Handbook of Zoology

Founded by Willy Kkenthal

Editor-in-chief Andreas Schmidt-Rhaesa

Gastrotricha, Cycloneuralia and Gnathifera

Edited by Andreas Schmidt-Rhaesa

DE GRUYTER

Unauthenticated | 10.248.254.158
Gastrotricha, Cycloneuralia
and Gnathifera

Volume 2:
Nematoda

Edited by Andreas Schmidt-Rhaesa

DE GRUYTER

Unauthenticated | 10.248.254.158
Scientific Editor
Andreas Schmidt-Rhaesa
University Hamburg
Martin-Luther-King Platz 3
20146 Hamburg, Germany

Copyright 2014 by Walter de Gruyter GmbH, Berlin/Boston

ISBN 978-3-11-027375-5
e-ISBN 978-3-11-027425-7
Typesetting: Compuscript Ltd, Shannon, Ireland
ISSN 21934231
Printing and Binding: Hubert & Co. GmbH & Co. KG, Gttingen
Library of Congress Cataloging-in-Publication Data
A CIP catalogue record for this book is available from the Library of Printed in Germany
Congress. www.degruyter.com
Bibliographic information published by the Deutsche
National bibliothek

The Deutsche Nationalbibliothek lists this publication in the

Deutsche Nationalbibliografie; detailed bibliographic data are
available in the Internet at http://dnb.dnb.de.

Unauthenticated | 10.248.254.158
Fonseca, Gustavo; Bezerra, Tania Nara. 2014. Order Araeolaimida De Coninck, Schuurmans Stekhoven
1933. In: Schmidt-Rhaesa, A. (Ed), Handbook of Zoology. Vol. 2, Nematoda. De Gruyter, Hamburg,
pp.467486.
Gustavo Fonseca and Tania Nara Bezerra
7.15 Order Araeolaimida De Coninck &
Schuurmans Stekhoven, 1933
The order Araeolaimida has been classified based on
molecular grounds under the class Chromadorea (De
Ley & Blaxter 2004). This order contains five families:
Axonolaimidae, Bodonematidae, Comesomatidae, Con-
nickiidae and Diplopeltidae. In an earlier classification,
these families belonged to the superfamily Axonolaimoi-
dea under the order Monhysterida (Lorenzen 1981, 1994).
Although the family Bodonematidae is not mentioned in
previous classifications (Lorenzen 1981, 1994, De Ley &
Blaxter 2004), this family was placed in the superfamily
Axonolaimoidea by Jensen (1991a). Today the position
of the family Comesomatidae within the order Araeolai-
mida is questionable. A recent study based on the small
subunit ribosomal DNA (SSU rDNA) suggested that the
family Comesomatidae does not form a sister group with
Axonolaimidae, but belongs to the order Monhysterida
(Meldal etal. 2007). The relationships between monhys-
terids and comesomatids remain uncertain (De Ley &
Blaxter 2004, Meldal et al. 2007), and so far there is no
morphological character to support the monophyly of
both groups in a single clade. In the present chapter,
Comesomatidae will be treated as a sister group of
Axonolaimidae, as suggested in the last systematic
classification based solely on morphological charac-
ters (Lorenzen 1981, 1994). It is important to emphasize,
however, that this classification might change in the near
future. In this chapter, 47 genera and 397 species were
considered as valid in the order Araeolaimida. These
genera are mainly marine with only the genera Cylind-
rolaimus de Man, 1880 (Siddiqi 2004) and Edalonema
[(Holovachov & Sturhan 2003) Andrssy, 2007] and three
species of Parodontophora inhabiting limno-terrestrial
environments (Wu etal. 2000).
Diagnosis (emended from Lorenzen 1981, 1994):
There is no morphological apomorphy to establish the
monophyly of the order. The most important characte-
ristic is that the amphids are always loop-shaped, spiral,
bowl-shaped or pore-like, but this feature also occurs
within Linhomoidae. Additional features that distingu-
ish the order are: cuticle smooth or striated and, in most
of the Comesomatidae, covered by transverse rows of
punctuations. The labial sensilla are papilliform or not
visible. Outer labial and cephalic sensilla are separated
into two circles (six + four), though some exceptions
Brought to you by | Universidade Federal de So Paulo UNIFESP
occur in the family Comesomatidae. The cephalic setae
are generally longer than the outer labial. The buccal
cavity varies from minute in Connickiidae and Bodone-
matidae, to cup-shaped in some of the Comesomatidae
and tubular, equipped with odontia or teeth in some
members of Axonolaimidae and Comesomatidae. The
buccal cavity may have in some cases three, six or nine
tooth-like structures on its front edge. Pharyngeal tubes
occur frequently within the Axonolaimidae and Comeso-
matidae. The pharynx is usually cylindrical and seldom
widens at the end, forming a bulb. Females have two out-
stretched ovaries or, in some cases in the Axonolaimidae,
only one posterior. The males have two testes or only one
anterior. Ventrally precloacal papillae occur, above all, in
the Comesomatidae and only seldom in Axonolaimidae.
A diagnostic key for the five valid families is presented
in Fig. 7.129.
7.15.1Family Axonolaimidae
Filipjev, 1918
Diagnosis (emended from Lorenzen 1981, 1994, Tarjan &
Kuong 1988): Araeolaimida. There is no morphological
apomorphy to establish the monophyly of this family. The
six movable tooth-like structures located at the front in the
buccal cavity, referred as odontia, are very characteristic
of the family. However, this feature cannot be considered
an apomorphy of the family because it also occurs in some
genera of the Comesomatidae (e.g., Hopperia, Vasostoma).
The six odontia are also insignificant in the genera Axono-
laimus, Ascolaimus and Apodontium. All members of this
family have the buccal cavity cuticularized in a columnar
or conical form. Additional features are used to identify
the members of this family. Posterior to the buccal cavity
in the pharynx, there are one ventral and two subdorsal
pharyngeal tubes. The cuticle is smooth, in most cases,
and seldom striated. The six + four anterior sensilla stand
in two separate circles, the posterior four being larger
than the anterior. The amphids are mostly curved in a
loop-shape (inverted U- or O-shaped) and longitudinally
elongated or, in the case of Apodontium, pore-shaped.
The shape of the amphids may change with the life stage
468 7.15 Order Araeolaimida De Coninck & Schuurmans Stekhoven, 1933
Fig. 7.129: Diagnostic key for all valid families belonging to
Araeolaimida.
(Lorenzen 1973). For instance, juveniles stage I of the genus
Odontophora have amphids formed by one spiral loop with
a circular outline, whereas in later stages, the loop is com-
plete and longitudinally elongated. There are usually two
testes and two ovaries, but one gonad is also possible (e.g.,
Aequalodontium gemellum and in some species of Apodon-
tium). In most species, the anterior gonad lies to the left
or to the right of intestine, and the posterior gonad lies on
the side opposite to it. Spicules are curved and apophyses
are generally present. The tail is conical (except in Pseudo-
lella, which is conico-cylindrical). The caudal glands open
through a common pore. The family comprises 13 valid
genera and 116 valid species. A diagnostic key for all valid
genera is presented in Fig. 7.130.
7.15.1.1Genus Aequalodontium Smolyanko &
Belogurov, 1993 (Fig. 7.131)
Diagnosis (adapted from Smolyanko & Belogurov 1993):
Axonolaimidae. Cuticle finely striated. Inner and outer
labial sensilla papilliform. Cephalic setae 2.8 times the
corresponding body diameter. Four subcephalic setae of
the same length of the cephalic seta and adjacent to them.
Four pairs of somatic setae in one circle posterior to the
amphids. Loop-shaped amphids at the level of the base
of the stoma. Buccal cavity conical, equipped with six
pairs of tooth-like structures. Excretory pore anterior to
the nerve ring. One anterior testis outstretched. Spicules
of equal size and ventrally curved. Gubernaculum with
Brought to you by | Universidade Federal de So Paulo UNIFESP
dorso-caudal apophyses. Precloacal supplements absent.
Telamon paired lying above the tip of the spicules.
Monospecific: A. gemellum Smolyanko & Belogurov,
1993
7.15.1.2Genus Apodontium Cobb, 1920
(Fig. 7.131)
Diagnosis (emended from Tarjan & Khuong 1988,
Smolyanko & Belogurov 1993): Axonolaimidae. Cuticle
striated. Anterior sensilla arranged in six + six + four
pattern, with the cephalic setae usually being the
longest. Four subcephalic setae posterior to the amphids.
Amphids round, consisting of a single loop and mostly
indistinct. Buccal cavity funnel- or conical-shaped and
equipped with six odontia. Each odontia has the anterior
edge serrated into three smaller teeth. Pharynx cylindri-
cal without a bulb. Females monodelphic, opistodelphic.
Gonad outstretched. Males with a single outstretched
testis. Spicules hookless, curved, with poorly developed
capitulum. Telamon and supplements absent.
Number of species: 2
Type species: A. pacificum Cobb, 1920
7.15.1.3Genus Ascolaimus Ditlevsen, 1919
(Fig. 7.131)
Diagnosis (emended from Tarjan & Khuong 1988,
Warwick etal. 1988): Axonolaimidae. Cuticle finely stri-
ated. Amphids formed by one-loop spiral with a circular
outline; located behind the cephalic setae at the base of
the stoma. Somatic setae cover the body. Buccal cavity
conical without odontia. Pharynx cylindrical, slightly
swollen at the posterior end. Ovaries outstretched. Spicu-
les ventrally curved, usually with a hook at the posterior
tip. Gubernaculum with two apophysis oriented cau-
dally. Ascolaimus resembles Axonolaimus, except in the
rounded rather than elongated amphids, and a swollen
posterior section of the tail. However, several species
designated as Axonolaimus come close to this condition.
Number of species: 3
Type species: A. elongatus [(Btschli, 1874) De
Coninck & Schuurmans Stekhoven, 1932] Bussau, 1990
7.15.1.4Genus Axonolaimus De Man, 1889
(Fig. 7.131)
Diagnosis (emended from Tarjan & Khuong 1988, Warwick
et al. 1998): Axonolaimidae. Cuticle finely striated or
 7.15.1Family Axonolaimidae Filipjev, 1918 469

