Meloidogyne arenaria (peanut root-knot nematode)

Pictures

Picture

Title

Caption

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Symptoms on roots

Meloidogyne arenaria (peanut root-knot nematode); galled root system with three females dissected free from root tissue.

©J.D. Eisenback

Symptoms on roots

Meloidogyne arenaria (peanut root-knot nematode); galled root system of tobacco.

©J.D. Eisenback

Damage to root system

Meloidogyne arenaria (peanut root-knot nematode); washed infected root system. The formation and function of the root system is abnormal in infected plants.

©Ed McGawley

Symptoms on pods

Meloidogyne arenaria (peanut root-knot nematode); damage symptoms on infected groundnut pods.

©Pat Phipps

Light micrographs

Meloidogyne arenaria (peanut root-knot nematode); detail of anterior end of female (left) and male (right).

©J.D. Eisenback

Male and female stylets

Meloidogyne arenaria (peanut root-knot nematode); scanning electron micrographs of extracted stylet of female (top) and male (bottom).

©J.D. Eisenback

Nematodes invading root tip

Meloidogyne arenaria (peanut root-knot nematode); light micrograph of second-stage juveniles (stained red) invading root tip.

©J.D. Eisenback

Female with egg mass

Meloidogyne arenaria (peanut root-knot nematode); light micrograph of female with egg mass in a galled root (stained red).

©J.D. Eisenback

SEM of female

Meloidogyne arenaria (peanut root-knot nematode); scanning electron micrograph of whole female.

©J.D. Eisenback

Perineal patterns

Meloidogyne arenaria (peanut root-knot nematode); left) light micrograph of perineal pattern of female. Right) light micrograph of perineal pattern.

©J.D. Eisenback

Identity

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Preferred Scientific Name

Meloidogyne arenaria (Neal, 1889) Chitwood, 1949

Preferred Common Name

peanut root-knot nematode

Other Scientific Names

Anguillula arenaria Neal, 1889
Heterodera arenaria (Neal, 1889) Marcinowski, 1909
Meloidogyne arenaria arenaria (Neal, 1889) Chitwood, 1949
Meloidogyne arenaria thamesi Chitwood in Chitwood et al., 1952
Meloidogyne thamesi (Chitwood et al., 1952) Goodey, 1963
Tylenchus arenarius (Neal, 1889) Cobb, 1890

International Common Names

English: groundnut root knot nematode; root-knot nematode disease
Spanish: nematodo nodulador del cacahuete
French: nématode galligène de l'arachide; nodosite des racines

Local Common Names

Germany: Erdnusswurzelgallen-Aelchen
Japan: Nekobu-sentyubyo

EPPO code

MELGAR (Meloidogyne arenaria)
MELGTH (Meloidogyne thamesi)

Taxonomic Tree

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Domain: Eukaryota
Kingdom: Metazoa
Phylum: Nematoda
Family: Meloidogynidae
Genus: Meloidogyne
Species: Meloidogyne arenaria

Notes on Taxonomy and Nomenclature

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The taxonomy of the peanut root-knot nematode was greatly clarified by Chitwood in 1949; however, one morphological variant that he described has caused confusion in the literature because it has been cited as either subspecies M. arenaria thamesi or elevated to species M. thamesi. Because this variant has been extremely difficult to identify and does not correlate with other taxonomic characters, the two rankings are considered as synonyms of M. arenaria (Eisenback and Triantaphyllou, 1991). The morphological variation of this species has been described for seven populations representing the two host races, as well as the two cytological races (Cliff and Hirschmann, 1985).

Two host races of M. arenaria have been recognized: race 1 infects and reproduces on groundnut, whereas populations of race 2 do not (Taylor and Sasser, 1978). It is ironic that many populations of the nematode with the common name peanut root-knot nematode do not infect peanut.

Two cytological races of M. arenaria have been described (Triantaphyllou, 1963, 1979). The most common populations belong to race A and are triploid (3n = 51-56). Race B populations are less common and are diploid (2n = 34-37). There is no correlation between host race and cytological race.

Description

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The body of the female is pearly white and pear-shaped, 500-1000 µm long by 400-600 µm wide. The conical neck of the female is in line with the spherical portion of the body. The stylet is robust, 13-17 (16) µm long and characteristically shaped with large, posteriorly sloped, tear-drop-shaped knobs. The distance of the dorsal oesophageal gland orifice to the base of the stylet is comparatively long (3-7 (5) µm). The overall morphology of females of M. arenaria is similar to other species within the genus.

