Invasive Species Compendium

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Ardisia crenata
(coral berry)

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Datasheet

Ardisia crenata (coral berry)

Summary

  • Last modified
  • 13 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Host Plant
  • Preferred Scientific Name
  • Ardisia crenata
  • Preferred Common Name
  • coral berry
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • Ardisia crenata is an evergreen shrub, native to east and southeast Asia and parts of India, which grows as an understorey forest species. It has attractive red berries, which has led it to be widely used as an...

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Pictures

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PictureTitleCaptionCopyright
Ardisia crenata (coral berry or Hilo holly); flowering habit in scrub border. Mealani Ag Station, Waimea, Hawaii, USA. July, 2012.
TitleHabit in scrub border
CaptionArdisia crenata (coral berry or Hilo holly); flowering habit in scrub border. Mealani Ag Station, Waimea, Hawaii, USA. July, 2012.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Ardisia crenata (coral berry or Hilo holly); flowering habit in scrub border. Mealani Ag Station, Waimea, Hawaii, USA. July, 2012.
Habit in scrub borderArdisia crenata (coral berry or Hilo holly); flowering habit in scrub border. Mealani Ag Station, Waimea, Hawaii, USA. July, 2012.©Forest Starr & Kim Starr - CC BY 4.0
Ardisia crenata (coral berry or Hilo holly); flowering habit in scrub border. Mealani Ag Station, Waimea, Hawaii, USA. July, 2012.
TitleFlowering habit
CaptionArdisia crenata (coral berry or Hilo holly); flowering habit in scrub border. Mealani Ag Station, Waimea, Hawaii, USA. July, 2012.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Ardisia crenata (coral berry or Hilo holly); flowering habit in scrub border. Mealani Ag Station, Waimea, Hawaii, USA. July, 2012.
Flowering habitArdisia crenata (coral berry or Hilo holly); flowering habit in scrub border. Mealani Ag Station, Waimea, Hawaii, USA. July, 2012.©Forest Starr & Kim Starr - CC BY 4.0
Ardisia crenata (coral berry or Hilo holly); fruiting habit. Iao Tropical Gardens of Maui, Maui, Hawaii, USA. May, 2012.
TitleFruiting habit
CaptionArdisia crenata (coral berry or Hilo holly); fruiting habit. Iao Tropical Gardens of Maui, Maui, Hawaii, USA. May, 2012.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Ardisia crenata (coral berry or Hilo holly); fruiting habit. Iao Tropical Gardens of Maui, Maui, Hawaii, USA. May, 2012.
Fruiting habitArdisia crenata (coral berry or Hilo holly); fruiting habit. Iao Tropical Gardens of Maui, Maui, Hawaii, USA. May, 2012.©Forest Starr & Kim Starr - CC BY 4.0
Ardisia crenata (coral berry or Hilo holly); fruit and leaves. Iao Tropical Gardens of Maui, Maui, Hawaii, USA. May, 2012.
TitleFruit and leaves
CaptionArdisia crenata (coral berry or Hilo holly); fruit and leaves. Iao Tropical Gardens of Maui, Maui, Hawaii, USA. May, 2012.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Ardisia crenata (coral berry or Hilo holly); fruit and leaves. Iao Tropical Gardens of Maui, Maui, Hawaii, USA. May, 2012.
Fruit and leavesArdisia crenata (coral berry or Hilo holly); fruit and leaves. Iao Tropical Gardens of Maui, Maui, Hawaii, USA. May, 2012.©Forest Starr & Kim Starr - CC BY 4.0
Ardisia crenata (coral berry or Hilo holly); seedlings. Nuuanu, Oahu, Hawaii, USA. August, 2006.
TitleSeedlings
CaptionArdisia crenata (coral berry or Hilo holly); seedlings. Nuuanu, Oahu, Hawaii, USA. August, 2006.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Ardisia crenata (coral berry or Hilo holly); seedlings. Nuuanu, Oahu, Hawaii, USA. August, 2006.
SeedlingsArdisia crenata (coral berry or Hilo holly); seedlings. Nuuanu, Oahu, Hawaii, USA. August, 2006.©Forest Starr & Kim Starr - CC BY 4.0

