Physalis peruviana L., Solanaceae

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Present on Pacific Islands?  yes

Primarily a threat at high elevations?  no

Common name(s): [more details]

Chinese: deng long guo

English: bladderberry, cape gooseberry, goldenberry, gooseberry tomato, obra cabbage, Peruvian cherry, Peruvian ground-cherry

Fijian: botebote yadra, kospeli, maulanggua, mbotembote yandra, tukiyandra

French: alkékenge du Pérou, capuli, coqueret comestible, coqueret du Pérou, coqueret du Peru, groseiller canaque, groseiller du cap

Hawaiian: pa‘ina, pohā

I-Kiribati: te baraki, te bin

Mangarevan: maru

Maori (Cook Islands): tupera, tūpere, tūpere

Nauruan: oatamo, watamo

Niuean: manini, manini fua lalahi

Other: goundou-goundou, winebusupén

Spanish: alquequenje, capulí, uvilla, uvilla, uvilla

Tahitian: tupere

Tongan: ku‘usi

Habit:  shrub

Description:  "Soft-wooded, short-lived shrubs up to ca. 1 m tall, straggly with age, all parts densely pubescent with erect, simple or glandular hairs up to 1 mm long.  Leaves simple, alternate, usually geminate, 1 larger than the other, ovate-acuminate, often 6 cm long, 4 cm wide, margins entire or rarely with a few blunt lobes, apex acuminate, base cordate, petioles 2-3 cm long.  Flowers perfect, actinomorphic, solitary in the leaf axils, pedicellate; calyx connate in lower ", 5-lobed, veins often prominent, the lobes acumunate-triangular, ca. 1 cm long, distinct at apex; corolla yellow with well-defined purplish brown spots at base, 15-20 mm in diameter, the limb rotate or shallowly 10-lobed, the tube swollen into shallow nectary pouches between the filaments, densely pubescent with pale yellowish dendritic hairs below the spots and around the nectaries; style 5-7 mm long.  Berries pale yellow, drying pale brown, aromatic, succulent, globose, 1.5-2 cm in diameter, enclosed in the inflated calyx 3-3.5 cm long.  Seeds numerous, pale brown, discoid, 1.75-2 mm long, minutely shallowly reticulate, embryo curved, endosperm present" (Wagner et al., 1999; p. 1265).

Habitat/ecology:  "Forests and forest edges, riparian habitats, disturbed sites.  This large herb forms dense thickets that crowd out native vegetation.  Littlee is known on the ecology of this species"  (Weber, 2003; p. 322).

In Hawai‘i, "naturalized in disturbed sites in mesic to wet forest, diverse mesic forest, and subalpine woodland, 450-2,020 m" (Wagner et al., 1999; p. 1265).  In Fiji, "seen at elevations from near sea level to 900 m in gardens and also naturalized in forest along trails and streams, in clearing, and in cultivated areas" (Smith, 1991; p. 29).  In Niue, "a common weed in some plantations" (Sykes, 1970; p. 197).  "Uncommon in plantations and waste places but sometimes collected for food" (Whistler, 1988; p. 36). In Tahiti, "frequent in all the cool valleys to 800 m" (Welsh, 1998; p. 273). In Tonga, "occasional as a plantation weed" (Yuncker, 1959; p. 237). In New Caledonia, "peu cultivé malgré les fruits comestibles; largement répandu comme mauvaise herbe" (MacKee, 1994; p. 131).

Propagation:  Seed

Native range:  South America (Smith, 1991; p. 29).

Presence:

