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Plasmodium relictum

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Datasheet

Plasmodium relictum

Summary

  • Last modified
  • 01 October 2018
  • Datasheet Type(s)
  • Invasive Species
  • Vector of Animal Disease
  • Preferred Scientific Name
  • Plasmodium relictum
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Protista
  •     Phylum: Protozoa
  •       Subphylum: Apicomplexa
  •         Order: Haemospororida
  • Summary of Invasiveness
  • P. relictum is a common mosquito-transmitted blood protozoan of wild birds that has a worldwide distribution. It has been reported from at least 411 avian species from 67 avian families and is considered to be re...

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Pictures

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PictureTitleCaptionCopyright
Composite figure for comparison: See individual figures for specific caption data.
TitleHistology - Composite comparison figure
CaptionComposite figure for comparison: See individual figures for specific caption data.
CopyrightU.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Composite figure for comparison: See individual figures for specific caption data.
Histology - Composite comparison figureComposite figure for comparison: See individual figures for specific caption data.U.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Early (left (a)) and late (right (b)) trophozoites (arrowed) of Plasmodium relictum in erythrocytes of an endangered Puaiohi (Myadestes palmeri).  Golden brown or black pigment granules (P) are produced as a by-product of haemoglobin digestion and are usually not visible in young trophozoites (left).
TitleHistology
CaptionEarly (left (a)) and late (right (b)) trophozoites (arrowed) of Plasmodium relictum in erythrocytes of an endangered Puaiohi (Myadestes palmeri). Golden brown or black pigment granules (P) are produced as a by-product of haemoglobin digestion and are usually not visible in young trophozoites (left).
CopyrightU.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Early (left (a)) and late (right (b)) trophozoites (arrowed) of Plasmodium relictum in erythrocytes of an endangered Puaiohi (Myadestes palmeri).  Golden brown or black pigment granules (P) are produced as a by-product of haemoglobin digestion and are usually not visible in young trophozoites (left).
HistologyEarly (left (a)) and late (right (b)) trophozoites (arrowed) of Plasmodium relictum in erythrocytes of an endangered Puaiohi (Myadestes palmeri). Golden brown or black pigment granules (P) are produced as a by-product of haemoglobin digestion and are usually not visible in young trophozoites (left).U.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Multinucleated meront (or schizont) of Plasmodium relictum in erythrocyte of an endangered Puaoihi (Myadestes palmeri).  Red, grape-like nuclei (arrowed) surround a central residual mass of pigment (P).  As the meront matures, nuclei will bud into developing merozoites that are released when the infected host cell ruptures.  Each merozoite can invade another erythrocyte to start another round of asexual reproduction.
TitleHistology
CaptionMultinucleated meront (or schizont) of Plasmodium relictum in erythrocyte of an endangered Puaoihi (Myadestes palmeri). Red, grape-like nuclei (arrowed) surround a central residual mass of pigment (P). As the meront matures, nuclei will bud into developing merozoites that are released when the infected host cell ruptures. Each merozoite can invade another erythrocyte to start another round of asexual reproduction.
CopyrightU.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Multinucleated meront (or schizont) of Plasmodium relictum in erythrocyte of an endangered Puaoihi (Myadestes palmeri).  Red, grape-like nuclei (arrowed) surround a central residual mass of pigment (P).  As the meront matures, nuclei will bud into developing merozoites that are released when the infected host cell ruptures.  Each merozoite can invade another erythrocyte to start another round of asexual reproduction.
HistologyMultinucleated meront (or schizont) of Plasmodium relictum in erythrocyte of an endangered Puaoihi (Myadestes palmeri). Red, grape-like nuclei (arrowed) surround a central residual mass of pigment (P). As the meront matures, nuclei will bud into developing merozoites that are released when the infected host cell ruptures. Each merozoite can invade another erythrocyte to start another round of asexual reproduction.U.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Mature macrogametocyte (arrow) in erythrocyte of an endangered Puaiohi (Myadestes palmeri). Female (macrogametocytes) and male (microgametocytes) develop from merozoites and are infectious to mosquitoes.  Macrogametocytes typically stain dark blue and have a single, small compact central nucleus and a number of pigment granules (P). Among members of the subgenus Haemamoeba, gametocytes are typically round or oval and exceed the size of the host cell nucleus (N). The host cell nucleus is usually displaced to one side of the erythrocyte and may be partially or completely turned.
TitleHistology
CaptionMature macrogametocyte (arrow) in erythrocyte of an endangered Puaiohi (Myadestes palmeri). Female (macrogametocytes) and male (microgametocytes) develop from merozoites and are infectious to mosquitoes. Macrogametocytes typically stain dark blue and have a single, small compact central nucleus and a number of pigment granules (P). Among members of the subgenus Haemamoeba, gametocytes are typically round or oval and exceed the size of the host cell nucleus (N). The host cell nucleus is usually displaced to one side of the erythrocyte and may be partially or completely turned.
CopyrightU.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Mature macrogametocyte (arrow) in erythrocyte of an endangered Puaiohi (Myadestes palmeri). Female (macrogametocytes) and male (microgametocytes) develop from merozoites and are infectious to mosquitoes.  Macrogametocytes typically stain dark blue and have a single, small compact central nucleus and a number of pigment granules (P). Among members of the subgenus Haemamoeba, gametocytes are typically round or oval and exceed the size of the host cell nucleus (N). The host cell nucleus is usually displaced to one side of the erythrocyte and may be partially or completely turned.
