Chloris gayana
Family :
Famille :
Poaceae
Synonym(s) :
Synonyme(s) :
Chloris abyssinica Hochst. ex A. Rich. (USDA-ARS-NPGS 2023)
Chloris glabrata Andersson (Rojas-Sandoval 2020)
Chloris repens Hochst. (Rojas-Sandoval 2020)
Eustachys gayana (Kunth) Mundy (Rojas-Sandoval 2020)
Common Name(s) :
Nom(s) commun(s) :
Rhodesgrass
(English) (AOSA 2023; USDA-AMS 2023)
Rhodes grass (English) (Wiersema and León 1999; USDA-ARS-NPGS 2023; USDA-NRCS 2023)
Abyssinian grass (English) (Rojas-Sandoval 2020)
Herbe de Rhodes (French) (Wiersema and León 1999; USDA-ARS-NPGS 2023)
Hunyani grass (English) (Rojas-Sandoval 2020)
Rhodasgras (German) (Wiersema and León 1999; USDA-ARS-NPGS 2023)
Erba di Rhodes (Italian) (USDA-ARS-NPGS 2023)
Capim-de-Rhodes (Portuguese) (Wiersema and León 1999; USDA-ARS-NPGS 2023)
Grama de Rodas (Spanish) (Wiersema and León 1999; USDA-ARS-NPGS 2023)
Grama Rhodes (Spanish) (Rojas-Sandoval 2020)
Pasto de Rodas (Spanish) (Wiersema and León 1999; USDA-ARS-NPGS 2023)
Zacate conejo (Spanish) (Rojas-Sandoval 2020)
Zacate gordura (Spanish) (Rojas-Sandoval 2020)
Fei zhou hu wei cao (非洲虎尾草 Chinese) (efloras 2023)
Chloris gayana (rhodesgrass) spikelet consisting of attached floret group and upper (2nd) glume. Lower (1st) glume missing. Scale in mm.
Chloris gayana (rhodesgrass) spikelet consisting of attached floret group and upper (2nd) glume. Lower (1st) glume missing. Scale in mm.
Chloris gayana (rhodesgrass) attached floret group consisting of one fertile floret and two sterile florets (lateral view) removed from spikelet glumes and exposing callus hairs of basal floret. Scale in mm.
Chloris gayana (rhodesgrass) caryopsis in lateral view. Scale in mm.
Chloris gayana (rhodesgrass) caryopsis in hilum view. Scale in mm.
Chloris gayana (rhodesgrass) caryopsis in embryo view. Scale in mm.
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Overview
Aperçu
Regulation :
Remarques Réglementation:
- Quarantine lists of countries e.g. Mexico *may be updated without notice
Regulation Notes:
Chloris gayana is on the Honduras list of harmful organisms (USDA-PCIT-PExD 2023) (*may be updated without notice).
Distribution :
Répartition :
Chloris gayana is native to parts of Africa: Algeria, Botswana, Egypt, Ethiopia, Kenya, Lesotho, Mali, Morocco, Mozambique, Namibia, Niger, Senegal, Somalia, South Africa, Sudan, Tanzania, Tunisia, Uganda, Zimbabwe, Canary Islands, and Madeira Islands (USDA-ARS-NPGS 2023). It has since been introduced, cultivated, and naturalized in other parts of Africa and areas of Oceania, Mexico, Central America, South America, the United States of America, southern Europe, as well as temperate and tropical Asia (Loch et al. 2004; Rojas-Sandoval 2020; USDA-ARS-NPGS 2023).
Habitat and Crop Association :
Habitat et Cultures Associées :
Chloris gayana grows in open woodland, grassland, savannas, and along river and lakes (Cook et al. 2020; Rojas-Sandoval 2020). In tropical and subtropical regions C. gayana grows near sea level to 2000 m in elevation but can also survive in dry and semi-arid regions (Rojas-Sandoval 2020). It is adapted to a wide range of soil types and conditions, preferring sandy or red volcanic soils or clay loams that are well drained and have a pH range of 5.5 to 7.5 (Rojas-Sandoval 2020). C. gayana is drought tolerant, salt tolerant, and can tolerate short term seasonal waterlogging (Loch et al. 2004; Cook et al. 2020; Rojas-Sandoval 2020).
