Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 October 2023 | 15(10): 24063–24078
ISSN 0974-7907
(Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.8420.15.10.24063-24078
#8420 | Received 23
February 2023 | Final received 05 June 2023 | Finally accepted 27 September
2023
A compendium of Aphelenchoides (Fischer, 1894) (Nematoda: Tylenchina: Aphelenchoidea)
nematodes with the description of a new species from Manipur, India
Loukrakpam Bina Chanu
1 & Naorem
Mohilal 2
1,2 Department of Zoology, Manipur
University, Canchipur, Manipur 795003, India.
1 bina.chanu@gmail.com (corresponding
author), 2 nmohilal@manipuruniv.ac.in
ZooBank: urn:lsid:zoobank.org:pub:517493EB-10A5-4707-A288-4557240ED5EB
Editor: Biplob K. Modak, Sidho Kanho
Birsha University, West Bengal, India.
Date of publication: 26 October 2023 (online & print)
Citation: Chanu, L.B. & N. Mohilal
(2023).
A compendium of Aphelenchoides (Fischer, 1894)
(Nematoda: Tylenchina: Aphelenchoidea)
nematodes with the description of a new species from Manipur, India. Journal of Threatened Taxa 15(10):
24063–24078. https://doi.org/10.11609/jott.8420.15.10.24063-24078
Copyright: © Chanu & Mohilal 2023. Creative Commons Attribution 4.0
International License. JoTT allows unrestricted use, reproduction, and
distribution of this article in any medium by providing adequate credit to the
author(s) and the source of publication.
Funding: The study was made under the funding agency of SERB, New Delhi with file no. SB/FT/LS-113/2013.
Competing interests: The authors declare no competing interests.
Author details: L. Bina chanu is a guest faculty at the Department of Life Sciences (Zoology), Manipur University, Canchipur. She is working on biodiversity of nematode parasites including ectoparasites. N. Mohilal is a professor, Department of Life Sciences (Zoology), Manipur University, Canchipur. He works on the biodiversity of soil nematodes and their bio-control methods using medicinal plants, biodiversity of Annelids, snakes and birds.
Author contributions: LBC conducted the survey, collected samples, processed the samples, extracted nematodes, identified the nematodes, did statistical calculations and wrote the manuscript.
NM helped in designing the survey and collection, in the identification of the nematodes, and in final proofing of the manuscript.
Acknowledgements: The authors are thankful to the Head of Department,
Department of Zoology, Manipur University, Canchipur,
Manipur, India for providing necessary laboratory facilities and the Science
& Engineering Board, New Delhi for the funding with the no.
SB/FT/LS-113/2013.
Abstract: The present compendium is based
on the findings of a research work on the survey of nematodes belonging to the
family Aphelenchoidea in the northeastern states of
India and the literature available on this particular species, mainly from
Manipur. During the study, a total of 12 Aphelenchoides
spp. were found, among which six species were reported for the first time from
Manipur. A new species, Aphelenchoides oryzae is also described in the present article. The
present study will help in making us understand the biodiversity status of Aphelenchoides nematodes in the region.
Diagnosis of the species and illustrations along with dichotomous keys are
provided in the manuscript.
Keywords: Aphelenchid, Coconut tree, fungivore, food-web,
soil fertility, soil dwelling, Pinus sp., Morus
sp., Orange plant and species richness.
INTRODUCTION
Forest conservation improves
ecosystem functions and will help to protect natural biodiversity. Plant
communities are the critical indicators for forest restoration. Below-ground
diversity relates very closely with above-ground biodiversity. Plant parasitic nematodes
are found in every soil of varied ecosystems. Nematodes are often sensitive to
habitat disturbance, showing the characteristic sequence of recolonization
after disturbance. Furthermore, they are represented in a wide array of trophic
groups as herbivores (Tylenchid), bacterivores &
fungivores (Aphelenchid), omnivores (Dorylaimid), and predators (Mononchid)
reflecting resource availability and changes of environmental conditions in the
soil providing information on succession and changes in decomposition pathways
in the soil food – web, nutrient status & soil fertility, acidity, and the
effects of soil contaminants (Yeates & Bongers 1999).
Bacterial as well as fungal
feeding nematodes like aphelenchids have a high
carbon: nitrogen (C:N) ratio (±5.9) than their
substrate (±4.1), consuming bacteria, they take in more N than necessary for
their body structure. The excess nitrogen is excreted as ammonia. The bacterial
and fungal feeding nematodes’ community in the top 15 cm of the field soil
mineralizes N at rates increasing to 1.01µg - N g – soil -1d - 1 in the
rhizosphere (Ferris et al. 1995; 1996; 1997).
Aphelenchid nematodes have diverse habitats.
Several aphelenchids are associated with insects,
some spending a part of their life - cycle in insects besides being
phytophagous, while others are mycetophagous. Some
forms of aphelenchids are true plant parasites and
are, therefore, economically significant. Of the available aphelenchid
nematodes, three species are major pests of agricultural and horticultural
crops, i.e., white-tip nematode Aphelenchoides
besseyi; Red-ring nematode of coconut Rhadinaphelenchus cocophilus,
and pine-wilt nematode Bursaphelenchus xylophilus respectively in the world. So far, 138
species of Aphelenchoides (Fischer,
1894) have been identified, of which India contributed more than 12 species.
