Lythrum maritimum (pukamole)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Plant Type
- Distribution Table
- History of Introduction and Spread
- Habitat List
- Hosts/Species Affected
- Biology and Ecology
- Latitude/Altitude Ranges
- Soil Tolerances
- Notes on Natural Enemies
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Environmental Impact
- Threatened Species
- Risk and Impact Factors
- Uses List
- Detection and Inspection
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Lythrum maritimum Kunth
Preferred Common Name
Other Scientific Names
- Lythrum albicaule Bertero
- Lythrum album Kunth
- Lythrum campestre Griseb.
Local Common Names
- USA/Hawaii: ninika
Summary of InvasivenessTop of page
L. maritimum is a perennial shrub or subshrub, known to be native to South America. It is widespread in the Hawaiian Islands, where it has only relatively recently been classed as a noxious weed, being a threat to native plant species and habitats. Although some authorities still treat it as a Hawaiian endemic, the status of L. maritimum as native was only questioned and changed to naturalized in the 1990s. Even in 2010, in a survey of rare plants in Oahu Forest National Wildlife Refuge, there was uncertainty whether L. maritimum should be classed as naturalized or endemic, and no weed risk assessment of the species was given (Imada et al., 2011). This confusion over its status, along with the enormity of the problem of introduced alien species in Hawaii, appears to explain why so little research has been undertaken on its invasiveness and control.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Spermatophyta
- Subphylum: Angiospermae
- Class: Dicotyledonae
- Order: Myrtales
- Family: Lythraceae
- Genus: Lythrum
- Species: Lythrum maritimum
Notes on Taxonomy and NomenclatureTop of page
The genus Lythrum, which comprises 25-36 species of herbaceous annuals and perennials, is probably best known for its introduced and invasive members, including the noxious L. salicaria (purple loosestrife) and, in Hawaii, L. maritimum.
The name Lythrum maritimum was validly published by K. S. Kunth in 1823 in Nova Genera et Species Plantarum 6: 194 (although many authorities have 1824 as the date of publication, see http://www.biodiversitylibrary.org/item/11238#page/200/mode/1up for a copy of the original publication).
There are no infraspecific taxa. Accepted synonyms for L. maritimum are L. albicaule, L. album and L. campestre (The Plant List, 2013). However, significant differences between L. maritimum and L. album have been noted by some authors; chromosome numbers are given as n = 20 for L. maritimum and n = 10 for L. album (Graham and Cavalcanti, 2001), and Fuentes et al. (2013) listed both species in their database of alien species in Chile, with L.maritimum as a perennial shrub and L. album as an annual herb.
In Hawaii, where L. maritimum is most widespread and problematic, its common names include pukamole and ninika.
DescriptionTop of page
From Hillebrand (1888), Wagner (1990) and Wagner (1999):
L. maritimum is a low prostrate perennial, herbaceous but often with a woody base, growing up to 30-45 cm long. It is glabrous and many-branched. Leaves are linear-oblong or lanceolate, 7-33 mm long, on very short petioles, acute, obtuse at the base. Flowers are single, on short bibracteolate peduncles 25-38 mm long. The calyx tube is 76 mm long, with 12 ribs and 6 membranous deltoid teeth, the 6 accessory teeth being more than twice as long as these and stiff subulate. Flowers have 6 purplish or pink, obovate petals about 4 mm long. The 6 stamens are enclosed, or nearly so, and the style is exserted, with a thick globose stigma. The capsule is about three-quarters the length of the calyx, 2-celled, with numerous, minute, obconical, angular seeds, 0.5 mm long. The bracts are subulate.
Plant TypeTop of page Herbaceous
DistributionTop of page
Most Lythrum species are distributed in the Northern Hemisphere, but L. maritimum has a native range extending above and below the Equator. It is widespread in South America, where it is largely considered a native species. Tropicos lists L. maritimum as present in Argentina, Bolivia, Chile, Colombia, Peru, Uruguay and Venezuela (Missouri Botanical Garden, 2015). It is also native to Brazil (JSTOR Global Plants, 2015). Pensiero et al. (2005) recorded L. maritimum and classed it as native in the Castellanos and Vera departments of Santa Fe province in northeast Argentina.
The first record of L. maritimum in Chile is from 1974 (Ugarte et al., 2011). It is treated as naturalized rather than native there. Although L. maritimum and the synonymous L. album are both listed in Chile, both are classed as non-invasive alien species. L. maritimum is reported as a perennial shrub, whereas L. album as an annual herb (Fuentes et al., 2013).