Fig. 7.130: Diagnostic key for all valid genera

belonging to the family Axonolaimidae.

smooth. Anterior end with four cephalic setae. Four sub- oriented caudally. Several postcloacal setae in the ventral
cephalic setae posterior to the amphids. Amphids loop- side.
shaped, longitudinally elongated, placed at the base of the Number of species: 34
stoma. Buccal cavity conical without odontia. Ovaries out- Type species: A. spinosus (Btschli, 1874) De Man,
stretched. Testis outstretched. Spicules arcuate. Apophyses 1889

Brought to you by | Universidade Federal de So Paulo UNIFESP

470 7.15 Order Araeolaimida De Coninck & Schuurmans Stekhoven, 1933

Fig. 7.131: Anterior and posterior end of all valid genera of the family Axonolaimidae. Drawings were modified from the following
publications: Aequalodotium gemellum (Smolyanko & Belogurov 1993); Apodontium pacificum (Cobb 1920); Axonolaimus spinosus
(Btschli 1874); Axonolaimus helgolandicus (Platt 1973); Ascolaimus elongatus (Gerlach 1953); Margonema ringens (Cobb 1920);
Namibnema papillata (Vincx & Furstenberg 1989); Nicascolaimus punctatus (Riemann 1986); Odontophora polynesea (Boucher 1973);
Odontophoroides monhystera (Boucher & Hellout 1977); Parascolaimus amphidoporus (Wieser 1959); Parodontophora xenotricha
(Boucher 1973); Pseudolella bengalensis (Timm 1963); Synodontium fecundum (Cobb 1920); and Synodontoides procerus (Gerlach 1957).

