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WEBBIA
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A synopsis of the native Combretaceae in the

Malay Peninsula
Citation: Ian M. Turner (2020) A synop-
sis of the native Combretaceae in the
Malay Peninsula. Webbia. Journal of
Plant Taxonomy and Geography 75(2): Ian M. Turner
263-280. doi: 10.36253/jopt-8891
Singapore Botanical Liaison Officer, Royal Botanic Gardens, Kew, Richmond, Surrey
Received: May 14, 2020 TW9 3AE, UK; Herbarium, Singapore Botanic Gardens, National Parks Board, 1 Cluny
Road, Singapore 259569
Accepted: July 6, 2020
E-mail: i.turner@kew.org
Published: November 18, 2020

Copyright: © 2020 Ian M. Turner. This is Abstract. A synopsis of the native species of Combretaceae in the Malay Peninsula
an open access, peer-reviewed arti- (Peninsular Thailand, Peninsular Malaysia and Singapore) is presented. A total of 29
cle published by Firenze University species in four genera (Combretum, Getonia, Lumnitzera and Terminalia) are recog-
Press (http://www.fupress.com/webbia) nised. Keys to genera and species are included with synonymy and typification. In total
and distributed under the terms of the 63 lectotypifications, three second-step lectotypifications and 11 neotypifications are
Creative Commons Attribution License, proposed in the paper.
which permits unrestricted use, distri-
bution, and reproduction in any medi-
Keywords: Calycopteris, Combretum, Getonia, Lumnitzera, Quisqualis, Terminalia.
um, provided the original author and
source are credited.

Data Availability Statement: All rel-

evant data are within the paper and its
Supporting Information files. INTRODUCTION

Competing Interests: The Author(s)

The Combretaceae include about 500 species of predominantly trees and
declare(s) no conflict of interest.
scandent shrubs found across most of the equatorial belt (Stace 2007). The
Editor: Peter C. Boyce family is well defined and its circumscription and higher-level classification
is uncontentious. It is numerically dominated by two large genera Com-
bretum and Terminalia which together make up about 97% of the extant spe-
cies (Stace 2010). Combretaceae species are found in tropical rain forests, dry
forests, savanna woodlands, savannas, littoral and mangrove vegetation. They
are less common in montane forests.
In South-East Asia the family is a member of the native flora and its spe-
cies are encountered in the lowlands. For the Malay Peninsula, the taxonomy
of the family was covered by Ridley (1922) and Exell (1954). Since then there
have been revisions of Combretum and Terminalia in Thailand (Nanakorn
1985, 1986), and of the arborescent taxa in Peninsular Malaysia (Kochum-
men 1972).
Recent activity in Combretaceae systematics has largely concerned
generic circumscription and the general trend has been for the subsumation
of small genera into the big two (Combretum and Terminalia). This can be
epitomised by a comparison between the system proposed by Exell and Stace
(1966) that recognised 20 genera and a recent summary (Byng 2014) where
this number had dropped to 10. These changes are less evident in South-East

Webbia. Journal of Plant Taxonomy and Geography 75(2): 263-280, 2020

ISSN 0083-7792 (print) | ISSN 2169-4060 (online) | DOI: 10.36253/jopt-8891
264 Ian M. Turner

Asia where the only major change is the inclusion of (=) Kleinia Crantz, Inst. Rei Herb. 2: 488. 1766, nom.
Quisqualis in Combretum (Maurin et al. 2020). illegit., non Kleinia Mill. (1754), nec Kleinia Jacq. (1760).
The purpose of this study is to provide a taxonomic Type: Kleinia quadricolor Crantz.
overview of the family as represented in the native flo-
ra of the Malay Peninsula including keys to genera and (=) Cacoucia Aubl., Hist. Pl. Guiane 1: 450, pl. 179. 1775.
species, a nomenclatural synopsis and details of typi- – Hambergera Scop., Introd. (1777) 106. 1777, nom. ille-
fication. The Malay Peninsula is taken here to include git. superfl. – Schousboea Willd., Sp. Pl. 2: 578. 1799,
Peninsular Thailand, Peninsular Malaysia and Singa- nom. illegit. superfl. – Combretum subgenus Cacoucia
pore. The research was conducted with reference to the (Aubl.) Exell & Stace, Bol. Soc. Brot., Sér. 2, 40: 10. 1966.
collections in herbarium of the Royal Botanic Gardens Type: Cacoucia coccinea Aubl.
Kew (K), some specimens on loan from SING, visits to
BM and access to various internet databases (e.g. JSTOR (=) Cristaria Sonn., Voy. Ind. Orient. 2: 247. 1782, nom.
Global Plants) and virtual herbaria (e.g. Berlin, Geneva, rejic. – Poivrea Comm. ex DC., Prodr. 3: 17. 1828, nom.
Paris). illegit. superfl. – Combretum sect. Poivrea G. Don, Gen.
Hist. 2: 665. 1832.
Type: Cristaria coccinea Sonn.
KEY TO GENERA OF COMBRETACEAE IN THE
MALAY PENINSULA (=) Sphalanthus Jack, Malayan Misc. 2(7): 55. 1822. –
Quisqualis sect. Sphalanthus (Jack) Exell, J. Bot. 69: 121.
1. Mangrove trees with reddish twigs, fleshy leaves with a 1931.
gland on the abaxial surface just below the apical notch; Type: Sphalanthus confertus Jack.
flowers and fruits bearing a pair of bracteoles adnate to
the hypanthium.......................................................Lumnitzera
(=) Gonocarpus Ham., Prodr. Pl. Ind. Occid. 39. 1825,
Trees and scandent shrubs of seashores and inland for- nom. illegit., non Gonocarpus Thunb. (1783).
ests, not true mangroves, twigs green or brown, not nota- Type: Gonocarpus jacquinii Ham., nom. illegit. superfl.
bly reddish, leaves chartaceous to coriaceous, not fleshy,
lacking an apical gland; flowers and fruits without adnate
bracteoles................................................................................... 2
(=) Forsgardia Vell., Fl. Flumin. 4: 152, t. 13. 1829.
Type: Forsgardia laevis Vell.
2. Scandent shubs (except C. quadrangulare); flowers pet-
alous......................................................................... Combretum (=) Chrysostachys Pohl, Pl. Bras. Icon. Descr. 2: 66, t. 143.
1830.
Erect trees and treelets; flowers apetalous............................ 3
Type: Chrysostachys ovatifolia Pohl.
3. Sepals accrescent in fruit..............................................Getonia
(=) Sheadendron G.Bertol., Mem. Reale Accad. Sci. Ist.
Sepals not persisting in fruit...................................Terminalia
Bologna 2: 574. 1850.
Type: Sheadendron butyrosum G.Bertol.

1. Combretum Loefl., Iter Hispan. 308. 1758, nom. cons. (=) Calopyxis Tul., Ann. Sci. Nat., Bot., sér. 4, 6: 86. 1856.
– Combretum sect. Calopyxis (Tul.) Jongkind, Bull. Mus.
Type: Combretum fruticosum (Loefl.) Stuntz Natl. Hist. Nat., B, Adansonia Sér. 4, 17(3-4): 193. 1995.
Type: Calopyxis sphaeroides Tul. (lectotype selected by
(≡) Aetia Adans., Fam. Pl. 2: 84, 513. 1763, nom. illegit. Stace (2010: 60).
superfl.
(=) Embryogonia Blume, Mus. Bot. 2: 122. 1856.
(=) Grislea L., Sp. Pl. 1: 348. 1753, nom. rejic. Type: Embryogonia acuminata (Roxb.) Blume (lectotype
Type: Grislea secunda L. selected by Stace (2010: 60).

(=) Quisqualis L., Sp. Pl. (ed. 2) 1: 556. 1762. – Udani (=) Bureava Baill., Adansonia 1: 71. 1860. 71, nom. rejic.
Adans., Fam. Pl. 2: (22). 1763, nom. illegit., superfl.. Type: Bureava crotonoides Baill.
– Combretum sect. Quisqualis (L.) Stace, Fl. Neotrop.
Monogr. 107: 159. 2010. (=) Thiloa Eichler, Flora 49: 149. 1866. – Combretum sect.
Type: Quisqualis indica L. Thiloa (Eichler) Stace, Fl. Ecuador 81: 13. 2007.
A synopsis of the native Combretaceae in the Malay Peninsula 265

Type: Thiloa glaucocarpa (Mart.) Eichler (lectotype 6. Flowers 5-merous; fruits with 5 wings or 5 longitudinal
selected by Stace, Fl. Ecuador 81: 11. 2007). ridges......................................................................................... 7

Flowers 4-merous; fruits with 4 wings or 4 longitudinal

(=) Campylogyne Welw. ex Hemsl., Hooker’s Icon. Pl. 26: ridges............................................................... 1.8 C. nigrescens
t. 2550. 1889. – Quisqualis sect. Campylogyne (Welw. ex
Hemsl.) Exell, J. Bot. 69: 120. 1931. 7. Leaves opposite with hairy domatia in axils of lateral
Type: Campylogyne exannulata (O.Hoffm.) Hemsl. nerves below; fruits with 5 wings ............ 1.12 C. roxburghii

Leaves often ternate without domatia; fruits with 5 longi-

(=) Campylochiton Welw. ex Hiern, Cat. Afr. Pl. 2: 353. tudinal ridges...............................................1.15 C. trifoliatum
1901.
Type: Campylochiton platypterus (Welw.) Hiern. 8. Trees, young twigs sharply 4-angled.. 1.11 C. quadrangulare