L. maritimum is found on the Hawaiian islands of Kauai, Oahu, Molokai, Lanai, Maui and Hawaii (Wagner et al., 1990; Wester, 1992). US collecting expeditions encountered the species on the coast of Oahu, near Honolulu, and at Waimea on Hawaii (Gray, 1854).
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|
|Mexico||Present, Localized||Native||Grisebach (1875); Hemsley (1888)||In Veracruz state|
|United States||Present||CABI (Undated)||Present based on regional distribution.|
|-California||Present||Native||JSTOR Global Plants (2015)|
|-Hawaii||Present, Widespread||Introduced||Invasive||Wagner et al. (1990); Wester (1992); Wagner et al. (1999)||On all of the main Hawaiian Islands except Niihau and Kahoolawe|
|Argentina||Present||Native||Pensiero et al. (2005); Missouri Botanical Garden (2015)||Reported in Santa Fe province|
|Bolivia||Present||Native||Ritter (2004); Missouri Botanical Garden (2015)|
|Brazil||Present||Native||JSTOR Global Plants (2015)|
|Chile||Present||Introduced||Ugarte et al. (2011); Fuentes et al. (2013); Missouri Botanical Garden (2015)|
|Colombia||Present||Native||Missouri Botanical Garden (2015)|
|Peru||Present, Widespread||Native||Gray (1854); Missouri Botanical Garden (2015)|
|Uruguay||Present||Native||Missouri Botanical Garden (2015)|
|Venezuela||Present||Native||Missouri Botanical Garden (2015)|
History of Introduction and SpreadTop of page
L. maritimum was first recorded in Hawaii in 1794, suggesting it probably arrived in the islands much earlier and probably accidentally. Based on its several Hawaiian names and use in traditional medicine, it was for a long time regarded as an indigenous species (Bohm, 2012). Its native status was questioned by Wagner et al. (1990) in the Manual of the Flowering Plants of Hawaii, the recognised standard work on the flora of Hawaii. By the mid-1990s, it was becoming accepted that L. maritimum was a naturalized alien (Wagner et al., 2012). It is now listed by the US federal government as a noxious weed and invasive in Hawaii (USDA–NRCS, 2015).
HabitatTop of page
All members of Lythrum are facultative wetland indicator species, so it is unusual that L. maritimum in its native range is associated with native vegetation of the deserts along South America's western coast (5-30ºS), where it is largely restricted to fog-zone locations or lomas (small hill) formations, separated by hyper-arid habitat where virtually no plants exist (Dillon et al., 2011). However, these desert areas do have wet soils around springs and are subject to coastal fogs (Rundel et al., 1996). There are also brackish marshes, such as around the Reñaca estuary in central Chile (San Martín, 2001).
L. maritimum is also found in wetter habitats in its native range. Ritter (2004) listed it as a wetland species in its native Bolivia. L. maritimum was encountered in riverbed habitat in Callao and Lima in Peru in the course of US expeditions from 1838 to 1842 (Gray, 1854). Grisebach (1875) found L. maritimum in Veracruz in southeast Mexico, associated with coastal dunes interrupted by lagoons, whose waters were fringed or populated with a number of species including L. maritimum.
In Hawaii, where L. maritimum is adventive, it is associated with a range of habitats. It can be found in mesic, open, disturbed habitats, especially in pastures, on windward coastal cliffs, in margins of wet forest, and on igneous formations, from sea level up to 2450 m elevation, on all of the main Hawaiian Islands except Niihau and Kahoolawe (Wagner et al., 1999). It has been observed colonizing a lava field 9 years after the eruption of Mt. Kilauea in 1959 (Sauer, 1988), and it has been found along a pathway on the summit of Mt. Haleakala on Maui (Starr and Starr, 2005).
The wet cliffs along the windward sides of the Hawaiian Islands, which are fluted, basalt cliffs up to 1000 m high, with almost no soil, provide crevices in which L. maritimum grows (Mueller-Dombois and Fosberg, 1998). L. maritimum has been recorded growing on the Pohakuao cliffs of Kaua'i at elevations of 365-460 m (Wood et al., 2002).
L. maritimum can also be found in grasslands. With the eradication in 1987 of feral pigs in Hawaii Volcanoes National Park on the eastern flank of Mauna Loa (1400-1950 m elevation), vegetation recovery was monitored in mountain parkland ecosystems, especially grasslands. Over the period 1985-92, L. maritimum increased its percentage cover in the aalii (Dodonaea viscosa)/velvet grass (Holcus lanatus) community from 0% in 1985 to 0.3, 0.4, 2.2 and 3.2% in 1986, 1987, 1989 and 1992, respectively. It was not present at all in the Deschampsianubigena/velvet grass community nor sweet vernal grass (Anthoxanthum odoratum) community, and showed very low or no presence, depending on year, in the pukiawe (Styphelia tameiameiae)/Deschampsia and velvet grass/Deschampsia communities (Tunison et al., 1994).
