REINWARDTIA
Vol. 20. No. 1. pp: 9‒16
DOI: 10.14203/reinwardtia.v20i1.4022
9
AN UPDATED SYNOPSIS OF LUDWIGIA (ONAGRACEAE) IN MALESIA
Received March 28, 2021; accepted April 12, 2021
I. M. TURNER
Singapore Botanical Liaison Officer, Royal Botanic Gardens Kew, UK. Singapore Botanic Gardens, National Parks
Board, 1 Cluny Road, Singapore 259569, Republic of Singapore.
E-mail: I.Turner@kew.org
ABSTRACT
TURNER, I. M. 2021. An updated synopsis of Ludwigia (Onagraceae) in Malesia. Reinwardtia 20(1): 9–16. — A
synopsis of the seven native or naturalised species of Ludwigia (Onagraceae) occurring in Malesia is presented. The
newly established Ludwigia leptocarpa is included. The paper incorporates a key to the species, synonymy and
typification. Lectotypes are designated for 25 names, including five at the second step.
Key words: Ludwigia leptocarpa, South-East Asia, typification, weeds.
ABSTRAK
TURNER, I. M. 2021. Pembaruan sinopsis Ludwigia (Onagraceae) di Malesia. Reinwardtia 20(1): 9–16. —
Tulisan ini menyajikan sinopsis tujuh jenis asli atau jenis yang bernaturalisasi dari Ludwigia (Onagraceae) yang
terdapat di Malesia, termasuk di dalamnya yang baru dikukuhkan yaitu Ludwigia leptocarpa. Tulisan ini
menggabungkan kunci jenis, sinonim dan tipifikasi. Ditetapkan pula lektotipe untuk 25 nama, termasuk lima pada
tahapan kedua.
Kata kunci: Asia Tenggara, gulma, Ludwigia leptocarpa, tipifikasi.
INTRODUCTION
Ludwigia is a genus of 82 species distributed
through the tropical, subtropical and warm
temperate regions of the world; most diverse in the
New World (Wagner et al., 2007). Ludwigia is
sister to the rest of the Onagraceae and placed in
its own subfamily: Jussiaeoideae. The genus has
been divided into 23 sections (Wagner et al.,
2007), but recent molecular analyses found limited
support for this system of infrageneric division
(Liu et al., 2017).
The revision of Ludwigia (Onagraceae) in the
Old World by Raven (1964) has stood the test of
time very well; and his account for Flora
Malesiana (Raven, 1977) equally so. Raven (1977)
recognised six species for Malesia. While this is a
very small number compared with the size of
the complete flora of the region, the species are
quite important as weeds, particularly in rice-
growing areas (Kostermans et al., 1987). A newly
established invasive species and some further
clarification of typification make it worthwhile in
publishing an updated synopsis of the genus for
Malesia.
MATERIALS AND METHODS
The paper is largely based on study of material
in the herbarium of the Royal Botanic Gardens
Kew (K). In addition, on-line information was
searched for information on type holdings. Recent
literature was scanned and relevant references are
cited under the description and distribution
heading for each species.
RESULTS AND DISCUSSION
LUDWIGIA L.
Ludwigia L., Sp. Pl. 1 (1753) 118, as Ludvigia. ––
Type: Ludwigia alternifolia L., lectotype designat-
ed by Britton & Brown, Ill. Fl. N. U.S. ed. 2. 2
(1913) 586.
Isnardia L., Sp. Pl. 1 (1753) 120. – Dantia Boehm.
in Ludwig, Def. Gen. Pl. (1760) 388, nom. illegit.
(superfl.). – Quadricosta Dulac, Fl. Hautes-
Pyrénées (1867) 329, nom. illegit. (superfl.). ––
Type: Isnardia palustris L.
Jussiaea L., Sp. Pl. 1 (1753) 388. – Ludwigia sect.
Jussiaea (L.) Baill., Hist. Pl. 6 (1877) 463. ––
Type: Jussiaea repens L., lectotype designated by
Hitchcock & Greene, Prop. Brit. Bot. (1929) 153.
Cubospermum Lour., Fl. Cochinch. (1790) 258,
275. –– Type: Cubospermum palustre Lour.
Prieuria DC, Prodr. 3 (1828) 58. –– Type: Pri-
euria senegalensis DC.
Adenola Raf., Aut. Bot. (1840) 36. –– Type:
Adenola grandiflora (Michx.) Raf.
Diplandra Raf., Aut. Bot. (1840) 35, nom. illegit.,
non Diplandra Bertero (1830), nec Diplandra
Hook. & Arn. (1838). –– Type: Diplandra
decurrens (Walt.) Raf., lectotype designated by
Pennell, Bull. Torrey Bot. Club 48 (1921) 92.
Corynostigma C.Presl., Epim. Bot. (1851 [‘1849’])
218. –– Type: Corynostigma jussiaeoides C.Presl.
mailto:I.Turner@kew.org
REINWARDTIA 10 [VOL.20
Nematopyxis Miq., Fl. Ned. Ind. 1(1) (1856) 626,
630. –– Type: Nematopyxis prostrata (Roxb.)
Miq.
Oocarpon Micheli, Flora 57 (1874) 303. –– Type:
Oocarpon jussiaeoides Micheli
Ludwigia [unranked] Ludwigiantha Torr. &
A.Gray, Fl. N. Amer. 1 (1840) 526. – Ludwigi-
antha (Torr. & A.Gray) Small, Bull. Torrey Bot.
Club 24 (1897) 178. –– Type: Ludwigia arcuata
Walt.
Jussiaea sect. Fissendocarpa Haines, J. Proc.