Climbers or scandent shrubs, young twigs more or less

(=) Meiostemon Exell & Stace, Bol. Soc. Brot., sér. 2, 40: terete.......................................................................................... 9
18. 1966.
Type: Meiostemon tetrandrus (Exell) Exell & Stace 9. Lower lamina with a dense covering of predominantly
whitish scales, tertiary venation obscure............................ 10
Notes Lower lamina with scales absent or with sparse covering of
The lectotypifications of Calopyxis and Thiloa were first predominantly reddish scales, tertiary venation visible.... 11
proposed by Exell (1931) but the choices were clearly 10. Inflorescences subcapitate..........................................................
mechanical, so the later designations by Stace (2007, ..........................................1.10a C. punctatum var. punctatum
2010) are accepted here.
Inflorescences elongate...............................................................
........................................1.10b C. punctatum var. squamosum
Key to Combretum taxa 11. Petals inconspicuous, to 1.5 mm long; fruits with 4 longi-
tudinal blunt-topped ridges..................... 1.1 C. acuminatum
1. Floral bracts persisting to anthesis and beyond; upper hyp-
anthium tubular; petals becoming pink or red.................... 2 Petals conspicuous at anthesis, more than 1.5 mm long;
fruits with 4 distinct wings or if with longitudinal ridges
Floral bracts generally dropping before anthesis; upper then ridges sharp-edged (C. tetralophum).......................... 12
hypanthium salveriform to infundibuliform; petals white
or green..................................................................................... 5 12. Sepals 1.5 mm or longer and petals exceeding length of
sepals; fruit with 4 longitudinal sharp-edged ridges ............
2. Upper hypanthium 5-8 cm long...................... 1.5 C. indicum .................................................................... 1.14 C. tetralophum
Upper hypanthium 1-2 cm long............................................. 3 Sepals to 1 mm long, or if longer then petals shorter than
sepals; fruits with 4 papery wings........................................ 13
3. Sepals recurved at anthesis, with filiform tips 1-2 mm long.
1.4 C. densiflorum 13. Inflorescences capitate (also detectable as knobbly ends to
infructescences 3-4 mm long).................. 1.13 C. sundaicum
Sepals not recurved at anthesis, filiform tips to 1 mm long..
.................................................................................................... 4 Inflorescences elongate.......................................................... 14
4. Leaves with tertiary venation visible adaxially; floral bracts 14. Flowers and inflorescence rhachis densely pale erect hairy;
to 2 mm long with 1 central longitudinal nerve..................... fruits sparsely hairy, scales absent................ 1.7 C. latifolium
............................................................... 1.6 C. langkawiense
Flowers and inflorescence rhachis densely scaly; fruits gla-
Leaves with tertiary venation obscure adaxially; flo- brous but with scales present................................................ 15
ral bracts 1 cm long or more with at least 5 longitudinal
nerves...............................................................1.2 C. caudatum 15. Twigs drying reddish brown; leaves with red-brown scales
conspicuous on lower lamina; flowers generally more than
5. Indumentum including glandular hairs but not scales; 4 mm long; fruits generally more than 3 cm across...............
flowers 5-merous; fruits with 5 wings or 5 longitudinal .............................................................................1.3 C. chinense
ridges (except C. nigrescens which has 4-merous flowers
and 4-winged fruits)................................................................ 6 Twigs drying whitish; leaves without conspicuous red-
brown scales on lower lamina; flowers generally less than 4
Indumentum including scales but not glandular hairs; mm long; fruits to 2.5 cm across.............1.9 C. porterianum
flowers 4-merous; fruits with 4 wings or 4 longitudinal
ridges......................................................................................... 8
266 Ian M. Turner

1.1 Combretum acuminatum Roxb., Fl. Ind. 2: 228. 1832 1.2 Combretum caudatum (Craib) O.Maurin & Boatw.,
Phytotaxa 451: 232. 2020
Type: [unpublished illustration] Icones Roxburghianae
2225 (K, lectotype selected by du Puy and Telford (1993: Bas.: Quisqualis caudata Craib, Bull. Misc. Inform. Kew
245). 1930: 164. 1930.
Type: Thailand, Ranawng, Kao Pawta, Chongdong, 21
(≡) Embryogonia acuminata (Roxb.) Blume, Mus. Bot. 2: January 1929, A.F.G. Kerr 16797 (K barcode K000786136,
123. 1856. lectotype selected by Maurin et al. (2020: 232); isolec-
totypes ABD, BK barcode BK257801, BM barcode
(=) Combretum costatum Roxb., Fl. Ind. 2: 227. 1832. BM000947035).
Type: [unpublished illustration] Icones Roxburghianae
2478 (2476 on drawing) (lectotype CAL, selected by Notes
Gangopadhyay and Chakrabarty (1997: 299). Maurin et al. (2010) included this species in their anal-
ysis that showed it to be sister to the other ‘Quisqualis’
(=) Combretum neurophyllum Miq., Fl. Ned. Ind. 1(1): species sampled and to be part of a subclade of Com-
608. 1856. bretum subgenus Cacoucia.
Type: Java, Tjitjiringin, J. Hasskarl s.n. (L barcode
L0844403, lectotype designated here).
1.3 Combretum chinense Roxb. ex G.Don, Trans. Linn.
(=) Combretum sarcopterum Thwaites, Enum. Pl. Zeyl. Soc. 15: 417, 432-433. 1827
415. 1864.
Type: Sri Lanka, Reigam Corle, Thwaites CP 3715 (CAL, Type: Calcutta Botanical Garden, W. Roxburgh s.n. (G
lectotype selected by Gangopadhyay and Chakrabarty barcode G00177910, lectotype selected by Nanakorn
(1997: 299). (1986: 174).

(=) Combretum stenopetalum Van Heurck & Müll.Arg., (=) Combretum chinense Roxb., Fl. Ind. 2: 230. 1832, as
Observ. Bot. Descript. Pl. Nov. Herb. Van Heurckiani 2: ‘chinensis’, nom. illegit., non C. chinense Roxb. ex G.Don
225. 1871. (1827).
Type: W. Griffith (syntypes BR, G). Type: India, Calcutta Botanical Garden, W. Roxburgh
s.n. (G barcode G00177910, lectotype selected by Nana-
(=) Combretum vanheurckii Müll.Arg., Observ. Bot. korn (1986: 174).
Descript. Pl. Nov. Herb. Van Heurckiani 2: 227. 1871.
Type: W. Griffith 725 (G barcode G00177909, lectotype (=) Combretum chinense Roxb. var. pubescens King, J.
selected by Gangopadhyay and Chakrabarty (1997: 299); Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: 339. 1897.
isolectotype BR). Type: Peninsular Malaysia, Perak, Sungai Ryah, October
1880, Dr King’s Collector 889 (K barcode K001129737,
(=) Terminalia borneensis Ridl., Bull. Misc. Inform. Kew lectotype designated here).
1938: 283. 1938, nom. illegit., non T. borneensis Slooten (1919).
Type: Borneo, Bangarmassing, J. Motley 380 (K barcode (=) Combretum annulatum Craib, Bull. Misc. Inform.
K000786585, lectotype designated here). Kew 1929: 113. 1929.
Type: Thailand, Chumpawn, Ta Ngaw, 17 January 1917,
Notes
A.F.G. Kerr 11488 (K barcode K000786590, lectotype
Nanakorn (1986: 196) referred to a specimen in G-DC selected by Gangopadhyay and Chakrabarty 1997: 314;
from the distribution of the East India Company Her- isolectotypes ABD, BK barcode BK257787, BM barcode
barium under number 4014D as type of Combretum BM000947049, TCD barcode TCD0015621).
acuminatum Roxb. Some error crept in here – Nana-
korn’s proposed type is a specimen of Anogeissus acumi- (=) Combretum annulatum Craub var. orbiculare Craib,
nata (Roxb. ex DC.) Wall. ex Guillem. & Perr. rather Fl. Siam. Enum. 1(4): 614. 1931.
than Combretum, and is certainly not original material Type: Thailand, Puket, Ranawng, Kampuam, 29 Janu-
for Roxburgh’s name in Combretum. Therefore, the later ary 1929, A.F.G. Kerr 16881 (K (2 sheets) barcodes
typification by du Puy and Telford to a Roxburgh draw- K000786589, K001129727, lectotype designated here;
ing is accepted here. isolectotypes ABD, BM barcode BM000947050).
A synopsis of the native Combretaceae in the Malay Peninsula 267

Notes 1.5 Combretum indicum (L.) DeFilipps, Useful Pl. Dom-

inica 277. 1998
There has been considerable confusion about the
name Combretum chinense. It was first published by
Bas. Quisqualis indica L., Sp. Pl., ed. 2. 1: 556. 1762.
William Roxburgh in his Hortus Bengalensis (Rox-
Type: [published illustration] “Quis qualis” in Rumphi-
burgh 1814) but was a nomen nudum in the absence of
us, Herb. Amboin. 5: t. 38. 1747 (lectotype selected by
any description or reference to a published description.
Merrill (1917: 390).
As with many Roxburgh names, it was first validated
by another author before it appeared with a description
(=) Kleinia quadricolor Crantz, Inst. Rei Herb. 2: 488.
in one of the publications of Roxburgh’s works after his
1766.
death. George Don described Combretum chinense Roxb.
Type: [published illustration] “Quis qualis” in Rumphi-
in 1827 (Don 1827), before Roxburgh’s own description
us, Herb. Amboin. 5: t. 38. 1747 (lectotype selected by
was finally published in 1832 (Roxburgh 1832). Don
Maurin et al. (2020: 233).
referred to a Roxburgh specimen in the herbarium of
A.B. Lambert. There is a relevant Roxburgh specimen
(=) Quisqualis glabra Burm.f., Fl. Ind.: 104, t. 28, f. 2.
now in G that appears to be annotated by Lambert. Exell
1768.
(1954) argued that this was not Don’s type as it did not
Type: Java, 1758, Anon. s.n. (G-PREL barcode
match all the details in Don’s description. He therefore
G00805923, lectotype selected by Maurin et al. (2020:
treated C. chinense Roxb. ex G.Don as a confused name
233).
and referred the Malesian material to Combretum yun-
nanese Exell. Nanakorn (1985) disagreed, and effectively
(=) Quisqualis pubescens Burm.f., Fl. Ind.: 104. 1768.
lectotypified both the Don and Roxburgh names to the
Type: [published illustration] “Quis qualis” in Rumphi-
G specimen. I find that the Malay Peninsula material is
us, Herb. Amboin. 5: t. 38. 1747 (lectotype selected by
a reasonable match to the Roxburgh lectotype (image
Merrill (1917: 390).
available online) and therefore employ C. chinense Roxb.
ex G.Don for the specimens.
(=) Quisqualis ebracteata P.Beauv., Fl. Oware 1: 56, 57, t.
35. 1806.
Type: Oware or Bénin, Palisot de Beauvois s.n. (G bar-
1.4 Combretum densif lorum (Wall. ex Planch.)
code G00446747 [2 sheets], lectotype selected by Maurin
I.M.Turner, Phytotaxa 451: 232. 2020
et al. (2020: 233).
Bas.: Quisqualis densiflora Wall. ex Planch., Fl. Serres 6:
(=) Quisqualis obovata Schumach. & Thonn., Beskr.
348. 1851.
Guin. Pl. 218-219. 1827.
Type: Peninsular Malaysia, Penang, G. Porter s.n. [EIC
Type: Guinea, P. Thonning 315 (C barcode C10004457,
4011] (K-W barcode K001117910, lectotype selected
lectotype selected by Hepper (1976: 40).
by Maurin et al., 2020: 232; isolectotypes BR barcode
BR0000005639281, GZU barcode GZU000273166, K
(=) Quisqualis indica Lour., Fl. Cochinchin. 274. 1790,
barcode K000786138, M barcode M0146742, P (×3) bar-
nom. illegit., non Q. indica L. (1762).
codes P01901448, P01901447, P01901446, PH barcode
– Quisqualis loureiroi G.Don, Gen. Hist. 2: 667. 1832, as
PH00023582
‘Loureiri’.
Type: J. de Loureiro s.n. (BM barcode BM000947033, lec-
(=) Sphalanthus confertus Jack, Malayan Misc. 2(7):
totype selected at the first step by Exell 1931: 124, and at
55–56. 1822. – Quisqualis conferta (Jack) Exell, J. Bot. 69:
the second step by Maurin et al. (2020: 233).
122. 1931.
Type: W. Jack s.n. (G barcode G00446750, lectotype
(=) Quisqualis villosa Roxb., Fl. Ind. 2: 426. 1832. – Quis-
selected by Exell (1931: 122).
qualis indica var. villosa (Roxb.) Kurz, Forest Fl. Burma
Notes 1: 467. 1877.
Ty pe: India, W. Roxburgh s.n. (BR barcode
The oldest name for this species is Sphalanthus BR000000583537, lectotype selected by Maurin et al.
confertus Jack, but Combretum confertum (Benth.) (2020: 233).
M.A.Lawson, the accepted name of an African species,
blocks the transfer of Jack’s name to Combretum.
268 Ian M. Turner

