JOURNAL OF NEMATOLOGY

VOLUME 25

SEPTEMBER 1993

NUMBER 3

Journal of Nematology 25(3):315-331. 1993.

T h e Society of Nematologists 1993.

Feeding Habits in Soil Nematode Families and

Genera--An Outline for Soil Ecologists
G. W.

YEATES, 1 T .

BONGERS,2 R. G. M. DE GOEDE, 3 D. W.
S. S. GEORGIEVA5

FRECKMAN, 4 AND

Abstract: Because research on nematode involvement in trophic interactions, foodweb structure,

and biodiversity is constrained by lack of an overview of nematode feeding habits, this outline
presents a consensus of current thought on nematode feeding habits. T h e source of food is fundamental to trophic interactions and provides the basis for our definitions of the essential feeding
types: 1) plant feeder, 2) hyphal feeder, 3) bacterial feeder, 4) substrate ingester, 5) predator of
animals, 6) unicellular eucaryote feeder, 7) dispersal or infective stage of parasites, and 8) omnivore.
Lists of families and genera with their presumed feeding types are given. Major gaps in knowledge
of feeding in the smaller tylenchids and many dorylaims are noted.
Key words: bacterial feeding, feeding habit, foodweb, fungal feeding, nematode, omnivore, predator, soil ecology, trophic interaction.

With the increasing interest of soil ecologists in the role of nematodes in ecosystem processes (roles such as nutrient cycling, biological control and economic crop
loss), there is an unmet need for a concise
summary of current knowledge of nematode feeding habits. T h e analysis of available information is made increasingly difficult by changes in nematode systematics
and recent contributions to nematode ecology. When all species of nematodes can be
confidently assigned to feeding groups,

Received for publication 12 January 1993.

Review.
z Landcare Research, Private Bag 31902, Lower Hurt, New
Zealand.
2 Nematology Department, Wageningen Agricultural University, P.O.B. 8123, 6700 ES Wageningen, The Netherlands.
3 Biological Station of Wageningen Agricultural University, Kampsweg 27, 9418 PD Wijster, The Netherlands.
Department of Nematology, University of California,
Riverside, CA 92521. Present Address: Natural Resource
Ecology Laboratory, Colorado State University, Fort Collins,
CO 80523.
5 Nematology Laboratory, Department of Zoology, Faculty
of Biology, University of Sofia, Sofia 1421, Bulgaria.
We are grateful to B. Sohlenius, V. R. Ferris, D. C. Coleman, S. Bostrom, R. K. Niles, K. Prejs, D. Wardle, and P.
Arpin for their comments on the manuscript, and to D.J.
Carlisle, E. M. Courtright, and T. Gates of D.W.F.'s lab for
proofreading and library research. Support from the Foundatiott for Research, Science and Technology (NZ) to G.W.Y.
and from the National Science Foundation (USA) Grant BSR
8818049 to D.W.F. is gratefully acknowledged.

there will follow a better understanding of

the role of nematodes in soil and how
changes in environmental factors influence the composition o f the n e m a t o d e
fauna.

T h e first c o m p r e h e n s i v e review o f
n e m a t o d e feeding habits was given by
Neilsen (77). In an attempt to produce
functional groups based on feeding habits,
Paramonov (84) applied to n e m a t o d e s
terms such as "pararhizobes" (occur in the
rhizosphere and sometimes damage
plants) and "dyssaprobes" (feed in decomposing material but may enter healthy tissue). Both Wasilewska (121) and Yeates
(127) grouped plant and soil nematodes by
feeding habits. The classification of Tylenchida advocated by Siddiqi (99) has a
strong "feeding habit" component. Recent
ecological studies have revealed that feeding-habit groupings may not be sharply
delimited. For example, abundant populations o f Aphelenchoides, Tylenchus, Tylencholaimus, and Ditylenchus were discovered
that could only be classified as "root/fungal
feeding nematodes" (105), as well as "predacious" mononchids that multiplied using
bacteria as a food source (130). These examples demonstrate the apparently arbitrary nature of traditional nematode feed-

T h e J O U R N A L OF NEMATOLOGY for J u n e (25:101-313) was

issued 11 J u n e 1993.