Identity

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Preferred Scientific Name

  • Ardisia crenata Sims

Preferred Common Name

  • coral berry

Other Scientific Names

  • Ardisia bicolor Walker
  • Ardisia crenulata Lodd.
  • Ardisia crispa var. taquetii Lév
  • Ardisia densa Miq.
  • Ardisia elegans Andrews
  • Ardisia konishii Hayata
  • Ardisia kusukusensis Hayata
  • Ardisia labordei Lév
  • Ardisia lentiginosa Ker Gawl.
  • Ardisia linangensis Hu
  • Ardisia miaoliensis Lu
  • Ardisia mouretii Pit.
  • Bladhia crenata (Sims) Hara
  • Bladhia crispa var. taquetii (Lév) Nakai
  • Bladhia kusukusensis (Hayata) Nakai
  • Bladhia lentiginosa (Ker Gawl.) Nakai
  • Bladhia lindleyana (Dietr.) Nakai
  • Bladhia punctata (Lindl.) Nakai
  • Tinus densa (Miq.) Kuntze

International Common Names

  • English: Australian holly; Christmas berry; coral ardisia; coral berry tree; coral bush; hen's eyes; Hilo holly; scratchthroat; spiceberry
  • French: arbre à noël; arbre de noël; ardisie crénelée; baie corail; bois de noël
  • Chinese: zhu sha gen

Local Common Names

  • Germany: gewürzbeere; spitzenblume
  • Japan: manryo
  • South Africa: koraalbessieboom (Afrikaans)

Summary of Invasiveness

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Ardisia crenata is an evergreen shrub, native to east and southeast Asia and parts of India, which grows as an understorey forest species. It has attractive red berries, which has led it to be widely used as an ornamental plant in shady conditions. It has consequently been transported globally from its native range. Artificial selection for traits - such as enhanced berry production - through cultivation as an ornamental has increased its invasiveness (Kitajima et al., 2006), and it tends to displace native species. It has a high risk of spread once present, and is widely listed as an invasive species, for example in parts of the USA, a number of Pacific Islands, and Australia. In its native range it is also invasive and of environmental concern in the Philippines, Japan and Taiwan (Space et al., 2009), but in Singapore, it is listed as critically endangered (Chong et al., 2009).

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Primulales
  •                         Family: Primulaceae
  •                             Genus: Ardisia
  •                                 Species: Ardisia crenata

Notes on Taxonomy and Nomenclature

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Ardisia crenata was named in 1817 by John Sims. There are 30 synonyms listed for this species, of which four are different forms, two are subspecies, and seven are varieties. A. crenata is the accepted basionym for Bladhia crenata, and it has one homonym named by William Roxburgh in 1824 (Missouri Botanical Garden, 2013; The Plant List, 2013).

In China A. crenata is known as zhu-sha-gen, which means ‘red sand roots’ (Kitajima et al., 2006). The popular common names of ‘coral’ or ‘Christmas’ berry are a result of its attractive red drupe fruit, and shade tolerance in gardens (Niu et al., 2012).

Description

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A. crenata is an evergreen subshrub growing up to 1.8 m (6 ft) tall, but more commonly between 0.5-1 m in height, growing in multi-stemmed clumps. Leaves are alternate, <21 cm long, dark green above and waxy, glabrous, with crenate (scalloped) margins and calluses (bacterial nodules) in the margin notches. Flowers are white to pink in stalked axillary clusters, usually drooping below the foliage. Flowers are small and bisexual, with petaloid parts pink-white, and yellow anthers (Langeland and Burks, 1998). Wagner et al. (1999), cited in PIER (2013), say: 'inflorescences simple and umbellate or cymose, principal rays 2-4 cm long, pedicels 0.5-1 cm long, elongating in fruit; sepals oblong-ovate to lanceolate, ca. 1.5 mm long, glabrous, sparsely glandular punctate; corolla lobes white or pinkish, ovate, glandular punctate'. The fruit is a dark red globose drupe, 5-8 mm in diameter, containing one seed (Chimera and Drake, 2010; Langeland and Burks, 1998).