Brought to you by | Universidade Federal de So Paulo UNIFESP


7.15.1.5Genus Margonema Cobb, 1920
(Fig. 7.131)
Diagnosis (emended from Tarjan & Khuong 1988): Axo-
nolaimidae. Cuticle smooth, with few somatic setae.
Amphids circular to elliptical, mostly indistinct. Buccal
cavity conical, weakly sclerotized and without odontia.
Pharynx without terminal bulb. Female amphidelphic,
gonads outstretched. Testis single, outstretched. Spicu-
les L-shaped. Apophyses oriented dorsocaudally. Several
precloacal supplements present. Tail conoid, tapering
posteriorly.
Number of species: 2
Type species: M. ringens Cobb, 1920
7.15.1.6Genus Namibnema Vincx &
Furstenberg, 1989 (Fig. 7.131)
Diagnosis (emended from Vincx & Furstenberg 1989): Axo-
nolaimidae. Cuticle striated with punctuations between
the annules; lateral field provided with longitudinal rows
of more pronounced punctuations. Inner and outer labial
sensilla papilliform and hardly recognizable. Cephalic
sensilla setiform, 22.5 times longer than the correspon-
ding body diameter. Lateral post-amphidial setae present.
Amphids loop-shaped and longitudinally elongated.
Buccal cavity large and conical, composed of six plates
or odontia, ending in 12 small teeth. Pharynx muscular
ending in a pyriform terminal bulb. Ovaries didelphic,
anterior to the right and posterior to the left of intestine.
Anterior testis outstretched, situated to the left of intes-
tine, posterior testis with reflexed tip and situated to
the right of intestine. Spicules without a capitulum and
slightly ventrally bent. Dorsal apophyses present. Several
cup-shaped precloacal supplements present.
Monospecific: N. papillata Vincx & Furstenberg, 1989
7.15.1.7Genus Nicascolaimus Riemann,
1986 (Fig. 7.131)
Diagnosis (emended from Riemann 1986): Axonolaimi-
dae. Cuticle clearly striated, each ring having a distinct
transverse row of punctuations, the dots being more
widely spaced in the lateral region. Lateral differentiation
clearer in the most anterior and posterior regions. Large
ring-shaped cuticular pores are present in somewhat irre-
gular arrangement in sublateral and submedian rows in
the anterior region. Inner labial and outer labial sensilla
poorly recognizable. Cephalic setae twice as long the head
Brought to you by | Universidade Federal de So Paulo UNIFESP
7.15.1Family Axonolaimidae Filipjev, 1918 471
diameter. A pair of subcephalic setae in the lateral field
just posterior to the amphids. Somatic setae limited to the
anterior and caudal region. Amphids round, consisting of
one loop. Buccal cavity conical with six poorly developed
odontia. Pharyngeal marginal tubes located immediately
behind the stoma. Pharynx, slight swelling posteriorly.
Two outstretched testes. Spicules without a capitulum and
ventrally curved. Apophyses dorsal, placed perpendicular
to the spicules. Precloacal supplements pore-shaped.
Monospecific: N. punctatus Riemann, 1986
7.15.1.8Genus Odontophora Btschli,
1874 (Fig. 7.131)
Diagnosis (emended from Tarjan & Khuong 1988): Axo-
nolaimidae. Anterior end round. Cuticle smooth or finely
striated. Cephalic setae long, usually followed by adjacent
subcephalic setae. Amphids loop-shaped, longitudinally
elongated. Buccal cavity funnel-shaped, provided with six
odontia in the anterior portion. Pharynx gradually enlar-
ged posteriorly. Ovaries outstretched. Two testes. Spicules
arcuate. Apophyses present. Pore-like precloacal supple-
ments present or absent.
Number of species: 36
Type species: O. marina Btschli, 1874
7.15.1.9Genus Odontophoroides Boucher &
Hellout, 1977 (Fig. 7.131)
Diagnosis (emended from Boucher & Hellout 1977,
Lambshead 1982): Axonolaimidae. Similar to Synodon-
tium except for the clearly visible shepherd crook-shaped
amphids and prominent teeth in the buccal cavity. Buccal
cavity conical with six relatively large bicuspides odontia.
With or without lateral subcephalic setae. Pore of the
ventral gland posterior to the buccal cavity. One posterior
ovary. Spicules ventrally curved with a small capitulum.
Precloacal supplements setiform.
Number of species: 3
Type species: O. monhystera Boucher & Hellout,
1977
7.15.1.10Genus Parascolaimus Wieser,
1959 (Fig. 7.131)
Diagnosis (emended from Tarjan & Khuong 1988): Axono-
laimidae. Cuticle striated or smooth. Six labial claws,
probably representing modified labial papillae. Cervical
472 7.15 Order Araeolaimida De Coninck & Schuurmans Stekhoven, 1933
setae arranged in two longitudinal sublateral rows on
both sides of the body; somatic setae present. Amphids
with one loop; circular or oval. Buccal cavity conical.
Ovaries outstretched. Testis reflexed. Spicules curved.
Gubernaculum with apophyses oriented dorsally and
telamon. Supplements present or absent.
Number of species: 5
Type species: P. tau Wieser, 1959
7.15.1.11Genus Parodontophora Timm,
1963 (Fig. 7.131)
Diagnosis: Axonolaimidae. Similar to Odontophora except
that the buccal cavity is much deeper and cylindrical,
equipped with six large odontia. Cuticle smooth. Sub-
cephalic setae, when present relatively short. Amphids
loop-shaped with a longer ventral branch. Two gonads
outstretched. Precloacal supplements usually present.
Number of species: 21
Type species: P. paragranulifera Timm, 1963
7.15.1.12Genus Pseudolella Cobb, 1920
(Fig. 7.131)
Diagnosis (emended from Tarjan & Khuong 1988): Axono-
laimidae. Cuticle smooth. Head tapering rounded. Cepha-
lic setae representing one-fourth of the corresponding
body diameter. Amphids usually very long and shaped
like a shepherds crook. Stoma cylindrical with sclero-
tized walls flaring out, then arching sharply at the base.
Odontia present in the anterior part of the buccal cavity.
Pharynx with a posterior bulb. Ovaries outstretched.
Two testes. Spicules curved. Apophyses oriented dorso-
caudally. Tail conico-cylindrical.
Number of species: 8
Type species: P. granulifera Cobb, 1920
7.15.1.13Genus Synodontium Cobb, 1920
(Fig. 7.131)
Diagnosis (emended from Tarjan & Khuong 1988): Axo-
nolaimidae. Cuticle striated. Four cephalic setae and two
lateral subcephalic setae. Somatic setae present. Amphids
obscure consisting of a large internal pouch (shaped like
a shepherds crook) with a small circular aperture (just
a pore). Buccal cavity conical, equipped with six minute
bifid odontia. Pharynx enlarged posteriorly with a light
basal swelling. One opisthodelphic, outstretched ovary.
Brought to you by | Universidade Federal de So Paulo UNIFESP
Vulva before the mid-body region. Testes opposite and
outstretched. Spicules slightly curved. Apophyses present.
Supplements present.
Number of species: 4
Type species: S. fecundum Cobb, 1920
7.15.1.14Genus Synodontoides Hopper,
1963 (Fig. 7.131)
Diagnosis (emended from Tarjan & Khuong 1988): Axono-
laimidae. Similar to Apodontium except for the smooth or
finely striated cuticle, four cephalic setae and two subce-
phalic setae situated laterally at the posterior portion of
the buccal cavity. Six labial claws present. Amphids
obscure. Bucal cavity without odontia. Two ovaries and
two testes.
Monospecific: S. procerus (Gerlach, 1957) Hopper,
1963
7.15.2Family Bodonematidae
Diagnosis (emended from Jensen 1991a): Araeolaimida.
A characteristic pharynx differentiates into a long, tubular
anterior region with sclerotized walls and surrounded
by musculature, a mid-pharynx region developed into
bulbs with sclerotized inner walls, and a posterior region
without sclerotization, consisting of a narrow portion and
terminally with a pyriform bulb. Gonads opposite and out-
stretched. This family is monospecific.
7.15.2.1Genus Bodonema Jensen, 1991
(Fig. 7.132)
Diagnosis (emended from Jensen 1991a): Bodonematidae.
Body stout, ending in a short terminal tail. Cuticle smooth
without somatic setae. Anterior sensilla arranged in sepa-
rated circles (six + six + four) with 2 m long setae at the two
most posterior circles. Amphids ventrally wound in one
spiral forming a circle. Buccal cavity minute, continuing
in a long tubular portion with sclerotized walls. No teeth
observed. Mid-pharynx formed by four bulbs with strongly
sclerotized walls. Posterior part of the pharynx (posterior
to the nerve ring) narrow, terminating in a pyriform bulb
without sclerotized inner walls. Anteriormost portion of
intestine differentiated in four enlarged non-granular
cells (progaster). Ovaries opposite and outstretched, ante-
rior to the left of intestine, posterior to the right, or both
branches to the left. Testes opposite and outstretched,

Fig. 7.132: Anterior and posterior end of Bodonema vossi modified
from Jensen (1991a).
anterior branch to the left of intestine, posterior to the
right. Tail conical with a round tip.
Monospecific: B. vossi Jensen, 1991
7.15.3Family Comesomatidae
Filipjev, 1918
Diagnosis (emended from Jensen 1979, Platt 1985): Araeo-
laimida. Cuticule striated and/or ornamented with dots in
transverse rows, usually differentiated laterally, someti-
mes with weakly sclerotized body pores. Anterior sensilla
usually in three crowns, two crowns may also occur (e.g.,
Brought to you by | Universidade Federal de So Paulo UNIFESP
7.15.3Family Comesomatidae Filipjev, 1918 473
Laimella, Metacomesoma). Amphids multispiral. Buccal
cavity in two compartments with a globular, cup-shaped
or shallow anterior portion. Posterior portion of the buccal
cavity either a narrow, weakly sclerotized, collapsed tube
or a dilated, cylindrical to conical portion with sclero-
tized walls and projections at the borders between the
two portions. Pharynx with three marginal tubes. Pha-
ryngeal gland opening at the bottom of the buccal cavity.
Excretory pore behind the nerve ring. Female reproductive
system didelphic-amphidelphic with outstretched ovaries
(reflexed ovaries occur in one Hopperia species), sperma-
theca present. Male gonads diorchic with opposite and
outstretched testes. Copulatory apparatus with weakly to
strongly sclerotized spicules and gubernaculums; paired
apophyses of variable position. Precloacal supplements
usually tubular. Tail conico-cylindrical with tip slightly
dilated, nonsclerotized, symmetrical and with three ter-
minal setae. So far, there are 20 genera assigned to this
family and a total of 164 valid species. A diagnostic key for
all valid genera is presented in Fig. 7.133.
7.15.3.1Subfamily Comesomatinae
Filipjev, 1918
Diagnosis (emended from Jensen 1979): Comesomati-
dae. Cuticle with transverse annules or punctuations.
Lateral setae in second crown sometimes longer than the
other outer labial setae and when longer may be situated
between the second and third crowns. Posterior portion
of the buccal cavity cylindrical, dilated or collapsed, pro-
vided with projections at border to anterior portion. Spi-
cules long. Gubernaculum either tube-like with a solid
median piece or only consisting of the median piece;
when present, lateral guiding pieces small and solid. Apo-
physes slender, directly dorsal and parallel to spicules or
apparently lacking.
7.15.3.1.1Genus Comesoma Bastian, 1865 (Fig. 7.134)
Diagnosis (emended from Jensen 1979): Comesomatinae.
Anterior sensilla in three distinctly separated crowns,
outer labial lateral setae longer than the other four outer
labial setae. Subcephalic setae in one or more crowns,
each with four setae. Posterior portion of the buccal cavity
collapsed, rather weakly sclerotized and with three small
blunt projections at the border to anterior portion. Spicu-
les long and slender. Apophyses plate-like or indistinct.
Number of species: 11
Type species: C. vulgare Bastian, 1865
474 7.15 Order Araeolaimida De Coninck & Schuurmans Stekhoven, 1933

Fig. 7.133: Diagnostic key for all valid genera belonging to the family Comesomatidae.

