Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide







  • Last modified
  • 16 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Kappaphycus
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Rhodophyta
  •       Class: Rhodophyceae
  •         Order: Gigartinales
  • Summary of Invasiveness
  • Kappaphycus alvarezii and Kappaphycus striatum have been introduced to numerous countries around the world for commercial aquaculture for the carrageenan industry (...

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Fixed off-bottom farming with long straight lines in Uzi cotonii farm in Tanzania.
TitleFixed off-bottom farming
CaptionFixed off-bottom farming with long straight lines in Uzi cotonii farm in Tanzania.
CopyrightErick Ask
Fixed off-bottom farming with long straight lines in Uzi cotonii farm in Tanzania.
Fixed off-bottom farmingFixed off-bottom farming with long straight lines in Uzi cotonii farm in Tanzania.Erick Ask
Harvesting of Kappaphycus in Bali, Indonesia.
CaptionHarvesting of Kappaphycus in Bali, Indonesia.
CopyrightErick Ask
Harvesting of Kappaphycus in Bali, Indonesia.
HarvestingHarvesting of Kappaphycus in Bali, Indonesia.Erick Ask
Harvesting of Kappaphycus in Songo-Songo, Tanzania.
CaptionHarvesting of Kappaphycus in Songo-Songo, Tanzania.
CopyrightErick Ask
Harvesting of Kappaphycus in Songo-Songo, Tanzania.
HarvestingHarvesting of Kappaphycus in Songo-Songo, Tanzania.Erick Ask
Raft farming of Kappaphycus in Madura, Indonesia.
TitleRaft farming of algae
CaptionRaft farming of Kappaphycus in Madura, Indonesia.
CopyrightErick Ask
Raft farming of Kappaphycus in Madura, Indonesia.
Raft farming of algaeRaft farming of Kappaphycus in Madura, Indonesia.Erick Ask
Harvesting of Kappaphycus in Bali, Indonesia.
TitleHarvesting algae
CaptionHarvesting of Kappaphycus in Bali, Indonesia.
CopyrightErick Ask
Harvesting of Kappaphycus in Bali, Indonesia.
Harvesting algaeHarvesting of Kappaphycus in Bali, Indonesia.Erick Ask


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Preferred Scientific Name

  • Kappaphycus

International Common Names

  • English: agar-agar
  • Chinese: chilin-t'-sai

Local Common Names

  • India: Indian carrageen
  • Indonesia: agar-agar besar; agar-agar pulau
  • Philippines: tambalang

Summary of Invasiveness

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Kappaphycus alvarezii and Kappaphycus striatum have been introduced to numerous countries around the world for commercial aquaculture for the carrageenan industry (Zemke-White and Ohno, 1999; Zemke-White and Smith, 2006). Cultivation most often occurs in open culture where cuttings of the algae are secured onto rope lines which are strung above the benthos in shallow reef flat environments. Despite the widespread introduction of these species few reports have indicated that they have established natural populations in the areas surrounding farm plots. However, in the Hawaiian Islands where several carrageenophtyes were introduced in the 1970s evidence suggests that many species have since spread and become naturalized on the reefs in Kane’ohe Bay, Oahu (Smith et al., 2002; Conklin and Smith, 2005). These species all reproduce through vegetative propagation thus facilitating spread and dispersal. Despite the evidence that these species have established wild populations and may even be reproducing sexually (J Smith, National Center for Ecological Analysis and Synthesis, UC Santa Barbara, personal communication, 2008), there are little data to indicate the ecological impacts that they may be having on the reef communities.

