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Phytophylactica 24: 61-73 (1992)

MONONCHIDA (NEMATODA) OF SOUTHERN AFRICA: GENERA MONONCHUS


BASTIAN, 1865, CLARKUSJAIRAJPURI, 1970 AND COOMANSUSJAIRAjPURI
& KHAN, 1977
SANDRA DE BRUIN and J. HEYNS. Department of Zoology, Rand Afrikaans University, P.O. Box 524,
Johannesburg, 2000, South Africa

ABSTRACT
Key words: Clarkus, Coomansus, key, Mononchus, southern Africa, taxonomy
Descriptions and illustrations are given of Mononchus aquaticus Coetzee, 1968; Mononchus trunca-
tus Bastian, 1865; Clarkus sheri (Mulvey, 1967) Jairajpuri, 1970; Clarkus papillatus (Bastian, 1865)
Jairajpuri, 1970 and Coomansus parvus (de Man, 1880) Jairajpuri & Khan, 1977. Mononchus bel/us
Andrassy, 1985 is considered a synonym of Mononchus truncatus. An identification key is given for the
southern African species.

Uittreksel
MONONCHIDA (NEMATODA) VAN SUlDELIKE AFRIKA: GENERA MONONCHUS BAS-
TIAN, 1865, CLARKUSJAlRAJPURl, 1970 EN COOMANSUS JAlRAJPURl & KHAN, 1977
Beskrywings en lyntekeninge van Mononchus aquaticus Coetzee, 1968, Mononchus truncatus Bas-
tian, 1865, Clarkus sheri (Mulvey, 1967) Jairajpuri, 1970, Clarkus papillatus (Bastian, 1865) Jairajpuri,
1970 en Coomansusj'arvus (de Man, 1880) Jairajpuri & Khan, 1977 word gegee. Mononchus bellus
Andrassy, 1985 wor beskou as 'n sinoniem van Mononchus truncatus. 'n Indentifikasiesleutel vir die
spesies van suidelike Afrika word gegee.

INTRODUCTION labial papillae, buccal cavity elongate-cylindroid,


Coetzee (1968) was the first to report on Monon- spicules and accessory pieces extremely long and
chus species occurring in southern Africa. She slender, tail elongate with rounded terminus and
described four new species namely: M. pretoriensis caudal glands, ducts and spinneret well developed
(= Coomansus pretoriensis) from Pretoria, Trans- (Jairajpuri, 1970).
vaal, M. parvulus (= Coomansus parvulus) , now re- Mononchus truncatus Bastian, 1865 and Monon-
garded as a synonym of Coomansus parvus (de Man, chus aquaticus Coetzee, 1968 are two closely related
1880), from Fouriesburg, Orange Free State, M. jug- species.
alis (= Clarkus jugalis) , now regarded as a synonym
of Clarkus sheri from Wellington, Cape Province Mononchus truncatus was first described by Bas-
and M. aquaticus from Gobabeb, Namibia. Further tian (1865) from Falmouth, England. Clark (1960)
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species reported by Coetzee were M. truncatus and gave a redescription of this species, designating a
M. papillatus (= Clarkus papillatus) from various neotype from specimens collected at the type local-
localities in South Africa. ity. He synonymized several species with M. trunca-
Since the publication by Coetzee (1968), further tus and also mentioned that Andrassy (1958) was the
Mononchidae specimens from South Africa, first to draw attention to the variability of this spe-
Botswana, Swaziland and Namibia accumulated in cies.
the nematode collections of the Rand Afrikaans Mononchus aquaticus was described by Coetzee
University and the National Collection of Nema- (1968) from the Kuiseb River bed in Gobabeb,
todes, Biosystematics Division, Plant Protection Namibia. Specimens were also rerorted from the
Research Institute. All of this material, as well as the Cape Province, Natal and Transvaa in South Africa,
types and other specimens used by Coetzee, were mostly found in water or in the vicinity of water.
available for study. According to Coetzee, M. aquaticus is very closely
MATERIALS AND METHODS
related to M. truncatus Bastian, 1865 except for a
smaller stoma, more posteriorly situated amphids
Specimens were killed by gentle heat, fixed in and a stouter tail. In M. aquaticus the indentation in
FAA and processed into glycerine according to the anterior subventral wall of the stoma is normally
Thorne's slow method and mounted on permanent situated anterior to, or at the same level as the dorsal
slides. Measurements and drawings were done with tooth apex and the refractive transverse thickening is
the aid of a Zeiss Standard 18 research microscope, usually posterior to this level.
equipped with a drawing tube. The body and all
curved structures were measured along the median Jairajpuri (1970) identified ten female specimens
line. Buccal cavity measurements do not include the from India as M. aquaticus. He states that if the
lip, vestibule leading into the cavity, the anterior variation of M. truncatus as given by Andrassy
oblique plates, or the cavity walls. Cavity width was (1958) and Mulvey (1967) is accepted, there are no
taken at the broadest part. Measurements of the characters left which distinguish M. aquaticus from
dorsal tooth apex and the subventral ribs were taken M. truncatus, and it could be regarded as a synonym
from the anterior vertical plates and calculated as a of M. truncatus.
percentage of the buccal cavity length.
Baqri & Jairajpuri (1972) gave a very comprehen"
sive description, as well as numerous illustrations of
TAXONOMY M. aquaticus. They had at their disposal three
Genus Mononchus Bastian, 1865 female paratypes as well as all the Indian specimens
Mononchus (senso stricto) with type species of!M. aquaticus, the neotype of M. truncatus, as well
Mononchus truncatus, has poorly developed lips and as l,qme additional specimens of M. truncatus from
MJllvey. After comparing the various populations of
M. ituncatus and M. aquaticus, they confirmed the
Received 24 April 1991; accepted for publication 28 October 1991 validity of M. aquaticus as a separate species.
61
MONONCHIDA OF SOUTHERN AFRICA: GENERA MONONCHUS, CLARKUS AND COOMANSUS

