© Institute of Parasitology, Biology Centre CAS ;+'- E; +&F doi: '+ '99''H ;+'- +&F http://folia.paru.cas.cz
Research Article Studies on ascaridid, oxyurid and enoplid nematodes
František Moravec1 and Liesl L. Van As;
1 ! " 5 $ D5 6 7 = ; D ! K ? ! M " ! "
Abstract: ! N[ R ! 7 @ U> ? V ! [ U$ 6 V ! Falcaustra similis W ;++9 N ! R N7 ! R Cucullanus sp. (only female) (Seuratoidea), Cith- ariniella longicaudata sp. n., Synodontisia annulata N7 U> R Contracaecum @ @ of Galeiceps sp. (both Ascaridoidea) and Eustrongylides @ ND ! R X V Citharinella longicaudata (type host Schilbe intermedius 6 R ! 7 '+< 3! Synodontisia annulata (type host S. intermedius) by the shape of cephalic papillae, body length of gravid females N9 <
X ! [ " $ V R 7 @ ! $ V 6 W @[ V $ 7 [ 7 N;+'- M R 7 @ and maintained unfed in plastic tanks for at least two days to al@ ! N" R ! [ U$ V ! X [ V $ 6 V 7 ! 7 ";;; V ! '} ! 5 N{ { W R ;++'M;++& saline. Nematodes removed from the digestive tract and viscera This paper is a continuation of these studies, based on the V [> 9} '+} 7 ! same nematode collection, now providing results of the ! > ! ! V systematic evaluation of nematodes of the orders Ascarid@ cleared gradually in glycerine. Drawings were made with the aid U> ? D ! >@ of a Zeiss microscope drawing attachment. Specimens used for aminations (using both light and scanning electron micro@ ! N"? R V @[> '} @ scopy) of these materials revealed the presence of several ! ! > N 7 R @ V > @ @ @ V 7 @$ V $ X = V > ! ?U{ " @*9+' results are presented herein. electron microscope at an accelerating voltage of 4 kV (GB low mode). All measurements are in micrometres unless otherwise MATERIALS AND METHODS indicated. Only small numbers of the available specimens could 'EF ! 7 [ be used in the descriptions and there is no relationship to the the families Alestidae, Cichlidae, Clariidae, Mochokidae and !7 M ! ! " 7 V U$ 6 N! ! V 7 V [
! " $ &' &*+ +- 5 $ D5 6 7 89;+ &< *** -9&;= > 89;+ &< -&'+&<<= ?@! ! B Zoobank number for article: 7 $ 7 D?E&E'&*@F*E @9D9<@ E <@'*
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. '+ '99''H ;+'- +&F Moravec and Van As: Fish nematodes from Botswana
[ ! 7 consists of short, muscular pharyngeal portion, long cor@ ! ! N > D ! R \ ! ! @7 7 7 7 [ ! ring encircling oesophageal corpus at about 1/4 of way N;++FR X ! [ V N > 5 ;+'-R @7 7 > N ' R X 7 > RESULTS Male N- ! R { 7 < <-M'; &E !! ! > ! ! V E&FM*
Fig. 1. Falcaustra similis W ;++9 ! Schilbe intermedius 6 A M ! V= B, C M @ V = D M ! V= E M = F, G M ! V = H M ! V
;+'- E; +&F ; 'F '+ '99''H ;+'- +&F Moravec and Van As: Fish nematodes from Botswana
A
C D
Fig. 2. Falcaustra similis W ;++9 ! Schilbe intermedius 6 ! A M V= B M ! V= C, D M ! V Abbreviations: M ! = 7 M 7 = M 7!