Asiat. Soc. Bengal 15 (1919) 314. – Fissendocar-
pa (Haines) Bennet, J. Bombay Nat. Hist. Soc. 67
(1970) 125. –– Type: Jussiaea fissendocarpa
Haines
Key to Ludwigia species in Malesia
1a. Sepals generally 5, sometimes more …..……2
1b. Sepals generally 4…………………………...3
2a. Plant with creeping or floating stems; petals
white with a yellow base; pedicel of capsule
2.5–5.5 cm long; each seed firmly embedded
in a block of endocarp……...1. L. adscendens
2b. Plant erect; petals entirely yellow; pedicel of
capsule to 2 cm long; each seed surrounded
by an easily detached horseshoe-shaped
section of endocarp……........3. L. leptocarpa
3a. Sepals less than 5 mm long……………….....4
3b. Sepals more than 6 mm long………………...6
4a. Capsules with 4 longitudinal rows of seeds
clearly discernible when dry…7. L. prostrata
4b. Capsules more or less terete when dry, longi-
tudinal rows of seeds not discernible …….... 5
5a. Capsules with two sorts of seed, free in apical
portion, embedded in corky endocarp below
…………..………………. 2. L. hyssopifolia
5b. Capsules with seeds all free ..….5. L. perennis
6a. Bracteoles ca. 8 mm long, capsules obcon-
cal………………………….....6. L. peruviana
6b. Bracteoles minute, ca. 0.5 mm long, capsule
cylindrical with 8 longitudinal ridges or ribs
……………………………… 4. L. octovalvis
1. LUDWIGIA ADSCENDENS (L.) H.Hara
Ludwigia adscendcens (L.) H.Hara, J. Jap. Bot. 28
(1953) 291. – Jussiaea adscendens L., Mant. Pl.
(1767) 69. –– Type: India. Mysore, Hassan
District, Belur-Gendehally Road, 17 September
1969, C. Saldanha 15027 (Neotype MO [MO-
2334663], designated by Raven et al. in Jarvis
(ed.), Order out of Chaos (2007) 605).
Jussiaea repens L., Sp. Pl. (1753) 388, non Lud-
wigia repens J.R.Forst. (1771). –– Type:
[published illustration] ‘Nir-carambu’ in Rheede,
Hort. Malab. 2: t. 51. 1679 (Lectotype designated
by Raven in Jarvis & al. (ed.), Regnum Veg. 127
(1993) 58).
Cubospermum palustre Lour., Fl. Cochinch.
(1790) 275. –– Type: Loureiro s.n. (Lectotype BM
[BM000793045], designated by Merrill, Trans.
Am. Phil. Soc. 24 (1935) 289).
Jussiaea fluviatilis Blume, Bijdr. Fl. Ned. Ind.
(1827) 1132. –– Type: Java, Blume 1447
(Lectotype L [L0008654], designated by Raven,
Reinwardtia 6 (1964 [‘1963’]) 387 at the first step
and at the second step here; possible Isolectotype
L0008653).
Jussiaea floribunda Griff., Not. Pl. Asiat. 4 (1854)
688. –– Type: Burma, Mergui, Sept 1834, Griffith
(not traced).
Jussiaea repens var. glaberrima Kuntze, Revis.
Gen. Pl. 1 (1891) 251. –– Type: Java, Batavia,
May 1875, Kuntze 4180 (Lectotype NY
[NY00232243], designated here).
Jussiaea repens var. pilosa Kuntze, Revis. Gen. Pl.
1 (1891) 251. –– Type: Cambodia, March 1875,
Kuntze s.n. (Lectotype NY [NY00232245],
designated here).
Jussiaea repens forma albiflora Hochr., Candollea
2 (1925) 479. –– Type: Java, Preanger, étangs tièd-
es de Tjipanas près Garroet, 26 October 1904,
Hochreutiner 2208 (Lectotype G [G00013806],
designated by Raven, Reinwardtia 6 (1964
[‘1963’]) 388).
Description and distribution. Raven (1964: 387),
Vu & Vidal (1973: 26), Khan & Shamsunnahar
(1977: 3), Raven (1977: 104), Kostermans et al.
(1987: 368), Thompson (1990: 227), Wagner
(1995: 341), Barua (2010: 60), Boufford & Raven
(2014: 602), Ummul-Nazrah (2017: 55).
2. LUDWIGIA HYSSOPIFOLIA (G.Don) Exell
Ludwigia hyssopifolia (G.Don) Exell, Garcia de
Orta 5 (1957) 471. – Jussiaea hyssopifolia G.Don,
Gen. Hist. 2 (1832) 693. –– Type: St Thomas [São
Tomé], G. Don 42 (Holotype BM
[BM000528814]).
Jussiaea linifolia Vahl, Eclog. Amer. 2 (1798) 32.
– Fissendocarpa linifolia (Vahl) Bennet, J.
Bombay Nat. Hist. Soc. 67 (1970) 126. – Ludwigia
linifolia (Vahl) R.S.Rao, Fl. Goa, Diu, Dadra &
Nagarhaveli 1 (1985) 179, nom. illegit., non Poir.
(1814). –– Type: America meridionali, von Rohr
188 (Lectotype C [C10016131], designated by
Raven, Reinwardtia 6 (1964 [‘1963’]) 385).
Jussiaea micrantha Kunze, Index Seminum (LZ,
Lipsiensis) 1847 (1848) 2; Linnaea 24 (1851) 177.
– Ludwigia micrantha (Kunze) H.Hara, J. Jap. Bot.
28 (1953) 293. – Lectotype (designated by Jonck-
er, Fl. Suriname 3(2) (1942) 19): Surinam, Kegel
909 (GOET042401).
Jussiaea weddellii Micheli, Flora 57 (1874) 301,
as Weddelii. –– Type: Brazil, Weddell 2334
(Lectotype P [P01819501], designated by Raven,
TURNER: An updated synopsis of Ludwigia in Malesia
2021] 11
Reinwardtia 6 (1964 [‘1963’]) 385).
Jussiaea fissendocarpa Haines, J. Proc. Asiat. Soc.
Bengal 15 (1920 [‘1919’]) 313. –– Type: India,
Bihar, Purneah, 21 December 1918, H.H. Haines
4541 (Lectotype CAL [CAL0000010226],
designated here; Isolectotype K [K001129806]).
Description and distribution. Raven (1964: 385),
Vu & Vidal (1973: 24); Khan & Shamsunnahar
(1977: 4), Raven (1977: 104), Kostermans et al.
(1987: 370), Thompson (1990: 227), Wagner
(1995: 349), Boufford & Raven (2014: 603),
Ummul-Nazrah (2017: 56).
Notes. Raven (1964) cited the type of Jussiaea
weddellii as 3331. I take this to be a typographical
error for 2334.
