiaty1187 (BRI!, K!, MPU!, NOU107484!, P!); Prony, Îlot Casy, 5 m, 22°21'20.84"S, 166°50'46.77"E, 14 Aug 2020, Gâteblé 1224 (BRI!, K!, MPU!, NOU107486!, P!); Prony, Sep 1910, Godefroy s.n. (P03292558image!, left plant); Ouipouin, 21°41'18.96"S, 165°59'25.08"E, 14 Dec 2018, Laudereau 1236 (NOU091404!); Sommet de la Table Unio (1000 m), 21 Sep 1965, MacKee 13414 (P04427666image!†); Plateau sommital de la Table Unio, 1000 m, 14 Nov 1970, MacKee 22908 (NOU072308!, NSW935348, P04427668image!†); Mont Nakada, 1000 m, 21°37'50"S, 166°3'35"E, 18 Apr 2001, Munzinger & McPherson 814 (P00217314image!); Ouaménie, 1 Jul 2006, Munzinger et al. 3488 (NOU013890!); Poya, sud-est de Mépouiri, 10 m, 9 Jul 1991, Veillon 7390 (NOU072081!); Colline à M’bée, 1855–1860, Vieillard 816 (P03292447image!†).
Haute Tipindje, Contrefort Sud du massif Oua Tilou, 400 m, 28 Jun 1970, MacKee 22128 (P03292503image!†). We were not able to conclusively identify this specimen as the image apparently bears no cypsela. Regarding morphology and ecology, it may be a L. sinuosa but further field work is needed to acquire certitude on that locality.
Lagenophora sinuosa is an endemic species to New Caledonia. It grows only on mainland Grande Terre from the southern tip to Kaala-Gomen as the northernmost locality. As with L. sublyrata, the species has a relatively broad altitudinal distribution ranging from 2 to 1000 m above sea level (Fig.
Both flowers and fruits were recorded from February through November from herbarium specimens.
The specific epithet sinuosa refers to the sinuate leaf margins, by which the species differs from L. sublyrata. Some immature plants or populations bear crenate leaves in natural conditions, but showed ability to produce deeply lobed leaves in greenhouse conditions.
The species is largely distributed on the mainland, though often neglected by collectors, perhaps because it is an inconspicuous herb and maybe considered as an “exotic” or weedy species. The number of localities where it occurs may then be underestimated through herbarium records. The ecology of the species being rainforests floors and maquis on both ultramafic and non-ultramafic substrates at low to medium altitudes tend to consider the invasive introduced Rusa deer (Rusa timorensis) as the major threat both by grazing and by trampling. The fire threat is another issue, especially for open maquis populations. Nevertheless, with over ten localities (sensu
Lagenophora sinuosa has variable leaf shapes. Although its leaf margins are usually deeply lobed, there are two populations growing in the understory of coastal Araucaria forests showing crenate leaf margins. These two populations were considered at one stage as a different species. However, further examinations of their fertile aspects showed there were no significant differences between these two and all other populations. Moreover, some individuals of two of the populations that were cultivated in greenhouse conditions with fertilizers can occasionally produce lobed leaves. Therefore, these two populations have been included in L. sinuosa.