Brought to you by | Universidade Federal de So Paulo UNIFESP

 7.15.3Family Comesomatidae Filipjev, 1918 475

Fig. 7.134 Part I: Anterior and posterior end of all valid genera of the family Comesomatidae. Drawings were modified from the
following publications: Comesoma bermudense (Jensen & Gerlach 1976); Comesomoides deconincki (Gourbault 1980); Metacomesoma
cyatholaimoides (Wieser 1954); Paracomesoma dubium (Kreis 1929); Paracomesoma inaequale (Jensen & Gerlach 1976); Asymmelaimus
vietnamicus (Dihn Tu etal. 2008); Dorylaimopsis punctata (Timm 1961); Expressonema granulate (Smolyanko & Belogurov 1991); Hopperia
indiana and Kenyanema monorchis (Muthumbi etal. 1997); Metasabatieria primigera (Timm 1961); Vasostoma spiratum (Wieser 1954);
anterior end and juvenile posterior end of Paramesonchium seriale (Wieser 1954); and male posterior end of Paracomesoma belgicum
(Jensen 1976).

Brought to you by | Universidade Federal de So Paulo UNIFESP

476 7.15 Order Araeolaimida De Coninck & Schuurmans Stekhoven, 1933

Fig. 7.134 Part II: Anterior and posterior end of Cervonema allemetricum (Wieser 1954); pharyngeal bulb of Cervonema papillatum (Jensen
1988); Laimella longicauda (Hopper 1967); Scholpaniella pontica (Sergeeva 1973); Minolaimus lineatus, Minolaimus cervoides, Pierrickia
decasetosa (Vitiello 1970); Actarjania splendens (Hopper 1967); Setosabatieria hilarula (Platt 1985); Sabatieira praedatrix (De Man 1907);
Sabatieria armata (Gerlach 1952); Sabatieria pulchra (Tchesunov 2000); Sabatieria celtica (Boucher 1973); and Sabatieria proabyssalis
(Vitiello & Boucher 1971).

Brought to you by | Universidade Federal de So Paulo UNIFESP


7.15.3.1.2 Genus Comesomoides Gourbault, 1980
(Fig. 7.134)
Diagnosis (emended from Gourbault 1980, Platt 1985):
Comesomatinae. Cuticle punctuated with lateral differen-
tiation of larger dots irregularly spaced. Anterior sensilla
separated in three circles with the cephalic setae longer
than the others. Amphids spiral with two turns. Anterior
portion of the buccal cavity cup-shaped, posterior portion
weakly sclerotized and tubular. Spicules elongated. Dorsal
apophyses present. Precloacal supplements present.
Monospecific: C. deconincki Gourbault, 1980
7.15.3.1.3 Genus Metacomesoma Wieser, 1954
(Fig. 7.134)
Diagnosis (emended from Jensen 1979): Comesomatinae.
Anterior sensilla in two crowns. Outer labial setae as long
as the cephalic setae. Posterior portion of the buccal cavity
collapsed and rather weakly sclerotized. Spicules long.
Gubernaculum tube-like, surrounding the distal part of
the spicules. Apophyses usually lacking.
Number of species: 2
Type species: M. cyatholaimoides Wieser, 1954
7.15.3.1.4 Genus Paracomesoma Hope & Murphy, 1972
(Fig. 7.134)
Diagnosis (emended from Jensen 1979): Comesomatinae.
Anterior sensilla in three distinctly separated crowns. Pos-
terior portion of the buccal cavity cylindrical, with strongly
sclerotized walls, and provided with three or six thorn-like
projections at border to anterior portion (six projections
in P. inaequale, each provided with a small subterminal
tooth). Spicules long. Gubernaculum plate-like without
dorsal apophysis.
Number of species: 8
Type species: P. dubium Filipjev, 1918
7.15.3.2Subfamily Dorylaimopsinae
De Coninck, 1965
Diagnosis (emended from Jensen 1979): Comesomatidae.
Cuticle usually differentiated laterally. Anterior sensilla
in three distinctly separated crowns. Buccal cavity stron-
gly sclerotized, anterior portion shallow, sometimes with
small pointed projections at border to the vestibulum,
posterior portion dilated, cylindrical or conical, with
strongly sclerotized walls and three thorn-like projections
Brought to you by | Universidade Federal de So Paulo UNIFESP
7.15.3Family Comesomatidae Filipjev, 1918 477
at the border between the two portions. Spicules
sometimes differentiated proximally, medially or distally.
Gubernaculum apophyses usually directed caudally,
occasionally dorsocaudally.
7.15.3.2.1 Genus Asymmelaimus Dihn Tu, Vu Than,
Smol & Vanreusel, 2008 (Fig. 7.134)
Diagnosis (emended from Dihn Tu et al. 2008): Dorylai-
mopsinae. Anterior end and buccal cavity asymmetrical.
Cuticle with transverse rows of punctuations; lateral dif-
ferentiation of one to two longitudinal rows of larger dots;
two rows in the pharyngeal region; one row in the mid-
body region. Amphids large and multispiral with 2 turns
Q. Buccal cavity shifted to the dorsal side; anterior part
cup-shaped with three minute teeth; posterior part tubular
with weakly sclerotized walls. Two outstretched gonads in
males and females. Spicules massive and curved. Apohy-
ses oriented caudally. Tail conico-cylindrical.
Monospecific: A. vietnamicus Dihn Tu, Vu Than,
Smol & Vanreusel, 2008
7.15.3.2.2 Genus Dorylaimopsis Ditlevsen, 1918
(Fig. 7.134)
Diagnosis (emended from Jensen 1979): Dorylaimopsinae.
Cuticle with transverse rows of punctuations, laterally
differentiated as a ridge with longitudinal rows of coarse
dots. Posterior portion of buccal cavity cylindrical, and
provided with three thorn-like projections at border to
anterior portion. Spicules slender, varying from sinusoid
to slightly bent or curved, usually differentiated. Apo-
physes directed caudally and parallel to the body axis or
dorsocaudally.
Number of species: 19
Type species: D. punctata Ditlevsen, 1918
7.15.3.2.3 Genus Expressonema1 Smolyanko &
Belogurov, 1991 (Fig. 7.134)
Diagnosis (emended from Smolyanko & Belogurov, 1991,
Hope & Zhang 1995): Dorylaimopsinae. Cuticle with trans-
verse rows of punctuations without lateral differentiation.
1The position of the genus Expressonema within the subfamily Dory-
laimopsinae is based mainly on the shape of the posterior portion of
the buccal cavity, although the shape of the spicules and the reduced
gubernaculum are similar to those belonging to Comesomatinae.
478 7.15 Order Araeolaimida De Coninck & Schuurmans Stekhoven, 1933
Buccal cavity conical, armed with nine odontia. Spicules
long, thin, slightly knobbed; gubernaculum reduced; apo-
physes dorsal (parallel to spicules) and small.
Monospecific: E. granulate Smolyanko & Belogurov,
1991
7.15.3.2.4 Genus Hopperia Vitiello, 1969 (Fig. 7.134)
Diagnosis (emended from Jensen 1979, 1992): Dorylaimop-
sinae. Cuticle punctuated laterally with transverse rows
of coarse dots. Posterior portion of buccal cavity cylind-
rical, provided with three thorn-like projections at border
to anterior portion. Reflexed ovaries may occur. Spicules
bent. Apophyses directed caudally or dorsocaudally.
Number of species: 13
Type species: H. massiliensis Vitiello, 1969
7.15.3.2.5 Genus Kenyanema Muthumbi, Soetaert &
Vincx, 1997 (Fig. 7.134)
Diagnosis (emended from Muthumbi et al. 1997): Dory-
laimopsinae. Cuticle punctuated in closely spaced
transverse rows, sometimes annules are visible. Lateral
differentiation with larger dots irregularly spaced. Inner
and outer labial sensilla indistinct. Cephalic sensilla
setiform. Amphids spiral with 1.52 turns. Buccal cavity
tubular without teeth. Ovaries amphidelphic and out-
stretched. Male reproductive system monorchic with
outstretched anterior testis. Spicules massive and curved
with a short central lamina. Gubernaculum with a long,
thin caudal apophyses. Tail short conical-cylindrical
with a swollen tip.
Monospecific: K. monorchis Muthumbi, Soetaert &
Vincx, 1997
7.15.3.2.6 Genus Metasabatiera Timm, 1961 (Fig. 7.134)
Diagnosis (emended from Jensen 1979): Dorylaimopsinae.
Cuticle undifferentiated laterally. Posterior portion of the
buccal cavity cylindrical, apparently unarmed. Spicules
bent. Apophyses directed dorsocaudally.
Monospecific: M. primigera Timm, 1961
7.15.3.2.7 Genus Paramesonchium Hopper, 1967
(Fig. 7.134)
Diagnosis (emended from Jensen 1979): Dorylaimopsinae.
Cuticle differentiated laterally with transverse or longitu-
Brought to you by | Universidade Federal de So Paulo UNIFESP
dinal rows of dots. Posterior portion of the buccal cavity
conical, provided with three ridges, each ending in an
acute projection at border to anterior portion. Spicules
bent. Apophyses directed dorsocaudally.
Number of species: 3
Type species: P. seriale Wieser, 1954
7.15.3.2.8 Genus Vasostoma Wieser, 1954 (Fig. 7.134)
Diagnosis (emended from Jensen 1979): Dorylaimopsi-
nae. Cuticle undifferentiated laterally. Posterior portion
of buccal cavity conical, provided with three small acute
projections at border to anterior portion. Spicules bent.
Apophyses directed dorsocaudally.
Monospecific: V. spiratum Wieser, 1954
7.15.3.3Subfamily Sabatieriinae
Filipjev, 1934
Diagnosis (emended from Jensen 1979): Comesomatidae.
Anterior sensilla in three separate crowns, second and
third crowns exceptionally very close together. Buccal
cavity rather weakly sclerotized; anterior portion glo-
bular to cup-shaped, sometimes very small; posterior
portion never strongly sclerotized and never cylindrical
or conical, but always a narrow collapsed tube. Spicules
bent, usually enlarged proximally. Apophyses usually
directed dorsocaudally and paired; if directed dorsally,
apophysis is single, small and rudimentary.
7.15.3.3.1 Genus Cervonema Wieser, 1954 (Fig. 7.134)
Diagnosis (emended from Jensen 1979): Sabatieriinae.
Cuticle finely striated. Anterior region narrow to elon-
gated. Outer labial and cephalic setae of equal length
and slender. Amphids with 4.257 turns, narrow spiral
and situated far behind the anterior end. Buccal cavity
minute. Posterior end of the pharynx formed by bundles
of muscles and pharyngeal glands and usually enlarged
but not forming a distinct muscular bulb. Copulatory
apparatus rather weakly sclerotized.
Number of species: 14
Type species: C. allemetricum Wieser, 1954
7.15.3.3.2 Genus Laimella Cobb, 1920 (Fig. 7.134)
Diagnosis (emended from Jensen 1979): Sabatieriinae.
Cuticle striated or with fine punctuations in closely

