Junker_115-128.indd INTRODUCTION The genus Tetrameres Creplin, 1846 are cosmopol- itan parasites, infecting a variety of aquatic and ter- restrial avian hosts. Females are usually located in the proventricular glands, and the males are found free in the lumen of the proventriculus (Ander son 1992). Several Tetrameres species have been recorded from the African continent, of which Tetrameres fis- sispina (Diesing, 1861) Travassos, 1914 that para- sitises ducks, pigeons and domestic chickens and Tetrameres americana Cram, 1927 that parasitises domestic chickens, turkeys and quails are the most commonly reported ones (Permin, Magwisha, Kassu- ku, Nansen, Bisgaard, Frandsen & Gibbons 1997; Poulsen, Permin, Hindsbo, Yelifari, Nansen & Bloch 2000). Tetrameres coccinea (Seurat, 1914) Travassos, 1914 from the Greater flamingo, Phoenicopterus ru- ber, Linnaeus, 1758, Cattle egret, Bubulcus ibis (Lin- naeus, 1758) and Eurasian spoonbill, Platalea leuco- rodia Linnaeus, 1758, as well as Tetrameres lhuillieri (Seurat, 1918) from the Rock partridge, Alectoris graeca (Meisner, 1804) and the Stock pigeon, Col- umba oenas Linnaeus, 1758 were recorded from Al- geria (Yamaguti 1961). Tetrameres nouveli (Seurat, 1914) Travassos, 1914 was present in the Black- winged stilt, Himantopus himantopus (Linnaeus, 1758) in Algeria (Yamaguti 1961), and in Nigeria Tetra meres plectropteri Thwaite 1926 was found in the Spur-winged goose, Plectropterus gambensis (Lin naeus, 1766) (Yamaguti 1961). 115 Onderstepoort Journal of Veterinary Research, 74:115–128 (2007) Tetrameres numida n. sp. (Nematoda: Tetrameridae) from Helmeted guineafowls, Numida meleagris (Linnaeus, 1758), in South Africa K. JUNKER and J. BOOMKER* Department of Veterinary Tropical Diseases, University of Pretoria Private Bag X04, Onderstepoort, 0110 South Africa ABSTRACT JUNKER, K. & BOOMKER, J. 2007. Tetrameres numida n. sp. (Nematoda: Tetrameridae) from Hel - meted guineafowls, Numida meleagris (Linnaeus, 1758), in South Africa. Onderstepoort Journal of Veterinary Research, 74:115–128 Tetrameres numida n. sp. from the proventriculus of Helmeted guineafowls, Numida meleagris, in South Africa is described from eight male and four female specimens. The new species shares some characteristics with other Tetrameres species, but can be differentiated by a unique combination of characters. It bears two rows of cuticular spines extending over the whole length of the body and pos- sesses two spicules. The left spicule measures 1 699–2 304 μm and the right one 106–170 μm. Caudal spines are arranged in three ventral and three lateral pairs and the tail is 257–297 μm long. Diagnostic criteria of some of the previously described species of the genus Tetrameres from Africa and other parts of the world have been compiled from the literature and are included here. Keywords: Helmeted guineafowls, nematodes, Tetrameres numida * Author to whom correspondence is to be directed. E-mail: joop.boomker@up.ac.za Accepted for publication date 4 April 2007—Editor 116 Tetrameres numida n. sp. (Nematoda: Tetrameridae) from Hel meted guineafowls in South Africa Both Tetrameres paradisea Ortlepp, 1932 and Tetra- meres prozeskyi (Ortlepp, 1964) were described from South African hosts. Tetrameres paradisea was recovered from a Stanley’s crane, Anthropoides paradisea (Lichtenstein, 1793) (Ortlepp 1932), and T. prozeskyi occurred in Red-billed hornbills, Tockus erythrorhynchus (Temminck, 1823) and Southern Yellow-billed hornbills, Tockus leucomelas (Lichten- stein, 1842) (= Tockus flavirostris leucomelas), re- spectively (Ortlepp 1964). Previous records of Tetrameres spp. from guinea- fowls pertain mostly to studies in North and West Africa, Tetrameres fissispina being recorded from Helmeted guineafowls in these countries (Fabiyi 1972; Vercruysse, Harris, Bray, Nagalo, Pangui & Gibson 1985). Appleton (1983) found Tetrameres sp. females in Crested guineafowls, Guttera edou- ardi (Hartlaub, 1867) (= Guttera pucherani), in Natal (now KwaZulu-Natal Province), South Africa, but because males were not present, the species could not be determined. We here describe a new species of the genus Tetra- meres from Helmeted guineafowls in South Africa for which we propose the name Tetrameres numida n. sp. With regards to the classification of the genus Tetra- meres we have followed that of Chabaud (1975), placing the genus into the subfamily Tetramerinae Railliet, 1915 within the family Tetrameridae Travas- sos, 1914, which is one of four families comprising the superfamily Habronematoidea. At the time the genus had been divided into the subgenera Tetra- meres s. str., Gynaecophila Gubanov, 1950, Pe- trow i meres Chertkova, 1953 and Gubernacules Ra- sheed, 1960. Chabaud (1975), arguing that this division could lead to errors and bore little phyloge- netic significance, chose not to retain these, but di- vided the genus Tetrameres into the two subgenera Tetrameres (Tetrameres) Creplin, 1846 and Tetra- meres (Micro tetrameres) Travassos, 1915. In the light of new findings, especially concerning the mor- phology of adults and larval stages of these two subspecies, Anderson (1992), while retaining their position within the subfamiliy, recognized Tetrameres Creplin, 1846 and Microtetrameres Travassos, 1915 as two distinct genera, a generic classification that had been suggested by Skrjabin (1969). We adopt his view in the present paper. MATERIAL AND METHODS Fifteen Helmeted guineafowls, Numida meleagris (Linnaeus, 1758), were collected on a farm 60 km to the west of Musina (Messina), Limpopo Province, South Africa (22°22.139’ S, 29°30.399’ E) between July 2005 and November 2006. Ten of these were mature guineafowls and five were young birds, about 6–10 months