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Plant Nematodes
of Agricultural
Importance
A Colour Handbook

John Bridge
Tropical Plant Nematology Advisor, Emeritus Fellow
CAB International UK Centre, Egham, Surrey, UK

James L. Starr
Professor, Department of Plant Pathology and Microbiology
Texas A&M University, College Station, Texas, USA

MANSON
PUBLISHING
Copyright © 2007 Manson Publishing Ltd

ISBN-10: 1-84076-063-X
ISBN-13: 978-1-84076-063-7

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted
in any form or by any means without the written permission of the copyright holder or in accordance with
the provisions of the Copyright Act 1956 (as amended), or under the terms of any licence permitting limited
copying issued by the Copyright Licensing Agency, 33–34 Alfred Place, London WC1E 7DP, UK.

Any person who does any unauthorized act in relation to this publication may be liable to criminal
prosecution and civil claims for damages.

A CIP catalogue record for this book is available from the British Library.

For full details of all Manson Publishing Ltd titles please write to:
Manson Publishing Ltd, 73 Corringham Road, London NW11 7DL, UK.
Tel: +44(0)20 8905 5150
Fax: +44(0)20 8201 9233
Website: www.mansonpublishing.com

Commissioning editor: Jill Northcott

Project manager: Ruth Maxwell/Paul Bennett
Copy-editor: Ruth Maxwell
Cover and layout design: Cathy Martin, Presspack Computing Ltd
Book layout: DiacriTech, Chennai, India
Colour reproduction: Tenon & Polert Colour Scanning Ltd, Hong Kong
Printed by: Grafos SA, Barcelona, Spain

Plant Protection Handbooks Series

Alford: Pests of Fruit Crops – A Colour Handbook
Alford: Pests of Ornamental Trees, Shrubs and Flowers – A Colour Atlas
Biddle & Cattlin: Pests and Diseases of Peas and Beans – A Colour Handbook
Blancard: Cucurbit Diseases – A Colour Atlas
Blancard: Tomato Diseases – A Colour Atlas
Blancard: Diseases of Lettuce and Related Salad Crops – A Colour Atlas
Fletcher & Gaze: Mushroom Pest and Disease Control – A Colour Handbook
Helyer et al: Biological Control in Plant Protection – A Colour Handbook
Koike et al: Vegetable Diseases – A Colour Handbook
Murray et al: Diseases of Small Grain Cereal Crops – A Colour Handbook
Wale et al: Pests & Diseases of Potatoes – A Colour Handbook
Williams: Weed Seedlings – A Colour Atlas
 Contents
Preface 4 Chapter 7 Tree, Plantation, and Cash Crops 97
Introduction 98
Chapter 1 Plant Nematode Biology and Banana and plantain: Musa spp. 98
Parasitism 5 Black pepper: Piper nigrum 106
Introduction 6 Citrus crops 108
Migratory ectoparasites 8 Coconut: Cocos nucifera;
Migratory endoparasites 11 Oil palm: Elaeis guineensis 111
Sedentary endoparasites 14 Cotton: Gossypium spp. 114
Tobacco: Nicotiana tabacum 120
Chapter 2 Grain Legumes 19 Coffee: Coffeae spp. 122
Introduction 20 Sugarcane: Saccharum officinarum 125
Soybean: Glycine max 20 Pineapple: Ananas comosus 128
Peanut (groundnut): Arachis hypogeae 27 Deciduous fruit and nut crops 130
Other beans and peas 32
Chapter 8 Collection, Extraction, and
Chapter 3 Vegetables 39 Preservation of Nematodes for Diagnosis 135
Vegetable crops 40 Introduction 136
Collecting samples 136
Chapter 4 Flowers 45 Care of samples after collection 139
Flower crops 46 Extraction of nematodes from soil and
plant samples 140
Chapter 5 Cereals 51 Staining nematodes in plant tissues 143
Rice: Oryza sativa 52 Nematode identification 144
Maize: Zea mays L. 61
Wheat: Triticum aestrivum 64 Bibliography 145

Chapter 6 Root and Tuber Crops 69 Glossary 146

Introduction 70
Potato: Solanum tuberosum 70 Index 149
Sweet potato: Ipomoea batatas 77
Yams: Dioscorea spp. 79
Cassava: Manihot esculenta 84
Taro: Colocasia esculenta 86
Ginger: Zingiber officinale 88
Carrot: Daucus carota 90
Sugar beet: Beta vulgaris 92
4

Preface
Soil and plant nematodes are one of the most nematodes. To be certain of the association between
numerous groups of organisms occurring in the soil. particular nematodes, the organisms have to be
They are microscopic animals and, with a few extracted from the soil, roots, or other plant material
exceptions, are not visible to the naked eye. The and identified microscopically.
majority of the soil-borne nematodes are not pests of This book is written to help people working with
crops and feed on other organisms, particularly plants to have an improved understanding of plant
bacteria and fungi. Those that are parasitic on crop nematode pests and enable them to give better
plants can be very damaging and, because of their informed diagnoses, where possible, of the damage
microscopic size, associating them with crop damage caused by the very wide range of nematodes that are
is therefore mainly dependent on determining the known to parasitize and injure plants. The colour
symptoms of their effects on plants or plant growth. photographs provide an extensive range both of
Some of the parasitic nematodes do produce symptoms of nematode injury and of nematodes
characteristic and recognizable symptoms of damage themselves, observed in the field or microscopically
but many of them only produce nonspecific in plant tissues, throughout the world. There is an
symptoms. The damage and the symptoms caused introductory chapter on biology and parasitism.
can be visible above-ground; however, apart from Parasitic nematodes are discussed under the crops
poor growth and yield of the plants, the specific that they are known to attack. Crop chapters are
nematode-induced damage symptoms can often only divided into grain legumes, vegetables, flowers,
be seen in the below-ground plant organs, mainly the cereals, root and tuber crops, and tree, plantation,
roots, rhizomes, bulbs, corms, and tubers. and cash crops. Concise information is provided for
When the above-ground symptoms are the result the nematodes in these crop sections on their
of nematodes feeding on roots below ground, these distribution, symptoms and diagnosis, management,
symptoms are often similar to those seen when there and identification. A final chapter outlines the
are water or nutrient deficiencies in the soil. These so- common methods used in nematology.
called poor soils or ‘tired soils’ are often a result of a A glossary of nematological terms and a biblio-
build-up of large populations of parasitic nematodes graphy of the most important nematology texts and
in the soil. The main above-ground symptoms of papers have also been produced to help the reader.
nematode root damage are poor or stunted growth, In the compilation of this book, we are grateful to
reduced foliage, twig/branch dieback, chlorosis or our nematological colleagues for their advice,
yellowing of leaves, poor fruit or seed production support, and encouragement. We also wish to thank
and, in extreme cases, wilting and early senescence or those who have generously allowed us to use their
death of plants. The below-ground symptoms are photographs; these persons are acknowledged
related to the health of the root systems and other specifically in the legends of the photographs. Lastly
structures which can be reduced in number, suffer we thank Monica and Marylou, our long-suffering
from necrosis and rot, or have abnormal growths wives, for their patience and support of our efforts.
such as swellings or galling. Generally, it is necessary
to examine roots and other plant tissues to establish John Bridge
a connection between damage symptoms and James L. Starr
 5

