Heliocarpus americanus (white moho)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Plant Type
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Latitude/Altitude Ranges
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Threatened Species
- Risk and Impact Factors
- Uses List
- Wood Products
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Heliocarpus americanus L. (1753)
Preferred Common Name
- white moho
Other Scientific Names
- Heliocarpus australis E. Watson
- Heliocarpus popayanensis Kunth
- Heliocarpus trichopodus Turcz.
International Common Names
- Spanish: calagua; caulote
Local Common Names
- Belize: broadleaf moho; high ridge moho; mountain moho
- Bolivia: balsa pancho; llausa mora; yarisa
- Colombia: balsa; burillo; burío
- Costa Rica: burillo; burío
- Ecuador: sanpan
- Mexico: janote; jonote
- Panama: majagüillo
- Peru: palo de balsa
- Venezuela: majagua
- HJCAM (Heliocarpus americanus)
Summary of InvasivenessTop of page
Heliocarpus americanus was introduced to the Hawaiian Islands in the early 20th century for use in reforestation projects. It has since escaped into the wild and is now present on several of the main islands. Although it has been reported as a pest, it has not been reported to have any direct adverse effects on native species, and its removal is not currently a conservation priority.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Spermatophyta
- Subphylum: Angiospermae
- Class: Dicotyledonae
- Order: Malvales
- Family: Tiliaceae
- Genus: Heliocarpus
- Species: Heliocarpus americanus
Notes on Taxonomy and NomenclatureTop of page
Heliocarpus americanus, known as “white moho”, is a species of tree in the family Malvaceae or Tiliaceae. (While many botanists have adopted the APG III system of classification for the orders and families of flowering plants which sinks the Tiliaceae into the Malvaceae (Angiosperm Phylogeny Group, 2009), the CAB Thesaurus continues to use the Cronquist system which treats them as separate families). Of the ten or so species in the genus (Robyns, 1964), H. americanus is the only one whose range extends beyond Mexico and Central America.
The nomenclature of this species is somewhat confused. The genus Heliocarpus was erected by Carl Linnaeus. Although, as E. E. Watson (1923) points out, the original description was in Linnaeus’ 1737 work Hortus Cliffortianus (as “Heliocarpos”), authorship is dated to its appearance in the 1753 Species Plantarum, as stated by Robyns (1964) and the International Plant Name Index (IPNI, 2012). Linnaeus described a single species of Heliocarpus, which he named “Heliocarpos americana” in Hortus Cliffortianus, but later revised to “Heliocarpus americana” in Species Plantarum and later works (and which was later emended to Heliocarpus americanus). Although Linnaeus gave the species’ native location simply as “crescit in America calidiori” (“grows in the warmer parts of the Americas”), Baker (1898) concluded that the specimens in question had come from Veracruz, Mexico, an opinion doubted by Watson (1923). Although there have been some differences in opinion, the Linnaean Plant Name Typification Project considers the material in the Clifford herbarium to be the type specimen, and this opinion is gaining acceptance.
In 1821, Carl Sigismund Kunth described a second species in the genus, Heliocarpus popayanensis, naming it after Popayán, Colombia, where the type specimen was collected, and differentiated that new species from Linnaeus’ species (Watson, 1923). A number of further species were described over the following decades, and by the time of Watson’s (1923) monograph, 22 species were recognised, including both H. americanus and H. popayanensis. Both of those two names have continued to be used for the widespread species, with authorship of H. americanus sometimes attributed to Watson rather than Linnaeus. H. americanus is now recognised as a widespread and phenotypically variable species, and H. popayanensis is often treated as a variety of this broader H. americanus, under the name Heliocarpus americanus var. popayanensis (Kunth) K. Schum.
DescriptionTop of page
Heliocarpus americanus is a medium to large tree, reaching 6–30 m tall (Robyns, 1964). Young stems and branches are densely covered with reddish hairs of various forms, but these are gradually lost, leaving older branches almost hairless (Watson, 1923; Lay, 1949; Robyns, 1964). The leaves of H. americanus show much greater variability in form than those of other species in the genus; in the central parts of its range, the leaves may have three distinct lobes, while towards the latitudinal limits of its range, the leaves become almost entirely undivided (Lay, 1949). The blade of the leaf is typically 16–20 cm long by 14–18 cm wide (Lay, 1949; Robyns, 1964), with pointed tips and a deeply cordate base. The petiole (leaf stalk) is usually 6–8 cm long (Lay, 1949; Robyns, 1964).
