The genus Salacia (Celastraceae: Salaciodeae) in peninsular India

The genus Salacia (Celastraceae: Salaciodeae) in peninsular India is revised, wherein ten species are recognised. Eight species are endemic to the region. Salacia macrophylla and S. reticulata are excluded from the revision as occurrence of both the species is doubtful in mainland India. Salacia macrosperma exhibits great variation in leaf shape, flower number, pedicel length and fruit texture, hence the morphologically similar species S. beddomei and S. vellaniana are relegated to its synonymy. Descriptions, photographs, and pen and ink as well as colour drawings of all recognised Salacia species are provided with an artificial key, updated nomenclature, and synonymy. Distribution and IUCN Red list categories and criteria are also discussed. Seven binomials, S. fruticosa, S. macrophylla, S. macrosperma, S. oblonga, S. ovalis, Tontelea prinoides and Johnia coromadeliana are lectotypified.


INTRODUCTION
Celastraceae consists of 96 genera and about 1300 species (Mabberley 2017), distributed in tropical, subtropical and temperate parts of the world. In India, the family (including Hippocrateaceae) is represented by 15 genera and about 114 species (Ramamurthy and Naithani 2000). The classification of Celastraceae has undergone several revisions based on molecular evidence and it is now recognized with four subfamilies (Celastroideae, Stackhousioideae, Hippocrateoideae and Salacioideae). Subfamily Salacioideae consists of six genera and most of the species within this subfamily are lianas. Cheiloclinium Miers, Peritassa Miers and Tontelea Miers are confined to South America, Salacighia Loes. and Thyrosalacia Loes. are native to tropical Africa, whereas one of the largest genera, Salacia L., is distributed in both the Old and New World tropics (Simmons 2004). Salacia is perhaps best known as the source of salacinol, an antidiabetic compound (Yoshikawa et al. 1997).
Among the Indian species, Salacia f loribunda Wight, S. jenkinsii Kurz and S. khasiana  Salacia is more diverse in peninsular India than other parts of the country. Of the ten species recorded so far, eight are endemic to this part of the country: S. agasthiamalana Udayan Sasidharan & Anaz (2015), is considered conspecific to S. macrosperma. In S. macrosperma, the fruits have been described unclearly. In the protologue, fruits were given as irregularly ovate, and in illustration fruit vary from globose to obovoid to ellipsoid, smooth (Wight 1845). The original material (housed at K) has broadly ellipsoid-oblong fruit. Nevertheless, most collections of S. macrosperma from peninsular India also bears globose to oblong fruits with a smooth to sparsely tuberculate exocarp. Hence, taxonomic delimitation of S. macrosperma from its congeneric S. beddomei Gamble and S. brunoniana is often difficult. Incidentally, both the latter species were described without fruit. In the case of S. beddomei, it was found to be conspecific with S. macrosperma due to similarities in leaf shape, size, flowers. On the other hand, S. brunoniana is retained as distinct species, as it differs from S. macrosperma by having serrate leaves, and few-flowered fascicles.
The claimed occurrence of S. macrophylla and S. reticulata in peninsular India by Ramamurthy and Naithani (2000) is based on misidentifications. The former was identified based on the work of Ding Hou (1964), which was based on Dalzell's collection from Konkan. The present author had an opportunity to scrutinize Dalzell's collection of S. oblonga at K which is erroneously determined by Ding Hou as S. macrophylla. Similarly, S. reticulata Wight (1840) does not occur in India. It was based on a collection of Colonel Walker from Ceylon (Sri Lanka). In the protologue, Wight was uncertain about its distribution in India. Subsequently, Gamble (1918) incorrectly included this species based on a collection of Rao (87440 at CAL), and misinterpreted the type specimen (C.P. 2720 at CAL) of the Sri Lankan endemic S. diandra Thwaites as S. reticulata (see .