Plant Type

Top of page Broadleaved
Perennial
Seed propagated
Shrub
Vegetatively propagated
Woody

Distribution

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The genus Ardisia comprises a large number of species, generally found in subtropical and tropical environments (Roh et al., 2005). A. crenata originates in Asia, specifically China, Japan, Korea, southeast Asia and parts of India (Niu et al., 2012; USDA-ARS, 2014).

It has been widely introduced, and in some countries it is invasive. In the USA it is listed as weedy or invasive in Florida and Hawai’i by the USDA (USDA-NRCS, 2013), and considered to be invasive by the Florida Exotic Pest Plant Council (Sellers et al., 2013) as well as being on the Florida noxious weed list (Center for Aquatic and Invasive Plants, 2013), with planting prohibited in Miami-Dade County, Florida (PIER, 2013). It is also listed as a Category 2 exotic plant that is a moderate problem in Georgia (Georgia Exotic Pest Plant Council, 2013), and is on ‘Watch List A: recently appearing in Alabama as free-living infestations’ (Alabama Invasive Plant Council, 2007). A risk assessment for A. crenata in Hawai’i produced a score of 10, which is a high risk value for invasive species (PIER, 2013).

A. crenata is also listed as a priority species for exclusion from Niue (Space et al., 2004), and as invasive and of environmental concern in the Cook Islands and Fiji (Space and Flynn, 2002a); within the Cook Islands it is recommended for eradication on Rarotonga, and should be subject to inter-island quarantine between the southern Cook Islands (Space and Flynn, 2002b).

In Australia A. crenata is listed an ‘emerging invader’ and is a category 1 species in the Conservation of Agricultural Resources Act, which means it is a prohibited weed that must be controlled in all situations (Nel et al., 2004); it has also been nominated as a ‘potential environmental weed’ in Australia (Csurhes and Edwards, 1998).

Distribution Table

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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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

CambodiaPresent2007Missouri Botanical Garden, 2013
ChinaPresentPresent based on regional distribution.
-AnhuiPresentNativeUSDA-ARS, 2013
-FujianPresentNativeUSDA-ARS, 2013
-GuangdongPresentNativeUSDA-ARS, 2013
-GuangxiPresentNativeUSDA-ARS, 2013
-HainanPresentNativeUSDA-ARS, 2013
-Hong KongPresent2003Missouri Botanical Garden, 2013
-HubeiPresentNativeUSDA-ARS, 2013
-HunanPresentNativeUSDA-ARS, 2013
-JiangsuPresentNativeUSDA-ARS, 2013
-JiangxiPresentNativeUSDA-ARS, 2013
-TibetPresentNativeUSDA-ARS, 2013
-YunnanPresentNativeUSDA-ARS, 2013
-ZhejiangPresentNativeUSDA-ARS, 2013
IndiaPresentNativeNiu et al., 2012; USDA-ARS, 2014In north according to one source; in south-west according to the other
JapanPresentNative Invasive Space et al., 2009; Niu et al., 2012
-HonshuPresentNativeUSDA-ARS, 2013Southern Honshu
-KyushuPresentNativeUSDA-ARS, 2013
-Ryukyu ArchipelagoPresentNativeUSDA-ARS, 2013
-ShikokuPresentNativeUSDA-ARS, 2013
Korea, DPRPresentNativeNiu et al., 2012; USDA-ARS, 2013
Korea, Republic ofPresentNativeNiu et al., 2012; USDA-ARS, 2013
MalaysiaPresentNativeUSDA-ARS, 2013
MyanmarPresentNativeUSDA-ARS, 2013
PhilippinesPresentNative Invasive Space et al., 2009; USDA-ARS, 2013
SingaporePresent, few occurrencesNativeChong et al., 2009Critically endangered
Sri LankaPresent1969Missouri Botanical Garden, 2013
TaiwanPresentNative Invasive Space et al., 2009; USDA-ARS, 2013
ThailandPresentNativeUSDA-ARS, 2013
VietnamPresentNativeUSDA-ARS, 2013