Recognizing the importance of
bacterial and fungal decomposition in forest ecosystems, survey for Aphelenchid nematodes is very important in every region of
the Earth. Recently, a survey was conducted on the Aphelenchoides
nematodes in the different ecosystems of the north-eastern region of India.
During the work, 12 Aphelenchoides were
encountered from Manipur, among which six species were recorded for the first
time from this particular region. The species encountered were Aphelenchoides aligarhiensis
(Siddiqi et al., 1967); A. baquei (Maslen,
1978); A. confusus (Thorne & Malek, 1968);
A. minor (Seth & Sharma, 1986); A. swarupi (Seth & Sharma, 1986); A. vigor (Thorne
& Malek, 1968); A. dhanachnadi (Chanu et al., 2012) and A. neoechinocaudatus
(Chanu et al., 2012); A. aerialis
(Chanu et al., 2015); A. longistylus
(Chanu & Mohilal,
2014); A. neominoris (Chanu
& Mohilal, 2014), and A. manipurensis
(Chanu &
Mohilal, 2018). These species are presented
along with their dichotomous keys.
METHODS
Study site
Manipur has a
geographical area of 22,327 km2, situated at an altitude of 790 m
with a longitude of 24.733333333333334 and a latitude of 93.9666666666667
(Figure 1). Soil samples were collected
from different localities of Manipur.
Methodology
For
collection of soil samples around the rhizospheric
region of a particular host, 500 g of soil around the plant from 8 different
sides were taken. The soil was mixed together thoroughly. From the thoroughly
mixed soil, again 500 g were taken, serving as the sample soil for a particular
host plant or tree. The samples were processed for extraction of nematodes
through the Cobb’s (1918) sieving and decanting method and Baermann’s
funnel technique. Collected nematodes through the process were fixed with warm
formalin alcohol (F.A) (4:1) for 24 hours and afterward, dehydrated under the Seinhorst (1952) dehydration techniques. Dehydrated
nematodes were mounted on clean non-greased slides with dehydrated glycerin as mountants. The specimens were studied, measurements taken and diagrams were drawn using a drawing tube attached
to a microscope.
RESULTS
Systematics
Aphelenchoides aeralis
Chanu et al., 2015
(Figure 2, Table 1 & 3)
Diagnosis
Female: Body contour S-shaped
with fine cuticular striations having a lateral field with two incisures. The
cephalic region set off with weak sclerotization, spear 15.3 µm long with small
basal thickenings. The median oesophageal bulb is
oblong with centered valvular apparatus. Oesophagous
overlap intestine. Nerve ring behind the esophagus-intestinal junction.
Excretory pore above nerve ring. Vagina with sphincter and raised vulval lips. Gonads monoprodelphic
and oocytes are arranged in single rows. Spermatheca is filled with sperms and
with uterine sac. Tail 42.5─48.2 µm long,conical,
and with small single mucro.
Male: Body more curved at tail
region. Testis long and outstretched. Spicule is typical of the genus. Dorsal
limb without knob. The capitulum and rostrum very well developed. Post anal
genital papillae, one pair situated above the tail terminus. Tail terminating
into a long spine-like mucro.
Remark: The morphometric details
of the present species conform well with those described by Chanu
et al. (2015).
Aphelenchoides aligarheinsis
Siddiqi et
al., 1967
(Table 1)
Diagnosis
Female: Body contour slightly
curved. Cuticle fine, striations about 1.7 µm at mid-body. Lip region set-off
round. Lateral fields with four incisures. Stylet slender 10.2 µm long with
weakly developed basal knobs. Oesophagous typical of
the genus. Excretory pore at level of the nerve ring. Ovary outstretched
and oocytes arranged in single row. Spermatheca oblong with discoidal
spermatozoa. The Uterine sac is five times body width in length. Tail 35.7–39.1
µm long, elongate-conoid, ventrally arcuate, rounded with spine like mucro.
Male: Not found.
Remarks: The morphometric details
of the present species conform well with those described by Chanu
et al., (2015).
Aphelenchoides baguei
Maslen, 1978
(Table 1)
Diagnosis
Female: Body contour slightly
ventrally curved. Fine cuticularisation with four
incisures throughout body length. Cephalic region set off flattened. Spear
10.2─12.8 µm long with a small basal knob. Oesophagous
typical of aphelenchoid. Mono-prodelphic
reproductive system and ovary outstretched. Vulva is a transverse slit with
slightly protruding lips. Oocytes in single row with a uterine sac. Tail
38.4–46.2 µm long, about 4─5 times anal body width long, terminus with a small
ventral mucro which is multi-papillate almost to its
tips.
Male: Not found.
Remarks: The morphometric details
of the present species conform well with those described by Chanu
et al. (2015).
Aphelenchoides confusus
Thorne &
Malek, 1968
(Table 1)
Diagnosis
Female: Body contour gradually
tapering near extremities. Cuticle with fine striations and lateral fields
marked by four fine lines. Cephalic region set-off. Spear 10.2─17 µm long. Oesophagous typical aphelenchoid
with massive valvular apparatus. Nerve ring is behind the oesophageal
bulb. Excretory pore at level of the nerve ring. Hemizonid
posterior to excretory pore.
