A revision of Gomphostemma (Lamiaceae)

Summary

The genus Gomphostemma Wall. ex Benth. is revised. Thirty-one species are recognised with three new species described: G. flexuosum Bongch. from north-eastern Thailand, southern Laos and central Vietnam, G. longipetalum Bongch. and G. repentum Bongch. from northern Myanmar. Thirty names are lectotypified. A key to species, distribution maps and conservation categories for all species are provided.

Introduction

Gomphostemma Wall. ex Benth. (subfamily Lamioideae, Lamiaceae) is found in tropical evergreen forest throughout India, China, Myanmar, Thailand, Laos, Vietnam, Cambodia, Malaysia, Indonesia, Brunei and the Philippines. This revision is the first taxonomic work since Prain (1891) to consider the genus throughout its distribution. Previous taxonomic studies of Gomphostemma based on political boundaries have limitations: widely distributed species, such as G. javanicum (Blume) Benth., G. parviflorum Wall. ex Benth. and G. strobilinum Wall. ex Benth., often have continuous variation in morphological characters when examined throughout their distribution but can have discontinuities in variation when viewed only in particular countries or states. This has led to inconsistent treatments of Gomphostemma in different regionally-based taxonomic treatments. Since Prain’s revision of 25 species (1891), the number of Gomphostemma species described has risen to 90 (Govaerts et al. 2018), of which we consider almost two-thirds are at unaccepted infraspecific ranks or forms of varieties: we recognise 31 species here.

Taxonomic history

The genus name Gomphostemma, derived from Greek, gompho- (nail) and -stemma (garland) (Short & George 2013), first appeared in the Wallich Catalogue in 1829 and included nine names which are invalid because they lacked diagnostic descriptions (Wallich 1829). Gomphostemma was first formally described by Bentham (1830a). Prior to formal description, the earliest record of a species mentioned by Bentham (1830a) used the name Prasium melissifolium Roxb., originally listed by Roxburgh (1814). In 1826, Blume described two new species from Java, Prasium javanicum Blume and P. phlomoides Reinw. ex Blume (Blume 1826). Later, these names were transferred to the genus Gomphostemma as synonyms of G. javanicum (Bentham 1835). The confusion between Gomphostemma and Prasium L. (both subfamily Lamioideae), was probably caused by their similar corolla shape and fleshy nutlets. Bentham (1830b) further revised Gomphostemma and published 11 taxa based on specimens in Wallich’s Catalogue. Bentham (1835, 1848) added further new species and finally provided a generic level account as part of a generic level treatment of the Lamiaceae (Bentham 1876).

The most complete monograph of the genus was published by Prain (1891) who recognised 25 species and eight varieties, and divided the genus into three. Prain did not specify a term for his infrageneric divisions but, to comply with the International Code of Nomenclature for Algae, Fungi and Plants (ICN) (Turland et al. 2018), subgeneric rank will be applied in this revision. This classification was based on corolla and nutlet characters: subgen. Pogosiphon Prain with an incurved corolla in which the inside of the corolla tube is hirsute, corolla lobes that are broad and 1 or 2 nutlets frequently abortive per flower; subgen. Stenostoma Prain in which the corolla is straight and the throat is narrower than other groups, corolla lobes that are narrow and rarely more than one nutlet develops per flower; and lastly subgen. Gomphostemma [Eugomphostemma — corrected according to ICN (Turland et al. 2018)], where the corolla tube and lobes are similar to subgen. Pogosiphon but the inside of the tube is glabrous and usually with four developed nutlets (Prain 1891).

Taxonomic efforts since the baseline work of Prain and Bentham in the 19^th Century have focused on particular geographic areas: these are summarised in Appendix 1, a list of species names, the areas in which they occur and the accounts in which they have been treated.

Phylogenetic studies

The genus Gomphostemma belongs to tribe Gomphostemmateae Scheen & Lindqvist, subfam. Lamioideae (Scheen et al. 2010; Bendiksby et al. 2011; Xiang et al. 2013; Li et al. 2016). The tribe was first described by Scheen et al. (2010) who included three genera, Gomphostemma, Chelonopsis Miq. and Bostrychanthera Benth., the latter subsequently reduced into Chelonopsis by Xiang et al. (2013). Li et al. (2016) used chloroplast DNA to reconstruct the phylogeny of the family with the result highly congruent to that of Scheen et al. (2010) and Xiang et al. (2013). However, an analysis with richer sampling of the Gomphostemmateae (Bongcheewin 2013) suggested the tribe was paraphyletic, with other genera of Lamioideae nested inbetween Gomphostemma and Chelonopsis, the monophyletic Gomphostemma being sister to a large Synandreae-Leucadeae clade which contains Chelonopsis. Roy & Lindqvist (2015) also found that a chloroplast DNA phylogeny supported the sister relationship between Gomphostemma and Chelonopsis but their phylogeny, based on pentatricopeptide repeat (PPR) nuclear DNA sequences, did not support the sister relationship but rather the two genera resolved as a grade with Chelonopsis as sister to the remaining Lamioideae. Further work is required to resolve the relationships of Gomphostemma more robustly.

Previous phylogenetic analyses focused on relationships within Gomphostemma, based on chemical and morphological characters, have not supported Prain’s infrageneric classification (Bongcheewin et al. 2014). In particular, subgenera Gompostemma and Stenostoma are not monophyletic. Although subgenus Pogosiphon is monophyletic and can be diagnosed by the presence of an annulate corolla, the sister group of this clade, which contains species of subgenera Gomphostemma and Stenostoma, cannot be diagnosed with a morphological apomorphy.

Taxonomic approach in this account

Here, we classify Gomphostemma into two informal groups based on the presence of an annulate corolla (Table 1): I) Glabrous group and II) Annulate group. The Glabrous group corresponds to Prain’s subgenera Gomphostemma and Stenostoma and contains 24 species. The Annulate group corresponds to Prain’s subgenus Pogosiphon and contains seven species (species no. 25 – 31 in the account).

Table 1. A list of informal groups of Gomphostemma recognised in the current revision.

Within the Glabrous group there are three informal subgroups based on corolla characters: i) narrow throat with a straight tube (species no. 1 – 9); ii) inflated throat with an incurved tube (species no. 10 – 23) and iii) uncertain: G. microcalyx Prain (species no. 24) has a variable corolla and could fit into either i) or ii).

The key to the species of Gomphostemma does not follow the informal groups, which rely on the corolla characters, to allow for an absence of flowering material.

Within the Annulate group the placement of Gomphostemma pedunculatum Benth. ex Hook.f. is problematic as the presence of an annulate corolla is variable in the species. Preliminary molecular phylogenetic work based on chloroplast (trnL-F, rpL32F-trnL(UAG)) and nuclear (ITS, ETS) markers (Bongcheewin 2013) suggests placement in the Annulate group, as presented here.

We have not formally recognised these groupings within Gompostemma due to the difficulty of placing problematic species such as G. pedunculatum and G. microcalyx.

Materials and Methods

The taxonomic study was based on herbarium collections which included c. 1700 specimens from the following herbaria: AAU, ABD, BCU, BK, BKF, BM, BO, C, CAL, CMUB, E, G, GH, HITBG, HN, IBSC, K, K-W, KEP, KUN, L, NY, OXF, P, PE, PNH, QBG, SING, SLR, TAI and TCD (Thiers 2019, continuously updated) and field observations in Thailand between September 2010 – January 2011 for Gomphostemma javanicum, G. parviflorum, G. strobilinum and G. pedunculatum. All specimens have been seen by the first author unless indicated otherwise. Acronyms of herbaria mentioned in Wu (1965), which are different from Thiers (2019, continuously updated), were checked from Fu (1993) as needed. The specimens represent complete species distributions as far as possible. Specimens examined are listed alphabetically by collector in Appendix 2.

Morphological terms for plant description follow Beentje (2012). Qualitative characters, e.g. colour, and quantitative characters, e.g. plant height, were taken from field observations and herbarium labels when necessary.

Collecting localities appear as written on herbarium labels: spelling of localities may be slightly varied, especially for historic collections. Any corrections and additions to localities are put in square brackets in the ‘Specimens Examined’ section below. Collections were georeferenced as much as possible. When coordinates were not available from herbarium labels, they were gathered from the Fuzzy Gazetteer (http://isodp.hof-university.de/fuzzyg/query) and Google Earth. Literature providing plant collecting localities was also consulted (e.g. Hamidah et al. 2011; Jacobs 1962). Coordinates were presented in decimal degrees. The online Wallich Catalogue (http://wallich.rbge.info/) was used to check the distribution of Wallich specimens.

Distribution maps were produced by using the SimpleMappr programme: http://www.simplemappr.net/. Conservation assessments were carried out using GeoCAT: http://geocat.kew.org/ (Bachman et al. 2011) and the IUCN Red List Categories and Criteria: Version 3.1 (IUCN 2012). Google Earth imagery was also checked. The assessments were based mostly on criterion B of the IUCN categories and criteria using extent of occurrence (EOO) and area of occupancy (AOO) based on a grid size of 2 km width plus evidence of, or inferring, decline in the habitat (Willis et al. 2003; IUCN 2012).

Morphology

General morphology

Gomphostemma species vary from annual to perennial herbs. Perennial herbs have woody stems at the base or woody rootstocks and are sometimes recorded as shrub-like or sub-shrubs.

Typically, Gomphostemma has a fibrous root system, although some perennial species have rootstocks. Tuber-like rhizomes are found only in G. mastersii Benth. ex Hook.f. (Fig. 1A) and G. ovatum.

Fig. 1

A – B Gomphostemma mastersii: A tuber-like roots; B corolla with inflated throat. C G. parviflorum: axillary inflorescence and corolla with narrow throat. D G. javanicum: fleshy nutlets. E – F G. leptodon: E axillary inflorescence; F flower, note calyx lobes longer than the tube and corolla with inflated throat. photos: a – b & d thammarat phutthai; c bhanubong bongcheewin; e – f maxim nuraliev.

Most species have deep or shallow longitudinal grooves on the faces of the quadrangular stems. Older stems are likely to have deeper grooves. Gomphostemma sulcatum C.Y.Wu was named on account of the longitudinal grooves on its stems [sulcatum, grooved], but this character is rather common in this genus.

Stems are typically erect, but a few species have decumbent stems, curving with wavy internodes, i.e. Gomphostemma aborense Dunn, G. chinense Oliv., G. curtisii Prain, G. flexuosum Bongch. (Fig. 2), G. melissifolium Wall. ex Benth., G. nutans Hook.f., G. repentum Bongch. (Fig. 3) and G. velutinum Benth. Stems rooting at lower nodes are found in G. curtisii, G. mastersii and G. ovatum Wall. ex Benth. These species have decumbent stems but G. parviflorum (Parnell et al. 95-156 (K)) has erect stems which root at the nodes. Stems are highly reduced in one form of G. strobilinum, which was previously recognised as a separate species, G. acaule Kurz ex Hook.f. [a-, without; -caule, stem] (Hooker 1885), the height of stem is variable throughout the range of distribution of G. strobilinum (Fig. 4).

Fig. 2

Gomphostemma flexuosum. A habit; B stellate hairs with a 1-celled stalk from lower leaf surface; C upper leaf surface indumentum; D flower, lateral view; E glandular hairs with a multi-celled stalk from corolla; F stellate hairs with a 1-celled stalk; G dissected calyx showing the outer surface on one lobe and the inner surface on the four others; H dendroid hair with a multi-celled stalk and stellate hair with a 1-celled stalk from calyx. All from Suddee et al. 955. drawn by andrew brown.

Fig. 3

Gomphostemma repentum. A habit; B stellate hair from lower leaf surface; C upper leaf surface indumentum showing 3-celled simple hairs and 1-celled simple hairs; D flower, lateral view; E calyx cut and opened to show ovary, style and stigma; F corolla cut and opened as flat as possible to show stamens and an annulus of simple hairs; G simple hairs from the annulus. All from Micholitz s.n. drawn by andrew brown.

Fig. 4

Morphological variation of Gomphostemma strobilinum. A from Larsen & S Larsen 34166; B from Sørensen et al. 5945b; C from Sørensen et al. 1053; D from van Beusekom et al. 3856; E from Suddee et al. 889; F from Hennipman 3183; G from Chantaranothai et al. 90/57. drawn by andrew brown.

Indumentum

Trichomes on stems, leaves and calyx can be seen using a 10× hand lens. There are three types of trichomes found in Gomphostemma: simple, stellate and dendroid, occurring as only one type or mixed. The term ‘tomentose’ used in the revision refers to any kind of hairiness. Simple 1 – 5-celled hairs are common.

Stellate hairs, where the branches radiate from the same point, can have a 1-celled stalk (Fig. 5U) or sessile hairs, branched from the base. Stellate hairs with a multi-celled stalk are found in Gomphostemma dolichobotrys Merr., G. flexuosum, G. pedunculatum, G. stellatohirsutum C.Y.Wu, G. hirsutum Walsingham and some collections of G. javanicum (Fig. 5T). They are easily visible using a 10× hand lens.

Fig. 5

Morphological variation of Gomphostemma javanicum. A – G leaves, A from Ridley s.n.; B from Ridley s.n.; C from Edaño 17836; D from Middleton et al. 1507; E from Burkill & Holttum 8650; F from Craib 733; G from Tsang 27020. H – P calyces, H from Henry 12011; J from Craib 733; K from Middleton et al. 1507; L from Tsang 27020; M from Larsen et al. 30855; N from Kerr 2054; P from Gardner & Khumchompoo ST 2117. NB. 4 outer surfaces area and 1 inner surface (on right) shown for each calyx. Only 1 outer surface (on left) showing indumentum, other 3 outer surfaces without hairs for clarity. Inner surface shows indumentum. Q – S verticils, Q from Hooker & Thomson s.n.; R from Middleton et al. 1507; S from Burkill & Holttum 8650. T – U type of hairs, T stellate hairs with a multi-celled stalk, from Gardner & Khumchompoo ST 2117; U stellate hairs with a 1-celled stalk, from Kerr 21078. drawn by andrew brown.

Dendroid hairs, where the branches are separate like the branches of a tree and with a base normally more than 1-cell long, are found in Gomphostemma dolichobotrys, G. heyneanum Wall. ex Benth., G. pedunculatum and G. strobilinum and some collections of G. crinitum Wall. ex Benth. Bifurcate hairs, which have one branch at the tip, are found sometimes, but this type has no particular pattern and is not useful for species circumscription in Gomphostemma. However, bifurcate hairs are informative for the infrageneric classification of the related genus, Chelonopsis, i.e. subgen. Chelonopsis and subgen. Aequidens (Xiang et al. 2010).

The density of indumentum on upper- and lower-side of leaves is normally different. On the upper side, the indumentum is much less dense. In all Gomphostemma, stellate hairs are found on both sides, except for the upper side of G. dolichobotrys and G. ovatum which have only simple hairs.

The colour of the leaf indumentum has been used for the delimitation of some species, Gomphostemma niveum Hook.f. and G. hainanense C.Y.Wu are whitish-tomentose on the lower-side. However, the colour of indumentum varies, for example the indumentum of G. javanicum can be golden yellow to ash grey.

Leaf morphology

Most leaves of Gomphostemma are petiolate except for G. flexuosum and G. niveum which have subsessile leaves. Petioles can be up to 150 mm long. Leaf shape is variable, e.g. G. strobilinum (Fig. 4) and G. javanicum (Fig. 5). Variegated leaves occur in some collections of G. strobilinum and G. pedunculatum. Leaf margins are generally variable but G. eriocarpon Benth. uniquely has a dentate margin with a glandular thickening at the tip of the serration, rather than the typical serrate margin. Some collections of G. javanicum and G. microcalyx have a subentire or shallowly serrate margin. The upper side has mostly darker colour than the lower.

Inflorescence

As is common in the Lamiaceae, the inflorescence is an indeterminate thyrse. Flowers are borne on opposite cymes usually subtended by bracts that are typically indistinguishable from the leaves, although they can be much smaller in species with congested inflorescences. Within the cymes the flowers are subtended by small bracteoles. The inflorescence is usually axillary (Fig. 1B & C) but can be terminal (Fig. 4) or basal. Cymes in the inflorescence are usually laxly disposed, but can be congested or very congested, giving the appearance of a raceme-like spike. Within cymes, the arrangement can be congested or lax, branched from the base or simple, and pedunculate or not. There is variation in length, the number of flowers per verticil and the density of inflorescence. The Glabrous group has axillary and basal inflorescences, but not terminal. Positions of inflorescences in the Annulate group can be terminal, axillary, or terminal and axillary. Terminal inflorescences are upright (Gomphostemma heyneanum, G. nutans, G. strobilinum).

Bracts

Bracts subtending the cymes are taxonomically informative for some Gomphostemma species. Gomphostemma aborense, G. hainanense, G. niveum, G. ovatum, G. stellatohirsutum and G. sulcatum have a cluster of small, leaf-like bracts subtending cymes. Large bracteoles within the cyme can exceed the calyx in G. curtisii, G. microdon Dunn and G. parviflorum. Other species have narrow and inconspicuous bracteoles. Bracteoles of G. curtisii are sometimes asymmetric at the base.

Prain (1891) recognised Gomphostemma wallichii Prain as a separate species from G. strobilinum due to its cordate bracts. However, bract and leaf shape are very variable and there is no clear discontinuity in shape within this group.

Calyx

The calyx is gamosepalous, 5-lobed. Prain (1891) used calyx lobe length, calyx indumentum and the presence of calyx ridges to distinguish some species. The relative length of calyx lobe and tube can help specific identification in the closely related species Gomphostemma crinitum, G. microcalyx, G. parviflorum and G. thomsonii Benth. ex Hook.f. (Fig. 6). However, calyx lobe length can be variable within a species: in G. javanicum the length of the calyx lobe varies continuously throughout its range (Fig. 5).

Fig. 6

Floral morphology of Gomphostemma crinitum, G. microcalyx, G. parviflorum and G. thomsoni. A – E flowering calyx, corolla and lower lip. A G. crinitum (Robinson s.n.); B G. parviflorum (from Garrett 997); C G. microcalyx [straight corolla form] (from Yii & Othman S46181); D G. microcalyx [incurved corolla form] (from de Wilde et al. 16949); E G. thomsonii (from Hooker & Thomson s.n. (holotype)). F – J fruiting calyx, dissected, drawn to show the outer surface on one lobe and the inner surface on the four others. F G. crinitum (from Lim FRI52830); G G. parviflorum (from Boyce 1026); H G. microcalyx (from Anderson S20955); J G. thomsonii (from Hooker & Thomson s.n. (isotype)). drawn by andrew brown.

Calyx indumentum was used by Prain at subgeneric level but this character is clearly affected by the degree of maturity of the calyx. The indumentum of the flowering and fruiting calyx is frequently different. Hence, referring to calyx characters, it is necessary to specify clearly either flowering or fruiting calyx when discussing the calyx indumentum.

Typically, Gomphostemma species have a 10-ridged calyx but the prominence of calyx ridges can vary. Gomphostemma javanicum has prominent ridges (Fig. 5) which can assist recognition of this species.

Corolla

The corolla is tubular and relatively large. In most species the corolla tube is exserted from the calyx and inflated. The presence or absence of a hairy annulus inside the corolla tube was used for subgeneric recognition by Prain (1891). The corolla lobes are 2-lipped with posterior lip 3-lobed and anterior lip 1-lobed.

The various colours of the corolla, including white, yellow, red, orange, blue and purple, are not taxonomically useful because colours can vary within the same species: colour pigments can be altered by ecological factors, e.g. acidic pH (Asen et al. 1972), UV light (Awad et al. 2000). For example, the corolla of Gomphostemma strobilinum can be yellow, creamy-white or red in natural habitats.

The stamens, ovary, style, stigma and disk, have no taxonomic value for the genus Gomphostemma as they have a very consistent morphology.

Pollen

Pollen grains of Gomphostemma are tricolpate but variable between species in shape and size and the exine sculpturing. Pollen characters do not support the subgeneric classification sensu Prain (1891) (Abu-Asab & Cantino 1994; Xiang et al. 2012) or the groups presented here.

Nutlet morphology

Generally, the nutlets of Gomphostemma are fleshy when fresh (Fig. 1D). No means of dispersal has been reported in the literature. Nutlet sculpture is unreliable when dry. Sometimes nutlets are covered by hairs but this character seems to vary with maturity. These observations support Ryding (1994) who suggested that hairs on the nutlets are variable and less informative for determining relationships between genera related to Gomphostemma (Ryding 1994). The number of nutlets was used to support infrageneric classification by Prain (1891) but this character is inconsistent, e.g. G. hemsleyanum Prain ex Collett & Hemsl. and G. strobilinum are described as having 2 – 3 mature nutlets, but 4 mature nutlets have been found. Nutlet shape can be oblong, ovate or globose. Nutlet colour varies in dried and fresh nutlets. From field observations in Thailand, nutlet colour may be useful. Gomphostemma parviflorum, G. javanicum and G. strobilinum have creamy-white, white and orange nutlets respectively. However, nutlet colours of herbarium specimens are always black or dark brown.

Chemistry and use

Few chemical studies of Gomphostemma have been done and these focus mainly on antimalarial activity (Zheng & Xing 2009; Acharya et al. 2008; Wang et al. 2007; Sathe & Kaushik 2009; Zhang et al. 2009; Dutta 2014). Bongcheewin et al. (2014) investigated flavonoid and phenolic acid compounds in Gomphostemma for the first time, concluding that chemical profiles of Gomphostemma are not informative for taxonomic classification due to their variable characters within species (see Bongcheewin et al. 2014).

The uses of Gomphostemma have been documented in several sources. A decoction of leaves of G. niveum is used for the treatment of tuberculosis, malaria fever, or insect bite in India (Dutta 2014). In Hainan Island, China, local healers use leaves of G. lucidum Wall. to treat sores (Zheng & Xing 2009). In the northern Andaman Islands, crushed leaf juice of G. javanicum is used for skin diseases (Rama Chandra Prasad et al. 2008). In northern Thailand, local healers use whole plants and roots of G. strobilinum to alleviate fatigue and to cure abscesses, respectively (Chuakul & Saralamp 2000; Chuakul et al. 1996). Gomphostemma crinitum and G. niveum have been used for the treatment of malaria in India (Acharya et al. 2008; Sathe & Kaushik 2009). Gomphostemma microdon has been recorded as a drug for use in menoxenia and as an antitussive drug (Zhang et al. 2009).

Distribution and conservation

Gomphostemma occurs in tropical areas of India, the eastern Himalaya, China, Southeast Asia and the Malesian region. The greatest number of species is in the eastern Himalaya with 14 species in total, six of which are endemic, namely G. aborense, G. longipetalum Bongch., G. melissifolium, G. niveum, G. ovatum and G. velutinum, This species diversity may be explained by the corresponding diversity of vegetation types (Singh & Singh 1987). The most widely distributed species is G. javanicum, found almost throughout the entire range of the generic distribution except South India. The Annulate group are only found in the continental area.

Most Gomphostemma grow in evergreen forests which are under threat because of human activities such as agriculture, industry or hydropower projects (Sodhi et al. 2010; Rao et al. 2013). A preliminary conservation assessement of each species is given in the account. One species (G. repentum) is Critically Endangered, four are Endangered, three are Vulnerable, 16 are of Least Concern and seven are Data Deficient.

Taxonomic treatment

Gomphostemma Wall. ex Benth. (Bentham 1830a: sub t. 1292); Bentham (in Wallich 1830b: 12, 1835: 646, in A. P. de Candolle 1848: 550); Hooker (1885: 696); Prain (1891: 245); Doan (1936: 1030); Mukerjee (1940: 203); Keng (1969: 79, 1978: 319); Murata (1971: 499, 1976: 188); Li & Hedge (1994: 70); Phuong (2000: 97); Khanam et al. (2003: 2); Walsingham & Bramley (2010: 479); Walsingham (2019: 194). Type: Gomphostemma javanicum (Blume) Benth, designated here. Bentham (1830a) did not nominate a type, nor has any subsequent author. Wallich (1829) clearly enumerated nine species of Gomphostemma. Bentham (1830a) indirectly included those and stated in the protologue of this genus “Besides the species enumerated by Dr. Wallich in his Catalogue of the East India Company’s collection, Prasium javanicum and phlomoides Blume should probably be referred to this genus”. Bentham transferred these species to Gomphostemma in 1835 (Bentham 1835). Therefore, the type species has to be chosen from 11 taxa as indicated by Bentham (1830a) but only two names were specifically mentioned by him. Gomphostemma javanicum has been selected because only this taxon was mentioned specifically when he validated the genus (Bentham 1830a), G. phlomoides being a synonym of G. javanicum. The general characters, indumentum, corolla and calyx morphologies and fleshy nutlets of this species are easy to recognise and relate directly to the protologue of Gomphostemma. This choice is also consistent with Prain’s infrageneric classification as G. javanicum belongs to his subgenus Eugomphostemma [Gomphostemma]. In the absence of any discussion by Wallich, this is a non-disruptive and conservative lectotypification.

Taitonia Yamam. (Yamamoto 1938: 277). Type: Taitonia callicarpoides Yamam.

Perennial, shrub-like, upright, decumbent or sprawling herbs. Stems erect, ascending or sprawling, sometimes rooting at nodes or woody at base, obtusely quadrangular, with creamy-white, coppery or pale yellow dendroid or stellate hairs. Leaves petiolate or subsessile, spreading, chartaceous or coriaceous; blades ovate, elliptic, lanceolate, subglobose, cordate, oblong, obovate or oblanceolate; apex acute or round; margin serrate or dentate; base cuneate, attenuate, cordate or obtuse; upper side with indumentum less dense than lower side; lower side with densely arranged stellate hairs with a 1-celled or multi-celled stalk or dendroid hairs with a multi-celled stalk, and usually with 1 – 4-celled simple hairs or sometimes bifurcate hairs. Inflorescence an indeterminate thyrse, axillary, terminal, terminal and axillary, or basal, lax, elongated or congested, unbranched or branched; verticils few – many-flowered; cymes, sessile or pedunculate; bracts persistent or deciduous, lanceolate, ovate, obovate, elliptic, oblanceolate, rhomboid, sometimes leaf-like; bracteoles linear, narrowly elliptic or narrowly ovate; florets pedicellate to sessile. Flowering calyx infundibular or tubular, outside tomentose, inside tomentose or glabrous; lobes equal or subequal, subulate, triangular, acute, or acuminate. Fruiting calyx cupular, campanulate or infundibular. Corolla white, creamy-white, yellow, orange, whitish-purple or scarlet, throat inflated or narrow; tube straight or incurved; lobes 2-lipped, spreading or scarcely so. Stamens included or slightly exserted from posterior lip, lower pair longer than upper pair, attached below the throat; filaments more-or-less fleshy, sometimes tomentose; anthers 2-celled, elliptic-oblong, 1 – 2 mm long, inclined under posterior lip, dorsifixed, longitudinally dehiscent. Disk lobed, tumescent behind. Style gynobasic, slender, glabrous, inclined under posterior lip; stigma apex bifid, subequal- or unequal-lobed. Nutlets 1 – 4, oblong, globose, elliptic or obovate, fleshy when fresh, smooth or wrinkled, glabrous or tomentose.

Key to the species of Gomphostemma

• 1. Inflorescence axillary, or basal with opposite cymes or short raceme-like flowering shoots without leaves borne laterally to the main leafy stem2

• 1. Inflorescence terminal at apex of leafy stem axis or terminal and axillary31

• 2. Inflorescence basal, opposite cymes inserted at the lower nodes from which leaves have mostly fallen3

• 2. Inflorescence axillary, opposite cymes inserted at the upper nodes at which leaves are usually present5

• 3. Stems tomentose with dendroid hairs and stellate hairs with a multi-celled stalk29. G. pedunculatum

• 3. Stems tomentose with stellate hairs with sessile or a 1-celled stalk, dendroid hairs absent...4

• 4. Calyx lobes with subulate apex; corolla 30 – 45 mm long; petioles 30 – 60 (– 90) mm long. Malay Peninsula, Sumatra, Borneo11. G. curtisii

• 4. Calyx lobes with acuminate apex; corolla 30 – 60 mm long; petioles 10 – 40 mm long. S China, Taiwan, N Vietnam10. G. chinense

• 5. Corolla tube with annulus of simple hairs inside6

• 5. Corolla tube glabrous inside8

• 6. Opposite cymes of inflorescence congested25. G. hemsleyanum

• 6. Opposite cymes of inflorescence elongated, raceme-like7

• 7. Erect herbs; stems robust; inflorescence with axis visble between cymes...27. G. microdon

• 7. Climbing herbs; stems slender; inflorescence with axis not visible between cymes30. G. repentum

• 8. Opposite cymes of inflorescence unbranched, elongate, raceme-like, lax with axis visible between flowers3. G. dolichobotrys

• 8. Opposite cymes of inflorescence branched then lax, or if unbranched then congested with axis not visible between flowers 9

• 9. Corolla throat inflated, tube abruptly dilated near the throat; nutlets 1 – 410

• 9. Corolla throat narrow, not enlarged, sometimes gradually dilated towards the throat; nutlets solitary23

• 10. Mature leaves whitish or creamy-white tomentose on the lower side, strongly discolorous . . . . 14. G. hainanense

• 10. Mature leaves pale green tomentose on the lower side, not or weakly discolorous11

• 11. Erect herbs up to 0.4 m or sprawling herbs12

• 11. Erect herbs 0.4 – 3 m, not sprawling16

• 12. Erect herbs up to 0.4 m; stems upright, sometimes flaccid13

• 12. Sprawling herbs; stems scrambling; internodes of leafy and flowering shoots with curving stem appearing wavy, always twisting15

• 13. Lower side of leaves dense tomentose; roots not tuber-like23. G. velutinum

• 13. Lower side of leaves sparse tomentose; roots tuber-like14

• 14. Upper side of leaves with sessile stellate hairs and 2 – 4-celled simple hairs all over; bracts deciduous . . . . . . . .18. G. mastersii

• 14. Upper side of leaves with 2 – 4-celled simple hairs, sometimes with sparse stellate hairs only on lower part of midrib; bracts persistent20. G. ovatum

• 15. Corolla 30 – 50 mm long; upper side of leaves tomentose with sessile stellate hairs or sparsely so to almost glabrous19. G. melissifolium

• 15. Corolla 15 – 20 (– 23) mm long; upper side of leaves scabrid with stiff hairs12. G. flexuosum

• 16. Flowering calyx densely covered with long simple hairs and long branched hairs15. G. hirsutum

• 16. Flowering calyx densely covered with sessile stellate or stellate hairs with a 1-celled stalk and/or short simple hairs17

• 17. Opposite cymes subtended by the upper leaves and cluster of small leaf-like bracts18

• 17. Opposite cymes subtended by the upper leaves only19

• 18. Stems densely covered by stellate hairs or dendroid hairs with a multi-celled stalk; upper side of leaves dense tomentose with 3 – 4-celled simple hairs, sometimes bifurcate hairs21. G. stellatohirsutum

• 18. Stems densely covered by stellate hairs with a 1-celled stalk, dendroid hairs absent; upper side of leaves tomentose with 1 – 2 (– 3)-celled simple hairs22. G. sulcatum

• 19. Fruiting calyx lobes longer than the tube at maturity20

• 19. Fruiting calyx lobes not longer than the tube at maturity21

• 20. Petioles 50 – 150 mm long; leaf margin serrate17. G. leptodon

• 20. Petioles 10 – 50 mm long; leaf margin subentire or obtusely serrate16. G. javanicum (1)

• 21. Margins of fruiting calyx lobes revolute in cross-section13. G. grandiflorum

• 21. Margins of fruiting calyx lobes flat in cross-section22

• 22. Flowering calyx 7 – 15 mm long; opposite cymes branched then lax or unbranched then congested with axis not visible between flowers, exceeding 20 mm wide when pressed, excluding corolla . . . . . 16. G. javanicum (2)

• 22. Flowering calyx 5 – 8 mm long; opposite cymes unbranched then very congested with axis not visible between flowers, up to 20 mm wide when pressed, excluding corolla24. G. microcalyx (2)

• 23. Mature leaves whitish or creamy-white tomentose on the lower side, strongly discolorous7. G. niveum

• 23. Mature leaves pale green tomentose on the lower side, not or weakly discolorous24

• 24. Bracts leaf-like, broadly ovate or broadly elliptic-ovate, 9 – 15 mm wide1. G. aborense

• 24. Bracts rhomboid, elliptic, elliptic-ovate, ovate, lanceolate, ovate-lanceolate, obovate, falcate, 2 – 8 mm wide . . . . . 25

• 25. Corolla tube included in the calyx5. G. keralense

• 25. Corolla tube exserted from the calyx26

• 26. Leaf margin dentate or dentate-serrate with glandular thickening at the tip of serration; flowering calyx glabrous inside4. G. eriocarpon

• 26. Leaf margin serrate, shallowly serrate or subentire, without glandular thickening at the tip; flowering calyx tomentose inside27

• 27. Flowering calyx 5 – 8 mm long; opposite cymes very congested, up to 20 mm wide when pressed, excluding corolla24. G. microcalyx (1)

• 27. Flowering calyx 8 – 20 mm long; opposite cymes congested or lax, exceeding 20 mm wide when pressed, excluding corolla28

• 28. Corolla 40 – 50 mm long6. G. longipetalum

• 28. Corolla 15 – 35 mm long29

• 29. Bracts lanceolate or elliptic-ovate, 2 – 3 mm wide, narrower than the calyx2. G. crinitum

• 29. Bracts ovate, ovate-lanceolate or obovate, 4 – 8 mm wide, broader than the calyx30

• 30. Petioles 20 – 40 mm long8. G. parviflorum

• 30. Petioles 50 – 140 mm long9. G. thomsonii

• 31. Plants climbing; stems slender, scrambling. . 28. G. nutans

• 31. Plants not climbing; stems robust, erect. . 32

• 32. Corolla tube 8 – 12 mm long, included in the calyx. . 26. G. heyneanum

• 32. Corolla tube 15 – 25 mm long, exserted from the calyx. 31. G. strobilinum

I) Glabrous group: i. corolla with straight tube and narrow throat (species no. 1 – 9).

1. Gomphostemma aborense Dunn (1920: 135); Mukerjee (1940: 210). Type: India, Eastern Himalaya, Arunachal Pradesh, outer Abor Hills, Janakmukh, at the mouth of the Yamne near Pongging, very common, 19 Dec. 1911, Burkill 37269 (holotype K [K000846334]).

Decumbent herb, up to 30 cm tall. Stems slender, ascending, sometimes rooting at nodes obtusely quadrangular with longitudinal grooves, with dense stellate hairs; internodes curving, always twisting. Leaves petiolate, chartaceous; blades broadly ovate, 5 – 8 × 3 – 5 cm, apex acute, margin serrate, base cuneate, with dense, stellate hairs on both sides, lower side pale green tomentose; petiolate 10 – 25 mm long. Inflorescence axillary with opposite cymes sessile, unbranched, congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils few-flowered; cluster of small leaf-like bracts, subsessile, persistent, broadly ovate or broadly elliptic-ovate, 10 – 20 × 9 – 15 mm, apex acute, margin serrate, base attenuate; bracteoles linear. Flowering calyx campanulate, 5 – 9 mm long, 10-ridged, outside with dense stellate hairs, inside dense simple hairs; tube 3 – 5 mm long; lobes triangular, 2 – 4 mm long. Fruiting calyx infundibular, 7 – 12 mm long; tube 4 – 6 mm long; lobes triangular, 3 – 6 mm long. Corolla yellow, 15 – 25 mm long, throat narrow, not enlarged, sometimes gradually dilated towards the throat, outside tomentose, inside glabrous; tube slender, straight, 15 – 20 mm long; posterior lip ovate-elliptic, 3 – 5 mm long, spreading, apex emarginate; anterior lip 3-lobed, spreading, slightly undulate, longer than posterior lip; lateral lobes bending downward. Staminal filaments glabrous. Style slender, 15 – 25 mm long, glabrous with apex subequally lobed. Nutlets solitary, obovate, 2 – 3 mm long, minutely glandular punctate.

distribution. India (Arunachal Pradesh). Map 1.

Map 1

Distribution range of Gomphostemma aborense, G. crinitum, G. dolichobotrys, G. eriocarpon and G. keralense.

specimens examined. arunchal pradesh. Nizamghat, 28°N 95°45'E, 29 Nov. 1928, Ward 8734 (K 2 sheets); outer Abor Hills, Janakmukh, 28.0619°, 95.2594°, at the mouth of the Yamne near Pongging, 19 Dec. 1911, Burkill 37269 (holotype K).

habitat. Open places, old, cultivated ground, secondary growth; alt. 300 – 900 m.

conservation status. Gomphostemma aborense is only known from two localities in India; no new collections have been made since 1928. Although the label of Burkill 37269 (K) mentioned that this species is very common, no new evidence to evaluate the decline of habitats is needed. The Data Deficient (DD) category is applied.

phenology. Flowering and fruiting in November – December.

note. Gomphostemma aborense is easily diagnosed by the curved internodes, stems appearing wavy and cymes subtended by upper leaves and clusters of small leaf-like bracts.

2. Gomphostemma crinitum Wall. ex Benth. (Bentham in Wallich 1830b: 12); Hooker (1885: 698); Prain (1891: 254); Mukerjee (1940: 209); Keng (1969: 84, 1978: 321); Murata (1976: 188); Li & Hedge (1994: 72); Phuong (2000: 103); Walsingham (2019: 196). Type: Burma, Tavoy [14.0333°, 98.2], 22 Oct. 1827, Gomez s.n. in Wall. Cat. 2159.2 (lectotype K [K000501085], selected here; isolectotype K-W [K001115348]).

Gomphostemma crinitum var. griffithii Prain (1891: 254). Type: Malaysia, Mt Ophir [Gunung Ledang, 2.3667°, 102.6167°], Chebu [Chabau] forest, Griffith 4042 (lectotype CAL [CAL0000007147], selected here; isolectotypes K [K000249839, K000249840], P).