Generic placement of the new species is subject to debate, and consideration was given to making it a new monotypic genus. However, as pointed out by
Comparison of the diagnostic characters of Lagenophora sinuosa and related genera with data modified from
Taxon | Marginal florets | Disc florets | Cypselae | Pappus |
---|---|---|---|---|
Lagenophora sinuosa | 1–2–seriate with ligule | 4–or 5–lobed, functionally male | ribbed, with glandular beak | absent |
Lagenophora (other species) | 2–5–seriate with ligule | 4– or 5–lobed, functionally male | smooth, with glandular beak | absent (except scales on disc florets of L. sublyrata) |
Brachyscome | uniseriate with ligule | 4– or 5–lobed, fertile or sterile | ribbed or not, no glandular beak | present |
Keysseria | 2– or 3–seriate with ligule | 4–lobed, sterile | smooth, with glandular beak | absent |
Myriactis | 2–5(–10) –seriate with ligule | 4– or 5–lobed, fertile or sterile | smooth, with glandular beak | absent |
Pytinicarpa | uniseriate with ligule | 5–lobed, sterile | ribbed, no glandular beak | present (barbellate) |
Solenogyne | 3– or 4–seriate without ligule | 4–lobed, functionally male | smooth, no glandular beak | absent |
Ixauchenus sublyratus
Cass. in F.Cuvier, Dict. Sci. Nat. 2nd ed. 56: 176. 1828. Type: New South Wales. Port Jackson, Nov.-Dec. 1819, C. Gaudichaud (Lectotype: P 00742955, image only extant, fide
Ixauchenus lyratus Less., Syn. Gen. Compos. 193. 1832, nomen nudum.
Lagenophora billardierei var. media DC., Prodr. 5: 307. 1836. Type: Nova Hollandia, 1823, F.W. Sieber 505 (Syntypes: G 00454010!, HAL, NY 00180436!).
Lagenophora billardierei var. glabrata DC., Prodr. 5: 307. 1836. Type: Nova Hollandia, without locality, 1816, from Lambert’s herbarium (Syntype: G 00454009!).
Lagenophora lanata
A.Cunn., Ann. Nat. Hist. 2: 126. 1838. Type: New Zealand. Between the Waitangy and Keri-Keri Rivers, 1834, R. Cunningham 437 (Lectotype: K000890104!, fide
Australia. New South Wales: Port Jackson, November–December 1819, C. Gaudichaud (Lectotype: P 00742955†, image only extant; designated by
Perennial rhizomatous herb ; roots fleshy, 0.2–1 mm diameter; no obvious stem; leaves and scapes firmly attached to rootstock. Leaves 4–9(–11), obovate, oblanceolate, elliptical or spathulate, 1–6 cm long by 0.6–1.6 cm wide (c. 2.5 ×longer than wide), sessile or with a winged petiole-like base to 1 cm long; leaf apex obtuse; leaf margins toothed, crenate to sinuate, with 2–10 teeth, each tooth c. 1 mm long; upper leaf surface green, with 2–7 trichomes per mm2, each 0.3–0.6 mm long; lower leaf surface pale green, with 3–7 trichomes per mm2, each 0.1–0.4 mm long; leaf margins with 6–12 trichomes per mm, each 0.1–0.4 mm long; net veins usually obscure on dried material on both surfaces. Scapes channelled or not, 1–6 per tuft, 5–11(–22) cm long, 0.5–0.8 mm diameter; bracts 2–4, upper ones c. 1.1 × 0.2 mm, lower ones 1.1–2.9 × 0.4 mm; trichomes c. 0.1 mm long, antrorse, more or less appressed; 10–30 trichomes per mm2 at midpoint of scape, slightly denser towards apex. Capitula 3.1–3.5 mm long, 1.8–4 mm diameter; involucral bracts 15–20 in 2–3 rows, glabrous, lanceolate, oblong to obovate, apex green or purple, obtuse to acute, ciliate on distal part, outer bracts 1.3–1.9 × 0.6 mm, inner bracts 2.1–2.7 × 0.7–0.8 mm. Receptacle convex, 0.6–0.8 mm diameter and 0.5–0.8 mm high. Ray florets 20–30 in 2 rows; tube c. 0.4 mm long, 0.1–0.2 mm wide, glandular pilose; style branches c. 0.4 mm long; ligules 1.8–2.6 mm × 0.5–0.6 mm, with obscure longitudinal veins, white, creamy, or purple with age, apex obtuse or bidentate. Disc florets 6–11, corolla light yellow, tubular, 1.5–1.8 mm long, outer surface with sparse glandular trichomes; corolla lobes 5, deltate, 0.2–0.3 × 0.4–0.5 mm; stamens 5, anthers c. 0.6 mm long; style branches 0.3–0.6 mm long; sterile ovary 0.6–0.7 mm long; pappus scales 1 or 2, c. 0.1 mm long. Cypselae laterally compressed, lanceolate or obliquely oblanceolate, 2.2–2.4 × 0.4–0.8 mm excluding beak, light brown to dark brown at maturity; cypsela edges more or less thickened, smooth; with 1–3 eglandular more or less caducous trichomes present usually at base of cypsela; cypsela glands confined to dorsal side of beak and adjacent area of cypsela; cypsela beak 0.4–0.5 mm long, with a thickened white annular collar at its apex, 0.15–0.2 mm diameter.
