J. Nematode Morphol. Syst.

, 10 (1): 1-10 (2007)

Nematodes of the order Rhabditida from India. Description of two

species of Mononchoides (Nematoda: Diplogastrina)

M. MAHAMOOD, I. AHMAD* AND A.A. SHAH

Section of Nematology, Department of Zoology, Aligarh Muslim University,
Aligarh – 202 002, India.
e-mail: ahmadirfi@yahoo.co.in

Summary.- A new and a known species of the genus Mononchoides Rahm, 1928 are described and illustrated. M. megaonchus
sp. n., collected from decaying banana rhizome, is characterized by a large dorsal tooth, long stoma, and absence of tooth,
teeth or denticles on the subventral walls in general. Males have stout spicules and a gubernaculum with distal sleeve
and nine pairs of genital papillae arranged in typical diplogastrid pattern: three precloacal and six postcloacal. Another
species which shares several common characters with M. striatus (Bütschli, 1876) Goodey, 1963 is also being reported from
India.

Key words: India, Mononchoides, morphology, new species, occurrence, taxonomy.

Resumen.- En esta contribución se describen e ilustran una nueva especie y otra ya conocida del género Mononchoides
Rahm, 1928. M. megaonchus sp. n., recogida en rizoma podrido de platanera, se distingue por tener un diente dorsal
grande, estoma largo, y ausencia de diente, dientes o dentículos en la pared subventral en general. Los machos poseen
espículas robustas y gubernáculo con manguito distal y nueve pares de papilas genitales situadas según el típico patrón
diplogástrido: tres precloacales y seis postcloacales. Otra especie que comparte varios caracteres con M. striatus (Bütschli,
1876) Goodey, 1963 se cita también de India.

Palabras clave: India, Mononchoides, morfología, nueva especie, registro, taxonomía.

INTRODUCTION Shikhobalova, Sobolev, Paramonov & Sudarikov, 1954

Rahm (1928) proposed the genus Mononchoides
with M. longicauda as its type species. Allgén (1947)
proposed the genus Glauxinema with G. filicaudatum as
its type species on the basis of the absence of claw-like
tooth on the right subventral wall and pyramidal tooth
on the left subventral wall. Andrássy (1984) proposed
the subfamily Glauxinematinae to accommodate
Glauxinema and Pareudiplogaster Paramonov, 1952.
Sudhaus and Fürst von Lieven (2003), in a detailed
study of Diplogastridae, synonymized eight genera
viz., Diplogaster (Loxolaimus) Rahm, 1928, Glauxinema
Allgén, 1947, Eudiplogaster Paramonov, 1952, Tridontus
Khera, 1965, Syedella Suryawanshi, 1971, Diplenteron
Andrássy, 1964, Pareudiplogaster Paramonov, 1952,
and Prosodontus Paramonov & Sobolev in Skrjabin,
with Mononchoides. During a survey of diplogastrid
nematodes from India two species of Mononchoides
were collected. Detailed description of both species
is presented.
MATERIALS AND METHODS
The nematodes were isolated by modified Cobb’s
sieving & decantation and modified Baermann’s funnel
technique and fixed in hot FA (Seinhorst, 1962). After
fixation, the nematodes were transferred to glycerine-
alcohol (5 parts: 95 parts 30% alcohol), dehydrated
by the slow method in a desiccator and mounted
in anhydrous glycerine by the wax ring technique.
M. MAHAMOOD ET AL.

TABLE I.– Measurements (in μm, except L in mm) of Mononchoides megaonchus sp. n.

Holotype Paratypes Paratypes

n 12 11
Character

L 1.37 1.34 ± 0.17 (1.11 – 1.66) 1.13 ± 0.18 (0.90 – 1.21)

a 31.4 30.9 ± 3 (26.0 – 37.8) 32.4 ± 6.8 (23.1 – 45.2)
b 7.2 6.8 ± 0.8 (5.7 – 8.1) 6.7 ± 0.8 (5.4 – 9.1)
c 2.1 2.1 ± 0.2 (1.8 – 2.6) 2.5 ± 0.3 (1.8 – 3.3)
c’ 25.3 23.1 ± 5.4 (15.0 – 34.6) 16.8 ± 4 (11 – 23.9)
V 32.8 34.3 ± 2.9 (29.8 – 39.6) -
Maximum body diameter 44 43 ± 5 (39 – 58) 37 ± 2 (34 – 40)
Length of stoma 44 43 ± 3 (39 – 49) 34 ± 1.5 (33 – 37)
Pharynx 189 194 ± 13 (175 – 218) 164 ± 6 (155 – 176)
Excretory pore from anterior end 141 140 ± 9 (121 – 152) 121 ± 5 (114 – 135)
Hemizonid from anterior end 133 136 ± 8 (122 – 147) 117 ± 5 (96 – 135)
Nerve ring from anterior end 116 123 ± 8 (112 – 133) 105 ± 6 (94 – 116)
Anterior gonad 155 157 ± 29 (102 – 197) -
Posterior gonad 136 146 ± 28 (97 – 194) -
VBD 44 43 ± 4 (39 – 56) -
Vulva – anus distance 282 266 ± 19 (234 – 301) -
Rectum 25 29 ± 3 (25 – 34) 29 ± 2.5 (25 – 32)
Tail 640 620 ± 137 (422 – 873) 429 ± 93 (300 – 582)
ABD 25 27 ± 2 (24 – 29) 27 ± 1 (25 – 28)
Phasmids from anus/cloaca 34 32 ± 3 (28 – 39) 28 ± 1.5 (24 – 29)
Testis - - 347 ± 55 (267 – 417)
Spicules length (along axis) - - 38 ± 1 (36 – 41)
Gubernaculum - - 20 ± 1 (18 – 21)