Perennial herb up to 2.5 m tall. Stems erect, woody at base, obtusely quadrangular with longitudinal grooves, with dense stellate hairs and sometimes dendroid hairs. Leaves petiolate, chartaceous; blades elliptic, elliptic-obovate or elliptic-ovate, 20 – 33 × 6 – 11 (–14) cm, apex acute, margin shallowly serrate, base attenuate, upper side dark green with sparse stellate hairs, lower side pale green tomentose with dense stellate hairs; petioles 20 – 50 mm long. Inflorescence axillary with opposite cymes, unbranched then congested with axis not visible between flowers or rarely branched, exceeding 20 mm wide when pressed, excluding corolla, inserted at the upper nodes at which leaves are usually present; verticils few – many-flowered; bracts lanceolate or elliptic-ovate, 15 – 30 × 2 – 3 mm, narrower than the calyx, apex acute, margin entire, base cuneate, upper side with sparse stellate hairs, lower side with dense stellate hairs; bracteoles linear, 10 – 20 × 1 – 3 mm. Flowering calyx tubular, 12 – 15 mm long, 10-ridged, outside with stellate hairs, inside tomentose with adpressed simple hairs; tube 4 – 5 mm long; lobes triangular, 8 – 10 mm long, apex filiform. Fruiting calyx cupular, 15 – 20 mm long; tube 5 – 7 mm long; lobes 10 – 13 mm long. Corolla bright yellow, 20 – 30 mm long, throat narrow, not enlarged, sometimes gradually dilated towards the throat; tube straight, 15 – 23 mm long, exserted from the calyx, outside with stellate hairs, inside glabrous; posterior lip elliptic-ovate, 5 – 7 mm long; anterior lip 3-lobed, longer than posterior lip with median lobe ovate, 5 mm long and lateral lobes ovate, apex round. Staminal filaments slightly fleshy. Style slender, slightly shorter than corolla, glabrous with apex unequally lobed. Nutlets solitary, globose or ovate, 4 – 7 mm long, glabrous. Fig. 6A, F.

distribution. Bangladesh, China (Yunnan), Burma, Thailand, Peninsular Malaysia, Indonesia (Sumatra). Map 1.

specimens examined. bangladesh. Chittagong: Rangamati, Sitapahar, Kaptai [22.5139°, 92.2056°], 8 Oct. 1998, Uddin & Ullah 3558 (ABD). china. Yunnan: Mong-hain or Gan-lan-ba [22.4833°, 100.5833°], Che-li Hsien, 860 m, Oct. 1936, Wang 79805 (KUN). myanmar. Tanintharyi: Eastern Tenasserim [11.8°, 99.25°], 50 m, 30 May 1932, Kerr 21614 (BK, BM, K); Tavoy, 22 Oct. 1827, Gomez s.n. in Wall. Cat. 2159.2 (lectotype, K [Herb. Benthamianum]; isolectotype K-W); Tavoy, Zimba Valley, 22 Nov. 1924, Parker 2255 (K); Tenasserim, Tavoy, along the W bank of Paungdaw R., 3 miles E of Paungdaw Power Station, 240 m [800 ft], 22 Aug. 1961, Keenan et al. 1098 (E, K). thailand. Chiang Rai: Trail to Doi Jong Summit, 20°37'67”N 98°94'68”E, Nov. 2011, Norsaengsri & Tathana 7371 (QBG); Kanchanaburi: Kwae Noi River Basin, near Neeckey, near Wangka [15.1°, 98.4667°], 150 m, 2 June 1946, den Hoed 912 (BK, K, L, P); Thung Yai Naraesuan Wildlife Sanctuary [15.3333°, 98.9167°], Lai Wo, Ban Saneh Pawng, 200 m, 16 June 1993, Maxwell 93-626 (BKF, CMUB); Phetchaburi: Kaeng Krachan Distr., Kaeng Krachan National Park, on steep slope down from road on the way to Phanoen Thung Ranger Substation, 12°48'N 99°24'E, 560 m, 9 May 2005, Middleton et al. 3323 (BKF, E); Kaeng Krachan, near Ban Krang Camp, 18^th km, 12°48'N 99°26'E, 350 m, 27 June 2000, Newman et al. 1104 (AAU, BKF, E); Prachuap Khiri Khan: Kao Luang [Khao Luang, 11.6°, 99.6667°], 700 m, 4 July 1926, Kerr 10832 (ABD, BK, BM, L); Ranong: Ban Racha Krude, 9°44'N 98°37'E, 50 – 100 m, 7 July 1992, Larsen et al. 43200 (AAU, BKF); Klong Naka Wildlife Sanctuary, stream branch about 200 m down stream from base camp, left branch, flowing from NW, 9°30'N 98°20'E, 100 – 600 m, 9 Jan. 1990, Hoover et al. 5512 (E); Kraburi [Kra Buri, 10.4°, 98.7833°], Klawng Wa, 50 m, 23 Dec. 1928, Kerr 16310 (ABD, BK, BM 2 sheets, L); Muang Distr., Ranong Hot Springs, 9°57'25”N 98°39'11”E, 40 m, 23 June 2006, Williams et al. 2080 (E); Surat Thani: Phanom, Khlong Phanom National Park, ‘Big Tree’ nature trail, S of park headquarters, 8°52'N 98°40'E, 150 m, 26 Nov. 2005, Gardner et al. ST 1972 (E 2 sheets); Phanom, Ban Phong Lom, Khao Mok, Khao Sok National Park [8.8333°, 98.7°], 50 – 150 m, 29 Aug. 1982, Shimizu et al. T-28921 (BKF); Takuapah, Takuapah-Surat Thani Road, 20 – 60 km from Takuapah, 8°53'N 98°21'E, 100 – 250 m, 14 July 1972, Larsen et al. 30948 (AAU, BKF, K, L); Phangnga: Khao Khlong Yang at Khao Phra Mi, 9°17'N 98°26'E, 100 – 300 m, 6 July 1972, Larsen et al. 30654 (AAU, BKF, K, L, P); Krabi: Tambon Kao Panom [8.25°, 99.0333°], 100 m, 1 April 1930, Kerr 18804 (BK, BM 2 sheets); Nakhon Si Thammarat: Kao Luang [8.5°, 99.6667°], 200 m, 28 April 1928, Kerr 15424 (BK, BM 2 sheets); Lansaka, Kiriwong [8.4333°, 99.7833°], 27 Jan. 1951, Smitinand s.n. (BKF); Trang: Kao Chong, 7°40'N 99°45'E, 200 m, 13 June 1974, Geesink et al. 7167 (AAU, BKF, K, L); Nayong, Khao Chong, trail from botanic garden headquarters to Ton Yai, 7°33'N 99°47'E, 80 m, 9 July 2000, Middleton et al. 362 (BKF); Pattani: Betong [5.75°, 101.0333°], 300 m, 3 Aug. 1923, Kerr 7480 (BK, BM, K, L); Yala. Betong, Bang Lang National Park, Ban Chulaphon Phattana 10, near rattan plantation project, 5°51'N 101°18'E, 600 m, 19 July 2004, Pooma et al. 4447 (BKF); Nikom Kua Long, 600 m, 12 Feb. 1972, Santisuk & Nimanong 377 (BKF); Than To, Bang Lang National Park, along stream to Halasa Waterfall, 6°04'N 101°5'E, 135 m, 12 Feb. 2004, Middleton et al. 3000 (BKF, E); Narathiwat: Bukit [6.1667°, 101.8333°], 200 m, 8 July 1923, Kerr 7119 (BK, BM, K). malaysia. Johor: Bukit Jitan [2.0667°, 102.9833°], Nov. 1900, Ridley 11131 (SING); Woods below Mt Ophir [2.3667°, 102.6167°], June 1892, Ridley s.n. (SING); Mt Ophir [Gunung Ledang, 2.3667°, 102.6167°], Chebu [Chabau] forest, Griffith 4042 [type of G. crinitum var. griffithii, lectotype CAL; isolectotypes K 2 sheets, P]; Sungai Linggiu logged forest via Kluang [1.8139°, 103.6942°], 150 m, 25 July 1991, Tay et al. 91-35 (SING); Kedah. Kedah, Kroh [5.7°, 100.9833°], 120 m [400 ft], 6 Aug. 1941, Nauen 38046 (K 2 sheets, GH, SING); Malacca: [2.2°, 102.2833°], Alvins 2130 (SING); Negeri Sembilan: Pasoh Forest [3.0333°, 102.3333°], 18 April 1978, Ang FRI 27573 (GH, L); Bukit Linggung, Ulu Bendol [2.7333°, 102.1333°], 1 Dec. 1922, Holttum 9836 (SING); Genting Peras [3.1333°, 101.9667°], May, Ridley 7600 (SING); Kelantan: Aring FR, Anthonysamy SA 1094 (KEP); Pahang: Bentong, Genting Sempah [3.5333°, 101.9167°], 375 m [1250 ft], 18 Jan. 1959, Wyatt-Smith 93195 (K, L); Bukit Sadam [Bukit Serdam, 3.8333°, 101.9167°], 120 m [400 ft], 6 Oct. 1931, Henderson 25061 (BK, BM, SING); Kuala Tembeling [4.0667°, 102.3167°], 20 Aug. 1928, Holttum 20531 (SING); Kwala Jahau, Aug. 1891, Ridley 2185 (K); Pulau Manis [3.55°, 103.2167°], 1891, Ridley s.n. (SING); Ulu Sungai, Sepia [4.6108°, 102.8839°], Jeram Perahu, 150 m [500 ft], 18 July 1970, Shah & Noor 1965 (C, GH, L, SING); Sungai Yu [4.5008°, 102.0028°], 1 Feb. 1924, Nur & Forworthy 11922 (SING); Penang [5.3333°, 100.25°], 1822, Wall. Cat. 2159.1 [lectoparatypes of G. crinitum, K-W]; ibid., Wall. Cat. s.n. (K); ibid., Wall. Cat. 2158C (K-W); Perak [5°, 101°], 300 m, July 1889, Wray 3775 (SING); Kampong Kota [4.4667°, 100.8°], Feb. 1889, Wray 3340 (SING); Batu Tegoh, Taiping [4.85°, 100.7333°], 19 Jan. 1922, Henderson 10058 (SING); Gunung Ijok [5.2°, 100.7167°], Sept. 1884, Scortechini 5 (lectoparatype of G. crinitum var. griffithii, CAL); Gopeng [4.4667°, 101.1667°], S of Ipoh, Gua Kandu, Davison 1 (SING); Gunong Kerbau [Korbu] [4.6833°, 101.3°], 1200 – 1350 m [4000 – 4500 ft], 14 March 1913, Robinson s.n. (BM); Gunong Raya [Korbu, 4.6833°, 101.2833°], 1200 m [4000 ft], 31 Jan. 1938, Strugnell 45854 (K); Hulu Kwansa [Kangsar], June 1884, Scortechini 928 (lectoparatypes CAL, SING); Hulu Perak, Lenggong, Lata Kekabu, along trail in recreational forest, after hanging bridge, 5°4.93'N 100°94.54'E, 171 m, 18 Dec. 2006, Lim et al. FRI 52830 (K, SING); Ipoh, near Rapat [4.5667°, 101.1167°], mouth of limestone caves, 600 m [2000 ft], 14 Jan. 1983, Davis 69298 (E); Kamuning [4.8333°, 101.0667°], Feb. 1904, Ridley s.n. (SING); Larut & Matang, Bukit Larut Forest Reserve [4.85°, 100.7833°], 310 m, 13 Aug. 1999, Saw FRI 44282 (K); Tanjung Rambutan [4.6667°, 101.1667°], 4 June 1930, Henderson 23773 (SING); Tambun limestone cliffs, 10 Sept. 1920, Burkill & Haniff 6298 (K, SING); Tong Temple, Chemor [4.7167°, 101.1167°], 4 miles N of Ipoh, base of limestone rocks, 23 Oct. 1958, Sinclair 9849 (E, GH, K, L, SING); Upper Perak, N of Lawin, S Kenerong [5.2667°, 101.05°], 20 June 1924, Burkill 12484 (SING); Upper Perak, Bintang Hijau Forest Reserve, Compartment 22, [5.2333°, 100.9667°], 24 Nov. 1966, Ismail 99812 (GH, K, L, SING); Selangor. Hulu Selangor, Gading Forest Reserve [3.6667°, 101.6333°], 450 m [1500 ft], 19 July 1969, Loh 13368 (K, L); 5 miles before Gap [3.6370°, 101.7379°] from Kuala Kubu Baru, 16 March 1985, Kiew 1629 (SING); Genting Bidai [3.3°, 101.8167°], Kloss s.n. (BM, K); Genting Sempah [3.35°, 101.7833°], 22 Oct. 1921, Hume 9263 (SING); Ulu Selangor [3.5667°, 101.5667°], 8 Oct. 1899, Goodenough s.n. (SING); Terengganu. Ulu Brang [4.85°, 102.9333°], 120 m [400 ft], July 1937, Moysey & Kiah 33705 (SING); Hulu Terengganu, Tasik Kenyir, Ex logging trail from Tanjung Bewah towards Gunung Taat, 4°50.74'N 102°42.97'E, 284 m, 7 Aug. 2007, Lim et al. FRI 56461 (K, SING). indonesia. sumatra. Riau: Indragiri, Pangian, Indragiri Kwantan Distr., 1 Oct. 1939, Buwalda 6258 (GH, K).

habitat. In primary or secondary, lowland dipterocarp forest, evergreen forest, on riverbanks; alt. 45 – 1350 m.

conservation status. Widespread; Least Concern (LC).

phenology. Flowering April – October. Fruiting September – March.

notes. Gomphostemma crinitum is recognised by filiform fruiting calyx lobes, much longer than the tube at maturity. It is morphologically similar to G. parviflorum in having a straight corolla and narrow corolla throat, but G. parviflorum does not have calyx lobes longer than the tube.

Prain named Gomphostemma crinitum var. griffthii based on some specimens from Peninsular Malaysia distinguished by their leaf blades, petiole and calyx; we consider them to fall within the variation in these characters in G. crinitum. Gomphostemma crinitum var. griffithii was named to honour William Griffith, a British colonial physician and botanist in 1891 when Prain acted as Superintendent of the Calcutta Botanic Garden, Calcutta: Griffith’s collection (Griffith 4042, C [CAL0000007147]) is selected as the lectotype.

3. Gomphostemma dolichobotrys Merr. (Merrill 1934a: 148); Keng (1978: 319); Walsingham (2019: 197). Type: Indonesia, Atjeh, Takengon, in primary forest, 1100 m, 7 Jan. 1932, Bangham & Bangham-Masters 728 (holotype GH [GH00001143]; isotype NY [NY00000533]).

Gomphostemma racemosum Steenis ex H.Keng (1969: 80). Type: Indonesia, Northern Sumatra, Atjeh, Gajolanden, 1000 m, 7 Feb. 1937, van Steenis 8684 (holotype L [L0003692]; isotypes BO n.v., K [K000824975], L [L0003691]).

Perennial herb up to 2 m tall. Stems erect, obtusely quadrangular with longitudinal grooves, with a mixture of coppery-coloured dense dendroid hairs with a 5 – many-celled stalk and stellate hairs. Leaves petiolate, chartaceous; blades broadly elliptic-ovate, 15 – 23 × 9 – 15 cm, apex acute, margin shallowly dentate, base cuneate; upper side dull green with 2 – 5-celled simple hairs, lower side pale green tomentose with dense stellate hairs and dendroid hairs confined on midrib and veins and with sparse pale yellow punctate glands; petioles 30 – 70 mm long. Inflorescence axillary, raceme-like with opposite cymes unbranched, lax with axis visible between flowers, 20 – 80 (– 100) mm long, inserted at the upper nodes at which leaves are usually present; verticils 3 – 6 (– 8)-flowered; peduncles (15 –) 30 – 60 mm long; bracts deciduous, not seen; bracteoles linear, narrowly elliptic or slightly falcate, 5 – 10 mm long. Flowering calyx tubular or narrowly infundibular, 6 – 10 mm long, 10-ridged, outside with a mixture of dense hairs, inside with adpressed simple hairs; tube 4 – 7 mm long; lobes 2 – 3 mm long, apex acute. Fruiting calyx tubular-infundibular, 7 – 11 mm long; tube 5 – 8 mm long; lobes 2 – 3 mm long. Corolla white, 15 – 20 mm long, throat narrow, gradually dilated towards the throat, outside tomentose with dense stellate hairs, inside glabrous; tube slender, straight, 8 – 15 mm long, hardly exserted from calyx; posterior lip hood-like, ovate, 2 – 5 mm long; anterior lip slightly longer than posterior lip, 3-lobed, subequal, ovate. Staminal filaments fleshy, tomentose. Style slightly fleshy, glabrous with apex subequally lobed. Nutlets solitary, 5 – 6 mm long.

distribution. Indonesia (Sumatra). Map 1.

specimens examined. indonesia. sumatra. Aceh: Gunung Leuser Nature Reserve, Climbing Gunung Leuser West top, from Penosan via Putjuk Angasan, 25 km SW of Blang Kedjeren [3.9976°, 97.2069°], 1300 – 1500 m, 31 March 1975, de Wilde & de Wilde-Duyfjes 15892 (L); Takengon, 1100 m, 7 Jan. 1932, Bangham & Bangham-Masters 728 (holotype GH; isotype NY); Gajolanden, 1000 m, 7 Feb. 1937, van Steenis 8684 (holotype of G. racemosum, L; isotypes K, L).

habitat. In primary forest, riverine forest and scrub edge; alt. 1000 – 1500 m.

conservation status. There are only three collections of Gomphostemma dolichobotrys collected from Gunung Leuser Nature Reserve in North Sumatra giving an EOO of 765 km² suggesting Endangered status. However, the localities are well-protected as a nature reserve and no evidence of habitat loss, hence, the Least Concern (LC) category is applied.

phenology. Flowering and fruiting January – March.

notes. Gomphostemma dolichobotrys is recognised by its erect stems, upright inflorescence with deciduous bracts, and corolla tube hardly exserted from the calyx. It is morphologically similar to G. microdon in having an unbranched, raceme-like inflorescence.

4. Gomphostemma eriocarpon Benth. (Bentham in Wallich 1830b: 12); Hooker (1885: 698); Prain (1891: 258); Mukerjee (1940: 210). Type: India, Courtallum [Courtallam, 8.9167°, 77.2833°], Herb. Wight s.n. (lectotype K [K000846338], selected here).

Herb up to 1.2 m tall. Stems ascending with creeping shoots, sometimes arising from leaf axils, obtusely quadrangular, tomentose with dense stellate hairs. Leaves petiolate, chartaceous; blades elliptic-ovate or elliptic, 7 – 18 × 3 – 8 cm, apex acute, margin dentate or dentate-serrate with glandular thickening at the tip of serration, base attenuate, upper side with stellate hairs and sparse 2 – 3-celled simple hairs, lower side pale green tomentose with dense stellate hairs; petiole 10 – 25 mm long. Inflorescence axillary with opposite cymes sessile, unbranched, congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils 2 – 4-flowered; bracts ovate, 7 – 10 × 2 – 5 mm, apex acute, margin entire or shallowly serrate at the apical part, base attenuate, stellate hairs on both sides; bracteoles narrowly lanceolate or narrowly elliptic, 5 – 10 mm long. Flowering calyx infundibular, 8 – 13 mm long, 10-ridged, outside with a mixture of dense stellate hairs and 3 – 4-celled simple hairs, inside glabrous; tube 4 – 6 mm long; lobes triangular at base tapering into the apex, 4 – 7 mm long. Fruiting calyx cupular, 10 – 15 mm long; tube 5 – 7 mm long; lobes triangular, 5 – 8 mm long, apex subulate. Corolla white, 20 – 30 mm long, throat narrow, not enlarged, sometimes gradually dilated towards the throat; tube slender, straight, 15 – 25 mm long, exserted from calyx, outside tomentose with stellate hairs, inside glabrous; lobes spreading; posterior lip ovate, 4 – 6 × 3 – 5 mm; anterior lip 3-lobed, subequal with round apex, shorter than posterior lip. Staminal filaments slender, tomentose. Style slender, glabrous, shorter than corolla with apex subequally lobed. Nutlets solitary, globose, 4 – 5 mm long, tomentose at apex.

distribution. India (Karnataka, Kerala, Tamil Nadu). Map 1.

specimens examined. india. Karnataka: Maisor & Carnatic, Thomson 119 (K [Herb. Hookerianum], P); Kerala: Quilon, Perundanaruvi Ranni, 125 m, 24 Nov. 1979, Chandra Bose 49190 (CAL); Madras, Travancore, 600 m [2000 ft], Nov. 1893, Lawson 89 (K); Travancore, Bourdillon 41 (K); Tamil Nadu: Anamally Hills [Anaimalai Hills, 10.3°, 77°], 1200 m [4000 ft], Beddome 244 (BM); Courtallum, Herb. Wight s.n. (lectotype, K [Herb. Benthamianum]); ibid., Herb. Wight s.n. (K); ibid., Herb. Madras in Wall. Cat. 2710A (K-W); ibid., Wight s.n. in Wall. Cat. 2710B (lectoparatype, K-W); Tinnevelly hills [8.8333°, 77.6667°], Beddome 243 (BM).

habitat. In evergreen forest (Sadidharan, 2004); alt. 125 – 1200 m.

conservation status. Gomphostemma eriocarpon has an EOO of 29,828.53 km² exceeding the threshold for Vulnerable, and it is found in several localities with no evidence of habitat loss, the Least Concern (LC) category is applied.

phenology. Flowering July – November. Fruiting August – November.

notes. Gomphostemma eriocarpon is recognised by a dentate leaf margin with the leaf tooth glandular-like at the apex and in having a solitary globose nutlet.

Bentham mentioned the collection from Courtallum. Wight’s collections from Courtallam kept in K were examined and this collection [K000846338] is selected as the lectotype.

5. Gomphostemma keralense Vivek., Gopalan & R. Ansari (Vivekananthan et al. 1983: 189). Type: India, Kerala, Idikki [Idukki] Distr., way to Mangaladevi Temple, 950 m, 10 Oct. 1976, Vivekananthan 48658 (holotype CAL (CAL0000006806); isotypes K [K000846335], MH n.v.).

Perennial herb. Stems erect, woody at base, branched, obtusely quadrangular with longitudinal deep grooves, with dense sessile stellate hairs. Leaves petiolate, chartaceous; blades elliptic or elliptic-ovate, 13 – 23 × 6 – 10 cm, apex acute, margin serrate-dentate, base attenuate, upper side with sparsely simple hairs and sessile stellate hairs, confined on midrib and veins; lower side pale green tomentose with sessile stellate hairs confined on midrib and veins; petioles 30 – 90 mm long. Inflorescence axillary with opposite cymes sessile, congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils 2 – 5-flowered; bracts rhomboid or elliptic, 5 – 10 × 4 – 7 mm, apex acute, margin shallowly serrate at the apical part and entire at the basal part, with sessile stellate hairs on both sides; bracteoles narrowly ovate, 5 – 9 mm long. Flowering calyx infundibular, 9 – 11 mm long, 10-ridged, outside creamy-white tomentose with sessile stellate hairs, inside with adpressed simple hairs; tube 4 – 5 mm long; lobes triangular, 5 – 6 mm long, apex acute. Fruiting calyx ovate-infundibular, 12 – 14 mm long; tube 6 – 7 mm long; lobes equal, triangular, 6 – 7 mm long, apex acuminate. Corolla 8 – 10 mm long, throat narrow, not enlarged, sometimes gradually dilated towards the throat, outside glabrous below and with sessile stellate hairs above, inside glabrous; tube slender, straight, 5 – 7 mm long, included inside calyx. Staminal filaments slightly fleshy. Style slender, as long as corolla. Nutlets solitary, elliptic, 6 mm long, sparsely tomentose.

distribution. India (Kerala). Map 1.

specimens examined. india. Kerala: Idukki Distr., Paermade [Peermade, 9.5667°N 76.9833°E], Meebold 12941 (K); Udumbansholay [Udumbanchola, 9.9°, 77.1667°], Meebold 13015 (K); way to Mangaladevi Temple, 950 m, 10 Oct. 1976, Vivekananthan 48658 (holotype CAL; isotypes K).

habitat. In evergreen or semi-evergreen forest; alt. 950 m.

conservation status. Gomphostemma keralense is known from three collections collected from Idukki Distr. and has an EOO of 451.01 km² and AOO of 12 km². Google Earth imagery shows some evidence of habitat loss. Hence, the Vulnerable (VU) category is applied (VU D2).

phenology. Flowering and fruiting in October.

notes. Gomphostemma keralense is recognised by the axillary inflorescence with a short corolla included in the calyx. It is morphologically similar to G. heyneanum in having an included corolla and is found in the same area of South India. But it differs by having axillary inflorescences.

6. Gomphostemma longipetalum Bongch. sp. nov. Type: Burma, Kachin, Mountains East of Fort Hertz, 27°30'N 97°50'E, 600 – 900 m [2000 – 3000 ft], 3 Sept. 1926, Ward 7359 (holotype K [K001081925]).

http://www.ipni.org/urn:lsid:ipni.org:names:77221537-1

Erect herb, up to 0.6 m tall. Stems ascending, woody at nodes, obtusely quadrangular, with longitudinal grooves, tomentose with dense stellate hairs with a 1-celled stalk or sessile stellate hairs, dendroid hairs absent; internodes straight. Leaves petiolate, spreading, chartaceous; blades elliptic or elliptic-ovate, 10 – 17 × 5 – 7 cm, apex acute, margin shallowly serrate, base attenuate, upper side coarse with 1 – 3-celled simple hairs, bifurcate hairs or stellate hairs, lower side pale green tomentose with dense stellate hairs; petioles 10 – 15 mm long. Inflorescence axillary, with opposite cymes sessile, unbranched then congested with axis not visible between flowers, exceeding 20 mm wide when pressed, excluding corolla, inserted at the upper nodes at which leaves are usually present; verticils many-flowered; bracts elliptic, 6 – 10 × 2 – 3 mm, apex acute, margin serrate, base attenuate, upper side with simple hairs and bifurcate hairs, lower side tomentose with stellate hairs; bracteoles narrowly elliptic, sometimes falcate or ensiform, 6 – 8 × 1 – 1.5 mm. Flowering calyx infundibular, 8 – 10 mm long, 5-ridged, outside with dense stellate hairs, inside tomentose with adpressed short simple hairs; tube 4 – 5 mm long; lobes 4 – 5 mm long, apex filiform. Fruiting calyx not seen. Corolla yellow, 40 – 50 mm long, throat narrow, not enlarged, outside tomentose with sessile stellate hairs and glandular-like simple hairs with a multi-celled stalk, inside glabrous; tube straight, 34 – 42 mm long, exserted from the calyx; posterior lobe hood-like, 6 – 8 × 3 – 5 mm; anterior lobes spreading; median lobe 5 – 7 mm long. Staminal filaments slender. Disk lobed, tumescent behind. Style slender with apex unequally lobed. Nutlets not seen. Fig. 7.

Fig. 7

Gomphostemma longipetalum. A habit; B inflorescence; C stellate hairs with a 1-celled stalk from lower leaf surface; D upper leaf surface indumentum; E flower, lateral view; F upper part of corolla opened to show stamens with hairy filaments and style; G stellate hairs with a 1-celled stalk from corolla; H glandular hairs with a multi-celled stalk from corolla. All from Ward 7359. drawn by andrew brown.

recognition. Gomphostemma longipetalum is distinguished from the straight corolla tube species of Gomphostemma in having a much longer corolla, 40 – 50 mm long, calyx lobes not longer than the tube and cymes inserted at the lower nodes where the leaves have fallen. This species could be confused with G. crinitum because of its straight corolla tube and narrow corolla throat but that species has a shorter corolla, 20 – 30 mm long, and calyx lobes much longer than the tube.

distribution. Myanmar. Map 2.

Map 2

Distribution range of Gomphostemma longipetalum, G. niveum, G. parviflorum and G. thomsonii.

specimens examined. myanmar. Kachin: mountains E of Fort Hertz, 27°30'N 97°50'E, 3 Sept. 1926, Ward 7359 (holotype K).

habitat. In mountains; alt. 600 – 900 m.

conservation status. Gomphostemma longipetalum is known only from the type specimen collected in 1926. Although the label mentioned that this species is ‘fairly abundant at one spot’, the Data Deficient (DD) category is applied due to the lack of further records.

phenology. Flowering September.

notes. The epithet longipetalum [longi- (in Greek compound), long; -petalum, petal] refers to its corolla length.

7. Gomphostemma niveum Hook.f. (Hooker 1885: 697); Prain (1891: 250); Mukerjee (1940: 207). Type: Bangladesh, East Bengal, Griffith 1015 (lectotype K [K000846344]), selected here; isolectotypes BM, E [E00301974]).

Herb up to 1.2 m tall. Stems erect, woody at base, obtusely quadrangular with longitudinal grooves, tomentose with dense whitish stellate hairs. Leaves subsessile, chartaceous; blades obovate or elliptic-obovate, 9 – 31 × 5 – 12 cm, apex acute, margin shallowly serrate, base attenuate; upper side glabrous; lower side whitish or creamy-white tomentose with dense sessile hairs; petioles 5 – 15 mm long. Inflorescence axillary with opposite cymes sessile, unbranched, congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils many-flowered; cluster of leaf-like bracts, sessile, deciduous, ovate or elliptic, (10 –) 15 – 30 × (4 –) 8 – 15 mm, apex acute, margin serrate at the apical part and entire at the basal part, base attenuate, upper side puberulent, lower side with white stellate hairs; bracteoles falcate or linear, 7 – 15 mm long. Flowering calyx infundibular, 8 – 10 mm long, 10-ridged, outside with stellate hairs, inside with adpressed shortly simple hairs; tube 4 – 5 mm long; lobes 4 – 5 mm long, apex acuminate. Fruiting calyx infundibular, 9 – 12 mm long; tube 4 – 6 mm long; lobes 5 – 6 mm long. Corolla yellow, 13 – 15 mm long, throat narrow, not enlarged, sometimes gradually dilated towards the throat, outside tomentose, inside glabrous; tube straight, 10 – 12 mm long, slightly or hardly exserted from calyx. Staminal filaments fleshy, slightly tomentose. Nutlets solitary, oblong or obovate, 4 – 5 mm long.

distribution. Bangladesh. Map 2.

specimens examined. bangladesh. East Bengal [now Bangladesh], Griffith 1015 (lectotype K; isolectotypes BM, E); ibid., Griffith s.n. (lectoparatype K).

habitat. In forest.

conservation status. Gomphostemma niveum is known only from the types and no new collection has been made since then. Data to evaluate decline of habitats is not available. Hence, the Data Deficient (DD) category is applied to this species.

notes. Gomphostemma niveum is recognised by having sessile or subsessile leaves which are white tomentose on the lower side and by a slightly or hardly exserted corolla tube from calyx. It is morphologically similar to G. sulcatum by having cluster of small leaf-like bracts but that species has a clearly exserted corolla tube.

Two of Griffith’s collections from ‘East Bengal’ are found at K, of which only one has the number 1015 [K000846344]; this is chosen as the lectotype.

8. Gomphostemma parviflorum Wall. ex Benth. (Bentham in Wallich 1830b: 12); Hooker (1885: 697); Prain (1891: 252); Mukerjee (1940: 208); Keng (1969: 83, 1978: 321); Murata (1976: 190); Li & Hedge (1994: 72); Phuong (2000: 102); Walsingham (2019: 203). Type: India, Sillet [Sylhet, 24.8833°, 91.85°], de Silva s.n. in Wall. Cat. 2158 (lectotype K [K000846340], selected here].

Gomphostemma multiflorum Benth. (Bentham 1848: 552). Type: India, Sillet, Wall. Cat. s.n. (holotype K [K000846341]).

Gomphostemma dichotomum Zoll. & Moritzi (Zollinger & Moritzi 1846: 54). Type: Indonesia, Java, Zollinger 2028 (holotype P [P00686242]; isotypes A n.v., BM 2 sheets, G [G00189004, G00188984], P [P0068250]).

Gomphostemma bartlettii Merr. (Merrill 1934b: 191). Type: Indonesia, Sumatra, Tapianoeli, between Panapparan and Pagar Batoe, Habinsaran, 17 May 1927, Bartlett 7935 (holotype NY; isotypes L, US).

Gomphostemma parviflorum var. farinosa Prain (1891: 253). Type: India, Assam, Goalpara [26.17°, 90.62°], 24 July 1808, Hamilton s.n. in Wall. Cat. 2158D [‘a’] (lectotype CAL [CAL0000007155], selected here; isolectotype K-W [K001115346]).

Gomphostemma salarkhaniana Khanam & M.A.Hassan (2003: 63). Type: Bangladesh, Sylhet, Tamabil, 13 Oct. 1973, Khan, Huq & Hassan 3296 (holotype DACB n.v.).

Perennial herb 1 – 2 m tall. Stems erect, obtusely quadrangular with longitudinal grooves, with dense dendroid hairs and stellate hairs. Leaves petiolate, chartaceous; blades elliptic-obovate, 20 – 40 × 10 – 20 cm, apex acute, margin shallowly serrate, base attenuate, upper side dark green with stiff stellate hairs, lower side pale green tomentose with dense stellate hairs with a 1-celled or multi-celled stalk; petioles 20 – 40 mm long. Inflorescence axillary, with opposite cymes pedunculate, branched then lax, exceeding 20 mm wide when pressed, excluding corolla, inserted at the upper nodes at which leaves are usually present; verticils many-flowered; peduncles 5 – 15 mm long; bracts obovate, 8 – 15 × 4 – 7 mm, broader than the calyx, apex acute, margin entire. Flowering calyx tubular-infundibular, 8 – 12 mm long, 10-ridged, outside with dense stellate hairs, inside tomentose with adpressed shortly simple hairs; tube 5 – 7 mm long; lobes triangular, 3 – 5 mm long, apex acuminate. Fruiting calyx infundibular, 8 – 13 mm long; tube 5 – 7 mm long; lobes triangular, 3 – 6 mm long, apex acuminate, sometimes subulate. Corolla bright yellow, 20 – 35 mm long, throat narrow, not enlarged, sometimes gradually dilated towards the throat, outside with simple hairs and glandular-like simple hairs, inside glabrous; tube slender, straight, 15 – 30 mm long, exserted from the calyx; posterior lip slightly hooded, elliptic-ovate, 5 – 6 mm long; anterior lip 3-lobed, slightly longer than posterior lip; median lobe elliptic, 4 – 6 mm long; lateral lobes elliptic. Staminal filaments fleshy, tomentose. Style slender, glabrous with apex unequally lobed. Nutlets solitary, white when fresh, elliptic-ovate or oblong, 4 – 5 mm long, glabrous. Fig. 1C & 6B, G.

distribution. India (Arunachal Pradesh, Nagaland, Sikkim, Meghalaya, West Bengal, Assam), Nepal, Bangladesh, China, (Yunnan), Myanmar, Vietnam, Thailand, Peninsular Malaysia, Indonesia (Java). Map 2.

specimens examined. india. West Bengal: Darjeeling, Ryang, 300 m [1000 ft], 14 Nov. 1870, Clarke 13763A (BM); Darjeeling, Bomunpokri Hill [27.0333°, 88.1667°], Aug. 1874, Gamble 3818A (K); Darjeeling, Badamtam [27.0333°, 88.1667°], 600 m [2000 ft], Aug. 1881, Gamble 9724 (lectoparatype of G. parviflorum var. farinosa, K); Jalpaiguri, Chapramari Forest, 27 July 2002, Bhattachcharya & Party 31167 (CAL); Jalpaiguri, Duars, Jan. – May 1895, Haines 516 (K); Kuntimari, Jalpaiguri, 14 Jan. 1939, Parkinson 3460 (GH); Sikkim: ibid., 1200 m [4000 ft], Hooker s.n. (K, L); Teesta Valley, 300 m [1000 ft], 20 Aug. 1874, Gamble s.n. (K 2 sheets); Teesta R., 300 m [1000 ft], 12 Nov. 1870, Clarke 13674A (lectoparatype of G. parviflorum var. farinosa, BM); Mongpo, 300 m [1000 ft], 5 Dec. 1884, Clarke 36174B (E); Terai, 1200 m [4000 ft], Sept. 1825, Hooker s.n. (K, TCD); Arunachal Pradesh: Namchik R. Valley, 23 mile mark on Ledo Road [27.2922°, 96.0908°], 11 Sept. 1945, Belcher 241 (K); Assam: Griffith 339 (K); ibid., Griffith s.n. (K); ibid., 1846, Griffith s.n. (K); ibid., July, Simons 46 (K); Goyalpara [Goalpara], 24 July 1808, Hamilton s.n. in Wall. Cat. 2158D [‘a’] (lectotype of G. parviflorum var. farinosa, CAL; isolectotype K-W); ibid., 23 Oct. 1808, Hamilton s.n. in Wall. Cat. 2158D [‘b’] (lectoparatypes of G. parviflorum var. farinosa, E, K-W); North Cachar, Haflong [25.1833°, 93.0333°], 750 m [2500 ft], 3 Aug. 1908, Craib 125 (K); Assam, Kohima, Naga Hills, 1500 m [5000 ft], 19 July 1950, Koelz 25506 (L); Assam, Badarpur [24.9°, 92.6°], Nov. 1907, Meebold 5814 (K); Assam, Shella, 24 miles from Maeosmai, K B J Hills Distr., 19 Dec. 1956, Panigrahi 4877 (CAL); Assam, Lushai Hills, Kaidun Kawn, 600 m [2000 ft], July 1927, Parry 283 (K); Assam, Baligan near Naga Hills, Nov. 1898, Prain's collector 390 (G 3 sheets); Assam, Jabrea Juyali Bam Garden [26.14°, 91.77°], Oct. 1898, Prain's collector 141 (E, GH, TCD); Nagaland: Mokokchcing [26.3333°, 94.5333°], 18 Sept. 1942, Bor 16665 (L); Naga Hills, Lumami, 1200 m [4000 ft], 1 July 1935, Bor 5075 (K); Kala, Naga Hills, 900 – 1200 m [3000 – 4000 ft], May 1882, Watt 7396 (E); Meghalaya: Jaintea, de Silva s.n. in Wall. Cat. 2158.2 [‘b’] (lectoparatype of G. parviflorum K-W); Khasia, Shillong [25.5667°, 91.8833°], 1290 m [4300 ft], 23 July 1886, Clarke 44358D, E & H (BM, G); Garo Hills [25.5°, 90.5°], Rewak, 360 m [1200 ft], Parry 925 (K); Cherrapoonjee, 1500 m [5000 ft], Sept. 1878, Gallatly 613 (lectoparatype of G. parviflorum var. farinosa BM); Nongpoh, Khasi Hills, 600 m [2000 ft], 26 June 1949, Koelz 23029 (E); Khasia, Maodadoom, 900 m [3000 ft], 3 Feb. 1886, Clarke 42921A & B (E, G); Khasia, 19 Oct. 1871, Clarke 16003 (SING); Khasia, Bor Panee [26.2167°, 92.5333°], 0 – 1200 m [0 – 4000 ft], 2 Oct. 1850, Hooker & Thomson s.n. (K); Khasia, Churra [25.5833°, 91.6333°], [0 – 1200 m 0 – 4000 ft], 11 Aug. 1850, Hooker & Thomson 2062 (K 2 sheets); ibid., 0 – 1200 m [0 – 4000 ft], 11 Aug. 1850, Hooker & Thomson s.n. (K); Khasia, Terai, 0 – 1200 m [0 – 4000 ft], 11 Nov. 1850, Hooker & Thomson s.n. (K); ibid., 0 – 1200 m [0 – 4000 ft], 16 Nov. 1850, Hooker & Thomson s.n. (K, TCD]; Mont Khasia [25.5833°, 91.6333°], 300 – 1200 m [1000 – 4000 ft], Hooker & Thomson s.n. (BM, E, G 3 sheets, L 2 sheets, P 2 sheets); Tamil Nadu: Medras, Herb. Heyne in Wall. Cat. 2158.1 (lectoparatype of G. parviflorum K-W). bangladesh. Sylhet: Wall. Cat. 2158 (lectotype of G. parviflorum K); ibid., July 1820, de Silva s.n. in Wall. Cat. 2158.2 [‘a’] (lectoparatypes of G. parviflorum K-W, L, NY 2 sheets); ibid., Wall. Cat. s.n. (holotype of G. multiflorum K; isotype K); Chittagong: Matamori [Matamuhory, Matamuhari, 21.65°, 92°], 29 Sept. 1920, Cowan 855 (E); Maghi, Fashaba [23.6244°, 91.0761°], Cowan 1341 (E); Barkal, 29 Feb. 1876, Lister 118 (BM); ibid., 0 – 300 m [0 – 1000 ft], 9 Jan. 1851, Hooker & Thomson 488 (K). china. Yunnan: Szemao [22.7667°, 101.0833°], 1200 m [4000 ft], Henry 12253A (K); Szemao, East Mountains, 1200 m [4000 ft], Henry 12253 (K); Ching-tung, Hsiao-ch'ang-chieh [24.2333°, 103.45°], 1100 m, 13 Dec. 1939, Li 1491 (KUN); en route from Xima to Yingjiang, 300 m, 24 Nov. 1993, Santisuk et al. 890 (BKF 2 sheets); Luchun, 23°22'51”N 102°57'23”E, 800 m, 18 Oct. 2000, Shui & Chen 13118 (KUN); Luchun, Huanglian Shan, Qi-ma-ba [22.7667°, 102.2°], 900 – 1200 m, 1 Nov. 1995, Wu et al. 1046 (KUN); Man-ya, Che-li Hsien [21.9833°, 100.8167°], 850 m, Sept. 1936, Wang 78775 (KUN). myanmar. Sagaing: Katha, Kadu Hill [24.1833°, 96.35°], 10 July 1911, Lace 5328 (K); Mandalay: Maymyo Plateau [22.0333°, 96.4667°], 26 July 1912, Lace 5860 (E 2 sheets, K). vietnam. Dien Bien: Muong phang [Dien Bien, 21.4431°, 103.1547°], 29 June 2005, Binh & Cuong 1536 (HN). thailand. Mae Hong Son: Khun Yuam, toward Mae Surin Waterfall, 18°52'N 98°5'E, 1500 m, 24 Sept. 1995, Larsen et al. 46873 (AAU); Chiang Mai: Doi Chiang Dao [19.3833°, 98.9°], 20 Dec. 1961, Bunchai 84 (BKF); ibid., 1010 m, 9 May 1954, Bunchai 250 (BKF); ibid., 1800 – 2150 m, 27 Oct. 1979, Shimizu et al. T-21197 (BKF); ibid., 1600 m, 17 Feb. 1958, Sørensen et al. 1295 (BKF, C 2 sheets); Doi Chiang Dao, at foot N of Ban Tam [19.3944°, 98.9267°], 440 m, 27 Aug. 1935, Garrett 997 (BKF, K 2 sheets, L, P, TCD); Doi Chiang Dao, Me Na Lao drainage, 640 m, 29 Dec. 1953, Garrett 1423 (K, L, P); Doi Chiang Dao, E side, Pa Blawng Caves area [19.3833°, 98.9°], 600 m, 15 Jan. 1989, Maxwell 89-44 (L); Doi Chiang Dao Wildlife Sanctuary, N side of Doi Luang, above Sop Huay Pah Dang-Huay Nah Lao Station, 975 m, 8 Oct. 1995, Maxwell 95-850 (BKF, CMUB); Doi Inthanon, Mae Pan Waterfall [18.5263°, 98.4557°], 1000 – 1060 m, 29 July 1988, Fukuoka T-62322 (BKF); Mae Rim, Pong Yaeng [18.8904°, 98.8296°], 1000 m, 9 Sept. 2009, Maxwell 09-259 (CMUB, QBG); Doi Ang Khang [19.8°, 98.9833°], 3 Feb. 1994, Tiptabiankarn 1372 (SLR); Chiang Rai: Khun Korn Waterfall [19.8821°, 99.6169°], 675 m, 11 Oct. 1997, KK 398 (BCU); Khun Lao, Doi Pee Pan Nam [19.15°, 101.1833°], 1000 m, 25 Sept. 2007, Watthana 2462 (QBG); Nan: Doi Phu Kha, 19°10'N 101°5'E, 1400 m, 11 Sept. 1995, Larsen et al. 46335 (AAU); Nan, Doi Phu Khan National Park, c. 15 km E after past highest point, 19°25'N 101°16'E, 1000 m, 16 Aug. 1995, Parnell et al. 95-156 (K, TCD); Pua, Doi Phu Kha National Park, Tam Pha Khaw, 1000 m, 5 July 2001, Srisanga 1924 (QBG); Phitsanulok: Phu Mieng Mountain [16.85°, 101.1167°], 700 m, 29 July 1966, Larsen et al. 1098 (AAU, BKF); Phetchabun: Nam Nao National Park [16.7385°, 101.5698°], trail W from Visitor Centre, 23 Sept. 1994, Boyce 1026 (K); Loei: Na Haew, 1000 m, 30 July 1995, Na Nakorn et al. 4020 (QBG). indonesia. java. Zollinger 2028 (holotype of G. dichotomum, P; isotypes BM 2 sheets, G, P).

habitat. Evergreen forest; alt. 440 – 1900 m.

conservation status. Widespread; Least Concern (LC).

phenology. Flowering June – January. Fruiting October – May.

notes. Gomphostemma parviflorum is recognisable by broad bracts broader than the calyx and calyx lobes not longer than the tube. It is morphologically similar to G. crinitum by having a straight corolla tube and a narrow corolla throat. Although the type specimen of G. salarkhaniana has not been seen, this species is similar to G. parviflorum based on the plant description from the protologue. Hence, G. salarkhaniana is reduced to a synonym of G. parviflorum.

The localities of Gomphostemma parviflorum mentioned in the protologue included Nepal, Sillet and Penang but, based on Wallich (1829), there is no collection collected from Nepal under the catalogue number 2158.

The evidence for matching Wall. Cat. 2158 housed at K (Herb. Benthamianum sheet) with either Wall. Cat. 2158.2 [‘a’] (K-W) or Wall. Cat. 2158.2 [‘b’] (K-W) is not clear. Hence, no isolectotype of Gomphostemma parviflorum is designated.

There is one specimen labelled as Zollinger 2028 (P [P00686249]) but it is Gomphostemma javanicum based on the calyx character, prominently 10-ridged.

9. Gomphostemma thomsonii Benth. ex Hook.f. (Hooker 1885: 698); Prain (1891: 251); Mukerjee (1940: 209). Type: India, Khasia Mts, Churra, 900 – 1500 m [3000 – 5000 ft], 11 Aug. 1850, Hooker & Thomson 2063 (lectotype K [K000846342], selected here; isolectotype K [K000846343]).