old (Siegfried 1966). Eight male Tetrameres sp. were recovered from the proventriculus, where they occurred free in the lu- men and four females were dissected from the prov- entricular glands. Two guineafowls harboured a sin- gle male each, two hosts harboured two and three males respectively, and from a single host one male and four females were recovered. All hosts were mature guineafowls. The worms were fixed in 70 % ethanol and cleared in lactophenol for identification. All measurements, unless otherwise indicated, are in micrometres. DESCRIPTION Tetrameres numida n. sp. (Fig. 1–3; Tables 1, 2) With characters of the genus. Sexual dimorphism marked. MALE: Body elongated, tapering towards both ends, posteriorly to a tail with a short, pointed tip. Cuticle with fine transverse striation and longitudinal cuticu- lar grooves. Total length 4.3–4.5 mm; maximum width 0.16–0.17 mm. Inconspicuous lateral alae ex- tending down the length of the body; parallell to these run two lateral rows of cuticular spines (Fig. 2F). One row of spines is situated dorsally, the sec- ond row ventrally to the lateral alae (Fig. 1B). A pair of deirids with apical spines is situated at approxi- mately the height of the second pair of cuticular spines at a distance of 139–204 from the apex (Fig. 1B). Cuticular spines start at 93–154 from the apex, numbering approximately 42–47 per row. The nerve ring and excretory pore are 215–284 and 236–331 from the anterior extremity, respectively. The excre- tory pore is slightly posterior to the nerve ring. The triangular mouth is bounded by a pair of trilobed pseudolabia. The inner surface of each lobe carries two to four tooth-like processes. The precise number is difficult to assess in our specimens (Fig. 1A, 2A). Depth of buccal capsule 16–28, inner diameter 8– 11. Oesophagus divided into muscular and glandu- lar portion, 232–401 and 734–984, respectively. Total length of oesophagus 1 023–1 318. Spicules unequal and dissimilar. Right spicule tubular, with slight bend and spatulate tip, 106–170 long (Fig. 1C, 2D). Left spicule long and thin, trough-shaped, with spatulate tip. Shaft slightly twisted at 100–120 from proximal end. Total length 1 699–2 304 (Fig. 1D–F, 2C, 2E). A gubernaculum is absent. Tail 117 K. JUNKER & J. BOOMKER FIG. 1 Tetrameres numida n. sp. Male. A. Apical view of the trilobed pseudolabia surrounding the triangular mouth. Note the tooth- like processes (scale bar = 10 μm). B. Ventro-lateral view of the anterior end (scale bar = 100 μm). C. Ventral aspect of the posterior end (scale bar = 100 μm). D. Lateral view of the proximal end of the left spicule showing the slight twist (scale bar = 100 μm). E. Ventral view of the proximal end of the left spicule (scale bar = 100 μm). F. Distal end of the left spicule, ventral view (scale bar = 100 μm). Female. G. Complete female (scale bar = 1 mm). H. Anterior extremity (scale bar = 100 μm) G H F E D C BA 118 Tetrameres numida n. sp. (Nematoda: Tetrameridae) from Hel meted guineafowls in South Africa length 257–297. Six pairs of caudal spines, three pairs each in two ventral and two lateral rows, re- spectively. One or two additional ventral spines may be present (Fig. 1C). FEMALE: Specimens in situ red. A minute head and tail of regular nematode shape, but often twisted or bent, emerge at opposite sides from the central part of the body which is distinctly globular and slightly bent along the axis (Fig. 1G–H, 3A, 3C). The cuticle bears marked transverse striation and four longitu- dinal cuticular grooves. The latter divide the body into four segments of which the two segments fol- lowing the outer curve are slightly longer (Fig. 1G). Much of the internal detail is obscured by the egg- FIG. 2 Tetrameres numida n. sp. Male. A. Head, apical view. B. Anterior extremity, ventral view. C. Left spicule, anterior end. D. Posterior extremity with right spicule and distal tip of left spicule. E. Tip of left spicule. F. Body spines (see arrow) FE DC BA 119 K. JUNKER & J. BOOMKER filled uterus coils surrounding a large sacular intes- tine. Body length 4.2–5.3 mm, maximum width 2.6– 3.5 mm. The following measurements were derived from a single specimen: The deirids are at 179 and 190 and the nerve ring at 215 from the apex, re- spectively. The excretory pore could not be located. Depth of buccal capsule 23, inner diameter 7. Muscular part of oesophagus 333, the distal part of the glandular oesophagus obscured by the uterus. Eggs are elongate with near parallel sides, polar filaments were not seen (Fig. 3D). Eggs containing fully developed larvae are 56–59 long and 31–34 wide. Anus and vulva appeared to be confined in body folds. Emerging from the last body fold is a tail approximately 336 long with a simple tip. SPECIFIC DIAGNOSIS: Tetrameres numida is differen- tiated from other members of the genus, by the pos- session of two rows of somatic spines and the ar- rangement of its caudal spines in two ventral and two lateral rows with usually three pairs of spines each, although deviation might occur. A short right and a long left spicule are present, ranging from 106–131 and from 1 699–2 304 in length, respec- tively. HOST: Numida meleagris (Linnaeus, 1758), Hel meted guineafowl. SITE: Males occur free in the lumen of the proven- triculus, females are situated in the proventricular glands. LOCALITY: Musina (Messina), Limpopo Province, South Africa (22°22.139’ S, 29°30.399’ E). ETYMOLOGY: The specific epithet numida refers to the host. Types deposited in the National Collection of Animal Helminths at the Onderstepoort Veterinary Institute, Pretoria, South Africa. Holotype male: T.2191, Alo- type female: T.2192, Paratype males: T.2193– T.2195. FIG. 3 Tetrameres numida n. sp. Female. A. Three