Nematology, 2009, Vol.

11(4), 541-550

Nematodes of the order Rhabditida from India. Description of

Diplogasteroides uriceus sp. n. and Rhabditidoides papillatus sp. n.
(Diplogastrina)
Md M AHAMOOD and Irfan A HMAD ∗
Department of Zoology, Aligarh Muslim University, Aligarh 202 002, India
Received: 20 September 2007; revised: 22 September 2008
Accepted for publication: 22 September 2008

Summary – Two new species of diplogastrid nematodes from India are described and illustrated. Diplogasteroides uriceus sp. n.
has a medium-sized body, monoprodelphic reproductive system lacking a post-uterine sac, anteriorly directed muscular vagina and
vulva located far posterior; long, slender, arcuate spicules; gubernaculum with proximal thread-like flexible part and nine pairs of
genital papillae of which four pairs are precloacal and five pairs postcloacal. The new species resembles D. crassus but can be most
easily differentiated by the structure of the cheilorhabdions, shape and structure of the gubernaculum, arrangement of the genital
papillae and the complete absence of a bursa. The new species also resembles D. proprius but can be differentiated by stoma structure,
more anteriorly located vulva, longer spicules, gubernaculum shape and tail form. Rhabditidoides papillatus sp. n. is characterised
by the presence of three pairs of post-deirids in the lateral fields, posteriorly curved tubular stoma, amphidelphic female reproductive
system, strongly developed, arcuate spicules with bilobed capitulum and a slender gubernaculum provided with a sleeve distally and
anteriorly directed bifid ends proximally. It can be distinguished from R. stigmatus by the structure of the spicule head, gubernaculum
shape and arrangement of the genital papillae; from R. mastigurus by spicule structure and the gubernaculum shape; and from R.
quercophilus by a more slender female body, stoma structure, absence of a male bursa, spicule form and arrangement of the genital
papillae.
Keywords – free-living nematodes, morphology, morphometrics, new species, taxonomy.

De Man (1912) described Diplogasteroides de Man,
1912 with D. spengelii de Man, 1912 as its type species.
Andrássy (1984) accepted three species under the genus
Diplogasteroides and considered Rhabditolaimus Fuchs,
1915, Masseyus Paramonov, 1964 and Fuchsnema An-
drássy, 1984 as valid genera. Sudhaus (1995) clarified
that the type species of Rhabditolaimus was R. halleri
and synonymised Fuchsnema with Rhabditolaimus. Later,
Sudhaus and Fürst von Lieven (2003) accepted that the
type species of Rhabditolaimus was R. leucarti Fuchs,
1914 and not R. halleri Fuchs, 1915, as had been sug-
gested by Sudhaus (1995). Nevertheless, they (l.c.) con-
sidered Rhabditolaimus Fuchs, 1915, Pseudodiplogaster
Takaki, 1941, Dirhabdilaimus Paramonov & Turlygina,
1955, Diplogasteroides (subgenus Neodiplogasteroides
Rühm, 1956), Masseyus and Fuchsnema as synonyms of
Diplogasteroides, accepting 27 nominal species in the
process. However, Andrássy (2005) distinguished Fuch-
snema from Rhabditolaimus by the prodelphic gonad and
extreme posterior position of the vulva. We believe that all
monophyletic groups should be placed under one genus
and, hence, accept the classification proposed by Sudhaus
and Fürst von Lieven (2003).
Rahm (1928) proposed Rhabditidoides Rahm, 1928
for the species R. longicauda Rahm, 1928. Andrássy
(1984) accepted four species under the genus Rhabdi-
tidoides, viz., R. mastigurus (Steiner, 1943) Andrássy,
1984, R. longicauda, R. longispiculus (Meyl, 1954) An-
drássy, 1984 and R. stigmatus (Steiner, 1930) Andrássy,
1984. Sudhaus and Fürst von Lieven (2003) moved R.
longispiculus to Diplogasteroides, transferred Rhabdito-
laimus leptosoma Völk, 1950 and Diplogasteroides quer-
cophilus Heindl-Mengert, 1956 to Rhabditidoides and
listed six species under the genus. In the present paper
one new species of Diplogasteroides and one of Rhabditi-
doides are proposed.