Herb up to 1 m tall. Stems erect, woody at base, obtusely quadrangular with longitudinal grooves, tomentose with dense stellate hairs; internodes straight. Leaves petiolate, chartaceous; blades elliptic-ovate, 17 – 26 × 9 – 15 cm, apex acute, margin serrate, base cuneate, upper side dull green with 2 – 3-celled simple hairs, bifurcate hairs or stellate hairs, lower side pale green tomentose with dense stellate hairs confined on midrib and veins; petioles 50 – 140 mm long, with dense stellate hairs. Inflorescence axillary with opposite cymes sessile, unbranched, congested with axis not visible between flowers, exceeding 20 mm wide when pressed, excluding corolla, inserted at the upper nodes at which leaves are usually present; verticils few – many-flowered; bracts deciduous, ovate or ovate-lanceolate, 5 – 12 × 4 – 8 mm, broader than the calyx; bracteoles linear, 6 – 10 mm long. Flowering calyx tubular-infundibular, 8 – 10 mm long, 10-ridged, outside with dense branched hairs, inside tomentose with simple hairs; tube 4 – 6 mm long; lobes narrow triangular, c. 4 mm long, apex acuminate. Fruiting calyx infundibular, 10 – 12 mm long; tube 7 – 8 mm long; lobes 3 – 4 mm long, apex subulate. Corolla pale yellow, 15 – 20 mm long, throat narrow, not enlarged, sometimes gradually dilated towards the throat, outside tomentose with simple hairs, inside glabrous; tube straight, 12 – 15 mm, exserted from the calyx; posterior lip ovate, 4 – 6 mm long, apex and margin entire; anterior lip 3-lipped, subequal, slightly longer than posterior lip. Staminal filaments slightly fleshy. Style slender. Nutlets solitary, oblong, 5 – 6 mm long, slightly wrinkled. Fig. 6E, J.

distribution. India (Meghalaya). Map 2.

specimens examined. india. Meghalaya: Khasia, Churra, 900 – 1500 m [3000 – 5000 ft], 11 Aug. 1850, Hooker & Thomson 2063 (lectotype K; isolectotype K); ibid., 1200 m [4000 ft], 15 June 1855, Hooker & Thomson s.n. (K); Khasia, 900 – 1500 m [3000 – 5000 ft], Hooker & Thomson s.n. (BM, C, G, L, P, TCD).

habitat. In evergreen forest; alt. 900 – 1500 m.

conservation status. Gomphostemma thomsonii is known from four collections, three localities with AOO of 16 km² with 2 km width grid size. The habitat in Sumatra is threatened by oil palm plantations (Fitzherbert et al. 2008). Hence, the taxon is given the status of Endangered (EN B2ab(ii)).

notes. Gomphostemma thomsonii is distinguished by having broad leaves, 90 – 150 mm wide and long petioles, 50 – 140 mm long. It is morphologically similar to G. parviflorum by having congested cymes and calyx lobes not longer than the tube at maturity but G. parviflorum has narrower leaves, 40 – 90 mm wide.

I) Glabrous group: ii. Corolla with inflated throat and an incurved tube (species no. 10 – 23)

10. Gomphostemma chinense Oliv. (Oliver 1884: sub t. 1468); Prain (1891: 267); Doan (1936: 1034); Li & Hedge (1994: 74); Phuong (2000: 110). Type: China, Lo-fan-shan Mts, East River [23.67°, 114.6°], Faber 1 (lectotype K [K000928868], selected here).

Taitonia callicarpoides Yamam. (Yamamoto 1938: 278). Type: Taiwan, Taito, Sinsuiei, 22°23'48”N 120°46'18”, 1200 m, 7 July 1937, Simidu s.n. (holotype TAI).

Gomphostemma callicarpoides (Yamam.) Masam. (Masamune 1942: 4). Type: as for Taitonia callicarpoides Yamam.

Gomphostemma formosanum Masam. (Masamune 1942: 4). Type: Taiwan, Hunkiko, Tainan-syu, 23°30'16”N 120°41'18”E, 3 Aug. 1941, Yosimura s.n. (holotype TAI).

Gomphostemma chinense var. cauliflorum C.Y.Wu & S.Chow (in Wu 1965: 149). Type: China, Hainan, Po-ting [18.6403°, 109.6994°], 700 m, 24 July 1935, How 73296 (holotype IBSC n.v.; isotype G [G00189003]).

Gomphostemma pseudocrinitum C.Y.Wu (1965: 144). Type: China, Kwangsi, Ling-lo (Ling-yun), Tsing-lung-shan [26.1356°, 110.8253°], 1130 m, 12 Aug. 1928, Ching 6851 (holotype PE).

Perennial herb, decumbent-like, up to 1 m tall. Stems ascending, woody at base, much-branched at base, obtusely quadrangular, tomentose with dense stellate hairs with a 1-celled stalk or sessile stellate hairs, dendroid hairs absent; internodes sometimes wavy. Leaves petiolate, chartaceous; blades elliptic, cordate, ovate or broadly elliptic-ovate, 4 – 15 × 3 – 9 cm, apex broadly acute or round, margin shallowly serrate, base cuneate or obtuse, upper side dark green with stellate hairs and 3 – 4-celled simple hairs, lower side pale green with dense stellate hairs; petiole 10 – 40 mm long. Inflorescence basal or near basal at the lower nodes from which leaves have mostly fallen, with opposite cymes pedunculate or subsessile, congested or slightly lax, branched; verticils 4 – 10-flowered; peduncles 5 – 60 mm long; bracts elliptic, ovate or lanceolate, 12 – 15 × 3 – 5 (– 8) mm, apex acute, margin spinulose or shallowly serrate, base cuneate, upper side glabrous or with stellate hairs, lower side with dense stellate hairs; bracteoles linear, 6 – 10 mm long. Flowering calyx campanulate, 8 – 10 mm long, 10-ridged; tube 4 – 6 mm long; outside with dense stellate hairs, inside with sparse adpressed simple hairs; lobes acute, c. 4 mm long, apex acuminate, outside with dense stellate hairs, inside with dense simple hairs. Fruiting calyx not seen. Corolla white, yellow or red, 30 – 60 mm long, throat inflated, abruptly dilated near the throat; tube slender, incurved, 20 – 50 mm long, outside with simple hairs, inside glabrous; posterior lip hooded, 7 – 8 × 6 – 7 mm, apex shallowly emarginate, margin entire; anterior lip 3-lobed, spreading, longer than posterior lip; median lobe 10 – 15 mm long, apex emarginate with lateral lobes spreading. Staminal filaments glabrous, tomentose at the apical part. Style slender, slightly shorter than corolla with apex unequally lobed. Nutlets not seen.

distribution. China (Guizhou, Guangxi, Hainan, Fujian, Guangdong, Hong Kong), Taiwan and Vietnam. Map 3.

Map 3

Distribution range of Gomphostemma chinense, G. curtisii, G. flexuosum, G. grandiflorum, G. hainanense and G. hirsutum.

specimens examined. china. Guizhou: Kouy Tcheou [Kweichow], Leveille s.n. (E, K); Guangxi: Ling-lo (Ling-yun), Tsing-lung-shan, 1130 m, 12 Aug. 1928, Ching 6851 (holotype of G. pseudocrinitum PE); Kiennan Distr., near Tung Lei Village, Sai Hang Cheung, 1 – 29 Aug. 1934, Lau 4095 (BM, G, GH); Yung Hsien [22.8667°, 110.55°], Ta Tseh Shan, 540 m, 12 Aug. 1933, Steward & Cheo 804 (GH, P); Hainan: Nov. 1954, Liang 68240 (IBSC); Lai area [19.8969°, 110.1108°], Hung Mo Shan and vicinity, 24 July 1929, Tsang & Fung 559 (GH); Po-ting, 700 m, 24 July 1935, How 73296 (isotype of G. chinense var. cauliflorum G); Fujian: Amoy [Xiamen, 24.46°, 118.0789°], inferior, June 1870, Swinhoe s.n. (lectoparatype of G. chinense K); Guangdong: Lo-fan-shan Mts, East R., Faber 1 (lectotype of G. chinense K); Cantonensis, Lo-fan-shan [23.6667°, 114.6°], Oct. 1883, Faber 22275 (BM); Shizing Xian, 400 m, 10 July 1982, Wang 366 (AAU); In montibus Lungtou schan 60 km ad orientem urbis Siudsao ("Schautschou") ad rivos montis Yiu Schan, 800 m, 6 Sept. 1917, Mell 941 (G); Lo Guai, North West R., 18 Aug. 1890, Ford 668 (P); Loh Fau Mt (Lofaushan) [23.6667°, 114.6°], 900 m, 24 Aug. 1917, Merrill s.n. (PHN); Mei (Kaying) Distr. [24.3°, 116.1167°], Yam Na Shan (Yit Nga Shan), 4 – 31 Aug. 1932, Tsang 21458 (BM, K, GH, P); S China, Swatow [Shantou, 23.3667°, 116.6667°], Thai-yong, 600 m [2000 ft], Aug. 1897, Dalziel s.n. (BM, E); Hong Kong: Hong Kong [22.3964°, 114.1095°], 21 July 2003, Lau 141 (K). taiwan. Chiayi: Hunkiko, Tainan-syu, 23°30'16”N 120°41'18”E, 3 Aug. 1941, Yosimura s.n. (holotype of G. formosanum TAI); Chisinliao (Hsihsinliao), 23°30'32”N 120°35'49”E, 15 Aug. 1988, Yen & Kao 10730 (TAI 5 sheets); Chunglun (Chunglun), 23°22'49”N 120°33'11”E, July 1994, Hsieh s.n. (TAI); Kaohsiung: Shanping (Sanping), 22°58'22”N 120°40'33”E, 10 Dec. 1968, Huang 4889 (TAI); Pingtung: Bankenseng Mts [Wanluan Rural Township, Wan-chin Shan (Mandarin) or Banking-sing (Taiwanese), 22.6074°, 120.6297°], Henry 1565 (K); Tainan: Kwantzuling [Kuantzuling (Kwansirei)], 23°20'32”N 120°29'33”E, 11 July 1984, Huang 1075 (TAI 2 sheets); Taitung: Chienching-Shenshan (Miharasi-Miyama), 22°42'11”N 120°56'29”E, 20 Aug. 1932, Hosokawa 5266 (TAI); Chinshuiyin (Sinsuiei), 22°23'48”N 120°46'18”E, 1200 m, 7 July 1937, Simidu s.n. (holotype of T. callicarpoides TAI). vietnam. Ha Tay: Tonkin, Mt Bavi [21.0781°, 105.3583°], 20 July 1886, Balansa 2937 (K, P); Lao Cai: Chapa [22.35°, 103.8667°], 1500 m, July 1927, Petelot 5115 (BM, P); ibid., 1800 m, July 1930, Petelot 5802 (BM, GH, P).

habitat. On wet forest floor; alt. 120 – 1020 m.

conservation status. Widespread and found in well-preserved areas in Vietnam, Hainan and Taiwan. The taxon has Least Concern (LC) status.

phenology. Flowering July – September. Fruiting September – December.

notes. Gomphostemma chinense is a very distinctive taxon defined by its decumbent habit and a glabrous inner corolla tube, curved internodes and stems appearing wavy. It is morphologically similar to G. pedunculatum by its basal inflorescence with cymes inserted at the lower nodes, above the ground, where leaves have fallen completely. This species has been described under several names. In Flora of Taiwan, Gomphostemma callicarpoides is recognised as an endemic species to Taiwan (Huang et al. 1998). Examination of Taiwanese specimens reveals that all Gomphostemma collected from Taiwan are G. chinense.

11. Gomphostemma curtisii Prain (1890: 315); Prain (1891: 266); Keng (1969: 85, 1978: 322); Murata (1976: 188); Walsingham & Bramley (2010: 482); Walsingham (2019: 196). Type: Malaysia, Perak, Waterloo, 600 m [2000 ft], Dec. 1887, Curtis 1310 (lectotype K [K000501083], selected here; isolectotype SING).

Cyrtandromoea repens Ridl. (Ridley 1911: 74). Type: Malaysia, Perak, Temengoh, July 1909, Ridley 14446 (holotype K [K000501084]; isotypes BM, SING [SING0091067]).

Perennial herb, decumbent-like, up to 1.6 m tall. Stems ascending, woody at base, obtusely quadrangular with longitudinal grooves, rooting at nodes, tomentose with sessile or a 1-celled stalk stellate hairs, dendroid hairs absent; internodes sometimes wavy. Leaves petiolate, chartaceous; blades ovate, elliptic-ovate or cordate, 5 – 20 (– 24) × 4 – 11 (– 16) cm, apex acute, margin serrate or shallowly serrate, base cuneate, upper side dark green with sessile stellate or stalked stellate hairs and sparse simple hairs, lower side pale green with stellate hairs; petioles 30 – 60 (– 90) mm long. Inflorescence basal, inserted at the lower nodes where leaves have mostly fallen, with opposite cymes subsessile, congested with axis not visible between flowers or sometimes lax, branched; verticils few – many-flowered; peduncles 0 – 15 (– 25) mm long; bracts ovate or obovate, sometimes asymmetric, keel-like, 7 – 20 × 3 – 10 mm, apex acute or acuminate, margin shallowly serrate at the apical part and entire at the basal part or entire, base attenuate, upper side with sparse stellate hairs at the upper part and glabrous at the basal part, lower side with dense stellate hairs. Flowering calyx dark red or reddish-brown, infundibular, 10 – 15 mm long, 10-ridged, outside with stellate hairs, inside with adpressed shortly simple hairs below and stellate hairs above; tube 4 – 7 mm long; lobes triangular, 6 – 8 mm long, apex subulate, margin slightly revolute. Fruiting calyx cupular, 13 – 23 mm long; tube 5 – 10 mm; lobes 8 – 13 mm long. Corolla white, creamy-white, pinky-white or pale yellow, 30 – 45 mm long, throat inflated, abruptly dilated near the throat; tube incurved, 25 – 35 mm long, outside tomentose with stellate hairs, inside glabrous; posterior lip spreading beyond anterior lip, ovate, 5 – 7 mm long, apex emarginate, margin slightly undulate; anterior lip subequally 3-lobed. Staminal filaments slightly fleshy. Style slender, glabrous, not long beyond anthers with apex subequally lobed. Nutlets 1 – 4, obovate, 4 – 6 mm long, slightly wrinkled.

distribution. Thailand, Peninsular Malaysia, Malaysian Borneo (Sabah, Sarawak), Indonesia (Sumatra). Map 3.

specimens examined. thailand. Songkhla: Had Yai Distr., Ton Nga Chang Fall [6.945°, 100.2259°], 150 m, Oct. 1993, Vichai s.n. (BKF); Yala: Banang Sta Distr., 100 m, 26 July 1923, Kerr 7346 (BK, BM, K 2 sheets); Ban Lae, 21 Nov. 1961, Chantanamuck 23 (BK, BKF); Bo Hin Forest, 18 Nov. 1961, Suvarnakoset 1692 (BKF); Narathiwat: Khao Nikom Vaeng, 14 Sept. 1966, Sangkhachand & Nimanong 1363 (BKF, K, L, P); Waeng, Hala Bala Wildlife Sanctuary, 5°48'N 101°49'E, 380 m, 15 Aug. 2006, Petrmitr 667 (CMUB). malaysia. Kelantan: Pergau, Sungei Long Intake I [5.7106°, 101.7836°], 750 m, 6 Oct. 1992, Karim & Shah 31 (GH, SING); Pahang: Bentung, Genting Highland, 3°26'N 101°40'E, 1850 m, 4 July 1997, Saw FRI 44870 (SING); Bukit Fraser [3.7167°, 101.75°], 8 Aug. 1937, Franck 1346 (C); Bukit Fraser, Sungei Yel, 810 m [2700 ft], 28 Aug. 1923, Nur 11100 (SING); Cameron Highlands [4.45°, 101.4667°], 1200 m [4000 ft], 8 April 1937, Henderson 32572 (BM, GH); Fraser’s Hill [3.7167°, 101.75°], 1200 m [4000 ft], 12 Aug. 1937, Corner 33178 (GH, K, SING); Pulau Tioman [2.8°, 104.1833°], Bukit Bagoh, 240 m [800 ft], 28 April 1927, Nur 18576 (K, SING); Pulau Tioman, Kampung Teke [2.8°, 104.15°] to Kampung Juara, 22 Sept. 2000, Kiew 5108 (SING); Telom [4.45°, 101.5833°], Nov. 1908, Ridley 13595 (BM, SING); Perak: Temengoh, July 1909, Ridley 14446 (holotype of Cyrtandromoea repens, K; isotype BM, SING); Gunong Pondok [4.7667°, 100.8333°], 19 Oct. 1924, Haniff 10337 (SING); Grik, Temengor Dam [5.4°, 101.3°], Sungei Sera, 325 m, 10 Nov. 1993, Saw FRI 39974 (K, L); Gunong Batu Puteh [4.2167°, 101.45°], Lower Camp, Wray 1233 (lectoparatypes of G. curtisii, K, SING); Maxwell Hill [4.85°, 100.7833°], 300 m, Oct. 1899, W. F. 173 (SING); Thaiping [Taiping, 4.85°, 100.7333°], near the cottage, Feb. – March, Ridley s.n. (SING); Waterloo [4.7667°, 100.8167°], 600 m [2000 ft], Dec. 1887, Curtis 1310 (lectotype of G. curtisii, K; isolectotype SING); Selangor: Ulu Langat, Gadoh 1688 (KEP); Sabah: Keningau Distr., Crocker range [5.4666°, 116.0613°], 21 Aug. 1986, Krispinus SAN 113847 (K); Crocker range Forest Reserve, mile 11, Keningau-Kimanis Road, 18 Nov. 1978, Gibot SAN 87949 (K, L); Crocker range, 19 km W of Keningau, mile 13.5, Kimanis Road, 1100 m, 23 Sept. 1983, Beaman et al. 7037 (K); mile 8 of Keningau-Kimanis Road, 30 Sept. 1981, Gibot & Sundaling SAN 94250 (K, L); Lahad Datu Distr., Hutan Simpan Sungai Segama, 28 March 1985, Madani & Ismail SAN 108657 (K); Tabin Wildlife Reserve near Wildlife Department's headquarters, 5°11'N 118°30'E, 150 m, 24 Feb. 2000, Kjeldsen et al. 34 (AAU, C); Nabawan Distr., Pensiangan, Sungei Sansiang, a tributary of the Sepulut R., Batu Punggul, 4°38'N 116°38'E, 325 m, 29 April 1984, Sands et al. 4049 (K); Ranau Distr., Kampung Menggis [5.3833°, 115.4167°], 11 Sept. 1996, Rumutom 393 (K). malaysia. sarawak. Mt Poi, 1929, Clemens & S. Clemens 20175 (K); Foot of Gunong Berumput, Poi Range, 9 Aug. 1962, Burtt & Woods B2744 (E); malaysia. unlocalised. Scortechini 924 (lectoparatype of G. curtisii SING). indonesia. sumatra. Aceh: Lawe Sigala, near Kutatjane, 23 March 1954, Alston 14680 (BM, GH); North Sumatra: NE of Mt Sibajak, near Bandarbaru, 950 – 1100 m, 9 Oct. 1928, Lorzing 14079 (K, L); By Sibolangit [3.3167°, 98.5833°], 525 m, 20 Jan. 1918, Lorzing 5498 (L); Coven Bandar, Caroe, Sibolangit, 1000 m, 3 Feb. 1917, Lorzing 4670 (L); E Coast [Siantar, Simeloengoen, 1.9667°, 98.7833°], [2 Aug. 1923], Yates 895 (BM, K); West Sumatra: Muaralabuh [-1.4833°, 101.05°], Peconina, 900 m, 15 Sept. 1929, Teruya 1406 (K, SING 2 sheets); Mt Sago [-0.3269°, 100.6706°], near Pajakumbuh, above Halaban Moutain Forest, 800 m, 3 Dec. 1955, Meijer 4504 (L); NW Kelling, Gunung Talamau, 600 m, 28 April 1917, Bünnemeijer 505 [L (L0796120)].

habitat. In lowland forest, montane dipterocarp forest, evergreen forest, stream sides, forest edges and roadsides; alt. 100 – 1350 m.

conservation status. Widespread; Least Concern (LC).

phenology. Flowering April – November. Fruiting October – March.

notes. Gomphostemma curtisii is a recognised by having subulate calyx lobes, broadly ovate or cordate leaf blades and axillary inflorescences subtended by leaves or in a lower, more basal position where leaves have fallen. Prain agreed with Mr W. B. Hemsley who remarked that this species is similar to G. pedunculatum by having pedunculate inflorescences and large bracts (Prain 1891).

12. Gomphostemma flexuosum Bongch. sp. nov. Type: Thailand, Chaiyaphum, Phu Khiew Wildlife Sanctuary, 10 km from Headquarters to main road, 800 m, 11 Oct. 1998, Suddee et al. 955 (holotype K [K001081924]; isotypes BKF, TCD).

http://www.ipni.org/urn:lsid:ipni.org:names:77221534-1

Sprawling herb. Stems slender, scrambling, obtusely quadrangular with shallowly longitudinal grooves, tomentose with stellate hairs; internodes of leafy and flowering shoots appearing wavy, always twisting. Leaves petiolate or subsessile, spreading, chartaceous; blades elliptic, elliptic-ovate or oblanceolate, 5 – 13 × 2.5 – 8 cm, apex acute, margin serrate to subentire, base cuneate or attenuate, upper side scabrid with stiff hairs, lower side pale green tomentose with stellate hairs with a 1-celled stalk; petiole 5 – 20 mm long. Inflorescence axillary with opposite cymes sessile, unbranched then congested with axis not visible between flowers; verticils 2 – 10-flowered, inserted at the upper nodes at which leaves are usually present; bracts oblanceolate, 6 – 15 × 2 – 5 mm, with dense stellate hairs on both sides; bracteoles linear or sometimes falcate. Flowering calyx infundibular, 7 – 10 mm long, 10-ridged, outside with dense stellate hairs with a 1-celled stalk and dendroid hairs, inside with simple hairs; tube 3 – 4 mm long; lobes narrowly triangular, 4 – 6 mm long, apex acuminate. Fruiting calyx infundibular, 10 – 12 mm long; tube 4 – 5 mm long; lobes triangular, 6 – 7 mm long. Corolla white, 15 – 20 (– 23) mm long, throat inflated, abruptly dilated near the throat, outside with sessile stellate hairs, inside glabrous; tube slightly incurved, 12 – 17 mm long; posterior lobe shorter than anterior lobes, outside with sessile stellate hairs and glandular-like simple hairs, inside sparsely tomentose near throat; anterior lobes spreading; median lobes broadly ovate, 4 – 5 mm long, apex slightly emarginate; lateral lobes ovate. Staminal filaments fleshy, glabrescent. Disk lobed, slightly tumescent behind. Style slender with subequally lobed apex. Nutlets solitary, immature seen. Fig. 2.

recognition. Gomphostemma flexuosum is recognised by having sprawling habit, flowering shoots appearing wavy with curving internodes, calyx lobes longer than the tube, short corolla, 15 – 20 (– 23) mm long and found in open and dry areas. It is morphologically similar to G. melissifolium by having an incurved corolla tube, coarse indumentum on the upper side of leaves and inflated corolla throat.

distribution. Vietnam, Laos, Thailand. Map 3.

specimens examined. vietnam. Dong Nai: Langahn [Long Khanh], Station de Langanh, Du Haut Danai, 1000 – 1200 m, 3 Oct. 1932, Poilane 21050 (P). laos. Champasak: Mekong, Se Don, 1866 – 1868, Thorel 2441 (P 6 sheets). thailand. Sakon Nakhon: Phu Phan National Park, 17°03'N 103°58'E, 220 m, 20 Oct. 1998, Suddee et al. 1014 (K); ibid., Chantaranothai et al. 728 (BKF); Chaiyaphum: Phu Khiew Wildlife Sanctuary, 10 km from Headquarters to main road, 800 m, 11 Oct. 1998, Suddee et al. 955 (holotype K; isotypes BKF, TCD); Nakhon Ratchasima: Pak Thong Chai, Sakaerat Forest Research Station, along Huai Krae Stream, 14°45'N 102°E, 400 m, 1 Nov. 1969, van Beusekom & Charoenpol 1992 (AAU, BKF, L); Ban Kaeng Khlo, 200 m, 17 Sept. 1971, Vidal 5118 (AAU, P); Buri Ram: Khao Phanom Rung, 14°32'N 102°56'E, 3 Oct. 1984, Murata et al. T-37345 (BKF); Saraburi: Muang, Sahm Lahn Forest, 150 m, 15 Dec. 1973, Maxwell 73-755 (AAU, BK); Ban Nawng Bua [14.5880°, 100.7634°], 2 Oct. 1927, Put 1136 (BK, BM, GH, L, SING).

habitat. In dry dipterocarp, fire-influenced open forest; alt. 150 – 1200 m.

conservation status. This species has been collected from several conservation areas in Thailand. Least Concern (LC).

phenology. Flowering September – December.

notes. The epithet flexuosum [flexuosum, zigzag] refers to its wavy stem.

13. Gomphostemma grandiflorum Doan ex Suddee & A.J.Paton (2006: 620); Doan (1936: 1034); Phuong (2000: 105). Type: Vietnam, Tonkin, Thua-luu (Thuu Thien-Hue), Lecomte & Finet 1320 (holotype P [P00344471]; isotype P [P00344472].

Herb up to 2 m tall. Stems erect, robust, obtusely quadrangular with longitudinal grooves, with stellate hairs; internodes straight. Leaves petiolate, coriaceous; blades ovate or ovate-oblong, 6 – 25 × 3 – 12 cm, apex acute, margin shallowly dentate-serrate, base cuneate or oblique, upper side tomentose with 2 – 4-celled simple hairs, sometimes bifurcate hairs, lower side pale green tomentose with dense stellate hairs; petioles 15 – 70 mm long. Inflorescence axillary with opposite cymes subsessile, congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present, subtended by the upper leaves only; verticils (5 –) 10 – 20-flowered; bracts deciduous, elliptic-ovate, 10 – 25 × 5 – 12 mm, apex acute, margin shallowly dentate at the apical part and entire at the basal part, base cuneate, with stellate hairs on both sides; bracteoles linear, 7 – 12 mm long. Flowering calyx infundibular, 10 – 12 mm long, thickly 10-ridged, outside with dense stellate hairs with a 1-celled stalk, inside glabrous below and adpressed simple hairs above; tube 5 – 7 mm long; lobes broadly triangular, c. 5 mm long, apex acute, margin slightly revolute. Fruiting calyx ovate-cupular, 15 – 18 mm long; tube 8 – 10 mm long; lobes broadly triangular, 7 – 8 mm long, apex acuminate, margin revolute in cross section. Corolla white, 30 – 50 mm long, throat much inflated, abruptly dilated near the throat, outside with stellate hairs, inside glabrous; tube very slender, incurved, 20 – 40 mm long; posterior lip obovate, 5 – 10 mm long, apex round; anterior lip 5 – 10 mm long; median lobe folding outward, obovate, 12 – 14 mm long, apex and margin undulate; lateral lobes broadly ovate, apex and margin undulate. Staminal filaments fleshy, tomentose. Style slender, 30 – 45 mm long, glabrous with apex unequally lobed. Nutlets 1 – 2, ovate or elliptic-ovate, 5 – 9 × 4 – 6 mm.

distribution. Vietnam. Map 3.

specimens examined. vietnam: Thua Thien Hue: Thua Luu [16.3°, 108°], 18 Nov. 1911, Lecomte & Finet 1320 (holotype P; isotype P); Thua Thien, Bach Ma [16.2°, 107.8667°], 20 Aug. 1941, Vidal 690A (P); Da Nang: Ba Na Nui Chua, 15°59'9.37'N 107°59'6.7”E, 21 Oct. 2008, Du et al. HNK 3193 (K); Tourane [Da Nang], Mt Bani [16.1224°, 107.9832°], May – July 1927, Clemens & S. Clemens 3885 (G, K, P); Quang Nam: Confin sud de la province de Quang Nam entre les villages Aboi de Maung Tra et Dai-lo, 1500 m, 27 Feb. 1941, Poilane 31881 (P); Kon Tum: NW Slope of Ngoc Linh mountain system [15.0667°, 107.9833°], 1600 – 1800 m, 23 Feb. 1995, Averyanov et al. VH 252 (AAU, P).

habitat. In hill evergreen forest, stream margins and open secondary forest in shady areas; alt. 1500 – 1800 m.

conservation status. Gomphostemma grandiflorum is known from a few collections but its habitat is clearly threatened by agriculture and rapid development projects in Vietnam (Paton et al. 2016): the EOO and AOO of this species are 1060.77 and 24 km², respectively. Hence, the Endangered category is applied. (EN B1ab(i, ii, ii) and B2ab(i, ii, iii)).

phenology. Flowering May – November. Fruiting October – February.

notes. Gomphostemma grandiflorum is a distinctive species by having large corolla, 30 – 50 mm long, and leaf 60 – 250 × 30 – 120 mm, calyx lobes not longer than the tube, a congested inflorescence and fruiting calyx lobes revolute in cross section. Gomphostemma leptodon also has similar morphological characters, large flowers and leaves, but its calyx lobes are mostly longer than the tube.

14. Gomphostemma hainanense C.Y.Wu (1965: 145); Li & Hedge (1994: 72). Type: China, Hainan, Po-ting, herb in forest, 660 m [2200 ft], 27 Aug. 1935, How 73517 (holotype PE; isotypes BM [BM000950531], G [G00188991], GH, IBSC).

Perennial herb up to 1.2 m tall. Stems erect, obtusely quadrangular with longitudinal grooves, with dense stellate hairs. Leaves petiolate, chartaceous; blades narrowly oblong, ovate-oblong or elliptic-oblong, 8 – 16 × 2 – 4 cm, apex acute, margin shallowly serrate-entire, base attenuate, upper side dull green with dense sessile stellate or stellate hairs, lower side whitish or creamy-white tomentose with dense stellate hairs; petioles (5 –) 10 – 20 mm long. Inflorescence axillary with opposite cymes sessile, unbranched then congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils few-flowered; cluster of small leaf-like bracts, deciduous, ovate-elliptic, 15 – 20 mm long, upper side dull green with dense sessile stellate, lower side with dense white or creamy-white stellate hairs; bracteoles linear or spathulate, 5 – 10 mm long. Flowering calyx infundibular, 7 – 10 mm long, inconspicuously 10-ridged, outside with dense stellate hairs, inside with simple hairs; tube 4 – 5 mm long; lobes narrowly triangular, 3 – 5 mm long, apex acute. Fruiting calyx 13 – 15 mm long; tube 5 – 6 mm long; lobes triangular, 8 – 9 mm long. Corolla pale yellow or white, 25 – 35 mm long, throat inflated, abruptly dilated near the throat, outside tomentose with stellate hairs, inside glabrous; tube slender, slightly incurved, 20 – 25 mm long; posterior lip 3 – 6 mm long, apex emarginate; anterior lip 3-lobed, unequal; median lobe elliptic, apex acute, margin undulate; lateral lobes undulate. Staminal filaments slightly fleshy. Style slender, slightly shorter than corolla, glabrous with apex subequally lobed. Nutlets solitary, ovate, 4 – 5 mm long, smooth with sparse short hairs.

distribution. China (Hainan), Vietnam. Map 3.

specimens examined. china. Hainan: Po-ting, 660 m [2200 ft], 27 Aug. 1935, How 73517 (holotype PE; isotypes BM, G, GH, IBSC, P). vietnam. Quang Ninh: Tien-yen [21.333°, 107.4°], Ho Yung Shan and vicinity, 13 Oct. – 22 Nov. 1940, Tsang 30705 (P); Nghe An: Vinh Province [now in Nghe An Province], Bu Khang [19.4333°, 105.1667°], 700 m, 11 Aug. 1929, Poilane 16692 (P 3 sheets); Da Nang. Tourane [Da Nang, 16.0678°, 108.2208°], Jan. 1837, Gaudichaud 127 (P); ibid., 900 m, 18 Sept. 1923, Poilane 8069 (P).

habitat. In mountain areas; alt. 660 – 700 m.

conservation status. Gomphostemma hainanense is known from four localities based on five collections. There is a decline of suitable habitats in Vietnam which is caused by agriculture and development projects (Paton et al. 2016). Because of the low AOO (20 km²) and only five collections, the Endangered (EN) category is applied (EN B2ab(ii)).

phenology. Flowering August – December. Fruiting October – January.

notes. Gomphostemma hainanense is recognisable by its leaves being whitish or creamy-white tomentose on the lower side and by their narrow shape. Doan (1936) recognised G. niveum in Flore Général de L’Indo-Chine but those specimens examined by Doan are actually G. hainanense. These two species have a similar white tomentose indumentum on the leaf lower surface. However, G. hainanense has narrow leaves, 20 – 40 mm wide, and its corolla is much exserted from the calyx while G. niveum has broader leaves, 50 – 120 mm wide, and the corolla is hardly exserted from the calyx.

15. Gomphostemma hirsutum Walsingham (in Walsingham & Bramley 2010: 480); Walsingham (2019: 199). Type: Malaysia, Sabah, Ranau Distr., Kampung Menggis, 29 Aug. 1995, Rumtom [Rumutom] 122 (holotype K [K000479761]; isotype KEP n.v.).

Herb up to 1 m tall. Stems dark brown or black, erect, obtusely quadrangular with longitudinal grooves, densely tomentose with a mixture of dendroid hairs with multi-celled stalk, stellate hairs with a 1-celled or multi-celled stalk and multi-celled simple hairs. Leaves petiolate, chartaceous; blades elliptic-obovate, narrowly elliptic or narrowly elliptic-oblanceolate, 12 – 26 × 3 – 8 cm, apex acute, margin shallowly dentate, base attenuate; upper side with simple hairs; lower side pale green tomentose with a mixture of branched hairs confined to midrib; petioles 10 – 30 mm long. Inflorescence axillary with opposite cymes sessile, unbranched then congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils 1 – 3-flowered; bracts narrowly ovate or straight falcate, 6 – 10 × 1 – 2 mm, upper side adpressed hirsute or sometimes with bifurcate hairs, lower side with stellate hairs and/or simple hairs; bracteoles linear, 3 – 6 mm long. Flowering calyx infundibular, 10 – 13 mm long, 10-ridged, outside with long simple hairs and long branched hairs, inside adpressed simple hairs; tube 5 – 7 mm long; lobes triangular, 5 – 6 mm long, apex acuminate. Fruiting calyx not seen. Corolla white or creamy-white, 25 – 45 mm long, throat inflated, abruptly dilated near the throat, outside tomentose with stellate hairs, inside glabrous; tube slender, incurved, 20 – 35 mm long; posterior lip ovate, 5 – 8 mm long, apex and margin slightly undulate; anterior lip 3-lobed, longer than posterior lip; median lobe 5 – 9 mm long, apex and margin slightly undulate; lateral lobes shorter than median lobe. Staminal filaments fleshy, sparse tomentose. Style slender, slightly shorter than corolla with apex subequally lobed. Nutlets not seen [round white berries: Darnton 185 (BM)].

distribution. Malaysian Borneo (Sabah). Map 3.

specimens examined. malaysia. sabah. Ranau Distr.: 510 m [1700 ft], 16 Feb. 1954, Darnton 185 (BM); Bukit Kulung near Bukit Hampuan at SE base of Mt Kinabalu, 5°59'N 116°41'E, 750 m, 8 Dec. 1983, Beaman 7778 (US); near Poring Hot Springs, Kipungit-Langanan Falls trail, 6°3'N 116°42'E, 650 – 750 m, 6 Aug. 1990, Beaman 10893 (K); Kampung Takutan [6.1167°, 116.7167°], 4 Sept. 1995, Tadong 524 (K); Kampung Menggis, 29 Aug. 1995, Rumutom 122 (holotype K); Lahad Datu Distr.: Borneo Rainforest Lodge, Hornbill trail, Borneo Rainforest Lodge, 4°59'40”N 117°50'11”E, 6 July 2006, de Kok et al. SAN 147234 (K); Danum Valley Road, near km 45 [5.4174°, 118.3927°], 10 June 1989, Campbell EC9 (E).

habitat. In shade and hilly areas of secondary forest, disturbed primary rainforest, lowland forest, dipterocarp forest; alt. 510 – 750 m.

conservation status. Endangered (EN B1ab(iii)) (Walsingham & Bramley 2010).

phenology. Flowering July – September. Fruiting August – February.

notes. Gomphostemma hirsutum is a distinctive species recognised by having long white simple hairs, stellate hairs, and dendroid hairs on the flowering calyx, narrow bracts, 1 – 2 mm wide, subtending cymes. It is endemic to Borneo (Sabah). However, the Chinese species, G. stellatohirsutum C.Y.Wu is similar to the Borneo species by having dark or dark brown stems with long dendroid hairs but lacks long simple hairs on the flowering calyx.

16. Gomphostemma javanicum (Blume) Benth. (Bentham 1835: 650); Prain (1891: 261); Keng (1969: 86, 1978: 322); Murata (1976: 188); Phuong (2000: 111); Walsingham & Bramley (2010: 483); Walsingham (2019: 199). Type: as for Prasium javanicum.

Prasium javanicum Blume (1826: 840). Type: Indonesia, Java, Mont. Gede [-6.7994°, 106.9689°], Blume s.n. (lectotype L (L0003700), selected by Keng (1969); isolectotypes 4 sheets [L0003696, L0003697, L0003698, L0003699]).

Prasium phlomoides Reinw. ex Blume (1826: 840). Type: Indonesia, Java, Reinwardt s.n. (lectotype L [L0003694], selected by Keng (1969); isolectotypes 2 sheets L [L0003693, L0003695]).

Gomphostemma lucidum Wall. ex Benth. (Bentham in Wallich 1830b: 12). Type: India, Pundua [Pandua, 25.1167°, 91.7333°], May 1822, de Silva s.n. in Wall. Cat. 2156.2 (lectotype K [Herb. Benthamianum], selected here; isolectotype K-W [K001115340]).

Gomphostemma oblongum Wall. ex Benth. (Bentham in Wallich 1830b: 12). Type: Burma, Tavoy, 12 Aug. 1827, Gomez s.n. in Wall. Cat. 2154 (lectotype K [Herb. Benthamianum, K000501086]; isolectotype K-W [K001115337]).

Gomphostemma phlomoides (Reinw. ex Blume) Benth. (Bentham 1835: 649). Type: as for Prasium phlomoides.

Gomphostemma philippinarum Benth. (Bentham in A.P. de Candolle 1848: 551). Type: Philippines, Island of Bohol [9.8333°, 124.1667°], 1841, Cuming 1840 (K [Herb. Benthamianum, K000856198] lectotype selected here; isolectotypes C, E [E00301969], G [G00189007], K [Herb. Benthamianum, K000856200]).

Gomphostemma oblongum var. helferi Hook.f. (Hooker 1885: 697). Type: Burma, Tenasserim, Helfer 4043 (holotype K [Herb. Hookerianum, K000846319]).

Gomphostemma scortechinii Prain (1890: 315). Type: Malaysia, Perak, Gunong Tujok, Sept. 1884, Scortechini 1225 [CAL [CAL0000007151], lectotype selected here; isolectotypes 3 sheets CAL [CAL0000007150, CAL0000007152, CAL0000007153], K [K00050182], SING).

Gomphostemma scortechinii var. helferi Prain (1891: 260). Type: as for G. oblongum var. helferi Hook.f.

Gomphostemma rugosum Prain (1891: 262). Type: Burma, Tenasserim, Chapadong [Chappedong] Hill, 23 Feb. 1827, Wall. Cat. 2156.1 (K-W [K001115339], lectotype selected here; isolectotype CAL n.v.).

Gomphostemma lucidum Wall. ex Benth. var. canescens Prain (1891: 263). Type: India, Lakhimpur, Griffith 4038 (K [Herb. Hookerianum], lectotype selected here; isolectotypes C, CAL n.v., P [P04061943]).

Gomphostemma intermedium Craib (1910: 23). Type: Thailand, Chiang Mai, Doi Sootep, in evergreen jungle, 730 m, 17 July 1909, Kerr 733 (K, lectotype selected here; isolectotypes ABD, BM).

Gomphostemma cinereum Elmer (1919: 3086). Type: Philippines, Luzon, Province of Laguna, Los Banos (Mt Maquiling), along stream beds of dense forests half way up the mountain, June – July 1917, Elmer 17836 (K [K000856203], lectotype selected here; isolectotypes 2 sheets BM [BM000950534, BM000950535], L [U0226559], NY [00000532] , P, US [00119424]).

Gomphostemma oblongum var. setosum Ridl. (Ridley 1923a: 653). Type: Malaysia, Selangor, Semangkok Pass [3.6833°, 101.75°], 22 Jan. 1921, Ridley s.n. (holotype K [K000501134]).

Gomphostemma lacteum Ridl. (Ridley 1924: 300). Type: Malaysia, Pahang, Pine Tree Hill [3.7167°, 101.6833°], near Fraser Hill, 1440 m [4800 ft], 27 Aug. 1923, Burkill 11075 (holotype K [K000501135]; isotype SING).

Gomphostemma lucidum var. intermedium (Craib) C.Y.Wu (1965: 145). Type: as for G. intermedium Craib.

Gomphostemma arbusculum C.Y.Wu (1965: 146). Type: China, Yunnan, Lan-tsang, Lan-tsang Hsien [22.5333°, 99.9333°], 1800 m, May 1935, Wang 76475 (holotype PE; isotype A n.v.).

Gomphostemma latifolium C.Y.Wu (1965: 147). Type: China, Yunnan, Jinghong, Che-li Hsien, 950 m, Aug. 1936, Wang 75550 (holotype PE; isotype A n.v.).

Gomphostemma javanicum f. phlomoides (Reinw. ex Blume) Murata (1976: 189). Type: as for Prasium phlomoides Reinw. ex Blume.

Gomphostemma javanicum var. helferi (Hook.f.) Murata (1976: 189). Type: as for G. oblongum var helferi Hook.f.

Gomphostemma javanicum subsp. lucidum (Wall. ex Benth.) Murata (1976: 189). Type: as for G. lucidum Wall. ex Benth.

Gomphostemma javanicum var. rugosum (Prain) Murata (1976: 190). Type: Gomphostemma rugosum Prain.

Gomphostemma javanicum var. brevidens Murata (1989: 101). Type: Thailand, Chiang Mai, Doi Inthanon, in evergreen forest, 1800 – 2500 m, 3 Oct. 1971, Murata et al. T-15988 (holotype KYO [KYO00020216]; isotypes BKF [SN029708], KYO [KYO00020217]).