whole specimens, approximately 4 mm in length. Note the globular shape. B. Anterior extremity. C. Posterior end. Note the digested blood showing as dark smudge. D. Egg containing fully developed larva DC BA 120 Tetrameres numida n. sp. (Nematoda: Tetrameridae) from Hel meted guineafowls in South Africa T A B L E 1 T h e m o rp h o lo g ic a l ch a ra ct e ri st ic s o f T e tr a m e re s n u m id a s p . n . m a le s fr o m H e lm e te d g u in e a fo w ls , co m p a re d t o T e tr a m e re s p a ra d is e a O rt le p p , 1 9 3 2 a n d t o T e tr a m e re s p ro - ze sk yi ( O rt le p p , 1 9 6 4 ), a ll d e sc ri b e d f ro m S o u th A fr ic a n h o st s. A ll m e a su re m e n ts in m ic ro m e tr e s u n le ss o th e rw is e in d ic a te d M o rp h o lo g ic a l c ri te ri a G F M 1 /N 4 T .2 1 9 1 T .2 1 9 3 T .2 1 9 4 T .2 1 9 5 G F M 1 1 /1 G F M 1 2 /1 T e tr a m e re s p a ra d is e a T e tr a m e re s p ro ze s k y i S o u rc e T h is p a p e r T h is p a p e r T h is p a p e r T h is p a p e r T h is p a p e r T h is p a p e r T h is p a p e r O rt le p p ( 1 9 3 2 ) O rt le p p ( 1 9 6 4 ) B o d y le n g th ( m m ) 4 .3 4 .4 4 .4 4 .3 4 .3 n 4 .5 5 .8 1 .3 – 2 .4 B o d y w id th m a xi m u m n n 1 6 0 1 6 0 1 6 4 1 7 0 1 6 2 1 4 0 6 0 -7 0 D is ta n ce a p e x to f ir st s o m a tic s p in e n 1 2 6 & 1 1 7 9 6 & 1 0 0 1 0 2 & 9 3 1 0 5 & 9 4 1 3 1 & 1 5 4 9 6 & 1 1 3 n n D is ta n ce a p e x to d e ir id s n 1 7 4 & 1 8 0 1 3 9 & 1 4 9 1 7 9 & 1 7 2 1 6 5 & 1 7 7 1 7 4 & 1 8 1 1 7 5 & 2 0 4 8 5 ~ 5 0 – 6 0 D is ta n ce a p e x to n e rv e r in g n 2 5 6 2 1 5 2 3 4 2 4 4 2 8 4 2 6 4 n ~ 1 5 0 – 1 6 0 D is ta n ce a p e x to e xc re to ry p o re 2 6 8 3 0 7 2 3 6 2 8 7 2 9 6 3 3 1 3 1 6 n n D e p th o f b u cc a l c a p su le 2 2 2 5 2 8 2 3 2 1 2 2 1 6 2 5 5 .0 – 7 .0 W id th o f b u cc a l c a p su le ( in n e r) n 1 0 1 0 8 8 1 1 8 1 2 1 1 .0 – 1 3 .0 M u sc u la r o e so p h a g u s n 3 5 1 3 0 4 2 3 2 2 6 0 4 0 1 4 0 0 3 1 0 1 6 0 – 2 1 0 G la n d u la r o e so p h a g u s n 7 3 4 7 6 9 9 8 4 7 8 1 8 1 2 9 1 8 9 0 0 3 0 0 – 4 0 0 O e so p h a g u s to ta l l e n g th n 1 0 8 5 1 0 7 3 1 2 1 6 1 0 2 3 1 2 1 3 1 3 1 8 1 2 1 0 n L e n g th o f ta il 2 8 4 2 9 7 2 8 7 2 5 7 2 9 6 n 2 9 0 1 1 5 1 4 0 – 1 6 0 L e n g th o f ri g h t sp ic u le 1 3 1 1 3 0 1 0 6 1 1 0 1 3 1 1 2 0 1 7 0 A b se n t U su a lly a b se n tb L e n g th o f le ft s p ic u le 1 9 8 8 2 1 0 3 2 3 0 4 2 1 6 9 1 6 9 9 n 2 2 0 4 6 9 0 ; 5 0 4 – 6 2 6 a 2 3 0 – 2 6 0 n D a ta n o t a va ila b le a R a n g e g iv e n b y M o llh a g e n ( 1 9 7 6 ) in C re m o n te e t a l. (2 0 0 1 ) b A r ig h t sp ic u le w a s p re se n t in t h re e o f m o re t h a n 3 0 m a le s 121 K. JUNKER & J. BOOMKER T A B L E 2 A c o m p a ri so n o f m o rp h o lo g ic a l c h a ra ct e ri st ic s o f so m e s p e ci e s o f th e g e n u s T e tr a m e re s C re p lin , 1 8 4 6 S p e c ie s B o d y le n g th o f m a le ( m m ) N u m b e r o f ro w s o f s o m a ti c s p in e s L e n g th o f ro w s o f s o m a ti c s p in e s N u m b e r o f s p ic u le s S p ic u le l e n g th ( m m ) A rr a n g e m e n t o f c a u d a l s p in e s o r p a p il la e P o la r fi la m e n ts o n e g g s S o u rc e T e tr a m e re s a m e ri ca n a C ra m , 1 9 2 7 5 – 5 .5 4 n 2 L e ft : 0 .2 9 – 0 .3 1 ; ri g h t: 0 .1 – 0 .1 3 5 v e n tr a l p a ir s, n o la te ra l p a ir s n S ch m id t (1 9 6 2 ); G ib b o n s e t a l. (1 9 9 6 ) T e tr a m e re s a ra lie n si s E fim o v & R ijo w a , 1 9 3 9 2 .5 5 4 W h o le b o d y le n g th 2 L o n g : 0 .9 1 3 ; s h o rt : 0 .2 2 2 v e n tr a l p a ir s a n d 2 su b la te ra l r o w s w ith 6 a n d 7 s p in e s, re sp e ci tv e ly . T w o la te ra l t a il p a p ill a e a ls o p re se n t n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s a u st ra lis J o h n st o n & M a w so n , 1 9 4 1 7 .8 – 9 .0 2 W h o le b o d y le n g th 2 L o n g : 5 .8 – 6 .3 ; sh o rt : 0 .8 5 t o 6 s m a ll sp in e s n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s b iz iu ra e J o h n st o n & M a w so n , 1 9 4 1 4 .2 – 4 .4 4 W h o le b o d y le n g th 2 L o n g : 0 .2 5 – 0 .2 6 ; sh o rt : 0 .0 7 n n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s ca lid ri s M a w so n , 1 9 6 8 2 .2 – 2 .5 4 /2 4 r o w s a n te ri o rl y, f ro m g la n d u la r o e so p h a g u s o n w a rd s o n ly 2 2 L e ft : 0 .7 5 – 1 .0 ; ri g h t: 0 .0 8 – 0 .0 9 5 v e n tr a l p a ir s, 2 la te ra l p a ir s O n ly m a le s kn o w n M a w so n ( 1 9 6 8 ) T e tr a m e re s ca rd in a lis Q u e n tin & B a rr e , 1 9 7 6 4 .2 – 4 .9 5 2 W h o le b o d y le n g th 2 L e ft : 0 .3 6 5 – 0 .4 0 0 ; ri g h t: 0 .0 6 5 – 0 .0 8 5 a 4 – 5 p a ir s o f p o st cl o a ca l s p in e s P re se n t Q u e n tin & B a rr e (1 9 7 6 ) T e tr a m e re s cl a d o rh yn ch i M a w so n , 1 9 6 8 2 .0 – 2 .9 4 W h o le b o d y le n g th 1 L e ft : 1 .0 – 1 .3 7 3 s u b ve n tr a l p a ir s, 3 su b la te ra l p a ir s P re se n t M a w so n ( 1 9 6 8 ); P e n ce e t a l. (1 9 7 5 ); C re m o n te e t a l. (2 0 0 1 ) T e tr a m e re s co lo ra d e n si s S ch m id t, 1 9 6 2 2 .0 5 4 W h o le b o d y le n g th 2 L e ft : 0 .7 7 7 ; ri g h t: 0 .0 6 7 4 v e n tr a l p a ir s, 3 la te ra l p a ir s P re se n t S ch m id t (1 9 6 2 ) T e tr a m e re s co n fu sa T ra va ss o s, 1 9 1 9 4 .0 – 5 .0 4 n 2 L o n g : 0 .2 9 1 ; sh o rt : 0 .0 6 8 3 v e n tr a l p a ir s, 3 la te ra l p a ir s S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s co rd o n ife re n s R a sh e e d , 1 9 6 0 n 4 n n L e ft s p ic u le : 0 .4 0 n n P e n ce e t a l. (1 9 7 5 ) T e tr a m e re s cr a m i S w a le s, 1 9 3 6 2 .9 – 4 4 n 2 L e ft : 0 .2 7 – 0 .3 5 ; ri g h t: 0 .1 3 6 – 0 .1 8 5 n n S ch m id t (1 9 6 