Vulva with protuberant labia and
vagina directed forward. Uterus spheroid-shaped, filled with sperm. Ovary
outstretched and post uterine sac collapsed. Tail 20.4–25.3 µm long without a mucro.
Male: Not found.
Remarks: The morphometric details
of the present species conform well with those described by Chanu
et al. (2015).
Aphelenchoides dhanachandi
Chanu et al., 2012
(Figure 3, Table 1)
Diagnosis
Female: Body contour slender
ventrally curved. Lateral filed marked by three incisures. The body cuticle is
fine. The cephalic framework is high. Spear 13.6–15.3 µm long, slender with
indistinct basal knobs. Oesophagous typical with
tamarind seed-shaped median bulb. Nerve ring behind the median bulb, 59.5–69.0
µm long. Excretory pore at the level of the nerve ring. Oesophageal
gland lying dorsally along the intestine.
Monoprodelphic reproductive system and oocytes
arranged in a single row with uterine sac. Vulva protrudes in some species.
Tail 62.9–98.0 µm long, highly curved ventrally tapering into a pointed
terminus.
Male: Not found.
Remarks: The morphometric details
of the present species conform well with those described by Chanu
et al. (2012).
Aphelenchoides longistylus
Chanu & Mohilal, 2014
(Figure 4, Table 1 & 3)
Diagnosis
Female: Body contour slightly
curved with fine annulation. Lateral fields with four longitudinal lines merge
into two lines at around the tail region. The cephalic region indistinctly set
off with six equal lips. Spear 24.22 µm long with indistinct basal knobs. Procorpus straight, median bulb spherical to pyriform in
shape. Excretory pore at the base of the median bulb. The vulva is a transverse
slit. The monoprodelphic reproductive system and
oocytes are arranged in a single row. Uterine sac well developed. The tail
gradually tapers into a cylindrical tube, terminating in a ventral prong tip.
Male: Slightly smaller than
female. Tail slender with single terminal mucro.
Spicules absent about 24.22 µm long. Testis single, 335.62─342.45 µm long.
Remarks: The morphometric details
of the present species conform well with those described by Chanu
et al. (2014).
Aphelenchoides manipurensis Chanu
& Mohilal, 2018
(Figure 5 & 6, Table 2 & 3)
Diagnosis
Female: Body contour cylindrical,
ventrally arcuate, the cephalic framework set off and flat. Cuticle marked by
fine annulus. Lateral fields with two incisures. Spear short, 10.33–13.84 µm
long with small rounded basal knobs. The median bulb is spherical and basal bulb
bifurcated. Excretory pore at 51.9–76.12 µm from anterior end of body. Nerve
ring behind median bulb, ovary single, outstretched, and oocytes arranged in
single row reaching the basal bulb. Spermatheca elongated oval with sperms.
Uterine sac filled with sperms and ventral rounded tip. Vulva protuberant and
vagina at right angle to the body. Tail curved ventrally with rounded tip with
a small mucro at tip.
Male: Body ventrally curved. Tail
conoid with mucronated lip. Testis 138.4–190.3 µm. spermatocytes in single row,
spicules simply arcuate, rostrum rounded, and prominent apex. Three pairs of
sub ventral papillae present towards tip of spicule. Bursa and gubernaculum
absent.
Remarks: The morphometric details
of the present species conform well with those described by Chanu
& Mohilal (2018).
Aphelenchoides minor Seth & Sharma, 1986
(Table 2)
Diagnosis
Female: Body contour straight to
slightly curved ventrally. Lateral fields with three incisures. Cephalic
framework set off without annulation. Spear 4.8–8.2 µm long with indistinct
basal thickenings. Oesophagous is aphelenchoid
type with squarish muscular median bulb with a flat base and crescentric wave. Excretory pore at level of nerve ring.
Vulva, a transverse slit with prominent lips. Oocytes are arranged in single
row with a uterus. Tail 22.4–30.8 µm long, rounded with ventral mucro.
Male: Not found.
Remarks: The morphometric details
of the present species conform well with those described by Chanu
et al. (2018).
Aphelenchoides neoechinocaudatus
Chanu et al., 2012
(Figure 7, Table 2)
Diagnosis
Female: Body contour is
slender with three lateral incisures. The cephalic region is slightly set off.
Stylet slender 11.9 µm long without basal swellings. Oesophagous
typical, median bulb elongated pear-shaped. Nerve ring 66.3─68.0 µm long.
Excretory pore at 62.9–64.4 µm, at the level of the nerve ring. The oesophageal gland was dorsal to the intestine. Reproductive
system monoprodelphic and oocytes arranged in a
single row, and uterine sac well developed. Tail 45.9–54.4 µm long, short, and
pointed with a ventral mucro.
Male: not found
Remarks: The morphometric details
of the present species conform well with those described by Chanu
et al. (2012).
Aphelenchoides neominoris
Chanu & Mohilal, 2014
(Figure 8, Table 2)
Diagnosis
Female: Body contour straight,
tapering towards both extremities with four incisures in lateral field.