Perennial herb 0.5 – 2 m tall. Stems erect, upright, woody at base, obtusely quadrangular with longitudinal grooves, with dense stellate hairs with a 1-celled or multi-celled stalk and sometimes dendroid hairs. Leaves petiolate, chartaceous or coriaceous; blades oblong-ovate, elliptic, elliptic-oblong, oblong, elliptic-ovate, elliptic-obovate or obovate, 7 – 30 × 3 – 13 cm, apex acute or broadly acuminate, margin entire, subentire, serrate, obtusely serrate or crenate, base cuneate, attenuate or obtuse, upper side tomentose with a mixture of soft hairs, lower side pale green tomentose with a mixture of dense branched hairs, sometimes with robust venation, rugose-like; petioles 10 – 50 mm long, with stellate hairs and/or dendroid hairs. Inflorescence axillary with opposite cymes sessile or sometimes pedunculate, branched then lax or unbranched then congested with axis not visible between flowers, exceeding 20 mm wide when pressed, excluding corolla, inserted at the upper nodes at which leaves are usually present; verticils few – many-flowered, subtended by upper leaves only; pedicels 0 – 10 (– 15) mm long; bracts deciduous, slightly falcate, ovate, narrowly obovate, obovate or spathulate, 5 – 15 × 3 – 7 mm, upper side with shortly simple hairs, lower side with stellate hairs; bracteoles linear, 4 – 10 mm long. Flowering calyx infundibular, 7 – 15 mm long, prominently 10-ridged, outside dense tomentose with stellate hairs with a 1-celled stalk or sessile stellate hairs or short simple hairs, inside with short simple hairs; tube 5 – 10 mm long; lobes triangular, 2 – 5 mm long, apex acute or acuminate. Fruiting calyx broadly infundibular or cupular, 10 – 20 mm long; tube 6 – 15 mm long; lobes 4 – 8 mm long, longer than the tube at maturity or not longer than the tube; apex acute or acuminate, margin flat in cross section. Corolla white, pale yellow, pale pink or scarlet, 25 – 40 (– 50) mm long, throat inflated, abruptly dilated near the throat, outside tomentose with stellate hairs or simple hairs, inside glabrous; tube incurved, 15 – 35 mm long; posterior lip ovate, 3 – 5 mm long, apex emarginate or undulate, margin undulate or entire; anterior lip 3-lobed, spreading beyond posterior lip, 7 – 15 mm long; median lobe tongue-like, spreading, broadly ovate, apex and margin undulate, base constricted. Staminal filaments fleshy, tomentose. Style fleshy, glabrous, shorter than corolla with apex subequally lobed. Nutlets 1 – 4, elliptic or oblong, 4 – 6 × 3 – 4 mm, sparse tomentose or glabrescent at apex, wrinkled. Fig. 1D & 5.

distribution. India (Arunachal Pradesh, Assam, Meghalaya, Andaman and Nicobar Islands), Bangladesh, China (Yunnan, Hainan, Guangdong), Burma, Thailand, Vietnam, Laos, Cambodia, Peninsular Malaysia, Malaysian Borneo (Sabah, Sarawak), Brunei, Indonesia (Sumatra, Java, Sulawesi), Philippines (Luzon, Mindanao, Visayas, Sulu Archipelago). Map 4.

Map 4

Distribution range of Gomphostemma javanicum.

specimens examined. india. Arunachal Pradesh. Nizamghat [28.2667°, 95.7°], Ward 8734 [in parte] (K); Lohit Valley, 28°N 96°35'E, 600 – 900 m [2000 – 3000 ft], 3 Dec. 1926, Ward 7689 (K); Lakhimpur, Griffith 4038 (isolectotypes of G. lucidum var. canescens, C, K [Herb. Hookerianum], P); Assam. Griffith 290 (BM), 1014 (BM) & 1850 (K [Herb. Hookerianum]); ibid., Masters s.n. (lectoparatype G. lucidum var. canescens, K [Herb. Hookerianum]); Seelsagar, Borpatta [Barpeta, 26.5333°, 91°], 90 m [300 ft], 13 Oct. 1885, Clarke 40763A (BM); Meghalaya. Jaintea, 1200 m [4000 ft], 19 Oct. 1867, Clarke 5950 (K); Khasia, 1200 – 1500 m [4000 – 5000 ft], Hooker & Thomson s.n. (G, K 2 sheets [Herb. Hookerianum], P); Khasia Hills, Griffith s.n. (L, P); Khasia, Churra, 1200 – 1500 m [4000 – 5000 ft], 13 Aug. 1850; Hooker & Thomson 2078 (K [Herb. Hookerianum]); Khasia, Mammai, 1050 m [3500 ft], 9 May 1886, Clarke 43730 (G); Khasia, Maodadoom, 900 m [3000 ft], 3 Feb. 1886, Clarke 42936 (BM); Cherrapunjee, 1200 m [4000 ft], 12 May 1952, Koelz 29855 (E); andaman and nicobar islands. South Andaman, Ranga Chang [11.5833°, 92.75°], 1 April 1891, Prain s.n. (G); South Andaman, Bajajagdah Hill, 18 Aug. 1894, King's Collector s.n. (P); Middle Andamans, camp no. 15, 24 km from Rangat [12.50994°, 92.912°], 25 m, 3 Nov. 1977, Bhargava & Nooteboom 6311 (CAL, L); Mt Harriet, Port Blair [11.6667°, 92.75°], 20 Sept. 1884, Kunstler 454 (BM, K). bangladesh. Sylhet. Pundua, May 1822, de Silva s.n. in Wall. Cat. 2156.2 (lectotype of G. lucidum, K [Herb. Benthamianum]; isolectotype K-W); unlocalised. Griffith 4038 (C, K [Herb. Hookerianum], P). china. Yunnan. Foo-ning [23.6247°, 105.6272°], Jar-gui, 700 m, 31 May 1940, Wang 89448 (KUN); Hekou, Nanxi [22.6403°, 103.9328°], Machang to Liangzi, 700 m, 6 May 2004, Shui & Chen 40883 (KUN); Kuen-ger, Che-li Hsien [21.9833°, 100.8167°], 1200 m, Oct. 1936, Wang 79289 (KUN); Lan-tsang Hsien [22.5333°, 99.9333°], 1800 m, May 1936, Wang 76475 (holotype of G. arbusculum, PE); Mengla [21.4667°, 101.5833°], 1500 m [5000 ft], Henry 11046 (K); Meng-soong, Dah-meng-lung, Che-li Hsien, 1300 m, Sept. 1936, Wang 77982 (KUN); Ping-bien Hsien [22.9°, 103.6667°], Hwang-tian-bien, 1500 m, 7 Oct. 1939, Wang 82349 (KUN); Szemao [22.7667°, 101.0833°], 1350 m [4500 ft], Henry 12609 (K); Jinghong, Che-li Hsien, 950 m, Aug. 1936, Wang 75550 (holotype of G. latifolium, PE); Hainan. Lam Ko, Cheung Wo and vicinity, 8 Aug. 1927, Tsang 400 (GH); ad pagum Fan lun, territ, indigenarum Lai diatorum, 15 Nov. 1882, Henry 153 (BM); Hong Fa, July 1893, Chinese Collector 464 (K); Lai area, Hung Mo Shan and vicinity, 4 June 1929, Tsang & Fung 247 (GH, K); Ling Shui [18.5036°, 110.0342°] (Ling-tui), Chim Shan, Fan Maan Ts'uen and vicinity, 3 – 20 May 1932, Fung 20110 (GH, K, P); Lingshuixian, Nanqiaoqu Shuangximu zi Dacong, 650 m, Anonymous s.n. (AAU); Loktung, 2 July 1936, Lau 27392 (GH, KUN); Po-ting [18.6403°, 109.6994°], 180 m [600 ft], 14 April 1935, How 71890 (GH, P); Tai Pin, foot of Loi Mother Mt, 290 m, 20 July 1935, Gressitt 1062 (BM, E, G, GH); Tamm-chau [19.7113°, 109.3147°], Nodoa and vicinity, 20 July 1927, Tsang 109 (G, K); Tingon [19.6997°, 110.3183°], 31 Oct. 1936, Lau 28123 (KUN 2 sheets); Yaichow, 16 July 1933, Liang 62115 (E, K); Ban-chiou-chian, Che-li Hsien [21.9833°, 100.8167°], 840 m, Oct. 1936, Wang 79725 (KUN); Ching-tung [24.4667°, 100.9°], 1700 m, 30 Oct. 1939, Li 950 (KUN); Guangdong. 28 July 1937, Liang 69878 (IBSC); Fang Ch'eng, T'aan Faan, Kung P'ing Shan and vicinity, 10 – 18 Sept. 1936, Tsang 26825 (C, E, GH, IBSC, K, P). myanmar. Tanintharyi. Eastern Tenasserim [11.8°, 99.25°], 50 m, 31 May 1932, Kerr 21645 (BK, BM, K, L); Tenasserim [12.0833°, 99.0167°], Griffith 4041 (K [Herb. Hookerianum]); ibid., Helfer 4041 (K [Herb. Hookerianum], P); ibid., Helfer 4043 (holotype of G. oblongum var. helferi, K [Herb. Hookerianum]); Tavoy, 12 Aug. 1827, Wall. Cat. 2154 (holotype of G. oblongum, K [Herb. Benthamianum]; K-W); Tenasserim, Tavoy, Hill W of Paungdaw Power Station, 14°N 98°30'E, 630 – 690 m [2100 – 2300 ft], Oct. 1961, Keenan et al. 1605 (E); Mon. Chapadong Hill, 23 Feb. 1827, Wall. Cat. 2156.1 (isolectotype of G. rugosum, K-W); Mandalay. Maymyo [22.0333°, 96.4667°], 1110 m [3700 ft], 31 July 1925, Parkinson 1595 (K); Shan. S Shan States, Keng Tung [Keng Tang, 21.3°, 99.65°], 1500 m [5000 ft], June 1909, MacGregor 594 (E). vietnam. Da Nang. Sur de Tourane, June 1909, D'alleizette s.n. (L); Ba Na pres Tourane, 1500 m, 8 July 1923, Poilane 6931 (P); Ravine du Massif Mautaguong de la pres qu' la de Tourane, May – June, Couderc s.n. (P 2 sheets); Dak Lak. Krong Pak, Hoa lo, 23 Dec. 1979, Phuong 699 (HN 2 sheets); Dong Nai. ad Gia lau me in prov Bien-hoa [10.95°, 106.8167°], Sept. 1869, Pierre s.n. (K, P); Ha Tay. Mont Bavi, 300 m, May 1887, Balansa 2934 (K, P); Mont Bavi, pres de la pagode de Dein Touan, June 1887, Balansa 2933 (K, P 3 sheets); Hoa Binh. Mai chau, Pa co, Ban Phuong, Khoi, Binh, 14 June 1999, Phuong 1761 (HN 4 sheets); Hang Tram [20.417°, 105.7°], 5 May 1970, Lien s.n. (HN); Quang Ninh. Ha-coi [21.4401°, 107.7619°], Chuk-phai, Taai Wong Mo Shan and Vicinity, 16 – 22 Oct. 1936, Tsang 27020 (C, E, K, P); Ha-coi, Shui Mei Village, NE of Chuk-phai, Taai Wong Mo Shan and vicinity, 23 June – 31 Aug. 1939, Tsang 29398 (BKF, C, E, G, K, L, P, SING); Lam Dong. Di Linh, Dinh Trang Thuong [11.7628°, 107.9503°], 16 May 1980, Nguyen 28 (HN); Duc Trong Distr., Xa Hiep An (Elephant Mountain), 11°50'35”N 108°25'43”E, 1500 – 1600 m, 31 Aug. 2001, Gardner et al. 80 (E); Lao Cai. Kilo 16, route de Ta Phinh Chapa [Sa Pa], 1450 m, 3 Aug. 1926, Poilane 12804 (P 3 sheets); Chapa [Sa Pa, 22.35°, 103.8667°], 1400 – 1500 m, 6 Dec. 1913, Chevalier 29410 (P 2 sheets); Nghe An. Co Ba (Ke Nhe), Chevalier 32420 (P 2 sheets); Thua Thien-Hue. Hue and vicinity, May – July 1927, Clemens & S. Clemens 3695 (G 2 sheets, K, P); Son La. Ngoc Chien, Ban Pieng, 21°35'45.11”N 104°15'34.4”E, 3 Oct. 2008, Du et al. HNK 2302 (K). laos. Vientiane. Pu Bia [19°, 103.1667°], 1500 m, 15 April 1932, Kerr 21078 (K 2 sheets, P); Champasak. Khong [14.1167°, 105.85°], Khon Island, Southern tip, Hill at Hang Khong Island, Mekong R., 125 m, 14 Oct. 1997, Maxwell 97-1167 (CMUB); Pak Song [15.1833°, 106.2333°], July 1949, Vidal 1044B (P); Salavan. Ban Munh Pak Di [15.4333°, 106.2°] & Ban Honh, 24 Sept. 1928, Poilane 15718 (P); unlocalised. Cahn Trap, Spire 1353 (P 2 sheets). cambodia. Angkor [13.4167°, 103.8667°], Godefroy-Lebeuf s.n. (K); montibus Tamir in Prov [Province] Tpong, 800 m, May 1870, Pierre 1181 (P). thailand. Chiang Mai. Chom Thong, Doi Song Mea, 1500 m, 21 Aug. 1999, Watthana et al. 599 (QBG); Chom Thong, Mae Soi Subdistr., Mae Soi Ridge, near Ban Bah Gluay (Meo Village), 1475 m, 12 Aug. 1991, Maxwell 91-737 (E, L, P); Doi Angka [Doi Inthanon, 18.5833°, 98.4833°], 1200 m, 18 July 1922, Kerr 6347 (BK, BM, K); ibid., 1800 – 2500 m, 3 Oct. 1971, Murata et al. T-15988 (holotype of G. javanicum var. brevidens, KYO; isotype BKF); ibid., 2550 m, 14 July 1990, Phengklai et al. 7505 (BKF 2 sheets); Doi Angka [Doi Inthanon], Mae Wang drainage [18.6125°, 98.7750°], 1900 m, 29 July 1927, Garrett 409 (BM 2 sheets, K, SING); Doi Inthanon, along the road to Mae Chaem, km 1 – 2, 18°31'N 98°29'E, 2000 m, 20 Sept. 1995, Larsen et al. 46729 (AAU, BKF); Doi Inthanon, en route to Mae Jaem [Mae Chaem] km 38, 1800 m, 6 Jan. 1998, Srisanga et al. 55 (QBG); Doi Inthanon, S side of Wachirathan Waterfalls [18.5418°, 98.5992°], 700 – 860 m, 1 Aug. 1988, Tsugaru T-61811 (BKF, GH, L); Doi Sootep, 720 m [2400 ft], 17 July 1909, Kerr 733 (holotype of G. intermedium, K; isotypes ABD, BM, K 2 sheets); Doi Suthep, 18°50'N 98°55'E, 1500 m, 17 Aug. 2004, Nielsen et al. 1714 (BKF); Doi Suthep, E side, Pah Laht [18.8014°, 98.9238°], 600 m, 25 July 1997, Maxwell 87-726 (BKF, L); Doi Suthep, E side, Mohntatahn Waterfalls [18.8167°, 98.9°], 600 m, 19 Nov. 1987, Maxwell 87-1465 (L); Huai Chang Kien [18.8458°, 98.8886°], 900 m [3000 ft], 25 July 1921, Kerr s.n. (ABD 2 sheets); Fang, 19°57'N 99°11'E, 600 m, 24 July 1968, Larsen et al. 2636 (AAU, L, P); Chiang Dao [19.3667°, 98.9667°], 1020 m, 25 June 1956, Bunchai 205 (ABD, BKF); Mae On, Doi Lahn, W side, above Mae Lio Village, Mae Lio Valley; Huay Gayo Subdistr., 1025 m, 14 July 2005, Maxwell 05-426 (BKF); Queen Sirikit Botanic Gardens, Huai Puu [18.8991°, 98.8603°], 900 m, 27 Aug. 1994, BGO staff 1464 (QBG); Me Sa [18.9032°, 98.9495°], July 1915, Kerr s.n. (BM); Ban Mae Saa Mai, 1000 m, 14 Aug. 1998, Pongamornkul 267 (QBG); Chiang Rai. Chiang Rai, Khun Korn Waterfalls [19.8821°, 99.6169°], 675 m, 11 Oct. 1997, KK 400 (BCU); Mae Suai Distr., Ban San Sa-ard, 16 July 1967, Sangkhachand 911 (BK); Wat Phra Thard Doi Tung [20.3241°, 99.8328°], 1315 m, 22 June 2002, Chamchumroon 1595 (BKF); Wiang Pa Pao Distr., Mae Chadee [19.1833°, 99.5167°], 1200 m, 31 Sept. 1993, Vial-Debas 56 (CMUB); Doi Luang National Park, [19.2°, 99.7167°], 1175 m, 8 Sept. 1997, Maxwell 97-982 (BKF, CMUB); Phayao. Doi Luang National Park, E side, 650 m, 22 Dec. 1997, Petrmitr 182 (CMUB); Doi Luang National Park, E side, Jahm Bah Tawng Falls, 575 m, 25 July 1997, Petrmitr 42 (CMUB, L); Nan. Bo Kleu, Doi Phu Kha National Park, Sapan Waterfalls, 19°12'N 101°12'E, 850 m, 6 July 2001, Srisanga 1953 (QBG); Doi Phu Kha National Park, 19°13'N 101°06'E, 1650 m, 1 July 1999, Srisanga et al. 759 (QBG); Nam Hang [18.3333°, 100.6167°], 600 m, 11 July 1926, Winit 1745 (BK, BKF, BM); Lamphun. Khun Tan [18.4708°, 99.2847°], 1000 m, 31 Oct. 1977, Chermsirivathana s.n. (BK); Lampang. Me Ngow, 440 – 460 m, 26 Aug. 1922, Winit 744 (BK, BKF, BM); Muang Pan, Chae Sorn National Park, along Mae Mawn Stream, near Jae Sawn Waterfalls, 550 m, 22 July 1996, Panatkool 132 (CMUB, L); Wang Nua [19.15°, 99.6167°], 650 m, 26 July 1996, Maxwell 96-1022 (CMUB); Tak. Doi Mussor, 900 m, 7 Dec. 1960, Smitinand 7038 (BKF); Yanhee Dam [17.25°, 98.9667°], Youngboonkird s.n. (ABD); Phitsanulok. Phu Miang [16.85°, 101.1167°], 2 Oct. 1968, SP et al. 22 (BKF); Thung Salaeng Luang, en route to Thung Nang Phaya, 850 m, 6 June 1982, Santisuk s.n. (BKF); Kamphaeng Phet, km 81 in Khlong Lan Natinal Park, 911 m, 12 April 1999, Chayamarit et al. 1537 (BKF 2 sheets); Dong Lan Waterfalls [16.1296°, 99.2814°], 300 m, 25 Nov. 1977, Phengklai et al. 3915 (BKF 2 sheets); Klong Lan, Mae Wong National Park [16.1317°, 99.2821°], 1290 m, 14 Aug. 1998, van de Bult 83 (BKF, CMUB); Phetchabun. Lom Kau, Pa-nam-nao, 600 m, 25 May 1953, Smitinand & Suvarnakoset 562 (BKF); Nam Nao National Park, 16°48'N 101°23'E, near check point of road to National Park, 280 – 350 m, 28 Oct. 1984, Murata et al. T-51688 (BKF); Sridit Waterfalls [16.6279°, 100.9397°], 380 m, 19 July 1986, Thaithong et al. 257 (BCU 3 sheets); Loei. Na Haew, 700 – 800 m, 31 July 1997, Na Nakorn et al. 9473 (QBG); Loei, Na Haew, Lao Gor Hok [17.4971°, 100.8781°], 820 m, 28 Aug. 2008, van de Bult 989 (CMUB); Loei, Wang Saphung, Sithan, 300 m, 21 Nov. 1958, Smitinand 4896 (BKF); Udon Thani. Nong Saeng, NE Arboretum [17.1675°, 102.7806°], Sidajium 31 (BKF); Buengkan. Bungkha, Phu Wua Wildlife Sanctuary [18.1539°, 103.9510°], 300 m, 7 May 1997, Pooma 1596 (BKF, CMUB); Sakon Nakhon. Phu Phan National Park, 17°N 104°E, 380 m, 13 Nov. 1984, Murata et al. T-49727 (BKF); Mukdahan. Dong Bug Ee, 150 m, 16 Dec. 1963, Smitinand 8449 (BKF); Chaiyaphum. Ban Nam Phrom, 15°40'N 102°E, 600 m, 23 May 1974, Geesink et al. 6870 (AAU, BKF, C, K, L); Nakhon Ratchasima. Khao Lotung, S of Pak Thong Chai, 800 m, 9 Aug. 1968, Larsen et al. 3181 (AAU, BKF, L); Khao Yai National Park, en route from Haew Suwat Waterfalls to Haew Pathun Waterfalls, 600 – 700 m, 26 Sept. 1982, Konta et al. T-29549 (GH, KUN); Khao Yai National Park, Kong Keo [14.4402°, 101.3740°], 850 m, 9 July 1966, Larsen et al. 160 (AAU, BKF); Pak Thong Chai, Sakae Rat [14.6°, 102.0333°], 2 Oct. 1968, Hambananda 378 (BKF); Salika Forest, 40 km SE from Pak Thong Chai, 14°40'N 102°2'E, 400 m, 25 Oct. 1971, van Beusekom & Geesink 3352 (BKF, C, L, P); Uthai Thani. Klong Pluu, 500 – 600 m, 22 Oct. 1993, Tiptabiankarn 1282 (SLR); Kanchanaburi. Kwae Noi Basin, near Ka Tha Lai, Ban Paung R. Valley, 300 m, 2 – 4 June 1946, Kostermans 809 (L); Sangkhlaburi, Kaoyai, 3 Aug. 1968, Sangkhachand 1460 (BK); Si Sawat, Erawan National Park [14.3833°, 99.1167°], 1 km N of Khao Kro Krae Mountain, 780 m, 1 Nov. 2003, van de Bult 697 (BKF, CMUB); Thong Pha Phum National Park, 14°40'N 98°30'E, 900 m, 22 Aug. 2004, Nielsen et al. 1862 (AAU, BKF, SING, TCD); Phetchaburi. Kaeng Krachan National Park, Khao Phanoen Thung [12.8665°, 99.3729°], 900 m, 22 Aug. 1998, Sasirat 49 (QBG); Prachuap Khiri Khan. Kao Luang [11.6°, 99.6667°], 800 – 900 m, 4 July 1926, Kerr 10829 (BK, BM, L 2 sheets); Bang Saphan [11.2°, 99.5167°], 0 – 150 m, 20 Aug. 1967, Shimizu & Nalampoon T-7801 (BKF); Sa Kaeo. Khao Takrup, 13°26'N 101°57'E, 200 m, 10 Sept. 1999, Middleton et al. 226 (BKF, K); Chon Buri. Khao Khieo opened zoo [13.2149°, 101.0566°], 300 m, 14 Aug. 2000, Phengklai et al. 12774 (BKF); Klong Yai, Huay Hindat [13.2167°, 101.1167°], 120 m, 12 Oct. 1922, Winit 56 (BKF); Sriracha [13.1667°, 100.9333°], 14 Nov. 1926, Put 442 (BK, BM 2 sheets, L); Sriracha, Nawng Kaw [13.15°, 100.8°], 30 m [100 ft], 18 Sept. 1911, Kerr 2054 (BM, K, P, TCD); Sriracha, Nong Nam Kheo [13.2333°, 101.1167°], 120 – 180 m [400 – 600 ft], 9 Nov. 1926, Collins 1483 (BM); Rayong. Klaeng, Khao Cha Mao National Park [12.9125°, 101.7221°], 50 – 150 m, 23 Nov. 1979, Shimizu et al. T-23429 (BKF); Chanthaburi. Khao Soi Dao, 12°56'N 102°13'E, 150 m, 9 Sept. 1999, Middleton & Pholsena 185 (BKF); Laem Sing, Pliew [Phliu] Waterfalls [12.5333°, 102.2°], 150 m, 13 Aug. 1973, Maxwell 73-422 (AAU, BK); Pong Nam Ron, Kao Krachai [12.6333°, 102.2°], 12 Sept. 1964, Chantanamuck 868 (BK); Trat. Dan Chumpon [12.45°, 102.6167°], 20 Dec. 1929, Kerr 18070 (BM); Krat, Kao Saming [12.35°, 102.45°], 25 Nov. 1924, Kerr 10016 (BK, BM 2 sheets); Nong Bon, 29 Nov. 1971, Vidal 5772 (AAU, P); Pong Song Salung, 5 Nov. 1964, Suvarnakoset 2200 (BKF); Chumphon. Langsuan, Tako [Takho, 10.0833°, 99.1°], 13 June 1928, Put 1632 (BK, BM 2 sheets, PHN); Phato, Nam Tok Ngao National Park, E side of Khao Nom Sao Mt, 9°54'N 98°44'E, 700 m, 17 Dec. 2005, Gardner & Khumchompoo ST2117 (E); Phato, Nam Tok Ngao National Park, trail to Haew Prow Waterfalls, 9°54'N 98°44'E, 400 m, 14 Dec. 2005, Sidisunthorn ST 2144 (K); Ranong. Kra Buri, Thungraya Nasak Wildlife Sanctuary, along Bok Krai R. from headquarters, 10°22'N 98°51'E, 200 m, 28 Aug. 2002, Middleton et al. 1437 (BKF); Surat Thani. Kanchanadit [9.1667°, 99.4667°], 5 m, 2 Aug. 1927, Kerr 13115 (BK, BM 2 sheets, L, PHN); Khao Sok, 8°55'N 98°40'E, 100 – 200 m, 25 Oct. 1990, Larsen et al. 40899 (AAU); Na Sarn, Pa Mai Daeng, 15 Aug. 1955, Snan 399 (BKF); Viphavadi, Khlong Yan Wildlife Sanctuary, trail from Headquarters, 9°9'N 98°53'E, 200 m, 31 Aug. 2002, Middleton et al. 1507 (BKF, CMUB, E, K, P); Phangnga. Khao Phra Mi, 9°17'N 98°26'E, 1100 m, 10 July 1972, Larsen et al. 30855 (BKF, K, L, P); Kuraburi, Song Pee Nong Fall, 50 m, 14 Aug. 1977, Santisuk 1171 (BKF 2 sheets); Phuket. 7 June 1930, Kiah 24251 (BK, K, SING); Krabi. Bang Kram Forest Reserve, 18 km E of Khlong Thom, 7°55'N 99°15'E, 75 m, 14 July 1992, Larsen et al. 43393 (AAU); Khao Phanom Bencha National Park, 8°14'N 98°55'E, 250 m, 17 July 2000, Middleton et al. 514 (BKF); Klong Tawm, Kao Pra Pang Kram Wildlife Sanctuary, 50 m, 13 Jan. 2006, Maxwell 06-011 (CMUB); Nakhon Si Thammarat. at middle elevation of Khao Luang, 650 – 950 m, 24 Aug. 1967, Iwatsuki et al. T-8461 (AAU, BKF, E); Kao Luang [8.5°, 99.6667°], 200 m, 27 April 1928, Kerr 15418 (BK, BM 3 sheets, L); Khao Na Ron, 23 July 1952, Suvarnakoset 379 (BKF); Kuan Mud, Khao Yod Pain, 400 m, 30 Aug. 1954, Suvarnakoset 830 (BKF); Lansaka, SE side of Khao Luang National Park, Kamjan, 8°28'N 99°41'E, 24 March 1993, Chantaranothai et al. 1367 (K); Ron Phibun hill [8.15°, 99.85°], 480 m [1600 ft], Feb. 1922, Smith 416 (BK, BM, K 2 sheets); Trang. 28 July 1904, Ridley 12602 (SING); Trang, Khao Chong, 15 km E of Trang, 7°30'N 99°45'E, 13 Nov. 1990, Larsen et al. 41307 (AAU); Satun. Tarutao National Park, Trail from Puntay Bay to Talo Wao, 13 Jan. 1981, Congdon 1054 (AAU); Kuan Karong, 40 km NW of Satun, 6°51'N 100°01'E, 15 Oct. 1970, Charoenphol et al. 3824 (AAU, BKF, L); Songkhla. Ban Prakawp [6.5667°, 100.6667°], 100 m, 18 July 1928, Kerr 15845 (BK, BM 2 sheets, L, PHN); Rattaphum [7.1333°, 100.2667°], 21 Dec. 1965, Youngboonkird 265 (BK); Rattaphum, Boriphat Waterfalls [6.9949°, 100.1514°], 120 – 150 m, 18 Dec. 1979, Shimizu et al. T-27663 (BKF); Ton Nga Chang [6.9477°, 100.2340°], 500 m, 21 Aug. 1992, Niyomdham 3116 (BKF); Narathiwat. Bachaw [Bacho, 6.4167°, 101.5833°], 400 m, 13 July 1923, Kerr 7173 (BK, BM, K); Waeng [5.9333°, 101.9°], 22 Nov. 1971, Anonymous 277 (BKF); Waeng, Hala Bala Wildlife Sanctuary, 5°47.787'N 101°50.217'E, 60 m, 14 Oct. 2005, Chongko & Boonkongchart 387 (CMUB). malaysia. Johor. Bukit Soga, Batu Pahat, Ridley 11130 (K, SING); Kedah. Kedah peak [Gunung Jerai, 5.7833°, 100.4333°], 450 m [1500 ft], 11 Dec. 1915, Robinson & Kloss 6144 (K); Gunung Baling [5.6667°, 100.9167°], 270 m [900 ft], 5 Aug. 1941, Nauen 38018 (SING); Kedah, Langkawi, Gunong Raya [6.3333°, 99.6667°], 500 m, 19 June 1932, Kerr 21706 (BK, BM, K, L, SING); Kelantan. Channing Woods [5.4667°, 102.2°], Feb. 1917, Ridley s.n. (K); Gua Musang-Kuala Betis track [4.9°, 101.7833°], 14 July 1935, Henderson 29658 (K, SING); S end Bukit Batu Papan, Sungai Lebir [4.9167°, 102.4167°], 8 July 1935, Henderson 29618 (K, SING); Ulu, Gua Musang [4.85°, 101.95°], 90 m [300 ft], 5 Aug. 1962, UNESCO Limestone Expedition 355 (K, L, PHN, SING); Negeri Sembilan. Pasoh Forest Reserve, 2°58'N 102°19'E, 19 Oct. 1981, Rogstad 585 (GH); Pahang. Bentong [3.5333°, 101.9167°], 15 July 1924, Best 14117 (SING); Cameron Highland, Tanah Rata, Robinson Falls, in shade [4.4667°, 101.3833°], 1425 m] [4750 ft], 29 Aug. 1970, Chung 169 (AAU); Kemanshul Division, Bentong, Sabai estate [3.3333°, 102.0833°], 27 Jan. 1958, Shah 166 (GH, K, SING); Fraser’s Hill [3.7167°, 101.75°], 1200 m [4000 ft], 15 Aug. 1937, Corner 33212 (GH, K 2 sheets, SING); Pine Tree Hill, near Fraser’s Hill, 1440 m [4800 ft], 27 Aug. 1923, Burkill 11075 (holotype of G. lacteum, K; isotype SING); Gunung Senyum [3.7°, 102.45°], 30 – 150 m [100 – 500 ft], 29 July 1929, Henderson 22234 (SING); Kota Gelanggi [3.9167°, 102.4833°], Aug. 1891, Ridley s.n. (SING); Kuala Tahan [4.3833°, 102.4°], Feb. 1921, Seimund 921 (SING); Pulau Tawar Wood [3.8833°, 102.4333°], 1891, Ridley 2148 (BM, K, SING); Tahan Valley [4.3847°, 102.4014°], 1891, Ridley s.n. (SING); Sungei Redap [4.4936°, 102.4903°], Ulu Sungai, Tembeling Pahang, 30 – 60 m [100 – 200 ft], 6 March 1968, Shah 1599 (L, SING); Sungei Yu [4.5833°, 101.9333°], 15 Aug. 1964, Hardial & Noor 109 (GH, K, L 2 sheets, SING); Telom [4.45°, 101.5833°], Nov. 1908, Ridley 13698 (BM, SING); Penang. Koh Pennan [Penang, 5.3667°, 100.25°], 25 May 1913, Robinson s.n. (K, P); Perak. Tapah [4.1833°, 101.2667°], Nov. 1908, Ridley 14061 (BM, SING); Upper Perak, Grik [5.4167°, 101.1333°], 26 July 1936, Corner 31627 (GH, L, SING 2 sheets); Gunong Tujok, Sept. 1884, Scortechini 1225 (holotype of G. scortechinii, CAL photo; isotype CAL 3 sheets, K, SING); Selangor, mile 15 Pahang Track [3.6486°, 101.7413°], July 1897, Ridley 8564 (BM, K, SING); Genting Highlands [3.4°, 101.8°], 26 Dec. 1966, Stone 6590 (L); Gua Batu Woods [3.25°, 101.65°], Dec. 1896, Ridley 8221 (SING 3 sheets); Semangkok Pass, 22 Jan. 1921, Ridley s.n. (holotype of G. oblongum var. setosum, K); Semenyih [2.9333°, 101.8333°], 14 July 1921, Hume 7965 (SING); Sabah. Keningau, Crocker range Forest Reserve [5.4666°, 116.0613°], 900 m, 9 Nov. 1976, Sundaling SAN 83957 (K, L); Keningau, Crocker range, mile 16, Kimanis Road, 21 Aug. 1986, Krispinus SAN 113840 (K, L); Kota Belud, Mt Kinabalu, Tenompok area toward Kadamaian [5.7974°, 116.4077°], 1500 m, 5 Jan. 1995, Beaman et al. 11173 (K); Lamag, Pua Din Kok logging area, Tongod [5.2667°, 116.9667°], 20 March 1980, Madani SAN 92027 (L); Mt Kinabalu, Dalas, hill trail, 900 m [3000 ft], 17 Aug. 1931, Clemens & S. Clemens 26116 (BM, K); Mt Kinabalu, ridge W of camp, 7 Feb. 1933, Clemens & S. Clemens 31502 (BM); Mt Kinabalu, Mesilau camp [6.0134°, 116.5966°], 1500 m [5000 ft], 19 Jan. 1964, Chew & Corner 4008 (GH, K, L, SING, US); Mt Kinabalu, Penataran basin, 1200 m [4000 ft], 25 July 1933, Clemens & S. Clemens 34145 (BM, G, GH, L); Nabawan, km 55, Jalan Nabawan/Sepulut, Ulu Sungai Nabawan [5.0453°, 116.4462°], 9 Feb. 1990, Krispinus SAN 128284 (K, L); Penibukan, Mt Kinabalu, 1200 m [4000 ft], 16 March 1933, Clemens & S. Clemens 35013 (BM); Ranau, Kinabalu National Park, Boundary Rentis, 1554 m, 22 Jan. 1976, Stevens et al. 616 (E, L); Ranau, Kinabalu National Park, Kiau View trail, 5°59'N 116°31'E, 19 July 2006, Bramley et al. 147246 (K); Ranau, Pinosuk Plateau, Mt Kinabalu golf course site, 1650 – 1750 m, 14 July 1984, Beaman et al. 10666 (K); Ranau, W Mesilau R., Valley above waterworks dam, 1600 – 1700 m, 19 Feb. 1984, Beaman et al. 8680 (K); Sandakan, VJR mile 42, Sungai Babakang [5.1167°, 118.9667°], 10 Dec. 1985, Tuyuk et al. SAN 113666 (K, L 2 sheets); Sipitang, Meligan, base of sandstone cliff [4.6833°, 115.7°], 19 March 1999, Kiew 4599 (SING); Sipitang, Sungai Ruan between Kampung Long Pasia and Ruan Watgerauo, 1300 m, 17 April 2000, Lim et al. SAN 143266 (K, SING); Sipitang, upper Sungai Miau near Mt Muruk Miau, 900 m [3000 ft], 13 April 2000, Pius & Dauni SAN 143153 (E, K); Tambunan, Crocker Range, km 64 on Kota Kinabalu, 5°46'N 116°1'E, 1250 m, 14 Nov. 1983, Beaman et al. 7495 (L); Tawau, Gunung Rara Forest Reserve, [4.4611°, 118.1247°], 2 April 1997, Pereira et al. 432 (K); Tawau, Tawau Hill Park [4.4356°, 117.8044°], 21 April 1992, Berhaman & Madani SAN 134509 (K); Tenom area [5.1333°, 115.95°], 1800 m [6000 ft], 1963 – 1964, Comber 140 (K); Tenompok, Mt Kinabalu, 1500 m [5000 ft], 2 March 1932, Clemens & S. Clemens 26116 (BO, K 2 sheets); Sandakan, Sebuga [5.8333°, 118.0333°], 4 Dec. 1920, Ramos 793 (PHN); Tenom, Hutan Simpan Mandalom, 10 Aug. 1993, Mantor SAN 136707 (E); Sarawak. Marudi, Pulong Tau National Park (Western Part), 3°40'38”N 115°17'11”E, 11 May 2007, de Kok et al. S97828 (K); Baram (R.), Tutoh, Sungai Melinau [4.0333°, 114.7833°], 5 Oct. 1971, Anderson S31794 (L); Bukit Berar, Gunung Mulu National Park, 4^th division, near Sungai Berar [3.05°, 115.8°], Ching S39552 (L); Bukit Rawan, Tebakang area, 1^st division, 635 m, 7 April 1983, Awa & Paie S 45593 (K, L); Gunong Balapau [3.1333°, 114.4667°], Ulu Tinjar, 600 – 700 m, 2 Feb. 1932, Richards 2403 (K); Lio Matu [3.1667°, 115.2333°], Baram, 3 Nov. 1914, Moulton 47 (PHN, SING); Lundu, Gunong Gading [1.7217°, 109.8145°], 840 m [2800 ft], 5 Aug. 1962, Burtt & Woods B2687 (E); Mulu National Park, trail to Camp I between Sungai Melinau and Sungai Melinau Paku, 4°N 114°50'E, 100 m, 31 March 1978, Stone 13571 (K). brunei. Temburong. Amo, Apoi Forest Reserve, Apan, beyond Sungai Tulan, 4°31'N 115°11'E, 70 m, 17 July 1993, Cowley et al. 98 (K); Amo, Apoi Forest Reserve, Temburong R. Catchment, Apan, Sungai Tulan, 4°32'N 115°11'E, 50 – 80 m, 15 July 1993, Sands et al. 5822 (K 2 sheets). philippines. Luzon. Albay. Mayon Volcano [13.25°, 123.6833°], 10 June 1953, Mendoza PNH 18370 (L); Aurora. Quezon, Baler, Dibutunan, 2 km from coast, 100 m, March 1948, Fox 5025 (GH); Benguet. Loher 4215 (K, US 2 sheets); ibid., Dec. 1908, Ramos 5714 (P); Mt Simacoco, Oct. 1921, Ramos & Edaño 40340 (US); Mt Tonglon [16.5°, 120.6667°], Aug. 1906, Curran 5014 (US); Mt Santo Tomas [16.3333°, 120.5667°], 1800 m [6000 ft], June 1904, Elmer 6534 (K, US); Bulacan. Angat [14.9333°, 121.0333°], Feb. 1919, Ramos & Edaño 34127 (BM); Cagayan. Gonzaga [18.2667°, 122°], Oct. 1929, Edaño 78203 (SING); Camarines Sur. Maagnas [13.7167°, 123.6667°], 27 – 28 Aug. 1908, Robinson 6364 (L); Panagan R. [13.5833°, 123.4667°], Dec. 1928, Edaño 76369 (G); Catanduanes. 14 Nov. – 11 Dec. 1917, Ramos 30280 (K, P, US); Cavite. km 75 of Cavite-Batangas Road, 13 Dec. 1959, Mendoza & Steiner PNH 41555 (L); Ifugao. Bontoc, Mt Polis [16.9667°, 121.0333°], Feb. 1920, Ramos & Edaño 37617 (K, P, US); Banaue, Bayninan [16.9167°, 121.0833°], 1200 m [4000 ft], 25 March 1963, Conklin & Buwaya 79601 (PHN); Ilocos Norte. Mt Pico de Loro [18.5167°, 120.8833°], 400 m, 8 March 1953, Edaño 18024 (K, L); Mt Quebrada [18.4833°, 120.8°], 750 m, 26 Feb. 1953, Edaño 17836 (G, GH, K, L); Bangui to Claveria, Aug. 1918, Ramos 33009 (SING); en route from Mt Bubonbilit to miner's camp in Mt Sicapoa range [18.0167°, 120.9333°], 1400 – 1900 m, 2 Dec. 1975, Iwatsuki et al. P-607 (L); en route from Neuva Era to Mt Semenublan [17.9°, 120.7167°], 450 – 800 m, 26 Nov. 1975, Iwatsuki et al. P-181 (AAU, K, L, PHN); Mt Bubungkayo, en route from Santo Nino to Miner's camp, 750 – 1300 m, 1 Dec. 1975, Iwatsuki et al. P-344 (K, L, PHN); Kalinga. Lubuagan, Mt Masingit [17.45°, 121.15°], Feb. 1920, Ramos & Edaño 37494 (K, US); Laguna. Jan. – Feb. 1916, Mabesa 25009 (K, US); Los Banos, Mt Maquiling [14.1358°, 121.19°], June – July 1917, Elmer 17836 (isotypes of G. cinereum, BM 2 sheets, K, L, NY, P, US); Mt Banajao [Banájao, 14.0675°, 121.4925°], May 1917, Ocampo 27975 (US); Mountain Province. Bontoc [17.0833°, 120.9667°], May 1913, Vanoverbergh 701 (C, G, GH, L, P); Mt Data [16.85°, 120.8667°], 2300 m, 9 April 1946, Alcasid 1894 (GH); Pampanga. Camp Stotsenburg [15.1833°, 120.5167°], Mt Pinatubo, May 1927, Elmer 22371 (BM, C, G, K, L, P, SING); Pangasinan. Mt San Isidro [15.9833°, 120.1167°], Nov. 1917, Fenix 29945 (BM, L, US); Quezon. Tayabas, Mt Banahao [14.0667°, 121.4833°], Oct. 1904, Whitford 975 (US); Tagkawayan [13.9667°, 122.5333°], 15 Sept. 1993, Barbon et al. 12269 (K); Rizal. Morong [14.5333°, 121.2167°], Vidal 3470 (GH, K); Bosoboso [14.6083°, 121.2589°], Sept. 1904, Ahern's collector 1866 (K, P, SING, US); Mt Susong-Dalaga [14.6333°, 121.3167°], Aug. 1917, Ramos & Edaño 29402 (K, P, US); San Andales, Dec. 1926, Edaño 48787 (BM); Sorsogon. Irosin, Mt Bulusan [12.7690°, 124.0565°], Dec. 1915, Elmer 15290 (BM, G, GH, K, L, P, PHN, US 2 sheets); Lake Agangay [Lake Aguingay, 12.7493°, 124.0731°], 19 June 1958, Sinclair & Edaño 9639 (E, SING); Mt Bulusan [12.77°, 124.05°], 650 m, 5 Aug. 1947, Sulit 2657 (GH, SING); Zambales. Mt Pinatubo [15.1367°, 120.3561°], 900 m, 9 July 1948, Fox 4814 (GH). Mindanao. Agusan Del Norte. Agusan, Cabadbaran [9.1167°, 125.65°], Aug. 1912, Elmer 13469 (BM, E, G 2 sheets, GH, K, L, P, PHN, US); Bukidnon. Aug. 1912, Fenix 15742 (SING, US); Mt Candoon, June – July 1920, Ramos & Edaño 38749 (K 2 sheets); Intavas, Impasug-ong, Mt Kitanglad [8.1161°, 124.9233°], 14 July 1991, Madulid et al. 3103 (K); Camiguin. March – April 1912, Ramos 14673 (BM, L); Davao Del Sur. Davao, Mt Apo [6.9875°, 125.2708°], May 1903, DeVore & Hoover 370 (US); Mindanao, Todaya [6.9597°, 125.3478°], Mt Apo, July 1909, Elmer 11146 (BM, E, G, GH, K, L, US 2 sheets); South Cotabato. Koronadal [6.4667°, 124.8833°], April 1932, Ramos & Edaño 84990 (GH); Mt Matutum [6.3606°, 125.0747°], 2100 m [7000 ft], 29 April 1932, Ramos & Edaño 85050 (GH). Visayas. Bohol. 1841, Cuming 1840 (holotype of G. philippinarum, K [Herb. Benthamianum]; isotypes C, E, G, K [Herb. Benthamianum]); Capiz. Panay, Jamindan [11.3667°, 122.4333°], April – May 1918, Ramos & Edaño 30961 (P); Iloilo. Panay, 1 – 11 Jan. 1913, Robinson 18189 (K, US); Leyte, 2 July 1913, Wenzel 220 (E, G 2 sheets, US); Dagami [11.0667°, 124.85°], Aug. 1918, Ramos 15187 (P); Ormoc, Lake Danao [10.8681°, 124.8558°], 666 m, 20 March 1950, Edaño 11894 (BM, GH, L, PHN, SING); Negros Occidental. Canlaon National Park, Mt Canlaon [10.3911°, 123.03°], 1500 m, 21 March 1992, Stone et al. 6266 (K); SW face of Mt Canlaon, 1200 m, 30 March 1992, Stone et al. 6450 (K); Negros Oriental. Cuernos de Negros [Horns of Negros, 9.2428°, 123.1794°], May 1948, Edaño 7145 (GH, L, SING); Dumaguete, Mt Cuernos, June 1908, Elmer 10391 (G, E); Sibulan, Lake Danao [9.3506°, 123.1833°], 976 m, 21 Aug. 1948, Edaño 6671 (GH); Northern Samar. Samar, Catubig R. [12.56°, 125.0167°], Feb. – March 1916, Ramos 24358 (US); Southern Leyte. Leyte, Cabalian [10.2833°, 125.1833°], Dec. 1922, Ramos 41489 (US); Western Samar. Samar, Mt Purog [11.8333°, 125°], 300 m, 19 Dec. 1951, Edaño 15389 (GH); Mt Capotoan [12.1528°, 124.9350°], 133 m, 10 Jan. 1951, Edaño 15533 (PHN); Mimaropa Region. Palawan. Busuanga Island [12.0833°, 120.0833°], Sept. 1922, Ramos 41251 (G, US); Aborlan, Malasgao R. [9.3092°, 118.4950°], 160 m, 10 March 1951, Edaño 14077 (BM, GH, L); Penigisan, 650 m, 20 Sept. 1961, Olsen 299 (C); Taytay, Bambanan, Mt Capoas [10.8°, 119.2833°], 150 m, 11 June 1993, Reynoso et al. 11232 (GH). Sulu Archipelago. Basilan. Aug. 1912, Reillo 15411 (BM, K, US). indonesia. sumatra. Aceh. Gunung Leuser Nature Reserve, Upper Mamas R. Valley Expedition, c. 15 km W of Kutacane, 3°25'N 97°40'E, Camp Aceh, 1250 m, 26 June 1979, de Wilde & de Wilde-Duyfjes 19014 (BO, K, L); Gayo & Alas Lands, Mt Losir [Mount Leuser], bivouac 9 on the Lau Alas R., 1000 m, 7 Feb. 1937, van Steenis 8685 (L); Gajolanden from Paloh to Kongke, 1200 m, 4 March 1937, van Steenis 9483 (L); Takengeun [4.6333°, 96.8333°], 1080 m [3600 ft], 7 Jan. 1932, Bangham & Bangham-Masters 736 (GH); North Sumatra. E of Mt Sibajak, 1300 m, 4 Jan. 1929, Lorzing 14909 (L); E of Mt Sibayak, upper Petani valley, 1300 m, 4 Jan. 1929, Lorzing 14918 (K, L 2 sheets, PHN, SING); W slope of Mt Sibayak near Berastagi, 3°15'N 98°29'E, 1000 m, 7 July 1988, Kessler 89 (L 2 sheets); Berastagi Woods, Karo Palateau, 1500 m [5000 ft], Feb. 1921, Ridley s.n. (K); Karo, 3°14'N 98°24'E [3.2333°, 98.4°], 1400 m, 28 Feb. 1999, Argent & Iqbar 99126 (E, GH); Coren Bandar, Caroe [3.1°, 98.5°], 950 m, 30 Dec. 1916, Lorzing 4616 (L); E Coast, vicinity of Berastagi, Karo Plateau [3.1833°, 98.5167°], 30 Jan. 1927, Bartlett 6527 (G, K, L); E Coast, Asahan, 1918, Bartlett & La Rue 224 (GH, L, US); E Coast, Asahan, Vicinity of Hoeta Bagasan, 7 Sept. 1934 – 4 Feb. 1935, Boeea 6469 (US); E Coast, Soebaktk, Karoland [3.1°, 98.5°], 21 June 1928, Boeea 760 (GH); Kaban Djahe [3.1°, 98.5°], 1420 m, 19 May 1939, Pooll s.n. (SING); Kabandjahe, 1200 m, 20 Jan. 1919, Lorzing 6178 (L); Permantang Siantar [2.95°, 99.05°], 6 April 1954, Alston 15219 (BM, BO); Sibalangit, Bandarbaroe, 900 m, July 1918, Lorzing 5803 (L); Sibolangit, Karo by Brastagi, 1350 m, 27 Aug. 1918, Lorzing 5949 (K, L); Berastagi, 1000 – 1300 m, 29 Aug. 1971, Iwatsuki et al. S-470 (K 2 sheets, L 2 sheets); E Coast, Hitean Haloban, 17 – 24 May 1933, Boeea 4317 (GH, US); E Coast, Vicinity of Hoeta Bagasan, Asahan, 6 Sept. 1934 – 4 Feb. 1935, Boeea 6414 (GH); E Coast, Vicinity of Tomoean Dolok, Asahan, 1000 m, 1 – 20 Aug. 1936, Boeea 9896 (GH); West Sumatra. Gunung Malintang [-0.3333°, 100.6667°], 1100 m, 18 July 1918, Bünnemeijer 3599 (L); Gunung Merapi [-0.3833°, 100.4667°], 1500 m, 18 Sept. 1918, Bünnemeijer 4692 (L); Gunung Singalang [-0.39°, 100.3308°], 2300 m, 27 May 1918, Bünnemeijer 2631 (K, L, SING); Kerintji, Sungei Kering, 2 March 1954, Alston 14057 (BM); Gunung Talamau, 600 m, 28 April 1917, Bünnemeijer 505 [L (L0796107)]; slope of Kerintji, Gunung Labu, 1 March 1954, Alston 13966 (BM); Koerintjipiek [Gunung Kerintji, -1.6967°, 101.2656°], 1100 m, 10 Feb. 1920, Bünnemeijer 8230 (K); S of Gunung Sago, en route from Kampun Puncak Pato to the summit, Batu Sangkar, Kabupaten Tanah Datar, 100°40'E 0°22'S, 1300 m, 10 March 1989, Nagamasu 3796 (BO); Jambi. Korinchi, Siolak Daras [Sioelakdaras, -1.9167°, 101.3°], 2970 m [9900 ft], 16 March 1914, Kloss s.n. (SING); Sandaran Agong, Korinchi, 735 m [2450 ft], 1 June 1914, Robinson & Kloss 193 (BM); Sungei Tanduk, Kayoe Aro Estate [-1.7976°, 101.2734°], 6 March 1954, Alston 14265 (BM, BO); South Sumatra. Bukit Palelawan Natural Reserve, 800 – 1450 m, 10 – 14 Feb. 1983, Afriastini 715 (K, L); Palembang, Gunung Dempo [-4.0158°, 103.1283°], 2000 m, 6 April 1933, de Voogd 1551 (L, SING); Lampung. Hoedjoeng [Hoodjoong Village, Blalamo, Blalauw or Belalau, -5.0833°, 104.2167°], 990 m [3300 ft], 1880, Forbes 1910 (BM, L). riau islands. Anamba Islands, Siantan, near Terempak, 60 – 90 m [200 – 300 ft], 2 April 1928, Henderson 20180 (K, SING). java. Banten. Udjung Kulon Nature Reserve [-6.7467°, 105.3336°], Mt Pajung, trail Tjibunar-Tjiuluran, 300 – 480 m, 31 Dec. 1963, Wirawan 178 (K, L); West Java. Preanger, Tangkoeban Prahoe [Tangkuban Perahu, -6.7667°, 107.6°], 1600 m, 5 March 1912, Backer 2430 (L); Preanger Region, Gunung Palasari, 1550 m, 13 Aug. 1923, Wisse 1011 (L); Preanger, Tjibodes [Tjibodas, -7.3833°, 107.5167°], 2400 m, 1 Nov. 1898, Koorders 32022β (L); Preanger, Tjidadap [-7.0167°, 107.2833°], Tjibeber, 1000 m, 26 April 1917, van den Brink 2189 (L); Preanger, Gunung Mandalagiri [-7.4167°, 107.8°], 1700 m, 25 March 1920, Lam 83 (L); Preanger, Tjadas-Malang, 1000 m, 12 April 1917, van den Brink 2968 (L); Preanger, Tjidadap, Guenoeng Besser [-6.6275°, 106.9414°], 1000 m, 19 May 1917, Winckel s.n. (L); Preanger Takoka [-7.0833°, 106.9667°], 1000 m, 27 Feb. 1894, Koorders 15012β (L); Preanger, Gunung Papandajan by Garoet, 1800 m, Koorders 40500β (L); Preanger, Mt Malabar [-7.1309°, 107.6610°], 1750 m, 24 July 1904, Hochreutiner 1396 (G); Preanger, Wates [-6.1658°, 107.1517°], Lembang, 1500 m, Feb. 1938, van de Pijl 673 (L 2 sheets); Buitenzorg [Bogor, -6.5897°, 106.7914°], 4 May 1895, Hallier s.n. (L); Gede, Blume s.n. (lectotype of G. javanicum, L; isolectotype L 4 sheets); Preanger, Gunung Gedeh, 19 Sept. 1911, Backer 3304 (L); Gunung Gedeh boven Tjibeureum, 1917, Hallier 392 (L); Gunung Gedeh, Kandang Badak, Lorzing 2239 (L); Gunung Gedeh by Tjibodas, 1917, Hallier 478 (L); Gunong Kendeng by Nirmala [-6.7333°, 106.5°], 1250 m, 1913, Backer 10976 (L); Gunong Salak [-6.7139°, 106.7350°] Boven Pasir Pogor, 1200 m, 1913, Backer 9375 (L); Gunong Tjibodas by Tjampea, 11 June 1896, Hallier s.n. (L 2 sheets); Gunong Pangrango [-6.7694°, 106.9644°], 2250 m [7500 ft], 2 April 1928, Best s.n. (SING); Mt Gedeh, Tjipanas sous Kandangbadak, 2100 m, 22 April 1904, Hochreutiner 1065 (G); Pasir Lectoeng by Tjidadap, 1000 m, 20 Dec. 1917, Winckel s.n. (L); Gunung Parang [-6.5950°, 107.3431°], 650 m, 29 July 1920, van den Brink 4936 (L); Tjampea, Buitenzorg, 200 – 250 m, 25 Sept. 1904, Hochreutiner 1900 (G); Tjidadap by Tjibeber, 900 m, May 1917, van den Brink s.n. (L, SING); Papandayan Volcano [-7.3333°, 107.7333°], 11 Feb. 1915, Ridley s.n. (K); Kerawang [-6.3°, 107.2950°], de Monchy 19 (L); Malabar [-7.2183°, 107.5886°], Anonymous s.n. (L); Tjibodas, Pangerango [-6.7667°, 106.95°], 1900 m, 13 Feb. 1969, Nooteboom 899 (L 3 sheets); Top van de Gunung Poeloesari [Pulusari], 1850 m, 21 April 1940, Coert 1547 (L); Dago [-6.8731°, 107.6197°], 16 April 1941, Holstvoogd 508 (L); Djampang tengah, Tjiletuk, on the road to Djampangkulon, W Bodjonglopang, 650 m, Nov. 1970, Kostermans 23940 (L); Lengkong, forest remnant SW of Skabumi, 800 m, 28 Jan. 1987, van Balgooy 4535 (L); W Banten, NW slope of Mt Pulosari, 750 m, 11 Feb. 1954, Adelbert 460 (K, L, US); Jawa Barat, Leuweung tengah, Ranca upas, Rasamala Forest, 1800 m, 13 Feb. 2000, Hoover & Wiriadinata 31026 (GH); Mt Halimun National Park, Cikaniki Research Station, 6°45'S 106°32'5”E, 900 – 1000 m, 20 May 2002, Wiriadinata et al. 10709 (GH); Central Java. Magelang [-7.4667°, 110.2167°], 19 June 1928, Rant s.n. (L); en route from Baturaden to Gunung Slamet, 1000 – 2000 m, 18 Aug. 1973, Murata et al. J-853 (L); Gunung Lawoe [-7.6333°, 111.1833°], 700 m, 1917, Backer 6731 (L); Gunung Sembung [-7.3833°, 110.0667°], 1300 m, 1913, Backer 12217 (L); Goendik [-7.2°, 110.9°], 150 m, 1917, Backer 6531 (L); Goenoeng Tenger [-7.2336°, 110.3069°], 29 March 1893, Hallier D280 (G); Moentilan [-7.75°, 109.95°], 350 m, 4 March 1914, Backer 12052 (L); Mt Sindoro [-7.2833°, 109.9833°], 1600 m, 3 Nov. 1904, Hochreutiner 2346 (G 2 sheets, L); Pemalang, 50 m, 14 Jan. 1918, Backer 23351 (L); Res Semarang, Gunung Telomojo [-7.3611°, 110.3994°], 12 April 1899, Koorders 36049β (L); Gunung Prahoe [-7.2°, 109.9167°], 1900 m, 1917, Backer 21878 (L); Sarie, 11 May 1843, Zollinger 144 (P); Yogyakarta. Djokjakarta [-7.8°, 110.3667°], 150 – 200 m, 1912, Backer 2517 (L); East Java. 1868, Teysmann s.n. (P); Besoeki [-8.2167°, 111.7833°], 50 m, 1 April 1929, Backer 37120 (L); Besoeki, Poeger Watangan [-8.0167°, 112.9167°], 26 Feb. 1898, Koorders 30015β (L); Besoeki, Kp Agoeng, Gungung Ringit [-7.7167°, 113.8333°], 700 – 1200 m, 8 March 1940, Buwalda 7538 (GH, K, L, SING); Besoeki, Sitoebondo [-7.7°, 114.0044°], 13 Nov. 1895, Koorders 20938β (K, L, P); Jdjen Plateau, Res Besoeki [-8.2167°, 111.7833°], 1700 m, 12 Nov. 1893, Koorders 14853β (L); Jang Plateau [-7.9833°, 113.6667°], 1971, Hoogerwerf 360 (L); Jdjen Plateau [-7.9833°, 113.6667°], Nov. 1938, Kostermans s.n. (L); Kediri, Gadoengan Pare, 350 – 500 m, 2 July 1896, Koorders 22937β (K, L, P); Kediri, near Ngandjuk, 25 May 1924, Rant s.n. (L); Purwodadi Garden [-8.25°, 112.4°], 31 March 1965, Kostermans s.n. (L); Bondowoso [-7.9167°, 113.8167°], 360 m [1200 ft], 30 March 1845, Zollinger 2010 (G, GH, P 2 sheets); Madioen, Ngebel, 850 m, 26 May 1896, Koorders 23258β (K, L, P); Madioen, Gunung Wilis [-7.8667°, 111.8°], 1300 – 1400 m, 27 May 1895, Koorders 23274β (L); Goenoeng Malang [-8.1167°, 112.8667°], 14 March 1909, Backer s.n. (L); Kawa Maruck Wood Forest, 8 Feb. 1915, Ridley s.n. (BM 2 sheets, K); Lumbing [Lumbang, -7.8333°, 113.0667°], 1500 m [5000 ft], Aug. 1875, Kuntze 5615 (K); Res Pasoeroean, Gunung Ardjoeno [-7.75°, 112.5667°], Sept. 1917, Koorders 38195β (L); Res Semarang [-7.0114°, 110.4°], 9 April 1897, Koorders 27205β (K, L); Lawoe, Sido Rampug, Gandangal, 1300 – 1400 m, Nov. 1907, Elbert 143 (L); Madura [-7.3333°, 108.65°], 1814, Reinwardt s.n. (L); Moenoeng Forestry [-7.5167°, 112.1°], 5 May 1927, Coert 476 (L); unlocalised. Java, Reinwardt s.n. (lectotype of G. phlomoides, L; isolectotypes L 2 sheets). kalimantan. West Kalimantan. Siantan [-0.0333°, 109.35°], 200 m, 2 April 1928, van Steenis 950 (L, SING); Central Kalimantan. Kalimantan Timur, Gunung Loan Api, NW of Long Bawan, Krayan, 3°55'N 115°40' E, 900 – 1250 m, 11 July 1981, Kato et al. B-8173 (BO); East Kalimantan. Malinau, Pujungan, Lg Belaka, Sungai Lurah, Kayan Mentarang National Park [3.2°, 115.5°], 200 – 500 m, 14 May 2002, Koizumi & Lawe 689 (BO); W Koetai, 200 m, 26 Aug. 1925, Endert 2891 (L); Loa Haur, W of Samarinda [-0.5°, 117.15°], 60 m, 12 May 1952, Kostermans 6824 (L); South Kalimantan. Kalimantan Selantan, Gunung Besar, [-2.7167°, 115.6167°], 800 – 1300 m, 19 Feb. 1979, Murata et al. B-3902 (L); Kalimantan Selatan, Gunung Halauhalau [Gunung Besar], Pegunungan Meratus, Barabai, 1400 m, 23 Oct. 1972, Dransfield 2868 (L). sulawesi. Central Sulawesi. Sopu Valley, 80 km SE of Palu, 0°30' – 1°30'S 119°30' – 120°30'E, 1000 m, 27 April 1979, van Balgooy 3032 (GH, K, L); South Sulawesi. Bonthain [-5.5333°, 119.9333°], Teysmann 13722 (L); S of Gunung Lompobatang, above Desa Bonto Lojang, Kab. Bantaeng. 5°24'S 119°56'E, 900 m, 9 Feb. 2000, Argent et al. 18 (GH, K); North Maluku. Ternate [0.7833°, 127.3667°] or Tidore Island, Curtis s.n. (K).

