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Zootaxa 3646 (4): 301–335 ISSN 1175-5326 (print edition)
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Copyright © 2013 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3646.4.1
http://zoobank.org/urn:lsid:zoobank.org:pub:DDE05DCD-A443-499D-9F38-8C3B43592694
Descriptions of ten known species of the superfamily Mononchoidea

(Mononchida: Nematoda) from North India with a detailed
account on their variations
QUDSIA TAHSEEN1,3, MOHAMMAD ASIF1, MALKA MUSTAQIM1, SHIKHA AHLAWAT1

& WIM BERT2
1
Nematode Research Laboratory, Department of Zoology, Aligarh Muslim University, Aligarh, India
2
Department of Biology, University of Ghent, Belgium
3
Corresponding author. E-mail:qtahseen@yahoo.com
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302
Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302
Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303
Clarkus papillatus (Bastian, 1865) Jairajpuri, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303
Prionchulus muscorum (Dujardin, 1845) Wu & Hoeppli, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 306
Mylonchulus minor (Cobb, 1893) Andrássy, 1958 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308
Mylonchulus lacustris (Cobb in Cobb, 1915) Andrássy, 1958 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 311
Mylonchulus obtusicaudatus (Daday, 1899) Andrássy, 1958 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 316
Mylonchulus hawaiiensis (Cassidy, 1931) Andrássy, 1958 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 318
Mylonchulus vasis Yeates, 1992 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 321
Mylonchulus contractus Jairajpuri, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 323
Sporonchulus vagabundus Jairajpuri, 1971. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 323
Sporonchulus ibitiensis (Carvalho, 1951) Andrássy, 1958 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 327
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 330
Acknowledgement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 332
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 332
Abstract
Ten species of the superfamily Mononchoidea from fourteen populations collected during a survey of terrestrial and fresh-
water habitats of North India, are described and illustrated. The species include Clarkus papillatus (Bastian, 1865) Jaira-
jpuri, 1970, Prionchulus muscorum (Dujardin, 1845) Wu & Hoeppli, 1929, Mylonchulus contractus Jairajpuri, 1970, M.
hawaiiensis (Cassidy, 1931) Andrássy, 1958, M. lacustris (Cobb in Cobb, 1915) Andrássy, 1958, M. minor (Cobb, 1893)
Andrássy, 1958, M. obtusicaudatus (Daday, 1899) Andrássy, 1958, M. vasis Yeates, 1992, Sporonchulus ibitiensis (Car-
valho, 1951) Andrássy, 1958 and S. vagabundus Jairajpuri, 1971. Mylonchulus vasis Yeates, 1992 is reported for the first
time from India. The variable, as well as the relatively consistent characters are discussed in different populations to assess
their role in the diagnosis of species.
Key words: Description, LM, SEM, mononchs, taxonomy, Clarkus, Mylonchulus, Prionchulus, Sporonchulus
Accepted by P. Mullin: 25 Feb. 2013; published: 6 May 2013 301

Introduction
Mononchs represent a group of predatory nematodes having large buccal cavities usually armed with teeth
(Andrássy, 1993). The mononchs are largely terrestrial nematodes, with very few species inhabiting freshwater or
marine habitats. Some species are cosmopolitan in distribution while others prefer relatively specific habitats
(Andrássy, 2006).
The first mononch species was reported by Dujardin in 1845 as Oncholaimus muscorum Dujardin, 1845.
However, the type genus Mononchus was proposed twenty years later by Bastian (1865), who described five new
species and also transferred the species of Dujardin under this genus. Cobb (1916) split Mononchus into five
subgenera, viz, Mononchus Bastian, 1865, Prionchulus Cobb, 1916, Mylonchulus Cobb, 1916, Iotonchus Cobb,
1916 and Anatonchus Cobb, 1916, and later, in 1917, he proposed the subgenus Sporonchulus Cobb, 1917. The
family Mononchidae Chitwood, 1937 was initially represented by only two genera, viz., Mononchus Bastian, 1865
(with subgenera Mononchus, Mylonchulus, Anatonchus and Sporonchulus) and Prionchulus (Cobb, 1916) Wu &
Hoeppli, 1929. It was later raised to superfamily level by Clark (1961). Jairajpuri (1969) proposed the order
Mononchida and included the family Mononchidae under the superfamily Mononchoidea Chitwood, 1937,
represented by Mononchus, Prionchulus and a later established genus Clarkus Jairajpuri, 1970. Wu & Hoeppli
(1929) gave Prionchulus generic rank, while the subgenera Mylonchulus and Sporonchulus were raised to generic
level by Altherr (1953) and Pennak (1953), respectively. Jairajpuri & Khan (1982) recognized three families under
Mononchoidea viz., Mononchidae Chitwood, 1937, Mylonchulidae Jairajpuri, 1969 and Cobbonchidae Jairajpuri,
1969. They (l. c.) considered two subfamilies under Mononchidae, Mononchinae Chitwood, 1937 for the genera
Mononchus and Paramononchus Mulvey, 1978 and Prionchulinae Andrássy, 1976 for the genera Prionchulus,
Clarkus, Coomansus Jairajpuri & Khan, 1977 and Actus Baqri & Jairajpuri, 1973. However, Andrássy (1993)
considered Mononchinae and Cobbonchinae Jairajpuri, 1969 under family Mononchidae and placed Mononchus,
Actus, Clarkus and Prionchulus together under subfamily Mononchinae.
During a nematode faunal survey of North India from soil and freshwater habitats, ten species, representing
five genera of mononchs, were identified from fourteen populations. The genera Clarkus, Prionchulus,
Mylonchulus and Sporonchulus belong to the non-tuberculate (pharyngo-intestinal junction without tubercles)
superfamily Mononchoidea and are the representatives of Mononchidae, Prionchulidae and Mylonchulidae sensu
Ahmad & Jairajpuri (2010). The species found: Clarkus papillatus (Bastian, 1865) Jairajpuri, 1970a, Prionchulus
muscorum (Dujardin, 1845) Wu & Hoeppli, 1929, Mylonchulus contractus Jairajpuri, 1970b, M. hawaiiensis
(Cassidy, 1931) Andrássy, 1958, M. lacustris (Cobb in Cobb, 1915) Andrássy, 1958, M. minor (Cobb, 1893)
Andrássy, 1958, M. obtusicaudatus (Daday, 1899) Andrássy, 1958, M. vasis Yeates, 1992, Sporonchulus ibitiensis
(Carvalho, 1951) Andrássy, 1958 and S. vagabundus Jairajpuri, 1971are described hereunder. The species
Mylonchulus vasis is reported for the first time from India.
Material and methods
The soil and water samples were processed using Cobb’s (1918) sieving and decantation and modified Baermann’s
(1917) funnel techniques. The nematodes were extracted and fixed in hot formalin-glycerol fixative, dehydrated by
the slow evaporation method (Seinhorst, 1959) and mounted in anhydrous glycerine. Permanent mounts were
prepared using the paraffin wax-ring method (de Maesneer & d’Herde, 1963). The measurements were taken with
an ocular micrometer. LM photographs were taken with a Jenoptik ProgRes digital camera mounted on an
Olympus BX-51 DIC microscope. For Scanning Electron Microscopy (SEM), the specimens were fixed in 2%
glutaraldehyde, post-fixed in 2% osmium tetroxide, dehydrated in alcohol series and critical point dried using CO2.
The mounted nematodes were coated with 10 nm gold before viewing at 10 kV with an XL30 FEG scanning
electron microscope.
302 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

Systematics
Clarkus papillatus (Bastian, 1865) Jairajpuri, 1970

(Fig. 1)
=Mononchus papillatus Bastian, 1865

=Mononchus (Prionchulus) papilIatus Bastian, 1865 (Schneider, 1939)
=Prionchulus papillatus (Bastian, 1865) Schneider, 1939
=Mononchus cristatus Bastian, 1865
=Clarkus koreanus Choi, Khan & Lee, 1999
Measurements. Table 1
Description. Adult: Body medium-sized, ventrally curved upon fixation. Cuticle smooth 2–3 µm thick in
different body regions. Body pores not visible. Lip region set off, ca 2.4–2.6 times as wide as high, distinctly wider
than adjoining body, with very small or inconspicuous labial and cephalic sensilla. Amphids cup-shaped with oval
apertures, 3–4 µm across, located anterior to dorsal tooth or 10–11 µm from anterior end. Buccal cavity barrel-
shaped tapering at base about 2–3 times as long as wide with heavily cuticularised parallel vertical and oblique
walls; vertical walls consisting of a tooth-bearing dorsal wall and two toothless sub-ventral walls. Dorsal tooth
prominent, 3–4 µm in thickness, situated in the anterior half of buccal cavity or about 82–83% of buccal capsule
from its base; anterior margin of the tooth somewhat straight. Subventral walls of buccal cavity with a longitudinal
ridge extending from the point opposite to dorsal tooth’s apex to posterior part of vertical plates. Pharyngeal sleeve
surrounding stoma at 1/3–1/4 of its length from base. Pharynx cylindroid, muscular ca 26–28% of body length.
Nerve ring at 31–38% of pharyngeal length. Excretory pore 35–42% of pharyngeal length from the anterior end.
Orifices of the pharyngeal glands: DO at ca 53–58%, SV1O1 and SV1O2 at ca 72–75%, and SV2O1 and SV2O2 at ca
94–97% of pharyngeal length. Pharyngo-intestinal junction non-tuberculate. Intestinal lumen wide, intestinal wall
made up of large polygonal cells. Rectum ca 0.8–1.1 times anal body diameter long. Tail conoid, ventrally arcuate
with acute terminus. Caudal glands absent, rudimentary in one specimen; spinneret absent.
Female: Reproductive system didelphic, amphidelphic; ovaries reflexed, of robust size, about as wide as
corresponding body diameter, with 7–10 oocytes arranged alternately or in single row in each ovary; and distal end
with a cluster of proliferating oogonial cells. Oviduct narrow connected to uterus without any sphincter in between.
Uterus muscular, accommodating usually one egg at a time, occasionally with two large-sized, smooth-shelled
eggs, 76–80 µm long × 48–49 µm wide. Vagina extending more than half of corresponding body diameter with
small, moderately sclerotised pars refringens. Vulva an oval slit of 9–12 µm diameter.
Male: Not found.
Locality and habitat. Soil containing Clarkus papillatus was collected from around the roots of Pinus sp.
from Dehradun, Uttarakhand, India, located at 30.3157°N 78.0336°E coordinates.
Voucher specimens. One female on slide Clarkus papillatus (Bastian, 1865) Jairajpuri, 1970 no. D13/1
deposited in the Nematode Collection, Department of Zoology, Aligarh Muslim University, Aligarh, Uttar Pradesh,
India. One female deposited at USDANC, Beltsville, MD, USA.
Salient characters. Medium-sized nematodes; buccal cavity barrel-shaped, dorsal wall bearing a medium-
sized tooth; each subventral wall provided with non-denticulate ridge. Pharyngo-intestinal junction non-
tuberculate. Female genital system amphidelphic. Tail short, conoid, ventrally arcuate with acute terminus. Caudal
glands rudimentary or absent; spinneret absent.
Remarks. This is the most widely distributed species of the genus, and has been reported from all continents
(Andrássy, 2006) except Antarctica. The present specimens show morphological as well as morphometric
characteristics similar to specimens of Clarkus papillatus described earlier (Jairajpuri, 1970a; Andrássy, 2006);
however, the uterine eggs were relatively smaller (76–80 × 48–49 µm vs 80–92 × 32–44 µm) than those reported
for C. papillatus apud Andrássy (2006). The lip region was variable from rounded to angular. The buccal cavity
also showed variation in shape with the vertical walls straight to slightly curved. The inconspicuous caudal glands
and absence of spinneret are typical features of this species, although the tails exhibited variation in degree of
ventral curvature of the two specimens studied.
MONONCHOIDEA FROM INDIA Zootaxa 3646 (4) © 2013 Magnolia Press · 303
TABLE 1. Morphometrics of females of Clarkus papillatus (Bastian, 1865) Jairajpuri, 1970, Prionchulus muscorum (Dujardin, 1845) Wu & Hoeppli, 1929,
Sporonchulus vagabundus Jairajpuri, 1971 and S. ibitensis (Carvalho, 1951) Andrássy, 1958. Measurements are in μm and in the form: mean ± SD (range).
Characters (n) Clarkus papillatus (2) Prionchulus muscorum Sporonchulus vagabundus (6) S. vagabundus(4) S. ibitensis (1)
(5) Sonbhadra population Dehradun population
Body length 1080±254 (900–1260) 1750±80 (1410–2020) 1023±49 (956–1076) 1009±66 (909–1090) 1103
Body diameter at vulva 55±14.8 (45–66) 92±6.2 (85–100) 38±1.9 (36–41) 45±4.3 (42–53) 40
a 19.5±0.7 (19.2–20.2) 18.9±1.6 (16.6–20.4) 26.4±1.3 (25.1–29) 22.1±1.8 (20.2–24.1) 27.5
b 3.7±0.1 (3.6–3.7) 3.8±0.2 (3.4–4.1) 3.9±0.1 (3.8–4) 3.9±0.2 (3.6–4.2) 4.0
c 16.9±1.6 (13.5–18.1) 10.7±1.2 (8.9– 12.1) 51.3±2.8 (47–55) 20.3±0.9 (19.2–21.8) 20.4
c’ 2.1±0.1 (2.1–2.3) 3±0.2 (2.6–3.2) 2±0.1 (1.9–2.2) 2±0.2 (1.7–2.4) 2
304 · Zootaxa 3646 (4) © 2013 Magnolia Press