315

316 Journal of Nematology, Volume 25, No. 3, September 1993

ing groups. Moreover, feeding habits of
m a n y n e m a t o d e s have b e e n i n f e r r e d
rather than confirmed by maintenance
over many generations under biologically
defined conditions.
Following Petersen and Luxton (86), we
use "grazing foodweb" and "detritus foodweb" as terms for communities based on
living green plants and dead organic matter, respectively. Their comments on the
merging of the two webs are particularly
relevant for nematodes, which are so often
abundant at interfaces between living and
dead material (as in the rhizosphere).
This paper is not a literature review and
does not present new results; rather we
present a basis for advancing understanding of the role of nematodes in soil ecology. The paper originated in discussions at
the S e c o n d I n t e r n a t i o n a l N e m a t o l o g y
Congress and lists nematode families and
genera with our assessment of current understanding of their feeding habits. We
hope that this paper will serve as a framework for ecologists to use independently of
taxonomic philosophies. Andr~ssy (2,3),
L t r e n z e n (58), Maggenti (62), and Siddiqi
(99) all have differing approaches to the
general classification of nematodes. The
most recent taxonomic overview is contained in the Manual o f Agricultural
Nematology (76). T h e generic makeup of
the families we use is compatible with these
recent works; because they often assign
differing taxonomic ranks to these groups,
we do not list authorities.
CONFLICTING RESULTS AND OBSERVATIONS

Soil ecologists are primarily concerned

with relationships between biological poputations and the soil environment, whether
it be a high-input agroecosystem, natural
ecosystem, or an area managed for sustained production. When these populations are cultured singly or together in the
laboratory, many possible interactions and
their consequences are reduced (52); thus,
results are difficult to extrapolate to field
conditions. T h e following are examples of
such difficult extrapolations.

1. Although normally regarded as bacterial feeders, Chiloplacus (Acrobelidae)

and Rhabditis (Rhabditidae) have been cultured on the fungi Phoma sp. and Pythium
middletonii, respectively (39,92). In both
cases, the "primary" food source is bacterial; the fact that the apparent secondary
food belongs to the same trophic level of
the "detritus foodweb" as the primary food
shows the value of acknowledging such
broad foodwebs. However, our interpretation of this dual feeding habit is i) bacterial
contamination of the nematode cultures or
ii) direct uptake o f nutrients by nematodes, as occurs in axenic culture of Caenorhabditis elegans (21,70,132).
2. H o o p e r and Cowland (45) cultured
the foliar nematode Aphelenchoides ritzemabosi, which normally feeds in the "grazing
foodweb," on fungi ("detritus foodweb"),
reinforcing observations that fungal feeding is the n o r m a l situation in A p h e lenchida. Just as plant root cells are fed on
in a variety of ways (see "plant feeding"), so
are fungi (1).
3. It is difficult to extrapolate data from
closely controlled monoxenic cultures of
Pratylenchus and Radopholus on carrot discs
to the heterogenous environment of field
populations.
4. Although certain mononchids successfully cultured on bacteria contain living bacteria within the intestines (6), we
question whether sufficient aggregations
of bacteria exist under field conditions for
bacterial ingestion to be of significant nutritional importance to the large mononchids.
5. The intestine of mononchids, rhabditids, dorylaims, etc., often appear pigmented, but such pigmentation has not
been observed in Tylenchida or plantfeeding Trichodorus and Longidorus (76).
There has been no attempt to relate this
pigmentation to feeding habits.
6. Axenic culture of Caenorhabditis elegans and other "bacterial feeding" nematodes (21,70,73,132) highlights problems
of interpretation, but there is evidence that
development may be slower under axenic
conditions (20). Bacterial feeding is re-