Pacific
Country/Terr./St. & Island group	Location	Cited status & Cited as invasive & Cited as cultivated & Cited as aboriginal introduction?	Reference & Comments
Australia (Pacific offshore islands) Lord Howe Island	Lord Howe Island	introduced	Pickard, J. (1984) (p. 208)
Australia (Pacific offshore islands) Lord Howe Island	Lord Howe Island	introduced invasive	Orchard, Anthony E., ed. (1994) (p. 21) "An occasional weed of wasteland and open areas". Vouchers cited: G. Uhe 1282 (K), L.A.S. Johnson & A.N. Rodd 1228 (NSW), J. Pickard 1480 (NSW), J. Pickard 2828 (NSW)
Australia (Pacific offshore islands) Norfolk Islands	Norfolk Island	introduced invasive	Orchard, Anthony E., ed. (1994) (p. 10) "An occasional weed of wasteland and open areas". Vouchers cited: P.S. Green 1394 (A), W.R. Sykes NI 606 (CHR)
Chile (offshore islands) Juan Fernández Islands	Isla Más a Tierra (Robinson Crusoe Island)	introduced	Skottsberg, Carl (1953) (p. 223)
Chile (offshore islands) Juan Fernández Islands	Isla Más a Tierra (Robinson Crusoe Island)	introduced invasive cultivated	Atkinson, Rachel/Sawyer, John (2011)
Chile (offshore islands) Juan Fernández Islands	Isla Más a Tierra (Robinson Crusoe Island)	introduced invasive	Danton, Philippe/Perrier, Christophe/Martinez Reyes, Guido (2006) (p. 556)
Chile (offshore islands) Juan Fernández Islands	Isla Más Afuera (Alejandro Selkirk Island)	introduced	Skottsberg, Carl (1953) (p. 223) Widely spread.
Chile (offshore islands) Juan Fernández Islands	Isla Más Afuera (Alejandro Selkirk Island)	introduced invasive cultivated	Atkinson, Rachel/Sawyer, John (2011)
Chile (offshore islands) Juan Fernández Islands	Isla Más Afuera (Alejandro Selkirk Island)	introduced invasive	Danton, Philippe/Perrier, Christophe/Martinez Reyes, Guido (2006) (p. 556)
Commonwealth of the Northern Mariana Islands (US) Northern Mariana Islands	Saipan Island	introduced	Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce (1979) (p. 245)
Cook Islands Southern Cook Islands	‘Atiu Island		McCormack, Gerald (2011)
Cook Islands Southern Cook Islands	Mangaia Island		McCormack, Gerald (2011)
Cook Islands Southern Cook Islands	Ma‘uke Island		McCormack, Gerald (2011)
Ecuador (Galápagos Islands) Isabela Group	Isabela Island	introduced invasive cultivated	Charles Darwin Foundation (2008)
Ecuador (Galápagos Islands) Isabela Group	Volcán Sierra Negra, Isabela Island	introduced invasive cultivated	Charles Darwin Foundation (2008)
Ecuador (Galápagos Islands) San Cristóbal Group	San Cristóbal Island	introduced invasive cultivated	Charles Darwin Foundation (2008)
Ecuador (Galápagos Islands) Santa Cruz Group	Santa Cruz Island	introduced invasive cultivated	Charles Darwin Foundation (2008)
Fiji Fiji Islands	Taveuni Island	introduced invasive	Smith, Albert C. (1991) (p. 29) Voucher cited: Seemann 338
Fiji Fiji Islands	Viti Levu Island	introduced invasive	Smith, Albert C. (1991) (p. 29) Vouchers cited: Greenwood 802, Parks 20575, Gillespie 4304, Gibbs 707, DA 16704, Smith 8692, DA 11437
French Polynesia Gambier Islands	Mangareva (Gambier) Island	introduced invasive	Florence, J./Chevillotte, H./Ollier, C./Meyer, J.-Y. (2011) Adventice
French Polynesia Gambier Islands	Mangareva (Gambier) Island	introduced	St. John, Harold (1988) (p. 23) Vouchers cited: St. John 14861 (BISH), St. John 14874 (BISH)
French Polynesia Marquesas Islands	Hiva Oa Island	introduced invasive	Lorence, David H./Wagner, Warren L. (2008)
French Polynesia Marquesas Islands	Hiva Oa Island	introduced invasive	Florence, J./Chevillotte, H./Ollier, C./Meyer, J.-Y. (2011) Adventice
French Polynesia Marquesas Islands	Nuku Hiva (Nukahiva) Island	introduced invasive	Lorence, David H./Wagner, Warren L. (2008)
French Polynesia Marquesas Islands	Nuku Hiva (Nukahiva) Island	introduced invasive	Florence, J./Chevillotte, H./Ollier, C./Meyer, J.-Y. (2011) Voucher cited: J. Florence 6899 (PAP) Adventice
French Polynesia Society Islands	Raiatea (Havai) Island		Welsh, S. L. (1998)
French Polynesia Society Islands	Tahiti Island		Welsh, S. L. (1998)
French Polynesia Society Islands	Tahiti Island	introduced invasive	Florence, J./Chevillotte, H./Ollier, C./Meyer, J.-Y. (2011) Voucher cited: J. Florence 6964 (PAP) Adventice
French Polynesia Austral (Tubuai) Islands	Tubuai Island	introduced invasive	Florence, J./Chevillotte, H./Ollier, C./Meyer, J.-Y. (2011) Adventice