CHAPTER 1

Plant Nematode
Biology and
Parasitism
• INTRODUCTION • SEDENTARY ENDOPARASITES
• MIGRATORY ECTOPARASITES Meloidogyne spp.; Nacobbus spp.;
Belonolaimus spp.; Xiphinema, Longidorus, Globodera and Heterodera spp.;
and Paralongidorus spp.; Anguina, Rotylenchulus spp.; Tylenchulus sp.
Aphelenchoides, and Ditylenchus spp.
• MIGRATORY ENDOPARASITES
Pratylenchus spp.; Radopholus spp.;
Hoplolaimus spp.; Ditylenchus spp.
 6 P LANT N EMATODE B IOLOGY AND PARASITISM

Introduction
Plant parasitic nematodes are principally aquatic 50 eggs/female to those that may produce more
animals requiring free moisture for activity; they than 1000 eggs/female.
inhabit the moisture films surrounding soil particles All crop plants are susceptible to at least one
and the moist environment of plant tissues. Nearly nematode species. Thus the potential exists for
all plant parasites spend a portion of their life cycle nematode parasitism in all climates on any crop.
in the soil. Most nematodes are adapted to the sub- The degree of damage caused by nematodes to the
tropical to tropical climates, but some are adapted to crop in any given field is closely related to the
the cooler climates of the more northerly and nematode population density, especially for annual
southerly latitudes or to higher elevations. Any crops. Precise data are lacking on the distribution
climate that supports higher plants will also support of most agriculturally important nematodes, but
a population of plant parasitic nematodes that are the distribution of the cyst nematodes on potato in
similarly adapted to that climate. Like most Europe and on soybean in the central USA, and of
invertebrate organisms, their level of activity is root knot and reniform nematodes on cotton in the
closely linked to the environmental conditions, southern USA is well documented. In each of these
especially temperature. cases, there are regions in which more than 50% of
Most nematodes and the problems they cause are the fields are infested with the problem nematode
typically associated with coarsely textured soil with species. In many other fields, even though
relative large pore spaces. Soils with sand contents potentially damaging species are present, their
of greater than 60% fulfill these conditions, as do numbers and reproductive potential in that
soils with high concentrations of organic matter or environment is insufficient to cause measurable
with low sand concentrations but with a high yield losses. With perennial crops, even low initial
degree of aggregation of silt and clay such that pore numbers of nematodes under conducive conditions
spaces are increased. However, substantial can increase sufficiently to cause substantial yield
nematode damage has been observed in nearly all suppression.
soils types. Although most nematodes are root parasites,
The nematode life cycle is relatively simple, there are nematodes adapted to parasitism in nearly
consisting of the egg, four juvenile stages, and the all plant tissues and organs. Plant parasites can be
adults. The length of the life cycle varies greatly conveniently classified based on their mode of
among the different genera, ranging from a few parasitism (Table 1). The symptoms of nematode
days to nearly 1 year under optimal environmental damage vary greatly and may be quite indistinct.
conditions and a favourable host. Nematodes It has been documented that with some crops a
reproduce both sexually and by various asexual, suppression of yield occurs prior to the expression of
parthenogenetic mechanisms. Males are common diagnostic symptoms.
in species reproducing sexually, but are generally This text provides general information on the
rare or unknown in species reproducing partheno- biology and parasitic habits of the most agriculturally
genetically. Reproductive potential also varies important genera of plant parasitic nematodes and
greatly with genera, with some producing less than specific information on the most important species.
I NTRODUCTION
 I NTRODUCTION 7

TABLE 1 Parasitic habits and examples of nematode genera

1. Ectoparasites: generally the nematodes remain on the surface of the plant tissues, feeding by inserting the stylet
into cells that are within reach
• Foliar ectoparasites: ectoparasites feeding generally on epidermal plant cells of young leaves, stems, and flower
primordia often enclosed by other foliage (Anguina, Aphelenchoides, Ditylenchus spp.)
• Root ectoparasites:
•• Ectoparasites with short stylets feeding mainly on outer root cells and root hairs (Tylenchorhynchus,
Trichodorus, Paratrichodorus, some Helicotylenchus spp.)
•• Ectoparasites with long stylets that can be inserted deep into root tissues normally at the growing tip (some
can become relatively immobile) (Belonolaimus, Cacopaurus, Criconemoides, Dolichodorus,
Hemicriconemoides, Hemicycliophora, Longidorus, Paralongidorus, Paratylenchus, Xiphinema spp.)

2. Migratory endoparasites: all stages of the nematodes can completely penetrate the plant tissues, remaining mobile
and vermiform and feeding as they move through tissues; they often migrate between the soil and roots
• Foliar endoparasites: endoparasites in stems, leaves, flower primordia, or seeds (Aphelenchoides,
Bursaphelenchus [Rhadinaphelenchus] cocophilus, Bursaphelenchus xylophilus, Ditylenchus angustus,
Ditylenchus dipsaci)
• Below-ground endoparasites: all stages of endoparasites are found throughout different tissues in roots, corms,
bulbs, tubers, and seeds (peanuts) (Aphasmatylenchus, Ditylenchus [some], Helicotylenchus [some],
Hirschmanniella, Hoplolaimus, Pratylenchoides, Pratylenchus, Radopholus, Rotylenchus [some], Scutellonema spp.)

3. Sedentary endoparasites: immature female or juvenile nematodes completely enter the plant tissues, develop a
permanent feeding site, become immobile, and swell into obese bodies. Expansion of plant tissues (galling) can
occur around nematodes (Achlysiella, Globodera, Heterodera, Meloidogyne, Nacobbus, Punctodera spp.)

4. Semi-endoparasites: immature female or juvenile nematodes only partially penetrate the roots, leaving the posterior
half to two-thirds of the body projecting into the soil. Nematodes become immobile in a fixed feeding site and the
projecting posterior of the body becomes enlarged. (Some migratory nematodes can also be found in a semi-
endoparasitic position on the roots.) (Rotylenchulus, Sphaeronema, Trophotylenchulus, Tylenchulus spp.)

I NTRODUCTION
 8 P LANT N EMATODE B IOLOGY AND PARASITISM

Migratory ectoparasites
This group includes a large number of genera and species. Many ectoparasitic species are only
rarely associated with crop damage. Among the migratory ectoparasites most noted as
aggressive pathogens (those that typically cause extensive damage to the host) of several crop
species are the sting nematode (Belonolaimus longicaudatus), the sheath nematode
(Hemicycliophora arenaria), the dagger nematodes (Xiphinema spp.), the needle nematodes
(Longidorus spp., Paralongidorus spp.), and the stubby root nematodes (Paratrichodorus and
Trichodorus spp., 7). Various stunt nematodes (Tylenchorhynchus (1), Quinisulcius, and
Merlinius spp.) and ring nematode (Criconemoides spp.) (2) are less commonly associated with
substantial yield losses but can be economically important on selected crops. Some
Helicotylenchus (3) and Rotylenchus species feed as ectoparasites. All life stages of the migratory
ectoparasites, except the egg, are parasitic. Root feeding nematodes with short stylets feed on the
epidermal and outer cortical cells often at the root tips (4), those with longer stylets feed deeper
in the cortex. Eggs are laid singly in the soil and the life cycle is usually straightforward, with
each stage having to feed on a suitable host in order to develop to the next stage (5).
Reproduction is by both sexual and asexual systems.

1 2

1 Females and egg of Tylenchorhynchus annulatus. 2 Criconemoides female.

3 4
ECTOPARASITES
M IGRATORY

3 Helicotylenchus indicus female. 4 Tylenchorhynchus maximus feeding as an ectoparasite

on a root tip.
 M IGRATORY E CTOPARASITES 9

(J1)
J4
J2 J3 Mature Eggs in soil
female

Feed on
surface
of root All stages feed
Eggs laid in soil

5 Life cycle of the ectoparasite Longidorus spp.

Belonolaimus spp. 6

The sting nematode, Belonolaimus longicaudatus, is

a migratory ectoparasite with a long feeding stylet
(6, 25) that allows it to pierce roots and feed on the
cortical tissues several cell layers deep in the roots.
The sting nematode does not induce any specialized
feeding sites, but causes extensive cellular damage
and necrosis of cortical tissues. Distribution of this
species is strongly influenced by soil texture and
temperatures. The nematode is found only in the soil 6 Anterior end of a female Belonolaimus longicaudatus.
in warm climates and requires very sandy soils
SPP.