A
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)

FIG. 1 Mononchus aquaticus Coetzee, 1968 B. Head region


A, Band D. Windhoek specimens C. Heat-relaxed specimens
A. Posterior reproductive branch D. Female tail

Andrassy (19M5) described a new species Mono- Mononchus aquaticus Coetzee, 1968 (Fig. 1)
chus bel/us from Puerto Rico, Argentina and Hung- For list of synononyms, see Andrassy (1985) and for
ary. According to him M. bellus is closely related to measurements see Table 1
both M. truncatus and M. aquaticus, differing in the
~osition of the dorsal tooth apex (M. bel/us = 30-33 M. aquaticus seems to be a cosmopolitan species
Yo; M. truncatus = 22-28 % and M. aquaticus = and has subsequent to the reports by Coetzee
1968j' Jairajpuri (1970) and Baqri & Jairajpuri

!
19-23 %) and in the level of the subventral
transverse ribs in the buccal cavity (situated before 1972 also been recorded by Zullini from Italy
the dorsal tooth aI?ex in M. bel/us). M. aquaticus can 1971 and Mexico (1973); Grootaert (1976) from
also be easily distmguished from the other two spe- the North-western Sahara; Jairajpuri & Khan (1982)
cies by the much smaller buccal cavity. Loof (1990) from India; Winiszewska (1985) from Poland and
regarded M. bel/us Andrassy, 1985 as a possible syn- Chaves (1990) from Argentina. Grootaert & Maer-
onym of M. truncatus, with which we tend to agree. tens (1976) established a population in a laboratory
The position of the dorsal tooth apex in M. truncatus in Belgium, and Small & Grootaert (1977) described
is quite variable viz. 19-33 % (Table 2) (19 % = males from this cultivated population.
calculated from Fig. 2B of Andrassy, 1985) and the Twenty-six female specimens from Windhoek,
measurements of M. bel/us given by Andrassy Namibia and Leslie, South Africa were identified as
(1985), falls within this range. The position of the M. aquaticus. These are in full agreement with pre-
subventral ribs is also more or less the same in both vious descriptions of this species.
M. truncatus and M. bel/us (Table 2). It is the varia-
tion in the position of the dorsal tooth apex which Description
gives rise to the ribs being either anterior to, or pos- Female: Cuticle finely striated, mesocuticle some-
terior to the tooth apex. Thus, in specimens with the times irregularly separated from rest of cuticle. Late-
dorsal tooth apex more posterior, the ribs will be ral field 36 (28-47) % of corresponding body width;
anterior to the tooth apex (as in M. bel/us). In speci- no lateral, dorsal or ventral pores seen. Lip region
mens with a more anterior tooth apex, the ribs will continuous with adjoining body; papillae distinct.
be posterior to the apex. However, we refrain from Amphid aperture small, anterior to dorsal tooth
making a formal synonymization, since we have not apex 10,1 (7-13) 11m from anterior end. Fovea
seen types or other specimens of M. bel/us. cupshaped in specimens from Windhoek, mostly
62
SANDRA DE BRUIN & J. HEYNS

TABLE 1 Morphometric data of two populations of Mononchus aquaticus Coetzee, 1968


Leslie Windhoek

L(mm) 1,46 (1,31-1,80) 1,90 (1,70-2,20)


a 37,7 (31,4-41,1) 37,3 (34,7-40,7)
b 4,6 (4,3-5,0) 4,8 (4,3-5,3)
c 9,2 (7,7-10,6) 9,5 (8,9-10,3)
c' 5,6 (4,3-6,7) 5,9 (5,0-6,3)
V% 50,3 (49-53) 50,4 (48-54)
Lip region width (Ilm) 21,9 (20-24) 24 (22-26)
Buccal cavity length (Ilm) 31,1 (30-32) 32,3 (29-36)
width (Ilm) 11,5 (11-13) 11,5 (11-13)
Indentation of anterior subventral vertical plate % 29,6 (28-31) 28,8 (25-34)
Nerve ring from anterior end (Ilm) 104,6 (98-109) (n=9) 127 (115-137)
Extretory pore from anterior end (Ilm) 140 (127-150)(n=lO)
Lateral fIeld % 37 (32-47) 34,8 (28-42)
Tail length (Ilffi) 157,8 (140-180) 201 (165-240)
Rectum length (Ilm) 24,5 (18-29) 28,4 (26-30) (n= 12)

indistinct in specimens from Leslie; amphidial canal specimens differ from the redescription by Clark
inconspicuous. Buccal cavity narrow; walls about 2 (1960) only in the more anteriorly situated amphids.
11m thick, anteriorly finely striated. Indentation of This discrepancy is resolved by the findings of Coo-
anterior subventral vertical plate posterior to dorsal mans & Khan (1981), who restudied the neotype and
tooth. Dorsal tooth al?ex at 19,9 (16-22) % of buccal found that the amp hid is in a far more anterior posi-
cavity length. One pair of geusids sometimes visible, tion than shown by Clark, viz. just behind the lateral
opening subventrally into buccal cavity. Oesophagus lip.
strongly muscular; oesophageal gland outlets distinct Three female and four juvenile specimens from
in most specimens, gland nuclei however mostly Botswana were identified as M. truncatus. Also
inconspicuous, except in a few specimens from available for study were a few females from Bar-
Windhoek. One pair of endolids seen in the lumen berton and Hoedspruit in the Transvaal, previously
of one specimen. Nerve rin~ distinct; excretory pore identified by Coetzee (1968) as this species, and
just posterior to nerve ring m specimens from Wind- Parys in the Orange Free State. All these specimens
hoek, inconspicuous in specimens from Leslie. agree well with the redescription by Clark (1960),
Vulva transverse with moderately sclerotized lips. except that lateral fields are distinct in the present
No advulval papillae present. Vagina with sclero- specimens. In Table 2 the measurements of the pre-
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)