(=) Quisqualis sinensis Lindl., Edwards’s Bot. Reg. 30: t. (=) Combretum extensum Roxb., Fl. Ind. 2: 229. 1832,
15. 1844. nom. illegit., non C. extensum Roxb. ex G.Don (1827).
Type: [published illustration] ‘Quisqualis sinensis’ in Type: [unpublished illustration] Icones Roxburghianae
Lindley, Edwards’s Bot. Reg. 30: t. 15. 1844 (lectotype No. 1830 (K, lectotype designated here).
selected by Maurin et al. (2020: 233).
(=) Combretum macrophyllum Roxb., Fl. Ind. 2: 231.
(=) Quisqualis spinosa Blanco, Fl. Filip., ed. 2: 254, 1845. 1832.
Type: Philippines, Palawan, Taytay, 9 May 1913, Mer- Type: not traced.
rill: Species Blanconae No. 582 (US barcode US00623716,
neotype selected by Maurin et al. (2020: 233); isoneo- (=) Combretum rotundifolium Roxb., Fl. Ind. 2: 226-227.
types BM, K). 1832, nom. illegit., non. C. rotundifolium Rich. (1792). –
Combretum cyclophyllum Steud., Nomencl. Bot., ed. 2, 1:
(=) Quisqualis longiflora C.Presl, Epimel. Bot. 216. 1851. 400. 1840.
Type: Martaban Helfer (not traced). Type: W. Roxburgh s.n. (BM barcode BM01382528, lecto-
type designated here).
(=) Quisqualis grandiflora Miq., J. Bot. Neerl. 1: 119.
1861. (=) Combretum wightianum Wall. ex Wight & Arn.,
Type: China austr., R. Krone 146 (U barcode U.1217341, Prodr. Fl. Ind. Orient. 316. 1834.
lectotype selected by Maurin et al. (2020: 233). Type: EIC 4007 (CAL, lectotype selected by Gangopad-
hyay and Chakrabarty (1997: 299).
(=) Quisqualis indica var. oxypetala Kurz, Forest Fl.
Burma 1: 467. 1877. (=) Combretum formosum Griff., Not. Pl. Asiat. 4: 682.
Type: Burma, Ava, Kakhyen Hills (not traced). 1854, nom. illegit., non C. formosum G.Don (1827).
Type: Burma, Mergue, in sylvis Kulweng, Dec 1834 (not
traced).
1.6 Combretum langkawiense O.Maurin & Christenh.,
Phytotaxa 451: 233. 2020 (=) Combretum horsfieldii Miq., Fl. Ned. Ind. 1(1): 609.
1856.
Replaced synonym: Quisqualis densiflora var. parvifolia Type: Java, Horsfield s.n. (holotype K barcode
Ridl., Fl. Malay Penins. 1: 711. 1922. K0011297290).
Type: Peninsular Malaysia, Langkawi, Dayang Bunt-
ing, November 1916, H.C. Robinson 6198 (K barcode (=) Combretum latifolium var. ellipticum Miq., Fl. Ned.
K000786137, lectotype selected by Maurin et al. (2020: Ind. 1(1): 609. 1856, as ‘elliptica’.
233); isolectotype BM barcode BM000947036 (fragment Type: Java, Horsfield s.n. (holotype K barcode
ex K). K001129730).

(≡) Quisqualis parvifolia (Ridl.) Exell, J. Bot. 69: 123. (=) Combretum leucanthum Van Heurck & Müll.Arg.,
1931. Observ. Bot. Descript. Pl. Nov. Herb. Van Heurckiani 2:
240. 1871.
Type: Ceylon, Thwaites C.P. 1601 (syntypes BR, G).
1.7 Combretum latifolium Blume, Bijdr. (13): 641. 1826
(=) Combretum platyphyllum Van Heurck & Müll.Arg.,
Type: C.L. Blume s.n. (NY barcode NY00245959, lecto- Observ. Bot. Descript. Pl. Nov. Herb. Van Heurckiani 2:
type designated at the first step by Gangopadhyay and 242. 1871.
Chakrabarty (1997: 311), and at the second step here). Type: EIC 3995 (lectotype G, selected by Gangopadhyay
(≡) Embryogonia latifolia (Blume) Blume, Mus. Bot. 2: and Chakrabarty (1997: 299).
122. 1856.
Notes
(=) Combretum extensum Roxb. ex G.Don, Trans. Linn. In the protologue for Combretum latifolium Blume
Soc. 15: 422. 1827. gave the locality ‘in sylvis montosis Salak’. I have failed
Type: India orientalis, W. Roxburgh s.n. (G barcode to locate any Blume specimens that mention Mt Salak,
G00446751, lectotype designated here). so follow Gangopadhyay and Chakrabarty (1997: 311)
A synopsis of the native Combretaceae in the Malay Peninsula 269

who selected unlocalised Blume material in NY as the sheet no. S09-25745, SINGx2 barcode SING0055423, U
type. As there are actually two specimens present in NY, barcode U0226823, US(2 sheets) barcodes US00810961
a second-stage lectotypification is made here. Gangopad- US00810960, Z barcode Z-000002869).
hyay and Chakrabarty (1997: 311) designated a Horsfield
specimen in CAL as lectotype for C. horsfieldii, but the (=) Combretum elmeri Merr. var. glabrescens Merr., Univ.
set of Horsfield specimens seen by Miquel when work- Calif. Publ. Bot. 15: 215. 1929.
ing on Flora van Nederlandsch Indië are in Kew and Type: Borneo, British North Borneo [Sabah], Elphin-
the single specimen present is therefore the holotype of stone Province, Tawao [Tawau], October 1922-March
Miquel’s name. 1923, A.D.E. Elmer 21561 (UC barcode UC290635,
lectotype designated here; isolectotypes A barcode
A00068598, BISH barcode BISH1001095, BR barcode
1.8 Combretum nigrescens King, J. Asiat. Soc. Bengal, BR0000005629206, C barcode C10009531, CAS bar-
Pt. 2, Nat. Hist. 66: 340. 1897 code CAS0032976, GH barcode GH00068597, HBG
barcode HBG514718, K barcode K000786581, L bar-
Type: Peninsular Malaysia, Perak, October 1882, Dr code L0009753, M barcode M0146600, MICH barcode
King’s Collector 3469 (K barcode K000786600, lectotype MICH1192150, MIN barcode MIN1002830, NY bar-
designated here; isolectotype BM). code NY00245958, S sheet no. S09-25747, SING bar-
code SING0055422, U barcode U0226815, US barcode
(=) Combretum kunstleri King, J. Asiat. Soc. Bengal, Pt. US00117555. Z barcode Z-000002870).
2, Nat. Hist. 66: 340. 1897. – Combretum nigrescens var.
kunstleri (King) Ridl., Fl. Malay. Penins. 1: 710. 1922. (=) Combretum kunstleri var. rectinerve Craib, Fl. Siam.
Type: Peninsular Malaysia, Perak, Larut, October 1884, Enum. 1: 617. 1931.
Dr King’s Collector 6664 (K barcode K000786597, lecto- Type: Thailand, Pattani, Bukit, A.F.G. Kerr 7126 (K bar-
type designated here; isolectotype BM). code K000786588, lectotype designated here; isolec-
totypes ABD, BK barcode BK257790, BM barcode
(=) Combretum scortechinii King, J. Asiat. Soc. Bengal, BM000947045).
Pt. 2, Nat. Hist. 66: 341. 1897.
Type: Peninsular Malaysia, Perak, Goping, April 1885,
Fr. B. Scortechini 1942 (K barcode K000786602, lecto- 1.9 Combretum porterianum (C.B.Clarke) Wall. ex
type designated here). Craib., Fl. Siam. Enum. 1(4): 618. 1931