However, species of Kappaphycus can regularly be seen growing on or over reef building corals. Due to the extremely large size of these algae they seem to be able to outcompete or smother many native reef species. Because of the concern that has been generated in Hawai’i about the potential ecological impacts of these species various seaweed removal activities have taken place. Researchers initially conducted removal by hand but have since moved to removal with a large underwater vaccum/suction pump operated from a floating barge. While the “super sucker” (Pala, 2008) is mainly used on 2 other highly invasive algal species Graciliaria salicornia and Eucheuma denticulatum (also introduced for aquaculture) occasionally events also target Kappaphycus. While Kappaphycus spp. are clearly successful invaders on Hawaii’s reefs they are not among the most problematic marine invasive species. Little information exists on the invasion potential of these species for the rest of the world.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Rhodophyta
  •             Class: Rhodophyceae
  •                 Order: Gigartinales
  •                     Family: Solieriaceae
  •                         Genus: Kappaphycus


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The thallus of Kappaphycus alvarezii ranges from 24 to 48 cm. The branches are cartilaginous and pliable, ranging from 8 to 12 cm in length with unilateral to irregular branching type. Branch diameter ranges from a few mm at the branch tips to greater than 1 cm in older tissue. Generally the branches are smooth and the thallus can be short with many branches to much larger with long smooth branches. The diameter of the different types of vegetative cells are as follows: 2-4 µm outer cortex, 30-240 µm inner cortex and, 25-40 µm medulla. Thylles of the medulla are present but rhizoids are absent.


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Algaebase reports that Kappaphycus has been recorded from the Philippines, Malaysia, Cuba, China, Vietnam, Micronesia, Fiji, Hawaii and Tahiti . However many of these locations are not part of the native distribution but are rather the result of aquaculture introductions. See Zemke-White and Smith (2006) for more information.

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


ChinaPresentIntroduced1985Wu et al., 1988
IndiaLocalisedIntroduced1989Mairh et al., 1995Saurashtra region
IndonesiaPresentNative1984Soerjodinoto, 1969
JapanLocalisedNative1983Mairh et al., 1986
MalaysiaPresentNative1978Doty, 1980
MaldivesLocalisedIntroduced1986de Reviers, 1989
PhilippinesLocalisedNative1971Doty and Alvarez, 1975
VietnamLocalisedIntroduced1993Ohno et al., 1996


DjiboutiPresentIntroduced1973Braud et al., 1974
KenyaPresentIntroduced1996Ask et al., undated
MadagascarPresentIntroduced1998Ask et al., undated
TanzaniaPresentIntroduced1989Lirasan and Twide, 1993

North America

MexicoPresentIntroduced1980sMuñoz et al., 2004
USAPresentPresent based on regional distribution.
-CaliforniaAbsent, formerly presentIntroducedDoty and Norris, 1985
-FloridaIntroduced, establishment uncertainIntroducedDawes, 1989
-HawaiiPresentIntroduced1970 Invasive Doty, 1985a; Doty, 1985b; Doty and Norris, 1985; Eldredge, 1994Multiple introductions to Kane’ohe Bay and Honolulu Harbor, Oahu

Central America and Caribbean

CubaPresentIntroduced1991Serpa-Madrigal et al., 1997

South America

BrazilPresentIntroduced1995de Paula et al., 1999
VenezuelaPresentIntroduced1996Rincones and Rubio, 1999


Cook IslandsIntroduced, establishment uncertainIntroduced1980sEldredge, 1994Aitutaki
FijiPresentIntroduced1976Prakash, 1990; Eldredge, 1994Suva, Mana, Telau, Bau, Rakiraki
French PolynesiaPresentIntroduced1985Doty and Norris, 1985
GuamLocalisedIntroduced1985Doty and Norris, 1985
Johnston IslandPresentRussell, 1982
KiribatiPresentIntroduced1977Russell, 1982; Luxton and Luxton, 1999Kiritimati, Fanning & Butaritari Atolls
Marshall IslandsPresentIntroduced1990Eldredge, 1994Majuro lagoon, Mili and Lildep
Micronesia, Federated states ofPresentIntroducedDoty and Norris, 1985Pohnpei, Kosrae
SamoaPresentIntroduced1975Doty and Santos, 1978Upolo
Solomon IslandsLocalisedIntroduced1987Eldredge, 1994Vonavona, Munda, Ontong Java
TongaLocalisedIntroduced1982Eldredge, 1994Vava'u