It is a considered a weed of native bushland, rainforest margins, agricultural fields, orchards, vineyards, planted pastures, parks, garden landscapes, roadsides, and other disturbed places (Rojas-Sandoval 2020). Once established, C. gayana can form nearly pure stands that overtake native vegetation (Rojas-Sandoval 2020).
Economic Use, cultivation area, and Weed Association :
Utilisation économique, zone de culture et association de mauvaises herbes :
Chloris gayana is used for silage, hay, and green chop for livestock feed, and for stabilization and revegetation of disturbed sites (Loch et al. 2004; Rojas-Sandoval 2020). This species is one of the most important perennial pasture grasses, widely cultivated throughout the tropical and subtropical regions of Africa, tropical and temperate Asia, Fiji, Brazil, Argentina, and Uruguay (USDA-ARS-NPGS 2023).
General Information
RENSEIGNEMENTS GÉNÉRAUX
Chloris gayana is C4 grass native to many tropical and subtropical regions of Africa and is cultivated as a forage in many parts of the world. As a warm-season pasture grass, C. gayana was first cultivated in South Africa around 1895, in Australia around 1902, and the United States a year later (Loche et al. 2004).
This naturally variable species consists of diploid and tetraploid types; the diploid types are generally subtropical in origin and cold tolerant, while the tetraploid types are generally tropical (Loche al. 2004). From these types, numerous cultivated varieties have been developed that serve a wide range of purpose for pasture, hay, and silage, as well as for soil stabilization and reclamation. (Loch et al. 2004). C. gayana is a salt-tolerant species occurring naturally on saline soils, using various physiological mechanisms to deal with salt toxicity, and excreting sodium ions via salt glands on its leaves (Loch et al. 2004).
C. gayana is considered one of the best C4 grasses for grazing horses because it has a low oxalate content, which helps limit the binding of dietary calcium in the digestive tract leading to calcium deficiency associated with skeletal malformations and other symptoms related to big head disease of horses (Loch et al. 2004; KER 2020).
C. gayana spreads by seed as well as vegetatively by stolons (Rojas-Sandoval 2020). In Australia, a yield of 100 to 200 kg of cleaned seed per hectare has been recorded depending on the variety and environmental conditions (Loch et al. 2004). Diploid cultivars produce more caryopses and show little or no seed dormancy compared to tetraploid cultivars (Loch et al. 2004). C. gayana has escaped cultivation to become naturalized in many areas. The seeds spread easily by wind and water and are usually introduced as a seed lot contaminant (Rojas-Sandoval 2020).
.Identification
Identification
-
Spikelet
Size
- Spikelet length: 3 – 5 mm (Baldwin et al. 2012).
- Lower glume (1st) length: 1 – 3 mm (Baldwin et al. 2012).
- Upper glume (2nd) length: 2 – 4.5 mm (Baldwin et al. 2012).
Shape
- Spikelet laterally compressed; spikelet body oval shaped in lateral view.
- Glumes narrowly teardrop shaped (lanceolate), pointed, keeled, usually not awned, shorter than florets (Barkworth et al. 2003).
Surface Texture
- Glumes thin and papery with short stiff hairs on keel.
Colour
- Spikelet light to dark yellowish-brown coloured.
Other Features
- Spikelet consists of one fertile and (1)2 – 4 staminate or sterile florets; the basal floret is fertile, rarely upper florets may be fertile.
- The florets of the spikelet disarticulate as a unit below the basal floret (glumes remain attached to the plant).
Chloris gayana (rhodesgrass) spikelet consisting of attached floret group and upper (2nd) glume. Lower (1st) glume missing. Scale in mm.
-
Fertile floret (basal floret in spikelet)
Size
Floret measurements taken in lateral view:
- Fertile floret (basal) length*: 3.0 – 4.0 mm (average 3.5 mm); width*: 0.4 – 1.0 mm (average 0.8 mm);
- Fertile floret (basal) awn length*: 2.0 – 6.4 mm (average 3.7 mm).
- Rachilla of basal floret about one-fourth the length of the floret.
*Note: minimum and maximum based on a random selection of 10 florets in normal range of this species using image measurement protocol (ISMA 2020). CDA-S-22525
Fertile floret measurements from the literature:
• Basal (fertile) floret length: 2.5–4.2 mm; width: 0.7–1.0 mm (Barkworth et al. 2003).