Brief taxonomic history of Salacia in India
'Courondi' was the earliest recorded vernacular name for a species of Salacia, used by Van Rheede (1683) in his Hortus Malabaricus and later identified as Salacia chinensis (Ding Hou 1964, Nicolson andSuresh 1986), although it should rather be S. fruticosa (see the note under S. fruticosa in the present contribution). The name 'Courondi' was validated as a new genus Courandi Adanson (1763), but later rejected against the earlier Linnaean Salacia (Dandy 1969, Nicolson andSuresh 1986). Roxburgh (1820) Wight, S. reticulata, S. verrucosa and S. macrosperma Wight. Bentham and Hooker (1862) in Genera Plantarum reduced Hippocrateaceae to the tribes Celastreae and Hippocrateae. Kurz (1872Kurz ( , 1875  Ding Hou (1964) revised the Malaysian Celastraceae and recorded 29 species, of which five species share a distribution with Andaman and Nicobar archipelago. With the addition of S. korthalsiana Miquel and S. tortuosa Srivastava 1993, Lakshminarasimhan andRay 1994), the Andaman and Nicobar Islands now harbours seven Salacia species: S. platyphylla, S. nicobarica, S. korthalsiana, S. chinensis, S. reticulata, S. salacioides and S. verrucosa, with the first two endemic to these islands.

Biogeography and comparative morphology of Indian Salacia
Salacia chinensis, S. macrosperma and S. oblonga occur on the plateaus, along forest fringes and in undergrowth. Salacia chinensis also occurs in dryland, and riverine forests of peninsular India. Salacia fruticosa and S. malabarica grow along forest margins and in coastal scrubland forest. Salacia gambleana occurs along forest margins and often shares the habitat with S. macrosperma. Salacia agasthiamalana and S. wayanadica are known only from the type localities. The former grows on the slopes in evergreen grassland at an altitude of 900-1100 m and is narrowly distributed in Agasthiamala Biosphere Reserve and adjoining areas, whereas the latter has a sporadic distribution along the forest margin in Wayanad District, Kerala. Similarly, S. megacarpa is known from wet evergreen forests bordering Kerala and Karnataka. Except for its type collection from Nilgiris (Tamil Nadu), S. brunoniana is known only from evergreen forests of Palode, Thiruvananthapuram, Kerala.
Salacia agasthiamalana is a shrub or small shrub, whereas S. chinensis is often recorded as small tree to scandent shrub. All the other Salacia species grows either as scandent shrubs or lianas, straggling over small trees. Branchlets are usually grey to dark brown or black, smooth to lenticellate; rusty, abruptly yellowstriated in S. oblonga (Figure 2A-C). Leaf shape ( Figure  2D) varies from obovate, oblanceolate, ovate to ellipticoblong. S. agasthiamalana has small, elliptic to obovate leaves in comparison with the rest of the species in peninsular India. S. gambleana has distinct oblanceolate leaves. The leaf margin also varies from entire to serrate, cuneate, sinuate, undulate, whereas the leaf apex varies from acute to acuminate or obtuse. The midrib and lateral nerves are usually conspicuous, rarely inconspicuous as in S. macrosperma, with the lateral nerves varying from 6 to 12 pairs. Some of the African species are characterised by resinous or latex threads, particularly when breaking of leaves and petiole (Robson et al. 1994), which is absent in peninsular Indian Salacia. The inflorescence usually consists of solitary to many flowered ramiflorous and axillary fascicles ( Figure 2E), whereas pedunculate cymes ( Figure 2F) are characteristic of S. fruticosa, S. oblonga and S. megacarpa.
The flowers ( Figure 3A-G) are moderately sized (7-15 mm in diam.), usually salver-shaped (urceolate in S. oblonga); pedicel length varies from 0.3 (S. oblonga) to 2 (S. malabarica) cm; bracts are often conspicuous and sepaloid; buds are usually globose, with calyptriform sepals, ovoid to conical in S. chinensis. The sepals are usually fused at the base, with the sepal lobes  laciniate to fimbriate in S. agasthiamalana, S. macrosperma, and S. gambleana, glabrous or marginally or apically erose in S. megacarpa, S. chinensis, S. oblonga and S. brunoniana, sparsely ciliate in S. malabarica, apically bearded in S. wayanadica, inconspicuous, undulate, or crisp, white-margined in S. fruticosa. The petals are primarily green to pale green, elliptic to oblong, often with hyaline margins, revolute, obovate to oblanceolate in S. chinensis and S. megacarpa, orbicular and orange to ochreous in S. fruticosa. Disc shape, size and colour varies with flower maturity, oblate and green to pale green in S. chinensis, otherwise conical or lenticular, entire, or faintly lobed, green with orange shades or ochreous. There are three stamens, the filaments often becoming reflexed or revolute; anthers bi-thecate, globose to ellipsoid, thecae confluent by oblique slits in S. megacarpa ( Figure 5A), longitudinal slits ( Figure 5B) in S. fruticosa and S. oblonga. Ovoid, yellow to ochrayellow anthers and thecae confluent by transverse slits ( Figure 5C) are most common in the remaining species of peninsular India; pollen simple, tricolporate; ovary immersed or embedded in the disc. Fruits (Figures 6, 7) usually a globose or ellipsoid berry, with pulpy mesoand endocarp and 1-(S. chinensis) to 8-seeded (dry indehiscent and prolate in S. megacarpa); exocarp smooth, or keeled, or verrucose, or tuberculate, thin (S. chinensis), otherwise leathery or tough; seeds ellipsoid to ovoid or oblate to obconical, usually dorsiventrally compressed, with surface smooth or reticulate.