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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
California 1985 Research (pathway cause)Unknown No No Doty (1985a)
China 1985 Aquaculture (pathway cause)Unknown No No Wu et al. (1988)
Djibouti 1973 Unknown No No Braud et al. (1974)
Fiji mid 1970s Unknown No No Prakash (1990)
Florida 1988 Research (pathway cause)Unknown No No Dawes (1989)
Hawaii 1971, 1985 Unknown No No Doty (1985a); Doty (1985b)
India 1989 Unknown No No Mairh et al. (1995)
Indonesia 1985 Aquaculture (pathway cause)Unknown Yes Yes Soerjodinoto (1969)
Japan 1983 Unknown No No Mairh et al. (1986)
Kiribati 1977, 1981 Aquaculture (pathway cause)Unknown No No Russell (1982)
Malaysia 1978 Aquaculture (pathway cause)Unknown Yes Yes Doty (1980)
Philippines 1971 Aquaculture (pathway cause)Unknown Yes Yes Doty and Alvarez (1973)
Tanzania 1989 Aquaculture (pathway cause)Unknown Yes Yes Lirasan and Twide (1993)
Vietnam 1993 Unknown No No Ohno et al. (1995); Ohno et al. (1996)

Habitat List

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Coral reefs Present, no further details Harmful (pest or invasive)
Benthic zone Present, no further details Harmful (pest or invasive)

Biology and Ecology

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Reproductive Biology

The life cycle of Kappaphycus spp. is triphasic and comprises the tetrasporophyte (2n), carposporophyte (2n) and gametophyte (n) phases. The vegetative and reproductive structures of tetrasporic and gametophytic populations and their occurrence in farming sites in the Philippines have been reported.

K. alvarezii produces zonate tetraspores and the tetrasporophytes have been found in K. alvarezii cultured species. Tetrasporophytes dominate natural beds near farms in Tawi-Tawi, Philippines whereas in Danajon Reef, Philippines, only K. striatum var. dichotomous (or ‘sakol’) and E. denticulatum from farms were found to have more tetrasporophytes. Male gametophytes and structures are rare to unknown in these genera.

Clone cultivation, however, is presently very useful for farming; further research, should therefore include the use of sporelings for culture as in other economically important seaweeds such as Porphyra and Laminaria. Sporeling cultivation could provide the possibility of other farming manipulations and increase genetic variation.


Light response

Light response curves have been plotted for the commercial eucheumatoids. Photosynthetic rates vary for tissue types in E. striatum var. tambalang with younger tissues having higher rates. In addition, there are UV-absorbing compounds in E. striatum that absorb at 333 nm, that increase in quantity with exposure to higher levels of UV. This photopigment is destroyed when exposed to twice the level of UV found in nature.

The different colour types of K. alvarezii have different photosynthetic responses related to the photosynthetic pigments characteristic of each type. Ecotypic differentiation and variation in photosynthetic efficiency have been shown for E. denticulatum colour types red, brown and green, which were not shown by K. alvarezii when nitrogen was limiting (Ask and Azanza, 2001).

Temperature response

Dawes (1989) studied the temperature responses with acclimation from 22 to 25°C of K. alvarezii and reported no acclimation of the species to 18ºC either through gradual or abrupt transfers. K. striatum (var. tambalang and elkhorn) and E. denticulatum have maximum photosynthetic rates at 30°C with inhibition at 35-40°C. An optimum photosynthetic rate was obtained between 30 and 35°C for E. denticulatum.

In tropical areas, Trono and Ohno (1989) reported that rapid growth and high biomass production by Kappaphycus occurs during months characterized by warmer temperature, i.e. 25-30°C.

