Summary
The dipterocarp tribe Shoreae, perhaps more than any other members of this elegant family of often giant emergent trees, is familiar to all who visit the once ubiquitous lowland forests of tropical Asia. Timbers of the genus Shorea comprised the bulk of hardwood traded on international markets for thirty years, since the nineteen seventies. Distinct species groups came to be recognised by taxonomists: for their characteristic androecia, and sometimes differences in bark and wood anatomy, while generic status for some was then proposed on embryological evidence. A new molecular phylogeny of the family, and this tribe, substantially confirms the embryological case. This paper formalises resultant nomenclatural changes. Those of us who recall the majestic forested former landscapes of the Sunda lands may regret this decision. But now, with landscapes irretrievably changed by serried ranks of oil palm and forest degradation, the case for recognition of the proposed new entities can guide conservation planners in recognising and making the case for permanently conserving surviving undisturbed stands of exceptional composition or diversity.
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Introduction
Linnaeus (1753, 1771) was the first to document genera in our family, the South Asian Vatica and Vateria, both trees of moderate stature and persistent but unremarkable calyces. The great German botanist of the Indies, K. L. Blume (1825), recognised a new family, Dipterocarpaceae, for the first of the emergent-crowned forest giants whose large fruit spiral down on sepals expanded into two oar-like wings, Dipterocarpus C.F.Gaertn. Further generic descriptions also relied on characters of the fruit calyx, whether the winged sepals are equal (Dryobalanops C.F.Gaertn.), unequal with three longer (Parashorea Kurz, Shorea Roxb. ex C.F.Gaertn., Doona Thwaites, Pentacme A.DC.; or three shorter therefore wingless (Anisoptera Korth., Upuna Symington, Cotylelobium Pierre, many Vatica L., Hopea Roxb.); or with all five shorter than the ripe nut (Vatica, Vateria, Vateriopsis F.Heim). Species were found which conformed to characters of winged genera but whose fruit lacked wings, and on that basis alone were identified as novel genera (Balanocarpus Bedd., Isoptera Scheff. ex Burck). F. Heim (1892) was first to discriminate supraspecific taxa on multiple characters (stamens, embryo, petiole anatomy), and to define distinct genera sharing the fruit calyx of Shorea, and sections within Shorea itself (Brachypterae, Pachycarpae), which Brandis (1895) expanded in his comprehensive revision of then known Asian taxa.
The number of known species expanded thereafter, notably from the work of D. F. van Slooten at the then Netherland Indies herbarium at Buitenzorg, Java (Bogor, Indonesia). Forestry botanical researchers were first to describe dipterocarps as living organisms, differentiating taxa on stature, and form including bark and buttresses and sometimes juveniles: first in the Philippines and later peninsular Malaya (peninsular Malaysia) Foxworthy (1911, 1938), then comprehensively expanded and rigorously researched by C. F. Symington (1939, 1943), focusing on Malayan species. Symington, aided by comparative wood anatomical research by Desch (1936, 1941), recognised three major timbers among botanically heterogeneous Malayan Shorea, botanically defined by bark morphology and wood anatomy as well as floral especially stamen characters: balau, comprising species with the characters of the Indian type species, S. robusta C.F.Gaertn., meranti pa’ang (white meranti), meranti damar hitam (yellow meranti) and the large but more heterogeneous group red meranti. Ashton (1963, 1967, 1972, 1978, 1980, 1982) accepted Symington’s criteria, reducing Pentacme (Ashton 1978) and Doona (Ashton 1980), which share the fruit calyx of Shorea, to sections within it, and providing formal botanical status to the timber groups (Ashton 1963) while adding two not occurring in Peninsular Malaysia: Pachycarpae F.Heim and Rubella P.S.Ashton. Full diagnoses of supraspecific taxa are provided in Ashton 1982.
Ashton divided Dipterocarpoideae into two tribes: Dipterocarpeae, which included Dipterocarpus, with imbricate fruit calyx, and Shoreae, which included Dryobalanops, with fruit calyx imbricate at base where sepals become incrassate and concave.
Meanwhile Maury (1978; Maury-Lechon 1979; Maury-Lechon & Curtet 1998) had proposed a novel reclassification within what Ashton (1982) recognises as Tribe Shoreae, principally based on examination of embryology within ripe fruits of forty species within it, raising Ashton’s infrageneric taxa, or returning them back to generic rank. The two classifications are compared by Ashton et al. (2021, Table 1). Maury recognised three subtribes within tribe Shoreae: a heterogeneous type subtribe including Hopea, Neobalanocarpus P.S.Ashton, Shorea, Richetia F.Heim and Rubroshorea as genera; monotypic Parashorineae; and Anthoshorineae including Anthoshorea Pierre, Doona and Pentacme as genera. The monotypic section Neohopea P.S.Ashton was not included in the study. Maury was unable to detect embryological characters diagnostic of Symington’s Red Meranti subgroup of his Shorea, for which she accepted Meijer’s subgenus Rubroshorea which is based solely on the pink to red-brown inner bark and wood, characters absent in a few species where they are yellow-brown, yet present in S. guiso (Blanco) Blume, section Shorea, subsection Shorea.
Ashton remained reluctant to accept the substantial name changes and recombinations implied: first because they need confirmation from more than the forty studied out of the c. 200 now recognised before embryological characters are confirmed as consistent within supraspecific taxa, including those now recognised within the heterogeneous red merantis; that they are correlated with other diagnostic characters; and lastly but importantly, because during the quarter century, 1970 – 2000, when dipterocarp timber dominated the international timber trade, and when there was still hope that dipterocarp forests could still be managed sustainably, nomenclatural fragmentation of the most widely known timber genus would have led to confusion among that wide community of foresters, silviculturists, loggers, and business interests who relied on easy and reliable identification of field groups and often species.
Thanks now to the ubiquitous replacement of mixed dipterocarp forest by oil palm in the aseasonal wet lowlands of tropical Asia, the world in which we live has changed immeasurably. Advances in molecular systematics have provided solid confirmation of much of Géma Maury’s pioneering work.