Guam Guam Island	Guam Island	introduced	Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce (1979) (p. 245)
State of Hawaii Hawaiian Islands	Hawai‘i (Big) Island	introduced invasive cultivated	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (pp. 1265-1266)
State of Hawaii Hawaiian Islands	Kaua‘i Island	introduced invasive cultivated	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (pp. 1265-1266)
State of Hawaii Hawaiian Islands	Lāna‘i Island	introduced invasive cultivated	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (pp. 1265-1266)
State of Hawaii Hawaiian Islands	Maui Island	introduced invasive cultivated	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (pp. 1265-1266)
State of Hawaii Hawaiian Islands	Moloka‘i Island	introduced invasive cultivated	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (pp. 1265-1266)
State of Hawaii Hawaiian Islands	O‘ahu Island	introduced invasive cultivated	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (pp. 1265-1266)
Japan (offshore islands) Bonin (Ogasawara) Islands	Bonin (Ogasawara) Islands	introduced invasive	Toyoda, Takeshi (2003) (p. 288) Naturalized
Japan (offshore islands) Bonin (Ogasawara) Islands	Bonin (Ogasawara) Islands	introduced	Kato, Hidetoshi (2007)
Kiribati Tungaru (Gilbert) Islands	Butaritari Atoll	introduced	Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce (1979) (p. 245)
Nauru Nauru Island	Nauru Island		Swarbrick, John T. (1997) (p. 95)
New Caledonia Îles Loyauté (Loyalte Islands)	Îles Loyauté (Loyalty Islands)	introduced invasive	Tassin, Jacques (2005)
New Caledonia Îles Loyauté (Loyalte Islands)	Île Lifou	introduced invasive	MacKee, H. S. (1994) (p. 131) Vouchers cited: Deplanche s.n., Schmid 1328 (NOU)
New Caledonia Îles Loyauté (Loyalte Islands)	Îles Ouvéa (Ouvea Atoll)	introduced invasive	MacKee, H. S. (1994) (p. 131) Voucher cited: Däniker 1997
New Caledonia New Caledonia Archipelago	Île Grande Terre	introduced invasive	Tassin, Jacques (2005)
New Caledonia New Caledonia Archipelago	Île Grande Terre	introduced invasive	MacKee, H. S. (1994) (p. 131) Vouchers cited: Balansa 525, Brousmiche 416, Cribs 1224, Gaillard in MacKee 26921
New Caledonia New Caledonia Archipelago	Île des Pins (Isle of Pines)	introduced invasive	Tassin, Jacques (2005)
New Caledonia New Caledonia Archipelago	Île des Pins (Isle of Pines)	introduced invasive	MacKee, H. S. (1994) (p. 131) Vouchers cited: MacKee 15801, MacKee 23509
New Zealand (offshore islands) Kermadec Islands	Raoul Island	introduced invasive	Sykes, W. R. (1977) (p. 143)
New Zealand (offshore islands) Kermadec Islands	Raoul Island	introduced invasive	Webb, C. J./Sykes, W. R./Garnock-Jones, P. J. (1988) (p. 1234) "Open habitats such as roadsides, waste places, beaches, freshwater mudflats and cultivated ground".
Niue Niue	Niue Island	introduced invasive	Sykes, W. R. (1970) (p. 197) Voucher cited: CHR 170265
Niue Niue	Niue Island	introduced invasive	Whistler, W. A. (1988) (p. 36)
Niue Niue	Niue Island	introduced invasive	Yuncker, T. G. (1943) (p. 105) Vouchers cited: Yuncker 9759 (BISH), Yuncker 10088 (BISH) "Occasional in waste areas and plantations".
Niue Niue	Niue Island	introduced invasive	Space, James C./Waterhouse, Barbara M./Newfield, Melanie/Bull, Cate (2004) (p. 45)
Palau Palau (main island group)	Koror Island		Beleu National Museum (Republic of Palau. Koror.) (1966) (voucher ID: BNM 753) Taxon name on voucher: Physalis peruviana L.
Philippines Philippine Islands	Philippine Islands	introduced invasive	Merrill, Elmer D. (1923) (p. 423) Thoroughly established in open waste places.
Pitcairn Islands Pitcairn Islands	Pitcairn Island	introduced	St. John, H. (1987) (p. 49) Vouchers cited: R. A. Young 30, St. John 14973
Samoa Western Samoa Islands	Western Samoa Islands		Whistler, W. A. (1988) (p. 36)
Tonga Tongatapu Group	‘Eua Island		Whistler, W. A. (1992) (p. 73) Vouchers cited: Parks 16327, Soakai 371
Tonga Tongatapu Group	Tongatapu Island		Whistler, W. A. (1992) (p. 73) Vouchers cited: Soakai 250, Soakai 742
Tonga Tongatapu Group	Tongatapu Island	introduced invasive	Yuncker, T. G. (1959) (p. 237) Voucher cited: Yuncker 15087
Tonga Vava‘u Group	Vava‘u Island	introduced invasive	Yuncker, T. G. (1959) (p. 237) Cited: Crosby.
Tonga Vava‘u Group	Vava‘u Island		Burkill, I. H. (1900) (p. 49)
Tuvalu Tuvalu (Ellis) Islands	Tuvalu Islands		Swarbrick, John T. (1997) (p. 95)

