Baqri&Coomans 1973 - Dorylaimida Descritos Por Stekhoven y Teunissen 1938

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    FONDATION POUR VORISER

    L

    RCHRC SCIETIFQU EN FRIUE

    RE CH ERCHS NTREPRSS

    PAR 'NSTTUT NATONA

    POUR A CONS ERVATON E A NATURE

    E

    PL Z0i

    Exploration u Par Nonal des Virunga

    F DE WTTE (1933-1935) et H D (1935-1936)

    F 1

    X Y

    KV & U (1938) and KV (944)

    1- DYLIMID, LIID and LGIID

    BY

    Q. . BAQRI A COOMANS Get

    *BRUXES973

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    PAC NATNA S VUNGA

    MSSNS G. F. WTT (1933-1935) et H AMAS (19351936)ascicul 1

    A TAXONOMIC REVISONOF THE NEMATODE SPECIES DESCRIBED BY

    S. STEKHOVEN & TEUNSSEN 38 and S. SEKHOVEN 44

    FROM NATIONAL VIRUNGA PARK(ZAiRE REPUBLIC)

    I - DORIIDE, ORELIIDEand LOIDORIE

    BY

    Q H BAQR and A. COOMAN (Gent)

    INTRODUCTION

    Ths s the frst paper deag wth a revson of the taxonomc worpubshed by CHUURMANS EKHOVEN & TEUNISSEN 1938) and CHUURMANSEKHOVEN (1944). The nematode speces descrbed n both publcatons werecolected from the Natonal Abert Park, Congo, respectvely by G. F DE WIEn 1933-35 and by H. AMAS n 193536

    The 1938 pubcaton was composed of two parts, the frst deang wthtaxonomy and dstrbuton, the second wth bology. In the taxonomcal part,84 speces were reported from 39 sampes colected at 0 dfferent ocates.Out of these, 28 speces were descibed as new (ncludng one new genus). In the1944 pubcaton 12 speces were dentfed from 27 samples, but descrptonsand ustratons were gven for only 7 speces, two of these were new

    Some of the new speces have been descrbed on a sngle adult specmen, onjuvenles or even on a sngle juvenle Other descrptons were based on a

    In his publication, the name Congo is to be replaced by "Zare and he

    name "Par Naional Albet (P.N.A.) by "Par Naional des Virunga (PN.V.) .

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    4 PARC NAONAL ES VRUNA

    complex of different species part from this, most of the illustrations and des-criptions are inadequate because some important characters have been ignoreand obvious mistakes were made Therefore it is necessary to reillustrate andredescribe the species listed in both publications

    In the present paper the representatives of the families DRYLAAE andPORCELAAE are revised, in two subsequent papers those of the familiesNOLAAE and BELONRAE will be considered The representativesof the family NORAE have been redescribed earlier by & TARJAN(1963) and TARJAN & (1963) Here only the occasional finding of oneLongdorus species is reported

    It should be stressed that the above mentioned families are considered here asbelonging to the superfamily DRYLAOEA part from YOLAOEA

    no other superfamily is recognised under Dorylaimina cf also DE NNCK,1965 and ANS & , 1970). Furthermore the family ROORYLAAEis not accepted. The main reason for this attitude is that the authors feel thatthe recent trend to raise the rank of many taxa in the Dorylamna is notjustified detailed discussion on this problem is however beyond the scopeof this paper

    nless otherwise stated the type specimens are at present deposited in theollection of the " Institut royal des ciences naturelles de Belgique , autierstraat 31, Brussels It is, however not excluded that they will be transferred

    later on, either to the " Institut des ars ationaux du ongo or to the" Muse royal de frique entrale

    MATERIALS ND METHODS.

    The whole material was deposited i the oyal nstitute of atura ciencesat Brssels nstitut royal des ciences naturelles de Belgique) Through thecourtesy of rof W A the material was made available for present study

    The nematodes were preserved in formalin and stored in glass tubes Forevery sample, each speies was kept in a separate tube Most of the tubes werenumbered and catalogued with fu informations corresponding to the publications,

    but on some tubes the details about sample number and locality were missingThe number of specimens was often mentioned, but occasionally a lower ofhigher number was found in the tubes Few tubes were found empty omenematodes were in poor condition or vey dark

    The dark specimens had to remain several years in glycerine for clearing,others were processed according to the einhorst (1962) as modified by

    DE GRSSE (1969) ll specimens have been mounted in dehydrated glycereon aluminium slides camera lucida has been used for the diagrams andfor the measurements GERAER's (1961) formula has been used for correctingthe body width measurements in flattened specimens ositions of sophagealgland nuclei and outlets are given according to & ANS (1970)

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    PARC NAONAL DES VIRUNA 5

    umerical data are presented mostly in tabelized form. The folowing terminology and abbreviations have been used in the tabes :

    L, a, b, c, , T, G G2: usual modified de Man formula.

    cut m : thickness of cutice at mid-body in microns.

    cut t : maximum thickness of cuticle on tai.

    lat b. p. tota number of lateral body pores.

    lat. b. p. oes. : lateral body pores in oesophageal region.

    lat b. p. ca: lateral body pores between cardia and vulva region.

    lat. b p prer. : latera body pores between vuva and intestine-prerectumjunction

    lat b. p. prer+

    r : lateral body pores in prerectum and rectum region.c. p.: caudal pores

    vent. b p. : ventral body pores

    dors b. p: dorsal body pores

    A: width at neck base divided by lip region width

    B: ip region width divided by lip region height. Height of the lip regionmeasured from anterior extremity to the labia constriction or to the amphidial aperture in the case of no offset lip region.

    amph w: width og amphidial sit in microns

    sens. : distance of sensilla pouches from the amphidial sit in microns

    od. st. : length of odontostyle in microns

    od. st ap. : length of odontostyle aperture in microns

    g. r. : guiding ring from anterior extremity in microns

    od ph : length of odontophore in microns.

    prer. : length of prerectum in microns.

    r : length of rectum in microns

    an dia.: body diameter at anus in microns.

    t. length of tail in micros

    nrv r. : distance from anterior extremity to the middle of nerve rg in microns

    egg : length and width of the egg.

    spic : length of spcules aong the curved median ine in microns

    1. g. p. : length of latera guiding pieces in microns

    supp : number of ventromedian supplements.

    s v. p. : number of subventral papilae

    cop. mus. : number of copulatory muscles.

