Front Cover
 Title Page
 Table of Contents
 Host-parasite relationships by...
 General control considerations
 Trends in host-parasite relati...
 Appendix I. Plant parasitic nematodes...
 Appendix II. Common names of tropical...
 Back Cover
 Historic note

Group Title: Bulletin - Agricultural Experiment Stations, University of Florida ; 823 (technical)
Title: Plant parasitic nematodes associated with tropical and subtropical fruits
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00027461/00001
 Material Information
Title: Plant parasitic nematodes associated with tropical and subtropical fruits
Series Title: Bulletin Agricultural Experiment Station, University of Florida
Physical Description: 49 p. : ; 23 cm.
Language: English
Creator: McSorley, R. ( Robert )
Publisher: Agricultural Experiment Stations, Institute of Food and Agricultural Sciences, University of Florida
Place of Publication: Gainesville Fla.
Publication Date: 1981
Subject: Tropical fruit -- Diseases and pests -- Tropics   ( lcsh )
Plant nematodes -- Tropics   ( lcsh )
Plant parasites -- Tropics   ( lcsh )
Genre: government publication (state, provincial, terriorial, dependent)   ( marcgt )
bibliography   ( marcgt )
non-fiction   ( marcgt )
Bibliography: Bibliography: p. 16-26.
Statement of Responsibility: Robert McSorley.
 Record Information
Bibliographic ID: UF00027461
Volume ID: VID00001
Source Institution: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: aleph - 000401356
oclc - 09178097
notis - ACE7184

Table of Contents
    Front Cover
        Front Cover
    Title Page
        Title Page
    Table of Contents
        Table of Contents
        Page 1
    Host-parasite relationships by crop
        Page 2
        Page 3
        Page 4
        Page 5
        Page 6
        Page 7
        Page 8
        Page 9
        Page 10
        Page 11
        Page 12
    General control considerations
        Page 13
    Trends in host-parasite relationships
        Page 14
        Page 15
        Page 16
        Page 17
        Page 18
        Page 19
        Page 20
        Page 21
        Page 22
        Page 23
        Page 24
        Page 25
        Page 26
    Appendix I. Plant parasitic nematodes associated with tropical and subtropical fruits listed by host plant
        Page 27
        Page 28
        Page 29
        Page 30
        Page 31
        Page 32
        Page 33
        Page 34
        Page 35
        Page 36
        Page 37
        Page 38
        Page 39
        Page 40
        Page 41
        Page 42
        Page 43
        Page 44
        Page 45
        Page 46
    Appendix II. Common names of tropical fruits
        Page 47
        Page 48
        Page 49
    Back Cover
        Page 50
    Historic note
        Page 51
Full Text



Robert McSorley

Agricultural Experiment Stations
Institute of Food and Agricultural Sciences
University of Florida, Gainesville
F. A. Wood, Dean for Research

Bulletin 823 (technical)


Robert McSorley
Assistant Professor
University of Florida
Agricultural Research and Education Center
Homestead, Florida

Introduction ............................................

Host-Parasite Relationships by Crop ........................... ..
Aegle marmelos ..... ........... ....................
Annona spp. .............................................
Artocarpus spp .......................................

Carica papaya .............................
Diospyros spp ...........................
Ficus carica ..............................
Litchi chinensis .............................
M alpighia glabra ..........................
M angifera indica ...........................
M anilkara achras ..........................
M onstera deliciosa .........................
Olea europaea .............................
Passiflora edulis ...........................
Persea americana ..........................
Phoenix dactylifera .........................
Psidium guajava ...........................
Punica granatum ...........................
Tamarindus indica .........................

General Control Considerations ................ .

Trends in Host-Parasite Relationships .............





Bibliography .............................................. 16

Appendix 1: Plant Parasitic Nematodes
Associated with Tropical and Subtropical
Fruits by Host Plant ...................................... 27

Appendix II: Common Names of Tropical Fruits ..................... 47

Trade names of chemicals, where used, are for the purpose of providing spe-
cific information. No endorsement of products is intended, nor any criticism of
products not named. No registrations are implied herein.




Florida is one of the few areas in the United States where tropical and sub-
tropical fruits can be grown successfully, and acreage of the minor subtropical
fruits is increasing in the southern part of the state. During the 1978-79 growing
season, there were an estimated 7,700 bearing acres of avocado (Persea ameri-
cana Mill.) and 1,750 acres of mango (Mangifera indica L.) in Florida (174).
Some commercial production exists for bananas and plantains (Musa spp.), car-
ambola (Averrhoa carambola L.), papaya (Carica papaya L.), mamey sapote
(Calocarpum sapota (Jacq.) Merr.), sapodilla (Manilkara achras (Mill.) Fosb.),
lychee (Litchi chinensis Sonn.), and several others (174).
Since prevention of infestation is the most important aspect of nematode con-
trol on tropical fruits in Florida and elsewhere, awareness of the actual and po-
tential nematode problems is essential. However, reports of plant-parasitic
nematodes on tropical fruits, where available, are widely scattered throughout
the literature, and have not been assembled into a unit. It is anticipated that a
review of published literature on the subject will not only aid in making control
decisions, but will serve as a starting point for future research on these nematode
associations. This review includes not only subtropical and tropical fruits grown
in Florida, but has been expanded to include those grown in other areas as well.
Nematode problems on several of the major tropical and subtropical fruits
have been reviewed elsewhere. These include bananas and plantains (15, 169,
223), citrus and citrus relatives (30, 52, 66, 85, 154, 155), coconut (67), and
pineapple (4, 5). In many cases, host-parasite relationships are similar; for ex-
ample, kumquat (Fortunella japonica (Thunb.) Swingle) is also a host of Tylen-
chulus semipenetrans Cobb (29) and Radopholus similis (Cobb) Thorne (66).
The present work seeks to organize nematological information on the many other
species of tropical fruits. Nematode problems of temperate tree fruits have been
reviewed elsewhere (133).
Certain nematodes, such as Aphelenchus avenae Bastian, are often listed in
the literature as plant-parasitic. However, since this nematode is reported to be
a fungal feeder (152), it is inappropriate to discuss it as a plant parasite of trop-
ical fruit trees. Therefore, associations of this type have not been included. In
many cases, plant-parasitic nematodes (of higher plants) are simply reported as
associated-with.a given fruit tree _Ithoutf ny additional mention-of damage
symptoms, ecology, or control. A list of such associations by crop is provided
in Appendix I.


Aegle marmelos L. Correa Bael
Bael has been reported as a host for both Radopholus similis (69, 159) and
Tylenchulus semipenetrans (209). The life cycle of Xiphinema insigne Loos in
soil around the roots of A. marmelos has been studied in India (89). Highest
populations were observed from June to September, followed by a decline to the
lowest levels in March and April. Reproduction occurred only once per year,
during the months of May to August. This was the warmest part of the year, with
mean seasonal temperatures of 300 C.

Annona muricata L. Soursop
Annona reticulata L. Bullock's Heart, Custard Apple
Annona squamosa L. Sugar apple, Sweetsop, Custard

Caveness (26) has investigated the nematode parasites of several of the annon-
aceous fruits. Annona muricata was considered an excellent host for three dif-
ferent Helicotylenchus species, including H. cavenessi Sher. Annona squamosa
was also a good host for H. cavenessi, but A. reticulata was considered a poor
host for it. The Annona species tested were not suitable hosts for Scutellonema
clathricaudatum Whitehead, Xiphinema sp., orHeterodera sp.

Artocarpus atilis (Park.) Fosberg Breadfruit
(Syn. A. communis Forst.;
A. incisa L.f.)
Artocarpus heterophyllus Lam. Jackfruit
(Syn. A. integrifolia L.)
Little information exists about nematode problems of the breadfruit tree, al-
though Meloidogyne spp. have been reported to be associated with this crop
(72). Breadfruit is an excellent host for several species of Helicotylenchus, with
10,000-fold population increases reported after 13 months in a shadehouse test
(26). Caveness (26) rated breadfruit to be a good host for Rotylenchulus renifor-
mis, a moderate host for Scutellonema clathricaudatum, and a poor host for
Xiphinema ifacolum Luc.
Several species of plant parasitic nematodes have been found associated with
the closely-related jackfruit, Artocarpus heterophyllus (177, 183, 218).

Carica papaya L.