MATERIALS AND METHODS
Multiple accessions of Salacia species were collected from different states of peninsular India. Photographs of habit, inflorescences, flowers, fruit, and seeds were taken using Nikon D7000 and Canon 700D digital cameras with different optics. Microscopic study was carried out with a Leica Stereo Microscope (DM500). Herbarium specimens were prepared based on the methods proposed by Forman and Bridson (1992). The processed and identified specimens were deposited in Naoroji Godrej Centre for Plant Research (NGCPR) Herbarium and duplicates presented to CAL and BSI. Existing herbarium specimens from the study area (peninsular India) were consulted at BSI, CAL, CALI, CMPR, FRLH, K, MH, PBL, RHT and TBGRI. Images of herbarium specimens were virtually consulted using JSTOR Global Plants (https://plants.jstor.org/) and herbaria webpages from A, BISH, BM, E, F, FR, G, GH, NY, P, U and US. The provisions of the ICN (Turland et al. 2018) was applied to update the nomenclature and for purposes of typification.
Descriptions were based on observation of morphology, study of type specimens, the protologue and herbarium specimens, whereas the descriptive terminology follows Ding Hou (1964) and Robson et al. (1994). The colour illustrations were prepared by the author from photographs, microscope images and herbarium specimens using water colour on hot press paper and processed in Adobe Photoshop 7.0. Conservation status was assigned, and threat assessment carried out for global level using IUCN Red list categories and criteria (IUCN 2019) based on population size, threats, and the estimation of the geographical range size (EOO and AOO), by using GeoCat (Moat 2007). The population trend and size data were produced from field observations, available literature, previous assessments and from information on herbarium specimen labels. Distribution maps were prepared for endemic species using DIVA GIS 7.5, with herbarium and field data being used to plot the distribution points.

General Distribution
About 200 species distributed in tropical and subtropical Africa, America and Asia, with few species in the temperate parts of Asia and America (Ding Hou 1963;Mabberley 2017). India with 21 species, ten of which are reported from peninsular India. nan S004 (holotype,MH n.v.;isotypes,K n.v.,SKC n.v.,SNCH n.v.).

Phenology
Flowers in March, fruits remain till June.

Distribution and habitat
Endemic to peninsular India, the species is known from two localities Pongalapara, Agasthiamalya, Thiruvananthapuram in Kerala and Peyaru to Panimalai, in Tirunelveli, Tamil Nadu ( Figure 18). It grows on slopes of evergreen grassland at the altitudinal range of 900-1100 m Rajasekar et al. 2018).