N ' M ; MDR " N9HEER 9&-M-&+ N ' ; MDR ^ 7 @ D ! ! @ ! V '&;M'<+ X lection of the Institute of Parasitology, Biology Centre of the &E*M-&+ ! " 5 $ D5 N Female (5 gravid specimens): Length of body @<'FR '' +EM'9 +E !! ! > ! ! V *&9MF'' { ;*M&F Remarks. To date, ten species of Falcaustra { 'F'9 = V ';FM'9' { 7 7 ! [ F. guiersi Vas@ ' EEM' <* !!= > 'F*& F. hexapapillata N` 'FE;R F. petrei &EM<' ' ;;M' &F !! @7 7 '99 7 7 N` 'F*+R F. piscicola N{ V 'F+*R F. similis ;-;M;F'= V > @7 7 7 7 W ;++9 F. straeleni ! @6 EEMF+ FFM';+ ''*M';F ;;-M;E9 'FE' F. sudanensis N` 'FE;R F. tchadi Vassiliadès 9&-M9F+ > ' ;FM' E& !! X 'F*& F. therezieni 'F*F F. verbekei <&+M' '*+ ! > ! W ! @6 'FE' N ` 'FF* E FEM* *+ !! ! > ! N -;ME<} W ;++9R ! 7 R= ! V @ ! ! ! [ ! ? ! @V EFMF+ (although F. therezieni, F. guiersi and F. straeleni were & E+MEE N '?R= ! [ R V F. si- ! & 7 ! X 99;M--< milis is known only from a siluriform host, Synodontis ni- bearing dorsolateral pair of large phasmids (Fig. 1D). gromaculatus (Mochokidae). " $ Synodontis nigromaculatus Boulenger, The nematodes of the present material are morpho@ 7 'FF !! S. vanderwaali Skelton et White, body logically and biometrically very similar to F. similis, both ;++ !! N7 $ R [ Schilbe forms occur in siluriform hosts in the same locality, and, intermedius 6 N" 7 R 7 ;;< !! N therefore, they are considered to belong to this species. Siluriformes). Small differences in the arrangement of postanal papillae Site of infection: Intestine. ! V [ Locality: U$ 6 N" $ V ! ! '< ;E+-"= variabilty, which is usually broad in cosmocercoids. Al@ ;' -9;&?R V N 'F ;++'R though one host of these nematodes belongs to a different Prevalence: S. nigromaculatus ;E} N'< [ H*+ [ siluriform family than the type host of F. similis (Schilbei@ > ! R= S. vanderwaali -} N'H;+R= S. intermedius E} dae vs Mochokidae), it is not surprising, because also an@
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Fig. 3. AMD M ! ! Schilbe intermedius 6 NA M V= B, C M @ V = D M VR EMG M Eustrongylides ! @ ! Hydrocynus vittatus Castelnau (E M V= F M V= G M VR
$ NSynodontis spp.), Procamal- <9+ = -' lanus laeviconchus N] '
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A
C D
Fig. 4. Atractidae gen. sp. female from Schilbe intermedius 6 ! A, B M 7 V = C M 7 V N VR V= D M V Abbreviations M ! = 7 M = M
Remarks. Since only a single nongravid female was absent. Cephalic end somewhat asymmetrical in lateral 7 [ V ! 7 view. Oral aperture dorsoventrally elongate, surrounded The morphology and measurements of this specimen are by raised narrow membranous ala (collarette) supported very similar to the representatives of Labeonema Puylaert, 7 V ! 7 N - M? E M ? R 'F*+ N R V [ Four submedian cephalic papillae and pair of lateral am@ N 'F*+ ` 'FF* N -D E M R U ! W ;++9 ` 7$ ;++
;+'- E; +&F - 'F doi: 10.14411/fp.2015.039 Moravec and Van As: Fish nematodes from Botswana
Fig. 5. Cucullanus sp. female from Tilapia sparmanni Smith. A, B – anterior end of body, lateral and dorsoventral views, respec- tively; C – oesophastome, lateral view; D – cephalic end, apical view; E – oesophastome, dorsoventral view; F – deirid, lateral view; G – vulva, lateral view; H – tail, lateral view; I – sculpture of egg surface; J – egg.
