Non Insect Pests of Crops Synoptic Outline K Roy 29-5-23

Non-Insect Pests of Crops and their Management (EC-364)
Synopsis of Class Lectures-2023

What is nematology?
By convention ‘Nematology’ refers to only Plant Nematology or Agricultural Nematology or

Phytonematology, and deals exclusively with those forms of nematodes which parasitize
plants and are of economic importance to agriculture, horticulture and forestry.
HISTORY OF NEMATOLOGY
The history of Nematology perhaps evolved with the development of mankind.
Ancient History
Ancient history corresponded to the discovery of human and animal parasitic nematodes.
 Vedas (6000-4000BC) are the earliest religious scripture of human history. There are
occasional references of nematodes in the Rig, Yajur and Atharv Vedas under the
Sanskrit name of krimin or krimi meaning worm (Ray, 1992).
 Vedic people knew about krimis perpetuating through eggs and also have their
‘friends’ i.e. phoretic hosts like flies and mosquitoes.
 In Vedaskrimis are divided into two broad groups- Drastam (visible) and Adrastam
(invisible).
 Charak recognized 20 different organisms as krimis in his Samhita. There were

accurate descriptions of intestinal helminths of human beings and animals.
 The next oldest reference to parasitic nematodes is found in ‘Huang Ti Nei Ching’ or
‘The Yellow Emperor’s Classic of Internal Medicine’ from China (2700BC).
 Ebers’ Papyrus during 1553-1550 BC indicated that Ascaris lumbricoides and

Dracunculus medinensis were known at that time.
 More definite references of roundworms and threadworms can be found in Greek

writings; Hipporcrates (400 BC) and Aristotle (350 BC) mention them. During 100-
1200AD, many other nematode parasites of animals and birds were discovered.
Early History
The early history is replete with the discoveries of several free-living and plant parasitic
nematodes.
o 1656 AD-Borellus discovered first free-living nematode ‘vineager eels’ (=Turbatrix

aceti) which once were practically present in all vinegar.
o 1743- A Catholic clergyman, Turbeville Needham discovered Anguina tritici -a

causal agent of ear cockle disease of wheat. That was the first record of plant parasitic
nematode.
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Dr. Kusal Roy, Associate Professor
Department of Ag. Entomology, BCKV, Mohanpur, Nadia
o 1855- R. J. Berkeley observed ‘vibrios’ (Meloidogyne) from green house grown
cucumber at the Rothamsted Experimental Station, England.
o 1859- The large scale monoculture of sugarbeet in Europe for sugar production was
facing economic crisis due to ‘beet tired soil’, a condition referred to as crop failure
due to unknown reason. Schacht in 1859 ascribed the decline in sugarbeet condition to
a cyst nematode, later named as Heterodera schachtii by Schmidt in 1871.
o 1891- Ritzema Bos discovered the foliar nematode, Aphelenchoides fragariae on

strawberry.
o 1892- Liebscher reported pea cyst nematode, Heterodera goettingiana on pea.
o 1913-14 Kuhn observed cyst-forming nematode attacking potatoes in Germany, but

he mistook it for H. schachtii. Zimmermann (1914) definitely establish the fact that
what was thought to be H. schachtii was a serious pest of potato. Massee (1913) had
already reported it as a common parasite of potatoes in Scotland. Wollenweber (1923)
proposed the name Heterodera rostochiensis (=Globodera rostochiensis) for this
nematode pest.
Modern History
 1941- Discovery of potato cyst nematode, Globodera rostochiensis in Nassau County,

Long Islands, USA. Though this nematode was already known from Europe, and its
role in potato production was well established, its discovery in USA led to
enforcement of quarantine regulations to check its further spread. Massive funds were
pumped in for the containment of this dreaded nematode which helped popularizing
the importance of nematodes to agriculture. Intensive studies conducted on this
nematode led to the understanding of some of the very basic principles of nematode
sampling methods, its biology and ecology.
 1943, 1945- Discovery of DD (dichloropropane-dichloropropene) mixture by Carter

in 1943 and EDB (ethylene dibromide) by Christie in 1945. These two chemicals are
the biproduct of petroleum industry, facilitated the demonstration of nematode
pathogenicity, crop loss estimations due to nematodes, and undertaking large scale
operational control.
 1951- Christie and Perry demonstrated the pathogenic potential of several

ectoparasitic nematodes, e.g., Xiphinema, Longidorus, Trichodorus, Belonolaimus,
Dolichodorus, Hemicycliophora etc.
 1953- Discovery of burrowing nematode, Radopholus similis as the causal agent of

‘spreading decline’ of citrus in Florida (USA) and ‘yellows’ disease of pepper in
Bangka Islands, Indonesia. Out of 22 million vines of pepper planted on this island,
only 2 million survived, thus this nematode made a severe dent in the economy of
Indonesia, which was based on the export of pepper.
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 1955- Lesion nematode, Pratylenchus penetrans discovered as the causal agent of
‘peach replant problem’ in Canada.
 1956- First ever independent Journal on PPN, Nematologica started in Europe.
 1958- First report of transmission of viruses by nematodes (Xiphinema index

transmitting ‘fanleaf grapevine virus’ in grapevine) in the USA by Hewitt, Raski and
Goheen.
 1971- Bursaphelenchus xylophilus (vectored by insects) found to be responsible for

mysterious ‘pine wilt’ disease in Japan.
Born in USA, he served in many capacities before adopting nematology as a profession. After
doing M. Sc. He taught chemistry in school for 6 years. He moved to Germany for his Ph. D.
and then moved to Australia where he had to sell soap and watches for sometime. Ultimately,
he got the job of consultant pathologist. About, 1900, he became associated with the
Experimental Station to the Hawaiian Sugar Planters’ Association, where he investigated the
nematode found in sugarcane fields. In 1907, he joined USDA and by 1910, he succeeded in
focusing his attention to Nematology.
Contribution of NA Cobb—
 His book ‘Contribution to a Science of Nematology’ published in 1913 is the most

outstanding.
 His work still rated as the highest standard, particularly the finest drawings of new
nematode species he described.
 devising many techniques which are routinely practised today, e.g. sieving method for
nematode extraction from soil, mounting of nematodes on double cover-slip
aluminium slide, preparation of en-face views, new camera lucida apparatus for
making nematode illustartion, etc.
 description of many minute sensory organs of nematode, like amphids, phasmids,

deirids, cephalids, papillae
 discovery of many nematode species
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 separation of FL and PPN from Helminthology to Nematology, term proposed by him
in 1914 and establishment of Division of Nematology in USDA.
Development of Nematology in India
Period from 1901-1958 was marked by some important records of nematode parasites from
various parts of the country. The organized work on PPN in India was started only during
sixties. A few important events in the history of Nematology in India are cited below-
 1901- Barber reported RKN infecting tea in south India- first ever report of a PPN
from India.
 1913-19- Butler reported ‘ufra’ disease of rice (causal agent Ditylenchus angustus)
from Bengal.
 1919- Milne recorded seed gall nematode (Anguina tritici) of wheat in Punjab.
 1926-1934- Ayyar discovered RKN on vegetables and other crops in south India.
 1936- ‘White tip’ disease of rice caused by Aphelenchoides besseyi reported by

Dastur from Central Province (Presently MP).
 1958- ‘Molya’ disease of wheat and barley (causal agent- Heterodera avenae)
recorded in Rajasthan by Basudeva.
 1959-61- Siddiqi reported PPN from UP including citrus nematode, Tylenchulus

semipenetrans on citrus.
 1961- F G W Jones reported Golden Nematode of potato (Globodera rostochiensis)

from Nilgiri hills in Tamil Nadu.
 1966- Nair, Das and Menon reported burrowing nematode, Radopholus similis on
banana in Kerala.
 1966- Division of Nematology established at the IARI, New Delhi.
 1967-68- First South-East Asia Post Graduate Nematology Course held at Aligarh
Muslim University, Aligarh and IARI, New Delhi in collaboration with the
International Agriculture Center, Wageningen, Netherlands. 1969- Nematological
Society on India (NSI) founded, 1st All India Nematology Symposium held at IARI,
New Delhi.
 1971-Indian Journal of Nematology (IJN) commenced publication.
 1977- AICRP on Nematode Pests of Cropping Systems started functioning.
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Economic Importance of Nematode in Agriculture
 Distribution-
 Symptom-
 Spread-
 Association with other disease causing agents-
 Crop Loss-
Table 1: Crop losses due to important plant parasitic nematodes in India
Crop Nematode species Loss (%)

Rice Meloidogyne graminicola 11-73
Hirschmanniella oryzae 30 – 87
Hirschmanniella mucronata 43
Wheat Anquina tritici 80
Heterodera avenae 80
Maize Heterodera zeae 12-26
Meloidogyne incognita 6
Finger millet Meloidogyne incognita 4.8
Potato Globodera rostochiensis Total crop failure
Tobacco Meloidogyne incognita 50
Jute Meloidogyne incognita
21.35
M. javanica
Cotton Meloidogyne incognita 17.7 – 19.9
Chickpea Pratylenchus thornei 27
Cowpea Meloidogyne incognita 28.6
Rotylenchulus reniformis 10.5-14.5
Pea Meloidogyne incognita 20
Blackgram Meloidogyne incognita 8.7
Groundnut Meloidogyne arenaria 51
Okra Meloidogyne incognita 28.-90
Brinjal Meloidogyne incognita 27-33.7
Tomato Meloidogyne incognita 48
Frenchbean Meloidogyne incognita 43.5
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Banana Radopholus similis 12

Citrus Tylenchulus semipenetrans 15
Tuberose Aphelenchoides besseyi 59
In India, the annual crop loss due to cereal cyst nematode, Heterodera avenae in wheat and
barley was estimated to be Rs. 32 million and 25 million, respectively in Rajasthan alone.
An annual crop loss of Rs. 20 million was assessed in coffee due to lesion nematode,
Pratylenchus coffeae in an area of about 1000 ha in Karnataka.
Crop loss in Nutshell
World :Avoidable yield loss due to nematode is approximately 12% in the world while
developed country assumes 9% as against 15% in the developing country (Sessar &
Freckman,1987).
India :21068.73 million rupees in 24 different crops (Jain et al.,2007)
Classification of Plant Nematodes based on Feeding Habits
Nematodes may be grouped by feeding habit as:
• Endoparasitic– entire body inside the root
• Ectoparasitic– entire body outside the root
• Semi-endoparasitic- part of body inside root
By movement when feeding, they are called:
• Sedentary – mostly immobile during their life
• Migratory – mobile for all their life.
List of Economically Important Plant Parasitic Nematodes:
Common Name Scientific Name

1. Root-knot Nematode Meloidogyne spp.
2. Cyst Nematode Globoderarostochiensis
Heterodera spp.
3. Burrowing Nematode Radopholus similis
4. Lesion Nematode Pratylenchus spp.
5. Tuberose Foliar Nematode or Rice Aphelenchoides besseyi
White tip Nematode
6. Citrus Nematode Tylenchulus semipenetrans
7. Reniform Nematode Rotylenchulus reniformis
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8. Rice Stem Nematode Ditylenchus angustus
9. Spiral Nematode Helecotylenchusmulticinctus
10. Dagger Nematode Xiphenema spp.
11. Stunt Nematode Tylenchorhynchus brevilineatus
Generalized Symptoms of Nematode Infestation
A. Below ground feeders or soil inhabiting nematodes-
They produce two types of symptom viz. below ground symptom and above ground
symptom. Above ground symptoms are non specific and often misinterpreted with nutrient
deficiency or some other malady.
Above Ground/Aerial Symptom Below Ground Symptom