HistologyMature macrogametocyte (arrow) in erythrocyte of an endangered Puaiohi (Myadestes palmeri). Female (macrogametocytes) and male (microgametocytes) develop from merozoites and are infectious to mosquitoes. Macrogametocytes typically stain dark blue and have a single, small compact central nucleus and a number of pigment granules (P). Among members of the subgenus Haemamoeba, gametocytes are typically round or oval and exceed the size of the host cell nucleus (N). The host cell nucleus is usually displaced to one side of the erythrocyte and may be partially or completely turned.U.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Mature microgametocyte (arrow) in erythrocyte of an endangered Puaiohi (Myadestes palmeri). Female (macrogametocytes) and male (microgametocytes) develop from merozoites and are infectious to mosquitoes.  Microgametocytes typically stain pink or light blue, depending on staining, and have a single, large diffuse central nucleus and a number of pigment granules (P). Among members of the subgenus Haemamoeba, gametocytes are typically round or oval and exceed the size of the host cell nucleus (N). The host cell nucleus is usually displaced to one side of the erythrocyte and may be partially or completely turned.
TitleHistology
CaptionMature microgametocyte (arrow) in erythrocyte of an endangered Puaiohi (Myadestes palmeri). Female (macrogametocytes) and male (microgametocytes) develop from merozoites and are infectious to mosquitoes. Microgametocytes typically stain pink or light blue, depending on staining, and have a single, large diffuse central nucleus and a number of pigment granules (P). Among members of the subgenus Haemamoeba, gametocytes are typically round or oval and exceed the size of the host cell nucleus (N). The host cell nucleus is usually displaced to one side of the erythrocyte and may be partially or completely turned.
CopyrightU.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Mature microgametocyte (arrow) in erythrocyte of an endangered Puaiohi (Myadestes palmeri). Female (macrogametocytes) and male (microgametocytes) develop from merozoites and are infectious to mosquitoes.  Microgametocytes typically stain pink or light blue, depending on staining, and have a single, large diffuse central nucleus and a number of pigment granules (P). Among members of the subgenus Haemamoeba, gametocytes are typically round or oval and exceed the size of the host cell nucleus (N). The host cell nucleus is usually displaced to one side of the erythrocyte and may be partially or completely turned.
HistologyMature microgametocyte (arrow) in erythrocyte of an endangered Puaiohi (Myadestes palmeri). Female (macrogametocytes) and male (microgametocytes) develop from merozoites and are infectious to mosquitoes. Microgametocytes typically stain pink or light blue, depending on staining, and have a single, large diffuse central nucleus and a number of pigment granules (P). Among members of the subgenus Haemamoeba, gametocytes are typically round or oval and exceed the size of the host cell nucleus (N). The host cell nucleus is usually displaced to one side of the erythrocyte and may be partially or completely turned.U.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Giemsa-stained blood smear from an Iiwi (Vestiaria coccinea) with an acute infection with Plasmodium relictum. The normal cellular makeup of the blood is severely altered, with mature erythrocytes being replaced by immature erythrocytes (reticulocytes, R) and erythrocyte precursors (EP) as the bird struggles to compensate for destruction of red blood cells by the parasites. Most of the red blood cells are parasitized (P) with trophozoites and meronts.
TitleHistology
CaptionGiemsa-stained blood smear from an Iiwi (Vestiaria coccinea) with an acute infection with Plasmodium relictum. The normal cellular makeup of the blood is severely altered, with mature erythrocytes being replaced by immature erythrocytes (reticulocytes, R) and erythrocyte precursors (EP) as the bird struggles to compensate for destruction of red blood cells by the parasites. Most of the red blood cells are parasitized (P) with trophozoites and meronts.
CopyrightU.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Giemsa-stained blood smear from an Iiwi (Vestiaria coccinea) with an acute infection with Plasmodium relictum. The normal cellular makeup of the blood is severely altered, with mature erythrocytes being replaced by immature erythrocytes (reticulocytes, R) and erythrocyte precursors (EP) as the bird struggles to compensate for destruction of red blood cells by the parasites. Most of the red blood cells are parasitized (P) with trophozoites and meronts.
HistologyGiemsa-stained blood smear from an Iiwi (Vestiaria coccinea) with an acute infection with Plasmodium relictum. The normal cellular makeup of the blood is severely altered, with mature erythrocytes being replaced by immature erythrocytes (reticulocytes, R) and erythrocyte precursors (EP) as the bird struggles to compensate for destruction of red blood cells by the parasites. Most of the red blood cells are parasitized (P) with trophozoites and meronts.U.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Oocysts of Plasmodium relictum on the outer midgut wall of Culex quinquefasciatus.  The oocysts are spherical in shape and at maturity contain thousands of elongate sporozoites (arrows).  These are released into the haemocoel when the oocysts rupture, travel to the salivary glands, and are injected into a new host when the mosquito takes another blood meal.
TitleHistology
CaptionOocysts of Plasmodium relictum on the outer midgut wall of Culex quinquefasciatus. The oocysts are spherical in shape and at maturity contain thousands of elongate sporozoites (arrows). These are released into the haemocoel when the oocysts rupture, travel to the salivary glands, and are injected into a new host when the mosquito takes another blood meal.
CopyrightU.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Oocysts of Plasmodium relictum on the outer midgut wall of Culex quinquefasciatus.  The oocysts are spherical in shape and at maturity contain thousands of elongate sporozoites (arrows).  These are released into the haemocoel when the oocysts rupture, travel to the salivary glands, and are injected into a new host when the mosquito takes another blood meal.