habitat. Shade and damp areas in primary or secondary evergreen forest, 0 – 1800 m.

conservation status. Widespread; Least Concern (LC).

phenology. Flowering and fruiting all year round.

notes. Gomphostemma javanicum is the most common species in the genus with the widest distribution and significant morphological variation (see Fig. 5). Polymorphic characters of this species include the varied size of the leaf, calyx and corolla. However, intermediate forms among those variable characters are seen when specimens are examined from across the range of distribution.

Gomphostemma javanicum is close to G. leptodon by having a prominently 10-ridged calyx but the margin of the fruiting calyx lobes of G. javanicum is not or hardly revolute in cross section unlike G. leptodon.

17. Gomphostemma leptodon Dunn (1913: 170); Doan (1936: 1036); Li & Hedge (1994: 72); Phuong (2000: 106). Type: Vietnam, Dong-dang [21.0667°, 106.95°], among bushes, Feb. 1886, Balansa 1000 (K, lectotype selected here; isolectotypes G, P [P00686236]).

Shrub-like herb up to 3 m tall. Stems erect, upright, robust, obtusely quadrangular with longitudinal grooves, with dense stellate hairs and sometimes dendroid hairs. Leaves petiolate, coriaceous, blades broadly ovate, 18 – 32 × 10 – 23 cm, apex acute, margin serrate, base cuneate, upper side tomentose with stellate hairs, lower side pale green tomentose with stellate hairs and robust venation; petioles quadrangular, 50 – 150 mm long, with dense stellate hairs. Inflorescence axillary with opposite cymes subsessile, branched then lax or rarely congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils many-flowered, subtended by upper leaves only; peduncles 5 – 12 (– 15) mm long; bracts deciduous, ovate or lanceolate, 5 – 15 × 4 – 10 mm, apex acute, margin serrate, base cuneate, with stellate hairs on both sides; bracteoles narrowly lanceolate or linear, 5 – 10 mm long. Flowering calyx infundibular, 8 – 13 (– 15) mm long, prominently 10-ridged; tube 4 – 6 mm long, outside with dense stellate hairs and glandular-like simple hairs, and minutely glandular punctuate, inside with short simple hairs; lobes spreading, lanceolate-ovate, 4 – 7 mm long, outside densely tomentose with short stellate hairs, glandular-like simple hairs and minutely glandular punctuate, inside adpressed simple hairs. Fruiting calyx infundibular-cupular, 10 – 17 (– 20) mm long; tubes 4 – 7 mm long; lobes triangular, 6 – 10 (– 13) mm long, apex acuminate, margin revolute in cross section. Corolla white or yellow, 25 – 50 (– 60) mm long, throat inflated, abruptly dilated near the throat, outside sparsely simple hairs, inside glabrous; tube slender, incurved, 20 – 40 (– 50) mm long; posterior lip flat, 5 × 13 mm, apex and margin emarginate; anterior lip 10 mm long, median lobe obovate, apex and margin margin slightly undulate. Staminal filaments fleshy. Style slender, glabrous, shorter than corolla. Nutlets 3 – 4, globular or obovate, 4 – 5 mm long. Fig. 1E – F.

distribution. China (Guangxi), Vietnam. Map 5.

Map 5

Distribution range of Gomphostemma leptodon, G. mastersii, G. melissifolium and G. ovatum.

specimens examined. china. Guangxi. Lungchow [22.3478°, 106.8578°], Mt A-chin, Feb., Morse 257 (lectoparatypes K 2 sheets). vietnam. Ha Nam: Kiên Khê [20.5031°, 105.8808°], Dông Bâu, 11 Nov. 1883, Bon 2272 (P 2 sheets); Kiên Khê, Dong Ham, 10 Jan. 1891, Bon s.n. (P); Ha Tay: Tu Phap, Feb. 1887, Balansa 2931 (P 2 sheets); Bavi Distr., Ba Vi National Park, 21°3.834'N 105°21.719'E, 600 m, 6 March 2007, Nguyen et al. HNK 1256 (de Kok 1203) (K); Pagode Dein Touan, sur la Mont Bavi, 25 Jan. 1889, Balansa 2930 (P); Hoa Binh: Canton de Ki Son [Ky Son, 20.8824°, 105.357°], 50 m, 9 Dec. 1926, Petelot 5002 (BM, P); Mai Chau, Van Mai [20.5867°, 105.04°], 21 Dec. 1999, Phuong 2460 (HN 5 sheets); Tan Lac Distr., Quyet Chien Municipality, Hung Village, 20°32'58”N 105°11'56”E, 700 – 940 m, 30 Nov. 2003, Wu et al. 355 (HN); Yen Thuy, Yen Tri, 20°22.40'N 105°37.90'E, 140 m, 4 Jan. 2001, Loc et al. 10510 (P); Cho-bo (Riviere Noire) [20.7667°, 105.1667°], 16 Nov. 1887, Balansa 2932 (P); Lao Cai: Sa Pa Distr., Khoang Village, 22°22'28”N 103°47'41.5”E, 1700 m, 24 April 1986, Phuong 142 (HN, K); Nam Dinh: Co Nghia, 28 Nov. 1913, Chevalier 29211 (P); Nghe An: Con Cuong, Yen Khe Commune, Trung Huong Hamlet, Cay Da Valley, 18°58'10.4”N 104°48'8.8”E, 15 Oct. 2008, Du et al. HNK 2947 (K); Quy Hop Distr. [19.333°, 105.167°], Chau Thai Commune, Ban Nha Village, 17 March 2007, Nguyen et al. HNK 1672 (K 2 sheets); Pu Mat National Park, Khe Kem Waterfall, 18°58'10.4”N 104°48'8.8”E, 13 Oct. 2008, Du et al. HNK 2821 (K); Ninh Binh. Cuc Phuong National Park, 20°21'N 105°35.77'E, 330 m, 22 Nov. 1998, Soejarto et al. 10360 (L, P); Phu Yen. Cho Ganh [13.2833°, 109.25°], Feb. 1923, Petelot 868 (P 2 sheets); Quang Binh. Dong Hoi Distr., Phong Nha-Ke Bang National Park [17.5372°, 106.1513°], Son Trach commune, 250 m, 22 March 2007, Nguyen et al. HNK 1813 (K 2 sheets); Quang Ninh. Dong-dang, Feb. 1886, Balansa 1000 (lectotype K; isolectotypes G, P); Thanh Hoa. Ba Thuoc, Pu Loung Nature Reserve [20.4667°, 105.1727°], 20 Nov. 2002, Phuong 5795 (HN); Thanh Hoa, Ben En National Park [19.6167°, 105.525°], 13 Jan. 2006, Sam HVS 065 (L).

habitat. In primary lowland, evergreen forest, limestone mountains, 50 – 1700 m.

conservation status. Widespread; Least Concern (LC).

phenology. Flowering and fruiting October – April.

notes. Gomphostemma leptodon is recognised by a large corolla, almost 25 – 50 (– 60) mm long and robust venation, especially on the lower side. It is morphologically similar to G. grandiflorum by having fruiting calyx lobes revolute in cross section but G. leptodon has the fruiting calyx lobes longer than the tube at maturity.

18. Gomphostemma mastersii Benth. ex Hook.f. (Hooker 1885: 699); Prain (1891: 268); Mukerjee (1940: 213); Murata (1976: 190); Walsingham (2019: 201). Type: India, Khasia, Nunklow [Nongkhlaw, 25.6833°, 91.6333°], 600 – 1200 m [2000 – 4000 ft], 21 July 1850, Hooker & Thomson 1814 (K [Herb. Hookerianum, K000928172], lectotype selected here; isolectotype K [Herb. Hookerianum, K000928173]).

Gomphostemma furfuraceum Hallier f. (1898: 351). Type: Indonesia, Sumatra orient, Deli [3.75°, 98.6833°], Kampong Pakam, Jaheri 1895 (holotype G).

Gomphostemma parvum Merr. (Merrill 1934b: 192). Type: Indonesia, Sumatra, East Coast, along Aek Panialalangan, Pargambrian, Asahan [2.3725°, 99.2193°], 270 – 360 m, 22 May 1927, Bartlett 8176 (holotype NY [00000534] photo; isotype US [00119425]).

Perennial herb up to 0.4 m tall with tuber-like roots, sometimes rooting at nodes. Stems upright, sometimes flaccid or more than one stem arisen from the same rootstock, obtusely quadrangular with longitudinal grooves, with dense sessile stellate hairs. Leaves petiolate, chartaceous; blades ovate, lanceolate-ovate, cordate or elliptic-oblong, 5 – 18 × 2 – 8 (– 11) cm, apex acute or broadly acute, margin shallowly serrate or serrate, base symmetrical or oblique, obtuse, attenuate or cordate, upper side with sessile stellate hairs and 2 – 4-celled simple hairs all over, lower side pale green tomentose with sparse sessile stellate hairs; petioles 15 – 60 mm long, strong or flaccid. Inflorescence axillary with opposite cymes sessile, unbranched then congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils few-flowered, subtended by upper leaves; bracts deciduous, ovate, obovate or oblanceolate, 9 – 20 × 3 – 8 mm, with sessile stellate hairs on both sides; bracteoles linear, falcate or lanceolate-oblong, 5 – 15 × 1 – 2 mm. Flowering calyx infundibular, thin, 10 – 15 mm long, 10-ridged, outside with dense sessile stellate hairs, inside with adpressed simple hairs; tube 4 – 6 mm long; lobes triangular, 6 – 9 mm long. Fruiting calyx cupular, 10 – 17 mm long; tube 4 – 6 mm long; lobes 6 – 10 mm long, apex subulate. Corolla creamy-white or bright yellow, 40 – 50 mm long, throat inflated, abruptly dilated near the throat, outside with sparse simple hairs, inside glabrous; tube incurved, 33 – 40 mm long; posterior lip broadly ovate, 7 – 10 × 8 – 12 mm, apex emaginate; median lobe tongue-like, apex retuse; lateral lobes ovate, apex and margin entire or undulate. Staminal filaments slightly tomentose. Style slender, glabrous with apex unequally lobed. Nutlets 2 – 4, ovate-elliptic or globose. Fig. 1A – B.

distribution. India (Assam, Meghalaya, Mizoram), Thailand, Indonesia (Sumatra). Map 5.

specimens examined. india. Assam. Masters 340 (lectoparatype of G. mastersii, K [Herb. Hookerianum]); Upper Assam, Jenkins 340 (K [Herb. Benthamianum]); Cachar [25.0833°, 92.9167°], Keenan s.n. (K); Cachar, Doarbund Pass, 22 July 1873, Keenan s.n. (K); N Cachar Hills, Haflong [25.1833°, 93.0333°], 750 m [2500 ft], 1 Aug. 1908, Craib 102 (K); Meghalaya. Khasia, Nunklow, 600 – 1200 m [2000 – 4000 ft], 21 July 1850, Hooker & Thomson 1814 (lectotype of G. mastersii, K [Herb. Hookerianum]; isolectotype K [Herb. Hookerianum]); Shillong [25.5667°, 91.8833°], Eastern Circle, Nongstoin Village between I.B. and Siem’s House, 21 June 1958, Panigrahi 16570 (CAL, L); Khasia, below Nongkhlaw, 600 – 1200 m [2000 – 4000 ft], 11 July 1850, Hooker & Thomson s.n. (lectoparatype of G. mastersii, K [Herb. Hookerianum]); Khasia, Borpani [Bor Panee] R. [26.2167°, 92.5333°], 600 – 1200 m [2000 – 4000 ft], 16 July 1850, Hooker & Thomson s.n. (lectoparatype of G. mastersii, K [Herb. Hookerianum]); Khasia, Churra, 600 – 1200 m [2000 – 4000 ft], 17 June 1850, Hooker & Thomson 976 (lectoparatype of G. mastersii, K [Herb. Hookerianum]); Khasia, Churra, 600 – 1200 m [2000 – 4000 ft], 15 Aug. 1850, Hooker & Thomson s.n. (lectoparatype of G. mastersii, K [Herb. Hookerianum]); Khasia, Jasper Hill, 600 – 1200 m [2000 – 4000 ft], 20 June 1850, Hooker & Thomson s.n. (lectoparatype of G. mastersii, K [Herb. Hookerianum]); Mont Khasia [25.5833°, 91.6333°], 1200 m [4000 ft], Hooker & Thomson s.n. (P); Mizoram. Aijal, Lushai Hills [23.1667°, 92.8333°], 900 m [3000 ft], Aug. 1926, Parry 58 (K); Tuipuibari, near old forest, 190 m, 12 Dec. 1995, Lalramnghinglova 8049 (CAL). bangladesh. Sylhet. de Silva s.n. in Wall. Cat. 2157 (K-W in parte ‘B’). thailand. Surat Thani. Ban Kawp Kep [8.8333°, 99.5°], 20 m, 7 Aug. 1927, Kerr 13189 (ABD, BK, BM 2 sheets, PHN); Phangnga. 5 Dec. 1918, Haniff & Nur 3867 (K, SING); Khao Khlong Yang at Khao Phra Mi, 9°19'N 98°26'E, 100 – 300 m, 6 July 1972, Larsen et al. 30688 (AAU, BKF, K); Nakhon Si Thammarat. Khao Luang [8.3704°, 99.7364°], 400 m, 28 Jan. 1966, Hansen & Smitinand 12142 (BKF, C, E, L 2 sheets, SING); Khao Luang National Park, Karom Waterfall, 300 m, 11 July 2000, Chamchumroon 835 (BKF); Phatthalung. Sii Ban Phot, Khao Pu Khao Ya National Park, 7°40'N 99°52'E, 140 m, 14 July 2000, Middleton et al. 446 (BKF); Khao Phu Khao Ya, 7°40'N 99°5'E, 100 – 150 m, 16 Nov. 1990, Larsen et al. 41512 (AAU); Trang. Chawng [Chong, 7.5°, 99.9167°], 100 m, 14 April 1928, Kerr 15760 (BM); Khao Chong [7.56°, 99.78°], 200 m, 12 Aug. 1975, Maxwell 75-785 (BK); Satun. Kwan Dohn [Khuan Don], Talay Ban [Thale Ban] National Park [6.4728°, 100.1339°], 100 m, 2 July 1985, Maxwell 85-662 (L); Yala. Banang Sta [6.2667°, 101.25°], 100 m, 26 July 1923, Kerr 7350 (BK, BM, K 2 sheets). indonesia. sumatra. Aceh. Gunung Leuser Nature Reserve, camp Simpang and vicinity, 3 – 5 km upstream Lau Ketambe, c. 35 km NW of Kutacane [3.6570°, 97.6132°], 400 – 600 m, 15 July 1972, de Wilde & de Wilde-Duyfjes 13615 (L); Gunung Leuser Nature Reserves, Ketambe Research Station and vicinity, Alas R. valley, c. 35 km NW of Kutacane, 3°40'N 97°40'E, 300 – 350 m, 11 July 1979, de Wilde & de Wilde-Duyfjes 18618 (BO, K, L); Aceh Tenggara, Ketambe, primary forest, 340 m, Feb. 1980, Rochadi 645 (BO); North Sumatra. Deli [3.75°, 98.6833°], Kampong Pakam, 6 March 1896, Jaheri 1895 (holotype of G. furfuraceum, G); E Coast, Adian Rindang, Asahan, vicinity of Hoeta Tomoean Dolok [Toemoean Dolok, 2.6667°, 99.1667°], 17 Nov. – 10 Dec. 1935, Boeea 8795 (GH); E Coast, along Aek Panialalangan, Pargambiran, Asahan, 270 – 360 m, 22 May 1927, Bartlett 8176 (holotype of G. parvum, NY photo; isotype US photo); E Coast, along the Toba trail N of the Asahan R., between Adian Langge and Si Martoloe, 21 – 25 April 1927, Bartlett 7370 (US).

habitat. In primary evergreen forest on limestone or thick leafy humus covered ground, or near rivers, 300 – 1200 m.

conservation status. Widespread; Least Concern (LC).

phenology. Flowering May – January. Fruiting November – April.

notes. Gomphostemma mastersii is recognisable by having stellate hairs on both sides of the leaves and its calyx lobes normally longer than calyx tube. It is one of only two species of this genus with tuber-like roots, the other being G. ovatum (more details in notes under G. ovatum).

Hooker cited a few collections in Herb. Hookerianum. A collection with two sheets, Hooker & Thomson 1814, is selected as the lectotype.

19. Gomphostemma melissifolium Wall. ex Benth. (Bentham in Wallich 1830b: 12); Hooker (1885: 698); Prain (1891: 270); Mukerjee (1940: 213). Type: India, Sillet, de Silva s.n. in Wall. Cat. 2157 (K [Herb. Benthamianum, K000928179], lectotype selected here; isolectotypes BM, G [G00189095], K [Herb. Benthamianum, K000928177], K [Herb. Hookerianum, K000928178], K-W [K001115341, in parte]).

Sprawling herb. Stems slender, scrambling, internodes of leafy and flowering shoots with curving stem appearing wavy, always twisting; obtusely quadrangular with longitudinal grooves, with stellate hairs. Leaves petiolate, chartaceous; blades elliptic or ovate, 5 – 8 × 2.5 – 4 cm, apex acute, margin serrate or shallowly serrate, base attenuate or obtuse, upper side tomentose with sessile stellate hairs confined on midrib or sparsely so to almost glabrous; lower side pale green tomentose with stellate hairs confined on midrib and veins; petioles 10 – 30 mm long, with dendroid hairs. Inflorescence axillary with opposite cymes sessile, unbranched then congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils 1 – 2-flowered, subtended by upper leaves; cluster of small leaf-like bracts deciduous, elliptic, lanceolate or obovate, 7 – 15 × 4 – 8 mm, apex acute, margin entire or shallowly serrate, base attenuate or obtuse, upper side with simple hairs, lower side with sessile stellate hairs; bracteoles linear or narrowly elliptic, 4 – 7 mm long. Flowering calyx infundibular, 10 – 12 mm long, 10-ridged, outside with dense sessile stellate hairs, inside with simple hairs; tube 5 – 7 mm long; lobes triangular, c. 5 mm long. Fruiting calyx not seen. Corolla whitish-purple, 30 – 50 mm long, throat inflated, abruptly dilated near the throat, outside with simple hairs at the upper part, inside glabrous; tube slender, incurved, 20 – 40 mm long; posterior lip ovate, apex emaginate; anterior lobes spreading beyond posterior lip; median lobe broadly ovate, tongue-like. Staminal filaments fleshy, tomentose. Style slender, glabrous with apex unequally lobed. Nutlets scarlet, solitary, 3 – 5 mm long.

distribution. India (Sikkim, Assam), Bangladesh. Map 5.

specimens examined. india. Sikkim. 600 m [2000 ft], Hooker s.n. (K 2 sheets [Herb. Hookerianum], P); Assam. Cachar [25.0833°, 92.9167°], Keenan s.n. (K); ibid., 20 Nov. 1889, Prazer s.n. (K). bangladesh. Sylhet. Sillet, de Silva s.n. in Wall. Cat. 2157 (lectotype K [Herb. Benthamianum]; isolectotypes BM, G, K [Herb. Benthamianum], K Herb. Hookerianum], K-W [in parte]); Sylhet Station, 24 Nov. 1872, Clarke 17935A (K); ibid., 24 Nov. 1872, Clarke 17935B (BM).

habitat. In hill evergreen forest, 600 m.

conservation status. Gomphostemma melissifolium is known from five localities with AOO of 20 km² and EOO of 37,692.17 km². No new specimens have been collected since 1872, and the original collecting localities are hard to pinpoint. Further information gathering is required to determine if there is evidence of decline of habitat or populations. The Data Deficient (DD) category is applied for this species.

phenology. Flowering and fruiting November.

notes. Gomphostemma melissifolium is a distinctive species recognised by having a sprawling habit and long corolla, 30 – 50 mm. It is morphologically similar to G. flexuosum by having coarse indumentum on the upper side of leaves and curving stems.

20. Gomphostemma ovatum Wall. ex Benth. (Bentham in Wallich 1830b: 12); Hooker (1885: 699); Prain (1891: 269); Mukerjee (1940: 213). Type: Nepal, Chandaghery [Chandagiry], Aug. 1821, Wall. Cat. 2155 (K [Herb. Benthamianum, K000928175], lectotype selected here; isolectotypes E [E00301973], G [G00189096], K [Herb. Hookerianum, K000928176], K-W [K001115338], NY [00000529], OXF).

Gomphostemma ovatum Wall. ex Benth. var. flaccidum Prain (1891: 270). Type: India, Sikkim, Rangiet, 2500 m, May 1862, Anderson 1093 (CAL [CAL0000025030], lectotype selected here).

Perennial herb up to 0.3 m tall, with tuber-like roots. Stems upright, sometimes flaccid, obtusely quadrangular with longitudinal grooves, with dense sessile stellate hairs. Leaves petiolate, chartaceous; blades ovate, lanceolate-ovate or elliptic, 5 – 20 × 2 – 10 cm, apex acute or broadly acute, margin shallowly serrate or serrate, base cuneate, attenuate or obtuse, upper side with 2 – 4-celled simple hairs, sometimes sparse stellate hairs confined only on lower part of midrib, lower side pale green tomentose with sparse sessile stellate hairs confined on leaf edge, midrib and veins; petioles 15 – 80 mm long, strong or flaccid. Inflorescence axillary with opposite cymes sessile, unbranched then congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils few-flowered, subtended by the upper leaves and cluster of small leaf-like bracts; bracts persistent, ovate, elliptic-ovate, spathulate or oblanceolate, 10 – 25 × 5 – 10 mm, apex acute, margin shallowly serrate, base cuneate; bracteoles linear, falcate or narrowly lanceolate-oblong, 6 – 10 × 1 – 2 mm. Flowering calyx infundibular, 8 – 12 mm long, 10-ridged, outside with dense sessile stellate hairs and shortly simple hairs, inside with densely adpressed simple hairs; tube slender, 4 – 6 mm long; lobes lanceolate-triangular, 4 – 6 mm long, apex acute. Fruiting calyx not seen. Corolla yellow, 25 – 40 mm long, throat inflated, abruptly dilated near the throat, outside with glandular-like simple hairs, inside glabrous; tube incurved, 30 – 40 mm long; posterior lip broadly ovate, 5 – 7 × 5 – 8 mm, apex and margin undulate; anterior lip 3-lobed, longer than posterior lip; median lobe tongue-like, apex and margin slightly undulate; lateral lobes ovate, apex and margin entire or undulate. Staminal filaments fleshy. Style slender, glabrous with apex subequally lobed. Nutlets not seen.

distribution. India (West Bengal, Sikkim, Meghalaya), Nepal. Map 5.

specimens examined. india. West Bengal. Darjeeling Distr. [27.0333°, 88.1667°], 600 m [2000 ft], 26 Aug. 1870, Clarke 12498A (K); Jalpaiguri Distr. [26.5167°, 88.7333°], Banya-6, 7 Dec. 1995, Chandra & Mandal 1066 (CAL); Sikkim. 900 – 1200 m [3000 – 4000 ft], July, Hooker s.n. (K [Herb. Hookerianum]); Rangiet, 2500 m, May 1862, Anderson 1093 (lectotype of G. ovatum var. flaccidum, CAL photo); Terai, 900 m [3000 ft], Aug. 1827, Hooker 71 (K [Herb. Hookerianum]); Meghalaya. Jaintea, Sept. 1852, Simons s.n. (lectoparatype of G. ovatum var. flaccidum, CAL photo); ibid., June 1850, Simons s.n. (lectoparatype of G. ovatum var. flaccidum, CAL photo); Cherapunjee, Khasia Hills, 1200 m [4000 ft], 5 July 1952, Chand 5984 (E); Khasia, 4000 m, Mann s.n. (lectoparatype of G. ovatum var. flaccidum, CAL photo); Laitlynkot [25.45°, 91.8333°], Khasi Hills, 1800 m [6000 ft], 13 July 1949, Koelz 23191 (E); unlocalised. 750 m [2500 ft], 28 June 1903, Younghusband s.n. (K); Sittong, 3000 m, 30 July 1876, King s.n. (lectoparatype of G. ovatum var. flaccidum, CAL photo). nepal. Chandaghery [Chandagiry, 27.7167°, 84.7833°], Aug. 1821, Wall. Cat. 2155 (lectotype K [Herb. Benthamianum]; isolectotypes E, G, K [Herb. Hookerianum], K-W, NY photo, OXF).

habitat. In hill evergreen forest, 600 – 1200 m.

conservation status. Gomphostemma ovatum is known from eight localities (eight collections) with AOO of 24 km² and EOO of 53,454.76 km². The habitat, hill evergreen forests, is in decline due to tourism activity and urbanisation. Hence, the Vulnerable (VU) category is applied.

phenology. Flowering and fruiting June – December.

notes. There are two species of Gomphostemma which have tuber-like roots: G. ovatum and G. mastersii. Floral parts of both are similar to some extent. Moreover, these species occur in similar areas in the Bengal Province. Hooker (1885) and Prain (1891) suggested reducing G. mastersii to a synonym of G. ovatum. However, the leaf indumentum on the upper side of both taxa is different from G. ovatum by having stellate hairs on the lower part of the midrib and 2 – 4-celled simple hairs on the upper side but G. mastersii having stellate hairs on both sides.