H. americanus is gynodioecious (that is, individuals are either female or hermaphroditic), producing flowers in terminal cymes, each cyme comprising 12–16 light green flowers (Lay, 1949; Robyns, 1964). The pistillate (exclusively female) inflorescences are typically larger than the hermaphroditic inflorescences, at 14 × 20 cm, compared to 12 × 14 cm in hermaphrodites (Lay, 1949; Robyns, 1964). The hermaphroditic flowers are around 10 mm in diameter, with four sepals, four petals and 12–16 stamens; the pistillate flowers have four sepals, no petals, and numerous staminodes in place of the stamens (Lay, 1949; Robyns, 1964).
The fruit of H. americanus, like that of other species in the genus, is a small seed, around 3–4 mm long by 2–3 mm wide, with a conspicuous fringe of plumose bristles, each 4–7 mm long, which form a halo around the seed. Linnaeus named the genus after this feature (helios = “sun”; carpus = “seed”), remarking “who could ever behold an almost rounded fruit, bordered with a halo of rays, without thinking of the sun as conceived by the painters?” (Lay, 1949). This distinguishes Heliocarpus from the closely related Triumfetta, in which the seed is entirely covered with bristles, forming a burr (Lay, 1949).
Plant TypeTop of page Broadleaved
DistributionTop of page
Heliocarpus americanus has a broad native distribution in the Neotropics, stretching from Mexico in the north, through Central America, and along the edges of the Andes to Argentina and northern parts of Paraguay in the south (Lay, 1949). It also extends across the mountains of northern South America, reaching as far east as Venezuela and Trinidad (Lay, 1949). It is typically found at elevations above 1000m – sometimes exceeding 2000m – in the tropical parts of its range, but descends to lower elevations in more temperate regions (Lay, 1949).
It has been introduced to Hawaii – see ‘History of Introduction and Spread’ section.
Distribution TableTop of page
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.Last updated: 10 Jan 2020
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Belize||Present||Native||USDA-ARS (2012); Smithsonian Tropical Research Institute (2013)|
|Costa Rica||Present, Widespread||Native||Lay (1949); USDA-ARS (2012); Smithsonian Tropical Research Institute (2013)|
|El Salvador||Present||Smithsonian Tropical Research Institute (2013)|
|Guatemala||Present||Smithsonian Tropical Research Institute (2013)|
|Honduras||Present||Smithsonian Tropical Research Institute (2013)|
|Martinique||Present||Smithsonian Tropical Research Institute (2013)|
|Mexico||Present, Localized||Native||Lay (1949); USDA-ARS (2012); Grandtner and Chevrette (2013); Smithsonian Tropical Research Institute (2013)||Central and south-eastern parts of the country|
|Nicaragua||Present||Smithsonian Tropical Research Institute (2013)|
|Panama||Present, Widespread||Native||Lay (1949); USDA-ARS (2012); Grandtner and Chevrette (2013); Smithsonian Tropical Research Institute (2013)|
|Trinidad and Tobago||Present, Widespread||Native||Lay (1949); Grandtner and Chevrette (2013)|
|United States||Present||CABI (Undated)||Present based on regional distribution.|
|-Hawaii||Present, Widespread||Introduced||1924||Invasive||Woodcock (2003); Grandtner and Chevrette (2013)||Found on at least four of the main islands|
|Argentina||Present, Localized||Native||USDA-ARS (2012); Lay (1949); Grandtner and Chevrette (2013)||In 6 of 24 provinces, all in the north: Catamarca, Jujuy, Misiones, Salta, Santiago del Estero and Tucuman|
|Bolivia||Present, Widespread||Native||Lay (1949); USDA-ARS (2012); Grandtner and Chevrette (2013); Smithsonian Tropical Research Institute (2013)|
|Brazil||Present, Localized||Native||Lay (1949); USDA-ARS (2012); Smithsonian Tropical Research Institute (2013)|
|-Acre||Present, Localized||Native||Lay (1949); Grandtner and Chevrette (2013)|
|-Minas Gerais||Present||Native||Grandtner and Chevrette (2013)|
|-Parana||Present||Native||Grandtner and Chevrette (2013)|
|-Rondonia||Present||Native||Grandtner and Chevrette (2013)|
