WEED SPECIES AS HOSTS OF MELOIDOGYNE: A REVIEW

J. R. Rich,1* J. A. Brito,2 R. Kaur2, and J. A. Ferrell3

1University of Florida, IFAS North Florida Research and Education Center, Quincy, FL 32351 USA; 2Division of Plant Industry, Gainesville, FL 32614; 3Department of Agronomy, University of Florida, Gainesville, FL 32611. *Corresponding author: [email protected]

ABSTRACT

Rich, J. R., J. A. Brito, R. Kaur, and J. A. Ferrell. 2008. Weed species as hosts of Meloidogyne: A review. Nematropica 39:157-185. Over 226 weed species belonging to 43 botanical families have been studied for host suitability to different root-knot nematodes worldwide. Meloidogyne incognita was reported to reproduce on the largest number of weeds with over 138 weedy plant hosts throughout the world. Four other Meloidog- yne spp., M. javanica, M. arenaria, M. hapla, and M. graminicola were found infecting 49, 48, 27, and 24 weed species, respectively, while M. mayaguensis, M. chitwoodi, and M. floridensis have been reported to reproduce on 24, 13 and 12 weed species, respectively. These data indicate that little is known about the exact weed host range of many species within the genus Meloidogyne. However, weed host status of Meloidogyne spp. reported herein indicates that weeds are a major reservoir of root-knot nematodes and should be considered as factors affecting the success of integrated nematode management pro- grams. Key words: host range, Meloidogyne arenaria, M. chitwoodi, M. coffeicola, M. exigua, M. floridensis, M. graminicola, M. hapla, M. incognita, M. javanica, M. konaensis, M. mayaguensis, M. naasi, M. paranaensis, M. triticoryza, root-knot nematodes, weeds.

RESUMEN

Rich, J. R., J. A. Brito, R. Kaur, and J. A. Ferrell. 2008. Especies de malezas como hospedantes de Me- loidogyne: Revisión. Nematropica 39:157-185. Se ha estudiado la susceptibilidad de más de 226 especies de malezas, pertenecientes a 43 familias botánicas, a diferentes nematodos agalladores. Se ha registrado a Meloidogyne incognita como la espe- cie con la capacidad de reproducirse en la mayor cantidad de malezas, con más de 138 especies ve- getales registradas como hospedantes en el mundo. Se han registrado cuatro otras especies de Meloi- dogyne spp., M. javanica, M. arenaria, M. hapla, y M. graminicola infectando 49, 48, 27 y 24 especies de malezas, respectivamente, mientras que M. mayaguensis, M. chitwoodi, y M. floridensis se han registrado en 24, 13 y 12 especies de malezas, respectivamente. Estos datos indican que se conoce poco acerca del rango de malezas hospedantes de muchas especies de Meloidogyne. No obstante, la lista de malezas susceptibles a Meloidogyne spp. que incluimos en esta revisión indica que las malezas son un importan- te reservorio de nematodos agalladores y que deben considerarse como factores que afectan el éxito de programas de manejo integrado de nematodos. Palabras clave: malezas, Meloidogyne arenaria, M. chitwoodi, M. coffeicola, M. exigua, M. floridensis, M. gra- minicola, M. hapla, M. incognita, M. javanica, M. konaensis, M. mayaguensis, M. naasi, M. paranaensis, M. triticoryza, nematodos agalladores, rango de hospedantes.

INTRODUCTION crop plants for water, soil nutrients, and light and also interfere with distribution of Weeds are a major constraint in agricul- irrigation water and efficient fertilizer tural production since they compete with application (Anderson, 1996). Competi-

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158 NEMATROPICA Vol. 39, No. 2, 2009

tion for resources with crop plants, particu- anica represent 95% of all infestations in larly in the early stages of growth, can agricultural lands (Hussey and Janssen, reduce crop yields more than 70% (Bar- 2002). The interaction of root-knot nema- rentine 1974; Bendixen, 1986a; Smith todes and weeds can magnify problems cre- 1968; Thurlow and Buchanan 1972). Due ated by both pests in agricultural to their ubiquitous presence before, dur- production systems compared to each act- ing, and after crop cycles, weeds serve as ing alone. For example, the four weed spe- reservoirs for plant pathogens and nema- cies, Ageratum conzyoides, Amaranthus todes that will cause damage in future spinosus, Eleusine indica and Portulaca olera- crops (McWhorter 1989; Quénéhervé et al., cea are considered among the world’s worst 1995; Quénéhervé et al., 2006). weeds and are multiple hosts of Meloidogyne The problem of weed hosts to plant- as well (Holm et al., 1977). Cyperus rotundus, parasitic nematodes is particularly severe in Amaranthus spp., Chenopodium album, and the subtropical and tropical environments Digitaria spp. are also frequently encoun- where weeds grow year round (Crane et al., tered weed hosts of root-knot nematodes 2008). Previous research has shown many (Myers et al., 2004). The weed species common agricultural weeds to be excellent Myrica faya (syn. Morella faya) is a major hosts of plant-parasitic nematodes (Asmus threat to the natural ecosystem in Hawaii and Andrade, 1997; Bendixen et al., 1986b; (Vitouseck and Walker, 1989) and was 1988a, b, c; Clements et al., 2004; Diop et found to be a host of Meloidogyne spp. al., 2000; Hogger and Bird, 1976; Sellami et (Duffy and Gardner, 1999). Since weeds al., 1999). Weed species enable plant-para- are ubiquitously present and many are sitic nematodes to survive in the absence as good hosts of root-knot nematodes, con- well as the presence of a crop, providing a trolling weeds would be a good initial step nematode inoculum source for the follow- in reducing root-knot nematode popula- ing season. For example, plant-parasitic tions worldwide (Das et al., 1998). nematodes including Aphelenchoides besseyi, The definition of a weed is ‘any Ditylenchus angustus, Heterodera orizicola and unwanted plant’, essentially meaning any M. graminicola were found to reproduce on plant which interferes with growth and weeds that grow in fallow fields between production of a desired plant species rice crops, which perpetuate nematode (Anderson, 1996). Using this definition, problems (Rao et al., 1986). the concept of weeds can be broadened to The root-knot nematodes, Meloidogyne include volunteer crops, such as peanut spp., have been reported to survive and plants growing before or during a cotton even thrive on weeds (Davidson and Town- crop. While these situations are important shend, 1967; Luc et al., 2005; Zhang and in rotations designed to minimize nema- Schmitt, 1994). This genus is considered tode damage, the weeds reported herein the most important among plant-parasitic generally do not include agricultural crop nematodes (Sasser and Freckman, 1987), plants that may be ‘out of place’. Weed host mainly due to the wide host range which is status descriptions as related in this article, known to exceed 3000 wild and cultivated e.g. good or moderate host, in most plant species (Hussey and Janssen, 2002). instances are terms used by individual To date, about 96 nominal species of authors to define degree of nematode Meloidogyne have been described (Brito et reproduction or galling on a particular al., 2008), but within the genus, M. weed. Also, some plants that are listed may arenaria, M. hapla, M. incognita, and M. jav- or may not be weeds in the classical sense