Papaya is a host of the reniform nematode, Rotylenchulus reniformis (26,
118). This nematode has been reported on papaya from many parts of the world,
including Hawaii (76, 116, 118), Florida (134), Cuba (42, 43), Puerto Rico (7,
8), Trinidad (202), Brazil (177), Liberia (42), Nigeria (26), India (199), Aus-
tralia (34), and Brunei (18, 199). Rotylenchulus parvus (Williams) Sher had
been identified from Kenya (42), and unidentified Rotylenchulus species have
been found associated with this crop in Thailand (218) and Florida (219).
Rotylenchulus reniformis is associated with poor growth and reduced yield in
Puerto Rico (7), and has been associated with damaged plants in Trinidad, along
with occasional tree death and toppling (202). In Brunei, plants are reported to
have been killed by a combination of R. reniformis and Phytophthora nicotianae
var. parasitica (18). The reniform nematode feeds on phloem cells in papaya
roots, and induces formation of giant cells in the phloem. The nematode may
also feed in the root cortex, or it may cause mechanical damage to cortical cells
as it moves toward the phloem. As a result of this damage, cortical cells break
down and a cavity is formed (204). Infected roots become brown and necrotic,
thus providing suitable sites for fungal attack.
Foliar applications of systemic nematicides (phenamiphos and oxamyl) have
been tested for control of R. reniformis on papaya in Puerto Rico (7). However,
the materials tested showed some phytotoxicity. Lange (116) tested several pre-
plant soil fumigants for control of R. reniformis. In one experiment, use of da-
zomet at 200 or 400 pounds active ingredient (a.i.) per acre (A), DD at 200 or
400 pounds a.i./A, or DBCP at 70 or 140 pounds a.i./A led to significant (P
= 0.05) yield increases, compared to unfumigated control plots for 15-month-
old plants. After 29 months however, yield differences among treatments were
no longer significant. In a second experiment, significant yield differences at 15
months compared to unfumigated controls were observed with dichoropropene
at 200 pounds a.i./A, DBCP at 35 or 70 pounds a.i./A, or methyl bromide (8%)
at 208 pounds a.i./A, but not with 440 pounds a.i./A of SMDC or 300 pounds
a.i./A of dazomet. Counts of R. reniformis in the soil of the fumigated plots
were reduced after two and four months compared to the untreated plots, but
after six months, numbers in the fumigated plots were comparable to or had ex-
ceeded the numbers in the untreated plots.
Root knot nematodes are a frequently reported problem of papaya. Meloido-
gyne incognita (Kofoid & White) Chitwood has been identified from Hawaii
(76), Puerto Rico (7), Brazil (41, 177), and India (22). Meloidogyne javanica
(Treub) Chitwood is reported from many areas, including Brazil (40, 41, 122),
Israel (143), Italy (82), Rhodesia (129), and Madagascar (124). Other species
associated with C. papaya are M. arenaria (Neal) Chitwood in Rhodesia (129)
and M. hapla Chitwood in Brazil (41) and Hawaii (76). Meloidogyne spp. have
also been reported on papaya in Florida (224), Trinidad (202), Brazil (41), Peru
(24), Israel (142), Thailand (218), Taiwan (78), and Oman (222).


Meloidogyne javanica has been reported to seriously damage papayas in the
field (122), and has damaged the plant in a greenhouse test as well (82). Meloi-
dogyne incognita, which, along with Rotylenchulus reniformis, is associated
with poor growth and declining yield in Puerto Rico, has been controlled to
some extent by foliar application of systemic nematicides (7). Wolfe and Lynch
(224) recommend planting in non-infested sites, use of sterilized soil in the
seedbed, heavy mulching, and optimum cultural practices for Florida papaya
production. Searching for resistance in interspecific hybrids may prove difficult,
since closely related species such as Carica quercifolia Solms and C. candamar-
censis Hook are also susceptible (20, 142). It may be that preplant soil fumi-
gants, such as those used by Lange (116) against Rotylenchulus reniformis,
would also be helpful if papayas must be planted in sites infested by root knot
Other plant-parasitic nematodes reported from papaya include Hemicrico-
nemoides mangiferae Siddiqi (172), which reproduced on papaya in an inocu-
lation study (171). In shadehouse tests, C. papaya was described as an excellent
host for four species of Helicotylenchus, a good host for Tylenchorhynchus mar-
tini Fielding, and a moderate host for aPratylenchus species (26). It was a poor
host for Hoplolaimus pararobustus (Schuurmans Stekhoven & Teunissen) Sher,
Scutellonema clathricaudatum, and Peltamigratus nigeriensis Sher. Papaya is
not a host of Radopholus similis (140, 205).
Other nematodes found associated with damaged or decaying papaya roots
have included Discomyctus longicaudatus (Imamura) Thorne (88), Seinura au-
rangabadensis Suryawanshi (210), and S. tandoni Singh (203), although these
are not reported as plant parasites here (Appendix I).

Diospyros kaki L.f. Japanese Persimmon
Diospyros lotus L. Persimmon
Diospyros virginiana L. Common Persimmon, American

The persimmons are hosts to several important plant-parasitic nematodes.
Meloidogyne spp. are reported from both Diospyros kaki andD. virginiana (72,
78), and both plants are also hosts ofRadopholus similis (14, 17, 29, 159). Cohn
and Minz (31) found Tylenchulus semipenetrans in the roots of several persim-
mon varieties which had been grafted onto American persimmon rootstock. The
affected plants were stunted and exhibited premature defoliation. Females and
egg masses of T semipenetrans have also been found in roots of D. lotus (150).

Ficus carica L. Fig
Root knot nematode infections are a frequently reported problem of fig (103,
141, 142, 201, 214). Taylor (214) demonstrated the pathogenicity of Meloido-
gyne spp. to fig by comparing seedlings grown in nematode-infested soil to

those grown in uninfested soil. By the time of transplanting (four months later),
significant height differences between the two groups occurred. All of the in-
fected seedlings later died in the field, while control plants grew normally. In a
second field experiment, inoculation of seedlings with infested soil resulted in
a high mortality compared to uninoculated controls. Meloidogyne species re-
ported on Ficus carica include M. arenaria (83, 91, 129, 143), M. incognita
(46, 50, 121, 143, 173),M. incognita acrita (72, 129, 143, 161, 208), andM.
javanica (72).
The severe damage to fig trees caused by root knot and other nematodes has
been described (103). However, figs may produce satisfactorily if growing con-
ditions are made as favorable as possible (35, 103). Mulching may also be ben-
eficial (35). Krezdorn and Adriance (103) report that figs may grow well if
planted near buildings, since soil beneath buildings contains no vegetation and
hence, few nematodes. Preplant fumigation is also recommended to allow trees
to become established before nematode populations increase (103). Populations
ofM. arenaria in soil around the roots of four-year-old fig trees were reduced by
the systemic activity of phenamiphos applied to the trunk (83). The activity of
various formulations of DBCP against M. incognita has also been investi-
gated (50).
Early tests in California revealed that of 120 fig varieties tested, all were sus-
ceptible to Meloidogyne spp. (35). More recently, resistance to M. incognita
acrita has been found in individual seedlings of the cultivars 'Hunt' and 'Ce-
leste' in Louisiana (161). Some tolerance to this nematode also occurs in three
cultivars 'Azendjar', 'Taranimt', and 'Zidi' introduced into California
(208). OtherFicus species show a wide range of susceptibility to root knot nem-
atodes (36). For this reason, it may be possible to use species showing immunity
or tolerance in developing interspecific hybrids with nematode resistance (208).
Ficus racemosa L. and F gnaphalocarpa Steudel ex Miquel have been recom-
mended as root knot resistant rootstocks forE carica (36).
The cyst nematode, Heteroderafici Kirjanova, has been reported from Ficus
carica in several parts of the world (13, 51, 193, 201). Under greenhouse con-
ditions, the F carica varieties 'Brown Turkey', 'Calimyrna', 'Kadota', and
'Mission' were shown to be susceptible, as well as the closely-related E elastica
Roxb. (193).
Ficus carica is the type host ofXiphinema index Thorne and Allen, described
from the soil of declining trees in California (217). This nematode is also widely
distributed on figs in Italy (127). Fig is one of only a few good hosts ofX. index
(162). Damage symptoms included necrosis and loss of lateral roots, angling of
main roots, and terminal swelling and necrosis, resulting in lateral root prolif-
eration. The root tip was the preferred feeding site. The feeding behavior of X.
index on E carica root tips has been described in detail (226). Initial feeding
occurred in the region of cell elongation, and was followed by gall formation and
cell modification (225, 227, 228). Developmental times for the various stages in
the life cycle ofX. index onE carica have also been investigated (160).

Damage to fig roots by "Anguillulina pratensis (DeMan) Goodey" has been
described in California (216). This nematode was probably Pratylenchus vulnus
Allen & Jensen (191). Injury ranged from root lesions to extensive root injury
and was accompanied by reduced top growth. Root damage to fig due to P mu-
sicola (Cobb) Filipjev has also been described (206).
Paratylenchus hamatus Thorne & Allen was described from the rhizosphere
of declining fig trees in California (217). Symptoms included light color of the
foliage, leaf drop, undersized or fallen fruit, and a slow decline of the entire
Martin et al. (131) report the presence of various species of nematodes in the
fruit ofFicus spp. and in the wasps which pollinate these fruit. The feeding hab-
its and effects of these nematodes are unknown.