Africa

MadagascarPresentIntroduced Invasive Lorence and Sussman, 1986
MauritiusPresentIntroduced Invasive Lorence and Sussman, 1986Specifically recorded at Brise Fer
RéunionPresentIntroduced1856 Invasive Macdonald et al., 1991; Kueffer and Mauremootoo, 2004; Baret et al., 2006
SeychellesPresentIntroduced Invasive Weber, 2003; USDA-ARS, 2013
South AfricaPresentIntroduced Invasive Nel et al., 2004

North America

USAPresentPresent based on regional distribution.
-AlabamaPresentIntroduced Invasive Miller, 2003; Alabama Invasive Plant Council, 2007
-CaliforniaPresent1999Missouri Botanical Garden, 2013
-FloridaPresentIntroduced Invasive Miller, 2003; Clewell and Tobe, 2011; USDA-NRCS, 2013
-GeorgiaPresentIntroducedUSDA-NRCS, 2013
-HawaiiPresentIntroduced Invasive Space and Flynn, 2002a; Stone et al., 1992; Wagner et al., 1999; Staples et al., 2000; PIER, 2013; USDA-NRCS, 2013
-LouisianaPresentIntroduced Invasive Miller, 2003; USDA-NRCS, 2013
-MissouriPresent1972Missouri Botanical Garden, 2013
-TexasPresentIntroduced1997Niu et al., 2012

Central America and Caribbean

Puerto RicoPresentIntroducedAcevedo-Rodríguez and Strong, 2012

Oceania

AustraliaPresentIntroducedRandall, 2007Naturalized
-New South WalesLocalisedIntroduced Invasive Csurhes and Edwards, 1998Naturalized in remnant rainforest in north-east New South Wales
-QueenslandPresentIntroduced Invasive Csurhes and Edwards, 1998
Cook IslandsPresentIntroduced Invasive Space and Flynn, 2002b; PIER, 2013Rarotonga Island
FijiPresentIntroduced Invasive Space and Flynn, 2002a
New ZealandPresentIntroduced1996 Invasive Heenan et al., 2002

History of Introduction and Spread

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A. crenata has historically been spread through international trade from its native range - China, Japan, Korea, parts of India and southeast Asia - as an ornamental species. The fine-scale and detailed history of its introduction is unknown, but the source of populations in the USA is thought to be southeast China via the Korean market (Niu et al., 2012).

It was introduced to the USA (Florida) in the early 20th century as an ornamental, and was noted as escaping into moist woodland in 1982. It has been reported in Florida natural areas in Alachua, Flagler, Gadsden, Highlands, Hillsborough, Leon, Liberty, Marion, Martin, and Orange counties (Langeland and Burks, 1998). It has spread from Florida into other southern states, including to Texas in 1997 (Niu et al., 2012). It was first recorded in Hawai’i in 1930, brought in as an ornamental species. It is now present on four of the Hawaiian islands (PIER, 2013), and is listed as an important alien invasive plant species in Hawai’i Volcanoes National Park. Here, it is invasive in the lowlands, but is localised and controlled (Stone et al., 1992). There is high risk of spread once A. crenata is present, and in the USA, it is thought that 11% of the country is suitable for its establishment (USDA-APHIS, 2012).

A. crenata was introduced to Mauritius around 1800, and it is now one of the most problematic invasive woody plant species in upland forests (Kueffer and Mauremootoo, 2004). It was noted as an escapee into wet forests in Mauritius over 60 years ago (Langeland and Burks, 1998). In 1856 it was introduced to Réunion Island; the reason is unknown, but it was probably its popularity as an ornamental plant. It is now a widespread species in Réunion occurring in all habitats from lowlands to mountains, but particularly in lowland rainforest and leeward mountain rainforest (Baret et al., 2006; Kueffer and Lavergne, 2004).

