Desmodium incanum
(creeping beggerweed)

Pictures

Picture	Title	Caption	Copyright
Title Flowers Caption Desmodium incanum (creeping beggerweed, Spanish clover); flowers. Makawao, Maui, Hawaii, USA. April, 2008. Copyright ©Forest Starr & Kim Starr - CC BY 4.0	Flowers	Desmodium incanum (creeping beggerweed, Spanish clover); flowers. Makawao, Maui, Hawaii, USA. April, 2008.	©Forest Starr & Kim Starr - CC BY 4.0
Title Flowering habit Caption Desmodium incanum (creeping beggerweed, Spanish clover); flowering habit. Makawao, Maui, Hawaii, USA. April, 2008. Copyright ©Forest Starr & Kim Starr - CC BY 4.0	Flowering habit	Desmodium incanum (creeping beggerweed, Spanish clover); flowering habit. Makawao, Maui, Hawaii, USA. April, 2008.	©Forest Starr & Kim Starr - CC BY 4.0
Title Leaves Caption Desmodium incanum (creeping beggerweed, Spanish clover); close-up of leaves. Kipahulu LZ Haleakala National Park, Maui, Hawaii, USA. February, 2009. Copyright ©Forest Starr & Kim Starr - CC BY 4.0	Leaves	Desmodium incanum (creeping beggerweed, Spanish clover); close-up of leaves. Kipahulu LZ Haleakala National Park, Maui, Hawaii, USA. February, 2009.	©Forest Starr & Kim Starr - CC BY 4.0
Title Flowers and developing seedpods Caption Desmodium incanum (creeping beggerweed, Spanish clover); flowers and developing seedpods. Kipahulu LZ Haleakala National Park, Maui, Hawaii, USA. February, 2009. Copyright ©Forest Starr & Kim Starr - CC BY 4.0	Flowers and developing seedpods	Desmodium incanum (creeping beggerweed, Spanish clover); flowers and developing seedpods. Kipahulu LZ Haleakala National Park, Maui, Hawaii, USA. February, 2009.	©Forest Starr & Kim Starr - CC BY 4.0
Title Burs stuck to trousers and boots Caption Desmodium incanum (creeping beggerweed, Spanish clover); burs stuck to trouser legs and boot laces. Piiholo, Maui, Hawaii, USA. August, 2013. Copyright ©Forest Starr & Kim Starr - CC BY 4.0	Burs stuck to trousers and boots	Desmodium incanum (creeping beggerweed, Spanish clover); burs stuck to trouser legs and boot laces. Piiholo, Maui, Hawaii, USA. August, 2013.	©Forest Starr & Kim Starr - CC BY 4.0

Identity

Preferred Scientific Name

• Desmodium incanum DC.

Preferred Common Name

• creeping beggerweed

Other Scientific Names

• Aeschynomene incana (SW.) G. Mey.
• Aeschynomene spicata Poir.
• Desmodium canum (J.F. Gmel.) Schinz & Thell.
• Desmodium frutescens Schindl.
• Desmodium frutescens var. amblyophyllum (Urb.) Schindl.
• Desmodium incanum var. supinum (DC.) Hooker & Arnott
• Desmodium portoricense (Spreng.) G.Don
• Desmodium supinum (Sw.) DC.
• Desmodium supinum var. amblyophyllum Urb.
• Hedysarum canescens Mill.
• Hedysarum canum J.F. Gmel.
• Hedysarum incanum Sw.
• Hedysarum racemiferum J.F. Gmel.
• Hedysarum racemosum Aubl.
• Hedysarum supinum Sw.
• Meibomia adscendens var. incana (Sw.) Kuntze
• Meibomia cana S. F. Blake
• Meibomia incana (Sw.) Vail
• Meibomia supina (Sw.) Britton

International Common Names

• English: beggar's lice; creeping beggarweed; kaimi-clover; Spanish clover; tick trefoil; wild peanut; zarzabacoa comun
• Spanish: pega-pega; zarzabacoa comun
• French: colle-colle; gros treff; gros trefle; herb gallon; pois madame
• Portuguese: amor-de-campo; carrapicho-beico-de-boi; pega-pega

Local Common Names

• Cook Islands: piripiri
• Dominican Republic: amor seco
• Lesser Antilles: sweethearts
• Madagascar: famalakantsy; pelatsifotra; savisaha; tialamba
• Marshall Islands: lattil pako
• USA/Hawaii: ka'imi