HistologyOocysts of Plasmodium relictum on the outer midgut wall of Culex quinquefasciatus. The oocysts are spherical in shape and at maturity contain thousands of elongate sporozoites (arrows). These are released into the haemocoel when the oocysts rupture, travel to the salivary glands, and are injected into a new host when the mosquito takes another blood meal.U.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Hematocrit tubes from a wild Apapane (Himatione sanguinea) with a naturally acquired infection with Plasmodium relictum (a) and an uninfected canary (b) for comparison.  After blood was collected into the heparinized tubes, they were sealed at one end with clay and centrifuged at high speed to separate erythrocytes and white blood cells from plasma.  Among most passerines, the hematocrit or volume of packed cells is about 50%.  Birds with acute malaria typically have much lower hematocrits as a result of red blood cell destruction.
TitleHematocrit tubes
CaptionHematocrit tubes from a wild Apapane (Himatione sanguinea) with a naturally acquired infection with Plasmodium relictum (a) and an uninfected canary (b) for comparison. After blood was collected into the heparinized tubes, they were sealed at one end with clay and centrifuged at high speed to separate erythrocytes and white blood cells from plasma. Among most passerines, the hematocrit or volume of packed cells is about 50%. Birds with acute malaria typically have much lower hematocrits as a result of red blood cell destruction.
CopyrightU.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Hematocrit tubes from a wild Apapane (Himatione sanguinea) with a naturally acquired infection with Plasmodium relictum (a) and an uninfected canary (b) for comparison.  After blood was collected into the heparinized tubes, they were sealed at one end with clay and centrifuged at high speed to separate erythrocytes and white blood cells from plasma.  Among most passerines, the hematocrit or volume of packed cells is about 50%.  Birds with acute malaria typically have much lower hematocrits as a result of red blood cell destruction.
Hematocrit tubesHematocrit tubes from a wild Apapane (Himatione sanguinea) with a naturally acquired infection with Plasmodium relictum (a) and an uninfected canary (b) for comparison. After blood was collected into the heparinized tubes, they were sealed at one end with clay and centrifuged at high speed to separate erythrocytes and white blood cells from plasma. Among most passerines, the hematocrit or volume of packed cells is about 50%. Birds with acute malaria typically have much lower hematocrits as a result of red blood cell destruction.U.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Hawaii Amakihi with an acute infection with Plasmodium relictum. When the abdominal cavity is opened, enlargement and discoloration of the liver (arrow) is immediately apparent and, in combination with a stained blood smear, is an important diagnostic criterion for acute malarial infection.
TitlePathology
CaptionHawaii Amakihi with an acute infection with Plasmodium relictum. When the abdominal cavity is opened, enlargement and discoloration of the liver (arrow) is immediately apparent and, in combination with a stained blood smear, is an important diagnostic criterion for acute malarial infection.
CopyrightU.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Hawaii Amakihi with an acute infection with Plasmodium relictum. When the abdominal cavity is opened, enlargement and discoloration of the liver (arrow) is immediately apparent and, in combination with a stained blood smear, is an important diagnostic criterion for acute malarial infection.
PathologyHawaii Amakihi with an acute infection with Plasmodium relictum. When the abdominal cavity is opened, enlargement and discoloration of the liver (arrow) is immediately apparent and, in combination with a stained blood smear, is an important diagnostic criterion for acute malarial infection.U.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Spleens (top) and livers (bottom) from a canary with an acute infection with Plasmodium relictum (left) and an uninfected canary (right). Both the liver and spleen increase dramatically in size during acute infections and become darkly discoloured by deposition of pigment in tissue macrophages.
TitlePathology
CaptionSpleens (top) and livers (bottom) from a canary with an acute infection with Plasmodium relictum (left) and an uninfected canary (right). Both the liver and spleen increase dramatically in size during acute infections and become darkly discoloured by deposition of pigment in tissue macrophages.
CopyrightU.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
Spleens (top) and livers (bottom) from a canary with an acute infection with Plasmodium relictum (left) and an uninfected canary (right). Both the liver and spleen increase dramatically in size during acute infections and become darkly discoloured by deposition of pigment in tissue macrophages.
PathologySpleens (top) and livers (bottom) from a canary with an acute infection with Plasmodium relictum (left) and an uninfected canary (right). Both the liver and spleen increase dramatically in size during acute infections and become darkly discoloured by deposition of pigment in tissue macrophages.U.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
The southern house mosquito (Culex quinquefasciatus) is a proven natural vector of Plasmodium relictum in Hawaii and other parts of the world.
TitleNatural vector
CaptionThe southern house mosquito (Culex quinquefasciatus) is a proven natural vector of Plasmodium relictum in Hawaii and other parts of the world.
CopyrightU.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii
The southern house mosquito (Culex quinquefasciatus) is a proven natural vector of Plasmodium relictum in Hawaii and other parts of the world.
Natural vector The southern house mosquito (Culex quinquefasciatus) is a proven natural vector of Plasmodium relictum in Hawaii and other parts of the world.U.S. Geological Survey/Pacific Island Ecosystems Research Center, Hawaii

