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-
chus species occurring in southern Africa. She
described four new species namely: M. pretoriensis
(= Coomansus pretoriensis) from Pretoria, Trans-
vaal, M. parvulus (= Coomansus parvulus) , now re-
garded as a synonym of Coomansus parvus (de Man,
1880), from Fouriesburg, Orange Free State, M. jug-
alis (= Clarkus jugalis) , now regarded as a synonym
of Clarkus sheri from Wellington, Cape Province
and M. aquaticus from Gobabeb, Namibia. Further
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)
spicules and accessory pieces extremely long and
slender, tail elongate with rounded terminus and
caudal glands, ducts and spinneret well developed
(Jairajpuri, 1970).
Mononchus truncatus Bastian, 1865 and Monon-
chus aquaticus Coetzee, 1968 are two closely related
species.
Mononchus truncatus was first described by Bas-
tian (1865) from Falmouth, England. Clark (1960)

species reported by Coetzee were M. truncatus and
M. papillatus (= Clarkus papillatus) from various
localities in South Africa.
Since the publication by Coetzee (1968), further
Mononchidae specimens from South Africa,
Botswana, Swaziland and Namibia accumulated in
the nematode collections of the Rand Afrikaans
University and the National Collection of Nema-
todes, Biosystematics Division, Plant Protection
Research Institute. All of this material, as well as the
types and other specimens used by Coetzee, were
available for study.
MATERIALS AND METHODS
Specimens were killed by gentle heat, fixed in
FAA and processed into glycerine according to
Thorne's slow method and mounted on permanent
slides. Measurements and drawings were done with
the aid of a Zeiss Standard 18 research microscope,
equipped with a drawing tube. The body and all
curved structures were measured along the median
line. Buccal cavity measurements do not include the
lip, vestibule leading into the cavity, the anterior
oblique plates, or the cavity walls. Cavity width was
taken at the broadest part. Measurements of the
dorsal tooth apex and the subventral ribs were taken
from the anterior vertical plates and calculated as a
percentage of the buccal cavity length.
TAXONOMY
Genus Mononchus Bastian, 1865
Mononchus (senso stricto) with type species
Mononchus truncatus, has poorly developed lips and
Received 24 April 1991; accepted for publication 28 October 1991
61
gave a redescription of this species, designating a
neotype from specimens collected at the type local-
ity. He synonymized several species with M. trunca-
tus and also mentioned that Andrassy (1958) was the
first to draw attention to the variability of this spe-
cies.
Mononchus aquaticus was described by Coetzee
(1968) from the Kuiseb River bed in Gobabeb,
Namibia. Specimens were also rerorted from the
Cape Province, Natal and Transvaa in South Africa,
mostly found in water or in the vicinity of water.
According to Coetzee, M. aquaticus is very closely
related to M. truncatus Bastian, 1865 except for a
smaller stoma, more posteriorly situated amphids
and a stouter tail. In M. aquaticus the indentation in
the anterior subventral wall of the stoma is normally
situated anterior to, or at the same level as the dorsal
tooth apex and the refractive transverse thickening is
usually posterior to this level.
Jairajpuri (1970) identified ten female specimens
from India as M. aquaticus. He states that if the
variation of M. truncatus as given by Andrassy
(1958) and Mulvey (1967) is accepted, there are no
characters left which distinguish M. aquaticus from
M. truncatus, and it could be regarded as a synonym
of M. truncatus.
Baqri & Jairajpuri (1972) gave a very comprehen"
sive description, as well as numerous illustrations of
M. aquaticus. They had at their disposal three
female paratypes as well as all the Indian specimens
of!M. aquaticus, the neotype of M. truncatus, as well
as l,qme additional specimens of M. truncatus from
MJllvey. After comparing the various populations of
M. ituncatus and M. aquaticus, they confirmed the
validity of M. aquaticus as a separate species.