IUCN status
Both the known locations of Salacia agasthiamalana are adjoining, Pongalapara is a protected area (part of Agasthiamala Biosphere Reserve), where no threat has been recorded so far. No information is available for the population at Panimalai. Hence, due to insufficient data on the population size and threat the species is assessed here as Data Deficient (DD).
Type material of S. agasthiamalana at the herbaria listed in the protologue could not be traced. The specimen at MH (Udayan & Regy Yohannan 00S9), collected from the type locality, was used here to describe and illustrate the species (flower and fruits characters obtained from the protologue and Rajasekar et al. 2018).

Phenology
Flowers in January to March, fruiting remains till June.

Distribution and habitat
Endemic to the Western Ghats of Tamil Nadu and Kerala ( Figure 18). Evergreen and fringing forest.

IUCN status
Salacia brunoniana is known from Tamil Nadu (Nilgiris), and Kerala (Thiruvananthapuram). At present there is no detailed information available on species distribution and population, therefore, considered here as Data Deficient (DD).

Notes
The distribution report of S. brunoniana in Maharashtra (Dalzell and (Almeida 1990) from Amboli, Sindhudurg Maharashtra. Similarly, its claimed occurrence in Odisha (Ramamurthy & Naithani 2000), is based on it being mistaken for S. chinensis. Perhaps, they have referred to Panigrahi 23710 (CAL) labelled as S. brunoniana, which is an incomplete specimen of S. chinensis. Collections (Nazarudeen 13846, Shareef 70649: TBGRI) from Vittalapacha, Kerala well agrees with the protologue of S. brunoniana except in the number of flowers. S. brunoniana planted at JNTBGRI was originally brought from Vittalapacha, Kerala, which has distinctly sinuate, crenate, or serrate leaves, 2-8-flowered fascicle and broadly oblong-obovate petals. The species shares similarities in habit and flowering characters with S. macrosperma and S. gambleana, but these differs from S. brunoniana by characteristic laciniate, or fringed sepals and entire to serrate leaves.

Phenology
Flowers from January, fruits may last till June.

Distribution and habitat
Peninsular India, also in Andaman and Nicobar Islands, Assam, Odisha, and West Bengal; Sri Lanka and throughout southeast Asia. In open dryland forest, along the coastal lowland forest, as a small tree on open plateaus, in riverine forest and dense deciduous woodland.

IUCN status
As the species has a wide distribution range from Indo-Malaya to China, it is classified here as Least Concern (LC). Rare occurrence of S. chinensis in semievergreen forests of the Western Ghats (Southern India and Sri Lanka) mentioned by Kamat et al. (2020) is an error; its status as Endangered by Majid et al. (2016) is also an error. However, some individuals in the field (Shimoga, Amba, and Amboli) were observed with cut marks which may indicate its use for medicinal purposes among the local inhabitants. Salacia chinensis is the preferred species for the extraction of salacinol, an important antidiabetic compound. Often, it is found growing in areas of human habitation resulting on it being impacted by human activities such as road clearing, road widening, and overharvesting of its edible fruits. Often well grown plants have been uprooted from the wild, mainly for the roots as a source of salacinol (Patwardhan et al. 2014), which is non-renewable. However, considering its wide distribution, these threats do not yet pose a significant risk to the sustenance of the species in the wild. A detailed ecological assessment is required, throughout its range of distribution, to understand the population, regeneration, and threat.

Notes
The scandent habit of S. chinensis can often be seen along the riverbanks and the erect shrubby habit is abundant on the plateaus in Maharashtra, Karnataka, Kerala, and Tamil Nadu. It can be recognised by the few-flowered axillary and ramiflorous fascicles, smaller globose, 1-seeded fruit with thin exocarp and hyaline, pulpy endocarp.
Collections from the Andaman and Nicobar Islands are more likely to be of S. platyphylla, as the latter differs from S. chinensis in its broader leaves and 2-seeded berries. Salacia chinensis is similar to S. megacarpa in having obovate to oblanceolate petals, but the latter is clearly distinguished by its branched cymes and ellipsoid to prolate, ridged, dry fruits.