(=) Combretum glandulosum Slooten, Bijdr. Combret. Bas.: Combretum chinense var. porterianum C.B.Clarke
Flacourt. Ned.-Ind. 39. 1919, nom. illegit., non C. glan- in J.D.Hooker, Fl. Brit. India 2: 457. 1878.
dulosum F.Hoffm. (1889). – Combretum adenophorum
Slooten, Bull. Jard. Bot. Buitenzorg, sér. 3, 6: 56. 1924. (≡) Combretum wallichii DC. var. porterianum
Type: Sumatra, 1880 H. Forbes 3100 (L barcode (C.B.Clarke) M.Gangop. & Chakrab., J. Econ. Taxon.
L.2500736, lectotype designated here; isolectotypes BM Bot. 17: 681. 1993.
barcode BM000947044, L barcode L.2500737). Type: EIC 4000 (CAL, lectotype selected by Gango-
padhyay and Chakrabarty 1997: 324; isolectotypes K
(=) Combretum elmeri Merr., Univ. Calif. Publ. Bot. 15: barcode K000786603, K-W barcode K001117875, M
214. 1929. M0146610).
Type: Borneo, British North Borneo [Sabah], Elphinstone
Province, Tawao [Tawau], October 1922-March 1923,
A.D.E. Elmer 21440 (UC barcode UC290634, lectotype 1.10 Combretum punctatum Blume, Bijdr. 640. 1826
designated here; isolectotypes A barcode A00068596,
BISH barcode BISH1001094, BM(2 sheets), BR barcode Type: Java, G. Parang, C.L. Blume s.n. (L barcode
BR0000005629534, C barcode C10009529, CAS bar- L0844301, lectotype designated here).
code CAS0033025, F barcode F0054644F, GH barcode
GH00068595, HBG barcode HBG514719, K barcode Notes
K000786582, L barcode L0009752, M barcode M0146601, In the protologue, Blume referred to ‘in montosis
MO barcode MO-176152, MSC barcode MSC0091506, Parang, et ad pedem montium Salak, Tjerimai etc’. Gan-
NY barcode NY00245957, PH barcode PH00006505, S gopadhyay and Chakrabarty (1997) designated a Blume
270 Ian M. Turner

specimen in NY as the lectotype, but it is not annotated K000786592, K000786593, lectotype designated here;
with any of the locations mentioned by Blume. There are isolectotypes BK barcode BK257797, BM barcode
specimens with these localities stated in L. Therefore, a BM000947046).
lectotype is here designated from one of the specimens
in L to supersede the designation of Gangopadhyay and (=) Combretum squamosum var. dissitum Craib, Flor.
Chakrabarty. Siam. Enumerat. 1: 620. 1931.
Type: Thailand, Maharat, Lampang, Me Tia, 28 April
1923, Winit 860 (K barcode K001129728, lectotype des-
1.10a Combretum punctatum var. punctatum ignated here).

1.10b Combretum punctatum var. squamosum (Roxb. Notes

ex G.Don) M.Gangop. & Chakrab., J. Econ. Taxon. Bot. I prefer to recognise the infraspecific taxa of Com-
17: 680. 1993 bretum punctatum at the rank of variety following Gan-
gopadhyay and Chakrabarty (1997) and Chen and Tur-
Bas.: Combretum squamosum Roxb. ex G.Don, Trans. land (2007), rather than subspecies (Exell 1954).
Linn. Soc. 15: 419, 438. 1827.
Type: Roxburgh in herb. Linn. Soc. (not traced).
1.11 Combretum quadrangulare Kurz, J. Asiat. Soc.
(≡) Poivrea squamosa (Roxb. ex G.Don) Walp., Repert. Bengal, Pt. 2, Nat. Hist. 43(3): 188. 1874
Bot. Syst. 2: 64. 1843.
Type: Burma, Tenasserim, Helfer s.n. [Kew distrib. no.
(≡) Combretum punctatum subsp. squamosum (Roxb. ex 2181] (lectotype CAL, selected by Gangopadhyay and
G.Don) Exell, Fl. Males. ser. 1 4: 539. 1954. Chakrabarty (1997: 317).

(=) Combretum squamosum Roxb., Fl. Ind. 2: 231. 1832, (=) Combretum quadrangulare var. lanceolatum Gagnep.,
nom. illegit., non C. squamosum Roxb. ex G.Don. Fl. Gen. 2: 746. 1920.
Type: Cambodia, Prèy-kdey, Kg Speu, 25 April 1909,
(=) Combretum distillatorium Blanco, Fl. Filip. 295. 1837. Magnen et al. s.n. [Châtillon leg.] (P barcode P05046162,
Type: Philippines, Luzon, Bulacan, Maon River, Mer- lectotype designated here).
rill: Species Blancoanae 847 [M. Ramos leg.] (US barcode
US00623715, neotype designated here; isoneotype BM).
1.12 Combretum roxburghii Spreng., Syst. Veg. 2: 231.
(=) Combretum lepidotum C.Presl, Abh. Königl. Böhm. 1825
Ges. Wiss., ser. 5, 3: 572. 1845
Type: ?Burma, J.W. Helfer 116 (PR sheet no. 615825, lec- Replaced synonym: Combretum decandrum Roxb.,
totype designated here). Pl. Coromandel 1: 43, t. 59. 1796, nom. illegit., non C.
decandrum Jacq. (1760).
(=) Combretum squamosum var. luzonicum C.Presl, Abh. Type: W. Roxburgh s.n. [EIC 4009[A]] (K-W barcode
Königl. Böhm. Ges. Wiss., ser. 5, 6: 576. 1851. K001117895, lectotype designated here).
Type: Philippines, H. Cuming 1122 (syntype BM).
(≡) Poivrea roxburghii (Spreng.) DC., Prodr. 3: 18. 1828.
(=) Combretum squamosum var. aureum C.B. Clarke in
Hooker, Fl. Brit. Ind. 2: 456. 1878. (≡) Pentaptera roxburghii (Spreng.) Tul., Ann. Sci. Nat.,
Type: Burma, Prome Hills, 1827, N. Wallich 252 [EIC Sér. 4 6: 84. 1856.
3988[A] p.p.] (K barcode K001117838l, lectotype desig-
nated at the first step by Gangopadhyay and Chakrabar- (=) Combretum klossii Ridl., J. Fed. Malay States Mus. 10:
ty (1997: 314) and at the second step here). 90. 1920.
Type: Thailand, Koh Pipidon [Ko Phi Phi Don], C.B.
(=) Combretum foliatum Craib, Bull. Misc. Inform. Kew Kloss 6556 (K barcode K000786591, lectotype designated
1930: 163. 1930. here; isolectotype SING barcode SING0055423).
Type: Thailand, Puket, Pang-nga, Nop Pring, 25 Feb-
ruary 1929, A.F.G. Kerr 17239 (K (2 sheets) barcodes
A synopsis of the native Combretaceae in the Malay Peninsula 271

Notes 1.15 Combretum trifoliatum Vent., Choix Pl. t. 58. 1808

Roxburgh’s name was an illegitimate later homonym
Type: Java, La Haye s.n. (G barcode G00177911, lectotype
when it was published. Gangopadhyay and Chakrabarty
selected by Lecompte (1969: 40).
(1997) took up Combretum album Pers. as the correct name.
This appears in parentheses after C. decandrum in Persoon’s
(≡) Cacoucia trifoliata (Vent.) DC., Prodr. 3: 22. 1828.
Synopsis plantarum (Persoon 1805). The nature of Persoon’s
parenthetical names is controversial. In the Preface (p.
(=) Combretum lucidum Blume, Bijdr. Fl. Ned. Ind. 641.
ix), Persoon wrote ‘E contrario in parenthesi nomen quasi
1826. – Cacoucia lucida (Blume) Hassk., Flora 27: 607.
secundarium passim addidi, quo idea plantae forte distinc-
1844. – Embryogonia lucida (Blume) Blume, Mus. Bot.
tior concipi possit.’ It is difficult to equate Persoon’s ‘quasi-
Lugd.-Bat. 2: 122. 1856.
secondary’ names with replacement or alternative names as
Type: Java, Pamanoekang, C.L. Blume 1543 (L barcode
currently employed, so Combretum album Pers. is not con-
L0844262, lectotype designated here).
sidered validly published. Combretum roxburghii is therefore
the earliest replacement name for C. decandrum Roxb.
(=) Terminalia lancifolia Griff., Not. Pl. Asiat. 4: 685.
1854.
Type: Moulmein ad marginem sylvarum (not traced).
1.13 Combretum sundaicum Miq., Fl. Ned. Ind., Eerste
Bijv. 327. 1861
2. Getonia Roxb., Pl. Coromandel 1: 61, t. 87. 1798.
Type: Sumatra, prov. Lampong, Tiga-nennin, HB 4330
(U barcode U0001193, lectotype designated here).
Type: Getonia floribunda Roxb.
(=) Combretum oliviforme A.C.Chao, Acta Phytotax. Sin.
(≡) Calycopteris Poir., Encycl., Suppl. 2: 41. 1811, nom.
7: 244. 1958, ‘olivaeforme’.
illegit. superfl.
Type: China, Hainan, Bak Sa, 23 July 1936, S.K. Lau
27571 (holotype IBSC barcode IBSC0004026; isotypes Notes
A barcode A00244267, IBK barcode IBK00127780, NAS
barcode NAS00047141, PE barcode PE00994471. Chen and Turland (2007: 315) explained why Caly-
copteris Lam. was not validly published and hence Geto-
(=) Combretum oliviforme A.C.Chao var. yaxianense nia is the correct name for the genus.
Y.R.Ling, Acta Phytotax. Sin. 19: 388. 1981.
Type: China, Hainan, [Ya Xian], 11 August 1933, C.
Wang 33616 (holotype IBSC barcode IBSC0004027; iso- 2.1 Getonia floribunda Roxb., Pl. Coromandel 1: 61, t.
type A barcode A00068593). 87. 1798