(typically with greater than 85% sand content).

During the warmest parts of the year, population
B ELONOLAIMUS

densities may decline in the upper soil profile, with

most nematodes being found 15–30 cm deep. The
reproductive potential of the sting nematode is
moderate and rarely do population densities exceed
100 nematodes/100 cm3 soil.
 10 P LANT N EMATODE B IOLOGY AND PARASITISM

Xiphinema, Longidorus, and Anguina, Aphelenchoides, and

Paralongidorus spp. Ditylenchus spp.

The dagger (Xiphinema) and especially the needle Some species of the genera Anguina (90),
nematodes (Longidorus and Paralongidorus spp.) Aphelenchoides (72), and Ditylenchus are migratory
(137) are among the largest plant parasitic nematodes. ectoparasites of the foliage, feeding on stems, leaves,
They typically feed at the root tip, with several and inflorescences; they can also be found
nematodes often feeding on a single root tip (5). These endoparasitically in some plant structures. Anguina
feeding activities often result in a swollen root tip, tritici is the infamous ear cockle nematode of wheat;
such that the symptoms may be confused with root the two most destructive and widely distributed
galls caused by the root knot nematodes. Similar to species of Aphelenchoides are A. besseyi, the rice white
the root knot and other sedentary nematodes, host tip nematode, and A. ritzemabosi, the chrysanthemum
cells fed upon by Xiphinema and Longidorus often leaf nematode. A. ritzemabosi invades leaves through
become enlarged, multinucleate, and with a more the stomata to feed endoparasitically on mesophyll
dense cytoplasm than adjacent cells. The long stylets cells; otherwise they are ectoparasites on the surface of
of these nematodes permit them to feed on cells in the leaves and buds. Eggs are laid in the leaf tissues and the
interior of the root apex while the nematode body is life cycle is very short taking only 10 days. All stages
exterior to the root. These nematodes have relatively are vermiform and migratory. Ditylenchus species
long life cycles, depending on environmental often have both ectoparasitic and endoparasitic
conditions and susceptibility of the host plant. Females feeding phases on their hosts (page 13).
of Xiphinema spp. range from 1 to over 10 mm in Aphelenchoides are relatively long thin nematodes
length. They have life cycles typically requiring 1 year similar in many respects to Ditylenchus. The
from egg to egg for X. americanum and up to 3 years A. ritzemabosi female is around 1 mm in length with
for X. diversicaudatum. Several species of these genera a posterior vulva and long tapering tail. Stylets are
are also important because they vector plant viruses, small, 12 μm long, with tiny basal knobs. Males are
such as the grapevine fanleaf virus by X. index and present and morphologically similar to the female
raspberry ringspot virus by L. elongatus. This group apart from sexual organs.
of nematodes encompasses a large number of different
species, the taxonomy of which is dynamic, and
SPP.

accurate identification of species is difficult. They

parasitize a large number of vegetable crops, field
PARALONGIDORUS

crops, and perennial fruit crops. Many species are

important in the cooler climates of northern Europe
and North America.

7
AND
X IPHINEMA , L ONGIDORUS ,

7 Head of Trichodorus sp. showing curved stylet.

 M IGRATORY E NDOPARASITES 11

Migratory endoparasites
Many of the important parasites of plants are migratory endoparasites that remain vermiform
throughout their lives. The species well documented as causing crop damage mainly
belong to the genera Pratylenchus, Radopholus, Hirschmanniella (124, 125),
Hoplolaimus, Scutellonema (113), Aphasmatylenchus, and Ditylenchus, and
also some Helicotylenchus (3) and Rotylenchus.

Pratylenchus spp. 8

The lesion nematodes, Pratylenchus spp. (8, 82, 150,

154), feed primarily on cortical tissues of smaller,
nonsuberized roots. They may also infect tubers,
peanut pods, and other below-ground organs.
Migration through host tissues and feeding activities
results in destruction of host cells with formation of
necrotic lesions. All life stages, except the J1, are
parasitic and can be found in host roots and
surrounding soil (9). Eggs are deposited singly in
infected tissues and the soil. Sexual reproduction is
common for some species such as P. penetrans but Eggs laid 9
in soil
rare for others such as P. brachyurus. The optimum
temperature for development is 26°C. As an
endoparasite, population densities of Pratylenchus Eggs
laid in
spp. are typically much greater in plant roots than in roots
J2
the surrounding soil, and this is a common feature of J3
these migratory endoparasites; therefore, it is J4
essential that samples for diagnosis include roots Some
nematodes
from symptomatic plants. Soil samples alone may Mature leave roots
fail to indicate the presence of damaging population female

densities of these nematodes. Feeding

causes
necrosis
SPP .
P RATYLENCHUS

8 Pratylenchus zeae female.

All stages
invade roots
and feed in
9 Life cycle of migratory endoparasites Radopholus and
cortex
Pratylenchus spp.
 12 P LANT N EMATODE B IOLOGY AND PARASITISM

Radopholus spp. 10

There are several Radopholus species damaging

crops and they are similar to the lesion nematodes in
behaviour and appearance (10, 143). The burrowing
nematode, Radopholus similis (143) is economically
the most important. Burrowing nematodes are
aggressive pathogens of such crops as anthurium,
citrus (spreading decline disease), banana and
plantain (toppling disease), and black pepper
(yellows disease). Distinct races (banana race and
citrus race) based on host preferences are recognized 10 Radopholus citri female.
within this species. The nematode has a relatively
large host range, numbering several hundred plant
species. Like the lesion nematodes, the burrowing
nematode’s feeding activities result in the
development of necrotic lesions in the infected host
tissues (9). All life stages are parasitic and the
nematode reproduces sexually. Eggs are laid within
infected host tissues (corms, roots, and tubers) and
embryonic development is completed in a few days.
The entire life cycle can be completed in 3 weeks
under optimal conditions. The nematode is restricted
to tropical climates.

Hoplolaimus spp. 11

The lance nematodes Hoplolaimus columbus,

H. pararobustus, H. seinhorsti, and H. galeatus
(11, 24) are relatively large migratory parasites that
feed both endo- and ectoparasitically. All life stages
are infective and can be found within the plant roots,
feeding on the cortical cells and in the surrounding
soil. Feeding activities typically result in cellular
necrosis (43). Eggs are deposited singly in roots and
the soil. Males of H. columbus are very rare in nature,
whereas males are common for H. galeatus. Diagnosis 11 Hoplolaimus seinhorsti females.
can be made from either root or soil samples.
SPP.
R ADOPHOLUS
 M IGRATORY ENDOPARASITES 13

Ditylenchus spp. region and is distinguished by its larger size.