∗ Corresponding author, e-mail: ahmadirfi@gmail.com

© Koninklijke Brill NV, Leiden, 2009 DOI:10.1163/138855409X12465362560476

Also available online - www.brill.nl/nemy 541
M. Mahamood & I. Ahmad
Materials and methods
Nematodes were extracted by sieving and decantation
and modified Baermann funnel techniques and fixed in
hot 4% formaldehyde solution. Following fixation they
were transferred to glycerin/alcohol (5 parts glycerin : 95
parts 30% alcohol) and dehydrated slowly over a period
of 3 weeks in a desiccator. Morphological observations
and drawings were made using an Olympus BX 50 DIC
microscope and photographs taken with an Olympus C
5050 digital camera.
Diplogasteroides uriceus* sp. n.
(Figs 1, 2)
M EASUREMENTS
See Table 1.
D ESCRIPTION
Female
Body tapering gradually anteriorly, more rapidly in
post-vulval region. Cuticle with fine transverse and lon-
gitudinal striations. Lateral fields with two lines. Am-
phidial apertures oval, at level of gymnostom. Lip region
5-7 μm diam., continuous with body contour. Lips six,
separate, labial papillae well developed. Stoma long, tubu-
lar. Cheilostom cuticularised somewhat barrel-shaped,
cheilorhabdial flaps protruding beyond labial contour.
Posteriorly, cheilorhabdions overlapping anterior part of
gymnostom. Gymnostom long, tubular, cuticularised,
walls anisotopic, dorsal wall shorter than subventrals.
Stegostom anisomorphic. Dorsal tooth supported by two
pointed smaller teeth at base. Subventral walls lacking
tooth, teeth or denticles. Procorpus muscular, median bulb
ovoid. Isthmus gradually expanding to form ovoid basal
bulb. Corpus 56-62% of total pharyngeal length. Nerve
ring encircling isthmus at midpoint. Hemizonid slightly
anterior or at level of nerve ring. Excretory pore lo-
cated posterior to hemizonid. Intestinal lumen with dark
granules. Reproductive system monoprodelphic, posterior
branch completely absent. Ovary reflexed, flexure on right
side of intestine, oocytes small, numerous, arranged in
multiple rows in germinal zone. Oviduct long, narrow,
gradually expanding posteriorly to form a spermatheca,
* The specific epithet is derived from the high content of uric
acid in the sample from which it was isolated.
542
mostly filled with sperm. Uterus with muscular and glan-
dular regions. Uterine lumen narrow in glandular region.
Vagina muscular, anteriorly directed with cuticularised lu-
men. Vulva located far posterior; vulval lips slightly or
very strongly protruding; posterior lip usually larger than
anterior. Vulva located 0.7-1.2 vulval body diam. from
anus. Phasmids ca one anal body diam. posterior to anus.
Rectum 1-2 anal body diam. long. Tail slender, elongate,
2.8-5.2 times vulva-anus distance long.
Male
Body smaller than female, strongly curved in posterior
region in fixed specimens. Anterior region similar to that
of female except for presence of four cephalic setae.
Shape and position of amphidial apertures as in female.
Testis single, ventral, proliferation zone reflexed on right-
hand side of body. Spicules slender, arcuate, 1.8-2.5
anal body diam. long. Gubernaculum 39-46% of spicule
length, slender, with sleeve in distal part. Proximally with
a thread-like appendage. Nine pairs of genital papillae
comprising four precloacal and five postcloacal pairs.
Genital papillae formula: v1, v2, (v3, v4)/ad, ph, (v5, v6,
v7), pd. Tail divisible into two parts, a conoid anterior part
and a narrower, elongate, posterior section.
T YPE HABITAT AND LOCALITY
Collected from chicken manure in Haldwani, Uttar
Pradesh, India.
T YPE MATERIAL
Holotype female, ten paratype females and ten paratype
males on slides Diplogasteroides uriceus sp. n./1-8; de-
posited in the nematode collection of the Department of
Zoology, Aligarh Muslim University, Aligarh, India.
D IAGNOSIS AND RELATIONSHIPS
Diplogasteroides uriceus sp. n. is characterised by a
medium-sized body; monoprodelphic reproductive sys-
tem lacking a post-uterine sac; anteriorly directed mus-
cular vagina and posteriorly located vulva; long, slender,
arcuate spicules; gubernaculum with proximal thread-like
flexible part and nine pairs of genital papillae, four pairs
of which are precloacal and five pairs postcloacal.
The new species resembles D. crassus Körner, 1954
in body posture, in having a monoprodelphic gonad,
absence of a post-uterine sac, in the general shape of
stoma and in general morphometric values. However, the
Nematology
 New species of Diplogastrina from India