Weed hosts of Meloidogyne: Rich et al. 159

but rather alternative hosts to Meloidogyne Meloidogyne chitwoodi spp. which have been included for thor- The Columbia root-knot nematode was oughness. A number of references were originally found in the Pacific Northwest found which described a weed or Meloidog- USA infecting potato (Golden et al., 1980). yne genus without giving a species name. It has subsequently been reported in a These references were not included in this number of other countries including article. All weeds described as hosts for Argentina, Belgium, Germany, Mexico the individual Meloidogyne spp. are listed in Netherlands, Portugal, and South Africa Table 1, but references to this table are not (Powers et al., 2005). In an extensive host made again in the text of this article. range study, O’Bannon et al. (1982) found WEEDS AS HOSTS OF MELOIDOGYNE SPP. that 53 of 68 weed and cultivated plant spe- cies were hosts to M. chitwoodi. Among the Meloidogyne arenaria weed species listed as hosts were Cirsium vulgare, Dactylis glomerata, Panicum capillare, The peanut root-knot nematode is pre- Setaria viridis, and Sonchus asper. Similarly, dominately found in warm-temperate and Griffin et al. (1984) reported several range tropical environments and is highly viru- grasses as hosts to M. chitwoodi, and these lent on numerous ornamental, vegetable, included Agropyron desertorum, Bromis iner- and agronomic crops. In weed host status mis, Dactylis glomerata, and Pascopyrum studies in North Carolina USA, Chenopo- smithii. Kutywayo and Been (2006) listed dium album, Euphorbia maculata, and Vicia Capsella bursa-pastoris, Senecio vulgaris, and villosa were described as good hosts while Solanum nigrum as hosts of M. chitwoodi, and Amaranthus hybridus, A. palmeri, Ambrosia Solanum sarrachoides was also reported as a artemisiifolia, Ipomoea hederacea var. integrius- host of this nematode (Boydston et al., cula, Setaria lutescens, Sida spinosa, Portulaca 2008). oleracea, Rumex acetosella and Rumex crispus were moderate hosts of M. arenaria race 2 Meloidogyne coffeicola (Tedford and Fortnum, 1988). Brito et al. (2008) identified Amaranthus spinosus, Ipo- The common name for this nematode moea triloba, Jacquemontia tamnifolia, Macrop- is the coffee root-knot nematode although tilium lathyroides, Portulaca oleracea, and another nematode, M. exigua, also has the Physalis angulata as weed hosts of M. same common name. This nematode has arenaria in Florida. Meloidogyne arenaria was only been found in Brazil and has a very observed on Solanum viarum, which is an narrow host range. Coffea arabica is the invasive perennial weed of pastures present major plant host of M. coffeicola, and it has predominantly in the southeastern USA only been reported infecting two weeds, (Church and Rosskopf, 2005). Their work Eupatorium pauciflorum and Psychotria nitid- indicated that this weed species is a poten- ula (Jaehn et al., 1980; Lordello and tial reservoir for M. arenaria in Florida and Lordello, 1972). throughout the southern USA. Other weeds reported as hosts of M. arenaria Meloidogyne exigua include Abutilon theophrasti, Amaranthus ret- roflexus, A. spinosus, Cnidoscolus stimulosus, This coffee root-knot nematode was Dichondra repens, Echinochloa muricata, Ipo- first described by Goeldi (1887) from cof- moea triloba, Leontodon hispidus, and Phyto- fee in Brazil, and now is wide spread in lacca americana (Kaur et al., 2007). central and south America. The nematode 160 NEMATROPICA Vol. 39, No. 2, 2009 Mh , graminicola . M – ., 1988

., 1999 , 1984 ., 1999 ., 1995 al , 1985

., 2004 ., 2004 ., 2004 Mg al ., 2008 et al ., 2007

; et mous, 2007). et al et et al. et al et al et al et al. et al et al ducted to determine current triticoryzae. Source/Reference . elasquez-Valle, 2001 elasquez-Valle, edford and Fortum, 1988 M Gowda Amin and Budai, 1994 Kaur Ehwaeti Brito Mani and Hinai, 1996 Mani and Hinai, 1996 Mamaril and Aberto, 1989 Khan Roese and Oliveira, 2004 Griffin Khan Mani and Hinai, 1996 Khan Sellami Mani and Hinai, 1996 V T Lima Lordello floridensis - . Mt M ; -

Mf ; z . sp paranaensis . M spp. worldwide. – Mp ; Meloidogyne Mi Mi Ma, Mf, Mi, Mj, Mm Mi Ma, Mi Mi Mi Mi, Mj Mg Mp Mc Mg Mi Mg Mj Mi Mi Ma, Mi Me Mj Mi, Mj Mani and Hinai, 1996 Meloidogyne M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name elvet-leaf Malvaceae umbleweed Amaranthaceae M Prickly chaff-flower Amaranthaceae V Australian acalypha Euphorbiaceae unknown Euphorbiaceae Kapok bushGoat weed Amaranthaceae Crested wheatgrassQuackgrass Asteraceae Poaceae Poaceae Sessile joyweed Amaranthaceae Prostrate pigweed Amaranthaceae T Smooth pigweed Amaranthaceae Low amaranth Amaranthaceae - Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the Achyranthes aspera names, however when noted, the current synonymns were listed in parenthesis. Abutilon theophrasti Acalypha australis Acalypha setosa Aerva javanica Ageratum conyzoides desertorumAgropyron repens Agropyron Alternanthera sessilis Amaranthus angustifolius T Scientific name x y z Amaranthus graecizans Amaranthus hybridus Amaranthus deflexus – Weed hosts of Meloidogyne: Rich et al. 161 Mh , graminicola . M –

Mg ., 1999 ., 1999 ., 1999 ., 1999 ; mous, 2007). al ., 2004 ., 2006

., 2008 ., 2008 ., 2007 ., 2007 et et al et al et al ducted to determine current triticoryzae. et al et al Source/Reference . et al et al et al et al M floridensis - edford and Fortum, 1988 edford and Fortum, 1988 edford and Fortum, 1988 . Asmus and Andrade, 1997 Sellami Souza Amin and Budai,1994 T 1967 Davidson and Townshend, Sellami Mani and Hinai, 1996 Myers T Sellami Kaur Sperandio and Amaral, 1994 Roese and Oliveira, 2004 Kaur Brito 1967 Davidson and Townshend, Ehwaeti Davidson and Townshend, 1967 Davidson and Townshend, T Brito Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M – Mp ; Meloidogyne Mj Mi Mm Mi, Ma Ma, Mi Mi Mi Mi Mi Ma, Mi Mi, Mj Ma, Mf, Mi, Mj, Mm Mg Mp Ma, Mf, Mi, Mj, Mm Ma, Mi Mi Mi Mi Ma, Mi Mi M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name M Livid amaranthPalmer amaranthRedroot amaranth Amaranthaceae Amaranthaceae Amaranthaceae Slender amaranth Common ragweed Amaranthaceae Scarlet pimpernel Asteraceae Primulaceae Spiny amaranth Amaranthaceae chamomile, mayweedThymeleaf sandwort Asteraceae Caryophyllaceae Carpetgrass Poaceae Garden yellow rocketCommon beggarticks Brassicaceae Asteraceae - Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the names, however when noted, the current synonymns were listed in parenthesis. x y z Scientific name Amaranthus lividus (syn. blitum) Amaranthus palmeri Amaranthus retroflexus Amaranthus viridis artemisiifoliaAmbrosia Anagallis arvensis T Amaranthus spinosus Anthemis cotula serpyllifolia Arenaria Axonopus affinis vulgaris Barbarea Bidens alba – 162 NEMATROPICA Vol. 39, No. 2, 2009 Mh , ., 1977 graminicola et al . M – ., 1982

., 1988 ., 1996 Mg ., 1995 ., 1996 et al ., 1984 ; mous, 2007). ., 2004 ., 2004 ., 2008 et al ., 1998 et al et al et al et al al ducted to determine current

et al et al triticoryzae. et al Source/Reference et . M floridensis - Lordello Asmus and Andrade 1997 Willers, 1997 Belair and Benoit, 1996 Gowda Khan Roese and Oliveira, 2004 Khan Griffin Belair and Benoit, 1996 Leroux Belair and Benoit, 1996 Kim Davidson and Townshend, 1967 O’Bannon Kutywayo and Been, 2006 Belair and Benoit, 1996 Rodriguez-Kabana Amin and Budai,1994 Gowda . Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M – Mj Mj Mm MmMh Mi Mg Brito Mp Mg Mc Mh Mh Mh Mi, Mh Mi Mc Mc Mh Ma Mi Mi Mp ; Meloidogyne M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name all beggarticks Asteraceae M T Hairy beggarticks Asteraceae LadainaCaucasian bluestem Poaceae Rubiaceae AlexandergrassBrowntop millet Poaceae Poaceae Smooth Broom Poaceae Nodding beggarticks Asteraceae beggarticks Devil’s Asteraceae CheatShepherd’s-purse Brassicaceae Poaceae Sicklepod Fabaceae Carrot bur parsley Celosia Apiaceae Amaranthaceae - Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne (syn. Urochloa plantaginea) (syn. Urochloa . incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the Bidens vulgata Bidens pilosa Borreria hispida (syn. Spermacoce hispida) Bothriochloa intermedia Brachiaria plantaginea ramosa) Brachiaria ramosa (syn. Urochloa Bromus inermis Bromus names, however when noted, the current synonymns were listed in parenthesis. Bidens cernua Bidens frondosa x y z Bromus secalinus Bromus Capsella bursa-pastoris Cassia obtusifolia (syn. Senna obtusifolia) Scientific name Caucalis platycarpos (syn. lappula) Celosia argentea T – Weed hosts of Meloidogyne: Rich et al. 163 Mh , , 2008 et al. graminicola ., 1977 . M et al –