Habitat ListTop of page
|Terrestrial – Managed||Rail / roadsides||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Terrestrial ‑ Natural / Semi-natural||Natural forests||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Natural grasslands||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Rocky areas / lava flows||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Scrub / shrublands||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Coastal areas||Principal habitat||Natural|
|Coastal dunes||Principal habitat||Natural|
|Salt marshes||Principal habitat||Natural|
|Rivers / streams||Principal habitat||Natural|
Hosts/Species AffectedTop of page
On the dry cliffs of the Hawaiian islands Lanai and Maui, the endangered native species Bidens campylotheca subsp. pentamera, Phyllostegia haliakalae and Pleomele fernaldii [Chrysodracon fernaldii] are threatened by the presence of L. maritimum and other understorey and subcanopy species (US Fish and Wildlife Service, 2012a). L. maritimum is one of the alien species threatening the sole surviving population of Dubautia plantaginea subsp. humilis on the wet cliffs of Maui (US Fish and Wildlife Service, 2012b). On Hawaii, L. maritimum has invaded the habitat of the critically endangered Clermontia pyrularia (State of Hawaii, Division of Forestry and Wildlife, 2015).
Biology and EcologyTop of page
The chromosome number is n = 20, although n = 10 has been recorded for the synonymous L. album (Graham and Cavalcanti, 2001).
L. maritimum is pollinated by insects and is a cross fertilizing species with hermaphrodite flowers.
Physiology and Phenology
Seeds exhibit physiological dormancy, and have been found to require greenhouse temperatures for germination (Lilleeng-Rosenberger, 2005).
L. maritimum is associated with a number of plant communities on the Hawaiian Islands, including Dodonaea viscosa/Holcus lanatus grasslands (Tunison et al., 1994). It is eaten by introduced alien pheasants in Hawaii (Schwartz and Schwartz, 1951).
ClimateTop of page
|A - Tropical/Megathermal climate||Preferred||Average temp. of coolest month > 18°C, > 1500mm precipitation annually|
|Af - Tropical rainforest climate||Preferred||> 60mm precipitation per month|
|BW - Desert climate||Preferred||< 430mm annual precipitation|
Latitude/Altitude RangesTop of page
|Latitude North (°N)||Latitude South (°S)||Altitude Lower (m)||Altitude Upper (m)|
Soil TolerancesTop of page
Special soil tolerances
Notes on Natural EnemiesTop of page
Pathway CausesTop of page
Pathway VectorsTop of page
Impact SummaryTop of page
Environmental ImpactTop of page
L. maritimum has been reported in numerous habitats in the Hawaiian Islands, including national parks, where it is displacing native vegetation. In a survey in 1987 it was found in Hakalau Forest National Wildlife Refuge on Hawaii (Stone et al., 1991). It is also known from Volcanoes National Park on Hawaii and Haleakala National Park on Maui (Johnson, 1999).
Of particular concern are the effects of invasion by L. maritimum and other exotic plant species on cliff habitats, and the consequent displacement of very specialized endemic plant taxa. For example, on the dry cliffs of Lanai and Maui the endangered native species Bidens campylotheca subsp. pentamera, Phyllostegia haliakalae and Pleomele fernaldii [Chrysodracon fernaldii] are threatened by the presence of L. maritimum and other understorey and subcanopy species, such as Ageratina adenophora, Hypochoeris radicata, Lapsana communis, Prunella vulgaris and Rubus spp. Non-native grasses that also threaten this ecosystem include Andropogon virginicus, Anthoxanthum odoratum, Dactylis glomerata and Holcus lanatus. These non-native plant species pose serious and ongoing threats to all three of these native species that depend on this ecosystem (US Fish and Wildlife Service, 2012).
Similarly, one of the major threats to the Hawaiian endemic Dubautia plantaginea subsp. humilis, a short-lived perennial shrub found on wet, barren, wind-blown cliffs between 350 and 400 m altitude in the Iao Valley on western Maui, is displacement by introduced invasive plant species, including L. maritimum (US Fish and Wildlife Service, 2012).
