Desmanthus pernambucanus (L.) Thell., Fabaceae

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Present on Pacific Islands?  yes

Primarily a threat at high elevations?  no

Risk assessment results:  See Desmanthus virgatus for risk assessments for a similar species.

Common name(s): [more details]

English: pigeon bundleflower

Other: anil, jureminha

Habit:  herb

Description:  "Erect or decumbent and scrambling perennial herbs or subshrubs, 0.5 to 2 m tall, little branched from the base.  Taproot cylindrical, woody, gnarled, bark brown, corky.  Young stems angled, red or green with golden corky ridges, sparsely pubescent with tangled white hairs; older stems terete, red or brown, shiny, glabrous.  Stipules persistent, 3.0-6.1 mm long, setiform with broad, nerved auricles at the base, one of which curves abaxially under the petiole, usually puberulent with white hairs.  Leaves 4.4-11.1 cm long, petiole 6-16 mm long, rachis 10-49 mm long, sparsely pubescent; pinnae 2-4 pairs, 15-54 mm long; nectary borne between the lowest pair of pinnae elliptical, sessile, crateriform or flattened, 1.0-3.3 mm wide, oblong to linear, rounded-oblique basally, mucronulate to rounded apically, glabrous, finely ciliate along the margin with white hairs, sometimes minutely denticulate where the hairs arise, venation obscure except excentric midvein.  Heads 1 per leaf axil, borne on peduncles 1.0-4.5 cm long.  Bract subtending each flower 1.0-2.1 mm long, 0.4-0.7 mm wide, deltate, pale green with red tips, membranous with an opaque midvein, peltate and pedicellate at the center of the head, sessile and often fused into a whorl at the base of the head, strongly 1-nerved, abaxially pubescent.  Floral buds obovate, apically rounded.  Heads 0.7-1.0 cm long containing 8-13 sterile, functionally male, and perfect flowers, rarely the male flowers absent.  Sterile flowers 2-4, calyx 0.9-1.7 mm long, 0.45-1.10 mm wide, 5-lobed, widely obconic, pale green with white margins; petals 1.6-2.7 mm long, 0.40-0.65 mm wide, obovate to oblanceolate, membranous, faintly 1-nerved, white or pale green with white margins; staminodia 10, 4.7-10.0 mm long, the same width as the filaments of the functional stamens to 3 times as wide, white.  Male flowers 0-4, lacking an ovary, otherwise with a perianth and androecium like that of the perfect flowers.  Perfect flowers 4-8, calyx 2.0-3.0 mm long, obconic, the tube 1.7-2.8 mm long, 0.8-1.5 mm in diameter, rimmed with free acute lobes 0.3-0.5 mm long, pale green with white margins, glabrous; petals 3.0-4.0 mm long, 0.5-0.89 mm wide, oblanceolate, green with red or purple tips, glabrous; stamens 10, rarely 5, 5.0-7.4 mm long, anthers eglandular; ovary 1.5-2.5 mm long, linear, glabrous, style 3.0-5.1 mm long, not exserted beyond the stamens.  Fruiting peduncles 2.4-5.6 cm long bearing 1-7 pods; legumes dehiscent along both sutures, the sutural ridges curling back over the valves causing the inner surface of the pods to be slightly convex; pod 4.5-8.5 cm long, 3.2-3.9 mm wide, narrow, linear, straight, tip mostly acute, rarely attenuate with a beak to 2 mm long, valves chartaceous, glabrous or rarely pubescent, nearly black at maturity, somewhat wrinkled with raised reticulate veins, the sutural ridges lighter in color than the valves.  Seeds 13-22 per pod, 2.4-3.2 mm long, 1.9-2.4 mm wide, obliquely inserted, ovate and flattened or rhomboidal and 4-angled, black or red-brown; pleurogram 0.7-1.0 mm wide, 0.7-1.5 mm deep, deeply U-shaped, symmetrical." (Luckow, 1993).