Cephalic framework smooth and set-off. Spear 8.65 µm long with distinct stylet
knobs. Procorpus zig-zag, coiled, strongly rounded
corpus with sclerotized plates and elongated gland lobe, dorsal to the
intestine. The excretory pore is close to the nerve ring. Vulval
lips protrude with an inclined vagina. Monoprodelphic
reproductive system, oocytes are arranged in a single row reaching up to the oesophageal bulb. Spermatheca large elongated and uterine
sac empty. The anterior lip of the anus protrudes, the tail bluntly rounded,
12.11–25.95 µm in length, with a small hair-like mucro.
Male: Not found.
Remarks: The morphometric details
of the present species conform well with those described by Chanu
& Mohilal (2014).
Aphelenchoides swarupi
Seth &
Sharma, 1986
(Table 2)
Diagnosis
Female: Body contour cylindrical
to slightly curved, with three incisures in the lateral lines. Cephalic region
set off without annules. Spear 8.2–10.2 µm long with
basal thickenings, procorpus muscular. Nerve ring at
72.8 µm from anterior body. Excretory pore ventral at level of nerve ring. Tail
32.8–37.4 µm long, bluntly rounded with a ventral mucro.
Vulva a transverse slit with vulval lips. Post vulval uterine sac well developed.
Male: Not found.
Remarks: The morphometric details
of the present species conform well with those described by Chanu
et al. (2015).
Aphelenchoides vigor Thorne & Malek,1968
(Table 2)
Diagnosis
Female: Body cylindrical with
coarse annulations. Lateral filed with two incisures. The cephalic framework is
set off by constriction. Spear 11.9–13.6 µm long with distinct knobs. Nerve
ring at 74.8–88.4 µm long and excretory pore at 78.2 µm from the anterior body
region. Vulva sclerotized, overlapping with jointed flap. Ovary outstretched,
tubular uterine sac filled with sperms. Tail 15.3–35.7 µm long, arcuate, blunt
tip without mucro.
Male: Not found.
Remarks: The morphometric details
of the present species conform well with those described by Chanu
et al. (2015).
The hosts and localities of all
the species are mentioned in Table 4.
Aphelenchoides oryzae sp. nov.
(Figure 9, Table 5)
urn:lsid:zoobank.org:act:223FC21F-A734-4794-B702-B396FA7F0D49
Material examined
Holotype:
Collected on August, 2015 from paddy fields (Oryza sativa L.
growing field) by L. Bina chanu, from Thoubal Khekman, Thoubal District, Manipur
with a longitude of 24.5036 and latitude of 93.9116. The specimen is
deposited on nematode collection of Parasitology Section, Centre for Advanced
Study in Life Sciences, Manipur University, Canchipur,
Manipur under the Voucher no. ZoDMU_MN02 with holotype female on slide FTY4
♀1.
Paratype: Females on slides FTY4 ♀2-12 and males on the slides FTY4
♂1-7, same data as holotype.
Descriptions
Holotype female: Body straight, cylindrical, and robust upon
fixation, 685.08 µm long. Lip region offset with rounded sides & flattened
anteriorly, 5.19 µm wide and 1.7 µm high, and smooth in appearance. Body
elongate with fine transverse annulations, 0.8 µm at mid body region. Lateral
lines extend almost to tail tip with two ridges having four evenly spaced lines
in the middle of the body.
Stylet is slender, 17.3 µm long,
the conus slightly shorter than the shaft with indistinct swellings. Median oesophageal bulb rounded to slightly oval with the
refractive thickenings usually placed centrally, 13.84µm high and 8.65µm across
in length. Excretory pore and nerve ring, one to two body widths posterior to
median bulb, excretory pore at 72.66µm from the anterior end and nerve ring at
74.39µm from the anterior end of the body. Oesophageal
glands overlap the intestine.
Vulva is at about two thirds of
the body length from the anterior end. Reproductive system with a single
anterior ovary, oocytes arranged in single rows, with spermatheca, 29.41µm long
filled with sperms and a prominent post-vulval
uterine sac which usually extends just half the distance from the vulva to the
anus. Post uterine sac is about 46.71µm in length.
Tail convex-conoid, 19.03µm long
and straight, usually 10.35 µm anal body widths long with a rounded tail tip
bearing a small terminal mucro.
Paratype males: Lip
region, stylet and oesophagous similar to female.
Tail ends curls ventrally through 45─90° when killed by heat and usually with
simple terminal mucro. Spicules well developed;
15.57–17.3 (16.43 ± 0.86) µm long, the dorsal limb smoothly curved in its proximal
half but flattened to concave tip; the ventral limb appears much weaker than
the dorsal limb. The rostrum and apex are moderately developed; a tangent drawn
from the apex to the spicule tips is separated from the tangent from the apex
through rostrum.
Etymology: The species name is derived
from the host plant.
Diagnosis and Relationships
Aphelenchoides oryzae
sp. nov. is characterized by narrow cylindrical body, adults
being 569.17─750.82 (675.04±53.74)µm long with a
stylet about 17.3 µm long with indistinct basal swellings, a prominent median
bulb and short end bulb, 4 lateral lines throughout the body, and tail
straight, convex-conoid with a simple terminal mucro.
Males are common and functional
with prominal spicules with dorsal limb flattened to
indent in its distal half and the tip curled ventrally. Lateral fields of
adults usually with four lines.
