TAXON 00 (00) • 1–13 Patil & al. • The identity of Zingiber neesanum NOMENCLATURE Fixing stray traditions in gingers: The identity and nomenclatural history of Zingiber neesanum and other entwined names Sameer Patil,1 Rajendra D. Shinde,2 Jana Leong-Škorničková3 & Rushabh Chaudhari2,4 1 Botanical Survey of India, Northern Regional Centre, 192 Kaulagarh Road, Dehradun – 248195, Uttarakhand, India 2 Blatter Herbarium (BLAT), St. Xavier’s College (Autonomous), Mumbai – 400001, Maharashtra, India 3 The Herbarium, Singapore Botanic Gardens, National Parks Board, 1 Cluny Road, Singapore – 259569 4 Naoroji Godrej Centre for Plant Research (NGCPR), Shirwal, Satara – 412801, Maharashtra, India Address for correspondence: Rushabh Chaudhari, rushabhchaudhari@gmail.com DOI https://doi.org/10.1002/tax.12567 Abstract The identities of two historical names, Zingiber neesanum (≡ Alpinia neesana) and Z. macrostachyum are re-investigated. Both have been considered to represent the same taxonomic entity by various workers in the past 160 years. Yet, based on traditional usage, they were applied to two distinct taxa occurring in the Western Ghats (India), both superficially similar by having long-stalked inflorescences, but much distinct in flower colour and overall morphology. Alpinia neesana was originally described only from fruiting material, while Z. macrostachyum was described from flowering material of a species with white flowers. Dalzell’s presumption that his taxon was identical to Graham’s, and unambiguous inclusion of A. neesana in his description made his name nomenclaturally superfluous and illegitimate. After realising the illegitimacy of Dalzell’s name, a new combination was created in Zingiber for A. neesana. The name Z. neesanum continued to be applied sensu Dalzell to the white-flowered species mainly in the central and southern Western Ghats, but it was also applied to a yellow-flowered species in the northern Western Ghats, for which another name, Z. diwakarianum, was proposed recently. Despite the absence of any original material relating to Graham’s or Dalzell’s descriptions, re-collections from the type localities prove beyond any reasonable doubt that the yellow-flowered species is the true Z. neesanum, making Z. diwakarianum a synonym, and that the existing name Z. anamalayanum has to be applied to the white-flowered species, originally described by Dalzell as Z. macrostachyum. The significance of authentic primary data sources concerning botanical history such as correspondence and manuscripts of collectors and botanists, and the importance of making these resources widely accessible through digitisation are highlighted. Keywords Alpinia neesana; Dalzell; Graham; Kew rule; typification; Western Ghats; Zingiber anamalayanum; Zingiber diwakarianum; Zingiber macrostachyum; Zingiberaceae ■ INTRODUCTION Zingiber Mill. (Miller, 1754) (Zingiberales: Zingiberaceae) is a genus of rhizomatous herbs widely distributed in tropical to subtemperate Asia with highest diversity in the monsoonal parts of Asia (Kishor & Leong-Škorničková, 2013; Bai & al., 2020). Several species are grown worldwide for their culinary, medicinal, and ornamental values with Ginger of commerce or Common ginger (Z. officinale Roscoe) being the best-known and economically most important example. Slightly over 300 names have been published in Zingiber so far (Govaerts & al., 2020), corresponding to 150–220 species (Kishor & LeongŠkorničková, 2013; Bai & al., 2020), making it one of the largest genera in Zingiberaceae. A detailed introduction to the genus and the sectional classification currently followed are given in Bai & al. (2015a) and not repeated here. No recent comprehensive revision of Zingiber in India exists, although the last decade has seen a steady rise of newly described species as well as new distributional records, increasing the total number of species to c. 36 (Bhargava & Nair, 1979; Bhaumik & Pathak, 2008; Sabu & al., 2009, 2013; Kishor & Leong-Škorničková, 2013; Kumar & al., 2013, 2015; Thongam & Konsam, 2014; Talukdar & al., 2015; Singh & Singh, 2016; Prabhukumar & al., 2016; Mibang & Das, 2016; Odyuo & Roy, 2019; Odyuo & al., 2019a,b; Thachat & al., 2020; Jayakrishnan & al., 2021). Despite the continuing description of new species, the identities of many Zingiberaceae species described in early history, including widely cultivated species, remain obscure (e.g., Leong-Škorničková & al., 2008a,b; Droop & al., 2013; Bai & al., 2020). The main reasons include scanty original descriptions and missing or poorly preserved herbarium material. As aptly pointed out by Turland (2019: 65), in the absence of type specimens, mere tradition determines the application of names, creating an unstable nomenclatural situation, as traditions may stray over time. Work on living flowering material is critical in Zingiberaceae because the diagnostic characters, especially those linked with flowers and rhizomes, do not preserve well in dry specimens (Burtt & Smith, 1976; Williams, 2004; Škorničková & Sabu, 2005). Numerous synonyms and new combinations, solely based on descriptions, or icons and illustrations, have resulted in many errors and further Article history: Received: 8 Mar 2021 | returned for (first) revision: 23 Apr 2021 | (last) revision received: 12 May 2021 | accepted: 14 May 2021 Associate Editor: Jefferson Prado | © 2021 International Association for Plant Taxonomy. Version of Record 1 TAXON 00 (00) • 1–13 Patil & al. • The identity of Zingiber neesanum aggravated an already ambiguous situation in many taxa concerning their identity and nomenclature (Veldkamp, 2012). Description of new species based on fruiting material must also be strongly discouraged because this practice has led to wrong generic placements in the past that can only be corrected after re-collection of living material from the type localities, e.g., Zingiber ellipticum (S.Q.Tong & Y.M.Xia) Q.G. Wu & T.L.Wu (≡ Plagiostachys elliptica S.Q.Tong & Y.M. Xia) (Wu & al., 1996). Our taxonomic and nomenclatural studies of Zingiber in the Western Ghats (India) have therefore not surprisingly revealed that the identities of some early-described names from this region remain unclear. Here we focus on re-investigating the identity of an enigmatic species from the northern Western Ghats described by Graham (1839) as Alpinia neesana J.Graham. The description was based on fruiting material and describes a species with an inflorescence that arises separately from the leafy shoot and is two feet (c. 60 cm) tall or more. In 1852, Dalzell described a Zingiber species with longstalked lateral inflorescences and white flowers with labellum ornamented with purple lines (here called the white-flowered species). He was convinced that it was the same as Graham’s A. neesana, as he explicitly stated “This is also mentioned in Graham’s Catalogue under No. 1455 as Alpinia Mesana from the native name “Meesum;” neither had the flowers of this