In a recent study on three plastid regions (rbcL, trnK-matK-trnk, trnT-trnL-trnF; Heckenhauer et al. 2017), Dipterocarpus is placed as sister to tribe Shoreae (Fig. 1). Although this relationship is only moderately supported in this study, it clearly separates Dipterocarpus from the remaining genera of tribe Dipterocarpeae containing the following genera: Anisoptera, Cotylelobium, Stemonoporus, Upuna, Vateria, Vateriopsis and Vatica. This was observed in earlier molecular studies (e.g. Kajita et al. 1998; Yulita et al. 2005; Gamage et al. 2006) and has been recently resolved with support based on the analyses of plastome and nuclear cistron (NRC) data (Cvetković et al. 2022).
Schematic presentation of molecular phylogenetic trees obtained from plastid (left) and nuclear (right) data. Species were collapsed to major clades. Neobalanocarpus was not included in the nuclear data set. green: tribe Dipterocarpeae, red: tribe Shoreae; *: the plastid data set included not only Vatica but also Anisoptera, Cotylelobium, Stemonoporus, Upuna, Vateria, Vateriopsis and Vatica; **: genus Shorea sensu Ashton. For detailed non-collapsed phylogenetic trees including support values for each clade see: Heckenhauer et al. 2017, 2018, 2019.
Interestingly, while most members of Dipterocarpaceae are pollinated by pollenivores (bees, thrips (Ashton 2014)), Dipterocarpus is mainly, perhaps solely, pollinated by nectarivorous Lepidoptera (Ghazoul 1997; Harrison 2005; Ashton 2014).
Regarding the tribe Shoreae, molecular analyses based on three plastid markers (Heckenhauer et al. 2017) found the monophyletic genera Hopea, Neobalanocarpus and Parashorea to be nested within the genus Shorea sensu Ashton, in broad agreement with Maury-Lechon. This also has been reported in earlier molecular analyses (e.g. Yulita et al. 2005; Gamage et al. 2006) and is strongly supported by next-generation sequencing plastome (whole plastid genomes, Heckenhauer et al. 2019; Cvetković et al. 2022) and nuclear data (RADseq-derived SNP data set, Heckenhauer et al. 2018; NRC data Cvetkovic et al. 2022). Thus, the genus Shorea sensu Ashton is polyphyletic in its current circumscription. Specifically, according to these molecular studies, the tribe Shoreae is divided into two highly supported major clades consisting of (1) Shorea section Doona, Shorea section Anthoshorea, Neobalanocarpus and Hopea and (2) of Shorea section Richetia, Shorea section Shorea, Parashorea and Shorea section Rubroshorea (Fig. 1); each as distinct microclades. Thus, molecular evidence divides genus Shorea sensu Ashton into five groups corresponding to Ashton’s section Doona, Anthoshorea, Rubroshorea, Shorea balau (selangan batu) and Richetia. These studies did not include Ashton’s sections Pentacme and Neohopea. The position of Parashorea within the major clade (2) remains unclear: In the plastome trees, Parashorea is sister to Shorea section Rubroshorea and Shorea section Shorea, whereas it forms a clade with Richetia in the RADseq trees (Fig. 1). This suggests that the origin of this genus may be associated with ancient hybridisation. Further, Neobalanocarpus was recovered as nested in Hopea based on analysis of NRC data (Cvetković et al. 2022). This is incongruent with the position retrieved using plastid data, where it is sister to Hopea (Fig. 1, Heckenhauer et al. 2017, 2019; Cvetković et al. 2022)
Molecular analyses support subgenus Rubroshorea Meijer as the largest microclade within the Shoreae clade, but lack consistent support for sections in Rubroshorea recognised by Symington and Ashton. The analysis reveals that, within Rubroshorea, the sections Brachypterae and Mutica especially fail to form monophyletic groups. This has also been observed for sect. Pachycarpae and sect. Shorea within Shorea s.str (Cvetkovic et al. 2022). Future molecular studies could shed light into these sectional classifications, as well as resolve the position of Neohopea, Pentacme and Parashorea.
Taxonomic implications
The inconsistency of placement of Dipterocarpus in molecular phylogenies is consistent with its unique morphology, and we agree with Cvetković et al. (2022) to isolate it in a monotypic tribe, requiring the renaming of the former Tribe Dipterocarpeae:
Tribe Dipterocarpeae
Buds, stipules, flowers and often leaves exceptionally large; stipule scars amplexicaul; leaf vernation plicate leaving opened leaves corrugate between veins; petiole vascular bundles in one or more complex rings; flower ovary and androecium enclosed but free from sepals united at base into a tube; petals large, lorate, not concave at base; ovary imbedded in receptacle at base; pollen gaping tricolporate, at least three times as large as in other Dipterocarpoideae, margins absent or much reduced, exine tilioid; vessels solitary, resin canals large, scattered, fibres with bordered pits. n = 11.
Tribe Vaticeae
Sepals free to an unswollen base and or united around a partially inferior ovary. Corolla free or loosely connate at base, spreading, not cupped or prominently overlapping forming a vase-like chamber at base. Vessels solitary, resin canals scattered, fibres with bordered pits. n = 11.
Tribe Shoreae
Fruit sepals expanded and imbricate at the incrassate cupped base, sepal vasculation more or less overlapping at margin in flower; petals at anthesis more or less strongly imbricate, cupped forming a vase-like chamber and contorted at base. Vessels grouped, resin canals in tangential bands. N = 7.
Published taxonomic subtribes of the tribe Shoreae are not consistently supported in the molecular phylogenies. Heckenhauer et al. (2017, 2019) did consistently recognise two major clades within Shorea sensu Ashton (1980, 1982); but we know of no morphological or anatomical criteria by which these two clades can be grouped separately as diagnosable taxonomic entities. One of these clades also includes the established genera Hopea and Neobalanocarpus together with Ashton’s sections Doona, Anthoshorea and Pentacme on Maury-Lechon’s embryological evidence; while the other bifurcates as Ashton’s section Richetioides (Symington’s yellow meranti timber group, earlier recognised as a genus, Richetia (Heim 1892)); and as a second minor clade which includes the type section of Shorea, and Ashton’s five sections within Symington’s red meranti timber group already published as the genus Rubroshorea Meijer.
We therefore propose not to recognise Heckenhauer et al.’s two clades as taxonomically definable entities. Based on recent phylogenomic analyses of Cvetković et al. (2022), Hopea Roxb. and Neobalanocarpus P.S.Ashton are retained as established genera. The former genera Doona Thwaites, Pentacme A.DC., Richetia F.Heim (1892) are resurrected. Symington’s and Ashton’s Shorea section Anthoshorea F.Heim is raised to generic rank. We suggest to retain Neohopea P.S.Ashton as a distinct section within Shorea Roxb. ex C.F.Gaertn. which otherwise retains only the type section. Rubroshorea Meijer is here recognised as a genus as a stimulus for others to identify morphological or anatomical characters by which it can be diagnosed in addition to the molecular findings; the five sections within it are retained as such, with their species awarded new binomials.