Fig. 1. Diplogasteroides uriceus sp. n. A: Female (entire); B: Male (entire); C: Anterior end (lateral); D: Anterior end (dorsoventral);
E: Pharyngeal region; F: Female reproductive system (entire); G: Female anal region; H: Male posterior end showing retracted
filamentous part of gubernaculum; I: Male posterior end showing extended filamentous part of gubernaculum.

Vol. 11(4), 2009 543

M. Mahamood & I. Ahmad

Fig. 2. Diplogasteroides uriceus sp. n. A: Anterior end (dorsal view); B: Anterior end (lateral view); C: Vulva (not protruded); D:
Vulva (protruded); E: Male posterior end (arrow indicates straight filamentous part of gubernaculum); F: Male posterior end (arrow
indicates curved filamentous part of gubernaculum). (Scale bars: A-F = 20 μm.)

new species can be differentiated by the structure of
the cheilorhabdions (long and posteriorly overlapping the
gymnorhabdions vs small with a distinct space between
cheilorhabdions and gymnorhabdions), shape and struc-
ture of the gubernaculum (with a proximal flexible thread-
like process and distally with a band-like sleeve vs prox-
imal flexible thread absent and sleeve differently shaped)
and in the arrangement of the genital papillae (v2 ca half a
spicule length anterior to cloaca and (v3, v4) grouped to-
gether precloacally vs v2d only slightly anterior to cloaca
and v4 ad-cloacal). In addition, smaller differences oc-
cur in female body length (L = 0.68-0.86 mm vs 0.91-
1.14 mm), length of spicules (47-54 vs 33-44 μm), female
a ratio (17.7-23.2 vs 14.1-15.6), arrangement of genital
papillae (v3, v4 pre-cloacal, placed close to each other on
the subventral wall vs v3 lateral and pre-cloacal, v4 post-