Reproduction is by cross-fertilization and males are
Ditylenchus species can have an ectoparasitic phase common. Eggs are deposited singly in infected tissues.
but are largely migratory endoparasites that infect the In the later stages of disease, the nematodes aggregate
bulbs (tulips, narcissus, onion, and garlic), stems into a mass often referred to as ‘nematode wool’ (41).
(lucerne and clover species), inflorescences (rice), and With slow drying, the J4 stage can enter an anhy-
pods and seeds (peanut) of host plants. All life stages drobiotic state and can survive for years in this
(J2 through to adults) are infective, but the J4 stage is condition. The survival potential and large host range
the most common primary inoculum as this stage has enables long-term persistence of populations in
the greatest potential for survival of adverse infested fields.
conditions. The nematodes feed primarily on All stages of D. dipsaci are vermiform. The female
parenchyma tissues, secreting large amounts of cell is 1 mm in length with a posterior ovary and a
wall degrading enzymes that cause much tissue tapering, pointed tail. It has a flattened head and a
destruction. Although many Ditylenchus species are small, delicate stylet about 10 μm long. The male is
able to utilize a variety of fungi as hosts, one of the similar to the female, apart from sexual structures.
most important plant pest species of the genus, D. angustus is a tropical species and an important
D. dipsaci, appears to be a strict obligate parasite of pest of rice in south and southeast Asia causing ufra
plants. The host range of D. dipsaci includes such disease (see Chapter 5). It feeds ectoparasitically on
diverse crops as oat, lucerne (alfalfa), clovers, tulip, stems but also feeds within the inflorescence.
narcissus, onion, garlic, broadbean, strawberry, and D. africanus is an economically important pest of
many weed species. Many populations of D. dipsaci peanuts in southern Africa, feeding on underground
exhibit distinct host preferences. A ‘giant race’ exists growing pegs, pods, and seeds.
that is particularly important in the Mediterranean

SPP.
D ITYLENCHUS
 14 P LANT N EMATODE B IOLOGY AND PARASITISM

Sedentary endoparasites
Meloidogyne spp. 12

Root knot nematodes (Meloidogyne spp.) are all

sedentary endoparasites. The wide host ranges of the Eggs
four most common species (M. arenaria, M. hapla, M.
incognita, and M. javanica) combined includes nearly
2nd stage
all crop plants. The total number of described species juveniles (J2s)
is approaching 100, thus many of the populations
formerly believed to be one of these four common
species are now recognized as a distinct new species. J2 juveniles
Many species have more restricted or less well invading
root Eggsac
described host ranges. Only the second juvenile stage
(J2) (158) is infective, with root tips being the primary
infection court (12). The J2 migrate through cortical Swollen
tissues and establish a permanent feeding site in the mature
females
region of differentiation of vascular tissues. In
addition to roots, the nematodes may also infect other
below-ground tissues such as potato tubers, peanut
pegs and pods, and rhizomes, corms, or bulbs of
various crops. The feeding site of each nematode
consists of several host cells that are induced by the
nematode to become enlarged, specialized feeding
Swelling and developing
cells called giant cells. The giant cells are plant transfer 3rd and 4th stages in root
cells with a dense cytoplasm; they are multinucleate
and have elevated rates of metabolism. The J2 12 Life cycle of the sedentary endoparasite
develops from the typical vermiform-shaped, pre- Meloidogyne spp.
parasitic stage to a swollen, sausage-shaped advanced
J2, and moults to the third stage (J3), fourth stage (J4),
and finally the adult female which undergoes a period
of rapid growth to achieve the typical rounded, pear 13
shape (12, 13, 157). Egg production begins shortly
after the final moult, and up to 1000 eggs can be
produced by each female and deposited into a
gelatinous matrix. This egg mass usually ruptures
through the root cortex and is visible on the root
SPP.

surface. Egg masses are initially light brown in colour

but become progressively darker with age.
M ELOIDOGYNE

Meloidogyne spp. are called root knot nematodes

because of the characteristic galling of host roots at
each infection site. The galls are due primarily to
hyperplasia and hypertrophy of cortical tissues 13 Stained, swollen Meloidogyne incognita female in root.
 S EDENTARY E NDOPARASITES 15

surrounding the nematode and the giant cells. Root Nacobbus spp.
galling begins within a few days of infection due to a
proliferation of cortical cells surrounding the The false root knot nematodes, Nacobbus spp., so-
developing nematode. For crop species with succulent called because the root galls caused by these
roots (e.g. tomato, melons, and cucurbits), small root nematodes are very similar to those caused by
galls are visible within a few days of infection. Galls Meloidogyne spp., have a sedentary and swollen,
may be indistinct early in the season and for crops mature female stage (14, 102). Other stages,
with fibrous or woody roots such as cotton, including the immature female stage, behave as
groundnut, and cereal grain crops, but on other crops migratory endoparasites moving between soil and
they can be very large especially in the latter part of roots (101). Therefore, all stages can be found in
the growing season. Galls on leguminous crops can the root, and all stages except the mature, swollen
be distinguished from Rhizobium nodules, in that the female, can be found in the soil. Males are
galls are an integral part of the root and cannot be common. The swollen female is elongated rather
removed without severing the root. Rhizobium than round and is often mistaken for a developing
nodules project from the root surface and can be Meloidogyne female in roots. Eggs are laid in a
easily removed without severing the roots. Nodules gelatinous matrix extruded from the root. The
can be infected by the nematodes, with galls and nematode is found in cooler areas, for example at
nodules being indistinguishable. high altitudes in the Andes, although can be found
Under favourable conditions most Meloidogyne as concomitant populations with Meloidogyne,
spp. can complete their life cycles in 4–5 weeks, with making their identification or diagnosis very
the production of several hundred eggs by each difficult. Galls are small and rounded, often
female; some species such as M. graminicola have a occurring at regular intervals along the root giving
shorter life cycle of less than 3 weeks. Soil population the appearance of rosary beads, and hence the
densities can increase more than 100-fold during the common name of rosary bead nematode or
growing season. The optimal temperature for ‘rosario’. Several Nacobbus species are referred to
development of M. arenaria and M. javanica is 28°C, in the literature and the genus taxonomically is in a
whereas the optimum for M. hapla is 24°C. state of flux; the species most commonly reported
Additionally, most populations of M. hapla can is N. aberrans.
survive at least brief periods of soil temperatures of
less than 0°C, whereas M. arenaria and M. javanica
do not survive freezing temperatures.
Because most stages of development of
Meloidogyne spp. (juvenile J3 and J4 stages and the
adult females) are present only in host tissues, the J2
and adult males are the only stages of the nematode
present in the soil. Beginning about 6 weeks after 14

planting, egg masses containing hundreds of eggs

each can be easily observed on roots with a hand lens
or a dissecting microscope. Juveniles can be
extracted from the roots, and vermiform adult males
can be extracted from soil or infected roots late in the
growing season when population densities are high.
Rounded to pear-shaped adult females can be
SPP.

dissected from infected roots. Correct identification

N ACOBBUS

of the Meloidogyne species is critical for effective

management based on crop rotation or host
resistance. 14 Swollen Nacobbus aberrans female.
 16 P LANT N EMATODE B IOLOGY AND PARASITISM

Globodera and Heterodera spp. 15

The cyst nematodes, Globodera and Heterodera

spp., are sedentary endoparasites that have a definite
survival stage, the cyst, which is the hardened dead
female body containing eggs (12, 15, 23, 73, 74, 93,
136). With many of the cyst nematodes, root
exudates from host plants are needed to stimulate
the eggs to hatch and emerge from the cysts. In
Heterodera spp., eggs are also laid in a gelatinous egg
mass. The second-stage juveniles of both Globodera
and Heterodera (16A, B, 17) which emerge from the 15 Heterodera sacchari: brown cysts and white female.
eggs are the infective stages and invade roots near the
root tip, inducing a feeding site of syncytial nurse be extracted from the soil by flotation. It is difficult
cells. Nematodes moult through the third and fourth to identify the species, or even the genus, without
stages and develop to maturity in the root. The examination of the mature cysts. The potato cyst
females swell and burst through the epidermal layers nematodes, G. rostochiensis and G. pallida, are
of the root and become visible on the root surface, found in cool temperate climates and generally only
first as round, white females and then brown cysts have one life cycle per growing season. Most of the
(17, 22, 33, 73, 135). No root galling occurs. Males Heterodera species are found in the warm temperate
and J2 can be found in the soil, as can the cysts when to tropical regions and have a number of generations
they eventually break away from the root. Cysts can per cropping season.

