Rhabditida: Diplogasteridae) from Termites
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J. Helminthol. Soc. Wash. 57(1), 1990, pp. 26-30 Redescription of Chroniodiplogaster aerivora (Cobb) gen. n., comb. n. (Rhabditida: Diplogasteridae) from Termites GEORGE O. POINAR, JR. Department of Entomology, University of California, Berkeley, California 94720 ABSTRACT: The nematode originally described as Diplogaster aerivora by Cobb is redescribed in the new genus Chroniodiplogaster. Diagnostic characters of the new genus include (1) the presence of 9 genital papillae, 3 of which are closely associated in a triplet set at the base of the tail, (2) 2 separate bursae, 1 which extends ventrolaterally along the tail, and the second, which is associated with the triplet papillae, and (3) a dorsal metarhabdion with a large mobile tooth and subventral metarhabdions with variably sized teeth. Dauer stages of C. aerivora occurred in the heads of worker termites of Reticulitermes tibialis. On occasion, the nematodes would enter the body cavity and kill the termites. KEY WORDS: Chroniodiplogaster aerivora gen. n., comb, n., Diplogasteridae, Rhabditida, Nematoda, termite, parasite. In 1916, Cobb described Diplogaster labiata aerivora, the present author redescribes the and Diplogaster aerivora in a paper by Merrill species and places it in a new genus, Chroniodip- and Ford (1916) who described the associations logaster. of these 2 nematodes with insects. In this study, the latter species was reported by Merrill and Materials and Methods Ford (1916) to occur in the heads of the termite, Populations of M. aerivora studied here were isolated Leucotermes lucifugus, with insect death result- from the heads of worker termites (R. tibialis) (Banks) ing from high numbers of nematodes. Along with in Colorado in 1987 by John L. Capinera. They were maintained on nutrient agar plates and taxonomic his description, Cobb also reported that D. aeriv- studies were made on adults and dauer juveniles re- ora fed on grasshopper eggs. Banks and Snyder moved from these cultures. All nematodes were killed (1920) reported juveniles of D. aerivora in the in hot (55°C) water, fixed in TAP, and processed to heads of active, normal-appearing Reticulitermes glycerin for measurements and drawings. Four of Cobb's original slides (nos. 1-4) containing adult male and flavipes and adult nematodes in sick and dead female M. aerivora were received from A. Morgan insects. Golden, Nematology Laboratory, U.S. Dept. of Agri- Davis (1919) found Mesodiplogaster (=Diplo- culture, Beltsville, Maryland, for comparative purpos- gaster) aerivora in dead and dying larvae of the beetle, Phyllophaga sp., and suggested that the nematodes were the cause of death. Cobb iden- tified Mesodiplogaster aerivora as the nematode Results responsible for natural mortality of corn ear- Populations of nematodes removed from the worm larvae, Heliothis obsoleta (Winburn and heads of R. tibialis in Colorado agreed with the Painter, 1932). Lim et al. (1981) isolated M. ae- description of M. aerivora by Cobb (in Merrill rivora from both living and dead grubs of the and Ford, 1916) and were similar to Cobb's orig- June beetle, Phyllophaga anxia. Recently, the inal specimens of M. aerivora. Therefore the Col- present author received a culture of M. aerivora orado population was considered to be M. aeriv- that was isolated from Reticulitermes tibialis in ora Cobb. Colorado. This reexamination also revealed certain char- Recognition of M. aerivora is made difficult acters that made it difficult to place M. aerivora by the absence of detailed illustrations and quan- in any of the existing genera of Diplogasteridae. titative data in the original description. Neither These characters consist of 9 pairs of genital pa- Weingartner (1955) nor Goodey (1963) treated pillae including a posterior set of 3, of which 2 M. aerivora in their taxonomic presentation of are always reduced, a gubernaculum with ventral the family Diplogasteridae and the species M. processes, which encloses the distal portion of aerivora has been treated under the genera Diplo- the spicule shafts, a long narrow bursa associated gaster, Micoletzkya, and Pristionchus. with the 3 ventrolateral papillae, and a separate After a detailed morphological study of M. short bursa associated with the posterior set of 26 Copyright © 2011, The Helminthological Society of Washington 27 3 unequal papillae. The genus Diplogaster extending ventrolaterally along tail region and Schultze lacks a bursa, the gubernaculum does second smaller ventral 1 associated with the trip- not encircle the spicule shafts, and there are 8 let papillae at base of tail. pairs of genital papillae without a posterior set The genus Chroniodiplogaster can be separat- of 3 (Goodey, 1963). The genus Micolctzkya