    sperm: length of sperms in microns

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    6 PARC NAIONAL DES VIRUNGA

    DESCRIPTIONS

    n the folowing table the species described by s. EKHOVEN & TEUNISSEN(1938) and EKHOVEN (1944) are isted on the eft side under the namementioned in those pubications A singe asterisk refers to the 1944-, a doubeasterisk to the 1938paper. n the right side the present status of the species

    is given

    Status of Dorylaimidae and Aporcelaimidaedescribed by s. SEKHOVEN EUNIEN 19 38 (**) and S STEKHOVEN 1 944 (*)

    Past

    * Dorylaimus filiformis ATIAN, 1 865

    * Dorylaimus stagnalis DuJARIN, 1845

    * Dorylaimus longicaudatus iTHLI,1874

    * Dorylaimus ruwenzorii DE CONINK,1935

    * Dorylaimus effilatus S STEKHOVEN &EUNIEN, 1938

    Dorylaimus filiformis ATIAN, 1 865

    * Dorylaimus parafecundus DE CONINK,1935

    * Dorylaimus parhomalopapillatus SSTEKHOVEN, 1944

    Dorylaimus flavomaculatus VON INTOW 1876

    * Dorylaimus granuliferus COBB, 1893

    * Dorylaimus iners AIAN, 1865

    * Longidorus multipapillatus S STEKHOVEN EUNIEN, 1938

    * Dorylaimus obtusicaudatus ATIAN,1865

    ot incuded in the present study :

    Present

    1 Prodorylaimus paralongicaudatus (MIO-LETZKY, 1925) NRY, 1959

    2 Prodory laimus spec.

    Dorylaimus stekhoveni sp n

    Dorylaimus spec

    Mesodorylaimus ruwenzorii (DE CONINK,1935) NRY, 1959

    Mesodorylaimus effilatus (S STEKHOVEN EUNIEN, 1938) comb n

    Mesodorylaimus kamandeanus sp n

    esodorylaimus spec

    Laimydorus parhomalopapillat1 (S STEKHOVEN, 1944) comb n

    Laimydorus spec

    Eudorylaimus granuliferus (COBB, 1893) NRfY, 1959

    Eudorylaimus spec

    Longidorella multipapillata (S STEKHOVEN EUNIEN, 1938) SIIGI, 1962

    1 Aporcelaimellus otusicaudatus (ATIAN, 1 865) LTHERR, 1 96 8

    2 Aporcelaimellus kikereensis sp n

    3 Discolaimus major HORNE, 1939

    * * Dorylaimus metobtusicadatus EKHOVEN & TEUNSSEN, 1938 n this

    species wi be reported esewhere

    * * Dorylaimus infeundus OBB. pecimen ost.

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    PARC NATIONAL DES VIRUNGA 7

    ** Dorylaimus lentifer . TEKHOVEN & EUNISSEN 1938 pecimen osten. et species inquirendae probably beonging to Aporcelaimidae.

    * * Dorylaimus maximodorus TEKHOVEN & EUNISSEN, 1938 pecimen ostn view of the body ength (314 mm) and the fact that it apparenty isa first stage juvenie (cf Fig 71 of s. TEKHOVEN & EUNISSEN, 1938)this species may beong to Ischiodorylaimus, hence becomes Ischiodorylaimusmaximodorus ( TEKHOVEN & EUNISSEN 1938) comb n t shoud

    however be considered as species inquirenda til topotypes have been found

    ** Dorylaimus spec pecimen ost

    * * Dorylaimus stagnalis JARDIN pecimen ost

    * * Aporcelaimus eurydorus (ITLEVSEN) pecimen ost

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    8 PARC NATIONAL DES VRUNGA

    D E

    F

    J

    FIG. 1 - Prodorylamus paralongicaudatus (MoLETZKY 1925).

    A and B : Anerr regns; C : surface vew f anerr regn; D and :E n f a c e vew; F : Crss secn just psterr lp regn; G : ntre

    mae; H : Femae ral regn; I : Basa expanded prtn f espags, esphageal gand nucle and rfces : Oespag-ntestinal junctn, carda;K : Female pserr gnad; L : Spcar regn; M : Psterr bdy regn

    f mae; N : Spermatza.

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    PARC NATIONAL DES VIRUNGA 9

    Family DORYLAIMIDAE DE MAN,

    1876.Genus PRODORYLAIMUS NDRSSY, 1959

    s TEKHOVEN & EUNISSEN (1938) reported Dorylimus filiformis BASTIAN,1865 from varous locaities. For present study two females of Sesero-popuationone female and one male of Tshumba-popuaton, one partally dstorted femaleand seven juveniles of Tshengelero-popuation and one totaly distorted femaleof Mubiliba-popuaton were availabe. None of these specimens representsParadorylaimus filiformis (BASTIAN 865) DRSSY 1969 but ey representtwo distinct species of the genus Prodorylaimus NDRSSY, 1959

    The two females from Sesero-population are described below as Prodory-laimu spec. the remaining specimens have been identified as Prodorylaimusparalongicaudatus (MICOLETZKY, 1925) NDRSSY 1959.

    Prodorylaimus paralongicaudatus (MICOLETZKY, 925) NDRSSY, 1959.

    Syn. Dorylaimus filiformis apud s. TEKHOVEN & EUNISSEN, 1938 partim

    Ne Dorylaius filiformis BASTIAN, 1865= Paradorylaimus filiformis (BASTIAN, 1865) NDRSSY, 1969

    M e a s u r e m e n t s . Refer to Table 1 (Fig 1)

    D e s c r i p t i o n . Femae : Body ventrally curved in the posterior half,tapering graduay towards both extremities. Cuticle finely striated transversally.Latera chords about /5th of the body width near midde.

    Lip region narrower than adjoning body, slghty marked by a depression.Lps bearing the usual number of papae, n two circlets (10+6) the submedan papillae occupy a anterior position (Fig 1 D) Amphids deepstirrupshaped; st occupyng about haf of the corresponding body width. Sensilarpouches not seen. Odontostyle about two lip regon widths long, its lumensightly wider than corresponding body cuticle; aperture ess than half the odotostye length Gudng rng 2- 3 ip region widths from anterior end

    Foremost portion of oesophagus forming an eipsoida, distincty offset

    swelling The constriction separating this sweling from the remainng part ofthe oesophags has been called " postextension constricton here it wll becaled " postodontophore constriction Basa expanded portion of oesophagsoccupying 47-50 % of the neck region Positions of oesophageal gland nucleiand orifices as follows :

    D51-53.6DN=5457

    DO-DN = 2.9-3 .4

    K=79-83

    S SN = 70-71S SN = 74-75

    SN = 82-83S=83-84

    K' = 8185

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    10 PAC NATIONAL DES VIRUNGA

    FIG 2 - Prodorylaimus spec.