2 ); G ib b o n s e t a l. (1 9 9 6 ) T e tr a m e re s cr a m i a si a tic a R yj ik o v, 1 9 6 3 3 .2 5 – 3 .6 4 W h o le b o d y le n g th 2 L o n g : 0 .2 3 8 – 0 .2 5 4 ; sh o rt : 0 .0 9 9 – 0 .1 0 6 5 v e n tr a l p a ir s, 3 la te ra l p a ir s n S kr ja b in & S o b o le v (1 9 6 3 ) 122 Tetrameres numida n. sp. (Nematoda: Tetrameridae) from Hel meted guineafowls in South Africa S p e c ie s B o d y le n g th o f m a le ( m m ) N u m b e r o f ro w s o f s o m a ti c s p in e s L e n g th o f ro w s o f s o m a ti c s p in e s N u m b e r o f s p ic u le s S p ic u le l e n g th ( m m ) A rr a n g e m e n t o f c a u d a l s p in e s o r p a p il la e P o la r fi la m e n ts o n e g g s S o u rc e T e tr a m e re s cy g n i R yj ik o v & K o zl o v, 1 9 6 0 n 4 n 2 L e ft : a b o u t o n e h a lf th e le n g th o f th a t o f T . tin a m ic o la 3 r o w s o f 5 c a u d a l p a p ill a e n P e n ce e t a l. (1 9 7 5 ) T e tr a m e re s d u b ia T ra va ss o s, 1 9 1 7 b 1 .3 5 – 2 .2 8 4 /2 D o rs o la te ra l r o w s re a ch o n ly t h e le ve l o f th e p o st e ri o r e n d o f th e g la n d u la r o e so p h a g u s 2 L o n g : 0 .7 1 – 0 .7 7 ; sh o rt : 0 .0 6 – 0 .0 8 4 v e n tr a l p a ir s, 3 la te ra l p a ir s P re se n t M a m a e v (1 9 5 9 ) ci te d b y S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s fe rm in i V ig u e ra s, 1 9 3 5 2 .5 n n 2 L o n g : 0 .0 7 3 ; sh o rt : 0 .0 2 3 3 p a ir s o f p o st cl o a ca l sp in e s n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s fis si sp in a (D ie si n g , 1 8 6 1 ) T ra va ss o s, 1 9 1 4 3 .0 – 6 .0 n n 2 L e ft : 0 .8 2 – 1 .5 ; ri g h t: 0 .2 8 – 0 .4 9 8 p a ir s o f p o st a n a l sp in e s n G ib b o n s e t a l. (1 9 9 6 ) 3 .2 – 3 .9 4 n 2 L o n g : 0 .3 7 – 0 .4 9 ; sh o rt : 0 .1 6 5 – 0 .1 9 8 3 v e n tr a l p a ir s, 5 la te ra l p a ir s n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s g a le ri cu la tu s O sc h m a ri n , 1 9 5 6 3 .4 4 W h o le b o d y le n g th 2 L o n g e r: 0 .4 5 0 ; sh o rt : 0 .0 8 6 P re se n t n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s g ig a s T ra va ss o s, 1 9 1 9 7 .5 4 W h o le b o d y le n g th 2 L o n g : 0 .7 4 ; sh o rt : 0 .0 1 6 T a il p a p ill a e h a ve n o t b e e n f o u n d n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s g lo b o sa ( V o n L in st o w , 1 8 7 9 ) 3 .6 – 3 .7 5 4 W h o le b o d y le n g th , sp a r s e r in p o st e ri o r h a lf 2 /1 L o n g : 0 .3 ; s h o rt sp ic u le r u d im e n ta ry S m a ll sp in e s p o st e ri o r to c lo a ca n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s g ru si S h u m a ko vi ts h , 1 9 4 6 3 .4 5 – 4 .4 0 2 2 d is tin ct r o w s, b u t sp in e s sc a tt e re d a n te ri o r to n e rv e r in g a n d p o st e ri o r to a n u s 1 0 .6 3 8 – 0 .7 8 3 S e ve ra l i rr e g u la r ro w s o f sp in e s n S kr ja b in & S o b o le v (1 9 6 3 ); B u sh e t a l. (1 9 7 3 ); P e n ce e t a l. (1 9 7 5 ) T e tr a m e re s g u b a n o vi S h ig in , 1 9 5 7 6 .6 7 2 W h o le b o d y le n g th , st a rt in g a t tr a n si tio n fr o m m u sc u la r to g la n d u la r o e so p h a g u s 2 L o n g : 3 .9 9 6 ; sh o rt : 0 .1 3 1 4 v e n tr a l p a ir s o f co n ic a l p a p ill a e , 3 la te ra l p a ir s o f st a lk e d p a p ill a e n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s h a g e n b e ck i T ra va ss o s & V o g e ls a n g , 1 9 3 0 3 .1 – 3 .4 2 ? R o w s o f cu tic u la r sp in e s a lo n g la te ra l fie ld s (2 r o w s ill u st ra te d ) L o n g s p ic u le : t h in a n d e n d in g a s a s p u r, p ro xi m a l 0 .0 7 – 0 .0 8 tw is te d . S h o rt s p ic u le 0 .0 3 2 – 0 .0 4 4 v e n tr a l p a ir s, 2 la te ra l p a ir s n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s lh u ill ie ri ( S e u ra t, 1 9 1 8 ) n 4 n 1 0 .4 8 n P re se n t O rt le p p ( 1 9 6 4 ) T A B L E 2 (c o n t. ) 123 K. JUNKER & J. BOOMKER S p e c ie s B o d y le n g th o f m a le ( m m ) N u m b e r o f ro w s o f s o m a ti c s p in e s L e n g th o f ro w s o f s o m a ti c s p in e s N u m b e r o f s p ic u le s S p ic u le l e n g th ( m m ) A rr a n g e m e n t o f c a u d a l s p in e s o r p a p il la e P o la r fi la m e n ts o n e g g s S o u rc e T e tr a m e re s lo b ib yc is M a w so n , 1 9 6 8 1 .5 4 /2 4 r o w s a n te ri o rl y, f ro m n e rv e r in g o n w a rd s o n ly 2 1 L e ft : 0 .7 3 6 s u b ve n tr a l p a ir s O n ly m a le kn o w n M a w so n ( 1 9 6 8 ) T e tr a m e re s m e g a p h a sm id ia ta C re m o n te , D ig ia n i, B a la & N a vo n e ( 2 0 0 1 ) 1 .9 4 – 2 .0 3 4 W h o le b o d y le n g th 1 L e ft : 0 .9 6 – 1 .2 2 6 s u b ve n tr a l p a ir s, 2 la te ra l p a ir s n C re m o n te e t a l. (2 0 0 1 ) T e tr a m e re s m ic ro p e n is T ra va ss o s, 1 9 1 5 4 .0 – 5 .0 2 W h o le b o d y le n g th 2 L o n g : 0 .3 5 5 ; sh o rt : 0 .0 5 6 2 v e n tr a l p a ir s n O rt le p p ( 1 9 3 2 ); S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s m ic ro sp in o sa V ig u e ra s, 1 9 3 5 3 .0 2 W h o le b o d y le n g th 2 L o n g : 1 .1 3 5 ; sh o rt : 0 .0 6 5 5 v e n tr a l p a ir s A b se n t S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s m o h te d a i B h a le ra o a n d R a o , 1 9 4 4 4 .2 7 – 5 .8 4 /2 S u b m e d ia n s p in e s e n d p o st e ri o r to m id d le o f g la n d u la r o e so p h a g u s 2 L o n g : 0 .3 9 7 – 0 .4 3 0 ; sh o rt : 0 .1 4 2 – 0 .1 6 0 5 s u b ve n tr a l p a ir s n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s n o u ve li (S e u ra t, 1 9 1 4 ) 1 .0 – 2 .4 4 W h o le b o d y le n g th 1 L e ft : 3 5 0 – 5 8 0 c 3 o r 4 s u b ve n tr a l p a ir s, 2 o r 3 s u b la te ra l p a ir s P re se n t O rt le p p ( 1 9 3 2 ); M a w so n ( 1 9 6 8 ); C re m o n te e t a l. (2 0 0 1 ) 2 .1 6 4 W h o le b o d y le n g th 1 0 .4 8 0 ; se co n d s p ic u le ru d im e n ta ry ( S e u ra t 1 9 1 4 , ci te d b y S kr ja b in & S o b o le v 1 9 6 3 ) 4 v e n ra l a n d 3 l a te ra l p a ir s ill u st ra te d ; a cc o rd - in g t o t e xt 2 p a p ill a e i n p o st e ri o r th ir d o f ta il P re se n t S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s n u m e n ii M a m a e v, 1 9 5 9 1 .6 4 – 2 .4 4 /2 D o rs o la te ra l r o w s re a ch o n ly t h e le ve l o f th e p o st e ri o r p a rt o f th e o e so p h a g u s 2 L o n g : 1 .0 8 – 1 .2 4 ; sh o rt : 0 .0 8 – 0 .1 0 4 v e n tr a l p a ir s, 3 la te ra l p a ir s A b se n t S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s n u m id a n . sp . 