Lagenophora (Asteraceae, Astereae) in New Caledonia

Abstract 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.


Introduction
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 (Nesom and Robinson 2007). Presently, there are 12 species in Australia (Wang and Bean 2019), nine species in New Zealand (Breitwieser et al. 2012), three species in southern South America (Cabrera 1966;Pruski 2017), two in New Guinea (Wang and Bean 2020) and two in New Caledonia (Munzinger et al. 2020+). The genus also extends to Indonesia, and as far as Sri Lanka and southern Japan (Walker 1976; Bean 2019, 2020). The genus is considered non-monophyletic as species of Solenogyne Cass. are nested between the two main clades of Lagenophora retrieved by Sancho et al. (2015).
The genus was apparently first recorded in a publication for New Caledonia by Schlechter (1906) who identified his own specimen (Schlechter 14804) as L. billardierei Cass. An earlier specimen (Pancher 473, P03292495) collected during the 1860s was, however, also identified as a Lagenophora by its collector. Other early specimens from this period by Balansa, Baudouin, Deplanche, Germain and Vieillard were either unidentified at the genus level or identified as Strongylosperma reptans Benth. [now synonymous to Leptinella reptans (Benth.) D.G.Lloyd & C.J.Webb]. In his catalogue of the plants of New Caledonia, Guillaumin (1911: 177) identified most of these early specimens as L. billardierei. Later, Moore (1921: 345) also identified some Compton specimens as L. billardierei and described a new species under L. neocaledonica S.Moore. Guillaumin maintained the two aforementioned names under Lagenophora in his revision of New Caledonian Asteraceae (Guillaumin 1937) and in his flora of New Caledonia (Guillaumin 1948) while the name L. neocaledonica was excluded from Lagenophora by Cabrera (1966: 307) and then placed in the synonymy of Pytinicarpa sarasinii (Däniker) G.L. Nesom by Nesom (2001). In addition, Cabrera (1966) listed two species of Lagenophora for New Caledonia viz. L. lanata A.Cunn. for the Schlechter specimen and L. gracilis Steetz for all the other specimens, L. billardierei being synonymous with L. stipitata (Labill.) Druce, an Australian taxon not present in New Caledonia. Wang and Bean (2019) put L. lanata into synonymy under L. sublyrata (Cass.) A.R.Bean & Jian Wang ter and narrowed the definition of L. gracilis as a Western Australian endemic species. After searches and studies over years, we recognise two species for New Caledonia -one is the indigenous L. sublyrata, the other one being a new endemic species described here.

Materials and methods
This revision is based on morphological examination of Lagenophora material from the following herbaria: BRI, CANB, L, MEL, NOU, NSW and P (acronyms following Thiers 2020+). Images of type specimens held at BM, FI, G, HAL, K, M, NY, P and W have also been examined. An exclamation mark (!) is used for the specimens physically seen while "image!" is used for specimens seen only by means of digitized images. In January 2017, a whole shipment of loaned Lagenophora specimens from P was destroyed at the Australian border before they could be examined by the borrowing botanists. These specimens were examined by means of scans and are marked with a † symbol. Some nomenclatural typifications were made subsequent to this event, including that of L. sublyrata, discussed here (Bean and Wang 2017;Bean and Jabbour 2018). Morphological descriptions and terminology follow Harris and Harris (2001). In addition to dried specimens, live specimens were collected by the first two authors and Christian Laudereau and subsequently cultivated at the nursery of Institut Agronomique néo-Calédonien in Païta, Port-Laguerre, to make finer measurements and facilitate observations. As much as possible, localities recorded on older specimens were visited to recollect the plants and to assess the ecology where they were found. Most measurements are based on live cultivated material under binocular Olympus SZ2-ILST equipped with a camera. When dried material was used, the dimensions are based on material (i.e. florets) reconstituted with boiling water. Specimen locations were mapped with QGIS 3.10 (QGIS Development Team 2020) to generate the distribution maps and to help for IUCN (2019) evaluation assessments. The occurrence data have been uploaded to the Global Biodiversity Information Facility (GBIF) via the Pensoft Data Hosting Center at the GBIF's Integrated Publishing Toolkit (IPT) (https://doi.org/10.15468/zkjcfx).
Distribution and habitat. 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. 2). It inhabits the wet forests and open scrublands (maquis minier), mainly on serpentinic alluvium but has been recorded also on peridotitic derived soils at higher altitude and on vertisols at low altitudes.
Phenology. Both flowers and fruits were recorded from February through November from herbarium specimens.
Etymology. 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. Conservation status. 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 IUCN 2019) recorded, L. sinuosa does not meet the requirements for a threatened species and qualifies for the Least Concern (LC) status.
Notes. 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 Saldivia et al. (2020), rank redundancy with monotypic genera is already relatively high (36%) in Asteraceae. Thus, considering existing genera in the region, this new species is here included in the genus Lagenophora because of the involucres with 1 to 2-seriate ray florets, the disc florets with 4-5 corolla lobes, and cypselae with a glandular beak. The new species is atypical in the genus because of its distinctive but variable leaf shapes and more importantly its cypselae that have longitudinal ribs on both surfaces. Its characteristics were also compared with other genera in the subtribe Lagenophorinae, including the closely related genera in the region, viz. Pytinicarpa G.L.Nesom, Solenogyne Cass., Keysseria Lauterb. and Myriactis Less., or the morphologically similar Brachyscome Cass. in Brachyscominae. In Table 1 also a comparison with other members of the genus is made. Further molecular based studies among these related genera, and including New Caledonian samples, are highly recommended to clarify its position. Description. 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 mm 2 , each 0.3-0.6 mm long; lower leaf surface pale green, with 3-7 trichomes per mm 2 , 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 mm 2 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.
Phenology. 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.
Conservation status. 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 IUCN 2019) recorded, L. sublyrata does not meet the requirements for a threatened species and qualifies for the Least Concern (LC) status.
Notes. 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 Hopkins and Bradford 2009;Morat 2010). An additional problem is the mounting of several plants on the same sheet, probably done by Sébastien-René Lenormand at Institut Botanique de Caen (CN) before the New Caledonian collection was sent to P.