been observed at the time of publication” (p. 342; note the misspelling of neesana and mesana). Dalzell, most probably exercising the flexibility of the informal Kew rule (Jackson, 1887; Stafleu, 1966; Stevens, 1991), named this species Z. macrostachyum Dalzell. This treatment was reiterated in Dalzell & Gibson (1861) and also followed by subsequent workers (Horaninow, 1862; Baker, 1892; Schumann, 1904; Cooke, 1907; Fischer, 1928) who treated Z. macrostachyum as the correct name for A. neesana. Santapau (1953, 1967) also used the name Z. macrostachyum but, contrary to previous workers, described his plants as having cadmium-yellow flowers on a scape up to 1 m in length. Particularly in his work on floristics of Khandala (Santapau, 1953), he noted that dried specimens of this and many other species of the Zingiberaceae are most difficult to examine and that his observations were based on a very large number of living plants. He provided a detailed description of the plant he studied from Khandala (Maharashtra) but expressed doubt about the conspecific treatment of these two names. Ramamoorthy (1976), who treated the Zingiberaceae in the Flora of Hassan District (Karnataka), realised the superfluous and illegitimate status of Z. macrostachyum (Art. 52.1, 52.2, 52.3 of the ICN, Turland & al. 2018) and proposed a new combination Zingiber neesanum (J.Graham) Ramamoorthy in Saldanha & Nicolson (1976). Thereafter, wide discrepancies occurred in the application of Z. neesanum in regional floristic works. In most of these publications, the descriptions, specimens cited and associated photographs do not match each other well. The numerous authors of subsequent regional floristic works may be informally divided into two main camps. While both groups agreed that the 2 species described by Graham as Alpinia neesana and by Dalzell as Z. macrostachyum were conspecific and continued to use the combination Zingiber neesanum, botanists particularly from the northern Western Ghats applied it to the species with cadmium-yellow flowers consistent with Santapau’s description of Z. macrostachyum (e.g., Bole & Almeida, 1986; Kulkarni, 1988; Lakshminarasimhan, 1996: 84; Yadav & Sardesai, 2002: pl. 44; Datar & Lakshminarasimhan, 2013: pl. 25(f)). On the other hand, botanists mainly from the central and southern Western Ghats applied it to a species with white flowers, consistent with Dalzell’s description of Z. macrostachyum (e.g., Bhat, 1993; Mohanan & Sivadasan, 2002; Sabu, 2003, 2006; Punekar & Lakshminarasimhan, 2011; Singh, 2011; Pushpakaran & Gopalan, 2014). Several works even avoided stating the flower colour altogether (e.g., Bole & Almeida, 1986; Kothari & Moorthy, 1993). Singh (2011) added to this already confusing situation when he stated that Santapau (1953) erroneously applied the name Zingiber macrostachyum to a species with cadmiumyellow flowers, and coined for it a new name, Z. diwakarianum R.Kr.Singh, while continuing to use Z. neesanum for the white-flowered species, following Sabu (2003). Here we re-investigate the identity, taxonomy, and nomenclatural history of two old names, Zingiber neesanum (≡ Alpinia neesana) and Zingiber macrostachyum, and two recent names, Z. diwakarianum and Z. anamalayanum Sujanapal & Sasidh., to determine to how many and to which species they correspond. To achieve that, we: (a) investigate the identity of the taxon described as A. neesana to confirm its generic placement and establish its correct name; (b) investigate the identity of Z. macrostachyum and determine whether or not it is conspecific with A. neesana; (c) review descriptions of Z. diwakarianum and Z. anamalayanum, and all morphologically similar species from Z. sect. Zingiber in the Indian subcontinent to determine other possible names in existence corresponding to these taxa; and (d) revise the synonymy and typification for all names involved, and provide updated descriptions and distributions of the recognised taxa. ■ MATERIALS AND METHODS This study largely follows the methods outlined in previous studies dealing with the identity of early-described Zingiberaceae, all of which relied on a combination of exhaustive herbarium and reference research combined with collection of new flowering material from the type localities and nearby areas (e.g., Leong-Škorničková & al., 2008a,b; Bai & al., 2020). All relevant names were identified by an extensive search of databases, namely IPNI (2020), Zingiberaceae Resource Centre (Newman & al., 2021), World Checklist of Selected Plant Families (Govaerts & al., 2020), and Tropicos (2021), and literature pertaining to Zingiber, with particular focus on the Indian subcontinent. The working methods of the authors of names and collectors of original material were investigated through Taxonomic Version of Record TAXON 00 (00) • 1–13 Patil & al. • The identity of Zingiber neesanum literature (Stafleu & Cowan, 1976–1988), biographies, and other publications dealing specifically with Indian botanical history (e.g., Burkill, 1954–1963; Noltie, 2002). Correspondence was consulted using the Directors’ Correspondence Project, Kew through JSTOR Global Plants (Global Plants, 2021). Original material, as well as all other specimens connected to the respective names, were searched for and examined at the following herbaria: BLAT, BM, BSI, CAL, CALI, DD, E, FRLH, K, L, MH, P, PDA and SUK. Highresolution images of specimens were sourced from websites or through the curators of the following herbaria: CGE, HBG, JCB, LE, M, MEL and W. Living flowering material collected from the type localities as well as other locations within Western Ghat region was examined during 2000–2020. The general descriptive terminology is based on Beentje (2010), and the style and level of detail follow the recent works of Bai & al. (2015a,b, 2016, 2018a,b, 2020). A distribution map of taxa based on 95 data points derived from fieldwork and herbarium records was created using QGIS v.2.18. Photoplates were created in Adobe Photoshop v.22.2.0. ■ RESULTS Search of original material of Alpinia neesana. — Alpinia neesana was described by John Graham (*1805–†1839) in A catalogue of the plants growing in Bombay and its vicinity (Graham, 1839). Graham was a native of Dumfriesshire, Scotland and arrived in India in 1828, where he held the post of “Deputy postmaster general” until his death on 28 May 1839 at Khandala, India (Preface in Graham, 1839). Graham briefly compared A. neesana to the preceding species in his catalogue, A. nimmonii J.Graham (≡ Zingiber nimmonii (J.Graham) Dalzell), stating that it “Has much the appearance of the last, only it is smaller and the leaves are of a darker green, with the margins frequently reflected.” He also states that “The flowers of neither species [i.e., A. nimmonii and A. neesana] have yet been observed.” The descriptive part states: “The scape, or culm, (for it is jointed, compressed and furnished with sheathing leaves like the Gramineae,) rises