The perineal pattern may be characteristic for the species, but some populations may contain individual variants that restrict the usefulness of this character. Likewise, other species of Meloidogyne may also have perineal patterns that closely resemble that of M. arenaria. The perineal pattern of M. arenaria may be very similar to that of M. incognita and other Meloidogyne species. Patterns that contain short, lateral incisures resemble that of M. javanica, and patterns that are rounded to hexagonal, often containing wings, are like that of M. hapla. The perineal pattern of M. arenaria has a low and rounded dorsal arch, but in some individuals it may be high and squarish. The striae are coarse and smooth to wavy, and some striae may bend toward the vulva. The most useful character of the perineal pattern is the lines in the lateral areas of the dorsal arch that sharply curve toward the tail terminus and meet the ventral striae at an angle. These striae become forked and the distance between them increases near the lateral areas which are often demarcated, but not delineated by distinct lateral incisures. Very short lateral incisures may be present very near the tail terminus. Some perineal patterns of M. arenaria form one or two 'wings' that extend laterally and are marked by fusion of the striae in the dorsal and ventral arches.

Males of M. arenaria are long (0.9-2.3 mm) and narrow (27-48 µm). Although the shape of the head is a useful morphological character, it is similar to several other less common species. The labial disc and medial lips form a smooth, posteriorly sloping head cap. The head annule is smooth and usually not marked with additional head annulations. Both the head annule and the body annulations are in the same contour. The stylet is long (20-28 µm) and robust with a bluntly pointed tip. The wide, cylindrical shaft gradually merges with the large, rounded, slightly tear-drop-shaped knobs. The dorsal oesophageal gland orifice to the base of the stylet is long to very long (4-8) 6 µm. The overall morphology of males of M. arenaria is similar to other species within the genus.

Second-stage juveniles of M. arenaria are long (398-605 (504) µm) and slender (13-18 (15) µm). The tail is moderately long (44-69 (56) µm), and the poorly defined hyaline tail terminus is moderately long (6-13 (9) µm), with a finely rounded to pointed tip. The stylet of second-stage juveniles is moderately long (10-12 (11) µm), and the dorsal oesophageal gland orifice distance to the stylet base is moderately long (3-5 (4) µm). The morphology of the second-stage juvenile of M. arenaria is similar to many other species of Meloidogyne and requires critical evaluation by light and scanning electron microscopy in order to be differentiated.

Distribution

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M. arenaria is widely distributed around the world in tropical, subtropical and temperate climates where the average temperature in the warmest month is 36°C or lower and the average temperature in the coldest month is at lowest 3°C. The principal limiting factor in the distribution of M. arenaria seems to be an average temperature in the coldest month of 3°C (Taylor et al., 1982). This species seems to be rare or absent in areas with an average annual temperature of less than 15°C and most common in climates with average annual temperatures of 18-27°C (Taylor et al., 1982). This root-knot nematode is most common where the annual precipitation averages 1000-2000 mm (Taylor et al., 1982).

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.