Type: [published illustration] Roxburgh, Pl. Coromandel

1.14 Combretum tetralophum C.B.Clarke in Hooker, Fl. 1: t. 87. 1798 (lectotype designated here).
Brit. India 2: 454. 1878
(≡) Calycopteris floribunda (Roxb.) Lam. ex Poir., Encycl.
Type: Peninsular Malaysia, Malacca, 1845, W. Grif- Suppl. 2: 41. 1811.
fith s.n. (K barcode K000786605, lectotype selected by
Lecompte (1969: 49). (=) Getonia nitida Roth, Nov. Pl. Sp. 217. 1821. – Geto-
nia nutans Roxb. ex DC., Prodr. 3: 15. 1828, nom. illegit.
(≡) Combretum tetragonocarpum var. tetralophum superfl. – Calycopteris nutans Kurz, J. Asiat. Soc. Bengal,
(C.B.Clarke) M.Gangop. & Chakrab., J. Econ. Taxon. Pt. 2, Nat. Hist. 46(2): 59. 1877, nom. illegit. superfl.
Bot. 17: 697. 1993. Type: B. Heyne s.n. [EIC 4013A] (K-W barcode
K001117915, neotype designated here, excluding material
(=) Combretum wrayi King, J. Asiat. Soc. Bengal, Pt. 2, on sheet annotated ‘c’ in pencil).
Nat. Hist. 66: 339. 1897.
Type: Peninsular Malaysia, Perak, Matang, July 1888, L. (=) Getonia nutans var. brachystachya DC., Prodr. 3: 15.
Wray 2504 (K barcode K000786607, lectotype designated 1828.
here; isolectotype BM barcode BM000947040). Type: ex herb. Thibaud (holotype G-DC IDC800 475/24).
272 Ian M. Turner

(=) Getonia nutans Roxb., Fl. Ind. 2: 428. 1832, nom. ille- 3.1 Lumnitzera littorea (Jack) Voigt, Hort. Suburb. Cal-
git., non G. nutans Roxb. ex DC. (1828) cutt. 39. 1845
Type: W. Roxburgh s.n. [EIC 4012[A]] (K-W barcode
K001117911, lectotype designated here excluding mate- Bas.: Pyrrhanthus littoreus Jack, Malayan Misc. 2(7): 57.
rial labelled ‘B’ in pencil on the sheet). 1822.
Type: Sumatra, W. Jack s.n. (E barcode E00273921, lecto-
(=) Calycopteris nutans Kurz var. roxburghii Kurz, J. Asi- type designated here).
at. Soc. Bengal, Pt. 2, Nat. Hist. 46(2): 59. 1877.
Type: not traced. (≡) Lumnitzera coccinea Wight & Arn., Prodr. Fl. Ind.
Orient. 316. 1834, nom. illegit. superfl.
(=) Calycopteris nutans Kurz var. glabriuscula Kurz, J.
Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 46(2): 59. 1877. (≡) Bruguiera littorea (Jack) Steud., Nomencl. Bot., ed. 2,
Type: not traced. 1: 231. 1840.

Notes (=) Laguncularia coccinea Gaudich., Voy. Uranie, Bot.

Gangopadhyay and Chakrabarty (1997: 296) referred (2): 68. 1826.
to both the published plate (t. 87) and the unpublished Type: Marianas, C. Gaudichaud-Beaupré s.n. (P barcode
drawing as the type of Getonia floribunda. As both ele- P01901322, lectotype designated here).
ments are original material, the typification is not effec-
tive. A new lectotypification to the published plate is (=) Laguncularia purpurea Gaudich., Voy. Uranie,
therefore made here. There are some Roxburgh speci- Bot. (12): 481. 1830. – Lumnitzera purpurea (Gaudich.)
mens extant, but as this name was published early in C.Presl, Repert. Bot. Syst. 1: 155. 1833.
Roxburgh’s career, it is difficult to be certain that collec- Type: Marianas, C. Gaudichaud-Beaupré s.n. (P barcode
tions were made before the publication date as Roxburgh P01901322, lectotype designated here).
very rarely provided dates (or much other information)
on his specimens. (=) Lumnitzera pedicellata C.Presl, Reliq. Haenk. 2: 23.
1830. – Laguncularia haenkei Endl., Ann. Wiener Mus.
Naturgesch. 1: 180. 1836, nom. illegit. superfl. – Lagun-
3. Lumnitzera Willd., Neue Schriften Ges. Naturf. Fre- cularia pedicellata (C.Presl) Steud., Nomencl. Bot., ed. 2,
unde Berlin 4: 186–187. 1803 2: 5. 1840.
Type: Marianas, T.P.X. Haenke s.n. (PRC barcode
Type: Lumnitzera racemosa Willd. PRC450418 lectotype designated here; isolectotypes
HAL barcode HAL118786, LE barcode LE00016815, PR
(=) Pyrrhanthus Jack, Malayan Misc. 2(7): 57. 1822. sheet nos. 25245a, 25245b).
Type: Pyrrhanthus littoreus Jack
(=) Petaloma coccineum Blanco, Fl. Filip. 345. 1837, as
(=) Problastes Reinw., Syll. Pl. Nov. 2: 10. 1825 [‘1828’]. ‘coccinea’.
Type: Problastes cuneifolia Reinw. Type: Philippines, Luzon, Tayabas Province, Mt Kinata-
kian, 6 April 1913, Merrill: Species Blancoanae No. 521
(=) Pokornya Montrouz., Mem. Acad. Lyon 10: 201. 1860. [L. Escritor leg.] (US barcode US 00623717, neotype des-
Type: Pokornya ettingshausenii Montrouz. ignated here; isoneotypes BM, K barcode K001129749).

(=) Lumnitzera pentandra Griff., Not. Pl. Asiat. 4: 684.

Key to Lumnitzera species 1854. Icon. Pl. As. T. 644A 1854
Type: Mergue, Inter Rhizophora, Madamacca, Sept 1834,
1. Inflorescences terminal, flowers pedicellate, petals red; sta- Griffith (not traced).
mens at least ×2 as long as erect petals............ 3.1 L. littorea
Notes
Inflorescences axillary; flowers more or less sessile, petals
white, stamens scarcely exceeding length of erect petals...... Gangopadhyay and Chakrabarty (1997: 326)
...........................................................................3.2. L. racemosa selected a specimen from K-W under number 4018A
(K001117945) as lectotype of Pyrrhanthus littoreus Jack.
However, this specimen was collected by Wallich from
A synopsis of the native Combretaceae in the Malay Peninsula 273

Pulau Dinding in 1822. It seems very unlikely Jack saw March 1914, Merrill: Species Blancoanae No. 303 (US
the specimen, but the typification would stand as a neo- barcode US00623718, neotype designated here; isoneo-
typification. However, there is a Jack specimen clear- types BM, K barcode K001129748).
ly labelled in E. This is here designated as lectotype to
supersede the earlier designation. (=) Pokornya ettingshausenii Montrouz., Mem. Acad.
Laguncularia coccinea Gaudich. was first validated Lyon 10: 201. 1860, ‘Ettingshanseni’.
in part 2 of the Voyage … l’Uranie rather than part 11 as Type: Ile d’Art, Montrouzier (not traced).
generally cited: “une combrétacée du genre Laguncularia
( L. coccinea), planche 104, et qui est le kada-kandel de (=) Lumnitzera edulis Blume ex Laness., Pl. Util. Col.
Rheed. Mal. 6, planche 37, à fleurs rouge-ponceau très- Franç.: 256. 1886.
vif, désigné ici sous le nom de guia-guia.” The Rheede Type: New Caledonia, Marais de Gatope, 1861-1867, E.
element actually refers to the white-flowered Lumnitzera Vieillard 2571 (K barcode K001129747, lectotype desig-
racemosa, but Gaudichaud-Beaupré’s mention of “very nated here).
bright poppy-red flowers” seems sufficient to distin-
guish the species as this is a highly distinctive character. (=) Lumnitzera racemosa var. pubescens Koord. & Valet.,
There also seems to have been confusion over the chosen Meded. Lands Planten. 61: 33, 34. 1903
epithet with L. purpurea being used in part 12, though Type: Java, Ins. Karimon, Koorders (not traced).
apparently referring to the same taxon. These two names
are lectotypified to the same specimen from Guam here.
4.1 Terminalia L., Syst. Nat. (ed. 12) 2: 665, 674. 1767,
nom. cons.
3.2 Lumnitzera racemosa Willd., Neue Schriften Ges.
Naturf. Freunde Berlin 4: 187. 1803 Type: Terminalia catappa L.

Type: India, J.G. Klein s.n. (B-W barcode B-W08141-010, (=) Buceras P.Browne, Civ. Nat. Hist. Jamaica 221. 1756,
lectotype selected by Wickens (1973: 93); isolectotype nom. rejic.
DNA (fragment). Type: Bucida buceras L.

(=) Funckia karakandel Dennst., Schlüssel Hortus Malab. (=) Bucida L., Syst. Nat. (ed. 10) 2: 1025, 1368. 1759, nom.
32. 1818. cons. (against Buceras P.Browne) and nom. rejic. (against
Type: [published illustration] Rheede, Hort. Malab. 6: Terminalia L.).
t. 37. 1685 (lectotype selected by Gangopadhyay and Type: Bucida buceras L.
Chakrabarty (1997: 326).
(=) Adamaram Adans., Fam. Pl. 2: 445. 1763, nom. rejic.
(=) Problastes cuneifolia Reinw., Syll. Pl. Nov. 2: 10. 1828. Type: Terminalia catappa L. (neotype selected by Exell
Type: [published illustration] Rheede, Hort. Malab. 6: (1931: 125).
t. 37. 1685 (lectotype selected by Gangopadhyay and
Chakrabarty (1997: 329). (=) Myrobalanifera Houtt., Nat. Hist. 2(2): 485, t. 10, f. 2.
1774.
(=) Bruguiera madagascariensis DC., Prodr. 3: 28. 1828. Type: Myrobalanifera citrina Houtt.
Type: [published illustration] Rheede, Hort. Malab. 6:
t. 37. 1685 (lectotype selected by Gangopadhyay and (=) Pamea Aubl., Hist. Pl. Guiane 946, t. 359. 1775, nom.
Chakrabarty (1997: 326). rejic. (against Buchenavia Eichler).
Type: Pamea guianensis Aubl.
(=) Petaloma alternifolium Roxb., Fl. Ind. 2: 373. 1832, as
‘alternifolia’. (=) Tanibouca Aubl., Hist. Pl. Guiane 448. 1775.
Type: [published illustration] Rheede, Hort. Malab. 6: Type: Tanibouca guianensis Aubl.
t. 37. 1685 (lectotype selected by Gangopadhyay and
Chakrabarty (1997: 329). (=) Kniphofia Scop., Intr. Hist. Nat.: 327. 1777, nom. rejic.
(against Kniphofia Moench).
(=) Petaloma album Blanco, Fl. Filip. 344. 1837, as alba. Type: Terminalia catappa L. (neotype designated here).
Type: Philippines, Luzon, Province of Rizal, Maricaban,
274 Ian M. Turner