Litchi chinensis Sonn. Litchi, Lychee
Serious nematode problems have been reported on Litchi chinensis in South
Africa. A tree decline characterized by leaf yellowing and tip burn, poor flower
and fruit set, dieback of twigs and branches, and eventual tree death, was ob-
served in Transvaal and Natal in 1969 (137). In some orchards, up to 40% of the
trees had been killed, with another 35% declining. Pathogenic fungi were not
found, but an initial survey (137) revealed high numbers (up to 2100 per 100
cubic centimeters (cm3) of soil) of aHemicriconemoides species, later identified
as H. mangiferae Siddiqi (75), and of the dagger nematode,Xiphinema brevi-
colle Lordello & da Costa (up to 1000/cm3 of soil). A more extensive survey
(138) revealed that large populations of one or both of these nematodes were
found where decline symptoms were seen. There appeared to be a direct rela-
tionship between the number of nematodes present and poor tree condition, al-
though populations dropped when the tree was almost dead. Decline symptoms
were associated with dieback of the root system, and trees replanted in decline
areas showed symptoms in seven to eight years (138).
Xiphinema brevicolle penetrated the root tissue while feeding, causing some
cell disruption (136). However, Hemicriconemoides mangiferae was observed
to penetrate deeply into the root and could completely enter the cortex (136).
Such penetration led to cell destruction, death of affected tissue, sloughing of
the root cortex, and a typical "stubby root" appearance (138). Milne et at.
(138) considered H. mangiferae to be the major nematode pathogen because of
its larger numbers and deeper penetration of the root tissue.
Treatment of infected trees with DBCP at 28 or 45 liters per hectare (L/ha)
improved tree health, while treatment with ethoprop did not (138). Since both
H. mangiferae andX. brevicolle can survive in the soil of an air layer, the use of
soil from infested groves in making air layers should be avoided (138).
Meloidogyne javanica has also been found on lychee roots in South Africa,
and caused galling, cell modification, and tissue disruption in an inoculation test
(136). Conflicting reports exist concerning the status of Radopholus similis on

L. chinensis. In an early study (54), it was found that roots of lychee trees
planted in spreading decline areas or in mixed plantings with infested citrus
were free of R. similis. In a related inoculation test, citrus became infected with
R. similis, while lychee did not (54). Yet recent reports (140, 153, 159) indicate
thatL. chinensis is a host of the citrus biotype of R. similis, but a non-host of the
banana biotype.

Malpighia glabra L. Barbados cherry, Acerola

Root knot nematodes are an important economic pest of Barbados cherry (6,
77, 117). Ayala (6) states that the plant can be almost destroyed by Meloidogyne
incognita in home gardens and commercial plantings in Puerto Rico. M. incog-
nita acrita causes severe damage in Florida, especially on sandy soils (117).
One of the best control methods is use of sterilized soil in propagation. Heavy
mulching and regular fertilization and irrigation may also alleviate the problem
to some extent. Fumigation of the planting site was also recommended (117).
Use of Malpighia suberosa L. as a root-knot tolerant rootstock for Barbados
cherry has been attempted, but the plants grafted onto M. suberosa have not
yielded well (117).
Malpighia glabra was also found to be a host of Radopholus similis (14). Ro-
tylenchulus reniformis has been reported from two related plants, M. infestis-
sima (A. Juss.) Rich. and M. punicifolia L. in Puerto Rico (8). Meloidogyne
javanica has been reported on an unidentified species of Malpighia (72).

Mangifera indica L. Mango

Hemicriconemoides mangiferae is one of the most widely reported plant-par-
asitic nematodes on mango. It has been reported from India (92, 167, 196), Pak-
istan (171, 172), Nigeria(158), and South Africa (75, 139). In an inoculation
study (171), H. mangiferae was found to be pathogenic to Mangifera indica at
a level of six nematodes per cm3 of soil. This nematode has been observed feed-
ing on mango roots, and together with Xiphinema brevicolle, is associated with
root damage and tree decline in South Africa (139). Application of DBCP or
phenamiphos to infected trees reduced populations ofH. mangiferae 12 and 24
months after treatment, but phenamiphos was not effective againstX. brevicolle
(139). Population fluctuations of H. mangiferae, Helicotylenchus erythrinae
(Zimmerman) Golden, and Hoplolaimus indicus Sher around mango roots were
significantly correlated with soil moisture at several sites in India (92). Mini-
mum soil populations occurred during the hot, dry months of April through
June. In experiments with mango seedlings grown in pots, optimum soil mois-
ture for H. mangiferae was found to be 25%, and optimum soil temperatures
were 20-250C (92).
A yield response has been obtained by using DBCP to control Hoplolaimus
columbus Sher and Xiphinema spp. in Egypt (176). Mangifera indica was rated

as an excellent host for Helicotylenchus cavenessi, H. pseudorobustus (Steiner)
Golden, Tylenchorhynchus martini, Xiphinema ebriense Luc, X. ifacolum, and
X. nigeriense Luc, but a poor host for Hemicycliophora cocophillus (Loos)
Goodey, Hoplolaimus pararobustus (Schuurmans Stekhoven & Teunissen) Sher,
and Scutellonema clathricaudatum (26). Radopholus similis has not been found
in mango roots in Florida (233), and the plant is not a host of the citrus race
(153), although it is a host of the banana race (140, 153). Pratylenchus brach-
yurus (Godfrey) Filipjev & Schuurmans Stekhoven has been reported on mango
from a number of locations throughout the world (77, 139, 179, 233), but its
effects on the plant are not fully known. Numerous Xiphinema species are also
associated with mango (Appendix I).

Manilkara achras (Mill.) Fosb. Sapodilla
(syn. Achras zapota L.)
The nematode fauna of the sapodilla in Pakistan has been investigated by
Saeed (171). Hemicriconemoides mangiferae was found to be the most frequent
plant parasite associated with sapodilla, occurring in 85% of the soil samples.
Pratylenchus spp., Helicotylenchus spp., and Xiphinema americanum were also
Plant-parasitic nematodes associated with sapodilla in Pakistan showed two
seasonal peaks (March/April and October/November), and favored a soil tem-
perature of 19-220C and a soil moisture of 12-16% (171). Horizontal and ver-
tical distribution of the plant parasites corresponded to the distribution of sapodilla
roots. Maximum numbers of H. mangiferae were found at a depth of 9 to 18
inches. Non-parasitic nematodes, such as Psilenchus hilarulus deMan, were
common near the soil surface where few roots occurred.
In inoculation studies, H. mangiferae, Helicotylenchus indicus, and Paraty-
lenchus spp. increased their populations on sapodilla. Hemicriconemoides man-
giferae was pathogenic to Manilkara achras at a rate of six nematodes per cm3
of soil. Treatment of infected trees in the field with DBCP reduced populations
of H. mangiferae, but they built up again to 85% of the original levels within 10
months (171).
The plant-parasitic nematodes associated with the sapodilla in Thailand and
the Philippines have also been reported (218).

Monstera deliciosa Liebm. Ceriman
Although this plant produces an edible fruit, it is more often encountered as
an ornamental or a houseplant. Several species of root knot nematodes have been
reported on it (38, 72). Monstera deliciosa has been damaged by Meloidogyne
arenaria when grown as an ornamental in a greenhouse in Belgium. Aldicarb
was an effective control. In addition, this plant is a host of Radopholus similis

Olea europaea L.

The nematode parasites of the olive have been reviewed by Lamberti (104).
Major nematode problems include Meloidogyne spp., Pratylenchus vulnus, and
Tylenchulus semipenetrans. The effects of several root knot nematode species on
olive were studied in a pathogenicity test (109). The varieties 'Ascolano' and
'Manzanillo' were highly susceptible to M. incognita, but resistant to M. aren-
aria and M. hapla. The varieties 'Ascolano' and 'Sevillano' were susceptible to
M. javanica, and the top weights of inoculated plants were reduced when com-
pared to uninoculated controls, but 'Manzanillo' was tolerant.
The pathogenicity ofPratylenchus vulnus to olive was demonstrated when top
weight of 'Manzanillo' olive was significantly reduced by inoculation (108).
Top weight of 'Ascolano' olive was reduced in plants inoculated with Tylen-
chulus semipenetrans (110), while 'Manzanillo' olive was somewhat tolerant
and did not show significant reductions after inoculation, despite development
of mature females on the root system. Recent work (9) indicates that 'Manza-
nillo' olive can be used as one of the differential hosts in distinguishing among
the four biotypes of T semipenetrans occurring in California. One or more ad-
ditional biotypes are present in Italy (115).
Helicotylenchus dihystera and Xiphinema elongatum Schuurmans Stekhoven
& Teunissen were associated with declining olive trees in Egypt (49), and were
found to damage plants and to reproduce on olive in a pathogenicity test. The
olive is also a suitable host forX. americanum (105). Recently, the feeding hab-
its of Gracilacusperatica Raski on olive roots has been described (84), and Hel-
icotylenchus oleae Inserra, Vovlas, and Golden has also been observed feeding
on olive roots (86).