Mg ; , 1999 mous, 2007). , 1999 ., 1999 , 2006 ., 2004; Brito , 2008 ducted to determine current triticoryzae. et al. ., 1998 Source/Reference et al. et al . al et al et al.

et al M et floridensis - . edford and Fortum, 1988 edford and Fortum, 1988 Mt M Davidson and Townshend, 1967 Davidson and Townshend, Rodriguez-Kabana T 1967; Davidson and Townshend, Ehwaeti Amin and Budai,1994 Belair and Benoit Amin and Budai,1994 Sellami et al., 1999 Mani and Hinai, 1996 Sellami Brito Mani and Hinai, 1996 Souza T Myers Sellami Mani and Hinai, 1996 Amin and Budai, 1994 Belair and Benoit, 1996 Kim ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M – Mi Ma Ma, Mi Mi Mi Mh Mi, Mh, Mn Mi Mi, Mj Mi Mm Mi Mm Ma, Mi Mi Mi Mi Mn Mh Mi, Mh Mp ; Meloidogyne M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name M - Common lambs-quarters Chenopodiaceae Field sandburBig chickweed Poaceae Caryophyllaceae Oak-leaf goosefootGiantseed goosefoot Chenopodiaceae Chenopodiaceae Nettleleaf goosefoot Chenopodiaceae Garden spurgeGround spurge Euphorbiaceae Euphorbiaceae Rhodes grass Poaceae Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the names, however when noted, the current synonymns were listed in parenthesis. Chenopodium album Cenchrus incertus (syn. Cenchrus spinifex) Cerastium vulgatum (syn. fontanum) x y z Scientific name Chenopodium glaucum Chenopodium hybridum (syn. simplex) Chenopodium murale Chamaesyce hirta Chamaesyce prostrata T Chloris gayana – 164 NEMATROPICA Vol. 39, No. 2, 2009 Mh , graminicola . M –

., 1982 ., 1999 Mg ., 1997 ., 1978 ., 1999 ., 1999 ., 1972 ; ., 1996 et al mous, 2007). et al , 2006 ., 2004 ., 2007 et al et al et al et al et al ducted to determine current et al triticoryzae. et al. et al Source/Reference . et al M floridensis - ovlas and Inserra, 1979 edford and Fortum, 1988 edford and Fortum, 1988 edford and Fortum, 1988 . Gowda Kaur Moraes Souza Roese and Oliveira, 2004 Aragon Mani and Hinai, 1996 Sellami Belair and Benoit, 1996 Kutywayo and Been, 2006 O’Bannon V T T Sellami Roese and Oliveira, 2004 1967 Davidson and Townshend, T Myers Thomas Schroeder Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M – Mp ; Meloidogyne Mi Ma, Mf, Mi, Mj, Mm Me Mm Mp Me Mj Mi, Mj Mh Mc Mc Mn Ma, Mi Ma, Mi Mi Mp Mi Ma, Mi Mi Mi Mi M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name ellow nutsedge Cyperaceae M Jasmin del rioSpurge nettle Capparidaceae Euphorbiaceae CitronmelonDevil nettle Cucurbitaceae Benghal dayflowerSpreading dayflower Commelinaceae Field bindweed Commelinaceae Euphorbiaceae Convolvulaceae Oxeye daisyCanadian thistleBull thistle Asteraceae Asteraceae Asteraceae CrownvetchShowy crotalariaBermudagrass Poaceae Fabaceae Fabaceae Hound’s TongueHound’s Y Boraginaceae - Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the Cleome viscosa Cnidoscolus stimulosus Citrullus vulgaris (syn. Citrullus lanatus) Cnidoscolus urens Commelina benghalensis Commelina diffusa Convolvulus arvensis names, however when noted, the current synonymns were listed in parenthesis. Chrysanthemum leucanthemum (syn. Leucanthemum vulgare) Cirsium arvense Cirsium vulgare x y z Coronilla scropioides Coronilla spectablis Crotalaria Cynodon dactylon Scientific name Cynoglossum officinale Cyperus esculentus T – Weed hosts of Meloidogyne: Rich et al. 165 Mh , graminicola . M –

Mg ., 2004 ., 1999 ., 1999 ., 1999 ., 2004 ; ., 1984 mous, 2007). ., 2004 ., 2004 al

., 2004 ., 2004 al et al et

et al et al et al ducted to determine current et al et et al triticoryzae. Source/Reference et al et al . M floridensis - edford and Fortum, 1988 . Davis and Webster, 2003 Davis and Webster, Meyers Dabur Aragon et al., 1978 Zhang and Schmitt, 1994 Mani and Hinai, 1996 Gaur and Sharma 1998 Sellami Myers et al., 2004 2003 Davis and Webster, Thomas Khan Dabur Roese and Oliveira, 2004 Brito et al., 2008; Griffin Khan Sellami Mani and Hinai, 1996 T Ehwaeti Davidson and Townshend, 1967 Davidson and Townshend, Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M – Mi Mi Mg Me Mk Mj Mt Mi Mi Mi Mi Mg Mg Mp Mg Mc Mg Mi Mj Ma, Mi Mi Mi Mp ; Meloidogyne M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name Rice flat sedge Cyperaceae Purple nutsedge Cyperaceae Orchardgrass Poaceae Crowfootgrass Poaceae Wild carrot Umbelliferae Sacred dacturaJimsonweed Solanaceae Solanaceae Herb sophia Brassicaceae M - Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the names, however when noted, the current synonymns were listed in parenthesis. x y z Cyperus iria Cyperus rotundus Dactylis glomerata Dactyloctenium aegyptium Daucus carota – Datura metel (syn. inoxia) Dactura stramonium Scientific name T Descurainia sophia 166 NEMATROPICA Vol. 39, No. 2, 2009 Mh , graminicola . M –

., 1988 Mg ., 2004 ., 2004, ., 2004 ., 2004 ., 2004 ., 2004 ; mous, 2007). et al ., 2007 ., 2007 al al al

et et al et et et al et al ducted to determine current et al et al triticoryzae. Source/Reference . M edford and Fortum, 1988 edford and Fortum, 1988 edford and Fortum, 1988 floridensis - Dabur Kaur Asmus and Andrade, 1997 T Amin and Budai, 1994 Noling and Gilreath, 2002a Khan Khan Roese and Oliveira, 2004 Sperandio and Amaral, 1994 T Gaur and Sharma, 1998 Dabur Kaur Brito et al., 2008 T Lordello Sperandio and Amaral, 1994 Gaur and Sharma, 1998 Dabur Mani and Hinai, 1996 Dabur . Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M – Mp ; Meloidogyne Mg Ma, Mf, Mi, Mj, Mm Mj Ma Mn Mi Mg Mp Mg Ma, Mi Mt Mg Ma, Mi Mm Ma, Mi Mj Mg Mt Mg Mj Mg M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name M Kleberg’s bluestemKleberg’s Dichondra Poaceae Convolvulaceae Jamaican crabgrassLarge crabgrass Poaceae Poaceae Barnyard-grass Poaceae Rough barnyard- grass Poaceae EcliptaGoosegrass Poaceae Asteraceae Jungle-rice Poaceae Eclipta Asteraceae - Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the Scientific name names, however when noted, the current synonymns were listed in parenthesis. Dichanthium annulatum Dichondra repens x y z Digitaria horizontalis Digitaria sanguinalis T Echinochloa crus-galli Echinochloa muricata Eclipta prostrata Eleusine indica Echinochloa colona Eclipta alba (syn. prostrata) – Weed hosts of Meloidogyne: Rich et al. 167 Mh , graminicola . M –