    A : Entire female; B : Basal expanded portion of oesophagus, oesophageal

    gland nuclei and orifices; C : Femae tai region; D : Femae anterior goad;

    E : Surface view of anterior region; F : Anterior regio.

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    PARC NATIONAL DES VIRUNGA 11

    Cardia eongate-cood, less than half of the correspondng body width long.

    A narrow oesophago-intestinal disc present. Prerectum 2.53 anal body widthsong, with a sngle median pece. Lateral guiding peces more or less rod

    Vulva transverse. Vagna more than /3rd of corresponding body widthlong, with weakly sclerotized distal part appearng as two pieces in lateral viewGonads amphdephic. Uterus much onger than oviduct. Ovaries with oocytesmosty arranged in doube row.

    Tail very long, fiiform with acute terminus; about 23-24 anal body widtslong; with 3 cda pores on each sde. Male : Smilar to femae in generashape and morphology except for the more curved posterior half. Mae gonadstypical. Spermatozoa ova in shape. Spices about 1.9 ana body widthsong, with a single median piece. Lateral guiding pieces more or ess rod

    shaped In addition to the adana pair, 20 ventromedian supplements present,spaced closely at regular intervas. Eleven irreglary spaced subventra papilaereach from near the anus up to the foremost supplement.

    Prerectum about 4 anal body widths ong startig from halfway the supplement region. Tal similar to female tou possiby shorter (exact length notknown because of broken tp !); with 8 cauda pores on each sde.

    L o c a 1 i t i e s a n d h a b t a t s . Sample no. 377 : colected froTshumba, Mushari regon, SouthWest of Rutshuru; Alt 2,100 m near the vage and Alt. 1 ,700 m in the ava plain. Volcanic shnge amost without organccomponents.

    Sampe no. 5 33 : collected from Tshengelero, swamp, North of Munagana;Alt 1,750 m.

    Sample no. 1518 : colected from Mubiiba, smal wood in the ava plain ofthe Nyamuragira vocano; At. 2,000 m.

    R e m a r k . There is no record of specimens from Tshuba bys. TEKHOVEN & EUNISSEN (1938, p. 114 or p 137). The male ad femaledescribed and reported here have been found in a tube abeed as Dorylaimusfiliformis, Tshumba, sample no. 377.

    Prodorylaimusspec.

    Syn. Dorylaimus filiformis apud s. TEKHOVEN & EUNISSEN, 1938, partim.

    (Fig. 2.)

    M e a s u r e m e n t s . Refer to Table 1.

    D e s c r i p t i o n . Female : Body irregulary curved, gradually taperingtowards both extremites Cuticle finey striated transversally Lateral chordsabout 1/5-1/4th of the body width near middele. Lateral body pores faint.

    Lip region narrower than adjoining body and marked y a epression Lipsbearing the usual number of paplae in two circets (10+ 6). Amphids deep,stirrupshaped; slit about half of the corresponding body width. Positon of

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    12 PARC NATIONAL DES VIRUNGA

    F l

    JB

    FIG 3. - Dorylaimus stekhoven n sp

    A : Entire female; B : Anteior region; C : Surface view of anterio egion;

    D : Femae anterior gonad; E : Basal expanded portion of oesophagus, oeso

    phageal gland nucei and orifices; F : Vuva! egion; G : En f a c e view;

    H : Coss section just below ip egion; I : Coss section through basa

    expanded portion of oesophagus; J : Coss section through mid-body

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    PARC NATIONAL DES VIRUNGA

    sensilar pouches not certain because of the presence of two swellings (Fig. E).

    Odontostyle 1.8-19 lip region widths long its lumen wider than the correspondingbody cuticle; aperture slitly less than half of the odontostyle lengthPostodontophore constriction present Basal expanded portio of oesophags

    occupying about 49-5 % of neck region Positios of oesophageal andnuclei and orifices a follows

    DO=486-56DN=53-55

    DO-DN= 1719

    K=8586

    S0 SN =69-71S0 SN2= 737 5

    S2N=84-85S20=8586

    K'=87

    Cardia conoid with broadly rounded tip. A very narrow oesophago-intestinaldisc present Prerectum about 45 anal body widths and rectum about 15 analbody widths long.

    Vulva transverse Vagina slightly less than half of the corresponding bodywidth with weakly sclerotized distal part appearing two pieces in lateral viewUterus very long with " Z-organ composed of weakly sclerotized apophysesOviduct very short. In one female the ovaries are narrow with smal oocytesarranged in a double and even tripe row (Fig D); in the other specimen theovaries are wider with oocytes arranged in a double and sine row. Spindleshaped sperms are present in the uteri of one female.

    Tail long, filiform with acute terminus; 9-1 anal body widths long; 4 caudlores on each side

    Male : not found.

    L o c a 1 i t y a n d h a b i t a t Sesero Mountain North of the Mikenovolcano bambou and mixed wood region of Gashole-Sesero ontais; Alt m.

    R e m a r k - Prodorylaimus spec. comes very close to P dahli (LTHERR196) NDRSSY, 1964 in many respects excepts in having a stouter body(a=554 in P. dahli) Since we have only two females and males were notpresent in the collection the species . which may be new is not named.

    Genus DORYLAIMUS uJARDIN, 1845.

    Dorlaimus stekhoveni sp n

    Syn Dorylamus stagnalis apud S TEKHOVEN 1944.

    Ne Dorylaimus stagnais uJARDN 1845

    (Figs. 3 and 4)

    Dorylaimus stekhoven n. sp was collected in large number from differentlocalities and was identified by S TEKHOVEN (1944) as D stagnalis uJARDN,1845

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    14 PARC NATIONAL DES VIRUNGA

    FIG 4 - Dorylamus stekhoveni n. s

    A : Male tai region; B : Femae tail region; C : Surface view of posteriorbody regon of male, sowing supplements subventral papilae, latera body

    pores; D Posteror end of male body, showing ejacuatory gands; E Spiculesand atera guidn pieces; F : Posterior body region of mae

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    PARC NATONAL DES VIRUNGA 15

    For the present study, many specimens were availabe fro Gando (sampesnos. 42, 46 and 49), one female from Kisenyi (sampe no 261) and oe juvenile

    from Ondo (sample no 287).D. stagnalis was also reported by S. TEKHOVEN & EUNISSEN (1938) but

    these specimens (1 0, 2 uv.) could not be traced in the colection.