The following species are now recognised in Shorea sensu Ashton (1982), with new binomial combinations where required. Full citations are provided in Ashton 1980 (for Malesia), 1982 (for Sri Lanka) and 2004 (for Borneo post-1982).
Species which retain their present binomial:
Shorea Roxb. ex C.F.Gaertn. (Gaertner 1805: 47, t. 186, f. 1). Type species: Shorea robusta C.F.Gaertn. (Gaertner 1805: 48, t. 186, f. 1).
vernacular names. Sal (India), balau (Peninsular Malaysia), selangan batu (Borneo Malay), tekam (Iban).
Section Shorea
Shorea astylosa Foxw. (Foxworthy 1918: 188).
Shorea atrinervosa Symington (1939: 363).
Shorea brunnescens P.S.Ashton (1967: 283).
Shorea calcicola P.S.Ashton (2004: 239 – 240).
Shorea ciliata King (1893: 118).
Shorea collina Ridl. (Ridley 1910: 182).
Shorea crassa P.S.Ashton (1962: 271).
Shorea domatiosa P.S.Ashton (1962: 285).
Shorea dyerii Thwaites (1885: 204).
Shorea exelliptica Meijer (1963: 323).
Shorea falcata J.E.Vidal (1962: 325).
Shorea falcifera Dyer ex Brandis (1895: 86).
Shorea falciferoides Foxw. (Foxworthy 1918: 189).
Shorea geniculata Symington ex P.S.Ashton (1962: 291).
Shorea guiso (Blanco) Blume (1856: 34).
Shorea havilandii Brandis (1895: 82).
Shorea hypoleuca Meijer (1963: 329).
Shorea inappendiculata Burck (1887: 206).
Shorea leptoderma Meijer (1963: 331).
Shorea lissophylla Thwaites (1864: 402).
Shorea lumutensis Symington (1939: 364).
Shorea lunduensis P.S.Ashton (1967: 284).
Shorea malibato Foxw. (Foxworthy 1913: 1955).
Shorea materialis Ridl. (Ridley 1910: 183).
Shorea oblongifolia Thwaites (1858: 36).
Shorea obscura Meijer (1963: 333).
Shorea obtusa Wall. ex Blume (1856: 32, t. 8).
Shorea ochrophloia Strugnell ex Symington (1935: 268).
Shorea pallescens P.S.Ashton (1972, publ. 1973: 360).
Shorea robusta C.F.Gaertn. (Gaertner 1805: 48, t. 186, f. 1).
Shorea scrobiculata Burck (1887: 207).
Shorea seminis (de Vriese) Slooten (1929: 204).
Shorea submontana Symington (1939: 368).
Shorea sumatrana (Slooten) Desch (1934: 201).
Shorea superba Symington (1960: 491).
Shorea thorelii Pierre ex Laness. (Lanessan 1886: 302).
Shorea tumbuggaia Roxb. (Roxburgh 1814).
Shorea section Barbata (Symington ex P.S.Ashton) P.S.Ashton & J.Heck. stat. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217905-1
Shorea section Shorea subsection Barbata Symington ex P.S.Ashton (1963: 266).
Shorea asahi P.S.Ashton (1962: 279).
Shorea biawak P.S.Ashton (1962: 281).
Shorea glauca King (1893: 117).
Shorea ladiana P.S.Ashton (1962: 295).
Shorea laevis Ridl. (Ridley 1922: 232).
Shorea maxwelliana King (1893: 114).
Shorea micans P.S.Ashton (1978: 38).
Species returned to an earlier generic status, or designated a new one:
Pentacme A.DC. (De Candolle 1868: 626). Shorea section Pentacme (A.DC.) P.S.Ashton (1978: 36). Type species: Pentacme siamensis (Miq.) Kurz (1870: 66).
vernacular name. Temak batu (Peninsular Malaysia).
Pentacme siamensis (Miq.) Kurz (1870: 66). Shorea siamensis Miq. (Miquel 1864: 214).
Pentacme paucinervis Brandis (1895: 73).
Shorea contorta S.Vidal (1883: 15). Hololectotype: Philippines, Luzon, Rizal Province, Vidal 987 (K, designated here).
Neohopea (P.S.Ashton) P.S.Ashton, stat. nov. Type species: Shorea isoptera P.S.Ashton, Gard. Bull. Sing. 19: 273 (1962).
http://www.ipni.org/urn:lsid:ipni.org:names:77217906-1
Shorea section Neohopea P.S.Ashton, Gard. Bull. Singapore 20: 266 (1963).
vernacular name. Selangan batu main bulu ayam.
Neohopea isoptera (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217907-1
Shorea isoptera P.S.Ashton, Gard. Bull. Singapore 19: 293 (1962).
Doona Thwaites (1858: 35). Shorea section Doona (Thwaites) P.S.Ashton (1972: 360). Type species Doona congestiflora Thwaites (1858: 35).
vernacular name. Dun (Sri Lanka).
Doona affinis Thwaites (1858: 35). Shorea affinis (Thwaites) P.S.Ashton (1972, publ. 1973: 361).
Doona congestiflora Thwaites (1858: 35). Shorea congestiflora (Thwaites) P.S.Ashton (1972, publ. 1973: 362).
Doona cordifolia Thwaites (1858: 35). Shorea cordifolia (Thwaites) P.S.Ashton (1972, publ. 1973: 362).
Doona disticha (Thwaites) Pierre (1890: t. 237). Shorea disticha (Thwaites) P.S.Ashton (1972, publ. 1973: 362). Basionym: Vateria disticha Thwaites (1864: 404).
Doona gardneri Thwaites (1858: 35). Shorea gardneri (Thwaites) P.S.Ashton (1972, publ. 1973: 362).
Doona macrophylla Thwaites (1864: 402). Shorea megistophylla P.S.Ashton (1972, publ. 1973: 362) nom. nov.
Doona ovalifolia Thwaites (1864: 404). Shorea ovalifolia (Thwaites) P.S.Ashton (1972, publ. 1973: 363).