., 1999 ., 1996 Mg ., 1996 ., 1996; al ., 2006 ; ., 1980 , 2008 mous, 2007).

al ., 2008 ., 2008

et ., 1998 ., 1998 et al et al et et al et al ducted to determine current triticoryzae. et al et al Source/Reference et al. et al et al . M edford and Fortnum, 1988 floridensis - Belair and Benoit, 1996 Belair and Benoit, 1996 Lordello and Lordello, 1972 Jaehn Mani and Hinai, 1996 Gowda T Mani and Hinai, 1996 Leroux Kim Kim Gowda Ehwaeti Mani and Hinai, 1996 Brito Sperandio and Amaral, 1994 Roese and Oliveira, 2004 Brito Nisha and Sheela, 2002 Roese and Oliveira, 2004 Souza Brito . Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M Mh Mh Mco Mco Mi, Mj Mi Ma Mi Mh Mh Mh Mi Mi Mj Mi Mg Mp Mi, Mj Mp Mm Mf – Mp ; Meloidogyne M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name ormseed wallflower Brassicaceae M W Spotted joepyeweed unknown Asteraceae Asteraceae unknownGarden spurge Euphorbiaceae Euphorbiaceae Spotted spurgePetty spurge Euphorbiaceae Euphorbiaceae Horseweed Asteraceae Fleabane Asteraceae Red tasselflower Asteraceae Hairy willowherb Onagraceae American burnweed Asteraceae Cupid’s-shaving-brush Asteraceae Cupid’s-shaving-brush - Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the Scientific name Erysimum cheiranthoides Eupatorium maculatum Eupatorium pauciflorum Euphorbia granulata Euphorbia hirta (syn. Chamaesyce hirta) T Euphorbia maculata (syn. Chamaesyce maculata) Euphorbia peplus Erigeron canadensis (syn. Conyza canadensis) Erigeron Erigeron annuus Erigeron Emilia sonchifolia Epilobiurn hirsutum Erechtites hieracifolia Erechtites names, however when noted, the current synonymns were listed in parenthesis. Emilia fosbergii x y z – 168 NEMATROPICA Vol. 39, No. 2, 2009 Mh , graminicola . M –

., 1988 ., 1988 Mg ., 1999 ., 1999 ; mous, 2007). al ., 2006 , 2008

., 1985 ., 1985 et al et al et et al ducted to determine current triticoryzae. et al Source/Reference et al et al . et al. M floridensis - edford and Fortum, 1988 . Sperandio and Amaral, 1994 Mani and Hinai, 1996 Sellami Davidson and Townshed, 1967; Davidson and Townshed, T Roese and Oliveira, 2004 Souza Lordello Amin and Budai,1994 Sheridan and Grbavas, 1979 Belair and Benoit, 1996 Desaeger and Rao, 2000 Mani and Hinai, 1996 Ehwaeti 1967 Davidson and Townshend, 1967 Davidson and Townshend, Campos and Villain, 2005 Lima Lordello Brito Roese and Oliveira, 2004 Lima Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M Mg Mi Mi Mi, Mi Mp Mm Me Ma Mn Mh Mj Mj Mi Mi Mi Mp Me Mj Mi Mp Mj – Mp ; Meloidogyne M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name M - Globe fringerushClustered yellowtopsWhite ramping fumitory Cyperaceae Compositae Fumariaceae Spotted spurgeIndiantree Euphorbiaceae Euphorbiaceae MannagrassLow cudweedunknownunknownMouseear hawkweed Poaceae Asteraceae Foxtail barleySpotted St. Johnswort Asteraceae Garden balsam Asteraceae Guttiferae Blue morning-glory Boraginaceae Poaceae unknown Convolvulaceae Balsaminaceae Convolvulaceae Mulberryweed Moraceae Hairy crabweedSmallflower galinsoga Asteraceae Asteraceae Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the Scientific name Fimbristylis miliacea Flaveria trinervia Fumaria capreolata names, however when noted, the current synonymns were listed in parenthesis. T Euphorbia supina (syn. Chamaesyce maculata) Euphorbia tirucalli Glyceria fluitans Gnaphalium uliginosum Gutenbergia cordifolia bacciferumHeliotropium Hieracium pilosella jubatum Hordeum Hypericum punctatum Impatiens balsamina Ipomoea acuminata (syn. indica) Ipomoea aristolochiaefolia x y z Fatoua villosa Galinsoga ciliata (syn. quadriradiata) Galinsoga parviflora – Weed hosts of Meloidogyne: Rich et al. 169 Mh , , 2008 , 1977 , 1977 , 1977 ., 1977 ., 1977 et al. graminicola et al et al. et al. et al. et al . M –

., 1999 ., 2006 Mg ., 1996 al ; , 2008 mous, 2007).

al al ., 2007; Brito ., 2007 ., 2007 ., 2007

et et et ducted to determine current triticoryzae. et al. et al et al et al et al Source/Reference . M edford and Fortum, 1988 floridensis - Roese and Oliveira, 2004 Rodriguez-Kabana T Rodriguez-Kabana Rodriguez-Kabana Rodriguez-Kabana Stanley Kaur Kaur Rodriguez-Kabana Gowda Davis and Webster, 2003 Davis and Webster, Brito Davidson and Townshed, 1967 Davidson and Townshed, Mani and Hinai, 1996 Kaur Ehwaeti Kaur Davidson and Townshend, 1967 Davidson and Townshend, . Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M – Mp ; Meloidogyne Mp Ma Ma, Mi Ma Ma Ma Mf Ma,Mi, Mm, Mf, Mj Mf, Mm Ma Mi Mi Ma Mi Mi Mf Mi Ma, Mf, Mi, Mj, Mm Mi M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family Convolvulaceae Convolvulaceae Convolvulaceae M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name M - Morning-gloryIvyleaf morningglory Convolvulaceae Convolvulaceae Pittted morningglory Convolvulaceae Bigroot morningglory Convolvulaceae Cypressvine moring-glory Convolvulaceae Three-lobed morning- glory Heavenly blue morning- glory Smallflower morning- glory lettuce Asteraceae Henbit deadnettleProcumbent launaeaLionsear Lamiaceae Asteraceae Lamiaceae Bristly hawkbit Asteraceae Common motherwort Lamiaceae Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the names, however when noted, the current synonymns were listed in parenthesis. Ipomoea grandifolia Ipomoea hederacea x y z Ipomoea lacunosa Ipomoea pandurata – Ipomoea quamoclit Ipomoea triloba Ipomoea violacea Jacquemontia tamnifolia Lactuca runcinata Scientific name Lamium amplexicaule Launaea procumbens Leonotis nepetaefolia Leontodon hispidus T Leonurus cardiaca 170 NEMATROPICA Vol. 39, No. 2, 2009 Mh , graminicola . M –