spaced transverse rows. Outer labial and cephalic setae
very close together. Buccal cavity poorly developed,
lumen of the anteriormost part of the pharynx distinctly
sclerotized. Tail conical-cylindrical with a flageliform pos-
terior portion.
Number of species: 7
Type species: L. longicauda Cobb, 1920
7.15.3.3.3 Genus Minolaimus2 Vitiello, 1970 (Fig. 7.134)
Diagnosis (emended from Hope & Zhang 1995): Saba-
tieriinae. Cuticle with transverse rows of punctuations
and lateral differentiation formed by longitudinal rows.
Cuticular pores might be present. Outer labial and
cephalic sensilla setiform of the same length, and nearly
at the same level on the head. Amphids multispiral with
at least six turns. Buccal cavity minute. Posterior end
of the pharynx formed by bundles of muscles and pha-
ryngeal glands and usually enlarged, but not forming
a distinct muscular bulb. Precloacal supplements cup-
shaped. Tail conical cylindrical, with the cylindrical
portion elongated.
Number of species: 2
Type species: M. lineatus Vitiello, 1970
7.15.3.3.4 Genus Pierrickia Vitiello, 1970 (Fig. 7.134)
Diagnosis (emended from Jensen 1979): Sabatieriinae.
Cuticle punctuate. Amphids with more than four spirals,
situated close to the front end. Outer labial setae and
cephalic setae equal in length. Buccal cavity very small.
Apophyses when present directed dorsocaudally (guber-
naculum and apophyses apparently lacking in P. aequalis).
Number of species: 4
Type species: P. decasetosa Vitiello, 1970
7.15.3.3.5 Genus Sabatieria Rouville, 1903
Diagnosis (emended from Jensen 1979): Sabatieriinae.
Cuticle striated or with transverse punctuation, lateral
differentiation as larger regular or irregular punctuations
may occur. Anterior sensilla arranged in three crowns.
Cephalic setae longer than the outer labial. Anterior
2This genus has been transferred from Cyatholaimidae to the
Comesomatidae by Hope and Zhang (1995) based on the pattern of
the cuticle, amphids, anterior sensilla, shape of the buccal cavity
and shape of the posterior end of the pharynx.
Brought to you by | Universidade Federal de So Paulo UNIFESP
7.15.3Family Comesomatidae Filipjev, 1918 479
portion of buccal cavity globular to cup-shaped, posterior
portion narrow as a collapsed tube, weakly sclerotized;
sometimes with small projections of the wall at the border
between the two portions. Male excretory system with two
additional uninucleate subventral gland cells far behind
the cardia. Spicules usually enlarged proximally, apophy-
ses usually directed dorsocaudally or caudally.
Number of species: 573
Type species: S. cettensis Rouville, 1903
According to Platt (1985) Sabatieria is separated into
five groups:
Praedatrix (Fig. 7.134): Sabatieria. Most species have
lateral cuticle differentiation of larger, more widely
spaced dots and amphids with three turns, but two or
four turns may also occur. Spicules without a central list
separating from the proximal projection. Simple tubular
or pore-like supplements and straight gubernaculum
apophyses.
Armata (Fig. 7.134): Sabatieria. Similar to Praedatrix
group except for the elongated cephalic (>1.7 head dia-
meters) and cervical sensilla, and usually slender bodies
(body length divided by maximum body width, more than
65). Amphids usually with three turns but four turns may
also occur. Simple tubular supplements.
Pulchra (Fig. 7.134): Sabatieria. Short paired cervical
setae. Among the genus, these setae appear to be shorter
and stouter. Amphids with three to four turns. Disposition
of the precloacal supplements are unique to this group.
Precloacal supplements conspicuous and relatively few in
number (five to nine). Gubernaculum with median pieces.
Celtica (Fig. 7.134): Sabatieria. Amphids with three
turns. Body shape relatively large and stout. All have
curved apophyses, conspicuous supplements.
Ornata (Fig. 7.134): Sabatieria. Amphids usually with
three turns. Similar to the Celtica group except for the pre-
sence of a posterior group of more closely situated prec-
loacal supplements.
7.15.3.3.6Genus Scholpaniella Sergeeva, 1973
(Fig. 7.134)
Diagnosis (emended from Jensen 1979): Sabatieriinae.
Cuticle with transverse rows of punctuations. Anterior
sensilla in three separated crowns; setae of the third crown
longer than those of the second crown. Spicules bent, pro-
ximally enlarged, distally tapering. Gubernaculum and
3The total number of Sabatieria species was based on the last
review made by Platt (1985) and all the articles publish thereafter.
480 7.15 Order Araeolaimida De Coninck & Schuurmans Stekhoven, 1933
apophyses apparently reduced to small sclerotized pieces
with the apophyses directed dorsally. Tail short, with the
dorsal margin curved.
Monospecific: S. pontica Sergeeva, 1973
7.15.3.3.7 Genus Actarjania Hopper, 1967 (Fig. 7.134)
Diagnosis (emended from Platt 1985): Sabatierinae.
Similar to Sabatieria except for the longitudinal rows of
lateral punctuations and sublateral rows of cervical setae.
Monospecific: A. splendens Hopper, 1967
7.15.3.3.8 Genus Setosabatieria Platt, 1985 (Fig. 7.134)
Diagnosis (emended from Platt 1985): Sabatierinae.
Similar to Actarjania, except for a lack of cuticular punc-
tuations and the presence of sublateral rows of long cer-
vical setae.
Number of species: 5
Type species: S. hilarula (De Man, 1922) Platt, 1985
Brought to you by | Universidade Federal de So Paulo UNIFESP
7.15.4Family Coninckiidae
Lorenzen, 1981
Diagnosis (emended from Lorenzen 1981, 1994, Tchesu-
nov 2000): Araeolaimida. No apomorphy can be cited
to establish the monophyly of the family. Nevertheless,
the combination of curved amphids into an O-shape,
the equal length of the outer labial and cephalic setae
and a tiny buccal cavity are unique within the order.
This family is also recognized by the smooth or striated
cuticle. Widening of the pharynx at the posterior end.
Two outstretched ovaries and two testes facing opposite
directions, anterior gonad lies to the left of intestine and
the posterior to the right.
7.15.4.1Genus Coninckia Gerlach, 1956
(Fig. 7.135)
Diagnosis: Coninckiidae. Anterior sensilla arranged
in three circles. Inner labial sensilla papilliform or
Fig. 7.135: Anterior and posterior end of
C. macroseta (modified from Tchesunov
2000) and C. seta (modified from Wieser &
Hooper 1967).
 7.15.5Family Diplopeltidae Filipjev, 1918 481