1. Stunting e.g. Globodera rostochiensis Root gall/knot e.g. Meloidogyne spp. on
in potato, Heterodera avenae in wheat veggies,Xiphinema diversicaudatum on
rose, Hemicycliophora arenaria on citrus
and lemon, Nacobbus batatiformis on
sugarbeet, Subanguina radicicola on
cereals and grasses.
2. Yellowing e.g. Pratylenchus coffeae on Stubby Root e.g. Trichdorus christiei on
coffee, Tylenchulus semipenetrans on maize Coarse Root e.g. Belonolaimus
orange & lemon longicaudatus(sting nematode) on maize
3. Wilting e.g. Meloidogyne Root Lesions e.g. Radopholus
incognita(Southern root knot nematode) similis/Pratylenchus spp./ Helicotylenchus
in jute, brinjal, tomato, pulses, beans multicinctus on banana
etc.
4. Dieback e.g. Tylenchulus semipenetrans Rots of fleshy parts e.g. Ditylenchus
on orange & lemon destructor causes potato rot
5. Excessive root
branching/bearding/witches broom/
hairy root/coral roote.g. northern root-
knot nematode, Meloidogyne hapla on
tomato
B. Above ground feeders or Aerial nematodes-
They produce specific above ground symptoms
Above Ground/Aerial Symptom
1. Dead or Devitalized Buds e.g. Aphelenchoides frageriae onstrawberry,
Aphelenchoides besseyi ontuberose
2. Crinkled and Distorted Stem & Foliage e.g. Anguina tritici on wheat,
Aphelenchoides besseyi onrice, Ditylenchus angutus on rice
3. Galls
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a) Seed Gall e.g. Anguina spp. on cereals and grasses

b) Leaf Gall e.g. Anguina balsamophila on balsam
c) Stem Gall e.g. Subanguina agropyri on grass
4. Necrosis and Discolouration e.g. red ring disease of coconut palm caused by
Rhadinaphelenchus cocophilus
5. Leaf Lesions e.g. Aphelenchoides ritzemabosi onchrysanthemum

Ufra disease of rice caused by stem nematodeDitylenchus angustus
Thor Ufra Ripe Ufra
Leaf blade of the flag leaf is twisted, curled & If, Panicle comes out of the leaf sheath of
deformedPanicle can not come out of the leaf flag leaf that are chaffy and sterile
sheath of flag leaf
Nematode diseases vectored by insect. Red ring nematode of coconut Bursaphelenchus

(=Rhadinaphelenchus) cocophilus vectored endophoretically by Palm weevil Rhynchophorus
palmarum .
Pine wilt nematode-Bursaphelenchus xylophilus vectored ectophoretically by Pine sawyer
beetle-Monochamus alternatus in Japan and M. caroliensis in USA.
Reference book for further reading-

1. Text Book on Introductory Nematology-HK Bajaj & RK Walia
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Morphology of Nematodes
WHY MORPHOLOGY?
 Nematode body organization is an architectural marvel. It possesses all the vital
systems of higher invertebrates packed intricately even in tiny plant parasitic forms.
 Nematode taxonomy is based mainly on their morphology, hence a sound knowledge
about the structure of nematodes is absolutely essential for their identification.
Body Size
The animal parasitic nematodes are quite large in size and easily visible to the naked
eye.The common intestinal roundworm, Ascaris lumbricoides is about 30cm long, while the
guinea worm, Dracunculus medinensis may reach up to 1m in length. The largest ever known
nematode, Placentonema gigantissima is about 8m long is parasitic in the placenta of sperm
whale. Greeffiella minutum, a marine nematode, measuring 82µm is the shortest nematode
recorded so far.
Plant parasitic tylenchid nematodes generally range between 0.3 to 2.0mm in length,
rarely up to 3.0 mm. Paralongidorus epimikis is the longest plant parasitic nematode,
reaching up to 1.2cm length.
Body shape
Nematode body is typically vermiform, thread-like and tubular, tapering towards both
ends. The principal axis is longitudinal. Males and females generally look alike. The body
may be fusiform (spindle-shaped), pear-shaped, lemon-shaped or variations of saccate.
Sexual dmorphism is also visible in some plant nematodes like Radopholous, Meloidogyne,
Heterodera/ Globodera, Rotylenchulus, Tylenchulus, Cricinematids etc.
Radopholus male (left) and female (right)

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Nematodes are elongate, cylindrical, unsegmented worms. In elongate-cylindrical
species the anterior end is usually bluntly rounded with the oral aperture (mouth opening)
terminal. The posterior end may be bluntly rounded or pointed; sometimes it is tapered to a
point or it may be long and filiform. The opening of the excretory system (excretory pore) is
located in the anterior portion of the body in a ventro-median position. The opening of the
female reproductive system (the vulva) is usually midway in the body in a ventro-median
position. The opening of the digestive system (anus) is ventro-median near the posterior end.
In males the digestive and the reproductive systems join posteriorly to form a cloaca which
opens on the ventral side near the terminal end. The following terms are used to refer to areas
of the nematode: adanal (located near the anus), caudal (located near the tail), cervical (in
the neck region).
Fig. Plant Feeding Nematodes

Body regions, colouration and segmentation
The body regions of a nematode cannot be easily divided into head, thorax and abdomen.
Often, the anterior proximity of the body bearing the mouth and lips is called head and it is
continuous with the main body. Sometimes, it is set-off by a constriction. The portion of the
body beyond anus (cloacal aperture in males) is known as tail. The tail shape is highly
variable (cylindrical, conoid, filiform etc.) even among plant nematodes. Longitudinally, the
body can be divided into 4 regions viz. ventral, dorsal and two laterals. Nematodes are
appendageless. The body is not metamerically segmented internally. Nematodes are
colourless and transperant in general. Some nematodes may appear variously tainted
depending on food material in the gut.
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Variations in Head shapes
Set-off /offset Continuous head (Helicotylenchus)

head(Hoplolaimus)
Variations in tail shapes
Cylindrical Filiform Elongate conoid Digitate

Nematode Body organization and symmetry
The nematode body organization is basically tubular and consists of 2 tubes. Outer tube is
wider and larger, and forms the body wall. It encloses a narrower and shorter inner tube,
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alimentary canal. Two tubes join at the anterior extremity to form the lip region, and the
posterior region through the rectum.
Pseudocoelom/ body cavity
Space between two tubes; it lacks mesodermal epithelial lining, filled with pseudocoelomic
fluid which exerts tremendous pressure on the body wall to give it a rounded outline in a
cross-section.Reproductive, excretory and nervous system are located within the
pseudocoelom.
 Nematodes lack special organs for respiration and circulation.

 The nematode body is basically bilaterally symmetrical, i.e. if the body is cut into 2
equal halves through a dorso-ventral (sagittal) plane, the two halves are mirror images
of each other. Other organs, however, also shown radial symmetry or asymmetry.
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A. The Outer Body Tube (Body Wall)
The body wall consists of 3 layers, the cuticle which may be considered to be an
exoskeleton, underneath the cuticle is the hypodermis (epidermis), and the muscle layer
composed of a single layer of elongate spindle shaped muscles.
Fig. Cross section of nematode through mid-body region

Cuticle:
It is a non-cellular, non- living layer forming the outer covering of the nematode. It is tough
but elastic. Cuticle is shed-off at each moult and a new one is formed. Cuticle itself is a
multi-layered structure, which can be basically divided into 3 regions, viz. cortex or the
cortical layer, matrix or the homogeneous layer, and fibrillar or the basal layer. These
layers are complicated chemically and are rich in structural proteins such as keratin,
collagen, matricin etc. Chitin is not found in nematode cuticle. Only nematode egg-shell
contain chitin.
Functions:
It functions as an exosceleton, regulates permeability and body growth, and helps in
locomotion. It also plays an important role in respiration and excretion.
Cuticular ornamentation or marking
The cuticle is beset with various types of cuticular markings. It may be smooth or marked
with transverse grooves (striation) known as striae. The raised area between two striae is
called interstrial region. In Criconematids these striae are very deep with very wide interstrial
zone i.e. called annulation. Annulation divides body into superficial segmentation called
annuli (sing. annule). The posterior margin of these annuli may bear spines, scales etc.
arranged in definite patterns. This is called cuticular ornamentation.
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Smooth Striated Annulated
Cuticular scales (left) and spines (right)

Lateral field marked with lateral lines or incisures
The striation becomes discontinuous on the two lateral sides, which is little bit raised and
running nearly the entire length of the body. This is called lateral alae or lateral ridge. Each
lateral alum constitutes a lateral field and is further marked by longitudinal grooves i.e. called
lateral lines or incisures. Number of lateral lines may vary from 0-16 and is specific. Lateral
field is also called as seam.
Variations in number of Lateral lines, lateral field areolated (G, H)
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Bursa or Caudal alae: Winged like lateral cuticular extension in the tail.
Fig. Caudal alae – Peloderan (left) and Leptoderan (right)
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Hypodermis/Epidermis
Underlying the cuticle is the hypodermal layer. It may be syncitial or made up of a single
layer of cells. Although hypodermis is a continuous layer but it is generally thickened in the
dorsal, ventral and lateral sides to form 4 hypodermal chords. The two lateral hypodermal
chords are more conspicuous. The main nerve chords and excretory canals are generally
located in the hypodermal chords.
Function:Secretion of cuticle
Muscle layer: There are two types of muscles in the nematode body- somatic and specialized
muscles.
o Somatic muscles: A single layer of spindle shaped cells attached to the hypodermis in the
4 inter-chordal zones.
Function:i) Locomotion and ii) Reservoir of stored food material
Function:Govern the movement of those specific organs only.

Muscle layer has two distinct zones
o Fibrillar zone- Contractile portion that contains myofibrils
o Sarcoplasmic zone- Non-contractile portion that contains nucleus and cell cytoplasm
Nematode muscles are unique- Innervation processes (cytoplasmic extension) from non-
contractile portion of muscle cells join the nerve chord. The innervation processes from
dorsal half join the dorsal nerve chord, and those from ventral half join the ventral nerve
chord. The muscle cells of dorsal half work in unison as muscles of ventral half, thus
regulating the bending of nematode body in a dorso-ventral plane while moving.
Based on the shape of cells and arrangement of contractile elements (myofibrils), the muscle
cells are classified into 3 types, i.e. platymyarian, coelomyarian and circomyarian.
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Platymyarian - Contractile elements restricted to wide base of cell touching the hypodermis
Coelomyarian – Contractile elements extending up the sides of the cells which have a
narrow base
Circomyarian – Contractile elements present on whole of the cell cicumference, enclosing

the sarcoplasm in between
When only 2-5mmuscle cells are present in each inter-chordal zone, the arrangement is
known as meromyarian; if they are more than 5, it is called polymyarian. Usually
platymyarian cells are meromyarian and coelomyarian cells are polymyarian.
B. The Inner Body Tube (Alimentary Canal)
The inner body tube forms the alimentary canal or the gut, and it can be distinguished into 3
zones- the stomodaeum (fore gut), the mesenteron (mid-gut), and the proctodaeum (hind-gut).
Stomodaeum and proctodaeum are lined with cuticle internally.
Stomodaeum-Oral aperture
Enface view of nematode showing arrangement of lips and cephalic setae (A) and
papillae (B) in 2-3 circles
Stomodaeum-Stoma
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Cephalic Sclerotization & Stylet
Fig. Stomatostylet in Tylenchids
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Fig. Odotostylet (left) and Onchiostylet (right)
Fig. Cross section of oesophageal lumen
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Variations in structure of oesophagi – A. Monochid, B. Dorylaimid, C. Rhabditid, D.