21. Gomphostemma stellatohirsutum C.Y.Wu (1965: 148); Li & Hedge (1994: 73). Type: China, Yunnan, Mon-hai [Fo-hai, 21.9667°, 100.4667°], 2100 m, July 1936, Wang 77368 (holotype PE; isotype A n.v.).

Perennial herb up to 1 m tall. Stems erect, robust, obtusely quadrangular with longitudinal grooves, dark brown, with dense stellate or dendroid hairs with a multi-celled stalk. Leaves petiolate, chartaceous; blades elliptic, elliptic-obovate, 10 – 15 (– 23) × 5 – 9 (– 11) cm, apex acute, margin shallowly serrate, base attenuate, upper side tomentose with dense 3 – 4-celled simple hairs, sometimes bifurcate hairs, lower side pale green tomentose with dense stellate hairs; petioles 20 – 50 mm long, with dense stellate hairs. Inflorescence axillary with opposite cymes sessile, unbranched then congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils 3 – 6-flowered, subtended by the upper leaves and cluster of small leaf-like bracts; bracts leaf-like, deciduous, ovate or elliptic-obovate, 13 – 25 × 5 – 14 mm, apex acute, margin serrate, base attenuate, upper side with 3 – 4-celled simple hairs, lower side with dense stellate hairs; bracteoles narrowly ovate, linear, 5 – 10 (– 14) mm long. Flowering calyx narrowly infundibular, 8 – 10 mm long, 10-ridged, outside brown-yellow tomentose with stellate hairs with a 1-celled stalk, inside glabrous below; tube 6 – 7 mm long; lobes narrowly triangular, 2 – 3 mm long, apex acute, with stellate hairs on both sides. Fruiting calyx infundibular, 13 – 15 mm long; tubes 6 – 7 mm long; lobes narrowly triangular, 7 – 8 mm long, apex acuminate. Corolla white or pale yellow, 25 – 40 mm long, throat inflated, abruptly dilated near throat, outside with dense stellate hairs, inside glabrous; tube slightly incurved, 23 – 30 mm long; posterior lip ovate, 5 mm long, apex obtuse; anterior lip 3-lobed, spreading; median lobe ovate, shorter than lateral lobes. Staminal filaments fleshy, sparsely tomentose. Style slender, glabrous, shorter than corolla with apex unequally lobed. Nutlets solitary, obovate, 4 – 5 × 5 – 6 mm.

distribution. China (Yunnan). Map 6.

Map 6

Distribution range of Gomphostemma stellatohirsutum, G. sulcatum, G. velutinum and G. microcalyx.

specimens examined. china. Yunnan. Mon-hai [Fo-hai], 2100 m, July 1936, Wang 77368 (holotype PE); Caiyanghe Nature Reserve [22.5333°, 101.1667°], 1340 m, 11 July 2009, Xiang 375 (KUN).

habitat. In hill evergreen and forest, 2100 m.

conservation status. Gomphostemma stellatohirsutum is known only from the type. The Data Deficient (DD) category is applied.

phenology. Flowering and fruiting in July.

notes. Gomphostemma stellatohirsutum is a distinctive species recognised by its flowering calyx indumentum of dense stellate hairs and cymes subtended by cluster of small leaf-like bracts, 5 – 14 mm wide. It is morphologically similar to G. hirsutum from Sabah, both species having dark brown stems with long branched hairs.

22. Gomphostemma sulcatum C.Y.Wu (1965: 149); Li & Hedge (1994: 74). Type: China, Yunnan, Ping-bien, 8 Dec. 1934 [22 Aug. 1934], Tsai 61603 (holotype PE; isotypes A n.v., IBSC).

Perennial herb up to 1 m tall. Stems erect, obtusely quadrangular with longitudinal grooves, tomentose with dense stellate hairs with a 1-celled stalk, dendroid hairs absent. Leaves petiolate, chartaceous; blades elliptic or elliptic-ovate, 18 – 28 × 7 – 15 cm, apex acute, margin shallowly serrate, base attenuate, upper side tomentose with 1 – 2 (– 3)-celled simple hairs, lower side pale green tomentose with dense stellate hairs; petioles 20 – 50 mm long. Inflorescence axillary with opposite cymes sessile, unbranched then congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils many-flowered, subtended by the upper leaves and cluster of small leaf-like bracts; bracts leaf-like, ovate, 20 – 30 × 8 – 15 mm, apex acute, margin shallowly serrate, base obtuse, upper side dark green with 1 – 2 (– 3)-celled hairs, lower side pale green with dense stellate hairs; bracts narrowly lanceolate, linear or falcate, 5 – 20 mm long. Flowering calyx tubular-infundibular, 9 – 14 mm long, 10-ridged with veinlets at each sinus; tube 5 – 8 mm long, outside dense tomentose with short simple hairs, inside glabrous; lobes triangular, 4 – 6 mm long, apex acuminate, with stellate hairs on both sides. Fruiting calyx not seen. Corolla yellow, 30 – 40 mm long, throat inflated, abruptly dilated near the throat, outside tomentose, inside glabrous; tube slender, slightly incurved, 25 – 33 mm long; posterior lip ovate, 5 – 6 mm long, apex and margin undulate; anterior lip 3-lobed, spreading beyond posterior lip; median lobe obcordate, base constricted; lateral lobes with round apex. Staminal filaments fleshy, tomentose at apex. Style slender, glabrous, inclined under posterior lip with apex subequally lobed. Nutlets not seen.

distribution. China (Yunnan). Map 6.

specimens examined. china. Yunnan. Ping-pien Hsien [22.9°, 103.6667°], 1100 m, 22 Aug. 1934, Tsai 61603 (holotype PE, isotype IBSC).

habitat. In hill evergreen forest, 1100 m.

conservation status. Gomphostemma sulcatum is known only from the type. Hence, the Data Deficient (DD) category is applied to this species.

phenology. Flowering August.

notes. Gomphostemma sulcatum is a distinctive species defined by having a corolla much exserted from the flowering calyx and the calyx glabrous inside. It is morphologically similar to G. niveum by having a cluster of small leaf-like bracts exceeding the calyx and congested cymes. This species is found close to the border between China and Vietnam. This species may be present in Vietnam as well but further botanical collection in Northern Vietnam is needed to confirm this.

23. Gomphostemma velutinum Benth. (Bentham in Wallich 1830b: 13); Hooker (1885: 699); Prain (1891: 269); Mukerjee (1940: 212). Type: India, Sillet, Jentya Moutain, Jan. 1829, Bruce s.n. in Wall. Cat. 2709 (K [Herb. Benthamianum, K000928174], lectotype selected here; isolectotype K-W [K001116878]).

Erect herb up to 0.4 m tall, without tuber-like roots. Stems slender, upright, sometimes rooting at nodes, obtusely quadrangular with shallowly longitudinal grooves, internodes curving, densely tomentose with stellate hairs and sometimes dendroid hairs. Leaves petiolate, chartaceous; blades ovate, elliptic, elliptic-globose or elliptic-ovate, 2 – 15 × 2 – 8 cm, apex acute, margin serrate, base attenuate or obtuse, upper side densely tomentose with a mixture of hairs, lower side pale green tomentose with dense stellate hairs; petioles 10 – 50 mm long. Inflorescence axillary with opposite cymes sessile, unbranched then congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils 1 – 2-flowered, subtended by the upper leaves; bracts lanceolate or elliptic-oblong, 4 – 10 × 3 – 5 mm long, with dense simple hairs and stellate hairs on both sides; bracteoles linear, 2 – 4 mm long. Flowering calyx infundibular, 9 – 13 mm long, 10-ridged; tube 5 – 6 mm long, outside with simple hairs and stellate hairs, inside with adpressed simple hairs; lobes triangular, 4 – 7 mm long, outside with a mixture of hairs, inside with simple hairs. Fruiting calyx not seen. Corolla white 25 – 30 mm long, throat inflated, abruptly dilated near the throat, outside with sparse simple hairs, inside glabrous; tube slightly incurved, 15 – 25 mm long; posterior lip obovate 10 × 7 mm, apex and margin undulate; anterior lip spreading, as long as posterior lip; median lobe elliptic-obovate; lateral lobes with round apex. Staminal filaments slightly fleshy. Style slender with apex subequally lobed. Nutlets not seen.

distribution. India (Assam, Meghalaya), Bangladesh. Map 6.

specimens examined. india. Assam. Mach s.n. (K [Herb. Hookerianum]); Meghalaya. Jentya Mt [Jaintia], Jan. 1829, Bruce s.n. in Wall. Cat. 2709 (lectotype K; isolectotype K-W); Khasia, Churra, 13 June 1850, Hooker & Thomson 636 (K); Khasia, 1200 m [4000 ft], Hooker & Thomson 48 (P); Mont Khasia, 1200 m [4000 ft], Hooker & Thomson s.n. (BM, C, E, G, L 2 sheets, P); Cherrapunjee, Khasi Hills, 1200 m [4000 ft], 6 July 1952, Chand 6005 (E).

habitat. In mixed forest or evergreen forest, 400 – 1200 m.

conservation status. Gomphostemma velutinum is known from five localities and has an EOO of 5020 km² exceeding the threshold for Endangered. Google Earth imagery shows the evidence of habitat loss of habitats due to urbanisation. Hence, the Vulnerable (VU) category is applied (VU D2).

phenology. Flowering and fruiting June – January.

notes. Gomphostemma velutinum is recognised by having a soft indumentum on both sides of leaves and slender stems. It is morphologically similar to G. ovatum by having upright stems and curving internodes.

I) Glabrous group: iii. Intermediate between groups i and ii, due to variation in corolla (species 24 only)

24. Gomphostemma microcalyx Prain (1890: 316, 1891: 251); Keng (1969: 82, 1978: 321); Walsingham & Bramley (2010: 483); Walsingham (2019: 202). Type: Malaysia, Ulu Bubong in dense bamboo forest, 120 – 180 m [400 – 600 ft], July 1886, Kunstler 10445 (lectotype K [K000249838], selected by Keng (1969)).

Herb up to 1.5 m tall. Stems upright, obtusely quadrangular with longitudinal grooves, tomentose with stellate hairs with a 1-celled stalk, dendroid hairs absent; internodes sometimes wavy. Leaves petiolate, chartaceous; blades narrowly ovate-elliptic or elliptic, 13 – 23 × 4 – 9 cm, apex acute, margin shallowly serrate or subentire, base attenuate, upper side with stellate hairs, lower side pale green tomentose with dense stellate hairs; petioles 25 – 40 (– 70) mm long, with dense stellate hairs. Inflorescence axillary with opposite cymes unbranched then very congested with axis not visible between flowers, up to 20 mm wide when pressed, excluding corolla, always inserted at the upper nodes at which leaves are usually present; verticils few-flowered, subtended by the upper leaves only; bracts falcate, 2 – 3 mm wide, upper side with stellate hairs, lower side glabrous or sparsely tomentose; bracteoles linear, 4 – 7 mm long. Flowering calyx infundibular, 5 – 8 mm long, 10-ridged, outside tomentose with dense stellate hairs with a 1-celled stalk, inside tomentose with adpressed shortly simple hairs; tube 3 – 5 mm long; lobes ovate-triangular tapering into the apex, 2 – 3 mm long, apex acuminate. Fruiting calyx cupular, 6 – 10 mm long; tube 4 – 6 mm long; lobes triangular, 2 – 4 mm long, apex acuminate, margin flat in cross section. Corolla orange, incurved or straight, 15 – 20 mm long in straight form, 25 – 30 mm long in incurved form, throat narrow in straight form or inflated in incurved form; tube exserted from the calyx, slender, not enlarged or sometimes gradually dilated towards the throat in straight form or abruptly dilated near the throat in incurved form, 10 – 15 mm long in straight form, 20 – 25 mm long in incurved form, outside tomentose with simple hairs or short glandular-like hairs, inside glabrous; posterior lip ovate-elliptic, 3 – 4 mm long for straight form, 5 – 6 mm long for incurved form, apex slightly emarginate; anterior lip 3-lobed, slightly shorter than posterior lip for incurved form or slightly longer than posterior lip for straight form; lateral lobes spreading, folded downward. Staminal filaments yellow, fleshy, tomentose at the upper part, included in straight form or slightly exserted in incurved form from underneath posterior lip. Style yellow, slender, shorter than corolla with apex unequally lobed. Nutlets solitary, globose, 5 mm long. Fig. 6C – D, H.

distribution. Peninsular Malaysia, Malaysian Borneo (Sabah, Sarawak), Indonesia (Sumatra, Java, Kalimantan). Map 6.

specimens examined. malaysia. Negeri Sembilan. Genting Peras [3.1333°, 101.9667°], 7 July 1985, Kiew 1866 (SING); Perak. Bujong Malacca [4.3333°, 101.2°], Sept., Ridley s.n. (SING); Larut [4.8°, 100.75°], 540 – 600 m [1800 – 2000 ft], Aug. 1881, Kunstler 2155 (P); ibid., June 1884, Scortechini 942 (lectoparatype CAL photo); Larut Hills [4.85°, 100.7833°], 1891, Ridley s.n. (SING); Larut [4.8°, 100.75°], 540 – 600 m [1800 – 2000 ft], Aug. 1881, Kunstler 2155 (CAL, K); Maxwell Hill, 4^th mile [4.85°, 100.7833°], 600 m [2000 ft], 5 Dec. 1965, Shah & Sidek 1135 (GH, K, L, SING); Tapa [Tapah, 4.1833°, 101.2667°], Wray 835 (lectoparatypes CAL photo, G, SING 2 sheets); Ulu Bubong, 120 – 180 m [400 – 600 ft], July 1886, Kunstler 10445 (lectotype K); Selangor. Ulu Bubong, 120 – 180 m [400 – 600 ft], July 1886, Kunstler 10445 (lectotype K); Gombak, Ulu Gombak [3.3167°, 101.7667°, descent to river at 22^nd mile, 420 m [1400 ft], 14 April 1962, Burtt & Woods B1611 (E 2 sheets); Sabah. Tawau, Tawau Hill Park, 4°23'N 117°53'E, 30 June 2006, Julia et al. SAN 147037 (K); ibid., 347 m, 30 June 2006, Suzana et al. SAN 147801 (K); Tawau Hill Park, trail to Air Panas, 4°23'N 117°53'E, 29 June 2006, de Kok et al. SAN 147206 (K); Sarawak. Matang [1.5833°, 110.2333°], July 1905, Ridley s.n. (K, SING); Gunung Berloban, 10^th km of Tebakang-Tebedu Road [1.1145°, 110.4612°], 120 m, 1 July 1983, Yii & Othman S46181 (K, L); Bintulu, Tatau [2.8833°, 112.85°], Bukit Sarang, 90 m [300 ft], 14 March 1965, Anderson S20955 (K); Bintulu, Ulu Sungai Kakus, Bukit Sarang, Batu Anyi, 2°65'N 100°00.30'E, 8 Oct. 2004, Rantai et al. S93323 (SING); Bintulu, Ulu Sungai Kakus, Bukit Sarang, Sungai Sarang [6.6°, 116.5333°] to Batu Rusa, 16 July 2006, Skornickova et al. 170 (SING); Gunong Medang [1.5922°, 110.2055°], 580 m, 8 Aug. 1966, Ashton & Hou 608 (L); Kuching Distr., Bukit Baju on the Padawan Road near Kampung Tiang Ba Kap, 1°10'N 110°16'E, 100 – 200 m, 10 May 1996, Beaman et al. 12070 (K); Kuching, 1^st Division, Bau, Gunung Ropih [1.3833°, 110.1333°], 25 June 2002, Connie et al. 3705 (SING); Kuching, Tiang Bekap, Mt Mentawa, 1°12'N 110°23'E, 75 m [250 ft], 25 July 1963, Chew 660 (L, SING); Lundu, Sungei Lundu near foot of Gunong Perigi [1.7167°, 109.8167°], 6 Aug. 1962, Burtt & Woods B2695 (E); Serian Distr., Ulu Kedup, Gunong Selabor, 120 m [400 ft], 29 Sept. 1964, Anderson S20828 (K, L); Ulu Baram, Kuba’an path from Bario [3.75°, 115.45°], 1080 m [3600 ft], 22 June 1964, Anderson S20013 (K). indonesia. sumatra. Aceh. North Sumatra, Atjeh, Gunung Leuser Nature Reserve, Ketambe, valley of Lau Alas, near tributary of Lau Ketambe, c. 35 km NW of Kutatjance, 200 – 400 m, 17 May 1972, de Wilde & de Wilde-Duyfjes 12027 (K, L 2 sheets, US); Gunung Leuser Nature Reserve, Gunung Ketambe and vicinity, 8 – 15 km SW [3.6346°, 97.5878°] from the mouth of Lau Ketambe, c. 40 km NW of Kutatjane, Camp 5, 1700 m, 6 Aug. 1972, de Wilde & de Wilde-Duyfjes 14079 (K, L); Gunung Leuser Nature Reserve, Climbing Gunung Mamas, c. 10 – 12 km SW from the mouth of Lau Ketambe, c. 30 km NW of Kutatjane, 1200 – 1800 m, 17 May 1975, de Wilde & de Wilde-Duyfjes 16949 (K, L); Gunung Leuser Nature Reserve, Ketambe Research Station and vicinity, Alas R. Valley, c. 35 km NW of Kutacane, 3°40'N 97°40'E, c. 5 km S of Ketambe, along Alas R., 300 – 350 m, 9 June 1979, de Wilde & de Wilde-Duyfjes 18018 (BO, K, L); Gunung Leuser Nature Reserve, Sikundur Forest Reserve, c. 75 km NW of Medan, 3°55'N 98°5'E, Besitang R., 50 – 100 m, 3 Aug. 1979, de Wilde & de Wilde-Duyfjes 19300 (BO, K, L); Aceh Selatan, W of Blangkajeren, 4°11'N 96°57'E, 700 m, 24 Feb. 1999, Argent & Iqbar 99104 (GH); Bampo-serba, 7 July 1939, Pooll s.n. (SING); Kloet Nature Reserve, S Kloet, along the Krung Lembang, vicinity of Pucuk Lembang, 3°05'N 97°25'E, 60 m, 10 July 1985, de Wilde & de Wilde-Duyfjes 19879 (BO, L 2 sheets); Lawe Sigala, near Kutatjane, 23 March 1954, Alston 14643 (BM, L); lower elevation of Gunong Kemiri, [3.7333°, 97.5°], 600 – 900 m, 21 Aug. 1971, Iwatsuki et al. S-755 (K); North Sumatra. E Coast, Aek na Gerger, between Djoema Tombak and Taratak, Tanah Djawa [Tanah Jawa], Simeloengoen [Simalungun], 7 – 8 June 1927, Bartlett 8241 (US); E Coast, Asahan, Bandar Poeloeh [Bandarpoelau, 2.6833°, 99.5167°], 16 Aug. 1925, Yates 1738 (K); E Coast, Asahan, Silo Maradja, June 1927, Bartlett 8680 (US); E Coast, Pargambiran, 18 – 30 Oct. 1933, Boeea 5815 (GH, L); E Coast, Vicinity of Hoeta Bagasan, Asahan, 7 Sept. 1934 – 4 Feb. 1935, Boeea 6833 (GH); Sibolangit [3.3167°, 98.5833°], 400 m, 25 June 1918, Lorzing 5769 (L); West Sumatra. Island of Sipora [-2.1833°, 99.6333°], 28 Oct. 1924, Kloss 14803 (K, SING); Afd L. Koto, Soengai Koeriman, 1000 m, 28 June 1918, Bünnemeijer 3479 (L); Jambi. Batang Soengai [-1.3667°, 102.4167°], 200 m, Sept. 1925, Posthumus 943 (L, SING); unlocalised. South Sumatra, 1881 – 1882, Forbes 1579 (BM, GH, L). java. unlocalised. Anonymous s.n. (L 2 sheets). kalimantan. West Kalimantan. W Borneo, auf dem Bukit Mehipit [Mehebit, -0.5147°, 112.6408°], 500 m, 28 Dec. 1924, Winkler 1113 (E, L); Central Kalimantan. Borneo, Bukit Raya, 0°40'S 112°44'E, 400 m, 22 Jan. 1983, Nooteboom et al. 4557 (BO, GH, L); Bukit Raya, Tumbang Tubus, 0°45'S 112°51'E, [-0.75°, 112.85°], 150 m, 5 Jan. 1983, Veldkamp 8027 (L); Bukit Raya and upper Katingan, Mendawai R. area, upper Samba R., 60 – 80 km NW of Tumbang Samba, 0°30'S 112°50'E, 150 m, 25 Nov. 1982, Mogea & de Wilde 3655 (BO, L 2 sheets); Tumbang Tapi, 1°1'S 113°E, 100 m, 22 Jan. 1983, Veldkamp 8372 (BO, L 2 sheets); South Kalimantan. Kalimantan Selatan, Gunung Besar [-2.7167°, 115.6167°], 800 – 1300 m, 17 Feb. 1979, Murata et al. B-4040 (BO); unlocalised. Borneo, Sungei Pary, 1896 – 1897, Nieawenhuis & Jaheri 1391 (BO).

habitat. In primary or disturbed forest, evergreen forest, dipterocarp forest, 60 – 1800 m.

conservation status. Widespread; Least Concern (LC).

phenology. Flowering May – January. Fruiting November – March.

notes. Gomphostemma microcalyx has two forms of corolla: straight or incurved, both with a small calyx. However, these two forms overlap in distribution and intermediate forms are observed (Fig. 6).

II) Annulate group (species 25 – 31)

25. Gomphostemma hemsleyanum Prain ex Collett & Hemsl. (Hemsley 1890: 116); Prain (1891: 250); Mukerjee (1940: 207). Type: Burma, Meiktila [20.8667°, 95.8667°], Sept. 1888, Collett 887 (CAL [CAL0000020512] photo, lectotype selected here; isolectotype K [K000846320]). (Another syntype, Collet 17, has not been seen.)

Herb up to 0.6 m tall. Stems erect, obtusely quadrangular with longitudinal grooves, with dense stellate hairs and sometimes dendroid hairs. Leaves subsessile, chartaceous; blades elliptic or elliptic-ovate, 4 – 15 × 1 – 7 cm, apex acute, margin deeply crenate or serrate, base attenuate; upper side dark green with bifurcate hairs, 1 – 2 (– 3)-celled simple hairs and sometimes stellate hairs; lower side pale green with dense stellate hairs; petiole 0 – 10 mm long. Inflorescence axillary with opposite cymes sessile, congested with axis not visible between flowers, inserted at the upper nodes at which leaves are usually present; verticils few-flowered; bracts elliptic or elliptic-obovate, 10 – 15 × 3 – 6 mm, upper side with simple hairs, lower side with dense stellate hairs; bracteoles narrowly ovate, 5 – 8 × 0.5 – 1 mm. Flowering calyx infundibular, 8 – 12 mm long, 10-ridged, outside with dense stellate hairs, inside with adpressed simple hairs; tube 5 mm long; lobes triangular, 3 – 7 mm long, apex acute. Fruiting calyx ovate-infundibular, 11 – 18 mm long, outside with dense stellate hairs, inside with adpressed simple hairs; tube 5 – 7 mm long; lobes triangular, 6 – 11 mm long, 3 – 4 mm wide at base, apex acute. Corolla 10 – 15 mm long, throat narrow, gradually dilated towards the throat, outside glabrous, inside with an annulus of simple hairs; tube slender, incurved, 5 – 7 mm long, included in calyx. Staminal filaments fleshy. Style slightly fleshy, linear with apex unequally lobed. Nutlets 4, elliptic-oblong or elliptic, 3 – 4 × 2 – 3 mm long, wrinkled.

distribution. Burma. Map 7.

Map 7

Distribution range of Gomphostemma hemsleyanum, G. heyneanum, G. microdon, G. nutans, G. pedunculatum and G. repentum.

specimens examined. myanmar. Meiktila, Sept. 1888, Collett 887 (lectotype CAL photo; isolectotype K).

habitat. In mountain areas.

conservation status. Known only from the type collection made in September 1888. Hence, the Data Deficient (DD) category is applied to this species.

phenology. Flowering and fruiting September.

notes. Gomphostemma hemsleyanum is recognised by having an axillary inflorescence, no soft hairs on leaves and is found in North Burma

Keng (1969) recognised a specimen, Backer 8241 (BO), from East Java as Gomphostemma hemsleyanum probably by comparing the specimen with an illustration [Ann. Roy. Bot. Gard. (Calcutta) 3 (1891), plate 82] from Prain’s revision (1891). Unfortunately, this collection has not been examined.

The epithet hemsleyanum is dedicated to Mr W. B. Hemsley who assisted Prain preparing a revision of Gomphostemma (Prain 1891).

26. Gomphostemma heyneanum Wall. ex Benth. (Bentham in Wallich 1830b: 12); Prain (1891: 248); Mukerjee (1940: 206). Type: India, 22 Sept. 1816, Herb. Heyne in Wall. Cat. 2152.1 (K [Herb. Benthamianum], lectotype selected here; isolectotype K-W [K001115333]).

Gomphostemma strobilinum var. heyneanum (Wall. ex Benth.) Hook.f. (Hooker 1885: 696). Type: as for G. heyneanum Wall. ex Benth.

Gomphostemma heyneanum var. rottleri Prain (1891: 249). Type: India, Madras, Nilgiris, Mudumalai Forest, 600 m [2000 ft], Aug. 1886, Gamble 17983 (K, lectotype selected here; isolectotypes, BM, CAL n.v.).

Herb up to 1 m tall. Stems robust, erect, obtusely quadrangular with longitudinal grooves, with a mixture of dense dendroid hairs and stellate hairs. Leaves petiolate, chartaceous; blades ovate or elliptic-ovate, 10 – 18 × 7 – 10 cm, apex acute, margin serrate-dentate, base attenuate, upper side dark green with a mixture of hairs, lower side pale green tomentose with dense dendroid hairs; petioles 10 – 40 mm long. Inflorescence upright, terminal or terminal and axillary, 40 – 100 mm long, with opposite cymes subsessile, congested or lax at the lower verticil; verticils few-flowered; subtending bracts present; bracts ovate or lanceolate, 10 – 25 × 5 – 20 mm, apex acute, margin entire or shallowly serrate, base cuneate or cordate, upper side with stellate hairs, lower side with dense stellate hairs; bracteoles linear, falcate-like, narrowly spathulate or narrowly elliptic-ovate, 5 – 18 mm long. Flowering calyx infundibular, 10 – 13 mm long, 10-ridged, with dense stellate hairs on both sides; tube 5 – 8 mm long; lobes spreading, triangular-ovate, 5 – 6 mm long, apex acute. Fruiting calyx infundibular, 10 – 15 mm long; tube 5 – 9 mm long; lobes triangular-ovate, 5 – 6 mm long, 3 – 4 mm wide at base, apex slightly acuminate, margin slightly folded outward. Corolla yellow or blue-purple, 13 – 20 mm long, throat narrow, gradually dilated towards the throat; tube incurved, 8 – 12 mm long, included in calyx, outside sparse simple hairs, inside with an annulus of simple hairs; posterior lip hooded, ovate, 5 – 8 mm long, apex emarginate, margin entire or slightly undulate; anterior lip spreading, platform-like, 5 – 10 mm long, apex emarginate or undulate; median lobe tongue-like, apex emarginate, margin entire or undulate, base constricted. Staminal filaments fleshy, tomentose. Style slightly fleshy, glabrous with apex subequally lobed. Nutlets 3 – 4, obovate, 3 – 4 mm long, wrinkled.

distribution. South India, Vietnam. Map 7.

specimens examined. india. Karnataka. Bababudda [Bababuddan Hill, 13.5°, 75.7333°], Sept. 1816, Herb. Rottlerianum s.n. (K); Coorg [Kodagu Distr., 12.4208°, 75.7397°], Wottecolli, 18 Oct. 1913, Bourne 6242 (K); Mysore, Hassan [12.95°, 76.0833°], Bannuhalla, 4 Sept. 1970, Jarrett et al. 650 (K); Kerala. Idikki Distr., Thekkady [9.6°, 77.2°], 875 m, 8 Oct. 1976, Vivekananthan 48614 (CAL); Palghat [10.77°, 76.65°], Thadikundu, 925 m, 9 Nov. 1976, Vajravelu 48896 (CAL); Kottayam near Thekkadi, around Periyar Lake [9.4667°, 77.1667°], 800 m, 9 Sept. 1973, Cook et al. 180 (K); Calicut [Kozhikode, 11.25°, 75.77°] Distr., Nilamboor [Nilambur], 300 m, 3 Nov. 1976, Suresh 22326 (L); Tamil Nadu. Madras, Anamallai Hills [Anaimalai Hills, 10.3°, 77°], 900 m [3000 ft], Sept. 1884, Gamble 14628 (K); Anamallay Forests, Beddome s.n. (BM); Madras, Nilgiris, Mudumalai Forest, 600 m [2000 ft], Aug. 1886, Gamble 17983 (isolectotypes of G. heyneanum var. rottleri, BM, K); Courtallum [Courtallam, 8.9167°, 77.2833°], 4 Oct. 1814, Anonymous s.n. [‘Stachys indica’] in Wall. Cat. 2152B (K, K-W); ibid., Herb. Madras [‘Stachys indica’] in Wall. Cat. 2152C (CAL, K-W); Madras [11°, 78°], Herb. Stocks s.n. (K, K [Herb. Hookerianum]); unlocalised. South India: Paulghautcherry, Aug. 1844, Wight s.n. (K); 22 Sept. 1816, Wall. Cat. 2152.1 (lectotype of G. heyneanum, K [Herb. Benthamianum]; isolectotype K-W). vietnam. Ninh Thuan. Phanrang [Phan Rang], Ca Na [11.3333°, 108.9°], 700 m, 13 Nov. 1923, Poilane 8618 (P).

habitat. In shade, on loam and humus, wet deciduous forest, 600 – 925 m.

conservation status. Widespread and in protected areas throughout South India; Least Concern (LC).

phenology. Flowering and fruiting July – November.

notes. Gomphostemma heyneanum can be distinguished by having a terminal infloresecence and soft hairs on both sides of leaves.

27. Gomphostemma microdon Dunn (1913: 170); Doan (1936: 1031); Li & Hedge (1994: 71); Phuong (2000: 100). Type: China, Yunnan, Szemao [22.7667°, 101.0833°], forest west, 1350 m, Henry 12501 (holotype K [K000928870]; isotypes A n.v., K [K000928871], NY [00000711]).

Gomphostemma deltodon C.Y.Wu (1965: 143). Type: China, Yunnan, Jinghong, Che-li Hsien [21.9833°, 100.8167°], 950 m, Aug. 1936, Wang 75545 (holotype PE; isotype A n.v.).

Erect herb up to 1 m tall. Stems robust, obtusely quadrangular, with dense stellate hairs; internodes straight. Leaves petiolate, spreading, chartaceous; blades elliptic, elliptic-ovate or obovate, 13 – 23 × 3 – 9 cm, apex acute, margin serrate, base cuneate, upper side dark green with sessile stellate hairs, lower side pale green tomentose with dense stellate hairs and dendroid hairs confined on midrib and veins; petioles 20 – 35 mm long. Inflorescence axillary, raceme-like, elongated, lax with axis visible between cymes, unbranched, 50 – 80 mm long with opposite cymes reduced, inserted at the upper nodes at which leaves are usually present; verticils 1 – 2-flowered, subtended by the upper leaves; peduncles 40 – 70 mm long, with dense stellate hairs; bracts exceeding calyx, elliptic or broadly ovate, 15 – 20 × 6 – 9 mm, apex acute, margin shallowly serrate, base cuneate, with stellate hairs on both sides; bracteoles obovate, 7 – 10 × 1 – 2 mm, with dense stellate hairs on both sides. Flowering calyx greyish-light green, infundibular, 5 – 8 mm long, 10-ridges; tube 4 – 6 mm long, outside with dense stellate hairs, inside glabrous; lobes short triangular, 1 – 2 mm long, apex acute, margin marginate, outside with dense stellate hairs, inside with adpressed simple hairs. Fruiting calyx infundibular, 5 – 7 mm long; tube 4 – 6 mm long; lobes triangular, c. 1 mm long, apex acute, margin marginate. Corolla yellow or reddish-yellow, (15 –) 20 – 25 (– 35) mm long, throat inflated, abruptly dilated near the throat, outside with stellate hairs, inside with an annulus of simple hairs; tubes slender, slightly incurved, 20 – 25 mm long; posterior lip semi-circular, 2 – 3 mm long; anterior lip 3-lobed, spreading; median lobe broadly ovate, 7 – 10 × 5 – 10 mm, slightly undulate; lateral lobes ovate, apex round. Staminal filaments slightly fleshy. Style slender, glabrous with apex unequally lobed. Nutlets 1 – 4, elliptic-ovate, 4 – 5 mm long, tomentose.

distribution. China (Yunnan), Vietnam, Thailand, Laos. Map 7.

specimens examined. china. Yunnan. Menglun, Mengla [21.4667°, 101.5833°], 17 Sept. 1983, Sino-American Expedition to Yunnan 191 (KUN 2 sheets); Che-li Hsien, 950 m, Aug. 1936, Wang 75545 (holotype of Gomphostemma deltodon, PE); Szemao, W Forest, 1350 m [4500 ft], Henry 12501 (holotype of G. microdon, K; isotypes K, NY). vietnam. Hoa Binh. Mai Chau, Hang Kia [20.667°, 105°], 3 Oct. 2000, Phuong 3519 (HN 3 sheets); Son La. Phu Yen Distr. [21.1997°, 104.7883°], 9 Aug. 2009, Cuong & Cuong 2095 (HN 2 sheets); Moc Chau [21.833°, 104.75°], 13 April 1988, Phuong 29 (HN 4 sheets); unlocalised. Lemarie s.n. (P 2 sheets); Balansa s.n. (P). laos. Sayabouly. Pak Lai [18.2°, 101.4167°], 1866 – 1868, Thorel 3362 (P 2 sheets); unlocalised. Counillon s.n. (P).

thailand. Chiang Mai. Jawm Tong [Chom Thong], Mae Soi Subdistr. [18.2767°, 98.6376°], Mae Soi Valley, 425 m, 11 Nov. 1990, Maxwell 90-1258 (AAU, L); Nan. Phu Ka [19.1992°, 101.0769°], 400 m, 28 Aug. 1938, Somkid 171 (BKF).

habitat. In disturbed deciduous forest, limestone area, 400 – 1350 m.

conservation status. Gomphostemma microdon has been regularly collected from the field. However, an AOO of 44 km² falls under EN and the EOO of 236,784.14 km² under LC. The Least Concern (LC) category is applied.

phenology. Flowering and fruiting April – November.

notes. Gomphostemma microdon is a remarkable species recognised by having short calyx lobes, 1 – 2 mm long, and spreading inflorescence with bracts exceeding calyx. It is morphologically similar to G. dolichobotrys and G. repentum by having an unbranched, raceme-like inflorescence. Wu circumscribed G. deltodon on the basis of its slightly larger calyx (Wu 1965). However, calyx length is variable throughout the distribution of this species.

28. Gomphostemma nutans Hook.f. (Hooker 1885: 697); Prain (1891: 248); Mukerjee (1940: 206). Type: India, Khasia, Oct. 1849, Simons s.n. (K [Herb. Hookerianum, K000846323], lectotype selected here).

Climbing herb. Stems slender, scrambling, obtusely quadrangular with longitudinal grooves, with stellate hairs and sometimes dendroid hairs. Leaves petiolate or subsessile, chartaceous; blades ovate or elliptic-ovate, 4 – 12 × 2 – 6 cm, apex acute, margin serrate or obtusely serrate, base attenuate, cuneate or obtuse, upper side with stellate hairs and sometimes simple hairs, lower side with dense stellate hairs; petiole 5 – 20 mm long. Inflorescence terminal or terminal and axillary, 20 – 110 mm long, congested; verticils few-flowered; subtending bracts present, spathulate, obovate, lanceolate or elliptic-lanceolate, 10 – 20 × 4 – 10 mm, apex acute, margin entire or serrate, base attenuate, with dense stellate hairs on both sides; bracteoles linear, 3 – 10 mm long. Flowering calyx infundibular, 10 – 15 mm long, 10-ridged, outside with dense stellate hairs, inside glabrous below and densely tomentose above; tube 3 – 7 mm long; lobes spreading, lanceolate-ovate, 7 – 8 mm long, apex acute or slightly acuminate. Fruiting calyx infundibular, 11 – 15 mm long; tube 4 – 7 mm long; lobes lanceolate-ovate, 7 – 8 mm long, apex acute, margin sometimes slightly revolute. Corolla yellow, 20 – 35 mm long, throat inflated, abruptly dilated near the throat; tube slender, incurved, 15 – 30 mm long, outside tomentose, inside with an annulus of simple hairs; posterior lip ovate, 5 – 6 mm long; anterior lip 3-lobed, spreading, 8 – 10 mm long, margin undulate. Staminal filaments fleshy, tomentose. Style slightly fleshy, glabrous, shorter than corolla with apex subequally lobed. Nutlets 1 – 4, obovate or ovate, 3 – 4 mm long, wrinkled.

distribution. India (Assam, Nagaland, Meghalaya, Manipur), Burma. Map 7.

specimens examined. india. Assam. Bamanigaon [Bamungaon, 26.9167°, 94.1667°], Sal Forest, near sea level, 3 Dec. 1949, Koelz 24266 (E); Nagaland. Naga Hills, Narum, 1500 m [5000 ft], Dec. 1907, Meebold 7405 (K); Meghalaya. Khasia, Oct. 1849, Simons s.n. (lectotype K [Herb. Hookerianum)]; ibid., Lobb s.n. (K [Herb. Hookerianum]); Khasia, Nunklow, 1050 m [3500 ft], 2 Nov. 1871, Clarke 16502 (lectoparatype K [Herb. Hookerianum]); Khasia, Nya Bungalhi, 750 m [2500 ft], 3 Oct. 1885, Clarke 40645A (K); Manipur. Mairing [24.75°, 94.3667°], Naga Hills, 1500 m [5000 ft], Dec. 1907, Meebold 7187 (E); Kanglatongbi [Kanglatongi, 24.9833°, 93.9°], 900 m [3000 ft], 5 Oct. 1945, Bullock 635 (K); On the way to Nungba [24.75°, 93.4333°], 900 m [3000 ft], 4 Jan. 1882, Watt 6664 (K). myanmar. Sagaing. Tamu, Upper Chindwin [22.6°, 95.1°], Dec. 1907, Meebold 7598 (E, K); near Kendat [23.7333°, 94.4333°], Sept. 1890, Prazer 374 (K); Suigon, Katha [24.1833°, 96.35°], 20 Nov. 1908, Lace 4411 (E 2 sheets, K).

habitat. In hill evergreen forest, 750 – 1500 m.

conservation status. Gomphostemma nutans has AOO of 40 km². Google Earth imagery shows the evidence of habitat loss due to agricultural activity but the habitats in India remains intact. Hence, the Least Concern (LC) category is applied.

phenology. Flowering and fruiting September – January.

notes. Gomphostemma nutans is easy to recognise by its climbing habit with slender stem. The specific epithet ‘nutans’ which means nodding may derive from the specimen mounted in an artificial position, rendering the inflorescence in a nodding state. It is a similar case for the species name of Clinacanthus nutans (Burm.f.) Lindau (Bongcheewin et al. 2019). The inflorescence may be upright in nature.

29. Gomphostemma pedunculatum Benth. ex Hook.f. (Hooker 1885: 696); Prain (1891: 266); Mukerjee (1940: 212); Li & Hedge (1994: 74); Phuong (2000: 108). Type: India, Khasia Mts, Nurtiung [25.5667°, 92.2167°], 900 m, Hooker & Thomson s.n. (holotype K [Herb. Hookerianum, K000846336]).

Gomphostemma inopinatum Prain (1900: 172), synon. nov. Type: Burma, Kachin Hills [26.6833°, 97.2333°], Langkon Post, 900 m [3800 ft], 24 March 1898, Muqim (Prain’s Collector) s.n. (holotype CAL [CAL0000020516]; isotype K [K000846333]).

Gomphostemma nayarii A.S.Chauhan (1986: 182).Type: India, Meghalaya, Khasi Hill, Nongstoin, 5 Sept. 1985, Chauhan 81993A (holotype CAL n.v., ASSAM n.v.).