Feeding Habits of Soil Nematodes: Yeates et al. 317

garded here as the principal source of nutrition for such nematodes u n d e r field
conditions. Physical crushing of bacteria
has been demonstrated in the pharyngeal
bulb of Acrobeloides nanus, but bacterial
feeding nematodes may defecate living
bacteria and there is no general knowledge
as to whether some nematodes actually kill,
rupture, or lyse bacteria or merely remove
adhering organic c o m p o u n d s (126). In
several situations, there may be a degree of
direct nutrient uptake through the cuticle
or epidermis (33,50,79,132).
7. A l t h o u g h identification to family
level is usually adequate, identification to
species is sometimes necessary to accurately assign nematodes to trophic groups.
For example, Ditylenchus dipsaci is an economically important plant feeding species,
but most Ditylenchus spp. are hyphal feeding.
8. T h e delicate-speared T y l e n c h i d a e
and Psilenchidae remain problematic. Although Sohlenius et al. (105) regarded several groups of them as "root/fungal feeding nematodes," the literature is conflicting and we have been unable to form a
consensus about the classification of these
two families. We have generally listed
them as feeding on root epidermal cells
and root hairs. Specific studies of their
feeding in the rhizoplane would provide
valuable information.
FEEDING TYPES OF NEMATODES IN PLANT

AND SOIL SYSTEMS

Ideally the feeding habits of each nematode species should be determined in each
particular ecological setting. Because this
task is impractical, we offer the following
outline for general use. Regardless of the
trophic group, we strongly advocate app e n d i n g species lists a n d d e p o s i t i n g
voucher specimens to make reinterpretation possible.
1. Plant feeding: This involves feeding
on vascular plants; a tylenchoid stomatostyle or dorylaimoid odontostyle is always
present. The posthatching life stages of
most species are migratory. In sedentary

species, the feeding site of the female may

be undifferentiated, uninucleate, or polynucleate. Males of sedentary species sometimes have a degenerate stylet or reduced
oesophagus, but data on the nutrition of
such males is inadequate. Plant feeders
may be polyphagous or show host specificity. Migratory species may generally be
classified as ecto- or endoparasites. Feeding sites may be root-hair, epidermal, cortical, or vascular.
Although apparently not actively feeding, the migratory phases of Pratylenchus,
infective second-stage Heterodera, and resistant stages of Paratylenchus are each an
essential part of the life-cycle of these obligate plant feeders and are thus included
in this category. This group includes algal
feeders that have a narrow stylet and do
not ingest chloroplasts, in contrast to types
that swallow unicellular algae more or less
intact.
The group may be subdivided into the
following six groups: la) sedentary parasites (e.g., females of Heterodera, Globodera,
Meloidogyne, Verutus, Sphaeronema); lb)
m i g r a t o r y e n d o p a r a s i t e s (e.g., Pratylenchidae, some Anguinidae); lc) semiendoparasites (e.g., Hoplolaimidae, Telotylenchus); ld) ectoparasites (e.g., Dolichodoridae, Cephalenchus, Criconematidae,
Hemicycliophoridae, Paratylenchidae,
Trichodoridae, Pungentus, Longidoridae);
le) epidermal cell and root hair feeders
(e.g., T y l e n c h i d a e , Psilenchidae, Atylenchidae); and lf) algal, lichen (algal or
fungal component), or moss feeders that
feed by piercing (e.g., Tylenchus, Laimaphelenchus, Anguinidae).
2. Hyphalfeeding: This involves penetration of fungal hyphae by a stomatostyle or
odontostyle (stylet or spear). In addition to
obligate hyphal feeders, this group includes the alternate life cycle of some invertebrate parasites (e.g., Deladenus) (75).
Yeasts may be included as a food source
under this heading, except when ingested
whole (see "unicellular eucaryote feeding"). Feeding on hyphae of saprophytic
fungi has ecological implications quite different from feeding on mycorrhizal fungi.