Doona trapezifolia Thwaites (1858: 35). Shorea trapezifolia (Thwaites) P.S.Ashton (1972, publ. 1973: 363).
Doona venulosa Thwaites (1864: 402). Shorea worthingtonii P.S.Ashton (1972, publ. 1973: 363).
Doona zeylanica Thwaites (1851: t. 12; 1852: 7). Shorea zeylanica (Thwaites) P.S.Ashton. (1972, publ. 1973: 363).
Anthoshorea Pierre, Fl. Forest. Cochinchine (1891) ante t. 250; Pierre ex F.Heim, Rech. Dipterocarp. (1892: 41) nom. illeg. supfl. (nomen in syn. sub Shorea henryana). Type: Anthoshorea harmandii Pierre.
vernacular names. Meranti pa’ang (Peninsular Malaysia), raruk (Iban).
Anthoshorea agamii (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217908-1
Shorea agamii P.S.Ashton, Gard. Bull. Singapore 19: 270 (1962).
Anthoshorea assamica (Dyer) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217909-1
Shorea assamica Dyer, Fl. Brit. India [J. D. Hooker] 1: 307 (1874).
Anthoshorea bentongensis (Foxw.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217910-1
Shorea bentongensis Foxw., Malayan Forest Rec. 10: 169 (Foxworthy 1932).
Anthoshorea bracteolata (Dyer) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217911-1
Shorea bracteolata Dyer, Fl. Brit. India [J. D. Hooker] 1 (2): 305 (1874).
Anthoshorea confusa (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217912-1
Shorea confusa P.S.Ashton, Gard. Bull. Singapore 31: 44 (1978).
Anthoshorea cordata (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217913-1
Shorea cordata P.S.Ashton, Gard. Bull. Singapore 22: 285 (1967).
Anthoshorea dealbata (Foxw.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217914-1
Shorea dealbata Foxw., Malayan Forest Rec. 10: 192 (Foxworthy 1932).
Anthoshorea farinosa (C.E.C.Fisch.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217915-1
Shorea farinosa C.E.C.Fisch., Bull. Misc. Inform. Kew. 1926: 461 (Fischer 1926).
Anthoshorea gratissima (Wall. ex Kurz) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217916-1
Shorea gratissima (Wall. ex Kurz) Dyer, Fl. Brit. India [J. D. Hooker] 1: 307 (1874). Hopea gratissima Wall. ex Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 42 (2): 61 (1873).
Anthoshorea henryana (Pierre ex Laness.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217917-1
Shorea henryana Pierre ex Laness., Pl. Util. Col. Franç. (Lanessan 1886: 302).
Anthoshorea hulanidda (Kosterm.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217918-1
Shorea hulanidda Kosterm., Bot. Jahrb. Syst. 104: 196 (Kostermans 1983).
Anthoshorea hypochra (Hance) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217919-1
Shorea hypochra Hance, J. Bot. 14: 242 (1876).
Anthoshorea javanica (Koord. & Valeton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217920-1
Shorea javanica Koord. & Valeton, Bull. Inst. Bot. Buitenzorg 2: 3 (Koorders & Valeton 1899).
Anthoshorea lamellata (Foxw.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217921-1
Shorea lamellata Foxw., Malayan Forest Rec. 10: 278 (Foxworthy 1932).
Anthoshorea montigena (Slooten) P.S.Ashton & J.Heck., comb. nov. Hololectotype, designated here: Indonesia, Java, Cult. Hort. Bog. VIII.D25 ex Kajeli, Buru, (BO).
http://www.ipni.org/urn:lsid:ipni.org:names:77217922-1
Shorea montigena Slooten, Reinwardtia 2: 57 (1952).
Anthoshorea ochracea (Symington) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Malaysia, Sarawak, Kapit Distr., Ulu Balleh, Merirai, Spurway S167 (KEP) (Ashton 2004: 298).
http://www.ipni.org/urn:lsid:ipni.org:names:77217923-1
Shorea ochracea Symington, Gard. Bull. Straits Settlem. 8: 285 (1935).
Anthoshorea polita (S.Vidal) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217924-1
Shorea polita S.Vidal, Sin. Gen. Pl. Leños. Filip., Atlas (1883: xv, t. 15, f. D).
Anthoshorea resinosa (Foxw.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217925-1
Shorea resinosa Foxw., Malayan Forest Rec. 10: 234 (Foxworthy 1932).
Anthoshorea retinodes (Slooten) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217926-1
Shorea retinodes Slooten, Bull. Jard. Bot. Buitenzorg, sér. 3, 18: 243 (1949).
Anthoshorea roxburghii (G.Don) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217927-1
Shorea roxburghii G.Don, Gen. Hist. 1: 813 (1831).
Anthoshorea stipularis (Thwaites) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217928-1
Shorea stipularis Thwaites, Enum. Pl. Zeyl. (1858: 36).
Anthoshorea symingtonii (G.H.S.Wood) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217929-1
Shorea symingtonii G.H.S.Wood, Gard. Bull. Singapore 17: 493 (1959, publ. 1960).
Anthoshorea virescens (Parijs) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77217930-1
Shorea virescens Parijs, Repert. Spec. Nov. Regni Veg. 33: 244 (1933).
Rubroshorea (Meijer) P.S.Ashton & J.Heck., stat. nov. Type species: Shorea parvifolia Dyer (1874: 305).
http://www.ipni.org/urn:lsid:ipni.org:names:77298305-1
Shorea subgen. Rubroshorea Meijer, Acta Bot. Neerl. 12: 322 (1963).
vernacular names. Meranti (general); seraya (Sabah, Kelantan); lop, perawan (Iban); lauan, luan (Tagalog).
Rubroshorea section Brachyptera (F.Heim) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298306-1
Shorea sect. Brachyptera F.Heim, Rech. Dipterocarp. (1892: 46).
Rubroshorea almon (Foxw.) P.S.Ashton & J.Heck., comb. nov. Lectotype: Philippines, Negros Occidental, Cadiz, Whitford F.B.11647 (K, isolectotype L) (Ashton 2004: 223).
http://www.ipni.org/urn:lsid:ipni.org:names:77298307-1
Shorea almon Foxw., Philipp. J. Sci. 67: 313 (Foxworthy 1938).
Rubroshorea andulensis (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298308-1
Shorea andulensis P.S.Ashton, Gard. Bull. Singapore 19: 275 (1962).