"Desmanthus pernambucanus differs from the other species of the virgatus complex in having fewer pinnae, and fewer, larger leaflets than either D. leptophyllus or D. virgatus, and in the nyctinastic leaf movements.  Desmanthus virgatus has shorter petioles and usually more pinnae, fruit dehiscence in which the valves become convex rather than concave, and generally a smaller stature"  (Luckow, 1993; pp. 113-117).

Habitat/ecology:  In its native range, "moist roadsides, ditches, and abandoned pastures, also common in coastal thickets and along the edges of marshes; on sandy, rocky, limestone and saline soils; 0-1500 m"  (Luckow, 1993; pp. 113-117). In Hawai‘i, "naturalized in dry, open, disturbed areas, 0-215 m" (Wagner et al., 1999; pp. 665; 1876). In Fiji (as Desmanthus virgatus), "perhaps originally introduced for cultivation but now sparingly naturalized along shores and in waste places near sea level" (Smith, 1985; p. 68).  In American Samoa (as Desmanthus virgatus), "uncommon near the airport on Tutuila" (Whistler, 1988; p. 23). In New Caledonia (as Desmanthus virgatus), "a peut-être quelque valeur fourragére mais se rencontre surtout en dehors des pâturages" (MacKee, 1994; p. 76).

Propagation:  Seed

Native range:  "Native to most islands in the Caribbean..., also in South America in Guyana, Surinam, Tobago, Trinidad, and along the coasts of northeastern Brazil; one collection from Uruguay; probably introduced and widely naturalized on islands in the Pacific and Indian Oceans, in southern Africa, southeast Asia, and Florida, U.S.A."  (Luckow, 1993; pp. 113-117).

Presence:

