Nematology 00 (2016) 1-18

brill.com/nemy

Description and molecular characterisation of two new species

of Dorylaimidae (Dorylaimida: Nematoda) from Iran,
with a compendium of all the nominal species
of Dorylaimus Dujardin, 1845
Maria Teresa V INCIGUERRA 1 , Ali E SKANDARI 2, , Mirella C LAUSI 1 ,
Ramezan A SGHARI 2 and Giancarlo R APPAZZO 1
1

Department of Biological, Geological and Environmental Sciences, University of Catania, via Androne 81,
95124 Catania, Italy
2
Department of Plant Protection, Faculty of Agriculture, University of Zanjan, 45371-38791 Zanjan, Iran
Received: 7 April 2015; revised: 9 December 2015
Accepted for publication: 9 December 2015; available online: ???

Summary Two new species of Dorylaimidae from Iran are described using morphology and molecular data. Dorylaimus elegans
sp. n. is characterised by 4.41-5.92 mm body length, lip region truncate, distinct from adjoining body by a depression, cuticle with
50-55 longitudinal ridges, odontostyle 42-51 m long, thicker than cuticle at its level, didelphic female reproductive apparatus, vulva
a small sub-equatorial pore, a variable number of advulval papillae (0-6), both pre- and post-vulval, present in most specimens, female
tail elongate, 5.6-7.6 anal body diam. long, with ventrally hooked terminus. Male with 26-30 ventromedian supplements arranged
very close to each other, copulatory hump weakly developed, and a convex-conoid tail with blunt terminus. Prodorylaimus reyesi sp.
n. is characterised by 2.78-3.33 mm body length, lip region slightly angular and well set off from adjoining body by a depression,
odontostyle 26-31 m long, thicker than cuticle at its level, didelphic female reproductive apparatus, vulva longitudinal, a variable
number of advulval papillae (0-3), both pre- and post-vulval, present in most specimens, tail in both sexes convex-conoid in its anterior
part, then narrowing abruptly to a filiform hyaline appendix, longer in female (c = 11.7-15.5) than in male (c = 7-7.7), and 19-21
contiguous ventromedian supplements. A compendium of the main morphometric characters of Dorylaimus and a key to species are
also provided.
Keywords Dorylaimus elegans sp. n., key, morphology, morphometrics, Prodorylaimus reyesi sp. n., taxonomy.

During a recent nematological survey carried out in

the Jangal-e Abr (Cloud Forest), Iran, two undescribed
species of the family Dorylaimidae de Man, 1876 were
isolated from the rhizosphere of hornbeam (Carpinus
betulus L.). This forest is located 45 km north-east
of Shahrood county, Semnan province, Iran, and is a
continuum of the northern forests found in the south of
the Alborz mountain range. The Abr forest area is overcast
and foggy in most seasons. It is one of the oldest forests
in Iran, a remnant of the third geological age.
The objectives of the present study were to describe
the two new species of Dorylaimidae found, which are
described and figured herein as Dorylaimus elegans sp. n.
and Prodorylaimus reyesi sp. n., and to prepare a new key
Corresponding

and a compendium containing morphometric and related

details to facilitate the identification of the Dorylaimus
species presently considered valid.
Members of Dorylaimus Dujardin, 1845 and Prodorylaimus Andrssy, 1959 are interesting free-living dorylaims occurring throughout the world. Nematodes belonging to Dorylaimus live in limnic habitats or in moist soil
and moss (Andrssy, 1959, 1988). Andrssy (1969) revised the taxonomy of the genus and gave a key to 16
valid species. Mulvey & Anderson (1979) also reviewed
the genus, provided a comprehensive key, and tabulated
diagnostic characters for 22 known species. In the latest
revision of the genus, Andrssy (1988) provided a key for
the identification of the 24 valid species of Dorylaimus

author, e-mail: eskandari.a@znu.ac.ir

Koninklijke Brill NV, Leiden, 2016

DOI 10.1163/15685411-00002960

M.T. Vinciguerra et al.

known at that time. Andrssy (2009) gave an updated list

of the Dorylaimus species, although since then, a number of new species have been described, the genus currently containing 33 species considered valid. Therefore,
and also because of the large morphological diversity of
the species and of new data provided for some species,
we decided to produce a new key and a diagnostic compendium to all the nominal species of this genus. In the
case of Prodorylaimus, a revised key is not currently required since recently Vinciguerra & Orselli (2011) provided one together with a table of the main morphometric
characters of each species.
The Iranian nematode fauna belonging to the Dorylaimidae is poorly known and as far as we know, this is the first
report of species of the aforementioned genera from Iran.

Materials and methods

Nematodes were extracted from soil samples by the tray
method (Whitehead & Hemming, 1965), then killed by
gentle heat and fixed in a FGA (formaldehyde, glycerin
and acetic acid = 4:1:1) solution and finally processed to
anhydrous glycerin (De Grisse, 1969). Permanent slides
were made and examined using a Leica model DMRB
light microscope. Drawings and measurements were made
using a drawing tube attached to the microscope and
with the aid of Adobe Photoshop CS5; microphotographs
were taken with a Leica EC3 camera. The positions
of pharyngeal gland nuclei were calculated following
Andrssys (1998) formula.
M OLECULAR CHARACTERISATION
DNA extraction and PCR
For the molecular study, a single specimen was handpicked into distilled water, its identity confirmed under
the light microscope, and then placed into a small drop of
AE buffer (10 mM Tris-Cl, 0.5 mM EDTA; pH 9.0) and
crushed into multiple pieces on a microscope slide with
a pipette tip (Zeng et al., 2012). The suspension (DNA
samples) of microscope slide was collected by pipette and
stored at 20C until used as a PCR template. The forward D2A (5 -ACAAGTACCGTGAGGGAAAGTTG-3 )
and reverse D3B (5 -TCGGAAGGAACCAGCTACTA3 ) primers were used for amplification and sequencing
of the fragment of the 28S rDNA gene (Nunn, 1992).
Twenty-five l of the PCR mixture contained 0.4 U Taq
DNA polymerase (CinnaGen), 1 l MgCl2 , 1 l dNTPs,
3 l of each primer (10 pmol l1 ), 2.5 l of 10 buffer,
2

