Dichanthium annulatum
(Kleberg's bluestem)

Pictures

Picture	Title	Caption	Copyright
Title Flower spike Caption Dichanthium annulatum; flower spike. Summer Island, New South Wales, Australia. May 2010. Copyright ©Harry Rose (Macleay Grass Man)/via flickr - CC BY 2.0	Flower spike	Dichanthium annulatum; flower spike. Summer Island, New South Wales, Australia. May 2010.	©Harry Rose (Macleay Grass Man)/via flickr - CC BY 2.0
Title Flower spike Caption Dichanthium annulatum; close-up of flower spike. Summer Island, New South Wales, Australia. May 2010. Copyright ©Harry Rose (Macleay Grass Man)/via flickr - CC BY 2.0	Flower spike	Dichanthium annulatum; close-up of flower spike. Summer Island, New South Wales, Australia. May 2010.	©Harry Rose (Macleay Grass Man)/via flickr - CC BY 2.0
Title Spikelet with long awn Caption Dichanthium annulatum; spikelet with long awn. Summer Island, New South Wales, Astralia. June 2013. Copyright ©Harry Rose (Macleay Grass Man)/via flickr - CC BY 2.0	Spikelet with long awn	Dichanthium annulatum; spikelet with long awn. Summer Island, New South Wales, Astralia. June 2013.	©Harry Rose (Macleay Grass Man)/via flickr - CC BY 2.0
Title Fruits Caption Dichanthium annulatum (Kleberg's bluestem); fruits. Copyright This image is not copyrighted and may be freely used for any purpose (original by Steve Hurst/USDA-NRCS PLANTS Database).	Fruits	Dichanthium annulatum (Kleberg's bluestem); fruits.	This image is not copyrighted and may be freely used for any purpose (original by Steve Hurst/USDA-NRCS PLANTS Database).

Identity

Preferred Scientific Name

• Dichanthium annulatum (Forssk.) Stapf

Preferred Common Name

• Kleberg's bluestem

Other Scientific Names

• Andropogon annulatus Forssk.
• Andropogon garipensis Steud.
• Andropogon ischaemum Roxb. ex Wight
• Andropogon obtusus Nees ex Hook. & Arn.
• Andropogon scandens Roxb.
• Bothriochloa tuberculata W.Z. Fang
• Gymnandropogon annulatum (Forssk.) Duthie
• Lepeocercis annulata (Forssk.) Nees
• Sorghum annulatum (Forssk.) Kuntze

International Common Names

• English: angleton grass; bluestem; diaz bluestem; Hindi grass; Kleberg bluestem; marvel grass; ringed dichanthium; sheda grass; two-flowered goldenbeard; vunda blue grass
• Spanish: climacuna; pajón; pitilla; yerba de las travesías; yerba de vía
• Chinese: shuang hua cao

Local Common Names

• Cuba: pitilla americana
• India: delhi grass; jargu grass; karad
• Philippines: lindi
• Thailand: ya hindi; ya waen
• Vietnam: song tha'o to

Summary of Invasiveness

D. annulatum is a perennial grass widely naturalized in tropical and subtropical regions of the world where it has been intentionally introduced. This grass species is used for grazing and for hay and silage (Barkworth et al., 2003; Cook et al., 2005; FAO, 2014). It has escaped from cultivation and has become a weed in the United States, Australia, Mexico, Central America and the West Indies (Villaseñor and Espinosa-García, 2004; Chacon and Saborío, 2012; Gonzalez-Torres et al., 2012; Randall, 2012; USDA-NRCS, 2014). D. annulatum has been intentionally introduced as a pasture grass because of its capability to establish on a wide variety of soils (including poorly drained soils), and its salinity and drought tolerance (i.e., 6- 8 months dry seasons; Cook et al., 2005). It competes aggressively with other plants and grows forming dense stands (Vibrans, 2011). D. annulatum also tolerates seasonal burning (Cook et al., 2005).

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Monocotyledonae
•                     Order: Cyperales
•                         Family: Poaceae
•                             Genus: Dichanthium
•                                 Species: Dichanthium annulatum

Notes on Taxonomy and Nomenclature

The genus Dichanthium of the family Poaceae comprises 20 species native to the Old World, but hybrids between different species have been reported (Saxena and Chandra, 2006). The name Dichanthium comes from the Greek “dicha” = in two, as in two separate things, and “anthos” = flower, as a reference to the presence of homogamous and heterogamous spikelets (Barkworth et al., 2003). The species D. annulatum is highly apomictic and it can form hybrids with other species such as D. fecundum (Saxena and Chandra, 2006). Thus, a considerable morphological variation has been reported for this species (Agarwal et al., 1999).