|-Sao Paulo||Present||Native||Grandtner and Chevrette (2013)|
|Ecuador||Present, Localized||Native||In 12 of 24 provinces, including coastal, Andean and Amazon regions|
|Paraguay||Present, Localized||Native||only in the north of the country|
|Peru||Present, Localized||Native||USDA-ARS (2012); Lay (1949); Grandtner and Chevrette (2013); Smithsonian Tropical Research Institute (2013)||In 12 of 26 regions|
History of Introduction and SpreadTop of page
H. americanus was introduced by the Hawaiian Sugar Planters’ Association to the Hawaiian Islands before 1930 (Motooka et al., 2003), with the first recorded planting occurring in 1924 (Woodcock, 2003). It was introduced as an ornamental tree as part of a reforestation scheme, and was first recorded as naturalised in 1941 (Wester, 1992). In 1985, it was recorded from three of the high islands of Hawaii – Hawaii (Big Island), Oahu and Kauai – and to be a pest on both Hawaii and Oahu (Smith, 1985). In 2007, it was also discovered on the island of Lanai (Oppenheimer, 2007).
The recent, and presumably inadvertent, spread of H. americanus to the island of Lanai indicates that the species has a significant dispersal ability, even over relatively large distances (Lanai is several miles from either Maui or Molokai). This is likely to be through wind-dispersal of the seeds (Smith, 1985), which are relatively small and bear a ring of long bristles (see ‘Description’ section).
IntroductionsTop of page
Risk of IntroductionTop of page
Given its anemochory (wind dispersal), and history of spread even between islands, it is likely that H. americanus will continue to spread over suitable habitat throughout the high islands of Hawaii. It is unlikely to spread over longer distances unless it is deliberately introduced.
HabitatTop of page
Heliocarpus americanus grows natively on steep slopes, either in cloud forest or rain forest, or along streams at the edge of forestry. It can also be a pioneer tree in more open areas (Gargiullo et al., 2008). It is found at elevations between 1000 m and 2000 m in the core of its range, but occurs at lower elevations towards its latitudinal extremes; in Argentina, it is generally found below 1000 m (Lay, 1949). In Hawaii, it is found at low to mid elevations, in disturbed parts of wetter mesic forests (Motooka et al., 2003). On the island of Hawaii (Big Island), it is found in native forest (Imada et al., 2007), while on Lanai, it has only been discovered in alien forest (Oppenheimer, 2007).
Habitat ListTop of page
|Terrestrial – Managed||Disturbed areas||Present, no further details||Natural|
|Terrestrial ‑ Natural / Semi-natural||Natural forests||Present, no further details||Natural|
|Riverbanks||Present, no further details||Natural|
Biology and EcologyTop of page
The flowering time of H. americanus varies across its range. In northern parts, it flowers in December and January and retains its fruit into mid-March; further south, it flowers from May to June, and the fruit last until the end of September (Lay, 1949).
ClimateTop of page
|Af - Tropical rainforest climate||Preferred||> 60mm precipitation per month|
Latitude/Altitude RangesTop of page
|Latitude North (°N)||Latitude South (°S)||Altitude Lower (m)||Altitude Upper (m)|
Means of Movement and DispersalTop of page
The major vector for transporting H. americanus over large distances has been mankind. The planting of H. americanus in the Hawaiian Islands was abundant and widespread. More recently, however, the species has demonstrated the ability to disperse itself, both in escaping from plantations, and in colonising new islands. It is known to be among the earlier colonisers of disturbed ground in parts of its native range (Grau et al., 1997; Gargiullo et al., 2008), and shows similar characteristics in Hawaii, outside its native range (Motooka et al., 2003). This dispersal ability is largely attributable to the seeds, which are well adapted for wind-dispersal (anemochory), through the ring of large bristles that project around the edge of the seeds (Smith, 1985).