4 .3 – 4 .4 2 W h o le b o d y le n g th 2 L e ft : 1 .6 9 9 – 2 .3 0 4 ; ri g h t: 0 .1 0 6 – 0 .1 3 1 3 v e n tr a l p a ir s, 3 la te ra l p a ir s A b se n t T h is p a p e r T e tr a m e re s o xy la b ia tu s O sc h m a ri n , 1 9 5 6 5 .0 n W h o le b o d y le n g th 2 L o n g : 0 .9 4 0 ; sh o rt : 0 .1 2 5 E xt e n d p o st e ri o rl y to m id d le o f ta il, g e tt in g ve ry s m a ll n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s p a ra a ra lie n si s O sc h m a ri n , 1 9 5 6 1 .7 1 4 W h o le b o d y le n g th 1 0 .4 0 5 – 0 .4 2 0 n n S kr ja b in & S o b o le v (1 9 6 3 ); M a w so n ( 1 9 6 8 ); M o llh a g e n ( 1 9 7 6 ) in C re m o n te e t a l. (2 0 0 1 ) T A B L E 2 (c o n t. ) 124 Tetrameres numida n. sp. (Nematoda: Tetrameridae) from Hel meted guineafowls in South Africa S p e c ie s B o d y le n g th o f m a le ( m m ) N u m b e r o f ro w s o f s o m a ti c s p in e s L e n g th o f ro w s o f s o m a ti c s p in e s N u m b e r o f s p ic u le s S p ic u le l e n g th ( m m ) A rr a n g e m e n t o f c a u d a l s p in e s o r p a p il la e P o la r fi la m e n ts o n e g g s S o u rc e T e tr a m e re s p a ra d is e a O rt le p p , 1 9 3 2 5 .8 2 W h o le b o d y le n g th 1 L e ft : 0 .6 9 d 3 v e n tr a l p a ir s, 3 d o rs o -e xt e rn a l p a ir s A b se n t O rt le p p ( 1 9 3 2 ) T e tr a m e re s p a ra d o xa (D ie si n g , 1 8 3 5 ) 1 2 – 1 5 2 n 2 L o n g : 3 .0 o r lo n g e r ; sh o rt : 0 .4 8 0 D ra sh e ( 1 8 8 4 ) ill u st ra te d a v e ry s m a ll p a ir o f ve n tr a l p a p ill a e a n d 3 a n d 4 la te ra l p a p ill a e r e sp e ct iv e ly n S kr ja b in & S o b o le v (1 9 6 3 ), D ra sh e ( 1 8 8 4 ) ci te d b y S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s p a tt e rs o n i C ra m , 1 9 3 3 4 .2 – 4 .6 2 W h o le b o d y le n g th 1 1 .2 – 1 .5 n n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s p a u ci sp in a S a n d g ro u n d , 1 9 2 8 n 2 F e w , o n ly in p o st e ri o r 2 /3 2 L e ft : 0 .3 2 8 – 0 .3 7 1 ; ri g h t: 0 .0 1 2 – 0 .1 5 4 e 3 c a u d a l p a p ill a e n B u sh e t a l. (1 9 7 3 ); Q u e n tin & B a rr e ( 1 9 7 6 ) 3 .1 – 4 .5 1 1 r o w in m e d ia n ve n tr a l f ie ld , n o t m o re th a n 2 5 s p in e s, o n ly in p o st 2 /3 2 L o n g : 0 .3 2 8 – 0 .3 7 1 ; sh o rt : 0 .1 5 4 3 c a u d a l p a p ill a e n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s p a vl o vs ki i I yg is , 1 9 6 5 n 4 n 1 n 4 v e n tr a l p a ir s, 4 la te ra l p a ir s n P e n ce e t a l. (1 9 7 5 ) T e tr a m e re s p a vo n is T sc h e rt ko va , 1 9 5 3 4 .7 n Ir re g u la r a n d d e n se a n te ri o rl y, in m id d le a n d p o st e ri o r p a rt a lm o st in vi si b le 2 L o n g : 0 .4 3 ; sh o rt : 0 .1 0 5 4 r o w s o f sp in e s, a n d 3 p a p ill a e : 1 la te ra l p a ir , 1 u n p a ir m e d ia n p a p ill a n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s p h a e n ic o p te ru s A li, 1 9 7 0 n 4 n 2 n n n P e n ce e t a l. (1 9 7 5 ) T e tr a m e re s p le ct ro p te ri T h w a ite , 1 9 2 6 n n n n L e ft : 0 .8 5 n n O rt le p p ( 1 9 6 4 ) T e tr a m e re s p ro ze sk yi (O rt le p p , 1 9 6 4 ) 1 .3 – 2 .4 4 W h o le b o d y le n g th 1 L e ft : 0 .2 3 – 0 .2 6 f 3 v e n tr a l p a ir s, 3 la te ra l p a ir sg n O rt le p p ( 1 9 6 4 ) T e tr a m e re s p u ch o vi G u sh a n sk a ja , 1 9 4 9 3 .8 6 – 4 .3 3 9 2 W h o le b o d y le n g th 1 0 .3 0 7 – 0 .3 0 9 ; se co n d sp ic u le r u d im e n ta ry : 0 .0 0 8 n n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e rs r yj ik o vi C h u a n , 1 9 6 1 4 .5 4 W h o le b o d y le n g th 2 L o n g : 0 .2 0 8 ; sh o rt : 0 .0 6 2 4 v e n tr a l p a ir s, 3 la te ra l p a ir s n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s sa kh a ro w i P e tr o w , 1 9 2 6 9 .4 7 4 n 2 L e ft : 0 .1 9 5 ; ri g h t: 1 .0 2 1 n n S kr ja b in & S o b o le v (1 9 6 3 ) T A B L E 2 (c o n t. ) 125 K. JUNKER & J. BOOMKER S p e c ie s B o d y le n g th o f m a le ( m m ) N u m b e r o f ro w s o f s o m a ti c s p in e s L e n g th o f ro w s o f s o m a ti c s p in e s N u m b e r o f s p ic u le s S p ic u le l e n g th ( m m ) A rr a n g e m e n t o f c a u d a l s p in e s o r p a p il la e P o la r fi la m e n ts o n e g g s S o u rc e T e tr a m e re s sc o lo p a ci d is M a w so n , 1 9 6 8 1 .0 6 – 1 .8 4 /2 4 r o w s a n te ri o rl y, f ro m e n d o f o e so p h a g u s o n ly 2 r o w s 2 L e ft :0 .7 0 – 0 .8 5 ; ri g h t: 0 .0 7 – 0 .1 0 5 4 s u b ve n tr a l p a ir s, 3 su b la te ra l p a ir s P re se n t M a w so n ( 1 9 6 8 ) T e tr a m e re s so m a te ri a e R yj ik o v, 1 9 6 3 4 .8 4 N o s p in e s in t h e m id d le p a rt o f th e b o d y 2 L o n g : 0 .5 7 6 ; sh o rt : 0 .0 8 6 5 v e n tr a l p a ir s, 4 la te ra l p a ir s n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s sp ir o sp ic u lu m P in to & V in ce n te , 1 9 9 5 2 .5 2 – 4 .0 6 n T h in ly d is p e rs e d a n d p o o rl y d e ve lo p e d 2 L e ft : 0 .8 