(=) Chuncoa Pav. in Juss., Gen. Pl. 76. 1789. – Gimberna- (=) Pteleopsis Engl., Abh. Königl. Akad. Wiss. Berlin
tia Ruiz & Pav., Fl. Peruv. Prodr. 138, t. 36. 1794, nom. 1894: 25. 1894.
illegit. superfl. Type: Pteleopsis variifolia Engl.
Type: Chuncoa amazonia J.F. Gmel.
(=) Finetia Gagnep., Notul. Syst. (Paris) 3: 278. 1916.
(=) Badamia Gaertn., Fruct. Sem. Pl. 2: 90. 1790. Type: Finetia rivularis Gagnep.
Type: Badamia commersonii Gaertn.
(=) Terminaliopsis Danguy in Lecomte et al., Madag.
(=) Myrobalanus Gaertn., Fruct. Sem. Pl. 2: 90. 1790. – Bois Analamaz. 110. 1922; Bull. Mus. Natl. Hist. Nat.
Terminalia section Myrobalanus (Gaertn.) DC., Prodr. 3: 29(1): 108. 1923.
12. 1828. Type: Terminaliopsis tetrandrus Danguy
Type: Myrobalanus bellirica Gaertn. (lectotype selected
by Pfeiffer (1874: 394).
Key to Terminalia species
(=) Catappa Gaertn., Fruct. Sem. Pl. 2: 206. 1791. – Ter-
minalia sect. Catappa (Gaertn.) DC., Prodr. 3: 10. 1828. 1. Leaves spirally arranged, clustered at swollen ends of twigs.
Type: Catappa benzoin Gaertn. .................................................................................................... 2

Leaves generally opposite to subopposite, not clustered at

(=) Hudsonia A. Rob. ex Lunan, Hort. Jamaic. 2: 310. twig ends, twig ends not notably swollen............................. 8
1814, nom. illegit., non Hudsonia L. (1767).
Type: Hudsonia arborea A.Rob. ex Lunan. 2. Leaves generally exceeding 10 cm wide; petiole to 12 mm
long...................................................................... 4.3 T. catappa
(=) Fatraea Juss., Dict. Sc. Nat. 16: 206. 1820. Leaves rarely exceeding 10 cm wide; petiole more than 12
Type: Fatraea buxifolia Juss. mm long.................................................................................... 3

(=) Ramatuela Kunth, Nov. Gen. Sp. 7: 253, t. 656. 1825. 3. Leaves with apex obtuse to acuminate.................................. 4
– Terminalia sect. Ramatuela (Kunth) Alwan & Stace, Leaves with apex rounded to emarginate.............................. 7
Ann. Missouri Bot. Gard. 76(4): 1126. 1989, ‘Ramatuella’.
Type: Ramatuela argentea Kunth 4. Leaves with venation distinctly raised above in dry leaves;
fruits without wings, densely velvety pubescent.....................
..............................................................................4.1 T. bellirica
(=) Conocarpus sect. Anogeissus DC., Prodr. 3: 16. 1828.
– Anogeissus (DC.) Wall. ex Guill. & al., Flor. Seneg. Leaves with venation more or less flush to sunken above in
Tent. 1: 279. 1832. dry leaves; fruits winged or if not winged then glabrous to
Type: Conocarpus acuminatus Roxb. ex DC. tomentose, but not densely velvety pubescent..................... 5

5. Petioles not exceeding 2.5 cm long; fruits fleshy without

(=) Pentaptera Roxb. ex DC., Prodr. 3: 14. 1828. wings............................................................. 4.6 T. foetidissima
Type: Pentaptera arjuna Roxb. ex DC. (lectotype desig-
nated here). Petioles generally more than 2.5 cm long; fruits dry with 2
wings.......................................................................................... 6
(=) Vicentia Allemão, Pl. Novas Brasil cum tab. 1844. 6. Leaves glaucous below; fruits densely hairy with wings
Type: Vicentia acuminata Allemão about as wide as long..................................... 4.7 T. glaucifolia

(=) Chicharronia A.Rich., Hist. Phys. Cuba, Pl. Vasc. 529, Leaves not glaucous below; fruits sparsely hairy with
wings distinctly wider than long............ 4.2 T. calamansanai
t. 43. 1845. – Terminalia sect. Chicharronia (A.Rich.)
Alwan & Stace, Fl. Neotrop. Monogr. 107: 187. 2010. 7. Leaves glaucous below; petioles 3 cm long or more; fruits
Type: Chicharronia intermedia A.Rich. winged..................................................... 4.10 T. subspathulata

Leaves not glaucous below; petioles to 2.5 cm long; fruits

(=) Buchenavia Eichler, Flora 49(11): 164. 1866, nom.
not winged....................................................4.9 T. phellocarpa
cons.
Type: Bucida capitata Vahl (lectotype selected by Stace 8. Inflorescences axillary spikes; fruits generally with 2 wings.
(2007: 41). ......................................................................... 4.8 T. mucronata

Inflorescences panicles, fruits with 0 or 3 wings................. 9

A synopsis of the native Combretaceae in the Malay Peninsula 275

9. Leaves with pair of glands on lamina margin near base laurinoides (Teijsm. & Binnend.) C.B.Clarke in Hooker,
rather than on petiole; fruits with three wings...................... Fl. Brit. India 2: 445. 1878. – Myrobalanus laurinoides
.............................................................................4.11 T. triptera (Teijsm. & Binnend.) Kuntze, Revis. Gen. Pl. 1: 237. 1891.
Leaves with pair of glands generally visible on petiole rath- Type: Java, J.E. Teijsmann s.n. (L barcode L0843169, lec-
er than lamina; fruits without wings................................... 10 totype designated here).

10. Leaves to 6 cm wide, glabrous; fruits ellipsoid to fusiform

apex beaked, endocarp with 5-angled sides...... 4.5 T. citrina
4.2 Terminalia calamansanai (Blanco) Rolfe, J. Linn.
Leaves generally more than 6 cm wide, hairy below; fruits Soc., Bot. 21: 310. 1884
globose to ovoid, apex rounded, not beaked, endocarp
more or less round in cross section, not 5-angled................. Bas.: Gimbernatia calamansanai Blanco, Fl. Filip., ed. 2
...............................................................................4.4 T. chebula 266. 1845.
Type: Philippines, Mindanao, Butuan Subprovince,
October 1913, D.P. Miranda For. Bur. 20781 [Mer-
rill: Species Blancoanae No. 605 p.p. (fruiting mate-
4.1 Terminalia bellirica (Gaertn.) Roxb., Pl. Coroman-
rial only)] (US barcode US00623709, neotype designated
del 2: 54, t. 198. 1805, as ‘bellerica’
here excluding flowering material mounted on the same
sheet).
Bas.: Myrobalanus bellirica Gaertn., Fruct. Sem. Pl. 2: 90,
t. 97, f. 2. 1790.
(=) Pentaptera pyrifolia C.Presl, Epimel. Bot. 215. 1851.
Type: [published illustration] Gaertner, Fruct. Sem. Pl. 2:
– Terminalia pyrifolia (C.Presl) Kurz, Prelim. Rep. For.
t. 97, f. 2. 1790 (lectotype selected by Wickens (1990: 7).
Veg. Pegu App. A: lix. 1875.
Type: Moulmein, J.W. Helfer (not traced).
(≡) Terminalia myrobalana B.Heyne ex Roth, Nov. Pl. Sp.
378. 1821, nom. illegit. superfl.
(=) Terminalia papilio Hance, J. Bot. 15: 333. 1877.
Type: Vietnam, Ins. Phu Kok, L. Pierre s.n. [Herb. Hance
(=) Myrobalanus taria Buch.-Ham., Trans Linn. Soc.
19745] (BM, lectotype designated here).
London 17: 159. 1835 [‘1837’]. – Terminalia taria Buch.-
Ham., Trans Linn. Soc. London 17: 159. 1835 [‘1837’].
(=) Terminalia bialata var. cuneifolia C.B.Clarke in
Type: India, Mysore, F. Buchanan-Hamilton s.n. (BM,
Hook.f., Fl. Brit. India 2: 449. 1878.
lectotype designated here).
Type: Burma, Ava, 22 October [1826], N. Wallich 315
[EIC 3972, p.p.] (K-W barcode K001117791, lectotype
(=) Myrobalanus tania Buch.-Ham. ex Kostel., Allg.
designated here).
Med.-Pharm. Fl. 4: 1497. 1835.
Type: [published illustration] ‘Tani’, Rheede, Hort. Mal-
(=) Terminalia blancoi Merr., Philipp. J. Sci., C 4: 645.
ab. 4: t. 10. 1683 (lectotype designated here).
1909.
Type: Philippines, Luzon, Province of Rizal, February
(=) Terminalia punctata Roth, Nov. Pl. Sp.: 381. 1821.
1904, Decades of Philippine Forest Flora No. 2 [Ahern’s
Type: J.P. Rottler s.n. (K barcode K000786123, neotype
Collector leg.] (NY barcode NY00245999, lectotype des-
designated here).
ignated here; isolectotype US barcode US00117589).
(=) Terminalia attenuata Edgew., Trans. Linn. Soc. Lon-
(=) Terminalia calamansanai var. platypteris Merr.,
don 20: 46. 1846.
Philipp. J. Sci., C 4: 646. 1909.
Type: India, Himalaya, altit. 4000 ped., near Jugutgurh,
Type: Philippines, Luzon, Province of Zambales,
M.P. Edgeworth 6 (K barcode K000786121, lectotype des-
Mar 1904, W.M. Maule For. Bur. 371 (NY barcode
ignated here).
NY00246001, lectotype designated here; isolectotype US
barcode US00117606).
(=) Terminalia gella Dalzell, Hooker’s J. Bot. Kew Gard.
Misc. 3: 227. 1851, as Gella.
(=) Terminalia calamansanai var. acuminata Merr.,
Type: Crescit rar in Concano australiore (not traced).
Philipp. J. Sci., C 4: 646. 1909.
Type: Philippines, Luzon, Prov. Principe, Baler,
(=) Terminalia laurinoides Teijsm. & Binnend. in Miquel,
August-October 1903, E.D. Merrill 1067 (NY barcode
Fl. Ned. Ind. 1: 600. 1856. – Terminalia bellirica var.
276 Ian M. Turner