Phenology
Flowers in January, fruits may remain till end of August, rarely till November.

Distribution and habitat
Endemic to the Western Ghats of Karnataka, Kerala, and Tamil Nadu (Figure 18). It prefers semi-evergreen to evergreen fringing forest, also known from coastal scrubland.

Notes
Salacia fruticosa can be recognised in the field by its scandent habit, ovate-elliptic, crenate leaves, acuteacuminate apex, pedunculate dichotomously branched cymes, orbicular petals, and smooth, orange fruits. It was first described as Tonsella fruticosa in Wallich Catalogue (no. 4223), based on Heyne's collection and later validated by Lawson citing additional collections by Rottler, Hoenacker, and Kurz. The collection by Heyne here designated as lectotype was cited in Wallich Cat. 4223 and agrees with the protologue.

Phenology
Flowers between January to April, fruits may remain till July.

Distribution and habitat
Endemic to the Western Ghats of Karnataka and Maharashtra (Figure 18). Evergreen forest, fringing forest, at 550 m a.s.l.

IUCN status
S. gambleana is occasionally found in the forests of north Karnataka and western Maharashtra. The species occurs at considerable number of localities and no threat is known at present. Hence, it is assessed here as List Concern (LC). However, more detailed study on population size and threats are needed to have precise IUCN status. Although the species was assessed as Vulnerable (VU) by Kamat et al. (2020), no data on population size and threat were provided by them to support their assessment.

Note
Salacia gambleana is mostly confused with S. macrosperma due to the similar habit and floral characters. However, the latter can be differentiated by its ellipticoblong leaves, and 6-12-paired lateral nerves. I have examined several collections (from Maharashtra, Karnataka, and Kerala) at CAL, BSI, and MH that were wrongly identified as S. macrosperma and S. beddomei. The oblanceolate leaves and 6-8-paired lateral nerves are amongst the more reliable characters to separate S. gambleana from allied species. Most collections from Agumbe, Karnataka clearly belong to S. gambleana. Wight, Spicil. Neilgh. i. 28. t. 32. October 1845;Icon. Pl. Ind. Orient. 3(3): 5, t. 962. November 1845. (Figure 13

Phenology
Flowers between February to April and fruits may remain till August.

Distribution and habitat
Endemic to the Western Ghats of Goa, Maharashtra, Karnataka, Kerala, and Tamil Nadu (Figure 19). Straggling over small trees in semi evergreen fringed forest, along roads, often grows as large shrubs with distal scandent branches along the edges of plateaus.

IUCN status
S. macrosperma is widely distributed in the Western Ghats, and a good and stable population trend was observed during the field surveys. Also, no major threats or future obstacles were detected. Hence, it is assessed here as Least Concern (LC).

Specimens examined
In addition to Salacia gambleana (see the note under S. gambleana), S. macrosperma also resembles S. salacioides in having entire leaves, axillary fascicles and ovoid fruits. However, the latter differs by its drooping branches, stipulate, broadly lanceolate leaves and orbicular petals. Salacia macrosperma is an extremely variable species, particularly in its leaf size, number of lateral nerves, and smooth to faintly tuberculate fruits. Some variants earlier described as separate species have been compared based on their protologues, and live collections of S. macrosperma from Kerala. The analysis reveals the description of S. macrosperma to be a perfect match for S. beddomei and S. vellaniana. Salacia beddomei Gamble (1916) was described based on Beddome's collection from Anamalai Hills, Tamil Nadu. The species was differentiated from S. macrosperma by having larger leaves with prominent lateral nerves and reddish brown, fimbriate sepal margins. I had an opportunity to observe the original material of S. beddomei at K (K000669921) and MH (MH00001914), and to compare the type of S. macrosperma. After studying the protologue, illustration made by Gamble on the type sheet revealed that the characters used to describe S. beddomei matches those of S. macrosperma, hence it is here relegated to synonymy. Unfortunately, Gamble (1916) had not seen the fruits for S. beddomei and this might have led him to misinterpret the species. Salacia vellaniana from Vellanimala, Thrissur, Kerala (Udayan et al. 2013) is based on characters such as smaller leaves, short petiole, few-flowered cymes, colour of the petals and sepals, and warty fruits. All these characters are within the variation range for S. macrosperma. Gamble in Bull. Misc. Inform. Kew 1916(5): 133. 1916 Type: India, Kerala, Kollam District, Travancore Hills, Nov. 1887 T. F. Bourdillon 18 (MH! barcode MH00001913), lectotype designated by Jadhav et al. (2016).