Identity

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Preferred Scientific Name

  • Plasmodium relictum (Grassi & Feletti, 1891)

Other Scientific Names

  • Haemamoeba majoris Laveran, 1902 (partim)
  • Haemamoeba relicta Grassi and Feletti, 1891
  • Haemoproteus alaudae Celli and Sanfelice, 1891 (partim)
  • Plasmodium alaudae (Celli and Sanfelice, 1891) emend. Garnham, 1966
  • Plasmodium biziurae (Gilruth, Sweet and Dodd, 1910) emend. Coatney and Roudabush, 1936
  • Plasmodium capistrani Russell, 1932
  • Plasmodium chloropsidis Mello, 1935
  • Plasmodium grassii Labbé, 1894, emend. Coatney and Roudabush, 1936
  • Plasmodium inconstans Hartman, 1927
  • Plasmodium maior Raffaele, 1930
  • Plasmodium majoris Lühe, 1906
  • Plasmodium muniae (Das Gupta and Siddons, 1941) emend. Garnham, 1966
  • Plasmodium paddae Brumpt, 1935
  • Plasmodium passeris Johnston and Cleland, 1909
  • Plasmodium pericrocoti Chakravarty and Kar, 1945
  • Plasmodium ploceii Chakravarty and Kar, 1945
  • Plasmodium praecox var. muniae Das Gupta and Siddons, 1941
  • Plasmodium relictum biziurae Gilruth, Sweet and Dodd, 1910, emend. Garnham, 1966
  • Plasmodium relictum capistranoae Russell, 1932, emend. Garnham, 1966
  • Plasmodium relictum spheniscidae Fantham and Porter, 1944, emend. Garnham, 1966
  • Plasmodium relictum var. spheniscidae Fantham and Porter, 1944
  • Proteosoma biziurae Gilruth, Sweet and Dodd, 1910
  • Proteosoma grassii Labbé, 1894

Summary of Invasiveness

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P. relictum is a common mosquito-transmitted blood protozoan of wild birds that has a worldwide distribution. It has been reported from at least 411 avian species from 67 avian families and is considered to be relatively non-pathogenic and non-invasive under most circumstances and in most geographic areas. However, recent experimental studies have shown that it causes severe disease in some species of wild birds (Palinauskas et al., 2008, 2011). It was first reported as an invasive pathogen in the Hawaiian archipelago (Warner, 1968) and is currently recognized as a threat in New Zealand (Tompkins and Gleeson, 2006) and probably in the Galapagos Islands (Levin et al., 2009). It has been reported from Bermuda, the Northern Mariana Islands, the Cook Islands, the Society Islands, and the Marquesas Islands of the east-central Pacific (Beadell et al., 2006; Ishtiaq et al. 2006), but its potential threat to the avifauna of these island groups has not been investigated. The broad host range of the parasite and ability to use a wide range of mosquito species as vectors make it exceptionally invasive when introduced to new areas. Some lineages are currently restricted in their geographical range of transmission but might become invasive in future; for example GRW4, which has caused problems in Hawaii, is brought to Europe by migrating birds but not currently transmitted there (Palinauskas et al., 2007).

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Protista
  •         Phylum: Protozoa
  •             Subphylum: Apicomplexa
  •                 Order: Haemospororida
  •                     Family: Plasmodiidae
  •                         Genus: Plasmodium
  •                             Species: Plasmodium relictum

Distribution

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Reports of P. relictum that have accumulated over the past century are based mostly on morphology of erythrocytic stages. Earlier reports are based primarily on microscopy and may underestimate the true prevalence of the parasite in many host species (Fallon and Ricklefs, 2008), particularly since the morphological stages that are necessary for identification may be absent or in extremely low numbers in chronic, low intensity infections. Many recent studies have attempted to "type" parasites based on sequence of conserved mitochondrial DNA (Beadell et al., 2006; Marzal et al., 2011), and molecular markers for identification of this parasite have been developed (Beadell et al., 2006; Palinauskas et al., 2007), but are of low sensitivity for the detection of mixed infections of different Plasmodium spp. -- a combination of optical microscopy and PCR-based tools is recommended in studies of distribution of malaria parasites in wildlife (Valkiunas et al., 2006). In addition, most reports in migratory species do not distinguish where transmission of the parasites occurs. Transmission may take place either in breeding or wintering grounds, or in some cases in both locations (Perez-Tris and Bensch, 2005); lineage GRW4, the one that has caused problems for native Hawaiian birds, is present in northern Europe in birds that have migrated from Africa, but is not transmitted there (Palinauskas et al., 2007). Many early reports of P. relictum are not backed by museum voucher specimens and need to be verified by more detailed field studies. The patchy distribution by countries and host species is most likely a sampling artifact. The parasite has been reported in all zoogeographic regions of the world with the exception of Antarctica and is a relatively common parasite of passerine birds.

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

ArmeniaPresentNative Not invasive Karaseferyan, 1959
AzerbaijanPresentNative Not invasive Zeiniev, 1975
Georgia (Republic of)PresentNative Not invasive Burtikashvili, 1978
IndiaPresentNative Not invasive Beadell et al., 2006; Ishtiaq et al., 2007; Marzal et al., 2011Lineages GRW4, SGS1
IndonesiaPresentNative Not invasive McClure et al., 1978
IraqPresentNative Not invasive Shamsuddin and Mohammad, 1981
IsraelPresentNative Not invasive Bensch et al., 2000; Beadell et al., 2006; Martinsen et al., 2006; Marzal et al., 2011Lineages GRW4, SGS1
JapanPresentNative Not invasive Beadell et al., 2006; Ejiri et al., 2008Lineage GRW4
KazakhstanPresentNative Not invasive Yakunin and Zhazyltaev, 1977
Korea, Republic ofPresentNative Not invasive Beadell et al., 2006Lineages GRW4, SGS1
KyrgyzstanPresentNative Not invasive Kairullaev and Yakunin, 1982
MalaysiaPresentNative Not invasive Laird, 1998
MyanmarPresentNative Not invasive Beadell et al., 2006
PakistanPresentNative Not invasive Lari, 1959
PhilippinesPresentNative Not invasive McClure et al., 1978
SingaporePresentNative Not invasive Beadell et al., 2006
TaiwanPresentNative Not invasive Manwell et al., 1976
TajikistanPresentNative Not invasive Subkhonov, 1972
ThailandPresentNative Not invasive McClure et al., 1978
TurkeyPresentNative Not invasive Marzal et al., 2011Lineages SGS1 and GRW11