Passiflora edulis Sims Passionfruit
A recent survey of Passiflora edulis (var. 'Flavicarpa') in Fiji (102) revealed
thatRotylenchulus reniformis was present in 16 of 19 sites sampled, and that at
11 of these sites, numbers were greater than 1000/200 cm3 of soil. In a patho-
genicity test, R. reniformis populations increased on passionfruit, and inocu-
lated plants showed a significant reduction in vine weight compared to
uninoculated controls. Thus, there is the likelihood that R. reniformis has con-
tributed to the passionfruit decline in Fiji (102). Rotylenchulus reniformis has
been found on dead seedlings in Brazil (180), and has also been reported on
P edulis in several other areas (8, 19, 77) and on P seemanni Griseb (118).
Meloidogyne spp. have been reported on P edulis in several instances (48,
142, 156, 180). Species found include M. arenaria (72), M. incognita (72), M.
incognita acrita (72, 129), and M. javanica (124, 128, 129, 143). A Meloido-
gyne species was often found associated with passionfruit in South Africa (48).
In tests with P edulis (var. 'Flavicarpa'), fumigation of seedbeds with methyl
bromide or DD resulted in improved plant growth. Yield increases were ob-


trained with pre-plant application of ethoprop, DBCP, or EDB (48). Incidence of
Meloidogyne spp. on passionfruit in Fiji was low, however (102). Kirby (102)
performed pathogenicity tests with three species of Meloidogyne M. aren-
aria, M. incognita, and M. javanica on P edulis (var. 'Flavicarpa'). Al-
though root galling was observed with all three Meloidogyne species in tests,
vine weights in inoculated plants did not differ significantly from uninoculated
controls, and nematode reproduction did not occur on passionfruit. Thus, P ed-
ulis is apparently a non-host for these three species of Meloidogyne, although it
is susceptible to galling from the initial attack. It may be possible that such initial
damage may have resulted in a yield loss which was avoided by the soil treat-
ments used in the South African test (48). Meloidogyne spp. have also been re-
ported from several other Passiflora species (72, 142, 201), including
P quadrangularis L. (72).

Persea americana Mill. Avocado
One of the earliest reports of nematode damage to the avocado is the descrip-
tion of the root lesions attributed to Radopholus similis and a Pratylenchus spe-
cies in Florida (53). These cortical lesions ranged from small brown or black
spots to depressed lesions girdling the roots. Both nematode species were found
in these lesions within the roots of avocados planted in a citrus spreading decline
site (53). Radopholus similis alone was found in avocado roots in a second site,
consisting of a mixed citrus and avocado planting, where both tree crops exhib-
ited decline symptoms (53).
A more extensive survey revealed that R. similis was found in 26 of 61 groves,
all on sandy soils along the Gulf Coast or in Central Florida, but not along the
Florida east coast. Pratylenchus brachyurus was generally distributed through-
out the avocado growing areas of the state (233). Pratylenchus brachyurus was
also found in 40.9% of the samples in a survey of avocados in the Cerrado region
of Brazil (178). When avocado seedlings were inoculated with R. similis and
P brachyurus, R. similis increased to higher population levels than did P brach-
yurus (231). Young and Ruehle (232) recommended that a barrier strip be
established in groves infested with R. similis. Avocado is susceptible to both the
citrus and banana biotypes ofR. similis (153), although it is probably a poor host
for the banana biotype (140).
Of the several nematodes associated with avocados in California(187), plant
damage has been attributed only to Pratylenchus vulnus. In a greenhouse test in
California (192), avocado seedlings inoculated with either 100 or 1000 P vulnus
each were compared to uninoculated controls. One year after inoculation, the
dry weights of the uninoculated control plants were significantly greater than
either of the two groups of inoculated plants. Tree growth was also increased
through preplant fumigation with DD, compared to unfumigated field plots.
Several other nematodes are occasionally associated with damage in avoca-
dos. Minz (144) reported that avocados were sometimes killed by Xiphinema

americanum. Several species of fungi have been observed in association with
Meloidogyne spp. in avocado roots (126). Rotylenchulus reniformis has been re-
ported from avocado in several areas (26, 42, 157, 177, 178, 182, 184). Pea-
cock (157) observed a light infestation on young plants, and Caveness (26) found
that avocado was a good host for R. reniformis. Helicotylenchus cavenessi in-
creased 1000-fold on avocado in a shadehouse test (26), while a moderate in-
crease was observed for Scutellonema clathricaudatum, and slight increase for
Helicotylenchus microcephalus Sher.
Several nematicides have recently been tested on young avocado trees against
mixed populations of plant-parasitic and free-living nematodes, although no
material provided consistent control of all species assayed (62).

Phoenix dactylifera L. Date Palm
Root knot nematodes (Meloidogyne spp.) have been reported from date palm
in the United States (21, 151), Israel (142), and Algeria (113). Several species
of Meloidogyne have been identified from the roots of date palm in Israel, in-
cluding M. arenaria (146), M. hapla (72), M. incognita (146), M. incognita
acrita (72), and M. javanica (72). Damage to date palm seedlings by M. javan-
ica has been described (25). The effects on date palm of three populations of M.
incognita from three different locations (Algeria, Florida, and Italy) and three
populations of M. javanica were examined in a greenhouse test (112). All pop-
ulations caused galling on the four varieties tested ('Bou Feggous', 'Deglet
Noor', 'Kab Kab', and 'Medjool'), although 'Deglet Noor' appeared slightly
less susceptible than the other varieties (112). Because of the susceptibility of
date palm seedlings to root knot nematodes, it is recommended that they be
grown in fumigated or nematode-free soil (151).
A survey of date palms in Algeria revealed that Pratylenchus penetrans was
widespread, occurring at 11 of the 14 oases sampled (113). Fusarium spp. were
also isolated from date palm roots during this survey, and the plant parasites
Xiphinema elongatum Schuurmans Stekhoven & Teunissen andLongidorus con-
goensis Aboul-Eid also were frequently detected. Pratylenchus penetrans had
previously been associated with root damage at the Zelfana oasis in Algeria
(107). Phoenix dactylifera has been recently reported as a host of Radopholus
similis (205).

Psidium guajava L. Guava
Both Meloidogyne arenaria (68, 130) and M. incognita (68, 229) have been
reported to cause galling on Psidium guajava. Damage to roots caused by Me-
loidogyne spp. in Florida has been described (170), and dieback and unsatisfac-
tory yield have been reported on sandy soils. Heavy fertilization, irrigation, and
nutritional sprays have helped to alleviate the problem. In Puerto Rico, trees are
reported to have been killed by root knot nematodes (6). In Cuba, both P gua-

java and P molle Bertol. were affected by both M. arenaria and M. incognita
(68). However, P friedrichsthalianum (Berg.) Nied. showed a high degree of
resistance to both species and was recommended as possible rootstock.
Host-parasite relationships for several other nematodes on guava have also
been investigated. Guava is a host for Hemicriconemoides mangiferae (171,
172), although levels as great as six nematodes per cm3 of soil were not patho-
genic. Caveness (26) considered guava to be only a moderate host of Helicoty-
lenchus cavenessi, H. pseudorobustus, and Tylenchus sp., and a poor host for
Scutellonema clathricaudatum, Hemicycliophora oostenbrinki Luc, and Hoplo-
laimus pararobustus. Psidium guajava is a host of Radopholus similis (17, 29,
159), as is the closely-related P cattleianum Sabine (14, 29, 159). Extremely
high counts (2000/100 cm3 of soil) of the stunt nematode Tylenchorhynchus cy-
lindricus Cobb have been associated with P guajava in Iran (1).

Punica granatum L. Pomegranate
Several species of root knot nematodes have been reported infesting pome-
granate, including Meloidogyne incognita (177, 201, 229), M. incognita acrita
(72), and M. javanica (124, 143, 201). However, nematode damage and yield
losses caused by nematodes on this plant have not been described. Soil around
the roots of Punica granatum is the type habitat for the free-living nematode,
Diphtherophora granata Husain & Khan (81).

Tamarindus indica L. Tamarind
In an inoculation test in Pakistan, Saeed (171) found that Hemicriconemoides
mangiferae multiplied on Tamarindus indica and was pathogenic to the plant at
a level of six nematodes per cm3 of soil. The tamarind is also a reported host of
Radopholus similis (205). Several other species of plant parasitic nematodes
have been found associated with T indica in the state of Sao Paulo, Brazil (149).

Control of nematodes on established fruit trees is difficult. Although chemical
control of nematodes on tropical and subtropical fruits has been investigated in
several cases mentioned here, no chemicals are currently registered in the United
States for nematode control on the plants mentioned, with two exceptions. Fen-
sulfothion is registered for use on Surinam cherry (Eugenia uniflora L.) and Na-
tal plum (Carissa grandiflora D.C.) when these plants are grown as ornamentals
(148), but not on fruit raised for consumption. When established trees do be-
come infected with plant-parasitic nematodes, mulching may be beneficial, and
acceptable yields may sometimes be obtained by maintaining a high level of fer-
tility and using supplementary irrigation. In very serious infestations, it may be
necessary to eradicate infected trees to prevent the spread to other susceptible
Since control of nematodes on established tropical fruit trees is so difficult,
it is important to take preventive measures prior to planting. In general, land that
is infested with a given nematode should be avoided when planting trees which
are hosts of that nematode. If such a site must be used, preplant fumigation may_
be helpful in getting the tree established. Several of the most serious nematode
parasites of tropical fruits, including Radopholus similis, Meloidogyne spp.,
Tylenchulus semipenetrans, Hemicriconemoides spp., Pratylenchus spp., and
Rotylenchulus reniformis, are wholly or partially endoparasitic in roots and thus
may be transmitted on infected plants'. Knowledge of the source and nematolog-
ical condition of the planting material can avoid the introduction of these para-
sitic nematodes into uninfested areas. Sterile mixes should be used when growing
seedlings for transplanting.
In general, nematode resistant varieties of tropical and subtropical fruits are
not yet available, although some resistance has been noted in certain fig vari-
eties. There is, however, the possibility of grafting fig or guava onto resistant
rootstocks of related species, and this procedure has been attempted with Bar-
bados cherry as well. A range of responses has been noted in olive, depending
on the nematode species and the variety of olive used. Although Passiflora ed-
ulis is reported as resistant to several Meloidogyne species, itis nevertheless sub-
ject to galling and it is not clear how much plant damage may occur from this in
a given situation.