Pacific
Country/Terr./St. & Island group	Location	Cited status & Cited as invasive & Cited as cultivated & Cited as aboriginal introduction?	Reference & Comments
American Samoa Tutuila Islands	Tutuila Island		Whistler, W. A. (1988) (p. 23) As Desmanthus virgatus but likely this species per Luckow (1993).
Commonwealth of the Northern Mariana Islands (US) Northern Mariana Islands	Agrigan (Agrihan) Island	introduced	Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce (1979) (p. 92) As Desmanthus virgatus but likely this species per Luckow (1993).
Commonwealth of the Northern Mariana Islands (US) Northern Mariana Islands	Agrigan (Agrihan) Island	introduced invasive	Fosberg, F. R./Falanruw, M. V. C./Sachet, M.-H. (1975) (p. 27) As Desmanthus virgatus but likely this species per Luckow (1993). Voucher cited: Falanruw 2196 (US)
Commonwealth of the Northern Mariana Islands (US) Northern Mariana Islands	Rota Island	introduced	Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce (1979) (p. 92) As Desmanthus virgatus but likely this species per Luckow (1993).
Commonwealth of the Northern Mariana Islands (US) Northern Mariana Islands	Saipan Island	introduced	Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce (1979) (p. 92) As Desmanthus virgatus but likely this species per Luckow (1993).
Federated States of Micronesia Chuuk Islands	Fefan Island	introduced	Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce (1979) (p. 92) As Desmanthus virgatus but likely this species per Luckow (1993).
Federated States of Micronesia Chuuk Islands	Fefan Island	introduced invasive	Space, James C./Waterhouse, Barbara/Denslow, Julie S./Nelson, Duane/Mazawa, Thomas R. (2000) (pp. 6, 27) As Desmanthus virgatus but likely this species per Luckow (1993).
Federated States of Micronesia Chuuk Islands	Tol Island	introduced	Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce (1979) (p. 92) As Desmanthus virgatus but likely this species per Luckow (1993).
Federated States of Micronesia Chuuk Islands	Tol Island	introduced invasive	Space, James C./Waterhouse, Barbara/Denslow, Julie S./Nelson, Duane/Mazawa, Thomas R. (2000) (pp. 6, 27) As Desmanthus virgatus but likely this species per Luckow (1993).
Federated States of Micronesia Chuuk Islands	Weno (Moen) Island	introduced	Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce (1979) (p. 92) As Desmanthus virgatus but likely this species per Luckow (1993). Voucher cited: Cherrier in MacKee 44080
Federated States of Micronesia Chuuk Islands	Weno (Moen) Island	introduced invasive	Space, James C./Waterhouse, Barbara/Denslow, Julie S./Nelson, Duane/Mazawa, Thomas R. (2000) (pp. 6, 27) As Desmanthus virgatus but likely this species per Luckow (1993). Voucher cited: Cherrier in MacKee 44080
Federated States of Micronesia Kosrae Island	Kosrae Island	introduced invasive	Space, James C./Waterhouse, Barbara/Denslow, Julie S./Nelson, Duane/Waguk, Erick E. (2000) (p. 6) As Desmanthus virgatus but likely this species per Luckow (1993).
Federated States of Micronesia Kosrae Island	Kosrae Island		Lorence, David H./Flynn, Tim (2005) (p. 15) As Desmanthus virgatus but likely this species per Luckow (1993). Voucher cited: Lorence 8135A (PTBG)
Federated States of Micronesia Kosrae Island	Kosrae Island	introduced invasive	Josekutty, P. C./Wakuk, E. E./Joseph, M. J. (2002) (p. 62) As Desmanthus virgatus but likely this species per Luckow (1993).
Federated States of Micronesia Pohnpei Islands	Pohnpei Island		Swarbrick, John T. (1997) (p. 91) As Desmanthus virgatus but likely this species per Luckow (1993).
Federated States of Micronesia Pohnpei Islands	Pohnpei Island		Lorence, David H./Flynn, Tim (1998) (p. 21) As Desmanthus virgatus but likely this species per Luckow (1993).
Federated States of Micronesia Yap Islands	Yap (Waqab) Island	introduced invasive	Fosberg, F. R./Falanruw, M. V. C. (1980) (p. 205) As Desmanthus virgatus but likely this species per Luckow (1993). Voucher cited: Falanruw 3212 (US)
Fiji Fiji Islands	Ovalau Island	introduced invasive	Smith, Albert C. (1985) (p. 68) As Desmanthus virgatus but likely this species per Luckow (1993). Voucher cited: DA 17038 Sparingly naturalized.
Fiji Fiji Islands	Vanua Mbalavu Island	introduced invasive	Smith, Albert C. (1985) (p. 68) As Desmanthus virgatus but likely this species per Luckow (1993). Vouchers cited: Tothill 135, DA 10221, DA L.15791 Sparingly naturalized.
Fiji Fiji Islands	Viti Levu Island	introduced invasive	Smith, Albert C. (1985) (p. 68) As Desmanthus virgatus but likely this species per Luckow (1993). Vouchers cited: Greenwood 1184, DA 4043, DA 4042 Sparingly naturalized.
French Polynesia Marquesas Islands	Hiva Oa Island	introduced invasive	Lorence, David H./Wagner, Warren L. (2008)
French Polynesia Marquesas Islands	Nuku Hiva (Nukahiva) Island	introduced invasive	Lorence, David H./Wagner, Warren L. (2008)
French Polynesia Marquesas Islands	Tahuata (Tuhuata) Island	introduced invasive	Lorence, David H./Wagner, Warren L. (2008)
French Polynesia Marquesas Islands	Ua Huka (Huahuna, Uahuka) Island	introduced invasive	Lorence, David H./Wagner, Warren L. (2008)
French Polynesia Society Islands	Huahine Island		Fosberg, F. R. (1997) (p. 50) As Desmanthus virgatus but likely this species per Luckow (1993).