544 Nematology
 New species of Diplogastrina from India

Table 1. Morphometrics of Diplogasteroides uriceus sp. n. and Rhabditidoides papillatus sp. n. Measurements are in μm and in the
form: mean ± s.d. (range).
Character Diplogasteroides uriceus sp. n. Rhabditidoides papillatus sp. n.
Female Male Female Male
Holotype Paratypes Paratypes Holotype Paratypes Paratypes
n – 10 10 – 8 10
L 834 777 ± 68 675 ± 40 783 856 ± 83 700 ± 66
(687-864) (623-732) (733-952) (627-861)
a 19.1 20.2 ± 1.5 22.7 ± 2.2 32.3 29.7 ± 3 29.7 ± 2.7
(17.7-23.2) (19.5-25.9) (26-35) (24.5-34.1)
b 5.9 5.6 ± 0.3 5 ± 0.2 5 5.5 ± 0.5 4.9 ± 0.3
(5.1-6.1) (4.6-5.4) (4.6-6.4) (4.5-5.8)
c 6.7 6.5 ± 1.1 10.2 ± 1.5 3.8 3.9 ± 0.4 4.1 ± 0.2
(5.3-9.6) (7.7-12.5) (3.5-4.4) (3.6-4.5)
c 6 8 ± 1.6 2.8 ± 0.7 15 13.7 ± 1.9 8.3 ± 1.2
(4.6-11.2) (2.2-4.5) (10.6-15.6) (6.6-9.7)
V 82.2 79.5 ± 2.2 – 42.3 42.5 ± 1.6 –
(76.3-84.4) (41.1-46.5)
Max. body diam. 44 38 ± 3 30 ± 3.5 24 29 ± 2 23 ± 2
(36-45) (28-35) (24-31) (21-29)
Length of stoma 14 15 ± 0.5 14 ± 0.5 17 18 ± 1 17 ± 1
(15-16) (13-15) (16-19) (16-18)
Pharynx 140 138 ± 8 133 ± 8 154 154 ± 10 139 ± 7
(126-151) (121-148) (138-170) (128-149)
Excretory pore from anterior end 113 112 ± 8 108 ± 7 117 123 ± 8 114 ± 7
(99-121) (96-114) (110-136) (102-123)
Hemizonid from anterior end 108 108 ± 7 102 ± 6 112 118 ± 8 109 ± 6
(98-118) (91-109) (106-131) (98-118)
Nerve ring from anterior end 109 106 ± 9 97 ± 6 108 110 ± 7 101 ± 4
(94-108) (87-108) (96-121) (92-107)
Median bulb length 18 18 ± 1 17 ± 1 20 20 ± 2 18 ± 1
(16-20) (15-19) (18-25) (16-19)
Basal bulb length 20.3 21 ± 2.5 19 ± 1 19 21 ± 2 18 ± 1
(19-25) (17-22) (19-23) (17-19)
Anterior gonad length 298 324 ± 40 – 99 116 ± 18 –
(258-367) (92-139)
Posterior gonad length – – – 94 113 ± 15 –
(90-138)
Vulval body diam. 35 33 ± 3.5 – 24 29 ± 2 –
(27-37) (24-32)
Vulva to anus distance 25 37 ± 6 – 247 276 ± 37 –
(21-45) (227-325)
Rectum 17 21 ± 3.5 21 ± 2 19 20 ± 2 15 ± 1
(17-27) (17-24) (17-22) (14-17)
Tail 123 121 ± 21 67 ± 10 204 216 ± 30 168 ± 17
(73-152) (55-87) (165-252) (147-199)
Anal/cloacal body diam. 20 15 ± 1 24 ± 2.5 14 16 ± 1 20 ± 1
(14-17) (19-26) (14-17) (17-24)
Phasmids from anus/cloaca 15 14.5 ± 1.5 12 ± 2 19 23 ± 2 24 ± 1
(13-17) (10-16) (19-25) (23-27)

Vol. 11(4), 2009 545

M. Mahamood & I. Ahmad
Table 1. (Continued).
Character Diplogasteroides uriceus sp. n.
Female
Holotype Paratypes
Testis – –
Spicule length (arc) – –
Gubernaculum length – –
cloacal) and in the form of the male tail (complete absence
of bursa vs presence of sub-terminal bursa).
The new species also resembles D. proprius (Andrássy,
1973) Sudhaus & Fürst von Lieven, 2003, which was also
collected from a chicken house, in a value, body length,
shape of spicules and arrangement of genital papillae.
However, the present species can be differentiated from
D. proprius by the structure of stoma (cheilostom over-
lapping gymnostom vs no overlap, dorsal tooth present on
a prominent bulge vs stegostomal bulge not prominent),