Tylenchid, E. Neotylenchid, F. Hoplolaimid, G. Croiconematid, H. Aphlenchid
The Mesenteron and Proctodaeum
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Reproductive System
Most nematode species are dioecious, with separate male and female individuals, though
some, such as Caenorhabditiselegans, are androdioecious, consisting of hermaphrodites and
rare males.
Female reproductive System
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Male Reproductive System
Spicule
The copulatory apparatus generally consists of two equal spicules, cuticularized, tubular
structures. The spicules are rarely absent (e.g. marine Monoposthia), may be dimorphic,
partly fused (in rhabditids) or reduced to a single spicule (as in some mermithids). Spicules
are formed by gradual invagination of the posterior wall of thespicular pouches, which
originate from the spicular primordium, specialized cells of the dorsal wall of the cloaca.
Excretory System
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Nitrogenous waste is excreted in the form of ammonia through the body wall, and is not
associated with any specific organs. However, the structures for excreting salt to maintain
osmoregulation are typically more complex.
Nervous System
CNS: The nerve ring usually encircles the isthmus of the pharynx, rarely the intestine. The
nerve ring is connected to several ganglia, longitudinal nerves running anteriorly towards the
anterior sensilla in the head region, while posteriorly four large nerves, the largest being the
ventral nerve, run through each of the four epidermal chords, with four smaller nerves, two
laterodorsal and two lateroventral, running adjacent.
PNS: The peripheral nervous system consists of a lattice-like network of nerves that connect
setae and papillae. It coordinates impulses from these somatic sense organs, both with each
other and with the central nervous system.
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Sense Organs
Other Systems
Definition of Nematode
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Population: A geographically localized group of individuals of same kind at a particular time
represent a population. So, some of the all populations of same kind will form a species.
Species concept:
A) Genetic species concept: Groups of interbreeding natural population having similar
genotype but are reproductively isolated from other such group is called a species.
The above mentioned definition of species solely applied to a bisexual species only. Thus
genetic species concept has seldom been confused practically due to our incomplete
knowledge of reproductive biology, geographical distribution and speciation process in
nematode. For this reason morphological species concept or phenetic species concept is
solely applicable for nematode as they may reproduce both parthenogenetically or bisexually.
B) Phenetic or Morphological species concept: A group of natural population having a high
degree of similarity in their phenotype or morphology is called a species. It is applicable both
for sexually and parthenogenetically reproducing individual. This concept is dominating in
nematology as it is applicable both ways.
Speciation: A species through evolution may become a different new and distinct species. As
the characters of nematode or organism varies from time to time. Speciation is a process of
multiplication by which one genetically connected population divides into two such
populations between which genetic interchange is not possible. Speciation in parasitic
nematodes is dependent upon their hosts.
Reproductive isolation: Reproductive isolation (inability to interbreed) is the most important
boundary between two species. This is the main factor which maintains the distinctness of the
species.
The information on reproductive biology is available in the two races of burrowing nematode
Radopholus similis. These two races are morphologically indistinguishable. Banana race
which attack banana only having haploid chromosome n=4; but citrus race attacks both
banana and citrus where n=5. Both the races have different protein pattern. Male of banana
race was found attracted to the female of citrus race but does not copulate. This observation
forms the basis to discard racial designation and an elevation of citrus race of Radopholus to
a species rank as Radopholus citrophilus (causal agent of citrus spreading decline disease).
Taxonomy and Sytematics:
The word taxonomy is derived from two Greek words taxis means arrangement or
division and nomos means law. It is the science of classification according to a pre-
determined system, with the resulting catalog used to provide a conceptual framework for
discussion, analysis, or information retrieval. In theory, the development of a good taxonomy
takes into account the importance of separating elements of a group (taxon) into subgroups
(taxa) that are mutually exclusive, unambiguous, and taken together, include all possibilities.
In practice, a good taxonomy should be simple, easy to remember, and easy to use.
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Taxon / Taxa:
In biology, a taxon (plural taxa; back-formation from taxonomy) is a group of one or more
populations of an organism or organisms seen by taxonomists to form a unit at any level of
hierarchic classification. A taxon is usually known by a particular name and given a
particular ranking, especially if and when it is accepted or becomes established. Texon is not
a species; but any given species is a taxon.
Definition of taxonomy
o The science dealing with the description, identification, naming and classification of
organisms in an ordered system based on similarities of structure, origin and natural
relationships especially of evolutionary trends is called taxonomy.
o The science or practice of classification or arranging organisms following law and
principles is called taxonomy.
Levels of taxonomy
-taxonomy: -taxonomy is concerned with identification, description or characterization,

and naming (i.e. nomenclature) of organisms. This is descriptive taxonomy.
-taxonomy: It focuses on placing organisms within hierarchical groups that show their
relationships to other organisms. In other words, arrangement of species into a
natural system of lower and higher taxa is called -taxonomy. This is
classificatory taxonomy (i.e. Classification).
-taxonomy: Dealing with the various biological aspect of taxa ranging from the study of
intraspecific population to studies of speciation and of evolutionary rates and
trends. Here, evolutionary characters and variations among organisms are
studied. This is called systematics or evolutionary taxonomy.
Role of taxonomy:
i) It gives the vivid picture of existing organism’s diversity.
ii) It also provides much of information permitting reconstruction of phylogeny of life.
iii) It supplies classification which is of hierarchic and explanatory value in most
branches of biology.
Classification:
The arrangement of animals and plants in taxonomic groups according to their
observed similarities or relationships (including at least kingdom and phylum in animals,
division in plants, and class, order, family, genus, and species). Such relationships may ne
phenetic (morphological), phyeltic (evolutionary development of a species) or both.
Types of classification: Classification is of two types viz.,
i) Artificial classification- it is exercised to fulfill our particular needs such as

indexing, cataloging, aid of teaching and identification as because our knowledge
is insufficient
26
ii) Natural classification- it is done on the basis of their phylogenetic relationship and
tries to show lineage.
Why classification of nematodes keeps on changing?
Classification is always changing due to updating of new information about the nematodes.
We are trying to reach near to natural classification exploiting modern techniques.
How can we classify nematodes?
We try to classify nematodes mostly based on morphological characters (phenetics) because
there is no fossil record or paleontological record of the nematode.
Nematode taxonomy= Morphotaxonomy (taxonomy based on morphology and

morphometric or allometric characters or traits).
Phylogenetic continuity: Finding continuity between the gaps of the different groups.
Systematics
Definition: Systematics deals with the kind and diversity of the organism and of any and all
relationships among them (Sympson, 1961).
Systematics is now a days called biosysatematics. Biosysatematics indicates the biological
content. Europeans tend to use the terms "systematics" and "biosystematics" for the field of
the study of biodiversity as a whole, whereas North Americans tend to use "taxonomy" more
frequently.
Systematist- A scientist who is specialized in systematics, must, therefore, be able to use

existing classification systems, or at least know them well enough to skillfully
justify not using them.
Difference between taxonomy and systematics
Taxonomy Systematics
Dealing with naming and description of Whole diversity of the population and the
individuals and populations. variations between the populations.
27
Systematic ranks having different suffixes attached to the stem
Taxonomic category Suffix
Kingdom -
Phyllum -a
Class -ea
Subclass -ia
Order -ida
Infraorder -omorpha
Suborder -ina
Superfamily -oidea
Family -idae
Subfamily -inae
Tribe -ini
Genus -
Species -
Subspecies -
Binomial Nomenclature: Nomenclature is derived from two latin words ‘nomen’ means
name and ‘clare’ means to call. International Code of Zoological Nomenclature (ICZN) is an
authoritative document that contains the valid rules of zoological nomenclature. Carolus
Linnaeus is the father of set of rules of ‘binomial nomenclature’. These rules were published
in the book ‘Critica Botanica’ and ‘Philosophia Botanica’. The scientific name of an
organism is designated by both a generic and a species name. Binomial nomenclature should
have 3 pre-requisites (i) uniqueness, (ii) universality, and (iii) stability. The purpose of
binomial nomenclature is to provide a label for taxa at all levels in order to facilitate
communication among the biologists.
 ICZN- It contains 1 preamble, 86 articles, 5 appendices, 1 official glossary, and a detailed
index.
 Starting point of zoological nomenclature-1st January, 1758.
28
Classification of Plant Nematodes
Kingdom: Animalia
Phyllum: Nematoda Potts, 1932
Class: Secernentea (=Chromadorea Inglis, 1983)
Sub-class: Diplogasteria (= Chromadoria Pearse, 1942)
Order: 1. TylenchidaÖrley, 1880
2. Aphelenchida Siddiqi, 1980
Class: Adenophorea (=Enoplea Inglis, 1983)
Subclass: Dorylaimia Inglis, 1983
Order: 1. Dorylaimida Pearse, 1942
Subclass: Enoplia Pearse, 1942
Order: 1. Triplonchida Cobb, 1920
Order 1: TylenchidaÖrley, 1880
Suborder: Tylenchina Thorne, 1949
Superfamily: 1. Tylenchoidea Örley, 1880
Family: 1. Tylenchidae (e.g. Tylenchus, Filenchus, Coslenchus, Malenchus)
2. Tylenchorhynchidae (e.g. Tylenchorhynchus, Merlinius)
3. Dolichodoridae (e.g. Dolichodorus)
4. Belonolaimidae (e.g. Belonolaimus)
5. Pratylenchidae (e.g. Pratylenchus, Radopholus, Hirschmanniella)
6.Hoplolaimidae (e.g. Aorolaimus, Helicotylenchus, Hoplolaimus,
Peltamigratus, Rotylenchus, Rotylenchulus,
Scutellonema)
7. Psilenchidae (e.g. Psilenchus)
8. Tylodoridae (e.g. Tylodorus)
Superfamily: 2. Atylenchoidea
Family: 1. Atylenchidae (e.g. Eutylenchus, Atylenchus)
2. Meloidogynidae (e.g. Meloidogyne, Meloinema, Bursadera)
Super Family: 3. Heteroderoidea
Family:1. Heteroderidae (eg. Cactodera, Globodera, Heterodera, Punctodera,
Meloidogyne)
2. Nacobbidae (e.g. Nacobbus)
Super Family: 4. Criconematoidea Taylor, 1936 (1914)
Family: 1. Cricoematidae (e.g. Criconema, Criconemoides, Hemicriconemoides,
Hemicycliophora)
Family: 2. Paratylenchidae (e.g. Paratylenchus)
Family: 3. Tylenchulidae (e.g. Tylenchulus, Trophotylenchulus, Sphaeronema)
Family: 4. Tylenchocriconematidae (e.g. Tylenchocriconema)
Super Family: 5. Neotylenchoidea Thorne, 1941
Family: 1. Neotylenchidae (e.g. Neotylenchus)
Family: 2. Paurodontidae (e.g. Paurodontus)
29
Family: 3. Nothotylenchidae (e.g. Nothotylenchus)
Family: 4. Ecphyadophoridae (e.g. Ecphyadophora)
Super Family: 6. Sphearularioidea Lubbock, 1861
Family: 1. Sphearulariidae (e.g. Sphearularia, Tripius)
Family: 2. Anguinidae (e.g. Anguina, Ditylenchus, Subanguina)
Family: 3. Halenchidae (e.g. Halenchus)
Order 2: Aphelenchida Siddiqi, 1980
Sub-order: Aphelenchina Geraert, 1966
Super Family: 1. Aphelenchoidea Fuchs. 1937 (Thorne, 1949)
Family: 1. Aphelenchidae (e.g. Aphelenchus)
Super Family: 2. Aphelenchoidoidea Skarbilovich, 1947 (Siddiqi, 1980)
Family: 1. Aphelenchoididae (e.g. Aphelenchoides, Bursaphelenchus)
Order 3: Dorylaimida Pearse, 1942
Suborder: Dorylaimina Pearse, 1942
Superfamily: Dorylaimoidea de Man, 1876 (Thorne, 1935)
Family: 1. Longidoridae Thorne, 1935 (Meyl, 1961)
(e.g. Longidorus, Paralongidorus, Xiphinema)
Order 4: Triplonchida Cobb, 1920
Suborder: Diptherophorina Coomans& Loof, 1970
Superfamily: Doiptherophoroidea Micoletzky, 1922
Family: 1. Trichodoridae (e.g. Trichodorus, Paratrichodorus)
Difference between the class Secernentea and Adenophorea

Secernentea/ Phasmidea/ Chromadorea Adenophorea/Aphasmidea/ Enoplea
1. Phasmid present 1. Phasmid absent
(exception-Aphasmatylenchus)
2. Amphid labial in position 2. Amphid post labial in position
3. Lateral gland opening into excretory duct 3. Lack lateral canal, excretory duct end in a
cell (renette cell)
4. Hypodermal and caudal gland absent 4. Hypodermal and caudal gland present
5. Males are monorchic, bursa generally 5. Males are diorchic, bursa generally absent
present e.g. Tylenchida, Rhabditida e.g. Dorylaimida, Mononchida, Mermithida
# adeno means gland, phorea means bearing

Difference between the orderTylenchida and Aphelenchida
Tylenchida Aphelenchida
1. All the three oesophageal glands do not 1. All the three glands open in the median
open in the median bulb. DOG orifice is bulb. DOG orifice is located in the
30
located at the base of stylet in procorpus median bulb/ metacorpus.
region.
2. Amphideal aperture is lateral in position. 2. Amphideal aperture is latero-dorsal in
position.
3. Median bulb with semi-lunar/crecentic 3. Median bulb strongly developed, offset,
valve plate. rectangular with crecentic valve plate. It
covers almost entire width/cross-section
of the body.
4. Caudal alae/bursa is without bursal rays. 4. Caudal alae is with bursal rays or absent.
5. Spicules are arcuate/ not rose-thorn 5. Spicules are arcuate or rose- thorn
shaped. shaped.
6. Post anal genital papillae/caudal papillae 6. Post anal genital papillae/ caudal papillae
absent. present.
7. Anus inconspicuous, pore like, directed 7. Anus conspicuous, directed backward and
outwards. anterior tip hanging type.
Difference between the orderDorylaimida and Triplonchida
Dorylaimida Triplonchida
1. Stylet (odontostyle) hollow, withlumen 1. Stylet (onchiostyle) an elongated
tooth ora complex structure
2. Amphidial duct long 2. Amphidial duct short
3. Males with two testes (diorchic) 3. Males with one testes (monorchic)
31
Genus- Meloidogyne (Goeldi, 1892)