Examination of the plant-nematode associations in Appendix I reveals general
trends in the host preferences of certain nematodes. This is particularly impor-
tant in the case of those genera containing the most dangerous plant parasites
on tropical fruits, namely Hemicriconemoides, Meloidogyne, Pratylenchus,
Radopholus, Rotylenchulus, Tylenchulus, and Xiphinema.
Unfortunately, associations with Pratylenchus spp., Rotylenchulus spp., and
Xiphinema spp. are quite generally distributed through most families of tropical
fruits, so it is difficult to detect any preferences of these nematodes for any one
particular group. With the remaining four genera of serious parasites (Hemi-
criconemoides, Meloidogyne, Radopholus, Tylenchulus), some trends are ap-
parent. In general, parasitism by Tylenchulus semipenetrans is evident in four
families of tropical fruits the Rutaceae, Rosaceae (Eriobotrya japonica),
Oleaceae (Olea europaea), and Ebenaceae (Diospyros spp.). Exceptions in sev-
eral other plant families are limited. Among the Rutaceae considered here, Ae-
gle marmelos and Clausena lansium are also hosts of Radopholus similis, as are
Citrus spp. A deviation in this usual pattern for the Rutaceae occurs with Cas-
imiroa edulis, since Meloidogyne spp. are the principal plant parasites reported
on this plant. However, C. edulis has received little study overall, and its fauna
is still largely unknown.
In the Ebenaceae, represented here by Diospyros spp., susceptibility to R.
similis and T semipenetrans is also evident, but several reports of Meloidogyne
spp. are also known. These three nematodes are also listed as associates of
Eriobotrya japonica (Rosaceae). In the olive (Oleaceae), there is also suscep-
tibility to T semipenetrans and Meloidogyne spp., but R. similis has not been
reported, nor have Hemicriconemoides spp., despite relatively widespread sam-
pling of this crop.
Some tropical fruits in certain families are host to Hemicriconemoides spp.,
Meloidogyne spp., and Radopholus similis. Some examples are Malpighiaceae
(Malpighia glabra), Sapindaceae (Litchi chinensis, Euphoria longana), and
Myrtaceae (Psidium guajava). Other less-studied Myrtaceae, such as Eugenia
spp. Feijoa sellowiana, and Syzygium spp. are hosts of R. similis as well.
Similar host-parasite relationships are apparent in the Caricaceae (Carica
spp.), the closely-related Passifloraceae (Passiflora spp.), and the Moraceae
(Artocarpus spp.). Meloidogyne spp. and Rotylenchulus spp. are reported fre-
quently on fruit trees in these families. Few reports of damage to tropical fruits
by Rotylenchulus reniformis exist, but the most convincing evidence comes from
nematode infections of Passiflora edulis and Carica papaya. Tylenchulus sem-

ipenetrans and Radopholus similis have not been reported from the tropical
fruits in these three families.
While it is evident that there may be patterns in the host-parasite relationships
for some groups of tropical fruits, these patterns are only barely discernible at
this point. Little nematological information exists on a great many of the species
of tropical fruits, and as these plants become more widely sampled, such pat-
terns in plant parasitism may become more evident. It is essential for those
working with nematodes on tropical fruits to take care to report actual nema-
tode-fruit associations, and not chance associations resulting from weeds and
other plant species in proximity to the trees. If the latter is the case, then most
plant-parasitic nematode species can eventually be "associated with" tropical
and subtropical fruits.

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Nematode Associate References

Actinidia chinesis Planch. Kiwi Fruit
Meloidogyne spp. 72, 201
Paratrichordorus christei (Allen) Siddiqi 201
Pratylenchus neglectus (Rensch) Filipjev & Schuurmans Stekhoven 201
P vulnus Alien & Jensen 201

Aegle marmelos (L.) Correa Bael
Radopholus similis (Cobb) Thorne 69, 159
Tylenchulus semipenetrans Cobb 209
Xiphinema insigne Loos 89

Annona cherimola Mill. Cherimoya
Criconemoides sp. 63
Gracilacus enata Raski 166
Helicotylenchus anhelicus Sher 190, 201
H. dihystera (Cobb) Sher 201
Heliocotylenchus spp. 63, 144,201
Hoplolaimus sp. 63
Paratylenchus sp. 201
Pratylenchus vulnus 201
Trichodorus sp. 63
Tylenchorhynchus sp. 63

Annona muricata L. Soursop
Helicotylenchus cavenessi Sher 26
H. microcephalus Sher 26
H. pseudorobustus (Steiner) Golden 26
H. retusus Siddiqi & Brown 218
Hemicriconemoides cocophillus (Loos) Chitwood & Birchfield 218
H. squamosum (Cobb) Siddiqi & Goodey 218
Hemicycliophora penetrans Thorne 218
Rotylenchulus sp. 218
Scutellonema clathricaudatum Whitehead 26
Xiphinema americanum Cobb 218
Xiphinema sp. 26

Nematode Associate

Annona reticulata L. Bullock's Heart, Custard Apple
Helicotylenchus cavenessi 26
Meloidogyne incognita (Kofoid & White) Chitwood 177

Annona squamosa L. Sugar Apple, Sweetsop, Custard Apple
Helicotylenchus cavenessi 26
H. dihystera 65, 177, 218
Heterodera sp. 26
Macroposthonia sp. 177
Rotylenchulus sp. 218
Scutellonema clathricaudatum 26
Trophurus indicus Chawla, Bhamburkar, Khan, & Prasad 28
Tylenchorhynchus delhiensis Chawla, Bhamburkar, Khan, & Prasad 28
Tylenchorhynchus martini Fielding 218

Annona spp.
Helicotylenchus dihystera 177
Helicotylenchus sp. 144
Xiphinema sp. 144

Artocarpus atilis (Park.) Fosberg Breadfruit
Syn. A. communis Forst.; A. incisa L.f.
Helicotylenchus cavenessi 26
H. concavus Roman 218
H. dihystera 26, 177
H. microcephalus 26
H. pseudorobustus 26
Meloidogyne incognita acrita Chitwood & Oteifa 72
Meloidogyne sp. 72
Peltamigratus holdemani Sher 177
Rotylenchulus reniformis Linford & Oliveira 26
Scutellonema clathricaudatum 26
Tylenchorhynchus triglyphus Seinhorst 218
Xiphinema ifacolum Luc 26

Artocarpus heterophyllus Lam. Jackfruit
Syn. A. integrifolia L.
Dolichodorus sp. 177
Helicotylenchus dihystera 177, 183
Helicotylenchus sp. 101
Hemicriconemoides birchfieldi Edward, Misra, & Singh 60
H. communis Edward & Misra 57
H. mangiferae Siddiqi 200
Hoplolaimus seinhorsti Luc 218
Meloidogyne incognita 177
Meloidogyne spp. 72, 183
Peltamigratus holdemani 177


Nematode Associate

Peltamigratus sp. 177, 183
Pratylenchus zeae Graham 218
Pratylenchus sp. 101
Quinisulcius acutus (Allen) Siddiqi 218
Rotylenchulus reniformis 177, 183, 199
Rotylenchulus sp. 218
Trichodorus monohystera Allen 177
Trichodorus sp. 183
Tylenchorhynchus martini 218
T triglyphus 218
Xiphinema americanum 218
X. setariae Luc 177, 183

Artocarpus odoratissima Blanco Marang

Helicotylenchus dihystera 218
Hoplolaimus seinhorsti 218
Rotylenchulus sp. 218

Averrhoa carambola L. Carambola
Helicotylenchus dihystera 184
Helicotylenchus sp. 78
Meloidogyne sp. 78,149
Rotylenchulus sp. 78
Tylenchorhynchus crassicaudatus Williams 27
Tylenchorhynchus sp. 78
Xiphinema elongatum Schuurmans Stekhoven & Teunissen 149

Calocarpum sapota (Jacq.) Merril Mamey Sapote
Helicotylenchus dihystera 177
Paratylenchus spp. 177
Pratylenchus sp. 184
Rotylenchulus reniformis 184
Xiphinema spp. 177, 184

Campomanesia maschalentha Berg.