Pathway CausesTop of page
ImpactTop of page
Although H. americanus has become naturalised outside its native range and is spreading in its introduced range, no significant impacts of its presence or its spread have been reported. It is considered a “problem invader” – one of the most significant woody alien species in the Hawaiian islands – but no details are available of the likely effects of its continued spread.
Threatened SpeciesTop of page
|Threatened Species||Conservation Status||Where Threatened||Mechanism||References||Notes|
|Phyllostegia mollis (Waianae Range phyllostegia)||USA ESA listing as endangered species||Hawaii||Competition (unspecified)||US Fish and Wildlife Service, 2009|
|Schiedea hookeri (sprawling schiedea)||CR (IUCN red list: Critically endangered); USA ESA listing as endangered species||Hawaii||Competition - monopolizing resources; Ecosystem change / habitat alteration||US Fish and Wildlife Service, 2011a|
|Urera kaalae||CR (IUCN red list: Critically endangered); USA ESA listing as endangered species||Hawaii||Ecosystem change / habitat alteration; Pest and disease transmission||US Fish and Wildlife Service, 2011b|
Risk and Impact FactorsTop of page Invasiveness
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Highly adaptable to different environments
- Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
- Pioneering in disturbed areas
- Tolerant of shade
- Highly mobile locally
- Long lived
- Fast growing
- Has high reproductive potential
- Ecosystem change/ habitat alteration
- Competition - monopolizing resources
- Competition (unspecified)
- Pest and disease transmission
- Rapid growth
- Highly likely to be transported internationally deliberately
UsesTop of page
Heliocarpus americanus has several small-scale economic uses. Its timber is similar to that of balsa (Ochroma pyramidale, Malvaceae–Bombacoideae), and is used for construction, in papermaking, as firewood, and for floats and bottle-stoppers, as well as being a substitute for balsa in modelling (Lay, 1949; Keyes-Hennin et al., 2000). The bark of various Heliocarpus species is used to produce a fibre, which is used for making ropes and baskets (Lay, 1949; Keyes-Hennin et al., 2000). Bark decoctions have been used against sickness in cattle and on sores (Lay, 1949).
In their native range, Heliocarpus species including H. americanus also provide ecological services such as hosting epiphytes, including orchids, or aiding in honey production or in the cultivation of mushrooms or larvae (Keyes-Hennin et al., 2000). Its rapid growth and ecological versatility make H. americanus a useful tree for reforestation (Keyes-Hennin et al., 2000), but also make it a potentially damaging invasive species (Motooka et al., 2003).
Uses ListTop of page
- Source of medicine/pharmaceutical
Wood ProductsTop of page
Similarities to Other Species/ConditionsTop of page
H. americanus is the only species of Heliocarpus in most of its native range, and in its invasive range. The distinctive form of the fruit in this genus makes it unlikely to be confused with any other taxon except for pre-reproductive individuals.
Prevention and ControlTop of page
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.
There are no reports of management plans for controlling H. americanus outside its native range (Motooka et al., 2003). Seedlings are occasionally uprooted on sight, but not as part of any systematic eradication programme (Imada et al., 2007). It might be possible to eradicate the species from Hawaii, but since it is already widespread and poses little obvious threat to the existing biota, conservation efforts are focussed initially on removing other species instead (Imada et al., 2007). No biological control agents against H. americanus have been suggested (Smith, 1985); its foliage appears to be unpalatable to axis deer (Axis axis) (Oppenheimer, 2007).
Gaps in Knowledge/Research NeedsTop of page
Despite H. americanus being generally considered a major invasive tree species in the Hawaiian Islands, there has been very little research into its impacts on the natural ecosystems there. Its response to fire is unknown, and there have been no investigations into biological control (Smith, 1985).
ReferencesTop of page
Angiosperm Phylogeny Group, 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society, 161(2):105-121. http://www.blackwell-synergy.com/loi/boj
Baker EG, 1898. Two old American types. Journal of Botany, 36(424):129-132.
Gargiullo M, Magnuson B, Kimball L, 2008. A field guide to plants of Costa Rica. New York, NY, USA: Oxford University Press, 542 pp.