Rubroshorea balangeran (Korth.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298310-1
Shorea balangeran (Korth.) Burck, Ann. Jard. Bot. Buitenzorg 6: 214 (1887). Hopea balangeran Korth., Verh. Nat. Gesch. Ned. Bezitt., Bot. 2: 74 (Korthals 1841).
Rubroshorea bullata (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298311-1
Shorea bullata P.S.Ashton, Gard. Bull. Singapore 19: 283 (1962).
Rubroshorea carapae (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298312-1
Shorea carapae P.S.Ashton, Gard. Bull. Singapore 22: 294 (1967).
Rubroshorea coriacea (Burck) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298314-1
Shorea coriacea Burck, Ann. Jard. Bot. Buitenzorg 6: 214 (1887).
Rubroshorea fallax (Meijer) P.S.Ashton & J.Heck., comb. nov. Lectotype: Malaysia, Sabah, Beaufort Hill, G. H. S. Wood SAN A1734 (hololectotype K; isolectotypes L, KEP) (Ashton 2004: 256).
http://www.ipni.org/urn:lsid:ipni.org:names:77298315-1
Shorea fallax Meijer, Acta Bot. Neerl. 12: 335 (1963).
Rubroshorea flaviflora (G.H.S.Wood ex P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298316-1
Shorea flaviflora G.H.S.Wood ex P.S.Ashton, Gard. Bull. Singapore 19: 289 (1962).
Rubroshorea flemmichii (Symington) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298317-1
Shorea flemmichii Symington, Gard. Bull. Straits Settlem. 10: 278 (1939).
Rubroshorea inaequilateralis (Symington) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Brunei, Kuala Belait, Zainal F.M.S. 30351 (KEP) (Ashton 2004: 270).
http://www.ipni.org/urn:lsid:ipni.org:names:77298318-1
Shorea inaequilateralis Symington, Gard. Bull. Straits Settlem. 8: 281 (1935).
Rubroshorea johorensis (Foxw.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298323-1
Shorea johorensis Foxw., Malayan Forest Rec. 10: 236 (Foxworthy 1932).
Rubroshorea kunstleri (King) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Malaysia, Peninsular Malaysia, Perak Distr., Larut, Kunstler 3674 (K) (Ashton 2004: 276).
http://www.ipni.org/urn:lsid:ipni.org:names:77298324-1
Shorea kunstleri King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 62: 116 (1893).
Rubroshorea monticola (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298325-1
Shorea monticola P.S.Ashton, Gard. Bull. Singapore 19: 297 (1962).
Rubroshorea pachyphylla (Ridl. ex Symington) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298326-1
Shorea pachyphylla Ridl. ex Symington, J. Malayan Branch Roy. Asiat. Soc. 19: 163 (1941).
Rubroshorea palembanica (Miq.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298327-1
Shorea palembanica Miq., Fl. Ned. Ind., Eerste Bijv. 3: 487 (Miquel 1861).
Rubroshorea palosapis (Blanco) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298328-1
Shorea palosapis (Blanco) Merr., Sp. Blancoan.: 271 (Merrill 1918). Dipterocarpus palosapis Blanco, Fl. Filip., ed. 2: 312 (1845). Isolectotype: ‘Philippines’, Cuming 892 (K). Merrill (1918) chose this collection as lectotype; the hololectotype was presumably in the Manila herbarium, now destroyed.
Rubroshorea parvistipulata (F.Heim) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298329-1
Shorea parvistipulata F.Heim, Bull. Mens. Soc. Linn. Paris 2: 974 (1891).
Rubroshorea pauciflora (King) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298330-1
Shorea pauciflora King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 62: 116 (1893).
Rubroshorea platyclados (Slooten ex Foxw.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298331-1
Shorea platyclados Slooten ex Foxw., Malayan Forest Rec. 10: 214 (Foxworthy 1932). Hololectotype: Peninsular Malaysia, Selangor, Ulu Gombak, Symington KEP 24416, (KEP) (Ashton 2004: 315).
Rubroshorea polysperma (Blanco) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298332-1
Shorea polysperma (Blanco) Merr., Publ. Bur. Sci. Gov. Lab. 27: 22 (Merrill 1905). Mocanera polysperma Blanco, Fl. Filip. [F. M. Blanco] (1837: 448).
Rubroshorea pubistyla (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298333-1
Shorea pubistyla P.S.Ashton, Gard. Bull. Singapore 22: 297 (1967).
Rubroshorea scaberrima (Burck) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Indonesia, Java, Cult. Hort. Bogor. s.n., tree VIII D 54 (this is a gardens tree number, no collector cited) (BO) (Ashton 2004: 328).
http://www.ipni.org/urn:lsid:ipni.org:names:77298334-1
Shorea scaberrima Burck, Meded. Lands Plantentuin 3: 22 (1886).
Rubroshorea selanica (Blume) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298335-1
Shorea selanica (Lam.) Blume, Mus. Bot. 2: 33 (1856). Dammara selanica Lam., Encycl. [J. Lamarck & al.] 2 (1): 259 (Lamarck 1786).
Rubroshorea smithiana (Symington) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298369-1
Shorea smithiana Symington, Gard. Bull. Straits Settlem. 9: 345 (1938).
Rubroshorea venulosa (G.H.S.Wood ex Meijer) P.S.Ashton & J.Heck., comb. nov. Lectotype: Malaysia, Sabah, Sipitang Distr., Menggalong Forest Reserve, G. H. S. Wood SAN 15134 (hololectotype K; isolectotypes KEP, SAN) (Ashton 2004: 344).
http://www.ipni.org/urn:lsid:ipni.org:names:77298370-1
Shorea venulosa G.H.S.Wood ex Meijer, Acta Bot. Neerl. 12: 342 (1963).
Rubroshorea waltonii (G.H.S.Wood ex Meijer) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298371-1
Shorea waltonii G.H.S.Wood ex Meijer, Acta Bot. Neerl. 12: 3442 (1963).
Rubroshorea section Rubroshorea subsection Auriculatae (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov. Type species: Rubroshorea macroptera (Dyer) P.S.Ashton & J.Heck.
http://www.ipni.org/urn:lsid:ipni.org:names:77298377-1
Shorea section Mutica subsection Auriculatae P.S.Ashton, Gard. Bull. Singapore 22: 300 (1967).