2 – 1 .0 8 ; ri g h t: n n n P in to & V ic e n te (1 9 9 5 ) T e tr a m e re s sk rj a b in i P a n o w a , 1 9 2 6 2 .6 4 W h o le b o d y le n g th 2 L o n g : 1 .5 4 3 ; sh o rt : 0 .1 0 3 N o t fo u n d n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s te tr ic a T ra va ss o s, 1 9 1 7 2 .6 4 D is sa p e a r n e a r la st q u a rt e r o f b o d y le n g th 2 L o n g : 0 .2 ; sh o rt : 0 .0 2 2 4 la te ra l p a ir s, 4 su b la te ra l p a ir s n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s tim o p h e e w o i T ra va ss o s, 1 9 5 0 4 .7 n W h o le b o d y le n g th 2 L o n g : 0 .4 2 1 ; sh o rt : 0 .1 8 9 n n S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s tin a m ic o la P e n ce , M o llh a g e n & P re st w o o d , 1 9 7 5 6 .5 2 4 V e n tr a l r o w s w h o le b o d y le n g th , d o rs a l ro w s e n d 1 .0 2 m m fr o m a p e x 2 L e ft : 2 .2 6 ; ri g h t: 0 .2 0 7 5 s u b ve n tr a l p a ir s, 3 ve n tr o -l a te ra l p a ir s A b se n t P e n ce e t a l. (1 9 7 5 ) T e tr a m e re s u xo ri u s M a m a e v, 1 9 5 9 n 4 n 2 L e ft : 2 .1 – 2 .3 h ; ri g h t: 0 .0 8 8 4 v e n tr o la te ra l p a ir s, 2 su b d o rs a l p a ir s A b se n t M a m a e v (1 9 5 9 ); P e n ce e t a l.( 1 9 7 5 ) 4 .7 6 – 5 .0 4 /2 D o rs o la te ra l r o w s re a ch o n ly t h e b e g in n in g o f th e g la n d u la r o e so p h a g u s 2 L o n g : 2 .1 – 2 .2 4 ; sh o rt : 0 .0 8 6 – 0 .0 8 8 4 v e n tr o la te ra l p a ir s, 2 su b d o rs a l p a ir s A b se n t S kr ja b in & S o b o le v (1 9 6 3 ) T e tr a m e re s vi e tn a m e n si s F a n th e V ie t, 1 9 6 8 n 4 n 2 L e ft : 1 .2 8 ; ri g h t: 0 .1 4 8 5 v e n tr a l p a ir s (l a te ra l a b se n t) n F a n t h e V ie t (1 9 6 8 ) in H e l - m in th o lo g ic a l A b st ra ct s (1 9 7 0 ), P e n ce e t a l. (1 9 7 5 ) T A B L E 2 (c o n t. ) n N o in fo rm a tio n a t o u r d is p o sa l a T h e o ri g in a l r e a d s 6 5 -3 5 0 μ m . W e c o n si d e r th is a t yp in g e rr o r a n d in cl u d e t h e r a n g e o f si n g le m e a s- u re m e n ts p ro vi d e d b y Q u e n tin & B a rr e ( 1 9 7 6 ) b S kr ja b in & S o b o le v (1 9 6 3 ) a ls o i n cl u d e a d e sc ri p tio n a ft e r C ra m ( 1 9 2 7 ), w h ic h d iff e rs s lig h tly f ro m th a t o f M a m a e v (1 9 5 9 ) c C re m o n te e t a l. (2 0 0 1 ) g iv e a r a n g e o f 0 .3 1 2 – 0 .5 8 7 m m d C re m o n te e t a l. (2 0 0 1 ) q u o te M o llh a g e n ( 1 9 7 6 ) g iv in g a r a n g e o f 0 .5 0 4 – 0 .6 2 6 m m e T h e le n g th p ro vi d e d b y Q u e n tin & B a rr e ( 1 9 7 6 ) is 1 2 – 1 5 4 μ m . W e c o n si d e r th is a n e rr o r. S kr ja b in & S o b o le v g iv e t h e w id th o f th e r ig h t sp ic u le a s 1 2 μ m f A cc o rd in g t o O rt le p p ( 1 9 6 4 ) in t h re e o f a b o u t 3 0 m a le s a r ig h t sp ic u le w a s p re se n t g C re m o n te e t a l. (2 0 0 1 ) q u o te M o llh a g e n ( 1 9 7 6 ) a s T . p ro ze sk yi h a vi n g v a ry in g c a u d a l p a p ill a e ( 3 /0 , 3 /3 , 4 /1 , 4 /2 ) h C a lc u la te d f ro m a 1 :2 4 t o 1 :2 6 r a tio b e tw e e n r ig h t a n d le ft s p ic u le 126 Tetrameres numida n. sp. (Nematoda: Tetrameridae) from Hel meted guineafowls in South Africa DISCUSSION Some of the main morphological characteristics of many of the species belonging to the genus Tetra- meres are listed in Table 2. Of the Tetrameres species with two rows of cuticular spines, Tetrameres pattersoni Cram, 1933, T. para- disea and Tetrameres grusi Shumakovitsh, 1946 have only one spicule and the spicule measure- ments of the latter two species differ distinctly from those in our specimens (Ortlepp 1932; Schmidt 1962; Bush, Pence & Forrester 1973). Tetrameres gubanovi Shigin, 1957 bears two rows of body spines, but has seven pairs of caudal papil- lae (Pence et al. 1975), as opposed to six pairs of caudal spines in T. numida n. sp. The use of the term caudal spines or caudal papil- lae is not always clear. Pence et al. (1975) use the term caudal papillae for several species in their pub- lication. They list T. paradisea as well as T. prozes- kyi as having caudal papillae, but in the original de- scriptions Ortlepp (1932, 1964) clearly refers to cuticular spines. Thus, Pence et al. (1975) seem to use the term indiscriminately. Mawson (1968), how- ever, describes T. nouveli as having caudal spines, but points out that in Tetrameres lobibycis Mawson, 1968 the spines are more like elongate papillae, and refers to Tetrameres calidris Mawson, 1968 and Tetrameres scolopacidis Mawson, 1968 as having papillae. The left spicules of Tetrameres cardinalis Quentin & Barre, 1976 and Tetrameres paucispina Sandground, 1928 are much shorter than those measured in our specimens (Quentin & Barre 1976). Tetrameres mi- cropenis Travassos, 1915 has been recovered from ciconiiform hosts, Nyctanassa violacea (Linnaeus, 1758) and Cochlearius cochlearia (Linnaeus, 1766) (Yamaguti 1961), whose geographic distribution is restricted to North and South America (Lepage 2006). Tetrameres fissispina has been recorded from guin- eafowls in Africa (Fabiyi 1972; Vercruysse et al. 1985) and, like T. americana, has a high prevalence in domestic chickens, whose nematode fauna is similar to that of guineafowls (Mukaratirwa, Hove, Es mann, Hoj, Permin & Nansen 2001; Magwisha, Kassuku, Kyvsgaard & Permin 2002). Tetrameres fissispina distinguishes itself from the new species by its shorter spicules and the larger number of cau- dal spines. Tetrameres americana differs not only in the spicule size and the number and arrangement of caudal spines, but also in its four rows of somatic spines (Schmidt 1962; Gibbons, Jones & Khalil 1996). The head of the female and the apical view of the head of the male of T. numida