3 l of template DNA and 11.1 l ddH2 O. The PCR

amplification profile consisted of 5 min at 95C; 35 cycles
of 30 s at 94C, 45 s at 55C and 2 min at 72C, followed
by a final step of 10 min at 72C. Four l of the PCR
product was run on a 1% TBE buffered agarose gel (80 V,
40 min) and stained with ethidium bromide. Gels were
photographed under UV light.
Sequence analysis
Sequences from the 28S rDNA locus, also known as
LSU (Large Sub-Unit), were obtained from D. elegans
sp. n. and P. reyesi sp. n. by sequencing PCR-amplified
fragments in both directions (Bioneer). For each species,
the consensus sequence was obtained by aligning forward
and reverse reads. The consensus span 793 bp for P. reyesi
sp. n. and 805 bp for D. elegans sp. n. Data were deposited
in GenBank under accession numbers KP954677 for D.
elegans sp. n. and KP954678 for P. reyesi sp. n.
Phylogenetic analysis
Homologous sequences were retrieved by Taxonomy
browser at NCBI (www.ncbi.nlm.nih.gov/Taxonomy) using the keyword Dorylaimoidea. This choice allowed the
inclusion of data from both Dorylaimus and Prodorylaimus. However, a large cohort of genera was also included, many of which are not very close to either of the
former two genera. It should be noted that the NCBI taxonomy database is not an authoritative source for nomenclature or classification. Data obtained included mostly
SSU (Small Sub-Unit, i.e., 18S rDNA) and a reduced
number of LSU (Large Sub-Unit, i.e., 28S rDNA) sequences.
In order to keep data consistency without including data
from relatively distant genera, we adopted a two steps
approach. In a first step, a reference tree was constructed
using only 28S sequences already available in taxonomy
databases. The resulting tree allowed us to recognise a
small cluster of genera, such as Dorylaimus, Labronema
Thorne, 1939, Prodorylaimus, Ecumenicus Thorne, 1974
and, in an adjacent group, Paraxonchium Krall, 1958,
Eudorylaimus Andrssy, 1959 and Opisthodorylaimus
Ahmad & Jairajpuri, 1982 (data not shown). More distant
entries were then excluded from the comparison, with the
exception of Sectonema Thorne, 1930, which was retained
as outgroup.
A total of ten sequences was selected for comparative
analysis. They included LSU sequences, ca 1100 bp long,
from the following species: Calcaridorylaimus castaneae
Nedelchev, Elshishka, Lazarova, Radoslavov, Hristov &
Nematology

Two new species of dorylaim from Iran

Peneva, 2014 (KF717498.1), Dorylaimus stagnalis Dujardin, 1845 (AY592994.1), Ecumenicus monohystera (de
Man, 1880) Thorne, 1974 (AY593013.1), Ecumenicus sp.
JH-2004 (AY593014.1), Eudorylaimus centrocercus (de
Man, 1880) Andrssy, 1959 (AY593007.1), Labronema
vulvapapillatum (Meyl, 1954) Loof & Grootaert, 1981
(AY592996.1), Opisthodorylaimus sylphoides (Williams,
1959) Carbonell & Coomans, 1986 (AY593008.1), Paraxonchium laetificans (Andrssy, 1956) Altherr & Loof,
1969 (AY593001.1), Prodorylaimus sp. HHBM-2007a
(EF207241.1), and Sectonema barbatoides Heyns, 1965
(AY593030.1). In a second phase, the sequences of D.
elegans sp. n. and of P. reyesi sp. n. were introduced
into the multi-alignment, with duplicated entries from
the same species being omitted. Multi-alignments were
done with ClustalW, giving a total of 790 sites including gaps, or with MUSCLE, which yielded 796 sites. For
phylogenetic analysis, the software pRANK was adopted
since its algorithm includes an evaluation of conserved
and variable sites on the basis of structural conservation (Lytynoja & Goldman, 2005). It was used online
at EBI (www.ebi.ac.uk/Tools/webservices/services/msa).
The package MEGA6 (Tamura et al., 2013) was extensively used on site. Genetic distances were inferred by
using the Maximum Likelihood method based on the
Kimura 2-parameter model (Kimura, 1980) and they were
expressed in the units of the number of base substitutions
per site. The rate variation among sites was modelled with
a gamma distribution (shape parameter = 2). Phylogeny
reconstruction was done with Maximum Likelihood, and
tested with 1000 bootstrap iterations (Felsenstein, 1985).

Results
Dorylaimus elegans* sp. n.
(Figs 1, 2)
M EASUREMENTS
See Table 1.
D ESCRIPTION
Adults
Large body appearing more or less straight, sometimes
C-shaped, after fixation. Cuticle two-layered, with very
* The specific epithet refers to the elegance and beauty of these
animals.