., 2000 ., 2002 Mg , 1972 ., 1999 ., 1999 ., 1999 ., 1999 ., 2006 ; ., 1995 mous, 2007). ., 2004 al al , 2008 , 2008 , 2008

al ., 1985 et al

et al al et et

et al et al et al. et ducted to determine current et al et triticoryzae. Source/Reference et al . et al. et al. et al. M floridensis - olvas and Inserra, 1979 olvas and Inserra, 1979 . Davidson and Townshend, 1967 Davidson and Townshend, Sellami V Stanley Moraes Lima Gaur and Sharma, 1998 Santiago Ehwaeti Sellami Gowda Brito Lorenzo Dabur Brito Davidson and Townshend, 1967 Davidson and Townshend, Amin and Budai,1994 Belair and Benoit, 1996 V 1967 Davidson and Townshend, Ehwaeti Brito Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M – Mi Mj Mn Mf Mj Me Mt Mp Ma, Mi Mi Mi Ma, Mi Mj Mg Mi Mi Mi Mh Mn Mi Mi Mi Mp ; Meloidogyne M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name ellow sweetclover Fabaceae Common mallowCheeseweed mallow Malvaceae Malvaceae MelilotEnglish watercress Brassicaceae Fabaceae Siberian motherwort Lamiaceae Sprangletop Poaceae Mate Aquifoliaceae Bird’s -foot trefoilBird’s Fabaceae High mallow Malvaceae Thumba plantWild bushbean Lamiaceae Fabaceae Y Catnip Labiatae Scentless chamomile Pine-apple weed Asteraceae California burcloverBlack medic Fabaceae Asteraceae Fabaceae M - Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the Malva neglecta Malva parviflora Melilotus sulcata Nasturtium officinalis Scientific name names, however when noted, the current synonymns were listed in parenthesis. Leonurus sibiricus Leptochloa coloniculus Llex paraguariensis x y z Lotus corniculatus Malva sylvestris Lucas aspera lathyroides Macroptilium Melilotus alba (syn. officinalis) T Nepeta cataria Matricaria indora (syn. Tripleurospermum perforata) Matricaria indora (syn. Tripleurospermum Matricaria matricarioides (syn. discoidea) Medicago hispida (syn. polymorpha) Medicago lupulina – Weed hosts of Meloidogyne: Rich et al. 171 Mh , graminicola . M – ., 1982

., 1999 Mg ., 1996 ., 1996 et al ., 1984 , 2006 ; mous, 2007). ., 2004 al al ., 2008 ., 2008 ., 2007

et al et et et al et al. ducted to determine current et al triticoryzae. et al et al et al Source/Reference . M edford and Fortum, 1988 edford and Fortum, 1988 floridensis - Gowda Gowda T 1967 Davidson and Townshend, O’Bannon Sellami Sperandio and Amaral, 1994 Sperandio and Amaral, 1994 Griffin Mani and Hinai, 1996 Khan Kaur Brito Zhang and Schmitt, 1994 T Souza Belair and Benoit, 1996 Mani and Hinai, 1996 Gaur and Sharma, 1998 Mani and Hinai, 1996 Brito . Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M – Mp ; Meloidogyne Mi Mi Ma Mi Mc Mi Mg Mg Mc Mi, Mj Mg Ma, Mf, Mi, Mj, Mm Mi Mk Ma, Mi Mm Mh Mi Mt Mi Ma M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family Onagraceae M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name M orpedograss Poaceae ellow woodsorrel Oxalidaceae - Basil, besobla Labiatae Basil Labiatae Common eveningprim- rose Y Witchgrass Poaceae Wild-proso millet Poaceae Annual bluegrass Poaceae T wheatgrass Poaceae Wild groundcherry gooseberry Solanaceae American pokeweedArtillery plant Phytolaccaceae Urticaceae Hilograss, sour paspalum Poaceae Bahiagrasspassion flower Poaceae Passifloraceae Wild parsnipNapiergrassLittleseed canarygrassunknown Poaceae Cutleaf groundcherry, Apiaceae Poaceae Solanaceae Euphorbiaceae Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the names, however when noted, the current synonymns were listed in parenthesis. Ocimum americanum x y z Ocimum canum Scientific name Oenothera biennis (syn. Oxalis stricta) Oxalis europaea Panicum capillare Panicum miliaceum T Poa annua – Panicum repens Pascopyrum smithii Physalis minima Phytolacca americana Pilea microphylla Paspalum conjugatum Paspalum notatum Passiflora mucronata Pastinaca sativa Pennisetum purpureum Phalaris minor Phyllanthus maderaspatensis Physalis angulata 172 NEMATROPICA Vol. 39, No. 2, 2009 Mh , graminicola . M –

., 1988 Mg ., 1999 ; ., 2002 mous, 2007). ., 2004 ., 1980 , 2008 , 2008 , 1985 et al et al et al ducted to determine current triticoryzae. et al et al Source/Reference et al . et al. et al. M floridensis alker - edford and Fortum, 1988 . Belair and Benoit, 1996 1967 Davidson and Townshend, Sheridan and Grbavac, 1979 Sheridan and Grbavac, 1979 Belair and Benoit, 1996 Mani and Hinai, 1996 T Belair and Benoit, 1996 Mani and Hinai, 1996 Sellami W Lima Asmus and Andrade, 1997 Lordello Brito 1967 Davidson and Townshend, 1967 Davidson and Townshend, Jaehn Roese and Oliveira, 2004 Roese and Oliveira, 2004 Myers Brito Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M – Mp ; Meloidogyne Mh Mi Mn Mn Mh Mi Ma, Mi Mh Mi Mi, Mj Mj Me Mj Mj Mm Mi Mi Mco Mp Mp Mi Ma, Mi, Mm, Mj, M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family Labiatae M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name M Wild buckwheatLadysthumb Polygonaceae Pale smartweed Polygonaceae Showy purslane Polygonaceae Common purslane Portulacaceae Portulacaceae wild poinsettia Euphorbiaceae Rough cinquefoilCommon shelfheal, allheal Rosaceae unknown Rubiaceae Wild radishBrazil pusleyFlorida pusley Brassicaceae Rubiaceae Rubiaceae - Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the Polygonum convolvulus Polygonum persicaria Polygonum scabrum (syn. Polygonum lapathifolium Portulaca grandiflora Portulaca oleracea Scientific name names, however when noted, the current synonymns were listed in parenthesis. Poinsettia heterophylla (syn. Euphorbia heterophylla) Potentilla norvegica Prunella vulgaris Psychotria nitidula x y z T Raphanus raphanistrum brasiliensis Richardia scabra Richardia – Weed hosts of Meloidogyne: Rich et al. 173 Mh , graminicola . M –

., 1982 Mg ; ., 1999 ., 1999 ., 1999 mous, 2007). ., 1972 ., 1996 et al , 2006 ., 2006 ., 2007 ., 1998 ducted to determine current et al et al et al et al et al triticoryzae. Source/Reference al et al et al. .

et al et M floridensis - . edford and Fortum, 1988 edford and Fortum, 1988 edford and Fortum, 1988 edford and Fortum, 1988 Belair and Benoit, 1996 T 1967 Davidson and Townshend, T Sellami Ciancio and Giudice, 1995 Mani and Hinai, 1996 Sellami Souza Kutywayo and Been, 2006 Moraes Kaur Souza Khan and Murmu, 2004 T Kim 1967 Davidson and Townshend, O’Bannon T Sellami Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M – Mp Mh Ma, Mi Mi Ma, Mi Mj Mj Mj Mi Mm Mc Mj Mf Mi,, Mj, Mm Mm Mj Ma, Mi Mi, Mh Mi Mc Ma, Mi Mi ; Meloidogyne M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name M ellowcress marsh Brassicaceae ellow foxtail Poaceae Y Red sorrelCurly dockLily of the valley vine Polygonaceae Solanaceae Polygonaceae Bristly foxtail Poaceae Common groundsel candlestickEmperor’s Asteraceae Fabaceae SicklepodCoffee sennaPrickly sesbaniaY Foxtail grass Fabaceae Fabaceae Fabaceae Giant green foxtail Poaceae Poaceae Cuban jute Malvaceae Mj Asmus and Andrade 1997 Southern sida Malvaceae Mi Gowda - Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the names, however when noted, the current synonymns were listed in parenthesis. Rorippa islandica (syn. palustris) Rumex acetosella Rumex crispus origanifolia Salpichroa Setaria verticillata x y z Senecio vulgaris Senna alata Senna obtusifolia Senna occidentalis Sesbania aculeate Setaria lutescens Setaria media Setaria viridis Scientific name – T Sida rhombifolia Sida acuta 174 NEMATROPICA Vol. 39, No. 2, 2009 Mh , ., 1977 graminicola . et al M –

., 1982 ., 1998 Mg , 1978 ., 1999 ., 1999 , 2004 ., 1999 ; et al mous, 2007). al al ., 2006

., 2008 et al et et ., 1998 et al. et al et al. ducted to determine current triticoryzae. al et al Source/Reference