• Basal (fertile) floret length: 2 – 4 mm; width: less than 1 mm wide awn length: 1 – 7.5 mm (Baldwin et al. 2012).
• Basal (fertile) floret awn length: 1 – 10 mm (Cook et al. 2020).Fertile floret awn measurements from the literature:
• Basal (fertile) floret awn length: 1.5 – 6.5 mm (Barkworth et al. 2003).
• Basal (fertile) floret awn length: 1 – 7.5 mm (Baldwin et al. 2012).Shape
- Fertile floret laterally compressed, strongly keeled.
- Fertile floret oval to egg-shaped, slightly bulged out along keel (gibbous), longitudinally grooved on lateral faces.
- Lemmas of fertile florets entire or minutely bilobed at tip, awn extending from below the lemma tip.
- Palea inconspicuous, concealed by folded lemma and upper florets, two-keeled.
- Callus short, point of attachment oval.
Surface Texture
- Fertile floret lemma tough, firm, but flexible (cartilaginous), three-nerved.
- Fertile floret lemma with appressed silky hairs in lateral grooves and on either side of the lower half of the keel.
- Hairs along marginal nerves very long and straight, appearing more concentrated near lemma tip.
- Upper one-third of lemma back with short, stiff upward pointing hairs.
- Palea membranous.
- Closely spaced hairs along the palea keels.
- Rachilla of basal floret mostly smooth.
- Callus hairy around margin.
- Awns covered with short, stiff hairs.
Colour
- Fertile florets light or dark yellowish-brown coloured.
Chloris gayana (rhodesgrass) attached floret group consisting of one fertile floret and two sterile florets (lateral view) removed from spikelet glumes and exposing callus hairs of basal floret. Scale in mm.
-
Sterile florets (upper florets in spikelet)
Size
Floret measurements taken in lateral view:
- Second floret (sterile or staminate) length*: 2.4 – 3.5 mm (average 2.9 mm); width*: 0.4 – 0.7 mm (average 0.5 mm).
- Second floret awn length*: 0.9 – 5.3 mm (average 2.8 mm).
- Third or fourth sterile florets are progressively smaller than the second floret.
*Note: minimum and maximum based on a random selection of 10 florets in normal range of this species using image measurement protocol (ISMA 2020). CDA-S-22525
Floret measurements from the literature:
• Second floret length 2.2 – 3.2 mm; width 0.3 – 1 mm; distal florets progressively smaller (Barkworth et al. 2003).
• Second floret length: 1.8 – 3.2 mm (Baldwin et al. 2012).Awn measurements from literature
• Second floret awn 0.8 – 3.2 mm long; distal florets awn-tipped or unawned (Barkworth et al. 2003).
• Second floret awn length: 0.8 – 3 mm (Baldwin et al. 2012).
• Second floret awn length: 1.5 – 5.5 mm (Cook et al. 2020).Shape
- Second floret oblong to teardrop shaped.
- Lemmas of second florets entire or minutely bilobed at tip, awn extending from below the lemma tip.
- Uppermost floret(s) may be truncate at tip and may be short awned or unawned.
- Palea inconspicuous, concealed by folded lemma.
Surface Texture
- Second floret lemmas tough, firm, but flexible (cartilaginous), three-nerved.
- Upper one-third of lemma back with short stiff upward pointing hairs.
- Palea membranous.
- Awns covered with short, stiff hairs.
Colour
- Second and distal florets light or dark yellowish-brown coloured.
Chloris gayana (rhodesgrass) attached floret group consisting of one fertile floret and two sterile florets (lateral view) removed from spikelet glumes and exposing callus hairs of basal floret. Scale in mm.
-
Caryopsis
Size
- Caryopsis length*: 1.3 – 2.0 mm (average 1.8 mm); width (embryo view)*: 0.3 – 0.6 mm (average 0.4 mm); thickness (lateral view)*: 0.3 – 0.6 mm (average 0.5 mm).
*Note: minimum and maximum based on a random selection of 10 caryopses in normal range of this species using image measurement protocol (ISMA 2020). CDA-S-22525
Caryopsis measurements from the literature:
• Caryopsis length: 1.0 – 1.5 mm; width: about 0.5 mm (Barkworth et al. 2003).