    M e a s u r e m e n t s . Refer to Table 2.

    D e s c r i p t i o n . Female Body straight or ventrally curved posterior tovuva, sightly tapering towards both extremities. Cutice finey strated transversaly, marked wth 41 longitudinal ines in the middee and at base of oesophagus (Fig 3 I, J). Lateral chords about 1/4th1/3rd of the corresponding bdywidth near middle

    Lip region slightly narrower than adjoining body, slightly marked by adepression. Lips amalgamated, bearing the usua number of papiae in twocirclets (10+ 6); subdivided four sects (Fig. 3 G) Oral aperture hexagonabut dorsoventray eongated. Amphids stirrup-shaped, sit about half of thecorresponding body wdth. Odontostye 2326 ip region widths long; itslumen about as wide a the corresponding body cutice; aperture ess than hafthe odontostyle ength Guiding ring 1 .619 lip region widths from anteriorextremity.

    Basa expanded portion of oesohagus 5255 % o neck regon. Positionsof oesophageal gland nucei and orifices as follows :

    DO = 46.949.0DN=480-509

    DODN = 1 . 1- 19

    K= 7177

    S, SN = 6668S, SN = 73-76

    SN=8385S = 8486

    K' = 7378

    Cardia cyidrcal to tongueshaped, surrounded by intestina tissue. Prerectum 58 ana body widths ong; cels usualy wth wel developed microvili.Rectum 1 .317 ana body widths long, separated by a wel deveoed sphincterfrom the prerectum.

    Vuva transverse. Vagina 4256 ong, extending less than halfway the

    correspondng body width, with scerotized distal part appearing as two refractivepieces in atera view. Goads amphidelphic. Usualy a female contains 23 eggs,but in one femae there are 1 1 eggs in the anterior uterus and 12 in the posteror.

    Tail elongate, taperng gradually, 57 anal body widths long; wth 23 caudalores on each side.

    Mae : Similar to female in genera shape and morphoogy except for themore ventraly curved posteror part of the body and the differently shaped tail.Male gonads typica Spermatozoa eiptical Three pairs of ejacuatory gandsare visible in the prerectum region; the posterior part of the ducts could not bediscerned; because of distortion the six glands may be visible from one side ofthe body (Fig. 4 D).

    2

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    16 PARC NATIONAL DES VIRUNGA

    B

    U

    l 0

    " ,

    FIG. 5 - Dorylaimus spec

    A Anterior region; B : Surface vew of anteror region; C Poseror body

    region of female; D : Basal expaded porion of oesophagus, oesophageal gland

    nce and orifices; E : Cross secion through mid-body; F Surface viewin mid-body, ongitudnal lnes and aera body pores; G Vulval regon

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    PARC NAIONAL DES VIRUNGA

    picules 224 anal body widths long, with a simple median piece ateral

    guiding pieces more or less rodshaped n addition to the adanal pair, 55-62contiguous and regularly spaced ventromedian supplements are present ubventral papillae spaced irregularly, reacng anterior to junction between intestine and prerectum; one postanal subventral papilla also present umerouslateral body pores occur the caudal region (Fig. 4 )

    rerectum starting anterior to the copulatory muscles; 9- 3 anal body widthslong Tail convexconoid with rounded terminus, about 3/4th of anal bodywidth long; with 02 caudal pores on each side

    i f f e r e n t i a d i a g n o s i s - Dorylaimus stekhoveni n sp comesclose to D. tepidus NDRSSY, 959 but diers from it in having more longi

    tudinal lines (32 in D. tepidus), comparatively longer odontostyle (21-22 lipegion widths long in D. tepidus); longer spicules (10006 in D. tepidus) andmore supplements (35-39 in D. tepidus). t also resembles D. montanusEANSKI, 923 but difers from this species by body length ( = 3.2-48 mmin D. montanus) an number of supplements (2535 in D. montanus).

    T y p e s p e c i m e n s olotype female along with one paratype femaleon slide Dorylaimus stekhoveni / aratype males and emales on slidesD. stekhoveni /2-31

    T y p e 1 o c a i t y a n d h a b i t a t pecimens collected by AAS

    from the upper lake at ando

    0 t h e r o c at i e s ne female from isenyi, dward ake; onejuvenile female from the bank of ndo river

    Dorylaimus spec.

    yn. Dorylaimus longicaudatus apud . EKHOVEN & TEUNISSEN, 938 partim

    e Dorylaimus longicaudatus BSCHLI, 874=Prodorylaimus longicaudatus (BSCHLI 874), RSSY, 959

    (Fig 5.)

    . EKHOVEN & TEUNISSEN (938) reported Dorylaimus longicaudatus fromala, Tshumba (sample no. 377), ibga (sample no 075), asenze (sampleno 204), utshuru, wepa river (sample ) and windi (sample ) Figre64 B in their publication represents a male tail of a specimen that probably

    belongs to the genus Prodorylaimus.nortunately only one specimen (a female from ibga) wa present in

    the collection This female belongs to the genus Dorylaimus s in somother instances, EKHOVEN & TEUNISSEN (938) identified here a complexof diferent speces belonging to different genera as one species

    M e a s u r e m e n t s efer to Table 2

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    8 PARC NATIONAL DES VIRUNGA

    D e s c r i p t i o n Female : Body slightly curved ventrally slightly tape

    ring towards both extremities Cuticle finely strated transversally marked with44 lontudinal lines in the middle of the body Here and there some faint linesvisible (Fig 5 F); in transverse section they appear as minor projections betweentwo major ones (Fig 5 E) Lateral chords 1/5th of body width near middle

    Lip region wider than adoining body offset by a moderate constriction.Lips subangular in superficial view; bearing the usual number of papillaeAmphids cupshaped, slit less than half of the corresponding body widthOdontostyle 21 lip region widths long; its lumen about equally wide as thecorresponding body cuticle; aperture slightly more than /3rd of odontostylelength Closed part of odontophore well developed, appearing in lateral viewa a 27 long sclerotized rod at the ventral base of the odontostyle (Fig 5 A)

    Guiding rng 13 lip region widths from anterior extremityBasal expanded portion of oesophagus occupying about 56 % of neck region

    Except for the dorsal oesophageal gland nucleus no gland nuclei could beobserved but gland orifices are well visible :

    D0=47DN=483

    DDN= 13

    S0=6S0=66

    K'=737

    S0=78S0=785

    Cardia about half of the corresponding body wdth long conoid envelopedby intestinal tissue its anterior fourth is darker than the rest Prerectum about5 anal body widths long Rectum less than /2 anal body widths long

    Vulva transverse Vagina about half the corresponding body width withweakly sclerotized distal part Gonads amphidelphic details not clear

    Tail elongate gradually tapering tip ventrally bent and forming a hook;bout 6 anal body widts long; with 4 caudal pores on each side.