tized pieces. Vulval glands as reported by Baqri & sent specimens are compared with those of Clark,
Jairajpuri (1972) not observed. Sphincter muscle Coomans & Khan and Mononchus bel/us of
present between uterus and oviduct, but difficult to Andrassy and Chaves.
discern. Ovaries well developed with numerous
oocytes. No eggs or sperm present in reproductive Description
branches. Tail elongate-conoid in anterior half,
curved ventrally in posterior half ending in a Female: Body almost straight when heat-relaxed,
rounded terminus. Three caudal glands mostly con- only tail tip slightly ventrally curved. Cuticle with
spicuous in specimens from Windhoek; arranged in fine transverse striations; mesocuticle sometimes
tandem and ending in an ampulla through a common irregularly separated from rest of cuticle. Lateral
duct. Spinneret terminal, seemingly armed. One field about one fourth to one third of corresponding
pair of lateral caudal papillae seen. Intestine four or body width. Hypodermal pores not seen, except in
six cells in circumference, filled with numerous one specimen where ventral pores are present just
yellow granules. Rectal glands sometimes evident. antenor to anus. Lip region slightly offset; 26
(23-28) !lm wide. Amphids 9 (8-10) 11m from ante-
Male: Not found. rior end in specimens from Botswana and Hoed-
Material examined spruit; indistinct in other specimens. Fovea
cupshaped; amphidial canal sometimes visible. Buc-
The present specimens are from a water-furrow cal cavity elongate-oval; walls about 3 !lm thick, pos-
next to the road en route to Leslie in the Transvaal terior vertical plates finely striated. Dorsal tooth
and among the roots of grass at the Daan Viljoen apex at about one third of buccal cavity length.
Nature Reserve, Windhoek in Namibia, collected by Indentation of anterior subventral vertical plates
I. Botha, 1 February, 1972 and by A. Coomans and anterior to dorsal tooth apex. One pair of geusids
J. Heyns, 22 July, 1986 respectively. opening subventrally into buccal cavity. Oesopha-
Transvaal specimens on slides 10181, 10185, geal gland nuclei and outlets mostly distinct. Nerve
10186, 10189-10192, 10194 in the National Collec- ring located 148 (143-155) 11m from anterior end in
tion of Nematodes, Biosystematic~ Division, Plant specimens from Botswana; obscure in specimens
Protection Research Institute, Pre\. :a; Namibian from Barberton, Hoedspruit and Parys. Excretory
specimens on slides RAU 2974-297 I 1,979, 2981, pore not seen.
2983-2985 in the collection of the Rano Afrikaans Vulva a transverse slit with moderately developed
University, Johannesburg. sclerotized pieces. Vagina with sclerotized l?ieces.
Mononchus truncatus Bastian, 1865 (Fig. 2) Uterus short; no eggs in the Botswana specimens;
one egg present in either one uterus or both uteri in
For list of synononyms, see Andrassy (1985) most of the specimens from Barberton and Hoed-
Coetzee (1968) identified specimens from various spruit. Sphincter between uterus and oviduct quite
localities in South Africa as M. truncatus. She gave distinct. Ovary well developed with numerous 00-
no morphometrical data, but mentioned that these cytes. Female tail conoid at first, tapering gradually
63
MONON CHIDA OF SOUTH ERN AFRICA : GENER A MONON CHUS,
CLARKU S AND COOMA NSUS

A o
c

F
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)

FIG. 2 Mononchus truncatus Bastian, 1865 C. Tail terminus


A-C and E. Botswana specimens D. Head region (Hoedsp ruit specimen)
A. Head and neck region E. Female tail
B. Reproductive system F. Heat-relaxed specimens

64
TABLE 2 Morphometric data of several populations of Mononchus truncatus Bastian, 1865 and Mononchus bel/us Andrassy, 1985
Mononchus truncatus Mononchus bel/us
According to Clark According to Coomans According to Andrassy According to
(1960) & Khan (1981) Present specimens (1985) Chaves (1990)

Falmouth Chippen- Falmouth Mount Kenya Baberton &


Parys Botswana P~~
Rico A rgentma
. H ungary A '
rgentma
ham Fen Hoedspruit

Neotype 9<j><j> 2 <j> <j> Neotype 8<j><j> 3<j><j> 3<j><j> 3<j><j> 4 jj ? <j> <j>' ? <j> <j>' ? <j> <j>' 5<j><j>

L(mm) 1,92 1,86 (1,65-2,14) 1,67-1,85 2,16(1,81-2,39) 2,10(2,05-2,15) 1,83(1,76-1,94) 1,91 (1,66-2,10) 1,41 (1,25-1,65) 1,58--1,611,54-1,621,60--1,79 2,05(1,9-2,2)
a 28 26--40" 33-36 29,7 (25,2-35,6) 37,6 (37,2-38) 35,3 (32,4-40,1) 33,5 (31,3-36,1) 34,4 (32,4-37,5) 29-34 30-31 28--30 33,4 (30,9-35,9)
b 3,9 3,4-4,3' 3,6-3,8 3,7 (3,3-4,1) 3,9 (3,6-4,1) 4,0 (4,0-4,1) 3,85 (3,6-4,0) 3,7 (3,6-3,8) 3,5-4,0 3,9-4,0 4,0-4,2 4
c 8 6,4-8,6' 5,8-6,4 6,3 (5,~,8) 8,9 (8,2-9,5) 7,7 (7,l-S,l) 8,4 (8,l-S,6) 8,0 (7,H,5) 6,1-6,8 6,8--7,0 6,8--8,4 8,7 (7,5-10,8)
~ c' [6,9]b 8,0 (6,8--9,0) 8,0 (7,0-9,5) 7,6 (7,3-s,O) 6,8 (6,6-7,0) 6,7 (6,3-s,2) 7,8--8,2 6,7 6-8 6,3 (5,3-7,3)
V% 55,5 48,4-55,5' 53,4-54,3 52 (49,5-54) 54-55 (n=2) 53,7 (53-55) 53 (52-54) 54-56 51-53 53-54 54,5 (53-56)
Lip region width (11m) [27,6] [29,3] [34,5] 26-27 (n=2) 24,7 (23-26) 26 (23-28) [29] 26,6 (26-28)
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)