 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
A, Band D. Windhoek specimens
A. Posterior reproductive branch
Andrassy (19M5) described a new species Mono-
chus bel/us from Puerto Rico, Argentina and Hung-
ary. According to him M. bellus is closely related to
both M. truncatus and M. aquaticus, differing in the
~osition of the dorsal tooth apex (M. bel/us = 30-33
Yo; M. truncatus = 22-28 % and M. aquaticus =
19-23 %) and in the level of the subventral
transverse ribs in the buccal cavity (situated before
the dorsal tooth aI?ex in M. bel/us). M. aquaticus can
also be easily distmguished from the other two spe-
cies by the much smaller buccal cavity. Loof (1990)
regarded M. bel/us Andrassy, 1985 as a possible syn-
onym of M. truncatus, with which we tend to agree.
The position of the dorsal tooth apex in M. truncatus
is quite variable viz. 19-33 % (Table 2) (19 % =
calculated from Fig. 2B of Andrassy, 1985) and the
measurements of M. bel/us given by Andrassy
(1985), falls within this range. The position of the
subventral ribs is also more or less the same in both
M. truncatus and M. bel/us (Table 2). It is the varia-
tion in the position of the dorsal tooth apex which
gives rise to the ribs being either anterior to, or pos-
terior to the tooth apex. Thus, in specimens with the
dorsal tooth apex more posterior, the ribs will be
anterior to the tooth apex (as in M. bel/us). In speci-
mens with a more anterior tooth apex, the ribs will
be posterior to the apex. However, we refrain from
making a formal synonymization, since we have not
seen types or other specimens of M. bel/us.
62
B. Head region
C. Heat-relaxed specimens
D. Female tail
Mononchus aquaticus Coetzee, 1968 (Fig. 1)
For list of synononyms, see Andrassy (1985) and for
measurements see Table 1
M. aquaticus seems to be a cosmopolitan species
and has subsequent to the reports by Coetzee
1968j' Jairajpuri (1970) and Baqri & Jairajpuri
!
1972 also been recorded by Zullini from Italy
1971 and Mexico (1973); Grootaert (1976) from
the North-western Sahara; Jairajpuri & Khan (1982)
from India; Winiszewska (1985) from Poland and
Chaves (1990) from Argentina. Grootaert & Maer-
tens (1976) established a population in a laboratory
in Belgium, and Small & Grootaert (1977) described
males from this cultivated population.
Twenty-six female specimens from Windhoek,
Namibia and Leslie, South Africa were identified as
M. aquaticus. These are in full agreement with pre-
vious descriptions of this species.
Description
Female: Cuticle finely striated, mesocuticle some-
times irregularly separated from rest of cuticle. Late-
ral field 36 (28-47) % of corresponding body width;
no lateral, dorsal or ventral pores seen. Lip region
continuous with adjoining body; papillae distinct.
Amphid aperture small, anterior to dorsal tooth
apex 10,1 (7-13) 11m from anterior end. Fovea
cupshaped in specimens from Windhoek, mostly

TABLE 1 Morphometric data of two populations of Mononchus aquaticus Coetzee, 1968
L(mm)
a 37,7 (31,4-41,1)
b 4,6 (4,3-5,0)
c 9,2 (7,7-10,6)
c'
V%
Lip region width (Ilm)
Buccal cavity length (Ilm)
width (Ilm)
Indentation of anterior subventral vertical plate %
Nerve ring from anterior end (Ilm)
Extretory pore from anterior end (Ilm)
Lateral fIeld %
Tail length (Ilffi)
Rectum length (Ilm)
indistinct in specimens from Leslie; amphidial canal
inconspicuous. Buccal cavity narrow; walls about 2
11m thick, anteriorly finely striated. Indentation of
anterior subventral vertical plate posterior to dorsal
tooth. Dorsal tooth al?ex at 19,9 (16-22) % of buccal
cavity length. One pair of geusids sometimes visible,
opening subventrally into buccal cavity. Oesophagus
strongly muscular; oesophageal gland outlets distinct
in most specimens, gland nuclei however mostly
inconspicuous, except in a few specimens from
Windhoek. One pair of endolids seen in the lumen
of one specimen. Nerve rin~ distinct; excretory pore
just posterior to nerve ring m specimens from Wind-
hoek, inconspicuous in specimens from Leslie.