V 62.7±0.5 (62.3–63.0) 61.3±0.9 (59.8–62.7) 60.1 ± 1.0 (59.7– 60.6) 61.9±1.7 (59.2–64.2) 61
G1 23.5±1.1 (22.7–24.2) 20.0±0.7 (18.9–20.0) 13.0±1.6(11.1–13.2) 10.1±0.8 (9.0–10.9) 14.1
G2 23.9±2 (22.5–25.3) 19.7±2.0 (17.7–22.5) 12.1±1.3(11.1–13.7) 9.6±0.8 (8.9–10.9) 11.5
Lip height 6.5±2 (5–8) 18.4±1.3 (17–21) 5±0 (5–5) 6.8±1.1 (5–8) 4
Lip diameter 25±1.4 (24–26) 41.2±2.78 (38–45) 21±1.0 (20–22) 22.5±0.5 (22–23) 22
Stoma length 25.5±3.5 (23–28) 45±1.0 (44–47) 26±0.8 (25–27) 25±0.7 (24–26) 22
Stoma diameter 14.5±2.1 (13–16) 27.8±0.7 (27–29) 12.5±0.1 (12–13) 11.7±0.4 (11–12) 11
Dorsal tooth position 5.5±0.7 (5–6) 15±0 (15–15) 6±0 (6–6) 6.5±0.5(6–7) 6
Pharynx length 292.5±60.1 (250–335) 460.2±37.3 (415–522) 259.8±14.5 (234–275) 254.2±25.1 (232–297) 272
Nerve ring from ant. end 97.5±3.5 (95–100) 147.5±15.9 (125–163) 109±20.4 (90–130) 80.2±10.6 (70–98) 95
Excretory pore from ant. end 113.5±9.2 (107–120) 163.3±16.4 (145–185) 116.1±1.4 (115–118) 101.2±7.5 (96–110) 109
Rectum length 26±5.7 (22–30) 41.7±2.7 (39–45) 23.7±0.9 (23–25) 26.0±1.7 (25–28) 22
Anal body diameter 30±2.8 (28–32) 54.6±4.9 (46–59) 24.5±0.8 (24–26) 25±2.1 (22–28) 27
Tail length 63.5±9.2 (57–70) 169.4±13.8 (107–187) 51.3±2.8 (47–55) 49.7±3.1 (45–54) 57
Vulva–anus distance 336.5±87 (275–398) 515.2±82.7 (460–601) 355.6±15.6 (335–380) 354±46.3 (285–383) 380
TAHSEEN ET AL.
FIGURE 1. Clarkus papillatus (Bastian, 1865) Jairajpuri, 1970 female. A–C: Anterior end. D: Pharyngo-intestinal junction. E:
Anterior genital branch. F: Posterior genital branch. G: Vulval region. H: Intestinal lumen showing two cellular bodies. I, J:
Posterior body region (Scale bar = 10 µm).
Prionchulus muscorum (Dujardin, 1845) Wu & Hoeppli, 1929
(Fig. 2)
=Oncholaimus muscorum Dujardin, 1845

=Mononchus muscorum (Dujardin, 1845) Bastian, 1865
=Mononchus (Prionchulus) muscorum (Dujardin, 1845) Bastian, 1865 (Cobb, 1916)
=Mononchus bastiani de Man, 1876
=Mononchus (Prionchulus) longicollis Cobb, 1917
=Mononchus papillatus macrodon Fuchs, 1930
=Prionchulus medius Eroshenko, 1975
=Prionchulus paucidentatus Zell, 1985
=Prionchulus ferox Winiszewska & Susulovsky, 2003
=Prionchulus olexandri Winiszewska & Susulovsky, 2003
=Prionchulus prasadi Saha, Lal & Singh, 2006
=Prionchulus ukhrum Mohilal & Dhanachand, 1997
=Prionchulus pachydermis Khan, Choi, Lee & Choi, 2000
Measurements. Table 1.
Description. Adult: Body medium to large-sized, straight to open ‘C’-shaped on fixation, strongly curved in
tail region, broad at anterior end but tapering at posterior extremity. Body pores indistinct. Cuticle smooth, about
3.0–3.5 µm thick in different body regions. Lip region slightly offset from the body contour, about 2.0–2.5 times as
wide as high; labial sensilla prominent, slightly raised. Amphids cup-shaped, 5–6 µm across, located anterior to
dorsal tooth, at 18–23 µm from anterior end of body and 32–34 µm from base of buccal cavity. Buccal cavity large,
elongate barrel-shaped, about 1.6 times as long as wide, with thick, heavily cuticularised vertical and oblique walls.
Dorsal wall bearing a sharp, pointed, 10–12 µm long and 5–6 µm wide dorsal tooth, directed forward, located in
the anterior half of buccal cavity at 73–75% from its base; each sub-ventral wall bearing a longitudinal denticulate
ridge with 11–16 closely placed denticles. Pharyngeal collar surrounding buccal cavity at 1/3–1/4 of its length from
base or up to the level of oblique plates. Pharynx cylindroid, muscular, 25–30% of body length. Orifices of
pharyngeal glands, DO located at ca 54–55%, SV1O1 and SV1O2 at ca 67–68%, SV2O1 and SV2O2 at ca 94–96% of
pharyngeal length from anterior end. Nerve ring at 30–32% of pharyngeal length, conspicuous in few specimens
only. Excretory pore at 34–36% of pharyngeal length. Pharyngo-intestinal junction non-tuberculate. Intestine with
narrow lumen, in some specimens containing cuticularised structures of devoured nematode prey. Rectum 1.2–1.3
times the anal body diameter. Tail conoid, strongly curved ventrally ending in acute terminus. Caudal glands and
terminal opening absent.
Female: Reproductive system didelphic, amphidelphic; ovaries reflexed with oocytes arranged in a single row,
except at the tip. Vagina about half of corresponding body diameter with ovoid, dorso-ventrally compressed pars
refringens. Vulva post-equatorial, a transverse slit with slightly cuticularised lips.
Male: Not found.
Locality and habitat. Soil sample containing Prionchulus muscorum was collected from around grasses
(Agrostis sp.), Dehradun, Uttarakhand, India, located at 30°20'38"N 77°59'52"E coordinates.
Voucher specimens. Four females on slide Prionchulus muscorum (Dujardin, 1845) Wu & Hoeppli, 1929 no.
D12/1–2 deposited in the Nematode Collection, Department of Zoology, Aligarh Muslim University, Aligarh, Uttar
Pradesh, India. One female deposited at USDANC, Beltsville, MD, USA.
Salient characters. Medium to large-sized species; buccal cavity of moderate size with parallel vertical walls;
dorsal wall bearing medium-sized tooth; each subventral vertical wall provided with denticulate ridge having 11–
16 denticles; Pharyngo-intestinal junction non-tuberculate; female reproductive system amphidelphic; tail short
conoid with rudimentary or no caudal glands; spinneret absent.
Remarks. P. muscorum (Dujardin, 1845) Wu & Hoeppli, 1929 is a widely distributed species reported from a
large number of countries of Europe, Asia, Africa, America and Australia (Andrássy, 2006), predominantly from
soil and occasionally from moss as well as aquatic habitats. The species seems to be facultatively parthenogenetic
as it is often represented by females in populations, with males extremely rare or absent.
The present specimens conformed well in most morphological and morphometric characteristics to P.
muscorum (apud Jairajpuri, 1970a; Andrássy, 1985) except in having a smaller a value (16–20 vs 21–33). Most
characters were in total agreement with those specified for P. muscorum apud Arpin et al. (1984) viz., dorsal tooth

FIGURE 2. Prionchulus muscorum (Dujardin, 1845) Wu & Hoeppli, 1929 female. A–D: Anterior end. E–H: Subventral wall
with denticulate ridge. I: Body region showing excretory pore. J–L: Pharyngo-intestinal junction. M: Vulval region. N, O:
Posterior body region. P: tail tip. (Scale bar = 10 µm).
located at 73–75% of the length of the buccal cavity from its base, smooth-shelled intra-uterine eggs, buccal cavity
with prominently ridged subventral walls having 11–16 denticles and a relatively slender, conoid tail. However,
presence of additional features considered to be specific for P. punctatus raise questions regarding the reliability
and credence of these characters in diagnosis. The buccal cavity was observed to be relatively small and
rectangular in some individuals (Fig. 2A); the thickness of the ridge as well as the arrangement of denticles
exhibited variation (Fig. 2E–H). The tail in a few individuals was observed to be short and conoid with a slightly
swollen (Fig. 2N), fusiform terminus (Fig. 2O). Thus the only character for P. muscorum that appears
differentiating and reliable is the smooth (not echinulate) egg shell surface. The intraspecific variations in P.
muscorum as also reported by Clark (1960), Mulvey (1967) and Arpin et al. (1984) can be due to the cosmopolitan
nature of species that occurs in different biogeographical locations.
Mylonchulus minor (Cobb, 1893) Andrássy, 1958

(Fig. 3)
=Mononchus minor Cobb, 1893

=Mononchus (Mylonchulus) minor Cobb, 1893 (Cobb, 1916)
=Mononchus (Mylonchulus) brachyuris microdenticulatus minor Cobb, 1893 (Micoletzky, 1922)
=Mylonchulus sculentus Jain, Saxena & Sharma, 1993
Description. Adult: Body medium-sized, ventrally arcuate, more strongly curved in posterior half after
fixation, tapering slightly towards extremities. Cuticle smooth, 1.5–2 µm thick. Body pores indistinct. Lip region
truncate to angular, demarcated from adjoining body, about 2.5–3 times as wide as high. Labial sensilla prominent,
slightly raised. Amphids cup-shaped with slit-like aperture, 4–7 µm across, located 9–12 µm from the anterior end.
Buccal cavity funnel-shaped, about 1.2–1.8 times as long as wide with sclerotized vertical and oblique walls.
Dorsal vertical wall bearing anteriorly directed, 8–9 µm long massive tooth, 5–6 µm in thickness with tip located
5–6 μm from anterior end of buccal capsule or at ca 70–80% from its base. Each subventral vertical wall provided
anteriorly with 6 transverse rows of rasp-like denticles (n=6) with anterior row relatively more prominent; number
of denticulate rows was found to be five in two specimens. A small tooth present on each subventral wall posterior
to denticles. Pharyngeal sleeve surrounding buccal cavity at 1/3–1/4 of its length from base. Pharynx cylindroid,
muscular, 30–33% of body length. Orifices of the pharyngeal glands, DO located at ca 58–67%, SV1O1 and SV1O2
at ca 69–73%, SV2O1 and SV2O2 at ca 92–93% of pharyngeal length from anterior end. Nerve ring encircling
pharynx at ca 27–33% and excretory pore faintly visible at ca 34–39% of pharyngeal length from anterior end.
Pharyngo-intestinal junction non-tuberculate with cardial flap of variable length ranging from 9–15 µm. Intestine
with polygonal cells. Rectum 0.8–1.1 times as long as anal body diameter. Tail conoid, ventrally bent at half of
length in most specimens, tapering into a rounded terminus. Caudal glands well developed, arranged in tandem,
opening to exterior through a narrow tubular duct; spinneret terminal.
Female: Reproductive system didelphic, amphidelphic with relatively compact, reflexed ovaries having 8–11
oocytes arranged in single row except the distal proliferation zone; proximal end of ovary containing a large grown
oocyte. Oviduct joining ovary sub-terminally, 35–77 µm long. Sphincter not observed at oviduct-uterus junction;
uterus devoid of sperms. Vagina extending inward about 1/3–1/4 of the corresponding body diameter with well
developed, almond-shaped pars refringens. A smooth-shelled intra-uterine egg observed in one female. Vulva
postmedian and transverse.
Male: Not found.
Locality and habitat. Moss containing Mylonchulus minor, scraped from around the roots of Bambusa sp.
from Haridwar, Uttar Pradesh, India located at 29°57′22″N 78°10′12″E coordinates.
Voucher specimens. Seven females on slide Mylonchulus minor (Cobb, 1893) Andrássy, 1958, no. D20 /1–4
Salient characters. Small to medium-sized species with buccal cavity funnel-shaped, about 1.2–1.8 times
long as wide; dorsal tooth large, situated in anterior half of buccal cavity; each subventral wall usually bearing 6
transverse rows of denticles; subventral tooth very small; female genital system amphidelphic; tail conoid,
ventrally bent at half of its length with rounded terminus; caudal glands arranged in tandem; spinneret terminal.