Last updated: 25 Feb 2021

Continent/Country/Region	Distribution	Reference	Notes
Africa
Algeria	Present, Few occurrences	CABI and EPPO (2003)
Côte d'Ivoire	Present	CABI and EPPO (2003) CABI (Undated)
Egypt	Present	Taylor et al. (1982) CABI and EPPO (2003) Ibrahim et al. (2010)
Ethiopia	Present	Awol Seid et al. (2019)
Gambia	Present, Widespread	CABI and EPPO (2003) CABI (Undated)
Ghana	Present, Widespread	CABI and EPPO (2003) CABI (Undated)
Liberia	Present, Few occurrences	CABI and EPPO (2003)
Libya	Present	CABI and EPPO (2003) CABI (Undated)
Madagascar	Present	CABI and EPPO (2003)
Malawi	Present	CABI and EPPO (2003) CABI (Undated)
Mauritius	Present	CABI and EPPO (2003)
Morocco	Present	Taylor et al. (1982) CABI and EPPO (2003)
Mozambique	Present, Widespread	Martin and Armstrong (1975) CABI and EPPO (2003)
Nigeria	Present	CABI and EPPO (2003) CABI (Undated)
São Tomé and Príncipe	Present	CABI and EPPO (2003)
Senegal	Present	CABI and EPPO (2003) CABI (Undated)
South Africa	Present	Wyk (1985) CABI and EPPO (2003)
Sudan	Present	CABI and EPPO (2003) CABI (Undated)
Tanzania	Present	CABI and EPPO (2003)
Tunisia	Present	Chihani-Hammas et al. (2018)
Uganda	Present	CABI and EPPO (2003)
Zimbabwe	Present	CABI and EPPO (2003)
Asia
Armenia	Present	CABI and EPPO (2003)
Azerbaijan	Present	Treskova et al. (1979) CABI and EPPO (2003)
Bangladesh	Present	Choudhury (1981) CABI and EPPO (2003)
China	Present	Shaoshing and Ziming (1991) CABI and EPPO (2003) Wang et al. (2012) Xu et al. (2012) Fu et al. (2013) Wang XiZhuo et al. (2013) Zeng JianMin et al. (2018)
-Anhui	Present	CABI and EPPO (2003) Wang et al. (2012) Wang XiZhuo et al. (2013)
-Fujian	Present	Shaoshing and Ziming (1991) CABI and EPPO (2003)
-Guangdong	Present	CABI and EPPO (2003) Xu et al. (2012)
-Hainan	Present	CABI and EPPO (2003) Fu et al. (2013)
-Hebei	Present	CABI and EPPO (2003)
-Henan	Present	Yang (1984) CABI and EPPO (2003)
-Hunan	Present	CABI and EPPO (2003)
-Inner Mongolia	Present	CABI and EPPO (2003)
-Jiangsu	Present	Sun et al. (1991) CABI and EPPO (2003)
-Shaanxi	Present	Mao Qi et al. (2007)
-Shandong	Present	CABI and EPPO (2003)
-Shanxi	Present	CABI and EPPO (2003)
-Yunnan	Present	CABI and EPPO (2003) Zeng JianMin et al. (2018)
-Zhejiang	Present	Huan (1983) CABI and EPPO (2003)
India	Present	CABI and EPPO (2003) Poornima et al. (2017) CABI (Undated)
-Andhra Pradesh	Present	CABI and EPPO (2003)
-Chhattisgarh	Present	Sahu et al. (2011)
-Delhi	Present	CABI and EPPO (2003)
-Gujarat	Present	CABI and EPPO (2003)
-Haryana	Present	CABI and EPPO (2003)
-Karnataka	Present	CABI and EPPO (2003)
-Madhya Pradesh	Present	CABI and EPPO (2003)
-Maharashtra	Present	CABI and EPPO (2003)
-Punjab	Present	CABI and EPPO (2003) Hussain et al. (2015) Jabbar et al. (2016)
-Tamil Nadu	Present	Kumar et al. (1987) CABI and EPPO (2003) Poornima et al. (2017)
-Uttar Pradesh	Present	Abdullaeva (1986) CABI and EPPO (2003) CABI (Undated)
-West Bengal	Present	Sen and Dasgupta (1979) CABI and EPPO (2003)
Indonesia	Present	CABI and EPPO (2003)
-Java	Present	Hadisoeganda (1981) CABI and EPPO (2003)
-Sumatra	Present, Widespread	Hadisoeganda (1981) CABI and EPPO (2003)
Iran	Present	Taylor et al. (1982) CABI and EPPO (2003)
Iraq	Present	CABI and EPPO (2003) CABI (Undated)
Japan	Present	Taylor et al. (1982) CABI and EPPO (2003)
-Honshu	Present	CABI and EPPO (2003)
-Kyushu	Present	CABI and EPPO (2003)
-Ryukyu Islands	Present	CABI and EPPO (2003)
-Shikoku	Present	CABI and EPPO (2003)
Jordan	Present	Karajeh et al. (2005)
Lebanon	Present	CABI and EPPO (2003) CABI (Undated)
Malaysia	Present	CABI and EPPO (2003) CABI (Undated)
Maldives	Present, Widespread	CABI (Undated)	Original citation: Kozhokaru et al., 1979