Buccal cavity:

fi~l f~l fi~l


length (11m) [54,5] 46,3 (44-48) 43 (42-45) 47 (44-49) (35-36)' 4O-47d 46,2 (44-49)
width (11m) [I 8] 16,7 (15-20) 14,3 (14-15) 14,7 (14-15) 11' 18--21d 21,2 (19-23)

f~~l 25,7 ~24-27~ 23,3 (22-26~


f~~l
Subventral ribs % [27J 23-25 (n=2) 25,6 (25-26f [271
Dorsal tooth apex % 28 29,2 (2 -33) 27,3 (26-29) 25,3 21-29 27,7 (27-28 26,6 (25-29 ' 30-33d 29,2 (26-32)
Lateral field % 25 (n=2) 24-33 (n=2) 28--31 (n=2)
Tail length (11m) [243] 346 (280-391) 220-251 (n=2) 239 (234-250) 228 (204-250) 220-258d 235,6 (199-258)

• -No average given


• [ ]-Calculated from the original illustrations
Vl
, -n = 3
d -Probably composite figures for all localities
~
t:I
::0
:>
t:I
tTl
ttl
::0
c:::
Z
fI:o
~

~
~
Vl
MONONCHIDA OF SOUTHERN AFRICA: GENERA MONONCHUS, CLARKUS AND COOMANSUS

A B
u.
<i C
c
0.
ID
Cl

D!:\
a
0 ° O(J
nE
E
a
l2
E
:I
0
on
E
:I

~C C
(J

G
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)

FIG. 3 Clark us sheri (Mulvey, 1967)Jairajpuri, 1970 D. Sperm cells


B, D, F-G. Ventersdorp specimens E. Heat-relaxed specimens
A. Head and neck region (Pinetown specimen) F. Male posterior region
B. Head G. Female tail
C. Anterior reproductive branch (Newcastle specimen)

to become more cylindrical, ending in a rounded the roots of a sausage tree (Kigelia africana) stand-
terminus. Three caudal glands present, arranged in ing on the edge of the water on Boba Island, Oka-
tandem. Ampulla distinct; spinneret terminal. Two vango, Botswana, collected by J. Heyns and A. Coo-
sublateral papillae present just anterior to tail termi- mans, 1 August 1989; from the bank of the Queens
nus. Coelomocytes sometimes seen in tail region. River en route to Barberton, Transvaal, collected by
Intestine six or eight cells in circumference. Rectum J. Heyns, 3 June, 1963; from soil among the roots of
31 (29-33) 11m long. lemon trees near the "Blyde Trading Store"
Males: Not found. between Hoedspruit and the Strydom tunnel,
Hoedspruit, Transvaal, collected by J. Heyns, G.
Juveniles: General morphology similar to adults. Lagerwey and C. van Zyl, September, 1963 and
Material examined Parys, Orange Free State, collected by J. Heyns,
The present specimens are from wet soil among September, 1963.

66
TABLE 3 Morphometric data of Clarkus sheri (Mulvey, 1967) and Clarkusjugalis (Coetzee, 1968)

Clarkus sheri Clarkus jugalis


According Accordin~ to
Present specimens According to Mulvey (1967) to Zullini Jairajpun & According to Coetzee
(1977) Khan (1982) (1968)

Ventersdorp, Pretoria, Newcastle & Several


Piggs Peak Fauresmith California Mexico India Wellington
B1ydepoort Pinetown localities

2 <? <? 5 <? <? 200 3 <? <? 4 <? <? Holotype Aliotype P~~a<y~es 1<? ? <? <? Holotype
ParalYR.es
12 '+' '+' 19 <? <?

L(mm) 2,25-2,49 2,1 (1,78-2,31) 2,13-2,24 1,81 (1,52-1,96) 1,95 (1,68-2,20) 1,8 1,8 1,8 (1,6--2,1) 2,10 1,9-22 1,7 1,7 (1,5-1,9) 1,6--2,3
a 34,0-36,1 27,2 (20,9--33,9) 31,8-32,5 28,6 (24,1-35,0) 24,5 (20,9--25,6) 27 33 28 (24-30) 29 23--27 24 27 (21-29) 22-30
b 4,9-5,1 4,4 (3,7--4,8) 4,4--4,8 4,1 (4,0--4,3) 4,4 (4,2--4,5) 4,2 4,3 4,1 (3,9--4,3) 4,0 3,9--4,6 4 4 4,0--4,4
c 17,4-17,7 17,4 (15,5-18,6) 20,4-22,4 16,3 (14,9-18,1) 18,4 (17,1-19,1) 17 22 18 ~16--21) 21 14-17 16 16 (14-19) 13--19
c' 2,7-3,4 2,6 (2,4-3,2) 2,0-2,2 2,76 (2,7-2,8) 2,3 (2,0-2,5) [2,4]" 3,2] [3,0]
V% 57-61 61 (60-63)' 62,7 (61-64) 62,5 (61-65) 59 60 59-63) 63 58-61 61 62 57-64
~ Lip region: width (~m) 33--37 35,6 (30--40) 39--40 36,7 (33--40) 38,5 (38--40) [38] 32-35 [341
height (~m) 10-11 12,8 (12-13) 10-11 13,7 (13-14) 10 (n=3) yo,8l
12,1 [12,7] 10-15 [11,2]
length (~m) 38 (37--40)
[f~l
Buccal cavity: 40 40,6 (38--43) 40--43 39,3 (36--43) 35 33 2-38' 41 34-38
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)