TABLE 2. Morphometrics of Mylonchulus minor (Cobb, 1893) Andrássy, 1958, M. obtusicaudatus (Daday, 1899) Andrássy, 1958, M. vasis Yeates, 1992 and M. contractus
Jairajpuri, 1970b. Measurements are in μm and in the form: mean ± SD (range).
Characters M. minor female M. obtusicaudatus female M. vasis female M. contractus female M. contractus male
(n) (8) (6) (6) (10) (1)
Body length 1010±60 (930–1140) 1400.3±90.7 (1280–1620) 916.6±65.8 (839–1014) 762.6±43.6 (696– 822) 801
MONONCHOIDEA FROM INDIA

Body diameter at vulva 48.2±3.6 (43–53) 56.2±6.5 (46–70) 43.3±4.0 (38–50) 31.6±0.9 (30–33) 28
a 21.1±1.6 (19.0–24.0) 25.1±2.2 (20.6–28.1) 21.1±1.4 (19–23.3) 24.1±1.5 (21.6–26.4) 28.6
b 3.4±0.1(3.2–3.5) 3.9±0.2 (3.7–4.6) 3.1±0.1(3–3.3) 3±0.1 (2.9–3.1) 3
c 25.1±1.7 (22.6–27.1) 24.9±1.2(22.3–26.8) 26.5±2 (24.2–29.8) 27±2.5 (22.8–31.4) 24.2
c' 1.4±0.9 (1.2–1.5) 1.6±0.1(1.3–1.9) 1.1±0.1(1.0–1.3) 1.2±0.1 (1.1–1.4) 1.1
V /T 58.2±1.1(56.7–60.1) 62.0±0.8 (60.7–63.1) 61.8±3.2 (58.7–66.5) 59.7±1.1 (57.7–61.0) 45.5
G1 15.5±1.9 (12.0–18.1) 12.3±1.2 (10.8–14.9) 15.0±2.1 (13.3–18.2) 12.3±1.1 (11.3–13.5) –
G2 14±1.8 (12.2–17.7) 11.9±1.3 (9.1–13.7) 12.3±0.8 (11.4–13.5) 10.4±0.5(9.7–11.2) –
Lip height 9.8±0.3 (9–10) 10.9±0.5 (10–12) 9.5±0.5 (9–10) 7.7±0.4 (7–8) 7
Lip diameter 24.8±1.2 (23–27) 28±0.7 (27–29) 24.1±1.1 (23–26) 18.5±0.9 (17–20) 18
Stoma length 25±0.5 (24–26) 31.2±0.9 (30–33) 30.6±0.8 (30–32) 24.7±0.8 (23–26) 25
Stoma diameter 15.2±1.1(13–17) 19.2±0.7(18–20) 16.0±0.8 (15–17) 12.4±0.5 (12–13) 12
Dorsal tooth position 5.7±0.5 (5–6) 8.6±0.5 (8–9) 6.6±0.5 (6–7) 4.6±0.5 (4–5) 10
Pharynx length 294.5±11.2 (280–318) 355.8±15.9 (335–390) 290.8±12.9 (276–312) 248.5±13.3 (233–268) 263
Nerve ring from ant. end 89.8±7.2 (80–100) 99.7±5.8 (85–110) 94±3.5 (90–99) 82±4.3 (75–90) 85
Excretory pore from ant. end 109.2±8.6 (96–125) 125.3±17.1 (108–173) 98±1.6 (96–100) 92.6±3.8 (89–95) 93
Rectum length 24.6±1.3 (23–27) 28.7±2.9 (24–33) 23±1.8 (21–25) 19.7±1.4 (18–22) 24
Anal body diameter 29.1±1 (28–31) 34.9±1.9 (32–38) 28.1±2.3 (25–31) 21.9±0.7 (21–23) 28
Tail length 40.7±3 (37–47) 56.4±4.5 (50–62) 34.5±1.7 (32–37) 28.4±3.1 (24–34) 33
Vulva–anus distance 345.2±21.3 (307–398) 426.7±28.9 (388–476) 323.1±18.6 (300–345) 277.9±17.3 (250–295) –
Spicule length – – – – 36
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Gubernaculum length – – – – 23
309
FIGURE 3. Mylonchulus minor (Cobb, 1893) Andrássy, 1958, female. A–G: Anterior end with buccal armature. H: Anterior
end showing amphids. I–L: Pharyngo-intestinal junction. M–Q: Anterior genital branch. R, S: Vulval region. T: Posterior
genital branch. U–X: Posterior body region. (Scale bar = 10 µm).

Remarks. This is a widely distributed species of Mylonchulus, and has been reported from different parts of
the world (Ahmad & Jairajpuri, 2010). The present specimens showed morphological as well as morphometric
characteristics similar to M. minor (Cobb, 1893) Andrássy, 1958. The typical features of the individuals as stated
by Cobb (1916) were the funnel-shaped buccal cavity with a prominent dorsal tooth but very short or relatively
inconspicuous subventral teeth, likely to be overlooked. Intrapopulation variations were observed in the size of
subventral tooth which ranged from a slight protuberance (Fig. 3B, C) to a prominent triangular tooth (Fig. 3G),
subventral rasp-like denticles being either regularly arranged in rows or slightly dispersed, the cardial flap ranging
from insignificant (Fig. 3L) to visibly longer (Fig. 3I); and the ovaries slender (Fig. 3M) to prominently broad (Fig.
3Q). Contrary to the description of the species apud Cobb (1916) as having a conoid, arcuate tail bent near the
middle, the shape of tail in the present specimens varied from cylindroid (Fig. 3U), slightly bent to strongly bent
(Fig 3X).
Mylonchulus lacustris (Cobb in Cobb, 1915) Andrássy, 1958

(Figs. 4–6)
=Mononchus lacustris Cobb in Cobb, 1915

=Mononchus (Mylonchulus) lacustris Cobb in Cobb, 1915 (Cobb, 1917)
=Mononchus (Mylonchulus) brachyuris macrodenticulatus lacustris Cobb in Cobb, 1915 (Micoletzky, 1922)
=Mylonchulus sarmini (Azmi, 1991) Ahmad & Jairajpuri, 2010
=Mylonchulus yassini (Elbadri, Khan, Moon, Lee & Choo, 2008) Ahmad & Jairajpuri, 2010
=Mylonchulus lacumurus (Saha, Lal & Singh, 2004) Ahmad & Jairajpuri, 2010
Description. Adult: Body medium-sized, ventrally curved upon fixation, more strongly curved in posterior
region. Cuticle smooth, 2–3 µm thick at various body regions. Body pores indistinct. Lip region offset from
adjoining body, more than 3 times as wide as high. Labial papillae slightly projecting above labial contour.
Amphids cup-shaped with oval aperture, 4 µm across, located 10 µm from anterior end, ahead of dorsal tooth.
Buccal cavity oblong, goblet-shaped with parallel walls and narrow base about 1.6–2 times as long as wide with
heavily sclerotised vertical and oblique plates. Vertical walls comprising of a dorsal tooth bearing wall and two
sub-ventral denticulate walls each with 7 transverse rows of denticles (n= 6) and a prominent sub-ventral tooth.
One individual possessed 6 irregular or dispersed denticulate rows (Fig. 4C) in the buccal cavity. Dorsal tooth
medium-sized, 8–9 x 5–6 µm in dimension, anteriorly directed, with tip located at 5–6 µm from anterior end of
buccal capsule or at ca 74–80% from its base. Pharyngeal sleeve surrounding buccal cavity base at 1/3–1/4 of its
length. Pharynx cylindroid, muscular, 28–29% of body length. Ventral pharyngeal wall with conspicuous bead-like
(Fig. 6G) appearance (in individuals belonging to Ghazipur population). Orifices of pharyngeal glands: DO located
at ca 65–67%, SV1O1 and SV1O2 at ca 79–82%, SV2O1 and SV2O2 at ca 91–93% of pharyngeal length from anterior
end. Nerve ring surrounding pharynx at 28–31% of pharyngeal length from anterior end. Excretory pore
inconspicuous, in few specimens observed at ca 36–39% of pharyngeal length. Pharyngo-intestinal junction non-
tuberculate. Rectum ca 0.8–1.2 times anal body diameters long. Tail conoid, with slight ventral curvature,
gradually tapering to a cylindroid part before ending in a rounded terminus. Caudal glands well developed,
arranged in tandem, or slightly deviated from linear pattern; spinneret terminal.
Female: Reproductive system didelphic, amphidelphic with reflexed ovaries. Anterior genital branch a little
longer than posterior in most specimens; each ovary with a very large proximal oocyte and small oocytes in double
rows present distally. Vagina one-third of corresponding body width with well-developed, almond-shaped pars
refringens. Vulva a transverse slit, slightly posterior to middle of the body, vulval lips occasionally protruding (in
Aligarh population).
Male: Not found.
Locality and habitat: Aligarh population: The sample containing M. lacustris was collected from a small
ditch (alluvial soil) at the outskirts of Aligarh, Uttar Pradesh, India, at 27°53′N 78°05′E coordinates. Ghazipur
population: The sample containing M. lacustris was collected from a pit at Ghazipur, Uttar Pradesh, India, at
25°35′N 83°34′E coordinates. Udaipur population: The sample containing M. lacustris was collected from a field at
Udaipur, Rajasthan, India, at 24°35'11"N 73°43'20"E coordinates.
TABLE 3. Morphometrics of females of Mylonchulus lacustris (Cobb in Cobb, 1915) Andrássy, 1958 and M. hawaiiensis (Cassidy, 1931) Andrássy, 1958. Measurements
are in μm and in the form: mean±SD (range).
Characters (n) M. lacustris (2) M. lacustris (3) M. lacustris (2) M. hawaiiensis (10) M. hawaiiensis (3)
Aligarh population Ghazipur population Udaipur population Dehradun population Keetham population
Body length 1080±60 (1020–1140) 962±28 (828–1085) 970±49 (794–1147) 880±80 (710–950) 820±68 (750–890)
Body diameter at vulva 38±2.3 (36–40) 37±1.1 (35–39) 41±7 (36–46) 40±2.9 (35–45) 37±2.3 (35–39)
a 28.5±0.2 (28.3–28.7) 25±1.3 (23.6–26.3) 24.4±1.5 (23.3–25.5) 22.6±1.2 (20.4–24.0) 21.8±0.4 (21.1–22.8)
b 3.4±0.1 (3.4–3.5) 3.5±0.2 (3.2–3.5) 3.5±0.2 (3.3–3.7) 3.2±0.2 (3.0–3.6) 3.3±0.1 (3.2–3.5)
312 · Zootaxa 3646 (4) © 2013 Magnolia Press