In view of its association with
paddy plants, Aphelenchoides oryzae sp. nov. might be
confused with Aphelenchoides besseyi Christie, 1942 the rice nematode. However, Aphelenchoides oryzae sp.
nov. is separated from A. besseyi
Christie, 1942 in having a single, simple, tail mucro
instead of three─four processes as in A. besseyi. The present species also has longer oesophagus and stylet than A. besseyi
Christie, 1942 whereas oesophagus ranges from 64─68
µm and stylet 10.0–12.5 µm in A. besseyi Christie,
1942.
Aphelenchoides oryzae sp. nov.
differed from other species of Aphelenchoides
in having a female body length of 569.17─750.82 (675.04 ± 58.74) µm, with a
slender stylet length of 17.3 µm and a convex conoid tail with a simple
terminal mucro with four lateral lines along the whole body length.
The differences of characters
between closely related species of Aphelenchoides
is provided in Table 6. Based on these morphometric differences the present
species is reported as new to science.
The Shannon -
Wiener species diversity index and Evenness for all the mentioned species are
given in the table 7.
Conclusions
The richness of the species in
the region may be due to warm climatic conditions, suitable habitats, and hosts
as well as due to the absence of drastic changes in the climatic conditions
during the past few years. But the present work could not cover all the varied
ecosystems of Manipur. Since, nematodes are soil dwelling living around the
roots of plants as well as plant parasitic forms, there is potential for
availability of the organism in various other plant and tree varieties in
different ecosystems of Manipur. There are various forest ecosystems found in
Manipur with varied tree species along the Himalayan range. Further survey and
in-depth taxonomic works incorporating molecular taxonomic techniques can
reveal the rich diversity of the nematode group in Manipur.
Table 1. Morphometric data of female species
of Aphelenchoides spp. from Manipur.
Character |
A. aeralis Chanu et al., 2013 |
A. aligar-hiensis Siddiqi et al.,
1967 |
A. ba-guie Maslen, 1973 |
A. confusus Thorne &
Malek, 1968 |
A. dhanach-adi
Chanu et al., 2012 |
A.longistyl-us Chanu & Mohilal, 2014 |
L |
0.46–0.51 |
0.42–0.61 |
0.58–0.74 |
0.48–0.79 |
0.37–0.50 |
0.59–0.66 |
a |
32.1–33.8 |
25.1–30.2 |
35.2–38.2 |
28.6–46.6 |
25.2–36.1 |
34.4–35.85 |
b |
6.15–7.2 |
6.1–8.1 |
5.2–7.2 |
2.5–4.4 |
6.6–7.7 |
8.53–9.81 |
b´ |
3.66–4.1 |
- |
3.6–4.2 |
- |
3.3–7.4 |
6.03–6.49 |
c |
9.8–10.1 |
11.4–16.1 |
14.2–16.2 |
23.8–38.8 |
5.3–7.4 |
13.67–14.58 |
c´ |
5–6.4 |
2.5–3 |
3.8–4.4 |
2–2.4 |
6.6–9.2 |
5.6–6.14 |
V |
66.7–68.2 |
64.2–71.8 |
62.8–68 |
76.8–78.6 |
57.9–62.9 |
67.78–69.76 |
G1 |
26.9–28.2 |
30.6–34.8 |
42.4–48.6 |
29–42.6 |
13.6–15.3 |
24.2 |
G2 |
- |
- |
14–16.2 |
4.3–18.2 |
- |
39.39–43.08 |
Spear |
15.3 |
10.2 |
10.2–12.8 |
10.2–17 |
- |
- |
Oesophago-us |
125.8 |
73.1–81.6 |
146.4–152 |
134.7–241.4 |
95.2–129.0 |
98.61–102.07 |
Nerve ring |
68 |
|
|
90.1–119 |
56.1–73.1 |
- |
Excretory pore |
62.9 |
52.7 |
82.4–94.2 |
86.7–115.6 |
61.2–85.0 |
- |
Tail |
42.5 |
35.7–39.1 |
38.4–46.2 |
8.5–10.2 |
62.9–85.0 |
41.52–48.44 |
ABD |
8.5 |
11.9–15.3 |
|
|
8.5–10.2 |
6.92–8.65 |
PUS |
- |
35.7–37.1 |
92.4–96.4 |
20.4 |
- |
- |
Table 2. Morphometric data of female species
of Aphelenchoides spp. from Manipur.