French Polynesia Society Islands	Tahiti Island		Fosberg, F. R. (1997) (p. 50) As Desmanthus virgatus but likely this species per Luckow (1993).
Guam Guam Island	Guam Island	introduced invasive	Stone, Benjamin C. (1970) (p. 297) As Desmanthus virgatus but likely this species per Luckow (1993). Voucher cited: Stone 4182 (GUAM)
Guam Guam Island	Guam Island	introduced	Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce (1979) (p. 92) As Desmanthus virgatus but likely this species per Luckow (1993).
State of Hawaii Hawaiian Islands	Hawai‘i (Big) Island	introduced invasive	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (p. 665)
State of Hawaii Hawaiian Islands	Kaho‘olawe Island	introduced invasive	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (p. 665)
State of Hawaii Hawaiian Islands	Kaua‘i Island	introduced invasive	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (p. 665)
State of Hawaii Hawaiian Islands	Lāna‘i Island	introduced invasive	Oppenheimer, Hank L. (2003) (p. 12) Voucher cited: Oppenheimer H40128 (BISH, PTBG)
State of Hawaii Hawaiian Islands	Maui Island	introduced invasive	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (p. 665)
State of Hawaii Hawaiian Islands	Moloka‘i Island	introduced invasive	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (p. 665)
State of Hawaii Hawaiian Islands	Ni‘ihau Island	introduced invasive	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (p. 665)
State of Hawaii Hawaiian Islands	O‘ahu Island	introduced invasive	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (p. 665)
New Caledonia Îles Loyauté (Loyalte Islands)	Île Lifou	introduced	MacKee, H. S. (1994) (p. 78) As Desmanthus virgatus but likely this species per Luckow (1993). Voucher cited: MacKee 5238
New Caledonia Îles Loyauté (Loyalte Islands)	Île Walpole	introduced	MacKee, H. S. (1994) (p. 78) As Desmanthus virgatus but likely this species per Luckow (1993). Voucher cited: Cherrier in MacKee 44080
New Caledonia New Caledonia	New Caledonia Islands		Swarbrick, John T. (1997) (p. 91) As Desmanthus virgatus but likely this species per Luckow (1993).
New Caledonia New Caledonia	New Caledonia Islands	introduced invasive	Gargominy, Oliver/Bouchet, Philipe/Pascal, Michel/Jaffre, Tanguy/Tourneu, Jean-Christophe (1996) (p. 381) As Desmanthus virgatus but likely this species per Luckow (1993). Se rencontre surtout en dehors des pâturages.
New Caledonia New Caledonia Archipelago	Île Grande Terre	introduced	MacKee, H. S. (1994) (p. 78) As Desmanthus virgatus but likely this species per Luckow (1993). Vouchers cited: Le Rat 1993, Franc 1313, Däniker 409a, Catala 66, MacDaniels 2043, Sarlin 77 s.loc., MacKee 3562, MacKee 8047, MacKee 30103, MacKee 37327
Palau Palau (main island group)	Babeldaob Island	introduced	Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce (1979) (p. 92) As Desmanthus virgatus but likely this species per Luckow (1993).
Palau Palau (main island group)	Babeldaob Island	introduced invasive	Space, James C./Waterhouse, Barbara/Miles, Joel E./Tiobech, Joseph/Rengulbai, Kashgar (2003) (p. 85)
Palau Palau (main island group)	Babeldaob Island	introduced invasive	Space, James C./Lorence, David H./LaRosa, Anne Marie (2009) (p. 59)
Palau Palau (main island group)	Koror Island	introduced	Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce (1979) (p. 92) As Desmanthus virgatus but likely this species per Luckow (1993).
Palau Palau (main island group)	Koror Island	introduced invasive	Space, James C./Waterhouse, Barbara/Miles, Joel E./Tiobech, Joseph/Rengulbai, Kashgar (2003) (p. 80) As Desmanthus virgatus but likely this species per Luckow (1993).
Palau Palau (main island group)	Malakal Island	introduced invasive	Space, James C./Waterhouse, Barbara/Miles, Joel E./Tiobech, Joseph/Rengulbai, Kashgar (2003) (p. 80) As Desmanthus virgatus but likely this species per Luckow (1993).
Palau Palau (main island group)	Ngerkebesang Island	introduced invasive	Space, James C./Waterhouse, Barbara/Miles, Joel E./Tiobech, Joseph/Rengulbai, Kashgar (2003) (p. 80) As Desmanthus virgatus but likely this species per Luckow (1993).
Solomon Islands Solomon Islands	Solomon Islands		Swarbrick, John T. (1997) (p. 91) As Desmanthus virgatus but likely this species per Luckow (1993).
Solomon Islands Solomon Islands	Solomon Islands	introduced	Hancock, I. R./Henderson, C. P. (1988) (p. 65) As Desmanthus virgatus but likely this species per Luckow (1993).
United States (other Pacific offshore islands) Midway Atoll	Midway Atoll	introduced invasive	Wagner, Warren L./Herbst, Derral R./Sohmer, S. H. (1999) (p. 665)
United States (other Pacific offshore islands) Wake Islands	Wake Island	introduced	Fosberg, F. R./Sachet, Marie-Hélène/Oliver, Royce (1979) (p. 92) As Desmanthus virgatus but likely this species per Luckow (1993).
United States (other Pacific offshore islands) Wake Islands	Wake Island	introduced invasive	Fosberg, F. R./Sachet, M.-H. (1969) (p. 10) As Desmanthus virgatus but likely this species per Luckow (1993). Voucher cited: McFarlane 74 (US)
Vanuatu New Hebrides Islands	Vanuatu (Republic of)		Swarbrick, John T. (1997) (p. 91) As Desmanthus virgatus but likely this species per Luckow (1993).