Africa

AlgeriaPresentNative Not invasive Sergent and Sergent, 1904
BotswanaPresentNative Not invasive Beadell et al., 2006
CameroonPresentNative Not invasive Beadell et al., 2006
ChadPresentNative Not invasive Williams et al., 1977
ComorosPresentNative Not invasive Beadell et al., 2006
EgyptPresentNative Not invasive MOHAMMED, 1958; Marzal et al., 2011Lineages SGS1 and GRW11
Equatorial GuineaPresentNative Not invasive Beadell et al., 2006
EthiopiaPresentNative Not invasive Ashford et al., 1976
GabonPresentNative Not invasive Beadell et al., 2006
KenyaPresentNative Not invasive Bennett and Herman, 1976; Bensch et al., 2000; Perkins and Schall, 2002Lineage GRW4
MadagascarPresentNative Not invasive Beadell et al., 2006Lineage GRW4
MauritiusPresentNative Not invasive Peirce et al., 1977; Beadell et al., 2006Lineage GRW4
MayottePresentNative Not invasive Beadell et al., 2006
NigeriaPresentNative Not invasive Waldenström et al., 2002; Beadell et al., 2006; Hellgren et al., 2007Lineages GRW4, SGS1
RéunionPresentNative Not invasive Peirce et al., 1977; Beadell et al., 2006Lineage GRW4
Rodriguez IslandPresentNative Not invasive Beadell et al., 2006Lineage GRW4
Sao Tome and PrincipePresentNative Not invasive Beadell et al., 2006
SenegalPresentBennett et al., 1978
SeychellesPresentNative Not invasive Beadell et al., 2006Lineage GRW4 on Praslin, other lineages on Fregete
South AfricaPresentNative Not invasive Beadell et al., 2006; Durrant et al., 2007Lineage GRW4
TanzaniaPresentNative Not invasive Bennett and Herman, 1976
UgandaPresentNative Not invasive Bennett et al., 1974b
ZambiaPresentNative Not invasive Peirce, 1984
ZimbabwePresentNative Not invasive Beadell et al., 2006

North America

BermudaPresentIntroducedBeadell et al., 2006; Marzal et al., 2011Lineage GRW4; speculation that it may have played a role in decline of Bermudan birds
CanadaPresentNative Not invasive Bennett et al., 1974a; Bishop and Bennett, 1992Records from Newfoundland and Saskatchewan; likely widespread throughout Canada
MexicoPresentNative Not invasive Beltran and Pardinas, 1953; Marzal et al., 2011
USAWidespreadNative Not invasive Beadell et al., 2006; Kimura et al., 2006; Marzal et al., 2011Multiple records from throughout continental U.S.
-HawaiiWidespreadIntroduced1920s? Invasive Laird and Van Riper, 1981; Beadell et al., 2006Lineage GRW4. Common in wet habitats at elevations below 1500m on main Hawaiian Islands; not present in NW Hawaiian Islands with possible exception of Midway Atoll.

Central America and Caribbean

Dominican RepublicPresentNative Not invasive Beadell et al., 2006Lineage GRW4
GrenadaPresentNative Not invasive Beadell et al., 2006Lineage GRW4
JamaicaPresentNative Not invasive Bennett et al., 1980
PanamaPresentNative Not invasive Sousa and Herman, 1982

South America

BrazilPresentNative Not invasive Marzal et al., 2011Lineage GRW4
ColombiaPresentNative Not invasive Bennett and Borrero, 1976
GuyanaPresentNative Not invasive Beadell et al., 2006
UruguayPresentNative Not invasive Beadell et al., 2006
VenezuelaPresentNative Not invasive Gabaldon and Ulloa, 1980

Europe

BelarusPresentNative Not invasive Dyl'ko, 1966; Beadell et al., 2006Lineage GRW4
BelgiumPresentNative Not invasive Hellgren et al., 2007Lineage SGS1
BulgariaPresentNative Not invasive Shurulinkov and Golemansky, 2003; Hellgren et al., 2007; Zehtindjiev et al., 2008LIneages GRW4, SGS1
Czech RepublicPresentNative Not invasive Votýpka et al., 2003
Czechoslovakia (former)PresentNative Not invasive Kucera, 1978
FrancePresentNative Not invasive Beadell et al., 2006; Bonneaud et al., 2006; Loiseau et al., 2008; Marzal et al., 2011Lineage SGS1
GermanyPresentNative Not invasive Beadell et al., 2006
GreecePresentNative Not invasive PAPADAKIS, 1935
HungaryPresentNative Not invasive Szöllosi et al., 2009Lineages GRW4, SGS1
ItalyPresentNative Not invasive Corradetti, 1970; Beadell et al., 2006; Hellgren et al., 2007; Marzal et al., 2011Lineages GRW4, SGS1 and GRW11
LithuaniaPresentNative Not invasive Hellgren et al., 2007; Palinauskas et al., 2007; Marzal et al., 2011Lineages SGS1, GRW11
NorwayPresentNative Not invasive Beadell et al., 2006; Marzal et al., 2011Lineages GRW4, SGS1
PolandPresentSulgostowska and Czaplinska, 1987
Russian FederationPresentNative Not invasive Kobyshev et al., 1975; Beadell et al., 2006; Zehtindjiev et al., 2009; Marzal et al., 2011Multiple reports from throughout Russia, including lineages GRW4, SGS1 and GRW11
SpainPresentNative Not invasive Orbaneja Aguero S de, 1934; Beadell et al., 2006; Hellgren et al., 2007; Marzal et al., 2008; Marzal et al., 2011Lineages GRW4, SGS1 and GRW11
SwedenPresentNative Not invasive Hellgren, 2005; Beadell et al., 2006; Bensch et al., 2007; Yohannes et al., 2008; Marzal et al., 2011Lineages GRW4, SGS1
UKPresentNative Not invasive Beadell et al., 2006; Wood et al., 2007; Cosgrove et al., 2008Lineages SGS1, GRW11
UkrainePresentNative Not invasive Gluschchenko, 1963; Beadell et al., 2006; Hellgren et al., 2007; Marzal et al., 2011Lineages GRW4, SGS1