c 24.8±0.5 (24.3–25.3) 24±1.6 (22.3–25.6) 26.4±1.2 (25.5–27.3) 21.6±1.4 (19.1–23.3) 23.1±1.5 (22.2–24.9)
c' 1.7±0.1 (1.6–1.7) 1.7±0.2 (1.6–2) 1.3±0.2 (1.2–1.5) 1.5±0.1 (1.3–1.8) 1.4±0.1 (1.3–1.6)
V 57.5±0.1 (57.3–57.7) 57±0.8 (56.8–57.5) 58.9±2.1 (57.4–60.4) 58.7±0.9 (57.2–60.5) 59.3±0.9 (58.3–60.1)
G1 15.1±2.6 (12.5–17.8) 10.5±3 (7–12.5) 12.5±1.4 (11.5–13.5) 13.5±4 (8.2–14.6) 12.4±1.6 (11.2–14.3)
G2 14.7±3.0(11.6–17.7) 10.7±0.7 (10.2–11.6) 12.8±0.3 (12.4–13.2) 12.2±2.4 (9.4–14.6) 12.1±1.3 (11.2–14.3)
Lip height 8.5±0.5 (8–9) 6.6±1.2 (6–8) 6.5±2.1 (5–8) 9.1±1 (7–10) 6.6±0.5 (6–7)
Lip diameter 20.5±0.4 (20–21) 20.2±0.5 (20–21) 22±2.8 (20–24) 23.3±1 (21–25) 20±0 (20–20)
Stoma length 24.5±0.1 (24–25) 23±0 (23–23) 23.5±2.1 (22–25) 23.5±1.2 (21–25) 22.3±0.5 (22–23)
Stoma diameter 14.5±0.5 (14–15) 14±0 (14–14)1 14.5±0.7 (14–15) 14.2±0.7 (13–15) 12.6±0.5 (12–13)
Dorsal tooth position 6±0 (6–6) 5.3±0.5 (5–6) 6.5±0.7 (6–7) 6.2 ± 0.5 (6–7) 5.6±0.5 (5–6)
Pharynx length 313.5±23.2 (290–337) 266.1±19.4 (244–281) 274.1±53.7 (236–312) 269.9±16.4 (237–290) 245.2±19.0 (233–267)
Nerve ring from ant. end 95±5 (90–100) 86.6±5.7 (80–90) 85.9±14.1 (75–95) 86.7±5.5 (77–95) 75±8.6 (70–85)
Excretory pore from ant. end 12.5±9.5 (103–122) 96.6±6.5 (90–103) 96.1±9.8 (89–103) 102.1±6.9 (90–115) 83.3±10.2 (76–95)
Rectum length 20.4±1.5 (18–23) 17.3±1.5 (16–19) 19.5±1.4 (18–20) 22.4 ± 1.2 (20–24) 19.5±1 (18–20)
Anal body diameter 20.5±0.4 (20–21) 22±0 (22–22) 26±4.2 (23–29) 26.6±1.1 (24–28) 24.1±0.4 (24–25)
Tail length 43.5±1.3 (42–45) 40.0±4.3 (37–45) 37.5±12.4 (29–46) 41.1±3.6 (36–50) 35.6±3.7 (33–40)
Vulva–anus distance 417±38.3 (390–445) 372±51(320–422) 372±116.6 (290–455) 298±26.9 (245–337) 299±28.1 (267–320)
TAHSEEN ET AL.
FIGURE 4. Mylonchulus lacustris (Cobb in Cobb, 1915) Andrássy, 1958 female (Udaipur population). A–D: Anterior end
with buccal armature. E: Anterior end showing amphids. F: Pharyngeal region; G: Pharyngo-intestinal junction. H: Anterior
genital branch. I: Posterior genital branch. J: Vulval region. K, L: Posterior body region. (Scale bar = 10 µm).
FIGURE 5. Mylonchulus lacustris (Cobb in Cobb, 1915) Andrássy, 1958 female (Tanna population). A–D: Anterior end with
buccal armature. E, F: Pharyngo-intestinal junction. G, H: Anterior genital branch. I: Posterior genital branch. J: Intestinal
region. K, L: Posterior body region. (Scale bar = 10 µm).

FIGURE 6. Mylonchulus lacustris (Cobb in Cobb, 1915) Andrássy, 1958 (Ghazipur population). A–F: Anterior end with
buccal armature. G: Posterior pharyngeal region. H: Anterior genital branch. I: Posterior genital branch. J, K: Vulval region. L:
Posterior body region. (Scale bar = 10 µm).
Voucher specimens. Two females on slide Mylonchulus lacustris (Cobb in Cobb, 1915) Andrássy, 1958 no.
AL/1–2 deposited in the Nematode Collection, Department of Zoology, Aligarh Muslim University, Aligarh, Uttar
Pradesh, India.
Three females on slide Mylonchulus lacustris (Cobb in Cobb, 1915) Andrássy, 1958 no. Ghz/1-2 deposited in
the Nematode Collection, Department of Zoology, Aligarh Muslim University, Aligarh, Uttar Pradesh, India.
Two females on slide Mylonchulus lacustris (Cobb in Cobb, 1915) Andrássy, 1958 no. Ud/1-2 deposited in the
Nematode Collection, Department of Zoology, Aligarh Muslim University, Aligarh, Uttar Pradesh, India.
Salient characters. Small to medium-sized species with buccal cavity goblet-shaped, about 1.6–2.0 times as
long as wide; dorsal tooth large, situated in anterior half of buccal cavity; subventral wall usually bearing seven
transverse rows of denticles; subventral teeth prominent; female genital system amphidelphic; tail conoid with
slight ventral curvature, gradually tapering to a cylindroid part before ending into a rounded terminus.
Remarks. Justifying its name, the species M. lacustris has been widely reported from moist or limnic habitats
(Andrássy, 2006). The species comes close to M. minor. Mulvey (1961) differentiated them on the basis of the
shape and size of buccal cavity and the subventral tooth. Cobb (1917) redescribed and illustrated the species and
emphasized on the ventrally arcuate tail with three caudal glands arranged in tandem. The three M. lacustris
populations in the present study, showed a large, goblet-shaped buccal cavity usually with seven tranverse rows
(Fig. 4D, 6F) of rasping denticles apud Jairajpuri, 1970b. However, one specimen of Aligarh population possessed
five (Fig. 5C) rows (apud Andrássy 2006). In an unusual case a large denticle was observed in between the small
rasping denticles (Fig. 6D). The individuals of the Ghazipur and Udaipur populations possessed large and slender
ovaries, however, in one individual of Ghazipur population the posterior ovary was markedly reduced (Fig. 6I).
The arrangement of caudal glands was usually in tandem, however, a slight deviation from the pattern was
observed in a few specimens of the Udaipur population (Fig. 4L) very similar to that reported in a population from
Thailand apud Buangsuwon & Jensen (1966).
Mylonchulus obtusicaudatus (Daday, 1899) Andrássy, 1958

(Fig. 7)
=Mononchus obtusicaudatus Daday, 1899 (Cobb, 1916)
Description. Adult: Body medium-sized, ventrally arcuate with sharply curved ventro-posterior region.
Cuticle smooth 3–6 µm thick at different body regions. Body pores indistinct. Lip region offset, about 2.5–3.0
times as wide as high, lip sensilla slightly projecting above labial contour, almost indiscernible in some specimens.
Amphids cup-shaped, 4–5 µm across, located 9–14 µm from anterior end. Buccal cavity goblet-shaped, about 1.6–
1.8 times as long as wide with strongly sclerotized vertical and oblique plates. Dorsal wall bearing large, 11–12 µm
long and 7–8 µm wide, anteriorly-directed tooth, located at 8–9 µm from anterior end of buccal capsule or ca 76–
84% from its base. Each subventral wall with 6 transverse rows of prominent, rasp-like denticles: denticles of
anterior row regular and larger while those of posterior ones relatively smaller, irregularly arranged; one subventral
tooth present below the level of rasp-like denticles. Pharyngeal sleeve surrounding stoma at 1/3–1/4 of its length
from base. Pharynx cylindroid, muscular, 24–26% of body length. Outlets of pharyngeal glands, DO situated at ca
51–56%, SV1O1 and SV1O2 at ca 65–67%, SV2O1 and SV2O2 at ca 91–93% of pharyngeal length from anterior end.
Nerve ring encircling pharynx at 27–29% of pharyngeal length from anterior end. Excretory pore faintly visible
and about 31–35% of pharyngeal length from anterior end. Pharyngo-intestinal junction non-tuberculate, cardial
flap small conical to long tubular ranging from 17–25 µm in length. Intestine made of polygonal cells, intestinal
lumen wide. Rectum 0.6–0.9 times anal body diameter in length. Tail conoid, ventrally arcuate with rounded to
subclavate terminus. Caudal glands well developed, tandem in arrangement. Spinneret terminal in position.
Female: Reproductive system didelphic, amphidelphic with short, compact, reflexed ovaries. Oocytes
arranged in single tier except at distal end of ovary. Sphincter present at junction of oviduct and uterus. Vagina
about one-third of corresponding body diameter, provided with conspicuous, ovoid pars refringens.
Male: Not found.
Locality and habitat. Moist soil sample containing M. obtusicaudatus was collected from Keoladeo National
Park, Bharatpur, Rajasthan, India, at 27°13′N 77°29′E coordinates.
Voucher specimens. Five females on slide Mylonchulus obtusicaudatus (Daday, 1899) Andrássy, 1958, no.
8B /1-4 deposited in the Nematode Collection, Department of Zoology, Aligarh Muslim University, Aligarh, Uttar
Salient characters. Large-sized species with buccal cavity ca 1.6–1.8 times as long as wide; dorsal tooth
large, situated in anterior half of buccal cavity; each subventral wall bearing six transverse rows of denticles and a
relatively posterior tooth; female genital system amphidelphic with conspicuous, ovoid pars refringens; tail conoid,
ventrally curved with blunt terminus; caudal glands in tandem; spinneret terminal.

FIGURE 7. Mylonchulus obtusicaudatus (Daday, 1899) Andrássy, 1958, A–I: Anterior end with buccal armature. J: Anterior
end showing amphids. K–N: Pharyngo-intestinal junction. O–Q: Anterior genital branch. R, S: Vulval region. T–V: Posterior
body region. (Scale bar = 10 µm).
Remarks. The present population conforms well to M. obtusicaudatus (Daday, 1899) Andrássy, 1958 in
morphometric and morphological details. The buccal cavity showed intrapopulation variations in the shape ranging
from funnel-shaped to goblet-shaped. Usually six subventral rows of prominent rasping denticles present. The
subventral teeth in a few specimens appeared crooked in shape or the tip seemed to be broken off (Fig. 7H),
perhaps during the capture of prey. The cardial flaps also showed variation in shape and length (Fig. 7K–N) in
different individuals. In females the reproductive system generally comprised of two short compact ovaries,
oviducts demarcated from uterus by sphincters (Fig. 7O–Q) and pars refringens conspicuously rounded or ovoid
(Fig. 7R) in a few individuals. The obtuse or blunt tail tip, the signature feature of this species, showed some degree
of variation (Fig. 7T–V) from blunt to a more or less rounded tip.
Mylonchulus hawaiiensis (Cassidy, 1931) Andrássy, 1958

(Figs. 8, 9)
=Mononchus (Mylonchulus) hawaiiensis Cassidy, 1931 (Goodey, 1951)