Vol. 00(0), 2016

thin outer layer and thick inner one, 4.5-6.0 m thick

at level of odontostyle fixed ring and 6-8 m at midbody, bearing 50-55 longitudinal ridges (counted in cross
section). Lip region anteriorly truncate, 2.3-3.2 times as
broad as high, slightly set off from adjacent body by a depression, lips amalgamated. Two circles of six inner and
six outer prominent papillae present. Amphids stirrupshaped with a slit-like opening at base of lip region, occupying 50-58% of lip region diam. Odontostyle massive,
clearly thicker than cuticle, 1.7-1.9 lip region diam. long,
its aperture 27-33% of its length, its width 1/6-1/7 of its
length. Odontophore rod-like, simple. Guiding ring double, fixed ring distant from anterior end ca one lip region diam. One or two body pores present both dorsally
and ventrally at odontostyle level. Pharynx very gradually
expanding without a clear transition from a slender to a
wider part, apparently in two steps, second step forming
posterior 15% of pharynx, glandularium generally occupying >50% of neck length. Positions of nuclei as follows (n = 6): D: 47-52.5%, AS1: 49.4-56.3%, AS2: 54.456.7%, PS: 72.5-76.4%. Nerve ring located at 21-24% of
neck length. Cardia conoid. Prerectum 2.6-5.0 and rectum
1.1-2.0 anal body diam. long.
Female
Reproductive system didelphic-amphidelphic, well developed, with anterior and posterior branch of similar
length, anterior 582-762 m or 11.5-13.0% of body
length, posterior 603-814 m or 12-13% of body length.
Ovaries large, reaching sphincter level, anterior 196374 m, posterior 222-357 m long, oocytes arranged
first in two or more rows, then in a single row. Oviduct
197-298 m long, consisting of a slender part with prismatic cells and a weakly developed but rather long pars
dilatata, often containing sperm cells, followed by a
sphincter at its junction with uterus. Uterus 803-943 m
long, with a complex morphology consisting of a distal slender part, with narrow lumen, and a proximal part
with wide lumen, distal part, in some specimens, particularly full of sperm, proximally enlarging to form a short
swollen part containing sperm followed by a weak constriction, proximal part of uterus much wider, composed
of two parts: distal part of variable length, full of sperm
cells densely packed in all specimens, proximal part often containing one or two eggs per side or empty, no constriction present between these parts which appear continuous. Uterus sperm cells 7-9 m long, spindle-shaped.
Uterine eggs thick-walled, ovoid, 52-58 109-118 m
in size. Vagina occupying 50-56% of corresponding body
diam., composed of three parts: pars distalis 3-6 m long;
3

M.T. Vinciguerra et al.

Fig. 1. Dorylaimus elegans sp. n. A: Female body; B: Female genital apparatus; C: Female posterior region; D: Anterior end; E: Neck;
F: Male body; G: Male posterior end. (Scale bars: A, F = 400 m; B, C, E, G = 100 m; D = 20 m.)

Nematology

Two new species of dorylaim from Iran

Fig. 2. Dorylaimus elegans sp. n. A, D: Anterior end; B: Neck; C: Female posterior genital branch; E: Cardia region; F: Vulval region;
G, H: Male posterior end; I: Female posterior end. (Scale bars: A, E, F, G = 20 m; B = 200 m; C, H, I = 50 m; D = 5 m.) This
figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/
content/journals/15685411.

Vol. 00(0), 2016

M.T. Vinciguerra et al.

Table 1. Morphometrics of Dorylaimus elegans sp. n. All measurements are in m, except for L in mm, and in the form: mean s.d.
(range).
Character

Female

Male

Holotype

Paratypes

Paratypes

n
L (mm)

5.48

57.0

4.9

15.6

c

6.7

V/T

48.7

13
4.80 0.28
(4.41-5.09)
50.5 8.5
(44.0-69.0)
4.6 0.3
(4.3-5.0)
123.5 23.7
(96-166)
0.6 0.1
(0.5-0.7)
54.1 3.6
(49.0-60.5)
104 5.6
(93-111)
65 4.5
(59-71)
26 1.2
(25-28.5)
9.5 0.8
(8.5-11.0)
47 2.6
(42-51)
7.0 0.7
(6.0-9.0)
66 12
(40-68)
29 1.2
(27-31)
1073 37.2
(1026-1135)
553 55.7
(425-620)
245 11.1
(224-263)
159 14.9
(130-180)
258 53.5
(184-348)
98 11.6
(79-118)
41 5.2
(33-48)
100 5.4
(92-107)
45 3
(40-51)

Spicule length

7
5.50 0.41
(4.76-5.92)
50.0 5.2
(39.0-56.0)
5.0 0.2
(4.9-5.2)
15.0 1.2
(13.0-17.0)
7.0 0.9
(5.6-7.6)
48.0 2.4
(44.0-50.3)
109 6.1
(103-122)
54 2.2
(51-56)
26 1.3
(25-28)
10.0 1.3
(8.0-12.0)
47 1.7
(46-50)
7.0 0.5
(6.5-8.0)
52 5.4
(47-64)
26 1.6
(24-29)
1110 87.3
(971-1273)
608 51.8
(542-691)
288 69.5
(235-399)
150 34.3
(81-178)
207 70.8
(170-261)
71 9.3
(61-87)
376 29.8
(312-417)

Lateral guiding pieces

Mid-body diam.

95

Anal/cloacal body diam.

52

Lip region diam.

27

Lip region height

Odontostyle length

46

Odontostyle width

7.0

Odontophore length

47

Fixed ring from anterior end

30

Neck length

1122

Glandularium length

571

Nerve ring from anterior end

264

Excretory pore from anterior end

168

Pre-rectum length

203

Rectum length

79

Tail length

352

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Two new species of dorylaim from Iran

pars refringens with two triangular sclerotised pieces, 1022 m long; and pars proximalis conoid-cylindroid, 1629 m long. Vulva a small pore. Variable number (generally two or three, exceptionally none or 5-6) of advulval papillae present both anterior and posterior to vulva,
in some specimens some additional incomplete papillae
present. Prerectum 2.6-5.0 times as long as anal body
diam. Tail elongate, at first convex-conoid then abruptly
decreasing in diam., 7 (5.6-7.6) anal body diam. long, terminus often bent to form a small hook. Two pairs of caudal pores present at end of wider part of tail.

Holotype female, one female paratype and two male paratypes deposited in the collection of the Section of Animal Biology of the Department of Biological, Geological and Environmental Sciences, University of Catania,
Catania, Italy; one female paratype and one male paratype deposited in USDA Nematode Collection, Beltsville,
MD, USA. Seven female paratypes and ten male paratypes housed in the Department of Plant Protection, Faculty
of Agriculture, University of Zanjan, Zanjan, Iran.