setiform, outer labial and cephalic setiform. Amphids 7.15.5.1Genus Araeolaimus De Man,
with one or two turns relatively large (mostly as large 1888 (Fig. 7.137)
as the corresponding body diameter) situated on
punctuated plaques. Somatic setae present or absent. Diagnosis (emended from Warwick et al. 1988,
Buccal cavity minute and sometimes difficult to reco- Tchesunov & Miljutina 2008): Diplopeltidae. Cuticle
gnize. Spicules ventrally curved. Gubernaculum with smooth or finely striated. Amphids loop-shaped with a
medium pieces and posterior apophyses. Tail conical circular or slightly elongated outline. Buccal cavity ter-
cylindrical with a dilated tip. Sexual dimorphism minal, narrow and conical. Pharynx with a bulbus swel-
usually present. ling midway along its length. Ocelli usually present. Two
Number of species: 5 ovaries. Spicules curved and apophyses directed dorsally.
Type species: C. circularis Gerlach, 1956 Tail conical.
Number of species: 34
Type species: A. bioculatus De Man, 1876
7.15.5Family Diplopeltidae
Filipjev, 1918 7.15.5.2Genus Cylindrolaimus De Man,
It has been suggested that this family be split into two 1880 (Fig. 7.137)
subfamilies, Diplopeltinae and Cylindrolaiminae, with
Diagnosis (emended from Siddiqi 2004): Diplopeltidae.
the second formed by the only genus Cylindrolaimus.
Cuticle striated. Amphids form a spiral with one turn.
This separation has been proposed chiefly on the shape
Buccal cavity long and tubular, always terminal. Ovaries
of the buccal cavity (cylindrical versus not cylindrical)
outstretched with the anterior gonad usually to the right
(Lorenzen 1981, 1994). However, a cylindrical buccal
side of intestine and posterior to the left. Testis single out-
cavity is also present in Southerniella, Intasia, Meta-
stretched. One precloacal supplement usually present.
raeolaimoides and Edalonema. Another diagnostic
Tail conical.
characteristic of the subfamily Cylindrolaiminae is
Number of species: 22
the presence of one-loop amphids (Lorenzen 1981,
Type species: C. communis (De Man, 1880) Gerlach &
1994), though this characteristic also occurs in several
Riemann, 1973
members of Diplopeltinae (see diagnosis below). There-
fore, in the present chapter, both subfamilies were con-
sidered synonyms.
7.15.5.3Genus Campylaimus Cobb, 1920
Diagnosis (emended from Lorenzen 1981, 1994):
Araeolaimida. There is no apomorphy to support the (Fig. 7.137)
monophyly of this group. Cuticle may be smooth or stri-
Diagnosis (emended from Warwick et al. 1988,
ated. Labial sensilla usually papilliform or not visible.
Tchesunov & Miljutina 2008): Diplopeltidae. Cuticle stria-
Four cephalic sensilla always predominant. Amphids
ted. Cephalic sensilla short or poorly recognizable. Buccal
curve to a bowl shape and consist of a spiral with one
cavity small, displaced dorsally and weakly sclerotized.
loop. Ocelli occasionally present. Buccal cavity narrow
Amphids loop-shaped with the ventral limb extending
and tubular or very small. Tooth-like formation absent.
down the body to the posterior end. Ocelli absent. Two
Pharynx has one ventral and two subdorsal pharyngeal
ovaries. Apophyses plate-like or weak. Tail conical or
tubes. Bulb might be present. Ovaries outstretched (only
conical-cylindrical.
one anterior in Cylindrolaimus monhystera). There are
Number of species: 9
usually two testes. Posterior testis is either anteriorly
Type species: C. inaequalis Cobb, 1920
oriented and reflexed in the anterior section or is pos-
teriorly directed. Precloacal supplements mostly absent
(present in Intasia). Tail conical or conico-cylindrical.
Caudal glands opening together or separately. With the
7.15.5.4Genus Diplopeltis Cobb in Stiles &
exception of Cylindrolaimus and Edalonema, all genera Hassal, 1905 (Fig. 7.137)
are marine. In total, there are 12 genera and 111 valid
species. Diagnostic key for all valid genera is presented Diagnosis (emended from Vincx & Gourbault 1992,
in Fig. 7.136. Warwick et al. 1988): Diplopeltidae. Cuticle smooth

Brought to you by | Universidade Federal de So Paulo UNIFESP

482 7.15 Order Araeolaimida De Coninck & Schuurmans Stekhoven, 1933

Fig. 7.136: Diagnostic key for all

valid genera belonging to the family
Diplopeltidae.

or finely striated. Amphids large and oval lying on a throughout. Two ovaries or only one anterior. Spicules
well-developed cuticular plate that has a double border. thin and ventrally curved. Tail conical.
Four longitudinal rows of cervical setae. Ocelli present. Number of species: 6
Buccal cavity minute and terminal. Pharynx narrow Type species: D. cirrhatus (Eberth, 1863) Cobb, 1891

Brought to you by | Universidade Federal de So Paulo UNIFESP

 7.15.5Family Diplopeltidae Filipjev, 1918 483

Fig. 7.137: Anterior and posterior end of all valid genera of the family Diplopeltidae. Drawings were modified from the following
publications: Cylindrolaimus scleris (Siddiqi 2004); Edalonema camerunense (Holovachov & Sturhan, 2003); Araeolaimus elegans (Gerlach
1953); Campylaimus gerlachi (Timm 1961); Diplopeltis cirrhatus (Gerlach 1962); Diplopeltula brevicepis and Diplopeltula incise (Gerlach
1950); Intasia monohystera (Tchesunov & Miljutina 2008); Metaraeolaimoides oxystomina (Luc & De Coninck 1959); Morlaxia contuse; and
Morlaxia obscura (Vincx & Gourbault, 1988); anterior end from Paraeolaimus nudus (Lorenzen 1973) and posterior end (Gerlach 1951);
Pseudoaraeolaimus perplexus (Chitwood 1951); and Southerniella allometrica (Vitiello 1971).