Host: Banded tilapia, Tilapia sparmanni Smith (Cichlidae, Per- should be principally compared with the species of Cucul- ciformes), body length 150 mm. lanus V [ Site of infection: Intestine. To date, three species of Cucullanus have been described Locality: Okavango River (Samochima lagoon 18°25'48.9''S; ! V [ Cucullanus barbi Baylis, 21°53'41.8''E), Botswana (collected 11 December 2003). 1923 parasitising Barbus spp. (Cyprinidae) in the Sudan ' [ H9 [ > ! and Egypt (Baylis 1923a, Moravec 1974, Amin 1978, El- Deposition of voucher specimen: Helminthologi- Naffar et al. 1983); C. baylisi Campana-Rouget, 1961 from cal Collection of the Institute of Parasitology, Biology Centre Synodontis spp. (Mochokidae) in the Sudan, the Demo- ! " 5 $ D5 N cratic Republic of the Congo and Chad (Baylis 1923a, No. N-1083). Campana-Rouget 1961, Khalil 1969, Vassiliadès and Remarks. The genus Cucullanus Müller, 1777 includes X 'F*9 5 $ ;+''= a large number of species parasitising freshwater, brackish- specimens from S. schall ! [ C. clarotis Bay- V ! [ lis, 1923 by Baylis 1923a and Khalil 1969); and C. clarotis Because of their rather uniform morphology and the inad- from Clarotes laticeps (Rüppel) (Claroteidae) in the Sudan ! ! - (Baylis 1923a). Al-Bassel (2003) also reported Cucullanus cally impossible to make a detailed comparison between barbi Paperna, 1964 (a homonym to and probably identi- ! ! V cal with C. barbi Baylis, 1923 – see Moravec 1974) from these parasites according to their zoogeographical region Mormyrus kannume Forsskål (Mormyridae) in Egypt, but or their host groups (see e.g. Yooyen et al. 2011 for respec- [ 7 R X > In addition, two congeneric species, C. djilorensis Ndew, [ ! ? 6 Diouf, Bâ et Morand, 2014 and C. egyptae Abdel-Ghaffar, Africa, the present nematode specimen from T. sparmanni Bashtar, Abdel-Gaber, Morsy, Mehlhorn, Al Quraishy et
Folia Parasitologica 2015, 62: 039 Page 6 of 19 doi: 10.14411/fp.2015.039 Moravec and Van As: Fish nematodes from Botswana
A B
C D
E F G
Fig. 6. Cucullanus sp. female from Tilapia sparmanni Smith, scanning electron micrographs. A–C – cephalic end, sublateral, apical and ventral views, respectively; D – tail, ventrolateral view; E, F – circumoral denticles in different views; G – deirid. Abbreviations: a – amphid; b – cephalic papilla.
Mohammed, 2014, have recently been described from the cies can be distinguished from Cucullanus sp. by the cos- [ Mugil curema Valenciennes (Mugilidae) in picuously wide and short pseudobuccal capsule, similar to Senegal (Ndew et al. 2014) and the European eel Anguilla those in C. barbi, C. baylisi and C. djilorensis (the shape anguilla (Linnaeus) in the Gulf of Suez (Red Sea, Egypt) of the pseudobuccal capsule was neither described nor il- (Abdel-Ghaffar et al. 2014), respectively. lustrated for C. clarotis). Moreover, the roughly granular ? [ ! Cucullanus sp. of surface of eggs, as observed in Cucullanus sp., was not de- the present material remains unknown, the available female scribed for any of these species. Since this Cucullanus sp. can be easily distinguished morphologically from all of the also differs from other African congeners in the host order above-mentioned species of Cucullanus by the distinctly (Perciformes vs Anguilliformes, Cypriniformes, Mugili- elongate pseudobuccal capsule and, especially, by the loca- formes and Siluriformes), it is apparent that it represents > ! - a hitherto undescribed species. @ 5 > Two species of Cucullanus, morphologically somewhat 5 7 > similar to the present African Cucullanus sp., were de- deirids are located much more anteriorly, at the oesopha- 7 ! [ ! ! >- geal part of the body, in C. barbi, C. baylisi, C. clarotis ico: C. angeli 7 @ @ 5 and C. djilorensis= > V 2007 from Vieja intermedia (Günther) and C. oaxaquensis described for C. egyptae, but this poorly described spe- @ 5 " @ @