50-55 longitudinal ridges, odontostyle 47 (42-51) m

long, thicker than cuticle at its level, pharynx muscular,
enlarging very gradually, female reproductive apparatus
didelphic, vulva a small sub-equatorial pore, a variable
number of advulval papillae (0-6), both pre- and postvulval, present in most specimens, female tail elongate,
7.0 (5.6-7.6) anal body diam. long, with ventrally hooked
terminus. Male with 26-30 ventromedian supplements
arranged very close to each other, copulatory hump
weakly developed and a convex-conoid tail with blunt
terminus.
The new species is very similar to D. stagnalis in
the main morphological features and morphometrics, but
can be distinguished from it by the greater number of
cuticular ridges (50-55 vs 28-41), fewer supplements (2630 vs 30-40) and the presence of numerous advulval
papillae in most specimens vs a maximum of one per
side. Moreover, the molecular data clearly distinguish the
two species. Among the species with a similar number of
longitudinal ridges, D. elegans sp. n. can be compared to
D. pachys Andrssy, 1970; D. alaeus Thorne, 1939; D.
stenus Andrssy, 1970; D. specialis Andrssy, 2003 and
D. neominimus Gantait, Bhattacharya & Chatterjee, 2009.
It differs from D. pachys by the thinner cuticle (4.58.0 vs 14-18 m), shorter odontostyle (47 (42-51) vs 6063 m), and more posterior vulva (V = 48 (44-50) vs 3644); from D. alaeus it mainly differs by the longer tail
(c = 7.0 (5.6-7.6) vs 4.0) and by fewer ventromedian supplements (28-30 vs 50-54); from D. stenus by the differently shaped tail (first convex-conoid and then abruptly
narrowing vs conoid, regularly narrowing), shorter prerectum (2.6-5.0 vs 5.7-6.0 anal body diam.), rectum normal
vs distally swollen, advulval papillae generally present vs
absent, and presence of male; from D. specialis by the
rather shorter body (L = 5.5 (4.4-5.9) vs 5.6-6.9 mm), the
more set off lip region, the shorter odontostyle (47 (42-51)
vs 52-58 m), shorter prerectum (2.6-5.0 vs 5.8-9.0 anal
body diam.), vagina with sclerotised vs non-sclerotised
pieces, and fewer ventromedian supplements (28-30 vs
44); and from D. neominimus by the much longer body
(L = 5.5 (4.41-5.92) vs 1.9 (1.75-2.16) mm), vagina sclerotised vs not sclerotised, advulval papillae present vs absent, and more numerous ventromedian supplements (2830 vs 21-22).

D IAGNOSIS AND RELATIONSHIPS

N OTES ON Dorylaimus D UJARDIN , 1845

Dorylaimus elegans sp. n. is characterised by the 5.5

(4.41-5.92) mm body length, lip region slightly angular,
distinct from adjoining body by a depression, cuticle with

Dorylaimus is the oldest genus of Dorylaimoidea and

many species now attributed to other genera were originally described as its members. As previously stated, the

Male
Posterior part of body more curved than in female, tail
convex-conoid with blunt terminus, shorter than cloacal
body diam. Testes paired. 26-30 ventromedian supplements arranged very close to each other, posteriormost
85-95 m from cloacal opening, copulatory hump weakly
developed but always observable anterior to supplement
series. Spicules rather slender, narrowing more in posterior half, with ventral hump poorly developed and only
slightly bent, 1.5-1.6 cloacal body diam. long. Prerectum
2.6-5 cloacal body diam. long. Very numerous (at least
18, but number variable) pairs of caudal pores visible in
all specimens.
T YPE HABITAT AND LOCALITY
The new species was extracted from the rhizosphere of
hornbeam, in Jangal-e Abr (Cloud Forest), Shahrood, Iran
(GPS coordinates: 3645.342 N, 5502.004 E, altitude
2120 m a.s.l.).
T YPE MATERIAL

Vol. 00(0), 2016

M.T. Vinciguerra et al.

species list of Dorylaimus has been updated several times

but no comparative compendium of its species has been
published since the key provided by Andrssy (1988).
Therefore, we provide an updated list, a compendium of
all the described species of Dorylaimus with their main
morphometric characters and related bibliography (Table 2) and a key to their identification.

List of Dorylaimus species

T YPE SPECIES
Dorylaimus stagnalis Dujardin, 1845
OTHER SPECIES
D. afghanicus Andrssy, 1960
D. alaeus Thorne, 1939
D. asymphydorus Andrssy, 1969
D. baylyi Nicholas & Hodda, 2000
D. bengalensis Sen, Chatterjee & Manna, 2011
D. carinatus Thorne & Swanger, 1936
D. conicus Andrssy, 1981
D. crassus de Man, 1884
D. deaconi Botha & Heyns, 1991
D. elegans sp. n.
D. fodori Andrssy, 1988
D. geraerti Baqri & Jana, 1986
D. gigas Kleynhans, 1970
D. helveticus Steiner, 1919
D. lineatus Altherr in Altherr & Delamare-Deboutteville,
1972
D. macroproctus Altherr, 1963
D. macrosoma Jimnez-Guirado, 1988
D. montanus Stefanski, 1923
D. murlii Bohra & Sultana, 2008
D. neominimus Gantait, Bhattacharya & Chatterjee, 2009
D. numidicus Andrssy, 1988
D. pachys Andrssy, 1970
D. parvus Gagarin & Vu Than, 2003
D. popus Gagarin, 1981
D. siddiqii Ahmad & Jairajpuri, 1982
D. specialis Andrssy, 2003
D. stekhoveni Baqri & Coomans, 1973
D. stenus Andrssy, 1970
D. stephani Andrssy, 1969
D. tepidus Andrssy, 1959
D. thornei Andrssy, 1969
D. unicus Andrssy, 1970
8