at a little distance from the stem to a height of 2 feet or more, and is terminated with a closely imbricated spike, of dingy brown fleshy capsules, which are hid in bracts of a lighter colour.” The last part of the description gives the whereabouts of this species as “The hills at Wargaum [Vadgaon]; the borders of Lanowlee [Lonavala] grove, and below bushes and detached trees on the undulating ground about Kandalla [Khandala]. The table land of Mahableshwur [Mahabaleshwar]. (Dr. Murray.)” (Graham, 1839: 207). No specimens are cited in the protologue of Alpinia neesana and none of the authors dealing with this name since cited any type material. Sabu (2003) explicitly stated in his account of Zingiber neesanum: “Types not available”. Clues to the existence and possible location of Graham’s herbarium are scarce. Manudev & Nampy (2016) stated that there was no evidence that Graham maintained a herbarium at the time of his work on the plants of Bombay and vicinity. Leong-Škorničková & al. (2010), who dealt with typification of names relating to Indian Curcuma, did not discount the possibility that Graham had a herbarium but, despite extensive searches in 35 herbaria, none was found and two binomials attributed to J. Graham published in the same work as A. neesana were neotypified by material newly collected from the type localities. An interesting piece of correspondence between Graham and William Jackson Hooker (*1785–†1865) dated 6 June 1834 has, however, revealed that J. Graham certainly maintained a herbarium, which he transferred to a German baron Carl Hügel (Global Plants, 2021: letter KDCAS1119, digital image!). This transfer of his herbarium is confirmed by J. Nimmo (Global Plants, 2021: letter KDCAS1136, digital image!). Carl (Karl) Alexander Anselm Freiherr [= baron] von Hügel (*1794–†1870) was a German-born Austrian soldier, traveller, and botanist. As stated by Burkill (1954: 874), “Von Huegel was liberally supplied by the Bombay botanists with collections which were purchased from him by the Vienna Museum.” Hügel’s collections are held by W, with duplicates in BR, CGE, CN (herbarium transferred to P), HBG, K, LE, M, and MEL (Stafleu & Cowan, 1976–1988). We corresponded extensively with these herbaria but failed to trace any original material of Alpinia neesana that could be linked to J. Graham. Besides Graham, two more people are associated with the protologue, namely Dr. James Murray and Joseph Nimmo. It is evident from the protologue that the locality statement “The table land of Mahableshwur. (Dr. Murray.)” was contributed by Murray, most likely through personal communication. Dr. James Murray (*1802–†1855) was an Assistant surgeon in Indian Medical Service and later Superintendent of Mahabaleshwar from 1831 onwards (Parasnis, 1916). Murray shared friendship with Graham and also collected plants for him (Global Plants, 2021: letter KDCAS10318, digital image!). He also acted as one of the sources from whom Graham received materials for his catalogue (Preface in Graham, 1839). Murray took particular interest in the study of the local climate as well as the flora of Mahabaleshwar and seemed to rely on Graham as a source of knowledge in the botanical domain (Murray, 1837). Whether or not Murray maintained a herbarium is uncertain, but if he did, the location of this collection remains unknown. Graham did not see his publication through the press owing to his untimely death in 1839 after which his work was completed by his contemporary Joseph Nimmo (*Unknown–†1854). Graham supervised the printing of the first 200 pages, while pages 201–254 were supervised by Nimmo (Graham, 1839). Alpinia neesana was published on page 207, so we have also investigated Nimmo. Nimmo and Graham were close associates particularly with regards to this publication as evidenced from the preface notes as well as many sporadic additions provided by Nimmo in the descriptions throughout. These additions are exclusively marked by the annotation “N” (Global Plants, 2021: letter KDCAS10367, digital image!). No such Version of Record 3 TAXON 00 (00) • 1–13 Patil & al. • The identity of Zingiber neesanum annotation exists in the protologue of A. neesana. However, a letter from Nimmo to W.J. Hooker has revealed that Nimmo sent a few plants via parcel (whether living or in the form of an herbarium is ambiguous) to Hooker in 1834, and that some of these plants were collected at Khandala (one of the localities mentioned in the protologue of A. neesana) (Global Plants, 2021: letter KDCAS1138, digital image!). A slim chance therefore existed that this parcel might have contained material of A. neesana from or with a link to Graham. Hooker’s herbarium was purchased by K in 1867 (Stafleu & Cowan, 1976–1988). No herbarium specimen of A. neesana attributed to Nimmo or Graham was located in K. As our extensive search did not locate any original material, and the protologue does not contain an illustration, a neotype designation is necessary. Identity and generic placement of Alpinia neesana. — With no original material extant, the best way to understand the identity of Alpinia neesana, as well as to gather new material for neotypification, was through comprehensive fieldwork in the localities stated in the protologue. All localities (Vadgaon, Khandala, Lonavala, and Mahabaleshwar) were visited repeatedly at different seasons to search for a ginger species fitting the protologue. Our findings conclude that the only species with separate inflorescences composed of long peduncles (reaching up to 1 m) terminating in an imbricately bracteate spike, matching the description of A. neesana, is a locally common species with cadmiumyellow flowers, as described in detail by Santapau under the name Zingiber macrostachyum (Santapau, 1953, 1957, 1967) (see Fig. 1). The presence of a pulvinus, beak-shaped anther crest, and also the fact that the plant regularly enters dormancy during the dry periods of the year unambiguously place this species in Zingiber, and the erect inflorescence further places it in Zingiber sect. Zingiber. The correct name to be applied to this taxon is Z. neesanum (J.Graham) Ramamoorthy (Art 7.3 of the ICN, Turland & al., 2018), regardless of Ramamoorthy’s description, which merely follows Dalzell in featuring a species with white flowers. No white-flowered species fitting Graham’s description could be found in any of the localities cited in the protologue of Z. neesanum, suggesting that Dalzell’s taxon is a distinct entity. As already stated in the introduction, Ramamoorthy correctly recognised the illegitimacy of Z. macrostachyum, but did not realise that Dalzell’s and his descriptions refer to another taxon than Graham’s, which needs a name. Singh (2011) correctly recognised the heterogeneous application of Z. neesanum to two distinct taxa, but erred in interpreting the identity of white-flowered ginger as Z. neesanum. He described the cadmium-yellow-flowered ginger as Z. diwakarianum, which has to be treated as a heterotypic synonym of Z. neesanum. Zingiber neesanum (J.Graham) Ramamoorthy in Saldanha & Nicolson, Fl. Hassan Dist.: 769. 