A Key and Diagnostic Compendium to the Species of the Genus Hoplolaimus Daday, 1905 (Nematoda: Hoplolaimidae)
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Journal of Nematology 24(1):45-53. 1992. A Key and Diagnostic Compendium to the Species of the Genus Hoplolaimus Daday, 1905 (Nematoda: Hoplolaimidae) ZAFAR A. HANDOO AND A. MORGAN GOLDEN 1 Abstract: An identification key to 29 valid species of Hoplolaimus is given. A compendium of the most important diagnostic characters for use in identification of species is included as a practical alternative and supplement to the key. Diagnosis of Hoplolaimus is emended and lists of species of the genus, their synonymies, species inquirendae, nomina nuda, and species transferred to other genera are given. Hoplolaimus sheri, H. chambus, H. casparus, and H. capensis are recognized as valid species. Key words: Compendium, diagnostic, Hoplolaimidae, Hoplolaimus, identification, key, lance nema- tode, morphology, Nematoda, taxonomy. The genus Hoplolaimus was established lies were included within Hoplolaimidae: on the basis of a single female by Daday in Hoplolaiminae with eight valid genera, in- 1905 when he described H. tylenchiformis cluding Hoplolaimus, and Rotylenchinae from a mud hole on Banco Island in the with three genera. Paraguay river at Asuncion, Paraguay (1). The purpose of this study is to examine Sher (11) provided the first comprehen- in detail representative specimens and sive work on the genus, giving historical, published data on Hoplolaimus species, to morphological, and systematic sections determine the interrelationship of the spe- and also a key to eight valid species. cies, to define the valid and most signifi- Golden (4) included Hoplolaimus under the cant differentiating characters, and to pre- subfamily Hoplolaiminae Filipjev, 1934 pare a new key and compendium to facil- along with Aorolaimus Sher, 1963, Scutel- itate identification of the species. lonema Andr/tssy, 1958, and Peltamigratus Sher, 1963. Krall (6) reviewed all the pre- MATERIALS AND METHODS vious work done on the family Hoplolaim- Paratype specimens of 10 species (H. ae- idae Filipjev, 1934 and provided details on gypti, H. aorolaimoides, H. californicus, H. the morphology and development, phylo- clarissimus, H. columbus, H. concaudajuven- genetic relationship, and host-parasite re- cus, H. indicus, H. magnistylus, H. sheri, and lationship of the species. He accepted H. stephanus) and nontype specimens of three subfamilies within Hoplolaimidae: five other species (H. galeatus, H. pararobus- Hoplolaiminae Filipjev, 1934 with four tus, H. seinhorsti, H. seshadrii, and H. tylenchi- genera; Rotylenchinae Golden, 1971 with formis) were examined from the USDA three genera; and Rotylenchoidinae Nematode Collection at Beltsville, Mary- Whitehead, 1958 with two genera. He also land. Specimens of each species, which gave a key to 18 species of Hoplolaimus. ranged in numbers from 1 to over 50, were Fortuner (3) more recently presented a di- mounted in glycerine and accompanied by agnosis and systematic relationship of the pertinent records. Examinations were family Hoplolaimidae. Only two subfami- made with a compound light microscope, usually at highest magnification, and mor- Received for publication 12 April 1991. phometric data were obtained with an eye- i Microbiologist and Zoologist/Collaborator, USDA ARS, piece micrometer. In evaluation of the spe- Nematology Laboratory, Plant Sciences Institute, Beltsville Agricultural Research Center, 10300 Baltimore Ave., Belts- cies, our own data and the original descrip- ville, MD 20705-2350. tions of most species, as well as any We thank Donna M. S. Ellington, Support Scientist, Nema- tology Laboratory, USDA ARS, Beltsville, for technical assis- subsequent redescription or other relevant tance. data, were utilized. All measurements are 45 46 Journal of Nematology, Volume 24, No. 1, March 1992 in micrometers (~m) unless otherwise Lip region high, offset, with prominent stated. transverse and longitudinal striae (except H. cephalus); cephalic framework massive, SYSTEMATICS basal annule divided into squares. Styler massive, 31-61 txm, with compact tulip- Genus Hoplolaimus Daday, 1905 shaped basal knobs bearing anterior tooth- Syn. Nemonchus Cobb, 1913 like projections (except H. aorolaimoides Hoplolaimoides Shakil, 1973 with poorly developed anterior projec- Basirolaimus Shamsi, 1979 tions). Dorsal esophageal gland opening (Diagnostic data on females and male near base of spear knobs (25% or less of spicules in Table 2.) spear length). Excretory pore posterior to Emended diagnosis: Hoplolaiminae. Fe- hemizonid in species with four lines in lat- male. Large sized nematodes (1-2 mm). eral field (except H. sacchari), anterior to TABLE 1. Hoplolaimus species transferred to other genera with their present names and status. Species