Description

D. annulatum is a perennial grass with culms 25-100 cm tall, ascending, nodes pubescent. Sheaths lax, terete, 3-4 cm long, glabrous except finely pubescent on margins near throat; ligule a shallow membrane, approximately 1.5 mm long; blades linear-acuminate, 3-30 cm long, 2-7 mm wide, margins sparsely pubescent, apex attenuate, base subcordate. Inflorescences composed of (1) 2-15 subdigitate, short-pedunculate racemes, each raceme 3-7 cm long, peduncles glabrous; spikelet pairs subimbricate along a slender filiform axis, with 0-6 smaller homogamous spikelet pairs at base, the joints and pedicels flattened, margins setose; sessile spikelet narrowly oblong, 2-6 mm long, first glume cartilaginous, slightly concave, villous below the middle, pubescent with long bulbous-based hairs above the middle, obscurely nerved, margins inflexed, apex obtuse to subacute, second glume narrowly boat-shaped, 2-6 mm long, 3-nerved, scabrid on back of the midnerve toward apex, obscurely scaberulous over the back toward apex, hyaline, first lemma delicately hyaline, oblong-rounded, setose near tip, approximately 2 mm long, palea absent, second lemma linear, approximately 1.6 mm long, hyaline, with the central nerve excurrent as a geniculate, twisted awn 8-25 mm long; pedicellate spikelet with first glume narrowly ovate-truncate, approximately 3 mm long, 11-nerved, pectinate-setose along the back of the inflexed margins, midnerve setose on the back toward apex. Caryopsis oblong to obovate, dorsally compressed, approximately 2 mm long (Wagner et al., 1999).

Plant Type

Distribution

D. annulatum is native to Africa, tropical and temperate Asia, the Middle East, and Papua New Guinea (Clayton et al., 2014; USDA-ARS, 2014). Currently this species is widely naturalized in Australia, America, and islands in the Pacific Ocean (see distribution table for details; Randall, 2012; Clayton et al., 2014; PIER, 2014; USDA-ARS, 2014).

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

The date of the initial introduction of D. annulatum into new habitats is uncertain, but most likely it occurred during the late nineteenth or early twentieth century. For the West Indies, this species was first reported in 1915 in Cuba and in 1922 in Puerto Rico (US National Herbarium Collection). Between 1944 and 1954, the species D. annulatum and several cultivars were intentionally released in Texas, USA (Cook et al., 2005).

Risk of Introduction

The risk of introduction of D. annulatum is very high. This grass has been repeatedly intentionally introduced in tropical and subtropical regions to be used as a fodder, forage, hay, and silage crop (Cook et al., 2005; FAO, 2014). It has escaped from cultivation and rapidly naturalized into natural areas where it colonizes new areas forming dense stands and displacing native vegetation. It is apomictic and produces large numbers of wind-dispersed seeds (Vibrans, 2011). In addition, D. annulatum is fire and drought tolerant, which are features helping this species to outcompete native species (Cook et al., 2005; Queensland Department of Primary Industries and Fisheries, 2011).

Habitat

D. annulatum can be found growing in waste and disturbed sites, urban areas, dry coastal sites, arid wetlands, sand dune grasslands, and along roadsides near pastures and grazing areas where this species often escapes and become weedy (Wagner et al., 1999; Cook et al., 2005; Queensland Department of Primary Industries and Fisheries, 2011). It is adapted to grow in conditions ranging from moist tropical and subtropical regions to dry and arid habitats (Barkworth et al., 2003; Cook et al., 2005).

Habitat List

Hosts/Species Affected

In Australia, D. annulatum is competing aggressively with important native grasses such as the species Eriachne benthamii. In Texas (USA), this species has escaped from pastures and is outcompeting native grasses in bluestem coastal grassland communities (Queensland Department of Primary Industries and Fisheries, 2011).

Biology and Ecology

Genetics

The basic chromosome number in Dichanthium is 10, however the species D. annulatum shows different ploidy levels with chromosome number 2n = 2X= 20, 4X = 40 and 6X = 60 with tetraploids being the most common (Saxena and Chandra, 2006). 