1976 ≡ Alpinia neesana J.Graham, Cat. Pl. Bombay: 207. 1839 ≡ Zingiber macrostachyum Dalzell in Hooker’s J. Bot. Kew Gard. Misc. 4: 342. 1852, nom. superfl. & illeg. – Neotype (designated here): INDIA. Maharashtra, Pune District, Khandala, on the way to St. Mary’s villa, 13 Jul 2019, R. Chaudhari 1609 (BLAT!; isoneotype: MH barcode MH309948!). — [Fig. 1] = Zingiber diwakarianum R.Kr.Singh in Indian J. Forest. 34(2): 245. 2011 – Holotype: INDIA, Maharashtra, Satara Dist., Mahabaleshwar, 15 Jul 2008, R.K. Singh 184173A (CAL barcode CAL0000003998 [digital image!]; isotypes: BSI barcodes BSI0000000363 [R.K. Singh 184173B]! & BSI0000000364 [R.K. Singh 184173C]!), syn. nov. Notes. – Colour photographs of this species were published by Yadav & Sardesai (2002: pl. 44, top right), Datar & Lakshminarasimhan (2013: pl. 25f), and Lakshminarasimhan & al. (2019: pls. 20C, 21A) under the correct name Zingiber neesanum, though the descriptions and the specimens cited do not unequivocally correspond to this species. A historical colour painting published in Noltie (2002: 157, pl. 90, left element, identified in the text as Z. cylindricum Thwaites), an unpublished line drawing attributed to J.E. Stocks at K (SGCL 50) and an illustration in Lakshminarasimhan (1996: 84) based on K. Hemadri 99746 (BSI) also illustrate this taxon. Zingiber neesanum is endemic to the northern Western Ghats and is presently known to occur only in eight districts of Maharashtra (Ahmednagar, Kolhapur, Pune, Sangli, Satara, Sindhudurg, Ratnagiri, Thane; see Fig. 2). It occurs in patches along forest edges, in shady or open areas on slopes, and among bushes in semi-evergreen to evergreen forests, between 200 and 1200 m altitude. Flowering occurs from June to August, followed by fruiting, which lasts till December. A detailed description and a complete list of specimens examined are provided in Appendix 1. Identity of Zingiber macrostachyum Dalzell and its correct name. — Although Zingiber macrostachyum is an illegitimate name, it is important to investigate its identity and provide a correct name for this white-flowered taxon, to which most workers have erroneously applied the name Z. neesanum. Nicol Alexander Dalzell (*1817–†1878) was a Scottish botanist and forester, who held the post of the conservator of forests in Bombay between 1841 and 1870 (Stafleu & Fig. 1. Zingiber neesanum (J.Graham) Ramamoorthy, A, Habit in early monsoonal season; B, Leafy shoot in late monsoonal season; C & D, Inflorescences with flowers in side and front view; E, Plant in early fruiting stage; F, Detail of ligules (lateral view); G, Infructescence (c. 40 cm long); H, Dehisced capsules; I, Cross-section of rhizome; J, Young fruit and mature seeds (with and without aril; scale 5 mm); K, Flower dissection, from the bottom left: bract, bracteole, corolla lobes, labellum in side view, floral tube with ovary, calyx and anther attached; upper left: two labella showing variation in shape; upper right: stamen in front and back view and overy with epigynous gland (scale 10 mm). — Photographs: R. Chaudhari from neotype locality in Khandala. 4 Version of Record TAXON 00 (00) • 1–13 Patil & al. • The identity of Zingiber neesanum Fig. 1. Caption on page 4. Version of Record 5 TAXON 00 (00) • 1–13 Patil & al. • The identity of Zingiber neesanum Cowan, 1976–1988). He described Zingiber macrostachyum based on plants from Ram Ghat (Ramghat), the present region of Maharashtra-Goa state border, and near the Central Western Ghats. The Latin description provided corresponds to a plant with red pubescent stem, lanceolate acuminate leaves dark green above, paler and pubescent beneath, inflorescence with a long peduncle arising separately from the rhizome, spike consisting of obovate rusty red bracts with acute apex, white corolla, labellum 3-lobed with round and emarginate midlobe marked with diverging/fan-like (flabellatim) purple lines, obovate red pubescent capsules, dark purple seeds altogether hidden by white fimbriate aril, flowering spikes rusty red, one and a half feet long, fruiting ones much longer and bright red. No specimens are cited in the protologue of Zingiber macrostachyum. Dalzell’s collections are known to be deposited at CAL, DD, K, and W. No specimens were found by us in CAL and DD, and the curators at K and W also confirmed that there are no specimens attributed to Dalzell in their collections in connection with this name. A set of 42 watercolour drawings of plants with links to N.A. Dalzell exists at BM (Stafleu & Cowan, 1976–1988; Noltie, 2002), but these do not include any Zingiberaceae species (M. Nandikar & S. Bramhadande, pers. comm.). Fieldwork carried out in the type locality Ramghat and the nearby region has resulted in a collection of a ginger closely fitting the description of Zingiber macrostachyum (Fig. 3), which is clearly distinct from the species we have identified as Z. neesanum. Of 36 Zingiber species currently known to occur in India, only 11 belong to the nominal Z. sect. Zingiber, of which we have shown Z. diwakarianum to be synonymous with Z. neesanum. Of the remaining nine species, only three bear some resemblance to the species described by Dalzell as Z. macrostachyum by having fusiform spikes composed of bracts with acute apices. They are Z. purpureum Roscoe, Z. cylindricum, and Z. anamalayanum. The identity of Z. purpureum (often misidentified in the last 20 years as Z. montanum (J.Koenig) Link ex A.Dietr.) has been elaborated in detail by Bai & al. (2020). This species can be distinguished by its bright yellow and strongly aromatic rhizome and pale yellow flowers with no prominent red or purple striation, although a few purple spots rarely occur at the very base of the labellum in some populations, but are microscopic and obscured by the anther. Zingiber cylindricum was described originally from Sri Lanka. Sharma & al. (1984) listed this species as occurring in Karnataka without any details, but this occurrence was deemed doubtful by Bhat (1993), and we also have no proof that this species occurs in India. The distinct Fig. 2. Distribution of Zingiber neesanum and Z. anamalayanum based on 95 points derived from fieldwork and herbarium data. 6 Version of Record TAXON 00 (00) • 1–13 Patil & al. • The identity of Zingiber neesanum Fig. 3. Caption on page 8. Version of Record 7 TAXON 00 (00) • 1–13 Patil & al. • The identity of Zingiber neesanum feature of this species is closed leaf-sheaths, a character that only rarely occurs in the genus and is sufficient alone to eliminate this name from further consideration. This only leaves Z. anamalayanum, a species described from high-altitude grasslands (c. 1430–1600 m) of the southern Western Ghats that shares many characteristics with the species described by Dalzell as Z. macrostachyum, to be considered