Rubroshorea acuta (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298382-1
Shorea acuta P.S.Ashton, Gard. Bull. Singapore 19: 268 (1962).
Rubroshorea ferruginea (Dyer ex Brandis) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298383-1
Shorea ferruginea Dyer ex Brandis, J. Linn. Soc., Bot. 31: 91 (1895).
Rubroshorea macroptera (Dyer) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298384-1
Shorea macroptera Dyer, Fl. Brit. India [J. D. Hooker] 1: 308 (1874).
Rubroshorea myrionerva (Symington ex P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298385-1
Shorea myrionerva Symington ex P.S.Ashton, Gard. Bull. Singapore 19: 299 (1962).
Rubroshorea sagittata (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298386-1
Shorea sagittata P.S.Ashton, Gard. Bull. Singapore 22: 299 (1967).
Rubroshorea slootenii (G.H.S.Wood ex P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298387-1
Shorea slootenii G.H.S.Wood ex P.S.Ashton, Gard. Bull. Singapore 19: 312 (1962).
Section Rubroshorea, Subsection Rubroshorea. Type species (designated here): Rubroshorea parvifolia (Dyer) P.S.Ashton & J.Heck.
Section Mutica Brandis, J. Linn. Soc. Bot. 31: 100 (1895).
Rubroshorea acuminata (Dyer) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298388-1
Shorea acuminata Dyer, Fl. Brit. India [J. D. Hooker] 1: 305 (1874).
Rubroshorea argentifolia (Symington) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298389-1
Shorea argentifolia Symington, Gard. Bull. Singapore 17: 489 (1960).
Rubroshorea curtisii (Dyer ex King) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Malaysia, Peninuslar Malaysia, Penang Hill, Curtis 1394 (K) (Ashton 2004: 245).
http://www.ipni.org/urn:lsid:ipni.org:names:77298390-1
Shorea curtisii Dyer ex King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 62: 111 (1893).
Rubroshorea dasyphylla (Foxw.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298391-1
Shorea dasyphylla Foxw., Malayan Forest Rec. 10: 224 (Foxworthy 1932).
Rubroshorea foraminifera (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298392-1
Shorea foraminifera P.S.Ashton, Gard. Bull. Singapore 22: 295 (1967).
Rubroshorea hemsleyana (King) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Malaysia, Peninsular Malaysia, Perak, Larut, Kunstler 6670 (K) (Ashton 2004: 266).
http://www.ipni.org/urn:lsid:ipni.org:names:77298393-1
Shorea hemsleyana (King) King ex Foxw., Malayan Forest Rec. 10: 1967 (Foxworthy 1932). Balanocarpus hemsleyanus King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 62: 134 (1893).
Rubroshorea lepidota (Korth.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298394-1
Shorea lepidota (Korth.) Blume, Mus. Bot. 2: 32 (1856). Vatica lepidota Korth., Verh. Nat. Gesch. Ned. Bezitt., Bot. 3: 73 (Korthals 1841).
Rubroshorea leprosula (Miq.) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Indonesia, Sumatra, loc. incert., Teijsmann Herb. Bog. 1122 (L) (Ashton 2004: 282).
http://www.ipni.org/urn:lsid:ipni.org:names:77298395-1
Shorea leprosula Miq., Fl. Ned. Ind., Eerste Bijv. 3: 487 (Miquel 1861).
Rubroshorea macrantha (Brandis) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298396-1
Shorea macrantha Brandis, J. Linn. Soc., Bot. 31: 97 (1895).
Rubroshorea ovata (Dyer ex Brandis) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Malaysia, Peninsular Malaysia, Penang Hill, Curtis 201 (K) (Ashton 2004: 303).
http://www.ipni.org/urn:lsid:ipni.org:names:77298397-1
Shorea ovata Dyer ex Brandis, J. Linn. Soc., Bot. 31: 91 (1895).
Rubroshorea pallidifolia (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298398-1
Shorea pallidifolia P.S.Ashton, Gard. Bull. Singapore 22: 296 (1967).
Rubroshorea parvifolia (Dyer) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298399-1
Shorea parvifolia Dyer, Fl. Brit. India [J. D. Hooker] 1: 306 (1874).
Rubroshorea platycarpa (F.Heim) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298400-1
Shorea platycarpa F.Heim, Bull. Mens. Soc. Linn. Paris 2: 956 (1891).
Rubroshorea quadrinervis (Slooten) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Indonesia, West Kalimantan, Sambas, Sungei Akar, (collector not cited) bb. 29450 (BO) (Ashton 2004: 319).
http://www.ipni.org/urn:lsid:ipni.org:names:77298401-1
Shorea quadrinervis Slooten, Bull. Jard. Bot. Buitentzorg, ser. 3, 17: 220 (1942).
Rubroshorea retusa (Meijer) P.S.Ashton & J.Heck., comb. nov. Hololectotype: C. Kalimantan, Barito Ulu, Muara Tewe, Malinka bb. 29715 (BO) (Ashton 2004: 321).
http://www.ipni.org/urn:lsid:ipni.org:names:77298402-1
Shorea retusa Meijer, Acta. Bot. Neerl. 12: 340 (1963).
Rubroshorea revoluta (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298403-1
Shorea revoluta P.S.Ashton, Gard. Bull. Singapore 19: 304 (1962).
Rubroshorea rubra (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298404-1
Shorea rubra P.S.Ashton, Gard. Bull. Singapore 19: 309 (1962).
Rubroshorea rugosa (F.Heim) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298405-1
Shorea rugosa F.Heim, Bull. Mens. Soc. Linn. Paris 2: 973 (1891).
Rubroshorea scabrida (Symington) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298406-1
Shorea scabrida Symington, Gard. Bull. Straits Settlem. 8: 287 (1935).
Rubroshorea singkawang (Miq.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298407-1
Shorea singkawang (Miq.) Burck, Ann. Jard. Buitenzorg 6: 219 (1887). Hopea singkawang Miq., Fl. Ned. Ind., Eerste Bijv. 3: 489 (Miquel 1861).
Rubroshorea teysmanniana (Dyer ex Brandis) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298408-1
Shorea teysmanniana Dyer ex Brandis, J. Linn. Soc. Bot. 31: 100 (1895).
Rubroshorea uliginosa (Foxw.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298409-1
Shorea uliginosa Foxw., Malayan Forest Rec. 10: 210 (Foxworthy 1932).