Mongolia	Present	CABI and EPPO (2003)
Nepal	Present	Bhardwaj and Högger (1984) CABI and EPPO (2003)
North Korea	Present, Widespread	Choi (1981) CABI and EPPO (2003)
Pakistan	Present	CABI and EPPO (2003) Tariq et al. (2007) Jiskani et al. (2009) Hussain et al. (2015) Jabbar et al. (2016) CABI (Undated)
Philippines	Present	Taylor et al. (1982) CABI and EPPO (2003)
Saudi Arabia	Present	Ibrahim and Al-Yahya (2002) El-Sherbiny (2011)
South Korea	Present	Choi (1981) CABI and EPPO (2003)
Sri Lanka	Present	CABI and EPPO (2003) CABI (Undated)
Syria	Present, Widespread	Tayar (1982) CABI and EPPO (2003)
Taiwan	Present	Tsai (1981) CABI and EPPO (2003)
Tajikistan	Present	Kulichin (1981) CABI and EPPO (2003)
Thailand	Present, Widespread	Taylor et al. (1982) CABI and EPPO (2003)
Turkey	Present	Taylor et al. (1982) CABI and EPPO (2003) Akgul et al. (2010) Mennan et al. (2011) Akyazİ et al. (2012) Akyazİ et al. (2017) Aydınlı (2018)
Turkmenistan	Present	CABI and EPPO (2003) CABI (Undated)
Uzbekistan	Present	Abdullaeva (1986) CABI and EPPO (2003)
Vietnam	Present	CABI and EPPO (2003)
Europe
Belgium	Present	BGARC (1972) CABI and EPPO (2003)
Bulgaria	Present	Stoyanov (1980) CABI and EPPO (2003)
Federal Republic of Yugoslavia	Present	Grujicic (1975)
France	Present	CABI and EPPO (2003) Quénéhervé et al. (2011) CABI (Undated)
Germany	Present	Zunke (1981) CABI and EPPO (2003)
Greece	Present	CABI and EPPO (2003) Tzortzakakis et al. (2011) CABI (Undated)
Hungary	Present	Budai (1980) CABI and EPPO (2003)
Iceland	Present, Localized	CABI and EPPO (2003) CABI (Undated)
Ireland	Present	CABI and EPPO (2003)
Italy	Present	CABI and EPPO (2003) CABI (Undated)
-Sicily	Present	CABI (Undated)	Original citation: d'Errico and Ingenito (2003)
Moldova	Present	CABI and EPPO (2003)
Montenegro	Present	Pajovic et al. (2007)
Netherlands	Present	Brinkman (1975) CABI and EPPO (2003)
North Macedonia	Present	CABI and EPPO (2003)
Poland	Present, Localized	CABI and EPPO (2003)
Portugal	Present	Taylor et al. (1982) CABI and EPPO (2003)
Romania	Present, Localized	Romascu et al. (1974) CABI and EPPO (2003)
Russia	Present	Mar'enko (1984) CABI and EPPO (2003)
-Central Russia	Present	CABI and EPPO (2003)
-Eastern Siberia	Present, Localized	CABI and EPPO (2003)
Spain	Present	CABI and EPPO (2003) Castillo et al. (2001) Castillo et al. (2006) López-Pérez et al. (2011) CABI (Undated)
-Canary Islands	Present	Taylor et al. (1982) CABI and EPPO (2003)
Switzerland	Present	Vallotton (1981) CABI and EPPO (2003)
Ukraine	Present	Volodchenko (1975) CABI and EPPO (2003)
United Kingdom	Present	CABI and EPPO (2003)
North America
Belize	Present	Taylor et al. (1982) CABI and EPPO (2003)
Bermuda	Present	Taylor et al. (1982) CABI and EPPO (2003) CABI (Undated)
Costa Rica	Present	Lopez (1984) CABI and EPPO (2003)
Cuba	Present	CABI and EPPO (2003) CABI (Undated)
El Salvador	Present	CABI and EPPO (2003) Villain et al. (2013)
Guadeloupe	Present	Taylor et al. (1982) CABI and EPPO (2003)
Guatemala	Present	Villain et al. (2013)
Jamaica	Present, Widespread	CABI and EPPO (2003) CABI (Undated)
Martinique	Present, Widespread	CABI and EPPO (2003)
Mexico	Present	CABI and EPPO (2003) Martínez-Gallardo et al. (2019)
Puerto Rico	Present	Taylor et al. (1982) CABI and EPPO (2003)
Trinidad and Tobago	Present	Schotman (1989) CABI and EPPO (2003)
United States	Present	CABI and EPPO (2003) Johnson et al. (1996) Church and Rosskopf (2005) Kaur et al. (2007) Kokalis-Burelle et al. (2012)
-Alabama	Present	Walters and Barker (1994) CABI and EPPO (2003)
-Arizona	Present, Widespread	Walters and Barker (1994) CABI and EPPO (2003)
-Arkansas	Present, Widespread	Walters and Barker (1994) CABI and EPPO (2003)