New Zealand also has A. crenata populations, which were first recorded in 1996 (Heenan et al., 2002). In Australia it has become naturalised in remnant rainforest in NE New South Wales, and is also present in Queensland (Csurhes and Edwards, 1998).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Hawaii 1930 Ornamental purposes (pathway cause) Yes No Stone et al. (1992)
Mauritius 1800 Yes No Kueffer and Mauremootoo (2004) Deliberate introduction, probably as an ornamental species
Réunion 1856 Yes No Baret et al. (2006) Deliberate introduction, probably as an ornamental species
USA Japan 1900 Ornamental purposes (pathway cause) Yes No Kitajima et al. (2006)

Risk of Introduction

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Transportation by humans for ornamental purposes is the primary mechanism for establishment and subsequent invasion of A. crenata. Containing transmission by human endeavour is therefore the best way to restrict further introductions and establishment (Niu et al., 2012). Accidental introduction can also occur locally through incorrect disposal of garden waste (USDA-APHIS, 2012).

Focussing on the USA, USDA-APHIS (2013) state that A. crenata is a high-risk species for invasion in the USA with a 59.3% chance of being a major invader there (38.6% chance of being a minor invader, and 2.0% chance of being a non-invader). Hutchinson et al. (2011) also indicate that there is potential for A. crenata to spread throughout the southern states of the USA, from North Carolina to California, although this presumably does not apply to areas that would be too dry for it. Fox and Kitajima (2001), referring to Florida, comment that due to its intolerance of drier conditions, it is unlikely that it would invade habitats such as scrub or sand-hills.

A. crenata is recommended for inter-island quarantine between the southern Cook Islands (Space and Flynn, 2002b), and as a priority species for exclusion on Niue (Space et al., 2004). Planting of A. crenata is prohibited in Miami-Dade County, Florida (PIER, 2013), and it is on the Florida noxious weed list (Center for Aquatic and Invasive Plants, 2013).

Habitat

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 In general, A. crenata will grow in moist woodland in tropical and subtropical regions (USDA-APHIS, 2012), between 100-2400 m altitude (Weber, 2003).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
 
Terrestrial – ManagedUrban / peri-urban areas Present, no further details
Terrestrial ‑ Natural / Semi-naturalNatural forests Principal habitat Harmful (pest or invasive)
Natural forests Principal habitat Natural

Biology and Ecology

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Genetics

Artificial selection of A. crenata for attractive traits through cultivation for ornamental purposes has resulted in slower growth rates of cultivars, resulting in a greater ability to shade native plants, increased seed production through selection for attractive berry production, and greater ability to re-sprout, compared to wild ecotypes in its native range (Kitajima et al., 2006).

The genetic variation of A. crenata populations is variable: within populations the genetic diversity can be low, indicating the importance of inbreeding (Mu, 2010; Cheon et al., 2000), but in other locations the genetic diversity can be high both within and between populations (Zeng et al. 2012).

Reproductive Biology

A. crenata can reproduce sexually and asexually by seed, as well as from vegetative cuttings. It is a self-compatible species, but long-distance pollen dispersal is also important for the reproduction of the species as long-distance seed dispersal is inefficient for genetic mixing (Zeng et al., 2012). Sexual maturity is reached when plants are approximately 20 cm in height and start developing lateral branches. Each lateral branch has a  lifespan of two years: as a vegetative branch in its first year, and bearing flowers and fruits terminally in the second year (Kitajima et al., 2006). Bees and flies pollinate A. crenata but there are no specific pollinators outside of its native range (USDA-APHIS, 2012).

Plants can produce around 150 fruits per year (Fox and Kitajima, 2001), which have a long period of development (34 weeks) and remain attached for a long time (Lee et al., 2012). Seed germination is controlled by several environmental factors such as seed moisture content, temperature, and light; removal of the seed coat increases germination rates. The long period (>13 weeks for 80% seed germination) and long juvenile state (2-3 years) hinders and delays breeding of A. crenata (Tezuka et al., 2012; Roh et al., 2005). Optimum germination temperature is between 18-30°C. Young plants are slightly temperature-sensitive, but once established, plants can tolerate temperatures up to 37°C (Conover and Poole, 1989).