Rubroshorea section Ovalis (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298410-1
Shorea section Ovalis P.S.Ashton, Gard. Bull. Singapore 20: 268 (1963).
Rubroshorea ovalis (Korth.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298411-1
Shorea ovalis (Korth.) Blume, Mus. Bot. 2: 33 (1856). Vatica ovalis Korth., Verh. Nat. Gesch. Ned. Bezitt., Bot. 73 (Korthals 1841).
Ruborshorea section Pachycarpae (F.Heim) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77303213-1
Shorea section Pachycarpae F.Heim, Rech. Dipt. (1892: 44). Type species (designated here): R. macrophylla (de Vriese) P.S.Ashton & J.Heck.
Rubroshorea amplexicaulis (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298412-1
Shorea amplexicaulis P.S.Ashton, Gard. Bull. Singapore 19: 273 (1962).
Rubroshorea beccariana (Burck) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Malaysia, Sarawak, loc. incert., Beccari P.B. 1127 (BO) (Ashton 2004: 235).
http://www.ipni.org/urn:lsid:ipni.org:names:77298413-1
Shorea beccariana Burck, Ann. Jard. Bot. Buitenzorg 6: 213 (1887).
Rubroshorea macrophylla (de Vriese) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Indonesia, W Kalimantan, de Vriese s.n. (Rubroshorea H.L. no. 902146589), ‘Borneo’ (BO) (Ashton 2004: 287).
http://www.ipni.org/urn:lsid:ipni.org:names:77298414-1
Shorea macrophylla (de Vriese) P.S.Ashton, Gard. Bull. Singapore 20: 278 (1963). Basionym: Hopea macrophylla de Vriese, Minjak Tangkawang (1861: 28).
Rubroshorea mecistopteryx (Ridl.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298415-1
Shorea mecistopteryx Ridl., Bull. Misc. Inform., Kew 1925: 280 (Ridley 1925).
Rubroshorea pilosa (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298417-1
Shorea pilosa P.S.Ashton, Gard. Bull. Singapore 19: 304 (1962).
Rubroshorea pinanga (Scheff.) P.S.Ashton & J.Heck., comb. nov. Lectotype: Indonesia, Java, s.n., cult. in Hort. Bogor (hololectotype BO, isolectotypes L, U) (Ashton 2004: 313).
http://www.ipni.org/urn:lsid:ipni.org:names:77298418-1
Shorea pinanga Scheff., Tijdschr. Nederl. Ind. 31: 350 (Scheffer 1870).
Rubroshorea praestans (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298419-1
Shorea praestans P.S.Ashton, Gard. Bull. Singapore 22: 297 (1967).
Rubroshorea rotundifolia (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298420-1
Shorea rotundifolia P.S.Ashton, Gard. Bull. Singapore 22: 299 (1967).
Rubroshorea splendida (de Vriese) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298422-1
Shorea splendida (de Vriese) P.S.Ashton, Gard. Bull. Singapore 20: 279 (1963). Hopea splendida de Vriese, Minjak Tangkawang (1861: 28).
Rubroshorea stenoptera (Burck) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Indonesia, W Kalimantan, Sintang, de Vriese s.n. (L) (Ashton 2004: 338).
http://www.ipni.org/urn:lsid:ipni.org:names:77298423-1
Shorea stenoptera Burck, Meded. Lands Plantentuin 3: 11 (1886).
Rubroshorea woodii (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298424-1
Shorea woodii P.S.Ashton, Tree Fl. Sabah & Sarawak 5: 486 (2004).
Rubroshorea section Rubella (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298425-1
Shorea section Rubella P.S.Ashton, Gard. Bull. Singapore 20: 267 (1963). Type species: Rubroshorea rubella (P.S.Ashton) P.S.Ashton & J.Heck.
Rubroshorea albida (Symington) P.S.Ashton & J.Heck., comb. nov. Lectotype: Brunei, Kuala Belait, Zainal F.M.S. 30353 (hololectotype KEP, isolectotype L) (Ashton 2004: 221).
http://www.ipni.org/urn:lsid:ipni.org:names:77298426-1
Shorea albida Symington, Gard. Bull. Straits Settlem. 8: 283 (1935).
Rubroshorea dispar (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298427-1
Shorea dispar P.S.Ashton, Gard. Bull. Singapore 31: 45 (1978).
Rubroshorea elliptica (Burck) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298428-1
Shorea elliptica Burck, Ann. Jard. Bot. Buitenzorg 6: 215 (1887).
Rubroshorea negrosensis (Foxw.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298429-1
Shorea negrosensis Foxw., Philipp. J. Sci., C. 6: 274 (Foxworthy 1911).
Rubroshorea rubella (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298430-1
Shorea rubella P.S.Ashton, Gard. Bull. Singapore 19: 307 (1962).
Richetia F.Heim (1891: 975, nomen nudum; 1892: 50). Shorea section Richetioides F.Heim (1892: 48). Shorea subgen. Richetia (F.Heim) Meijer (1963: 322). Type species: Richetia coriacea F.Heim (1891: 975), designated here.
vernacular names. Meranti damar hitam (general); lun (Iban).
Richetia acuminatissima (Symington) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298433-1
Shorea acuminatissima Symington, Gard. Bull. Straits Settlem. 9: 340 (1938).
Richetia alutacea (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298435-1
Shorea alutacea P.S.Ashton, Gard. Bull. Singapore 22: 288 (1967).
Richetia angustifolia (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298436-1
Shorea angustifolia P.S.Ashton, Gard. Bull. Singapore 19: 277 (1962).
Richetia bakoensis (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298437-1
Shorea bakoensis P.S.Ashton, Gard. Bull. Singapore 22: 289 (1967).
Richetia balanocarpoides (Symington) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298438-1
Shorea balanocarpoides Symington, Gard. Bull. Straits Settlem. 9: 330 (1938).
Richetia blumutensis (Foxw.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298439-1
Shorea blumutensis Foxw., Malayan Forest Rec. 10: 236 (Foxworthy 1932).
Richetia chaiana (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298440-1
Shorea chaiana P.S.Ashton, Gard. Bull. Singapore 31: 42 (1978).
Richetia collaris (Slooten) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298441-1
Shorea collaris Slooten, Reinwardtia 3: 329 (1956).
Richetia conica (Slooten) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298442-1
Shorea conica Slooten, Reinwardtia 3: 336 (1956).
