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Full-Text (PDF) J. Crop Prot. 2020, 9 (2): 183-193______________________________________________________ Review Article An updated checklist of Meloidogyne Göldi, 1887 species, with a diagnostic compendium for second-stage juveniles and males Reza Ghaderi1* and Gerrit Karssen2 1. Department of Plant Protection, School of Agriculture, Shiraz University, 71441-65186, Shiraz, Iran. 2. National Plant Protection Organization, Wageningen Nematode Collection, 6700 HC Wageningen, The Netherlands. Abstract: The cosmopolitan root-knot nematodes of the genus Meloidogyne are considered as the most important plant-parasitic nematodes damaging almost all higher plant species. This paper provides an updated checklist of 105 valid species of Meloidogyne, along with their synonyms and non-valid members. Furthermore, species are grouped based upon their diagnostic characteristics of second-stage juveniles (tail shape, tail length, stylet length and dorsal gland orifice (DGO) distance from stylet base) and males (number of lateral incisures, stylet length, DGO and spicules length). Keywords: Identification, Meloidogyne, Morphology, root-knot nematodes, species list, taxonomy Introduction12 that time. Subsequent reviews on the genus systematics were carried out during 1976- Root-knot nematodes are included in the 2002, providing identification keys and genus Meloidogyne Göldi, 1887 and diagnostic compendia for its species (Esser constitute a relatively small, but important et al., 1976; Hewlet and Tarjan, 1983; polyphagous group of highly adapted Jepson, 1987; Eisenback and Triantaphyllou, Downloaded from jcp.modares.ac.ir at 13:58 IRST on Friday October 1st 2021 obligate plant parasites (Karssen et al., 1991; Karssen and Hoenselaar, 1999; 2013). Their taxonomic position at family Karssen, 2002). The list of known species, level has been the subject of discussion for with additional notes on the most important many years, and these nematodes have been species, has been published by different confused with cyst nematodes nematologists, with the number of valid (Heteroderidae Filipjev and Schuurmans species reaching54 in Jepson (1987), 68 in Stekhoven, 1941) for a long time, and Luc et al. (1988), 62 in Eisenback and assigning the same or separate families for Triantaphyllou (1991), 80 in Siddiqi (2000), cyst and root-knot nematodes was 81 in Karssen (2002), 88 in Andrássy controversial. (2007), 97 in Hunt and Handoo (2009) and Chitwood (1949) revised Meloidogyne 98 in Karssen et al. (2013). taxonomy for the first time and developed a The present study provides an updated list key for five species (and one subspecies) at including 105 valid species with their synonyms, as well as lists of species inquirendae and nominanuda. Moreover, a Handling Editor: Majid Pedram grouping of recognized species is presented __________________________ * Corresponding author, e-mail: [email protected] based on diagnostic characters of second- Received: 31 July 2019, Accepted: 26 November 2019 stage juveniles and males. Published online: 16 January 2020 183 An updated checklist of Meloidogyne ________________________________________________ J. Crop Prot. Diagnosis of Meloidogyne Göldi, 1887 lips. Six inner labial sensilla centered around Family Meloidogynidae (Skarbilovich, 1959) the stoma and one cephalic sensillum present Wouts, 1973 on each median lip. Large slit-like amphidial Subfamily Meloidogyninae Skarbilovich, 1959 openings located between labial disc and = Meloidogynini (Skarbilovich, 1959) Coomans, lateral lips. The latter reduced or absent in 1979 some species. Lip region sometimes set off = Meloidoderellinae Husain, 1976 and/or partly subdivided by transverse = Meloidoderellini (Husain, 1976) Coomans, 1979 incisures. Cephalic framework and straight Genus Meloidogyne Göldi, 1887 stylet well developed, the latter ranging in = Caconema Cobb, 1924 length between 10-33 µm. DGO at 1-13 µm = Spartonema Siddiqi, 1986 behind the stylet knobs. Metacorpus smaller = Hypsoperine (Spartonema) Siddiqi, 1986 than that in females. Secretory-excretory pore = Hypsoperine Sledge and Golden, 1964 and hemizonid between metacorpus level and = Hypsoperine (Hypsoperine) (Sledge and the ventrally-overlapping pharyngeal glands. Golden, 1964) Siddiqi, 1986 Hemizonid anterior or sometimes posterior to the pore. Pharyngeal glands nuclei reduced to Female. Sedentary, white to transparent, two. A single long testis usually present, rounded to pear-shaped with protruding neck. rarely two reduced ones were observed. Body length from 350 µm to 3 mm. Cuticle Lateral field in most species with four, up to annulated, posteriorly with a characteristic 15 incisures in certain species, outer bands unique pattern around the perineum called the often areolated. Tail short, bluntly