Reproductive Biology

D. annulatum is a highly apomictic species, but it also shows facultative apomixis and different levels of sexual reproduction. It is predominantly wind-pollinated (Ambasht and Maurya, 1970; Cook et al., 2005; Saxena and Chandra, 2006; Vibrans, 2011). 

Environmental Requirements

D. annulatum grows in dry to moist environments from sea level up to 1600 m asl with mean annual rainfall ranging from 300 to 1400 mm and annual mean temperatures of 12.5 to 27.5°C (Duke, 1983; FAO, 2014). It is well-adapted to drought and can grow in areas with dry seasons lasting up to 8 months (Cook et al., 2005). D. annulatum is also adapted to a wide variety of soil types from sandy to heavy soils with neutral to alkaline conditions. It can tolerate poor drainage, salinity, and seasonal burning but not waterlogged conditions (Cook et al., 2005).

Climate

Air Temperature

Rainfall

Rainfall Regime

Soil Tolerances

Soil drainage

• free

Soil reaction

• acid
• neutral

Soil texture

• heavy
• light
• medium

Special soil tolerances

• infertile
• saline

Natural enemies

Notes on Natural Enemies

D. annulatum is parasitized by Striga asiatica (Orobanchaceae) and several Claviceps species may generate a significant reduction in seed production. In addition, a wide number of fungal species have been reported affecting this species (Cook et al., 2005; FAO, 2014): many of these are listed in the Natural Enemies Table.

Means of Movement and Dispersal

D. annulatum spreads mainly by seeds, but also by stolons. It produces large numbers of seeds which are mainly wind-dispersed. Seeds can also be dispersed secondarily by animals such as birds and by cattle, agricultural machinery and as a contaminant in grass seeds, agricultural products, and soil (Cook et al., 2005; FAO, 2014; Prota4U, 2013).

Pathway Causes

Pathway Vectors

Impact Summary

Environmental Impact

D. annulatum behaves as a grass weed and competes aggressively once established, often suppressing other species and forming dense stands. It has the potential to alter ecosystem functions by altering fire regimes, hydrology cycles, nutrient cycles, and community composition because it is drought-resistant and tolerates seasonal burning and saline conditions (Cook et al., 2005; Vibrans, 2011; FAO, 2014; PIER, 2014). In Australia, this species is highly invasive in arid wetlands where it is competing aggressively with important native grasses such as the species Eriachne benthamii (Queensland Department of Primary Industries and Fisheries, 2011). In Texas (USA), this species has escaped from cultivation and is outcompeting native grasses in bluestem coastal grassland communities (Barkworth et al., 2003; Queensland Department of Primary Industries and Fisheries, 2011).

Threatened Species

Risk and Impact Factors

• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• Highly adaptable to different environments
• Is a habitat generalist
• Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
• Pioneering in disturbed areas
• Highly mobile locally
• Benefits from human association (i.e. it is a human commensal)
• Long lived
• Fast growing
• Has high reproductive potential
• Reproduces asexually

• Altered trophic level
• Ecosystem change/ habitat alteration
• Modification of fire regime
• Modification of hydrology
• Modification of nutrient regime
• Modification of successional patterns
• Monoculture formation
• Reduced native biodiversity
• Soil accretion
• Threat to/ loss of native species

• Competition - monopolizing resources
• Competition - smothering
• Competition - strangling
• Pest and disease transmission
• Herbivory/grazing/browsing
• Hybridization
• Rapid growth
• Rooting

• Highly likely to be transported internationally accidentally
• Highly likely to be transported internationally deliberately
• Difficult to identify/detect as a commodity contaminant

Uses

D. annulatum is one of the most commercialized fodder grasses, especially in India and SE Asia because it is adapted to intense grazing and tolerates drought and fire (Duke, 1983). It is often planted as pasture for grazing and as a hay and silage grass. D. annulatum is also used for controlling soil erosion on sloping fields and to improve degraded grasslands (Cook et al., 2005; Prota4U, 2013).

Uses List

Animal feed, fodder, forage

• Fodder/animal feed
• Forage

Environmental

• Erosion control or dune stabilization
• Soil improvement

Similarities to Other Species/Conditions

Dichanthium is closely related to Bothriochloa, but can be distinguished by its pedicels and rachis internodes being solid and lacking a median, purple line (eFloras, 2014).