Vulva transverse with moderately sclerotized lips.
No advulval papillae present. Vagina with sclero-
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)
tized pieces. Vulval glands as reported by Baqri &
Jairajpuri (1972) not observed. Sphincter muscle
present between uterus and oviduct, but difficult to
discern. Ovaries well developed with numerous
oocytes. No eggs or sperm present in reproductive
branches. Tail elongate-conoid in anterior half,
curved ventrally in posterior half ending in a
rounded terminus. Three caudal glands mostly con-
spicuous in specimens from Windhoek; arranged in
tandem and ending in an ampulla through a common
duct. Spinneret terminal, seemingly armed. One
pair of lateral caudal papillae seen. Intestine four or
six cells in circumference, filled with numerous
yellow granules. Rectal glands sometimes evident.
Male: Not found.
Material examined
The present specimens are from a water-furrow
next to the road en route to Leslie in the Transvaal
and among the roots of grass at the Daan Viljoen
Nature Reserve, Windhoek in Namibia, collected by
I. Botha, 1 February, 1972 and by A. Coomans and
J. Heyns, 22 July, 1986 respectively.
Transvaal specimens on slides 10181, 10185,
10186, 10189-10192, 10194 in the National Collec-
tion of Nematodes, Biosystematic~ Division, Plant
Protection Research Institute, Pre\. :a; Namibian
specimens on slides RAU 2974-297 I 1,979, 2981,
2983-2985 in the collection of the Rano Afrikaans
University, Johannesburg.
Mononchus truncatus Bastian, 1865 (Fig. 2)
For list of synononyms, see Andrassy (1985)
Coetzee (1968) identified specimens from various
localities in South Africa as M. truncatus. She gave
no morphometrical data, but mentioned that these
63
SANDRA DE BRUIN & J. HEYNS
Leslie Windhoek
1,46 (1,31-1,80) 1,90 (1,70-2,20)
37,3 (34,7-40,7)
4,8 (4,3-5,3)
9,5 (8,9-10,3)
5,6 (4,3-6,7) 5,9 (5,0-6,3)
50,3 (49-53) 50,4 (48-54)
21,9 (20-24) 24 (22-26)
31,1 (30-32) 32,3 (29-36)
11,5 (11-13) 11,5 (11-13)
29,6 (28-31) 28,8 (25-34)
104,6 (98-109) (n=9) 127 (115-137)
140 (127-150)(n=lO)
37 (32-47) 34,8 (28-42)
157,8 (140-180) 201 (165-240)
24,5 (18-29) 28,4 (26-30) (n= 12)
specimens differ from the redescription by Clark
(1960) only in the more anteriorly situated amphids.
This discrepancy is resolved by the findings of Coo-
mans & Khan (1981), who restudied the neotype and
found that the amp hid is in a far more anterior posi-
tion than shown by Clark, viz. just behind the lateral
lip.
Three female and four juvenile specimens from
Botswana were identified as M. truncatus. Also
available for study were a few females from Bar-
berton and Hoedspruit in the Transvaal, previously
identified by Coetzee (1968) as this species, and
Parys in the Orange Free State. All these specimens
agree well with the redescription by Clark (1960),
except that lateral fields are distinct in the present
specimens. In Table 2 the measurements of the pre-
sent specimens are compared with those of Clark,
Coomans & Khan and Mononchus bel/us of
Andrassy and Chaves.
Description
Female: Body almost straight when heat-relaxed,
only tail tip slightly ventrally curved. Cuticle with
fine transverse striations; mesocuticle sometimes
irregularly separated from rest of cuticle. Lateral
field about one fourth to one third of corresponding
body width. Hypodermal pores not seen, except in
one specimen where ventral pores are present just
antenor to anus. Lip region slightly offset; 26
(23-28) !lm wide. Amphids 9 (8-10) 11m from ante-
rior end in specimens from Botswana and Hoed-
spruit; indistinct in other specimens. Fovea
cupshaped; amphidial canal sometimes visible. Buc-
cal cavity elongate-oval; walls about 3 !lm thick, pos-
terior vertical plates finely striated. Dorsal tooth
apex at about one third of buccal cavity length.