Type species: Meloidogyne exigua
Important Indian species - M. incognita, M. javanica, M.
arenaria, M. indica, M. graminicola
Hosts: Polyphagous (approximately 3000)
Symptoms: Typical root galls, size and nature of galls depend on
host crops.
Type of parasitism: Sedentary endoparasite of roots
Brief History
1855- Reverend Miles Joseph Berkeley (clergyman) noted galls on

cucumbers in greenhouse in England - first official report.
1879- Cornu - described a root-knot nematode as Anguillula marioni.
1884- Muller decided root-knot nematode was the same as Greeff's root-galling nematode
and that both should be Heterodera radicicola.
1887- Goeldi of Brazil described a root knot on coffee as Meloidogyne exigua.
1932- Goodey decided that use of H. radicicola was incorrect according to International
Rules of Zoological Nomenclature (ICZN) as original H. radicicola was not a root-knot
nematode. He renamed it Heterodera marioni.
32
1949 - Chitwood removed them from Heterodera on the basis that they differed from cyst
nematodes. Since the oldest name for the genuswas Goeldi's Meloidogyne, that name had
precedence.
Chitwood described five species based on perineal patterns.
The name Meloidogyne is derived from two Greek words meaning "apple-shaped" and
"female".
Now host range test (J.N. Sasser), chromosome counts (Triantaphyllou), juvenile head
structure (Eisenback), protein (gel electrophoresis patterns -Esbenshade) and DNA patterns
are used to separate species. Many of these approaches came out of International
Meloidogyne Project.
Important Identifying Features:

Second stage Juveniles (J2) –
 Vermiform, straight to arcuate upon death. Lip region with
coarse annules (1-4), a distinct labial disc; framework lightly
sclerotized, lateral sectors wider than submedians.
 Stylet slender, <20 μm.
 Excretory pore posterior to hemizonid.
 Median bulb with large oval valve plates.
 Tail with conspicuous hyaline region, tip narrow and
irregular in outline.
J3 and J4 represent spike tailed (retain moulted
cuticle) non-feeding stages.
Male:
 Vermiform, up to 2 mm long, tail end twisted,

develops by metamorphosis within a swollen
larvae.
 Cuticle strongly annulated, lateral field with 4
incisures.
 Lip region not sharply offset with distinct labial
disc and 1-3 annules; lateral sectors wider than
submedians.
 Stylet robust, 18-25 μm long with large knobs.
 Oesophageal glands mostly ventral to intestine.
 Spicules slender, generally 25-35 μm long,
gubernaculum 7-11 μm long.
 Testis single or paired (in case of sex reversal).
 Tail rounded.
33
 Phasmids dot-like, near cloacal aperture which is subterminal.
 Bursa absent.
Female:
 Round to pear-shaped with short projecting neck, white, sedentary. No cyst stage.
 Vulva and anus close together, terminal; perineum with a fingerprint-like cuticular
pattern, not elevated.
 Phasmids dot-like, slightly above and on either side of anus. Cuticle striated.
 Stylet slender, generally 12-15 μm long, with small knobs.
 Excretory pore anterior to median bulb, often closely behind base of stylet.
 Ovaries paired, prodelphic, convoluted.
 Rectal glands six, large secrete gelatinous material in which eggs are deposited; eggs
not retained in body.
34
Cyst Forming Nematodes
Schematic diagram of vulval cone of a Heterodera sp. (Enfaceview)
35
Cyst bifenestrate- H.avenae / H. filipjevi Cyst semi- or ambifenestrate- H. cajani/ H.

graminis/ H. mothi/ H. Oryzicola / H. sorghi/
H. zeae 36
Heterodera Schmidt, 1871

Second stage juvenile:
 Body slender, straight to arcuate.

 Lip region generally continuous; labial disc
indistinct.
 Stylet less than 30 μm long.
 Oesophageal glands filling body cavity, pointed
with prominent terminal hyaline part.
 Phasmids punctiform.
Type species: Heterodera schachtii A. Schmidt, 1871
Mature female and cyst:
 Lemon shaped with a short neck and terminal cone, turns into hard-walled cyst,
brown to black in colour, with a lace-like or zig-zag pattern.
 Vulva terminal. Anus dorsally subterminal, near vulva but not on vulval lip.
 Vulval fenestration present, ambifenestrate, bifenestrate or very rarely

circumfenestrate; anal fenestration absent.
 Underbridge generally present. Bullae present or absent. Eggs retained in body;

in some cases egg mass also present.
Male:
 Body twisted. Lateral fields with four incisures, outer band often areolated.
 Lip region generally offset by a constriction, and with 3-6 annules; labial disc
indistinct; basal lip annule may or may not be indented.
37
 Spicules robust, over 30 μm long, with blunt bifid or single tip.
 Tail very short, rounded.
Globodera Skarbilovich, 1959
Second stage juvenile:
Labial disc subcircular; distinct lip sector present.
Stylet less than 30 μm long.
Oesophageal glands filling body cavity.
Tail concial, pointed, with terminal half hyaline.
Phasmids punctiform.
Type species: Globodera rostochiensis
Mature female and cyst:
 Spheroid with a short projecting neck,

terminal region not forming a cone. Cyst brown, surface with a lace-like pattern
 Vulva terminal, slit length less than 15 μm; tuberculate area near vulva present.
 Vulval fenestrae circumfenestrate; underbridge and bullae rarely present.
 Anus dorsally subterminal not on dorsal lip, separated from vulva by a short distance,
but both lie in a terminal vulval basin; no anal fenestra.
 All eggs retained in body.
Male:
 Body twisted; tail short, rounded, less than anal body width long.
 Lateral field with 4 incisures, outer bands often areolated.
 Lip region offset by constriction, with indistinct labial disc, and 3-7 annules. Spicules
over 30 μm long, distally pointed.
38

Globodera Heterodera
Cysts rounded, globose, without Cyst lemon-shaped, with vulval cones
vulval cones
Cysts circumfenestrate Cysts bi-, ambi- or circumfenestrate
Meloidogyne Globodera/Heterodera
Second stage juveniles with weak stylet and Second stage juveniles with strong stylet and
poorly/ weakly developed cephalic well sclerotized cephalic framework
sclerotization
Tail with very short hyaline region Tail with long hyaline region
Third and fourth stage juveniles without Third and fourth stage juveniles with fully
fully developed stylet and oesophagus; developed stylet and oesophagus; not
enclosed within spike tailed cuticle of second enclosed within spike tailed cuticle of second
stage. stage.
Females not forming cysts; with perineal Females transformed into cyst upon death;
pattern; excretory pore anterior to median without perineal pattern; excretory pore
bulb posterior to median bulb
Gall forming Non gall forming
39
40
Ditylenchus
Species-D. dipsaci, D. angustus, D. destructor, D. myceliophagus

Host plants: Onion, oat, tulip, rice, potato, mushroom
Symptoms: D. angustus-Ufra disease of rice, D. destructor- tiny chalky spots (dry rot) on
tubers of potato, D. dipsaci- bloat of onion.
Type of parasitism: Endoparsite and ectoparasite of above ground parts, roots, tubers, bulbs
Morphometric features
 Both sexes are vermiform, finely striated cuticle

 Lip region low and flattened, continuous or slightly
set off
 Low sclerotization, stylet weak <15μm with basal
knobs
 Lateral fields with 4-6 lines
 Basal bulb of esophagus clavate or expanded with a
short overlapping on intestine
 Vulva posterior, monodelphic-prodelphic sometimes
reflexed
 Male with bursa-terminal/sub-terminal,
 Tail elongate conoid, sub-cylindrical to filiform.
Ditylenchus dipsaci
41
Stunt nematode -Tylenchorhynchus
Species-T. cylindricus, T. claytoni, T. annulatus, T. brassicae, T. mashhoodi

Hosts: Polyphagous (groundnut, tobacco, cabbage, maize, rice etc.)
Symptoms: Non-specific, stunting of root systems, yellowing of foliage and stunted tops,
poor growth of maize, ‘Kalahasty’ disease in groundnut induced by T. brevilineatus.
Type of parasitism: Ectoparasite of roots
Taxonomic features
 Nematodes 0.7 - 1.5 mm long, straight
to slightly curved
 Typical tylenchoid esophagus
 Stylet varies in length(15-35um)
 Esophageal glands as pyriform basal
bulb or rarely overlapping intestine
 Vulva median in position, ovaries
paired
 Tail conoid-rounded
 Small phasmids on tail
 Lateral fields with 4 lateral lines.
Tylenchorhynchus cylindricus
Reniform nematode –Rotylenchulus
Species-Rotylenchulus reniformis, R. parvus, R. macrodoratus

Major crop hosts: Polyphagous (vegetables, pulse, fruits, flowers etc.)
Symptoms: Non-specific,poor stand of seedling, nutrient deficiency symptoms-drying of leaf
margins, shedding of leaves, pruning of roots, plant stunted with less vigour.
Type of parasitism: Semi-endoparasite of roots
Identifying features of Rotylenchulus
Body posture/ Habitus : Heat relaxed immature
female, male and juvenile are C-shaped
Juvenile: Esophageal glands elongate, overlapping
intestine ventrally and laterally. Tail tapering to
rounded tip, hyaline region shorter than adults.
Immature female: Vermiform, cephalic region
high, continuous, sclerotised. Dorsal esophageal
gland opens at one stylet length posterior to stylet
42
knobs. Esophageal glands overlap ventrally. Tail elongate- conoid with long hyaline terminal
region.
Mature female: Kidney-shaped, didelphic, ovaries reflexed or coiled. Tail- a short
projection.
Male: Stylet weaker than in females, esophagus reduced, sclerotisation weaker. Bursa low,
not enclosing tail tip. Tail elongate-conoid, terminal portion hyaline.
Lesion (meadow) nematode –Pratylenchus
Species-P. pratensis, P. penetrans, P. zeae, P. coffeae, P. indicus
Major crop hosts: Polyphagous (maize, tobacco, potato, banana, tea, coffee, etc.)
Symptoms: Lesions on roots, decay and rotting of roots due to cell death followed by
secondary infection of soil-borne pathogens; brown
root-rot of flue cured tobacco.
Type of parasitism: Migratory endoparasite of
roots
Identifying features of Pratylenchus
 Body - small, 0.4-0.8 mm in length, slightly
curved ventrally when relaxed
 Head- truncated, head framework
sclerotised, appearing as black line under
low magnification
 Stylet- short but robust with round knobs
 Esophageal glands overlap intestine ventrally
 Vulva posterior (75-85%), ovary-monodelphic-prodelphic, post uterine sac present
 Tail conoid-rounded or subcylindrical-smooth/crenate
 Males rare or absent, spicules - slender, arcuate, bursa extending to tail tip.
Burrowing nematode- Radopholus