    Male unknown

    D i f f e r e n t i a d i a g n o s i s - Dorylaimus spec. comes close toD montanus TEFANSKI, 923 in having 44 longitudinal lines but differs in theodontostyle being abot equally wide as the correspondg body cuticle (widerin D. montanus), larger odontostyle aperture (lss than 1/3rd in D montanus),differently shaped amphids and comparatvely longer oesophagus (b =43-56in D montanus) Dorylaimu spec may represent a new speces but as themateral is insufficient we prefer not to name it

    L o c a 1 i t y a n d h a b i t a t Kibga, southern slope of Visoke volcanoat the fringe of a bambou forest; Alt 2,4 m

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    PAR NATIONAL DES VIRUNGA 9

    THE GENEA MSODORYLIMUS AN LIMYDORS

    The genus Laimydorus was differentiated from Mesodorylaimus mainly onthe basis of the following characters : guiding ring double (L) against single (M),vulva longitudinal (L) against transverse (M), supplements contiguous (L)against mot (M), prerectum in the male extending well beyond the supplementregion (L) against not extending beyond this region (M).

    On this basis many species can be attributed to either of these genera Somespecies, however present a mixture of characters considered typical for bothgenera The diagnostic value of " single or " double giding ring has

    been discussed by ooDEY (96) and CooMANS (1964) Both genera have thefixed guiding at such a position that only a slight shift of the spear wil make theguiding ring appear as " double i those species were the " single conditionexists when the spear is fuly retracted Hence this character is not very reliableto differentiate both genera. The shape of the vuva slit has not been describedfor a number of species; combined with other characters it may be perhaps agood character The distribution of the supplements may in other dorylaimidgenera vary a lot and the reliability of this character for distingishing Laimydorusfrom Mesodorylaimus has to be further documented. It should be stressedthat some species currently considered under Mesodorylaimus possess contiguous

    supplements!

    The length of the male prerectum is often not reliable sinceseveral exceptions exist. Because of this some species from the present collection,beonging to one or the other of both genera were difficult t place. The namesproposed are thought to be the best compronise at present Specific problemswl be discussed under the headng " Remarks after each description

    Genus MESODORYLAIMUS NDRSSY, 959

    Mesodorylaimus ruwenzorii (DE CoNINK, 1935) NDRSSY, 1959

    Syn. Dorylaimus ruwenzorii DE CoNINK 1935.

    (Fig 6.)M e a s u r e m e n t s . Refer to Table 3.

    D e s c r i p t i o n . Male : Body ventraly curved slightly tapering towardsanterior end Cuticle fnely strated transversally Lateral chords narrow about/th of body width near middle. Only some body pores visible.

    Lip region slightly marked by a depression Lps rounded. Amphids deepstirrup-shaped; apertures less than half of the corresponding body width

    Odontostyle 13 lip region widths long ; its aperture about /3rd of thelength. Guiding ring less than one lip region width from anterior extremity;

    appearing " double because of protruded stylet.

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    20 PARC NA TION AL DES VIRUNGA

    FIG. 6 - Mesodorylaimu ruwenzorii (DE CoNNK 1935)

    A : Anterior region; B : Surface view of anterior region; C Basal expanded

    portion of oesophagus, oesophagea gland nucei and orifices; D ntire

    male; Posterior body region of ma

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    PARC NAIONAL DES VIRUNGA

    Basal expanded portion of oesophagus occupyng about 46 % of the neck

    region. Locations of oesophagea gand nuce and orfces as foows

    D0=56.DN=58

    DDN= 2.1K=8

    S0, SN76S0, SN 81

    S0, SN=9K'=81

    Exact shape of cardia not known since the testne is detached from theoesophagus and the carda partly damaged. Disc-ike porton present. Prerectumabout 5 anal body widths long; startng from opposite the first supplement.

    Gonads typical Spicules 8 ana body widths ong, with burcated distaend. Lateral gudng pieces short but wide. Suppements consist of an adanalpair and 7 regulary spaced ventromedians. Subventra papae spaced amostregularly in the region of the copuatory muscles.

    Ta dorsally convex conod with rounded terminus, about 3/4th of the analbody widths long; with 5 caudal pores on ech sde

    L o c a 1 i t y a n d h a b t a t . Visoke vocano, area wth Senecio,Lobelia ao around and insde te old crater; At 3,77 m

    Mesodorylaimus efilatus (. EKHOVEN & TEUNSSEN 938)NDRSSY, 1959.

    Syn. Dorylaimus effilatus S. EKHOVEN & TEUNISSEN, 1938.

    Laimydorus effilatus (S. EKHOVEN & TEUNISSEN, 938) DRSSY,1969.

    (Fig. 7)

    M e a s u r e m e n t s Refer to Tabe 3

    D e s c r i p t i o n . Femae : Body ventraly curved, tapering slghtlytowards both ends Cuticle fnely striated transversally. Lateral chords about /3rd of the body width near middle.

    Lip regon wider than adjoning body, offset by a constrcton Lps subanguar, with the usual number of paplae. Amphids stirrupshaped, aperturesoccupyng amost half of the correspondng body width. Sensllar pouchesnot seen.

    Odontostyle about 13 p regons wdths long; aperture somewhat more than1/3rd of the odontostye length. Gudg rng about 6 times the lip regionwidth rom anterior extremty; appearing " double because of the protruded

    position of the styet.

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    22 PARC NATIONAL DES VIRUNGA

    F

    u

    m w

    0l

    .

    .

    r- M

    FIG 7

    Mesodorylaimus effilatus (S STEKHOVEN & TEUNIEN 1938) ANRY, 959

    A : Anterior femae gonad; B Anterior region C : Surface view of anterior

    region D Basal expanded portion of oesophagus oesophageal gland nucei

    and orifices; : uva egion; F : Posteior body region of femae G : ntire

    femae.