Indentation of anterior subventral vertical plates
anterior to dorsal tooth apex. One pair of geusids
opening subventrally into buccal cavity. Oesopha-
geal gland nuclei and outlets mostly distinct. Nerve
ring located 148 (143-155) 11m from anterior end in
specimens from Botswana; obscure in specimens
from Barberton, Hoedspruit and Parys. Excretory
pore not seen.
Vulva a transverse slit with moderately developed
sclerotized pieces. Vagina with sclerotized l?ieces.
Uterus short; no eggs in the Botswana specimens;
one egg present in either one uterus or both uteri in
most of the specimens from Barberton and Hoed-
spruit. Sphincter between uterus and oviduct quite
distinct. Ovary well developed with numerous 00-
cytes. Female tail conoid at first, tapering gradually
 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
terminus. Three caudal glands present, arranged in
tandem. Ampulla distinct; spinneret terminal. Two
sublateral papillae present just anterior to tail termi-
nus. Coelomocytes sometimes seen in tail region.
Intestine six or eight cells in circumference. Rectum
31 (29-33) 11m long.
Males: Not found.
Juveniles: General morphology similar to adults.
Material examined
The present specimens are from wet soil among
the roots of a sausage tree (Kigelia africana) stand-
ing on the edge of the water on Boba Island, Oka-
vango, Botswana, collected by J. Heyns and A. Coo-
mans, 1 August 1989; from the bank of the Queens
River en route to Barberton, Transvaal, collected by
J. Heyns, 3 June, 1963; from soil among the roots of
lemon trees near the "Blyde Trading Store"
between Hoedspruit and the Strydom tunnel,
Hoedspruit, Transvaal, collected by J. Heyns, G.
Lagerwey and C. van Zyl, September, 1963 and
Parys, Orange Free State, collected by J. Heyns,
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
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)
FIG. 4 Clark us papillatus (Bastian, 1865) Jairajpuri, 1970
A, C, D, F, and G. Franschhoek specimens
A. Head and neck region
B. Head (dorso-ventral view) and neck region (Gra-
bouw specimen)
Botswana specimens on slides RAU 5271-5274 in
the nematode collection of the Rand Afrikaans Uni-
versity, Johannesburg; other specimens on slides
1763, 2184 and 2971 in the National Collection of
Nematodes, Biosystematics Division, Plant Protec-
tion Research Institute, Pretoria.
Genera Clarkus Jairajpuri, 1970 and Coomansus
Jairajpuri & Khan, 1977
Jairajpuri (1970) proposed a new genus Clarkus
68
LI~l~m~m~____________~! E
LI.!.>10"'O"'AJ"'m'--____________.J1 C, F, G
150A,lm A,B,D
C. Head of another specimen showing slight variation
D. Reproductive system
E. Heat-relaxed specimens
F. Slightly aberrant tail
G. Tail
for those species of Mononchus (sensu lato) with lips
and labial papillae prominent, buccal cavity barrel-
shaped; with or without a non-denticulate ventral
ridge, spicules and accessory pieces short and stout,
tails conoid, ventrally arcuate, caudal glands poorly
developed or absent and spinneret absent or
inconspicuous.
Jairajpuri & Kahn (1977) subdivided the genus
Clarkus into Clarkus (sensu stricto), represented by
the type species, C. papillatus, having a non-denticu-
 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
ridge.
CJarkus sheri (Mulvey, 1967) Jairajpuri, 1970 (Fig.