Measurements were made with an ocular micrometer
on an Olympus CH2 microscope. All morphological
observations, drawings and photographs were made
on an Olympus BX50 DIC microscope.
The terminology describing the parts of the
stoma follows De Ley et al. (1995), and that of other
structural details is according with Fürst von Lieven
and Sudhaus (2000).
DESCRIPTIONS
Mononchoides megaonchus sp. n.
(Figs 1 & 2)
Measurements: See Table I.
Female: Body almost straight upon fixation,
tapering towards both extremities, slightly anteriorly
and sharply posteriorly. Cuticle very thick, with 34
prominent longitudinal ridges and less prominent
transverse striations. Lip region low, continuous
with body contour, 12-16 μm wide. Lips six, fused,
each with a papilla. Amphidial openings elliptical, at
level of cheilostom, almost half as wide as lip region
width. Stoma wide anteriorly, narrow and slender
posteriorly, 2.9-3.8 times lip diameter long. Cheilostom
wide, strongly cuticularised. Cheilorhabdial plates
rod-like, posteriorly overlapping the gymnostom
walls. Cheilorhabdial filaments bifid apically,
extending beyond labial contour. Gymnostom wide,
anisotopic; dorsal wall shorter than subventrals,
heavily cuticualrised. Stegostom anisotopic and
anisomorphic; dorsal wall with a large, massive, claw-
like tooth extending across width of stoma. Right and
left subventral walls without tooth, teeth or denticles.
A strut about one-third to almost half length of the
tubular part of stoma present on the dorsal wall.
Procorpus muscular. Median bulb globular, 18-30
μm long. Isthmus long, gradually expanding to form
an oblong basal bulb. Anterior pharynx muscular,

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 Mononchoides species from India

FIGURE 1. Mononchoides megaonchus sp. n. A: Entire female. B: Entire male. C: Anterior end (female). D:
Pharyngeal region. E: Female reproductive system (anterior). F: Vaginal pouches. G: Female posterior
end. H: Male posterior end.

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M. MAHAMOOD ET AL.

FIGURE 2. Mononchoides megaonchus sp. n. A: Stoma. B: Male anterior end showing amphidial
aperture and cephalic setae. C: Anterior end showing cheilorhabdial flaps. D: Cuticle. E: Vulval
region. F: Phasmidial aperture. G: Female posterior region. H: Male posterior region.