Richetia coriacea F.Heim (1891: 975). Shorea richetia Symington (1938: 330).
Richetia cuspidata (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298443-1
Shorea cuspidata P.S.Ashton, Gard. Bull. Singapore 22: 290 (1967).
Richetia faguetiana (F.Heim) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Malaysia, Sarawak, Kuching Distr., Matang, Beccari P.B. 2491 (P) (Ashton 2004: 253).
http://www.ipni.org/urn:lsid:ipni.org:names:77298444-1
Shorea faguetiana F.Heim, Bull. Mens. Soc. Linn. Paris 2: 975 (1891).
Richetia faguetioides (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298445-1
Shorea faguetioides P.S.Ashton, Gard. Bull. Singapore 19: 287 (1962).
Richetia gibbosa (Brandis) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298446-1
Shorea gibbosa Brandis, J. Linn. Soc. Bot. 31: 99 (1895).
Richetia hopeifolia (F.Heim) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298447-1
Shorea hopeifolia (F.Heim) Symington, Gard. Bull. Straits Settlem. 7: 150 (1933). Cotylelobium hopeifolium F.Heim, Bull. Mens. Soc. Linn. Paris 2: 971 (1891).
Richetia iliasii (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298449-1
Shorea iliasii P.S.Ashton, Gard. Bull. Singapore 22: 291 (1967).
Richetia induplicata (Slooten) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298450-1
Shorea induplicata Slooten, Reinwardtia 3: 327 (1956).
Richetia kuantanensis (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298451-1
Shorea kuantanensis P.S.Ashton, Gard. Bull. Singapore 31: 41 (1978).
Richetia kudatensis (G.H.S.Wood ex Meijer) P.S.Ashton & J.Heck., comb. nov. Hololectotype: Malaysia, Sabah, Kudat Distr., Tamalang Forest Reserve, G. H. S. Wood SAN 15363 (K) (Ashton 2004: 275).
http://www.ipni.org/urn:lsid:ipni.org:names:77298452-1
Shorea kudatensis G.H.S.Wood ex Meijer, Act. Bot. Neerl. 12: 346 (1963).
Richetia laxa (Slooten) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298453-1
Shorea laxa Slooten, Reinwardtia 3: 345 (1956).
Richetia longiflora (Brandis) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298454-1
Shorea longiflora (Brandis) Symington, Gard. Bull. Straits Settlem. 9: 330 (1938). Hopea longiflora Brandis, J. Linn. Soc., Bot. 31: 63 (1895).
Richetia longisperma (Roxb.) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298455-1
Shorea longisperma Roxb., Fl. Ind. Carey ed. 2, p. 618 (Roxburgh 1832).
Richetia macrobalanos (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298456-1
Shorea macrobalanos P.S.Ashton, Gard. Bull. Singapore 22: 292 (1967).
Richetia maxima (King) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298457-1
Shorea maxima (King) Symington, Gard. Bull. Straits Settlem. 9: 339 (1938). Balanocarpus maximus King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 62 (2): 133 (1893).
Richetia mujongensis (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298458-1
Shorea mujongensis P.S.Ashton, Gard. Bull. Singapore 22: 292 (1967).
Richetia multiflora (Burck) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298459-1
Shorea multiflora (Burck) Symington, Gard. Bull. Straits Settlem. 9: 330 (1938). Doona multiflora Burck, Ann. Jard. Bot. Buitenzorg 6: 231 (1887).
Richetia obovoidea (Slooten) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298460-1
Shorea obovoidea Slooten, Reinwardtia 3: 332 (1956).
Richetia patoiensis (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298462-1
Shorea patoiensis P.S.Ashton, Gard. Bull. Singapore 19: 302 (1962).
Richetia peltata (Symington) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298463-1
Shorea peltata Symington, J. Malayan Branch Roy. Asiat. Soc. 19: 158 (1941).
Richetia polyandra (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298464-1
Shorea polyandra P.S.Ashton, Gard. Bull. Singapore 22: 286 (1967).
Richetia subcylindrica (Slooten) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298465-1
Shorea subcylindrica Slooten, Reinwardtia 3: 331 (1956).
Richetia tenuiramulosa (P.S.Ashton) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298466-1
Shorea tenuiramulosa P.S.Ashton, Gard. Bull. Singapore 31: 42 (1978).
Richetia xanthophylla (Symington) P.S.Ashton & J.Heck., comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77298467-1
Shorea xanthophylla Symington, Gard. Bull. Straits Settlem. 9: 342 (1938).
Species incertae sedis:
Shorea furfuracea Miq. (Miquel 1861: 488).
The lofty emergent canopy of dipterocarps by which the equatorial lowland forests of Asia were so elegantly distinguished has now become restricted by conversion and unmanaged logging to national parks and nature reserves, most small, in Malaysia and Sri Lanka but for larger swathes in Sabah; also the Temburong District of the Sultanate of Brunei which has now lost its isolation, having been connected to the rest of its nation by a 30 km bridge, seemingly in part over the mangrove which is itself the world’s most species rich, ostensibly to enhance development. Little unlogged forest remains elsewhere in the Indonesian Sunda heartlands, or Philippines. Many species are now reduced to clusters too small to ensure long term sustainment, and some may now be extinct. The priority must be in conservation research policy planning and execution. In this, the dipterocarps, with their easily identifiable coriaceous rot-resistant leaves, and their high habitat specificity, provide a unique field tool for this critical endeavour. The re-establishment of clearly recognisable generic boundaries within the tribe Shoreae provides a botanical icon comparable to the great terrestrial vertebrates to this quest.
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Acknowledgements
We dedicate this contribution in recognition of Géma Maury-Lechon, whose brilliant insights and proposals which have so long been underestimated as pioneer in our current understanding of dipterocarpoid systematics. Rosabelle Samuel continues to be generous with her wide knowledge of plant molecular systematics. We thank Tim Utteridge, Helen Hartley (International Plant Names Index, ipni.org) and Ruth Linklater for their valuable contributions to the nomenclature and editing of this article.
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Ashton, P.S., Heckenhauer, J. Tribe Shoreae (Dipterocarpaceae subfamily Dipterocarpoideae) Finally Dissected. Kew Bull 77, 885–903 (2022). https://doi.org/10.1007/s12225-022-10057-w
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DOI: https://doi.org/10.1007/s12225-022-10057-w