ed from previously described genera in the Dip- (Weingartner) contains 10 pairs of genital pa- logasteridae having a similarly shaped stomal pillae with the first 2 pairs adjacent (Goodey, opening with the dorsal metarhabdion contain- 1963). Members of the genus Mesodiplogaster ing a distinct movable tooth by the normal pres- Goodey (nee. Weingartner; see Loof [1976]) pos- ence of 2 separate bursae, a gubernaculum that sess 2 different stomal forms (Goodey, 1963). surrounds the terminal spicular tips, and the The genus Pristionchus Kreis, which was not presence of 9 basic genital papillae always in- mentioned by Goodey (1963), was originally de- cluding a postanal set of 3 (triplet). scribed as having a small postanal bursa sup- ported by 4 papillae (Kreis, 1933). However, Fe- Chroniodiplogaster aerivora (Cobb) 1916 dorko and Stanuszek (1971) show a long narrow comb. n. bursa running posteriorly from the anus to the The following description also incorporates tail spike in Pristionchus uniformis. They illus- features cited by Cobb (in Merrill and Ford, 1916) trate 10 genital papillae pairs but state that only in his original description. 9 pairs exist. DESCRIPTION: Adults: Cuticle transparent, The generic status of the Diplogasteridae is in with faint transverse striae and 24 longitudinal a state of confusion and the family needs to be striae; lateral lines present; anterior end truncat- revised. The present author prefers to redescribe ed; not offset, lips 6, fused at base, each bearing the species M. aerivora Cobb in a new genus in single inner labial papilla, submedial lips also the family Diplogasteridae, based on the discov- bearing 2 additional papillae, lateral lips with ery of new morphological characters not reported single additional papilla, amphids located at base or seriously considered previously. of lateral lips (on dorsal side), stoma walls highly In the quantitative portion of the description, cuticularized; dorsal metarhabdion with large all figures are given in micrometers unless oth- protruding movable tooth, the inner contour of erwise specified. The first number following the which fits into the contour of the protruding tooth character is the mean and numbers in parenthe- on the right subventral metarhabdion, which may ses indicate the range. be notched at apex; left subventral metarhabdion bearing a smaller, conoid-lacinate tooth, which Chroniodiplogaster gen. n. is closely associated with that of the right sub- (Diplogasteridae (Micoletzky) Steiner) ventral metarhabdion: pharynx extending from DESCRIPTION: Medium-sized nematodes with base of stoma, gradually widening, until reaching smooth cuticle; 6 lips united at base but distinct swollen metacorpus containing elongated val- at tip. Stoma massive, slightly longer than wide, vular apparatus, followed by isthmus surround- variable in shape but with only a single basic ed by nerve ring; pharyngeal base expanding into form, dorsal metarhabdion with large mobile pyriform bulb lacking distinct valve; excretory tooth, subventral metarhabdions variable; right pore opening in vicinity of basal bulb; small, subventral bearing tooth usually wide and point- 2-celled pharyngeal-intestinal valve present; in- ed or notched at tip; left subventral bearing testine forming narrow rectum before leading to smaller pyramidal or lacinate tooth; median bulb anus or cloacal vent. of pharynx distinct, with strong crescent-shaped FEMALE (Figs. 1, 2, 6, 8): Vulva located in valves; female didelphic; female and male tails midbody region, roughly spherical in shape in variable, short and acute or long and tapering or ventral view; gonads paired and opposite with long and offset leading to filamentous tip; spic- ovaries reflexed past vulvar opening; tail tapering ules large (length greater than body width at clo- gradually to fine hairline terminus; eggs ellipsoi- aca), slender, with small but distinct capitulum; dal with smooth and relatively thick shell; em- gubernaculum with proximal (terminal) portion bryonation occurs within female uterus. bent up and enclosing spicule tip region; with 9 MALE (Figs. 4, 5, 7): Similar to female in re- pairs of genital papillae including 3 pairs located spect to cuticle, stoma, pharynx and intestine; together in an off-ventral position near base of tail tapering abruptly posterior to cloaca; tail ta- tail constriction; 2 bursae normally present; 1 pering uniformly to small hairline terminus pos- Copyright © 2011, The Helminthological Society of Washington 28 JOURNAL OF THE HELMINTHOLOGICAL SOCIETY Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 57, NUMBER 1, JANUARY 1990 29 terior to triplet group of ventrolateral genital pa- 181); reflexion of testis, 155 (120-192); length pillae, faint constriction present just posterior to tail, 112 (88-138); width