Helicotylenchus dihystera 184
Rotylenchulus reniformis 184
Xiphinema sp. 184

Carica papaya L. Papaya

Helicotylenchus cavenessi 26
Helicotylenchus concavus 218
H. dihystera 26, 190, 218
H. erythrinae (Zimmermann) Golden 218
H. microcephalus 26
H. pseudorobustus 26


Nematode Associate

Helicotylenchus sp.
Hemicriconemoides mangiferae
Hemicycliophora belemis Germani & Luc
H. epicharoides Loof
H. nyanzae Schoemaker
H. oostenbrinki Luc
Hoplolaimus pararobustus (Schuurmans Stel

24, 78, 144, 202,218
171, 172
khoven & Teunissen) Sher 26

Longidorus sp. 218
Macroposthonia curvatum (Raski) de Grisse & Loof 218
Meloidogyne arenaria (Neal) Chitwood 129
M. arenaria thamesi Chitwood 72
M. hapla Chitwood 76, 77
M. incognita 7, 22, 72, 76, 177
M. incognita acrita 129, 186
M. javanica (Treub) Chitwood 40, 82, 122, 124, 129, 143
Meloidogyne spp. 24, 41, 78, 142, 202, 218, 222, 224
Pratylenchus sp. 77
Peltamigratus nigeriensis Sher 26
Pratylenchus coffeae (Zimmermann) Filipjev & Schuurmans Stekhoven 218
P zeae 218
Pratylenchus sp. 26, 77
Rotylenchulus parvus (Williams) Sher 42
R. reniformis 7, 8, 18, 26, 34, 42, 43, 76, 116, 118, 134, 177, 199, 202, 204
Rotylenchulus spp. 218,219
Scutellonema bradys (Steiner & Le Hew) Filipjev 188
S. clathricaudatum 26
Scutellonema sp. 218
Tylenchorhynchus martini 26, 218
Xiphinema americanum 144, 218

Carissa grandiflora D.C. Natal Plum

Meloidogyne spp.
Xiphinema americanum

72, 201

Carissa sp.
Paratylenchus nawadus Khan, Prasad, & Mathur

Casimiroa edulis Llave & Lex White Sapote
Meloidogyne arenaria
M. incognita acrita
M. javanica
Meloidogyne sp.
Scutellonema sp.

Casimiroa spp.
Criconemoides sp.
Meloidogyne spp.
Merlinius brevidens (Allen) Siddiqi


Nematode Associate

Pratylenchus vulnus 201
Xiphinenua americanum 201

Ceratonia siliqua L. Carob

Helicotylenchus dihystera 201
Meloidogyne spp. 72, 201
Pratylenchus vidlnus 201
Xiphinema americanum 201

Chrysophyllum cainito L. Star Apple

Helicotylenchus concavus 218
H. d;In ,,. ,, 184
H. microcephalus 26
Helicotylenchus sp. 218
Pratylenchus zeae 218
Radopholus similis 17, 29
Xiphinema sp. 184

Clausena lansium (Lour.) Skeels Wampi

Radopholus similis 159
Tylenchulus semipenetrans 72

Coccoloba uvifera (L.) Jacq. Sea Grape

Longidorus edmundsi Hunt & Siddiqi 79
Rotylenchulus reniformis 118

Cyphomandra betacea (Cav.) Sendt. Tree Tomato

Globodera rostochiensis (Wollenweber) Mulvey & Stone 72
Meloidogyne arenaria 129
M. hapla 129
M. incognita acrita 129
M. javanica 128

Diospyros ebenaster Retz. Black Sapote

Helicotylenchus concavus 218
Hemicycliophora penetrans Thorne 218
Xiphinema elongatum 218

Diospyros kaki L.f. Japanese Persimmon

Meloidogyne spp. 72, 78
Pratylenchos sp. 78
Radopholus similis 14, 29, 159
Rotylenchulus reniformis 177
Rotylenchulus sp. 78
Trichodorus sp. 78


Nematode Associate

Diospyros lotus L. Persimmon
Tylenchulus semipenetrans 9, 29, 150

Diospyros virginiana L. Common Persimmon, American Persimmon
Meloidogyne sp. 72
Radopholus similis 17, 29, 159
Tylenchulus semipenetrans 11, 31

Diospyros spp.
Criconemoides sp. 201
Helicotylenchus dihystera 201
Meloidogyne hapla 201
Meloidogyne spp. 201
Pratylenchus neglectus 201
P penetrans (Cobb) Filipjev & Schuurmans Stekhoven 201
P vulnus 201
Scutellonema brachyurum (Steiner) Andrassy 201
Trichodorus spp. 201
Tylenchorhynchus spp. 201
Tylenchulusfloridensis (Raski) Maggenti 72
T semipenetrans 201

Durio zibethinus Murr. Durian
Macroposthonia curvatum 218
Rotylenchulus sp. 218
Tylenchorhynchus martini 218

Eriobotryajaponica Lindl. Loquat
Criconemoides spp. 78, 215
Helicotylenchus dihystera 201
Helicotylenchus spp. 78, 113, 135
Longidorus sp. 215
Macroposthonia xenoplax (Raski) de Grisse & Loof 201
Meloidogyne spp. 20, 78, 201
Pratylenchus vulnus 201
Pratylenchus spp. 73, 78, 135
Radopholus similis 14, 29, 159
Rotylenchulus sp. 78
Rotylenchus sp. 201
Tylenchorhynchus clarus Allen 201
T claytoni Steiner 201
Tylenchorhynchus spp. 78, 135
Tylenchulus semipenetrans 201
Tylenchulus spp. 78
Xiphinema americanum 201
X. krugi Lordello 149
Xiphinema spp. 78, 135, 215


Nematode Associate

Eugenia alba Roxb.

Helicotylenchus dihystera 184
Pratylenchus brachyurus 184
Xiphinema sp. 184

Eugenia dombeyi Skeels Grumichama
Syn. E. brasiliensis Lam.
Helicotylenchus cavenessi 149
H. dihystera 184
H. multicinctus (Cobb) Golden 149
Radopholus similis 159
Rotylenchulus reniformis 184
Xiphinema krugi 149

Eugenia uniflora L. Surinam Cherry

Helicotylenchus dihystera 177
Macroposthonia spp. 177
Meloidogyne incognita 177
Paratylenchus spp. 177
Pratylenchus spp. 177
Trophurus spp. 177
Tylenchulus spp. 177

Eugenia spp.

Helicotylenchus dihystera 201
Hemicycliophora eugeniae Khan & Basir 93
Meloidogyne sp. 201
Xiphinema vuittenezi Luc, Lima, Weischer, & Flegg 201

Euphoria longana Lam. Longan

Criconemoides sp. 78
Helicotylenchos sp. 78
Hemicriconemoides mangiferae 200
Meloidogyne sp. 78
Paratylenchus sp. 78
Pratylenchus sp. 78
Radopholus similis 159
Rotylenchulus sp. 78
Rotylenchus sp. 78
Tylenchulus sp. 78

Feijoa sellowiana Berg Feijoa

Helicotylenchus dihystera 201
Radopholus similis 14, 29, 159


Nematode Associate

Feronia limonia (L.) Swingle Indian Wood Apple
Radopholus similis 69, 159

Ficus carica L. Fig
Criconemoides spp. 63, 121
Ditylenchus spp. 144
Helicotylenchus spp. 113
Hemicycliophora spp. 121
Heteroderafici Kirjanova 13, 51, 193, 201
Hoplolaimus sp. 144
Longidorus martini Merney 230
Longidorus sp. 71
Meloidogyne arenaria 83, 91, 129, 143
M. incognita 46, 50, 121, 143, 173
M. incognita acrita 72, 129, 143, 161,208
M. javanica 72
Meloidogyne spp. 103, 141, 142, 201,214
Merlinius brevidens 201
Paratrichodorus christei 103
Paratylenchus hamatus Thorne & Allen 201, 217
P nainianus Edward & Misra 55
Paratylenchus spp. 201
Pratylenchus coffeae 103
P minyus Sher & Allen 144
P musicola (Cobb) Filipjev 206
P neglectus 201
P scribneri Steiner 144
P thornei Sher & Allen 201
P vulnus 191,201,216
Pratylenchus spp. 141, 215
Quinisulcius capitatus (Allen) Siddiqi 201
Rotylenchulus macrodoratus Dasgupta, Raski, & Sher 221
Rotylenchus spp. 71, 144
Scutellonema truncatum Sher 189
Trichodorus porosus Allen 201
Trichodorus spp. 71, 121,141,144
Tylenchorhynchus spp. 63, 141, 144,201
Xiphinema americanum 201
X. index Thorne & Allen 113, 127,160,162,201,217,225,226,227,228
X. mediterraneum Martelli & Lamberti 39, 113
X. vuittenezi 201
Xiphinema spp. 71, 103, 141, 144,215

Flacourtia ramontchi L'Herit. Governor's Plum
Helicotylenchus dihystera 184
Peltamigratus sp. 184


Nematode Associate

Garcinia mangostana L. Mangosteen

Meloidogyne sp.
Peltamigratus sp.
Rotylenchulus reniformis
Tylenchorhynchus martini
Xiphinema elongatum
Xiphinema sp.

Genipa americana L. Genipa

Pratylenchus brachyurus
Xiphinema sp.

Harpephyllum caffrum Bernh. Kaffir Plum

Helicotylenchus dihystera

Lansium domesticum Jack Langsat

Macroposthonia curvatum
Meloidogyne sp.
Tylenchorhynchus triglyphus
Xiphinema elongatum

Litchi chinensis Sonn. Lychee, Litchi

Criconemoides sp.
Helicotylenchus dihystera
H. microcephalus
H. pseudorobustus
Helicotylenchus sp.
Hemicriconemoides litchi Edward & Misra
H. mangiferae
Hemicriconemoides sp.
Longidorus sp.
Meloidogyne javanica
Meloidogyne spp.
Paratylenchus sp.
Pratylenchus spp.
Radopholus similis
Rotylenchulus sp.
Trichodorus litchi Edward & Misra
Trichodorus sp.
Tylenchulus sp.
Xiphinema americanum
X. brevicolle Lordello & daCosta
X. orthotenum Cohn & Sher
Xiphinema sp.