width (~m) 16 16,9 (15-20) 15 15 (14-16) 19,3 (16--21) 20 20 19-23' 16--18


Dorsal tooth apex % 23 25,6 (23--29) 25-26 25 (23-27) 25 (22-27) 25-28' [27] 27-30 28 25-30'
Amphid from anterior cnd (~m) 11 (n=l) 13,8 (12-16) 10-11 15 (13-16) 10,8 (10-13) [12,5] 12-16 [9,6]
Nerve ring from antcrior cnd
(~m) 145 (n=l) 150-153 (n=2) 130-136
Lateral field % 26 (n=l) 25,2 (24-27) 26--27 29,7 (28-32) 24,3 (23--26)
(n=3)
Tail length (~m) 127-143 120,8 (110-134) 95-110 111,7 (96--131) 106 (98-115) 105 [90] [105,8] 110 100--110
Spiculum Icngth (~m) 85-94 80
Lateral guiding piecc Icngth
(~m) 20-22 19,31
Gubernaculum length (~m) 24 [ 18,2 C/l
Supplements 14 10 ;l>
Z
tj
" - No average given :0
;l>
" -Calculated from thc original illustrations tj
, -n=4 l'T1
tc
~
Z
R>
......
:I:
l'T1
~
C/l
MONONCHIDA OF SOUTHERN AFRICA: GENERA MONONCHUS, CLARKUS AND COOMANSUS

LI~l~m~m~____________~! E
LI.!.>10"'O"'AJ"'m'--____________.J1 C, F, G
150A,lm A,B,D
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)

FIG. 4 Clark us papillatus (Bastian, 1865) Jairajpuri, 1970 C. Head of another specimen showing slight variation
A, C, D, F, and G. Franschhoek specimens D. Reproductive system
A. Head and neck region E. Heat-relaxed specimens
B. Head (dorso-ventral view) and neck region (Gra- F. Slightly aberrant tail
bouw specimen) G. Tail

Botswana specimens on slides RAU 5271-5274 in for those species of Mononchus (sensu lato) with lips
the nematode collection of the Rand Afrikaans Uni- and labial papillae prominent, buccal cavity barrel-
versity, Johannesburg; other specimens on slides shaped; with or without a non-denticulate ventral
1763, 2184 and 2971 in the National Collection of ridge, spicules and accessory pieces short and stout,
Nematodes, Biosystematics Division, Plant Protec- tails conoid, ventrally arcuate, caudal glands poorly
tion Research Institute, Pretoria. developed or absent and spinneret absent or
inconspicuous.
Genera Clarkus Jairajpuri, 1970 and Coomansus Jairajpuri & Kahn (1977) subdivided the genus
Jairajpuri & Khan, 1977 Clarkus into Clarkus (sensu stricto), represented by
Jairajpuri (1970) proposed a new genus Clarkus the type species, C. papillatus, having a non-denticu-

68
SANDRA DE BRUIN & J. HEYNS

TABLE 4 Morphometric data of Clarkus papillatus (Bastian, 1865)


According to Clark (1960) Present specimens
Broadmoor Mont-aux-Sourccs Franschhoek Grabouw

Neotype 9Cjl Cjl 1Cjl 7Cjl Cjl 4Cjl Cjl

L(mm) 1,29 1,23 (1,01-1,30) 1,23 1,06 (0,96-1,22~ 0,96 (0,90-1 ,09~
a 29 22-29' 26,7 24,9 (22,3-27,4 25,1 (23,3-27,1
b 3,9 3,6-3,9' 3,7 3,6 (3,4-3,8) 3,6 (3,4-3,8)
c 19 14-19 14,6 14,5 (12,8-17,7) 15,1(13,8-16,0)
c' [2,5]" 2,9 3,0 (2,6-3,6) 2,9 (2,8-3,1)
V% 62,2 6O,O-Q7,7' 59 62 (61-{)3) 62,8 ~62-{)4)
Lip region width (/lm) [26,3] 25 24 (22-25) 24,3 22-26)
height (/lm)

W~t
9 8,1 (8-9) 7,8 (7-8)
Buccal cavity length (/lm) 26 24,7 (24-27) 24
width (/lm) 10 10,6 (9-14) 9,8 (9-10)
Dorsal tooth apex % 15-21' 15 17 (16-19) 17
Lateral field % 24 24,1 (22-30) 23 (21-26)
Tail length (J.lID) [93,3] 84 73,9 (69-83) 63,8 (60-68)

, -No average given


b [ ]-Ca\Culated from the original illustrations

late ridge and Coomansus which is without such a Ventral ridge distinct. One pair of geusids 0rening
ridge. subventrally into buccal cavity. Oesophagea gland
nuclei and outlets mostly evident. Nerve ring distinct
CJarkus sheri (Mulvey, 1967) Jairajpuri, 1970 (Fig. in specimens from Piggs Peak, Ventersdorp, Blyde-
3) poort and Pretoria; obscure in other specimens. Ex-
cretory pore and duct distinct, just posterior to nerve
Syn: Clark us jugalis (Coetzee, 1968) Jairajpuri, 1970 ring.
This species was first described by Mulvey (1967) Female: Vulva a transverse slit with slightly thick-
from Riverside, California, U.S.A. Coetzee (1968) ened lips. Vagina with well developed sclerotized
described a new species, Mononchus jugalis (now pieces and surrounding musculature. Reproductive
Clarkus jugalis) from several localities in South branches well developed in specimens from Piggs
Africa, with the type locality Wellington in the Cape Peak, Ventersdorp, Blydepoort and Pretoria, less
Province. In the diagnosis, she stated that Clarkus well developed in specimens from Pinetown and
jugalis is close to Mononchus papillatus (now Clar-
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)