Brought to you by | Universidade Federal de So Paulo UNIFESP

484 7.15 Order Araeolaimida De Coninck & Schuurmans Stekhoven, 1933
7.15.5.5Genus Diplopeltula Gerlach, 1950
(Fig. 7.137)
Diagnosis (emended from Vincx & Gourbault 1992,
Warwick etal. 1988): Diplopeltidae. Cuticle finely to coar-
sely striated, sometimes with longitudinal striae to alae.
Amphids loop-shaped elongated on a cuticularized plate.
Ocelli absent. Buccal cavity small or absent, situated ter-
minally or subterminally. Pharynx with or without a pos-
terior bulb. Two ovaries. Apophyses usually present or
inconspicuous. Tail conical or conical-cylindrical.
Number of species: 21
Type species: D. brevicepis Gerlach, 1950
7.15.5.6Genus Edalonema (Holovachov &
Sturhan, 2003) Andrssy, 2007
(Fig. 7.137)
Diagnosis (emended from Holovachov & Sturhan 2003):
Diplopeltidae. Cuticle coarsely annulated, with ten lon-
gitudinal ridges. Inner and outer labial sensilla papilli-
form. Cephalic sensilla setiform. Amphid a transverse
slit. Buccal cavity weakly cuticularized, tubular. Pharynx
mostly cylindrical, widening at the posterior end. Females
monodelphic, prodelphic, ovary branch outstretched.
Spermatheca an offset pouch, located on right side at ovi-
duct-uterus junction. Postvulval uterine branch long. Male
reproductive system diorchic, anterior testis outstretched,
posterior testis reflexed. Spicules thin and arcuate. Guber-
naculum plate-like and apophyses absent. One precloacal
sensillum present. Caudal glands and spinneret absent.
Tail enlogated conoid with a filiform posterior end.
Monospecific: E. camerunense Holovachov & Sturhan,
2003
7.15.5.7Genus Intasia Tchesunov &
Miljutina, 2008 (Fig. 7.137)
Diagnosis (emended from Tchesunov & Miljutina 2008):
Diplopeltidae. Cuticle finely striated without lateral dif-
ferentiation. Anterior sensilla visible as six outer labial
papillae and four cephalic setae. Amphidial fovea round,
formed by one loop. Buccal cavity terminal, oral opening
relatively small and posterior portion long and tubular
with slightly sclerotized walls. Pharynx evenly muscular
with a slight widening posterior to the buccal cavity and
a pyriform bulb at the posterior end. Female gonad pro-
delphic, outstretched with a posterior blind sack. Vulva
Brought to you by | Universidade Federal de So Paulo UNIFESP
placed posterior to the mid-body region. Spicules short
and arcuate. Gubernaculum with dorso-caudal apophy-
ses. Pore-like precloacal supplement present. Tail conical.
Monospecific: I. monohystera Tchesunov & Miljutina,
2008
7.15.5.8Genus Metaraeolaimoides De
Coninck, 1936 (Fig. 7.137)
Diagnosis (emended from Tchesunov & Miljutina 2008):
Diplopeltidae. Cuticle finely striated. Anterior body region
narrower than the rest of the body. Amphids loop-shaped
with a circular outline, situated far posterior to the ante-
rior end. Cervical setae usually present. Buccal cavity ter-
minal narrow cylindrical and weakly sclerotized. Pharynx
narrow slightly widening at the posterior end. Ocelli
present. Two ovaries. Apophyses present. Tail conical.
Monospecific: M. oxystomina De Coninck, 1936
7.15.5.9Genus Morlaxia Vincx & Gourbault,
1988 (Fig. 7.137)
Diagnosis (emended from Vincx & Gourbault 1988):
Diplopeltidae. Cuticle finely striated. Amphids spiral,
loop-shaped with a circular outline. Anterior body
end asymmetrical with the buccal aperture and buccal
cavity displaced to the ventral side of the body. Pharynx
narrow, ending with a posterior oval bulb. One ante-
rior ovary outstretched. Postvulva sac present. Spicu-
les ventrally curved. Apophyses directed dorsally. Tail
conical.
Number of species: 2
Type species: M. contusa Vincx & Gourbault, 1988
7.15.5.10Genus Pararaeolaimus Timm,
1961 (Fig. 7.137)
Diagnosis (emended from Lorenzen 1973, Jensen 1991b):
Diplopeltidae. Body shape stout. Cuticle smooth. Labial
sensilla indistinct. Cephalic sensilla papilliform. Amphids
relatively large, formed by one loop with a circular outline.
Ocelli absent. Buccal cavity terminal or slightly shifted
to the dorsal side, cup-shaped and sclerotized. Pharynx
cylindrical. Three to four pairs of ventral glands cells
arranged in tandem. Two outstretched gonads. Spicules
ventrally curved without caudal apophyses. Tail conical.
Number of species: 3
Type species: P. nudus (Gerlach, 1951) Timm, 1961