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    PRC NIONL DES VIRUNG 23

    Basal expanded porton of oesophagus occupyng about 42 % of the neckregon Locations of oesophagea gland nuclei and outlets as folows :

    D0=63DN=626

    DODN= 2.3

    K=82

    S0 SN=78S0, SN =82

    SN=92S0=94

    K=85

    Carda conod, more than haf of the corresponding body wdth long; anteriorthird darker than remaining part Two granular bodes surround the oesophagointestna juncton (Fg. 7 D) Intestina microvill wel deveoped Prerectum

    more than 2.5 ana body wdths ong. Rectum about 2 anal body wdths ong.Vulva a trasverse st Vagna extendng more than hafway into the body

    with scerotized dista part and wel developed sphincter. Two eggs present ineach uteus; few oval sperms present n both oviducts

    Tail dorsaly convex-conoid n its anterior thrd, then suddeny narrowing toa subacute tip; 2. 9 ana body wdths long. Two cauda pores on each side Twocells of unknown nature present n the tail, each with a nuceus having a smallnucleous (Fig 7 F)

    Mae : nknown

    D f f e r e n t i a I d i a g n o s i s . - Mesodorylaimus effilatus comes clseto M pseudobastiani L, 1969 but differs by its reativey more slender body(a=37-41 in M pseudobastiani), higher vaues for K and K' (K=61-78,K'=6778 n M pseudobastiai) and by the shape of vagnal sclerotization

    T y p e s p e c i m e n s . Although S. EKHOVEN & TEUNISSEN reportedaso a juvenie (from a dffernt localty !), their descripton and ustration isentrey based on the femae specimen. We therefore do not consder thejuvenie as belonging to the orignal type series, so that the female specimenis to be treated as the hootype. It is mounted on side Mesodorylaimus

    effilatus SST 31.

    T y p e 1 o c a t y a n d h a b t a t Kanyamenoni, lake, old crater inthe bambou forest, near Tshamugussa, n the direction of Musule vocano,Bweza region; Alt. 2,3 m

    R e m a r k s . NDRASSY (1959) transferred Dorylaimus effilatus to Mesodorylaimus. In 969 NDRASSY shfted the species to Laimydorus but mentionedoonEY in oODEY, 1963 as authoity for he transfer to MesodorylaimusThis s a lapsus snce oonEY (1963) dd not list the species and NDRASShimsef proposed the trsfer in 1959.

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    24 PARC NATIONAL DES VIRUNGA

    B c

    0D

    .-E

    B

    f0

    -5 AFH

    FIG 8 - Mesodorylaimus kamandeanus n. sp

    A : Basa expanded porton of oesophagus; B : Anterior region C Surface

    view of anteror regon D E n f a c e vew E. Entire femae; : Posterior

    body region of mae; G : emae posterior gonad; H : Posterior body reon

    of female

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    PARC NATIONAL DES VIRUNGA 2

    The position of the species is doubtful, bt since the fixed guide ring is less

    than a lip region width from the anterior extremity and becase of the transverseslitlike vlva it is left nder Mesodorylaimus

    Mesodorylaimus kamandeanus sp. n

    Syn. Dorylaimus filiformis apud S. TEKHOVEN, 1944.

    Ne Dorylaimus filiformis BASTIAN, 1865

    Paradorylaimus filiformis (BASTIAN 1865) AY, 1969

    (Fig 8.)

    S TEKHOVEN (194) identified these nematodes as D. filiformis He washaving 22 feales and 26 males from Kamande (sample no 1), bt for presentstdy only 6 females and 7 maes were available. These specimens are difficultto place since they share characters of Mesodorylaimus and Laimydorus

    M e a s r e m e n t s . Refer to Table 3.

    D e s c r i p t i o n . Female : Body ventrally crved n the posterior hafor irreglarly curved, slightly tapering towards both extremities. Ctice finelystriated ransversally Lateral chords 1/6th1/th of the body width near midbody Body pores not visible

    Lip egion contiguous with adjoining body Lips with the sal nmber of

    papilae in two circlets (6+1) Amphids gobet-shped, apertres occupyingabout 1/4th of the corresponding body width.

    Odontostyle 1.21.4 ip region widths long; apertre less than half of theodontostyle length. Guiding ring about or slitly less than one lip region widthbehind anterior extremity normally " single but appearing " doble whenthe stylet is protruded (Fig 8 B)

    /

    Basal expded portion of oesohagus occupying 354 % of the neckregion Locations of oesophageal gland nuclei and orices as follows

    D0=63.166.2DN=66.4-7

    DO-DN =3338

    K=79-8

    S0, SN7881S0, SN =818

    SN= 88-9S0= 89-91

    K'= 888

    Cardia conoid with ronded tip, more than 1/3rd of the corresponding bodywidth long, enveoped by intestinal tissue. A very narrow oesophagointestinaldisc is present Prerectm 3-.5 anal body widths long. In some specime thecells of the proximal part of the prerectum bear well developed microvilli.Rectum about 17-1.9 anal body widths long.

    Vva a transverse slit Vagina abot /3rd-1/2 of the corresponding bodywidth Uteri somewhat shorter than ovidcts. Eiptical sperms present in

    distal half of ters and proxima part of oviduct.

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    26 PARC NAIONAL DES VIRUNGA

    Tail elongate-conid, tapering gradally towards a sharp termins, abot

    6-8 abal body widths long; with 2 cadal pores on each side

    Male : Similar to female in general shape and morphology, except dissimilartail Gonads typical Spicles abot 14-18 anal body widths long, withsingle median piece Lateral giding pieces well developed. Spplementsconsist of an adanal pair and 18-21 contigosly arranged ventromedians Sbventral papillae more or less reglarly spaced

    Prerectm 6-8 anal body widths starting anteror to the region of thecopilatory mscles.

    Tail dorsally convex-conoid with rounded termins, abot 0814 anal bodywidths long; with 3 cadal pores on each side

    D i f f e r e n t i a d i a g n o s i s . Mesodorylaimus kamandeanus n spcan be differentiated from most oher M esodorylaimus species by the high nmberof contigos spplements and the long prerectm in the male and by itssmall size. It comes close to M. thermae (OBB in oEPPLI 1926) ooDEin oonEY 1963 bt differs in the nmber of cadal pores on the male tail(at least 6 pairs in M. thermae) and the differently shaped female tail.