3. LUDWIGIA LEPTOCARPA (Nutt.) H.Hara
Ludwigia leptocarpa (Nutt.) H.Hara, J. Jap. Bot.
28 (1953) 292. – Jussiaea leptocarpa Nutt., Gen.
N. Am. Pl. 1 (1818) 279. –– Type: Nuttall s.n.
(Lectotype PH [PH00016622], designated by
Brenan in Turrill & Milne-Redhead, Fl. Trop. E.
Africa, Onagraceae (1953) 16).
Jussiaea pilosa Kunth, Nov. Gen. Sp. 6 (1823)
101, t. 532. – Jussiaea variabilis var. pilosa
(Kunth) Kuntze, Revis. Gen. Pl. 1 (1891) 251. ––
Type: Colombia, Río Apuro, Bonpland 375
(Lectotype P-Bonpl. [P00679565], designated by
Brenan in Turrill & Milne-Redhead, Fl. Trop. E.
Africa, Onagraceae (1953) 16).
Jussiaea velutina G.Don, Gen. Syst. 2 (1832) 695.
–– Type: São Tomé, G. Don s.n. (Holotype BM
[BM000957914]).
Jussiaea surinamensis Miq., Linnaea 18 (1844)
370. –– Type: Surinam, Oranjewoud, langs de
zandweg, October 1842, Focke 721 (Holotype U
[U0005340]).
Jussiaea schottii Micheli, Flora 57 (1874) 302. ––
Type: Brazil, Rio de Janeiro, Schott 4192
(Lectotype W, designated by Raven, Reinwardtia 6
(1964 [‘1963’]) 375).
Jussiaea pilosa var. robustior Donn.Sm., Bot. Gaz.
16 (1891) 6. –– Type: Guatemala, Zacate-
pequez, April 1890, Donnell Smith 2123
(Lectotype US [US00123986], designated here;
Isolectotypes GH, K [K000533292], NY).
Jussiaea pilosa var. pterocarpa Hassler, Fedde
Rep. Sp. Nov. 12 (1913) 274. –– Type: Paraguay,
Gran Chaco, Santa Eliza, December 1903, Hassler
2708 (Lectotype B [B 10 0367937], designated
here; isolectotypes BM, K, MPU, NY, POM).
Jussiaea seminuda H.Perr., Not. Syst. 13 (1947)
146. –– Type: Madagascar, Marais de Mahabo,
June 1926, H. Perrier de la Bâthie 17641
(Lectotype P [P00412973], designated at the first
step by Raven, Reinwardtia 6 (1964 [‘1963’]) 376
and at the second step here; Isolectotypes K
[K000310868], P [P00412974, P00412975]).
Description and distribution. Raven (1964:
375).
Notes. The first report of this species being found
in Malesia came from Papua New Guinea. Tucker
(2017: 96) found that the species is invading
riparian areas, wetlands and swamps in Papua
New Guinea. I have also seen a specimen (FRI
84523) collected from Gunong Ledang, Johore,
Peninsular Malaysia. This species should certainly
looked out for in other parts of the Malesian
region.
4. LUDWIGIA OCTOVALVIS (Jacq.) P.H.Raven
Ludwigia octovalvis (Jacq.) P.H.Raven, Kew Bull.
15 (1962 [‘1961’]) 476. – Oenothera octovalvis
Jacq., Enum. Syst. Pl. (1760) 19. – Jussiaea
octovalvis (Jacq.) Sw., Observ. Bot. (1791) 142. –
Jussiaea octofila DC., Prodr. 3 (1828) 57, nom.
illegit. (superfl.). – Jussiaea suffruticosa var.
octofila H.Lev., Le Monde des Plantes (1894) 3. –
Ludwigia octovalvis var. octofila Alain, Bull.
Torrey Bot. Club 90 (1963) 191, nom. illegit.
(superfl.). –– Type: [published illustration]
Plumier, Pl. Am. (7) (1758) t. 175, f. 1 (Lectotype
designated by Bizzarri & Raven, Webbia 27
(1972) 490).
Jussiaea pubescens L., Sp. Pl. ed. 2 (1762) 555. –
Ludwigia pubescens (L.) H.Hara, J. Jap. Bot. 28
(1953) 293. –– Type: Anon. s.n. (Lectotype LINN
552.3, designated by Dorr & Wiersema, Taxon 59
(2010) 1573).
Jussiaea hirsuta Mill., Gard. Dict. ed. 8 (1768) no.
5. –– Type: Houston s.n. (Lectotype BM
[BM000957916], designated by Raven,
Reinwardtia 6 (1964 [‘1963’]) 357).
Jussiaea angustifolia Lam., Encycl. 3 (1789) 331.
– Jussiaea suffruticosa var. angustifolia (Lam.)
F.M.Bailey, Syn. Queensl. Fl. (1883) 197. –
Jussiaea suffruticosa forma angustifolia (Lam.)
Alston, Handb. Fl. Ceylon 6 (Suppl.) (1931) 131.
–– Type: Java, Anon. s.n. (Lectotype P-LA – IDC
6207 LM-243/5, designated by Raven,
Reinwardtia 6 (1964 [‘1963’]) 357).
Jussiaea hirta Lam., Encycl. 3 (1789) 331. ––
Type: Brazil, Commerson s.n. (Lectotype P-LA
LM-243/9, designated by Munz, Darwiniana 4
(1942) 238).
Jussiaea octonervia Lam., Encycl. 3 (1789) 332. –
Jussiaea suffruticosa var. octonervia (Lam.)
Bertoni, Descr. Fis. Econ. Paraguay (1910) 17. –
Jussiaea suffruticosa subsp. octonervia (Lam.)
Hassl., Bull. Soc. Bot. Genève ser. 2, 5 (1913)
271. –– Type: Martinique, Anon. 33b (Lectotype
P-LA LM-243/3, designated by Munz, Darwini-
REINWARDTIA 12 [VOL.20
ana 4 (1942) 242).
Jussiaea villosa Lam., Encycl. 3 (1789) 331. –
Jussiaea suffruticosa var. villosa (Lam.)