There is no evidence of A. crenata forming a soil seed bank, but seeds are able to survive up to two years in moist conditions at temperatures (e.g. 3°C) which are too cold for germination. In conditions suitable for germination, A. crenata has a high germination rate (>90% in one study), but also relatively high rates of seedling mortality (Fox and Kitajima, 2001).

A. crenata can also be propagated vegetatively through cuttings, which is faster than via seed: commercially viable plants take four years to produce from seed, and two via vegetative propagation (Roh et al., 2005).

Physiology and Phenology

Artificial selection for traits - such as enhanced berry production - through cultivation as an ornamental has increased the invasiveness of this species outside of its native range: exotic populations show increased seed production compared to native populations, due to the selection for enhanced berry production of ornamental plants (Kitajima et al., 2006).

There is no further evidence for differences between exotic and native populations, or phenological variation within populations.

Associations

A. crenata can be associated with root mycorrhizal fungi which have been shown to improve its growth rates and to alter its competitive interactions with native plant species (Bray et al., 2003).

Leaves have bacterial nodules in their margins as part of an obligate life-cycle symbiosis, where bacteria are maintained in the bud and re-infect each new leaf primordium as well as flowers and seeds, and are transmitted vertically to the next generation. It is thought that the nodules might contribute to growth capacity during different life stages, during resource shortage, or during exposure to specific herbivores or pathogens (Nakahashi et al., 2005).

Environmental Requirements

This species requires shade and is damaged by full sun. The optimal light conditions for growth are between 18-45% sunlight (Fox and Kitajima, 2001). Growth of A. crenata has also been shown to be positively correlated with soil phosphorous levels (Bray et al., 2003).

Climate

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ClimateStatusDescriptionRemark
Af - Tropical rainforest climate Preferred > 60mm precipitation per month
Aw - Tropical wet and dry savanna climate Preferred < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Preferred Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
36 20

Rainfall

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ParameterLower limitUpper limitDescription
Mean annual rainfall10002500mm; lower/upper limits

Soil Tolerances

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Soil drainage

  • free
  • impeded

Soil reaction

  • acid
  • alkaline
  • neutral
  • very acid
  • very alkaline

Soil texture

  • heavy
  • medium

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Haematonectria haematococca Pathogen Roots/Stems not specific N

Notes on Natural Enemies

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There is little literature regarding the natural enemies of A. crenata. Foot rot of A. crenata has been reported and first observed in Nantou (Taiwan) in 1997. It was caused by the fungus Fusarium solani [Haematonectria haematococca], with diseased plants showing basal stem rot, root crown rot and wilting. Infection resulted in plant death. Infection was promoted when the stem crown was wounded, but could be treated with anti-fungal agents (Fu and Chang, 1999).

Means of Movement and Dispersal

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Natural Dispersal (Non-Biotic)

The fruits and seeds of A. crenata are not adapted for wind dispersal, and there is no evidence that they are regularly dispersed by water although this could potentially occur.

Vector Transmission

The persistence of fruits on trees and high seedling density recorded around parent plants indicate that seed dispersal is low, although this may be compensated for by the high berry production of cultivars. Some seed dispersal is via birds, however; in Japan, part of the native range, all seeds are dispersed by birds including Ixos amaurotis (or Microscelis amaurotis) (Kitajima et al., 2006). In the USA gray catbirds (Dumetella carolinensis) are thought to be the most important avian consumers of Ardisia fruits, but outside the native range, dispersers are limited (Meisenburg, 2007); mockingbirds, cedar waxwings (Bombycilla cedrorum) and racoons (Procyon lotor) are also noted as dispersers in Florida (Langeland and Burks, 1998). Kitajima et al. (2006) also cite northern mockingbirds (Mimus polyglottos) as seed dispersers.

De-pulping of fruits by birds during digestion significantly increases seed germination rates: in one study, de-pulped fruits germinated 72% more than pulped fruits, and also germinated nearly 27% faster. This indicated that frugivores can significantly increase the spread of A. crenata by enhancing its germination rates (Chimera and Drake, 2010).

Accidental Introduction

Accidental introduction can occur via inappropriate disposal of garden waste (USDA-APHIS, 2012).