=Mylonchulus bareilliensis Sharma & Saxena, 1980
=Mylonchulus indicus Sharma & Saxena, 1980
Description. Adult: Body medium-sized, ventrally curved upon fixation, more in posterior region. Cuticle
smooth 2–3 µm thick; body pores indistinct. Lip region slightly offset, about 2.3–2.8 times as wide as high with
labial papillae projecting above labial contour. Amphids cup-shaped with oval aperture, 4–5 µm across, located
10–12 µm from anterior end. Buccal cavity goblet-shaped with narrow base about 1.5–1.8 times as long as wide
with heavily sclerotised vertical and oblique walls. Dorsal wall having massive dorsal tooth directed anteriorly, 9–
10 × 6–6 µm in dimension, located at ca 8–9 µm from anterior end of buccal capsule or at ca 69–79 % of its base.
Two subventral denticulate walls, each bearing 6 transverse rows of denticles and a subventral tooth posterior to the
last row of denticles. Pharynx cylindroid, muscular, ca 27–30% of body length. Pharyngeal sleeve surrounding
stoma at 1/3–1/4 of its length from base. Outlets of pharyngeal glands: DO situated at ca 52–58%, SV1O1 and
SV1O2 at ca 66–69%; SV2O1 and SV2O2 at ca 87–94% of pharyngeal length from anterior end. Pharynx highly
muscular, cylindrical in shape with clear, thick lumen. Nerve ring located at ca 28–32% of pharyngeal length from
the anterior end. Excretory pore faintly visible, approximately at ca 36–37% of pharyngeal length from anterior
end. Pharyngo-intestinal junction non-tuberculate, with variable cardial flap length Fig. 9I-L. Intestinal lumen
narrow. Rectum length 0.6–0.9 times anal body diameter. Tail abruptly curved ventrally, assuming an ‘L’ shape,
often appearing digitate with slightly clavate tip. Caudal glands well developed, arranged in tandem, occasionally
grouped; spinneret terminal.
Female: Reproductive system didelphic, amphidelphic with reflexed ovaries each having 7–8 developing
oocytes arranged alternately in proximal region and multiple proliferating oocytes at distal end. Vagina about one-
fourth of corresponding body diameter long with well developed pars refringens. Vulva a transverse slit.
Male: Not found.
Locality and habitat: Dehradun population: The soil sample containing M. hawaiiensis was collected from a
ditch at Dehradun, Uttarakhand, India, at 30°20'42"N 77°59'50"E coordinates.
Keetham population: Soil sample containing Mylonchulus hawaiiensis was collected from the bank of
Keetham Lake, Agra, Uttar Pradesh, India.
Voucher specimens. Nine females on slide Mylonchulus hawaiiensis (Cassidy, 1931) Andrássy, 1958, no.
DJ29/1-2 deposited in the Nematode Collection, Department of Zoology, Aligarh Muslim University, Aligarh,
Uttar Pradesh, India. One female deposited at USDANC, Beltsville, MD, USA.
Two females on slide Mylonchulus hawaiiensis (Cassidy, 1931) Andrássy, 1958, no. Kee/1-2 deposited in the
Nematode Collection, Department of Zoology, Aligarh Muslim University, Aligarh, Uttar Pradesh, India. One
female deposited at USDANC, Beltsville, MD, USA.
Salient characters. Small to medium-sized species with buccal cavity funnel-shaped, about 1.5–1.8 times as
long as wide; dorsal tooth large, situated in anterior half of buccal cavity; each subventral wall bearing six
transverse rows of denticles and a relatively posterior subventral tooth; female genital system amphidelphic; tail
conoid, ventrally arcuate, bent at middle with clavate terminus; caudal glands usually arranged in tandem;
spinneret terminal.

FIGURE 8. Mylonchulus hawaiiensis (Cassidy, 1931) Andrássy, 1958, (Keetham population). A–F: Anterior end with buccal
armature. G, H: Pharyngo-intestinal junction. I: Posterior genital branch. J: Anterior genital branch. K, L: Posterior body
region. (Scale bar = 10 µm).
FIGURE 9. Mylonchulus hawaiiensis (Cassidy, 1931) Andrássy, 1958, (Dehradun population). A–G: Anterior end with buccal
armature. H: Anterior end showing amphids. I–L: Pharyngo-intestinal junction. M, N: Anterior genital branch. O, P: Vulval
region. Q, R: Posterior genital branch. S–W: Posterior body region. (Scale bar = 10 µm).

Remarks.The species M. hawaiiensis has been reported from India, Egypt, Niger, Kenya, South Africa, El
Salvador, Argentina, Hawaii, Abuachapan and Costa Rica. The species is widely distributed in India. The present
populations showed variations in the size of ovaries (Fig. 8I–J; 9M, N, Q, R), cardial flaps (Fig. 8G; 9I–L) and the
shape of tail (Fig. 8K, L; 9S–W); moreover, the spinneret also appeared terminal with a slight shift toward ventral
side in some individuals as also illustrated (Fig. 2C) by Jairajpuri (1970b). The variations in the body length, shape
of tail, and the position of amphids have also been reported by Mulvey (1967), and Jairajpuri & Khan (1982).
Earlier the species was synonymised by Mulvey (1961) with M. incurvus (Cobb, 1917) Andrássy, 1958 but he later
revalidated it. Bruin & Heyns (1992) also considered it a possible synonym of M. minor.
Mylonchulus vasis Yeates, 1992

(Fig. 10)
Description. Adult: Body ventrally curved, C-shaped upon fixation. Cuticle smooth under light microscope,
2–3 µm thick; body pores indistinct. Lip region slightly demarcated, broader than adjoining body about 2.5–2.6
times as wide as body width at the neck base. Labial papillae well developed, slightly protruding above the labial
contour. Amphids cup-shaped with oval-shaped aperture near the base of the lips. Buccal cavity funnel-shaped or
conical with narrow tapering base nearly half as wide as the diameter of buccal cavity anteriorly. Dorsal tooth
massive, 11 × 7 µm in dimension, anteriorly directed with tip located at ca 6–7 µm from anterior end of buccal
capsule or at ca 62–70% from its base. Subventral walls usually with four transverse rows of rasping denticles; first
row with regularly arranged denticles, others with more or less dispersed arrangement. One specimen (Fig. 10E)
with five transverse denticule rows. Subventral teeth posterior to the rows of denticles. Pharynx muscular,
cylindroid with thick, wide lumen. Orifices of the pharyngeal gland DO located at ca at 50–55%, SV1O1 and SV1O2
at ca 64–68% and SV2O1 and SV2O2 at ca 94–97% of pharyngeal length from the anterior body end. Nerve ring
surrounding pharynx at ca 28–32% of pharyngeal length from the anterior end. Excretory pore not discernible.
Pharyngo-intestinal junction non-tuberculate with cardial flap of variable lengths. Intestinal lumen wide, made of
large polygonal cells. Rectum 0.8–1.0 times anal body diameter. Tail short cylindrical with digitate posterior part
and rounded to slightly swollen, blunt terminus. Caudal glands well developed, tandem in arrangement, rarely
grouped; spinneret terminal.
Female: Genital system didelphic, amphidelphic; ovaries reflexed with conspicuous constriction between the
large proximal oocyte and the rest of the oocytes. Oviduct leading to a prominent crustaformeria. Uterus devoid of
any intra-uterine eggs or sperms in the present specimens. Vagina extending inward up to 1/4 of corresponding
body diameter. Vulva a transverse slit without protruded lips.
Male: Not found.
Locality and habitat. Soil sample containing M. vasis was collected from around the roots of wild trees from
district Sonebhadra, Uttar Pradesh, India, at 24°41'3"N 83°4'0"E coordinates.
Voucher specimens. Five females on slide Mylonchulus vasis Yeates, 1992 no. BS01/1-4 deposited in the
Nematode Collection, Department of Zoology, Aligarh Muslim University, Aligarh, Uttar Pradesh, India. One
female deposited at USDANC, Beltsville, MD, USA.
Salient characters. Small to medium-sized species with buccal cavity goblet-shaped, about 2 times as long as
wide; dorsal tooth large, situated in anterior half of buccal cavity; subventral wall bearing four transverse rows of
denticles; subventral teeth present; female genital system amphidelphic; tail conoid, ventrally arcuate with rounded
to blunt, slightly swollen terminus; caudal glands in tandem; spinneret terminal
Remarks. The species M. vasis was reported from New Caledonia Island close to Australia. The presence of
the species in the Indian subcontinent is slightly intriguing in view of the discontinuity of habitats and many
geographical barriers in between. Nevertheless, the features of the present population totally conformed to the
species. Besides similar morphometric characteristics, the main feature viz., four rows of subventral rasping
denticles was also observed in present specimens with the exception of one individual possessing five rows. The
other feature, shape of the tail also corresponded with the tail shape of M. vasis apud Yeates (1992).
FIGURE 10. Mylonchulus vasis Yeates, 1992. A–G: Anterior end with buccal armature. H: Anterior end showing amphids. I,
J: Pharyngo-intestinal junction. K: Female genital system. L: Anterior genital branch. M: Vulval region. N: Posterior genital
branch. O: Intestinal region. P–R: Posterior body region. (Scale bar = 10 µm).

Mylonchulus contractus Jairajpuri, 1970
(Fig. 11)
Description. Adult: Body small-sized, C-shaped upon fixation, males with greater posterior curvature than
females due to presence of copulatory musculature. Cuticle smooth, 1–2 µm thick at different body regions. Body
pores indistinct. Lip region offset, angular in shape and distinctly wider than the adjacent body. Amphids cup-
shaped with oval-shaped aperture located at level of dorsal tooth apex. Labial papillae prominent and slightly
raised. Buccal cavity funnel-shaped with thick cuticularised walls. Dorsal wall of buccal cavity bearing a large,
anteriorly directed dorsal tooth, 7–8 µm long × 5–6 µm wide, located at ca 4–5 um from anterior end of buccal
capsule or at ca 67–75% from its base. Each subventral wall bearing 5 transverse rows of closely spaced, small-
sized denticles and a small subventral tooth posterior to them. Pharynx highly muscular, cylindrical with thick
lumen. Orifices of pharyngeal glands: DO located at ca 59–63%, SV1O1 and SV1O2 at ca 70–75% and SV2O1 and
SV2O2 at ca 95–97% of pharyngeal length from anterior body end. Nerve ring surrounding pharynx at 31–35% of
its length from the anterior end. Excretory pore faintly visible at 34–38% of pharyngeal length. Pharyngo-intestinal
junction non-tuberculate; cardia flattened to slightly overhanging. Intestinal lumen with traces of dead nematodes.
Intestine with wide lumen, bordered by prominent and conspicuous microvilli (Fig. 11J, O). Intestine contracted in
the region of gonads but some individuals showed a partial overlap. Rectum approximately the length of anal body
diameter. Tail short and curved ventrally at right angle terminating gradually to a narrow terminus. Caudal glands
well developed, grouped, with their duct opening through a subdorsal spinneret.
Female: Reproductive system didelphic, amphidelphic with both branches on one side of intestine. In most
specimens one genital branch well developed while other one reduced or rudimentary. Oviduct provided with
distinct pars dilatata. Vagina with thick, petal-shaped pars refringens.
Male: Genital system diorchic with opposed testes measuring 64–66 µm. Spicules ventrally arcuate (angular)
with slightly broader manubrium; gubernaculum simple trough-shaped. Ventromedian supplements 10 in number,
regularly spaced or equidistant, starting within the spicular range.
Locality and habitat: Moss containing M. contractus was collected from the bark of trees from district
Sonebhadra, Uttar Pradesh, India, located at 24°33'3"N 83°4'1"E coordinates.
Voucher specimens. Nine females and one male on slide Mylonchulus contractus Jairajpuri, 1970 no. BS05/1-
4 deposited in the Nematode Collection, Department of Zoology, Aligarh Muslim University, Aligarh, Uttar
Salient characters. Small to medium-sized species with buccal cavity funnel-shaped with thick cuticularized
walls; dorsal tooth large, situated in anterior half of buccal cavity; subventral wall bearing five transverse rows of
denticles; subventral teeth present; female genital system amphidelphic; intestine contracted in the region of gonad;
tail conoid, ventrally arcuate, bent at right angle; caudal glands usually grouped; spinneret subdorsal.
Remarks. The present specimens conform well to M. contractus in morphometrics and morphological
features. The species has so far been reported from India, Dominica and Costa Rica. A few minor differences that
may be noted including the greater number (= 10) of ventro-median supplements in the single male specimen of the
present population whereas only 9 supplements were reported in M. contractus apud Khan & Jairajpuri (1979).
Furthermore, the female reproductive system indicated a transition between amphidelphic and monodelphic
condition with either of the branches in rudimentary form in most specimens.
Sporonchulus vagabundus Jairajpuri, 1971

(Figs. 12, 13)
Description. Adult: Body J-shaped upon fixation, tapering gradually towards posterior extremity with long
conoid tail. Cuticle smooth, 3–4 µm thick. Lateral chord about one-third of body width at mid body. Body pores
indistinct. Lip region slightly rounded to truncate, distinct and wider than adjoining body ca three times as wide as
high and about two-thirds of the corresponding body diameter. Amphids cup-shaped with slit-like aperture, about
8–9 µm from anterior end located slightly posterior to labial base. Buccal cavity with converging, arched walls,
FIGURE 11. Mylonchulus contractus Jairajpuri, 1970. A–G: Anterior end with buccal armature. H: Anterior end showing
amphids. I, J: Pharyngo-intestinal junction. K–N: Female genital system. O: Intestinal region. P: Vulval region. Q–S: Caudal
region (female). T, U: Caudal region (male) with genital papillae indicated by arrows. (Scale bar = 10 µm).