• Caryopsis length: 1.0 – 2.0 mm; width: about 0.5 mm (Cook et al. 2020).Shape
- Caryopsis narrowly spindle-shaped; slightly three-sided in cross-section.
Surface Texture
- Caryopsis surface smooth.
Colour
- Caryopsis glossy amber to reddish-brown coloured.
Other Features
- Hilum nearly round, located near base of the caryopsis.
- The caryopsis will easily separate from the lemma and palea in some cultivars (Cook et al. 2020).
Chloris gayana (rhodesgrass) caryopsis in embryo view. Scale in mm.
-
Embryo
Size
- Embryo length*: 0.7 – 1.3 mm (average 1.1 mm).
- Embryo approximately three-fifths the length of the caryopsis, scutellum margin and root-shoot axis protruding beyond the main body of the endosperm.
*Note: minimum and maximum based on a random selection of 10 embryos in normal range of this species using image measurement protocol (ISMA 2020). CDA-S-22525
Shape
- Embryo narrowly teardrop shaped.
Endosperm
- Endosperm solid (Terrell 1971).
Other Features
- Embryo in lateral position (Martin 1942).
Chloris gayana (rhodesgrass) caryopsis in embryo view. Scale in mm.
Identification Tips
CONSEILS POUR L’IDENTIFICATION
- In the genus Chloris, disarticulation of the spikelet usually occurs above the glumes and the dispersal unit consists of a group of attached florets.
- The basal floret is fertile, and the upper florets are usually staminate or sterile.
- The basal floret is laterally compressed, awned (1.5 – 37 mm; Barkworth et al. 2003), and usually has long hairs along the lemma margins, especially near the tip of the lemma.
- The second floret in the spikelet is usually smaller in size, often differently shaped than the lower floret, and is usually long awned.
- Distal florets are usually greatly reduced in size, differently shaped from either of the lower or second florets.
- The closely related genus Eustachys has similar dispersal units, however the lemmas are unawned or are short awned (up to 1.2 mm long) (Barkworth et al. 2003).
Chloris gayana (rhodesgrass) attached floret group consisting of one fertile floret and two sterile florets (lateral view) removed from spikelet glumes and exposing callus hairs of basal floret. Scale in mm.
Additional Botany Information
AUTRES RENSEIGNEMENTS BOTANIQUES
Flowers/Inflorescence
- Inflorescence of numerous finger-like branches, 8 – 20 cm long, originating from a common point at the tip of the stem (looks like a feather duster); spikelets laterally compressed, closely packed in two rows on one side of each branch axis (Barkworth et al. 2003).
Vegetative Features
- Stoloniferous, tufted perennial.
- Stems 0.5 – 3 m tall or more, rooting at lower nodes.
- Cauline leaf blades up to 50 cm long and 2 – 20 mm wide, glabrous or scabrous.
- Stolons up to 3 m long, stolon leaves two to four per node and shorter than cauline leaves.
Descriptions based on Gould (1951), Barkworth et al. (2003), Loch et al. (2004), DiTomaso and Healy (2007), Rojas-Sandoval (2020).
Similar Species
ESPÈCES SEMBLABLES
Similar species are based on a study of seed morphology of various species, and those with similar dispersal units are identified. The study is limited by physical specimen and literature availability at the time of examination, and possibly impacted by the subjectivity of the authors based on their knowledge and experience. Providing similar species information for seed identification is to make users aware of similarities that could possibly result in misidentification.
Chloris barbata Sw. swollen windmill grass (Barkworth et al. 2003; USDA-NRCS 2023)
Spikelet with one fertile floret and two or three sterile florets; basal floret (fertile) 2 – 2.7 mm long, slightly compressed laterally, D-shaped to oval, marginal hairs near tip of lemma up to 1 mm long, keel with or without hairs; awn 4 – 7.7 mm long, callus hairy around margin; second floret 0.9 – 1.3 mm long, 0.4 – 0.9 mm wide, inflated (not compressed), usually without hairs, somewhat truncate at tip, awn 5 – 7 mm long; third floret, strongly inflated and nearly spherical, smaller than the second floret, long awned; fourth floret (if present) similar to third but smaller; caryopsis 1.1 – 1.4 mm long. (Measurements from Barkworth et al. 2003). In C. gayana the second and third florets are not inflated, and the hairs at the tip of the basal floret and around the callus are shorter than in C. barbata.