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	ILDIS Co-ordinating Centre (2009) As Desmanthus virgatus but likely this species per Luckow (1993).
Mauritius Mautitius Islands (Mauritius and Rodrigues)	Mauritius Island	introduced	ILDIS Co-ordinating Centre (2009) As Desmanthus virgatus but likely this species per Luckow (1993).
Mauritius Mautitius Islands (Mauritius and Rodrigues)	Rodrigues Island	introduced	ILDIS Co-ordinating Centre (2009) As Desmanthus virgatus but likely this species per Luckow (1993).
Seychelles Seychelles Islands	Groupe d'Aldabra	introduced	ILDIS Co-ordinating Centre (2009) As Desmanthus virgatus but likely this species per Luckow (1993). Voucher cited: Falanruw 3212 (US)
Seychelles Seychelles Islands	Seychelles Islands	introduced	ILDIS Co-ordinating Centre (2009) As Desmanthus virgatus but likely this species per Luckow (1993).

Also reported from
Country/Terr./St. & Island group	Location	Cited status & Cited as invasive & Cited as cultivated & Cited as aboriginal introduction?	Reference & Comments
United States (continental except west coast and Florida) United States (other states)	USA (Florida)	introduced invasive	Luckow, Melissa (1993) (p. 114) Naturalized.

Comments:  "In a recent monograph of Desmanthus (Luckow, 1993; pp. 113-117), species of the D. virgatus complex were reinterpreted.  The result is that the species naturalized on many Pacific islands, including the Hawaiian Islands, is now referred to D. pernambucanus" (Wagner & Herbst, 1995).

Quite weedy on Chuuk. Common along roadsides, extending into the forest.

First recorded in Guam (as Desmanthus virgatus) in 1964 per Stone (1970; p. 297).

Additional information:
Additional online information about Desmanthus pernambucanus is available from the Hawaiian Ecosystems at Risk project (HEAR).

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

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

References:

Fosberg, F. R. 1997. Preliminary checklist of the flowering plants and ferns of the Society Islands. Ed. by David R. Stoddart. U. Cal. Berkeley.

Fosberg, F. R./Falanruw, M. V. C. 1980. Noteworthy Micronesian plants 4. Micronesica 16(2):201-210.