Folia Parasitologica 2015, 62: 039 Page 7 of 19 '+ '99''H ;+'- +&F Moravec and Van As: Fish nematodes from Botswana
Fig. 7. Cithariniella longicaudata sp. n. from Schilbe intermedius 6 A M ! V= B, C M ! V = D M ! V= E M ! V= F M V= G M ! V= H M
6 ! ;+'+ ! Thorichthys helleri Steindachner (see shape of deirids (rounded in the former species vs sharp@ 7 @ @ 5 ;++* @ ly pointed in the latter), egg surface (roughly granular vs @ 5 ;+'+R ! smooth), environment (freshwater vs marine) and the host species has unusually rounded deirids and the latter species family (Cichlidae vs ! { 5 R possesses eggs with a roughly granular surface, the fea@ tures that are found in the present Cucullanus V Oxyuroidea 77 '
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A
C D
E F G
Fig. 8. Cithariniella longicaudata sp. n. female from Schilbe intermedius 6 ! A, B M V = C M = D M 7 V= E M V [ ! = F M @ $ 7 = G M V Abbreviations M ! = 7 M = M ! !
V E ! N 7 7 R N; @ > ! " V '+< V = teral and 4 ventrolateral) protruding out of mouth, 4 oval ! ;* N *? R ^ 7 ! @ cephalic papillae and pair of lateral amphids (Figs. 7C, &E 6 7 @ < DR V V & @ ; N' ' R 7 \ ! V @ $ @ N *? R " ! N * < R U ! EF N ''} R uniform in diameter, ending in globular bulb provided with and 540 from end of tail. Ventral precloacal pseudosucker valvular apparatus and separated from corpus by constric@ X E+F N ! R N * ^R V ?>@ ;'} 7 N *?R cretory pore situated relatively far below level of bulb (Fig. Female (4 gravid specimens): Length of body 7A,G). 9 ''M- 99 !! ! > ! ! V ;+9M&9+ { 7 @ Male N' ! R { 7 ; << !! ! > ! ! '< ? E99M*'< = V '-+ '; ? 9** -+&M-*' V 99- ! '; V = &EF V ;9 ! ;' &+ 7 7 '''M''* '+-M'+< V V ''< 7 7 *- V E& >@ > ;&&M;9+ <-*M' ''- @ '
;+'- E; +&F F 'F '+ '99''H ;+'- +&F Moravec and Van As: Fish nematodes from Botswana posterior to bulb, posterior ovary attaining level of vulva. ;+'+R V 7 C. longicaudata. It differs W ; 9
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Fig. 9. Synodontisia annulata sp. n. from Schilbe intermedius 6 A M ! V= B M ! V= C M ! V= D M ! V= E M F M ! V= G M ! V= H M
of the Institute of Parasitology, Biology Centre of the Czech @ 6 \ X @ ! " 5 $ D5 N @F+-R ! [ > ! ? ! X ! ` 7@ found probably belonged to Cithariniella khalili, which is ková, an outstanding Czech helminthologist. known to parasitise S. batensoda in Senegal (Koubková et Without providing morphometrical data, Dougnon et ;+'+R N;+';R C. petterae, a parasite of Distichodi@ dae, from Synodontis nigrita Valenciennes and S. schall Synodontisia annulata sp. n. F '+ N $ R " 7 [ known as the hosts of C. khalili, it is highly probable that, ZooBank number for species: in this case, C. khalili V ! [ C. petterae. 7 $ ';F?'+E<@ ;E @9*<9@ -&'@' ?D ++ <; ? N;+'&R Oxyuris equi N" $ '*<
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A
C D
E F
Fig. 10. Synodontisia annulata sp. n. female from Schilbe intermedius 6 ! A M V= B M V= C M V= D M V= E M V N @ V R= F M V Abbreviations M ! = 7 M = M = M ! !