Summary of Invasiveness

Originally from New World tropics and subtropics, D. incanum is a perennial shrub that was promoted as a forage crop around the world, with further accidental spread. Although it has some value as forage, it is no longer highly regarded. Its main value lies in its nitrogen-fixing ability (for intercropping). D. incanum is regarded as weedy both within and outside of its native range. It occupies roadsides, rough places, pastures, unplanted grasslands, volcanic soils and in dry to wet areas with low stature vegetation. It is an occasional problem in coffee, banana and rice crops. D. incanum is able to withstand and thrive with frequent herbicidal treatments, grazing, cultivation and fire. Its dispersal is mediated by sticky seeds, which can adhere to people, animals and machinery, making it a nuisance. It is able to compete and persist well with other dominant grasses, including invasive ones. Its spreading (scrambly) low stature habit helps it to both dominate other low vegetation and to persist within it. It could also carry pests and diseases of legume crops such as peanut. 

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Fabales
•                         Family: Fabaceae
•                             Subfamily: Faboideae
•                                 Genus: Desmodium
•                                     Species: Desmodium incanum

Notes on Taxonomy and Nomenclature

The most commonly used synonym in the literature is Desmodiumcanum but this was the result of the incorrect use of an illegitimate basionym (Nicolson, 1978). Sometimes the authority for D. incanum is given as (Sw.) DC, but this is also incorrect. That name refers to a different taxon which is either Desmodium mauritianum (ILDIS, 2013) or Desmodium ramosissimum; these being synonyms of each other.

Desmodium comes from the Greek word desmos which means a bond or chain. Kaimi (correctly Ka’imi) clover, the Hawaiian name for the plant, means 'seeker'. The other English names, which refer to lice and ticks, relate to the sticky seeds that stick to clothes and pets like those pests. The number of common names for the plant in Madagascar implies that it is a common, well-known plant there (ILDIS, 2013).

Description

The following description is from Wagner et al. (1999):

Prostrate, ascending, or erect subshrubs or shrubs up to 1 m tall; stems pubescent with spreading hooked and longer straight hairs, glabrate. Leaves trifoliolate, leaflets subcoriaceous, usually elliptic or narrowly elliptic, terminal one 4-9 cm long, 1.5-4.5 cm wide, lateral nerves conspicuous, upper surface with minute hooked and straight hairs, lower surface densely appressed pubescent, apex obtuse or often acute, petioles 1-4 cm long. Flowers numerous in racemose inflorescences 5-12 cm long, rachis densely pubescent with minute hooked hairs, pedicels 3-10 mm long, persistent after articles fall, pubescent with minute hooked hairs; corolla pink to purplish, 5-6 mm long. Pods stipitate, (3) 4-5 (-8)-jointed, (2-) 2.5-3 (-4) cm long, densely pubescent with hooked hairs, articles nearly semicircular, 4-5 mm long, ca. 3 mm wide.

The following description is from Acevedo-Rodríguez (2005):

Decumbent or scandent herb, much branched from the woody base, attaining 1-1.5 m in length. Stems more or less cylindrical, appressed pubescent or pilose, glabrescent with age. Leaves alternate, trifoliolate; leaflets 3-9 × 0.8-3.8 cm, elliptical, oblong, ovate, lanceolate, or exceptionally rounded, chartaceous, involute, the apex obtuse or acute, mucronate, the base obtuse or rounded, the margins entire; upper surface dark green, puberulent; lower surface grayish green, strigulose, with prominent venation; petiolules 1.5-3 mm long, pilose; petioles 0.7-3 cm long, pilose, canaliculate; rachis 3-10 mm long, pilose, canaliculate; stipules ovate to lanceolate, 6-10 mm long, persistent; stipels minute, subulate. Inflorescences of terminal pseudoracemes, 10-18 cm long, pubescent; bracts minute, persistent. Calyx green, campanulate, 2.5-3 mm long, strigulose; corolla pink or pink-violet, the standard ovate, 5-6 mm long, retuse, unguiculate at the base, the wings and the keel as long as the standard. Legume uncinate-pubescent, 2-4 cm long, crenate on the ventral suture, with 5-8 fertile segments. Seeds 2.5-3.5 mm long, oblong elliptical, light brown. 

Plant Type

Distribution

D. incanum is native to Mexico, Central America, the Caribbean and South America. Some accounts report that the species is native in Florida and Texas (Green et al., 2008; ILDIS, 2013), although important databases with expert input for the flora regard it as introduced (Wunderlin and Hansen, 2012; USDA-NRCS, 2013). It is not clear whether the plant is native in Galapagos (Simbaña and Tye, 2009).  