as the potential correct name. Sujanapal & Sasidharan (2010), in the original description of Zingiber anamalayanum, noted its close morphological affinities to Z. cylindricum, Z. montanum, Z. neesanum (a name that they applied to the white-flowered species sensu Z. macrostachyum Dalzell), and Z. zerumbet (L.) Sm., and provided a comparative table with 72 morphological characters, to distinguish Z. anamalayanum from these species. They also converted these characters into binary matrices and performed UPGMA and PCoA to support the recognition of the new taxon. Unfortunately, the authors did not state the origin of the data pertaining to these characters for any of the four comparative taxa in their table 1. It is clear that intraspecific variation among these species has not been adequately considered. In the discussion, the authors pointed out that Z. anamalayanum could be distinguished from Z. neesanum by the broad, coriaceous, sericeous leaves, the thick cylindrical spike, the free lateral lobes, and the highly wrinkled midlobe of the labellum. We are aware that plants of Z. anamalayanum are at first glance robust and with the characters as highlighted by Sujanapal & Sasidharan (2010). However, based on our field observations over many years in various habitats of the central and southern Western Ghats, we believe that the white-flowered species, which sets seeds freely, is very widespread and highly variable in terms of the height of the plant, size and shape of the leaves, ligules and overall robustness. The inflorescences are also highly variable with spike length ranging from 5 to 30 cm and peduncles from a few to 50 cm, being shorter during the flowering phase and slightly extending during fruiting. Plants growing at lower altitudes are more slender, with narrower leaf blades and with smaller spikes and shorter (rarely with almost negligible) peduncles, whereas those growing at higher altitudes and in more open habitats are increasingly robust with elongated spikes supported by long peduncles. The colour of the flowers ranges from white to cream-white with the margins of the labellum sometimes approaching pale yellow, while the fine ornamentation on the labellum ranges from pink-red, blood red to dark purple and may be limited to the central part of the labellum, or extend almost to the margins. The shape of the labellum and position and degree of connation of the lateral staminodes to the labellum also somewhat vary (Fig. 4). The plants we have observed and collected in the type locality of Z. macrostachyum (Fig. 3) exhibit characters more or less in the middle of the range of the variation, and all measurements largely or partially overlap with Z. anamalayanum. We have failed to find any discernible discontinuity in any of the characters that may warrant recognition of distinct specific or even subspecific units across the distribution range (Fig. 2). We are therefore of the opinion that the name Z. anamalayanum is the earliest validly published name for the white-flowered species. Future molecular-based studies will be undertaken to assess the broad species concept applied here. Zingiber anamalayanum Sujanapal & Sasidh. in Nordic J. Bot. 28: 289–293. 2010 – Holotype: INDIA. Kerala, Palakkad District, Nelliyampathy, Minampara 1450 m, 16 Aug 2008, P. Sujanapal & N. Sasidharan KFRI30829 (MH barcode MH00001411 [digital image!]; isotypes: CALI, KFRI, L not located). — [Figs. 3, 4] Notes. – Many authors have treated Zingiber anamalayanum under the name Z. macrostachyum. This includes the original description of Dalzell (1852), who thought that his white-flowered species was Graham’s Alpinia neesana, as well as those who followed his description (Dalzell & Gibson, 1861; Baker, 1892; Cooke, 1907; Fischer, 1928). Subsequently, all authors who followed Ramamoorthy’s (1976) treatment provided descriptions and/or photographs/illustrations of Z. anamalayanum under the incorrect name Z. neesanum (Saldanha & Dhawan, 1984: pl. 13; Manilal & al., 1988; Bhat, 2003; Sabu, 2003, 2006: 235, fig. 61 & pl. 17B & C.; Punekar & Lakshminarasimhan, 2011: pl. 112a & b; Datar & Lakshminarasimhan, 2013; Nayar & al., 2014). A photograph of Z. anamalayanum was also published by Yadav & Sardesai (2002: pl. 44, bottom left) under the name Z. purpureum and by Lakshminarasimhan & al. (2019: fig. 20B, flowering plant under the name Z. montanum and fig. 21B, fruiting plant under the name Z. neesanum). Zingiber anamalayanum is a widespread species occurring in many districts across five states of India: Maharashtra (Kolhapur), Goa (North Goa, South Goa), Karnataka (Belagavi, Chamarajanagar, Chikkamagaluru, Dakshin Kannada, Hassan, Kodagu, Mysore, Shimoga, Udupi, Uttara Kannada), Kerala (Idukki, Kannur, Kollam, Kottayam, Palghat, Pathanamthitta, Thiruvananthapuram, Thrissur, Wayanad), and Tamil Nadu (Coimbatore, Nilgiri, Tirunelveli) (Fig. 2). It occurs infrequently in fragmented patches along forest edges and in the undergrowth of dense shady areas rich with humus in semievergreen to evergreen forests, at 50 to 1600 m elevation. Flowering occurs between June and August, followed by fruiting till December. Detailed descriptions covering the morphological variation and a complete list of specimens examined are provided in Appendix 2. Fig. 3. Zingiber anamalayanum Sujanapal & Sasidh., A, Habit; B, Young infructescence; C, Flower, semi-front view; D, Indehised capsule within bracts; E, Flower in bract, side view; F, Rhizome and immature root tubers; G, Flower dissection, from left: bract, bracteole, floral tube with ovary, calyx and stamen attached, corolla lobes, labellum; right bottom: seeds with and without aril (scale 10 mm). — Photographs: A–D, R. Chaudhari; E–G, Sushant More from Ram Ghat. 8 Version of Record TAXON 00 (00) • 1–13 Patil & al. • The identity of Zingiber neesanum ■ CONCLUSION If Graham had described flowers of Alpinia neesana or Dalzell had simply described his Zingiber macrostachyum without attempting to judge whether it was identical with Graham’s A. neesana, much chaos might not have occurred. Due to the lack of nomenclatural types, subsequent workers carried forward locally established but taxonomically heterogeneous traditions in the application of these historical names of two distinct species that look superficially similar at the fruiting stage. Almost 160 years after this confusion arose, we show that the species first described by Graham from the northern Western Ghats as Alpinia neesana J.Graham is a species with cadmium-yellow flowers to which the correct name Zingiber neesanum (J.Graham) Ramamoorthy must be applied and that the recently coined but superfluous name Z. diwakarianum R.Kr.Singh has to be treated as a heterotypic synonym of it. The designation of a neotype collected in the flowering stage from the type locality of Z. neesanum will provide taxonomic clarity and stability for future studies. We also show that Z. macrostachyum Dalzell, a species