Species-Radopholus similis, R. citrophilus, R. colbrani
Major crop host: Black pepper, banana, citrus, coconut, ginger etc.
Symptoms: Non-specific, ‘slow wilt’ or ‘yellows’ in pepper, ‘toppling’ of banana and
‘spreading decline’ of citrus.
Type of parasitism: Migratory endoparasite of roots
Morphology of Radopholus
 Small (<1 mm), cylindrical, cephalic frame work low, continuous/set off, stylet well
developed both in female and juvenile
 Sexual dimorphism marked, males degenerated in anterior region (stylet and
esophagus), male head high rounded and reduced
 Lateral fields with 3-7 lines
 Esophageal gland overlap intestine dorsally
 Ovaries- amphidelphic, outstretched
 Female tail elongate conoid/sub-cylindroid
 Male tail with bursa-terminal to sub-terminal.
43
Radopholus similis
Awl nematode-Dolichodorus
Identifying Features
 Body large 1.5-3mm, head set-off with labial disc,

massive cephalic sclerotization
 Stylet 54-170µ with basal knob; conus long &
tubular
 Basal bulb overlap intestine slightly
 LF with 3 lines, areolated
 Ovary amphidelphic outstretched
 Female tail convex conoid to filiform, tail tip
conically pointed, post rectal intestinal sac usually
filling the tail
 In male bursa trilobbed enveloping tail
Type species: Dolichodorus heterocephalus
44
Citrus nematode- Tylenchulus semipenetrans
Symptoms: Slow decline, dirty appearance of root, reported to induce ‘citrus dieback’
symptoms.
Identifying Features
Mature female:
 Dorsal esophageal gland opening is 1/3 to 1/2 stylet length
behind the stylet knobs.
 Basal bulb set off from the intestine.
 Excretory gland large and opens in a pore.
 Post-vulval region elongate and tapering.
 Vulval lips bulging, post-vulval sac absent.
 Short blunt tail.
Immature female:
Straight to ventrally curved, ovary immature, excretory pore
posterior. Vulval lips thick, bulging, anus obscure.
Male:
Less than 0.5 mm long, stylet and oesophagus are partially degenerate. Excretory pore
posterior at 50-60%. Bursa absent.
Juvenile (J2):
Straight to slightly curved. Two lateral lines, narrow lateral field. Excretory pore
posterior at 50-60%. Lip framework moderately sclerotized. Basal bulb set off from
intestine. Anus obscure.
45
Aphelenchoides(Foliar nematode) Aphelenchus

Type Species-Aphelenchoides kuehnii Type Species - Aphelenchus aveane
Female Female
o Nematodes slender, 0.8-1.5 mm long. o Straight or ventrally arcuate when
Lip region is set off . fixed, 0.8- 1.4 mm long,lip region
o Stylet small, slender with or without continuous.
small knobs o Stylet weak, without basal swelling.
o Median bulb large and conspicuous o Median bulb rounded-rectangular in
o Oesophageal gland lobe originating from shape and more or less filling body
median bulb itself and overlapping diamaeter.
intestine dorsally and hanging. Isthmus o Oesophageal gland lobe originating
region is lacking. from a short isthmus and overlapping
o Vulva posterior (60-75%), ovary – intestine dorsally.
monoprodelphic with post uterine sac.
o Tail –conoid, with or without mucro o Vulva posterior, Ovary –
monoprodelphic with post uterine sac.
o Tail short-cylindrical, round end
Male
Walking stick / J-shaped, spicules “rose- Straight or ventrally arcuate, spicules
thorn” shaped, no bursa. arcuate, bursa covering entire tail, provided
with bursal rays
46
Aphelenchoides(Foliar nematode) Aphelenchus

Type Species-Aphelenchoides kuehnii Type Species - Aphelenchus aveane
47
1. ROOT KNOT NEMATODES (Meloidogyne spp.)
M. J. Berkeley in 1855, first reported the occurrence of root galls on glass house
grown cucumbers at Rothamstead Experimental Station in England. Soon after this nematode
was recorded by various other people who assigned it by different names. In India, Barber
(1901) recorded this nematode for the first time on tea from Kerala.
Root knot nematodes, Meloidogyne spp., are economically important plant parasite
and distributed worldwide. They are obligate parasites and parasitize thousands of different
plant species including monocotyledons, dicotyledon, herbaceous and woody plants. They
reduce the yields as well as the quality of the produce. Common name, root knot nematode is
given to the species under genus Meloidogyne due to characteristic symptom‘knot’ or ‘gall’,
induced by them on roots.
ECONOMIC IMPORTANCE
The extent of crop losses depends on the initial nematode population, susceptibility of
the crop, cropping sequence, age of the plant, soil fertility and involvement of secondary
plant pathogens.The precise estimates are difficult to make due to uneven distribution of
nematodes in field and non-availability of standard specific nematicides. On a world basis,
crop losses to the extent of 5% have been estimated due to root knot nematodes. Losses in
individual fields are, however, highly variable, may be as high as 25-50%; 28-90% losses in
okra, 48% in tomato and 27-33.7% in brinjal was estimated due to Meloidogyne incognita.
DISTRIBUTION
In a roving survey, Mukhopadhyay and Roy (2006) observed M. incognita and M.
javanicaasprominent plant parasite of solanaceous, leguminous, cucurbitaceous and
cruciferous vegetable crops in eastern and north eastern states of India. They are prevalent in
all the districts of West Bengal.
SYMPTOMS
1. Unthrifty patchy plant growth with leaf chlorosis