rounded. perineal pattern. Anus and vulva terminal, Bursa lacking. Phasmids near cloaca. phasmids near the anus, the latter usually Spicules slender, 18-47 µm long; covered with cuticular fold. The perineum gubernaculum crescentic, about 10 µm long. sometimes slightly elevated. Head not or slightly set off, cephalic framework distinct Second-stage juvenile. Vermiform, infective but delicate. Labial disc not, to slightly stage. Body length 250-600 µm. Cuticle raised, fused with median and lateral lips. annulated. Lip region structure as in males, Two slit-like amphidial openings and ten but much smaller and with weakly sclerotized small labial sensilla present. Stylet delicate, cephalic framework. Straight stylet, about 7- Downloaded from jcp.modares.ac.ir at 13:58 IRST on Friday October 1st 2021 ranging in length from 8-25 µm, cone in most 23 µm long. DGO 1-8 µm behind the knobs. species slightly curved dorsally, shaft straight Metacorpus not well developed. Hemizonid with three knobs. Dorsal pharyngeal gland anterior to, at the same level or posterior to orifice (DGO) 2-12 µm behind the knobs. the secretory-excretory pore. Three Secretory-excretory pore between lip region pharyngeal glands ventrally overlapping the and metacorpus level. Metacorpus relatively intestine. Rectum often inflated. Tail 13-115 large, pharyngeal glands variable in size and µm long, tapering towards hyaline tail part. shape, ventrally overlapping the intestine. Lateral field with four incisures. Reproductive system didelphic, the gonads long, partly convoluted. Most of the Third and fourth-stage juvenile. Sedentary unembryonated eggs deposited in an egg-sac, inside root and swollen, lacking stylet, develop formed by six large rectal glands and secreted within second-stage cuticle. through the anus. Male. Vermiform, non-sedentary, ranging in List of Meloidogyne species length between 600-2500 µm. Cuticle annulated. Lip region composed of a cap and Type species post-labial annulus (annuli). Head cap with Meloidogyne exigua (Göldi, 1887) Chitwood, rounded labial disc and four fused median 1949 184 Ghaderi and Karssen _______________________________________________ J. Crop Prot. (2020) Vol. 9 (2) = Heterodera exigua (Göldi, 1887) Marcinowski, M. cirricauda Zhang and Weng, 1991 1909 M. citri Zhang, Gao and Weng, 1991 M. coffeicolaLordello and Zamith, 1960 Other species = Meloidodera coffeicola (Lordello and Zamith, M. aberrans Tao, Xu, Yuan, Wang, Lin, Zhuo 1960) Kirjanova, 1963 and Liao, 2017 M. cruciani Garcia-Martinez, Taylor and Smart, M. acronea Coetzee, 1956 1982 = Hypsoperine acronea (Coetzee, 1956) Sledge M. cynariensis Fam-Tkhan-Bihn, 1990 and Golden, 1964 M. daklakensis Trinh, Le, Nguyen, Nguyen, = Hypsoperine (Hypsoperine) acronea Liébanas and Nguyen, 2018 (Coetzee, 1956) Siddiqi, 1986 M. decalineata Whitehead, 1968 M. actinidiae Li and Yu, 1991 M. dimocarpus Lio and Zhang, 2001 M. aegracyperi Eisenback, Holland, Schroeder, M. donghaiensis Zheng, Lin and Zheng, 1990 Thomas, Beacham, Hanson, Paes-Takahashi & M. dunensisPalomares-Rius, Vovlas, Troccoli, Vieira, 2019 Liébanas, Landa and Castillo, 2007 M. africana Whitehead, 1960 M. duytsi Karssen, van Aelst and van der M. aquatilis Ebsary and Eveleigh, 1983 Putten, 1998 M. arabicida López and Salazar, 1989 M. enterolobii Yang and Eisenback, 1983 M. ardenensis Santos, 1968 = M. mayaguensis Rammah and Hirschmann, = M. deconincki Elmiligy, 1968 1988 = M. litoralis Elmiligy, 1968 M. ethiopica Whitehead, 1968 M. arenaria (Neal, 1889) Chitwood, 1949 = M. brasilensis Charchar and Eisenback, 2002 = Anguillula arenaria Neal, 1889 M. fallax Karssen, 1996 = Tylenchus arenarius (Neal, 1889) Cobb, 1890 M. fanzhiensis Chen, Peng and Zhang, 1990 = Heterodera arenaria (Neal, 1889) M. floridensis Handoo, Nyczepir, Esmenjaud, Marcinowski, 1909 van der Beek, Castagnone-Sereno, Carta, = M. arenaria arenaria (Neal, 1889) Chitwood, Skantar and Higgins, 2004 1949 M. fujianensis Pan, 1985 = M. thamesi (Chitwood in Chitwood, Specht = M. fujianensis Cangsang, Jing and and Havis, 1952) Goodey, 1963 Shengyuan, 1988 Downloaded from jcp.modares.ac.ir at 13:58 IRST on Friday October 1st 2021 = M. arenaria thamesi Chitwood in Chitwood, M. graminicola Golden and Birchfield, 1965 Specht and Havis, 1952 M. graminis (Sledge and Golden, 1964) = M. thamesi gyulai Amin, 1993 Whitehead, 1968 = M. gyulai Amin, 1993 = Hypsoperine graminis Sledge and Golden, M. artiellia Franklin, 1961 1964 M. baetica Castillo, Vovlas, Subbotin and = Hypsoperine (Hypsoperine) graminis (Sledge Troccoli, 2003 and Golden, 1964) Siddiqi, 1986 M. brevicauda Loos, 1953 M. hainanensis Liao and Feng, 1995 M. californiensis Abdel-Rahman and Maggenti, M. haplaChitwood, 1949 1987 M. haplanaria Eisenback, Bernard, Starr, Lee M. camelliae Golden, 1979 and Tomaszewski,
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