N F ^ '+ R V N F R X 7 > ! @ lateral alae arise anteriorly somewhat anterior to level of $ N F DR > Male (1 specimen): Length of body including caudal of anus in females and to short distance posterior to median $ ; -E !! ! > ! ! V ;9- F ! N F^ '+ ?R U ? &9- = ;E' ;* V = 7 E ! @ $ ! N; ! '; ;' V = 7 7 E& -* V ; ' R ! > '
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Fig. 11. AMD M Galeiceps @ ! Hydrocynus vittatus Castelnau (A M V= B, C M V = D M VR= EMH M Contracaecum @ ! Hydrocynus vittatus (E M V= F M V= G M V= H M VR
A
Fig. 12. Contracaecum @ ! Hydrocynus vittatus Castelnau, scanning electron micrographs of cephalic end. A M V= B M ! ! [ Abbreviations M = M > = M = M tooth.
Female N; ! = 7 ! ! ' 7 '-M9- { @ nongravid specimen in parentheses): Body distinctly fusi@ E ';&M;+9 ! > ! = ! 7 = ! > ! ! V ;9M&E N F^ '+ R $ 9 <
;+'- E; +&F '& 'F '+ '99''H ;+'- +&F Moravec and Van As: Fish nematodes from Botswana
A
C D E
Fig. 13. Galeiceps @ ! Hydrocynus vittatus Castelnau, scanning electron micrographs. A M V= B M ! ! [ N V ! R= C M V= D M V= E M 7 V Abbreviations M = M > = M
7 7 F+MFE FFM'+< V > (type species) and S. okavangoensis Moravec et Van As, 'F+M;-< <*+ ! > ! @ ;++9 7 ! ! $ [ Synodontis spp. tively. Location of vulva not determined due to damage of N 'F*; W ;++9R ! @ '& ;+'- E; +&F '9 'F '+ '99''H ;+'- +&F Moravec and Van As: Fish nematodes from Botswana A C D Fig. 14. Eustrongylides ! @ ! Hydrocynus vittatus Castelnau, scanning electron micrographs. AMC M @ 7 V = D M V Abbreviations M > = 7 M = M ! = M ! Ascaridoidea 6 'F'- X [ Hydrocynus vittatus Castelnau (Alestidae, ! R 7 -+' !! 7 [ Clarias Anisakidae 6 'F'; stappersi N " ! R 7 ;;+ mm, rainbow happy, Sargochromis carlottae (Boulenger), body 'FF !! Serranochromis angusticeps Contracaecum sp. @ N R 7 ;E+ !! ! Ser- '' ?M '; ranochromis macrocephalus N R 7 ;E< !! and southern mouthbrooder, Pseudocrenilabrus philander (We@ Description N; ! ! Hydrocynus vitta- ber), body length 50 mm (all Cichlidae, Perciformes). tus R V 'F &M;' - !! Site of infection: Viscera. ' ;9M' && !! V X ! Locality: U$ 6 N $ '< ;-'F *"= 7 N ''? ^ '; R ;' -E'F F?R V N ;+ ;++'R rounded, bearing small ventral cuticular tooth and 4 ce@ Prevalence: H. vittatus F} N' [ H'' [ > ! R= phalic submedian cephalic papillae surrounding small C. stappersi 'HE= S. carlottae 'H-= S. angusticeps *} N'H'-R= = V $ @ S. macrocephalus F} N'H''R= P. philander 'HF N ''? ^ '; R ?