Pacific Rim
Country/Terr./St. & Island group	Location	Cited status & Cited as invasive & Cited as cultivated & Cited as aboriginal introduction?	Reference & Comments
Australia Australia (continental)	Australia (continental)	introduced invasive	Weber, Ewald (2003) (p. 322)
Chile (continental) Chile	Chile (Republic of)		Skottsberg, Carl (1953) (p. 223)
China China	China (People's Republic of)	introduced invasive	Zhengyi, Wu/Raven, Peter H./Deyuan, Hong (2011) Near roads, river valleys, cultivated and naturalized; 1200-2100 m. Fujian, Guangdong, Jiangsu, Yunnan.
Colombia Colombia	Colombia (Republic of)	native	U.S. Dept. Agr., Agr. Res. Serv. (2011)
Ecuador (Mainland) Ecuador	Ecuador (Republic of) (continental)	native	U.S. Dept. Agr., Agr. Res. Serv. (2011)
Indonesia Indonesia	Indonesia (Republic of)		Holm, Leroy/Pancho, Juan V./Herberger, James P./Plucknett, Donald L. (1979) (p. 279)
Japan Japan	Japan (country)	introduced	Mito, Toshikazu/Uesugi, Tetsuro (2004) (p. 188)
New Zealand New Zealand	New Zealand (country)	introduced invasive cultivated	Webb, C. J./Sykes, W. R./Garnock-Jones, P. J. (1988) (p. 1234) "Open habitats such as roadsides, waste places, beaches, freshwater mudflats and cultivated ground".
Perú Perú	Perú (Republic of)	native	U.S. Dept. Agr., Agr. Res. Serv. (2011)
Taiwan Taiwan	Taiwan Island	introduced invasive	Ching-I Peng, ed. (2011) "In disturbed uplands".