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 timing of the spread of the plant in the tropics including most oceanic islands is unclear. For a short period during the twentieth century it was promoted as a forage plant and may have been intentionally planted, but its use and promotion has diminished since (Jones and Bunch, 2003; ILDIS, 2013). With its seeds being such able hitchhikers on clothes, socks, legs, shoes, shoe-laces, animals, pets and livestock, in most case it is likely to have spread accidentally.

D. incanum was first noted on Round Island, Mauritius, in 1987, and was probably introduced during invasive animal and plant management efforts (C. Griffiths, University of Bristol, UK, personal communication, 2013; Griffiths et al., 2013). Today it is prolific and widespread on Round Island. Both the Aldabran giant and Radiata tortoises consume large quantities of this legume, and as a result it has changed its growth form, becoming more prostrate (and possibly occupying more space). Seed productivity, however, has fallen where tortoises continually graze this species (C. Griffiths, University of Bristol, UK, personal communication, 2013).

Introductions

Risk of Introduction

D. incanum is mostly spread via its sticky seeds and could easily be transported anywhere on people, camping equipment, pets, vehicles etc. It is known to be transported to new sites by conservation workers, for example on helicopter landing zones in relatively intact forest sites in Hawaii (Forest Starr, Starr Environmental, Hawaii USA, personal communication, 2013) and during invasive mammal eradication efforts on Round Island, Mauritius in 1987 (Griffiths et al., 2013). Spread via wool fleeces, hides or other animal products, is also possible.

Habitat

D. incanum is known to occur in disturbed places on both inhabited and uninhabited islands of the Pacific (Whistler, 1998; Starr and Starr, 2006; Starr et al., 2006; US Forest Service, 2014). It tends to grow alongside trails and fence lines, but does not do well under the forest canopy (J. Beachy and S. Kaye, Big Island Invasive Species Committee, Hawaii USA, personal communication, 2013).

The plant occurs in important habitats in Florida, including pine rockland (Green et al., 2008), dry prairies (Orzell and Bridges, 2006), Cedar Key State Reserve (Amoroso and Judd, 1995) and in various protected areas (e.g. Hubbard and Judd, 2013) including the Castillo de San Marcos National Monument (Zomlefer and Giannasi, 2005). It is a significant weed along roads, trails and disturbed areas in Guana Tolomato Matanzas National Estuarine Research Reserve (E. Montgomery, Hawaii Invasive Species Council, Hawaii USA, personal communication, 2013). 

Habitat List

Hosts/Species Affected

Banana, rice and pastures are the main crops for which D. incanum is a recognized weed. However, this species is usually one of a suite of problematic species rather than a principal problem (Bergin, 2003; Jones and Bunch, 2003; Isaac et al., 2007; Guglieri-Caporal et al., 2010; Mesquita et al., 2013). 

Host Plants and Other Plants Affected

Biology and Ecology

Genetics

The chromosome number is 2n = 22 (Young, 1940; Schifino-Wittmann, 2000). Hybrids with Desmodium uncinatum are possible (Chow and Crowder, 1973), but it is unclear whether they occur in the wild.

Reproductive biology

The plant is commonly pollinated by bees (or other generalist pollinators) and via self-fertilisation (Etcheverry et al., 2010). Flowers are diurnal and triggered explosively by tension created in the keel and wings of the flower. After the initial visit most of the pollen is released and few visits from bees are likely again (Alemán et al., 2014). Anthers of D. incanum dehisce in the mid-afternoon, with 14-15.2 hours of light in the glasshouse, and flowers can be tripped by insect pollinators. Even in the absence of pollinators, seed set can exceed 50% (Chow and Crowder, 1974). Reproduction is often via seeds, and dispersal via the stick tight adherence provided by its hairy Velcro-like seed pods (Mori and Brown, 1998). The plant can root from stems, and fragments of the tap root can resprout (University of Florida Nassau County Extension, 2013).

Chemical and mechanical scarification and heating of seeds improves germination rates which were about 25% in one study (Franke and Baseggio, 1998).

Physiology and phenology

In warm tropical areas, D. incanum flowers and fruits all year round. Elsewhere in subtropical to temperate sites it may flower and fruit only in the warm season. Under high levels of herbivory it can adopt a more spreading prostate habit (C. Griffiths, University of Bristol, UK, personal communication, 2013).

Associations

D. incanum has symbiotic associations with root-nodulating nitrogen-fixing rhizobial bacteria Bradyrhizobium elkanii (Kuykendall et al., 1992; Menna et al., 2006; 2009).