Characters |
A. manipure-nsis
Chanu & Mohilal, 2018 |
A. minor Seth & Sharma.
1986 |
A. neoechino-caudatus Chanu et al., 2012 |
A. neominoris Chanu & Mohilal, 2014 |
A. swarupi Seth & Sharma, 1986 |
A. vigor Thorne & Malek,
1968 |
Length |
0.294–0.461 |
0.28–0.36 |
0.53–0.60 |
0.35–0/43 |
0.52–0.68 |
0.44–0.49 |
a |
24.28–38.14 |
18.2–26.2 |
25.5–28.6 |
36.29–40.6 |
34.4–42.4 |
24.8–29.3 |
b |
6.3–8.65 |
4.8–7.2 |
8.3–9.0 |
4.23–6.51 |
5.8–8.2 |
2.7–4.8 |
b´ |
3.61–8.15 |
2.4–4.4 |
4.5–5.0 |
6.03–6.49 |
- |
5.6–8.6 |
c |
12.47–16.46 |
12.2–16.2 |
11.0–11.7 |
13.53–36.29 |
12.2–16.2 |
12.4–30.3 |
c´ |
3–4.25 |
2.2–2.6 |
5.3–5.4 |
1.75–3.75 |
2.2–2.6 |
1.5–3.5 |
V |
68.88–71.69 |
66.4–72.2 |
64.4–64.9 |
69.95–71.65 |
66.4–72.2 |
69.2–88.6 |
Stylet |
10.38–13.84 |
48.2–56.2 |
11.9 |
8.65 |
8.2–10.2 |
11.9–13.6 |
G1 |
24.34–33.01 |
4.2–5.2 |
- |
41.87–50.0 |
32.4–36.4 |
33.5–45.9 |
G2 |
- |
4.8–8.2 |
- |
- |
- |
10.5–13.3 |
Oesophagous |
- |
|
119.0122.4 |
67.47–83.04 |
- |
102–164.9 |
Nerve ring |
- |
- |
66.3–68.0 |
- |
- |
74.8–88.4 |
Exc. Pore |
- |
47.6–54.6 |
62.9–64.6 |
- |
- |
61.2–78.2 |
Tail |
20.76–31.14 |
22.4–30.8 |
45.9–54.4 |
12.11–25.95 |
32.3–37.4 |
15.3–35.7 |
ABD |
6.92–8.65 |
|
8.5–10.2 |
6.92 |
8.5–12.4 |
10.2 |
Table 3. Morphometric data of male species of
Aphelenchoides spp. from Manipur.
Characters |
A. aerialis Chanu et al., 2013 |
A. longistylus Chanu
& Mohilal, 2014 |
A. manipurensis Chanu & Mohilal, 2018 |
L |
0.46─0.51 |
0.562–0.62 |
264.69–320.05 |
a |
29─29.41 |
38.2─42.0 |
25.5–33.39 |
b |
5.9─6.92 |
7.24─8.33 |
25.59–41.52 |
b´ |
3.4 |
3.25–5.60 |
3.47–4.02 |
c |
8.7 |
13.54─17.45 |
12.0–13.91 |
c´ |
4.2 |
4–6.2 |
2.4─6 |
T |
46.71─47.8 |
92.61─102.84 |
50.89─61.09 |
Testis |
46.71─47.8 |
335.62─342.45 |
138.4–190.3 |
Spicule |
23.8–25.95 |
24.22 |
10.38─17.3 |
Tail |
51.0─55.36 |
41.52–46.23 |
20.76–25.95 |
ABD |
11.9–13.84 |
10.32 |
6.92–8.65 |
Stylet |
15 |
24.22 |
10.38─13.84 |
Table 4. Species of Aphelenchoides
spp. with their hosts and localities.
Species |
Host |
Locality |
A. aerialis Chanu et al., 2015 |
Pine tree, Pinus roxburghii Sarg |
Nongpok Sekmai, Thounal District |
A. aligarhiensis Siddiqi et al.,
1967 |
Pine tree, Pinus kesiya Royle |
Khuga Dam, Churchandpur District |
A. baguei
Maslen,
1973 |
Morus alba Lin, Pinus kesiya Royle |
Kakwa Naorem Leikai, Imphal West Distrcit; Keibul Lamjao, Bishnupur District |
A. confusus Thorne &
Malek, 1968 |
Morus alba Linn |
Matai garden, Imphal East District |
A. minor Seth & Sharma,
1986 |
Morus alba Linn |
Kalika Village, Irilbung, Kyamgei, Imphal East District; C. I. College, Bishnupur
district; Govt. Silkfarm Wangbal , Thoubal district; Regional Tasar
Research Station, Chingmeirong, Imphal
West district. |
A. swarupi
Seth &
Sharma, 1986 |
Morus alba Linn, Morus indica
Linn, Pinus roxburghii Sarg |
Regional Tasar
Research Station, Chingmeirong, Imphal
West District; Bishnupur ward no. 4, Bishnupur district; Nongpok Sekmai, Thoubal District |
A. vigor Thorne & Malek,
1968 |
Morus indica Linn |
Bishnupur ward no. 4, Bishnupur District |
A. dhanachandi Chanu et al., 2012 |
Mulberry plant, Morus alba L. |
Yurembam Rose Garden, Imphal West District |
A. neoechinocaudatus Chanu et al., 2012 |
Mulberry plant, Morus alba L. |
Yurembam Rose Garden, Imphal West District |
A. neominoris
Chanu & Mohilal, 2014 |
Orange plants |
Sibilong, Chandel district |
A. longistylus
Chanu & Mohilal, 2014 |
Coconut tree (Cacos nucifera) |
Ninghsing Khul, Jiri, Imphal West
District |
A. manipurensis Chanu & Mohilal, 2018 |
Rooten wood lock |
Manipur University campus, Canchipur. |
Table 5. Morphometric data of species of Aphelenchoides oryzae sp.
nov. All measurements in µm except L in mm.