-California	Present	Walters and Barker (1994) CABI and EPPO (2003)
-Florida	Present	Walters and Barker (1994) CABI and EPPO (2003) Church and Rosskopf (2005) Kaur et al. (2007) Kokalis-Burelle et al. (2012)
-Georgia	Present	Walters and Barker (1994) Johnson et al. (1996) CABI and EPPO (2003)
-Hawaii	Present, Widespread	Walters and Barker (1994) CABI and EPPO (2003)
-Kansas	Present	Walters and Barker (1994) CABI and EPPO (2003)
-Louisiana	Present, Widespread	Walters and Barker (1994) CABI and EPPO (2003)
-Mississippi	Present, Widespread	Walters and Barker (1994) CABI and EPPO (2003)
-New Jersey	Present	Walters and Barker (1994) CABI and EPPO (2003)
-New York	Present	Walters and Barker (1994) CABI and EPPO (2003)
-North Carolina	Present	Walters and Barker (1994) CABI and EPPO (2003)
-Oklahoma	Present, Widespread	Walters and Barker (1994) CABI and EPPO (2003)
-South Carolina	Present	Walters and Barker (1994) CABI and EPPO (2003)
-Texas	Present	Walters and Barker (1994) CABI and EPPO (2003)
-Virginia	Present	Walters and Barker (1994) CABI and EPPO (2003)
-West Virginia	Present, Widespread	Walters and Barker (1994) CABI and EPPO (2003)
Oceania
Australia	Present	Taylor et al. (1982) CABI and EPPO (2003)
-New South Wales	Present	CABI and EPPO (2003)
-Queensland	Present	CABI and EPPO (2003)
-South Australia	Present	CABI and EPPO (2003)
-Tasmania	Present	Pethybridge et al. (2008)
-Western Australia	Present	CABI and EPPO (2003)
Cook Islands	Present	CABI and EPPO (2003)
Fiji	Present	Taylor et al. (1982) CABI and EPPO (2003)
Niue	Present	CABI and EPPO (2003)
Papua New Guinea	Present	Bridge and Page (1984) CABI and EPPO (2003)
Samoa	Present	CABI and EPPO (2003)
Solomon Islands	Present	CABI and EPPO (2003)
Tuvalu	Present	CABI and EPPO (2003)
South America
Argentina	Present	Taylor et al. (1982) CABI and EPPO (2003)
Bolivia	Present	Taylor et al. (1982) CABI and EPPO (2003)
Brazil	Present	Taylor et al. (1982) CABI and EPPO (2003) Lima et al. (2005) Moura et al. (2010) Araújo Filho et al. (2012) Lima et al. (2013) Machado et al. (2013) Bellé et al. (2016) Kaspary et al. (2017) Neves et al. (2017)
-Alagoas	Present	Lima et al. (2013)
-Amazonas	Present, Widespread	Lemos and Ponte (1978) CABI and EPPO (2003)
-Bahia	Present	Freire and Ponte (1976) CABI and EPPO (2003)
-Ceara	Present	CABI and EPPO (2003) CABI (Undated)
-Goias	Present	Antônio (1982) CABI and EPPO (2003)
-Mato Grosso do Sul	Present	CABI and EPPO (2003)
-Minas Gerais	Present	CABI and EPPO (2003)
-Para	Present	Freire and Freire (1978) CABI and EPPO (2003)
-Paraiba	Present	CABI and EPPO (2003) CABI (Undated)
-Parana	Present	Antônio (1982) CABI and EPPO (2003) Moura et al. (2010) Machado et al. (2013)
-Pernambuco	Present	CABI and EPPO (2003)
-Rio de Janeiro	Present	Lima et al. (2005)
-Rio Grande do Norte	Present	CABI and EPPO (2003)
-Rio Grande do Sul	Present	CABI and EPPO (2003) Bellé et al. (2016) Kaspary et al. (2017) Neves et al. (2017) CABI (Undated)
-Santa Catarina	Present	CABI and EPPO (2003)
-Sao Paulo	Present	CABI and EPPO (2003) Araújo Filho et al. (2012) CABI (Undated)
-Sergipe	Present	CABI and EPPO (2003)
Chile	Present	CABI and EPPO (2003) CABI (Undated)
Colombia	Present	Taylor et al. (1982) CABI and EPPO (2003)
Ecuador	Present, Widespread	CABI and EPPO (2003) CABI (Undated)
Guyana	Present	Schotman (1989) CABI and EPPO (2003)
Paraguay	Present, Widespread	CABI and EPPO (2003) CABI (Undated)
Peru	Present	Taylor et al. (1982) CABI and EPPO (2003)
Suriname	Present	Taylor et al. (1982) CABI and EPPO (2003)
Uruguay	Present	Taylor et al. (1982) CABI and EPPO (2003)
Venezuela	Present	Crozzoli P. et al. (1991) CABI and EPPO (2003)