Chloris verticillata Nutt. windmillgrass (AOSA 2003), tumble windmill grass (Barkworth et al. 2003; USDA-NRCS 2023)
Spikelet with one fertile floret and one sterile floret; basal floret (fertile) 2 – 3.5 mm long, 1.5 – 1.9 mm wide, oval to narrowly teardrop shaped, keel hairless or with short appressed hairs, lateral surfaces of lemma without conspicuous longitudinal groove, with or without short hairs along lemma margins, lemma tip pointed, awn 4.8 – 9 mm long, callus margin with short hairs; second floret (sterile) 1.1 – 2.3 mm long, oblong and slightly inflated, tip truncate, may be bilobed; awn 3.2 – 7 mm long; caryopsis 1.3 – 1.5 mm long, 0.5 mm wide, oval shaped. (Measurements from Barkworth et al. 2003). C. gayana has conspicuous longitudinal grooves between the lemma keel and margins of the basal floret, whereas C. verticillata does not.
Chloris virgata Sw. feather fingergrass (Barkworth et al. 2003; AOSA 2023; USDA-NRCS 2023), feather windmill grass (Barkworth et al. 2003)
Spikelet with one fertile and one or two sterile florets; basal floret (fertile) 2.5 – 4.2 mm long, 0.5 – 1.5 mm wide, keel prominently bulged outward (gibbose), keel with or without hairs, lateral surfaces of lemma without longitudinal groove, lemma margins smooth to hairy in lower two-thirds and with very long hairs in upper one-third, hairs 1.5 mm or longer, awned from below the lemma tip, awn 2.5 – 15 mm long, callus margin with short hairs; second floret 1.4 – 2.9 mm long, 0.4 – 0.8 mm wide, open near tip but not inflated, slightly bilobed, awned from below tip, awn 3 – 9.5 mm long; third floret highly reduced in size, shorter than its supporting rachilla segment, unawned; caryopsis 1.5 – 2 mm long, 0.5 mm wide, oval shaped. (Measurements from Barkworth et al. 2003; DiTomaso and Healy 2007). In C. gayana, the lemma keel of the basal floret is only slightly curved, whereas in C. virgata the keel is gibbose, and the hairs near the tip of the basal floret are much shorter than in C. virgata.
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Chloris barbata
Chloris verticillata
Chloris virgata
Comparison Window
Fenêtre de comparaison
MAIN SPECIES
ESPÈCES PRINCIPALES
Chloris gayana
Chloris gayana
Poaceae
Chloris gayana (rhodesgrass) spikelet consisting of attached floret group and upper (2nd) glume. Lower (1st) glume missing. Scale in mm.
MAIN SPECIES
ESPÈCES PRINCIPALES
Chloris gayana
Chloris gayana
Poaceae
Chloris gayana (rhodesgrass) attached floret group consisting of one fertile floret and two sterile florets (lateral view) removed from spikelet glumes and exposing callus hairs of basal floret. Scale in mm.
MAIN SPECIES
ESPÈCES PRINCIPALES
Chloris gayana
Chloris gayana
Poaceae
Chloris gayana (rhodesgrass) caryopsis in lateral view. Scale in mm.
MAIN SPECIES
ESPÈCES PRINCIPALES
Chloris gayana
Chloris gayana
Poaceae
Chloris gayana (rhodesgrass) caryopsis in hilum view. Scale in mm.
MAIN SPECIES
ESPÈCES PRINCIPALES
Chloris gayana
Chloris gayana
Poaceae
Chloris gayana (rhodesgrass) caryopsis in embryo view. Scale in mm.
SIMILAR SPECIES
ESPÈCES SEMBLABLES
Chloris barbata
Chloris barbata
Poaceae
Chloris barbata (swollen windmill grass) attached floret group in palea view of basal floret and lemma view of second floret. Scale in mm.
SIMILAR SPECIES
ESPÈCES SEMBLABLES
Chloris barbata
Chloris barbata
Poaceae
Chloris barbata (swollen windmill grass) attached floret group (three florets, one fertile and two sterile) in lateral view. Scale in mm.
SIMILAR SPECIES
ESPÈCES SEMBLABLES
Chloris barbata
Chloris barbata
Poaceae
Chloris barbata (swollen windmill grass) caryopsis in embryo view. Scale in mm.