> Deposition of voucher specimens: ! @ below ventral cephalic tooth. Somewhat elevated region lection of the Institute of Parasitology, Biology Centre of the of cephalic papillae surrounded by several low, circular ! " 5 $ D5 N cuticular lamellae interrupted by narrow smooth lateral @9E ;+'- E; +&F '- 'F '+ '99''H ;+'- +&F Moravec and Van As: Fish nematodes from Botswana In Africa, morphologically similar larvae were reported, ! " 5 $ D5 N 7 @U V N;++ER ! Clarias @'+<*R gariepinus (Burchell) in South Africa. Larvae of Contracaecum 7 @ Remarks. By the general morphology, especially the ! N ! E+R V structure of the cephalic end, these larvae belong to the [ 7 ! genus Galeiceps 6 'F'E N Cloeoascaris Baylis, > ! D ! 6 7 ? 'F;&R V 7 ` !7 " " @ ! ! ! !! N 7 'FE*R " X M K !7 7V N @ the type species, G. cucullus N{ V ' ;+'- E; +&F 'E 'F '+ '99''H ;+'- +&F Moravec and Van As: Fish nematodes from Botswana ' +F !! ! > ! ! V > ! ability of Eustrongylides @ 7 7 N &? '9 DR V V > ! ! N '9 MDR U 7 W N'F99R Eustrongylides @ ! 7 '; are occasionally found as the parasites of humans after be@ ; = ! 7 ; ; 7 V V [ = ; 7 = ! V usually penetrate through the wall of the caecum into the $ @ $ ! ! !7 peritoneal cavity and can be removed only during surgery ! 5 @ N?7 'F REFERENCES ǃDždžǍ@^ljǂLJLJǂǓ ǂǔljǕǂǓ @6 ǃDždžǍ@^ǂǃdžǓ 6 ǐǓǔǚ ǏDždžǓǔǐǏ 6 ;+++ ! W 7 X ` džljǍljǐǓǏ Ǎ ¶ǖǓǂNJǔljǚ " ǐljǂǎǎdžDž " ;+'9 D ! X ! " ? 7 @ Cucullanus egyptae sp. nov. (Nematoda, Cucullanidae) infecting ] V » $ E-+ ? Anguilla anguilla ? ǏDždžǓǔǐǏ 6 ljǂǃǂǖDž ^ ]NJǍǍǎǐǕǕ " N? ) ;++F ! 6 ''& &9-*M&9E- Keys to the Nematode Parasites of Vertebrates. Archival Vol@ Ǎ@ ǂǔǔdžǍ D ;++& ! ! ] 9E& [ ! V ! ^ ǂnjdžǓ 6 'F<; " ! ? 6 F+ '&-M'&F Cosmocercidae (Nematoda: Cosmocercoidea): two new atractids ǎNJǏ U 'F*< ! ! [ @ ! 7 [ K E+ ;&F-M;9+; ? V Camallanus kirandensis Baylis, ǂǓǔǐǏ ǗdžǏǂǏǕ@UǍDždžǘǂLjdž ;++E ! 'F;< N ! R Bothriocephalus aegyptiacus 6 º of Clarias gariepinus N '<;;R ! 6 D ! 