Oceania

AustraliaPresentNative Not invasive Beadell et al., 2006Lineage GRW4
Cook IslandsPresentIntroduced1922?Ishtiaq et al., 2007Single report from introduced Common Myna (Acridotheres tristis) based on PCR diagnostics; distribution in indigenous birds unknown
French PolynesiaPresentIntroducedBeadell et al., 2006Lineage GRW4, single report based on PCR diagnostics from introduced birds on Moorea (Society Islands) and from indigenous birds on Nuku Hiva (Marquesas Islands)
New ZealandWidespreadIntroduced Invasive Beadell et al., 2006; Tompkins and Gleeson, 2006; Marzal et al., 2011Widespread in introduced passerines on North and South Islands. Lineage SGS1.
Northern Mariana IslandsPresentBeadell et al., 2006Single report based on PCR diagnostics from the indigenous Rufous Fantail (Rhipidura rufifrons). Not clear if it was introduced with non-native passerines.
Papua New GuineaPresentNative Not invasive Beadell et al., 2006
Solomon IslandsPresentNative Not invasive Laird, 1960

History of Introduction and Spread

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P. relictum was first detected in the Hawaiian archipelago in the 1930s in a Red-billed Leiothrix (Leiothrix lutea) and a Japanese White Eye (Zosterops japonicus) that were collected in Hawaii Volcanoes National Park on the Island of Hawaii (Baldwin, 1941; Fisher and Baldwin 1947). The parasite was subsequently reported in non-native forest birds on Kauai in the 1950s (Warner, 1968) and in native species on the island of Hawaii in the 1970s (van Riper et al., 1986). The precise date, point of entry, and details about how it spread in the islands are not known. Warner (1968) and van Riper et al. (1986) have argued that the parasite was introduced inadvertently during the early 1900s when passerine birds from Southeast Asia, North and South America, and Africa were introduced to replace native birds that were disappearing from the lowlands as a result of epizootics of introduced Avipoxvirus, habitat destruction, and predation from introduced cats, rats, and mongooses. The successful spread of the parasite was dependent on the introduction and spread of the mosquito vector, Culex quinquefasciatus, in forested habitats throughout the islands during the nineteenth century (Warner, 1968). While it is possible that P. relictum first reached the islands via migratory birds, absence of a suitable vector prior to the establishment of mosquitoes would have prevented spread to endemic forest birds. The parasite is currently widespread throughout the main Hawaii Islands of Kauai, Oahu, Molokai, Maui, and Hawaii at elevations below 1500 m. It is likely to be present on Lanai and Kahoolawe, but these areas have not been surveyed for avian diseases. Plasmodium relictum is likely to be absent from the low atolls and rocky islets of the Northwestern Hawaiian Islands, with the possible exception of Midway Atoll, where introduced mosquitoes and canaries (Serinus canaria) could maintain transmission of the parasite.

Little is known about the history of the parasite in other isolated island archipelagoes in the Pacific Basin. The importation and release of the Common Myna (Acridotheres tristis) in eastern and central Polynesia may have led to introduction of P. relictum to French Polynesia and the Cook Islands (Beadell et al., 2006; Ishtiaq et al., 2006), but natural occurrence of P. relictum in migratory birds in the region (Laird, 1960) and occurrence of indigenous mosquito vectors that could support transmission of the parasites throughout the tropical Pacific make it possible that the parasite is indigenous (Atkinson et al., 2006). However, presence of identical lineages of the parasite (GRW4) of apparent Old World origin in these areas makes it more likely that the parasites were introduced (Beadell et al., 2006).

An unidentified species of avian malaria currently threatens the endemic avifauna of the Galapagos Islands. Culex quinquefasciatus has recently become established in the archipelago (Whiteman et al., 2005) and a recent, unconfirmed report of a Plasmodium species in Galapagos Penguins (Levin et al., 2009) raises the possibility that malaria may spread throughout the archipelago as it did in the Hawaiian Islands.