Perennial, decumbent herb up to 0.6 m tall. Stems ascending, woody at base, obtusely quadrangular with longitudinal grooves, tomentose with dendroid hairs and stellate hairs with a multi-celled stalk. Leaves petiolate, slightly coriaceous, sometimes variegated; blades ovate or elliptic, 10 – 20 (– 35) × 6 – 15 (– 23) cm, apex acute, margin serrate, base attenuate, upper side with stellate hairs and 2 – 3-celled simple hairs, sometimes with only simple hairs, lower side with stellate hairs and dendroid hairs; petioles 40 – 90 mm long. Inflorescence pedunculate, basal, (20 –) 40 – 130 mm long with opposite cymes congested, inserted at the lower nodes from which leaves mostly fallen; verticils few-flowered; peduncles 15 – 150 mm long, with a mixture of dense branched hairs; bracts dark red or red-orange, obovate or ovate, 5 – 20 × 4 – 17 mm, apex acute, margin entire or shallowly serrate, base attenuate, upper side glabrous or glabrescent, lower side with dendroid hairs and simple hairs confined on margin; bracteoles narrowly lanceolate or oblanceolate, 4 – 10 mm long. Flowering calyx dark red or red-orange at the apical part and paler at the basal part, narrowly infundibular, 8 – 12 mm long, 10-ridged, outside with stellate hairs and simple hairs, inside glabrous below or with adpressed simple hairs above; tube 4 – 7 mm long; lobes lanceolate, 4 – 5 mm long, apex acute. Fruiting calyx infundibular, 13 – 15 mm long, inconspicuously ridged, outside with stellate hairs, inside with adpressed simple hairs; tube 7 – 10 mm long; lobes spreading, triangular, 5 – 6 mm long, apex acute. Corolla yellow or red-orange, 25 – 35 mm long, throat inflated, abruptly dilated near the throat; tube slender, incurved, 20 – 25 mm long, outside tomentose, inside usually glabrous, occasionally with an annulus of simple hairs; posterior lip ovate, 5 – 7 mm long, apex slightly emarginate, margin slightly undulate; anterior lip 3-lobed, subequal. Staminal filaments fleshy, tomentose at apical part. Style slender, slightly longer than corolla tube with apex unequally lobed. Nutlets red-orange, 1 – 4, oblong, 4 – 5 mm long, wrinkled.

distribution. India (Meghalaya), China (Yunnan), Burma, Vietnam, Thailand. Map 7.

specimens examined. india. Meghalaya. Khasia, 900 m [3000 ft], Hooker & Thomson s.n. (K [Herb. Hookerianum], L, P, TCD); Khasia, Bor Panee, 900 m [3000 ft], 2 Oct. 1850, Hooker & Thomson 2444 (K [Herb. Hookerianum]); Khasia, Nurtiung, 900 m [3000 ft], Hooker & Thomson s.n. (holotype of Gomphostemma pedunculatum, K [Herb. Hookerianum]). china. Yunnan. Che-li Hsien [21.9833°, 100.8167°], 850 m, Sept. 1936, Wang 78779 (KUN); Dehong, Yingjiang [24.7167°, 97.9333°], Tongbiguan, 1460 m, 14 July 2009, Xiang 374 (KUN); Gingtung [Ching-tung, 24.4667°, 100.9°], Pang-pieh, 1780 m, 1 Oct. 1939, Li 333 (KUN 2 sheets); Gingtung, Bin-bu, 2200 m, 10 Jan. 1940, Li 2886 (KUN 2 sheets); NW Hills of Yeng Zueh, 25°30'N 98°15'E, 2400 m [8000 ft], Aug. 1925, Forrest 27176 (E); S Hills of Yeng Zueh, 1800 m [6000 ft], July 1919, Forrest 18427 (E, K); Jinping [22.7583°, 103.1639°], Wutaishan, 1300 m, 10 July 1996, Wu et al. 3821 (KUN); Luchun, 23°22'51”N 102°57'23”E, 1100 m, 19 Oct. 2000, Shui & Chen 13283 (KUN); Szemao [22.7667°, 101.0833°], E Mts, 1350 m [4500 ft], Henry 12518A (K); Szemao, W Mts, 1500 m [5000 ft], Henry 12518 (E, K); Tehseling, Shunning [24.5833°, 99.9°], 2000 m, 11 Sept. 1938, Yu 17623 (E 2 sheets, KUN 2 sheets); Western Flank of the Shweli Salwin Divide, 2400 – 2700 m [8000 – 9000 ft], Dec. 1912, Forrest 9442 (E, K). myanmar. Kachin. Upper Burma, Kachin Hills, Langkon Post, 1140 m [3800 ft], 24 March 1898, Muqim (Prain’s Collector) s.n. (holotype of G. inopinatum, CAL photo; isotype K). vietnam. Thua Thien. Binh Tri Thien [now split into Thua Thien Provinc], A Luoi [16.167°, 107.333°], 9 Sept. 1980, Anonymous 659 (HN); Kon Tum. Dak Gley Distr., about 12 km to N of Dak Gley Town (24 km by road), near Mang Ken Village, 1100 – 1200 m, 16 Nov. 1995, Averyanov et al. VH 1710 (AAU). thailand. Chiang Mai. Doi Chiang Dao [19.3833°, 98.9°], 23 Oct. 1926, Put 427 (ABD, BK 2 sheets, BM 2 sheets, L, PHN); Mae Awn [Mae On], Doi Lahn, W Side, 1175 m, 23 Oct. 2005, Maxwell 05-594 (CMUB); Mae Rim, Queen Sirikit Botanical Gardens [18.8991°, 98.8603°], Huai Puu, 17 Nov. 2000, Glamwaewwong 4 (QBG); Mae Ton Luang, 18°58'N 98°20'E, 900 m, 25 Nov. 1993, Larsen et al. 44839 (AAU, K); Chiang Dao, Chiang Dao Wildlife Research Station [19.3622°, 98.9218°], 554 m, 23 Oct. 2010, Bongcheewin & Suddee 888 (BKF); Chiang Rai. Doi Tung, Wat Pra That Doi Tung [20.3241°, 99.8328°], 1375 m, 6 Sept. 2006, Maxwell 06-638 (CMUB, QBG); Khun Korn Waterfall [19.8821°, 99.6169°], 675 m, 11 Oct. 1997, KK 416 (BCU); Nan. Sapan Waterfall, few km N of Ban Bo Kluei, E of Road no. 1081, 19°12'N 101°13'E, 600 m, 7 Sept. 1995, Larsen et al. 46162 (AAU).

habitat. In hill evergreen forest, under shade, along stream, 600 – 1900 m.

conservation status. Widespread; Least Concern (LC).

phenology. Flowering July – December. Fruiting October – March.

notes. Gomphostemma pedunculatum is recognised by having dendroid hairs with a multi-celled stalk and a pedunculate inflorescence inserted at the lower nodes, where leaves have fallen completely (see Bongcheewin et al. 2015). Prain (1891) reported that the corolla tube of G. inopinatum showed an annulus inside. Specimen examination across the species distribution revealed that most characters of G. inopinatum are similar to G. pedunculatum so we place G. inopinatum into synonymy here.

30. Gomphostemma repentum Bongch. sp. nov. Type: Upper Burma, Karenni, Ywathit, Salween Valley, 600 m [2000 ft], 11 Jan. 1913, Micholitz s.n. (holotype K [K001081923]).

http://www.ipni.org/urn:lsid:ipni.org:names:77221538-1

Climbing herb. Stems slender, scrambling, obtusely quadrangular, with longitudinal grooves, with dense stellate hairs; internodes curving appearing wavy, always twisting. Leaves petiolate, chartaceous; blades ovate, 10 – 13 × 7 – 9 cm, apex acute, margin obtusely serrate, base attenuate or round, upper side with coarse 1 – 3-celled simple hairs, bifurcate hairs and stellate hairs confined on midrib; petioles 40 – 60 mm long, with dense stellate hairs. Inflorescence axillary, upright, 30 – 70 mm long, with opposite cymes elongate, raceme-like, unbranched then congested with axis not visible between cymes, inserted at the upper nodes at which leaves are usually present; verticils few-flowered; bracts broadly ovate, 10 – 15 × 10 – 13 mm, apex acute, margin serrate, base obtuse, upper side with a mixture of hairs, lower side with dense stellate hairs; bracteoles linear, 5 – 7 mm long. Flowering calyx infundibular, 10 – 12 mm long, 10-ridged, outside with dense stellate hairs, inside with adpressed simple hairs; tube 7 – 8 mm long; lobes triangular, 3 – 4 mm long, 1.5 – 2.5 mm wide at base. Fruiting calyx not seen. Corolla yellow, 20 – 23 mm long, throat inflated, abruptly dilated towards the throat, outside glabrous, inside with an annulus of simple hairs; tube incurved, 13 – 15 mm long, much exserted from the calyx; posterior lobe hood-like, obovate, 7 – 8 × 4 – 5 mm; anterior lobes spreading; median lobe with slightly bilobed apex; lateral lobes undulate. Staminal filaments fleshy, glabrescent. Disk lobed, tumescent behind. Style slender with equally lobed, cordate-like apex. Nutlets not seen. Fig. 3.

recognition. Gomphostemma repentum is similar to G. dolichobotrys and G. microdon by having an unbranched, elongated, raceme-like inflorescence, but it is a climbing herb with curving stems. Additionally, G. dolichobotrys differs by its bracts that are smaller than the calyx, whereas the bracts exceed the calyx in G. repentum; and the inflorescence of G. microdon has lax opposite cymes with the axis visible between flowers whereas the inflorescence of G. repentum has congested opposite cymes with the axis not visible between flowers. Gomphostemma repentum is found in Northern Burma which neighbours the areas where G. microdon occurs in China and Laos: G. dolichobotrys is found in Sumatra, Indonesia.

distribution. Burma. Map 7.

specimens examined. myanmar. Kayah. Upper Burma, Karenni [Kayah], Ywathit, Salween Valley, 600 m [2000 ft], 11 Jan. 1913, Micholitz s.n. (holotype K).

habitat. On mountain, 600 m.

conservation status. Gomphostemma repentum is known only from the type specimen. The Ywathit Dam is a mega-hydropower-project with 4,500 MW capacity on the Salween River which will affect the valley in which this species grows (Salween Watch Coalition, 2016). Furthermore, extensive logging concessions have been granted around the project site. Therefore, the Critically Endangered (CR) category is applied: CR B1ab(i,ii,iii) and B1ab(i,ii,iii).

phenology. Flowering January.

etymology. The epithet repentum [repens, creeping] refers to its habit.

31. Gomphostemma strobilinum Wall. ex Benth. (Bentham in Wallich 1830b: 12); Hooker (1885: 696); Prain (1891: 246); Doan (1936:1032); Mukerjee (1940: 205); Murata (1976: 190); Phuong (2000: 98). Type: Burma, Prome [Pyay, 18.8333°, 95.2333°], River Irawaddi, 9 Oct. 1826, Wall. Cat. 2151.1 (K [Herb. Benthamianum, K000846330], lectotype selected here; isolectotype K-W [K001115331]).

Gomphostemma strobilinum var. elatius Benth. (Bentham in Wallich 1830b: 12). Type: Burma, Taong Dong [21.9093°, 96.4575°], 12 Nov. 1826, Wall. Cat. 2151.2 (K [Herb. Benthamianum, K000846326], lectotype selected here; isolectotypes BM, K 2 sheets [Herb. Benthamianum, K000846327 & K000846328], K [Herb. Hookerianum], K-W [K001115332]). [Prain (1890) named G. wallichii: one of the specimens he examined was Wall. Cat. 2151.2 (should be at CAL n.v.). To avoid homotypic status, Wall. Cat. 2151.2 is selected as a lectotype of G. strobilinum var. elatius.]

Gomphostemma viride Wall. ex Benth. (Bentham in Wallich 1830b: 12). Type: Burma, Tavoy, 28 Aug. 1827, Gomez s.n. in Wall. Cat. 2153 (K [Herb. Benthamianum, K000846332], lectotype selected here; isolectotype K-W [K001115336]).

Gomphostemma acaule Kurz ex Hook.f. (Hooker 1885: 696). Type: Burma, Arracan, Boronga Island, on the Mingoo Hill, evergreen forest, 240 – 360 m [800 – 1200 ft], Oct. 1869, Kurz s.n. (holotype K [K000846325]; isotype K [K000846324]].

Gomphostemma strobilinum var. viride (Wall. ex Benth.) Hook.f. (Hooker 1885: 696). Type: as for G. viride Wall. ex Benth.

Gomphostemma wallichii Prain (1890: 314). Type: India, Assam, Naga Hills, Phesama [25.7075°, 94.1035°], Sept. 1886, Prain s.n. (K, lectotype selected here; isolectotype CAL n.v.).

Gomphostemma strobilinum var. acaule (Kurz ex Hook.f.) Prain (1891: 247). Type: as for G. acaule Kurz ex Hook.f.

Gomphostemma dentatum Muschl. (Muschler 1907: 270). Type: Thailand, Doi Sutep, c. 800 m, 29 July 1902, Hosseus 280 (holotype K; isotype BM).

Gomphostemma strobilinum var. variegatum Craib (1924: 96). Type: Thailand, Chiang Mai, Doi Sutep, evergreen jungle, 540 – 900 m, 13 Oct. 1909, Kerr 835 (K, lectotype selected here; isolectotype BM).

Gomphostemma lacei Mukerjee (1938: 305). Type: Burma, North Shan States, near Masam Falls, 15 Oct. 1911, Lace 6322 (holotype E [E00301978]).

Gomphostemma strobilinum subsp. acaule (Kurz ex Hook.f.) Murata (1976: 192). Type: as for G. acaule Kurz ex Hook.f.

Gomphostemma strobilinum subsp. wallichii (Prain) Murata (1976: 191). Type: as for Gomphostemma wallichii Prain.

Perennial herb or shrub-like up to 0.2 – 2 m tall. Stems robust, erect, obtusely quadrangular with longitudinal grooves, with a mixture of stellate hairs and dendroid hairs. Leaves petiolate, slightly coriaceous; blades ovate-lanceolate or ovate, 11 – 32 × 7 – 16 cm, apex acute, margin obtusely serrate or serrate, base attenuate, tomentose with a mixture of simple hairs and branched hairs on both sides; petioles 10 – 70 mm long, tomentose. Inflorescence upright, terminal or terminal and axillary, 70 – 200 mm long, with cymes congested or lax at the basal verticil; subtending bracts present, ovate or ovate-lanceolate, 20 – 60 × 10 – 40 mm, apex acute, margin serrate, base cordate, obtuse or cuneate; upper bracts green, scarlet or creamy-white, ovate, 16 – 20 × 9 – 18 mm, apex acute, margin entire, base cuneate or cordate, upper side with sessile stellate hairs, lower side with branched hairs; bracteoles falcate or narrowly elliptic-ovate, 5 – 10 mm. Flowering calyx green or dark red, infundibular, 10 – 20 mm long, 10-ridged, outside with stellate hairs, inside glabrous below and with simple hairs above; tube 6 – 10 mm long; lobes ovate or lanceolate, 4 – 10 mm long, apex acute or acuminate. Fruiting calyx infundibular, 10 – 20 mm long, 10-ridged, with stellate hairs on both sides; tube 5 – 10 mm long; lobes triangular-ovate, 5 – 10 mm long, margin slightly revolute. Corolla bright yellow, creamy-white or scarlet, 20 – 35 mm long, throat inflated, abruptly dilated towards the throat; tube incurved, 15 – 25 mm long, exserted from the calyx, outside with sessile stellate hairs, inside with an annulus of simple hairs; posterior lip ovate, 7 – 11 × 4 – 9 mm, apex slightly emarginate or entire, margin undulate; anterior lip spreading, 8 – 10 mm long, undulate; median lobe obovate, 4 – 5 mm long, apex emarginate, margin entire, or sometimes revolute in cross section; lateral lobes folded downward. Staminal filaments fleshy, glabrous. Style slender, slightly fleshy at the apical part with apex unequally lobed. Nutlets 3 – 4, elliptic-ovate, 4 – 5 mm long, glabrous, wrinkled. Fig. 4.

distribution. India (Manipur, Mizoram, Nagaland), China, Burma, Laos, Thailand. Map 8.

Map 8

Distribution range of Gomphostemma strobilinum.

specimens examined. india. Nagaland. Naga Hills, Kohima, 1200 m [4000 ft], 10 – 23 Sept. 1950, Chand 3628 (E, L); Naga Hills, Kohima, 1410 m [4700 ft], 31 Aug. 1935, Bor 6235 (K); Naga Hills, Kasom, 1200 m [4000 ft], 24 Aug. 1948, Mukerjee 3354A (CAL); Naga Hills, Phesama [25.7075°, 94.1035°], Sept. 1886, Prain s.n. (lectotype of G. wallichii, K); Naga Hills, Piffima [25.75°, 93.9667°], 900 m [3000 ft], 19 Oct. 1885, Clarke 40961 (K); Assam, Yarobami, 1500 m [5000 ft], 4 Sept. 1942, Bor 15948 (BKF); Manipur. Kohima Road, 900 m [3000 ft], Sept. 1891, Collett s.n. (K); Muneypoor, Karong, 1125 m [3750 ft], 14 Nov. 1885, Clarke 42012 (K); Mizoram. Lushai Hills, Kolasib [24.2311°, 92.6756°], 600 m [2000 ft], 22 Jan. 1953, Koelz 32535 (E). china. Yunnan. Menglian [24.6°, 98.8°], Lalei Huigu Village, 17 Oct. 1981, Tao & Li 39472 (KUN). myanmar. Kachin. Bhamo [24.2667°, 97.2333°], 105 m [350 ft], 19 Nov. 1911, Lace 5571 (E 2 sheets, K); Sagaing. Ruby Mines Distr., near Mogok [22.9167°, 96.5°], 1350 m [4500 ft], Oct. 1912, Lace s.n. (E); Upper Burma, Mingin [22.8833°, 94.5°], 1890, Prazer s.n. (K); Shan. Upper Burma, Southern Shan State, Taungyi [20.7833°, 97.0333°], 1893, Khalil s.n. (G 2 sheets, P2 sheets); Upper Burma, Fort Stedman [20.5833°, 96.95°], Nov. 1892, Huk s.n. (K, P2 sheets); South Shan States, Keng Tung [21.3°, 99.65°], 1200 m [4000 ft], Sept. 1909, MacGregor 754 (E); North Shan States, near Masam Falls, 15 Oct. 1911, Lace 6322 (holotype E); Shan Hills, Terai, 600 m [2000 ft], Nov. 1888, Collett 961 (K); SE Shan State, Meongpaon [21.3°, 99.65°], 1350 m [4500 ft], 3 Dec. 1890, Watt 10726 (E); Southern Shan, Loilem [20.9250°, 97.5639°], 1350 m [4500 ft], 17 Sept. 1911, Robertson 431 (K 2 sheets); Upper Burma, Southern Shan State, Laikaw, 1894, Khalil s.n. (L); Upper Burma, Kyali [23.7333°, 94.4333°], 1890, Prazer 419 (ABD, BM, K); Upper Burma, Southern Shan State, Tameinkhan [Thamakan or Thamakhan, 20.6833°, 96.7°], 1895, Khalil s.n. (ABD, TCD); Chin. W Central Burma, Mindat [21.3667°, 93.9833°], 1200 – 1500 m [4000 – 5000 ft], 30 July 1956, Kingdon-Ward 22543 (BM); Mandalay. Maymyo, Oct. 1888, Khan 214 (G); ibid., 1050 m [3500 ft], 30 Aug. 1908, Lace 4245 (E, K 2 sheets, TCD); ibid., 600 m [2000 ft], Jan. 1908, Meebold 8104 (K); Taong Dong, 12 Nov. 1826, Wall. Cat. 2151.2 (lectotype of Gomphostemma strobilinum var. elatius, K [Herb. Benthamianum]; isolectotypes BM, K 2 sheets [Herb. Benthamianum], K [Herb. Hookerianum], K-W); Thoudaung, Oct. 1934, Dickason 5216 (GH); Rakhaing. Arracan, Boronga Island [19.9667°, 93.1°], Mingoo Hill, 240 – 360 m [800 – 1200 ft], Oct. 1869, Kurz s.n. (holotype K; isotype K); Bago. Tharrawaddy [17.65°, 95.8°], 120 m [400 ft], Nov. 1901, Sohool 8 (G); Prome, R. Irawaddi, 9 Oct. 1826, Wall. Cat. 2151.1 (lectotype of G. strobilinum, K [Herb. Benthamianum]; isolectotype K-W); Yangon. Rangoon [16.75°, 96.25°], 11 Jan. 1854, McClelland s.n. (K [Herb. Hookerianum]); Mon. Moulmein [16.4914°, 97.6256°], 1859, Parish 78 (K [Herb. Hookerianum]); Tanintharyi. Tavoy, 28 Aug. 1827, Wall. Cat. 2153 (holotype of G. viride, K [Herb. Benthamianum]; isotype K-W); Tenasserim, Tavoy, 2.5 miles E of Paungdaw Power Station, 14°N 98°30'E, 390 m [1300 ft], 22 Aug. 1961, Keenan et al. 1036 (E); Tenasserim, Tavoy, NW of Paungdaw Village, 300 m [1000 ft], 23 Aug. 1961, Keenan et al. 1229 (E, K); Tenasserim, Tavoy, Hills W of Paungdaw Power Station, 660 m [2200 ft], Aug. 1961, Keenan et al. 898 (E); unlocalised. Meong, R. Irawaddi, 13 Sept. 1826, Wall. Cat. 2151.1 (lectoparatypes of G. strobilinum, G 2 sheets, K [Herb. Benthamianum], K-W). laos. unlocalised. Massie s.n. (P). thailand. Mae Hong Son. Huai Hee, Doi Pui [19.2167°, 98.0667°], 1500 m, 22 Oct. 2000, Suksathan 2825 (QBG); Khun Yuam, 18°15'N 98°E, 600 – 700 m, 5 Sept. 1974, Larsen et al. 34166 (AAU, BKF, K, L); Muang, Huai Bu Ling, 1125 m, 28 Oct. 2007, Maxwell 07-667 (CMUB, QBG); Pai, Ban Pee Lu, 24 Oct. 1996, Na Nakorn et al. 7769 (QBG 2 sheets); Pai, Ban Watchan, 1100 m, 3 Dec. 2007, Wongprasert 712-2 (BKF); Pai, Muang Sroi Waterfall, 800 m, 17 Jan. 1983, Koyama et al. T-32680 (BKF); Pang Ma Pha, Kew Lom [19.4547°, 98.3051°], 1300 m, 5 Sept. 1999, Suksathan 1730 (QBG); Pang Ma Pha, Tam Lawt, Doi Pah Baw, 725 m, 12 Nov. 2004, Palee 787 (CMUB); Pang Ma Pha, Tam Pla, 12 Dec. 2007, Tanaka et al. 8219 (QBG); Chiang Mai. Doi Chiang Dao, 19°25'N 98°55'E, 550 – 1100 m, 3 Dec. 1965, Hennipman 3183 (BKF, C, K); ibid., 1500 m, 2 Dec. 1961, Smitinand & Anderson 7260 (BKF); ibid., 1900 – 2200 m, 3 Dec. 1961, Smitinand & Anderson 7339 (BKF); Doi Chiang Dao, Khun Me Terhy, 1130 m, 31 Oct. 1955, Garrett 1464 (K, L); Doi Chiang Dao, Ridge above Ban Tam, steep slope above caves [19.3944°, 98.9267°], 500 – 600 m, 3 Jan. 1941, Garrett 1219 (E, K, L 2 sheets, P 2 sheets, TCD); Chiang Dao, Ban Bing Kong, 500 m, 29 Oct. 1987, Maxwell 87-1305 (L); Doi Chiang Dao Wildlife Sanctuary, 1700 m, Nov. 1995, Morsi 253 (CMUB); Doi Chiang Dao, SE foothills at Ban Yang Toong Brong Forest Station [19.3833°, 98.9°], 475 – 575 m, 11 Nov. 1989, Maxwell 89-1389 (L); Chiang Dao, Pa Sam Sob, 6 Nov. 1963, Chantanamuck 601 (BK); Chom Thong, Doi Hua Sua [18.4979°, 98.5565°], along the trail extending from the W part to the top in Doi Inthanon National Park, 1600 m, 17 Dec. 1998, Konta et al. 4636 (BKF); Doi Angka [Doi Inthanon, 18.5833°, 98.4833°], 1800 m, 17 July 1922, Kerr 6333 (BK, BM, K 2 sheets); ibid., 550 m, 20 July 1988, Phengklai et al. 6650 (BKF); Doi Inthanon National Park, around Wachirathan Waterfall [18.5418°, 98.5992°], 600 – 700 m, 6 Dec. 1984, Koyama et al. T-39944 (BKF); Doi Inthanon National Park, near Ban Sop Hat, 880 m, 6 Dec. 1984, Mitsuta et al. T-46480 (BKF); Doi Kar [Doi Inthanon], range NE of the Pah Ngeam, 1110 m, 1 Oct. 1910, Garrett 66 (lectoparatypes of G. strobilinum var. variegatum, BM, K, L); Doi Pha Mawn and spur connecting with Doi Angka, 1520 – 1610 m, 3 Sept. 1927, Garrett 435 (AAU, K 2 sheets); en route from Mae Klang Waterfall to Ban Yang, along Nam Mae Klang, on approach to Doi Inthanon, 350 – 600 m, 16 Dec. 1965, Tagawa et al. T-2295 (BKF, L); Changkien [18.8458°, 98.8886°], 1200 m, 26 Oct. 1975, Sadakorn 620 (BK); Doi Pui [17.3667°, 98.7167°], 27 Sept. 1997, BGO staff 9656 (QBG); ibid., 1450 – 1630 m, 16 Dec. 1984, Koyama & Nagamasu T-40151 (AAU, BKF, L); Doi Sutep-Pui National Park, N part, Doi Mawn Luang, above Nong Rai Village, 1475 m, 24 Dec. 1996, Maxwell 96-1684 (BKF, CMUB); Doi Sootep, 420 m [1400 ft], 12 Nov. 1910, Kerr 1541 (BM); ibid., 1050 – 1500 m [3500 – 5000 ft], 4 Sept. 1910, Kerr 1380 (BM, K, TCD); ibid., 540 – 900 m [1800 – 3000 ft], 13 Oct. 1909, Kerr 835 (lectotype of G. strobilinum var. variegatum, K; isolectotype BM); ibid., 600 – 900 m [2000 – 3000 ft], 6 Oct. 1912, Kerr 835A (lectoparatypes of G. strobilinum var. variegatum, BM, K); ibid., c. 800 m, 29 July 1902, Hosseus 280 (holotype K; isotype BM); Doi Sutep, below Puping Palace [18.8°, 98.9°], 1300 m, 31 Oct. 1987, Maxwell 87-1344 (L); Doi Saket [18.8744°, 99.1356°], 31 Oct. 1930, Put 3272 (BK, BM 2 sheets); Fang, Doi Ang Kang [19.8°, 98.9833°], 9 April 1975, Sutheesorn 3158 sht. 1 (BK); Hod, Baw Salee Subdistr., km 53, Hod-Mae Sariang Road, Highway no. 108, Ban Mae Sanam Mai, Mai Muang Nao, 950 m, 28 Sept. 2001, Sankamethawee 287 (CMUB, L); Jawm Tong, Mae Soi Ridge, Mae Soi Subdistr., W of Ban Bah Gluay [18.2768°, 98.6376°], 1625 m, 28 Sept. 1992, Maxwell 92-590 (CMUB, L, P); Mae Jam, Ban Wat Chan Subdistr., Huay Bong Village [19.0645°, 98.2963°], 1125 m, 6 Dec. 1998, Maxwell 98-1427 (BKF, CMUB, L 2 sheets); Mae Rim, Huai Mae Sa Noi [18.9122°, 98.9397°], 500 m, 21 Nov. 1996, Na Nakorn 7827 (QBG); Mae Rim, Mon Long [18.9194°, 98.8355°], 1320 m, 20 Nov. 2001, Glamwaewwong 55 (QBG); Mae Rim, Pong Yaeng [18.8831°, 98.8169°], 1008 m, 11 Jan. 2008, Jatupol 08-176 (CMUB, QBG, XTBG); Mae Sa, N of Chiang Mai, Rock Tower Mt, 18°55'N 98°50'E, 1300 – 1400 m, 18 Sept. 1995, Larsen et al. 46633 (AAU); Mae Song Sob [18.7903°, 98.9817°], 20 May 1995, BGO staff 5315 (QBG); Chiang Mai, Mae Tang, Doi Sahng Liang, S side, upper Gu Gahp Valley, above Pah Dang Village, 1275 m, 21 Sept. 1997, Maxwell 97-1028 (BKF, CMUB); Mae Tang, Doi Tung Jaw, 1200 m, 12 Dec. 1981, Thongisan 36 (CMUB); Mae Tang, Pa Pae, 1000 m, 9 Nov. 1980, Paisooksantivatana & Sangkhachand 378-80 (BK 2 sheets, CMUB); Mae Tang, Pa Pae, Pong Duat hot springs [19.2315°, 98.7009°], 850 m, 19 Nov. 1992, Maxwell 92-741 (CMUB, L); Mok Fah Waterfall, 40 km NE of Chiang Mai, along the road to Pai, Road no. 1095, 19°3'N 98°13'E, 500 m, 24 Nov. 1993, Larsen et al. 44788 (AAU); Muang Krong, Koon Krong [19.5143°, 98.802°], 21 Nov. 1963, Chantanamuck 702 (BK); Wang Tao, 800 m, 13 Feb. 1958, Sørensen et al. 1032 (C, KUN); Om Koi, Bo Luang Tableland, Ban Om Koi to Mae Tuen Watershed improvement station, 1340 m, 17 Oct. 1979, Shimizu et al. T-19136 (BKF); Omkoi, Mae Rah Ah Watershed Station, 1200 m, 13 Aug. 1999, Petrmitr 461 (CMUB, L 2 sheets); Pah Hom Pok [20.1167°, 99.2°], 1400 m, 24 Feb. 1958, Sørensen et al. 1560 (BKF, C, K, KUN); Huai Sai, Huai Nam Dang National Park. [19.3025°, 98.5985°], 1287 m, 20 Jan. 2002, Chayamarit et al. 3217 (BKF 2 sheets); Samoeng, Sanoengtai, Ban Pa Kar [18.9667°, 98.9667°], 1424 m, 8 July 2008, Jatupol 08-339 (QBG 2 sheets); Chiang Rai. Doi Luang National Park [19.2°, 99.7167°], 1175 m, 7 Sept. 1997, Maxwell 97-978 (CMUB); Khun Jae [19.0779°, 99.3978°], 925 m, Maxwell 97-1368 (CMUB, L); Khun Korn Waterfall [19.8821°, 99.6169°], 750 m, 12 Oct. 1997, KK 456 (BCU); Mae Fah Luang, W of Doi Pah Giah, 1200 m, 24 Oct. 2006, Palee 1068 (CMUB); Pahn, Doi Luang National Park, W side, summit area of Doi Mauk [20.1247°, 100.2961°], Sahn Salee, 1230 m, 6 Oct. 1998, Palee 420 (BKF, CMUB, L); Wiang Pa Pao, Mae Chadee [19.1833°, 99.5167°], 1400 m, 29 Sept. 1993, Vial-Debas 10 (CMUB); Thoeng, Ban Mai Ya, 9 Feb. 1970, Sutheesorn 1642 (BK); Phayao. Doi Luang National Park, E side, 610 m, 21 Dec. 1997, Petrmitr 174 (CMUB); ibid., 1200 – 1225 m, 28 Dec. 1997, Petrmitr 227 (CMUB); Nan. Chalerm Pra Kiat, Wiang Peian Waterfall, 450 m, 15 Nov. 2001, Srisanga 2288 (QBG); Lamphun. higher elevation of Doi Khun Tan, 1200 – 1375 m, 5 Sept. 1967, Tagawa et al. T-9288 (AAU, BKF, K); Khun Tan [18.4708°, 99.2847°], 1000 m, 31 Oct. 1977, Chermsirivathana 2264 (BK); Mae Ping National Park, Ko Thung Waterfall, 250 – 300 m, 18 Nov. 1997, Pongamornkul 11 (QBG); Me Kow, 380 m, 4 Nov. 1925, Winit 1525 (BK, BKF 2 sheets, BM, L); Lampang. Doi Khun Tan [18.4708°, 99.2847°], 800 – 850 m, 9 Jan. 1994, Koyama T-62057 (BKF); ibid., 1100 – 1300 m, 30 Oct. 1977, Na Songkhla et al. 349 (BCU 2 sheets); Me Saloi, 200 m, 28 Oct. 1925, Winit 1490 (BKF, BM); Muang Pan, Chae Sorn National Park [18.8364°, 99.4694°], 775 m, 24 Oct. 1995, Maxwell 95-996 (CMUB); Wang Nua, Doi Luang National Park, W side of and below Doi Nawk, 1150 m, 7 Nov. 1998, Petrmitr 323 (CMUB); Phrae. Song, Mae Yom National Park [18.63°, 100.1627°], 400 m, 14 Dec. 1993, Maxwell 93-1502 (CMUB); Tak. Wang Djao [Wang Chao, 16.7042°, 99.2428°], 100 m, 10 Oct. 1904, Hosseus 62 (BM, C, E, K, M, P 2 sheets); Raheng [Tak], Doi Tung Cha [17.2490°, 98.9379°], 500 m, 18 Nov. 1920, Kerr 4595 (BK, BM 2 sheets); Bhumibol Dam [17.25°, 98.9667°], Feb. 1960, Anonymous s.n. (BK); Doi Mussor, 1000 m, 9 Dec. 1961, Smitinand 7069 (BKF, K); Mae Sot Distr., Wat Tham Manorha, 16°46'48”N 98°39'12”E, 500 m, 11 Sept. 2009, Middleton & Triboun 4851 (E); Taksinmaharaj National Park [16.7808°, 98.9286°], 10 Oct. 1992, Herb Trip 282 (BCU); Tha Song Yang, Mae Hla, Mae Sot-Mae Sariang, road no. 105, km 56, 200 m, 22 Nov. 2005, Pooma 5783 (BKF, E); Umphang, Doi Hua Mot, [15.9615°, 98.8573°], 960 m, 5 Nov. 1998, Suddee & Puujaa 1117 (K, TCD); Sukhothai. Kirimat, Ramkamhaeng National Park [16.8709°, 99.6874°], 500 m, 1 Dec. 1987, Paisooksantivatana & Sangkhachand 2185-87 (BK); Muang Gow, near PraTah Due Nahm Doke So Gah Cha Nah Temple, 4 Nov. 1971, Maxwell 71-669 (AAU, BK, L); Phitsanulok. Kaeng Sopa [16.8834°, 100.6567°], 12 Dec. 1966, Sangkhachand 591 (BK); Thung Salaeng Luang National Park, about 20 km E of Phitsanulok, 16°49'N 100°36 – 38'E, 450 – 480 m, 21 Oct. 1984, Murata et al. T-38331 (BKF); Kamphaeng Phet. Kao Son, Ban Nam Tok, 200 m, 24 Nov. 1977, Phengklai et al. 3899 (BKF); Klong Lan Waterfall, 200 – 400 m, 25 Oct. 2001, Watthana 1508 (QBG); Mae Wong National Park [16.1317°, 99.2821°], 300 m, 15 Oct. 1998, van de Bult 139 (CMUB); Nakhon Sawan. Lat Yao, Kao Mae-kar-see, 13 Jan. 1975, Sutheesorn 3158 sht. 2 (BK); Phetchabun. Nam Nao National Park, trail to N of Park Headquarters, 800 m, 8 Dec. 1998, Suddee et al. 913 (K, TCD); Loei. Na Haew National Park, Tad Hueng Fall [17.5664°, 100.9919°], 620 m, 20 Sept. 2000, Wongprasert 009-8 (BKF); Phu Kradung [16.8833°, 101.75°], 14 Oct. 1967, Sangkhachand 1032 (BK); Phu Kradung, 16°52'N 101°48 – 50'E, 900 – 1200 m, 30 Oct. 1984, Murata et al. T-42115 (BKF); Khon Kaen. Chumphae, Dong Lan [16.8278°, 101.9932°], 300 m, 22 Nov. 1958, Smitinand 4918 (BKF); Pha Nok Khao [16.8450°, 101.9551°], 400 m, 9 Sept. 1963, Smitinand & Sleumer 1127 (BKF); Chaiyaphum. Phu Khieo, 16°28'N 101°45'E, 650 m, 9 Nov. 1984, Murata et al. T-50374 (BKF); Nakhon Ratchasima. Pak Chong [14.7042°, 101.4131°], 300 m, 31 Dec. 1923, Marcan 1558 (BM); Pakchong, Khao Yai National Park, Haew Suwat Waterfall [14.4339°, 101.4008°], 700 – 730 m, 19 Nov. 1982, Koyama et al. T-30079 (BKF); Uthai Thani. Klong Pluu, 500 – 600 m, 21 Oct. 1993, Tiptabiankarn 1248 (SLR); Larn Suk Distr., Huay Kha Kaeng Wildlife Sanctuary, on way to Khao Nang Rum Research Station, 300 m, 12 Nov. 1979, Shimizu et al. T-22394 (GH, K, L, SING); Nong Chang, Kao Phra-rhae, 29 Oct. 1974, Sutheesorn & Sangkhachand 3134 (BK); Kanchanaburi. Bo Phloi, Tham Tarn Lod National Park, Khao Kam Paeng, 8 Nov. 1979, Shimizu et al. T-22163 (BKF, L); Erawan National Park, 14°17'N 99°15'E, 400 m, 19 Nov. 1971, van Beusekom et al. 3856 (BKF, C, K, L, P); Si Sawat, Erawan National Park, summit of Khao Kro Krae Mt, 990 m, 1 Nov. 2003, van de Bult 701 (BKF, CMUB); Hard Palom [14.0476°, 99.2161°], 250 m, 25 Dec. 1961, Larsen 8953 (C); Huay Bankau, 14°55'N 98°45'E, 750 m, 7 Nov. 1971, van Beusekom et al. 3480 (BKF, C, L, P); Khao Kampaeng, Chalerm Rattanakosin National Park, 1240 m, 26 Sept. 2001, van de Bult 478 (BKF, CMUB); km 53 Thong Pha Phum to Sangkhlaburi, 300 m, 2 Oct. 1998, Suddee et al. 889 (K, TCD); Sai Yok [14.4167°, 98.85°], 250 m, 14 Dec. 1961, Larsen 8718 (C); Sai Yok, Wang Khamen [14.5833°, 99.0333°], 200 m, 23 Sept. 2000, Phengklai et al. 14058 (BKF); Thong Pha Phum, Ban Linthiu, 150 m, 26 Aug. 1999, Wongprasert 998-04 (BKF); Phetchaburi. Tung Luang [Thung Luang, 13.1°, 99.95°], under 50 m, 9 Nov. 1931, Kerr 20610 (BK, BM 2 sheets); Saraburi. Hin Lap [14.6630°, 101.1331°], 20 Aug. 1929, Put 2425 (BK, BM 2 sheets, L, PHN); Muak Lek, W of Forest Station, 300 m, 2 Dec. 1963, Smitinand & Sleumer 1323 (BKF, K, L, SING); Chon Buri. Khao Khieo opened zoo [13.2149°, 101.0566°], 300 m, Phengklai et al. 12796 (BKF); Sriracha [13.1667°, 100.9333°], 60 m [200 ft], 30 Sept. 1929, Collins 1038 (BK, BM, L); Rayong. Klaeng, Khao Cha Mao National Park [12.9125°, 101.7221°], 24 Nov. 1979, Shimizu et al. T-23517 (BKF); Phangnga. Kuraburi, Klong Saeng Wildlife Sanctuary [9.1667°, 98.5833°], 11 Sept. 1984, Fukuoka & Na Nakhon T-36105 (BKF).

habitat. In primary or disturbed lowland forest, evergreen forest, mixed deciduous forest, dipterocarp forest, on limestone, 60 – 2200 m.

conservation status. Widespread; Least Concern (LC).

phenology. Flowering May – December. Fruiting October – February.

notes. Gomphostemma strobilinum is widespread in continental South East Asia. Continuous morphological variation is observed in this species (Fig. 4), especially in Burma and Thailand where most specimens of it have been collected. Gomphostemma strobilinum has been described under several names depending on these variable characters, for example, G. wallichii (forms with cordate bracts), G. variegatum, G. acaule. The characters used to diagnose these forms include the height of plant, colour and shape of bracts, and leaf decoration, all found to be variable based on field observations and herbarium specimens seen in this study. Intermediate forms between previously recognised taxa are frequently seen.

This species is easily recognised by having upright, mostly terminal or terminal and axillary inflorescences, with the inflorescences terminating leafy shoots.

Excluded names

Gomphostemma flavescens (Zoll. & Moritzi) Miq. (Miquel 1859: 987). [a synonym of Cymaria dichotoma Benth.].

Gomphostemma macrophyllum Miq. (Miquel 1859: 987). [a synonym of Paraphlomis oblongifolius (Blume) Prain].

Gomphostemma membranifolium Miq. (Miquel 1859: 988). [a synonym of Paraphlomis javanica (Blume) Prain].

Gomphostemma petiolare Miq. (Miquel 1859: 987). [a synonym of Paraphlomis javanica].

Gomphostemma insuave Hance (1884: 231). [a synonym of Microtoena insuavis (Hance) Prain ex Briq.].

Gomphostemma luzonense Elmer (1908: 339). [a synonym of Paraphlomis javanica].

Gomphostemma sumatrense Ridl. (Ridley 1923b: 85). [a synonym of Paraphlomis javanica].

Appendices

Appendix 1

Gomphostemma accounts in India (Hooker 1885^A; Mukerjee 1940^B), Bangladesh (Khanam et al. 2003^C), China (Li & Hedge 1994^D), Burma (Kress 2003^E), Vietnam (Phuong 2000^F), Laos & Cambodia (Doan 1936^G), Thailand (Murata 1971^H & 1976 ^I) and the Malesian region (Keng 1969^J; Walshingham & Bramley 2010 ^K).

Table 2

NB. Taxon names printed in bold are accepted names. Gomphostemma dolichobotrys Merr., G. flexuosum Bongch., G. keralense Vivek., Gopalan & Ansari, G. longipetalum Bongch. and G. repentum Bongch., which are not included in the regional accounts, are omitted from Appendix 1.

Appendix 2. Alphabetical index to collections examined

Adelbert 460 (G. javanicum); Afriastini 715 (G. javanicum); Ahern's collector 1866 (G. javanicum); Alcasid 1894 (G. javanicum); Alston 13966 (G. javanicum); 14057 (G. javanicum); 14265 (G. javanicum); 14643 (G. microcalyx); 14680 (G. curtisii); 15219 (G. javanicum); Alvins 2130 (G. crinitum); Anderson 1093 (G. ovatum); S20013 (G. microcalyx); S20828 (G. microcalyx); S20955 (G. microcalyx); S31794 (G. javanicum); Ang FRI 27573 (G. crinitum); Anonymous s.n. [‘Stachys indica’] in Wall. Cat. 2152B (G. heyneanum); Anthonysamy SA 1094 (G. crinitum); Argent & Iqbar 99104 (G. microcalyx); 99126 (G. javanicum); Argent et al. 18 (G. javanicum); Ashton & Hou 608 (G. microcalyx); Averyanov et al. VH 252 (G. grandiflorum); VH 1710 (G. pedunculatum); Awa & Paie S 45593 (G. javanicum).