Salinity response

Ask and Azanza (2001) suggested that single and interactive effects of salinity with other factors on the commercial cultivation of Kappaphycus should be studied because in shallow floating forms or off bottom farms exposed at low tide the crops can be exposed to rapid decreases in salinity during tropical downpours. Noticeable drops in temperature and light levels also happen and plants located in the intertidal zone can experience rapid changes in salinity from freshwater runoff.


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A - Tropical/Megathermal climate Preferred Average temp. of coolest month > 18°C, > 1500mm precipitation annually
Am - Tropical monsoon climate Tolerated Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
Cf - Warm temperate climate, wet all year Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Cs - Warm temperate climate with dry summer Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Tolerated 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)
20 20

Air Temperature

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Parameter Lower limit Upper limit
Mean annual temperature (ºC) 20 30

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Illumination (Lux illuminance) >10,000 Harmful Adult
Illumination (Lux illuminance) 6000 Optimum Adult
Salinity (part per thousand) 32 36 Optimum
Salinity (part per thousand) <24 >45 Harmful Adult
Water temperature (ºC temperature) 22 29 Optimum 20-30 tolerated
Water temperature (ºC temperature) 25 30 Optimum Adult
Water temperature (ºC temperature) 35 40 Harmful Adult

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Chelonia mydas Herbivore Adult
Diadema setosum Herbivore Adult
Tripneustes gratilla Herbivore Adult

Impact Summary

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Animal/plant products Positive
Crop production Positive
Environment (generally) Positive
Fisheries / aquaculture Positive
Human health Positive
Other Positive
Tourism Positive
Trade/international relations Positive

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Highly adaptable to different environments
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Pioneering in disturbed areas
  • Long lived
  • Has high reproductive potential
  • Has propagules that can remain viable for more than one year
  • Reproduces asexually
Impact outcomes
  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Modification of natural benthic communities
  • Modification of successional patterns
  • Reduced native biodiversity
Impact mechanisms
  • Competition - monopolizing resources
  • Competition - shading
  • Competition - smothering
  • Competition - strangling
Likelihood of entry/control
  • Highly likely to be transported internationally deliberately
  • Difficult/costly to control

Gaps in Knowledge/Research Needs

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Species of Kappaphycus have been introduced to numerous countries around the world over the last 40 years yet very little is known about the potential ecological/environmental impacts that these species may be having on surrounding benthic communities. Additionally, very little information is known about the ecology of Kappaphycus spp. in their native range. Based on data and observations from exotic populations in the Hawaiian Islands it seems that these species do have the potential to become invasive. More detailed assessments need to be conducted around the world to determine if these species are regularly establishing wild populations and if so what are the impacts to these ecosystems? Is the benefit gained by farming outweighed by the cost of introducing non-native species?


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Zemke-White WL, Ohno M, 1999. World seaweed utilization: an end-of-century summary. Journal of Applied Phycology, 11(4):369-376.

Zemke-White WL, Smith JE, 2006. Environmental impacts of seaweed farming in the tropics. World Seaweed Resources [ed. by Critchley AT, Ohno M, Largo DB].

Zuccarello GC, Critchley AT, Smith J, Sieber V, Lhonneur GB, West JA, 2006. Systematics and genetic variation in commercial shape Kappaphycus and shape Eucheuma (Solieriaceae, Rhodophyta). Journal of Applied Phycology, 18(3/5):643-651.

Links to Websites

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GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gateway source for updated system data added to species habitat list.
Irish Seaweed Centre
Marine Algae of Hawaii


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22/04/2008 Updated by:

Jennifer Smith, National Center for Ecological Analysis & Synthesis, University of California, Santa Barbara,, 725 State St. Suite 300, Santa Barbara, CA 93101, USA

Main Author
Rhodora Azanza
The Marine Science Institute, College of Science, University of the Philippines, Velasquez Street, Diliman, Quezon City 1101, Philippines

Distribution Maps

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