FIGURE 12. Sporonchulus vagabundus Jairajpuri, 1971 (Sonbhadra population). A, F–H: Anterior end with buccal armature.
B–E: Subventral wall. I: Anterior end showing amphids. J–L: Pharyngo-intestinal junction. M, N: Anterior genital branch. O:
Vulval region. P, Q: Posterior genital branch. R–U: Posterior body region. (Scale bar = 10 µm).
FIGURE 13. Sporonchulus vagabundus Jairajpuri, 1971 (Dehradun population). A–F: Anterior end with buccal armature. G,
H: Subventral wall. I, J: Pharyngo-intestinal junction. K: Anterior genital branch. L: Posterior genital branch. M: Posterior
body region. N: Tail tip (Scale bar = 10 µm).

1.4–1.6 times as wide as long. Dorsal wall bearing a medium-sized, 4 µm long and 5 µm wide tooth, with apex
straight or perpendicular to longitudinal body axis; its tip at about 64–70 % of buccal cavity from its base.
Subventral walls with 18–24 irregularly placed denticles arranged roughly in 4 vertical rows. Pharynx muscular,
cylindrical with thick lumen. Nerve ring encircling pharynx at ca 33–40% and a faintly visible excretory pore at ca
42–49% of pharyngeal length from anterior end. Orifices of the pharyngeal glands, DO located at ca 53–66%,
SV1O1 and SV1O2 at ca 70–76%, SV2O1 and SV2O2 at ca 95–97% of pharyngeal length. Pharyngo-intestinal junction
non-tuberculate. Rectum 0.8–1.0 times anal body diameter long. Tail long, conoid, ventrally arcuate, tapering
evenly to a narrow rounded terminus. Caudal glands well developed, arranged in tandem. Spinneret terminal,
occasionally sclerotised.
Female: Genital system didelphic, amphidelphic; short and compact with reflexed ovaries measuring 110–139
µm. Each ovary with a large proximal oocyte, 4–5 ocytes singly or alternately arranged with very small
proliferating ones near distal tip. Oviduct with distinct pars dilatata. No trace of sperms either in uterus or in pars
dilatata. Vagina thick-walled, about two-fifths of the corresponding body diameter long; pars refringens vaginae
trapezoid, conspicuous, 2 × 2 µm in size. Vulva a transverse slit; vulval papillae absent.
Male: Not found.
Locality and habitat: A soil sample containing S. vagabundus was collected from Bina coal field at
Sonebhadra, Uttar Pradesh, India. S. vagabundus was also recovered from a soil sample collected from a ditch at
Dehradun, Uttar Pradesh, India, at 30°19'22"N 78°1'20"E coordinates.
Voucher specimens. Five females on slide Sporonchulus vagabundus Jairajpuri, 1971 no. BS-05/1-2
Three females on slide Sporonchulus vagabundus Jairajpuri, 1971 no. DJ/1-2 deposited in the Nematode
Collection, Department of Zoology, Aligarh Muslim University, Aligarh, Uttar Pradesh, India. One female
deposited at USDANC, Beltsville, MD, USA.
Salient characters. Medium-sized species with buccal cavity having converging and arched walls, about 1.4–
1.6 times as long as wide; dorsal tooth medium-sized, situated in anterior half of buccal cavity; subventral walls
bearing four vertical rows with a total of 18–24 denticles; subventral teeth more or less of equal size; female genital
system amphidelphic; tail conoid, ventrally arcuate, gradually tapering to a truncate terminus; caudal glands
arranged in tandem; spinneret conspicuous, terminal, usually sclerotised.
Remarks. S. vagabundus was first described by Jairajpuri, 1971 from Rishikesh, Saharanpur, Bareilly and
Aurangabad localities of India. It was later recorded from several other Indian localities (Jairajpuri & Khan, 1982).
The present specimens conformed well to those described from other localities in India except for having smaller
body length (0.9–1.0 µm vs 1.2–1.8 µm). The spinneret also varied from conspicuous and well-sclerotised to non-
sclerotised (Fig. 12R–U; 13 M, N). A great degree of intraspecific variability was observed in the shape of the
buccal cavity and in the arrangement of teeth (Fig. 12A–H; 13C, D; F–H).
Sporonchulus ibitensis (Carvalho, 1951) Andrássy, 1958

(Fig. 14)
=Mononchus ibitiensis Carvalho, 1951

=Sporonchuloides ibitiensis (Carvalho, 1951) Mohandas & Prabhoo, 1982
Description. Adult: Body ventrally curved upon fixation. Cuticle smooth, 2–3 µm thick. Body pores
indistinct. Lip region rounded, almost continuous with adjoining body, more than 2.5 times as wide as high. Lips
amalgamated, labial sensillla papilliform, not raised above the body contour. Amphids cup-shaped with slit-like
aperture, 3–4 µm across, located at level of dorsal tooth apex. Buccal cavity barrel-shaped, tapering at base, about
1.8 times as long as wide with thick vertical and oblique plates. Dorsal wall bearing a medium-sized, dorsal tooth
of 6 × 4 µm dimension, located at 6 µm from anterior end of buccal capsule or at 70% from its base, opposed by
four longitudinal rows of 24 subventral teeth. Posterior-most pair occasionally larger and separated by a wider gap
from anterior ones. Pharyngeal sleeve surrounding buccal cavity at 1/3–1/4 of its length from base. Pharynx
cylindrical, muscular, 24% of body length. Orifices of pharyngeal glands: DO located at 58%, SV1O1 and SV1O2 at
69%, SV2O1 and SV2O2 at 96% of pharyngeal length from anterior end. Pharyngo-intestinal junction non-
tuberculate with cardial flaps of variable lengths. Nerve ring at about 35% of pharyngeal length. Excretory pore
slightly posterior to nerve ring, at about 40% of the pharynx length. Intestine with narrow lumen. Rectum 1.1 times
anal body diameter long. Tail conoid, slightly ventrally arcuate, tapering to a narrow rounded terminus. Caudal
glands weakly developed, arranged in tandem, spinneret terminal, short, narrow and relatively inconspicuous.
FIGURE 14. Sporonchulus ibitensis (Carvalho, 1951) Andrássy, 1958. A–D: Anterior end with buccal armature (posterior
subventral tooth indicated by arrow). E: Anterior end showing amphids. F: Pharyngo-intestinal junction. G: Anterior genital
branch (ovarian sac indicated by arrow). H: Vulval region. I: Posterior genital branch (ovarian sac indicated by arrow). J:
Posterior body region. (Scale bar = 10 µm).

FIGURE 15. SEM micrograph of anterior end of Mylonchulus lacustris (A, B) and M. minor (C). Reconstruction of buccal
cavity of Clarkus (D), Prionchulus (F), Mylonchulus (H), Sporonchulus (J), Actus (L) modified from Borgonie et al. (1995).
LM micrograph of Clarkus (E), Prionchulus (G), Mylonchulus (I), Sporonchulus (K), Actus (M). (Scale bar = 10 µm).
Female: Reproductive system didelphic, amphidelphic with reflexed ovaries. Ovary with a large proximal
oocyte and oocytes of increasingly smaller size arranged in two rows towards the distal end. Long ovarian sac
present associated with each ovary (Fig. 14 G, I). Oviduct attached subterminally to the ovary, without distinct pars
dilatata. Crustaformeria and uteri well developed. Vagina about one-third of corresponding body diameter with
well developed globular pars refringens. Vulva slightly posterior to mid-body with simple vulval lips.
Male: Not found.
Locality and habitat: Soil samples containing Sporonchulus ibitensis were collected from a field in Ghazipur
district, Uttar Pradeshat 25°34'46"N 83°34'24"E coordinates.
Voucher specimens. One female on slide Sporonchulus ibitensis (Carvalho, 1951) Andrássy, 1958 no. GZ/1
India.
Salient characters. Medium-sized species with buccal cavity barrel-shaped, about 1.7 times as long as wide;
dorsal tooth medium-sized, situated in anterior half of buccal cavity; subventral walls bearing two vertical rows of
total 24 denticles; posteriormost subventral teeth larger and separated by a wider gap from anterior teeth; female
genital system amphidelphic; pars refringens vaginae globular; tail conoid, ventrally arcuate gradually tapering to
a narrow, rounded terminus; caudal glands arranged in tandem; spinneret terminal, short, narrow and
inconspicuous.
Remarks. This species has been reported from India, Sri Lanka, Nigeria, Zaire, Brazil, Argentina, Malaysia,
and Cameroon (Mulvey, 1963; Jairajpuri & Khan, 1982). The only specimen reported here conforms to the species
S. ibitensis in morphological and morphometric values. However, a difference was noted in the dorsal tooth that
had a flattened anterior with apex directed forward [vs apex directed anteriad apud Fig. 4, Mulvey (1963)]. Besides
other differentiating features, the pars refringens vaginae were observed to be more or less globular in the present
specimen (Fig. 14H) compared to the trapezoid pars refringens vaginae found in the populations of S. vagabundus
(Fig. 12O; 13L).
Discussion
The order Mononchida Jairajpuri, 1969 is a widely distributed group of nematodes, with species found in soil and
fresh water (Cobb, 1917), at great depths in lakes, at high altitudes on mountains, and in few instances, from
marine waters (Yeates et al., 1994). The group comprises large-sized predators that are k-strategists with long life
cycles (Yeates, 1967) and life spans, and that are very sensitive to slight changes in the environment (Bongers,
1990). They occur in greater numbers in undisturbed soils compared to arable land although they are found nearly
in all kinds of soils and known to adapt themselves to varied climatic conditions. The individuals are usually
gonochoristic but occasionally hermaphroditic (Cobb, 1917; Arpin & Kilbertus, 1981).
The species belonging to Mononchoidea Chitwood, 1937 possess a more or less narrow, barrel-shaped buccal
cavity with a tapering base and a non-tuberculate pharyngo-intestinal junction. Jairajpuri & Khan (1982)
considered three families under this superfamily viz., Mononchidae Chitwood, 1937, Mylonchulidae Jairajpuri,
1969 and Cobbonchidae Jairajpuri, 1969 whereas Andrássy (1993) accepted only two families viz., Mononchidae
and Mylonchulidae. Mononchidae sensu Andrássy (2006) further included two subfamilies, Mononchinae
Chitwood, 1937 and Cobbonchinae Jairajpuri, 1969. He (l.c.) considered Mononchus Bastian, 1865, Clarkus,
Prionchulus, Actus Baqri & Jairajpuri, 1973 and Sporonchulus along with six other genera under Mononchinae
based on the moderately-sclerotised buccal cavity, the anteriorly placed, medium-sized dorsal tooth, the subventral
walls with or without armature along the vertical plane (a simple or denticulate ridge or assorted denticles in a
vertical row). Cobbonchinae Jairajpuri, 1969 on the other hand comprised those species that possessed buccal
cavities with equal-sized dorsal tooth and subventral teeth. Jairajpuri and Khan (1982) considered Mononchinae
and Prionchulinae Andrássy, 1976 as subfamilies of Mononchidae with the genera Mononchus, Clarkus,
Coomansus Jairajpuri & Khan, 1977 and Actus placed under the former and Prionchulus under the latter subfamily.
In a later revision, Ahmad & Jairajpuri (2010) considered Actus to be close to Sporonchulus in the subfamily
Sporonchulinae Jairajpuri, 1969 of Mylonchulidae. Having subventral ridge and the tail with reduced or no
spinneret or caudal glands, Clarkus, Prionchulus, Coomansus and Parkellus Jairajpuri, Tahseen & Choi, 2001 are
the closely related taxa. The hypothesis is further supported by 18S rDNA studies (Holterman et al., 2006; van