n. sp. most closely resemble Tetrameres tinamicola Pence, Mollhagen & Prestwood, 1975. The authors of the latter spe- cies describe the male head as possessing a triangu- lar mouth surrounded by a pair of trilobed structures originating from the inner surface of the pseudola- bia. Each lobe bears a pair of tooth-like processes in T. tinamicola. Similar processes can be seen in our specimens, but it is difficult to determine their exact number. However, there seem to be three or four per lobe. Pronounced lateral alae, as illustrated by Pence et al. (1975), were not found in our speci- mens. Moreover, T. tinamicola has a total of four rows of cuticular spines and the deirids are without apical spines. While the length of the left spicule of both species is similar, the right spicule of T. numida is only approximately half the length of T. tinamicola. Ortlepp (1932) described the buccal capsule of T. paradisea as having trilobed structures showing two to three bright refringent markings towards its pos- terior border. This, as well as other features of our specimens such as the transverse grooves anterior to the cloaca and the size of the spines, appeared so similar to T. paradisea that we initially considered assigning them to T. paradisea, especially in view of the fact that both were recovered from South African hosts. Close examination has nevertheless revealed distinct differences between the two. Tetrameres paradisea possesses a single spicule, whereas in our males two spicules are consistently present. While the arrangement of caudal spines is nearly identical and both carry three pairs of ventral and three pairs of externo-dorsal or lateral spines, the tail of T. paradisea is considerably shorter than that of our specimens (see Table 1). Ortlepp (1932) described and illustrated two rows of body spines found in T. paradisea and he uses this criterion to distinguish his species from Tetrameres nouveli which he lists as possessing four rows of spines. Cremonte, Digiani, Bala & Navone (2001) record T. paradisea as having four rows of spines, but cite Mollhagen (1976) as describing the dorsal rows of spines as very short, ending at 94–155 from the anterior end. When comparing T. paradisea to T. prozeskyi, Ort- lepp (1964) lists the length of the left spicule of the former species as 0.48 mm, but his original descrip- tion of T. paradisea (Ortlepp, 1932) clearly states the length of the spicule as 0.69 mm. We list T. pro- 127 K. JUNKER & J. BOOMKER zeskyi as monospicular, which differentiates it from our bispicular specimens. As regards T. prozeskyi it should be borne in mind that Ortlepp (1964) found a well-chitinized right spicule in three of the more than 30 males he examined. In the summary of the description of Tetrameres car- dinalis Quentin & Barre, 1976, the range of the length of the right spicule is given as 65–350 μm (Quentin & Barre 1976). As this seems erroneous, we decid- ed to include the range provided in the same paper, namely 365–400, in Table 2. Similarly, we consider the first measurement these authors provide for the short spicule of T. paucispina as incorrect and be- lieve it should read 120 instead of 12. Apart from T. numida n. sp., only T. tinamicola and Tetrameres uxorius Mamaev, 1959 have a left spi- cule that reaches 2 mm in length, while in the re- maining Tetrameres spp. the long spicule usually does not exceed 1 mm (Mamaev 1959; Pence et al. 1975). Relative to body length, however, there are other species with long spicules, such as T. lobiby- cis where the single spicule reaches about half of the body length (1.5 mm) and T. scolopacidis where the spicule length reaches almost two thirds of the body length (1.06–1.8 mm) (Mawson 1968). To our knowledge, Tetrameres phaenicopterus Ali, 1970 is the only member of the genus Tetrameres possessing a gubernaculum (Pence et al. 1975) and Tetrameres greeni Mawson, 1979 is unique in the genus Tetrameres in that it has caudal alae (Mawson 1979). Tetrameres spirospiculum Pinto & Vicente, 1995 is distinguished from our specimens and all the other species of Tetrameres by the spiral shaped distal end of the longer of its two spicules (Pinto & Vicente 1995). The numbers of T. numida n. sp. recovered from the guineafowl hosts from Musina (Messina) were low, and the parasite was only found in the older birds, being absent in young adults. While it is possible that guineafowls are not the main host for this para- site, we attribute the low intensity of infection to the fact that the area had been experiencing a severe drought during the past years. This would decrease the survival rates of nematode eggs while at the same time causing the numbers of possible inter- mediate hosts necessary for the completion of the life-cycle to decline. While differences in the im- mune status between guineafowls of different age might play a role in the intensity of infection, we be- lieve that the presence of T. numida n. sp. in older hosts simply reflects the increased possibility of prior exposure to the parasite as a function of time. ACKNOWLEDGEMENTS The authors are indebted to Dr S. Sokolov of the Institute of Parasitology, Russian Academy of Sci- ence, Moscow, for obtaining the extensive chapter on the genus Tetrameres in “Principles of nema- todology XI” (Skrjabin & Sobolev 1963) and to Dr D.A. Apanaskevich, Georgia Southern University, for the translation from the original Russian into English. The authors thank Mr K. Meyer and Mr M. Storm, the previous and current owners of the farm Sandown, Musina (Messina), respectively, for plac- ing the guineafowls at our disposal and Mr H.E. Hattingh, University of Limpopo, for collecting them. Dr W.J. Luus-Powell, University of Limpopo, has kindly