Prevention and Control

Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

D. annulatum is susceptible to glyphosate (Cook et al., 2005).

References

Acevedo-Rodríguez P, Strong MT, 2012. Catalogue of the Seed Plants of the West Indies. Smithsonian Contributions to Botany, 98:1192 pp. Washington DC, USA: Smithsonian Institution. http://botany.si.edu/Antilles/WestIndies/catalog.htm

Agarwal DK, Sanjeev Gupta, Roy AK, Gupta SR, 1999. Study on agromorphological variation vis-à-vis geographical distribution in marvel grass (Dichanthium annulatum L. (Stapf.)). Plant Genetic Resources Newsletter, No. 118:27-29.

Ambasht RS, Maurya AN, 1970. Reproductive capacity of Dichanthium annulatum Stapf. in relation to biotic factors. Tropical Ecology, 11(2):186-193.

Amit Chauhan, Singh DK, Dhakre JS, 2005. Additions to the Poaceae of Ladakh, Jammu & Kashmir. Journal of Economic and Taxonomic Botany, 29(1):224-226.

Barkworth ME, Capels KM, Long S, Piep MB, 2003. Dichanthium. Flora of North America, volume 25. http://herbarium.usu.edu/webmanual/

Broome R, Sabir K, Carrington S, 2007. Plants of the Eastern Caribbean. Online database. Barbados: University of the West Indies. http://ecflora.cavehill.uwi.edu/index.html

Chacón E, Saborío G, 2012. Red Interamericana de Información de Especies Invasoras, Costa Rica ([English title not available]). San José, Costa Rica: Asociación para la Conservación y el Estudio de la Biodiversidad. http://invasoras.acebio.org

Chong KY, Tan HTW, Corlett RT, 2009. A checklist of the total vascular plant flora of Singapore: native, naturalised and cultivated species. Singapore: Raffles Museum of Biodiversity Research, National University of Singapore, 273 pp. http://lkcnhm.nus.edu.sg/nus/pdf/PUBLICATION/LKCNH%20Museum%20Books/LKCNHM%20Books/flora_of_singapore_tc.pdf

Clayton WD, Govaerts R, Harman KT, Williamson H, Vorontsova M, 2014. World Checklist of Poaceae. Richmond, UK: Royal Botanic Gardens, Kew. http://apps.kew.org/wcsp/

Cook B, Pengelly B, Brown S, Donnelly J, Eagle D, Franco A, Hanson J, Mullen B, Partridge I, Peters M, Schultze-Kraft R, 2005. Tropical Forages: an interactive selection tool. Brisbane, Australia: CSIRO, DPI&F (Qld), CIAT and ILRI. http://www.tropicalforages.info/

Cruz Trujillo GM, Rodríguez Delgado O, Wildpret de la Torre W, 2008. Dichanthium annulatum (Forssk.) Stapf: a new grass species for the Canary Islands. (Dichanthium annulatum (Forssk.) Stapf, una gramínea nueva para las Islas Canarias.) Botanica Macaronesica, No.27:137-140.

DAISIE, 2014. Delivering Alien Invasive Species Inventories for Europe. European Invasive Alien Species Gateway. www.europe-aliens.org/default.do

Duistermaat H, 2004. New grass (Poaceae) records for Singapore, including Panicum laxum new for Asia. Gardens' Bulletin (Singapore), 56(1/2):29-42.

Duke JA, 1983. Handbook of Energy Crops. Unpublished. Purdue University, West Lafayette, Indiana, USA: Centre for New Crops and Plant Products. World Wide Web page at http://www.hort.purdue.edu/newcrop/Indices/index_ab.html.

eFloras, 2014. Flora of China. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.efloras.org/flora_page.aspx?flora_id=2

FAO, 2014. Grassland species profiles. http://www.fao.org/ag/AGP/AGPC/doc/Gbase/Default.htm

Funk V, Hollowell T, Berry P, Kelloff C, Alexander SN, 2007. Checklist of the plants of the Guiana Shield (Venezuela: Amazonas, Bolivar, Delta Amacuro; Guyana, Surinam, French Guiana). Contributions from the United States National Herbarium, 584 pp.

González-Torres LR, Rankin R, Palmarola A (eds), 2012. Invasive plants in Cuba. (Plantas Invasoras en Cuba.) Bissea: Boletin sobre Conservacion de Plantad del Jardin Botanico Nacional, 6:1-140.

