Revision of the Genus Hemicriconemoides Chitwood & Birchfield, 1957 (Nematoda: Criconematidae)
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Revision of the Genus Hemicriconemoides Chitwood & Birchfield, 1957 (Nematoda: Criconematidae) D. R. DASGUPTA, 1 D. J. RASKI1 AND S. D. VAN GUNDY2 Abstract: In the present study, evaluations of the nominal species and descriptions (H. parvus, H. inter- medius, H. insignis, H. microdoratus and H. brevicaudatus) are made of five new species. H. squamosus is proposed as a species inquirenda; H. mangiferae is judged a valid species. H. litchi and H. birchfieldi are synonymized with H. mangi]erae and H. ureshinoensis with H. kanayaensis. H. strictathecatus is con- sidered a valid species. A key to the species of the genus is given. The limitation of use of male diag- nostic characters in a mixture of populations of Hemicriconemoides with some other criconematids is discussed. The life cycle of H. chitwoodi has four complete molts; one in the egg, three in soil, two of which have spines. The adult female has a sheath cuticle from an incomplete fifth molt. Chitwood and Birchfield (2) proposed and H. strictathecatus with squamosus. One the genus Hemicriconemoides to include of the generic characters of Hemicricone- those species "which fit neither the present moides and Hemicycliophora given by Sid- concepts of Criconemoides Taylor, 1936 nor diqi & Goodey (16) is: "spear base anchor- that of Hemicycliophora de Man, 1921." shaped" in Hemicriconemoides and "basal Three species ( Hemicriconemoides wessoni, knobs of spear spheroid" in Hemicyclio- H. bi]ormis and H. floridensis) were de- phora. On the basis of this difference in scribed by these authors and H. wessoni was knob shape, Hemicriconemoides biformis made genotype. Three species, identified and and H. floridensis were proposed to be re- described as members of the genus Cricone- moved from Hemicriconemoides and placed moides by Loos (12), were transferred to in Hemicycliophora. the new genus (2). Since then, eleven new In the present study, evaluations of the species have been described: H. chitwoodi, nominal species and descriptions of five new H. minutus and H. strictathecatus by Esser species are presented. A brief description of (9) ; H. kanayaensis by Nakasono & Ichinohe the developmental stages of H. chitwoodi is (13) ; H. mangiferae by Siddiqi (15) ; H. also included. obtusus by Colbran (4); H. communis and The authors wish to extend their thanks H. litchi by Edward & Misra (7); H. ure- shinoensis by Yokoo (20); H. pseudobrach- to Mr. R. P. Esser for providing some of yurum by De Grisse (5) and H. birchfieldi the paratypes and other collections of Hemi- by Edward et al. (8). criconemoides for this study. We also ac- Goodey ( 11 ) placed the genus Hemicrico- knowledge the help of Dr. S. A. Sher for nemoides in synonymy with Hemicyclio- making available the collections of Hemi- phora. Siddiqi & Goodey (16) resurrected criconemoides in the Department of Nema- the genus again. Iota squamosus Cobb, tology at the University of California, River- 1913 (3) was considered to be a member side. of Hemicriconemoides by Siddiqi & Goodey 16), and they synonymized H. rnangiJerae MATERIALS AND METHODS Nematodes in the nematode survey collec- Received for publication 8 October 1968. Department of Nematology, University of California, Davis, tions at Davis and Riverside were assembled California 95616, and 2 Department of Nematology, Uni- versity of California, Riverside, California 92502. Present for this generic study. These specimens in- address of senior author: Division of Nematology, Indian Agricultural Research Institute, New Delhi-12, India. clude representatives from six continents: 126 REVmION OF Hemicriconemoides • Dasgupta, Raski, Van Gundy 127 North America, South America, Asia, Aus- body, the cuticular sheath is closely ap- tralia, Africa and Europe. pressed to the inner cuticle only at the head, Some of the symbols used by De Grisse the vulva and sometimes at the terminus. (6) are added to the deManian formula. A lateral field is lacking. Timm (17) re- These are: R -- total number of sheath ported longitudinal lines on the cuticle of annules; R ex = annule on which excretory H. litchi. Examination of specimens from pore is located, beginning from anterior end; Nigeria identified as H. mangiferae, speci- R V = annule on which vulva is located, mens from Ceylon identified as H. gaddi and counting from terminus; R an = annule on specimens from the same locality and habitat which anus is located, counting from ter- from where H. litchi was reported by Timm minus; R Van = number of annules be- (17), revealed the presence of very faint tween vulva and anus; VL/VB = distance longitudinal lines which could only be seen from terminus to vulva divided by body with very careful observation. These lines width at vulva; V' = length of body from appeared to be subcuticular rather than cutic- vulva to terminus in microns. The measure- ular. The annules of the sheath and body ments given