Newcastle, difficult to discern in specimens from


kus papillatus). Jairajpuri (1970) and Andrassy Fauresmith. One egg measuring 81 x 50 11m present
(1983) however regard Clarkus jugalis as a synonym in posterior uterus of one Piggs Peak specimen.
of Clark us sheri, with which we are inclined to agree. Sphincter present between uterus and oviduct,
Zullini (1977) recorded one female specimen of although not always very distinct. Pars dilatata ovi-
Clarkus sheri from Fortin de Flores, Central Mexico. ductus well developed. Ovary with numerous
In this specimen the buccal cavity is slightly wider, oocytes. Tail conoid, ventrally arcuate. Caudal
the amphids situated slightly more posterior and the glands and spinneret absent. Caudal papillae com-
tail less arcuate. This is in agreement with the prising one to three lateral pairs. Hyaline core 7,6
description of C. jugalis by Coetzee (1968). (5-9) 11m long. Intestine six to fourteen cells in cir-
Two males and fourteen females from Piggs Peak, cumference. Rectum 33,8 (23-43) 11m long.
Ventersdorp, Pretoria, Blydepoort, Newcastle, Male: Diorchic, testes opposed and outstretched.
Pinetown and Fauresmith were available for study. Anterior testis 254-340 11m; posterior testis 231-270
These specimens agree well with previous descrip- 11m. Spermatozoa ovoid, about 4 11m in diameter.
tions C. sheri, except for a slightly wider lip region, Three ejaculatory glands observed. Supplements
longer spicules and slightly longer gubernaculum papilloid, non-contiguous. Spicules arcuate. Termini
and lateral guiding pieces. In Table 3 the mea- of lateral guiding pieces not clear, but seemingly not
surements of the present specimens are compared bifurcated as mentioned in original description.
with those of Mulvey, Zullini and J airajpuri & Khan Gubernaculum crescent-shaped. Rectal glands pre-
and Clarkus jugalis Coetzee, 1968. sent, ducts however indistinct. Tail shape similar to
that of female. Caudal glands and spinneret absent.
Description Caudal papillae comprising three lateral pairs. Hya-
Female and male: Cuticle with fine transverse line core 7-9 11m long. Intestine eight to ten cells in
striations, mesocuticle sometimes irregularly sepa- circumference. Rectum 33,2 (23-40) 11m long.
rated from rest of cuticle. Lateral field about one
quarter of corresponding body width. Hypodermal Material examined
pores indistinct except lateral pores sometImes seen The present specimens are from soil among grass
Just anterior to tail region in female; in male, ventral next to the road en route to Piggs Peak, two kilo-
pores are visible in the region just posterior to the metres after the Oshoek turnoff, Swaziland, col-
oesophagus and again towards the beginning of the lected by J. Heyns and M. Luc, November, 1984;
supplements. Lip region truncate, well developed from soil on the farm "Syferfontein", Ventersdorp;
and slightly offset. Position of amp hid apertures va- from soil underneath the dam wall on the plot of Mr
rying from base of lip region to about half-way be- M. G. Louw, Pretoria and from soil among the roots
tween beginning of anterior vertical plates and dor- of indigenous plants behind chalet 41 at the H. F.
sal tooth apex. Fovea cupshaped, amphidial canal Odendaal camp, Blydepoort, Transvaal, collected
inconspicuous. Buccal cavity oval-shaped, walls by E. van den Berg, 1 March, 1980 and 5 February,
about 2-3 11m thick; striations sometimes visible. 1981 and by J. Heyns and A. Coomans July, 1984
69
MONONCHIDA OF SOUTHERN AFRICA: GENERA MONONCHUS, CLARKUS AND COOMANSUS

c
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)

~ll~m~m~ ____________~1 E
5~O=u~m~________~1
Wi A,B
ISOAJm C,D

FIG. 5 Coomansus parvus (de Man, 1880) Jairajpuri & Khan, B. Reproductive system
1977 C. Tail
A-D. Oudtshoorn specimens D. Head
A. Head and neck region E. Heat-relaxed specimens

respectively; from soil among the roots of natural February, 1980.


vegetation on the farm "Eikenhof", Newcastle and Specimens from Piggs Peak and Blydepoort on
from soil among the roots of Tagetes minuta in the slides RAU 1583, 1589,4993 in the nematode collec-
"Wyebank" area, Pinetown, Natal, collected by tion of the Rand Afrikaans University, Johannes-
C. H. Hendriks, 12 October, 1978 and by K. P. N. burg; other specimens on slides 15358,15361,17390,
Kleynhans and D. P. Keetch, 16 March, 1981 18305,18310,18312,18535,19823,19907,19908 and
respectively; from soil among the roots of natural 19910 in the National Collection of Nematodes,
vegetation at the Philippolis turnoff in Fauresmith, Biosystematics Division, Plant Protection Research
Orange Free State, collected by E. van den Berg, 6 Institute, Pretoria.
70
SANDRA DE BRUIN & J. HEYNS