Hokche O, Berry PE, Huber O, 2008. Nuevo Catálogo de la Flora Vascular de Venezuela (New catalogue of the vascular flora of Venezuela). Caracas, Venezuela: Fundación Instituto Botánico de Venezuela, 860 pp.

MacKee HS, 1994. Catalogue of introduced and cultivated plants in New Caledonia. (Catalogue des plantes introduites et cultivées en Nouvelle-Calédonie.) Paris, France: Muséum National d'Histoire Naturelle, unpaginated.

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

Perkins AJ, 2014. Dichanthium annulatum (Forssk.) Stapf. in PlantNET. Sydney, Australia: The Plant Information Network System of The Royal Botanic Gardens and Domain Trust. http://plantnet.rbgsyd.nsw.gov.au

PIER, 2014. Pacific Islands Ecosystems at Risk. Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

Prota4U, 2013. PROTA4U web database. Grubben GJH, Denton OA, eds. Wageningen, Netherlands: Plant Resources of Tropical Africa. http://www.prota4u.org/search.asp

Queensland Department of Primary Industries and Fisheries, 2011. Special edition of Environmental Weeds of Australia for Biosecurity Queensland., Australia: The University of Queensland and Department of Primary Industries and Fisheries. http://keyserver.lucidcentral.org/weeds/data/03030800-0b07-490a-8d04-0605030c0f01/media/Html/Index.htm

Randall RP, 2012. A Global Compendium of Weeds. Perth, Australia: Department of Agriculture and Food Western Australia, 1124 pp. http://www.cabi.org/isc/FullTextPDF/2013/20133109119.pdf

Saxena R, Chandra AA, 2006. RAPD and cytological analyses and histological changes caused by moisture stress in Dichanthium annulatum accessions. Cytologia, 71:197-204.

Smith AC, 1979. Flora vitiensis nova. A new flora of Fiji. Hawaii, Pacific Tropical Botanical Garden, 1:495.

Stevens PF, 2012. Angiosperm Phylogeny Website. http://www.mobot.org/MOBOT/research/APweb/

US Fish and Wildlife Service, 1994. In: Aristida chaseae, Lyonia truncata var. proctorii, and Vernonia proctorii Recovery Plan. US Fish and Wildlife Service, 21 pp.

USDA-ARS, 2014. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

USDA-NRCS, 2014. The PLANTS Database. Baton Rouge, USA: National Plant Data Center. http://plants.usda.gov/

Vibrans H, 2011. [English title not available]. (Malezas de México. Listado alfabético por familia, géneroy especie.) . http://www.conabio.gob.mx/malezasdemexico/2inicio/paginas/listaplantas

Villaseñor JL, Espinosa-Garcia FJ, 2004. The alien flowering plants of Mexico. Diversity and Distributions, 10(2):113-123.

Wagner WL, Herbst DR, Sohmer SH, 1999. Manual of the flowering plants of Hawaii. Revised edition. Honolulu, Hawaii, USA: University of Hawaii Press/Bishop Museum Press, 1919 pp.

Whistler WA, 1988. Checklist of the weed flora of Western Polynesia. An annotated list of the weed species of Samoa, Tonga, Niue, and Wallis and Futuna, along with the earliest dates of collection and the local names. Technical Paper, South Pacific Commission, No. 194:69 pp.

Whistler WA, 1996. Botanical survey of Diego Garcia, Chagos Archipelago, British Indian Ocean Territory. Isle Botanica (online), 49 pp. http://www.zianet.com/tedmorris/dg/2005NRMP-Appendixe-botanicalsurvey.pdf

Zuloaga FO, Morrone O, Belgrano MJ, Marticorena C, Marchesi E, 2008. [English title not available]. (Catálogo de las Plantas Vasculares del Cono Sur (Argentina, Sur de Brasil, Chile, Paraguay y Uruguay).) Monographs in Systematic Botany from the Missouri Botanical Garden, 107:1-3348.