Risk of Introduction

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M. arenaria is a phytosanitary risk, but because it is so widespread it is not specifically quarantined.

Hosts/Species Affected

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The host range of M. arenaria is extremely large and includes members from many plant families including monocotyledons, dicotyledons, and herbaceous and woody plants. This root-knot species parasitizes most of the major food crops (vegetables, fruit trees, brambles and vines) and ornamental plants grown in tropical, subtropical and temperate climates.

For further details on hosts see Sasser (1952, 1954), Taylor et al. (1982) and Colbran (1958).

Host Plants and Other Plants Affected

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Plant name	Family	Context	References
Abelmoschus esculentus (okra)	Malvaceae	Main	Tariq-Khan et al. (2020)
Actinidia deliciosa (kiwifruit)	Actinidiaceae	Other
Amaranthus viridis (slender amaranth)	Amaranthaceae	Wild host
Angelica dahurica	Apiaceae	Other	Wang et al. (2013)
Anubias barteri	Araceae	Other
Arachis hypogaea (groundnut)	Fabaceae	Main
Atractylodes macrocephala	Asteraceae	Other	Wang et al. (2012)
Avena sativa (oats)	Poaceae	Main
Brosimum gaudichaudii	Moraceae	Habitat/association
Capsicum annuum (bell pepper)	Solanaceae	Unknown	López-Pérez et al. (2011)
Capsicum frutescens (chilli)	Solanaceae	Main
Chenopodium album (fat hen)	Chenopodiaceae	Wild host
Citrullus lanatus (watermelon)	Cucurbitaceae	Main
Clerodendrum inerme	Lamiaceae	Habitat/association
Coffea arabica (arabica coffee)	Rubiaceae	Main
Cucumis sativus (cucumber)	Cucurbitaceae	Main	López-Pérez et al. (2011); Tariq-Khan et al. (2020)
Cucurbita pepo (marrow)	Cucurbitaceae	Main	López-Pérez et al. (2011)
Cyperus esculentus (yellow nutsedge)	Cyperaceae	Wild host
Cyperus rotundus (purple nutsedge)	Cyperaceae	Wild host
Dahlia pinnata (garden dahlia)	Asteraceae	Habitat/association
Datura stramonium (jimsonweed)	Solanaceae	Wild host
Daucus carota (carrot)	Apiaceae	Main	López-Pérez et al. (2011)
Desmodium tortuosum (Florida beggarweed)	Fabaceae	Wild host
Deutzia gracilis (slender deutzia)	Hydrangeaceae	Wild host
Digitalis purpurea (foxglove)	Scrophulariaceae	Wild host
Dioscorea alata (white yam)	Dioscoreaceae	Other
Dioscorea cayenensis (Guinea yam)	Dioscoreaceae	Other
Eleusine indica (goose grass)	Poaceae	Wild host
Festuca arundinacea (tall fescue)	Poaceae	Wild host
Ficus carica (common fig)	Moraceae	Unknown	López-Pérez et al. (2011)
Glycine max (soyabean)	Fabaceae	Main
Gossypium hirsutum (Bourbon cotton)	Malvaceae	Main
Hibiscus cannabinus (kenaf)	Malvaceae	Unknown	López-Pérez et al. (2011)
Hibiscus tiliaceus (coast cottonwood)	Malvaceae	Unknown
Hordeum vulgare (barley)	Poaceae	Main
Ipomoea batatas (sweet potato)	Convolvulaceae	Main
Lactuca sativa (lettuce)	Asteraceae	Other	López-Pérez et al. (2011)
Lespedeza stipulacea (Korean lespedeza)	Fabaceae	Wild host
Lolium multiflorum (Italian ryegrass)	Poaceae	Wild host
Malus domestica (apple)	Rosaceae	Other
Malva parviflora (pink cheeseweed)	Malvaceae	Other	Kokalis-Burelle et al. (2012)
Medicago sativa (lucerne)	Fabaceae	Main
Melissa officinalis (Lemon balm)	Lamiaceae	Habitat/association
Morinda citrifolia (Indian mulberry)	Rubiaceae	Other	Fu et al. (2013)
Musa (banana)	Musaceae	Main
Musa acuminata (wild banana)	Musaceae	Unknown	López-Pérez et al. (2011)
Nicotiana tabacum (tobacco)	Solanaceae	Main	López-Pérez et al. (2011)
Olea europaea	Oleaceae	Unknown	López-Pérez et al. (2011)
Olea europaea subsp. europaea (European olive)	Oleaceae	Other
Oryza sativa (rice)	Poaceae	Main
Oxalis corniculata (creeping woodsorrel)	Oxalidaceae	Other
Passiflora edulis (passionfruit)	Passifloraceae	Main
Paulownia elongata (elongate paulownia)	Scrophulariaceae	Other
Petroselinum crispum (parsley)	Apiaceae	Other	Mennan et al. (2011)
Phaseolus vulgaris (common bean)	Fabaceae	Main	López-Pérez et al. (2011)
Phoenix dactylifera (date-palm)	Arecaceae	Other
Pimpinella anisum (aniseed)	Apiaceae	Habitat/association
Pisum sativum (pea)	Fabaceae	Main
Pongamia pinnata (Indian beech)	Fabaceae	Other
Prunus persica (peach)	Rosaceae	Main	López-Pérez et al. (2011)
Psidium guajava (guava)	Myrtaceae	Main
Raphanus sativus (radish)	Brassicaceae	Wild host
Ricinus communis (castor bean)	Euphorbiaceae	Wild host
Rosa multiflora (multiflora rose)	Rosaceae	Wild host
Rumex acetosella (sheep's sorrel)	Polygonaceae	Wild host
Saccharum officinarum (sugarcane)	Poaceae	Main
Salix babylonica (weeping willow)	Salicaceae	Unknown	López-Pérez et al. (2011)
Solanum lycopersicum (tomato)	Solanaceae	Main	López-Pérez et al. (2011); Tariq-Khan et al. (2020); Wang et al. (2013)
Solanum nigrum (black nightshade)	Solanaceae	Wild host
Solanum tuberosum (potato)	Solanaceae	Main	Johnson et al. (1996)
Solanum viarum (tropical soda apple)	Solanaceae	Other
Tectona grandis (teak)	Lamiaceae	Other	Araújo et al. (2012)
Trifolium pratense (red clover)	Fabaceae	Wild host
Trifolium repens (white clover)	Fabaceae	Main
Vicia sativa (common vetch)	Fabaceae	Unknown	López-Pérez et al. (2011)
Vigna unguiculata (cowpea)	Fabaceae	Main
Vitis vinifera (grapevine)	Vitaceae	Main	López-Pérez et al. (2011)
Zantedeschia aethiopica (calla lily)	Araceae	Other
Zea mays (maize)	Poaceae	Main	López-Pérez et al. (2011)

Growth Stages

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Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage

Symptoms

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Underground symptoms are primarily galls on roots, corms, tubers or peanut pods, abnormal formation and function of root system and giant cells blocking the vascular cylinder.