Threatened SpeciesTop of page
|Threatened Species||Conservation Status||Where Threatened||Mechanism||References||Notes|
|Bidens campylotheca subsp. pentamera (ko`oko`olau)||CR (IUCN red list: Critically endangered); USA ESA listing as threatened species||Hawaii||Competition - monopolizing resources||US Fish and Wildlife Service, 2012a|
|Chrysodracon fernaldii (hala pepe)||EN (IUCN red list: Endangered); USA ESA listing as endangered species||Hawaii||Competition - monopolizing resources||US Fish and Wildlife Service, 2012a|
|Clermontia pyrularia||CR (IUCN red list: Critically endangered)||Hawaii||Competition - monopolizing resources||Center for Plant Conservation, 2015; State and of Hawaii, Division of Forestry and Wildlife, 2015|
|Dubautia plantaginea subsp. humilis||National list(s)||Hawaii||Competition - monopolizing resources||US Fish and Wildlife Service, 2012a|
|Peristylus holochila (Hawai'i bog orchid)||CR (IUCN red list: Critically endangered); USA ESA listing as endangered species||Hawaii||Competition - monopolizing resources||US Fish and Wildlife Service, 2000|
|Phyllostegia haliakalae (Lanai phyllostegia)||CR (IUCN red list: Critically endangered); USA ESA listing as endangered species||Hawaii||Competition - monopolizing resources||US Fish and Wildlife Service, 2012a|
Risk and Impact FactorsTop of page Invasiveness
- Proved invasive outside its native range
- Has a broad 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
- Damaged ecosystem services
- Ecosystem change/ habitat alteration
- Reduced native biodiversity
- Threat to/ loss of endangered species
- Threat to/ loss of native species
- Competition - monopolizing resources
- Rapid growth
- Difficult/costly to control
UsesTop of page
L. maritimum has been used in traditional Hawaiian medicine as a cure for asthma. A mash is made from the bark of taproots mixed with plant parts from Sida fallax and ko kea (white-rinded sugarcane), and the liquid extracted from this is taken as a tonic (Krauss, 2001).
Uses ListTop of page
Detection and InspectionTop of page
Species within the genus Lythrum exhibit considerable variation, and there are few readily apparent differences among species.
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.
Manual weeding of L. maritimum and other invasive species has been successfully applied to the remaining population of Dubautia plantaginea subsp. humilis on wet cliffs in the Iao Valley on Maui (US Fish and Wildlife Service, 2012).
Case study: Volcanoes National Park, Hawaii
Although no specific control measures and actions have been reported for L. maritimum, it would have been included in general and affected by the following strategies and actions proposed and undertaken in Volcanoes National Park, Hawaii, in the 1990s: (1) controlling feral pigs and goats; (2) excluding fire; (3) controlling localized alien plants; (4) controlling all disruptive alien plants in special ecological areas (the most diverse and intact areas in the park); (5) confining one widespread species, fountain grass (Pennisetum setaceum), to the area it currently infests; (6) developing herbicidal control methods for target species; (7) developing biological controls for some widespread species; (8) mapping the distribution of important alien plants; (9) researching the ecology, seed biology and phenology of important alien plant pests; (10) educating the public to the importance of alien plant control, and (11) working with other agencies and groups in alien plant management. Control of new introductions and incipient infestations, and protection of some of the most diverse and intact areas in the park have been very successful strategies (Stone et al., 1992).
Gaps in Knowledge/Research NeedsTop of page
There seems little if any research into the biology, ecology, physiology, phenology, reproduction and dispersal of L. maritimum. It is seen as just one of many alien species in Hawaii when it comes to developing and assessing control measures, and no specific interventions have been reported for the species.
ReferencesTop of page
Anderson SJ; Stone CP; Higashino PK, 1992. Distribution and spread of alien plants in Kipahulu Valley, Haleakala National Park, above 2,300 ft elevation. 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 Cooperative National Park Resources Studies Unit, 300-338. http://manoa.hawaii.edu/hpicesu/book/1992_chap/14.pdf
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Gray A, 1854. Botany Phanerogamia., XV Philadelphia, USA: Sherman. 606.
Grisebach AHR, 1875. The vegetation of the Earth after its arrangement according to climate, Volume 2. (La végétation du globe d'après sa disposition suivant les climats, Tome 2)., Paris, France: Guérin. 488-489.
Hemsley WB, 1888. Biologia Centrali-Americana., I London, UK: R. H. Porter. 448.
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Pensiero JF, Gutiérrez HF, Luchetti AM, Exner E, Kern V, Brnich E, Oakley L, Prado D, Lewis JP, 2005. Vascular flora of the province of Santa Fe. (Flora vascular de la provincia de Santa Fe)., Santa Fe, Argentina: Universidad Nacional del Litoral. 404 pp.
Ritter N, 2004. Checklist of dicotyledonous species associated with Bolivian wetlands., http://www.botanize.com/bol_checklist/bolspecies.htm
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24/02/15 Original text by:
Andrew Praciak, consultant, Ireland
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