Distribution References

Acevedo-Rodríguez P, Strong M T, 2012. Catalogue of the Seed Plants of the West Indies. Washington, DC, USA: Smithsonian Institution. 1192 pp. http://botany.si.edu/Antilles/WestIndies/catalog.htm

Broome R, Sabir K, Carrington S, 2007. Plants of the Eastern Caribbean. Online database., Barbados: University of the West Indies. http://ecflora.cavehill.uwi.edu/index.htm

CABI, Undated. Compendium record. Wallingford, UK: CABI

CABI, Undated a. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI

Chacón E, Saborío G, 2012. [English title not available]. (Red Interamericana de Información de Especies Invasoras, Costa Rica)., San José, Costa Rica: Asociación para la Conservación y el Estudio de la Biodiversidad. http://invasoras.acebio.org

Chong K Y, Tan H T W, Corlett R T, 2009. A checklist of the total vascular plant flora of Singapore: native, naturalised and cultivated species. Singapore: Raffles Museum of Biodiversity Research, National University of Singapore. 273 pp. https://lkcnhm.nus.edu.sg/app/uploads/2017/04/flora_of_singapore_tc.pdf

Clayton WD, Govaerts R, Harman KT, Williamson H, Vorontsova M, 2014. World Checklist of Poaceae., Richmond, UK: Royal Botanic Gardens, Kew. http://apps.kew.org/wcsp/

DAISIE, 2014. Delivering Alien Invasive Species Inventories for Europe. http://www.europe-aliens.org/

Funk V, Hollowell T, Berry P, Kelloff C, Alexander S N, 2007. Contributions from the United States National Herbarium, Washington, USA: Department of Systematic Biology - Botany, National Museum of Natural History, Smithsonian Institution. 55, 584 pp.

González-Torres LR, Rankin R, Palmarola A, 2012. Invasive plants in Cuba. (Plantas Invasoras en Cuba). In: Bissea: Boletin sobre Conservacion de Plantad del Jardin Botanico Nacional, 6 [ed. by González-Torres LR, Rankin R, Palmarola A]. 1-140.

Hokche O, Berry PE, Huber O, 2008. New catalogue of the vascular flora of Venezuela. (Nuevo Catálogo de la Flora Vascular de Venezuela)., Caracas, Venezuela: Fundación Instituto Botánico de Venezuela. 860 pp.

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

Mito T, Uesugi T, 2004. Invasive alien species in Japan: the status quo and the new regulation for prevention of their adverse effects. In: Global Environmental Research, 8 171-191.

Perkins AJ, 2014. Dichanthium annulatum (Forssk.) Stapf. In: PlantNET The Plant Information Network System of The Royal Botanic Gardens and Domain Trust, Sydney, Australia: The Plant Information Network System of The Royal Botanic Gardens and Domain Trust. http://plantnet.rbgsyd.nsw.gov.au

PIER, 2014. Pacific Islands Ecosystems at Risk., Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

Smith AC, 1979. Flora vitiensis nova. A new flora of Fiji., 1 Hawaii, Pacific Tropical Botanical Garden. 495.

USDA-ARS, 2014. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysimple.aspx

USDA-NRCS, 2014. The PLANTS Database. Greensboro, North Carolina, USA: National Plant Data Team. https://plants.sc.egov.usda.gov

Villaseñor J L, Espinosa-Garcia F J, 2004. The alien flowering plants of Mexico. Diversity and Distributions. 10 (2), 113-123. DOI:10.1111/j.1366-9516.2004.00059.x

Wagner W L, Herbst D R, Sohmer S H, 1999. Manual of the flowering plants of Hawai'i, Vols. 1 & 2. Honolulu, USA: University of Hawai'i Press/Bishop Museum Press. 1918 + [1] pp.

Whistler W A, 1988. Checklist of the weed flora of Western Polynesia. An annotated list of the weed species of Samoa, Tonga, Niue, and Wallis and Futuna, along with the earliest dates of collection and the local names. In: Technical Paper, South Pacific Commission, 69 pp.

Whistler WA, 1996. Botanical survey of Diego Garcia, Chagos Archipelago, British Indian Ocean Territory., Isle Botanica. 49 pp. http://www.zianet.com/tedmorris/dg/2005NRMP-Appendixe-botanicalsurvey.pdf

Zuloaga FO, Morrone O, Belgrano MJ, Marticorena C, Marchesi E, 2008. [English title not available]. (Catálogo de las Plantas Vasculares del Cono Sur (Argentina, Sur de Brasil, Chile, Paraguay y Uruguay)). In: Monographs in Systematic Botany from the Missouri Botanical Garden, 107 1-3348.

Links to Websites

Contributors

11/02/14 Original text by:

Julissa Rojas-Sandoval, Department of Botany-Smithsonian NMNH, Washington DC, USA

Pedro Acevedo-Rodríguez, Department of Botany-Smithsonian NMNH, Washington DC, USA

Distribution Maps