New Caledonia. North Prov.: Mont Mi, 9 Mar 1869, Balansa 1023 (P03292499image!†); Col des Roussettes, 537 m, 15 Sep 1964, Blanchon 963 (NOU072076!); Ouaté, 21°9'19.98"S, 165°9'6.98"E, 15 Apr 2019, Laudereau 1268 (NOU091405!); Diahoué, 20°28'53.11"S, 164°41'33.29"E, 28 Jul 2019, Laudereau 1286 (NOU091406!); Mt Pouitchate between upper Tipindjé and upper Kamendoua above Ateu, 1000 m, 29 Aug 1956, MacKee 5139 (L1815294image!, P03292522image!†); Contrefort de la roche Ouaième, 400 m, 27 Dec 1964, MacKee 11865 (P03292502image!†); Haute Diahot, forêt de Tendé, exploitation forestière Frouin, 500 m, 31 Mar 1969, MacKee 20470 (NOU072078!, P03276833image!, P04234036image!); Tiwaka, Moindip, 550 m, 31 Mar 1974, MacKee 28455 (P04427679image!†); Col Maré, Amoa-Tiwaka, 500 m, 13 Aug 1977, MacKee 33612 (CANB718871.1!, NOU072079!, P04427665image!†); Pouébo, Ouangati, 700 m, 20 Oct 1978, MacKee (legit Cherrier)35947 (P04427672image!†); Néhoué, vallée de la Rade, 50 m, 8 Mar 1979, MacKee 36698 (NOU072077!, P04427673image!†); Piémont sud du Kantalupaik, 300 m, 20°51'6.012"S, 165°0'36"E, 1 Nov 2017, Pignal, Munzinger & Bruy 5263 (P01073109image!); Région de Pouembout au nord de Forêt Plate, 25 Mar 1981, Suprin 1079 (NOU072080!); Plateau de Tango, 650 m, 20 Oct 1981, Veillon 4555 (NOU072082!, P04427674image!†); Cap Tonnerre, 1861–1867, Vieillard 816 (P03292525image!†); sur la montagne à Balade, 1855–1860, Vieillard 817 (P03292446image!†, P03292497image!†); Gatope, 1861–1867, Vieillard 817 (P03292496image!†). South Prov.: Dumbéa, Baudouin 498 (P03292450image!†); Mont Mou summit, 3500 ft, 15 Mar 1914, Compton 574 (BM013867014image!, P03292498image!†); Mts Koghis, 300 m, 25 Jan 1927, Franc 486 (P03292501image!†); 1874–1876, Germain s.n.(P03292500image!†); Mé Aoui, 500 m, 8 Feb 1951, Guillaumin & Baumann-Bodenheim 10444 (P03292560image!†); Cultivated plant at IAC Port-Laguerre, 11 Jun 2020, originally collected at Cascade de Dogny, 915 m, 21°37'04.2"S, 165°53'18.8"E, 21 Jan 2019, Lannuzel 348 (NOU107487!); Cultivated plant at IAC Port-Laguerre, 5 Jan 2021, originally collected at Monts Koghis, sur le chemin du Pic Malaoui, 670m, 22°10'52.2"S, 166°30'43.2"E, 15 May 2020, Lannuzel 427 (NOU107488!); Ouipouin, 21°41'17.88"S, 165°59'13.56"E, 13 Dec 2018, Laudereau 1235 (NOU091403!); Vallée de la Thy, 400 m, 7 Jan 1956, MacKee 3741 (L1815296image!, P03292451image!†, P03292523image!†); Slope of Mt Koghi toward Vallée de la Thy (St Louis), 400–500 m, 24 May 1956, MacKee 4651 (L1815293image!, P03292520image!†); Col d’Amieu, Mont Pembai, 600 m, 15 Apr 1976, MacKee 31018 (NSW935344!, P04427663image!†); Dogny, la cascade, 26 Oct 2007, Munzinger 4621 (NOU030729!); Monts Koghis, propriété Lavoix, 11 Mar 1966, Nothis 67 (NOU072075!); Ile des Pins, 1860s, Pancher 473 (P03292495image!†); Auf den Hügeln bei Yaouhé, 150 m, 25 Sep 1902, Schlechter 14804 (L1815295image!, P03292494image!†).