3)
Syn: Clark us jugalis (Coetzee, 1968) Jairajpuri, 1970
This species was first described by Mulvey (1967)
from Riverside, California, U.S.A. Coetzee (1968)
described a new species, Mononchus jugalis (now
Clarkus jugalis) from several localities in South
Africa, with the type locality Wellington in the Cape
Province. In the diagnosis, she stated that Clarkus
jugalis is close to Mononchus papillatus (now Clar-
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)
Ventral ridge distinct. One pair of geusids 0rening
subventrally into buccal cavity. Oesophagea gland
nuclei and outlets mostly evident. Nerve ring distinct
in specimens from Piggs Peak, Ventersdorp, Blyde-
poort and Pretoria; obscure in other specimens. Ex-
cretory pore and duct distinct, just posterior to nerve
ring.
Female: Vulva a transverse slit with slightly thick-
ened lips. Vagina with well developed sclerotized
pieces and surrounding musculature. Reproductive
branches well developed in specimens from Piggs
Peak, Ventersdorp, Blydepoort and Pretoria, less
well developed in specimens from Pinetown and

Newcastle, difficult to discern in specimens from

kus papillatus). Jairajpuri (1970) and Andrassy
(1983) however regard Clarkus jugalis as a synonym
of Clark us sheri, with which we are inclined to agree.
Zullini (1977) recorded one female specimen of
Clarkus sheri from Fortin de Flores, Central Mexico.
In this specimen the buccal cavity is slightly wider,
the amphids situated slightly more posterior and the
tail less arcuate. This is in agreement with the
description of C. jugalis by Coetzee (1968).
Two males and fourteen females from Piggs Peak,
Ventersdorp, Pretoria, Blydepoort, Newcastle,
Pinetown and Fauresmith were available for study.
These specimens agree well with previous descrip-
tions C. sheri, except for a slightly wider lip region,
longer spicules and slightly longer gubernaculum
and lateral guiding pieces. In Table 3 the mea-
surements of the present specimens are compared
with those of Mulvey, Zullini and J airajpuri & Khan
and Clarkus jugalis Coetzee, 1968.
Description
Female and male: Cuticle with fine transverse
striations, mesocuticle sometimes irregularly sepa-
rated from rest of cuticle. Lateral field about one
quarter of corresponding body width. Hypodermal
pores indistinct except lateral pores sometImes seen
Just anterior to tail region in female; in male, ventral
pores are visible in the region just posterior to the
oesophagus and again towards the beginning of the
supplements. Lip region truncate, well developed
and slightly offset. Position of amp hid apertures va-
rying from base of lip region to about half-way be-
tween beginning of anterior vertical plates and dor-
sal tooth apex. Fovea cupshaped, amphidial canal
inconspicuous. Buccal cavity oval-shaped, walls
about 2-3 11m thick; striations sometimes visible.
69
Fauresmith. One egg measuring 81 x 50 11m present
in posterior uterus of one Piggs Peak specimen.
Sphincter present between uterus and oviduct,
although not always very distinct. Pars dilatata ovi-
ductus well developed. Ovary with numerous
oocytes. Tail conoid, ventrally arcuate. Caudal
glands and spinneret absent. Caudal papillae com-
prising one to three lateral pairs. Hyaline core 7,6
(5-9) 11m long. Intestine six to fourteen cells in cir-
cumference. Rectum 33,8 (23-43) 11m long.
Male: Diorchic, testes opposed and outstretched.
Anterior testis 254-340 11m; posterior testis 231-270
11m. Spermatozoa ovoid, about 4 11m in diameter.
Three ejaculatory glands observed. Supplements
papilloid, non-contiguous. Spicules arcuate. Termini
of lateral guiding pieces not clear, but seemingly not
bifurcated as mentioned in original description.
Gubernaculum crescent-shaped. Rectal glands pre-
sent, ducts however indistinct. Tail shape similar to
that of female. Caudal glands and spinneret absent.
Caudal papillae comprising three lateral pairs. Hya-
line core 7-9 11m long. Intestine eight to ten cells in
circumference. Rectum 33,2 (23-40) 11m long.