Characters |
Holotype female |
Paratype females |
Paratype males |
n |
1 |
18 |
7 |
Length |
685.08 |
569.17–750.82
(675.045±58.74) |
536.3–615.88
(576.09±39.79) |
a |
44 |
36.55–44 (42.39±2.93) |
34.44–39. 55
(36.99±2.65) |
b |
7.33 |
6.18–7.33
(6.72±0.51) |
5.74–5.83
(5.78±0.04) |
b´ |
|
|
- |
b1 |
9.65 |
9.65–11.12(10.26±0.55) |
7.75–9.12
(8.43±0.68) |
c |
36 |
16.45–36 (25.70±
8.59) |
15.5–19.77
(17.63±2.13) |
c´ |
1.83 |
1.83–4 (3.144±0.94) |
2.57─4 (3.28±0.71) |
T |
- |
- |
81.86–93.54
(94.24±0.69) |
V |
69.44 |
68.20–70.7
(69.65±0.89) |
- |
G1 |
49.24 |
29.95–49.24
(43.55±7.66) |
- |
Post. Uterine sac |
46.71 |
46.71–86.5 (62.97±
15.61) |
- |
PVS/ V-A% |
24.54 |
24.54–51.02
(34.65±14.15) |
- |
Oesophagous |
93.42 |
91.69–114.18 (101.2±8.91) |
91.69–114.18
(101.2±8.91) |
Stylet |
17.3 |
17.3 |
17.3 |
Lip width |
5.19 |
5.19 |
5.19 |
Lip height |
1.73 |
1.73 |
1.73 |
Median bulb length |
13.84 |
10.38–13.84
(12.68±1.28) |
10.38─13.84
(12.68±1.28) |
Median bulb diam. |
8.65 |
6.92–8.65 (7.49±
0.81) |
- |
Spicule |
- |
- |
15.57–17.3(16.43±0.86) |
Ovary |
337.35 |
224.9–337.35
(293.05±54.2) |
- |
PUS/ VBD |
3 |
3–6.25 (4.27±1.2) |
- |
Nerve ring |
74.39 |
69.2–76.12
(72.94±2.72) |
69.2–76.12
(72.94±2.72) |
Excretory pore |
72.66 |
72.66–77.85
(744.04±2.01) |
72.66–77.85
(744.04±2.01) |
Spermatheca |
29.41 |
29.41 |
- |
Rectum |
8.65 |
5.19–8.65 (7.26±
1.69) |
- |
Tail |
19.03 |
19.03–41.52 (29.41±
8.59) |
31.14–34.6
(32.87±1.73) |
ABD |
10.38 |
8.65–10. 38 (9.51±
0.94) |
8.65–12.11 (10.38±
1.73) |
Table 6. Characters differentiating Aphelenchoides oryzae
sp. nov. from other related Aphelenchoides
species.
Aphelenchoides spp. |
Character differentiation |
A. blastophthorus Franklin, 1952 |
Shorter stylet (15─16 µm) with
prominent knobs |
A. brassicae Edward & Misra, 1969 |
Excretory pore opposite median
bulb base, female tail, shorter (c´=3) with longer mucro,
spicules more smoothly curved. |
A. baguei Maslen, 1979 |
The female tail is ventrally
concave with a longer mucro, and spicules more
smoothly curved with a more prominent apex. |
A. hamatus Thorne & Malek, 1968 [After Vovlas,
1982] |
Shorter stylet (12─13 µm),
female tail ventrally curved with a ventral mucro,
spicule larger |
A. helophilus (de Man, 1880) Goodey,
1933 |
Female body length over 1mm,
stylet 14─16 µm with prominent knobs, spicules smoothly curved |
A. lanceolatus Tandon & Singh, 1974 |
Lip region continuous, stylet
shorter (12.5─13 µm), female body thinner (a=33), shorter post–vulval sac, spicules smoothly curved |
A. lichenicola Siddiqi & Hawksworth, 1982 |
Female body was shorter (L=610
µm), female tail longer (c´=3.5), spicules were characteristically swollen
near the distal end of the dorsal limb |
A. lilium Yokoo, 1964 |
Excretory pore a body width
posterior to nerve ring, shorter stylet (12.5 µm), shorter post–vulval sac, female tail ventrally curved, spicules
smoothly curved |
A. saprophilus Franklin, 1957 |
Shorter female body (L= 546
µm), ventrally curved female tail and larger spicules |
A. sexlineatus Eroskenko ,1967 |
Shorter stylet (9 µm), shorter
female body (L = 605─645 µm), longer post vulval
sac, female tail with a longer mucro. |
A. submerses Truskova, 1973 |
Lip region narrower than the
adjacent body; female tail more curved ventrally; excretory pore anterior to
median bulb |
A. suipingensis Feng & Li, 1986 |
Female with the thinner body
(a=32); excretory pore opposite median bulb base; female tail ventrally
curved with hair-like mucro |
A. Tumulicaudatus Truskova, 1973 |
Lip region not offset, post vulval sac shorter; female tail with characteristic
terminal swelling. |
A. nechaleos Hooper & Ibrahim, 1994 |
Female body longer (L=10.5–11.5
mm); shorter stylet (10.5–11.5 µm) and shorter tail (3.9–4.6 µm) |
A. paranechaleos Hooper & Ibrahim, 1994 |
Longer body (631–860 mm);
shorter stylet (9.5–10.5µm); thinner body (a=37–46) and shorter tail
(c´=2.6─3.6) |
Table 7. Shannon-Wiener species diversity
index and evenness of the species of Aphelenchoides.