2. Wilting during hotter part of the day
3. Knot or gall or forking of the root is the main diagnostic symptom of root-knot nematode
infestation (Fig. 1)
4. Intensity of fungal or bacterial wilt disease increases when occurs in association with
root-knot nematodes. Nitrogen fixation of leguminous plants is impaired severely.
LIFE CYCLE
Root-knot nematodes represent marked sexual dimorphism; the females are pyriform,
swollen or saccate, and the males are vermiform. The embryonic development results in a
first-stage juvenile which moults first time in the egg itself and comes out of egg shell as a
second-stage juvenile. This motile, vermiform, and infective stage migrates through the soil
and enter the root just behind the root cap of a suitable host plant and lie parallel to the steler
region.It moves through the plant tissue to a preferred feeding site and establishes a complex
host-parasite relationship with the plant. The second-stage juvenile becomes sedentary and
resume feeding on special nurse cells (giant cells), it undergoes more morphological changes.
It becomes flask-shaped and without further feeding moults three times into the third- and
48
fourth stage juvenile, and finally becomes an adult. Shortly after the last moult the saccate
adult female resumes feeding and continues to do so for the remainder of her life. The adult
male does not feed. It will leave the root and move freely in the soil. Depending on the mode
of reproductioni.e. amphimixis or parthenogenesis of a particular species, the male may
search for a female to mate, or remain in the soil and finally die. Root tissues around the
nematode and the giant cells undergo hyperplasia and hypertrophy resulting in the
characteristic root gall. Galls usually develop 12 days after juvenile penetration. The length
of the life-cycle of root-knot nematodes is greatly influenced by temperature. For
Meloidogyne incognita on tomato at approximately 29ºC temperature, the first adult female
appears in about 13-15 days after root penetration. The life span of egg-producing females
may extend from 2 to 3 months, but that of males be much shorter.M. incognita could
develop eight or more generations per year on tomato, with the length of the life cycle
dependent on soil temperature. One cycle was completed in 19 days at 30.6°C versus 43 days
at 21.8°C.
2. RENIFORM NEMATODE (Rotylecnhulus reniformis)
It was for the first time recorded in USA on cowpea (Linford and Oliveira,1940).
Seshadri and Sivakumar (1963) recorded it for the first time on cotton in India. Among all the
species of this genus, Rotylenchulus reniformis is economically important.After Meloidogyne,
the reniform nematode, R. reniformis, is the most important nematode affecting vegetable
crops. The nematode attacks over 100 plant species including many vegetable crops and is a
limiting factor in vegetable production, but is often neglected or overlooked where it occurs
concomitantly with Meloidogyne. Reniform nematodecan survive and reproduce well on
weed hosts likeOldanlandia corymbosa,Phyllanthus niruri, Euphorbia sp., Blainvillea
lattifolia, Vernonia cinerea, Mikania scandens and Trianthema portulacastrum.The
nematode has been detected in more than 36 countries (Heald and Thames, 1982). R.
reniformis was recorded as the most important nematode in vegetable crops of Tripura and
Nagaland.In West Bengal (India) it was most abundant on cucurbitaceous vegetable crops
followed by solanaceous and leguminous vegetable crops.
ECONOMIC IMPORTANCE
In an experimentwith initial population of 595–636 reniform nematodes per 200 cm3
of soil, 10.5-14.5% yield of green pod of cowpea could be saved by application of carbofuran
3G @ 2 kg a.i. ha-1 at the time of sowing (Roy et al., 2008).
SYMPTOMS
Above-ground symptoms include stunting and leaf curling (Singh and Khera,
1979).Root necrosis and cortical necrosis have been observed following infection.Cantaloupe
growing in heavily infested soilwas badly stunted and yields were greatlyreduced (Heald,
1975). Leaf chlorosis canbe produced (Bridge, 1983). Females and their adhering egg masses
can be easily observed under the dissecting microscope. Soil adhering to the gelatinous egg
masses often gives them a dark appearance, aiding in detection. Severely infested plants lack
feeder roots and appear sick.
49
DAMAGE THRESHOLD
An initial population density of 1000 nematodes per 500 cm3 of soil is considered as
the minimum density level of R. reniformis for expression of pathogenic effects on brinjal
(Roy et al., 2008).
LIFE CYCLE
Immature females penetrate the root and become sedentary. Galls are not produced.
The life cycle is completed on okra in 24-29 days (Sivakumar and Seshadri, 1971). The
existence of amphimictic and parthenogenetic races of R. reniformis has been demonstrated
by Hirschmann and Triantaphyllou (1964).
The reniform nematode can survive in moist soil in the absence of hosts for 7 months,
and for 6 months in dry soil. Stoyanov (1971) reported that R. reniformis was able to survive
29 months in the absence of host plants. The intensity of brinjal mosaic virus and okra yellow
vein mosaic was promoted on plants parasitized by R. reniformis. Charcoal rot caused by
Macrophomena phaseolina on cantaloupe was significantly higher when the roots were
infested with the reniform nematode.
3. CYST NEMATODES
The cyst nematodes comprise a group of different genera, i.e. Heterodera, Globodera,
Punctodera, Cactodera, Dolicodera, Afenestrata, Brevicephalodera etc. But only Heterodera
and Globodera are considered economically important in India. Cyst nematodes are host
specific.
SYMPTOMS
Cyst nematode infested plants are less vigorous, stunted or discoloured. Such plants
typically yield less than a healthy plant. Evidence of the mechanisms and economic impact of
cyst nematode damage to crops can be found in numerous sources. Many plant functions are
related to photosynthesis. The net photosynthetic rate of plants is probably suppressed by all
cyst nematodes, especially at high levels of infection. Globodera pallida suppresses
photosynthetic rates of potato at high inoculum densities. Moisture stress decreases the
photosynthetic rate of plants. If nematode damage is sufficient to decrease water uptake and
induce a moisture-stress condition in the plant, then photosynthesis is expected to decrease.
On the other hand, even if significant root damage occurs, but the plant is still able to obtain
adequate moisture (as might occur in fine textured soil); the overall damage might not be
great because the rate of photosynthesis would remain unaltered.
4.ROOT LESION NEMATODE (Pratylenchus spp.)
Root-lesion nematodes of the genus Pratylenchus rank third only to root-knot and cyst
nematodes in terms of their worldwide economic impact on crops. Pratylenchus species are
obligate biotrophic, soil-inhabiting parasites and are found in all agricultural regions of the
world.Lesion nematodes are important parasites of many crops and are known to form
disease complexes with many different soil-borne root rotting fungi, thereby increasing root
damage.
50
SYMPTOMS
Feeding by root-lesion nematodesproduces lesions on roots which initially appear as
water soaked areas on the root surface. These sites later become yellow and elliptical, and
eventually develop dark brown centres as nematodes continue to feed. Further feeding and
migration produces galleries within the cortex, and discreet brown lesions of necrosis usually
appear in two to four weeks but may form as fast as 96 hours after invasion. Lesions form
due to production of phytotoxic substances such as high concentrations of phenolic
compounds. The necrosis (tissue breakdown) which causes the lesions often appears to
advance ahead of the migrating nematode. The entire circumference of roots may be girdled
when too many nematodes invade a root in the same small area. The combination of
structural damage from feeding and physiological damage due to production of phytotoxic
compounds reduces the amount of functional root length, restricting adequate water and
nutrient uptake for the plant. In addition, the necrotic lesions provide opportunities for
invasion by other microbial pathogens in the soil.
Field symptoms of damage from root-lesion nematodes generally occur as circular to
irregular patches, perhaps 30-150 feet in diameter that have thin stand and stunted plants.
Presence of root-lesion nematodes can often be suspected by looking for reddish-brown
lesions on roots.
5.STEM AND BULB NEMATODE (Ditylenchus dipsaci)
The stem and bulb nematode Ditylenchus dipsaci is a migratory endoparasite that has
a five-stage life cycle and the ability to enter into a dormancy stage. D. dipsaci enters
through stoma or plant wounds and create galls or malformations in plant growth. This allows
for the entrance of secondary pathogens like fungi and bacteria. D. dipsaci is economically
detrimental because infected crops are unmarketable.Ditylenchus dipsaci is not prevalent in
India.
SYMPTOMS
Penetration of onion leaves by this endoparasite causes leaf deformation and leaf
swellings or blister-like areas on the surface. The leaves grow in a disorderly fashion and
often hang as if wilted. As the season progresses, they become chlorotic. Infected onions
become bloat (swollen) and the bulbs may rot during storage. The inner scales of the bulb are
usually more severely attacked than the outer scales. As the season proceeds, the bulbs
become soft, show browning of the scales in concentric circleswhen cut open. Conversely, D.
dipsaci on garlic does not induce deformation or swellings, but causes leaf yellowing and
death.
Small seeds exhibit no discernible symptoms of infestation while larger seeds (e.g.
Phaseolus vulgaris and Vicia faba) the skin may be shrunken and show discoloured spots.
6. POTATO ROT NEMATODE(Ditylenchus destructor)
D. destructor is also known as iris nematode, the potato tuber eelworm and the potato
tuber nematode. It is an endoparasitic, migratory nematode commonly found in areas such as
the United States, Europe, central Asia and Southern Africa.
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SYMPTOMS (D. destructor)
Common symptoms of infestation of D. destructoron garlic are discoloration and
rotting of the plant tissue. In potato symptoms are appeared as white spots under the skin.
Badly affected tubers have slightly sunken areas with cracked and papery skin, the tissue
below the skin is darkened and may have a mealy or spongy appearance. Symptoms may be
more visible after storage. There is in general a secondaryinvasion of fungi, bacteria and free-
living nematodes.
Infestations on flower bulbs and cormsusually begin at the base of the bulb and extend
up to the ﬂeshy scales with yellow to dark brown lesions. Secondary rotting may occur and
the bulbs can be destroyed. Specimens of D. destructor are accumulated on the boundary
between distinctly diseased parts and healthy sections, but are rarely isolated from completely
decayed tissues.
Damage to carrots appears as transverse cracks in the skin with white patches in the
sub-cortical tissue. The patches are easily seen in a transverse cut. Infested areas are subject
to secondary infections by fungi and bacteria resulting in decay and rot.
7.DAGGER NEMATODES (Xiphinema spp.)
The genus,Xiphinema represents ectoparasitic root nematodes commonly known as
the Dagger nematode. They are easily recognizable, due to a long body length and a long
stylet capable of reaching vascular tissue. They are most feared because of their capability to
act indirectly as virus vectors of various NEPOviruses. The virus is often vectored by specific
species and attaches to the interior cuticle lining where it can be transferred from infected to
uninfected root tissue as the nematode feeds and sheds.
SYMPTOMS
The symptoms that plants exhibit in response to the pathogenicity of Xiphinema
americanum are similar to those of other migratory ectoparasitic nematodes of roots. It is
common to see poor growth and or stunting of the plant, yellowing or wilting of the foliage,
and reduced root systems which can include root necrosis, lack of feeder or secondary roots,
and occasional tufts of stubby rootlets.
X. index was discovered as a vector of grapevine fan leaf virus disease in the 1950s. It
causes yellow mosaic and bands on the leaves, and may result in an 80% reduction in fruit
set, causing huge economic losses. It is spread by contaminated seed and the grafting of
infected rootstocks.X. americanum is an important transmitter of various plant viruses
including tomato ring spot nepovirus, tobacco ring spot nepovirus, peach rosette mosaic
nepovirus and cherry rasp leaf nepovirus.
8. CITRUS NEMATODE (Tylenchulus semipenetrans)
The citrus nematode,Tylenchulus semipenetrans was first discovered in California in
1913 by JR Hodges and was later described and named by NA Cobb that year. In India, its
occurrence was first reported by Siddiqi (1961) from Aligarh, Uttar Pradesh.It is a causal
agent of the slow decline disease of citrus and found inalmost all citrus growing areas across
the world. Their feeding strategy is semi-endoparasitic and has a very narrow host range
among commonly grown crops. These nematodes are considered as major plant-parasitic
52
nematode because they can cause 10-30% losses on citrus yield. They also parasitize other
hosts such as olive, grape, persimmon and lilac.
SYMPTOMS
Symptom development depends on overall conditionsof citrus orchard. Infested trees
growing under optimum conditions may yield somewhat less fruit while appearing quite
healthy. Under unfavourable conditions for tree growth, the effects of citrus nematode
parasitism are more evident. In new citrus plantings, symptom developmentprogresses slowly
as nematode populations develop to high levels. Symptoms include poor root development
accompanied with small chloroticleaves. Heavily infected feeder roots are slightly thicker
than healthy roots and have a dirty appearance due to soil particles that adhere to gelatinous
egg masses on the root surface. Feeder roots decay more rapidly due to lossof integrity at the
epidermis and invasion of feeding sites by secondary microorganisms. This may be expressed
as lesions on lightly infected roots, while heavy infections result in cortical sloughing and
rootdeath.The incidence of fungal pathogens Fusarium solani and F. oxysporium is enhanced
in the presence of citrus nematode resulting in root decay and lowering the tree vitality.
9. BURROWING NEMATODE (Radopholus similis)
Theburrowing nematode, R. similis, was firstobserved by Cobb in necrotic tissue of
theroots of Musa sp. sent to him in New South Wales from Fiji in July, 1891. The common
name ‘burrowing nematode’ was coined because it causes extensive cavity formation in roots
due to migratory behaviour. In India, it was introduced into Kerala along with banana suckers
from some other country. Since its first record in 1966, R. similis have been recorded from
Tamil Nadu, Karnataka, Andra Pradesh, Goa, Maharashtra, Gujarat, Madhya Pradesh,Assam
and Tripura. The host range of R. similis is very wide, yet it is a serious problem of banana,
pepper, coffee, tea, cocoa, coconut, arecanut, sugarcane, turmeric, ginger etc. A population of
R. similis (known as R. citrophilus) is confined to Florida state of USA, where it attacks
citrus only.
SYMPTOMS
The disease of banana caused by R. similisis known throughout the world by
differentnames, the most common are ‘black headtoppling disease’, ‘banana decline’,
‘rhizome rot’, ‘banana root rot’ and ‘toppling disease’.
The most obvious symptom of attack of R.similis on banana is the toppling over or
uprooting of plants especiallythose bearing fruit, but there is a range ingradation in the
severity of damage, fromthe lengthening of the vegetative cycle tothe drastic reduction in
bunch weight. Thisreveals two types of damage in banana plantations- (i) affecting
theanchorage of the plant and (ii)reduce the ability to take up waterand nutrients. Severaldark
red lesions appear on the outer part ofthe root penetrating throughout the cortexbut not in the
stele, adjacentlesions may coalesce and the cortical roottissue atrophies and later turns black.
Inheavy infestations, the lesion girdles theroots. Nematodes can migrate from infectedroots
into the corm, causing diffuse blacklesions which may then spread around the corm. Roots
grow out of thecorm become infected. Uprooting occurs commonly inwindstorms or if heavy
rains loosen thesoil. The mechanical stresses on the rootsystem are often increased by the
naturalangle of leaning which develops as fruitbunches grow. Nematode feeding and
53
movement causes severe necrosis and cavity formation within the cortex. The cavities
coalesce and break down leading to tunnel formation. Eggs are often laid in these cavities,
while nematodes move to adjacent healthy tissues. Three to four weeks after infection, deep
cracks appear on the root surface due to breakdown of tunnels. The presence of a number of
fungi in nematode induced lesions probablyhastens the destruction of roots andmay
contribute to toppling disease becausefungi colonize the stele which is not penetratedby R.
similis.R. similis interacts with Fusarium oxysporum f. sp. cubense and causes Panama wilt.
The incidence of wilt is doubled in presence of R. similis. The nematode also causes
resistance breakdown to Panama wilt in Lakatan bananas.
R. similiscauses ‘yellows’ disease in black pepper. The primary symptom of the
yellows (slowwilt) disease is the appearance of pale yellow or whitish yellow drooping leaves
onthe vines. The number of such leavesincreases gradually until large numbers ofleaves or
even the entire foliage becomes yellow. Yellowing is followedby shedding of leaves,
cessation of growth and dieback symptoms. Thesymptoms are very pronounced when
soilmoisture is depleted. In the very early stageof the disease in India, the symptoms
maydisappear with the onset of the south-westmonsoon, resulting in an apparentlyhealthy
appearance of such plants in thefollowing years because of new leaf growthand shedding of
yellowed leaves. This hasoften given an incorrect impression of thedisease being caused by
soil moisturestress rather than nematodes. However,within 3–5 years of initiation of
yellowing,all the leaves are shed and death of thevine takes place, and hence the name
‘slowwilt’ disease. In bearing vines, shedding of spikes (inflorescences) is a major
symptom.Large numbers of shed spikes are seen atthe base of the affected vines. In large
plantations,affected patches become conspicuousinitially as yellowed plants and laterwith
large numbers of barren standards that have lost the vines, or standardssupporting dead vines
without anyleaves. Young and old plants are affectedand the replanted vines normally
diewithin 2 years. The tender thin, white feeding rootsshow typical orange to purple
colouredlesions. Lesions are not clearly seen onolder roots, being brown in colour. Theroot
system exhibits extensive rotting andthe main roots are devoid of fine feederroots that rot
quickly. Extensive necrosisof larger lateral roots develops over time.
LIFE CYCLE
R. similis is a migratory endoparasitic species which is able to complete its life cycle
within the root cortex. The nematode completes its life cycle in about 21 days at 25°C. The
eggs hatch after 8–10 days and the juvenile stages are completed in 10–13 days. Females and
juveniles feed inside roots, especially near the tips. Males with their weak stylets do not feed.
Females lay two to six eggs per day for 2 weeks.
10. FOLIAR NEMATODE(Aphelenchoides spp.)
Aphelenchoides besseyi was described by Christie in 1942. This species is often
associated with rice plants. A. besseyi causes “white tip” disease of rice in Japan and the
southern United States. While the most economically significant plant host for A. besseyi is
rice, this nematode has also been recorded on many ornamental plant hosts. A. besseyi can be
distinguished from other Aphelenchoides species by having a lateral field with four lines, a
stellate mucro at the tail, and shorter postvulval uterine sac than A. ritzemabosi .
54
The first report of foliar nematode infestation on tuberose (Polianthestuberosa L.),
inducing foliar disease was in Hawaii (Holtzmann, 1968). In India, ‘floral malady’ of
tuberose induced by A. besseyi was reported for the first time from Ranaghat areas of the
Nadia district of West Bengal (Chakraborti and Ghosh, 1993). A. besseyi had been reported to
cause a serious foliar disease of tuberose in the Mekong Delta of Vietnam. Widespread
occurrence of foliar nematode infesting rice, tuberose, gladiolus and onion was reported from
West Bengal (Roy et al., 2010; Roy et al., 2011a & 2011b).
A. besseyi is often found in warmer climates whileA. ritzemabosi and A. fragariae are
more commonly associated with temperate climates, even though the two later species are
found in both tropical and temperate localities. While all three species infect a wide variety of
ornamental and field crop plants. BesidesJapan and India, it has been recorded from USA,
Italy, South-east Asian countries, Central and West Africa and South Australia. In India it is
prevalent in eastern and southern states.
SYMPTOMS
Infestation of A. besseyi on tuberose led to initial appearance of light chlorotic lesions
along the mid axis (rib) of the leaf starting from base extending upwards but never touches
the apex. Subsequently, it became yellowish and finally turned to brown or black, elongated
necrotic spot. Infected leaves later on die and abscise. Ectoparasitic feeding of A. besseyi on
the scape (floral stalk) leads to the formation of prickle like rough, irregular structure. Scape
becomes curled and twisted. Scapes of the infected plants are many times smaller than
healthy and normal scapes.Endoparasitic feeding of A. besseyi was observed on leaves, bracts
and tepals (petals) where brownish to black necrotic spots of varying shapes and sizes were
developed. Often infected plant bears small sized twisted flowers. Sometimes inflorescence
gets aborted completely. Nematode may survive on bulb, leaves, scapes, bracts, tepals and
anthers but not in the soil (Roy et al., 2010; Roy et al., 2011b).Second year onwards tuberose
crops suffered most from infestation of foliar nematodes; this may led to annual yield loss of
35% in 'Calcutta Double' as against 63% in ‘Calcutta Single’ cultivar. International
quarantine exists against A. besseyi, hence, their presence on cut flowers may limit the
potential of flower export.
Both A. fragariae and A. ritzemabosi are temperate climate species and are
widespread in Europe, Canada, Australia, New Zealand and South Africa. In India, these
have been reported from Jammu and Kashmir and Himachal Pradesh. The preferred hosts of
A. ritzemabosi are plants belonging to family Asteraceae or Compositae with chrysanthemum
as main host, while A. fragariaeis considered a problem on strawberry but its host range
extends to families Liliaceae, Ranunculaceae, Primulaceae, besides some ferns.
Both the species feed ectoparasitically on buds and endoparasitically on leaves.
During high humidity and rainy season, the nematodes ascend up the stem in a thin film of
water covering the plant, enter the leaves through stomatal openings and feed on mesophyll
tissues. Infection spreads from lower to upper leaves and symptoms appear as tiny brown
spots on leaves initially which enlarge to acquire inter-veinal angular spots. Nematode
feeding on buds results in a ‘blind’ plant (A. fragariae on strawberry) or undersized and
distorted flowers (A. ritzemabosi on chrysanthemum).
55
A. fragariaecauses ‘cauliflower’ disease in strawberry in the presence of bacterium
Clavibacter fasciens.The strawberry plants were stunted and deformed so that the crown and
lateral branches resembled like a cauliflower.
MANAGEMENT OF NEMATODES
Management of nematodes is difficult. The most reliable practices are preventive,