Indian Ocean
Country/Terr./St. & Island group	Location	Cited status & Cited as invasive & Cited as cultivated & Cited as aboriginal introduction?	Reference & Comments
La Réunion (France) La Réunion Island	La Réunion Island	introduced invasive	MacDonald, I. A. W./Thebaud, C./Strahm, W. A./Strasberg, D. (1991) (pp. 51-61)
Seychelles Seychelles Islands	Frégate Island		Robertson, S. A./Todd, D. M. (1983) (p. 61)

Control: 

Physical:  Hand pull or dig out seedlings and young plants.

Chemical:  Cut large plants or treat with herbicide.  "Probably susceptible to hormone-type herbicides, especially when young, and probably to tebuthiuron"  (Motooka et al., 2003).

Additional information:
Excerpt from the book "Weeds of Hawaii‘s Pastures and Natural Areas; An Identification and Management Guide" (Motooka et al., 2003). (PDF format).
Fact sheet from Environmental Bay of Plenty Regional Council, New Zealand.
Information from the Global Invasive Species Database.

Additional online information about Physalis peruviana is available from the Hawaiian Ecosystems at Risk project (HEAR).

Information about Physalis peruviana as a weed (worldwide references) may be available from the Global Compendium of Weeds (GCW).

Taxonomic information about Physalis peruviana may be available from the Germplasm Resources Information Network (GRIN).

References:

Atkinson, Rachel/Sawyer, John. 2011. Naturalized species in the Juan Fernández Archipelago, Chile. Unpublished spreadsheet.

Beleu National Museum (Republic of Palau. Koror.). 1966. Voucher specimen #BNM753(Blackburn E-108).

Burkill, I. H. 1900. The flora of Vavau, one of the Tonga Islands, with a short account of its vegetation by Charles Steele Crosby. Lin. Jour. Bot. 35:20-65.

Charles Darwin Foundation. 2008. Database inventory of introduced plant species in the rural and urban zones of Galapagos. Charles Darwin Foundation, Galapagos, Ecuador.

Charles Darwin Research Station. 2005. CDRS Herbarium records.

Ching-I Peng, ed. 2011. Digital flora of Taiwan (online resource).

Danton, Philippe/Perrier, Christophe/Martinez Reyes, Guido. 2006. Nouveau catalogue de la flore vaculaire de l'archipel Juan Fernández (Chile) [Nuevo catálogo de la flora vacular del Archipélago Juan Fernández (Chile)]. Acta Bot. Gallica 153(4):399-587.

Florence, J./Chevillotte, H./Ollier, C./Meyer, J.-Y. 2011. Base de données botaniques Nadeaud de l'Herbier de la Polynésie Française (PAP). (online resource).

Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce. 1979. A geographical checklist of the Micronesian dicotyledonae. Micronesica 15:1-295.

Heine, Heino. 1976. Acanthacées, Bignoniacées, Boraginacées, Solanacées. Flore de la Nouvelle-Calédonie et Dépendances. Muséum National d'Histoire Naturelle, Paris. Vol. 7, 212 pp.

Holm, Leroy/Pancho, Juan V./Herberger, James P./Plucknett, Donald L. 1979. A geographical atlas of world weeds. John Wiley & Sons, New York. 391 pp.

Kato, Hidetoshi. 2007. Herbarium records of Makino Herbarium, Tokyo Metropolitan University. Personal communication.

Lorence, David H./Wagner, Warren L. 2008. Flora of the Marquesas Islands. National Tropical Botanical Garden and the Smithsonian Institution. Online database.

MacDonald, I. A. W./Thebaud, C./Strahm, W. A./Strasberg, D. 1991. Effects of alien plant invasions on native vegetation remnants on La Reunion (Mascarenes Islands, Indian Ocean). Environmental Conservation 18 (1):51-61.

MacKee, H. S. 1994. Catalogue des plantes introduites et cultivées en Nouvelle-Calédonie. Muséum National d'Histoire Naturelle, Paris, 164 p.

McCormack, Gerald. 2011. Cook Islands Biodiversity Database, Version 2007.2. Cook Islands Natural Heritage Trust, Rarotonga.

Merrill, Elmer D. 1923. An enumeration of Philippine flowering plants, vol. 3 [reprint]. Bureau of Printing, Manila. 628 pp.

Mito, Toshikazu/Uesugi, Tetsuro. 2004. Invasive alien species in Japan: the status quo and the new regulation for prevention of their adverse effects. Global Environmental Research 8(2)/2004: 171-191.

Motooka, Philip/Castro, Luisa/Nelson, Duane/Nagai, Guy/Ching, Lincoln. 2003. Weeds of Hawaii‘s Pastures and Natural Areas; An Identification and Management Guide. College of Tropical Agriculture and Human Resources, University of Hawaii at Manoa. 184 pp.