position of vulva (V = 76-84 vs 87-88), length of spicules
(47-54 vs 40 μm), shape of gubernaculum (with a distal
sleeve vs without) and in the shape and length of the fe-
male tail (elongate slender vs conoid, c = 5.3-9.6 vs 17-
18).
Rhabditidoides papillatus* sp. n.
(Figs 3, 4)
M EASUREMENTS
See Table 1.
D ESCRIPTION
Female
Body almost straight in fixed specimens, tapering to-
wards both extremities. Cuticle with fine transverse and
longitudinal striations. Cuticular surface divided into
blocks. Punctations present, arranged in both transverse
and longitudinal rows. Lateral fields with three lines.
Deirids at level of nerve ring or slightly posterior. Three
additional pairs of post-deirids present – first pair slightly
* The specific epithet refers to the presence of post-deirids in the
lateral fields.
546
Rhabditidoides papillatus sp. n.
Male Female Male
Paratypes Holotype Paratypes Paratypes
310 ± 25 – – 288 ± 31
(274-363) (245-347)
50 ± 2 – – 25 ± 1
(47-54) (23-27)
21 ± 1 – – 15 ± 1
(19-22) (13-17)
posterior to level of cardia, second pair far posterior to
vulva at level of ovary of posterior gonad and third slightly
anterior to level of rectum. Lip region continuous with
body contour. Lips fused, labial papillae small, protrud-
ing only slightly from labial contour. Amphidial aper-
tures oval, at junction of cheilostom and gymnostom.
Stoma tubular, longer than wide, ventro-dorsally curved
in posterior region. Cheilorhabdions weakly cuticularised,
cheilorhabdial flaps extending up to anterior end. Gym-
nostom anisotopic and anisomorphic, subventral walls
more strongly curved and longer than dorsal. Stegostom
anisomorphic, dorsal wall with large, ventrally directed,
flap-like tooth, surface weakly cuticularised. Subventral
walls plate-like, without armature. Procorpus muscular,
median bulb with large valve plates. Isthmus long, basal
bulb ellipsoidal. Corpus 60-66% of pharyngeal length.
Nerve ring encircling isthmus in mid-region or in ante-
rior half. Hemizonid at level of nerve ring with excretory
pore located slightly posterior to hemizonid. Intestinal
cells large, granular, lumen generally narrow. Reproduc-
tive system amphidelphic, anterior branch on right, pos-
terior on left side of intestine. Ovaries reflexed, oocytes
arranged in more than one row in germinal zone. Oviduct
narrow, spermatheca irregularly lobular in form, gener-
ally wider than uterus, offset. Uterus with glandular and
muscular parts. Vagina thick-walled, almost half vulval
body diam. Vulval opening circular, radially ridged. Rec-
tum 1.0-1.8 anal body diam. long. Phasmids situated 1.2-
1.8 anal body diam. from anus. Tail long, filiform, 0.6-0.9
times vulva-anus distance long.
Male
Body strongly curved in posterior region. Anterior
region similar to that of female except for four small
cephalic papillae. Testis ventral, flexure on right-hand
side of body. Spicules well developed, arcuate, capitulum
Nematology
 New species of Diplogastrina from India