Environmental requirements

D. incanum does not tolerate long periods of wet conditions, preferring well-drained soils, and can withstand fairly arid conditions. It is often regarded as a poor competitor, except under disturbed conditions with high light levels. 

Natural Food Sources

Climate

Latitude/Altitude Ranges

Natural enemies

Notes on Natural Enemies

Several genera of fungi and other plant pathogens have been shown to attack D. incanum, and other diseases are known to infest Desmodium spp., but no information is given about these species (Lenné and Stanton, 1990). Viruses include peanut mottle virus (Demski et al., 1981) and peanut stunt cucumovirus (Gillaspie Jr and Ghabrial, 1998). As such, the plant could harbour a number of diseases and pests that attack peanut crops. It is resistant to root-knotting nematodes and is a suggested source of resistance genes (Quesenberry et al., 2008). A few species of pea or bean weevils (genus Meibomeus) are known to feed on D. incanum (Romero and Johnson, 2002). Any predators, diseases or pests of peas and beans could potentially impact this species. Some generalist insect predators of beans, such as Urbanus proteus, are known to attack D. incanum, and there are probably many more of these than are documented here (Capinera, 2011). The seed beetle Acanthoscelides puelliopsis is a seed predator of D. incanum, and is from a group that is specialized on plants in the subfamily Papilionoideae (Johnson and Siemens, 1995).

Pathway Causes

Pathway Vectors

Plant Trade

Economic Impact

D. incanum is frequently a significant component of productive grasslands, and is variously viewed as a secondary and valued forage or a weed both within and outside of its native range (Hacker et al., 1996; Jones and Bunch, 2003; Garbulsky and Deregibus, 2006; Guglieri-Caporal et al., 2010). Due to its nitrogen-fixing abilities it is possible that it has an impact on soil nutrient cycles (these could be viewed as positive or negative), but no studies quantify this, or any possible consequences to co-occurring species – not even in the studies of it as a forage. It is a common weed of slash and burn agriculture, and disturbed secondary vegetation (Dalle and de Blois, 2006; Mitja and Miranda, 2010; Randriamalala et al., 2012). It is widespread in many regions where it occurs, but rarely the most significant weed (e.g. Lárez Rivas, 2007). Some costs may be offset at the most minor scales by the benefits derived from its use as a diuretic and anti-inflammatory (Vargas et al., 1991).

On the Big Island of Hawaii (USA), ranchers on (windward) Big Island welcome D. incanum in the pasture. It disappears after about two years of continuous cattle grazing, but comes back vigorously when the field is rested. It does well in a mix with California grass (Urochloa mutica), which also gives way to Paspalum conjugatum after a year or two without steady rotation. D. incanum is also controlled by mowing on coffee farms in the South Kona area of Big Island. 

Some extension offices in universities and state government, as well as lawn management companies in North America, describe its identification, control and management as a lawn weed (e.g. University of Florida Extension Office, 2013).

Environmental Impact

Impact on Habitats

D. incanum is known to occur in disturbed places on both inhabited and uninhabited islands of the Pacific, and while it is often widespread, it is rarely more than locally abundant (Whistler, 1998; Starr and Starr, 2006; Starr et al., 2006; US Forest Service, 2014). The offshore islands and islets of Hawaii are mainly protected areas and provide a habitat for seabirds. It occurs in protected areas in Hawaii on Molokai at Kalaupapa National Historic Park (Medeiros et al., 1996) and on Maui in Haleakala National Park (Welton and Haus, 2008), but is simply listed with a suite of other invasive species and no negative impacts are attributed to it alone. This weed is occasionally controlled in Hawaii in conservation areas, where it is never a major component. It tends to grow alongside trails and fence lines, but does not do well under the forest canopy (J. Beachy and S. Kaye, Big Island Invasive Species Committee, Hawaii USA, personal communication, 2013). Its control is usually for its nuisance value and because conservation workers do not want to see it even as a trailside weed in more pristine areas. A congener Desmodium tortuosum is viewed as a more significant (and already widespread) problem in Hawaii (J. Beachy and S. Kaye, Big Island Invasive Species Committee, Hawaii USA, personal communication, 2013).