Orchard, Anthony E., ed. 1994. Flora of Australia. Vol. 49, Oceanic islands 1. Australian Government Publishing Service, Canberra.

Parham, J. W. 1972. Plants of the Fiji Islands. Revised edition. Government Printer, Suva. 462 pp.

Pickard, J. 1984. Exotic plants on Lord Howe Island: Distribution in space and time, 1853-1981. J. of Biogeography 11:181-208.

Robertson, S. A./Todd, D. M. 1983. Vegetation of Frégate Island, Seychelles. In: Sachet, M. H., D. R. Stoddart, and F. R. Fosberg. Floristics and ecology of Western Indian Ocean islands. Atoll Research Bulletin No. 273. Smithsonian Institution, Washington. 253 pp.

Skottsberg, Carl. 1953. The natural history of Juan Fernandez and Easter Island. Vol. II, Botany. Almqvist &l; Wiksells, Uppsala. 960 pp.

Smith, Albert C. 1991. Flora Vitiensis nova: a new flora of Fiji. National Tropical Botanical Garden, Lawai, Kauai, Hawaii. Volume 5. 626 pp.

Space, James C./Waterhouse, Barbara M./Newfield, Melanie/Bull, Cate. 2004. Report to the Government of Niue and the United Nations Development Programme: Invasive plant species on Niue following Cyclone Heta. UNDP NIU/98/G31 - Niue Enabling Activity. 80 pp.

St. John, H. 1987. An account of the flora of Pitcairn Island with new Pandanus species. Pacific Plant Studies 46. Honolulu, 65 pp. [Privately published].

St. John, Harold. 1988. Census of the Flora of the Gambier Islands, Polynesia. Pacific Plant Studies 43. Honolulu, Hawaii. 34 pp. [Privately published].

Swarbrick, John T. 1997. Weeds of the Pacific Islands. Technical paper no. 209. South Pacific Commission, Noumea, New Caledonia. 124 pp.

Sykes, W. R. 1970. Contributions to the flora of Niue. New Zealand Department of Scientific and Industrial Research Bulletin 200, Wellington. 321 pp.

Sykes, W. R. 1977. Kermadec Islands flora: an annotated checklist. New Zealand Department of Scientific and Industrial Research Bulletin 219, Wellington. 216 pp.

Tassin, Jacques. 2005. Jacques Tassin (IAC-CIRAD), personal communication.

Thaman, R. R. 1987. Plants of Kiribati: a listing and analysis of vernacular names. Atoll Res. Bull. 296:1-42.

Toyoda, Takeshi. 2003. Flora of the Bonin Islands. Aboc & Co. 522 pp.

U.S. Dept. Agr., Agr. Res. Serv. 2011. National Genetic Resources Program. Germplasm Resources Information Network (GRIN). Online searchable database.

Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. 1999. Manual of the flowering plants of Hawaii. Revised edition. Bernice P. Bishop Museum special publication. University of Hawai‘i Press/Bishop Museum Press, Honolulu. 1919 pp. (two volumes).

Webb, C. J./Sykes, W. R./Garnock-Jones, P. J. 1988. Flora of New Zealand, Volume IV: Naturalised pteridophytes, gymnosperms, dicotyledons. Botany Division, DSIR, Christchurch. 1365 pp.

Weber, Ewald. 2003. Invasive plants of the World. CABI Publishing, CAB International, Wallingford, UK. 548 pp.

Welsh, S. L. 1998. Flora Societensis: A summary revision of the flowering plants of the Society Islands. E.P.S. Inc., Orem, Utah. 420 pp.

Whistler, W. A. 1988. Checklist of the weed flora of western Polynesia. Technical Paper No. 194, South Pacific Commission, Noumea, New Caledonia. 69 pp.

Whistler, W. A. 1992. Flora of Tonga [unpublished checklist]. 107 pp.

Yuncker, T. G. 1943. The flora of Niue Island. Bishop Mus. Bull. 178. Bishop Museum Press, Honolulu. 126 pp.

Yuncker, T. G. 1959. Plants of Tonga. Bishop Museum Bull. 220. Bishop Museum Press, Honolulu. 343 pp.

Zhengyi, Wu/Raven, Peter H./Deyuan, Hong. 2011. Flora of China (online resource).