F.M.Bailey, Syn. Queensl. Fl. (1883) 197. –
Jussiaea suffruticosa forma villosa (Lam.) Alston,
Handb. Fl. Ceylon 6 (Suppl.) (1931) 131. – Lud-
wigia pubescens var. villosa (Lam.) H.Hara, J.
Jap. Bot. 28 (1953) 293. –– Type: Anon. s.n., India
(Lectotype P-LA LM-243/6 designated by Raven,
Reinwardtia 6 (1964 [‘1963’]) 362).
Epilobium fruticosum Lour., Fl. Cochinch. (1790)
226. – Jussiaea fruticosa (Lour.) DC., Prodr. 3
(1828) 57. –– Type: not traced.
Epilobium tetragonum Lour., Fl. Cochinch. (1790)
225, nom. illegit., non L. (1753). – Jussiaea
tetragona Spreng., Syst. Veg. 2 (1825) 231. ––
Type: not traced.
Jussiaea scabra Willd., Enum. Pl. 1 (1809) 449.
–– Type: Germany, Hort. Bot. Berol., Anon. s.n.
(Lectotype B-W [B -W 08139 -01 0], designated
by Raven, Reinwardtia 6 (1964 [‘1963’]) 362).
Jussiaea exaltata Roxb. ex Andrews, Bot. Repos.
10 (1811) t. 621, as ‘Jussieua’. – Jussiaea erecta
var. exaltata (Roxb. ex Andrews) Ridl., J. Bot. 59
(1921) 259. –– Type: [published illustration]
Andrews, Bot. Repos. 10 (1811) t. 621 (lectotype
designated here).
Jussiaea exaltata Roxb., Hort. Bengal. (1814) 33,
as ‘Jussieua’, nom. illegit., non Roxb. ex Andrews
(1811). –– Type: [published illustration] ‘Cattu-
carambu’, Rheede, Hort. Malab. 2 (1679) t. 50
(lectotype designated by Turner, Taxon 62 (2013)
158).
Jussiaea ligustrifolia Kunth, Nov. Gen. Sp. 6
(1823) 100. – Jussiaea suffruticosa var.
ligustrifolia (Kunth) Griseb., Mem. Amer. Acad.
Arts (n. s.) 8 (1860) 187. – Ludwigia pubescens
var. ligustrifolia (Kunth) H.Hara, J. Jap. Bot. 28
(1953) 293. – Ludwigia octovalvis var. ligustrifo-
lia (Kunth) Alain, Bull. Torrey Bot. Club 90
(1963) 191. –– Type: Mexico, Humboldt & Bon-
pland s.n. (Lectotype P-Bonpl. [P00679564],
designated by Raven, Reinwardtia 6 (1964
[‘1963’]) 358).
Jussiaea salicifolia Kunth, Nov. Gen. Sp. 6 (1823)
99. –– Type: Colombia, Guaduas, Humboldt &
Bonpland 1756 (Lectotype P-Bonpl. [P00679563],
designated at the first step by Raven, Reinwardtia
6 (1964 [‘1963’]) 358, and at the second step here;
Isolectotypes P [P00136827, P01819477]).
Jussiaea ovalifolia Sims, Bot. Mag. 52 (1824) t.
2530, as ‘Jussieua’. –– Type: [published
illustration] Sims, Bot. Mag. 52 (1824) t. 2530
(lectotype designated here).
Jussiaea angustifolia Blume, Bijdr. Fl. Ned. Ind.
(1827) 1132, nom. illegit., non Lam. (1789). –
Jussiaea blumeana DC., Prodr. 3 (1828) 55. ––
Type: Java, Buitenzorg, August [sine anno],
Blume 1050 (Lectotype L [L008658], designated
by Raven, Reinwardtia 6 (1964 [‘1963’]) 358;
possible Isolectotype L [L008659]).
Jussiaea burmanni DC., Prodr. 3 (1828) 57. ––
Type: not designated.
Jussiaea marcgravii DC., Prodr. 3 (1828) 58. ––
Type: [published illustration] Camaranbaya in
Marcgravius, Historia Naturalis Brasiliae Historiae
Plantarum 1 (1648) 30 (Lectotype, upper
illustration on page, designated here).
Jussiaea parviflora Camb. in St. Hil., Fl. Bras.
Merid. 2 (1829) 263. –– Type: Brésil, Province de
Minas Geraes, 1816-1821, A. de Saint Hilaire 692
(Lectotype P [P01819478], designated here, Iso-
lectotype P [P01819479]).
Jussiaea calycina C.Presl, Reliq. Haenk. 2 (1831)
34. –– Type: not traced.
Jussiaea hirsuta C.Presl, Rel. Haenk. 2 (1831) 34,
nom. illegit., non Mill. (1768), nec Vell. (1829). –
Jussiaea haenkeana Steud., Nom. ed. 2, 1 (1840)
836. –– Type: not traced.
Jussiaea macropoda C.Presl, Reliq. Haenk. 2
(1831) 35. – Jussiaea suffruticosa var. macropoda
(C.Presl) Munz, Darwiniana 4 (1942) 239. – Lud-
wigia pubescens var. macropoda (C.Presl) H.Hara,
J. Jap. Bot. 28 (1953) 293. – Ludwigia octovalvis
subsp. macropoda (C.Presl) P.H.Raven, Kew Bull.
15 (1962) 476. – Ludwigia octovalvis var. macrop-
oda (C.Presl) Shinners, Sida 1 (1964) 385. ––
Type: In Peruvia prope urbem Lima, T.P.X. Haen-
ke s.n. (Lectotype PRC [PRC450096], designated
here).
Jussiaea venosa C.Presl, Reliq. Haenk. 2 (1831)
33. –– Type: Mexico, T.P.X. Haenke s.n.
(Lectotype PR, designated by Raven, Reinwardtia
6 (1964 [‘1963’]) 358).
Jussiaea persicariifolia Schltdl., Linnaea 12
(1838) 271, as ‘persicariaefolia’. –– Type: not
traced.
Jussiaea persicariifolia Schltdl. forma major
Schltdl., Linnaea 12 (1838) 271. –– Type: Mexico,
near Hacienda de la Laguna, August 1829, Schiede
s.n. (Lectotype HAL [HAL0069536], designated
by Raven, Reinwardtia 6 (1964 [‘1963’]) 358).