16A 17

Cysts free
in soil

J2 juveniles
in soil

Cyst
attached
J2s invade to root
root Mature
female

16B
SPP.

3rd and
H ETERODERA

4th stages
develop in
root
AND
G LOBODERA

16 Second-stage juveniles of Heterodera sacchari. 17 Life cycle of cyst nematodes Globodera and
A: Anterior body; B: tail and posterior body. Heterodera spp.
 S EDENTARY E NDOPARASITES 17

Rotylenchulus spp. Eggs

18

The reniform nematodes, Rotylenchulus spp., are

sedentary semi-endoparasites. The life cycle of
reniform nematodes is unique in that the nematode
2nd, 3rd and 4th stage
proceeds from the freshly hatched J2 stage through juveniles in soil Swollen
four moults to the immature female in the soil without mature
female
feeding (18). This development occurs over 7–10 days
at temperatures of 25–30°C. The vermiform,
immature female is the only infective stage,
penetrating the cortex of roots of less than 2 mm
diameter with the anterior third of its body. The head
Eggsac
of the nematode is then adjacent to the vascular tissue,
where the nematode induces formation of
multinucleate syncytia that are the parasite’s
permanent feeding site. These syncytia are
functionally similar to the giant cells induced by
Immature
Meloidogyne spp. The posterior two-thirds of the female semi-
nematode’s body swells outside the root (19) and endoparasitic
and swelling
assumes the characteristic ‘renal’ shape within a few in root
days of root penetration (184). Reproduction occurs
following fertilization of the developing female by
males, which are vermiform and apparently develop
without feeding. Eggs are deposited into a gelatinous
matrix which surrounds the female’s body (185), 18 Life cycle of the sedentary, semi-endoparasite
with each egg mass containing 50–100 eggs. The Rotylenchulus spp.
nematode can complete several generations in a
growing season and population densities at crop
maturity often exceed 10,000 nematodes/500 cm3
soil. Both soil and root samples should be examined 19

for diagnosis. All life stages, except mature swollen

females, can be extracted from infested soil.
R. reniformis has a host range of hundreds of plant
species. These nematodes are also able to survive for
several months in very dry soils by a process of
anhydrobiosis, and the bodies of these dormant forms
typically assume tightly coiled positions in the dry soil.
SPP.

19 Rotylenchulus reniformis female, semi-endoparasitic on

R OTYLENCHULUS

root and beginning to swell.

 18 P LANT N EMATODE B IOLOGY AND PARASITISM

Tylenchulus sp. shape on the surface of the root (20, 165). Eggs are
laid in a gelatinous matrix around female bodies
The citrus nematode, Tylenchulus semipenetrans, is (166) and can cover parts of the root in a
also a sedentary semi-endoparasite, almost continuous layer. These matrices have soil particles
exclusively of citrus and related plants. The J2 adhering to them which obscures the nematodes on
partially penetrate the root and develop through the the roots. The nematodes are extremely small and
different stages to mature females, with the neck of are only normally visible when viewed using a
the nematode deep inside the root, and a series of microscope and when stained in a suitable stain (see
feeding or nurse cells are initiated around the head Chapter 8). Only J2 and males can be found in the
of the nematode. The remaining body of the soil, the juveniles often in very large populations;
nematode swells into a characteristic asymmetrical males are very difficult to identify.

20

20 Tylenchulus semipenetrans females protruding from citrus root.

SPP.
T YLENCHULUS
 19

CHAPTER 2

Grain Legumes

• INTRODUCTION • OTHER BEANS AND PEAS

• SOYBEAN (Glycine max) Heterodera goettingiana Liebscher;
Heterodera glycines; Meloidogyne incognita, Meloidogyne spp.; Ditylenchus dipsaci
M. arenaria, M. javanica, and M. hapla; (Kühn) Filipjev; Hoplolaimus seinhorsti
Pratylenchus agilis, P. alleni, P. brachyurus, Luc
P. penetrans, and P. sefaensis; Rotylenchulus
reniformis; Hoplolaimus columbus;
Belonolaimus longicaudatus
• PEANUT (GROUNDNUT) (Arachis
hypogeae)
Meloidogyne arenaria, M. hapla, and
M. javanica; Pratylenchus brachyurus
Godfrey; Aphelenchoides arachidis Bos;
Belonolaimus longicaudatus Rau;
Aphasmatylenchus straturatus Germani
 20 G RAIN L EGUMES

Introduction
There are more than 20 important legume species grown as major food crops in different
regions of the world. In addition to the widely grown soybean, Glycine max, and peanut
or groundnut, Arachis hypogeae, there are many other genera and species of legumes
commonly referred to as beans or peas that are vital sources of calories, proteins and oils.
Nematodes are economically important pests of all the legume crops.

Soybean Glycine max

Soybean is an important crop in North and South America, South Africa, and Asia. The most
serious nematode parasites of soybean are the cyst nematode, Heterodera glycines, and the root
knot nematodes, especially Meloidogyne incognita, M. arenaria, and M. javanica. These
nematodes are aggressive parasites able to cause significant yield losses and are widely
distributed in most areas of soybean production. Nematodes of lesser importance, due
primarily to their more restricted distributions, include the reniform nematode,
Rotylenchulus reniformis, the Columbia Lance nematode, Hoplolaimus columbus, several
lesion nematodes, Pratylenchus spp., and the sting nematode, Belonolaimus longicaudatus.

Heterodera glycines Carolina in 1954 and is now present from Florida in

the south to Minnesota in the north, and from New
Distribution Jersey in the east to Texas in the west. Most
The soybean cyst nematode, H. glycines, is widely infestations in the new world are believed to have
distributed. The first official report of the occurred with the importation of soybean seed or
GLYCINES

occurrence of the nematode was from Japan in soil from Asia (as a source of rhizobia inocula)
1915. It has subsequently been reported from during the late 19th and early 20th centuries. There
China, Korea, the former Soviet Union, and Taiwan is evidence, however, to suggest that the nematode
H ETERODERA

in Asia. In the new world, H. glycines is widely may also be native to North America as a parasite of
distributed in the USA, and is reported from weed hosts. The nematode occurs across a wide
Ontario province in Canada, from Argentina, range of temperate to sub-tropical environments
Brazil, and Columbia. In the USA, the nematode and in a wide range of soil types.
was first reported from a single location in North
 S OYBEAN 21