Material examined
The present specimens are from soil among grass
next to the road en route to Piggs Peak, two kilo-
metres after the Oshoek turnoff, Swaziland, col-
lected by J. Heyns and M. Luc, November, 1984;
from soil on the farm "Syferfontein", Ventersdorp;
from soil underneath the dam wall on the plot of Mr
M. G. Louw, Pretoria and from soil among the roots
of indigenous plants behind chalet 41 at the H. F.
Odendaal camp, Blydepoort, Transvaal, collected
by E. van den Berg, 1 March, 1980 and 5 February,
1981 and by J. Heyns and A. Coomans July, 1984
 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
from soil among the roots of Tagetes minuta in the
"Wyebank" area, Pinetown, Natal, collected by
C. H. Hendriks, 12 October, 1978 and by K. P. N.
Kleynhans and D. P. Keetch, 16 March, 1981
respectively; from soil among the roots of natural
vegetation at the Philippolis turnoff in Fauresmith,
Orange Free State, collected by E. van den Berg, 6
70
Specimens from Piggs Peak and Blydepoort on
slides RAU 1583, 1589,4993 in the nematode collec-
tion of the Rand Afrikaans University, Johannes-
burg; other specimens on slides 15358,15361,17390,
18305,18310,18312,18535,19823,19907,19908 and
19910 in the National Collection of Nematodes,
Biosystematics Division, Plant Protection Research
Institute, Pretoria.
 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
(Fig. 4)
For a detailed discussion of the taxonomic uncer-
tainties and nomenclatorial problems concerning
Bastian's species and the species identified and
described as Mononehus papillatus by Butschli in
1873, together with Dujardin's Oneholaimus museo-
rum (Prionehulus museorum), see Andrassy (1958)
and Clark (1960). Since we do not believe that it is
possible to determine with any certainty the identity
(and conspecifity or not of any of these species), we
accept Clark's (1960) redescriftion and designation
of a neotype (from the type 0 locality) for Monon-
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)
ity oval-shaped; walls about 1-2 Ilm thick, without
striations. Ventral ridge distinct. One pair of geusids
opening into buccal cavity, visible in some speci-
mens. Oesophagus muscular, more strongly so in
posterior third. All oesophageal ~land outlets as well
as dorsal gland nucleus visible m most specimens;
other gland nuclei however inconspicuous. One pair
of endolids seen in oesophageal lumen in two speci-
mens. In one specimen, two outlets and in another
three outlets can be seen anterior to the dorsal gland
outlet; one outlet associated with a small glandular
organ (Fig. 4 A & B). Nerve ring difficult to discern
in specimens from Grabouw, faintly visible in speci-
mens from Mont-aux-Sources and Franschhoek.

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

71
 MONONCHIDA OF SOUTHERN AFRICA: GENERA MONONCHUS, CLARKUS AND COOMANSUS
Division, Plant Protection Research Institute,
Pretoria.
Coomansus parvus (de Man, 1880) Jairajpuri &
Khan, 1977 (Fig. 5)
Syn. Coomansus parvulus (Coetzee, 1968) Jairajpuri
& Khan, 1977
Monochus parvus (now Coomansus parvus), was
first described by de Man (1880) from the Nether-
lands.
Coetzee (1968) described a new species
Mononchus parvulus (now Coomansus parvulus)
from Fouriesburg in the Orange Free State. Accord-
ing to her this species appears to be closely related to
C. parvus, but differs in the more strongly developed
dorsal tooth, which is also situated further forward
in the buccal cavity. According to Jairajpuri & Khan
(1982), C. parvus shows considerable variation in
body measurements, shape of buccal cavity and tail,
amphid position, nature and position of dorsal tooth
and in the thickness of the buccal cavity walls. We
are therefore in agreement with the synonomy of C.
parvulus with C. parvus, as first su~gested by Jairaj-
puri (1970) and endorsed by Jalrajpuri & Khan
(1982).
Five females from Transvaal and the Cape, identi-
fied as C. parvus, correspond well with previous de-
scriptions of this species. Measurements of the
present specimens are rresented and compared with
the type specimens 0 C. parvulus (c. parvus) of
Coetzee.