Grandtner MM, Chevrette J, 2013. Dictionary of Trees, Volume 2: South America. Nomenclature, taxonomy and ecology. Academic Press, 1176 pp.
Grau HR, Arturi MF, Brown AD, Acenolaza PG, 1997. Floristic and structural patterns along a chronosequence of secondary forest succession in Argentinian subtropical montane forests. Forest Ecology and Management, 95(2):161-171; 40 ref.
Imada C, Lau A, Frohlich D, Kennedy B, 2007. Botanical inventory of Board of Water Supply lands, Waihee and Kahaluu valleys, windward Oahu. Honolulu, HI, USA: Board of Water Supply, 37 pp.
IPNI, 2012. International Plant Names Index. Kew\Cambridge, MA\Canberra, UK\USA\Australia. Royal Botanic Gardens\The Harvard University Herbaria\Australian National Herbarium. http://www.ipni.org/
Keyes-Hennin MR, Morales-Ríos A, Maciel Gómez A, 2000. The white moho (Heliocarpus sp.) a tree with multiple uses from the eastern zone of the Gulf of Mexico, Mexico. (El Jonote (Heliocarpus sp.), árbol de usos múltiples de la zona oriental del Golfo de México, México.) Brenesia, 54:37-50.
Motooka P, Castro L, Nelson D, Nagai G, Ching L, 2003. Weeds of Hawaii's Pastures and Natural Areas; an identification and management guide. Manoa, Hawaii, USA: College of Tropical Agriculture and Human Resources, University of Hawaii.
Oppenheimer H, 2007. New plant records from Molokai, Lanai, Maui, and Hawaii for 2006. Bishop Museum Occasional Papers, 96:17-34.
Sherley G, 2000. Invasive species in the Pacific: a technical review and draft regional strategy. Apia, Samoa: South Pacific Regional Environment Programme, vi + 190 pp.
Smith CW, 1985. Impact of alien plants on Hawaii's native biota. Honolulu, HI, USA: University of Hawaii. In: Hawaii's terrestrial ecosystems: preservation and management [ed. by Stone, C. P. \Scott, J. M.]. Honolulu, HI, USA: University of Hawaii, 180-250.
Smithsonian Tropical Research Institute, 2013. STRI Herbarium. Balboa, Panama: Smithsonian Tropical Research Institute. http://biogeodb.stri.si.edu/herbarium/
USDA-ARS, 2012. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx
Watson EE, 1923. The genus Heliocarpus. Bulletin of the Torrey Botanical Club, 50(3):109-128.
Wester L, 1992. Origin and distribution of adventive alien flowering plants in Hawaii. Honolulu, HI, USA: University of Hawaii, pp. In: Alien plant invasions in native ecosystems of Hawaii: management and research [ed. by Stone, C. P. \Smith, C. W. \Tunison, J. T.]. Honolulu, HI, USA: University of Hawaii, 99-154.
CABI, Undated. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI
CABI, Undated a. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI
Grandtner MM, Chevrette J, 2013. Dictionary of Trees: South America. Nomenclature, taxonomy and ecology., 2 Academic Press. 1176 pp.
Smithsonian Tropical Research Institute, 2013. STRI Herbarium., Balboa, Panama: Smithsonian Tropical Research Institute. http://biogeodb.stri.si.edu/herbarium/
USDA-ARS, 2012. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysimple.aspx
OrganizationsTop of page
USA: Conservation Council for Hawai'i (CCH), P.O. Box 2923, Honolulu, HI96802, Hawaii, http://www.conservehi.org/
USA: Hawai'i Conservation Alliance (HCA), 677 Ala Moana Blvd, Suite 320, Honolulu, HI96813, Hawaii, http://hawaiiconservation.org/
USA: Hawaiian Ecosystems at Risk project (HEAR), P.O. Box 1272, Puunene, HI96784, Hawaii, http://www.hear.org/
ContributorsTop of page
12/10/12: Original text by:
Christopher J. Dixon, University of Oxford, Department of Plant Sciences, South Parks Road, Oxford, OX1 3RB, U.K.
Distribution MapsTop of page
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