. et al et M floridensis - edford and Fortum, 1988 . Zancada Roese and Oliveira, 2004 Rodriguez-Kabana Ciancio and Giudice, 1995 Mani and Hinai, 1996 Kim Curi, 1973 Aragon Ehwaeti Sellami T Davidson and Townshend, 1967 Davidson and Townshend, O’Bannon Davis and Webster, 2003 Davis and Webster, Ehwaeti Davidson and Townshend, 1967 Davidson and Townshend, Belair and Benoit 1996 Asmus and Andrade, 1997 Meyers Roese and Oliveira, 2004 Souza Brito Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M – Mi Mp Ma Mj Mi Mi, Mh Me Me Mi Mi, Mj Ma, Mi Mi Mc Mi Mi Mi Mh Mj Mi Mp Mm Mi, Mj, Mm Mp ; Meloidogyne M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name ater Parsnip Umbeliferae all tumble mustard Brassicaceae M Prickly sida Malvaceae Climbing nightshadeBlack nightshade Solanaceae Solanaceae Red campion Caryophyllaceae T W American black nightshade Solanaceae - Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the names, however when noted, the current synonymns were listed in parenthesis. Sida spinosa x y z Scientific name T Solanum dulcamara Solanum nigrum Silene dioica Sisymbrium altissimum Sium suave Solanum americanum – Weed hosts of Meloidogyne: Rich et al. 175 Mh , graminicola . M –

., 1982 , 2008 ., 1988 Mg ., 1999 ., 1999 ., 2000 ; et al mous, 2007). ., 2004 ., 2006 et al. ., 2004 ., 1985 ., 1985 ., 2008 et al al ., 1998

et al et al et al ducted to determine current et triticoryzae. al et al Source/Reference

et al et al et al . et al et M floridensis - edford and Fortum, 1988 . Amin and Budai,1994 Sellami Souza Sellami Church and Rosskopf, 2005 O’Bannon Roese and Oliveira, 2004 Lordello Lopez and Vilchez, 1991 Dabur Khan 1967 Davidson and Townshend, Kim Lima T Lima Brito Kutywayo and Been, 2006 Boydston Doucet Belair and Benoit, 1996 1967 Davidson and Townshend, Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M – Ma Mi, Mj Mm Mj Ma Mc Mp Mj Me Mg Mg Mi Mi, Mh Me Ma, Mi Me Ma Mc Mc, Mh Mh Mh Mi Mp ; Meloidogyne M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis . Common name aterleaf Portulacaceae ropical soda apple Solanaceae ussock dropseed Poaceae urkeyberry Solanaceae all meadow-rue Ranunculaceae M Common sowthistle Asteraceae W Saltcedar Tamaricaceae T Spiny sowthistle Asteraceae Wild sorghumJohnsongrass Poaceae Poaceae unknownCorn spurryT Common chickweed Apiaceae Caryophyllaceae Caryophyllaceae Fieldnettle betony Lamiaceae Common dandelion Asteraceae T Hairy nightshade Solanaceae T Field pennycress Brassicaceae - Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts alinum triangulare alinum triangulare amarix gallica araxacum officinale Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the Sonchus oleraceus T T Solanum viarum Sonchus asper Sorghum arundinaceum Sorghum halepense Spananthe paniculata Spergula arvensis diander Sporobolus Stellaria media Stachys arvensis T Scientific name names, however when noted, the current synonymns were listed in parenthesis. Solanum torvum Solanum sarrachoides (syn. Solanum physalifolium) x y z T Thalictrum pubescens Thlaspi arvense – 176 NEMATROPICA Vol. 39, No. 2, 2009 Mh , graminicola . M –

., 1999 ., 1999 Mg ., 2004 ; mous, 2007). ., 2008 ., 2008 al et al

et al et ducted to determine current triticoryzae. et al et al Source/Reference . M ovlas and Inserra, 1979 edford and Fortum, 1988 edford and Fortum, 1988 floridensis - Roese and Oliveira, 2004 Ehwaeti Dabur Brito Sellami Mani and Hinai, 1996 Belair and Benoit, 1996 1967 Davidson and Townshend, 1967 Davidson and Townshend, V T Mani and Hinai, 1996 T Brito . Mt M ; -

spp. worldwide. Mf ; z . sp paranaensis . Meloidogyne M Mp Mi Mg Mi Mi Mi, Mj Mh Mi Mi Mn Ma, Mi Mi, Mj Ma, Mi Mi, Mj – Mp ; Meloidogyne M. naasi – Mn ; Mco – M. coffeicola; Me exigua ; Plant Family M. chitwoodi y –

Mc Mm - M. mayaguensis ; ; arenaria

konaensis vain Verbenaceae . Common name r M Coatbuttons Asteraceae White clover Fabaceae Ve Fenugreek Fabaceae Cadillo Malvaceae Inflammation bushGreen field speedwell Asteraceae Purslane speedwell Scrophulariaceae Thymeleaf speedwell Scrophulariaceae Hairy vetch Scrophulariaceae AshwagandhaCommon cocklebur Fabaceae False hawksbeard Asteraceae Solanaceae Asteraceae - Mk ; Meloidogyne –

Ma

javanica . M spp.:

– Mj ; Meloidogyne incognita . M x – Mi ; hapla . nonia cinerea onica agrestis onica peregrina onica serpyllifolia r r r r ithania somnifera M able 1. (Continued) List of wild plant species, common names and botanical families occurring as hosts ridax procumbens rifolium repens rigonella polyceratia erbena officinalis oungia japonica Scientific names are listed as given by the authors although many of have been changed. No systematic attempt was con Common names listed herein were those given by the author or found at Weed Science Society of Americal website (Anony Abbreviations for the names, however when noted, the current synonymns were listed in parenthesis. Scientific name T T x y z V T T lobata Urena Ve Ve Ve Ve Vicia villosa W Xanthium strumarium Y – Weed hosts of Meloidogyne: Rich et al. 177

is highly damaging to coffee (Barbosa et al., Echinochloa colonum (Khan and Murmu, 2004), but only a few weed hosts have been 2004). Khan et al. (2004) studied the weeds described. Lima et al. (1985) found that harboring M. graminicola in India and weed hosts of M. exigua included Amaran- found 17 weeds associated with this nema- thus deflexus, Poinsettia heterophylla, Ipomoea tode in rice fields, eight of which were new acuminata, Leonurus sibiricus, Stachys arven- host records (Agropyron repens, Alternanthera sis, and Taraxacum officinale. Other weed sessilis, Bothriochloa intermedia, Brachiaria hosts of M. exigua include Commelina dif- ramosa, Dactyloctenium aegyptium, Digitaria fusa, Cyperus rotundus, and Solanum nigrum sanguinalis, Physalis minima, and Sporobolus (Aragon, 1978), Citrullus vulgaris (Moraes diander). The egg-laying behavior of et al., 1972) and Spananthe paniculata females of M. graminicola differed with the (Lopez and Vilchez, 1991). weed host species. In some of the weeds, the female nematode produced egg masses Meloidogyne floridensis within the galls, while others such as Cype- rus rotundus, Echinochloa colonum, Physalis This root-knot nematode was recently minima and Sporobolus diander had small egg described as a new species infecting peach masses on the root gall; however in Agera- in Florida, USA (Handoo et al., 2004) and tum conyzoides, heavy root galling and large is known to occur only in the United States egg masses were observed (Khan et al., (Brito et al., 2008). The nematode was 2004). This nematode was also found reported by Brito et al. (2008) to infect infecting Cyperus rotundus in Florida (Brito Emilia sonchifolia, while Stanley et al. (2006) et al., 2008). found that two other weeds, Ipomoea qua- moclit and Nasturtium officinalis, were hosts of M. floridensis. In another study, Kaur et al. Meloidogyne hapla (2007) found that Amaranthus spinosus, A. The northern root-knot nematode is an retroflexus, Cnidoscolus stimulosus, Dichondra important vegetable crop pest and is a repens, Ipomea triloba, I. violacea, Leonotis major limiting factor in temperate regions nepetaefolia and Phytolacca americana were of the world (Melakeberhan et al., 2007; good hosts. Mitkowski et al., 2002; Potter and Olthof, 1993; Vrain, 1982). Many weed species are Meloidogyne graminicola highly susceptible to M. hapla and serve as The rice root-knot nematode has been alternative hosts in agricultural areas (Ben- reported from several rice growing coun- dixen, 1986b; Edwards and Jones, 1984; tries of the world, especially in light tex- Gaskin and Crittenden, 1956). In green- tured soils and irrigated upland conditions house tests, Belair and Benoit (1996) where it causes damage to rice in both found high populations of M. hapla on nurseries and field plantings (Panwar and Bidens cernua, B. frondosa, B. vulgata, Matri- Rao, 1998; Rao et al., 1986). This nematode caria matricarioides, Pastinaca sativa, had been found infecting weed species Polygonum scabrum, Sium suave, and Thlaspi associated with rice cultivation including arvense. Moderate reproduction of M. hapla Cyperus deformis, C. iria, Dichanthium annula- was supported by Capsella bursa-pastoris, tum, Echinocloa colona, E. crus-galli, Eclipta Chrysanthemum leucanthemum, Gnaphalium alba, Melilotus alba, and Trigonella polyceratia uliginosum, Stellaria media, Thalictrum pubes- (Bajaj and Dabur 2000; Dabur et al., 2004; cens, and Veronica agresti while Chenopodium Sperandio and Amaral, 1994), and also album, C. glaucum, Erysimum cheiranthoides, 178 NEMATROPICA Vol. 39, No. 2, 2009