TABLE 5 Morphometric data of South African populations of Coomansus parvus (de Man, 1880)
Type specimens of C. parvulus (c. parvus) according to
Present specimens Coetzee (1968)
Pretoria Oudtshoom Fouriesburg
Holotype Paratype
L(mm) 0,90-0,95 1,05 (0,95-1,14) 0,9 0,8 0,86 (0,8-0,9)
a 22-25 26,6 (24,4-28,3) 22 22 21 (19-23)
b 3,3--3,4 4,0 (3,9-4,0) 4 4 4
c 11,5-12,3 12,1 (11,7-12,3) 13 14 12
c' 3,3--3,4 3,2 (3,1-3,4) [2,8]' [2,4-27]"
V% 64-67 61 (60--62) 63 62 62,3 (62-63~

~}}
Lip region width (J.lID) 22 22,7 (22-23) [20,6-22,2]

~
Buccal cavity
height (J.1m)
length (J.lID)
8
25
9 (7-11)
25,3 (24-26)
8'3-9'7f
23,6-25 "
width (J.1m) 10--11 9,7 (9-10) [12,5] 8,3--9,7
Dorsal tooth apex % 32-36 30,3 (27-35) 30 37 [24-29]
Lateral field % 29-30 33 (26-38)
Tail length (J.lID) 77-78 87 (78-93) [62,5] [59,7-65,2]"

• [ ]-Calculated from original illustrations


b n =2

CJarkus papiJIatus (Bastian, 1865) Jairajpuri, 1970 ity oval-shaped; walls about 1-2 Ilm thick, without
(Fig. 4) striations. Ventral ridge distinct. One pair of geusids
opening into buccal cavity, visible in some speci-
For a detailed discussion of the taxonomic uncer- mens. Oesophagus muscular, more strongly so in
tainties and nomenclatorial problems concerning posterior third. All oesophageal ~land outlets as well
Bastian's species and the species identified and as dorsal gland nucleus visible m most specimens;
described as Mononehus papillatus by Butschli in other gland nuclei however inconspicuous. One pair
1873, together with Dujardin's Oneholaimus museo- of endolids seen in oesophageal lumen in two speci-
rum (Prionehulus museorum), see Andrassy (1958) mens. In one specimen, two outlets and in another
and Clark (1960). Since we do not believe that it is three outlets can be seen anterior to the dorsal gland
possible to determine with any certainty the identity outlet; one outlet associated with a small glandular
(and conspecifity or not of any of these species), we organ (Fig. 4 A & B). Nerve ring difficult to discern
accept Clark's (1960) redescriftion and designation in specimens from Grabouw, faintly visible in speci-
of a neotype (from the type 0 locality) for Monon- mens from Mont-aux-Sources and Franschhoek.
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)

ehus papillatus Bastian 1865 (= Clarkus papillatus). Excretory pore seen just posterior to nerve ring in
For these and other reasons we do not agree with the specimen from Mont-aux-Sources.
actions taken by Zell (1985). Vulva a transverse slit with thickened lips. Vagina
In view of the past confusion concerning Bastian's muscular, strongly sclerotized pieces present in spec-
M. papillatus, Butschli's M. papillatus and Dujar- imens from Mont-aux-Sources and Franschhoek,
din's Oneholaimus museorum, we regard most of the less strongly sclerotized in specimens from Gra-
older distribution records for these species (dating bouw. Ovejector present. No sphincter muscle
from before about 1958) as uncertain. observed at uterus-oviduct junction. No sperm pre-
Coetzee (1968) identified specimens from the sent in reproductive branches. One egg present in
Cape and Transvaal as C. papillatus, but gave no each uterus in a specimen from Franschhoek, mea-
description or morphometrical data. Illustrations of suring 90 x 39 Ilm and 88 x 42 Ilm. Another speci-
the female head and tail correspond well with those men with one egg in the posterior uterus, measuring
of Clark (1960). 90 x 32 Ilm. Tail elongate-conoid, ventrally arcuate
Since Coetzee's recording of C. papillatus from with a rounded terminus. Caudal glands not seen;
spinneret absent. Caudal papillae difficult to dis-
South Africa, females from Mont-aux-Sources, cern. Hyaline core 4,7 (3-9) Ilm long. Intestine four
Franschhoek and Grabouw were identified as this to six cells in circumference. Rectum 22,6 (19-29)
species. These specimens agree well with previous Ilm long.
descriptions, except for a shorter tail and the pre-
sence of a lateral fIeld. Measurements of the {lresent Males: Not found.
specimens are presented and compared wIth the Material examined
neotype and other specimens of Clark in Table 4.
The present specimens are from Cape Province:
Description from soil in a vineyard on the farm "1' Arc
Female: Cuticle with distinct but fine striations: d'Orleans", Franschhoek, collected by J. Heyns, 30
mesocuticle mostly irregularly separated from rest of March, 1966; from soil in an apple orchard on the
cuticle in Franschhoek specimens. Lateral field farm "Weltevreden", Grabouw, collected by J. M.
about one quarter the corresponding body width; Giliomee, July and October, 1968. Natal: from soil
lateral pores visible in neck region of some speci- among the roots of grass under pine trees at the
mens, but no dorsal or ventral pores seen. Lip region camping site in Rugged Glen, Mont-aux-Sources,
slightly offset; papillae distinct. Amphid aperture collected by J. Heyns and M. Hutsebaut, 9
just anterior to dorsal tooth apex 9,7 (8-12) Ilm from September, 1988.
anterior end in specimens from Franschhoek and Natal specimens on slide RAU 4381 in the nema-
Mont-aux-Sources; indistinct in specimens from tode collection of the Rand Afrikaans University,
Grabauw, except in one, where it is situated just Johannesburg; specimens from Cape Province on
posterior to the dorsal tooth apex. Fovea cup- slides 7310, 7311, 7684, 8070, 8790, 8701, 8788 in the
shaped; amphidial canal inconspicuous. Buccal cav- National Collection of Nematodes, Biosystematics