Shannon-Wiener index is denoted by H and evenness by E. Total
number of species is 255 and the Shannon-Wiener species index is 3.783.
Species |
No. of individuals (n) |
Proportion, pi=n/N |
ln(pi) |
pi ×ln(pi) |
H = -∑[(pi)×ln(pi)] |
lnk |
E=H / ln(k) |
A. aeralis Chanu et al., 2015 |
20 |
20/40=0.5 |
-0.6931 |
-0.346 |
0.346 |
3.688 |
0.093 |
A. aligarheinsis Siddiqi et al.,
1967 |
5 |
5/35=0.1428 |
-1.9463 |
-0.277 |
0.277 |
3.555 |
0.077 |
A. baguei Maslen, 1973 |
18 |
18/90=0.2 |
-1.609 |
-0.321 |
0.321 |
4.499 |
0.071 |
A. confusus Thorne &
Malek, 1968 |
10 |
10/32=-0.3125 |
-1.1631 |
-0.321 |
0.321 |
3.465 |
0.092 |
A. dhanachandi Chanu et al., 2012 |
12 |
12/53=0.2264 |
-1.485 |
-0.336 |
0.336 |
2.484 |
0.135 |
A. longistylus Chanu & Mohilal, 2014 |
17 |
17/46=0.3695 |
-0.995 |
-0.367 |
0.367 |
3.850 |
0.095 |
A. manipurensis Chanu & Mohilal, 2018 |
38 |
38/58=0.6551 |
0.422 |
-0.277 |
0.277 |
4.060 |
0.068 |
A. minor Seth & Sharma,
1986 |
51 |
51/200=0.255 |
-1.366 |
-0.348 |
0.348 |
5.298 |
0.065 |
A.neoechinocaudatus Chanu et al., 2012 |
4 |
4/53=0.0754 |
-2.5849 |
-0.194 |
0.194 |
3.970 |
0.048 |
A. neominoris Chanu & Mohilal, 2014 |
11 |
11/47=0.2340 |
-1.452 |
-0.339 |
0.339 |
3.850 |
0.088 |
A. swarupi Seth & Sharma,
1986 |
36 |
36/143=0.251 |
-1.382 |
-0.346 |
0.346 |
4.962 |
0.069 |
A. vigor Thorne &
Malek, 1968 |
7 |
7/38=0.1842 |
-1.6917 |
-0.311 |
0.311 |
3.687 |
0.084 |
A. oryzae
sp. nov. |
26 |
26/43=0.4651 |
-0.7655 |
-0.356 |
0.356 |
5.366 |
0.074 |
Key to the species of Aphelenchoides spp. from Manipur
Cephalic
region set-off from body ………………………………………………………………………………………… 2
Cephalic
region not set-off from body ……………………….................... A. longistylus Chanu & Mohilal, 2014
Lateral
fields with 2 incisures ……………......…………………………………………………………………………. 3
Lateral
fields with 3─4 incisures ……………………………………………………………….......………………….
6
Basal
bulb elongated …………………………………………………………………………………………………… 4
Basal
bulb bifurcated ……………………………………………………………….. A. manipurensis Chanu
& Mohilal, 2018
Vagina
with lips …………………………………………………………………………………………. A. aerialis Chanu et al., 2015
Vagina
without lips ………………………………………………………………………....…………………………….………….… 5
Distinct
spear knob, spear 11.9–13.6 µm long, arcuated blunt tail
…………… A.
vigor Thorne & Malek, 1968
Indistinct
spear knob, spear 17.3 µm long, tail
convex-conoid with blunt tip ……. Aphelenchoides
oryzae sp. nov.
Lateral
fields with 3 incisures …………………………………………………………………….........…………………………. 7
Lateral
fields with 4 incisures …………………………………………………………………………………………………... 10
Tail
tip pointed …………………………………………………………………………………………………………………………… 8
Tail
tip bluntly rounded ……………………………………………………………………………………………………………… 9
Spear
4.8–8.2 µm, indistinct knob, tail 22.4─30.8 µm …………………………..…
A. minor Seth &
Sharma, 1986
Spear
8.2–10.2 µm, distinct knob, tail 32.8–37.0 µm long ……………………..
A. swarupi Seth & Sharma, 1986
Tail
with a mucro …………………………………………………………………… A. neoechinocaudatus
Chanu et al., 2012
Tail
without a mucro ………………………………………………………………………… A. dhanachandi
Chanu et al., 2012
With
simple single tail mucro
…………………………………………………………………………........………………………. 11
With
multi-papillated tail mucro
……………………………………………………………………… A. baguei Maslen,
1978
Uterine
sac collapsed/absent …………………………………………………………. A. confusus Thorne & Malek, 1968
Uterine
sac well-developed …………………………………………………………….....…………………………………...
12
Body
contour curved, weak knob, tail elongated conoid with spine like mucro …………………………………………………………………………………………………… A.aligarhiensis Siddiqi et al., 1967
Body
contour straight, tapering at extremities, knob distinct, tail blunt conoid
with hair like mucro …………………………………………………………………………………..…………..….. A. neominoris
Chanu & Mohilal,
2014
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