The plant occurs in important habitats in Florida, including pine rockland (Green et al., 2008), dry prairies (Orzell and Bridges, 2006), Cedar Key State Reserve (Amoroso and Judd, 1995)and in various protected areas (e.g. Hubbard and Judd, 2013)including the Castillo de San Marcos National Monument (Zomlefer and Giannasi, 2005). It is a significant weed along roads, trails and disturbed areas in Guana Tolomato Matanzas National Estuarine Research Reserve but doesn’t overly compete with important species, or in relatively intact native plant communities (E. Montgomery, Hawaii Invasive Species Council, Hawaii USA, personal communication, 2013). It is widespread in the lower two-thirds of the state of Florida and is also documented in several of the northern counties (Wunderlin and Hansen, 2012), and so its occurrence in protected areas is unsurprising.

Impact on Biodiversity

D. incanum has been listed as co-occurring with important species of conservation concern in the Galapagos (Linum cratericola), where it may or may not be native (Simbaña and Tye, 2009). However, it is not described as a threat to these species. Similarly, it is listed as an invasive species in cockaded woodpecker (Picoides borealis) habitat on land used by the military for training, but is likely to pose little or no threat to the woodpecker or the vegetation upon which the woodpecker depends (Guertin and Tess, 2006). It is listed as one of many non-native species co-occurring with the threatened endemic species in Florida including Harrisia aboriginum, Argythamnia blodgettii and Indigoferatrita subsp. scabra (Bradley et al., 2004; Hodges and Bradley, 2006). The presumed mechanism for an impact is via competition, but no clear indication of it being a significant threat is given. In Hawaii, there is one instance where D. incanum threatens conservation values, where it (along with other non-native species) has invaded the West Maui population of Tetramolopium sylvae (Hank Oppenheimer, Maui Nui Plant Extinction Prevention Program, Hawaii USA, personal communication, 2013).

Threatened Species

Social Impact

Mainly a scourge for parents who have to pick it out of their children’s socks (and their own), and it frequently attaches to shoe laces, and trouser bottoms. It is best described as a major nuisance.

Conservation managers with awareness of the issue try not to spread weeds of any kind to pristine areas, and the diligence required can be time-consuming and stressful.

Risk and Impact Factors

• Invasive in its native range
• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• 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

• Ecosystem change/ habitat alteration
• Increases vulnerability to invasions
• Modification of nutrient regime
• Negatively impacts agriculture
• Negatively impacts livelihoods
• Reduced amenity values
• Reduced native biodiversity
• Threat to/ loss of endangered species
• Threat to/ loss of native species

• Competition - monopolizing resources
• Competition (unspecified)
• Pest and disease transmission
• Herbivory/grazing/browsing
• Rapid growth
• Produces spines, thorns or burrs

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

Uses

Economic Value

In pastures it has been promoted as a useful nitrogen-fixer and a source of fodder with extra protein. It may be useful because it survives herbivory, some herbicide applications, and dry conditions (Smith, 1995; Schifino-Wittmann, 2000; Garbulsky and Deregibus, 2006; ILDIS, 2013; Tropical Forages, 2013). It is secondarily preferred by livestock, however, and therefore ranchers must invest effort in maintaining the preferred grasses (Bergin, 2003; Jones and Bunch, 2003).

Social Benefit

D. incanum is used as a diuretic, stomachic, febrifuge and hemostatic in Central America (Setyowati-Indarto and Brink, 1999).

Environmental Services

Despite its status as an invasive species in the Pacific (US Forest Service, 2014), the USDA continues to promote the planting of this species to establish ground cover, in the context of agricultural and biodiversity values and for soil conservation measures (USDA-NRCS, 2012).

Uses List

Animal feed, fodder, forage

• Fodder/animal feed

Environmental

• Erosion control or dune stabilization
• Land reclamation
• Revegetation

Diagnosis

No methods are available for testing and screening this species, apart from the usual use of plant identification tools such as keys. DNA barcoding would be possible, since several DNA barcoding regions have been sampled and are available on GenBank (NCBI, 2014).

Detection and Inspection

Inspection of sticky D. incanum seeds and seed pods on equipment and clothing, and removing or cleaning of these, could slow or prevent the spread of this species.

Similarities to Other Species/Conditions

Desmodium is a large genus (around 450 species). Congeners in its native range are described with keys (Acevedo-Rodríguez, 2005; Lárez Rivas, 2007; Dutra et al., 2009) and from flora descriptions outside its native range (e.g. Wagner et al., 1999).

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.