with white flowers described by Dalzell from a region with close proximity to the central Western Ghats, and a name that is superfluous and illegitimate, is not, contrary to wide belief, conspecific with Z. neesanum. We clarify that the correct name for this widely distributed and highly variable species is Z. anamalayanum Sujanapal & Sasidh. The results of herbarium studies and fieldwork show that Z. neesanum occurs in the northern Western Ghats, whereas Z. anamalayanum occurs in the central and southern Western Ghats. The distribution of the two species is confirmed to overlap in the southern region of the northern Western Ghats in Kolhapur District, and most likely also in the adjacent District of Sindhudurg (both districts in Maharashtra). Based on observations, this overlap may further extend to Goa and the northern region of the central Western Ghats, in particular to Belagavi and North Kanara districts (Karnataka). The historic information uncovered during our extensive search for any information relating to herbarium collections of botanists such as J. Graham and J. Nimmo highlights the significance of authentic primary data sources such as the “Directors’ Correspondence Project” at Kew, as well as the paramount importance of making such rich repositories more widely accessible through digitisation, allowing botanists across the globe to tap its vast potential. Fig. 4. Flower shape and colour variation documented from various localities of Zingiber anamalayanum Sujanapal & Sasidh. A, Kerala, Minampara, the type locality of Z. anamalayanum; B, Maharashtra, Ram Ghat, the locality connected to Dalzell’s description of Z. macrostachyum; C, Karnataka, Pushpagiri Wildlife Sanctuary; D, Kerala, cultivated at Calicut Botanic Garden. — Photographs: A, P. Sujanapal; B, S. More; C, S. Patil; D, J. Leong-Škorničková. Version of Record 9 TAXON 00 (00) • 1–13 Patil & al. • The identity of Zingiber neesanum ■ AUTHOR CONTRIBUTIONS SP, RDS, RC and JL-Š designed the present study; SP, RC and JL-Š drafted the manuscript and figures; SP, RC and JL-Š conducted fieldwork for this study. All authors have read, commented and approved the final manuscript. — SP, https://orcid.org/0000-0003-0598-2114, sameerpatil. c@gmail.com; RDS, https://orcid.org/0000-0002-9315-9883, rajendra. shinde@xaviers.edu; JL-Š, https://orcid.org/0000-0003-4490-3490, jana_skornickova@nparks.gov.sg; RC, https://orcid.org/0000-00 03-0278-6599 ■ ACKNOWLEDGEMENTS We are grateful to the curators and other staff members of the following herbaria for facilitating access to specimens in their care or providing digital images of the specimens as well additional information on some of the collections: BLAT (Mr. Praveen Kale), BM (Dr. Ranee Prakash, Dr. Norbert Holstein), BSI (Dr. J. Jayanti, Dr. Priyanka Ingle, Mr. Dinesh Shirodkar), CAL (Dr. Mahua Pal, Mr. Anand Kumar, Mr. K. Karthigeyan), CALI (Dr. Jayakrishna T.), CGE (Dr. Lauren M. Gardiner), DD (Mr. P.K. Verma), FRLH (Dr. Ravikumar K.), HBG (Mr. Matthias Schultz), JCB (Dr. Sankara Rao, Mr. Mayur Bhagwat), K (Ms. Alison Moore, Ms. Julia Buckley, Ms. Wiebke Hillebrecht), LE (Ms. Larisa Orlova), M (Dr. Hajo Esser), MEL (Ms. Pine Milne, Mr. Wayne Gebert), MH (Dr. Ravi Kiran A.), P (Dr. Thomas Haevermans), PDA (Dr. Cyril Wijesundara), SUK (Dr. Manoj Lekhak, Dr. Sharad Kambale) and W (Dr. Christian Bräuchler). We also thank Dr. John McNeill and Dr. Kanchi Gandhi for nomenclatural comments, Dr. Mark Newman for language revision of this manuscript, Dr. M. Sabu for discussions on Zingiber in South India and for supervising JL-Š during her research stay at Calicut University; Dr. Mayur Nandikar & Ms. Sneha Bramhadande for sharing their expertise on N.A. Dalzell’s plant illustrations at BM and other valuable suggestions to this manuscript; Mr. Sushant More for the help during the fieldwork and photography, Dr. Sujanapal for providing photos of Zinbiber anamalayanum from the type locality. RC is indebted to Rajdeo Singh, Maniruddin Dhabak, Sadanand Gupta and Shahid Nawaz for their continual support. The authors are thankful to the Director, Botanical Survey of India, Kolkata; the authorities of St. Xavier’s College (Autonomous), Mumbai; Blatter Herbarium (BLAT); Naoroji Godrej Centre for Plant Research (NGCPR), Shirwal, Satara for the continual encouragement and providing research facilities. The research of RC is supported by the Research Assistant fellowship (2019–20) at BLAT by the Bombay St. Xavier’s college society trust (Trust Regn. No. F-2164 (BOM)). The research of JL-Š is supported by the National Parks Board, Singapore. 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Taxon 53: 1027–1032. https://doi.org/10. 2307/4135569 Wu, Q.G., Liao, J.P. & Wu, T.L. 1996. A new combination of the genus Zingiber – Z. ellipticum (S.Q. Tong & Y.M. Xia) Q.G. Wu & T.L. Wu and the systematic evidence. Acta Phytotax. Sin. 34: 415–420. Yadav, S.R. & Sardesai, M.M. 2002. Flora of Kolhapur District. Kolhapur: Shivaji University. Appendix 1. Zingiber neesanum (J.Graham) Ramamoorthy: description and complete list of specimens examined. Description: Perennial rhizomatous herb, 1–2 m tall. Rhizome fleshy, composed of units upright and parallel to pseudostem (along with previous year rhizomes/ units), single unit ovoid to barrel-shaped, 2.5–6 × 0.5–2.5 cm, terete in transverse section, light brown externally, pale to dark sulphur-yellow internally, aromatic; roots numerous, 5–30 cm long, fleshy, tuberous; tubers fusiform, 3–4 × 1.2–1.8 cm, yellow internally. Leafy shoots 1–2 per plant, erect, 0.8–2 m long, composed of 10–25 leaves (leaf-blades aggregated on upper 2/3 of pseudostem), basal 1/3 enclosed in bladeless reddish-purplish to greenish pubescent sheaths with hyaline densely pubescent margin; ligules bilobed, lobes 2–6 mm long, obtuse at apex, membranous, pubescent to sericeous; petiole 3–4 mm long (consisting of pulvinus only), sparsely pubescent; lamina narrowly elliptic to narrowly oblong, 7–38 × 5.5–8.5 cm, smooth (not plicate), mid to dark green and glabrous adaxially, abaxially paler green and with sparse silky hairs (especially on midrib and lateral veins), base subacute to cuneate, apex acute to acuminate, margins undulate. Inflorescences 1–2 per plant, radical, erect; peduncle 20–90 cm long, covered with mostly bladeless sheaths (rarely one to a few leaf vestiges of more or less developed leaves appear), sheaths green sometimes with reddish or purplish tinge, puberulent, 8–8.5 cm long; spike cylindrical, tapering to a narrow apex; 5–50 cm long, fertile bracts each subtending one flower, ovate to obovate with acute and distinctly recurved apex, 3.5–4 × 1.5–2.5 cm, chartaceous, green to yellow-orange with more or less prominent red to purple tinge and yellow-green to yellow apex (turning dark orange-red in fruiting stage), minutely pubescent externally, glabrous internally, margin hyaline, sericeous; bracteoles ovate, 2.5–3.5 × 1–1.5 cm, yellowish-green to