75, 136,137,138,200,207
54, 153, 159
136, 137, 138,207


Nematode Associate

Malpighia coccigera L.
Helicotylenchus dihystera 184
Macroposthonia sp. 184
Meloidogyne sp. 184

Malpighia glabra L. Barbados Cherry, Acerola
Criconemoides sp. 6
Helicotylenchus sp. 6
Hemicriconemoides communis 57
Meloidogyne incognita 6, 77
M. incognita acrita 117
Radopholus similis 14, 29, 159
Rotylenchulus sp. 6

Maliphia infestissima (A. Juss.) Rich Cowhage Cherry
Rotylenchulus reniformis 8

Malpighia punicifolia L. West Indian Cherry
Rotylenchulus reniformis 8

Malpighia sp.
Meloidogyne javanica 72

Mangifera indica L. Mango
Criconema squamosum (Cobb) Taylor 167
Criconemoides sp. 78
Ditylenchus emus Khan, Chawla, & Prasad 94
Ditylenchus sp. 179
Helicotylenchus cavenessi 26
H. dihystera 177, 179, 184, 185, 190, 218
H. erythrinae 92
H. mangiferensis Elmiligy 64
H. microcephalus 190
H. multicinctus 65, 190
H. pseudorobustus 26
Helicotylenchus spp. 6, 23, 78, 101, 139, 144, 177, 179
Hemicriconemoides birchfieldi Edward, Misra, & Singh 60
H. communis 56, 57, 167
H. mangiferae 75, 92, 139, 158, 167, 171, 172, 196, 200
H. squamosus 72, 218
Hemicriconemoides sp. 23
Hemicycliophora cocophillus (Loos) Goodey 26
H. mangiferum Misra & Edward 147
Hirschmanniella sp. 78
Hoplolaimus columbus Sher 176


Nematode Associate

H. indicus
H. pararobustus
H. seinhorsti
H. tylenchiformis Daday
Hoplolaimus spp.
Longidorus brevicaudatum (Schuurmans Stekhoven) Siddiqi
Longidorus sp.
Macroposthonia citri (Steiner) deGrisse & Loof
M. curvatum
M. xenoplax
Macroposthonia sp.
Meloidogyne incognita
Meloidogyne spp.
Merlinius bijnorensis Khan
Paratylenchus sp.
Pratylenchus brachyurus (Godfrey) Filipjev &
Schuurmans Stekhoven
Pratylenchus sp.
Radopholus similis
Rotylenchoides brevis Whitehead
Rotylenchulus reniformis 23,
Rotylenchulus sp.
Rotylenchus sp.
Scutellonema brachyurum
S. clathricaudatum
S. mangiferae Khan & Basir
S. truncatum
S. unum Sher
Trichodorus sp.
Tylenchorhynchus martini
Tylenchorhynchus spp.
Tylenchulus sp.
Xiphinema americanum 77,
X. basiri Siddiqi
X. brevicolle

X. ebriense Luc
X. elongatum
X. ifacolum
X. lambertii Bajaj & Jairajpuri
X. nigeriense Luc
Xiphinema spp.

Manilkara achras (Mill.) Fosberg Sapodilla
Syn. Achras zapota L.

Helicotylenchus indicus
H. multicinctus
H. retusus Siddiqi & Brown
Helicotylenchus sp.
Hemicriconemoides cocophilus


125, 167, 179,219
144, 167, 195,218

171, 172


Nematode Associate

H. mangiferae
Hemicycliophora penetrans
Hoplolaimus columbus
H. seinhorsti
Macroposthonia curvatum
Paratylenchus spp.
Pratylenchus zeae
Rotylenchulus reniformis
Rotylenchulus sp.
Tylenchorhynchus martini
Xiphinema americanum
X. elongatum

Monstera deliciosa Liebm. Ceriman
Meloidogyne arenaria
M. incognita
M. incognita acrita
Radopholus similis

Myrcianthes edulis Berg.
Helicotylenchus dihystera
H. pseudorobustus
Xiphinema sp.

Myrciariajaboticaba Berg. Jaboticaba
Helicotylenchus dihystera
Macroposthonia spp.

Nephelium lappaceum L. Rambutan
Hemicriconemoides squamosus
Paratylenchus breviculus Raski

Olea europaea L. Olive
Criconemoides spp.
Dolichodorus heterocephalus Cobb
Gracilacus peratica Raski
Helicotylenchus digonicus Perry
H. dihystera
H. erythrinae
H. neopaxilli Inserra, Vovlas, & Golden
H. oleae Inserra, Vovlas, & Golden
Helicotylenchus sp.
Hoplolaimus sp.
Longidorus elongatus (deMan) Thorne & Swanger
Longidorus sp.
Meloidogyne arenaria

171, 172
171, 172
171, 218
171, 172

38, 72
159, 219

72, 74, 201


Nematode Associate

M. hapla 109, 145
M. incognita 109, 145, 201
M.javanica 49, 109, 114, 124, 143,201,212
Meloidogyne spp. 21,74, 142,201
Merlinius brevidens 201
Nacobbus dorsalis Thorne & Allen 201
Paratrichodorus christiei 201
Paratylenchus spp. 113, 201
Pratylenchus coffeae 37
P crenatus Loof 201
P neglectus 201
P penetrans 201
P thornei 201
P vulnus 3, 37, 106, 107, 123, 201
Rotylenchulus macrodoratus Dasgupta, Raski, & Sher 220, 221
R. macrosomus Dasgupta, Raski, & Sher 42
R. parvus 201
Rotylenchulus sp. 113
Rotylenchus robustus (DeMan) Filipjev 201
Trichodorus porosus 201
Trichodorus sp. 71
Tylenchorhynchus clarus 201
T cylindricus Cobb 201
Tylenchorhynchus sp. 49, 113
Tylenchulus semipenetrans 9, 10, 29, 49, 104, 112, 115,201
Xiphinema americanum 105, 201
X. elongatum 49
X. ingens Luc & Dalmasso 113
Xiphinema spp. 71

Passiflora edulis Sims Passionfruit
Dolichodorus minor
Dolichodorus sp.
Helicotylenchus dihystera
Helicotylenchus spp.
Meloidogyne arenaria
M. incognita
M. incognita acrita
M. javanica
Meloidogyne spp.
Peltamigratus holdemani
Pratylenchus sp.
Rotylenchulus reniformis
Trichodorus monohystera
Xiphinema parhtaliae Loof & Sharma
Xiphinema spp.

77, 177, 180
72, 103
72, 77, 102, 177
72, 129
102, 124, 128, 129, 143
48, 142, 156, 180
177, 180
8, 19,77, 102, 177, 180
177, 180


Nematode Associate

Passiflora quadrangularis L. Giant Granadilla

Meloidogyne incognita acrita
M. javanica

Passiflora spp.
Helicotylenchus c!, i~,. ,i
Helicotylenchus sp.
Meloidogyne incognita acrita
M. javanica
Rotylenchulus reniformis
Scutellonema sp.
Xiphinema americanum
X. surinamense Loof & Maas

Persea americana Mill. Avocado
Criconema sp.
Criconemoides informs (Micoletzky) Taylor
Criconemoides spp.
Ditylenchus spp.
Dolichodorus spp.
Helicotylenchus cavenessi
H. concavus
H. dihystera
H. erythrinae
H.flatus Roman
H. microcephalus
H. rotundicauda Sher
Helicotylenchus spp.
Iih ,, I. li.?rt, ,,' spp.
Hoplolaimus seinhorsti
Hoplolaimus sp.
Longidorus africanus Merny
Macroposthonia citri
M. mutabile (Taylor) deGrissee & Loof
M. sosamossi del Prado Vera
Macroposthonia spp.
Meloidogyne incognita
Meloidogyne spp.
Merlinius brevidens
Nothocriconema demani del Prado Vera
Paratrichodorus christei
Paratylenchus hamatus
Paratylenchus sp.
Peltamigratus holdemani Sher
Pratylenchus brachyurus
P neglectus
P penetrans
P thornei

72, 142, 201
118, 177

149, 177, 178, 182, 184, 190, 201
190, 201
126, 178, 182, 187,201,218


Nematode Associate

P vulnus
Pratylenchus sp.
Quinisulcius acti (Hopper) Siddiqi
Radopholus similus
Rotylenchulus reniformis
Rotylenchulus sp.
Rotylenchus breviglans Sher
Rotylenchus spp.
Scutellonema brachyurum
S. clathricaudatum
Trichodorus monohystera Allen
T porosus
Trichodorus spp.
Tylenchorhynchus clarus
T claytoni
T triglyphus
Tylenchorhynchus spp.
Tylenchulus semipenetrans
Xiphinema americanum
X. ensiculiferum (Cobb) Thorne
X. ifacolum
X. setariae
X. vuittenezi
Xiphinema spp.