NY00246000, lectotype designated here; isolectotype US (=) Terminalia latifolia Blanco, Fl. Filip. 376. 1837, nom.
barcode US00117607). illegit., non T. latifolia Sw. (1788).
Type: Philippines, Luzon, Pangasinan Prov., Umingan,
(=) Terminalia oryzetorum Craib, Bull. Misc. Inform. May 1914, Merrill: Species Blancoanae No. 197 (US bar-
Kew 1928: 69. 1928. code US00689474, neotype designated here).
Type: Thailand, Mûang Pichit, 25 March 1922, A.F.G.
Kerr 5671 (K barcode K000786200, lectotype designated (=) Terminalia paraensis Mart., Flora 24 Beibl. 2: 24. 1841.
here; isolectotypes ABD, BM barcode BM000947099, NY Type: Brazil, Pará, in sylvis Prov. Paraensis, C.F.P. Mar-
barcode NY00245995, P barcode P02286610). tius s.n. (M sheet no. M-0146722, lectotype designated
here; possible isolectotype M sheet no. M-0146723).
(=) Terminalia latialata C.T.White, J. Arnold Arbor. 10:
249. 1929. (=) Terminalia catappa var. chlorocarpa Hassk., Tijdschr.
Type: Papua New Guinea, Lower Vailala River, 15 Natuurl. Gesch. Physiol. 10: 145. 1843.
March 1926, L.J. Brass 1147 (A barcode A00068656, Type: non designatus.
lectotype designated here; isolectotypes BM barcode
BM000947096, BRI, K). (=) Terminalia catappa var. macrocarpa Hassk., Tijdschr.
Natuurl. Gesch. Physiol. 10: 145. 1843.
4.3 Terminalia catappa L., Mant. Pl. 1: 128. 1767 Type: [published illustration] Rumph., Herb. Amboin. 1:
t. 68. 1741 (lectotype designated here).
Type: Anon. s.n. (lectotype LINN 1221.1, selected by
Byrnes 1977: 38). (=) Terminalia catappa var. rhodocarpa Hassk., Tijdschr.
Natuurl. Gesch. Physiol. 10: 145. 1843.
(≡) Juglans catappa (L.) Lour., Fl. Cochinch. 2: 703. 1790. Type: non designatus.

(≡) Myrobalanus catappa (L.) Kuntze, Revis. Gen. Pl. 1: (=) Terminalia rubrigemmis Tul., Ann. Sci. Nat., Bot. sér.
237. 1891. 4, 6: 102. 1856.
Type: Madagascar (not traced).
(≡) Buceras catappa (L.) Hitchc., Rep. (Annual) Missouri
Bot. Gard. 4: 85. 1893. (=) Terminalia catappa var. pubescens Kurz, For. Fl. Brit.
Burma 1: 454. 1877.
(=) Phytolacca javanica Osbeck, Dagb. Ostind. Resa 276. Type: Andamans, W.S. Kurz (not traced).
1757.
Type: Java, Pulau Panaitan, west of Tg. Parat, 3 October (=) Terminalia burmanica King ex Prain, J. Asiat. Soc.
1951, J. van Borssum Waalkes 818 (K (2 sheets) barcode Bengal, Pt. 2, Nat. Hist. 73(5): 204. 1904.
K001129735, K001129736, neotype designated here; iso- Type: Burma, Sajein, August 1891, Abdul Huk s.n. (lecto-
neotype L L0843090). type CAL accession no. 163973-4, selected by Gangopad-
hyay and Chakrabarty (1997: 343).
(=) Terminalia moluccana Lam., Encycl. 1: 349. 1783.
Type: [published illustration] Rumph., Herb. Amboin. 1: Notes
t. 68. 1741 (lectotype selected by Merrill (1917: 390). Phytolacca javanica Osbeck is the oldest name avail-
able for this well-known seashore species but its trans-
(=) Terminalia subcordata Humb. & Bonpl. ex Willd., fer to Terminalia is blocked by T. javanica Miq. Osbeck
Sp. Pl., ed. 4, 4(2): 968. 1806. – Terminalia catappa var. encountered the species on Nieu Eyland (New Island),
subcordata (Humb. & Bonpl. ex Willd.) DC., Prodr. 3: now called Pulau Peucang, just off the South-West tip of
11. 1828. Java. In the absence of any original material, a specimen
Type: Cuba, Havana, F.W.H.A. Humboldt & A. Bonpland collected from Pulau Panaitan quite nearby in the Sunda
1329 (B-W barcode B-W18940000, lectotype designated Straits, is here designated as neotype for Osbeck’s name.
here; isolectotype P-Bonpl. barcode P00679491). A name often cited in synonymy of T. catappa is
Badamia commersonii Gaertn. However, the original
(=) Terminalia intermedia Bertero ex Spreng., Syst. Veg. figure and description, which refer to the fruit and seed
2: 359. 1825. alone, do not match T. catappa. The fruit is reported
Type: Guadeloupe, 1816-1818, C.G. Bertero s.n. (TO). as oblong ellipsoidal with a slight obtuse beak and an
A synopsis of the native Combretaceae in the Malay Peninsula 277

endocarp with six pronounced longitudinal ridges. It is (=) Terminalia reticulata B.Heyne ex Roth, Nov. Pl. Sp.:
based on a Commerson collection from Madagascar. The 381. 1821.
Tubingen herbarium has some material labelled Bada- Type: India, Cannanore, February 1852, R. Wight s.n.
mia commersonii from Gaertner’s herbarium which con- [Peninsula Indiae Orientalis No. 1012] (K barcode
sists of fruits of T. catappa that certainly differ from the K001129738, neotype designated here).
original description and figure. I am inclined to follow
Tulasne (1856: 92) who considered that Badamia comm- (=) Terminalia aruta Buch.-Ham. ex G.Don, Gen. Hist.
ersoni was a species similar to Terminalia exsculpta. 2: 659. 1832.
Gangopadhyay and Chakrabarty (1997: 343) desig- Type: Buchanan-Hamilton in herb. Lambert (not traced).
nated two plates from Rheede’s Hortus Malabaricus as
lectotype of Badamia commersonii Gaert., but as Gaertner (=) Terminalia gangetica Roxb., Fl. Ind. 2: 437. 1832.
did not cite Rheede, this cannot represent original mate- – Myrobalanus gangetica (Roxb.) Kostel., Allg. Med.-
rial. In order to encourage further work on clarifying the Pharm. Fl. 4: 1497. 1835.
true identity of the Gaertner species, below I designate the Type: not traced.
Gaertner figure as lectotype of Badamia commersonii.
Poiret published Myrobalanus terminalia, apparently (=) Terminalia tomentella Kurz, J. As. Soc. Bengal 42:
entirely based on a figure in Lamarck’s Tableau Encyclo- 80. 1873. – Terminalia chebula var. tomentella (Kurz)
pédique. This figure is clearly a reworking of Gaertner’s C.B.Clarke in Hooker, Fl. Brit. Ind. 2: 446. 1878. –
Badamia illustration, but as neither Poiret nor Lamarck Myrobalanus tomentella (Kurz) Kuntze, Revis. Gen. Pl.
referred to Gaertner or his name, Poiret’s name has to be 1: 237. 1891.
considered as nomenclaturally independent. Type: Burma, Kurz (not traced).

(=) Badamia commersonii Gaertn., Fruct. 2: 90, t. 97 f. (=) Terminalia glandulipetiolata De Wild., Pl. Bequaert.
1(a-g). 1790, as ‘Commersoni’. 4(3): 344. 1928.
Type: [published illustration] “Badamia Comersoni”, Type: Congo Belge, Eala, 29 December 1920, A.A.P.J.G.
Gaertner, Fruct. 2: t. 97 f. 1(a-g). 1790 (lectotype desig- Corbisier-Baland 1630 (BR barcode BR00000887320,
nated here). lectotype designated here; isolectotypes BR barcode
BR00000887353, BR00000887386, S acc. no. S09-25894).
(≡) Terminalia badamia DC., Prodr. 3: 12. 1828, nom.
illegit. superfl.
4.5 Terminalia citrina (Gaertn.) Roxb. in Fleming, Asi-
(=) Myrobalanus terminalia Poir. in Lamarck, Encycl., at. Res. 11: 183. 1810
Suppl. 3: 707. 1814, as Mirobolanus.
Type: [published illustration] Lamarck, Tabl. Encycl. 2, Bas.: Myrobalanus citrina Gaertn., Fruct. Sem. Pl. 2: 91,
5(1): Pl. 849 f. 2. 1799 (lectotype designated here). t. 97, f. 2n-s. 1790.
Type Cons.: Bangladesh, Mymensingh District, Gabtulli,
8 July 1872, C.B. Clarke 17257 (K K000608176).
4.4 Terminalia chebula Retz., Observ. Bot. 5: 31. 1788
(≡) Terminalia chebula Retz. var. citrina (Gaertn.) Gag-
Type: J.G. Koenig s.n. (lectotype LD acc. no. 1638155, nep., Fl. Gen. Indo-Chine 2: 753. 1920.
selected by Fischer (1932: 57). (=) Bucida comintana Blanco, Fl. Filip. 856. 1837. – Ter-
minalia comintana (Blanco) Merr., Philipp. J. Sci., C 4:
(≡) Myrobalanus chebula (Retz.) Gaertn., Fruct. Sem. Pl. 300. 1909.
2: 91. 1790. Type: Philippines, Luzon, Batangas Prov., Mt Macolot,
5 Feb 1915, E.D. Merrill: Species Blancoanae 780 [M.
(≡) Buceras chebula (Retz.) A.Lyons, Pl. Nam.: 71. 1900. Ramos leg.] (US barcode US00623713, neotype designat-
ed here; isoneotype BM).
(=) Myrobalanifera citrina Houtt., Handl. Pl.-Kruidk. 2:
486, t. 10 f. 2. 1774. (=) Embryogonia arborea Teijsm. & Binnend., J. Bot.
Type: [published illustration] Houttuyn., Handl. Pl.- Neerl. 1: 365. 1861. – Combretum arboreum (Teijsm. &
Kruidk. 2: t. 10, f. 2. 1774 (lectotype selected by Turner Binnend.) Miq., Ann. Mus. Bot. 4: 115. 1868. – Termi-
(2014: 11). nalia teysmannii Koord. & Valeton, Meded. Lands Plan-
278 Ian M. Turner

tentuin 61: 20. 1902, nom. illegit. superfl. – Terminalia (=) Terminalia ellipsoidea Merr.
arborea (Teijsm. & Binnend.) Koord., Exkursionsfl. Java Type: Philippines, Mindoro, Pinamalayan, May 1903,
2: 671. 1912. E.D. Merrill 2148 (K barcode K000786172, lectotype des-
Type: Java (not traced). ignated here; isolectotype US barcode US00117598).