within parentheses in the de- are 2-8 ~ wide in the middle of the body, scription of each holotype refer to the pop- usually smoothly flattened in the sheath and ulation range. The annule counts were made rounded on the body (not retrorse) and on the ventral side of body. No attempt was their number varies from 51 to 164. made to describe males in those populations The adult males lack a cuticular sheath. which contained a mixture of Hemicricone- The lateral field of the males usually has moides, Criconemoides, Paratylenchus and four incisures, although only two lines have Criconema. been reported in H. gaddi. Of the four Slides of type specimens are deposited at incisures, the outer two lines are more UCNC (University of California Nematode prominent than the two fine inner lines. Collection, Davis, California) and USDANC Occasionally males are found within larval (United States Department of Agriculture cuticles bearing twelve rows of lobed, scale- Nematode Collection, Beltsville, Maryland). like or conoid spines. The cuticular spines in the larvae of H. pseudobrachyurum bear MORPHOLOGY dentations on their edges. Species in the genus Hemicriconemoides LIP REGION: The lip region of Hemi- vary in length from 0.29-0.63 mm. Fixed criconemoides is markedly conoid ( H. kana- specimens are straight or slightly curved yaensis), rounded (H. gaddi) or truncate ventrally. The adult females of Hemicrico- (H. chitwoodi). The oral aperture is obscure nemoides have double cuticles; according to and appears to be a simple narrow slit. Fassuliotis (10) in H. chitwoodi the outer There is wide variation in shape and extent or sheath cuticle represents an incomplete of development of the labial disc between fourth molt. However, the present study of species. In some species the labial disc is developmental stages of this species indicated conspicuous and elevated, surrounding the four complete molts and an incomplete fifth oral opening, while in other species it is in- molt. Therefore, the sheath cuticle of Hemi- conspicuous. In lateral view the labial disc criconemoides is an incomplete fifth-molt appears more or less circular in form, cuticle. Except for H. obtusus, in which the rounded or flattened at the top. In en [ace cuticular sheath is closely appressed to the view the labial disc appears to be hexagonal inner cuticle throughout the entire length of or quadrangular in shape. In H. cocophillus, 128 Journal o[ Nematology, Vol. 1, No. 2, April 1969 besides the labial disc, there is a well- REPRODUCTIVE SYSTEM: The adult fe- developed amphidial disc surrounding the males of Hemicriconemoides have a monodel- labial disc. The amphid apertures are prom- phic, prodelphic and outstretched reproduc- inent, appearing as narrow darkened slits tive system. Only two specimens were found on the lateral margin of the labial disc of with a single flexure in the gonad anterior to the amphidial plate. The sclerotization of the spermatheca. The vulva is at 88-96% and the lip region is hexaradiate, widening at a vulvar sheath may be present or absent. the base. The papillae are not identifiable, The vulvar sheath may be narrow and very but there is a suggestion of fine internal fine (e.g., H. microdoratus) or broad and innervations in the sublateral and lateral formed of two to three annules (e.g., H. sectors. brachyurus). In some species a vulvar sheath LABIAL ANNULES: Labial annules are is present both in the sheath cuticle and in similar to body annules but are not definitely the inner body cuticle (e.g., H. brachyurus), set off. The base of the sclerotized labial while another has the vulvar sheath only framework is used to distinguish the demar- in the sheath cuticle (H. microdoratus n. cation between the body and labial regions, sp.). In ventral view the sheath appears as and the labial annules are counted to that a projection with rounded corners. The point. The lip region of H. obtusus is an vulva anteriorly leads to a narrow vagina of exception since the labial annules are def- variable length (8-14 t~), slightly curved, initely set off. Basically there are two labial which opens into the uterus, oviduct and annules, which vary considerably in shape ovary. The anterior end of the uterus is and size. Within one population of H. coco- offset, usually ventrally, into a rounded or phillus, most of the individuals have two well- oblong spermatheca, frequently filled with developed labial annules, an occasional speci- rounded spermatozoa measuring 2-3 tz. Ed- men may have three, while others show ward and Misra (7) illustrated spermatozoa almost complete suppression of the first as elongated rod-like structures, but we have labial annule. Usually the first annule is never observed this type. smaller than the second (e.g., H. mangife- Males have a monorchic reproductive rae), but the reverse is true for some species system. The spicules are slender and slightly (e.g., H. chitwoodi). The first annule may curved ventrally. The gubernaculum is small be angular or rounded, and is irregularly and trough-like.