Megen et al., 2009, Meldal et al., 2007). Mononchus, although resembling the above taxa in having a similar type
of buccal cavity with a moderately-developed dorsal tooth, was placed close to them thus validating their position
in Mononchidae. The genus Actus possesses a buccal cavity similar to that of Clarkus (Fig. 15E, M) with a well-
developed dorsal tooth and vertically aligned subventral armature, and also has a tail with a relatively reduced or
inconspicuous spinneret and caudal glands. It can be considered closer to Prionchulus based on the vertically
aligned subventral teeth and the relatively long, conoid tail with reduced or inconspicuous spinneret. Its separate
identity according to Olia et al. (2008) and the similar features, therefore, argue for a position closer to
Mononchinae and Prionchulinae within Mononchidae rather than placement within Mylonchulidae, which very
much differs in the nature of the dorsal tooth and horizontally aligned secondary armature; and a tail with
conspicuous spinneret and caudal glands (Fig. 15). The genus Sporonchulus (four vertical rows of subventral
denticles) shows similarity with Actus (two vertical rows of subventral denticles) on the basis of buccal cavity
elements as well as the tail terminus and instead of forming a separate group along with Actus and Granonchulus
Andrássy, 1958 within Mylonchulidae, may be considered forming a group close to Prionchulinae within
Mononchidae. Similar placement was also proposed by Siddiqi (1984) and by Andrássy (2006), who considered
Sporonchulus to be placed with other genera of Mononchinae.
Mylonchulus, the type genus of Mylonchulidae, contains a vast array of species. These species occur in various
terrestrial and limnic habitats and reported from every continent. The most abundant species are stated to be M.
brachyuris (Butschli, 1873) Altherr, 1953 and M. sigmaturus (Cobb, 1917) Altherr, 1953 reported from 42 and 33
countries respectively (Andrássy, 2006). Ahmad & Jairajpuri (2010) listed fifty-eight valid species under the genus
while reporting the presence of above sixty species. However, Andrássy (2006) considered about seventy-nine
species to be valid. The main differentiating features for species, as considered by different workers, include body
size, buccal cavity dimension, size of dorsal tooth, its position, number of rows of transverse denticles on the
subventral walls, vagina structure, tail length and shape, arrangement of caudal glands and spinneret position. The
body size ranges from 0.5–2.9 and most of the morphometric values and ratios are similar in species of similar size.
The morphological features also show a high degree of variation and overlap. Hence the salient features of most
species overlap and lack uniqueness. Several groups of species can be identified sharing common characteristics
viz., M. amurus Khan & Jairajpuri, 1979, M. exacutus Jensen & Mulvey, 1968 , M. insolitus Andrássy, 1968 and M.
oceanicus Andrássy, 1986 lack subventral teeth whereas M. kaszabi Andrássy, 1967, M. sexcristatus
(Merzheevskaya, 1951) and M. vulvapapillatus Altherr in Altherr & Delamare Deboutteville, 1972 possess vulval
papillae. Most of the species possess 5–6 transverse rows of subventral denticles, however, M. brassicus Soni &
Nama, 1980 and M. parabrachyurus (Thorne, 1924) Andrássy, 1958, display 6–7 rows, and others, including M.
dentatus Jairajpuri, 1970b and M. apapillatus Khan & Jairajpuri, 1979 have 10–15 rows. M. vasis and M.
neocontractus Patil & Khan, 1982 possess the minimum number of rows (4), however, one specimen in the present
population of M. vasis possessed five transverse rows of denticles. Keeping in view the extensive variation in the
species selected in the present study, we found that in M. minor most specimens possessed 6 rows of denticles with
the exception of two specimens with five rows (Fig. 3E). However, tail shape varied widely among the specimens
(Fig. 3U–X) and appears to be an unreliable differentiating feature. M. lacustris populations possessed an elongate
buccal cavity with 7 rows of denticles in all but one specimen which possessed six transverse rows (Fig. 4C). There
is also a disparity in the statements of Andrássy (2006) and Jairajpuri (1970b) regarding the number of denticles in
M. lacustris: 5–6 and seven, respectively. The arrangement of caudal glands is usually linear or tandem but has also
been reported to be grouped (apud Buangsuwon & Jensen, 1966). Likewise, tail shape, a character which has been
widely relied upon by many workers, displays a high level of inconsistency in addition to the confusion arising due
to disparity in interpretations from scientist to scientist. For instance, M. minor, M. lacustris and M. hawaiiensis
have been described with similar tail shapes (Fig. 26 C, D and Fig. 27 I) apud Ahmad and Jairajpuri, 2010; Fig. 1B
apud Coetzee, 1966 and Fig. 29 apud Mulvey, 1961; Fig. 3C apud Zullini et al., 2002). The tail of M. lacustris as
illustrated by different workers (Jairajpuri, 1970b Fig.1B; Coetzee, 1966 Fig. 1B) shows a high degree of variation.
Coomans et al. (1995) also suggested a great resemblance of their specimens of M. polonicus with M. lacustris.
Due to great degree of variational overlap many species of Mylonchulus are likely to be synonymous as also
reflected by the variable positions of M. sigmaturus in the dendrogram apud Holterman et al. (2006). Therefore, a
revision of the family based on morphological features and further supported by molecular characters
(mitochondrial DNA and 28S rDNA) is probably necessary.
Acknowledgement
The funding provided by the Ministry of Environment and Forests and Department of Science and Technology,
New Delhi is gratefully acknowledged.
References
Ahmad, W. & Jairajpuri, M.S. (2010) Mononchida: The Predaceous Nematodes. Nematology Monographs and Perspectives,
Vol.7. Brill Leiden-Boston, The Netherlands, 298 pp.
Altherr, E. (1953) Nématodes du sol du Jura vaudois et français. I. Bulletin Societe vaudoise des Sciences Naturelles, 65, 429–
460.
Altherr, E. & Delamare Deboutteville, C.L. (1972) Nématodes interstitiels des eaux douces des États-Unis d’Amérique (États
de Washington, du Colorado et du Massachusetts) récoltés par Cl. Delamare Deboutteville. Annales de Spéléologie, 27,
683–760.
Andrássy, I. (1958) Über das System der Mononchiden (Mononchidae Chitwood, 1937; Nematoda). Annales Histoirico-
Naturalis Musei Nationalis Hungarici, 50, 151–171.
Andrássy, I. (1967) Ergebnisse der Zoologischen Forschungen von Dr. Z. Kaszab in der Mongolei. 92. Weitere
Bodennematoden aus den Jahren 1964 und 1965. Opuscula Zoologica Budapest, 6, 203–233.
Andrássy, I. (1968) The scientific results of the Hungarian soil zoological expedition to the Brazzaville-Congo. 31. Nematoden
aus grundwasser. Annales Universitatis Scientiarum Budapestensis de Rolando Eötvös nominatae Sectio Biologica, 9–10,
1–27.
Andrássy, I. (1976) Evolution as a basis for the systematization of nematodes. London, UK, Pitman Publishing, 288 pp.
Andrássy, I. (1985) On the genera Mononchus Bastian, 1865 and Prionchulus (Cobb, 1916) Wu and Hoeppli, 1929 (Nematoda :
Mononchidae). Opuscula Zoologica (Budapest), 21, 9–22.
Andrássy, I. (1986) Fifteen new nematode species from the southern hemisphere. Acta Zoologica Hungarica, 32, 1–33.
Andrássy, I. (1993) A taxonomic survey of the family Mononchidae (Nematoda). Acta Zoologica Hungarica, 39, 13–60.
Andrássy, I. (2006) Free-living nematodes of Hungary. II. (Nematoda errantia). Pedozoologica Hungarica, 4, 496pp.
Arpin, P. & Kilbertus, G. (1981) Ultrastructure du contenu digestif et de l’épithelium intestinal chez quelques nématodes
prédateurs (Mononchida) et bactériophages. Revue de Nematologie, 4, 131–143.
Arpin, P., Samsoen, L., Ponge, J.F. & Khan, S.H. (1984) Ecology and systematics of the Mononchid nematodes from wood and
grassland areas in wet temperate climate. II. The genus Prionchulus Cobb 1916. Revue de Nématologie, 7, 215–225.
Azmi, M.I. (1991) Mylonchulus sarmini n. sp. (Mononchida: Nematoda) from India. Current Nematology, 2, 193–194.
Baermann, G. (1917) Eine einfache Methode zur Auffindung von Ankylostomum (Nematoden) Larven in Erdproben.
Geneeskunding Tijdschrift voor Nederlandsch-Indië, 57, 131–137.
Baqri, S.R. & Jairajpuri, M.S. (1973) Studies on Mononchida. V. The mononchs of El Salvador with descriptions of two new
genera, Actus and Paracrassibucca. Nematologica, 19, 326–333.
Bastian, H.C. (1865) Monograph on the Anguilullidae, free nematoids, marine, land and freshwater; with descriptions of 100
new species. Transactions Of The Linnean Society of London, 25, 73–184.
http://dx.doi.org/10.1111/j.1096-3642.1865.tb00179.x
Bongers, T. (1990) The maturity index: an ecological measure of environmental disturbance based on nematode species
composition. Oecologia, 83, 14–19.
http://dx.doi.org/10.1007/BF00324627
Borgonie, G., Claeys, M. & Coomans, A.V. (1995) Ultrastructure of the intestine of the bacteriophagous nematodes
Caenorhabditis elegans, Panagrolaimus superbus and Acrobeloides maximus (Nematoda: Rhabditida). Fundamental and
Applied Nematology, 18, 123–133.
Bruin, S. & de Heyns, J. (1992) Mononchida (Nematoda) from Southern Africa: genera Mylonchulus (Cobb, 1916) Altherr,
1953 and Granonchulus Andrássy, 1958. Phytophylactica, 24, 169–193.
Buangsuwon, D.K. & Jensen, H.J. (1966) A taxonomic study of Mononchidae (Enoplida: Nemata) inhabiting cultivated areas
of Thailand. Nematologica, 12, 259–274.
http://dx.doi.org/10.1163/187529266X00671
Bütschli, O. (1873) Beitrage zur Kenntnis der freilebenden Nematoden. Nova Acta Academiae Caesareae Leipold-Carolinae
Nat Curios, 36, 1–24.
de Carvalho, J.C. (1951) Uma nova espécie de Mononchus (Nematoda, Mononchidae). Bragantia, 11, 51–54.
http://dx.doi.org/10.1590/S0006-87051951000100007
Cassidy, G.H. (1931) Some mononchs of Hawaii. Hawaii Planters’ Record, 35, 305–339.
Chitwood, B.G. (1937) A revised classification of Nematoda, In: Papers in Helminthology, 30 years Jubileum of K.I. Skrjabin.
Moscow, USSR, Nauka, pp. 67–79.
Choi, Y.E., Khan, Z. & Lee, S.M. (1999) Descriptions of new species of predatory nematodes (Mononchida) from Korea.
Korean Journal of Soil Zoology, 4, 89–100.