Backer 2430 (G. javanicum); 2517 (G. javanicum); 3304 (G. javanicum); 6531 (G. javanicum); 6731 (G. javanicum); 9375 (G. javanicum); 10976 (G. javanicum); 12052 (G. javanicum); 12217 (G. javanicum); 21878 (G. javanicum); 23351 (G. javanicum); 37120 (G. javanicum); Balansa 1000 (G. leptodon); 2930 (G. leptodon); 2931 (G. leptodon); 2932 (G. leptodon); 2933 (G. javanicum); 2934 (G. javanicum); 2937 (G. chinense); Bangham & Bangham-Masters 728 (G. dolichobotrys); 736 (G. javanicum); Barbon et al. 12269 (G. javanicum); Bartlett 6527 (G. javanicum); 7370 (G. mastersii); 8176 (G. mastersii); 8241 (G. microcalyx); 8680 (G. microcalyx); Bartlett & La Rue 224 (G. javanicum); Beaman 7778 (G. hirsutum); 10893 (G. hirsutum); Beaman & Perkins 8486 (G. javanicum); Beaman et al. 7037 (G. curtisii); 7059 (G. curtisii); 7495 (G. javanicum); 8680 (G. javanicum); 10666 (G. javanicum); 11173 (G. javanicum); 12070 (G. microcalyx); Beddome 243 (G. eriocarpon); 244 (G. eriocarpon); Belcher 241 (G. parviflorum); Berhaman & Madani SAN 134509 (G. javanicum); Best 14117 (G. javanicum); BGO staff 73 (G. strobilinum); 1214 (G. javanicum); 1464 (G. javanicum); 3952 (G. javanicum); 4200 (G. strobilinum); 5315 (G. strobilinum); 5371 (G. strobilinum); 6929 (G. javanicum); 9656 (G. strobilinum); Bhargava & Nooteboom 6311 (G. javanicum); Bhattachcharya & Party 31167 (G. parviflorum); Binh & Cuong 1536 (G. parviflorum); Boeea 760 (G. javanicum); 4317 (G. javanicum); 5738 (G. microcalyx); 5815 (G. microcalyx); 6414 (G. javanicum); 6467 (G. javanicum); 6469 (G. javanicum); 6833 (G. microcalyx); 8795 (G. mastersii); 9848 (G. mastersii); 9896 (G. javanicum); Bold 1301 (G. javanicum); Bon 2272 (G. leptodon); 2807 (G. leptodon); Bongcheewin & Suddee 888 (G. pedunculatum); Boonkird 30 (G. strobilinum); Boonnab 18 (G. javanicum); Bor 5075 (G. parviflorum); 6235 (G. strobilinum); 15948 (G. strobilinum); Bourdillon 41 (G. eriocarpon); Bourne 6242 (G. heyneanum); 6271 (G. heyneanum); Boyce 1026 (G. parviflorum); Bramley et al. 147246 (G. javanicum); Britton 328 (G. javanicum); Bruce s.n. in Wall. Cat. 2709 (G. velutinum); Bullock 635 (G. nutans); Bunchai 84 (G. parviflorum); 205 (G. javanicum); 250 (G. parviflorum); 688 (G. strobilinum); 958 (G. strobilinum); 1000 (G. strobilinum); 1245 (G. strobilinum); 1315 (G. strobilinum); Bunchai & Nimanong 1417 (G. javanicum); Bünnemeijer 505 (L0796107, G. javanicum); 505 (L0796120, G. curtisii); 2747 (G. javanicum); 2631 (G. javanicum); 3479 (G. microcalyx); 3599 (G. javanicum); 3889 (G. javanicum); 4692 (G. javanicum); 4744 (G. javanicum); 8230 (G. javanicum); 8557 (G. javanicum); 8868 (G. javanicum); 9299 (G. javanicum); Bunpheng 208 (G. strobilinum); 256 (G. strobilinum); Burkill 11075 (G. javanicum); 12484 (G. crinitum); 37269 (G. aborense); Burkill & Haniff 6298 (G. crinitum); Burkill & Holttum 8650 (G. javanicum); Burtt & Woods B1611 (G. microcalyx); B2687 (G. javanicum); B2695 (G. microcalyx); B2744 (G. curtisii); Buwalda 6258 (G. crinitum); 7538 (G. javanicum).

Campbell EC9 (G. hirsutum); Chamchumroon 835 (G. mastersii); 1595 (G. javanicum); Chamchumroon & Puff 1131 (G. javanicum); Chand 3628 (G. strobilinum); 5984 (G. ovatum); 6005 (G. velutinum); 6105 (G. velutinum); Chandra & Mandal 1066 (G. ovatum); Chandra Bose 49190 (G. eriocarpon); Chantanamuck 23 (G. curtisii); 601 (G. strobilinum); 702 (G. strobilinum); 868 (G. javanicum); Chantaranothai et al. 90/57 (G. strobilinum); 728 (G. flexuosum); 1367 (G. javanicum); Charoenphol et al. 3824 (G. javanicum); 4871 (G. strobilinum); Chayamarit et al. 1537 (G. javanicum); 3217 (G. strobilinum); Chermsirivathana 2264 (G. strobilinum); Chevalier 29211 (G. leptodon); 29410 (G. javanicum); 32420 (G. javanicum); Chew 660 (G. microcalyx); Chew & Corner 4008 (G. javanicum); Chinese Collector 464 (G. javanicum); Ching 6851 (G. chinense); S39552 (G. javanicum); Chongko & Boonkongchart 387 (G. javanicum); Chung 169 (G. javanicum); Clarke 5950 (G. javanicum); 12498A (G. ovatum); 13674A (G. parviflorum); 13763A (G. parviflorum); 16003 (G. parviflorum); 16502 (G. nutans); 17935A (G. melissifolium); 17935B (G. melissifolium); 18337 (G. javanicum); 19090 (G. parviflorum); 36174B (G. parviflorum); 40645A (G. nutans); 40763A (G. javanicum); 40938A (G. strobilinum); 40961 (G. strobilinum); 41722 (G. strobilinum); 42012 (G. strobilinum); 42921A & B (G. parviflorum); 42936 (G. javanicum); 43730 (G. javanicum); 44358D, E & H (G. parviflorum); Clason 55B (G. javanicum); Clason-Laarman 97 (G. javanicum); Clemens & S. Clemens 3885 (G. grandiflorum); 3695 (G. javanicum); 20175 (G. curtisii); 26116 (G. javanicum); 30655 (G. javanicum); 31502 (G. javanicum); 34145 (G. javanicum); 35013 (G. javanicum); Coert 476 (G. javanicum); 1547 (G. javanicum); Collett 887 (G. hemsleyanum); 961 (G. strobilinum); Collins 990 (G. javanicum); 1038 (G. strobilinum); 1220 (G. strobilinum); 1483 (G. javanicum); Comber 140 (G. javanicum); Congdon 1054 (G. javanicum); Conklin & Buwaya 72663 (G. javanicum); 79601 (G. javanicum); Connie et al. 3705 (G. microcalyx); Cook et al. 180 (G. heyneanum); Corner 31627 (G. javanicum); 33178 (G. curtisii); 33212 (G. javanicum); Cowan 855 (G. parviflorum); 1341 (G. parviflorum); Cowley et al. 98 (G. javanicum); Craib 102 (G. mastersii); 125 (G. parviflorum); Cuming 1840 (G. javanicum); Cuong & Cuong 2095 (G. microdon); Curran 5014 (G. javanicum); Curtis 1310 (G. curtisii); 2166 (G. curtisii); 3191 (G. crinitum).

Danser 6156 (G. javanicum); Darnton 185 (G. hirsutum); Davis 69298 (G. crinitum); Davison 1 (G. crinitum); de Kok et al. S97828 (G. javanicum); SAN 147206 (G. microcalyx); SAN 147234 (G. hirsutum); de Monchy 19 (G. javanicum); de Silva s.n. in Wall. Cat. 2156.2 (G. javanicum); 2157 (G. mastersii); 2157 (G. melissifolium); 2158.2 [‘a’] (G. parviflorum); 2158.2 [‘b’] (G. parviflorum); de Voogd 1551 (G. javanicum); de Wilde & de Wilde-Duyfjes 12027 (G. microcalyx); 12358 (G. mastersii); 12844 (G. mastersii); 13483 (G. mastersii); 13615 (G. mastersii); 14079 (G. microcalyx); 14473 (G. mastersii); 15892 (G. dolichobotrys); 16593 (G. microcalyx); 16949 (G. microcalyx); 18018 (G. microcalyx); 18163 (G. mastersii); 18618 (G. mastersii); 19014 (G. javanicum); 19300 (G. microcalyx); 19879 (G. microcalyx); den Hoed 912 (G. crinitum); DeVore & Hoover 370 (G. javanicum); Dickason 5216 (G. strobilinum); 5223 (G. strobilinum); Dixen 70-1027 (G. strobilinum); Dransfield 2868 (G. javanicum); Du et al. HNK 2302 (G. javanicum); HNK 2821 (G. leptodon); HNK 2947 (G. leptodon); HNK 3193 (G. grandiflorum); HNK 3342 (G. grandiflorum).

Edaño 6671 (G. javanicum); 7145 (G. javanicum); 11894 (G. javanicum); 14077 (G. javanicum); 15389 (G. javanicum); 15440 (G. javanicum); 15533 (G. javanicum); 17836 (G. javanicum); 18024 (G. javanicum); 48787 (G. javanicum); 76369 (G. javanicum); 78203 (G. javanicum); Edaño & Gutierrez 37837 (G. javanicum); Elbert 143 (G. javanicum); Elmer 6534 (G. javanicum); 10391 (G. javanicum); 11146 (G. javanicum); 13469 (G. javanicum); 15290 (G. javanicum); 17836 (G. javanicum); 22371 (G. javanicum); Endert 2891 (G. javanicum).

Faber 1 (G. chinense); 22275 (G. chinense); Fenix 15742 (G. javanicum); 29945 (G. javanicum); Forbes 1579 (G. microcalyx); 1910 (G. javanicum); Ford 668 (G. chinense); Forrest 9442 (G. pedunculatum); 18427 (G. pedunculatum); 27176 (G. pedunculatum); Fox 4814 (G. javanicum); 5025 (G. javanicum); 12602 (G. javanicum); Franck 1346 (G. curtisii); Fukuoka T-62322 (G. parviflorum); Fukuoka & Ito T-35328 (G. javanicum); Fukuoka & Na Nakhon T-36105 (G. strobilinum); Fung 20110 (G. javanicum).

Gadoh 1688 (G. curtisii); Gallatly 613 (G. parviflorum); Gamble 3818A (G. parviflorum); 9724 (G. parviflorum); 14628 (G. heyneanum); 17983 (G. heyneanum); Gardner & Khumchompoo ST2117 (G. javanicum); Gardner et al. 80 (G. javanicum); 137 (G. javanicum); ST 1972 (G. crinitum); Garrett 66 (G. strobilinum); 409 (G. javanicum); 435 (G. strobilinum); 997 (G. parviflorum); 1219 (G. strobilinum); 1367 (G. strobilinum); 1423 (G. parviflorum); 1464 (G. strobilinum); Gaudichaud 127 (G. hainanense); Geesink et al. 6870 (G. javanicum); 7167 (G. crinitum); Gibot SAN 87949 (G. curtisii); Gibot & Sundaling SAN 94250 (G. curtisii); Glamwaewwong 4 (G. pedunculatum); 55 (G. strobilinum); 124 (G. parviflorum); Gomez s.n. in Wall. Cat. 2159.2 (G. crinitum); Gressitt 1062 (G. javanicum); Griffith 185 (G. parviflorum); 290 (G. javanicum); 339 (G. parviflorum); 1014 (G. javanicum); 1015 (G. niveum); 1850 (G. javanicum); 4038 (G. javanicum); 4039 (G. melissifolium); 4040 (G. parviflorum); 4041 (G. javanicum); 4042 (G. crinitum).

Haines 516 (G. parviflorum); Hallier D280 (G. javanicum); 392 (G. javanicum); 478 (G. javanicum); Hambananda 378 (G. javanicum); Hamilton s.n. in Wall. Cat. 2158D [‘a’] (G. parviflorum); 2158D [‘b’] (G. parviflorum); Haniff 10337 (G. curtisii); Haniff & Nur 3867 (G. mastersii); 7138 (G. javanicum); Hansen & Smitinand 12053 (G. crinitum); 12142 (G. mastersii); Hansen et al. 10773 (G. strobilinum); Hardial & Noor 109 (G. javanicum); Helfer 4041 (G. javanicum); 4043 (G. javanicum); Henderson 10058 (G. crinitum); 17972 (G. javanicum); 20180 (G. javanicum); 21770 (G. crinitum); 22234 (G. javanicum); 23773 (G. crinitum); 25061 (G. crinitum); 29618 (G. javanicum); 29658 (G. javanicum); 32572 (G. curtisii); Hennipman 3183 (G. strobilinum); 3911 (G. javanicum); Henry 153 (G. javanicum); 1565 (G. chinense); 11046 (G. javanicum); 12011 (G. javanicum); 12011A (G. javanicum); 12253 (G. parviflorum); 12253A (G. parviflorum); 12307 (G. javanicum); 12314 (G. javanicum); 12501 (G. microdon); 12518 (G. pedunculatum); 12518A (G. pedunculatum); 12609 (G. javanicum); 13594 (G. pedunculatum); Herb Trip 282 (G. strobilinum); Herb. Heyne in Wall. Cat. 2158.1 (G. parviflorum); Herb. Hongkong 464 (G. javanicum); Herb. Madras [‘Stachys indica’] in Wall. Cat. 2152C (G. heyneanum); Herb. Madras in Wall. Cat. 2710A (G. eriocarpon); Hochreutiner 144 (G. javanicum); 1065 (G. javanicum); 1396 (G. javanicum);1714 (G. javanicum); 1900 (G. javanicum); 2346 (G. javanicum); Holstvoogd 508 (G. javanicum); Holttum 9836 (G. crinitum); 20531 (G. crinitum); Hoogerwerf 360 (G. javanicum); Hooker 71 (G. ovatum); Hooker & Thomson 48 (G. velutinum); 488 (G. parviflorum); 636 (G. velutinum); 976 (G. mastersii); 1814 (G. mastersii); 2062 (G. parviflorum); 2063 (G. thomsonii); 2078 (G. javanicum); 2444 (G. pedunculatum); Hoover & Wiriadinata 31026 (G. javanicum); Hoover et al. 5512 (G. crinitum); Horsfield 38 (G. javanicum); 39 (G. javanicum); Hosokawa 5266 (G. chinense); Hosseus 62 (G. strobilinum); 63 (G. strobilinum); 280 (G. strobilinum); How 71890 (G. javanicum); 72022 (G. javanicum); 73296 (G. chinense); 73517 (G. hainanense); Huang 1075 (G. chinense); 4889 (G. chinense); 4896 (G. chinense); 4943 (G. chinense); Hume 7965 (G. javanicum); 8418 (G. javanicum); 9263 (G. crinitum).

Intawongsa 4 (G. javanicum); Ismail 99812 (G. crinitum); Iwatsuki et al. P-181 (G. javanicum); P-344 (G. javanicum); P-607 (G. javanicum); S-470 (G. javanicum); S-755 (G. microcalyx); T-8461 (G. javanicum).

Jaheri 1895 (G. mastersii); Jarrett et al. 650 (G. heyneanum); Jatupol 08-176 (G. strobilinum); 08-339 (G. strobilinum); 08-472 (G. strobilinum); Jenkins 340 (G. mastersii); Julia et al. SAN 147037 (G. microcalyx).

K.K. 1094 (G. strobilinum); Karim & Shah 31 (G. curtisii); Kato et al. B-8173 (G. javanicum); Keenan et al. 523 (G. crinitum); 863 (G. crinitum); 898 (G. strobilinum); 923 (G. crinitum); 949 (G. crinitum); 950 (G. javanicum); 1036 (G. strobilinum); 1098 (G. crinitum); 1105 (G. strobilinum); 1178 (G. javanicum); 1229 (G. strobilinum); 1244 (G. strobilinum); 1602 (G. crinitum); 1605 (G. javanicum); 1737 (G. javanicum); Kerr 733 (G. javanicum); 835 (G. strobilinum); 835A (G. strobilinum); 1380 (G. strobilinum); 1541 (G. strobilinum); 1541B (G. strobilinum); 2054 (G. javanicum); 4595 (G. strobilinum); 6333 (G. strobilinum); 6347 (G. javanicum); 7119 (G. crinitum); 7173 (G. javanicum); 7346 (G. curtisii); 7350 (G. mastersii); 7480 (G. crinitum); 10016 (G. javanicum); 10829 (G. javanicum); 10832 (G. crinitum); 13115 (G. javanicum); 13189 (G. mastersii); 15418 (G. javanicum); 15424 (G. crinitum); 15760 (G. mastersii); 15845 (G. javanicum); 16310 (G. crinitum); 18070 (G. javanicum); 18804 (G. crinitum); 20610 (G. strobilinum); 21078 (G. javanicum); 21614 (G. crinitum); 21645 (G. javanicum); 21706 (G. javanicum); Kessler 89 (G. javanicum); Khan 214 (G. strobilinum); Kiah 24251 (G. javanicum); 24261 (G. crinitum); Kiew 807 (G. crinitum); 1629 (G. crinitum); 1866 (G. microcalyx); 3317 (G. javanicum); 3329 (G. curtisii); 4599 (G. javanicum); 5108 (G. curtisii); Kingdon-Ward 22543 (G. strobilinum); 22561 (G. strobilinum); 22716 (G. strobilinum); Kjeldsen et al. 34 (G. curtisii); KK 398 (G. parviflorum); 400 (G. javanicum); 416 (G. pedunculatum); 456 (G. strobilinum); Kloss 14803 (G. microcalyx); Koelz 23029 (G. parviflorum); 23191 (G. ovatum); 24266 (G. nutans); 25506 (G. parviflorum); 29855 (G. javanicum); 32535 (G. strobilinum); Koizumi & Lawe 689 (G. javanicum); Kolur 7231 (G. javanicum); Konta et al. 4636 (G. strobilinum); T-29549 (G. javanicum); Koorders 14853β (G. javanicum); 15012β (G. javanicum); 20938β (G. javanicum); 22937β (G. javanicum); 23258β (G. javanicum); 23274β (G. javanicum); 27205β (G. javanicum); 29370β (G. javanicum); 29833β (G. javanicum); 29856β (G. javanicum); 30015β (G. javanicum); 31959β (G. javanicum); 32022β (G. javanicum); 36049β (G. javanicum); 36057β (G. javanicum); 38195β (G. javanicum); 40500β (G. javanicum); Kostermans 809 (G. javanicum); 6824 (G. javanicum); 23940 (G. javanicum); Koyama & Nagamasu T-40151 (G. strobilinum); Koyama & Phengklai T-39128 (G. strobilinum); T-39140 (G. strobilinum); T-39599 (G. strobilinum); Koyama et al. T-30079 (G. strobilinum); T-32680 (G. strobilinum); T-39537 (G. javanicum); T-39620 (G. strobilinum); T-39944 (G. strobilinum); T-61044 (G. javanicum); T-62057 (G. strobilinum); Krispinus SAN 113840 (G. javanicum); SAN 113847 (G. curtisii); SAN 128284 (G. javanicum); Kunstler 454 (G. javanicum); 2155 (G. microcalyx); 10445 (G. microcalyx); Kuntze 5615 (G. javanicum).

Lace 4245 (G. strobilinum); 4411 (G. nutans); 5328 (G. parviflorum); 5571 (G. strobilinum); 5860 (G. parviflorum); 6322 (G. strobilinum); Lalramnghinglova 8049 (G. mastersii); Lam 83 (G. javanicum); Lanjouw 31 (G. javanicum); Larsen 8718 (G. strobilinum); 8719 (G. sp. aff. flexuosum); 8953 (G. strobilinum); 8973 (G. sp. aff. flexuosum); Larsen & S. Larsen 34166 (G. strobilinum); Larsen et al. 160 (G. javanicum); 475 (G. javanicum); 1098 (G. parviflorum); 2636 (G. javanicum); 3181 (G. javanicum); 30654 (G. crinitum); 30688 (G. mastersii); 30855 (G. javanicum); 30948 (G. crinitum); 34166 (G. strobilinum); 40899 (G. javanicum); 41307 (G. javanicum); 41512 (G. mastersii); 43200 (G. crinitum); 43393 (G. javanicum); 44041 (G. mastersii); 44788 (G. strobilinum); 44839 (G. pedunculatum); 45733 (G. curtisii); 46162 (G. pedunculatum); 46335 (G. parviflorum); 46633 (G. strobilinum); 46688 (G. strobilinum); 46729 (G. javanicum); 46873 (G. parviflorum); Lau 141 (G. chinense); 4095 (G. chinense); 27392 (G. javanicum); 28123 (G. javanicum); Laux 12844 (G. heyneanum); Lawson 89 (G. eriocarpon); Lecomte & Finet 1320 (G. grandiflorum); Leschenault 259 (G. javanicum); Li 333 (G. pedunculatum); 950 (G. javanicum); 1491 (G. parviflorum); 2256 (G. javanicum); 2886 (G. pedunculatum); Liang 62115 (G. javanicum); 68240 (G. chinense); 69878 (G. javanicum); Lim et al. FRI 52830 (G. crinitum); FRI 56461 (G. crinitum); SAN 143266 (G. javanicum); Lister 118 (G. parviflorum); Loc et al. 10510 (G. leptodon); Loh 13368 (G. crinitum); Loher 4215 (G. javanicum); 6594 (G. javanicum); Lorzing 741 (G. javanicum); 832 (G. javanicum); 2239 (G. javanicum); 4616 (G. javanicum); 4670 (G. curtisii); 4780 (G. javanicum); 5498 (G. curtisii); 5769 (G. microcalyx); 5803 (G. javanicum); 5949 (G. javanicum); 6178 (G. javanicum); 14079 (G. curtisii); 14909 (G. javanicum); 14918 (G. javanicum).

Mabesa 25009 (G. javanicum); MacGregor 594 (G. javanicum); 754 (G. strobilinum); 755 (G. strobilinum); Madani & Ismail SAN 108657 (G. curtisii); Madani SAN 92027 (G. javanicum); Madulid et al. 3103 (G. javanicum); Maknoi 1036 (G. javanicum); Mantor SAN 136707 (G. javanicum); Marcan 1558 (G. strobilinum); Masters 340 (G. mastersii); Maxwell 71-669 (G. strobilinum); 72-322 (G. javanicum); 73-422 (G. javanicum); 73-755 (G. flexuosum); 74-784 (G. javanicum); 75-759 (G. javanicum); 75-785 (G. mastersii); 75-936 (G. javanicum); 75-1025 (G. strobilinum); 85-662 (G. mastersii); 86-37 (G. javanicum); 87-726 (G. javanicum); 87-1305 (G. strobilinum); 87-1344 (G. strobilinum); 87-1465 (G. javanicum); 89-44 (G. parviflorum); 89-1389 (G. strobilinum); 89-1435 (G. strobilinum); 90-1258 (G. microdon); 91-737 (G. javanicum); 92-590 (G. strobilinum); 92-724 (G. strobilinum); 92-741 (G. strobilinum); 93-626 (G. crinitum); 93-979 (G. strobilinum); 93-1502 (G. strobilinum); 95-850 (G. parviflorum); 95-996 (G. strobilinum); 95-1112 (G. strobilinum); 96-34 (G. strobilinum); 96-982 (G. javanicum); 96-1022 (G. javanicum); 96-1435 (G. strobilinum); 96-1684 (G. strobilinum); 97-978 (G. strobilinum); 97-982 (G. javanicum); 97-1028 (G. strobilinum); 97-1167 (G. javanicum); 97-1226 (G. strobilinum); 97-1315 (G. strobilinum); 97-1368 (G. strobilinum); 98-1427 (G. strobilinum); 05-426 (G. javanicum); 05-594 (G. pedunculatum); 05-734 (G. strobilinum); 06-011 (G. javanicum); 06-638 (G. pedunculatum); 07-667 (G. strobilinum); 09-259 (G. parviflorum); McKee 5976 (G. strobilinum); Meebold 5814 (G. parviflorum); 7187 (G. nutans); 7405 (G. nutans); 7598 (G. nutans); 8104 (G. strobilinum); 12941 (G. keralense); 12944 (G. keralense); 13015 (G. keralense); Meijer 4504 (G. curtisii); 4594 (G. curtisii); Mell 941 (G. chinense); Mendoza PNH 18370 (G. javanicum); Mendoza & Steiner PNH 41555 (G. javanicum); Middleton & Pholsena 185 (G. javanicum); Middleton & Triboun 4851 (G. strobilinum); Middleton et al. 226 (G. javanicum); 362 (G. crinitum); 446 (G. mastersii); 514 (G. javanicum); 1075 (G. crinitum); 1437 (G. javanicum); 1507 (G. javanicum); 3000 (G. crinitum); 3323 (G. crinitum); Mitsuta T-50214 (G. strobilinum); Mitsuta et al. T-46427 (G. strobilinum); T-46480 (G. strobilinum); Mogea & de Wilde 3655 (G. microcalyx); Morse 257 (G. leptodon); Morsi 253 (G. strobilinum); Moulton 47 (G. javanicum); 6701 (G. javanicum); Moysey & Kiah 33705 (G. crinitum); Mukerjee 3354A (G. strobilinum); Murata et al. B-3898 (G. javanicum); B-3902 (G. javanicum); B-3914 (G. javanicum); B-4016 (G. javanicum); B-4040 (G. microcalyx); B-4041 (G. javanicum); J-853 (G. javanicum); T-15084 (G. parviflorum); T-15128 (G. parviflorum); T-15372 (G. strobilinum); T-15513 (G. strobilinum); T-15514 (G. strobilinum); T-15536 (G. strobilinum); T-15988 (G. javanicum); T-16087 (G. javanicum); T-37345 (G. flexuosum); T-38331 (G. strobilinum); T-38714 (G. strobilinum); T-42115 (G. strobilinum); T-43064 (G. strobilinum); T-43088 (G. strobilinum); T-49727 (G. javanicum); T-50374 (G. strobilinum); T-51136 (G. javanicum); T-51530 (G. strobilinum); T-51688 (G. javanicum).

Na Nakorn 7827 (G. strobilinum); Na Nakorn et al. 3855 (G. javanicum); 4020 (G. parviflorum); 7769 (G. strobilinum); 9473 (G. javanicum); Na Songkhla et al. 349 (G. strobilinum); 417 (G. strobilinum); Nagamasu 3796 (G. javanicum); Nauen 38018 (G. javanicum); 38046 (G. crinitum); Neamsuvan 36 (G. crinitum); Newman et al. 1104 (G. crinitum); Nguyen 28 (G. javanicum); Nguyen et al. HNK 1256 (G. leptodon); HNK 1672 (G. leptodon); HNK 1813 (G. leptodon); Nieawenhuis & Jaheri 1391 (G. microcalyx); Nielsen et al. 1714 (G. javanicum); 1862 (G. javanicum); Niyomdham 3116 (G. javanicum); Niyomdham & Phudjaa 3132 (G. javanicum); Nooteboom 687 (G. strobilinum); 899 (G. javanicum); Nooteboom et al. 4557 (G. microcalyx); Norsaengsri & Tathana 7371 (G. crinitum); Nur 11100 (G. curtisii); 18576 (G. curtisii); 32839 (G. curtisii); Nur & Forworthy 11922 (G. crinitum).

Ocampo 27975 (G. javanicum); Olsen 299 (G. javanicum).

Paisooksantivatana & Sangkhachand 378-80 (G. strobilinum); 2185-87 (G. strobilinum); Palee 10 (G. strobilinum); 132 (G. javanicum); 137 (G. strobilinum); 420 (G. strobilinum); 787 (G. strobilinum); 1068 (G. strobilinum); Panatkool 132 (G. javanicum); Panigrahi 4877 (G. parviflorum); 16570 (G. mastersii); Parinya & Santi 302 (G. parviflorum); Parish 78 (G. strobilinum); Parker 2255 (G. crinitum); Parkinson 1595 (G. javanicum); 3460 (G. parviflorum); Parnell et al. 95-156 (G. parviflorum); Parry 58 (G. mastersii); 283 (G. parviflorum); 925 (G. parviflorum); Pereira et al. 432 (G. javanicum); Petelot 868 (G. leptodon); 5002 (G. leptodon); 5115 (G. chinense); 5117 (G. javanicum); 5290 (G. javanicum); 5802 (G. chinense); Petrmitr 42 (G. javanicum); 174 (G. strobilinum); 182 (G. javanicum); 227 (G. strobilinum); 323 (G. strobilinum); 327 (G. strobilinum); 461 (G. strobilinum); 667 (G. curtisii); Phengklai 1307 (G. crinitum); Phengklai & Tamura 10172 (G. strobilinum); Phengklai et al. 3915 (G. javanicum); 3899 (G. strobilinum); 3900 (G. strobilinum); 6650 (G. strobilinum); 6806 (G. strobilinum); 6816 (G. javanicum); 6868 (G. strobilinum); 6926 (G. javanicum); 7128 (G. javanicum); 7178 (G. javanicum); 7296 (G. javanicum); 7505 (G. javanicum); 12774 (G. javanicum); 12796 (G. strobilinum); 14058 (G. strobilinum); Phengnaren 4 (G. javanicum); Phuakam 65 (G. strobilinum); 87 (G. strobilinum); Phuong 29 (G. microdon); 142 (G. leptodon); 699 (G. javanicum); 1761 (G. javanicum); 2460 (G. leptodon); 3519 (G. microdon); 5795 (G. leptodon); Phusomsaeng & Pinnin 317 (G. mastersii); Pierre 1181 (G. javanicum); Pius & Dauni SAN 143153 (G. javanicum); Poilane 6931 (G. javanicum); 8069 (G. hainanense); 8618 (G. heyneanum); 8802 (G. heyneanum); 12381 (G. heyneanum); 12804 (G. javanicum); 15718 (G. javanicum); 16692 (G. hainanense); 21050 (G. flexuosum); 31881 (G. grandiflorum); Pongamornkul 11 (G. strobilinum); 267 (G. javanicum); 336 (G. strobilinum); Pooma 1596 (G. javanicum); 5783 (G. strobilinum); Pooma et al. 4447 (G. crinitum); Posthumus 943 (G. microcalyx); Prain's collector 141 (G. parviflorum); 390 (G. parviflorum); Praphat 373 (G. javanicum); Prazer 374 (G. nutans); 419 (G. strobilinum); Put 294 (G. strobilinum); 385 (G. strobilinum); 427 (G. pedunculatum); 442 (G. javanicum); 1136 (G. flexuosum); 1632 (G. javanicum); 1899 (G. strobilinum); 2425 (G. strobilinum); 3272 (G. strobilinum); 4394 (G. strobilinum); 4431 (G. strobilinum).

Ramos 793 (G. javanicum); 5714 (G. javanicum); 14673 (G. javanicum); 15187 (G. javanicum); 23481 (G. javanicum); 24358 (G. javanicum); 24403 (G. javanicum); 30280 (G. javanicum); 33009 (G. javanicum); 41251 (G. javanicum); 41489 (G. javanicum); 43199 (G. javanicum); Ramos & Edaño 29402 (G. javanicum); 30961 (G. javanicum); 34127 (G. javanicum); 37494 (G. javanicum); 37617 (G. javanicum); 38749 (G. javanicum); 40340 (G. javanicum); 75504 (G. javanicum); 84990 (G. javanicum); 85050 (G. javanicum); Rantai et al. S93323 (G. microcalyx); Reillo 15411 (G. javanicum); Reynoso et al. 11232 (G. javanicum); Richard 61 (G. javanicum); Richards 2403 (G. javanicum); Ridley 2148 (G. javanicum); 2149 (G. crinitum); 2185 (G. crinitum); 7600 (G. crinitum); 8221 (G. javanicum); 8564 (G. javanicum); 11130 (G. javanicum); 11131 (G. crinitum); 12602 (G. javanicum); 13595 (G. curtisii); 13698 (G. javanicum); 14061 (G. javanicum); 14446 (G. curtisii); Robertson 431 (G. strobilinum); Robinson 6364 (G. javanicum); 18189 (G. javanicum); Robinson & Kloss 187 (G. javanicum); 193 (G. javanicum); 6144 (G. javanicum); Rochadi 645 (G. mastersii); Rogstad 585 (G. javanicum); Rumutom 122 (G. hirsutum); 393 (G. curtisii).

S.P. 39 (G. mastersii); Sadakorn 620 (G. strobilinum); 625 (G. strobilinum); 626 (G. strobilinum); Sam HVS 065 (G. leptodon); Sands et al. 4049 (G. curtisii); 5822 (G. javanicum); Sangkhachand 447 (G. curtisii); 591 (G. strobilinum); 751 (G. crinitum); 911 (G. javanicum); 1032 (G. strobilinum); 1460 (G. javanicum); 1658 (G. javanicum); 2160 (G. mastersii); Sangkhachand & Nimanong 1363 (G. curtisii); Sankamethawee 285 (G. strobilinum); 287 (G. strobilinum); Santisuk 1171 (G. javanicum); 6860 (G. strobilinum); Santisuk & Nimanong 377 (G. crinitum); Santisuk et al. 890 (G. parviflorum); Sapiin 49 (G. javanicum); 285 (G. javanicum); 420 (G. javanicum); Sasirat 49 (G. javanicum); Saw FRI 39974 (G. curtisii); FRI 44282 (G. crinitum); FRI 44870 (G. curtisii); Scortechini 5 (G. crinitum); 924 (G. curtisii); 928 (G. crinitum); 942 (G. microcalyx); 1225 (G. javanicum); Seimund 921 (G. javanicum); Serm 99 (G. strobilinum); Shah 166 (G. javanicum); 1599 (G. javanicum); 2772 (G. javanicum); Shah & Noor 1965 (G. crinitum); Shah & Sidek 1135 (G. microcalyx); Shimizu & Hutoh T-10154 (G. parviflorum); Shimizu & Nalampoon T-7801 (G. javanicum); Shimizu et al. T-12355 (G. strobilinum); T-18591 (G. strobilinum); T-19131 (G. strobilinum); T-19136 (G. strobilinum); T-21197 (G. parviflorum); T-22163 (G. strobilinum); T-22394 (G. strobilinum); T-22588 (G. strobilinum); T-23429 (G. javanicum); T-23517 (G. strobilinum); T-27663 (G. javanicum); T-28921 (G. crinitum); T-29076 (G. javanicum); Shui & Chen 13118 (G. parviflorum); 13241 (G. parviflorum); 13283 (G. pedunculatum); 40883 (G. javanicum); Shui et al. 13524 (G. parviflorum); 13555 (G. parviflorum); Sidajium 31 (G. javanicum); Sidisunthorn ST 2144 (G. javanicum); Simidu 3655 (G. chinense); Simons 46 (G. parviflorum); Sinclair 9849 (G. crinitum); Sinclair & Edaño 9639 (G. javanicum); Sino-American Expedition to Yunnan 191 (G. microdon); Sitthikul et al. 1 (G. javanicum); Skornickova et al. 170 (G. microcalyx); Smith 416 (G. javanicum); Smitinand 652 (G. javanicum); 742 (G. crinitum); 3445 (G. javanicum); 4896 (G. javanicum); 4918 (G. strobilinum); 7038 (G. javanicum); 7069 (G. strobilinum); 8449 (G. javanicum); Smitinand & Anderson 7260 (G. strobilinum); 7339 (G. strobilinum); Smitinand & Sleumer 1127 (G. strobilinum); 1323 (G. strobilinum); Smitinand & Suvarnakoset 562 (G. javanicum); Smitinand et al. 7857 (G. strobilinum); Snan 294 (G. crinitum); 399 (G. javanicum); 616 (G. javanicum); Soejarto et al. 10360 (G. leptodon); Somkid 171 (G. microdon); Soradej 85 (G. javanicum); 148 (G. strobilinum); 283 (G. strobilinum); 319 (G. strobilinum); Sørensen et al. 881 (G. strobilinum); 1032 (G. strobilinum); 1036 (G. strobilinum); 1053 (G. strobilinum); 1295 (G. parviflorum); 1560 (G. strobilinum); 3787 (G. javanicum); 3916 (G. javanicum); 4209 (G. parviflorum); 4874 (G. strobilinum); 5408 (G. strobilinum); 5469 (G. strobilinum); 5910 (G. strobilinum); 5945 (G. strobilinum); 6563 (G. strobilinum); SP et al. 22 (G. javanicum); Spire 1353 (G. javanicum); Srisanga 1698 (G. parviflorum); 1924 (G. parviflorum); 1953 (G. javanicum); 1974 (G. javanicum); 2288 (G. strobilinum); Srisanga et al. 55 (G. javanicum); 759 (G. javanicum); Stevens et al. 616 (G. javanicum); Steward & Cheo 804 (G. chinense); Stone 6590 (G. javanicum); 13571 (G. javanicum); Stone et al. 6266 (G. javanicum); 6450 (G. javanicum); Strugnell 45854 (G. crinitum); Suddee & Puujaa 1117 (G. strobilinum); Suddee et al. 889 (G. strobilinum); 913 (G. strobilinum); 955 (G. flexuosum); 1014 (G. flexuosum); Suksathan 1730 (G. strobilinum); 2825 (G. strobilinum); Sulit 2657 (G. javanicum); Sundaling SAN 83957 (G. javanicum); SAN 84036 (G. curtisii); Suresh 22326 (G. heyneanum); Sutheesorn 1642 (G. strobilinum); 2241 (G. strobilinum); 2645 (G. javanicum); 3091 (G. strobilinum); 3158 sht. 1 (G. strobilinum); 3158 sht. 2 (G. strobilinum); Sutheesorn & Sangkhachand 3134 (G. strobilinum); Suvarnakoset 151 (G. crinitum); 379 (G. javanicum); 830 (G. javanicum); 952 (G. strobilinum); 988 (G. strobilinum); 1692 (G. curtisii); 2171 (G. mastersii); 2200 (G. javanicum); Suzana et al. SAN 147801 (G. microcalyx).

Tadong 524 (G. hirsutum); Tagawa et al. T-354 (G. strobilinum); T-2295 (G. strobilinum); T-5319 (G. crinitum); T-5335 (G. javanicum); T-9132 (G. strobilinum); T-9288 (G. strobilinum); T-9498 (G. strobilinum); T-9788 (G. javanicum); T-9789 (G. javanicum); T-9922 (G. parviflorum); T-17513 (G. strobilinum); Tamura T-60312 (G. javanicum); Tanaka et al. 8219 (G. strobilinum); Tao & Li 39472 (G. strobilinum); Tay et al. 91-35 (G. crinitum); Teruya 1406 (G. curtisii); Teysmann 13722 (G. javanicum); Thaithong 964 (G. strobilinum); Thaithong et al. 257 (G. javanicum); Thaworn 688 (G. javanicum); Thomson 119 (G. eriocarpon); Thongisan 36 (G. strobilinum); Thorel 2441 (G. flexuosum); 3362 (G. microdon); Tiptabiankarn 1248 (G. strobilinum); 1282 (G. javanicum); 1372 (G. parviflorum); Tsai 61603 (G. sulcatum); Tsang 109 (G. javanicum); 400 (G. javanicum); 21458 (G. chinense); 26825 (G. javanicum); 27020 (G. javanicum); 29398 (G. javanicum); 30705 (G. hainanense); Tsang & Fung 247 (G. javanicum); 465 G. javanicum); 559 (G. chinense); Tsugaru T-61811 (G. javanicum); Tuyuk et al. SAN 113666 (G. javanicum). Uddin & Ullah 3558 (G. crinitum); UNESCO Limestone Expedition 355 (G. javanicum).

Vajravelu 48896 (G. heyneanum); van Balgooy 3032 (G. javanicum); 4535 (G. javanicum); van Beusekom & Charoenpol 1992 (G. flexuosum); van Beusekom & Geesink 3352 (G. javanicum); van Beusekom et al. 3480 (G. strobilinum); 3501 (G. strobilinum); 3856 (G. strobilinum); van de Bult 83 (G. javanicum); 85 (G. javanicum); 139 (G. strobilinum); 478 (G. strobilinum); 697 (G. javanicum); 701 (G. strobilinum); 989 (G. javanicum); van de Pijl 673 (G. javanicum); van den Brink 2189 (G. javanicum); 2469 (G. javanicum); 2968 (G. javanicum); 3317 (G. javanicum); 4936 (G. javanicum); van Steenis 950 (G. javanicum); 8684 (G. dolichobotrys); 8685 (G. javanicum); 9483 (G. javanicum); Vanoverbergh 701 (G. javanicum); Veldkamp 8027 (G. microcalyx); 8372 (G. microcalyx); Vial-Debas 10 (G. strobilinum); 56 (G. javanicum); Vidal 690A (G. grandiflorum); 1044B (G. javanicum); 3470 (G. javanicum); 5118 (G. flexuosum); 5177 (G. parviflorum); 5772 (G. javanicum); Vivekananthan 48614 (G. heyneanum); 48658 (G. keralense).

W.F. 173 (G. curtisii); Wall. Cat. 2151.1 (G. strobilinum); 2151.2 (G. strobilinum); 2153 (G. strobilinum); 2154 (G. javanicum); 2155 (G. ovatum); 2156.1 (G. javanicum); 2158 (G. parviflorum); 2158C (G. crinitum); 2159.1 (G. crinitum); Wang 219 (G. pedunculatum); 33285 (G. javanicum); 34945 (G. javanicum); 75545 (G. microdon); 75550 (G. javanicum); 76475 (G. javanicum); 77368 (G. stellatohirsutum); 77982 (G. javanicum); 78214 (G. javanicum); 78775 (G. parviflorum); 78779 (G. pedunculatum); 79016 (G. parviflorum); 79289 (G. javanicum); 79725 (G. javanicum); 79805 (G. crinitum); 82349 (G. javanicum); 89448 (G. javanicum); Ward 7359 (G. longipetalum); 7689 (G. javanicum); 8734 (G. aborense); 8734 [in parte] (G. javanicum); Watt 6664 (G. nutans); 7396 (G. parviflorum); 10726 (G. strobilinum); Watthana 1508 (G. strobilinum); 2462 (G. parviflorum); Watthana et al. 436 (G. javanicum); 599 (G. javanicum); Whitford 975 (G. javanicum); Wiang 366 (G. chinense); Wight 631 (G. eriocarpon); 632 (G. eriocarpon); 2369 (G. eriocarpon); Wight s.n. in Wall. Cat. 2710B (G. eriocarpon); Williams 1357 (G. javanicum); Williams et al. 2080 (G. crinitum); Winit 56 (G. javanicum); 744 (G. javanicum); 1490 (G. strobilinum); 1525 (G. strobilinum); 1745 (G. javanicum); Winkler 1113 (G. microcalyx); Wirawan 178 (G. javanicum); Wiriadinata et al. 10709 (G. javanicum); Wisse 1011 (G. javanicum); Wongprasert 009-8 (G. strobilinum); 712-2 (G. strobilinum); 998-04 (G. strobilinum); Worawoat 86 (G. javanicum); Wray 835 (G. microcalyx); 1233 (G. curtisii); 3340 (G. crinitum); 3775 (G. crinitum); Wu et al. 355 (G. leptodon); 1046 (G. parviflorum); 3821 (G. pedunculatum); Wyatt-Smith 93195 (G. crinitum).

Xiang 374 (G. pedunculatum); 375 (G. stellatohirsutum).

Yates 895 (G. curtisii); 1738 (G. microcalyx); Yen & Kao 10730 (G. chinense); Yii & Othman S46181 (G. microcalyx); Youngboonkird 265 (G. javanicum); Yu 17623 (G. pedunculatum).

Zollinger 144 (G. javanicum); 1324 (G. javanicum); 2010 (G. javanicum); 2028 (G. parviflorum).