Fosberg, F. R./Falanruw, M. V. C./Sachet, M.-H. 1975. Vascular flora of the northern Marianas Islands. Smithsonian Contrib. Bot. 22:1-45.

Fosberg, F. R./Sachet, M.-H. 1969. Wake Island vegetation and flora, 1961-1963. Atoll Research Bulletin No. 123. Smithsonian Institution, Washington. 15 pp.

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

Gargominy, Oliver/Bouchet, Philipe/Pascal, Michel/Jaffre, Tanguy/Tourneu, Jean-Christophe. 1996. Conséquences des introductions d'espèces animales et végétales sur la biodiversité en Nouvelle-Calédonie. Rev. Ecol. (Terre Vie) 51:375-401.

Hancock, I. R./Henderson, C. P. 1988. Flora of the Solomon Islands. Research Bulletin No. 7. Ministry of Agriculture and Lands, Honiara. 203 pp.

ILDIS Co-ordinating Centre. 2009. International Legume Database & Information Service. Online searchable database.

Josekutty, P. C./Wakuk, E. E./Joseph, M. J. 2002. Invasive/weedy angiosperms in Kosrae, Federated States of Micronesia. In: Invasive species and their management. Micronesica Suppl. 6:61-65.

Lorence, David H./Flynn, Tim. 1998. Checklist of the plants of Pohnpei. Unpublished checklist. National Tropical Botanical Garden, Lawai, Hawai‘i. 21 pp.

Lorence, David H./Flynn, Tim. 2005. Checklist of the plants of Kosrae. Unpublished checklist. National Tropical Botanical Garden, Lawai, Hawai‘i. 23 pp.

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

Luckow, Melissa. 1993. Monograph of Desmanthus (Leguminosae-Mimosoideae). Syst. Bot. Monogr. 38.

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

Oppenheimer, Hank L. 2003. New plant records from Maui and Hawai‘i Counties. In: Evenhuis, Neal L. and Eldredge, Lucius G., eds. Records of the Hawaii Biological Survey for 2001-2002. Part 1: Articles. Bishop Museum Occasional Papers. 73:3-30.

Smith, Albert C. 1985. Flora Vitiensis nova: a new flora of Fiji. National Tropical Botanical Garden, Lawai, Kauai, Hawaii. Volume 3. 758 pp.

Space, James C./Lorence, David H./LaRosa, Anne Marie. 2009. Report to the Republic of Palau: 2008 update on Invasive Plant Species. USDA Forest Service, Hilo, Hawai‘i. (In press).

Space, James C./Waterhouse, Barbara/Denslow, Julie S./Nelson, Duane/Mazawa, Thomas R. 2000. Invasive plant species in Chuuk, Federated States of Micronesia.  USDA Forest Service, Honolulu. 41 pp.

Space, James C./Waterhouse, Barbara/Denslow, Julie S./Nelson, Duane/Waguk, Erick E. 2000. Invasive plant species on Kosrae, Federated States of Micronesia.  USDA Forest Service, Honolulu. 43 pp.

Space, James C./Waterhouse, Barbara/Miles, Joel E./Tiobech, Joseph/Rengulbai, Kashgar. 2003. Report to the Republic of Palau on invasive plant species of environmental concern. USDA Forest Service, Honolulu. 174 pp.

Stone, Benjamin C. 1970. The flora of Guam. Micronesica 6:1-659.

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

U.S. Dept. Agr., Nat. Res. Cons. Serv. 2009. The PLANTS Database. National Plant Data Center, Baton Rouge, LA 70874-4490 USA.

Wagner, Warren L./Herbst, Derral R. 1995. Contributions to the Flora of Hawai‘i. IV. New Records and name changes. In: Evenhuis, Neal L. and Miller, Scott, E., eds. Records of the Hawaii Biological Survey for 1994. Part 2: Notes. Bishop Museum Occasional Papers. 42:13-27.

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).

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