including sanitation and choice of plant varieties. We can reduce existing infestations through
fallowing, crop rotation, and soil solarization. However, these methods reduce nematodes
primarily in the top layer of the soil, hence they are effective only for about a year. They are
suitable primarily for annual plants or to help young woody plants to establish. Once
nematodes infest an area or crop, try to minimize damage by adjusting planting dates to
cooler times of the season when nematodes are less active. Try to provide optimal conditions
for plant growth including sufficient irrigation and soil amendments to make plants more
tolerant to nematode infestation.
All the control methods are directed to bring down the population of nematodes in
soil, low enough to raise the crop profitably. Usually a combination of the control methods is
needed to raise crops economically. Nematodes are omnipresent and can be transmitted
through a variety of methods. Irrigation water is the prime source of transfer followed by use
of infected composts, soil.
INTEGRATED MANAGEMENT OF NEMATODE PESTS OF VEGETABLE CROPS

A. Nursery management-
1. Plastic mulching
2. Rabbing the nursery bed by burning of rice husk @20kg/m2. Later mix the ash with the soil
and sow seeds one week later.
3. Nursery application of granular nematicide e.g. carbofuran 3G @ 0.3-0.5g a.i./m2
4. Seed soaking in nematicide solution for 1-2hrs/ seed dressing with carbosulfan 25DS @2-
3% a.i. w/w or Trichodermaviride orPseudomonas fluorescens @4g/kg seed.
B. Main field management-
1. Summer ploughing
2. Crop rotation with non host crop/ antagonistic crop
3. Intercropping with antagonistic crop
4. Apply organic cakes either in row or pit
5. Grow resistant variety
One of the best ways to manage nematodes is to use crop varieties that are resistant to
nematode injury. Tomato varieties with the code VFN (Verticillium, Fusarium, Nematodes)
on the seed packet or label are resistant to common root knot nematode species. Although
even resistant tomato varieties can still exhibit some root galling under high nematode levels,
they usually maintain their yield. For example in recent vegetable garden-type experiments
on root knot nematode soil, nematode-resistant tomatoes yielded almost 6 times more
tomatoes than a similar susceptible variety. An additional benefit of growing a resistant
variety is the nematode levels in the soil decline rather than increase, making it more feasible
to grow a susceptible crop the following season.
56
Crop cultivar having resistance to plant parasitic nematodes
Crop Resistant/Tolerant Variety Nematode Pest
Brinjal Jiant of Benaras, Gola, Black Beauty,
Meloidogyne spp.
PBR-91-2
Chilli Pusa Jawala, Suryamukhi, NP-46A,
Meloidogyne spp.
Biswa Sundari, Mohini
Tomato Hisar Lalit, Mangala Hybrid, Hisar-N-
1, Meloidogyne spp.
Hisar-N-2
Okra Contai Local Green, Baishali Badhu Meloidogyne spp.
Pumpkin Dasna, Jaipuri Meloidogyne spp.
Watermelon Sahjahanpuri Meloidogyne spp.
Cowpea GAU-1 Meloidogyne spp.
Potato Kufri Swarna, Kufri Thenmalai Globodera rostochiensis, G. pallida
6. Apply bioagents like Purpureocillium lilacinum (= Paecilomyces lilacinus) or T. viride or

P. fluorescens orIsaria fumosorosea or Pochonia chlamydosporia @ 2.5kg/ha
7. Apply chemical nematicides e.g carbofuran 3G or cartap hydrochloride 4G @ 1kg a.i./ha
MANAGEMENT OF TUBEROSE FOLIAR NEMATODE
o Fairly tolerant variety e.g. Phule Rajani and Prajwal can be grown.
o Prevent spreading of irrigation water from infected field to healthy field.Wider spacing
prevent dissemination of nematode from infected to healthy plant.
o Select nematode free bulb for planting.
o Infected plants and inflorescence should be either burnt or buried into field.
o Bulb Treatment: (i) Presoaking of bulbs in plain water for 4hrs to bring anhydrobiotic
form of nematode into hydrobiotic condition. Then treat the bulbs in monocrotophos
36%SL or cartap hydrochloride 50SP @ 750 ppm for 2hrs. Dry bulbs under shade and
plant thereafter.(ii) Hot water treatment of bulbs at 50-550C for 10-15 min can be
advocated. Here also bulb should be presoaked for 4hrs in plain water. It is rather tricky
to exercise because farmers face trouble to maintain 50-550C temperature constantly for
10-15 minutes. Here we can utilize paraffin wax whose melting point is closer to our
need. To do this, simply put a bit of wax into boiled water when it starts to melt then dip
the bulbs and intermittently add boiling water to maintain melting condition of wax. This
can assure that water was having incessantly 500C or slightly more temperature.
o Foliar spray: Fifteen days after sprouting of bulb prophylactic spraying withcartap
hydrochloride 50SP or carbosulfan 25 EC @ 250g a.i./hamay be recommended in
problematic areas. If infestation persists then 2-3 sprays at 10 days interval may be given.
Three to five sprays may berequiredfor the second and third year crop. Same nematicide
57
should not be repeated. Use sticker while spraying. Chlorfenapyr 10SC @ 75g a.i./ha can
also be used in place of carbosulfan orcartap hydrochloride.
MANAGEMENT OF BURROWING NEMATODE
The rhizomes are the source of primary inoculum, therefore, it is very important to use
nematode-free rhizomes for planting. Rhizomes can be disinfected with nematodes by
different methods, viz.
o Paring: Removing the necrotic tissues along with some adjacent healthy portion with the
help of a sterilized knife before planting.
o Paring and pralinage: Since the pared sets are more prone to secondary infection by
fungi etc. application of Bordeaux mixture + DBCP paste (20 kg hydrated lime, 20 kg of
copper sulphate, 1,288 ml of 70% DBCP and 455litres of water) on the cut surfaces
protects the rhizomes, besides killing the residual nematode inoculum.
o Paring and hot water treatment: The pared sets can be given hot water treatment at 53-
55ºC for 20-25 minutes before planting.
o Paring and nematicidal coating: The pared sets can be dipped in mud slurry followed by
sprinkling of carbofuran granules on the rhizome surface @ 1.2g a.i./ rhizome.
MANAGEMENT OF THE CITRUS NEMATODE
Management practices consist of exclusion, preventive measures and post-planting

nematicide applications.
Good sanitation practices are essential to avoid nematode infestations. Use certified
nematode-free material for planting. Rotation with annual crops for 1 to 3 years before
replanting citrus helps to reduce citrus nematode populations. Nematodes can easily be
removed from seedlings by dipping the roots in 45oC water for 25 min, this kills the
nematodes but does not harm the plant. The container-grown citrus can be treated with steam
and soil solarization.
Using a resistant rootstock is recommended whether or not nematodes are present.
Trifoliate orange (Poncirus trifolaiata) is known to be tolerant to citrus nematode. Troyer
citrange is also resistant to citrus nematode, but this nematode has resistance-breaking
biotypes that may develop on this rootstock after a period of time, thus increasing its
susceptibility. Currently, the use of resistant hybrid rootstock called Swingle citrumelo
(Citrus paradisi x P. trifoliata) and certified propagative citrus plants free from nematode
parasites of citrus are promising strategies for preventing the damage induced by Tylenchulus
semipenetrans to citrus. This practice has greatly reduced the spread of this parasite in new
Florida lands free from existing nematode infestations. Resistant (Ramsey) or moderately
resistant (vinfera Dog Ridge) grape rootstocks were also used successfully in California
vineyards.
If the site was previously infested with nematode pests of citrus, preplant nematicidal
application may be necessary to reduce nematode population levels. When replanting a citrus
orchard, a preplant treatment is recommended even if a resistant rootstock is used. Trees
planted on treated orchard sites are generally known to have improved growth and yields
compared to those on untreated sites.Tree basin area treatment with carbofuran @ 4 kg a.i.
58
per ha or combined application of neem cake @ 1 kg / tree + carbofuran @ 2 kg a.i./ha may
be advocated.Interculture of onion, garlic and Tagetes help to restrain population build up.
In established orchards, treat when sampling indicates more than 400 female citrus
nematodes are present in 1 gram of roots in February to April or more than 700 in 1 gram of
roots during May and June.
59
General Concepts of Nematode Management
For Nematode Management
 Correct identification of nematode species and
 Quantification nematode population
are necessary.
Nematode control- Complete eradication (invariably unattainable)
Nematode management- advocates keeping pests population below ETL, so that economic losses do
not occur
Tactics/Methods of nematode management-
A. Cultural methods-
1. Crop rotation
2. Fallowing and summer ploughing
3. Flooding
4. Organic manuring /soil amendments
5. Enemy plant/antagonistic crop
6. Trap crop
7. Propagation through healthy planting material
8. Removal and destruction of infested plants
9. Time of sowing
B. Physical methods-
1. Soil solarization
2. Hot water treatments
C. Biological methods
D. Host resistance/ resistant variety
E. Chemical methods
F. Regulatory methods
A. Cultural methods-
1. Crop rotation-Cultivation of non host/ poor host crop in between two susceptible ones, non host/
poor host crop –economic for the selected area and should not invite any new pest or pathogen. e.g.
crop rotation with mustard, sesame, rice, wheat and maize can bring down rkn population in veg.
crop below threshold level
2. Fallowing and summer ploughing
Use of field tillage operations to maintain a fallow condition within an agricultural field e.g. M.
graminicola use Cyperus as alternate host
3. Flooding
Extended periods of off-season field flooding (emission of lethal chemical e.g. H2S, fatty acids &
ammonia) can promote the declination of soil population density of nematodes –due to reduction of
oxygen content of the field (asphyxiation)e.g. Root-knot nematodes and Tylenchorhynchus sp. in
vegetable crops
Demerit: Uneconomic
4. Organic manuring/ soil amendments
FYM, vermi compost, neem cake, karanj cake, jatropha cake, castor cake, saw dust etc.- act in many
ways
 While decomposition –they release cetain substances which are toxic to nematodes
 Promote population of predatory nematode and parasitic fungi
 Availability of certain nutrient element increased
 Healthy plant can sustain more ppn population (carrying capacity)
60
5. Antagonistic crop/Enemy plant- Marigold, mustard, sesame, cabbage, onion, garlic
6. Trap crop e.g. Crotolaria juncea can trap M. incognita
7. Propagation through healthy planting material
Radopholus similis- Kerala to MP spread through infected rhizome
Tylenchulus semipenetrans, Pratylenchuscoffeae,
Globoderarostochiensis (infected saplings/ soil adhering tuber)
Earcockle / tundu disease of wheat can be eliminated, if gall free seeds are sown.
8. Removal and destruction of infested plants
9. Time of sowing - Wheat plants can escape Molya disease ( Heterodera avenae ) infestation if sown
fortnight earlier than usual sowing time.
B. Physical methods
1. Soil solarization- Can be exploited in tropical climate, where summer temp. becomes high (40-
42OC).
Summer ploughing can expose the nematodes to hot sun and cause desiccation leading to reduction of
nematode population, e.g. rkn and CCN
Plastic Mulching- transparent plastic of 25 micron thickness, soil should be moist
How it works?
 Augment soil temperature to lethal level
 Retention of soil moisture leads to the release of toxic subs. lethal to PPN
Useful for small area like nursery bed to raise nematode free seedling
2. Hot water treatment
O
Nematode Planting Stock Time (min) Temp ( C)
T. semipenetrans Citrus rooted 10 46.7
cuttings 25 45.0
R. similis Rhizome of banana 10-25 50.0-55.0
suckers
A. besseyi Rice seed 10 52.0
A. tritici Wheat seed 10 54.0
Meloidogyne sp. Sweet potato 65 46.7
Potato tuber 120 46.0
C. Biological Control
• Several kinds of parasite, predator and pathogens of PPN exist in agricultural field
and biological interactions continue in the nature.
• Few successful examples of introduced control (difficult to introduce organisms into
soil environment)
• Natural biological control agents present in most soils
1. Predators-
a. Predacious nematode- Mononchus papillatus feeding on Heterodera schachtii (sugar beet

cyst nematode) juveniles and males. Other genera of predacious nematode include
Anatonchus, Mylonchulus, Pristonchus, Iotonchus, Diplogaster, Labronema vulvapapillatus
etc.
61
• Constitute a large portion of the biomass in some soils
• They (Monochida, Dorylaimida, Diplogasteroidea) can be readily cultured on plates
Demerits:
• Inconsistent demonstration of plant parasitic nematode control