Phoenix dactylifera L. Date
Criconemoides sp.
Helicotylenchus dihystera
Helicotylenchus sp.
Hemicriconemoides mangiferae
Longidorus congoensis Aboul-Eid
Meloidogyne arenaria
M. hapla
M. incognita
M. incognita acrita
M. javanica
Meloidogyne spp.
Paratylenchus sp.
Pratylenchus penetrans
Pratylenchus sp.
Radopholus similis
Trichodorus sp.
Tylenchorhynchus goffarti Sturhan
T latus Allen
Xiphinema elongatum

Phyllanthus emblica L. Emblic
Hemicriconemoides birchfieldi
Radopholus similis

3, 187,192,201
29,53, 159,231,232,233
189, 201
62,126, 144, 187, 201
144, 187,201
177, 182
177, 182

112, 146
107, 113


Nematode Associate

Phyllanthus spp.
Meloidogyne incognita
M. incognita acrita
M. javanica
Radopholus similis

P'li, !,, angulata L.

Meloidogyne incognita acrita
M. javanica

P'.. -:,; v peruviana L. Cape Gooseberry

Meloidogyne javanica
M. hapla
M. incognita

Pouteria caimito (Ruiz & Pav.) Radlk. Abiu
Helicotylenchus dihystera
Pratylenchus brachyurus
Xiphinema sp.

Psidium guajava L. Guava
Belonolaimus sp.
Criconemoides sp.
Ditylenchus nanus Siddiqi
Dolichodorus minor
Helicotylenchus abunaamai Siddiqi
H. agricola
H, cavenessi
H. concavus
H .. 1.
H. multicinctus
H. pseudorobustus
Helicotylenchus spp.
Hemicriconemoides cocophillus
H. mangiferae
H. squamosus
Hemicriconemoides sp.
Hemicycliophora oostenbrinki
H. penetrans
Hemicycliophora spp.
Hoplolaimus parabustus Elmiligy
H. pararobustus
H. seinhorsti
Longidorus psidii Khan & Khan
Macroposthonia onoensis
Meloidogyne arenaria

72, 159

41, 130


177, 184

177, 184, 218
78, 177, 181,201,218
171, 172
68, 130


Nematode Associate

M. incognita
Meloidogyne spp.
Paralongidorus similis Khan, Chawla, & Prasad
Paratylenchus sp.
Peltamigratus holdemani
Pratylenchus impar Khan & Singh
Pratylenchus spp.
Radopholus similis
Rotylenchulus reniformis
Rotylenchulus sp.
Rotylenchus buxophilus Golden
R. helicus Husain & Khan
Rotylenchus sp.
Scutellonema clathricaudatum
S. orientalis Rashid & Khan
Trichodorus sp.
Tylenchorhynchus cylindricus
T martini
Tylenchorhynchus sp.
Tylenchulus sp.
Xiphinema americanum
X. neoelongatum Bajaj & Jairajpuri
X. setariae
Xiphinema spp.

Psidium cattleianum Sabine Cattley Guava
Paratylenchus nainianus
Radopholus similis

68, 229
8, 125, 177, 184
78, 177, 181

14, 29, 159

Psidium spp.
Hoplolaimus indicus

Punica granatum L. Pomegranate
Criconema punici Edward, Misra, Peter, & Rai 59
Ditylenchus minutus Husain & Khan 80
Helicotylenchus dihystera 149, 177, 184
H. erythrinae 201
H. pseudorobustus 149
H. punicae Swarup & Sethi 211
Helicotylenchus spp. 113,177
Macroposthonia macrolobata (Jairajpuri & Siddiqi) de Grisse & Loof 90
M. ornatum (Raski) deGrisse & Loof 149
Meloidogyne incognita 177,201,229
M. incognita acrita 72
M. javanica 124, 143,201
Meloidogyne spp. 78,142,201
Merlinius brevidens 201


Nematode Associate

Paratylenchus spp. 201
Pratylenchoides crenicauda Winslow 87
Pratylenchus brachyurus 184, 201
P neglectus 201
P vulnus 201
Pratylenchus spp. 201
Rotylenchulus ,.ni r.. *..,, 177, 184
Trichodorus spp. 201
Tylenchorhynchus clarus 201
Tylenchorhynchus spp. 177
Xiphinema spp. 177, 184

Rollinia deliciosa Safford Biriba, Corosol
Radopholus similis 72, 159

Rollinia sp.
Xiphinema krugi 149

Sandoricum koetjape Merr. Santol
Helicotylenchus concavus 218
Helicotylenchus sp. 218
Tylenchorhynchus triglyphus 218
Tylenchorhynchus sp. 218

Spondias cytherea Sonn. Golden Apple
Syn. S. dulcis Forst.
Helicotylenchus cavenessi 149
H ;;,.' ,, ',a 177
Helicotylenchus spp. 177
Meloidogyne incognita 177
M. javanica 149
Pratylenchus brachyurus 177

Spondias mombin L. Hog Plum
Syn. S. lutea L.
Meloidogyne sp. 72

Spondiaspurpurea L. Red Mombin
Pratylenchus brachyurus 218
Xiphinema americanun 218
X. elongatum 218


Nematode Associate

Syzygium cuminii Skeels Jambolan Plum
Syn. Eugeniajambolana Lam.
Helicotylenchus concavus 218
H. dihystera 184
Pratylenchus brachyurus 184
Radopholus similis 159, 219
Trichodorus porosus 184
Tylenchorhynchus chonai Sethi & Swarup 175
T triglyphus 218

Syzygiumjambos (L.) Alston Rose Apple
Radopholus similis 159

Syzygiumjavanicum (Lam) Merr. & Perry Wax Jambu
Syn. S. samarangense Blume
Helicotylenchus concavus 218
Tylenchorhynchus triglyphus 218
Xiphinema elongatum 218
X. insigne 218

Syzygium malaccensis L. Malay Apple
Helicotylenchus dihystera 177
Meloidogyne incognita 177
Pratylenchus brachyurus 177
Rotylenchulus reniformis 118
Tylenchorhynchus martini 177
Xiphinema spp. 177

Syzygium paniculatum Gaertn. Brush Cherry
Syn. Eugenia paniculata Banks
Radopholus similis 159

Tamarindus indica L. Tamarind
Helicotylenchus pseudorobustus 149
Hemicriconemoides mangiferae 171
Hoplolaimus galeatus (Cobb) Filipjev & Schuurmans Stekhoven 149
Longidorus elongatus 194
Meloidogyne sp. 72, 149
Radopholus similis 205
Tylenchorhynchus latus Allen 149
Xiphinema elongatum 149
X. krugi 149


Nematode Associate References

Zizyphusjujuba Mill. Chinese Jujube
Criconemoides sp. 78
Helicotylenchus sp. 78
Hemicriconenoides communis 57
Meloidogyne sp. 78
Pratylenchus sp. 78
Tylenchorhynchus sp. 78

7i,: 'imn, mucronata Willd.
Meloidogyne sp. 72

Zizyphus spp.
Paratylenchus neonanus Mathur, Khan, & Prasad 132


Common Name



American Persimmon



Barbados Cherry


Black Sapote


Brush Cherry

Bullock's Heart

Cape Gooseberry



Cattley Guava



Chinese Jujube

Common Persimmon


Cowhage Cherry

Custard Apple


Scientific Name

Pouteria caimito

Malpighia glabra

Diospyros virginiana

Persea americana

Aegle marmelos

Malpighia glabra

Rollinia deliciosa

Diospyros ebenaster

Artocarpus atilis

Syzygium paniculatum

Annona reticulata

Physalis peruviana

Averrhoa carambola

Ceratonia siliqua

Psidium cattleianum

Monstera deliciosa

Annona cherimola

Zi: '1ai:" jujuba

Diospyros virginiana

Rollinia deliciosa

Malpighia infestissima

Annona reticulata

Phoenix dactylifera

Common Name






Giant Granadilla

Golden Apple

Governor's Plum



Hog Plum

Indian Wood Apple



Jambolan Plum

Japanese Persimmon


Kaffir Plum

Kiwi Fruit






Malay Apple

Mamey Sapote


Scientific Name

Durio zibethinus

Phyllanthus emblica

Feijoa sellowiana

Ficus carica

Genipa americana

Passiflora quadrangularis

Spondias cytherea

Flacourtia ramontchi

Eugenia dombeyi

Psidium guajava

Spondias mombin

Feronia limonia


Artocarpus heterophyllus

Syzygium cuminii

Diosypyros kaki

Zizyphus spp.

li .p, ph li ,.,, caffrum

Actinidia chinensis

Lansium domesticum

Litchi chinensis

Euphoria longana

Eriobotrya japonica

Litchi chinensis

Syzygium malaccensis

Calocarpum sapota

Mangifera indica

Common Name




Natal Plum







Red Mombin

Rose Apple



Sea Grape


Star Apple

Sugar Apple

Surinam Cherry



Tree Tomato


Wax Jambu

West Indian Cherry

White Sapote

Scientific Name

Garcinia mangostana

Artocarpus odoratissima

Spondias spp.

Carissa 1'i: 'r,

Olea europaea

Passiflora edulis

Carica papaya

Diospyros spp.

Punica granatum

Nephelium lappaceum

Spondias purpurea


Sandoricum koetjape

Manilkara achras

Coccoloba uvifera

Annona muricata

Chrysophyllum cainito

Annona squamosa

Eugenia uniflora

Annona squamosa

Tamarindus indica

Cyphomandra betacea

Clausena lansium


Malpighia punicifolia

Casimiroa edulis

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costof 78 cenis a copy to provide information on plant parasitic nematodes
associated with tropical and subtropicalfruits.

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