facilitated the co-operation. Ms D.T. Durand, University of Pretoria, photographed the three com- plete females of T. numida. This research was made possible through a Claude Leon Foundation Post- doctoral Fellowship grant to the first author. REFERENCES ANDERSON, R.C. 1992. Nematode parasites of vertebrates, their development and transmission, 1st ed. Wallingford and New York: CAB International. APPLETON, C.C. 1983. Tetrameriasis in Crested guineafowl from Natal. Ostrich, 54:238–240. BERGAN, J.F., RADOMSKI, A.A., PENCE, D.B. & RHODES, O.E. Jr. 1994. Tetrameres (Petrowimeres) striata in ducks. Journal of Wildlife Diseases, 30:351–358. BUSH, A.O., PENCE, D.B. & FORRESTER, D.J. 1973. Tetra- meres (Gynaecophila) williamsi sp. n. (Nematoda: Tetra- meridae) from the White ibis, Eudocimus albus, with notes on Tetrameres (Tetrameres) grusi Shumakovich from Sand- hill crane, Grus canadensis. Journal of Parasitology, 59:788– 792. CREMONTE, F., DIGIANI, M.C., BALA, L.O. & NAVONE, G.T. 2001. Tetrameres (Tetrameres) megaphasmidiata n. sp. (Nematoda: Tetrameridae), a parasite of the Two-banded plover, Charadrius falklandicus, and White-rumped sandpi- per, Calidris fuscicollis, from Patagonia, Argentina. Journal of Parasitology, 87:148–151. FABIYI, J.P. 1972. Studies on parasites of the grey-breasted hel- met guineafowl (Numida meleagris galeata Pallas) of the Vom area of the Benue Plateau State, Nigeria. I. Helminth parasites. Bulletin of Epizootic Diseases of Africa, 20:235– 238. FAN THE VIET 1968. [Two new species of spirurates (Nematoda, Spirurata) from birds of Vietnam], in Helminthological Abstracts 1970, 39:187. GIBBONS, L.M., JONES, A. & KHALIL, L.F. 1996. Manual of the 8th international training course on identification of helminths of economic importance. St. Albans: International Institute of Parasitology. HELMINTHOLOGICAL ABSTRACTS 1970. Series A. Animal and human helminthology. Commonwealth Agricultural Bureaux. LEPAGE, D. 2006. Avibase hosted by Bird Studies Canada, Bird- Life International. www.bsc-eoc.org/avibase. 128 Tetrameres numida n. sp. (Nematoda: Tetrameridae) from Hel meted guineafowls in South Africa MAGWISHA, H.B., KASSUKU, A.A., KYVSGAARD, N.C. & PER- MIN, A. 2002. A comparison of the prevalence and burdens of helminth infections in growers and adult free-range chick- ens. Tropical Animal Health and Production, 34:205–214. MAMAEV, Y.L. 1959. New helminths from birds of eastern Siberia. Trudi Gelmintologicheskoi Laboratorii. Akademiya Nauk SSSR, 9:175–187. MAWSON, P.M. 1968. Nematodes from Australian waders. Para- sitology, 58:277–305. MAWSON, P.M. 1979. Some Tetrameridae (Nematoda: Spiru ri- da) from Australian birds. Transactions of the Royal Society of South Australia, 103:177–184. MUKARATIRWA, S., HOVE, T., ESMANN, J.B., HOJ, C.J., PERMIN, A. & NANSEN, P. 2001. A survey of parasitic nem- atode infections of chickens in rural Zimbabwe. Onderstepoort Journal of Veterinary Research, 68:183–186. ORTLEPP, R.J. 1932. A new species of Tetrameres (Tetrameres paradisea sp. nov.) from Stanley Cranes. 18th Report of the Director of Veterinary Services and Animal Industry, Union of South Africa, August, 1932. ORTLEPP, R.J. 1964. Some helminths recorded from Red- and Yellow-Billed hornbills from the Kruger National Park. Onder- stepoort Journal of Veterinary Reseach, 31:39–52. PENCE, D.B., MOLLHAGEN, T. & PRESTWOOD, A.K. 1975. Tetrameres (Tetrameres) tinamicola sp. n. from the Crested tinamou, Eudromia elegans, with comments on the subge- nus Petrowimeres (Nematoda: Tetrameridae). Journal of Parasitology, 61:825–829. PERMIN, A., MAGWISHA, H., KASSUKU, A.A., NANSEN, P., BISGAARD, M., FRANDSEN, F. & GIBBONS, L. 1997. A cross-sectional study of helminths in rural scavenging poultry in Tanzania in relation to season and climate. Journal of Helminthology, 71:233–240. PINTO, R.M. & VICENTE, J.J. 1995. Tetrameres (Tetrameres) spirospiculum n. sp. (Nematoda, Tetrameridae) from the buff-necked ibis, Theristicus caudatus caudatus (Boddaert) (Aves, Threskiornithidae). Memórias do Instituto Oswaldo Cruz, 5:615–617. POULSEN, J., PERMIN, A., HINDSBO, O., YELIFARI, L., NAN- SEN, P. & BLOCH, P. 2000. Prevalence and distribution of gastro-intestinal helminths and haemoparasites in young scavenging chickens in upper eastern region of Ghana, Africa. Preventative Veterinary Medicine, 45:237–245. QUENTIN, J.C. & BARRE, N. 1976. Description et cycle bio- logique de Tetrameres (Tetrameres) cardinalis n. sp. Annales de Parasitologie humaine et comparée (Paris), 51:65–81. SCHMIDT, G.D. 1962. Tetrameres coloradensis n. sp., a nema- tode parasite of the common snipe Capella gallinago delica- ta. Journal of Parasitology, 48:850–851. SCHMIDT, G.D. 1977. Observations on the type specimens of two species described by Lauro Travassos. Journal of Para- sitology, 63:343. SIEGFRIED, W.R. 1966. Growth, plumage development and moult in the Crowned Guineafowl Numida meleagris coro- nata Gurney. Department of Nature Conservation Investiga- tional Report No. 8. SKRJABIN, K.I. (Ed.). 1969. Key to parasitic nematodes. Vol. 1, Spirurata and Filariata. Translated and edited by M. Raveh, 1991. Leiden: E.J. Brill Publishing Company. SKRJABIN, K.I. & SOBOLEV, A.A. 1963. Principles of nematod- ology XI. Spirurata of animals and man and the diseases caused by them Part I (Spiruroidea). Moscow: Izdatelstv Aka- demii Nauk SSSR (Russian). VERCRUYSSE, J., HARRIS, E.A., BRAY, R.A., NAGALO, M., PANGUI, M. & GIBSON, D.I. 1985. A survey of gastrointesti- nal helminths of the common helmet guinea fowl (Numida meleagris galeata) in Burkina Faso. Avian Diseases, 29:742– 745. YAMAGUTI, S. 1961. Systema Helminthum. The nematodes of vertebrates. Vol. III, Parts I & II. New York: Interscience Pub- lishers.