Symptoms and diagnosis cyst nematode contributes to its major economic

Yield suppression of 10–15% without obvious importance. Losses in the single state of Iowa were
expression of other symptoms is often the first sign estimated at more than 5.8 ⫻ 109 kg valued at
that a field is infested with this nematode. Higher $13 million in 1997. Similar losses are estimated
levels of infection by the soybean cyst nematode throughout soybean production areas in the USA
typically cause a distinct chlorosis of the foliage, and wherever the nematode occurs on the crop.
followed by stunting of the plant (21). Affected Although damage thresholds vary with soil type,
portions of the field are often first observed as initial population densities of 100 eggs/500 cm3 soil
elliptical areas of symptomatic plants. The chlorosis may suppress yields by as much as 30%. Greatest
is prominent because the soybean cyst nematode damage is typically observed in more coarsely
inhibits the development and activity of N2-fixation textured, sandy soils.
nodules. In the most severe infestations, plant stands
are reduced. Although the nematode does not cause Management
even discrete root galls, some swelling of the roots at Management of H. glycines is achieved primarily
the site of infection may be observed. White swollen though the combined use of resistant soybean
females and mature brown cysts can be observed on cultivars and crop rotations. Several hundred
the root surface, especially with the aid of a hand lens soybean cultivars are available in the USA that have
(22). The presence of white females and/or mature resistance to one or more races of the soybean cyst
brown cysts on the roots of soybean is sufficient for nematode. However, there are several races for
diagnosis of the soybean cyst nematode. Cysts, which resistance is not yet available in agronomically
vermiform males and J2 individuals can also be acceptable cultivars. The former system of describing
extracted from soil samples throughout the year. races is being discontinued because it was inadequate
for describing variation in virulence within a
Economic importance population of H. glycines. A new HG Type scheme
H. glycines is an aggressive parasite that causes yield has been developed that more accurately describes
losses amounting to millions of dollars in the USA the ability of a given population to parasitize the
annually. The widespread distribution of the soybean available sources of resistance. As with other plant

21 22

GLYCINES

21 Damage to soybean caused by Heterodera glycines in 22 White and light brown coloured cysts of Heterodera
H ETERODERA

the USA. (Courtesy of D.P. Schmitt.) glycines exposed on infected soybean root. (Courtesy of
R.D. Riggs.)
 22 G RAIN L EGUMES

pathogens, continued use of a single source of Meloidogyne incognita, M. arenaria,

resistance to H. glycines will usually result in a shift M. javanica, and M. hapla
in the virulence characteristics of that population,
such that that source of resistance is no longer Distribution
effective. Thus it is critical that use of resistance be All four of the root knot nematodes, Meloidogyne
limited and used in combination with other species, are distributed world-wide in different
management tactics, especially crop rotation. agroecosystems. In warm temperate and tropical
Although H. glycines parasitizes numerous weed regions, M. incognita is the most commonly detected
species, it is only able to parasitize successfully species of root knot nematodes, followed by
relatively few crop species. Adzuki bean (Vigna M. javanica. M. arenaria is the least common species.
angularis) and common bean (Phaseolus vulgaris) These three species are rarely found where the mean
are also good hosts of the nematode in addition temperature for the coldest month is less than 1°C . M.
to soybean. Therefore, rotation of soybean with a hapla is more common in the cooler temperate zones,
wide range of crops, especially graminaceous, and is relatively rare in climates where the mean
solanaceous, and cruciferous crops, is effective in temperature of the warmest month exceeds 27°C. All
managing nematode populations. The most effective Meloidogyne species are favored by coarsely textured,
control is achieved with 2 years of either a nonhost sandy soils and are rarely found in finely textured soils
crop and/or an effective resistant soybean cultivar. with high percentages of silt and clay.
Good weed control to eliminate potential alternate
hosts is an important consideration. Symptoms and diagnosis
Symptoms caused by all Meloidogyne species on
Identification soybean are similar, with plants stunted and
Second-stage juveniles range in size from 0.375 mm chlorotic at high levels of infection. Root galling is
to 0.540 mm, with a prominent stylet (22–25 ␮m), a moderate to severe, varying with level of infection.
HAPLA

sclerotized lip region, a distinct oesophageal Generally, galling by M. arenaria and M. javanica is
intestinal overlap, and an acute tail terminus. The more prominent than that caused by M. incognita.
M.

cysts are lemon-shaped with protruding neck and Galls caused by M. hapla are small and are
JAVANICA , AND

vulva cone, and are of variable size (340–920 ␮m characterized by the presence of adventitious roots.
length ⫻ 200–560 ␮m width) (23). Chlorosis of the foliage is less pronounced with
Meloidogyne species than it is with H. glycines.
Root galls are the most diagnostic symptom of
these nematodes. Galls can be distinguished from
M.

Rhizobium root nodules in that nodules are easily

broken off the root and may have a pink (active) to
ARENARIA ,

greenish (inactive) interior. Nematode galls are an

23 integral portion of the root and cannot be removed
without severing the root. Nodules may be galled in
M.

later stages of crop development.

INCOGNITA ,

Economic importance
The widespread distribution of root knot nematodes
is a major reason for their economic importance.
M. javanica is generally considered to be more
M ELOIDOGYNE

aggressive than M. incognita which, in turn, causes

more damage than M. hapla. Aggressiveness of
M. arenaria varies with host race. Race 1, which also
23 Mature cyst of Heterodera glycines opened to expose parasitizes peanut, is weakly aggressive on soybean
eggs carried in the cyst. (Courtesy of R.D. Riggs.) whereas race 2 of M. arenaria (that does not
 S OYBEAN 23

parasitize peanut) is highly aggressive towards temperate climates, whereas P. brachyurus is most
soybean. Initial population densities of 150 common in warmer climates. Other species,
M. incognita/500 cm3 soil can result in a 10% yield including P. alleni (north-central USA) and
suppression. Greater losses would be expected from P. sefaensis (west Africa) have more restricted
M. javanica at similar initial population densities, distributions, and may be limited to the particular
whereas M. hapla would cause less yield suppression region from which they were initially described.
at this population density.
Symptoms and diagnosis
Management The primary symptom of damage is the formation of
Management options for root knot nematodes on elliptical, necrotic lesions on feeder roots, with a
soybean are dependent upon which species is suppression of root and shoot growth with high levels
present, and are made more difficult if the field is of infection (several hundred nematodes per g of root
infested with multiple species. Several crop rotation fresh weight). The amount of necrosis varies with
systems have been identified for the different different soybean cultivars and Pratylenchus spp.
Meloidogyne species. Cotton is an effective rotation combinations. Diagnosis based on root symptoms is
crop with soybean for all species except those difficult, and usually requires extraction of nematodes
populations of M. incognita that are parasitic on from root and soil samples. Collection of root samples
cotton (host races 3 and 4). Peanut is an effective is especially important because most of the nematodes
rotation crop for management of M. incognita, most are inside the roots during the growth of the crop.
populations of M. javanica, and M. arenaria race 2. Diagnosis based only on soil samples may greatly

SEFAENSIS
Maize and other cereals are effective in suppressing underestimate the nematode population density.
populations of M. hapla. Numerous soybean
cultivars are available that have various levels of Economic importance

P.
resistance to M. incognita, and a few cultivars have Lesion nematodes are considered of minor

PENETRANS , AND
resistance to M. arenaria and M. javanica. Many of importance to soybean production on a world-wide
these cultivars also are resistant to one or more races basis, but can cause substantial yield losses in
of the soybean cyst nematode. severely infested fields.Yield losses of 15–20% can
be expected in sandy and sandy loam soils.
Identification
Second-stage juveniles are readily extracted from Management

P.
infested soil. They have a slender shape (158) with a A few soybean cultivars with moderate levels of

BRACHYURUS ,
pointed tail terminus and a delicate stylet (10–12 ␮m resistance to P. brachyurus have been identified.
in length for all four species) and the oesophagus Crop rotations can be effective, but require accurate
has a distinct overlap of the intestine. Mature, pear- identification of the Pratylenchus spp. Maize can be
shaped females are present only in infected roots and an effective rotation crop for management of
are variable in size and may be as much as 1 mm in P. brachyurus but not P. penetrans. Cotton is a
P.
diameter (157). relatively poor host of both P. brachyurus and
ALLENI ,

P. penetrans.

Pratylenchus agilis, P. alleni,

P.