http://dx.doi.org/10.1163/187529260X00352
Clark, W.C. (1960) Redescription of Mononchus truncatus Bastian, M. papillatus Bastian and Prionchulus muscorum
(Dujardin) (Enoplida, Nematoda). Nematologica, 5, 184 –198.
Clark, W.C. (1961) The Mononchidae (Enoplida: Nematoda) of New Zealand IV. The genus Mylonchulus (Cobb, 1916)
Pennak, 1953. Nematologica, 6, 1–6.
http://dx.doi.org/10.1163/187529261X00216
Cobb, M.V. (1915) Some freshwater nematodes of Douglas Lake region of Michigen, USA. Transactions of the American
Microscopical Society, 34, 21–47.
http://dx.doi.org/10.2307/3221598
Cobb, N.A. (1893) Nematodes mostly Australian and Fijian. McLeay memorial volume. Linnean Society of New South Wales,
pp. 252–308.
Cobb, N.A. (1916) Notes on new genera and species of nematodes. Four subdivisions of Mononchus. Journal of Parasitology,
2, 195–196.
Cobb, N.A. (1917) The mononchs (Mononchus Bastian, 1865). A genus of free-living predatory nematodes. Soil Science, 3,
431–486.
http://dx.doi.org/10.1097/00010694-191705000-00004
Cobb, N.A. (1918) Filter-bed nemas: nematodes of the slow sand filter-beds of American cities. Contributions to a Society of
Nematology, 7, 189–212.
Coetzee, V. (1966) Species of the genera Granonchulus and Cobbonchus (Mononchidae) occuring in Southern Africa.
Nematologica, 12, 302–312.
http://dx.doi.org/10.1163/187529266X00725
Coomans, A., Rashid, F. & Heyns, J. (1995) On some predatory nematodes from the Okavango Delta, Botswana.
Hydrobiologia, 302, 119–131.
http://dx.doi.org/10.1007/BF00027037
von Daday, E. (1899) Új-guineai szabadon éló nematodok. Mathematikai és Természettudomanyi Értesitö Budapest, 17, 557–
572.
Dujardin, F. (1845) Histoire naturelle des helminthes ou vers intestinaux. Paris, France, Librairie Encyclopédique de Roret, 654
pp.
Elbadri, G.A.A., Khan, Z., Moon, I.S., Lee, D.W. & Choo, Y.H. (2008) Descriptions of three new species of soil nematodes
from Sudan. International Journal for Nematology, 18, 4–12.
Eroshenko, A.S. (1975) Ten new nematode species of the order Mononchida Jairajpuri, 1969 from coniferous forests of
Primorye. Institute of Biology and Pedology, Far-East Science Centre, USSR Academy of Sciences Proceedings, New
Series, 26, 152–169.
Fuchs, A.G. (1930) Neue an Borken-und Rüsselkäfer gebundene Nematoden, aus Moos und Walderde in Borken-und
Rüsselkäfergängen. Zoologische Jahrbücher Systematik, 59, 505–646.
Goodey, T. (1951) Soil and freshwater nematodes. London, UK, Methuen, 390 pp.
Holterman, M., Rybarczyk, K., van der Wurff, A., van den Elsen, S., van Megen, H., Mooyman, P., Bongers, T., Holovachov,
O., Bakker, J. & Helder, J. (2006) Phylum-wide analysis of SSU rDNA reveals deep phylogenetic relationships among
nematodes and accelerated evolution towards crown clades. Molecular Biology and Evolution, 23, 1792–1800.
http://dx.doi.org/10.1093/molbev/msl044
Jain, S.K., Saxena, R. & Sharma, R.K. (1993) Mylonchulus esculentus sp. n. (Nematoda: Mononchida) and occurrence of M.
lacustris in Rohilkhand Division, Uttar Pradesh, India. Annals of Plant Protection Sciences, 1, 85–88.
Jairajpuri, M.S. (1969) Studies on Mononchida of India. I. The genera Hadronchus, Iotonchus and Miconchus, and a revised
classification of Mononchida, new order. Nematologica, 15, 557–581.
http://dx.doi.org/10.1163/187529269X00894
Jairajpuri, M.S. (1970a) Studies on Mononchida of India. II. The genera Mononchus, Clarkus n. gen. and Prionchulus (Family
Mononchidae Chitwood, 1937). Nematologica, 16, 213–221.
http://dx.doi.org/10.1163/187529270X00225
Jairajpuri, M.S. (1970b) Studies on Mononchida of India. III. The genus Mylonchulus (Family Mylonchulidae Jairajpuri, 1969).
http://dx.doi.org/10.1163/187529270X00153
Jairajpuri, M.S. (1971) Studies on Mononchida of India. IV. The genera Sporonchulus, Bathyodontus and Oionchus
(Nematoda). Nematologica, 17, 407–412.
http://dx.doi.org/10.1163/187529271X00639
Jairajpuri, M.S. & Khan, W.U. (1977) Studies on Mononchida of India. IX. Further division of the genus Clarkus Jairajpuri,
1970 with the proposal of Coomansus n. gen. (Family Mononchidae Chitwood, 1937) and descriptions of two new species.
http://dx.doi.org/10.1163/187529277X00264
Jairajpuri, M.S. & Khan, W.U. (1982) Predatory nematodes (Mononchida) New Delhi, India, Associated publishing, 131pp.
Jairajpuri, M.S., Tahseen, Q. & Choi, Y.E. (2001b) Parkellus parkus gen. n., sp. n. and Miconchus koreanus sp. n.
(Mononchida), two new predaceous nematodes from Korea. International Journal of Nematology, 11, 98–103.
Jensen, H.J. & Mulvey, R.H. (1968) Predaceous nematodes (Mononchidae) of Oregon. Oregon Sate Monographs, Studies in
Zoology, 12, 57 pp.
Khan, Z., Choi, Y.E., Lee, S.M. & Choi, J.S. (2000) Descriptions of four new and an unknown species of predatory nematodes
(Mononchida) from Korea. Korean Journal of Soil Zoology, 5, 71–83.
Khan, W.U. & Jairajpuri, M.S. (1979) Studies on Mononchida of India XII. The genus Mylonchulus (Cobb, 1916) Altherr, 1953
with descriptions of three new species. Nematologica, 25, 406–418.
http://dx.doi.org/10.1163/187529279X00578
de Maeseneer, J. & D’ Herde, J. (1963) Métodes utilisées pour l’étude des anguillules libres du sol. Revue de Agriculture
Bureaux, 16, 441–447.
de Man, J.G. (1876) Onderzoekingen over vrij in de aarde levende Nematoden. Tijdschrift Nederlandsche dierkundige
Vereenigning, 2, 78–196.
van Megen, H., van den Elsen, S., Holterman, M., Karssen, G., Mooyman, P., Bongers, T., Holovachov, O., Bakker, J. & Helder,
J. (2009) A phylogenetic tree of nematodes based on about 1200 full length small subunit ribosomal DNA sequences.
http://dx.doi.org/10.1163/156854109X456862
Meldal, B.H.M., Debenham, N.J., De Ley, P., Tandingan, De Ley, I., Vanfleteren, J., Vierstraete, A., Bert, W., Borgonie, G.,
Moens, T., Tyler, P.A., Austen, M.C., Blaxter, M., Rogers, A.D. & Lambshead, P.J.D. (2007) An improved molecular
phylogeny of the Nematoda with special emphasis on marine taxa. Molecular Phylogenetics and Evolution, 42, 622–636.
http://dx.doi.org/10.1016/j.ympev.2006.08.025
Merzheevskaya, O.I. (1951) [Nematodes of peaty and mineral soils and their importance to agriculture.] Sbornik Nauchnykh
Trudov Akademiya Nauk Belorussko˘ı SSR, 2, 112–120.
Micoletzky, H. (1922) Die freilebende Erd-Nematoden. Archiv für Naturgreschichte, Abteilung A, 87, 1–650.
Mohandas, C. & Prabhoo, N.R. (1982) On a new genus, Sporonchuloides with notes on Sporonchulus (Cobb, 1917) Pennak,
1953 (Mononchida: Nematoda). Records of the Zoological Survey of India, 79, 359–362.
Mohilal, N. & Dhanachand, Ch. (1997) Three new species of mononchs (Nematoda: Mononchida). Indian Journal of
Mulvey, R.H. (1961) The Mononchidae: a family of predaceous nematodes I. Genus Mylonchulus (Enoplida: Mononchidae).
Canadian Journal of Zoology, 39, 665–696.
http://dx.doi.org/10.1139/z61-070
Mulvey, R.H. (1963) The Mononchidae: a family of predaceous nematodes V. Genera Sporonchulus, Granonchulus and
Prionchuloides n. gen. (Enoplida: Mononchidae). Canadian Journal of Zoology, 41, 763–774.
http://dx.doi.org/10.1139/z63-048
Mulvey, R.H. (1967) The Mononchidae: a family of predaceous nematodes VII. Genus Prionchulus (Nematoda:
Mononchidae). Canadian Journal of Zoology, 45, 941–953.
http://dx.doi.org/10.1139/z67-104
Mulvey, R.H. (1978) Predaceous nematodes of the family Mononchidae from the Mackenzie and Porcupine river systems and
Somerset Island, N.W.T., Canada. Canadian Journal of Zoology, 56, 1847–1868.
http://dx.doi.org/10.1139/z78-252
Olia, M., Ahmad, W., Araki, M., Minaka, N., Oba, H. & Okada, H. (2008) Actus salvadoricus Baqri & Jairajpuri (Mononchida:
Mylonchulidae) from Japan with comment on the phylogenetic position of the genus Actus based on 18S rDNA sequences.
Japanese Journal of Nematology, 38, 57–69.
http://dx.doi.org/10.3725/jjn.38.57
Patil, K.J. & Khan, E. (1982) Taxonomic studies on nematodes of Vidarbha region of Maharashtra, India IV. Sporonchulus
grandis sp. n. and Iotonchus shamimi sp. n. (Nematoda: Mononchida). Indian Journal of Nematology, 12, 161–166.
Pennak, R.W.P. (1953) Fresh-water invertebrates of the United States. New York, NY, USA, Ronald Press, 769 pp.
Saha, M., Lal, M. & Singh, M. (2004) Description of two new species of predatory nematodes belonging to Mylonchulidae and
Ironidae from Delhi, India. Annals of Plant Protection Sciences, 12, 171–176.
Saha, M., Lal, M. & Singh, M. (2006) Three new species of nematodes belonging to Mononchida Jairajpuri 1969, from
rhizosphere of mango in Western Uttar Pradesh, India. Annals of Plant Protection Sciences, 14, 177–183.
Schneider, W. (1939) Würmer oder Vermes, II. Fadenwünner oder Nematoden. I. Freilebende und Pflanzenparasitische
Nematoden. Die Tierwelt Deutschlands, 36, 1–260.
Seinhorst, W. (1959) A rapid method for the transfer of nematodes from fixative to anhydrous glycerin. Nematologica, 4, 67–
69.
http://dx.doi.org/10.1163/187529259X00381
Sharma, R.K. & Saxena, V. (1980) Two new species of the genus Mylonchulus (Cobb, 1916) Altherr, 1953 (Mononchida:
Nematoda) from north India. Instituto Lombardo Accademia di Scienze e Lettere, Rendiconti Classe di Scienze (B), 114,
18–26.
Siddiqi, M.R. (1984) Four new genera and four new species of mononchs (Nematoda). Pakistan Journal of Nematology, 2, 1–
13.
Soni, G.R. & Nama, H.S. (1980) On a new species of Mylonchulus (Cobb, 1916) Altherr, 1956 (Nematoda: Mylonchulidae).
Current Science, 49, 750–751.

Thorne, G. (1924) Utah nemas of the genus Mononchus. Transactions of the American Microscopical Society, 43, 157–171.
http://dx.doi.org/10.2307/3221665
Winiszewska, G & Susulovsky, A. (2003) Revision of the genus Prionchulus Cobb, 1916 (Nematoda: Mononchina). I.
Prionchulus muscorum (Dujardin, 1845) Cobb, 1916 and related species. Annales Zoologici, 53, 559–577.
Wu, H.W. & Hoeppli, R.J.C. (1929) Free living nematodes from Fookien and Chekiang. Archiv für Schiffs-und Tropen-
Hygienie, 33, 35–43.
Yeates, G.W. (1967) Studies on nematodes from dune sands. B. Oncholaimidae, Ironidae. Alaimidae and Mononchidae. New
Zealand Journal of Science, 10, 299–321.
Yeates, G.W. (1992) Nematodes from New Caledonia.I. Introduction and Mononchoidea. Fundamental and Applied
Yeates, G.W., Boag B. & Small, R.W. (1994) Species diversity and biogeography of Mononchoidea (Nematoda). Russian
Journal of Nematology, 2, 45–54.
Zell, H. (1985) Nematodes of a beechwood soil. III. Prionchulus muscorum (Nematoda: Mononchida). Carolinea, 42, 57–74.
Zullini, A., Loof, P.A.A. & Bongers, T. (2002) Free-living nematodes from nature reserves in Costa Rica. 2. Mononchina.
http://dx.doi.org/10.1163/156854102760082168

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Zootaxa 3646 (4): 301–335 ISSN 1175-5326 (print edition)

Descriptions of ten known species of the superfamily Mononchoidea

QUDSIA TAHSEEN1,3, MOHAMMAD ASIF1, MALKA MUSTAQIM1, SHIKHA AHLAWAT1

Accepted by P. Mullin: 25 Feb. 2013; published: 6 May 2013 301

Material and methods

302 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

Clarkus papillatus (Bastian, 1865) Jairajpuri, 1970

=Mononchus papillatus Bastian, 1865

304 · Zootaxa 3646 (4) © 2013 Magnolia Press

=Oncholaimus muscorum Dujardin, 1845

306 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

Mylonchulus minor (Cobb, 1893) Andrássy, 1958

=Mononchus minor Cobb, 1893

308 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

MONONCHOIDEA FROM INDIA

Zootaxa 3646 (4) © 2013 Magnolia Press ·

310 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

Mylonchulus lacustris (Cobb in Cobb, 1915) Andrássy, 1958

=Mononchus lacustris Cobb in Cobb, 1915

312 · Zootaxa 3646 (4) © 2013 Magnolia Press

314 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

Mylonchulus obtusicaudatus (Daday, 1899) Andrássy, 1958

=Mononchus obtusicaudatus Daday, 1899 (Cobb, 1916)

316 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

Mylonchulus hawaiiensis (Cassidy, 1931) Andrássy, 1958

=Mononchus (Mylonchulus) hawaiiensis Cassidy, 1931 (Goodey, 1951)

318 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

320 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

Mylonchulus vasis Yeates, 1992

322 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

Sporonchulus vagabundus Jairajpuri, 1971

324 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

326 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

Sporonchulus ibitensis (Carvalho, 1951) Andrássy, 1958

=Mononchus ibitiensis Carvalho, 1951

328 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

330 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

332 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

334 · Zootaxa 3646 (4) © 2013 Magnolia Press TAHSEEN ET AL.

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