Eupatorium maculatum, Polygonum convolvu- cosa and Sida acuta supported moderate lus, Portulaca oleracea, and Rorippa islandica galling and small to large galls, respectively. reportedly sustained low reproduction of Other weeds (Achyranthes aspera, Borreria this nematode. Roots of Eupatorium macula- hispida, Emelia sonchifolia, Euphorbia hirta, tum and Thalictrum pubescens exhibited no Lactuca runcinata, and Lucas aspera) also distinct galling caused by M. hapla but sup- were found to be hosts of M. incognita. In ported low and moderate populations of an extensive survey in the Sultanate of the nematode on the two weed species, Oman, Mani and Hinai (1996) observed 30 respectively. Interestingly, M. hapla weeds as hosts of M. incognita and M. javan- induced galls on Senecio vulgaris but no ica. Among those found as hosts of M. eggs or juveniles were produced. Some incognita were Achyranthes aspera, Aerva jav- non-host weed species of M. hapla were anica, Alternanthera sessilis, Amaranthus Amaranthus retroflexus, Ambrosia artemisiifo- hybridus, A. graecizans, Chamaesyce hirta, Che- lia, Echinochloa crus-galli, and Setaria viridis. nopodium murale, Datura metel, Portulaca Taraxacum officinale was also reported as a grandiflora, P. oleracea, Phyllanthus mad- good host of M. hapla (Doucet et al., 2000) eraspatensis, Solanum nigrum and Withania and poses a serious threat to agriculture somnifera. Cyperus esculentus and C. rotundus production in Argentina since it is widely were reported as hosts of M. incognita, and disseminated in the country. these weeds produced a pest complex that is a serious threat to cotton in Georgia Meloidogyne incognita (Bird and Hogger, 1973) and chile pepper production in New Mexico USA The southern root-knot nematode is (Schroeder et al. 1999; Thomas et al. 1997; widely distributed throughout the world 2004). Myers et al. (2004) found M. incog- and has a wide range of weed hosts (Alam nita associated with major weeds in Florida and Khan, 1976; Brito et al., 2008; Krishna- USA, especially, Cyperus esculentus, C. rotun- Murthy and Elias, 1968). In greenhouse dus, Cynodon dactylon, Richardia scabra and tests, 34 species of weeds in 32 genera Solanum americanum. In Mexico, M. incog- belonging to 19 botanical families were nita and unidentified Meloidogyne spp. were found to be good hosts of M. incognita found on Amaranthus graecizans in irrigated (Davidson and Townshend, 1967). In large and dry lands of the states of Aguascali- greenhouse tests, Gaskins (1958) found entes, San Luis Potosi and Zacatecas that 70 out of 103 weed species tested were (Velasquez-Valle, 2001). Nisha and Sheela hosts to Meloidogyne incognita as measured (2002) reported Emilia sonchifolia as a new by presence of mature females. Among the host of M. incognita in India and suggested most highly susceptible weeds were Ampela- that M. incognita was the major nematode mus albidus, Amaranthus graecizans, A. retrof- infesting crops and weeds in that country. lexus, Ipomoea hederacea, Lactuca scariola, Plantago lanceolata, Polygonum aviculare, P. Meloidogyne javanica persicaria, and Solanum dulcamara. Gowda et al. (1995) assessed the susceptibility of the The Javanese root-knot nematode is 22 common weeds found to grow in tropical and subtropical in distribution and tobacco fields in India and observed that has a broad host range (Taylor and Sasser, Celosia argentea and Ocimum canum were 1978). Asmus and Andrade (1997) found highly infected and showed gall sizes rang- that Amaranthus hybridus, Bidens pilosa, Digi- ing from small to medium while Cleome vis- taria horizontalis, Euphorbia heterophylla, Sida Weed hosts of Meloidogyne: Rich et al. 179

rhombifolia and Solanum americanum served reported as hosts of M. mayguensis (Souza et as reservoir hosts for M. javanica. Since this al., 2006). Similarly, 10 other weed species nematode multiplied on all the weed spe- were identified as good hosts of this nema- cies, control of weeds, especially A. hybri- tode including Abutilon theophrasti, Amaran- dus, E. heterophylla and S. americanum, was of thus retroflexus, A. spinosus, Cnidoscolus high importance for preventing multiplica- stimulosus, Dichondra repens, Ipomoea triloba, tion of M. javanica. Other weeds reported I. violacea, Leontodon hispidus, Phytolacca as hosts of M. javanica are Gutenbergia cordi- americana and Senna obtusifolia (Kaur et al., folia in Kenya (Desaeger and Rao, 2000), 2007). Meloidogyne mayaguensis was also Melilotus alba in Argentina (Lorenzo et al., found infecting Eclipta prostrate and Portu- 2002), Portulaca oleracea in Australia laca oleracea under field conditions (Brito et (Walker et al., 2002), and Sesbania aculeata al., 2008). in India (Khan and Murmu, 2004). Meloidogyne naasi Meloidogyne konaensis The barley root-knot nematode has The Kona coffee root-knot nematode only been studied as a host of a few weed was first found infecting coffee plantations species. It was found infecting Coronilla scro- in Hawaii (Eisenback et al., 1994) but also pioides, Medicago hispida, Melilotus sulcata, reproduced heavily on tomato over a wide and Vicia villosa in a wheat field in Italy temperature range (Zhang, 1994). Only (Vovlas and Insera, 1979) and was present one weed species, Cyperus rotundus, has on Glyceria fluitans and Polygonum persicaria been observed as host of this nematode in New Zealand (Sheridan and Grbavac, (Zhang and Schimtt, 1994). Further stud- 1979) and Chenopodium hybridum, C. album ies of host suitability of this nematode and Digitaria sanguinalis in Hungary (Amin among plants should be useful for crop and Budai, 1994). selection in areas adjacent to coffee plant- ings. Meloidogyne paranaensis This nematode was first found infect- Meloidogyne mayaguensis ing coffee plants in Parana state, Brazil, The guava root-knot nematode (Souza and as such, was named the Parana coffee et al., 2006) was originally described from root-knot nematode (Carneiro et al., eggplant (Solanum melongena) collected in 1996). The major weed hosts of M. para- Puerto Rico (Rammah and Hirschmann, naensis include Cyperus rotundus, Echi- 1988). Since then, this nematode has been nochloa colonum, Eleusine indica, Galinsoga found to be pathogenic to many economi- ciliata, Ipomoea grandifolia, Raphanus cally important crops including coffee, raphanistrum, Solanum americanum and Sor- guava, pepper, tomato, soybean and also ghum arundinaceum while Acanthospermum ornamental plants in several regions of the australe, Biden pilosa, Cenchrus echinatus, world (Brito et al., 2007; 2008; Carneiro et Conyza bonariensis, Desmodium tortuosum, al., 2001; Decker and Fuentes, 1989; Far- Digitaria horizontalis, Mimosa invisa, Sonchus gette et al., 1996; Molinari et al., 2005; Will- oleraceus and Spermacoce latifolia were non- ers, 1997). Biden pilosa was the first weed hosts of this nematode (Roese and species identified as a host of M. mayaguen- Oliveira, 2004). Impatiens balsamina was sis (Willers, 1997). Recently, 14 additional also found to be a host of M. paranaensis weed species in 12 botanical families were (Campos and Villain, 2005). 180 NEMATROPICA Vol. 39, No. 2, 2009