Jussiaea occidentalis Nutt.ex Torr. & A.Gray., Fl.
N. Am. 1 (3) (1840) 521. –– Type: USA, Arkansas,
Nuttall s.n. (Lectotype PH [PH00016619],
designated by Raven, Reinwardtia 6 (1964
[‘1963’]) 358).
Jussiaea linearis Hochst., Flora 27 (1844) 425,
nom. illegit., non Willd. (1799). – Jussiaea
angustifolia var. linearis Harv. in Harvey & Sond.,
Fl. Capensis. 2 (1862) 504. –– Type: South Africa,
Port Natal, Krauss 73 (Lectotype K [K000310870],
designated here).
Jussiaea sagrana A.Rich. in de la Sagra, Hist.
Phys. Cuba, Pl. Vasc. (1846) 534, as ‘sagraeana’. –
Ludwigia sagrana (A.Rich.) M.Gómez, Anales
Soc. Esp. Hist. Nat. 23 (1894) 66. –– Type: Ramón
de la Sagra s.n. (Lectotype P [P01819476],
designated by Raven, Reinwardtia 6 (1964
[‘1963’]) 358).
TURNER: An updated synopsis of Ludwigia in Malesia
2021] 13
Jussiaea velutina Kunze, Linnaea 20 (1847) 56,
nom. illegit., non G.Don (1832). –– Type: Austria,
Beschke, cultivated (LZ†).
Jussiaea costata C.Presl, Epim. Bot. (1849) 217.
–– Type: Philippines, Luzon, Laguna Prov., Cala-
wang, Cuming 655 (Lectotype BM
[BM000984335], designated here; isolectotypes K,
L).
Jussiaea junghuhniana Miq., Fl. Ned. Ind. 1 (1)
(1856) 627. –– Type: Sumatra, Padang, bij Lumut,
October [sine anno], Junghuhn s.n. (Lectotype L
[L0008657], designated by Raven, Reinwardtia 6
(1964 [‘1963’]) 363).
Jussiaea octonervia forma sessiliflora Micheli in
Martius, Fl. Bras. 13 (2) (1875) 171. – Jussiaea
suffruticosa var. sessiliflora (Micheli) Hassl., Bull.
Soc. Bot. Genève, ser. 2, 5 (1913) 271. – Ludwigia
pubescens var. sessiliflora (Micheli) H.Hara, J.
Jap. Bot. 28 (1953) 293. – Ludwigia octovalvis
subsp. sessiliflora (Micheli) P.H.Raven, Kew Bull.
15 (1962) 476. – Ludwigia octovalvis var.
sessiliflora (Micheli) Shinners, Sida 1 (1964) 385.
–– Type: Brazil, Rio de Janeiro, Burchell 927
(Lectotype K [K000911823], designated by Munz,
Darwiniana 4 (1942) 237; Isolectotype GH
[GH00054202]).
Jussiaea suffruticosa var. subglabra Thwaites ex
Trimen, Handb. Fl. Ceylon 2 (1894) 233. –– Type:
Ceylon, Thwaites 170 (not traced).
Jussiaea suffruticosa var. angustifolia Chod. &
Hassler, Bull. Herb. Boiss. sér. 2, 3 (1903) 909,
nom. illegit., non (Lam.) F.M.Bailey (1883). ––
Type: Paraguay, Igatimi, November 1898, Has-
sler 5541 (Holotype G [G00442815] 2 sheets).
Jussiaea suffruticosa var. sintenisii Urb., Symb.
Antill. 4 (1910) 469. –– Type: Puerto Rico, prope
Hato Grande in collibus graminosus ad montem
“Gregorio” versus, 31 August 1885, Sintenis 2719
(Lectotype B [B 10 0367942], designated by
Munz, Darwiniana 4 (1942) 242).
Jussiaea suffruticosa var. linearifolia Hassl., Rep-
ert. Spec. Nov. Regni Veg. 12 (1913) 277. – Jus-
siaea suffruticosa forma linearifolia (Hassl.)
Munz, Darwiniana 4 (1942) 243. – Ludwigia pu-
bescens var. linearifolia (Hassl.) A.Fern. &
R.Fern., Garcia de Orta 5 (1957) 115. –– Type:
Paraguay, Camp San Luis, Fiebrig 4128
(Lectotype G [G00442814], designated by Raven,
Reinwardtia 6 (1964 [‘1963’]) 359).
Jussaia suffruticosa var. hirta Ridl., Trans. Linn.
Soc. Bot. 9 (1916) 57. –– Type: New Guinea,
Camp VIII to IX, C. Boden Kloss s.n. (Lectotype
K, designated by Raven, Reinwardtia 6 (1963)
363; Isolectotype BM [BM013717213]).
Jussiaea suffruticosa var. hawaiensis Hochr., Can-
dollea 2 (1925) 479. –– Type: Iles Sandwich
[Hawaii], Kauai, Weimea-village, 21 April 1905,
Hochreutiner 3592 (Lectotype G [G00442813],
designated by Raven, Reinwardtia 6 (1963) 359.
Jussiaea suffruticosa var. samoensis Hochr., Can-
dollea 2 (1925) 479. –– Type: Hochreutiner 3280,
Samoa, île d’Upolu, lac de Lanuto, 22 March
1905 (Lectotype G [G00442809], designated
by Raven, Reinwardtia 6 (1964 [‘1963’]) 359.
Jussiaea clavata M.E.Jones, Contr. W. Bot. 15
(1929) 131. –– Type: Mexico, Acaponeta, Naya-
rit, 24 February 1927, M.E. Jones 22871
(Lectotype WIS [WISv0255953WIS], designated
here).
Jussiaea suffruticosa auctt. plur., non L.
Description and distribution. Raven (1964:
356), Vu & Vidal (1973: 20), Khan &
Shamsunnahar (1977: 6), Raven (1977: 101),
Kostermans et al. (1987: 372), Thompson (1990:
226), Wagner (1995: 349), Barua (2010: 65),
Boufford & Raven (2014: 603), Ummul-Nazrah
(2017: 58).
Notes. Raven (1964) considered the pantropical
Ludwigia octovalvis to be represented by four
subspecies, two of which (subsp. octovalvis and
subsp. sessiliflora (Micheli) P.H.Raven) occurred
in Malesia. However, Raven (1977) retracted this
infraspecific classification in favour of one broad-
ly defined and variable species. The synonymy
above only includes names relevant to the two
former subspecies recognised for Malesia. When
Bailey published Jussiaea suffruticosa var.
angustifolia and var. villosa he provided a brief
description of each variety, but did not cite any
potential basionyms for new combinations. Later
however, Bailey (1913: 215) did cite Jussiaea
angustifolia Lam. and J. villosa Wight & Arn. as
synonyms of his varieties. The first can be taken
directly as a basionym, the second provides a
direct citation of J. villosa Lam. I therefore
consider that it was Bailey’s intention to make
new combinations at varietal rank when he
published these names and take them as such
based on the Lamarck species names.
5. LUDWIGIA PERENNIS L.
Ludwigia perennis L., Sp. Pl. 1 (1753) 119. –
Jussiaea perennis (L.) Brenan, Kew Bull. 8
(1953) 163. –– Type: Herb. Hermann 2: 9, No. 66
(Lectotype BM [BM000621529], designated by
Brenan in Turrill & Milne-Redhead, Fl. Trop. E.
Africa, Onagraceae (1953) 13).
Jussiaea suffruticosa L., Sp. Pl. (1753) 388. –
Ludwigia suffruticosa (L.) M.Gómez, Anales Soc.
Esp. Hist. Nat. 23 (1894) 66, nom. illegit., non
Walter (1788). –– Type: [published illustration]
‘Carambu’ in Rheede, Hort. Malab. 2 (1679) t. 49
(Lectotype designated here).
Ludwigia oppositifolia L., Syst. Nat., ed. 12, 2
(1767) 125. –– Type: Herb. Hermann 2: 9, No. 66
(Lectotype BM [BM000621529], designated
here).
REINWARDTIA 14 [VOL.20
Jussiaea caryophyllaea Lam., Encycl. 3 (1789)
331. – Ludwigia caryophyllaea (Lam.,) Merr. &
F.P.Metcalf, Lingnan Sci. J. 16 (1937) 396. ––
Type: India, Sonnerat s.n. (Lectotype P-LA [IDC
243/7], designated by Raven, Reinwardtia 6 (1964
[‘1963’]) 367).
Ludwigia parviflora Roxb. Fl. Ind. 1 (1820) 440.
1820. – Isnardia parviflora (Roxb.) Kuntze,
Revis. Gen. Pl. 1 (1891) 250. –– Type:
[unpublished illustration] Icones Roxburghianae
1340 (Lectotype K, designated here).
Ludwigia lythroides Blume, Bijdr. Fl. Ned. Ind.
(17) (1826-1827) 1134. –– Type: Java, Batavia,
Blume 1132 (Lectotype L [L0008663], designated
by Raven, Reinwardtia 6 (1964 [‘1963’]) 367).
Isnardia multiflora Guill. & Perr., Fl. Senegamb.
Tent. (1838) 295. – Ludwigia multiflora (Guill. &
Perr.) Walp., Repert. Bot. Syst. 2 (1843) 75. ––
Type: Sénegambie, Richardtal, 1824, Leprieur s.n.
(Lectotype G [G00383323], designated by Brenan
in Turrill & Milne-Redhead, Fl. Trop. E. Africa,
Onagraceae (1953) 13; possible Isolectotype P
[P00412956]).
Isnardia diffusa Oken, Allg. Naturgesch. 3(3)
(1841) 1874. –– Type: [published illustration]
‘Carambu’ in Rheede, Hort. Malab. 2 (1679) t. 49
(Lectotype designated here).
Ludwigia gracilis Miq., Fl. Ned. Ind. 1 (1) (1856)
629. –– Type: Sumatra, Padang, Junghuhn s.n.
(Lectotype L [L0008662], designated by Raven,
Reinwardtia 6 (1964 [‘1963’]) 367).
Ludwigia nesaeoides H.Perr., Not. Syst. 13 (1947)
141. –– Type: Madagascar, Boina, near Marovo-
ay, May 1922, H. Perrier de la Bâthie 17965
(Lectotype P [P00412981], designated at the first
step by Raven, Reinwardtia 6 (1964 [‘1963’]) 368
and at the second step here; Isolectotype P
[P00412982]).
Ludwigia humbertii Robyns & Lawalrée, Bull.
Jard. Bot. État Bruxelles 18 (1947) 291. –– Type:
Congo, south of Lake Edward, Apr-May 1929, H.
Humbert 8709 (Lectotype BR
[BR000000897374], designated by Raven, Rein-
wardtia 6 (1964 [‘1963’]) 368; Isolectotype P
[P00412957]).
Description and distribution. Raven (1964:
367), Vu & Vidal (1973: 27), Khan & Shamsun-
nahar (1977: 7), Raven (1977: 103), Kostermans
et al. (1987: 374), Thompson (1990: 227), Wagner
(1995: 338), Barua (2010: 65), Boufford & Raven
(2014: 605), Ummul-Nazrah (2017: 50).
Notes. Jussiaea suffruticosa L. was, for many
years, the accepted name for what is now called
Ludwigia octovalvis. In the protologue Linnaeus
gave a brief description, ‘Jussiaea erecta villosa,
floribus tetrapetalis octandris pedunculatis’, cited
a few references and stated ‘Habitat in India’. The
only cited element eligible as type material is
Rheede’s Carambu, more specifically Rheede’s t.
49. This is generally identified as Ludwigia
perennis L. and was cited by Linnaeus in the proto-
logue of that name. Ludwigia perrenis is notable
for having four stamens, rather than the eight noted
in Linnaeus’s description of Jussiaea suffruticosa.
It has been suggested that Linnaeus really intended
to cite Rheede’s ‘Cattu-carambu’ (Rheede, Hort.
Malab. 2 (1679) t. 50) (Nicolson et al., 1988), but
there is no evidence of Linnaeus correcting himself
in this regard in later publications. Fortunately, as
Ludwigia suffruticosa Walt. makes the combina-
tion in Ludwigia for Jussiaea suffruticosa illegiti-
mate, there are no nomenclaturally disruptive con-
sequences from typifying Jussiaea suffruticosa on
t. 49. Given the lack of an alternative under current
rules, this is done here.
Linnaeus appears to have changed his mind
about the name Ludwigia perennis and used
Ludwigia oppositifolia from 1767, citing the
former as a synonym of the latter in 1774. While
Ludwigia oppositifolia L. has been referred to as
an illegitimate superfluous name it fails to meet the
requirements of the code for being superfluous
(Art. 52). In the protologue Linnaeus did not cite
Ludwigia perennis, or all the elements cited under
that name, or use the exact phrase name. There-
fore, as it is can clearly be inferred that Linnaeus
was basing L. oppositifolia on the same elements
as L. perennis, I here lectotypify it to the same
specimen as L. perennis.
6. LUDWIGIA PERUVIANA (L.) H.Hara
Ludwigia peruviana (L.) H.Hara, J. Jap. Bot. 28
(1953) 293. – Jussiaea peruviana L., Sp. Pl. (1753)
388. – Jussiaea grandiflora Ruiz & Pav., Anales
Inst. Bot. Cavanilles 14 (1956) 753, nom. illegit.,
superfl., non Michx. (1803). –– Type: [published
illustration] "Onagra Laurifolia, flore amplo,
pentapetalo" in Feuillée, J. Obs. 2 (1714) t. 9
(Lectotype designated by Raven, Reinwardtia 6
(1964 [‘1963’]) 346).
Oenothera hirta L., Syst. Nat. ed. 10, 2 (1759) 998.
– Jussiaea hirta (L.) Sw., Obs. Bot. (1791) 142,
nom. illegit., non Lam. (1789). – Ludwigia hirta
(L.) M.Gómez, Anales Soc. Esp. Hist. Nat. 23
(1894) 66. –– Type: [published illustration]
‘Oenothera hirsuta’ of Plumier in Burman, Pl.
Amer. 8 (1758) 167 (Lectotype designated by
Ramamoorthy & Zardini, Monogr. Syst. Bot.
Missouri Bot. Gard. 19 (1987) 29).
Jussiaea macrocarpa Kunth, Nov. Gen. Sp. 6
(1823) 102. – Jussiaea peruviana var. macrocarpa
(Kunth) Bertoni, Descr. Fis. Econ. Paraguay
(1910) 13. –– Type: Colombia, Guaduas,
Humboldt & Bonpland 1758 (Lectotype P-Bonpl.
[P00679566], designated at the first step by Raven,
Reinwardtia 6 (1964 [‘1963’]) 345), and at the
second step here; Isolectotype P [P00135180]).
TURNER: An updated synopsis of Ludwigia in Malesia
2021] 15
Jussiaea mollis Kunth, Nov. Gen. Sp. 6 (1823)
102. –– Type: Venezuela, Sucre, Bordones, Hum-
boldt & Bonpland 414 (Lectotype P-Bonpl.
[P00679567], designated by Raven, Reinwardtia 6
(1964 [‘1963’]) 345).
Jussiaea hirsuta Vell., Fl. Flumin. (1829) 186,
nom. illegit., non Miller (1768). –– Type:
[unpublished illustration] ‘Iussieva hir-
suta’ [original drawing for Vellozo, Fl. Flumin.
Icon. 4 (1831) t. 144] (Lectotype Biblioteca
Nacional Manuscript Section mss1198653,
designated here).
Jussiaea peruviana var. australis Hassl., Repert.
Spec. Nov. Regni Veg. 12 (1913) 369. –– Type:
Paraguay, in regione fluminis Alto Paraná,
1909/1910, Fiebrig 6063 (Lectotype B
[B100249031], designated here; isolectotypes G,
SI, US).
Jussiaea peruviana var. glaberrima Donn.Sm.,
Bot. Gaz. 16 (1891) 6. –– Type: Guatemala, De-
part. Zacatepequez, Dueñas, April 1890, J. Donnell
Smith 2130 (Lectotype US [US00123985], desig-
nated by Ramamoorthy & Zardini, Monogr. Syst.
Bot. Missouri Bot. Gard. 19 (1987) 30; Isolecto-
types GH, K [K000533294], NY).
Jussiaea peruviana f. hirsuta Hassl., Repert. Spec.
Nov. Regni Veg. 12 (1913) 269. –– Type: Para-
guay, in viciniis Caaguazú. Paraguay, Hassler
9167 (Lectotype B [B100249033], designated by
Raven, Reinwardtia 6 (1964 [‘1963’]) 345).
Jussiaea peruviana f. tomentosa Hassl., Repert.
Spec. Nov. Regni Veg. 12 (1913) 269. –– Type:
Paraguay, in altoplanitie et decliviis Sierra de Ma-
racayú, Hassler 5011 (Lectotype B [B100249032],
designated here; isolectotypes G, GH, P, POM).
Jussiaea speciosa Ridl., J. Bot. 59 (1921) 259. ––
Type: India, Madras, Vellangiri Hills, Devala, No-
vember 1886, Gamble 18361 (Lectotype BM
[BM01371212], designated by Raven, Reinwardtia
6 (1964 [‘1963’]) 345; Isolectotype K
[K001129807]).
Jussiaea sprengeri L.H.Bailey, Stand. Cycl. Hort.
3 (1915) 1730. –– Type: USA, in cultivation, John-
son s.n. (Lectotype POM, designated by Rama-
moorthy & Zardini, Monogr. Syst. Bot. Missouri
Bot. Gard. 19 (1987) 31).
Description and distribution. Raven (1964: 345),
Raven (1977: 100), Kostermans et al. (1987: 376),
Ramamoorthy & Zardini (1987: 29), Thompson
(1990: 226), Wagner (1995: 334), Barua (2010:
66), Boufford & Raven (2014: 606), Ummul-
Nazrah (2017: 60).
Notes. Raven (1964) cited the type of Jussiaea
mollis Kunth as no. 878. I take this as a
typographic error and correct it to 414. Although
Ummul-Nazrah (2017) noted the occurrence of
Ludwigia peruviana in Sabah and Sarawak, she did
not cite any specimens. I have seen collections
from Kalimantan (Leeuwenberg 13381, Ambri &
Arifin A.A.455, Arbainsyah AA1861) that confirm
the species is naturalized in Borneo.