Some plants on Vieillard 817, sur la montagne à Balade, 1855–1860 (P03292448image!†), see notes.
Lagenophora sublyrata is the most widespread species in the genus ranging from India and Sri Lanka to south-east Asia (e.g. Vietnam), China, Taiwan, Japan, Indonesia (e.g. Java), New Guinea, Australia and New Zealand. In New Caledonia, it has a widespread distribution (Fig.
Flowers and fruits have been recorded almost all year round with a peak of specimens in March but this could be an artefact. In cultivation, the species seems to flower throughout the year.
The species is relatively common on mainland, though often neglected by collectors, perhaps because it is inconspicuous or considered to be an exotic or weedy species. Hence, the number of localities where it occurs, judging by herbarium records, may be underestimated. The ecology of the species is rainforest on non-ultramafic substrates at low to medium altitudes. The invasive-introduced Rusa deer (Rusa timorensis) may represent a major threat through overgrazing or by trampling of herbaceous vegetation. Nonetheless, with over ten localities (sensu
Lagenophora sublyrata is a widespread species with variable leaf shape, indument and plant size. New Caledonia specimens are usually smaller in stature than typical plants from eastern Australia, but features of the roots, cypselae, scapes and involucral bracts are consistent with it. The specimen MacKee11865 bears two numbers on it; 11864 on the Paris herbarium label and on the wrapper and 11865 on a manuscript label by MacKee himself. MacKee’s field notebook (held at NOU herbarium) shows that 11864 is a Mitrasacme Labill., and 11865 is a Lagenophora sp. The correct collection number is therefore 11865. The specimen Vieillard 817 (P03292448image!†) is a mixed specimen with plants of both L. sublyrata and L. sinuosa. Vieillard used a confusing system of numbering of herbarium specimens and mixed specimens are well-known (see
We acknowledge the curators of CANB, L, MEL, NOU, NSW and P for the loan of herbarium specimens and especially Marc Pignal (P) and David Bruy, the curator of NOU. Ranee Prakash (BM) provided high resolution images of Compton specimens. Jérôme Munzinger and the P herbarium team are also acknowledged for the transmission of the destroyed herbarium list. Christian Laudereau (www.endemia.nc) and Dominique Fleurot (www.endemia.nc) shared their valued photos and observations, Daniel Lemaître for access to his estate and Giliane Karnadi-Abdelkader (IAC) for fieldwork. The authors are most thankful to the South and North Province authorities for providing collecting permits and to the South Province rangers Catherine Geoffray and Sophie Raillard for fieldwork on Îlot Casy. This study was partly supported through a grant aiming at identifying rare and threatened species in the Lagenophora/Pytinicarpa complex from the Kaala massif (funded by Société des Mines de la Tontouta,