Identification
AGILIS ,

P. brachyurus, P. penetrans, and Pratylenchus spp. are relatively short nematodes

P. sefaensis (0.39–0.80 mm in length), with a short robust stylet
(14–22 ␮m ). Species can be distinguished based on
P RATYLENCHUS

Distribution variation in the tail shape (conical to crenate or

The different species of lesion nematodes, irregular), the number of lip annules, and the position
Pratylenchus spp., are widely distributed especially of the vulva (in the posterior third of the body).
in coarsely textured, sandy soils. P. penetrans is
among the most commonly found species in
 24 G RAIN L EGUMES

Rotylenchulus reniformis effective than having the soil wetted periodically to

induce nematode activity.
Distribution
The reniform nematode, Rotylenchulus reniformis, Identification
is widely distributed in tropical and warm temperate Immature, vermiform females of R. reniformis
regions. These nematodes can be found on soybean extracted from the soil are 0.34–0.42 mm in length,
in soils of a wide range of textural classes and are whereas mature, swollen females (reniform ⫽ kidney-
commonly found in more finely textured, silty soils. shaped) from roots are 0.38–0.52 mm in length with
a vulva located in the posterior third of the body
Symptoms and diagnosis (184). The tail is conical, and the oesophagus
Reniform nematodes tend to be more uniformly overlaps the intestine, typically laterally. Males of R.
distributed over a field than most nematode species reniformis are 0.38–0.42 mm in length; the bursa
and, hence, the field may lack discrete, irregular areas envelopes the tail with readily observed spicules.
of symptomatic plants. The first symptom of damage
may be a suppression of yield, followed by slight to
severe stunting and chlorosis. Reniform nematodes
can predispose soybean to seedling diseases when
initial nematode population densities are relatively
high (greater than 1000 nematodes/500 cm3 soil at
planting). Roots of infected plants may be stunted, but
lack other diagnostic symptoms. Signs of the nematode
include the presence of immature and mature females
protruding from the surface of feeder roots and egg
masses covered with soil particles on the root surface
(184, 185 Chapter 7). When heavily infected roots are
washed gently with water, the soil particles adhering to
egg masses will give the roots a dirty appearance. Soil
population densities frequently exceed 10,000
individuals/500 cm3 soil at crop maturity.

Economic importance
Detectable yield suppression in a range of soil types
can be observed when initial population densities
of R. reniformis exceed 100 nematodes/100 cm3
soil.Yield suppression of more than 50% has been
observed when initial nematode populations exceed
1000 nematodes/100 cm3 soil.
RENIFORMIS

Management
Management has been primarily through use of
granular and fumigant nematicide applications.
Soybean cultivars with moderate levels of resistance
R OTYLENCHULUS

are available. Although R. reniformis has an

extensive host range, rotation with nonhosts such as
sorghum or peanut provide effective management.
Because of the ability of the nematode to survive
desiccation, fallowing during a dry season is less
 S OYBEAN 25

Hoplolaimus columbus prominent knobs that resemble a tulip flower (24).

The tail is blunt and the vulva is positioned near the
Distribution mid-body.
The lance nematode, H. columbus, has a limited
distribution, being found on soybean primarily in
sandy coastal plain soils of the southeastern USA,
and has also been reported from Egypt and 24

Pakistan.

Symptoms and diagnosis

H. columbus feeds both as an ectoparasite and an
endoparasite on the cortical tissues of soybean roots,
causing large necrotic lesions. Affected plants are
stunted with reduced root development, and typically
occur in irregular clusters within the field. Symptoms
are similar to those caused by Belonolaimus
longicaudatus. Extraction of nematodes from root
and soil samples is required for diagnosis.
24 Head of Hoplolaimus columbus from soybean.
Economic importance
Yield suppression in the range of 10–38% have been
documented in the southeastern USA, when
nematode population densities at planting were in
the range of 90–200 individuals/100 cm3 soil. Total
yield losses are much lower than for cyst or root knot
nematodes because of the limited distribution of
H. columbus.

Management
Although many common field crops are susceptible
to H. columbus, rotations with non- or poor hosts
such as peanut, sweet potato, tomato, pepper,
and other vegetables are effective. Additionally,
agronomic practices that reduce other stresses on
soybean and promote greater root development
(e.g. soil fertility, irrigation, and sub-soiling to
disrupt hardpan layers in the soil) are effective in
reducing yield losses due to this nematode.
Management with granular or fumigant nematicides
COLUMBUS

is rarely justified because of the relatively low profit

margins associated with this crop.

Identification
H OPLOLAIMUS

H. columbus is a relatively large robust nematode,

with mature females being 1.25–1.6 mm in length,
with a heavily sclerotized and prominent lip region.
They have a large, robust stylet (40–48 ␮m) with
 26 G RAIN L EGUMES

Belonolaimus longicaudatus Economic importance

Although B. longicaudatus is an aggressive
Distribution nematode that can cause substantial plant damage
The sting nematode, Belonolaimus longicaudatus, is where it occurs, it is of overall limited economic
distributed throughout the southeastern USA, and importance because of its restricted distribution. As
was recently reported from California on turf few as 10 nematodes/500 cm3 soil at planting are
grasses, and from the Bahamas, Bermuda, Puerto sufficient to cause moderate damage to soybean.
Rico, Costa Rica, and Mexico. Most surveys indicate
that less than 2% of the soybean fields in the Management
southeastern USA are infested with this nematode. Crop rotation with watermelon or tobacco is
effective in suppressing population densities of this
Symptoms and diagnosis nematode and increasing soybean yields. Although
B. longicaudatus induces symptoms similar to those some populations of B. longicaudatus are parasitic
of Hoplolaimus columbus, including primarily on peanut, peanut can be an effective rotation crop
stunted, chlorotic shoots with poorly developed in those fields with populations of the nematode that
root systems. Sunken necrotic lesions in the cortical do not reproduce on that crop. Control of
tissues may be visible. Affected plants are typically B. longicaudatus by nematicide application is
distributed in irregular clusters throughout the justified only in the most extreme cases.
field, with boundaries between healthy and stunted
plants usually well marked (25). Seedling death Identification
may occur at high initial population densities The female B. longicaudatus is a relatively long
(greater than 100 nematodes/100 cm3 soil), (2–3 mm in length) and slender nematode, with a
resulting in reduced plant populations. Population distinct offset lip region and a long, thin, flexible
densities rarely exceed 500 individuals/100 cm3 stylet (100–140 ␮m ) with rounded knobs (6, 132).
soil. Extraction of nematodes from soil is required The vulva is located near mid-body with two
for diagnosis. outstretched ovaries, and the female tail is
comparatively long with a bluntly rounded terminus.

25
LONGICAUDATUS
B ELONOLAIMUS

25 Damage to soybean caused by Belonolaimus longicaudatus in the USA.

 P EANUT ( GROUNDNUT ) 27

Peanut (groundnut) Arachis hypogeae

Peanut (groundnut) is a major food and cash crop in American, African, and Asian countries
and has numerous nematode parasites. It is unusual amongst the legumes in producing
the mature seed pods and seeds below ground. Among the most common and
economically important nematodes of peanut are the root knot nematodes
(Meloidogyne spp.), the lesion nematode (Pratylenchus brachyurus), the testa nematode
(Aphelenchoides arachidis), the sting nematode (Belonolaimus longicaudatus), the pod
nematode (Ditylenchus africanus), and Aphasmatylenchus straturatus.

typically becoming darkly pigmented (27). Galling of

Meloidogyne arenaria, M. hapla, and peanut roots by M. hapla is distinguished from that
M. javanica of the other two species in that there is prolific

Distribution
The root knot nematodes, Meloidogyne arenaria, 26

M. hapla, and M. javanica are all distributed world-

wide. All populations of M. hapla are parasitic on
peanut and are found parasitizing peanut in the
cooler, northern peanut productions areas of Virginia
and Oklahoma in the USA, the Punjab state in India,
and the Shandong province in China. M. arenaria is
the most common root knot species attacking peanut
in the more southern regions of the USA, in central

JAVANICA
and southern India, and in southern China. Only race
1 of M. arenaria is parasitic on peanut. Most
populations of M. javanica in the USA are not 26 Patches of stunted and chlorotic peanut plants infested

M.
parasitic on peanut, but some populations that are with Meloidogyne javanica in Egypt.
parasitic on peanut have been reported from a few

HAPLA , AND
fields in Georgia and Texas. M. javanica populations
parasitic on peanut are common in Egypt and India. 27

Symptoms and diagnosis

M.