(=) Terminalia citrina var. malayana Kurz, J. Asiat. Soc. (=) Terminalia borneensis Sloot., Bijdr. Combret. Fla-
Bengal, Pt. 2, Nat. Hist. 45: 130. 1876. court. Ned.-Ind. 14. 1919.
Type: India, Ins. Nicobar, Expedition Novara 75 [leg. A. Type: Borneo, O. Beccari P.B. 2593 (K barcode
Jelinek 132] (W sheet no. W 0080071, lectotype designat- K000786182, lectotype designated here; isolectotype
ed here). FI-B).

(=) Terminalia manii King, J. Asiat. Soc. Bengal, Pt. 2,

Nat. Hist. 66: 329. 1897. 4.7 Terminalia glaucifolia Craib, Bull. Misc. Inform.
Type: Nicobars, Dr King’s Collector 485 (CAL, lectotype Kew 1928(2): 68. 1928
selected by Gangopadhyay and Chakrabarty (1997: 348);
isolectotypes BM, K barcode K001129734). Type: Thailand, Doi Sootep, 20 August 1910, A.F.G. Kerr
1328 (BK, lectotype selected by Nanakorn 1985, p. 81;
(=) Terminalia multiflora Merr., Govt. Lab. Publ. Philipp. isolectotypes ABD, BM, K barcode K000786198, P).
27: 34. 1904.
Type: Philippines, Luzon, Province of Rizal, Bosoboso,
July 1903, E.D. Merrill 2796 (K barcode K000786176, 4.8 Terminalia mucronata Craib & Hutch., Bull. Misc.
lectotype designated here; isolectotypes NY barcode Inform. Kew 1909(9): 358. 1909
NY00246011, US barcode US00117611).
Type: Thailand, Chiengmai, Doi Sootep, 11 April 1909,
(=) Terminalia curtisii Ridl., Bull. Misc. Inform. 1931: A.F.G. Kerr 593 (K (4 sheets) barcodes K000786194,
449. 1931. K000786194, K000786195, K000786196, lectotype desig-
Type: Peninsular Malaysia, Penang, Waterfall, June 1896, nated here; isolectotypes BM barcode BM000947102, L
C. Curtis 3207 (K barcode K000786161, lectotype desig- barcode L0009802, P barcode P02286601, TCD barcode
nated here; isolectotypes K barcode K000786160, SING TCD0016741).
barcode SING0055417).
(=) Terminalia corticosa Pierre ex Craib & Hutch., Bull.
Misc. Inform. Kew 1909(9): 358. 1909.
4.6 Terminalia foetidissima Griff., Not. Pl. Asiat. 4: 685. Type: Vietnam, Baria, montibus Dinh, March 1862, L.
1854 Pierre 203 (P barcode P02286603, lectotype designated
here).
Type: Burma, Mergui, March 1835, W. Griffith s.n. (syn-
type CAL). (=) Terminalia thorelii Gagnep., Not. Syst. 3: 287. 1916.
Type: Cambodia, Ubon, January anno. 1866-1868, C.
(≡) Myrobalanus foetidissima (Griff.) Kuntze, Revis. Gen. Thorel 2824 (P barcode P02286602, lectotype designated
Pl. 1: 237. 1891. here).

(=) Terminalia sumatrana Miq., Fl. Ned. Ind., Eerste Bijv. (=) Parinari euadenia Kosterm., Bot. Tidsskr. 67: 321.
326. 1861. 1973.
Type: Sumatra, Palembang, Toeboean, Ogan oeloe, H.B. Type: Thailand, Chiengmai, Doi Sutep, 15 October 1958,
3814 [J.E. Teijsmann leg.] (U barcode U0001196, lecto- T. Sørensen et al. 5687 (holotype C C10017914).
type designated here; isolectotype L barcode L.2493804).
Notes
(=) Terminalia oocarpa Merr., Publ. Bur. Sci. Gov. Lab. The name Terminalia corticosa first appeared in
17: 32. 1904, as ovocarpa. Lanessan (1886: 315). The descriptive content of the
Type: Philippines, Luzon, Province of Bataan, Lamao account is slight – a size range of the tree and colour of
River, November 1903, P.T. Barnes For. Bur. 67 (K bar- the wood. This has been considered insufficient to vali-
code K000786170, lectotype designated here). date the name (Gagnepain 1916: 215, Lecompte 1969:
A synopsis of the native Combretaceae in the Malay Peninsula 279

88). Craib and Hutchinson described Terminalia mucro- Type: Vietnam, Delta du Mè-Không, 1875-1877, J.H.A.J.
nata in 1909. As well as a fairly detailed description, they Harmand s.n. (P barcode P04716917, lectotype designat-
included a diagnosis comparing Terminalia mucronata ed here; isolectotypes K barcode K001129732, P barcodes
and T. corticosa. This effectively validated T. corticosa P04716912, P04717014).
because the description of T. mucronata was sufficient
to validate that name. It is not clear on what Craib and (=) Terminalia obliqua Craib, Bull. Misc. Inform. Kew
Hutchinson based their concept of T. corticosa as I have 1912: 153. 1912.
not found any flowering specimens in K or BM that Type: Thailand, Sriracha, Nawng Kaw, 19 Septem-
they are likely to have seen. Possibly they visited Paris ber 1911, A.F.G. Kerr 2073 (K barcode K000786152,
or had material on loan. In order to maintain continu- lectotype designated here; isolectotypes BM barcode
ity, I designate the specimen chosen by Lecompte (1969) BM000947100, E barcode E00284575, K barcocde
as lectotype of T. corticosa Pierre ex Craib & Hutch. As K001129733, TCD barcode TCD0016766).
Lecompte did not cite Craib and Hutchinson’s publica-
tion, I here establish T. mucronata Craib & Hutch. as (=) Terminalia tripteroides Craib, Bull. Misc. Inform.
having priority over T. corticosa Pierre ex Craib & Hutch. Kew 1912: 152. 1912.
For Terminalia thorelii, the collection number 2823 Type: Thailand, Meh Ping, Muang Hawt, 6 September
was cited in the protologue. I have not traced this num- 1911, A.F.G. Kerr 2010A (K barcode K000786151, lec-
ber and presume it was an error for 2824. totype designated here; isolectotypes BM, E barcode
E00284577, K no barcode, L barcode L0009816, TCD
barcode TCD0016755).
4.9 Terminalia phellocarpa King, J. Asiat. Soc. Bengal,
Pt. 2, Nat. Hist. 66: 330. 1897 (=) Terminalia hainanensis Exell, Sunyatsenia 2: 1. 1934.
Type: China, Hainan, [Gnai Yuen,] 5 Jul 1933, F.C. How
Type: Singapore, Bukit Mandai, 1892, H.N. Ridley 3835 70938 (BM barcode BM000947030, lectotype designated
(K barcode K000786153, lectotype designated here). here; isolectotypes A barcodes A00068634, A00068635,
B barcode B 10 0272120, IBK barcode IBK00190688, K
barcode K000786127, NAS barcode NAS00047726, NY
4.10 Terminalia subspathulata King, J. Asiat. Soc. Ben- barcode NY00245994, P barcode P02286590, PE barcode
gal, Pt. 2, Nat. Hist. 66: 332. 1897 PE00994592, US barcode US00117594).

Type: Peninsular Malaysia, Perak, Kinta, Gopeng, July Notes

1883, Dr King’s Collector 4529 (K barcode K000786156, An entry for Terminalia inguovenulosa Pierre is
lectotype designated here; isolectotypes BM, SING (×2) found in Lanessan (1886: 315), but as the only descrip-
barcodes SING0055425, SING0055427). tion presented is a vague size of the tree, the name is
not validly published. Gagnepain (1916) indicated that
Lanessan’s name was a nomen nudum and that it was an
4.11 Terminalia triptera Stapf, Bull. Misc. Inform. Kew error for Terminalia nigrovenulosa. Gagnepain effectively
1895: 103. 1895 validated T. nigrovenulosa by providing a diagnosis com-
paring it with Terminalia oliveri Brandis. Terminalia
Type: Peninsular Malaysia, Langkawi, September 1890, triptera Stapf has priority over T. nigrovenulosa Pierre ex
C. Curtis 1684 (lectotype K barcode K000786162, lec- Gagnep. and is here accepted as the correct name for the
totype designated here; isolectotype SING barcode species.
SING0055430).

(=) Terminalia nigrovenulosa Pierre ex Gagnep., Not. ACKNOWLEDGEMENTS

Syst. 3: 285. 1914.
Type: Vietnam, ad pedes montis Mu Xoai, October 1866, I am very grateful to Otakar Šida (PR), Uta Grünert
L. Pierre 5011 (P barcode P02286589, lectotype selected (TUB) and Christian Bräuchler (W) for information on
by Lecompte (1969: 92). specimens.

(=) Terminalia nigrovenulosa Pierre ex Gagnep. var. gra-

cilior Gagnep., Fl. Gen. Indo-Chine 2: 756. 1920
280 Ian M. Turner

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