Cultural control and sanitary measures

Reducing grazing frequency can favour co-occurring pasture grasses, and the competitive interaction can reduce D. incanum dominance (Bergin, 2003). Partially contradicting, one observer suggested continuous grazing may effectively control D. incancum to low levels, but that after release from grazing pressure it 'comes back with a vengeance' (S. Kaye, Big Island Invasive Species Committee, Hawaii USA, personal communication, 2013). Tortoises were found to significantly reduce the cover of introduced plants compared to native species on Round Island in Mauritius (Griffiths et al., 2013), and D. incanum formed a significant part of their diet. It is not clear whether the cover of D. incanum was reduced by tortoise herbivory. In most cases it is regarded as tolerant of herbivory, disturbance and tillage (Dalle and de Blois, 2006; Guglieri-Caporal et al., 2010; Randriamalala et al., 2012).

Physical/mechanical control

D. incanum reproduces by seed, stolons and broken taproots. Complete removal of the roots is required for effective control. It withstands or benefits from fire, intense grazing and tillage (Dalle and de Blois, 2006; Guglieri-Caporal et al., 2010; Randriamalala et al., 2012).

Movement control

Conservation workers in Hawaii and elsewhere are in the practice of wearing clean, D. incanum seed-free clothes when going to field sites that may be relatively weed-free. For example, this would be important when flying into conservation areas by helicopter.

Chemical control

Often D. incanum is a lawn weed and is considered hard to control because the tap root can persist after herbicide treatments (University of Florida Nassau County Extension, 2013). A pre-emergent herbicide can be used in spring to help control D. incanum in lawns (University of Florida Extension Office, 2013). One control method requires that you locate the main stem, cut it close to the ground and immediately “paint” the cut stem with a glyphosate-type product (University of Florida Extension Office, 2013). Control with triclopyr can be as high as 90% and does not kill associated grasses such as Paspalum notatum (Freitas et al., 2003). Use of herbicides generally to control it, or other weeds will not necessarily cause its decline, and can promote its dominance (Reinert et al., 2004). 

Gaps in Knowledge/Research Needs

This weed may be under-reported where it is common and is probably in more sites than reported. Its pattern and history of spread could perhaps be ascertained via population genetic studies.

References

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Capinera JL, 2011. Bean Leafroller, Urbanus proteus (Linnaeus) (Insecta: Lepidoptera: Hesperiidae). Gainesville, Florida, USA: University of Florida. http://edis.ifas.ufl.edu/pdffiles/IN/IN12700.pdf

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Distribution References

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Jones RM, Bunch GA, 2003. Experiences with farm pastures at the former CSIRO Samford Research Station, south-east Queensland, and how these relate to results from 40 years of research. In: Tropical Grasslands, 37 151-164.

Josekutty PC, Wakuk EE, Joseph MJ, 2002. Invasive weedy angiosperms in Kosrae, Federated States of Micronesia. In: Micronesica Supplement, 6 61-65.

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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, 8 171-193.

Orchard AE, 1994. Flora of Australia., 50 Canberra, Australia: Australian Government Publishing Service. 606 pp.

Parodi LR, 1943. (La vegetacion del Departamento de San Martin en Corrientes (Argentina)). In: Darwiniana, 6 127-178.

PIER, 2013. Pacific Islands Ecosystems at Risk., Honolulu, Hawaii, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

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Ragone D, Lorence DH, 2003. Botanical and Ethnobotanical Inventories of the National Park of American Samoa., Hawaii, USA: Pacific Cooperative Studies Unit, Department of Botany, University of Hawaii. 91 pp.

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Robertson SA, Todd DM, 1983. Vegetation of Fregate Island, Seychelles. Floristics and ecology of Western Indian Ocean islands. In: Atoll Research Bulletin No. 273, 253 pp.

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Smith AC, 1979. Flora Vitiensis nova: A new flora of Fiji., I Lawai, Kauai, Hawaii, USA: National Tropical Botanical Garden. 494 pp.

Smith EG, 1995. Preliminary evaluation of tropical forage legumes in Suakoko, Liberia. In: Pasturas Tropicales, 17 34-37.

Space JC, Imada CT, 2004. Report to the Republic of Kiribati on invasive plant species on the islands of Tarawa, Abemama, Butaritari and Maiana., Honolulu, Hawaii, USA: USDA Forest Service and Bishop Museum. 103 pp.

Space JC, Waterhouse B, Miles JE, Tiobech J, Rengulbai K, 2003. Report to the Republic of Palau on invasive plant species of environmental concern., Honolulu, Hawaii, USA: USDA Forest Service. 174 pp.

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Links to Websites

Contributors

13/1/14 Original text by:

Christopher E. Buddenhagen, Florida State University, USA

Distribution Maps