71
MONONCHIDA OF SOUTHERN AFRICA: GENERA MONONCHUS, CLARKUS AND COOMANSUS

Division, Plant Protection Research Institute, Transvaal, collected by J. Heyns, 23 September,


Pretoria. 1971 and from soil at the "Herrie" monument in
Meiringspoort, Oudtshoorn in the Cape, collected
Coomansus parvus (de Man, 1880) Jairajpuri & by K. P. N. Kleynhans, 23 October, 1980.
Khan, 1977 (Fig. 5) On slides RAU 275 (Pretoria specimens) and
Syn. Coomansus parvulus (Coetzee, 1968) Jairajpuri 18160 (Oudtshoorn specimens) in the nematode
& Khan, 1977 collection of the Rand Afrikaans University and the
National Collection of Nematodes, Biosystematics
Monochus parvus (now Coomansus parvus), was Division, Plant Protection Institute, Pretoria,
first described by de Man (1880) from the Nether- respectively.
lands.
Coetzee (1968) described a new species KEY TO THE MONONCHUS, CLARKUS AND COOMAN-
Mononchus parvulus (now Coomansus parvulus) SUS SPECIES OF SOUTHERN AFRICA
from Fouriesburg in the Orange Free State. Accord- 1. Lips well developed; caudal glands and spin-
ing to her this species appears to be closely related to neret absent........... ............ .... ... ... . ........ 2
C. parvus, but differs in the more strongly developed Lips poorly developed; caudal glands and
dorsal tooth, which is also situated further forward splOneret present... .... ... .... ... .... .... .......... 5
in the buccal cavity. According to Jairajpuri & Khan
(1982), C. parvus shows considerable variation in 2. Buccal cavity with a non-denticulate ventral
body measurements, shape of buccal cavity and tail, ridge ................................................... 3
amphid position, nature and position of dorsal tooth Buccal cavity without a ventral ridge.......... 4
and in the thickness of the buccal cavity walls. We 3. Buccal cavity bigger (36-43 x 14-20 !lm);
are therefore in agreement with the synonomy of C. tail longer (95-143 Jlm) .......................... .
parvulus with C. parvus, as first su~gested by Jairaj- ......... .......... ........ Clarkus sheri (c. jugalis)
puri (1970) and endorsed by Jalrajpuri & Khan
(1982). Buccal cavity smaller (24-27 x 9-14 !lm);
tail shorter (60-83 !lm) ...... Clarkus papillatus
Five females from Transvaal and the Cape, identi-
fied as C. parvus, correspond well with previous de- 4. Body longer (1,6-1,8 mm); buccal cavity
scriptions of this species. Measurements of the larger (33 x 15,2 Jlm); tail longer (118 Jlm) ..
present specimens are rresented and compared with ............................. Coomansus pretoriensis
the type specimens 0 C. parvulus (c. parvus) of Body shorter (0,8-1,14 mm); buccal cavity
Coetzee. smaller (23,6-26 x 8,3-12,5 Jlm); tail shorter
(59,7-93 Jlm) ....................................... .
Description ....... .......... Coomansus parvus (c. parvulus)
Female: Cuticle with fine transverse striations, 5. Buccal cavity larger (42-49 x 14-20 Jlm); tail
mesocuticle sometimes irregularly separated from
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)

more slender............... Mononchus truncatus


rest of cuticle. Lateral field about one quarter to one
third of corresponding body width. Lateral hypoder- Buccal cavity smaller (29-36 x 11-13 Jlm);
mal pores viSible in neck region of one specimen tail stout ....... ........ .... Mononchus aquaticus
only. No dorsal or ventral pores seen. Lip region
slightly offset; papillae distinct. Amphid apertures ACKNOWLEDGEMENTS
and fovea indistlOct, except in one specimen where it The authors thank Dr. Esther van den Berg for
is situated almost at the beginning of the buccal the loan of specimens from the National Collection
cavity 8 !lm from anterior end. Buccal cavity oval- of Nematodes, Biosystematics Division, Plant Pro-
shaped; walls 1-2 !lm thick, without striations. One tection Research Institute, Pretoria. Financial sup-
pair of geusids opening subventrally into buccal port by the Foundation for Research Development
cavity. Oesophagus more strongly muscular in of the CSIR is gratefully acknowledged.
posterior third. Oesophageal gland nuclei indistinct,
except dorsal gland nucleus visible in one specimen REFERENCES
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chidae Chitwood, 1937; Nematoda). Annales Historico-Natu-
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outlets of first and second pair of gland nuclei ANDRAssY, I. 1983. The free-living nematode fauna of the Hor-
discern able in all specimens. Nerve ring difficult to tobagy National Park. Akademiai Kiado, Budapest: 31-46.
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Extretory pore not seen. Prionchulus (Cobb, 1916) Wu & Hoeppli, 1929 (Nematoda:
Vulva a transverse slit with thickened lips. Vagina Mononchidae). Opuscula Zoologica, Budapest 21: 9-21.
with sclerotized pieces. Ovejector present. Sphincter BASTIAN, H. C. 1865. Monograph of the Anguillulidae, or free
between uterus and oviduct observed in one speci- nematoids, marine, land, and freshwater; with descriptions of
10? new species. Transactions of the Linnean Society oj London
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uterus of one specimen, one egg measuring 31 x 99 BAQRI, S. Z. & JAIRAJPURI, M. S. 1972. Morphology of Monon-
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Material examined COOMANS, A. & KHAN, S. H. 1981. Mononchida from Mount
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72
SANDRA DE BRUIN & J. HEYNS

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Dodonaea 45: 162-170.
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genera Mononchus, Clarkus n. gen. and Prionchulus (family WINISZEWSKA, G. 1985. Six species of Mononchoidea (Nema-
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Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)

73