Phenology
Flowers from November, peak in February and fruits may remain till June.

Distribution and habitat
Endemic to the Western Ghats of Karnataka and Kerala ( Figure 19). It grows in evergreen and moist deciduous forest, coastline scrubland and fringing forest. Its occurrence in southeast Asia as recorded by Kamat et al. (2020) is wrong.

IUCN status
S. malabarica is sporadically distributed in coastal districts of Kerala and south Karnataka. The southernmost distribution is Kottoor Reserve Forests, whereas the northernmost distribution is known from Dakshin Kannada District. As much of the known populations falls within the Biosphere Reserves, hitherto no threat has been observed and the species is seemingly secured and protected. Hence, it is assessed here as Least Concern (LC). The IUCN status as Endangered (EN) by Kamat et al. (2020) is flawed and contradicts their remarks on the species' distribution.

Note
S. malabarica can be recognised by its ellipticoblong or obovate, distinctly serrate leaves, 2-12, pedicellate (1.5-2 cm long) flowers on axillary, ramiflorous, tuberculate, fascicles. It shares similarities with S. gambleana and S. macrosperma but differs in having broadly elliptic to oblong, serrate leaves and long pedicellate flowers. Udayan 04512 (CMPR) from Shendurney Wildlife Sanctuary, has entire to faintly serrate leaves, and c. 2.5 cm long fruiting pedicel, but it appears to agree with S. malabarica in most of the other diagnostic characters of the species.

Phenology
Flowers between January March, followed by fruiting till June.

Distribution and habitat
Endemic to the Western Ghats of Karnataka and Kerala ( Figure 19). It grows in open and semi evergreen forest, fringing forest, along the perennial streams.

ICUN status
The conservation status is assessed as Least Concern (LC). At present the species is known from two localities in Karnataka and Kerala. No threats have been observed as both the locations are well protected and forms part of a Biosphere Reserve.

Note
Lack of mucilaginous or fleshy seeds or endocarp in the fruits of Salacia megacarpa makes its distinct among its congeners. All the Salacia species in the study area have orange-ochreous, tawny fruits at the maturity, except the present species in which the fruits remain are green or greenish yellow at maturity. One of the largest fruits (c. 6 cm long) in the genus was described from the Malaysian S. maingayi M.A. Lawson (Ding Hou 1964). However, S. megacarpa fruits are claimed to be the largest in Salacia and is also distinct due to the presence of three keels running along the longitudinal axis of the fruit. Salacia megacarpa is similar to S. fruticosa with respect to its habit and inf lorescence but the later differs in having crenate leaf margins, orbicular petals and 2-or 3-seeded, orange berries and mucilaginous pulpy seeds. Salacia oblonga also has axillary and ramiflorous, dichotomously branched cymes. The latter can, however, be easily differentiated from the species described here by having subsessile, urceolate f lowers, globose and orange-red fruit. The Indo-Malaysian S. korthalsiana also has similar branched cymes but can be distinguished based on single-seeded fruits. The Southeast Asian S. maingayi is the only other species which has comparable sized large fruits; however, it can be distinguished based on 1 or 2 axillary flowers.

Phenology
Flowers between January to March, fruits may remain till end of July.