P. relictum was first reported from New Zealand in the early 1900s (Dore, 1920; Laird, 1950), but it is not clear whether it is indigenous or whether it was introduced when non-native House Sparrows (Passer domesticus), Blackbirds (Turdus merulus), Song Thrush (Turdus philomelos), Chaffinch (Fringilla coelebs), or Starlings (Sturnus vulgaris) were imported and released (Tompkins and Gleeson, 2006). Prevalence and distribution of the parasite are currently expanding southward in New Zealand as the range of the introduced mosquito, Culex quinquefasciatus, increases. While there were only a few historic reports of the parasite in native New Zealand birds (Dore,1920; Laird, 1950), there are increasing reports of morbidity and mortality among native species, particularly in zoos and captive propagation facilities (Tompkins and Gleeson, 2006; Alley et al., 2008; Derraik et al., 2008; Hale, 2008).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Cook Islands New Zealand 1920s Biological control (pathway cause) Yes No Ishtiaq et al. (2006) Accidental via Acridotheres tristis
French Polynesia   Biological control (pathway cause) ,
Pet trade (pathway cause)
Yes No Beadell et al. (2006) Possible accidental introduction via Acridotheres tristis, Zosterops sp. or Neochmia temporalis
Hawaii Africa 1900-1930 Pet trade (pathway cause) Yes No Warner (1968) Accidental in cage birds, undetermined species
New Zealand India 1895 Biological control (pathway cause) Yes No Ishtiaq et al. (2006) Unclear if P. relictum is indigenous or introduced with Acridotheres tristis or another passerine

Risk of Introduction

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Broad host range and laboratory evidence that a wide range of mosquito species can transmit this parasite (Huff, 1965) make risk of accidental introduction of P. relictum to new areas high. Most documented introductions to date have followed importation and release of naturally infected cage birds, although movement of infected mosquitoes through commercial air, land, and sea traffic may also be an important mechanism for dispersal (LaPointe, 2007).

Pathogen Characteristics

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Species of five different subgenera of Plasmodium infect birds. They are distinguished by morphological characteristics of the erythrocytic stages of the parasites, by morphological changes in their host cells, and by their preference for either mature erythrocytes or erythrocyte precursors (Valkiunas, 2005). Species that infect both wild and domestic birds are separated further by host range, developmental characteristics and vectors. Plasmodium relictum is currently placed in the subgenus Haemamoeba, which includes parasites with round gametocytes and meronts that exceed the host erythrocyte nucleus in size. The taxonomy of this group of parasites is currently in a state of flux, as more traditional morphological methods for distinguishing species are being challenged by new molecular data about the genetic diversity and host range of this group of organisms. One recent study has documented at least 38 different lineages of this parasite based on morphological characteristics and the sequence of conserved mitochondrial DNA (Beadell et al., 2006). One of these, lineage GRW4, has had significant effects on endemic Hawaiian birds and may threaten endemic and indigenous birds on other isolated island archipelagoes in the Pacific Basin, where it appears to have been introduced with non-native passerines. This lineage appears to be indigenous among island groups in the Indian Ocean off the eastern coast of Africa. Like some others, it is restricted in its geographical range of transmission (Marzal et al., 2011). Transmission of this lineage has not been reported in northern Europe where it is present in adult birds after their arrival from African wintering grounds, but is absent from juvenile birds; it might become invasive in the future (Palinauskas et al., 2007).

The conservation significance of other lineages of P. relictum is unknown. While it is possible that P. relictum may actually comprise a group of cryptic species that share many morphological traits, this question is currently unresolved and will ultimately depend on more detailed studies to relate genetic haplotypes with morphological and life history characteristics of these intracellular protozoans.

Hosts/Species Affected

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P. relictum is primarily a parasite of passerine birds, although species in at least 16 additional avian orders are susceptible to infection. Both penguins (Spheniscidae) and Hawaiian Honeycreepers (Drepanidinae) are particularly susceptible to infection with P. relictum, but the physiological basis for this high susceptibility is not understood. Reports of pathogenicity from infections with P. relictum are more common in zoo settings, where hosts are moved outside of their normal range and exposed to local vectors and transmission of the parasite. The impact of P.relictum on wild birds is poorly known in natural ecosystems, but experimental data show that lineage SGS1 is virulent and might cause severe disease and even mortality in wild birds, particularly during co-infections with other species of Plasmodium (Palinauskas et al., 2008; Palinauskas et al., 2011). Infection with the same lineage of P. relictum causes diseases of different severity in different host species (Palinauskas et al., 2008); this should be taken into consideration in conservation projects.

Climate

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ClimateStatusDescriptionRemark
A - Tropical/Megathermal climate Preferred Average temp. of coolest month > 18°C, > 1500mm precipitation annually
C - Temperate/Mesothermal climate Preferred Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
66 45

Air Temperature

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Parameter Lower limit Upper limit
Mean maximum temperature of hottest month (ºC) 14

Vectors and Intermediate Hosts

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VectorSourceReferenceGroupDistribution
Culex quinquefasciatusInsectWorld
Culex stigmatosomaInsectCalifornia
Culex tarsalisInsectCalifornia

Environmental Impact

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P. relictum has had environmental impacts in the Hawaiian archipelago where it has contributed substantially to loss of biodiversity in native forest bird communities on the main Hawaiian Islands. Loss of ecosystem services such as pollination and seed dispersal have been compensated in part by replacement of native forest birds with introduced species.

Impact of P.relictum on wild birds more generally is poorly investigated in natural ecosystems, but experimental data show that it is virulent and might cause severe disease and even mortality in some wild birds (Palinauskas et al., 2011).