Non-specific above-ground symptoms include patchy, stunted growth; discoloration and leaf chlorosis; excessive wilting during dry, hot conditions; stunting of whole plants; reduced yield and quality; and sometimes premature senescence or death.

Infected plants are often stunted and chlorotic. Small to large galls (2-200 mm in diameter) occur on the roots of infected plants. M. arenaria populations often produce many small bead-like galls that do not form short lateral roots (Eisenback et al., 1981). Small wart-like projections may occur on infected corms, tubers and peanut pods. Symptoms are similar to those produced by most of the other root-knot nematode species.

Root-knot is generally more severe in sandy soils and under adverse environmental conditions such as drought and high temperatures. M. arenaria can interact with fungi or bacteria to cause more severe symptoms, break resistance to the disease agent, or allow weakly parasitic and non-pathogenic organisms to cause disease.

List of Symptoms/Signs

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Sign	Life Stages	Type
Leaves / abnormal colours
Leaves / wilting
Roots / galls along length
Roots / reduced root system
Whole plant / dwarfing
Whole plant / early senescence

Biology and Ecology

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Life Cycle

The egg of a root-knot nematode develops into a vermiform first-stage juvenile that undergoes one moult into a second-stage juvenile. The second-stage juvenile hatches from the egg, moves freely in the soil, penetrates the root just behind the root cap, migrates intercellularly in the root and establishes a feeding site within the developing vascular cylinder. As it feeds on the nematode-induced giant cell system, the second-stage juvenile loses its mobility and begins to increase in girth. After it has imbibed a sufficient quantity of sustenance, the flask-shaped second-stage juvenile moults three times without feeding and matures into a saccate adult female. Females of M. arenaria reproduce by mitotic parthenogenesis; as soon as they are mature adults they begin producing eggs (Triantaphyllou and Hirschmann, 1960).

Male second-stage juveniles undergo a metamorphosis during the third moult into elongate vermiform fourth-stage juveniles. The fourth-stage juvenile male remains enclosed in the cuticle of the second and third stages where it moults again to form an adult vermiform male. The male escapes from the cuticles and the root system. It moves freely in the soil, not feeding, only mating with mature adult females. As populations of M. arenaria reproduce by mitotic parthenogenesis, males serve no reproductive function (Triantaphyllou and Hirschmann, 1960).

The length of one generation of M. arenaria is greatly affected by temperature. At very high temperatures (>29°C), the life cycle takes approximately 3 weeks, but at very cool temperatures it can be extended to 2-3 months.

Natural enemies

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Natural enemy	Type	Life stages
Anatonchus	Parasite	Juveniles
Arthrobotrys tortor	Predator	Juveniles
Aspergillus flavus	Antagonist
Aspergillus niger	Antagonist
Butlerius	Parasite	Juveniles
Catenaria anguillulae	Pathogen	Juveniles
Clonostachys rosea	Antagonist	Adults
Dactylella	Predator	Juveniles
Diplogaster	Parasite	Juveniles
Discolaimus	Parasite	Juveniles
Dorylaimus	Parasite	Juveniles
Fusarium oxysporum	Pathogen	Eggs
Gliocladium catenulatum	Antagonist	Adults
Hypoaspis aculeifer	Predator	Eggs
Mononchoides	Parasite	Juveniles
Mononchus	Parasite	Juveniles
Myrothecium verrucaria	Pathogen
Paecilomyces lilacinus	Parasite	Juveniles
Paecilomyces nostocoides	Pathogen
Pasteuria penetrans	Pathogen	Juveniles
Pseudopapulaspora kendrickii	Pathogen
Sarocladium strictum	Pathogen	Eggs
Seinura	Parasite	Juveniles
Trichoderma harzianum	Antagonist
Trichoderma koningii	Antagonist
Tripyla	Predator	Juveniles
Verticillium chlamydosporium	Parasite	Eggs
Verticillium lamellicola	Pathogen

Notes on Natural Enemies

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The immense reproductive potential of the root-knot nematodes make them difficult to control by biological methods. Of the 500-1500 eggs produced by a single female, only 2% of the offspring need to be successful parasites for the population to increase by a factor of 12 in just three generations (Taylor and Sasser, 1978). Under most growing conditions where M. arenaria occurs, more than three generations are completed during one season.

Pathway Vectors

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Vector	Notes	Long Distance
Clothing, footwear and possessions	Eggs and galls in soil.	Yes
Containers and packaging - wood	Eggs and galls in soil.	Yes
Land vehicles	Eggs and galls in soil.	Yes
Mail	Eggs and galls in soil.	Yes
Soil, sand and gravel	Eggs and galls in soil.	Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transport	Pest stages	Borne internally	Borne externally	Visibility of pest or symptoms
Bulbs/Tubers/Corms/Rhizomes	adults; eggs; juveniles	Yes	Yes	Pest or symptoms not visible to the naked eye but usually visible under light microscope
Growing medium accompanying plants	adults; eggs; juveniles		Yes	Pest or symptoms not visible to the naked eye but usually visible under light microscope
Roots	adults; eggs; juveniles	Yes	Yes	Pest or symptoms not visible to the naked eye but usually visible under light microscope
Seedlings/Micropropagated plants	adults; eggs; juveniles	Yes	Yes	Pest or symptoms not visible to the naked eye but usually visible under light microscope