7.15.5.11Genus Pseudaraeolaimus
Chitwood, 1951 (Fig. 7.137)
Diagnosis (emended from Chitwood 1951, Tchesunov &
Miljutina 2008): Diplopeltidae. Cuticle smooth or striated.
Amphids loop-shaped and slightly elongated longitudi-
nally. Four paramphidial setae present at the level of the
amphids. Ocelli present or absent. Buccal cavity narrow,
not sclerotized and terminal. Pharynx slender, ending
with posterior ventral apophyses containing five or six
pharyngeal glands, posterior to the cardia and in tandem,
with excretory gland. Two outstretched ovaries. Guberna-
culum without apophyses. Papilloide precloacal supple-
ment usually present. Tail conical.
Number of species: 2
Type species: P. perplexus (Chitwood, 1951) Gerlach &
Riemann, 1973
7.15.5.12Genus Southerniella Allgn,
1932 (Fig. 7.137)
Diagnosis (emended from Tchesunov & Miljutina 2008):
Diplopeltidae. Cuticle smooth or finely striated. Labial
sensilla indisctinct, cephalic sensilla papilliform or
setiform. Amphids round or cryptospiral, with a broken
outline. Ocelli absent. Buccal cavity weakly sclerotized,
narrow, cylindrical with a small aperture. Pharynx
cylindrical, gradually widening posteriorly. Terminal
bulb might be present. Ventral gland usually associa-
ted with other cells, up to four glands. Two outstretched
ovaries. Spicules with or without apophyses. Tail conical
cylindrical.
Number of species: 13
Type species: S. cylindricauda (Allgn, 1933) Gerlach &
Riemann, 1973
Acknowledgments
The authors would like to thank D. C. Vieira for assis-
ting with the figures. The first author is supported with a
post-doc grant (FAPESP 2010/054720) project (FAPESP
2009/140190).
Literature
For references regarding nematode species up to 1972,
see Gerlach, S. A. & Riemann, F. (1973/74): The Bremer-
haven checklist of aquatic nematodes, Vol. 4. A catalogue
Brought to you by | Universidade Federal de So Paulo UNIFESP
Literature 485
of Nematoda Adenophorea excluding the Dorylaimida.
Verffentlichungen des Instituts fr Meeresforschung,
Bremerhaven.
Andrssy, I. (2007): Free-living Nematodes of Hungary (Nematoda
errantia), Vol. II. Hungarian Natural History Museum and
Systematic Zoology Research Group of the Hungarian Academy
of Sciences, Budapest.
Boucher, G. (1973): Nmatodes libres marins des les hautes de
Polynsie. 1. Comesomatidae et Axonolaimidae. Cah. Pacif.
17: 205233.
Boucher, G. & Hellout, M. N. (1977): Nmatodes des sables fins
infralittoraux de la pierre Noire (Manche occidentale). III.
Araeolaimida et Monhysterida. Bull. Mus. Natn. Hist. Nat.
3: 85122.
Bussau, C. (1990): Freilebende Nematoden aus Kstendnen und
angrenzenden Biotopen der deutschen und dnischen Ksten.
II. Monhysterida, Enoplida und Trefusiida (Nematoda). Zool.
Anz. 225: 189209.
De Ley, P. & Blaxter, M. L. (2004): A new system for Nematoda.
In: Cook, R. & Hunt, D. J. (eds.) Proceedings of the Fourth
International Congress of Nematology, 813 June 2002,
Tenerife, Spain. Nematol. Monogr. Perspect 2: 633653.
Dihn Tu, N., Vu Thanh, N., Smol, N. & Vanreusel, A. (2008). New
genus Asymmelaimus gen. n., sp. n. and new marine nematode
species of the subfamily Dorylaimopsinae de Coninck, 1965
(Comesomatidae Filipjev, 1918) from Halong Bay, Vietnam.
Russ. J. Nematol. 16: 716.
Gourbault, N. (1980): Nematodes abyssaux (Campagne Walda du
N/O Jean Charcot). II Especes et genre nouveaux de
Comesomatidae. Bull. Mus. Natn. Hist. Nat. 3: 737749.
Holovachov, O. & Sturhan, D. (2003): Adelonema camerunense
gen. et sp. n. (Araeolaimida: Diplopeltidae) from rain forest
in Cameroon. Russ. J. Nematol. 11: 6366.
Hope, W. D. & Zhang, Z. (1995): New nematodes from the Yellow
Sea, Hopperia hexadentata n. sp. and Cervonema deltensis
n. sp. (Chromadorida: Comesomatidae), with observations
on morphology and systematic. Invert. Biol. 114: 119138.
Jensen, P. (1976): Free-living marine nematodes from a sublittoral
station in the North Sea off the Belgian Coast. Biol. Jaarb.
Dodonaea 44: 231255.
Jensen, P. (1979): Revision of Comesomatidae (Nematoda). Zool.
Scripta 8: 81105.
Jensen, P. (1988): Four new nematode species abundant in the
deep-sea benthos of the Norwegian Sea. Sarsia 73: 149155.
Jensen, P. (1991a): Bodonematidae fam. n. (Nematoda, Monhysterida)
accomodating Bodonema vossi gen. et sp. n. from the deep-sea
benthos of the Norwegian Sea. Sarsia 76: 1115.
Jensen, P. (1991b): Nine new and less known nematode species
from the deep-sea benthos of the Norwegian Sea.
Hydrobiologia 222: 5776.
Jensen, P. (1992): Description of the marine nematode Hopperia
australis sp. n. (Comesomatidae) from mangroves in Darwin,
Australia, with a pictorial key to Hopperia species. Cah. Biol.
Mar. 33: 495500.
Jensen, P. & Gerlach, S. A. (1976): Three new Nematoda-
Comesomatidae from Bermuda. Ophelia 16: 5976.
Lambshead, P. J. D. (1982): A new species of free-living nematode
from the Firth of Clyde, Scotland. Bull. Brit. Mus. Nat. Hist. 42:
149155.
486 7.15 Order Araeolaimida De Coninck & Schuurmans Stekhoven, 1933
Lorenzen, S. (1973): Freilebende Meeresnematoden aus dem
Sublitoral der Nordsee und der Kieler Bucht. Verff. Inst.
Meeresforsch. Bremerhaven 14: 103130.
Lorenzen, S. (1981): Entwurf eines phylogenetischen Systems
der freilebenden Nematoden. Verff. Inst. Meeresforsch.
Bremerhaven, Sonderband 7: 1472. (Engl. Translation:
Lorenzen, S. (1994). The Phylogenetic Systematics of Freeliving
Nematodes. Ray Society, London.)
Meldal, B. H. M., Debenham, N. J., De Ley, P., De Ley, I. T.,
Vanfleteren, J. R., Vierstraete, A. R., Bert, W., Borgonie,
G., Moens, T., Tyler, P. A., Austen, M. C., Blaxter, M. L., Rogers,
A. D. & Lambshead, P. J. D. (2007): An improved molecular
phylogeny of the Nematoda with special emphasis on marine
taxa. Mol. Phyl. Evol. 42: 622636.
Muthumbi, A. W., Soetaert, K. & Vincx, M. (1997): Deep-sea
nematodes from the Indian Ocean: new and known species of
the family Comesomatidae. Hydrobiologia 346: 2557.
Platt, H. M. (1973): Freeliving marine nematodes from Strangford
Lough, Northern Ireland. Cah. Biol. Mar. 14: 295321.
Platt, H. M. (1985): The freeliving marine nematode genus
Sabatieria (Nematoda: Comesomatidae). Taxonomic revision
and pictorial keys. Zool. J. Linn. Soc. 83: 2778.
Riemann, F. (1986): Nicascolaimus punctatus gen. et sp. n.
(Nematoda Axonolaimoidea) with notes on sperm dimorphism
in free-living marine nematodes. Zool. Scripta 15: 119124.
Sergeeva, N. G. (1973): New species of free-living marine nematodes
from the order Chromadorida in the Black Sea. Zool. Zh. 52:
12381241.
Siddiqi, M. R. (2004): Ten new species of Cylindrolaimus
de Man, 1880 (Nematoda: Araeolaimida). Int. J. Nematol.
14: 199212.
Smolyanko, O. I. & Belogurov, O. I. (1991): Description and
taxonomic position of Expressonema grandulata gen. and
sp. n. and structural analysis of the family Dorylaimopsinae
(Nematoda, Comesomatidae). Zool. Zh. 70: 117127.
Brought to you by | Universidade Federal de So Paulo UNIFESP
Smolyanko, O. I. & Belogurov, O. I. (1993): Apodontium bellum sp.
n. and Aequalodontium gemellum gen. n., sp. n. (Nematoda:
Axonolaimidae) with remarks on Apodontium morphology.
Russ. J. Nematol. 1: 4754.
Tarjan, A. C. & Kuong, N. B. (1988): A compendium of the family
Axonolaimidae (Nematoda). Cah. Biol. Mar. 28: 375393.
Tchesunov, A. V. (2000): Several new and known species from the
families Coninckiidae and Comesomatidae (Nematoda) in the
White Sea. Hydrobiologia 435: 4359.
Tchesunov, A. V. & Miljutina, M. A. (2008): A new freeliving
nematode Intasia monohystera gen. n., sp. n. (Nematoda,
Araeolaimida, Diplopeltidae) from the Barents Sea and the
White Sea, with a key to genera of Diploepltidae. Russ. J.
Nematol. 16: 3348.
Vincx, M. & Furstenberg, J. (1989): Namibnema papillata gen.
et sp. n. and Axonolaimus deconincki sp. n. (Nematoda,
Axonolaimoidea) from marine and estuarine beaches of
southern Africa. Zool. Scripta 18: 231237.
Vincx, M. & Gourbault, N. (1988): Two new species of Morlaxia gen.
n. (Nematoda: Diplopeltidae) with a subterminal ventral mouth.
Bull. Mus. Natn. Hist. Nat. 4: 711718.
Vincx, M. & Gourbault, N. (1992): Six new species of the genus
Diplopeltula (Nematoda: Diplopeltidae) with remarks on
the heterogeneity of the taxon. Hydrobiologia 230:
165178.
Warwick, R. M., Platt, H. M. & Somerfield, P. J. (1998): Freeliving
marine nematodes: Part III Monhysterids. Pictoral key to world
genera and notes for the identification of British species.
Synopses of the British Fauna (New Series) No. 53, Field
Studies Council, London.
Wu, J., Somerfield, P. J., Austen, M. C. & Liang, Y. (2000): The
freeliving nematode genus Parodontophora Timm 1963
(Nematoda: Axonolaimidae) is not exclusively marine:
Parodontophora limnophila sp. nov. from freshwater in
China. Hydrobiologia 431: 205210.