• Lack host specificity and thus will eat themselves
b. Predacious fungi- Some fungi belonging to the order Moniliales and Phycomycetes
predate on nematode, e.g. Arthrobotrys oligospora, Monacrosporium ellipsospora,
Dactylaria brochopaga etc.
• Predaceous fungi types
• Adhesive Networks
• Adhesive Knobs
• Nonconstricting Rings
• Constricting Rings
• Adhesive Conidia
c. Tardigardes-
d. Turbellarians (Flat worm)-
e. Collembola (apterygote insect)-
f. Mite-Pergalumna wilmanni is reported to feed on Tylenchorhynchus martini
g. Enchytraeids-
 Pot Worms, known as enchytraeids, are very small, little white worms that can reach
densities of 250,000 individuals/sq. m.
 Populations are more in acid soils.
 Feed on bacteria, fungi, nematode, organic matter and feces.
 They ingest small mineral particles and help in mixing organic matter into the mineral
soil.
 Have no enzymes for digesting polysaccharides and thus do not digest the organic
matter they ingest.
 Harmless to humans, worms and plants.
 Enchytraeids are used as fish food.
 Enchytraeids predate on nematodes.
 Pot Worms, known as enchytraeids, are very small, little white worms that can reach
densities of 250,000 individuals/sq. m.
 Populations are more in acid soils.
 Feed on bacteria, fungi, nematode, organic matter and feces.
 They ingest small mineral particles and help in mixing organic matter into the mineral
soil.
 Have no enzymes for digesting polysaccharides and thus do not digest the organic
matter they ingest.
 Harmless to humans, worms and plants.
 Enchytraeids are used as fish food.
 Enchytraeids predate on nematodes.
h. Protozoa-A large amoeboid organism Theratromyxa weberi preyed upon Globodera
rostochiensis.Theratromyxa weberi, an amoeba, was isolated and cultured in the laboratory to
62
determine the physical conditions that influence its predation on plant parasitic nematodes. In
greenhouse tests, the amoeba was a poor biological control agent of the root-knot nematode,
Meloidogyne incognita.
2. Parasite
a. Fungi- Paecilomyces lilacinus and Pochonia chlamydosporia canparasitize the eggs
of Meloidogyne spp. They are ovoparasite. Trichoderma viride, T. harzianum, and
Beuvaria bassiana are effective parasite of juveniles and females of root-knot
nematode.
b. VAM fungi like Glomus mossae and Glomus fasiculatum are also effective parasite of
soil borne nematodes.
Commercial biological control agents developed from fungal antagonists
Fungal species Type of fungi Product Name Country
Arthrobotrys superba Predacious Royal 300 France
Arthrobotrys irregular Predacious Royal 350 France
Paecilomyces lilacinus Egg-parasitic, Biocon, Bionematon, The Philippines, India
producing antibiotics Yorker
Paecilomyces lilacinus Egg-parasitic, Soybean root bio- China
producing antibiotics protectant
producing antibiotics
Myrothecium Producing antibiotics DiTera USA
verrucaria
Fungi - Nemout Saudi Arab
b. Bacterial Parasites of Nematodes: Pasteuria penetrans, Pseudomonas fluorescens,
Bacillus thuringiensis, B. firmus etc.
c. Protozoa
d. Rickettsias
e. Viruses
D. Resistant variety:
Crop Resistant Variety Nematode

Tomato Hisar Lalit, Hisar N-1, Hisar N-2, Sl-120, Meloidogyne spp.
Karnataka Hybrid, Mangala
Chilli Pusa Jawala, Suryamukhi, NP-46A, Viswa Meloidogyne spp.
Sundari
Okra Kanthi Local Green, Baishali Badhu Meloidogyne spp.
Brinjal Giant of Banaras, Black Beauty, Gola, HOE-101 Meloidogyne spp.
Potato Kufri Swarna (Multiple gene resistance)- S. Globodera pallida
vernei G. rostochiensis
Kufri Thenmalai (Single gene resistance)- S.
63
tuberosum ssp andigena
Nematicides are broadly grouped into two categories

1) Fumigants -halogenated hydrocarbon
2) Non-fumigants-organophosphate, carbamate, avermectin etc.
1) Fumigants-They are generally highly volatile compound and when applied in the soil turn
into gaseous phase. The vapours diffuse through pore spaces and cause toxicity to nematodes.
e.g. DD- mixture of 1,2-dichloropropane, 1,3-dicholoropropene
EDB-ethylene dibromide
MB-methyl bromide
DBCP-dibromo chloropropane
Fumigants are highly volatile compounds and phytotoxic, so it necessitates pre-plant
application. They may need special applicators and plastic covers to prevent the escape of
vapours into the atmosphere. Fumigants are banned for application in agricultural field of our
country.
Why fumigants are avoided?
 Doses are very high
 Difficult to inject (need special applicator)
 Carcinogenic and cause ground water contamination
 Problematic for large scale application and costly
2) Non-fumigants-
Organophosphate
Chemical name Commercial brand name
/Trade name
Ethoprof Mocap Not available in India
Fensulfothion/ Fensulphothion Dasanit Not available in India
Phenamiphos/Fenamiphos Nemacur Not available in India
Phorate Thimet Available; banned in Kerala
Thionazin Nemaphos Not available in India
Cadusaphos Rugby Not available in India
Carbamate
Aldicarb Temik Banned in India
Carbofuran Furadan, Tatafuran, Carbomain Available; banned in Kerala
Carbosulfan Marshal, Ayudh, Atank, Electra Available
Metham Sodium Vapam
Oxamyl Vydate
Avermectin
Abamectin Vertimec Available but has no label claim
Others
Cartap hydrochloride Caldan, Kritap, Bicon Available but has no label claim
Fluensulfone Nimitz Available in India
Fluopyram Velum Prime Available in India
Clothianidin -- Available but has no label claim
Advantage of non-fumigant nematicides over fumigants-
64
 Effective at low dosages
 Easy to handle and apply
 Less phytotoxic and suited for application in standing crop
 Systemic and slow release in action
 Relatively less volatile
 Effective against insect pests also
 Waiting period/half life is lesser as compared to fumigants
Economizing nematicide use/ eco-friendly application method of nematicides-
 Bare root dip treatment of seedling in systemic nematicides (carbofuran, carbosulfan,
oxamyl). It is very much effective against root-knot nematodes of transplanted
vegetable crops e.g. brinjal, tomato, chilli etc.
 Seed soaking and seed dressing or seed coating with nematicides are very much
effective in bold seeded crops.
 Nursery bed treatment for vegetable crops and rice with granular nematicide (e.g
carbofuran 3G @2 kg a.i./ha) is very much effective against root-knot nematodes.
 Row and spot or pit application in widely spaced crops.
F) Regulatory methods-
Legal enforcement of quarantine regulations has been employed in case of several nematode
diseases.
Quarantine-Rules and regulations which are enforced to prevent the entry of infected plant
material and contaminated soil into a state or country is called quarantine.
Quarantine
Domestic International
Quarantine Quarantine
Domestic Quarantine - Restriction prevailed between state within a country or one area to
another area within the state
e.g. on Globodera rostochiensis &G. pallida against import of potato seed tuber from Tamil
Nadu Nilgiri Region; Radopholus similis against import of banana sucker from Kerala and
MP
International Quarantine - Restriction prevailed between two countries e.g. Ditylenchus
dipsaci, Heterodera glycines, Heterodera schachtii and Aphelenchoides besseyi.
65

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Non-Insect Pests of Crops and their Management (EC-364)

Synopsis of Class Lectures-2023

By convention ‘Nematology’ refers to only Plant Nematology or Agricultural Nematology or

The history of Nematology perhaps evolved with the development of mankind.

 Charak recognized 20 different organisms as krimis in his Samhita. There were

 Ebers’ Papyrus during 1553-1550 BC indicated that Ascaris lumbricoides and

 More definite references of roundworms and threadworms can be found in Greek

o 1656 AD-Borellus discovered first free-living nematode ‘vineager eels’ (=Turbatrix

o 1743- A Catholic clergyman, Turbeville Needham discovered Anguina tritici -a

o 1891- Ritzema Bos discovered the foliar nematode, Aphelenchoides fragariae on

o 1892- Liebscher reported pea cyst nematode, Heterodera goettingiana on pea.

o 1913-14 Kuhn observed cyst-forming nematode attacking potatoes in Germany, but

 1941- Discovery of potato cyst nematode, Globodera rostochiensis in Nassau County,

 1943, 1945- Discovery of DD (dichloropropane-dichloropropene) mixture by Carter

 1951- Christie and Perry demonstrated the pathogenic potential of several

 1953- Discovery of burrowing nematode, Radopholus similis as the causal agent of

 1956- First ever independent Journal on PPN, Nematologica started in Europe.

 1958- First report of transmission of viruses by nematodes (Xiphinema index

 1971- Bursaphelenchus xylophilus (vectored by insects) found to be responsible for

 His book ‘Contribution to a Science of Nematology’ published in 1913 is the most

 description of many minute sensory organs of nematode, like amphids, phasmids,

 discovery of many nematode species

Development of Nematology in India

 1936- ‘White tip’ disease of rice caused by Aphelenchoides besseyi reported by

 1959-61- Siddiqi reported PPN from UP including citrus nematode, Tylenchulus

 1961- F G W Jones reported Golden Nematode of potato (Globodera rostochiensis)

 1966- Division of Nematology established at the IARI, New Delhi.

 1971-Indian Journal of Nematology (IJN) commenced publication.

 1977- AICRP on Nematode Pests of Cropping Systems started functioning.

 Association with other disease causing agents-

Table 1: Crop losses due to important plant parasitic nematodes in India

Crop Nematode species Loss (%)

Banana Radopholus similis 12

India :21068.73 million rupees in 24 different crops (Jain et al.,2007)

Classification of Plant Nematodes based on Feeding Habits

Nematodes may be grouped by feeding habit as:

• Endoparasitic– entire body inside the root

• Ectoparasitic– entire body outside the root

• Semi-endoparasitic- part of body inside root

By movement when feeding, they are called:

• Sedentary – mostly immobile during their life

• Migratory – mobile for all their life.

List of Economically Important Plant Parasitic Nematodes:

Common Name Scientific Name

A. Below ground feeders or soil inhabiting nematodes-

Above Ground/Aerial Symptom Below Ground Symptom

a) Seed Gall e.g. Anguina spp. on cereals and grasses

5. Leaf Lesions e.g. Aphelenchoides ritzemabosi onchrysanthemum

Nematode diseases vectored by insect. Red ring nematode of coconut Bursaphelenchus

Reference book for further reading-

Radopholus male (left) and female (right)

Fig. Plant Feeding Nematodes

Set-off /offset Continuous head (Helicotylenchus)

Cylindrical Filiform Elongate conoid Digitate

Pseudocoelom/ body cavity

 Nematodes lack special organs for respiration and circulation.

Fig. Cross section of nematode through mid-body region

Smooth Striated Annulated

Cuticular scales (left) and spines (right)