Plant parts not known to carry the pest in trade/transport
Bark
Flowers/Inflorescences/Cones/Calyx
Fruits (inc. pods)
Leaves
Stems (above ground)/Shoots/Trunks/Branches
True seeds (inc. grain)
Wood

Impact

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M. arenaria is an economically important plant pathogen that parasitizes thousands of plant species worldwide. The peanut root-knot nematode is a pest of major food crops and significantly reduces the quantity and quality of food and fibre production.

The average loss caused by root-knot nematodes is thought to be around 5%; however, in some fields the loss can be complete. In some areas of the world, root-knot nematodes are so common that galls on roots are considered normal. Often the damage caused by these nematodes is overlooked or the blame is placed on other agronomic problems. Stunted, unthrifty growth by infected plants is often attributed to vague agricultural ailments such as tired, poor, worn-out and exhausted land (Sasser and Carter, 1984).

Diagnosis

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Morphology of perineal patterns, shape and measurements of the stylet of the female, shape and measurements of the head and stylet of the male, and measurements of the second-stage juveniles are useful characters for species identification. Additional host range tests may be necessary to confirm the identification of the species and determination of the host race. Hosts of M. arenaria in the North Carolina differential host range test (Hartman and Sasser, 1984) include tobacco (Nicotiana tabacum cv. NC95), pepper (Capsicum annuum cv. California Wonder), tomato (Lycopersicon esculentum cv. Rutgers) and watermelon (Citrullus vulgaris [C. lanatus] cv. Charleston Gray). Host race 1 populations infect and reproduce on groundnut, whereas host race 2 populations do not.

A DNA probe that is specific for M. arenaria has been developed and may be useful for diagnosis of this species (Baum et al., 1994). Cytological and biochemical characterization provide additional characters for identification of M. arenaria (Triantaphyllou, 1979; Esbenshade and Triantaphyllou, 1989).

Detection and Inspection

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Galls formed on the roots of plants are very diagnostic for most species of root-knot nematode. Root-knot infected tubers, corms and groundnut pods can be examined for characteristic surface swellings, but may also be sliced into 1-2 mm sections and stained with hot acid fuchsin (Daykin and Hussey, 1984). Likewise, endoparasitic second-stage juveniles can be stained with acid fuchsin inside whole roots, placed between glass slides and examined with a dissecting microscope.

Similarities to Other Species/Conditions

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Many aspects of the morphology of M. arenaria are similar to other species within the Meloidogyne genus.

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Host-Plant Resistance

Much progress has been made in the use of resistant plants for reducing the damage caused by M. arenaria on various crop plants (Sasser and Kirby, 1979). Plants with some level of resistance include cultivars of cowpea, crown vetch, soyabean, passionfruit, okra, cassava, tomato, sweet potato, cucumber, guava rootstock, pepper, tobacco and various grasses (Sasser and Kirby, 1979). No useful source of resistance has been found in peanut.

Crop Rotation

Meloidogyne species are obligate parasites and populations decline rapidly in the absence of a host. Rotation of susceptible host crop plants with those that are immune or poor hosts is a useful way to reduce the effect that M. arenaria has on plant growth. Unfortunately, the non-host, when it does occur, is usually less profitable than the susceptible crop. M. arenaria has a very large host range and non-hosts or cultivars that have been reported resistant should be used with caution because of the innate variability that occurs in the root-knot nematodes. Other agronomic and economic factors are also important in the selection of a rotation crop. An adequate weed control programme is absolutely necessary for a crop rotation scheme to be effective because many weed species serve as suitable hosts (Taylor and Sasser, 1978).

Chemical Control

Nematicides have often been used for limiting the damage that nematodes cause on plants. Nematicides are usually used as a soil treatment before planting. However, a few nematicides can be applied after planting. These chemicals are relatively expensive and they require costly equipment and trained personnel to apply them.

High-value crops that are good hosts of M. arenaria can be protected with a soil fumigant. These chemicals volatilize and kill the nematodes on contact. Less valuable crops can be protected with cheaper non-fumigant nematicides that dissolve in water and act as nerve poisons. They prevent nematodes from feeding on plants for 2-3 weeks, but their effect is reversible. Often they do not kill the nematodes. Because they are water soluble, their effectiveness is dependent on an adequate amount of soil moisture. If an optimum amount of water is available, the optimum effect is achieved; if too much or too little water is present, very little control is achieved (Bunt, 1987).

Biological Control

Numerous attempts have been made to control root-knot nematodes with parasitic and predacious organisms or various organic amendments, with varying degrees of success. Naturally occurring organisms, such as Pasteuria penetrans, which are obligate parasites of Meloidogyne, may prove to be effective for biological control.

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