Distribution and habitat
Peninsular India (Goa, Karnataka, Kerala, Maharashtra, and Tamil Nadu) and Sri Lanka. Semi-evergreen forest, fringing forest, undergrowth forest, along the plateaux borders. The species occurrence in southeast Asia by Kamat et al. (2020) is erroneous.

IUCN status
Though the species is fairly widely distributed in peninsular India and Sri Lanka, an IUCN Red list category of Vulnerable (VU) [A2cd] is assigned to it (Ved et al. 2015), mainly due to decline in area, extend and habitat quality caused by over exploitation, agricultural invasion, and road construction. Nevertheless, the species needs to be reassessed based on the precise data on population and threat for the Sri Lankan part of its distribution. The Endangered (EN) category assessed by Kamat et al. (2020) is not supported. This is one of the most exploited species from the wild due to its use as an antidiabetic medicine. As the root are richer in the active compound, salacinol, mature plants are often uprooted from the wild. Salacia oblonga is in need of harvest management to ensure sustainability, as well as a mass cultivation programme as an immediate conservation measure. Sujana, Nagaraju, Ratheesh & Anil Kumar in Taiwania

Phenology
Flowers and fruit collected from January to August.

Distribution and habitat
Endemic to the Western Ghats of Kerala ( Figure  19). Climbing over small trees in semi evergreen forest, fringing forest, and along roads.

IUCN status
The species is recorded from only two localities in Wayanad District. Both the localities are adjacent to places of human habitation and threatened by habitat degradation. Few plants were found damaged, perhaps due to road clearing and widening, also confirmed by Sujana et al. (2015). The restricted AOO and EOO, continuing decline (observed) of quality of habitat and continuing decline (inferred) in the number of mature individuals suggests an assessment of Critically Endangered (CR) for this species [B1+2ab(iii,v); D]. Note S. wayanadica shares a similar habit and inflorescence with S. macrosperma, S. gambleana and S. malabarica. However, S. macrosperma differs by having acute to acuminate leaves, laciniate to fimbriate sepals and ovate to broadly elliptic petals. Similarly, S. gambleana differs by having oblanceolate leaves, whereas S. malabarica has serrate leaves, much longer pedicels, and glabrous sepals. Blume, Bijdr. Fl. Ned. Ind. 5: 221. 1825.

Distribution and habitat
India (Andaman and Nicobar Islands), Cambodia, Thailand, Hainan. Low land coastline forest.

Note
Blume (1825) described the species from Indonesia. Later, Ding Hou (1964) erroneously reported the occurrence of this species in peninsular India. It seems he misidentified the collections of S. oblonga (from Maharashtra and Karnataka, the then Bombay Presidency) by Dalzell, Law and Ritchie that are available at K. Subsequent authors, Ramamurthy & Naithani (2000) and Londhe (2000) included the taxon in Maharashtra following Ding Hou (1964). I could not locate any specimen of S. macrophylla to support its occurrence in mainland India. The original material collected by Blume and Maingay of S. macrophylla and S. ovalis is housed at L, K and CAL. For both the names there is no precise indication of type and hence warrant typification. Comparing the original material with protologue, the best specimen is designated here as lectotype following ICN Art. 9.3 (Turland et al. 2018). Wight, Ill. Ind. Bot. 1: 134. 1838.

Distribution and habitat
India (Andaman and Nicobar Islands) and Sri Lanka. Evergreen forest.

Note
Wight (1838) described the species based on Walker's collection from Sri Lanka. He was not certain about its occurrence in Malabar of South India. In the protologue, he mentioned that "the identity of the Malabar specimen is doubtful as flowers were not seen, leaves are less distinctly reticulate" and added characters of the fruit is doubtful. Gamble (1918) listed S. reticulata in Madras Presidency based on Rama Rao's collection (CAL 87440). Further details on the doubtful occurrence of S. reticulata in India and misapplication of S. reticulata to different elements by various authors in India can be found in . The occurrence recorded in peninsular India by Kamat et al. (2020) is an error.