Intentional Introduction

A. crenata has been intentionally transported as an ornamental species used in gardens, with subsequent escape into the wild (USDA-APHIS, 2012).

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Escape from confinement or garden escapeThis is the primary invasion pathway Yes USDA-APHIS, 2012
Garden waste disposalIncorrect disposal of garden waste has been implicated in new populations Yes USDA-APHIS, 2012
Ornamental purposesInternational trade as an ornamental species has resulted in global spread Yes Yes Niu et al., 2012

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Host and vector organismsBirds, e.g. Bombycilla cedrorum, Dumetella carolinensis, Ixos amaurotis, Mimus polyglottos Yes Kitajima et al., 2006; Langeland and Burks, 1998; Meisenburg, 2007

Impact Summary

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CategoryImpact
Cultural/amenity Positive
Economic/livelihood Positive
Environment (generally) Negative
Human health Positive and negative

Economic Impact

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There are no known economic impacts associated with A. crenata invasion apart from the costs associated with control. There is no strong evidence that this species is problematic in agricultural production systems, and because it does not tolerate strong sunlight, it would be unlikely to affect agriculture in open land (USDA-APHIS, 2012). Randall et al. (2007) do record it as a weed of agriculture, however. It has been implicated in the death of livestock in 2001 and 2007, but there is no published literature into the toxicity of the berries (Sellers et al., 2013).

Environmental Impact

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Impact on Habitats

A. crenata invades the understory of mesic hardwood forests forming dense patches (up to 300 stems per m2), and competitively displaces native plants through shading (Kitajima et al., 2006). Langeland and Burks (1998) found that presence of A. crenata reduces the already low levels of light reaching forest floors by an additional 70%, which shades out native seedlings.

In the USA, A. crenata invades forest margins and understories, and riverbanks in swamp forests, and also thrives in urban and suburban woodlands (USDA-APHIS, 2012). Clewell and Tobe (2011) state that in Northern Florida, native old-field pineland with associated hardwood undergrowth has been replaced by alien hardwood forest dominated by invasive species; A. crenata dominates the understory of this non-native forest and is the species with the most ground cover. Within the Hawai’i Volcanoes National Park, A. crenata is invasive in the lowlands, but is under control by park authorities (Stone et al., 1992).

In Mauritius, Lorence and Sussman (1986) report that dense thickets of A. crenata are likely to displace native forest species in time, and on Réunion Island it is ranked 10th among 26 non-indigenous woody plant species as having an impact, and innate ability to be a pest, of 64% of the maximum possible score (Tassin et al., 2006).

Impact on Biodiversity

There is a negative correlation between the abundance of native understorey plants and the presence of A. crenata (Fox and Kitajima, 2001). Interestingly, introduced A. crenata individuals have been shown to grow at higher densities and be more shade-tolerant in their new ranges compared to the native range, leading to suppression of local understorey plants and the formation of dense monocultures causing severe impacts on biodiversity (Niu et al., 2012).

Indirect impacts on biodiversity have been investigated on Réunion Island, where increasing numbers of exotic plant species, including A. crenata, have been hypothesised to be leading to increases in the number of exotic frugivorous bird species which could potentially impact native bird species (Kueffer and Lavergne, 2004).

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Silene ovata (fringed campion)USA ESA listing as endangered species USA ESA listing as endangered speciesFlorida; GeorgiaCompetition - monopolizing resourcesUS Fish and Wildlife Service, 1996

Social Impact

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There is little evidence of A. crenata having a detrimental social impact. It has been implicated in the death of livestock in 2001 and 2007, which indicates that the berries could be toxic to animals and humans; there is no published literature into the toxicity of A. crenata berries (Sellers et al., 2013).

Risk and Impact Factors

Top of page Invasiveness
  • Invasive in its native range
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Tolerant of shade
  • Has high reproductive potential
  • Has propagules that can remain viable for more than one year
  • Reproduces asexually
  • Has high genetic variability
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Modification of successional patterns
  • Monoculture formation
  • Negatively impacts animal health
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Competition - shading
Likelihood of entry/control
  • Highly likely to be transported internationally deliberately

Uses

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Economic Value

A. crenata is an ornamental plant of economic importance through sales and provision of jobs in cultivating the species. There is little literature on its economic value, but Wirth et al. (2004) estimate that in Florida, annual sales of more than $2 million result in $4.8 million of economic output, supporting 65 jobs.