Present name and status H. aberrans Whitehead, 1960 Scutellonema clathricaudatum (Whitehead, 1959) Germani et al., 1986 H. annulifer de Man, 1921 Criconema annuliferum (de Man, 1921) Micoletzky, 1925 H. aquaticus (Micoletzky, 1913) Hemicycliophora aquatica (Micoletzky, 1913) Loos, 1948 Menzel, 1917 H. bradys Steiner & LeHew, ScuteUonema bradys (Steiner & LeHew, 1933) Andr~issy, 1958 1933 H. gracilidens Sauer, 1958 Rotylenchus graciliden~ (Sauer, 1958) Sauer, 1958 H. quernei (Certes, 1889) Criconema giardi (Certes, 1889) Micoletzky, 1925 Menzel, 1917 (partially) H. quernei nec Certes, 1889 from Ogma menzeli (Stefanski, 1924) Schuurmans Stekhoven & Schneider, 1923; from Teunissen, 1938 Stefanski, 1924 H. quernei (Certes, 1889) Ogma menzeli (Stefanski, 1924) Schuurmans Stekhoven & Menzel, 1917 (partially) Teunissen, 1938 H. heideri (Stefanski, 1916) Criconemoides heideri (Stefanski, 1916) Taylor, 1936 (Now species Menzel, 1917 inquirenda) H. informis Micoletzky, 1922 Criconemella informi~ (Micoletzky, 1922) Luc & Raski, 1981 H. leiomeru~ de Guiran, 1963 Aorolaimus leiomerus (de Guiran, 1963) de Guiran & Sher, 1968 H. menzeli Stefanski, 1924 Ogma menzeli (Stefanski, 1914) Schuurmans Stekhoven & Teunissen, 1938 H. morgen6e (Hofm~inner in Criconemoides morgense (Hofm~inner in Hofm~nner & Menzel, HofmSnner & Menzel, 1914) 1914) Taylor, 1936 (Now species inquirenda) Taylor, 1936 H. murrayi (Southern, 1914) Ogma murrayi Southern, 1914 Menzel, 1917 H. octangularis (Cobb, 1914) Ogma octangularis (Cobb, 1914) Schuurmans Stekhoven & Menzel, 1917 Teunissen, 1938 H. rusti¢us (Micoletzky, 1915) Criconemella rustica (Micoletzky, 1915) Luc & Raski, 1981 Menzel, 1917 H. ru6ticus var. peruensis Steiner, Criconemella peruensis (Steiner, 1920) Luc & Raski, 1981 1920 H. similis (Cobb, 1918) Criconemoides similis (Cobb, 1918) Chitwood, 1949 (Now species Micoletzky, 1922 inquirenda) H. sinensis Rahm, 1937 Criconemoides sinensis (Rahm, 1937)Goodey, 1951 (Now species inquirenda) H. squamosus (Cobb, 1913) Hemicriconemoides squamosus (Cobb, 1913) Siddiqi & Goodey, 1963 Menzel, 1917 (Now species inquirenda) H. thienemanni Schneider, 1925 Hemicycliophora thienemanni (Schneider, 1925) Loos, 1948 H. uniformis Thorne, 1949 Rotylenchus robust~ (de Man, 1876) Filipjev, 1936 H. zavadskii Tulaganov, 1941 Criconemella zavadskii (Tulaganov, 1941) De Grisse & Loof, 1965 Hoplolaimus Key and Compendium: Handoo, Golden 47 hemizonid in species with no or fewer than Hoplolaimus californicus (Sher, 1963) four incisures. Esophagus well developed; Siddiqi, 1986 esophageal glands overlapping intestine 6. H. capensis Van den Berg & Heyns, 1970 dorsally and laterally, containing 3-6 nu- H. pararobustus by Siddiqi, 1986 clei. Epiptygma present, distinct or indis- 7. H. casparus Van den Berg & Heyns, 1970 tinct, single or double. Vulva median; two H. pararobustus by Siddiqi, 1986 outstretched genital branches. Tail short, 8. H. cephalus Mulk & Jairajpuri, 1976 terminus hemispherical to bluntly B. cephalus (Mulk & Jairajpuri, 1976) rounded, annulated. Phasmids large, Shamsi, 1979 scutellum-like, one prevulval, one postvul- H. cephalus (Mulk & Jairajpuri, 1976) val (in H. abelmoschi adjacent to vulval area, Luc, 1981 one anterior and one posterior; in H. B. cephalus by Siddiqi, 1986 puertoricensis prevulval; and in H. californi- 9. H. chambus Jairajpuri & Baqri, 1973 cus and H. tabacum postvulval). Lateral B. chambus (Jairajpuri & Baqri, 1973) field with 0-4 incisures. Shamsi, 1979 Male. When present, generally similar to H. chambus (Jairajpuri & Baqri, 1973) females (except for reproductive struc- Luc, 1981 tures) but slightly smaller. Tail short. Spi- H. columbus by Khan, 1978 in Siddiqi, cules well developed, arcuate with distal 1986 flanges, variable in size. Bursa extending 10. H. citri (M. L. Khan & S. H. Khan, to tail tip. Gubernaculum large, pro- 1985) n. comb. trusible, with titillae and telamon. B. citri M. L. Khan & S. H. Khan, 1985 11. H. clarissimus Fortuner, 1973 LIST OF VALID HOPLOLAIMUS SPECIES B. clarissimus (Fortuner, 1973) Shamsi, 1979 (Hoplolaimus species transferred to other H. clarissimus by Luc, 1981 genera with their present name and B. clarissimus by Siddiqi, 1986 status in Table 1.) 12. H. columbus Sher, 1963 B. columbus (Sher, 1963) Shamsi, 1979 Type species H. columbus by Luc, 1981 1. Hoplolaimus tylenchiformis Daday, 1905 B. columbus by Siddiqi, 1986 syn. Criconema tylenchiformis (Daday) 13. H. concaudajuvencus Golden & Minton, Micoletzky, 1917 1970 lapsus calami H. paradoxus Daday, 14. H. dimorphicus Mulk &Jairajpuri, 1976 1905 B. dimorphicus (Mulk & Jairajpuri, 1976) Shamsi, 1979 Other species (with synonyms) H. dimorphicus by Luc, 1981 2. Hoplolaimus abelmoschi Tandon & B. dimorphicus by Siddiqi, 1986 Singh, 1973 15. H. dubius Chaturvedi, Singh & Khera, Basirolaimus abelmoschi (Tandon & 1979 Singh, 1973) Siddiqi,