orange, sometimes with reddish tinge, minutely pubescent externally, glabrous internally. Flower exserted from the bracts, 5–7 cm long; calyx tubular, 10–15 mm long, unilaterally split to 1/3–1/2 of the total calyx length, apex with 2–3 teeth, hyaline, semi-translucent white to pale yellow, pubescent externally and glabrous internally; floral tube 22–28 mm long, yellow (lighter at base, darker distally), dorsal corolla lobe ovate to elliptic, 25–33 × 5–13 mm, acute and slightly hooded at apex, cadmiumyellow with 10–13 translucent veins; lateral corolla lobes narrowly ovate, slightly oblique with acute and slightly hooded apex, 25–30 × 5–7 mm, cadmiumyellow with 5–6 translucent veins; labellum always shorter than lateral corolla lobes, 16–17 × 3–5 mm, apex deeply cleaved forming two narrow lobes each measuring 10 × 1.5–2 mm, cadmium-yellow to orange usually with red-maroon blotches (rarely plain), each lobe 3-nerved (nerves visible only in dry material); lateral staminodes irregularly ovate with bluntly acute apex, connate to labellum in basal 2/3–4/5 turning upwards nearly enclosing the anther, 10–14 × 3–4 mm, cadmium-yellow to orange, glabrous. Stamen 13 mm long (with anther crest unmanipulated); filament c. 3 × 2 mm, cadmium-yellow to orange, glabrous; anther c. 5.5 mm (excluding anther crest), connective tissue cadmium-yellow to orange, glabrous; anther thecae c. 5 mm long, cream-white, dehiscing throughout entire length; pollen cream-white; anther crest bluntly triangular with minutely dentate apex, 3–3.5 m long, cadmium-yellow to orange. Style filiform, yellowish-white, glabrous; stigma narrowly funnel-shaped, c. 1.5 × 1 mm, yellowish-white, ostiole ciliate. Ovary cylindric, slightly swollen in the middle, trilocular, 3–4 × 2– 2.5 mm, yellow-green, densely pubescent; epigynous glands 2, 2 mm long, cream-white, apices blunt. Fruit septifragal capsule; ovoid to ellipsoid, bluntly trigonous, c. 1.5 × 1.2 cm, 3-valved, externally yellowish with more or less rich red tinge, valves internally dark pink to bright pink-red; seeds in clusters of 5– 12 in each locule, more or less globose, 3.5–4 × 3 mm, pale brown to purplish brown (mature; white with reddish tinge when young); aril laciniate, white to cream-white (turns yellow in dried material). Specimens examined: Maharashtra. Ahmednagar Dist.: Harishchandragad, Pimparicha uran, 28 Sep 1970, B.M. Wadhwa 127841 (BSI); Ratangarh, 2 Oct 1970, B.M. Wadhwa 128043A (BSI). Kolhapur Dist.: Amba Ghat, 5 Jul 1984, M.K. Mistry 949 (BLAT); Panhala, 15 Aug 2002, S.P. Gaikwad SPG-219 (SUK). Pune Dist.: Singhar, 31 Jul 1902, L.D. Garade 55 (BSI); Purandhar, 22 Sep 1902, R.K. Bhide 1015 (BSI); Khandala, 25 Jun 1903, L.D. Garade s.n. (BSI); Katraj Ghat, 29 Jul 1909, S.V. Shevade s.n. (BSI); Khandala, 29 Jul 1916, collector unknown [Blatt. Herb.] 25943 (BLAT); Khandala, Jun 1917, collector unknown [Blatt. Herb.] 27936 & [Blatt. Herb.] 27974 (BLAT); St. Xavier’s ravine, Khandala, Oct 1918, collector unknown [Blatt. Herb.] 28200 (BLAT); Khandala, 3 Aug 1908, collector unknown [Blatt. Herb.] 25955 (BLAT); Khandala, St. Xavier’s villa, 4–5 Jul 1942, H. Santapau 574 (BLAT); ibidem, 15 Jun 1943, H. Santapau 2215 & 2216 (BLAT); ibidem, 4 Aug 1944, H. Santapau 4632 (BLAT); ibidem, 14 Jun 1946, H. Santapau 9149 (BLAT); ibidem, 15 Jun 1946, H. Santapau 9148 (BLAT); ibidem, 16 Jun 1946, H. Santapau 9150 (BLAT); Khandala, Convo. House, 1 Oct 1944, H. Santapau 4998 (BLAT); Khandala, Battery Hill Plateau, 19 Aug 1945, H. Santapau 6992 (BLAT); Sinhagad, 26 Jun 1956, Puri & Party 2784 (BSI); ibidem, 9 Aug 1956, G.S. Puri 5683 (BSI); ibidem, 1956, R.M. Patil 7823 (BSI); Bhimashankar road, Poona, 8 Oct 1962, K.P. Janarshanan 81676 (BSI); Sinhagad site, Haveli Taluka, 13 Jun 1963, M.Y. Ansari 87379 (BSI); ibidem, 29 Aug 1963, M.Y. Ansari 87813 (BSI); Purandar, on the way to Vajragad near N.C.C. camp, 19 Jul 1963, R.S. Rolla 88632 (BSI); Durga Killa Plateau, 19 miles west of Junnar, 26 Jun 1964, K. Hemadri 94354 (BSI); Lonavala, Hill slopes near Bushi village, 27 Jun 1964, B.V. Reddi 97959A (BSI); ibidem, 28 Jun 1964, B.V. Reddi 97959B (BSI); near Ambavane village, 26 Jul 1964, B.V. Reddi 99307 (BSI); Ambavane, forest near rest house, 6 Sep 1964, B.V. Reddi 99015 (BSI); Donaje, Sinhagad hill range, 7 Aug 1964, M.Y. Ansari 97552 (BSI); Ganesh caves, 4 miles north of Junnar, 17 Aug 1964, K. Hemadri 99746 (BSI); Junnar, Ralegaon hill, 23 Sep 1965, K. Hemadri 107285 (BSI); Khandala, on the way to St. Mary’s villa, 13 Jul 2019, R. Chaudhari 1609 (BLAT, MH); ibidem, 13 Jul 2019, Khandala, Kune Plateau, 18 Aug 2019, R. Chaudhari 1610 (BLAT). 12 Version of Record TAXON 00 (00) • 1–13 Patil & al. • The identity of Zingiber neesanum Appendix 1. Continued. Sangli Dist.: Shirala, 5 Oct 2019, Shahid Nawaz S200 (BLAT). Satara Dist.: Near Nakinda village, Mahabaleshwar, 18 Jul 1959, P.V. Bole 1985 (BLAT); to Kate’s point, Mahabaleshwar, 1 Aug 1958, P.V. Bole 1691 (BLAT); Koyana, Kumbharli Ghat, 4 Aug 1979, R.K. Kochhar 158504 (BSI); ibidem, 1 Sep 1978, R.K. Kochhar 152571 (BSI); Koyana, Shirshingi, 7 Sep 1978, R.K. Kochhar 153674 & 154114 (BSI); Kas, Aug 91, M.P. Bachulkar-Cholekar MPB 5131 (SUK). Sindhudurg Dist.: Phonda Ghat, 18 Aug 1965, B.G. Kulkarni 105573 (BSI); ibidem, 4 Oct [year unknown], B.G. Kulkarni 121445 (BSI). Ratnagiri Dist.: On the way to Mandangad Fort, approx. 300 m [altitude], 6 Jul 1983, M.K. Mistry 24 (BLAT); Shirgaori-Shindi village path, 16 Aug 1983, M.K. Mistry 208 (BLAT); Chiplun, Parshuram temple, 17 Aug 1983, M.K. Mistry 208A (BLAT); above Murshi village, near Amba Ghat, 27 Oct 1983, M.K. Mistry 515 (BLAT); Amba Ghat, 5 Jul 1984, M.K. Mistry 949 (BLAT); near top of Gothane, s.d., M.K. Mistry 949X (BLAT); Gothane Plateau, 9 Aug 2008, M.M. Lekhak MML-116 (SUK). Thane Dist.: Washala Range, Vinchu cha nal, 20 Sep 1968, K.V. Billore 116785 (BSI); Washala Range, Ajoba Parvat, 17 Oct 1967, K.V. Billore 111933 (BSI); Exact location uncertain: Malabar, Concan, Stocks & Law [Herb. Stocks] 15? (K [K000815661]); Malabar, Concan, Stocks & Law [Herb. Stocks] 15 (K [K001222301]); Concan, Stocks & Law [Mr. Law] s.n. (K [K000815662]); Malabar, Concan, Stocks & Law 16 (P [P00451126]). Appendix 2. Zingiber anamalayanum Sujanapal & Sasidh.: description and complete list of specimens examined. Description: Perennial rhizomatous herb, 0.5–1.5 m tall. Rhizome composed of units upright and parallel to pseudostem, single unit 5–2.5 × 0.5–1 cm, light brown externally, cream-white to pale yellow internally; roots numerous, 5–17 cm long, tuberous; tubers fusiform to narrowly fusiform (globular when young), 2.5–3.5 × 1–1.5 cm, cream-white to pale yellow internally. Leafy shoots 1–2 per plant, erect to slightly arching, 0.5–2 m long, composed of 16–20 leaves (leafblades aggregated on upper 2/3 of pseudostem), basal 1/3 enclosed in bladeless reddish-purplish to greenish pubescent sheaths; hyaline margin of leaf sheaths densely pubescent; ligules bilobed, lobes 1–13 mm long, obtuse at apex, pubescent; petiole to 3 mm long (consisting of pulvinus only), lamina narrowly elliptic to elliptic-oblong, 7.2–30 × 2.9–8 cm, smooth (not plicate), dark green and glabrous adaxially, pale green with prominent lateral veins and pubescent abaxially (sometimes glabrous with pubescence confined to midrib), base rounded to subacute, apex acute to acuminate, margins slightly undulate. Inflorescences 1–2 per plant, radical, erect; peduncle (5–)10–40 cm, covered with bladeless sheaths c. 2.7–5 cm long (rarely one to a few leaf vestiges of more or less developed leaves appear), dull green to maroon, puberulent; spike narrowly fusiform to fusiform, 8–30 cm long; fertile bracts each subtending one flower, elliptic to obovate with rounded, obtuse or acute apex (never recurved), 2.9–4.5 × 1.8–3 cm, coriaceous, dull olive green with brown to maroon-red tinge to completely maroon-reddish, puberulent externally, glabrous internally (turn rusty brown-red to bright to dark crimson red in fruiting stage), margin hyaline, glabrous; bracteoles narrowly elliptic to obovate, 3–3.7 × 1.6–2.5 cm, pale green at base with maroon tinge at apex, puberulent externally, glabrous internally. Flower exserted from the bracts, 4–6.2 cm long; calyx tubular, 13–25 mm long, hyaline, semi-translucent white, glabrous externally and internally; floral tube 24–35 mm long, white; dorsal corolla lobe ovate to elliptic, 25–30 × 10–16 mm, white to cream-white with c. 10 translucent veins; lateral corolla lobes ovate to elliptic, slightly oblique with acute and hooded apex, 13–22 × 5–8 mm, white to cream-white with c. 4 translucent veins; labellum shorter to or longer than lateral corolla lobes, broadly obovate with emarginate apex, 18–27 × 11–26(–30) mm (excluding staminodes), white to cream-white; central part more or covered with pink-red, maroon or dark purple blotches (pattern spreading from base to apex and towards margins), margins crisp without any blotches; lateral staminodes elliptic to narrowly obovate, 8–17 × 6–8 mm, obtuse at apex, connate to adnate to posterior part of labellum in basal 1/2–4/5, white to cream-white, glabrous. Stamen 20–30 mm long (with anther crest unmanipulated); filament negligible, to 1 mm long, white to cream-white, glabrous; anther 10–13 mm (excluding anther crest), connective tissue white to cream-white, sometimes with pink-red to purple stripes dorsally, glabrous; anther thecae 9–12 mm long, cream-white, dehiscing along entire length; pollen cream-white, anther crest 13–14 mm long, white to cream-white. Style filiform, white, glabrous; stigma slightly thicker than style, narrowly cylindric to narrowly funnel-shaped, recurved, ostiole ciliate. Ovary cylindric to ellipsoid, 3–5 × 2–4 mm, trilocular, cream-white to greenish, minutely to densely pubescent; epigynous glands two, 3–5 mm long, cream-white. Fruit septifragal capsule, broadly ovoid to ellipsoid, 14–20 × 11–14 mm, 3-valved, externally green to dull green with reddish tinge to dark red or maroon, sparsely puberulous to glabrescent, valves internally red to dark red; seeds 6–12 in each locule, ovoid, 4–5 × 3 mm, dark purple (immature) to black (mature), surface with fingerprint-like striations; aril laciniate, white (turns yellow in dried material). Specimens examined: Maharashtra. Kolhapur Dist.: Waghotre to Patane road, Ram Ghat, 26 Aug 2019, R. Chaudhari 1611 (BLAT, BSI, CAL, SING); Amba, Jul 94, M.P. Bachulkar s.n. (SUK); Gaganbawada, s.d., Priti D. Mahekar SUK 398 (SUK). Goa. North Goa Dist.: Nagargao, 5 Oct 1964, R.S. Raghavan 103263A (BSI); South Goa Dist.: Molem–Belgaum rd 1769′ [1769 feet; ca. 540 m altitude], 17 Aug 1970, N.P. Singh 124203 (BSI); Nandran, Molem N.P., 19 Oct 2002, M.N. Datar 186917 (BSI); Karnataka. Belagavi Dist.: Jamboti, s.d., Arun N. Chandore ANC 1065 (SUK); Jamboti, 10 Aug 2008, Arun N. Chandore ANC 745 (SUK). Chamarajanagar Dist.: Biligiriranga Hills, 1300 m, Jun 1938, Edward Barnes 1882 (K [K001222300]); Chikkamagaluru Dist.: Malabar, Bababoodan Hill, Stocks & Law [Mr. Law] 15? (K [K001222305]); Bababudan Hills, 26 Jun 1978, C.J. Saldanha KFP1721 (CAL); Canara & Mysore, Stocks & Law [Mr. Law] 15? (K [K001222309]). Hassan Dist.: Achihalli [Achanahalli], 16 Oct 1969, C.J. Saldanha 17937 (E [E00389839]); ibidem, C.J. Saldanha 17937 (JCB); Kodagu Dist.: Sampaje Ghat, 27 Aug 1985, K.G. Bhat 1947 (BSI); ibidem 1948 (BSI); Mundrote, 22 Dec 1996, R.D. Kshirsagar 176155 (BSI); Kote betta, Pushpagiri WLS, 12°31′49″N, 75°45′42″E, 1400 m, 18 Jun 2017, Sameer Patil 207320 (BSI); ibidem, 12°32′20″N, 75°46′ 37″E, 1270 m, 19 Aug 2018, Sameer Patil 206767 (BSI). Shimoga Dist.: Someshwar Ghat 59th mile, 29 Oct 1960, R.S. Raghavan 67912 (BSI); Barakana near Agumbe, 16 Oct 1962, R.S. Raghavan 83269 (BSI); Hulical Ghat, 9 Oct 1962, R.S. Raghavan 83103 (BSI). Udupi Dist.: Udupi, 29 Aug 1976, K.G. Bhat 291 (BSI). Uttara Kannada Dist.: Shauli, 20 May 1890, W.A. Talbot s.n. (BSI); Tutty hully on Kali naddi, s.d., Ritchie 719 (K [K001222302]); Karwar, Jun 1918, T.R. Bell 4092 (K [K001222306]); ibidem, Jun 1918, T.R. Bell 4092 (K [K001222307]); Devimane, Kotagal, 400 m, 9 Dec 1994, K.V. Devar 7633 (FRLH). Kerala. Idukki Dist.: 13 km from Kulemavu, 23 May 1984, M. Sabu 37328 (CALI); Pindimedu – Pooyamkutty, 50 m, 7 Jul 1988, P. Bhargavan 87401 (CAL, MH); Neriyumangalam, 750 m, 6 Aug 2002, Prasanth Kumar M.G. 86384 (CALI). Kannur Dist.: on the way to Panoth, 278 m, 13 Nov 1978, V.S. Ramachandran 58713 (CAL, MH); Nedumpayil, 550 m, 14 Aug 1979, V.S. Ramachandran 63940 (MH). Kollam Dist.: Thenmala, s.d., (det. Sabu, M., 2003) 37372 (CALI). Kottayam Dist.: Pulluparai – Peermade, 500 m, 24 Sep 1964, K. Vivekananthan 21368 (MH); Vaguman, 1100 m, 17 Jun 2002, Prasanth Kumar M.G. 86360 (CALI); between Theekoy and Erattupetta, 17 Oct 1976, J.K. Manglay 6764 (CALI). Kozhikode Dist.: Kannoth, 8 Dec 1913, collector unknown 9481 (MH). Palakkad Dist.: Perambicolam [Parambikulam], 1050 m, Nov 1909, A. Meebold 73/12467 (K [K001222304]). Palghat Dist.: Silent Valley, 25 Apr 1980, V.J. Nair 67300 (CAL). Pathanamthitta Distr.: Angamoozhy forest, 250 m, 19 Aug 1988, R. Chandrasekaran 89210 (MH). Thiruvananthapuram Dist.: Kottur R.F., 200 m, 26 Sep 1973, J. Joseph 41986 (MH); Ponmudi, 1100 m, 16 Aug 1980, M. Mohanan 69228 (MH). Thrissur Dist.: Athirapally, Charpa, 300 m, 2 Jul 1994, N. Sasidharan 618A & 618B (FRLH). Wayanad Dist.: Thirunelly, s.d., [M. Sabu] 37351 (CALI). Tamil Nadu. Coimbatore Dist.: Topslip, Ulandy, 775 m, 4 Sep 1994, V.S. Ramachandran 17602 (FRLH). Nilgiri Dist.: Agnibetta, Mudumalai Tiger Reserve, 10 Jun 2012, B. Pushpakaran 449 (Karpagam University Herbarium). Tirunelveli Dist.: Mundomurlii, 26 Aug 1913, Calder & Ramaswami 183 (CAL, DD); Karippanthode, 28 Aug 1913, M. Rama Rao 1512 (CAL); Palaumi, 10 Sep 1913, M. Rama Rao 1751 (K [K001222303]). Exact location uncertain: Canara & Mysore, Herb. Ind. Or. Hook. f. & Thomson, Stocks & Law [Mr. Law] 15? (K [K001222309]). Version of Record 13