Description
Female: Cuticle with fine transverse striations,
mesocuticle sometimes irregularly separated from
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012)
rest of cuticle. Lateral field about one quarter to one
third of corresponding body width. Lateral hypoder-
mal pores viSible in neck region of one specimen
only. No dorsal or ventral pores seen. Lip region
slightly offset; papillae distinct. Amphid apertures
and fovea indistlOct, except in one specimen where it
is situated almost at the beginning of the buccal
cavity 8 !lm from anterior end. Buccal cavity oval-
shaped; walls 1-2 !lm thick, without striations. One
pair of geusids opening subventrally into buccal
cavity. Oesophagus more strongly muscular in
posterior third. Oesophageal gland nuclei indistinct,
except dorsal gland nucleus visible in one specimen
and nucleus of second pair distinct in another. Out-
let of dorsal gland nucleus seen in one specimen;
outlets of first and second pair of gland nuclei
discern able in all specimens. Nerve ring difficult to
see, 105 Jlm from anterior end in one specimen.
Extretory pore not seen.
Vulva a transverse slit with thickened lips. Vagina
with sclerotized pieces. Ovejector present. Sphincter
between uterus and oviduct observed in one speci-
men. One egg (proximally indented) present in each
uterus of one specimen, one egg measuring 31 x 99
Jlm in anterior uterus of another. Tail conoid, quite
strongly ventrally arcuate, with a rounded terminus.
No caudal glands or spinneret. Caudal parillae,
comprising one subdorsal and one subventra Eair,
seen only in one specimen. Hyaline core 4,4 (3-5)
Jlm long. Intestine four to six cells in circumference.
Rectum 21 (18-24) Jlm long.
Male: Not found.
Material examined
The present specimens are from soil among the
roots of shrubs, Menlo Park, Pretoria, in the
72
Transvaal, collected by J. Heyns, 23 September,
1971 and from soil at the "Herrie" monument in
Meiringspoort, Oudtshoorn in the Cape, collected
by K. P. N. Kleynhans, 23 October, 1980.
On slides RAU 275 (Pretoria specimens) and
18160 (Oudtshoorn specimens) in the nematode
collection of the Rand Afrikaans University and the
National Collection of Nematodes, Biosystematics
Division, Plant Protection Institute, Pretoria,
respectively.
KEY TO THE MONONCHUS, CLARKUS AND COOMAN-
SUS SPECIES OF SOUTHERN AFRICA
1. Lips well developed; caudal glands and spin-
neret absent........... ............ .... ... ... . ........ 2
Lips poorly developed; caudal glands and
splOneret present... .... ... .... ... .... .... .......... 5
2. Buccal cavity with a non-denticulate ventral
ridge ................................................... 3
Buccal cavity without a ventral ridge.......... 4
3. Buccal cavity bigger (36-43 x 14-20 !lm);
tail longer (95-143 Jlm) .......................... .
......... .......... ........ Clarkus sheri (c. jugalis)
Buccal cavity smaller (24-27 x 9-14 !lm);
tail shorter (60-83 !lm) ...... Clarkus papillatus
4. Body longer (1,6-1,8 mm); buccal cavity
larger (33 x 15,2 Jlm); tail longer (118 Jlm) ..
............................. Coomansus pretoriensis
Body shorter (0,8-1,14 mm); buccal cavity
smaller (23,6-26 x 8,3-12,5 Jlm); tail shorter
(59,7-93 Jlm) ....................................... .
....... .......... Coomansus parvus (c. parvulus)
5. Buccal cavity larger (42-49 x 14-20 Jlm); tail
more slender............... Mononchus truncatus
Buccal cavity smaller (29-36 x 11-13 Jlm);
tail stout ....... ........ .... Mononchus aquaticus
ACKNOWLEDGEMENTS
The authors thank Dr. Esther van den Berg for
the loan of specimens from the National Collection
of Nematodes, Biosystematics Division, Plant Pro-
tection Research Institute, Pretoria. Financial sup-
port by the Foundation for Research Development
of the CSIR is gratefully acknowledged.
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73