7. LUDWIGIA PROSTRATA Roxb.
Ludwigia prostrata Roxb., Fl. Ind. 1 (1820) 441. –
Nematopyxis prostrata (Roxb.) Miq., Fl. Ned. Ind.
1(1) (1856) 630. – Isnardia prostrata (Roxb.)
Kuntze, Revis. Gen. Pl. 1 (1891) 250. – Jussiaea
prostrata (Roxb.) H.Lév., Repert. Spec. Nov.
Regni Veg. 8 (1910) 138. –– Type: [unpublished
illustration] Icones Roxburghianae 1945
(Lectotype K, designated by Vu & Vidal, Fl.
Cambodge, Laos & Vietnam 14 (1973) 30).
Ludwigia diffusa Buch.-Ham., Trans. Linn. Soc.
London 14 (1824) 301. –– Type: India, Goyalpara,
2 September 1808, F. Buchanan-Hamilton s.n.
[EIC 6336A] (Lectotype K-W [K001123615],
designated here).
Ludwigia fruticulosa Blume, Bijdr. Fl. Ned. Ind.
(1827) 1133. – Nematopyxis fruticulosa (Blume)
Miq., Fl. Ned. Ind. 1 (1) (1856) 630. – Type: Java,
Blume s.n. (Lectotype L [L0008664], designated
by Raven, Reinwardtia 6 (1964 [‘1963’]) 374).
Ludwigia leucorhiza Blume, Bijdr. Fl. Ned. Ind.
(1827) 1133. –– Type: Java, circa Buitenzorg ad
flumen Tjeliwong, Blume s.n. (not traced).
Nematopyxis pusilla Miq., Fl. Ned. Ind. 1 (1)
(1856) 630. –– Type: Java, Horsfield s.n.
(Holotype K [K000742277]).
Description and distribution. Raven (1964:
374), Vu & Vidal (1973: 30), Khan & Shamsun-
nahar (1977: 9), Raven (1977: 103), Wagner
(1995: 339), Barua (2010: 68), Boufford & Raven
(2014: 607), Ummul-Nazrah (2017: 60).
Species excludenda
Ludwigia decurrens Walt.
Ramamoorthy & Zardini (1987: 95) reported a
collection of Ludwigia decurrens Walt. from near
Laguna in the Philippines (Parchs 6608 (MO)). I
have not found any more herbarium material of
this species from Malesia and it does not appear to
have naturalised in the Philippines (R. Joshi pers.
comm., R. Lubigan pers. comm.).
ACKNOWLEDGEMENTS
The following are thanked for their invaluable
assistance in searching for type material: Marc
Appelhans, Anand Kumar, Otakar Šida and Jim
Solomon. Dr. Nigel Tucker of Biotropica
Australia Pty. Ltd. generously supplied infor-
mation on Ludwigia leptocarpa in Papua New
Guinea. David Ples, Dr. Ravindra Joshi and Ross
Lubigan very kindly responded to my queries on
Philippines Ludwigia during a particularly
difficult period in that country. Prof. Marco Pelle-
REINWARDTIA 16 [VOL.20
grini’s assistance with typifying Vellozo names
was greatly appreciated. Charlie Jarvis provided
guidance with matters relating to the Linnaean
names.
REFERENCES
BAILEY, F. M. 1913. Comprehensive Catalogue
of Queensland Plants both Indigenous and
Naturalised. A. J. Cumming, Brisbane.
BARUA, I. C. 2020. The genus Ludwigia
(Onagraceae) in India. Rheedea 20: 59‒70.
BOUFFORD, D. & RAVEN, P. H. 2014.
Onagraceae. Fl. Thail. 11(4): 598‒607.
KHAN, M. S. & SHAMSHUNNAHAR. 1977.
Onagraceae. Flora of Bangladesh Fasc. 6: 1‒
10. Bangladesh National Herbarium and
Bangladesh Agricultural Research Council.
KOSTERMANS, A. J. G. H., WIRJAHARDJA, S.
& DEKKER, R. J. 1987. The weeds:
description, ecology and control. In:
SOERJANI, M., KOSTERMANS, A. J. G. H.
& TJITROSOEPOMO, G. (Eds.). Weeds of
Rice in Indonesia. Balai Pustaka, Jakarta.
Pp. 24‒565.
LIU, S.-H., HOCH, P. C., DIAZGRANADOS, M.,
RAVEN, P. H. & BARBER, J. C. 2017.
Multi-locus phylogeny of Ludwigia
(Onagraceae): insights on infra-generic
relationships and current classification of the
genus. Taxon 66: 1112‒1127.
NICOLSON, D. H., SURESH, C. R. & MA-
NILAL, K. S. 1988. An Interpretation of Van
Rheede’s Hortus Malabaricus. Regnum. Veg.
119: 1‒378.
RAMAMOORTHY, T. P. & ZARDINI, E. M.
1987. The systematics and evolution of Lud-
wigia sect. Myrtocarpus sensu lato
(Onagraceae). Monogr. Syst. Bot. Missouri Bot.
Gard. 19: 1‒120.
RAVEN, P. H. (1964 [‘1963’]). The Old World
species of Ludwigia (including Jussiaea), with a
synopsis of the genus (Onagraceae).
Reinwardtia 6: 327‒427.
RAVEN, P. H. 1977. Onagraceae. Fl. Males., ser.
I, 8: 98‒113.
THOMPSON, J. 1990. Onagraceae. Fl. Austral.
18: 215‒243.
TUCKER, N. I. J. 2017. Exotic Plants of the Kikori
River Basin. Second Edition. National
Quarantine Inspection Authority, Port Moresby.
UMMUL-NAZRAH, A. R. 2017. Onagraceae. Fl.
Penins. Malaysia, Ser. II, 6: 49‒62.
VU, V. C. & VIDAL, J. E. 1973. Onagraceae. Fl.
Cambodge, Laos, Vietnam 14: 17‒39.
WAGNER, W. L. 1995. Onagraceae. Rev. Handb.
Fl. Ceylon 9: 332‒350.
WAGNER, W. L., HOCH, P. C. & RAVEN, P. H.
2007. Revised classification of the Onagraceae.
Syst. Bot. Monogr. 83: 1‒240.