There is potential for A. crenata extracts to be developed as anti-herbivory compounds (Sandoval-Mojica and Capinera, 2011).

Social Benefit

In China, A. crenata is widely cultivated for its medicinal properties, which arise from the high concentration of triterpenoid saponins in the roots (Kitajima et al., 2006), which have been recorded in the origins of several traditional Chinese medicines (Zhao et al., 2012). A. crenata has several medicinal properties including utero-contraction, platelet aggregation and lowering blood pressure, cAMP inhibition, and antithrombin inhibition (Kobayashi and de Mejía, 2005). It has traditionally been used in Chinese medicine to treat respiratory tract infections, tonsillitis and menstrual disorders (Zaima et al., 2013), and parasitoses (Niu et al., 2012).

Uses List

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General

  • Ornamental

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical
  • Traditional/folklore

Diagnosis

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There is currently no information on laboratory techniques which could be used to identify this species.

Detection and Inspection

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Identification of the species in the field would be via detailed surveys and identification.

Similarities to Other Species/Conditions

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There are many species and cultivars of Ardisia, and identifying Ardisia crenata may be difficult when other Ardisia species and varieties are also present.

Prevention and Control

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Prevention

Limit planting and remove existing plants within the landscape. If possible, removal should occur before the plant produces seeds. As fruits containing seeds remain on plants for several months, care should be taken not to spread fruits and seeds during the removal process (Center for Aquatic and Invasive Plants, 2013).

Public awareness

Humans are the primary mechanism for long-distance dispersal of A. crenata; therefore constraining transmission by human endeavour is the best way to restrict introduction and establishment of populations (Niu et al., 2012).

Control

Cultural control

Cultural management is difficult once the plant has become established, but good ground cover would limit seedling establishment (Center for Aquatic and Invasive Plants, 2013).

Mechanical control

For small or isolated infestations, hand-pulling of plants is effective for seedling control. Larger plants can be cut or burned, but re-growth from roots and root crowns needs monitoring for at least a year (Center for Aquatic and Invasive Plants, 2013). Roots are unable to re-sprout if they are severed from the root crown; therefore root crown removal is essential to prevent re-growth after mechanical control (Fox and Kitajima, 2001).

Biological control

There are no known biological control agents for Ardisia crenata. However, because invading populations are thought to have originated in China, potential sources of biological control might be found there, within its native range (Niu et al., 2012).

Chemical control

Herbicides are effective, but the waxy leaves may prevent some uptake; care should be taken to avoid targeting desirable plants with herbicide applications (Center for Aquatic and Invasive Plants, 2013). One herbicide trial (Hutchinson et al., 2011) found that application to mature plants suppressed plant cover for 12 months, with highly variable seedling cover within the same period; monitoring and re-treatment at 12 months was recommended for further suppression, especially for emerging seedlings.

Gaps in Knowledge/Research Needs

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Overall there is a good level of information on A. crenata. Quantification of the impacts of invasion on biodiversity is lacking however. Further research into potential modes of biological control, and increasing public awareness on the impacts of this species, would be helpful.

References

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Links to Websites

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WebsiteURLComment
Global Invasive Species Databasehttp://www.issg.org/database/welcome
PIERhttp://www.hear.org/pier
UF-IFAS Center for Aquatic and Invasive Plantshttp://plants.ifas.ufl.edu
USDA PLANTS databasehttp://plants.usda.gov

Organizations

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USA: Center for Aquatic and Invasive Plants, University of Florida, Gainesville, Florida, http://plants.ifas.ufl.edu

USA: PIER (Pacific Island Ecosystems at Risk), University of Hawaii, Honolulu, http://www.hear.org/pier/index.html

Contributors

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11/04/13: Original text by:

Isabel Jones, consultant, UK

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