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53-59% of pharyngeal length. Nerve ring encircling
the isthmus in anterior half or mid-isthmus region.
Excretory pore al level of basal bulb expansion,
hemizonid anterior to excretory pore. Intestinal lumen
wide.
Reproductive system amphidelphic; anterior
branch on right and posterior on left side of intestine.
Ovary reversed; oocytes arranged in one or more
rows. Oviduct broad, spermatheca not set off from
the uterus. Uterus not easily distinguishable into
glandular and muscular parts. Vagina sclerotized. A
pair of pouches open at the vagina-uterus junction;
anterior pouch usually longer than posterior. Vulval
opening circular, pore-like. Rectum 0.9-1.4 anal body
diameters long. Phasmids very prominent, 1.1-1.5 anal
body diameter posterior to anus. Tail long, filiform,
1.7-3.0 times vulva-anus distance.
Male: Body smaller than female, curved upon
fixation. Anterior end similar to that of female except
for four cephalic setae. Testis single, reflexed ventrally
anteriorly. Spicules strong, stout, arcuate, 1.3-1.5 anal
body diameters long. Gubernaculum keel-shaped, 50-
55% of spicule length, distally attenuated into a beak-
like process, with sleeve. Genital papillae nine pairs:
three pairs precloacal, six pairs postcloacal. Genital
papillae formula: v1, v2, v3d / v4, ad, phasmids, (v5,
v6, v7), pd. Genital papillae v7 and pd very prominent.
Tail divisible into a short conoid part and a long
filiform part.
Diagnosis: Mononchoides megaonchus sp. n. is
characterized by a large dorsal tooth, long stoma,
absence of tooth, teeth or denticles on the subventral
walls, continuous oviduct, spermatheca and uterus,
stout spicules, and gubernaculum with distal
sleeve.
Relationships: The new species resembles
Mononchoides andrassyi Timm, 1961 in body length,
morphometric values, the structure of female
reproductive system, size of gubernaculum and
number of genital papillae. However, it can easily be
differentiated from M. andrassyi in the length of stoma
(39-49 vs 19-26 μm), size and clarity of amphidial
apertures (large elliptical vs faint reniform), size of
dorsal tooth (as wide as stoma vs smaller than the
width of stoma), shape of spicules (stout vs slender)
and in the shape of gubernaculum (with a distal sleeve
vs without a distal sleeve). The new species further
resembles M. splendidus (Körner, 1954) Goodey, 1963 in
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Mononchoides species from India
body length, value of a-ratio, b-ratio, shape of stoma,
oesophagus, female reproductive system, built and
length of spicules, and number and arrangement of
genital papillae. However, it can be differentiated from
M. splendidus in the value of c-ratio (1.8-2.6 vs 3.1-3.4),
position of vulva (V = 29.8-39.6 vs 41.1-42.1), length
of stoma (more than two times as long as lip width
vs less than two times as long as lip width), built of
spicules (stout vs not stout), shape of gubernaculum
(no projection on the anterio-dorsal side vs hook-
like projection on the anterio-dorsal side) and in the
arrangement of genital papillae (v3d posterior to v2
vs v3d anterior to v2).
Type habitat and locality: Decaying banana rhizome
collected from F-Sector, Itanagar, Arunachal Pradesh,
India.
Type material: Holotype female and paratype
females and males on slides Mononchoides megaonchus
sp. n. deposited in the nematode collection of
Department of Zoology, Aligarh Muslim University,
Aligarh, India. One paratype female and one male
Mononchoides megaonchus sp. n. at the Departamento
de Biología Animal, Biología Vegetal y Ecología,
Universidad de Jaén, Spain.
Etymology: The species name is derived from the
large-sized dorsal tooth.
Mononchoides sp.
(Figs 3 & 4)
Measurements: See Table II.
Female: Body almost straight upon fixation.
Cuticle with transverse striations and prominent
longitudinal ridges. Amphidial apertures elliptical,
at level of gymnostom. Lip region continuous with
body contour. Lip region 9-13 μm wide; lips six,
fused, each with a papilla. Stoma longer than wide.
Cheilostom wide, cuticularized. Cheilorhabdions
rod-like, filaments bifid apically, extending beyond
labial contour. Gymnostom anisotopic, subventral
walls longer than dorsal. Stegostom anisotopic and
anisomorphic. Dorsal wall with a claw-like tooth, right
subventral with a plate-like pyramidal tooth and left
subventral with a denticulate ridge. Posterior part of
stegostom tubular. Procorpus muscular, expanding
M. MAHAMOOD ET AL.

FIGURE 3. Mononchoides sp. A: Entire female. B: Entire male. C: Anterior end (female). D: Anterior end
(male). E: Pharyngeal region. F: Female reproductive system (anterior). G: Female posterior end. H:
Male posterior end.

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 Mononchoides species from India

FIGURE 4. Mononchoides sp. A: Pharynx. B: Amphidial aperture. C: Female

reproductive system (anterior). D: Female anal region. E: Male posterior
region.

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M. MAHAMOOD ET AL.

TABLE II: Measurements (in μm, except L in mm) of Mononchoides sp.

n 9 10
Character

L 928 ± 118 (865 – 1144) 848 ± 66 (724 – 989)

a 27.6 ± 1.6 (24.6 – 30.2) 27.8 ± 1.8 (24.1 – 30.7)
b 6.7 ± 0.3 (6.1 – 7.2) 6.3 ± 0.4 (5.3 – 7)
c 2.4 ± 0.1 (2.2 – 2.8) 2.6 ± 0.4 (2.2 – 3.5)
c′ 18.7 ± 1.8 (16.0 – 21.3) 15.6 ± 3 (10.5 – 21.1)
V 37 ± 1.7 (34.4 – 39.2) -
Maximum body diam. 37 ± 3.2 (32 – 43) 30.5 ± 2 (27 – 35)
Length of stoma 23 ± 2 (18 – 25) 19 ± 1 (18 – 21)
Pharynx 153 ± 11 (128 – 169) 131 ± 7 118 – 143 ()
Excretory pore from anterior end 121 ± 9 (100 – 130) 106 ± 7 (90 – 117)
Hemizonid from anterior end 114 ± 9 (93 – 123) 99 ± 6 (85 – 108)
Nerve ring from anterior end 106 ± 8 (87 – 114) 90 ± 5 (81 – 98)
Anterior gonad 125 ± 13 (97 – 142) -
Posterior gonad 119 ± 14 (91 – 137) -
VBD 37 ± 3 (32 – 42) -
Vulva – anus distance 241 ± 25 (196 – 269) -
Rectum 24 ± 1 (21 – 27) 23 ± 3 (17 – 29)
Tail 403 ± 46 (330 – 475) 318 ± 47 (204 – 369)
ABD 22 ± 2 (19 – 24) 20 ± 2 (17 – 25)
Phasmids from anus/cloaca 25 ± 2 (20 – 29) 24 ± 1 (21 – 27)
Testis - 251 ± 32 (204 – 369)
Spicules length (along axis) - 39.5 ± 2 (37 – 43)
Gubernaculum - 17 ± 2 (14 – 22)