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Charadrius obscurusEN (IUCN red list: Endangered) EN (IUCN red list: Endangered)New ZealandParasitism (incl. parasitoid); PathogenicReed, 1997
Chasiempis sandwichensisVU (IUCN red list: Vulnerable) VU (IUCN red list: Vulnerable)HawaiiParasitism (incl. parasitoid); PathogenicVanderwerf et al., 2006b
Dysmorodrepanis munroi (Akiapolaau)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered)HawaiiParasitism (incl. parasitoid); PathogenicAtkinson and LaPointe, 2009
Eudyptes chrysocome (southern rockhopper penguin)VU (IUCN red list: Vulnerable) VU (IUCN red list: Vulnerable); USA ESA listing as threatened species USA ESA listing as threatened speciesNew ZealandParasitism (incl. parasitoid); PathogenicSturrock and Tompkins, 2007
Hemignathus lucidus affinisCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)HawaiiParasitism (incl. parasitoid); PathogenicAtkinson and LaPointe, 2009
Hemignathus lucidus hanapepeCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)HawaiiParasitism (incl. parasitoid); PathogenicAtkinson and LaPointe, 2009
Loxioides bailleui (palila)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiParasitism (incl. parasitoid); PathogenicUS Fish and Wildlife Service, 2006; Atkinson and LaPointe, 2009
Loxops caeruleirostris (akekee)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiParasitism (incl. parasitoid); PathogenicAtkinson and LaPointe, 2009
Loxops coccineus coccineusEN (IUCN red list: Endangered) EN (IUCN red list: Endangered)HawaiiParasitism (incl. parasitoid); PathogenicAtkinson and LaPointe, 2009
Loxops coccineus ochraceusEN (IUCN red list: Endangered) EN (IUCN red list: Endangered)HawaiiParasitism (incl. parasitoid); PathogenicAtkinson and LaPointe, 2009
Melamprosops phaeosomaCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)HawaiiParasitism (incl. parasitoid); PathogenicAtkinson and LaPointe, 2009
Mohoua ochrocephala (Yellowhead)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered)New ZealandParasitism (incl. parasitoid); PathogenicDerraik et al., 2008
Myadestes lanaiensisCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)HawaiiParasitism (incl. parasitoid); PathogenicAtkinson et al., 2001b
Myadestes myadestinusEX (IUCN red list: Extinct) EX (IUCN red list: Extinct)HawaiiParasitism (incl. parasitoid); PathogenicAtkinson et al., 2001b
Myadestes obscurusVU (IUCN red list: Vulnerable) VU (IUCN red list: Vulnerable)HawaiiParasitism (incl. parasitoid); PathogenicAtkinson et al., 2001b
Myadestes palmeri (Puaiohi)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)HawaiiParasitism (incl. parasitoid); PathogenicAtkinson et al., 2001b
Oreomystis bairdi (akikiki)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiParasitism (incl. parasitoid); PathogenicUS Fish and Wildlife Service, 2006; Atkinson and LaPointe, 2009
Oreomystis mana (Hawaii creeper)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiParasitism (incl. parasitoid); PathogenicUS Fish and Wildlife Service, 2006; Atkinson and LaPointe, 2009
Palmeria dolei (crested honeycreeper)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiEcosystem change / habitat alteration; Parasitism (incl. parasitoid); PathogenicAtkinson and LaPointe, 2009; US Fish and Wildlife Service, 2011a
Paroreomyza maculata (Oahu creeper)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiParasitism (incl. parasitoid); PathogenicUS Fish and Wildlife Service, 2006; Atkinson and LaPointe, 2009
Philesturnus carunculatusNT (IUCN red list: Near threatened) NT (IUCN red list: Near threatened)New ZealandParasitism (incl. parasitoid); PathogenicHale, 2008
Pseudonestor xanthophrys (Maui parrotbill)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); National list(s) National list(s); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiParasitism (incl. parasitoid); PathogenicUS Fish and Wildlife Service, 2006; Atkinson and LaPointe, 2009; US Fish and Wildlife Service, 2011c
Psittirostra psittacea (Ou)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiParasitism (incl. parasitoid); PathogenicUS Fish and Wildlife Service, 2006; Atkinson and LaPointe, 2009; US Fish and Wildlife Service, 2009
Spheniscus mendiculus (Galapagos Penguin)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered)Galapagos IslandsParasitism (incl. parasitoid); PathogenicLevin et al., 2009
Megadyptes antipodes (yellow-eyed penguin)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); USA ESA listing as threatened species USA ESA listing as threatened speciesNew ZealandPathogenicUS Fish and Wildlife Service, 2010
Paroreomyza flammea (Molokai creeper)EX (IUCN red list: Extinct) EX (IUCN red list: Extinct); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiPathogenicUS Fish and Wildlife Service, 2006
Pterodroma sandwichensisVU (IUCN red list: Vulnerable) VU (IUCN red list: Vulnerable)HawaiiPathogenicUS Fish and Wildlife Service, 2011b
Zosterops rotensis (rota bridled white-eye)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesNorthern Mariana IslandsPathogenicUS Fish and Wildlife Service, 2007

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Reproduces asexually
  • Has high genetic variability
Impact outcomes
  • Changed gene pool/ selective loss of genotypes
  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Host damage
  • Negatively impacts animal health
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
Impact mechanisms
  • Interaction with other invasive species
  • Parasitism (incl. parasitoid)
  • Pathogenic
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field
  • Difficult/costly to control

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Links to Websites

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WebsiteURLComment
Global Invasive Species Databasehttp://www.issg.org/databaseThe GISD aims to increase awareness about invasive alien species and to facilitate effective prevention and management. It is managed by the Invasive Species Specialist Group (ISSG) of the Species Survival Commission.
Hawaiian Ecosystems at Risk Project (HEAR)http://www.hear.org/

Organizations

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USA: U.S. Geological Survey, Pacific Island Ecosystems Research Center, PO Box 44, Hawaii National Park, Hawaii 96718, http://biology.usgs.gov/pierc/

Contributors

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03/11/09 Original text by:

Carter Atkinson, USGS Pacific Island Ecosystems Research Center, Kilauea Field Station, P.O. Box 44, Bldg 343, Hawaii National Park, HI 96718, USA

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