Key to Dorylaimus species

(updated from Andrssy, 1988)
1 Number of longitudinal ridges >50 . . . . . . . . . . . . . . 2
Number of longitudinal ridges <45 . . . . . . . . . . . . . . 7
2 Cuticle very thick, 14-18 m at mid-body; odontostyle >60 m . . . . . . . . . . . . . . . . . . . . . . . . . . pachys
Cuticle not so thick, odontostyle <60 m . . . . . . . . 3
3 Body >6.4 mm long . . . . . . . . . . . . . . . . . . . . specialis
Body <6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Body length hardly reaching 2 mm . . . . neominimus
Body >4 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Tail relatively short (c = 4); ventromedian supplements 50-54 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . alaeus
Tail longer, c = 5 or more; ventromedian supplements 28-30 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Prerectum 5.7-6 anal body diam. long; female tail
gradually tapering . . . . . . . . . . . . . . . . . . . . . . . . . stenus
Prerectum <5 anal body diam. long; female tail
abruptly tapering . . . . . . . . . . . . . . . . . . . elegans sp. n.
7 Female tail conoid and very short; c = 2-4 . . . . . . . 8
Female tail attenuated to filiform; c > 4 . . . . . . . . . 9
8 Body length 3.6-5.2 mm; supplements 26-36. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . conicus
Body length 5.45-9.10 mm; supplements 72-86 . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . macrosoma
9 Body very slender (mean a = 55 or more) . . . . . . . 10
Body not so slender (mean a = up to 52) . . . . . . . . 12
10 Body length ca 4 mm, ventromedian supplements
22-25 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . baylyi
Body length  5 mm, ventromedian supplements 3339 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Body length ca 5 mm . . . . . . . . . . . . . . . . . . . . . tepidus
Body length 6.5-7.5 mm . . . . . . . . . . . . . . . . . . . . gigas
12 Odontostyle <45 m . . . . . . . . . . . . . . . . . . . . . . . . . 13
Odontostyle 50-60 m long . . . . . . . . . . . . . . . . . . . 23
13 Vulval region with 4-5 advulval papillae . . . . . . . . 14
Vulval region without papillae . . . . . . . . . . . . . . . . . 15
14 Lip region distinctly set off; longitudinal ridges 32
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . thornei
Lip region not or hardly set off, longitudinal ridges
38-42 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lineatus
15 Odontostyle 3 lip diam. long . . . . . . . . . . . . . . siddiqii
Odontostyle 2.0-2.5 lip diam. long . . . . . . . . . . . . . . 16
16 Body ca 2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Body 3-4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Nematology

Vol. 00(0), 2016

D. macroproctus

D. lineatus

D. helveticus

D. gigas

D. geraerti

D. fodori

D. elegans sp. n.

D. deaconi

D. crassus

D. conicus

D. carinatus

D. bengalensis

34-50

33-34

35-40

33

50-55

32-34

32-34

36

32-35

40-50

40-60
32?

Male
Female
Male
Female

Male
Female
Male
Female
Male
Female

Male
Female
Male
Female
Male
Female

Male
Female
Male
Female
Male
Female

Male
Female

2.6-4.0
4.4-5.1

2.5-2.9
6.5-7.5
5.2
4.2
3.6
2.4-4.3

2.67-3.78
4.76-5.92
4.41-5.09
3.6-4.2
3.34
2.8-3.5

2.2
4.5-5.2
3.6-4.6
4.2-4.8
3.6-4.2
2.80-3.92

3.75-3.87
2.6-3.5
2.4-3.0
2.3

26-42
48

32-33
58-62
52
28
28
28-43

31-47
39-56
44-69
37-41
39
32-39

28
44-50
44-48
24-27
25-26
39-47

46-66
29-34
30-31
39

56-86
14-17

68-83
13-16
122
24
67
11-15

87-140
13-17
96-166
15-16
76
12-13

50
37-44
85-94
14
62-70
11-19

77-99
13-15
71-82
8.3

36-40

46

45-51

39-41

0.9
5-7

46-50

38-41

40-43

47-51

44-45

35-42

40-43

45

10
0.8
3.7-6.7

5-6

0.6-1.5
5-8
0.5-0.7
5.0-5.3
0.9
6-11

1.0
2.3-3
0.7
5-6

5.3-7.6

0.8-1.3
4.2-4.7
0.7-0.8
8.6

40-42
57-60

50-52
50-52
48
48
40-42

0/1

0/0

0/0

2/3

0-5/0-5

0/0

0/0

1/1

0/0

0/0

40
53-55
53-55
48
48
39-48
40-48
44-50
42-51
40-42
40-42
43-47

0/0

0/0

43-46
44-46
41-47
40

Table 2. Main morphometric characters of Dorylaimus species ( measurements obtained from original drawings).
Species
Sex
Ridges L (mm)
a
c
c
V
Odontostyle Pre/postvulval
(m)
papillae
D. afghanicus
Female 32-34
3.4-4.0
38-42 12-15
5-6
37-39
46-50
0/0

46-50

Male

3.6
44
55
0.9
0/0
D. alaeus
Female 56-60
4.4
33
31
4
48
50

Male

4.2
36
71
0.9
D. asymphydorus Female 32-34
4.7-5.7
41-45 15-18
5-6
34-40
50-53
0/0
Male
5.1
44
55

50-53

D. baylyi
Female 28-32 3.42-4.35 48-61 15-20
6-7
42-53
43-46
0/0

85

81-84

128

100

71-86

92-107

88

92-98

140

53-61

73

Spicules
(m)

100

100

80

35-40

35-38

33

36-55

35-42

26-30

46

55

26-36

44

35-42

32-36

Andrssy (1960)

52

50-54

28

Altherr (1963)

Altherr &
DelamareDeboutteville
(1972)

Steiner (1919)

Kleynhans (1970)

Baqri & Jana

(1986)

Andrssy (1988)

This paper

Botha & Heyns

(1991)

de Man (1884)

Andrssy (1981)

Thorne & Swanger

(1936)

Sen et al. (2011)

Nicholas & Hodda

(2000)

Andrssy (1969)

Thorne (1939)

Reference

Suppl.

Two new species of dorylaim from Iran

10
23-32

44

32-34

54-56

32-34

52-56

30-35

31-33

34

50-60

32-35

41

56-58
40
36
32

32
28-29

Female

Male
Female
Male
Female

Male
Female

Male
Female
Male
Female
Male
Female

Male
Female
Male
Female

Male
Female
Male
Female

Male
Female

Male
Female
Female
Male
Female
Male
Female
Female
Male

D. macrosoma

D. stagnalis

D. stekhoveni

D. thornei
D. unicus

D. tepidus

D. stenus
D. stephani

D. specialis

D. siddiqii

D. popus

D. parvus

D. pachys

D. numidicus

D. neominimus

D. murlii

D. montanus

Ridges

Table 2. (Continued.)
Species
Sex

4.5-5.4
4.1-5.1
3.4-3.7
3.3-3.4
5.0-5.6
4.3-4.5
2.7-2.8
3.5-4.2
3.9

3.9-5.0
4.0-6.4

1.6-1.9
6.40-6.89
5.58
4.3-5.2

2.13-2.39
3.3-4.1
3.5-4.0
2.2

1.47-1.53
3.12-3.25
2.63-2.80
3.9-4.9
4.4
2.54-2.83

2.40-2.81
1.75-2.16

5.45-8.11
3.4-3.7
3.3-3.4
2.30-2.37

6.63-9.10

L (mm)

49-61
40-46
46-50
43-47
66-76
55-60
27-28
33-40
33

30-35
47-57

28-35
45-46
42
30-38

23-27
30-36
36-43
35-38

33-34
40-47
40-42
26-30
32
26-28

35
37-43

28-33
46-50
43-47
33-45

29-32

102-149
16-18
12-13
90-94
17-20
97-108
12-13
14-15
85

70-110
12-21

53-64
15
90
14-20

53-56
14-17
72-84
14

47
13-15
91-96
11-14
80
15-17

109-122
12-14

82-135
12-13
90-94
15-16

29-29

44-49

49

41-44

42-47

36-40

42-45
41-42

36-43

44-45
37-38

0.8
6.4-6.7
0.8
5-6
0.8
5-6
0.7
6-7
6-7
0.7
5-6
0.8
5.0-5.5
6-7
0.8

36-39

39-44

39-43

44

41-42

44-46

39-43

0.8-1.0
4.5-6
0.8
5-6

1.3
6-7
0.7
5-6
0.7
3.6-4.8

0.4
6.9-7.3

0.6-0.9
6-7
0.8
4.1-6.4

2.0-3.9

c

53-58
48-50
46-50
46-50
50-53
50-53
40
55-57
55-57

47-51
53-57

35-36
54-58
52
47-51

45-48
40-43
40-43
37

25-27
43-45
43-45
60-63
60-63
45-50

32
28-30

29-30

80
50

Odontostyle
(m)
71-89

0/0
0/0

1/1
0/0

0/0

0/0

0/0

0/0

0/0

0/0

0/0

0/0

0/0

0/0

0/0

Pre/postvulval
papillae
1/1

116-127

90

100

120

100

50

115

76-77

78-82

40-42

82-90

120

54-58

148-170

Spicules
(m)

55-62

39-41

35-39

46

30-40

31-34

44

47-49

26-27

21-22

22-27

33

24-28

72-86

39-41

Suppl.

Andrssy (1969)
Andrssy (1970)

Andrssy (1959)

Andrssy (1970)
Andrssy (1969)

Baqri & Coomans

(1973)

Thorne & Swanger

(1936); Mulvey &
Anderson (1979)

Andrssy (2003)

Ahmad &
Jairajpuri (1982)

Gagarin (1981)

Gagarin & Vu
Than (2003)

Andrssy (1970)

Andrssy (1988)

Gantait et al.
(2009)

Bohra & Sultana

(2008)

Stefanski (1923)

Jimnez Guirado
(1988)

Reference

M.T. Vinciguerra et al.

Nematology

Two new species of dorylaim from Iran

17 Spicule 80-90 m long; supplements 55 . . carinatus

Spicule 54-58 m long; supplements 24-28 . . murlii
18 Ventromedian supplements 46 . . . . . . . . . . . . . . fodori
Ventromedian supplements <42 . . . . . . . . . . . . . . . . 19

32 Body length 2.5-2.8 mm; ventromedian supplements

50-52 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . parvus
Body length >4 mm; ventromedian supplements 3645 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . stagnalis

19 Ventromedian supplements 32-42 . . . . . . . . . . . . . . 20

Ventromedian supplements 22-27 . . . . . . . . . . . . . . 22
20 Tail shorter (c = 4.2-4.7); ventromedian supplements 32-36 . . . . . . . . . . . . . . . . . . . . . . . . . bengalensis
Tail longer (c = 6-11); ventromedian supplements
36-42 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 Tail elongate-conoid gradually tapering . . . . deaconi
Tail convex-conoid, abruptly tapering . . . . . . geraerti
22 V = 36-39 . . . . . . . . . . . . . . . . . . . . . . . . . . . . numidicus
V = 44-49 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . popus
23 Cuticle very thick, 12-14 m at mid-body; spicule
140 m long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . crassus
Cuticle much thinner, <10 m; spicule 120 m
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 Longitudinal ridges 40-44 at mid-body . . . . . . . . . . 25
Longitudinal ridges 28-36 at mid-body . . . . . . . . . . 27
25 Lips well separate, angular; lip region well set off . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .stephani
Lips amalgamated, rounded; lip region not set off . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 Supplements 55-62; spicule length ca 120 m . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . stekhoveni
Supplements 25-35; spicule length ca 80 m . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . montanus
27 Odontostyle 3 lip diam. long . . . . . . . . . . . . . . . . . . . 28
Odontostyle not longer than 2.5 lip diam. . . . . . . . 29
28 Longitudinal ridges 28-29, irregularly spaced; V =
37-38 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . unicus
Longitudinal ridges 32-35, regularly spaced; V = 46
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . helveticus
29 Odontostyle 57-60 m long; one pre- and postvulval papillae present . . . . . . . . . . . . . . macroproctus
Odontostyle 45-53 m long; no advulval papillae . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30 Aperture occupying half odontostyle length; odontostyle with a dorsal fissure . . . . . . . . . asymphydorus
Aperture occupying one-third of odontostyle length;
no dorsal fissure in odontostyle . . . . . . . . . . . . . . . . . 31
31 Odontostyle markedly thicker than cuticle at same
level . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . afghanicus
Odontostyle as thick or thinner than cuticle at same
level . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Vol. 00(0), 2016

Prodorylaimus reyesi* sp. n.

(Figs 3, 4)
M EASUREMENTS
See Table 3.
D ESCRIPTION
Adults
Rather slender nematodes of relatively large size. Body
straight or slightly curved ventrally after fixation, tapering
towards both extremities, more so towards posterior end.
Cuticle 4-5 m thick posterior to lip region and 5 m
at mid-body. Two dorsal and two ventral pores present
posterior to lip region. Lip region slightly angular, 2.23.1 times as broad as high, well set off from adjoining
body by a depression, lips distinct, with prominent labial
and cephalic papillae. Amphidial opening 40-50% of
corresponding body diam.; fovea cup-like, opening at
base of lips. Odontostyle straight, 6-7 times as long
as wide and 1.4 times as long as lip region diam., its
aperture one third of its length. Odontophore rod-like, 1.31.6 times as long as odontostyle. Guiding ring double.
Pharynx consisting of a slender anterior part, gradually
expanding into a posterior enlarged section occupying
half of neck length. Positions of nuclei as follows (n = 3):
D: 55.2-57.5%, AS1: 39.6-48.6%, AS2: 40.2-51.3%, PS1:
56-66%, PS2: 56.4-66.1%. Nerve ring situated at 26-38%
of pharynx length. Cardia conoid, separated from pharynx
base by a thin disk-like structure. Tail convex conoid in its
anterior part, then narrowing abruptly to a filiform hyaline
appendix.
Female
Reproductive apparatus didelphic-amphidelphic, with
anterior and posterior branch of similar length, anterior
344-471 m or 12-15% of body length, and posterior
303-514 m or 12-16% of body length. Ovaries reflexed,
variable in length but generally relatively short, anterior
* The species is dedicated to Prof. Reyes Pea-Santiago, who has
made, and still provides, a great contribution to the knowledge
of Dorylaimida biodiversity and morphology.

11

M.T. Vinciguerra et al.

Fig. 3. Prodorylaimus reyesi sp. n. A: Anterior end; B: Female posterior genital branch; C: Neck; D: Male body; E: Female tail; F:
Female body; G: Male posterior end. (Scale bars: A = 20 m; B, C, E, G = 100 m; D, F = 400 m.)
12

Nematology

Two new species of dorylaim from Iran

Fig. 4. Prodorylaimus reyesi sp. n. A, C: Anterior end; B: Neck; D: Cardia region; E: Vulval region; F: Female posterior genital
branch; G: Male posterior end; H: Female posterior end. (Scale bars: A, D, E = 20 m; B, F-H = 50 m; C = 5 m.) This figure
is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/
journals/15685411.

Vol. 00(0), 2016

13

M.T. Vinciguerra et al.

Table 3. Morphometrics of Prodorylaimus reyesi sp. n. All measurements are in m, except for L in mm, and in the form: mean s.d.
(range).
Character

Female

Male

Holotype

Paratypes

Paratypes

n
L (mm)

3.33

37.4

5.5

5.4

c

14.6

V/T

55.0

8
3.07 0.25
(2.78-3.33)
36.9 1.3
(34.8-38.9)
5.3 0.1
(5.1-5.5)
5.7 0.6
(4.6-6.8)
13.2 1.6
(11.7-15.5)
46.4 5.4
(42.3-58.4)
83 7.2
(76-92)
41 1.2
(38-42)
20 0.8
(19-21)
7.6 0.7
(7.0-8.5)
28 1.3
(26-30)
4.2 0.4
(4.0-5.0)
40 2.5
(36-42)
19 0.7
(18-20)
571 41
(499-622)
283 22.3
(252-318)
49.3 1.9
(46.3-52.5)
178 26.8
(140-220)
88 23.9
(60-130)
88 10.7
(70-107)
65 4.5
(58-71)
546 68
(450-634)

3
3.18 0.10
(3.13-3.23)
32.5 2.1
(32-35)
5.3 0.3
(5.1-5.6)
8.5 0.3
(8.3-8.7)
7.3 0.5
(7.0-7.7)
49.1 1.4
(47.7-50.4)
90 7.7
(79-98)
46 4.0
(40-49)
20 0.9
(19-21)
8.2 0.2
(8.0-8.5)
30 1.0
(29-31)
4.5 0.9
(4.0-5.5)
43 4.9
(40-49)
18 0.9
(17-19)
572 41
(530-612)
260 24
(236-284)
45.4 0.9
(44.4-46.4)
215 15
(198-227)
127 15
(117-138)
208 13
(199-217)
72 11
(64-85)
373 3.0
(370-376)
75 1.6
(73-77)

Mid-body diam.

89

Anal/cloacal body diam.

42

Lip region diam.

21

Lip region height

7.5

Odontostyle length

29

Odontostyle width

5.0

Odontophore length

44

Fixed ring from anterior end

22

Neck length

604

Glandularium length

289

Glandularium (% of neck)

47.8

Nerve ring from anterior end

231

Excretory pore from anterior end

144

Pre-rectum length

93

Rectum length

67

Tail length

615

Spicule length

14

Nematology

Two new species of dorylaim from Iran

one 99-169 m long and posterior one 113-159 m long,

oocytes arranged in a single row except near ovary tip
where forming two or more rows, oviduct 109-190 m
long, ending with a pars dilatata, in some specimens appearing as a true spermatheca full of sperm, followed by a
sphincter at its junction with uterus. Uterus 241-328 m
long, bipartite, with a narrow distal part and a wider proximal part which in some specimens contains sperm massed
just after its enlargement. True spermatheca not differentiated. No uterine eggs observed. Vagina extending inwards for 33-43 m, or less than half corresponding body
diam., with pars proximalis anchor-shaped, 21-30 m
long, pars refringens with weakly sclerotised pieces appearing as a continuous ring, 8-11 m long and pars distalis 2-5 m long. Vulva longitudinal. Advulval papillae
present in some specimens, 0-2 anterior and 0-3 posterior
to vulva. Prerectum 2-2.5 anal body diam. long. Tail 1215 anal body diam. long, caudal pores generally hardly
visible, two subdorsal pairs in one specimen.
Male
Reproductive apparatus diorchic, testes opposed. Sperm
5-8 m long. Spicules stout, regularly narrowing at both
ends, with ventral hump poorly developed, 1.5-1.8 anal
body diam. long. In addition to adcloacal pair, a series
of 19-21 contiguous ventromedian supplements present,
sometimes not clearly distinguishable, starting well anterior to range of spicules. Prerectum 4-5 anal body diam.
long, twice as long as in female, its junction with intestine
anterior to supplement series or at level with two anteriormost supplements. Tail 7-8 anal body diam. long, much
shorter than in female, caudal pores well developed, one
pair subventral and one or two pairs subdorsal.
T YPE HABITAT AND LOCALITY
The new species was extracted from the rhizosphere of
hornbeam in Jangal-e Abr (Cloud Forest), Shahrood, Iran
(GPS coordinates: 3645.342 N, 5502.004 E, altitude
2120 m a.s.l.).
T YPE MATERIAL
Holotype female, one female paratype and two male
paratypes deposited in the collection of the Section
of Animal Biology of the Department of Biological,
Geological and Environmental Sciences, University of
Catania, Catania, Italy; one female paratype and one
male paratype deposited in USDA Nematode Collection,
Beltsville, MD, USA. Six female paratypes and two male
Vol. 00(0), 2016

paratypes housed in the Department of Plant Protection,

Faculty of Agriculture, University of Zanjan, Zanjan, Iran.
D IAGNOSIS AND RELATIONSHIPS
Prodorylaimus reyesi sp. n. is characterised by 3.1
(2.8-3.3) mm body length, lip region well set off from
adjoining body by a depression, lips amalgamated with
prominent labial and cephalic papillae, odontostyle 29
(26-31) m long, thicker than cuticle at its level, pharynx
muscular enlarging very gradually, female reproductive
apparatus didelphic, vulva longitudinal, a variable number
of advulval papillae (0-3), both pre- and post-vulval,
present in most specimens, tail in both sexes convexconoid in its anterior part, then narrowing abruptly to a
filiform hyaline appendix, longer in female (c = 13.2
(11.7-15.5)) than in male (c = 7.3 (7.0-7.7)), and 19-21
contiguous ventromedian supplements.
If compared to all the other species of Prodorylaimus,
this species resembles P. longicaudatoides Altherr, 1968,
particularly in the tail length, which is greater in the
female than the male. However, if compared to the
type specimens as re-described by Loof (1985) and to
the populations from Andalusia thoroughly described
by Abolafia et al. (1998), it differs in some important
characters: larger body (L = 3.1 (2.8-3.3) vs 2.2-2.8 mm
in the Spanish populations and 2.4-2.5 mm in the type
specimens); shorter odontostyle (29 (26-31) vs 32-34 m
in the type specimens and 32-38 m in the Spanish ones),
particularly when related to the animal size and lip breadth
(1.4 lip diam. in the new species vs 1.6 in the Spanish
populations); the prerectum length of the new species
is different in the two sexes, being longer in the male,
where it joins the intestine just anterior to the supplements
series or at the level of the two anterior supplements,
whereas in P. longicaudatoides it is of the same size in
both sexes and in the male joins the intestine halfway
along the supplement series. P. longicaudatoides is not
the only species with a different tail length between the
sexes, since this is also true for P. irminii Vinciguerra
& Orselli, 2011 and P. paralongicaudatus (Micoletzky,
1925) Andrssy, 1959. From the former species, P. reyesi
sp. n. differs in its larger body (L = 3.1 (2.8-3.3) vs 2.23
(2.02-2.50 mm)), longer tail (c = 13.2 (11.7-15.5) in
female and 7.3 (7.0-7.7) in male vs 3.5 (3.1-4.2) in female
and 2.2 (1.9-2.6) in male), longer odontostyle (29 (2631) vs 22.4 (19.5-25) m); and from the latter it mainly
differs by having a rather larger body (L = 3.1 (2.8-3.3) vs
1.83-2.57 mm) while the odontostyle is definitely shorter
(29 (26-31) vs 29-39 m) and by having lips distinct and
15

M.T. Vinciguerra et al.

Fig. 5. Phylogenetic tree obtained by the Maximum Likelihood

method. Bootstrap values more than 50% are indicated at branch
nodes. Newly obtained sequences are in bold.

well set off from neck vs lips partly amalgamated and

only slightly set off from neck. From all the other species
of the genus, the new species differs greatly in the main
morphometric characters.
P HYLOGENETIC ANALYSIS
Since sequence data for most species belonging to
Dorylaimus and Prodorylaimus are lacking, as well as for
other genera that appear to be close to them, molecular
analyses devoted to find the best phylogenetic placement
for both new taxa species may be expected to give
unsatisfactory or even misleading results. Basically, either
SSU or LSU data (or both) are available. We discarded
SSU data, since it has been shown that this locus was not
able to give reliable support to phylogenetic studies of
those groups (Holterman et al., 2008). Our analysis was
thus limited to the LSU (i.e., 28S rDNA) locus for which,
however, comparatively few sequences are available.
The optimal tree obtained using the Maximum Likelihood method (Tamura & Nei, 1993) is shown in Figure 5. This analysis involved a total of 12 nucleotide sequences (those of the new species, of the nine closest
taxa and of the outgroup). As can be seen, D. elegans
sp. n. was placed close to D. stagnalis (interspecific sequence variation 0.029) and their genetic differentiation
is well supported by a bootstrap value of 100; while P.
reyesi sp. n. was placed rather close to L. vulvapapilla16

tum (interspecific sequence variation 0.075) and Prodorylaimus HHBM-2007a (interspecific sequence variation
0.078), even though bootstrap values are lower than 70%
and therefore not so appropriate. Similar results were obtained with the Minimum Evolution method (Rzhetsky &
Nei, 1992), which follows an approach not based on an
explicit model of evolution (data not shown). The reason
why the bootstrap replicas gave overall unsatisfactory results in our analysis may reside in the fact that our sequences were somewhat shorter (700-800 bp) than the reference ones (ca 1100 bp).
Recently, Pea-Santiago & Alvarez-Ortega (2014),
based on molecular, morphological and embryological
data, transferred Labronema, together with other roundtailed genera previously attributed to the Qudsianematidae, to the Dorylaimidae, the latter formerly consisting
of only long-tailed dorylaims. Their action is supported
by the present analysis, although itself based on insufficient data, confirming the genetic relatedness between
Labronema on one side and Dorylaimus and Prodorylaimus on the other.

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