slightly posteriorly to form a 16-20 μm long median
bulb. Isthmus short. Anterior pharynx muscular,
61-68% of pharyngeal length. Nerve ring encircling
isthmus in the anterior half, 87-114 μm from anterior
end. Hemizonid slightly posterior to nerve ring, 93-
123 μm from anterior end. Excretory pore further
posterior to hemizonid.
Reproductive system amphidelphic, anterior
branch on right and posterior branch on left of
intestine. Ovary long, oocytes arranged in several rows
in germinal zone. Oviduct short, narrow, gradually
expanding to form a spermatheca. Spermatheca
elongate, separated from the uterus by a constriction.
Uterus divisible into a long glandular and a short
muscular part. Glandular part 2-3 times muscular part.
Vagina muscular. Vulval opening circular, pore-like.
A pair of dumb-bell shaped pouches present at level
of uterus. Rectum 1.0-1.3 anal body diameters long.
Phasmids prominent, 0.8-1.3 anal body diameters
posterior to anus. Tail long filiform.
Male: Body smaller than female. Anterior part
similar to that of females except for four additional
cephalic setae and narrow elliptical amphidial
apertures. Testis single, reflexed ventrally anteriorly.
Spicules free, arcuate, 1.7-2.1 anal body diameters long.
Gubernaculum 36-52% of spicules length, provided
with a flexible proximal appendage and a distal sleeve.
Genital papillae nine pairs: three pairs precloacal, six
pairs postcloacal. Genital papillae formula: v1, v2, v3d
/ v4, ad, phasmids, (v5, v6, v7), pd. Tail divisible into
a short conoid part and a long filamentous part.
Habitat and locality: Slurry from sewage of D.M.
College of Science, Imphal, Manipur, India.
Voucher specimens: Females and males on slide
Mononchoides sp. deposited in the nematode collection
of Department of Zoology, Aligarh Muslim University,
Aligarh. One female and one male M. sp./11 deposited
at the Departamento de Biología Animal, Biología
Vegetal y Ecología, Universidad de Jaén, Spain.

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Remarks: The present species shares several
characters of taxonomic importance with M. striatus
(Bütschli, 1876) Goodey, 1963 and M. splendidus
(Körner, 1954) Goodey, 1963. It resembles M. striatus
in morphometric characters and also in the shape
and size of spicules. However, there is a significant
difference in the length-width ratio of the tubular
part of stegostom (more than two times as long
as wide vs as long as wide) and in the structure of
gubernaculum, which is narrower, has a sleeve at
the distal end and a short flexible appendage at the
proximal end. In M. striatus the gubernaculum is
broader, the distal sleeve and the proximal thread-
like appendage were not depicted either by Butschlii
(1876) or Weingartner (1955). M. splendidus resembles
our species in the morphometrics, but is different in
that the buccal cavity lacks the claw-like tooth on
the right sub-ventral wall and denticles on the left
sub-ventral wall. The present species further differs
from M. splendidus in the shape and structure of
gubernaculum (slender with a filamentous appendage
proximally and a small sleeve distally vs robust with
a prominent hook proximally and a broad sleeve
distally). Slight differences also occur in the c-ratio
(2.2-2.8 vs 3.1-3.4) and position of vulva (V = 34-39 vs
41-42). Both these values are length-dependent and the
tail in the genus Mononchoides, as in some other genera
also, is extremely long and filamentous/whip-like
and is frequently broken. The breakage is sometimes
imperceptible and affects the c-ratio and V (as well
as b-ratio). Hence, we consider these differences to be
minor. Between the two species we believe that our
Mononchoides sp. is more closely related to M. striatus
than to M. splendidus. Nevertheless, certain pertinent
differences especially those of gubernaculum and
stegostom prohibit us in assigning these specimens
as M. striatus.
ACKNOWLEDGEMENT
This work was supported by a grant from the
Ministry of Environment and Forests, Government
of India, New Delhi, under the AICOPTAX
programme.
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Mononchoides species from India
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Accepted May 7, 2007