Fig. 3. Rhabditidoides papillatus sp. n. A: Male (entire); B: Female (entire); C: Female showing position of post-deirids; D: Anterior end
(dorsoventral); E: Anterior end (lateral); F: Pharyngeal region; G: Female reproductive system (anterior tract); H: Female posterior
end; I: Male posterior end.

Vol. 11(4), 2009 547

M. Mahamood & I. Ahmad

Fig. 4. Rhabditidoides papillatus sp. n. A: Anterior end (lateral); B: Anterior end (dorsoventral); C: Lateral field (arrows indicate
post-deirids); D: Lateral lines; E: Female reproductive system (anterior tract); F: Vulva; G: Female posterior end; H: Male posterior
end. (Scale bars: A-C, E-H = 20 μm; D = 30 μm.)

548 Nematology

bilobed. Gubernaculum slender, proximal end anteriorly
directed, bifid, 46-61% of spicule length. Sleeve present at
distal end. Nine pairs of genital papillae comprising three
pairs precloacal, six pairs postcloacal. Genital papillae
formula: v1, v2, v3d/v4, (ad, v5, v6, ph), pd, v7. Genital
papillae v1, v2 within range of retracted spicules. GP v4
far posterior to cloaca. Phasmids situated posterior to v6
and v7 at junction of conoid and filamentous parts of tail.
Tail comprising two parts; a short conoid part and a long
filamentous section.
T YPE HABITAT AND LOCALITY
Farmyard manure from Gursai, Mendhar, Poonch,
Jammu and Kashmir, India.
T YPE MATERIAL
Holotype female, eight paratype females and ten para-
type males on slides Rhabditidoides papillatus sp. n./1-8;
deposited in the nematode collection of the Department of
Zoology, Aligarh Muslim University, Aligarh, India.
D IAGNOSIS AND RELATIONSHIPS
Rhabditidoides papillatus sp. n. is characterised by,
in addition to the deirids, the presence of three pairs of
post-deirids in the lateral fields, posteriorly curved tubu-
lar stoma, lobular uterus, arcuate spicules with bilobed ca-
pitulum, slender gubernaculum with a distal sleeve and
anteriorly directed bifid proximal ends and nine pairs of
genital papillae.
The new species resembles R. stigmatus in body mor-
phometrics, general shape of stoma and number of geni-
tal papillae. However, the new species can be differenti-
ated by the structure of spicule head (bilobed vs round),
shape of gubernaculum (narrow with anteriorly directed
proximal bifid ends vs broad with anteriorly curved fine
proximal end) and arrangement of the genital papillae
(two pairs vs three pairs precloacal; GP v4 far posterior
to cloaca vs GP v4 just posterior to cloaca). Post-deirids
have not been reported in R. stigmatus and therefore their
presence or absence needs to be confirmed.
The present species also resembles R. mastigurus in
morphometric values but can be differentiated from it
in the spicule structure (head bilobed vs round) and in
the shape of gubernaculum (proximally bifurcate vs not
bifurcate; distal sleeve present vs absent). The pharyngeal
collar shown in the diagram of Steiner (1943) is doubtful.
It further resembles R. quercophilus (Heindl-Mengert,
1956) Sudhaus & Fürst von Lieven, 2003 in body length
Vol. 11(4), 2009
New species of Diplogastrina from India
and morphometric values and in gubernaculum shape.
However, it can be differentiated by a more slender female
body (a = 26-35 vs 15), stoma structure (cheilostom and
gymnostom without any thickening vs both cheilostom
and gymnostom with a thickening), male tail lacking a
rudimentary bursa vs rudimentary bursa present, shape
and form of spicules (head bilobed vs round; blade slender
with a blunt distal tip vs blade broad proximally, slender
distally with a finely pointed tip) and in the arrangement
of the genital papillae (v2 and v3d far from each other vs
v2 and v3d nearly at same level).
Acknowledgement
Financial assistance from the Ministry of Environment
and Forests, Government of India, New Delhi through
AICOPTAX is gratefully acknowledged.
References
A NDRÁSSY, I. (1973). 100 Neue Nematodenarten in der un-
garischen Fauna. Opuscula Zoologica, Budapest 11, 7-48.
A NDRÁSSY, I. (1984). Klasse Nematoda. Stuttgart, Germany,
Gustav Fischer Verlag, 509 pp.
A NDRÁSSY, I. (2005). The free-living nematode fauna of
Hungary (Nematoda errantia), I. In: Csuzdi, Cs. & Mahunka,
S. (Eds). Pedozoologica Hungarica, 3. Budapest, Hungary,
Hungarian Natural History Museum and Systematic Zoology
Research Group of the Hungarian Academy of Sciences,
518 pp.
H EINDL -M ENGERT, H. (1956). Die Nematodenfauna im
Schleimfluß lebender Laubbäume. Sitzungsberichte der
Physikalisch-medizinischen Sozietät zu Erlangen 77,
158-176.
KÖRNER , H. (1954). Die Nematodenfauna des vergehenden
Holzes und ihre Beziehungen zu den Insekten. Zoologische
Jahrbücher (Systematik) 82, 245-353.
DE M AN , J.G. (1912). Helminthologische Beiträge. Zoologi-
sche Jahrbücher (Systematik) Supplement 15, 439-464.
M EYL , A.H. (1954). Die bisher in Italien gefundenen freileben-
den Erd- und Süßwasser-Nematoden. Archivio Zoologico
Italiano 39, 161-264.
R AHM , G.F. (1928). Alguns nematodes parasitas e semipara-
sitas das plantas culturaes do Brasil. Archivos do Instituto Bi-
ológico de Defesa Agricola e Animal (Sao Paulo) 1, 239-251.
S TEINER , G. (1930). The nemic fauna of the slime flux of
the Carolina poplar. Journal of Agricultural Research 41,
427-434.
S TEINER , G. (1943). New nematodes associated with a disease
of the papaya in Chile. Boletin Departamento de Sanidad
Vegetal (Santiago) 3, 95-116.
549
M. Mahamood & I. Ahmad

S UDHAUS , W. (1995). What is the type species of Rhabdito- VÖLK , J. (1950). Die Nematoden der Regenwürmer und aas-
laimus? Nematologica 41, 641. besuchenden Käfer. Zoologische Jahrbucher (Systematik) 79,
S UDHAUS , W. & F ÜRST VON L IEVEN , A. (2003). A phyloge- 1-70.
netic classification and catalogue of the Diplogastridae (Se-
cernentea, Nematoda). Journal of Nematode Morphology and
Systematics 6, 43-90.

550 Nematology