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Reference(s)
Référence(s)
Association of Official Seed Analysts (AOSA). 2023. Rules for Testing Seeds, Vol. 3: Uniform Classification of Weed and Crop Seeds. Association of Official Seed Analysts, Wichita, KS, USA.
Baldwin, B. G., Goldman, D. H., Keil, D. J., Patterson, R., Rosatti, T. J., Wilken, D. H. (Eds.). 2012. The Jepson Manual: Vascular Plants of California. 2nd edition. University of California Press, Berkeley, CA.
Barkworth, M. E., Capels, K. M., Long, S., and Piep, M. B. 2003. Flora of North America, Vol. 25 Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford University Press.
Cook, B.G. Pengelly, B.C. Schultze-Kraft, R;. Taylor, M. Burkart, S. Cardoso Arango, J.A. González Guzmán, J.J. Cox, K. Jones, C., Peters, M. 2020. Tropical Forages: An Interactive Selection Tool. 2nd and Revised Ed. International Center for Tropical Agriculture (CIAT), Cali, Colombia and International Livestock Research Institute (ILRI), Nairobi, Kenya. https://www.tropicalforages.info/text/entities/chloris_gayana.htm Accessed January 1, 2024.
DiTomaso, J. M. and Healy, E. A. 2007. Weeds of California and Other Western States. Vol. 2: Geraniaceae – Zygophyllaceae. Agriculture and Natural Resources Publication 3488. University of California, Oakland, CA.
efloras. 2023. Chloris gayana Kunth. Flora of China. http://efloras.org/florataxon.aspx?flora_id=2&taxon_id=200025056 Accessed December 29, 2023.
Gould, F. W. 1981. Grasses of Southwestern United States. The University of Arizona Press, Tucson, AZ.
International Seed Morphology Association (ISMA). 2020. Method for seed size measurement. Version 1.0. ISMA Publication Guide. https://www.idseed.org/authors/details/method_for_seed_size_measurement.html.
Kentucky Equine Research (KER). 2020. Big Head Disease in Horses: Calcium Deficiency. Equinnews Nutrition and Health Daily. https://ker.com/equinews/big-head-disease-horses-calcium-deficiency/ Accessed December 31, 2023.
Loch, D.S., Rethman, N.F.G., Van Niekerk, W.A. (2004). Chapter 25 Rhodesgrass in L.E. Moser, B.L. Burson & L.E. Sollenberger (Editors) Warm-Season (C4) Grasses, Volume 45: pp. 833-872. https://doi.org/10.2134/agronmonogr45.c25.
Martin, A. C. 1946. The comparative internal morphology of seeds. The American Midland Naturalist 36(3):513-660.
Rojas-Sandoval, J. 2020. Chloris gayana (Rhodes grass). CABI Compendium. https://doi.org/10.1079/cabicompendium.13115.
Terrell, E. E. 1971. Survey of occurrences of liquid or soft endosperm in grass genera. Bull. Torr. Botan. Club 98(5):264-268.
United States Department of Agriculture-Agricultural Marketing Service (USDA-AMS). 2023. Federal Seed Act Regulations. https://www.ecfr.gov/current/title-7/subtitle-B/chapter-I/subchapter-K/part-201?toc=1 Accessed December 29, 2023.
United States Department of Agriculture-Agricultural Research Service-National Plant Germplasm System (USDA-ARS-NPGS). 2023. Germplasm Resources Information Network (GRIN Taxonomy). National Germplasm Resources Laboratory, Beltsville, Maryland.
https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomydetail?id=10260 Accessed December 29, 2023.
United States Department of Agriculture-Natural Resource Conservation Service (USDA-NRCS). 2023. The PLANTS National Plant Data Team, Greensboro, NC USA. Database http://plants.usda.gov Accessed December 29, 2023.
United States Department of Agriculture-Phytosanitary Certificate Issuance and Tracking System-Phytosanitary Export Database (USDA-PCIT-PExD). 2022. Phytosanitary import requirements of U.S.-origin commodities to foreign countries. https://pcit.aphis.usda.gov/PExD/faces/ViewPExD.jsf Accessed December 29, 2023.
Wiersema, J. H. and León, B. 1999. World Economic Plants: A Standard Reference. CRC Press, Boca Raton, FL.