    T y p e s p e c i m e n s Holotype female along with one paratype femalemonted on slide Mesodorylaimus kamandeanus SS/57. Paratype females andmales monted on slides Mesodorylaimus kamandeanus SS/58-63

    T y p e o c a i t y a n d h a b i t a t Kamand, Edward Lake; collectedby H AMAS in 1935

    R e m a r k s . This species shares the foowing characters with Mesodory-laimus : " single gdng ring, transverse vlva, small body size, fint bodypores; with Laimydorus it shares the contigosly arranged spplements and thelong prerectm in males. However as explained before some of hese charactershave dobtfl diagnostic vale This also appears fro a comparson with somerecently described spec by LF (1969) Mesodorylaimus cryptosperma ooF1969 possesses a longitdinal vlva, 2125 contigos spplements and a longprerectm in males, hence it fits the diagnosis of Laimydorus and is here transferred to that gens, becomng Laimydorus cryptosperma (ooF 1969) comb nMesodorylaimus litoralis ooF 1969 possesses also a longitdinal vlva and15-18 contigos spplements, bt has a short prerectm n the male and a" single giding ring Mesodorylaimus derni ooF 1969 has a longitdinalvlva and 1120 contigos spplements, bt again a short prerectm in themale and a " single giding ring. Some older species crrently considerednder Mesodorylaimus as eg M alpestris (THORNE 1939) NDRSSY 1959,M clavicaudatus (THORNE & WANGER 1936) NDRSSY 1959, M. tenellus(THORNE & WANGER 1936) NDRSSY, 1959 and M thermae (OBB in oEPPLI

    1926) OODEY in oonEY 1936 do possess contigosly arranged spplements

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    PARC NAIONAL DES VIRUNGA 27

    From the above it is clear tat a number of specie& do not fit well in either

    of both generic diagnoses d that, if both genera are real, they need a betterdefinition It is evident that this problem can only be solved by a careful studyof as many species as possible Therefore we propose to use the oldest name(Mesodorylaimus) for all the doubtful cases until this better definition is provided

    Mesodorlaimus spec

    Syn Dorylaimus parafecundus apud S EKHOVEN & TEUNISSEN, partim

    Ne Dorylaimus parafecundus DE oNNCK, 1935

    =Paradorylaimus parafecundus (DE oNNCK, 1935) NDRSSY 1969

    (Fig. 9.)

    S EKHOVEN & TEUNISSEN (1938) reported Dorylaimus parafecundus fromTshengelero (1 d) Gahinga volcano (11 o d 1 juv), Kanyabayongo (1 c,1 ) and Rutshuru (1 d). Of these specimens only the male from Tshengelero(sample no 533) is left

    In their illustration S EKHOVEN & TEUNISSEN (ec Fig 67) show a greatvariation in the number and arrangement of the ventromedian supplements,so that they were probably with a complex of different species Although their

    figure 67 shows a ventromedian series of 28 supplements it may have beendrawn from the Tshengelerospecimen or at least from a specimen conspecicwith it Finy it should be pointed out that their figure 67 D, though

    referred to as a female tail, shows an adanal supplement and the terminal partsof spiculeprotractors, hence represents a male tailThe male from Tshengelero is in rather bad condtion, especially in the

    region between odontophore and intestineprerectum junction It is neverthelessceary different from P. parafecundus as described by DE oNINCK (1935)

    M e a s u r e m e n t s Refer to Table 3.

    D e s c r i p t i o n - Male : Body curved dorsally posterior to the cardia andventrally at posterior extremity; tapering slightly towards anterior extremityCuticle finely striated transversally Body pores not visible Lateral chordsabout /8th of the body width near middle Lip region offset by a constrction

    Amphids stirrupshapedOdontostyle 17 lip region widths long; aperture less than 1/3rd of the

    odotostyle length Guiding ring idented, about one lip region width fromanterior extremity

    Details of oesophags, nerve ring and cardia not clear Prerectum slightlymore than 4 anal body widths long, starting from opposite the first ventramediansupplement

    Spicules about 2 anal bdy widths long Lateral guiding pieces weldeveloped Supplements consisting of an adanal pair and a series of 27 contiguous ventramedians Subventral papilae irregularly spaced, reaching beyond

    the prerectum

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    28 PARC NATIONAL DES VIRUNGA

    0C< u

    :00

    8

    FIG. 9 - Mesodorylaimus spec

    A : Anterio region; B : Surface view of anteio region; C : Posterior bodyregion of mae; D : Mae tail egion

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    PARC NATIONAL DES VIRUNGA 29

    Tai dorsaly convex-conoid with rounded terminus about 2/3rd of anal body

    width long with 11 caudal pores on each side.

    L o c a 1 i t y a n d h a b i t a t Collected from Tshengelero, a swampnorth of Munagana; Alt 1,75 m.

    R e m a r k s . This species shows characters of Laimydorus (many, contiguous supplements) and Mesodorylaimus (short prerectum, " singe guidingring) Its position is dubtfu aso since the female is unknown It is assignedtentatively to the oldest of both genera.

    GenusLAIMYDORUS IDDIQI,

    1969.Laimydorus parhomalopapillatus (. TEKHOVEN, 1944) n comb

    Syn. Dorylaimus parhomalopapillatus TEKHOVEN, 1944.

    (Figs. 10 and ll.)

    M e a s u r e m e n t s . Refer to Table 4.

    D e s c r i p t i o n . Female : Body ventraly curved posterior to base ofoesophagus, gradualy taperng towards both extremities. Cuticle finey striatedtransversaly, detached here and there from body. Lateral chor about 1/7thof the correspondng body width in the midbody region. Numerous lateral

    body pores along the dorsa and ventral sides of the lateral chords. In additionto these there are two irreguar rows of pores, one ateroventral, the otherlaterodorsal (Fig. 11 E) The lateroventral row consists of 3 pores, of which8 in the oesophageal region and 22 more posterior reaching till the end of theposterior gonad. The aterodorsal row conssts of 2 pores, of which 3 n theoesophageal region and 17 posterior to it, reaching up to the end of the psteriorgonad. The dorsa body pores exceptionay reach till below the vulva.