All three Meloidogyne species cause typical

ARENARIA ,

symptoms of nematode damage on the above-ground

portions of peanut, including a clustered arrangement
of stunted, chlorotic plants with premature
senescence (26). M. arenaria and M. javanica cause
M ELOIDOGYNE

similar symptoms with respect to root and pod

galling. Root galling is indistinct early in the growing
season, but becomes more pronounced as the crop
matures. Both M. arenaria and M. javanica can cause 27 Galling and darkening of peanut pods caused by
severe galling of the pegs and pods, with the pod galls Meloidogyne javanica.
 28 G RAIN L EGUMES

adventitious root development from the galls; are effective rotation crops for control of
M. hapla rarely forms galls on the pods. Care must be M. arenaria. In general, crop rotations are more
taken to avoid misdiagnosis of Rhizobium root effective if the non- or poor-host crop is grown for
nodules as nematode-induced galls. Soil population 2 years rather a single year of the alternative crop.
densities can be near the detection limits during early No cultivar with resistance to M. hapla is available,
portions of the growing season. In the middle to the but the first cultivar (cv ‘COAN’) with resistance to
later portion of the growing season, most of the M. arenaria and M. javanica was released in the USA
nematodes are developing inside the galled roots or in 1999 (28). Biological control of M. arenaria with
are present as eggs in the egg masses on the root the obligate bacterial parasite Pasteuria penetrans
surface. Numbers of J2 stages in the soil are the has also been effective, but is not yet commercially
highest in the later half of the growing season and available.
may exceed 1000 J2/100 cm3 soil.
Identification
Economic importance The second-stage juvenile (J2) can be identified to
Yield losses of more than 50% in heavily infested genus based on its acute tail, overall length of
fields have been documented for M. arenaria and 0.36–0.56 mm, with M. hapla being typically shorter
M. javanica. Losses due to M. hapla are usually less than M. javanica, which in turn is shorter than
and rarely exceed 25% of the yield potential. Several M. arenaria. The juveniles are also characterized by
studies have documented a pre-plant damage having a slender body, a distinct oesophagus overlap
threshold population density for M. arenaria and of the intestine, and an acute tail terminus. The
M. javanica of 1–10 juveniles/500 cm3 soil. The stylets are delicate with a length of 10–12 ␮m. The
damage threshold population density for M. hapla is mature females dissected from roots have a distinctly
greater at 16 juveniles/500 cm3 soil. Because of their rounded to pear-shaped bodies. The perineal
widespread distribution and high frequency of patterns (cuticular markings surrounding the anus
occurrence, the root knot nematodes are considered and vulva) are helpful in the identification of species.
to be very important pathogens of peanut. In some
production areas of the USA, nearly 30% of the
fields are infested with one or more of these
nematodes.
JAVANICA

Management
Root knot nematodes on peanut have been typically
M.

controlled through the use of nematicides in the

USA. The fumigant 1,3-dichloropropene is the most 28
HAPLA , AND

effect nematicide, followed by the nonfumigant

nematicides aldicarb and phenamiphos. A common
practice is to use either the fumigant 2–3 weeks prior
to planting or a nonfumigant at planting, followed
M.

by an application of aldicarb about 6 weeks after

ARENARIA ,

planting as the pegs are beginning to form. Several

crop rotations systems have been shown to be
effective for controlling root knot nematodes. Most
grass and grain species are poor hosts of M. hapla
M ELOIDOGYNE

and, therefore, are good rotation crops for this

species. Several crops, including cotton, bahiagrass
(Paspalum notatium), velvet bean (Mucuna pruriens 28 Comparison of the growth of susceptible and resistant
var. utilis), partridge pea (Chamaecrista fasciculata), peanuts growing in a field infested with Meloidogyne
and American jointvetch (Aeschynomene americana), arenaria.
 P EANUT ( GROUNDNUT ) 29

Pratylenchus brachyurus Godfrey 29

Distribution
Lesion nematodes (Pratylenchus spp.) are
cosmopolitan, and P. brachyurus is common in the
warm temperate and tropical regions. This species
has been specifically reported attacking peanuts
in Australia, Egypt, the USA, and Zimbabwe.
Additionally, P. coffeae has been reported in
association with peanut in India.

Symptoms and diagnosis

Diagnostic foliar symptoms are rare, but may include
stunting and chlorosis if the level of infection is 29 Lesions on peanut pods caused by Pratylenchus
extremely high. Root symptoms are the presence of brachyurus.
distinct necrotic lesions, often elliptical in shape,
ranging from a few millimetres to several centimetres
in length. Root lesions are most evident on smaller
diameter, feeder roots. P. brachyurus typically also for the following season. Both fumigant and
cause distinct necrotic lesions on the pods (29). These nonfumigant nematicides are effective in reducing
lesions are characterized by diffuse rather than sharply crop losses due to P. brachyurus, but are too costly for
delineated margins. Eggs are deposited singly in most peanut production systems. P. brachyurus has a
infected tissues and surrounding soil. The nematode wide host range including numerous weed and crop
may survive for 24 months in infected pods at room species, making development of effective crop rotation
temperature. Diagnosis requires identification of the systems difficult. The long-term survival of the
nematode in addition to observation of symptoms. nematode in infected pods further complicates
Because of the endoparasitic nature of these development of effective rotation systems. No cultivar
nematodes, detection of the nematode is best with resistance is available.
accomplished by extraction of root and pod samples.
Identification
Economic importance P. brachyurus can be identified based on its distinct
More than 90% of the total nematode population lip region with two annules, a robust stylet
are typically associated with the host tissues during (17–22 μm), and an overall length of 0.75 mm.
the cropping season and, immediately after harvest, Adult females, as with all Pratylenchus, have a single
affecting the yield quantity; P. brachyurus also ovary, with the vulva located posteriorly (80% of
G ODFREY
affects marketable value through the effects on pod body length). The tail terminus is typically blunt. The
appearances. Incidences of pod rot caused by soil- oesophagus overlaps the intestine ventrally. Males
borne fungi, especially Pythium spp. and are rarely observed.
BRACHYURUS

Rhizoctonia solani, are increased by concomitant

infection by the nematode. Aflotoxin contamination
of pods due to colonization of nematode-infected Aphelenchoides arachidis Bos
pods by Aspergillus flavus is also increased.
Distribution
P RATYLENCHUS

Management The peanut testa nematode, Aphelenchoides

Losses due to P. brachyurus can be reduced in severely arachidis, is known currently only from northern
infested fields by a 6-week fallow period during the Nigeria; however, there is concern that, because it is
early summer. Timely harvest with removal of infected a seed-borne pathogen, it may be spread to other
pods may also reduce nematode population densities peanut production areas.
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