Meloidogyne triticoryzae Digitaria sanguinalis was reported to be a host of M. incognita (Myers et al., 2004; Nol- This root-knot nematode was first ing and Gilreath, 2002), but no reproduc- described from India and was found infect- tion of M. incognita race 4 was reported in ing wheat and rice (Gaur et al., 1993). another study (Kaur et al., 2007). Weeds known to be good hosts of this nem- It is important to also remember that atode are Cyperus rotundus, Echinochloa some weeds are NOT hosts of plant-para- colonum, Echinochloa crus-galli, Leptochloa sitic nematodes, a fact that may be useful in coloniculus and Phalaris minor (Gaur and management programs. For example, Sharma, 1998). Rhoades (1976) reported that a summer cover crop of hairy indigo (Indigofera hir- DISCUSSION suta) was a non-host to Meloidogyne incognita and M. javanica as well as the sting nema- Information on the host range of tode (Belonolaimus longicaudatus). In field Meloidogyne spp. is very incomplete and experiments, hairy indigo was as effective sometimes contradictory; and many addi- as soil fumigation for control of those nem- tional studies are necessary to adequately atode species in subsequent vegetable crop describe this subject. For example, pub- production. lished research to determine weed hosts Knowledge about weeds as hosts of was only found on about 14 of the 96 nom- Meloidogyne spp., particularly weeds known inal species of Meloidogyne. In addition, to be highly symptomatic hosts of root-knot there are 3479 recognized weed species in nematodes, makes it possible to use exist- the Weed Science Society of America data- ing weeds to monitor fields for those nem- base (Anonymous, 2007), thus adding tre- atodes. This is especially important when mendously to the work for Nematologists. laboratory assay is impractical or when Weeds, like crop plants, differ widely in more data points on Meloidogyne infestation their host status to individual Meloidogyne are needed than can be derived by labora- spp. For example, Crotalaria spectabilis was tory soil assay alone. For instance, a wild found to be a good host of M. arenaria and Citrullus sp. has been used to monitor M. incognita (Tedford and Fortnum, 1988) Meloidogyne arenaria race 2 in North Flor- but immune to M. mayaguensis (Guimarães ida, USA, and several Leguminosae were et al., 2003). Two other factors, Meloidogyne used to index a Meloidogyne sp. infestation spp. race and weed biotypes, also compli- in fields to be planted to melons Cucumis cate the host status designation of weeds to melo in Guatemala where laboratory soil Meloidogyne spp.; therefore, localized obser- assay was unavailable (R. A. Dunn, pers. vations and tests will be needed in addition comm.). to simple host range tests (Griffin, 1982). The host status and ultimately the For example, different authors have found degree of nematode reproduction on differing results on host ranges of weeds weeds should be a major concern in inte- such Senna obtusifolia. Rodriguez-Kabana et grated nematode management recommen- al. (1977) found M. arenaria race 1 causing dations since weeds are almost universally severe root galling on this weed, but others present during crop growth and afterwards have reported that M. arenaria race 1 (Kaur in fallow periods. This statement seems et al., 2007) and M. arenaria race 2 (Tedford contrary to modern agricultural percep- and Fortnum, 1988) were not found to tions of many scientists and farmers who reproduce on Senna obtusifolia. Similarly, now consider that weeds are not major con- Weed hosts of Meloidogyne: Rich et al. 181

straints in commercial production due to vides further evidence that weed manage- the excellent control provided by herbi- ment both within and after the normal cides. However, weed control is often con- cropping cycle is an overlooked yet critical ducted relative to weed populations and component of nematode management sys- threshold levels established for weed/crop tems. Without a strong weed management competition. If weed populations are rela- program, the benefits of crop rotation for tively low, it may be viewed as unprofitable nematode management can be quickly to perform weed control, regardless of annulled by weed hosts of plant-parasitic whether these weeds host parasitic nema- nematodes. Recommendations for crop todes. More commonly, limited resource rotation by Nematologists should also growers, particularly those in the subtrop- include clear statements about the neces- ics and tropics, often are not able to man- sity of controlling weeds, particularly in age weeds as effectively during the those periods between cropping cycles. cropping season. Additionally, both large- scale commercial and limited resource LITERATURE CITED growers routinely have long fallow periods Alam, M. M., and A. Khan. 1976. New host records of between crops, when weeds continue to the root-knot nematode in North India. Indian grow. These fallow periods may last from 3- Phytopathology 28:540-541. 6 months and are natural in most crop pro- Amin, A. W., and C. S. Budai. 1994. Some weed host plants of the root-knot nematode, Meloidogyne duction cycles. For example, in northern species, in south-eastern Hungary. Pakistan Jour- Florida USA, thousands of acres of cotton nal of Nematology 12:59-65. and peanut are harvested in September Anderson, W. P. 1996. Weed Science: Principles and and October each year, and the land is applications. 3rd Ed. Pp. 3-13. Brooks Cole Pub- then not used for crop production for over lishing, St. Paul, Minnesota USA. Anonymous. 2007. Composite List of Weeds. Stan- 6 months until planting commences the dardized Plant Names Subcommittee, Weed Sci- following May. ence Society of America. http://www.wssa.net/ Hence, agricultural land devoted to Weeds/ID/WeedNames/namesearch.php annual crops remains idle for periods of Aragon, B. C. A., M. B. Gomes, and J. E. Caicedo. several months between crops, allowing 1978. Plantas de la zona cafeteria colombiana hospedantes de espécies de Meloidogyne goeldi, prolific weed growth. A solution that may 1887. Cenicafé 29:35-45. seem simple is to control weeds, particu- Asmus, G. L., and P. J. M. Andrade. 1997. Reproduc- larly in the off-cropping season. However, tion of Meloidogyne javanica in some weeds spe- this could lead to increased grower cost, cies frequently found in the western region of greater soil erosion potential, less nutrient Brazil. Comunicado-Tecnico-EMBRAPA Centro de Pesquisa Agropecuaria do Oeste 19:3. recycling, and lower soil organic matter lev- Bajaj, H. K., and K. R. Dabur. 2000. Cyperus deformis, a els. Research is necessary to develop nema- new host record of rice root-knot nematode, Me- tode management options such as loidogyne graminicola. Indian Journal of Nematol- selectively eliminating major weed hosts ogy 30:256. with herbicides, encouraging the growth of Barbosa, D. H. S. G., H. D. Vieira, R. M. Souza, A. P. Vi- ana, and C. P. Silva. 2004. Field estimates of cof- non-host weeds, or planting cover crops fee yield losses and damage threshold by Meloid- that could suppress weed populations. ogyne exigua. Nematologia Brasileira 28:49-54. Results from this review indicate that the Barrentine, W. L. 1974. Common cocklebur competi- presence of weeds any time during the year tion in soybeans. Weed Science 22:600-603. are likely to compromise carefully docu- Belair, G., and D. L. Benoit. 1996. Host susceptibility of 32 common weeds to Meloidogyne hapla in or- mented and effective rotation systems for ganic soils of southwestern Quebec. Journal of nematode management. This review pro- Nematology 28(4S):643-647. 182 NEMATROPICA Vol. 39, No. 2, 2009

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Received: Accepted for publication: 24/VIII/2009 10/XXI/2009 Recibido: Aceptado para publicacion: