Zootaxa 3499: 46–62 (2012) ISSN 1175-5326 (print edition)

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Copyright © 2012 · Magnolia Press
Article ISSN 1175-5334 (online edition)

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A new and three known species of the genus Tylencholaimellus Cobb in M.V.
Cobb, 1915 (Nematoda: Dorylaimida) from Changbai Mountain, China

MIN ZHANG1, SUMAYA AHAD2, M. BANIYAMUDDIN1,2, WENJU LIANG1 & WASIM AHMAD1,2,3
1
State Key Laboratory of Forest and Soil Ecology, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110164,
China
2
Section of Nematology, Department of Zoology, Aligarh Muslim University, Aligarh- 202 002 India
3
Corresponding author. E-mail: ahmadwasim57@yahoo.co.in

Abstract

A new and three known species of the genus Tylencholaimellus Cobb in M.V. Cobb, 1915 are described from Changbai
Mountain, Northeast China. Tylencholaimellus sinensis n. sp. species is characterized by having a medium sized body (L=
1.1–1.3 mm); outer cuticle with fine transverse striations; inner layer distinctly striated; radial refractive elements
abundant; labial disc absent; lip region offset by constriction; lips rounded and amalgamated; odontostyle 18–19 µm,
odontophore 7–8 µm, combined length 25–27 μm; anterior uterine sac 0.8–1.6 mid body diameters long; mono-
opisthodelphic female genital system with small anterior uterine sac; transverse vulva and short conoid tail.
Tylencholaimellus striatus Thorne, 1939, T. montanus Thorne, 1939 and T. cinctus Orr & Dickerson, 1965 are described
for the first time from this region of the world.

Key words: Changbai Mountain, China, known species, new species, taxonomy, Tylencholaimellus

Introduction

Changbai Mountain (126o 55’–129o 00’ E; 41o 23’–42o 36’ N) is located in the northeast part of China at the border
with North Korea. Altitudes range from 720 to 2691 m above sea level (a.s.l.), mean annual temperature from 3 to
-7oC, and mean annual precipitation from 679 to 1330 mm. Vegetation consists of deciduous forest, mixed
coniferous-broadleaved forest, dark-coniferous spruce-fir forest, dark coniferous spruce forest, birch forest and
alpine tundra. The plant species comprise European, Siberian and central Chinese species, and include 1,800
species of vascular plants (Zhang et al., 2003; Zhao et al., 2004; Tong et al., 2010). The variations in climate and
vegetation of Changbai Mountain make it a special habitat for soil-inhabiting nematodes. As no data on the
mematode fauna in this habitat are available, we plan to carry out a thorough systematic study of soil-inhabiting
nematodes from Changbai Mountain. The present paper is the first in a projected series, documenting three known
and a new species of the genus Tylencholaimellus Cobb in M.V. Cobb, 1915 from this region.

Material and methods

The nematodes were extracted from soil samples by Cobb’s (1918) sieving and decantation technique and by a
modified Baermann’s funnel technique. Nematodes so obtained were killed and fixed with hot 4% formalin,
dehydrated to glycerin by a slow evaporation method, and mounted in anhydrous glycerin on slides. Measurements
were taken using an ocular micrometer and line illustrations were made using a drawing tube attached to a Nikon
Optiphot-2 microscope. Photographs were taken with a digital camera attached to a Nikon Eclipse 80i DIC
microscope.

46 Accepted by P. Mullin: 5 Sept. 2012; published: 27 Sept. 2012

Tylencholaimellus sinensis n. sp.
(Figs. 1, 2)

Measurements: Table 1.

TABLE I. Tylencholaimellus sinensis n. sp.; all measurements in µm.

Characters Holotype female Paratype females
Mean ± SD (Range)
n 1 5
L 1242 1169± 53.9 (1103–1256)
Body width at neck base 42 42.4± 2.0 (39–45)
Body width at mid body 47 46.8± 2.9 (43–52)
Body width at anus 41 33.4± 4.3 (29–41)
a 28.6 25± 1.4 (24–27)
b 7.7 6.3± 0.3 (5.8–6.7)
c 44.8 39.5± 1.86 (36.7–42)
c` 0.88 0.89± 0.07 (0.78–0.96)
V 23.7 29.5± 1.2 (27.8–31.0)
G1 3.56 5.1± 1.2 (3.0–6.8)
G2 27.5 23.6± 7.2 (15.3–34.7)
Lip region width 15 14.6± 0.8 (14–16)
Lip region height 5 5.8±0.4 (5–6)
Amphid width 7 6.9±0.6 (6–8)
Odontostyle length 19.5 18.4± 0.48 (18–19)
Odontophore length 8 7.2± 0.4 (7–8)
Guiding ring from anterior end 12 12.2± 1.9 (10–15)
Nerve ring from anterior end 100 93.8± 6.6 (85–104)
Neck length 173 185± 16.9 (167–217)
Expanded part of Pharynx 32 27.2± 1.7 (25–30)
Cardia length 9 9± 1.2 (7–10)
Anterior genital branch 48 60.2± 13.7 (36–75)
Posterior genital branch 370 294±87 (192–390)
Vaginal length 22 19.2± 1.9 (17–22)
Vulva from anterior end 319 346± 22 (322–385)
Prerectum length 150 126± 21 (90–146)
Rectum Length 37 35.2± 2.9 (30–38)
Tail length 30 29.6±1.3 (28–32)

Description. Female: Medium sized nematodes, slightly curved upon fixation, tapering gradually towards the
anterior end. Cuticle with two distinct layers, 3–4 μm thick at mid body and 4–5 μm on tail. Outer cuticle finely
transversely straited, inner layer with distinct striations; radial refractive elements abundant. Lateral chords
occupying about one-sixth of the midbody diameter. Lateral body pores 6–10 in pharyngeal region, 6–9 between
pharynx and vulva and 31–41 between vulva and anus. Ventral body pore one, 11–19 μm behind the hemizonid.
Dorsal pores indistinct. Lip region offset from body by constriction, 2.3–3 times as wide as high or one third of the
body diameter at neck base. Lips rounded and amalgamated; labial and cephalic papillae indistinct. Amphid cup-
shaped with aperture located at the level of cephalic constriction and occupying about two-thirds of lip region
diameter. Stoma a truncate cone. Odontostyle 1.1–1.35 times the lip region width long and provided with dorsal

TYLENCHOLAIMELLUS FROM CHINA Zootaxa 3499 © 2012 Magnolia Press · 47

accessory piece covering entire dorsal arm; its aperture one-sixth of the odontostyle length. Odontophore with
basal knobs; 0.3–0.4 times the odontostyle length. Guiding ring simple, refractive at 0.6–1.07 times lip region
width from anterior end. Pharynx consists of a slender and weakly muscular anterior part and a short pear-shaped
constricted basal bulb, which occupies about 13.8–18.4% of the total neck length. Pharyngeal glands often visible.
Cardia tongue-shaped, about one-fifth of the corresponding body width long. Nerve ring located at 46–58% of the
neck length. Hemizonid posterior to nerve ring, located at 96–108 μm from anterior region. Genital system
monodelphic-opithodelphic. Ovary reflexed, measuring 72–149 µm long, not reaching the oviduct-uterus junction,
oocytes arranged in single row except near the tip. Oviduct joining the ovary subterminally, measuring 127–296
µm, its proximal and distal parts not differentiated. Oviduct-uterus junction marked by well developed sphincter.
Uterus short and tubular, measuring 54–75 µm. Sperm present throughout the genital tract. Anterior genital branch
reduced to a simple sac, 0.8–1.6 midbody diameter long. Vagina cylindrical, pars proximalis vaginae 11–13 µm
long, its wall encircled by muscles. Pars distalis vaginae short, 2.5–3 µm long with slightly curved walls; pars
refringens absent. Vulva apparently a transverse slit. Prerectum 2.5–4.6 anal body diameters long. Rectum
1.02–1.2 anal body widths long. Tail short conoid, 1.03–1.2 anal body widths long.
Male: Not found.
Type habitat and locality: Mixed coniferous broad-leaved forest, Changbai Mountain, Northeast China, P.R.
China (42°24′N, 128°06′E): 760m a.s.l.; annual mean temperature 2.6 ºC, annual mean precipitation 691 mm; tree
species were dominated by a mixture of coniferous Pinus koraiensis, broadleaved Quercus spp. and Acer spp. on
Albi-Boric Argosols.
Type specimens: Holotype female on slide Tylencholaimellus sinensis n. sp./1; paratype females on slides
Tylencholaimellus sinensis n. sp./2–3; deposited in the nematode collection of the Department of Zoology, Aligarh
Muslim University, India.
Etymology: The new species is named sinensis because of its origin in China.
Diagnosis and relationships: Tylencholaimellus sinensis n. sp. species is characterized by having a medium
sized body (L = 1.1–1.3 mm); outer cuticle with fine transverse striations, inner cuticle with distinct striations;
labial disc absent; lip region offset by constriction; lips rounded and amalgamated; odontostyle 18–19 µm,
odontophore 7–8 µm, combined length of spear 25–27 μm; anterior uterine sac 0.8–1.6 midbody diameters long;
mono-opisthodelphic female genital system with small anterior uterine sac; transverse vulva and short conoid tail.
In the presence of an offset lip region without labial disc, and comparatively long spear, the new species comes
close to T. polonicus Szczygiel, 1962; T. pluvialis Siddiqi, 1965; and T. sayi Goseco et al. 1975. However, it differs
from T. polonicus in having a truncate-conical stoma not swollen in the middle (vs stoma a truncate cone slightly
wider in its central portion); lip region offset by rather shallow constriction, lips more rounded with indistinct labial
and cephalic papillae (vs lip region offset by very deep constriction, lips more angular with very distinct labial and
cephalic papillae); both parts of pharynx separated by a constriction (vs both parts of pharynx separated by a more
or less marked depression); comparatively smaller pharyngeal bulb (vs pharyngeal bulb 30–37 µm long); larger
cardia (vs 3 µm Goseco et al. (1975) and 4.5–6.5 µm Peralta & Peña-Santiago, (1995), and deeper vagina (11–12
µm for Goseco et al. (1975), 12–17 µm in Peralta & Peña-Santiago (1995)).
From T. pluvialis, the new species differs in having larger body size (L= 1.10–1.25 vs 0.80–0.89 mm), in the
shape and size of lip region (vs lip region 9–10 µm wide with six inner liplets around oral aperture) and
comparatively longer tail (vs tail 14–26 µm long). Goseco et al. (1975) recorded few specimens of T. pluvialis from
Indiana. Their specimens distinctly differ from the original description in having shorter odontostyle (14.4 vs 18
µm) and shorter prerectum (1.5 vs 3 anal body width). Although the total length of spear was same in both the
populations, the ratio between odontostyle and odontophore length is distinctly different and hence the Indiana
population probably represents a different species.
From T. sayi Goseco et al. 1975, the new species distinctly differs in the shape and size of lip region (vs lip
region narrow, 9 µm wide with low lips and with distinct sclerotization of the oral opening); comparatively anterior
vulva (V= 27.8–31 vs 31–33); longer tail (tail 28–32 vs. 21–22 µm long, c = 36.7–42 vs 44–45) and longer
prerectum (90–106 vs 40–48 µm).

48 · Zootaxa 3499 © 2012 Magnolia Press ZHANG ET AL.

FIGURE 1. Tylencholaimellus sinensis n. sp. A: Entire female. B: Anterior end. C: Anterior end showing amphid. D:
Pharyngeal region. E: Expanded part of pharynx. F: Female genital system. G: Vulval region. H: Female posterior region.

TYLENCHOLAIMELLUS FROM CHINA Zootaxa 3499 © 2012 Magnolia Press · 49

FIGURE 2. Tylencholaimellus sinensis n. sp. Anterior region. B: Anterior end showing amphid. C: Expanded part of
pharynx. D: Vulval region. E: Female posterior region. F: Female posterior end(scale bar A–D, F,G: 20 µm; E: 50 µm).

50 · Zootaxa 3499 © 2012 Magnolia Press ZHANG ET AL.

Tylencholaimellus striatus Thorne, 1939
(Figs. 3, 4)

Measurements: Table 2.

TABLE 2. Tylencholaimellus striatus Thorne, 1939; all measurements in µm.

Characters Females
Mean ± SD (Range)
n 5
L (mm) 718±52 (674–791)
Body width at neck base 24.2±1.5 (23–27)
Body width at mid body 26.8±2.6 (24–31)
Body width at anus 15.6±1.3 (14–18)
a 26.9±1.5 (25–28)
b 5.5±0.3 (5–6)
c 27.6±2.3 (25.0–31.6)
c` 1.6±0.12 (1.5–1.8)
V 33.3±2.15 (30.7–37)
G1 1.06±0.10 (0.9–1.18)
G2 14.8±2.2 (10.75–17.2)
Lip region width 6.9±0.2 (6.5–7)
Lip region height 3.4±0.48 (3–4)
Amphid aperture 3.7±0.25 (3.5–4)
Odontostyle length 8.6±0.48 (8–9)
Odontophore length 7.6±0.5 (7–8)
Guiding ring from anterior end 6±06–6
Nerve ring from anterior end 70.6±3.5 (65–74)
Neck length 130±10.3 (113–143)
Expanded part of pharynx 27±2.3 (25–31)
Cardia length 5.8±1.1 (4–7)
Anterior genital branch 7.6±0.48 (7–8)
Posterior genital branch 107±21.9 (73–133)
Vaginal depth 11.2±0.4 (11–12)
Vulva from anterior end 239±25 (215–286)
Prerectum length 81.4±7.4 (73–94)
Rectum length 16±0.8 (15–17)
Tail length 26±0.8 (25–27)

Description. Female: Moderately slender nematodes of small size. Body cylindrical, slightly tapering towards
both extremities. Body slightly curved upon fixation. Cuticle with two distinct layers, 2 µm thick at mid body and
3 µm on tail. Outer cuticle with distinct transverse striations; inner layer thick with prominent transverse striations;
radial refractive elements abundant. Lateral body pores obscure. Lateral chords occupying about one-fifth to one-
fourth of the mid body diameter. Lip region slightly offset by a distinct constriction, 1.7–2.3 times as wide as high
and one-third of body diameter at neck base. Lips amalgamated, with rounded contour; its inner portion
transformed into a perioral disc. Amphid cup-shaped, opening at level of the cephalic constriction and occupying
one-half of the lip region diameter. Stoma a truncate cone. Odontostyle 1.1–1.3 times the lip region width long and
provided with dorsal accessory piece covering the entire dorsal arm; its aperture one-sixth of the odontostyle

TYLENCHOLAIMELLUS FROM CHINA Zootaxa 3499 © 2012 Magnolia Press · 51

FIGURE 3. Tylencholaimellus striatus Thorne, 1939. A: Entire female. B: Anterior region. C: Anterior end showing amphid.
D: Pharyngeal region. E: Expanded part of pharynx. F: Female genital system. G: Female posterior region.

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FIGURE 4. Tylencholaimellus striatus Thorne, 1939. A: Anterior region. B: Anterior end showing amphid. C: Expanded part
of pharynx. D: Female genital system. E: Vulval region. F: Female posterior region. G: Female posterior end (scale bar: A–B:
10 µm; C–G: 20 µm).

TYLENCHOLAIMELLUS FROM CHINA Zootaxa 3499 © 2012 Magnolia Press · 53

length. Odontophore with basal knobs; 0.7–1 times the odontostyle length. Guiding ring simple, refractive, at
0.8–0.9 times lip region width from anterior end. Pharynx consist of a slender and weakly muscular anterior part
and separated from basal bulb by a constriction. Pharyngeal basal bulb cylindroid, which occupies about 19.5–22%
of the total neck length. Pharyngeal glands often visible. Cardia rounded to hemispherical, surrounded by the
intestine. Nerve ring located at 50–57% of the neck length. Genital system monodelphic-opisthodelphic. Ovary
reflexed, measuring 51–107 µm long, and provided with oocytes arranged in single row except near the tip.
Oviduct joining the ovary subterminally, measuring 41–72 µm and consisting of a slender distal part and a poorly
developed pars dilatata. Oviduct-uterus junction marked by a weak sphincter. Uterus short and tubular, measuring
23–36 µm long. Sperms not seen either in oviduct or uterus. Anterior genital branch a rudimentary cell mass with
inconspicuous lumen. Vagina cylindrical, pars proximalis vaginae 5–6 µm long, its wall encircled by muscles.
Pars distalis vaginae short, 1.5–2.0 µm long with slightly curved walls; pars refringens absent. Vulva apparently a
transverse slit. Prerectum 4–6 anal body diameters long. Intestine-prerectum junction guarded by three cells.
Rectum 0.9–1.0 anal body widths long. Tail cylindroid and relatively long; outer layer of cuticle thickened in its
posterior portion forming a distinct hyaline layer.
Male: Not found
Habitat and locality: Dark-coniferous spruce-fir forest, Changbai Mountain, Northeast China (42°09′N,
128°08′E): 1250m a.s.l., annual mean temperature 0.3ºC, annual mean precipitation 811 mm; tree species
dominated by Abies nephrolepis and Picea jezoensis on Bori-Udic Cambosols.
Remarks: Thorne (1939) described T. striatus from Utah, U.S.A and he (1974) further reported this species
from South Dakota. Goseco et al. (1975) restudied the Thorne’s original specimens, provided a detailed description
and also designated lectotype and paralectotype. The morphometrics of the present population confirms well with
the type population as well as the South Dakota population, except for having outer portion of lips rather rounded
(vs lip region very angular); odontostyle slightly longer than odontophore (vs odontostyle as long as odontophore);
cardia rounded to hemispheroid (vs cardia flat), and slightly longer tail (vs c = 35). Thorne in both the descriptions
mentioned anterior uterine sac as absent while Goseco et al. in their redescription reported a small anterior uterine
sac of about 7 µm. In our present specimens a small anterior uterine sac could be recognized. Our present
population also agrees fairly well with Spanish population described by Peralta & Peña-Santiago (1995) except for
having pharyngeal bulb occupying about one-fifth of the total pharyngeal length (vs occupying one-fourth of the
total pharyngeal length), prerectum slightly long (vs 2–4 anal body diameters long); male absent (vs male present).
The present population also agrees fairly well with Indian population described by Mushtaq, Naz & Ahmad (2007).

Tylencholaimellus montanus Thorne, 1939

(Figs. 5, 6)

Measurements: Table 3.
Description. Female: Moderately slender nematodes of small size. Body cylindrical, curved ventrad upon
fixation, slightly tapering towards both extremities. Cuticle with two distinct layers, 2.0–2.5 µm thick at mid body
and 3 µm on tail. Outer cuticle with fine transverse striations; inner cuticle distinctly striated; radial refractive
elements abundant. Lateral, dorsal and ventral body pores indistinct. Lateral chords occupying about one-fifth of
mid body diameter. Lip region slightly offset by a weak constriction, 1.5–2.0 times as wide as high and one-third of
the body diameter at neck base. Lips amalgamated, with rounded contour; its inner portions transformed into a
perioral disc. Amphid cup-shaped, opening at level of the cephalic constriction and occupying about two-thirds of
the lip region diameter. Stoma a truncate cone. Odontostyle 1.2–1.5 times the lip region width long and provided
with dorsal accessory piece covering the entire dorsal arm; its aperture one-fifth of the odontostyle length.
Odontophore with basal knobs; 0.7–0.8 times the odontostyle length. Guiding ring simple, refractive, at 0.7–0.9
times lip region width from anterior end. Pharynx consists of a slender and weakly muscular anterior part separated
from a cylindrical basal bulb by a depression, which occupies about 18.6–23.8% of the total neck length.
Pharyngeal gland often visible. Cardia hemispherical, partially surrounded by the intestine. Nerve ring at 48–61%
of the neck length. Genital system monodelphic-opithodelphic. Ovary reflexed, measuring 40–90 µm long; oocytes
arranged in single row except near its tip. Oviduct joining the ovary subterminally, measuring 53–88 µm and
consisting of a slender distal part and a poorly developed pars dilatata. Weakly developed sphincter present at

54 · Zootaxa 3499 © 2012 Magnolia Press ZHANG ET AL.

oviduct-uterus junction. Uterus short and tubular, measuring 25–37 µm; sperms not present in either uterus or
oviduct. Anterior genital branch practically vestigial, almost always appearing as a small cell mass with
inconspicuous lumen. Vagina cylindrical, pars proximalis vaginae 9–10 µm long, its wall encircled by muscles.
Pars distalis vaginae short, 0.5–1.0 µm long with slightly curved walls; pars refringens absent. Vulva apparently a
transverse slit. Prerectum 3.4–4.7 times anal body diameter long. Intestine-prerectum junction guarded by three
cells. Rectum 0.9–1.2 anal body diameter long. Tail cylindroid and relatively long; outer layer of cuticle thickened
posteriorly forming a distinct hyaline layer.
Male: Not found.
Habitat and locality: Mixed coniferous broad-leaved forest, Changbai Mountain, Northeast China (42°24′N,
128°06′E): 760m a.s.l., annual mean temperature 2.6 ºC, annual mean precipitation 691 mm; tree species were
dominated by a mixture of coniferous Pinus koraiensis, broadleaved Quercus spp. and Acer spp. on Albi-Boric
Argosols.

TABLE 3. Tylencholaimellus montanus Thorne, 1939; all measurements in µm.

Characters Females
Mean ± SD (Range)
n 7
L (mm) 645±47.5(593–755)
Body width at neck base 22±0.5 (21–23)
Body width at mid body 23.8±0.6 (23–25)
Body width at anus 15±0.7 (14–16)
a 26.7±2.7 (24–33)
b 4.6±0.4 (4.0–5.2)
c 26.6±1.5 (25–30)
c` 1.5±0.04 (1.5–1.6)
V 35.8±2.0 (33–39)
G1 1.66±0.12 (1.5–1.9)
G2 15.4±1.8 (13–18)
Lip region width 6.57±0.31 (6–7)
Lip region height 3.8±0.2 (3.5–4.0)
Amphid aperture 3.8±0.2 (3.5–4.0)
Odontostyle length 9±0
Odontophore length 7. 14±0.34 (7–8)
Guiding ring from anterior end 5.14±0.34 (5–6)
Nerve ring from anterior end 75.8±5.2 (69–83)
Neck length 139±5.7 (132–147)
Expanded part of pharynx 30±2.5 (25–33)
Cardia length 4.3±0.45 (4–5)
Anterior genital branch 10.8±0.9 (9–12)
Posterior genital branch 99±10.5 (83–114)
Vaginal depth 13±0.92 (12–14)
Vulva from anterior end 231±16.5 (210–257)
Prerectum length 58.7±4.65 (51–66)
Rectum length 17.3±1.6 (15–20)
Tail length 24±0.9 (22.5–25)

TYLENCHOLAIMELLUS FROM CHINA Zootaxa 3499 © 2012 Magnolia Press · 55

FIGURE 5. Tylencholaimellus montanus Thorne, 1939. A: Entire female. B: Anterior region. C: Anterior end showing
amphid. D: Pharyngeal region. E: Expanded part of pharynx. F: Female genital system. G: Female posterior region.

56 · Zootaxa 3499 © 2012 Magnolia Press ZHANG ET AL.

FIGURE 6. Tylencholaimellus montanus Thorne, 1939. A: Anterior region. B: Anterior end showing amphid. C: Expanded
part of pharynx. D: Female genital system. E–F: Female posterior region (scale bar: 20 µm).

TYLENCHOLAIMELLUS FROM CHINA Zootaxa 3499 © 2012 Magnolia Press · 57

 Remarks: Thorne (1939) described this species from Utah and differentiated it from the closely related
species, T. striatus Thorne, 1939 mainly in having somewhat more slender body, less strongly striated subcuticle
and a more conoid rather than hemispheroid tail. Andrássy (1967) identified a single female specimen of this
species from Argentina which had a comparatively longer tail (2 anal body widths). Goseco et al. (1975) in their
revision of the genus Tylencholaimellus examined the type specimens, redescribed this species and also designated
a lectotype and paralectotype. In their description, the tail shape of the female is conoid (Figs. 10EF of Goseco et
al., 1975), the a ratio is 27–30 and they also mentioned the presence of body pores. Peralta & Peña-Santiago (1995)
recorded three populations of T. montanus from Spain. Their specimens had a values ranging from 19–28 and
rounded-conoid to cylindroid tails, mostly dorsally convex but sometimes ventrally almost straight. They
redescribed T. striatus also in the same paper and differentiated the two species mainly in the shape of lip region,
being scarcely offset in T. montanus, but more strongly offset in T. striatus, which also has a cylindroid tail with a
thick hyaline outer layer in the terminal portion. Mushtaq, Naz & Ahmad (2007) also mentioned a thick terminal
tail cuticle in T. striatus. The present Chinese specimens conform well to the earlier descriptions in their
morphometrics. The lip region is offset by a depression rather than constriction and the tail shape is more like
Spanish populations, and quite different from the original American specimens. In our specimens the prerectum is
also slightly longer (vs 31–56 µm or 2–3 anal body widths long). These differences could be due to geographical
variation.
The two species, T. striatus and T. montanus are morphometrically very similar to each other but still display
some definite morphological differences which clearly differentiate these two species. These include: the shape of
lip region, being scarcely offset in T. montanus (vs strongly offset), distinctly striated subcuticle (vs strongly
striated subcuticle, the striation being more pronounced in pharyngeal region), and tail shape. The amphids are
comparatively wide in T. montanus, almost occupying the entire lateral surface at that level (vs about two-thirds lip
region width in T. striatus).

Tylencholaimellus cinctus Orr & Dickerson, 1965

(Figs. 7, 8)

Measurements: Table 4.
Description. Female: Moderately slender nematodes of small size, slightly curved upon fixation, tapering gradually
towards the anterior end. Cuticle with two distinct layers, 2 μm thick at mid body and 3 μm on tail. Outer cuticle with fine
transverse striations, inner layer with distinctly striated with abundant radial refractive elements. Lateral chords occupying
about 22% of the midbody diameter. Lateral body pores distinct, 8 in pharyngeal region, 5 between pharynx to vulva, 23
between vulva to tail; dorsal and ventral body pores indistinct.
Lip region cape-like, offset from body by a less marked constriction, about twice as wide as high or one-third
as wide as body diameter at neck base. Lips rounded and amalgamated; labial and cephalic papillae indistinct.
Amphid cup-shaped with aperture located at the level of cephalic constriction and occupying about two-thirds of
lip region diameter. Stoma truncate cone, weakly sclerotized. Odontostyle 1.5 times the lip region width long and
provided with dorsal accessory piece covering the entire dorsal arm; its aperture one-seventh of odontostyle length.
Odontophore with basal knobs; 0.5 times the odontostyle length. Guiding ring simple, refractive, at about one lip
region width from anterior end. Pharynx consists of a slender and weakly muscular anterior part and a short pear-
shaped constricted basal bulb, which occupies about 14.6% of the total neck length. Pharyngeal gland often visible.
Cardia hemispherical, about one-fourth of the corresponding body width long. Nerve ring located at 61% of the
neck length. Genital system monodelphic-opithodelphic. Ovary reflexed, measuring 137 µm long, not reaching the
oviduct-uterus junction, oocytes arranged in single row except near the tip. Oviduct joining the ovary subterminally
and consisting of a slender distal part and a well developed pars dilatata. Oviduct-uterus junction marked by
poorly developed sphincter. Uterus short and tubular. Sperms present throughout the genital tract. Anterior genital
branch reduced to a simple sac, 1.15 mid body diameters long. Vagina cylindrical, pars proximalis vaginae 10 µm
long, its wall encircled by muscles. Pars distalis vaginae short, 1.5 µm long with slightly curved walls; pars
refringens absent. Vulva apparently a transverse slit. Prerectum 3 anal body diameter long. Rectum 0.95 anal body
width long. Tail short, 0.9 anal body width long, rounded-conoid.
Male: General morphology similar to female but the posterior body region is more ventrally curved. Diorchic
with opposed testes. Sperms spindle-shaped. Apart from the adcloacal pair only one ventromedian supplement is

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present, located outside the spicule range and separated from adcloacal pair by slightly more than two times the
distance of the adcloacal pair to the anus. Spicules 1.6–1.8 times anal body width long. Lateral guiding pieces one-
fifth of the spicule length. Tail slightly shorter than that of the female and with a more concave ventral side.

TABLE 4. Tylencholaimellus cinctus Orr and Dickerson, 1965

Characters Female Males
n 1 2
L (mm) 0.77 0.73, 0.66
Body width at neck base 28 24, 25
Body width at mid body 32 34, 27
Body width at anus 21 18, 16
a 24.3 21.7, 24.4
b 5.1 5, 5.6
c 41 35, 44
c` 0.90
V 28.6
G1 4.7
G2 29.7
Lip region width 9 9, 9
Lip region height 4 4,4
Amphid aperture 5 5, 5
Odontostyle length 14 12, 12
Odontophore length 7 6, 7
Guiding ring from anterior end 9 7, 7
Nerve ring from anterior end 91 86, 65
Neck length 150 148, 116
Expanded part of pharynx 22 24, 19
Cardia length 7 6, 8
Anterior genital branch 22
Posterior genital branch 232
Vaginal depth 13
Vulva from anterior end 223
Prerectum length 65 73, 70
Rectum length 20 31, 28
Tail length 19 21, 15
Spicule length 30, 27
Lateral guiding pieces 7, 8
Ventromedian supplements 1, 1

Habitat and locality: Mixed coniferous broad-leaved forest, Changbai Mountain, Northeast China (42°24′N,
128°06′E): 760m a.s.l.; annual mean temperature 2.6ºC, annual mean precipitation 691 mm; tree species were
dominated by a mixture of coniferous Pinus koraiensis, broadleaved Quercus spp. and Acer spp. on Albi-Boric
Argosols.
Remarks: Orr & Dickerson (1965) described this species from Kansas, USA. Goseco et al. (1975) provided a
redescription of this species based on the study of type material. Peralta & Peña-Santiago (1996) recorded this
species from several localities in Spain and provided a redescription. The present specimens from China conform
well to the earlier descriptions. Peralta & Peña-Santiago described the pharyngeal bulb as expanding abruptly

TYLENCHOLAIMELLUS FROM CHINA Zootaxa 3499 © 2012 Magnolia Press · 59

FIGURE 7. Tylencholaimellus cinctus Orr & Dickerson, 1965 . A: Entire female. B: Entire male. C: Anterior end. D:
Anterior end showing amphid. E: Pharyngeal region. F: Expanded part of pharynx. G: Female genital system. H: Female
posterior region. I: Male posterior region.

60 · Zootaxa 3499 © 2012 Magnolia Press ZHANG ET AL.

FIGURE 8. Tylencholaimellus cinctus Orr & Dickerson, 1965 . A: Anterior region. B: Anterior end. C: Anterior end
showing amphid. D–E: Expanded part of pharynx. F: Vulval region. G: Female genital system. H–I: Female posterior region.
J–K: Male posterior region (scale bar: 20 µm).

TYLENCHOLAIMELLUS FROM CHINA Zootaxa 3499 © 2012 Magnolia Press · 61

whereas in our specimens as well as the earlier description of Orr & Dickerson (1965) and Goseco et al. (1975) it
has been indicated as offset by constriction. In our single female specimen, the vulva is slightly anterior compared
to earlier descriptions (V= 28.5 vs 31–41). These minor differences could be considered as geographical variations.

Acknowledgments

This research was conducted at the Research Station of Changbai Mountain Forest Ecosystems, Chinese Academy
of Sciences. This work was financially supported by the National Natural Science Foundation of China (Nos.
31170577 and 31170484). WA and MB are thankful to the Chinese Academy of Science for inviting them for a
short duration to work at Institute of Applied Ecology, Chinese Academy of Sciences.

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