    Lip region wider than adjoining body, offset by a constriction. Lips anguarin superficia view and bearing the usual number of papilae Amphids deep,stirrup-shaped, slit occupying less than half of the corresponding bodywidthOdontostyle 2.3 lip region widths long aperture slighty ess than haf of odonto

    stye length. Guiding ring 1.4 lip region widths from anterior extremity.Basal expanded portion of oesophagus occupying 57 % of neck length.

    Only the dorsal and nucleus and outlet visibe

    D0=44.2 DN=45.9

    Cardia long conoid, about haf of the corresponding body width, enveopedby ntestinal tissue Prerectum about 7 anal body widths long proximal partwith prominent microvili. Rectum about .3 anal body widths long.

    Vuva appearing longitudinal Vagina about half of the corresponding bodywidth long, surrounded by a prominent sphincter. Poar bodies sometimes

    visible within the egg shell (Fig. 1 H).

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    30 PARC NATIONAL DES VIRUNGA

    -

    ;

    :>

    B

    -.

    ) "-

    .x-

    DE

    '

    '

    H

    FIG 10 - Laimydorus parhomalopapillatus (S STKOVEN, 1944).

    A : Anterior region; B : Surface view of anterior region; C : Female anterior

    gonad; D : esophago-intestina junction cardia; : Surface view posterior

    body region of mae showing suppements subventra papiae and aterabody pores; F : Basal expaded portion of oesophagus oesophageal gand

    nucei and orifices; G : ntire femae; H : uva region

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    PARC NAIONAL DES VRUNGA 3

    Tail elongate, tapering gradually, about 5 anal body widths long (tip broken)wth 5 caudal pores on each sde.

    Male Smlar to female in general shape and morphology, except for thedssmar tal. Lateral chords about 1/5th of body width in the mdbodyLateral body pores numerous, along he dorsal and ventral sde or n the middle(cf. Fg. 1 E of the lateral chord. In the regon of the prerectum the lateroventral series of pores s quite far from the lateral chords; long ducts connectthe pores with these chords.

    Oesophageal gland nucle and outlets well vsble, postons as follows

    DO=43.45.7DN=44.5474

    DODN 15-1.7

    K=68-8

    S0 SN=65-69S02, SN2=73-76

    S2N=84-86S20=85-87

    K'=718

    Male gonads typical Spcules about ana body widths long with a sinemedian piece. Lateral guding pieces well developed. Supplements consstng ofan adanal pair and 59 ventromedans. The ventromedan ones are arrangedn the folowing manner : first a group of 91 cosely spaced, then 78 morewdely spaced and fnaly again 9-1 closely spaced supplements. Subventralpapllae irregularly spaced, reachng below or above the ntestne-prerectum

    junctonPrerectum 8-5 anal body wdths long Tail dorsally convexconoid withbroadly rounded tp about 3/4th of the anal body wdth long; wth -1 caudalpores on each side.

    D i f f e r e n t a d i a g n o s s . - Laimydorus parhomalopapillatus comesclose to the type species L prolificus (THORNE & WANGER, 1936) SQ, 1969but dffers from t n the poston of he vulva (V 47 n L. prolificus) thelength of the odontostyle (about 36 in L prolificus) and the arrangementof the supplements It also resembles L. unipapillatus ADAY, 195) DRSSY,1969, but dffers by he odontostyle length (7 in L unipapillatus) dfferent

    arrangement of supplements and arato (a= 3 in L. unipapillatus).

    T y p e s p e c i m e n s Lectotype female mounted on slde Laimydorusparhomalopapillatus SS/49

    Paralectotypes two males mounted on sldes Laimydorus parhomalopapillatus SS/5-5.

    T y p e o c a t y a n d h a b t a t . From fresh water (N.A.P.), exactlocality not known.

    R e m a r k . S. EKHOVEN (1944) descrbed ths species a Dorylaimus

    parhomalopapillatus, with 444 longtudnal lines. The original descrption s

    3

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    32 PARC NATIONAL DES VIRUNGA

    c

    1

    0

    -

    C

    E

    --

    0

    , B

    100

    F 1 - Laimydous parhomalopapillatus (S STEKOVEN, 1944)

    A : Posterior body region o male; B Cross secion hrough copulaory musces

    region; C : Femae ail; D : Male ai; : Surace view in expanded portion of

    oesophags sowing arragemen of body pores.

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    PARC NATIONAL DES VRUNGA 33

    based on one female and two maes but in the tube with type specimens an

    additional posteror fragment of a male was fond. Snce longtudna linescoud not be detected on any of the specimens in latera vew a cross sectionwas made of the fragment This secton shows that longtudina lines are absent(Fig 7 B) hence the species does not beong in Dorylaimus It is here transferredto Laimydorus with which it agrees in many respects.

    Laimydorus spec.

    Syn Dorylaimus flavomaculatus apud S. TEKHOVEN, 1944

    Ne Dorylaimus flavomaculatus v. NSTOW 1876

    = Laimydoru flavomaculatus (v. NSTOW, 876) IDDIQI, 1969.

    (Fig. 12)

    S TEKHOVEN (944) reported two males and one femae of D. flavomacu-latu from Kimboko The single male present in the collection was in rathergood condition except for the head end.

    M e a s u r e m e n t s Refer to Tabe 4.

    D e s c r i p t i o n . - Male : Body ventray curved in posteror haf gradually taperng towads anterior end Cutcle finey striated transversaly Lateral chords about /5th of the body width near midde Body pores not visble.Lip region offset by slght constriction; about /3rd a wide as neck base Exactshape of amphids not known

    Odontostye probably 13 lip region wdths ong (junction with odontoporenot clear !) ; aperture probably half of the odontostye length. Two darkcooured elongate bodies present in the odntophore region (Fg 12 B) Guidingring about one ip region width from anterior extremity appearing " doube ,

    /

    but the stylet is protrded.Basal expanded portion of oesophags occupying 4 % of the neck region.

    Locations of oesophagea gand nucei and orifices as folows :

    D0=625

    DN=639DODN= 14

    K91

    S0 SN=81

    S0 SN=83SN=9S0=92

    K'=924

    Carda cardiform about 1/3rd of the corresponding body width Prerectum

    about 5 ana bdy wdths longGonads typical Spicules about 2.3 anal body wdths long Latera giding

    pieces wel developed Supplements consisting of an adana pair and a contigousseries of 13 ventromedians. Subventral papilae irregularly spaced

    Ta conod dorsally covex ventray concave; with rounded terminus and

    with 4 caudal pores on each side.

    3*

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