'F*- N R E9 FF&M''+' " U W 6 *& <*MF9 ;+'- E; +&F '* 'F '+ '99''H ;+'- +&F Moravec and Van As: Fish nematodes from Botswana ǂǚǍNJǔ 'F;& " ! ! Cucullanus from `ljǂǍNJǍ { 'F*9 " ! ! ! V [ [ " F '; ;&&M;&E 6 V V Cithariniella pet- ǂǚǍNJǔ 'F;&7 V ! terae ! ! 9< <''M<'< " F ; 9-FM9E& `ljǂǍNJǍ { ǐǍǍNJǏLj { 'FF* $ { ! @ ǂǚǍNJǔ 'F;< " ! V ! ! ! [ ! V " ? M { $ X $ " '+ 9 &*;M&<' 7 '<- ǂǚǍNJǔ 'F&+ " @D 'F;*M `ljǂǍNJǍ { XljǖǓǔǕǐǏ 'F*& " ! @ 'F;< ! ;e V [ M ; ''*M'&+ V V 6 K <* ;+FM;9< ǂǚǍNJǔ 'F9+ U V ! ! `ǐǖǃnjǐǗǤ ǂǓǖȪ W ǐDžǐǗǤ ;+'+ ! Citharin- " '' ;F 9+'M9'* iella N R ! V [ " V ǐǐǎnjdžǓ 'FF9 " V [ W $ ! 9* '+-M''9 ! ! [ ! 7 D ! `ǐǖǃnjǐǗǤ ǂǓǖȪ W ǐDžǐǗǤ NJǎnjǐǗǤ ;++< @ X U W 6 E' 'F*M'FF metric and molecular characteristics of Labeonema synodontisi ǂǃǂDzǂǔ@ ǂǓǓǂǏǛǂ ^ ǂǔǑdžǕǂ@ ǂǏDžǖNjǂǏǐ ;++* !7 N ! R ! ] [ V N ! R ! V [ 6 '+; '+'&M'+;+ Vieja intermedia N^ ' ;+'- E; +&F '< 'F '+ '99''H ;+'- +&F Moravec and Van As: Fish nematodes from Botswana Dždžǘ D ? DNJǐǖLJ ǥ X ǐǓǂǏDž " ;+'9 V "NJnjǐnjNJ Ǜdžǂnjǐ " DŽljdžLjdž ;+'& ? ! of Cucullanus (Nematoda: Cucullaninae, Cucullanidae) from parasitemia in Oreochromis niloticus from Lower New Calabar Mugil curema (Mugilidae) in Senegal (West Africa). Comp. 6 D " 6 ? <' '-M;; " ' ;E&M;E* UǕǂDŽljNJ ? U `ǷǓǏdžǓ ] ǗdžǏǂǏǕ@UǍDždžǘǂLjdž džǍǍǏdžǓ@ "ǑǓdžǏǕ 'F<; ! " ! ! !@ ǓǂǏnj NJǓǔǂ ;+'9 X ! ! 7 ! V [ V ! -E ;*-M spotted tilapia Oreochromis leucostictus NX V 'F&&R ;F- its parasite Contracaecum multipapillatum from Lake Naivasha, WǂǏ ]ǂdžǓdžǃdžnjdž D ljǂǃǂǖDž ^ ǂNJǏ U ^džǐǓLjdžǔ @ ` ! $ ? 'F<< D > > ! " 6 ;' *&&FM*&9F RU 7 9 ǂǑdžǓǏǂ 'F*9 7 '+ -'FM-;* with larvae of Eustrongylides (Dioctophymidae, Nematoda) in WǂǔǔNJǍNJǂDžǪǔ ^ 'F*- ! R [ ! ? $ E E*M*E 6 7 " ! " džǕǕdžǓ @ {dž džǍ 'FF; XV V Cuc- &* E+-ME'< ullanus N ! M R ! WǂǔǔNJǍNJǂDžǪǔ ^ XǓǐǏDŽǚ @ 'F*9 ! ! U V <* N" R ;+'M;+E 7 " džǕǕdžǓ @ WǂǔǔNJǍNJǂDžǪǔ ^ XǓǐǏDŽǚ @ 'F*; X @ &E E*+ME<' R > WǐǏ ǓǂǏDž X 'F99 7 Eus- ! ! 9* -EFM-*F trongylides. IV. Transmission to various coldblooded intermedi@ ǖǚǍǂdžǓǕ 'F*+ D Labeonema intermedia gen. ! " ] '' ;&M;* R Labeo N @ R N M »ǐǐǚdžǏ X ǐǓǂǗdžDŽ ]ǐǏLjǔǂǘǂDž ;+'' XV V ! M W ! R 6 K <; &*+M&*E of Cucullanus '*** N ! R ! ! @ "DŽljǖǖǓǎǂǏǔ "ǕdžnjljǐǗdžǏ 'F&* ! ?> @ [ X " *< '&FM'9F 7 ^ ] N'F&&M 'F&-R > 9 &M9+ 6 ;& 7 ;+'- - ;+'- 7 ;; ;+'- Cite this article as: W { { ;+'- " > ! N ! R ! [ U$ 6 V E; +&F ;+'- E; +&F 'F 'F