318 Journal of Nematology, Volume 25, No. 3, September 1993

There is need for research on such interactions to determine whether the same
nematode can feed on both saprophytic
and mycorrhizal fungi.
3. Bacterial feeding: This category includes species that feed on any procaryote
food source, whether through a narrow
(Rhabditis, Alaimus) or broad (Diplogaster)
mouth; axenic culture indicates that there
may be an element of absorption of nutrients (33,50,79,132). Species with a broad
mouth may ingest other types of food. The
soil stages of certain nematode parasites of
vertebrates and invertebrates that feed on
bacteria should be included. Although
morphological data (75) suggest that infective stages of Steinernema may be nonfeeding, they may be mass produced on
artificial media, providing their bacterial
symbionts are present (76). Some Rhabditidae and Diplogasteridae may use a
phoretic (transport) host, especially insects.
4. Substrate ingestion: This type of feeding occurs in at least diplogasterids and
Daptonema sp. Substrate ingestion may be
incidental to bacterial feeding, predation,
and unicellular eucaryote feeding in many
groups, because more than a pure food
source is ingested. Although soil nematodes may ingest and lyse bacteria, there is
no available evidence that digestion of
complex organic substrates occurs in the
gut of soil nematodes (a fact that may reflect technical difficulty). Consequently, no
genera have been given primary classification under this type. The expression "nonselective deposit feeding" used for aquatic
nematodes covers a similar situation.
5. Animal predation: Some species of
nematodes feed on invertebrates such as
protozoa, nematodes, rotifers, and
enchytraeids, either as "ingesters" (type
5a; e.g., Diplogaster, Mononchus, Nygolaimus)
or as "piercers" (type 5b), sucking body
fluids t h r o u g h a n a r r o w stylet (e.g.,
Seinura, Labronema, Laimaphelenchus). The
intestine of piercers never contains distinct
prey remnants.
6. Unicellular eucaryote feeding: A wide

range of nematodes reportedly feed on diatoms or other algae, but lack of marker
structures in the nematode food and the
presence of globules, pigmentation, or inclusions in nematode intestines make determination difficult. This feeding type includes ingestion of fungal spores and
whole yeast cells.

7. Dispersal or infective stages of animal

parasites: Other stages of animal parasitic
nematodes outside their alternate or definitive hosts may occur in the soil (e.g., Deladenus, Heterorhabditis) or vertebrates (e.g.,
Strongyloides); we include here entomogenous species. If they feed and contribute to
soil processes, these species should be included in other appropriate categories; if
they invade a host they leave the soil system (in the narrow sense); if they die in the
soil they contribute to nutrient pools. We
do not include in this category forms (especially Rhabditidae and Diplogasteridae)
that merely use animals as phoretic (transport) hosts.
8. Omnivorous: Although some nematodes appear normally to feed on a wide
range of foods (particularly combining
feeding types 2-6), it is best to restrict use
of this term to (a few) dorylaims (31,95);
when possible, nematodes should be classified in types 1-7. A similar procedure
was used for soil fauna by Petersen et al.

(87).
FEEDING TYPES IN NOMINAL ORDERS
AND FAMILIES

In the following systematic list of nematode families and feeding type, we give indicative genera for each family; the numbers refer to the eight feeding types. Numbers in p a r e n t h e s e s r e f e r to tentative
assignments or situations with alternative
food sources. For a given family or genus,
if two or more types are indicated, they are
presented in strictly numerical order. For
some groups, detailed information is given
(e.g., algal feeding may be a specialization
of unicellular eucaryote feeding). Table 1
contains an alphabetical list of nematode
genera and their feeding types.

Feeding Habits of Soil Nematodes: Yeates et al. 319

TABLE 1. Feeding types in nematode genera. Numbers refer to the eight feeding types outlined in the
text; numbers in parenthesis indicate a tentative assignment. The main food source is given first. For genera
maintained for several generations under defined conditions, unqualified reference is made to that publication
by number; for genera where the publication number is preceded by "see," a more general (or less specific)
support for allocation to the feeding type is indicated; for genera where we have not obtained a direct
reference to feeding activity, the family name is given.
Genus

Ablechroiulus
Achromadora
Acontylus
Acrobeles
Acrobeloides
Acrobelophis
Acrolobus
Acromoldavicus
Acrostichu3
Actinca
Actinolaimus
Aetholaimus
Aglenchus
Alaimus
Allantonema
Allodorylaimu~
Allotrichodorus
Amphidelus
Amplimerlinius
Anaplectus
Anatonchus
Anguina
Anomyctus
Antarctylus
Aorolaimus
Aphanolaimus
Apha3matylenchus
Aphelenchoides
Aphelenchus
Aporcelaimellus
Aporcelaimium
Aporcelaimus
Apratylenchoides
Aprutides
Aquatides
Atylenchus
Aulolaimus
Axonchium
Basiria
Bastiania
Bathyodontus
Belonolaimus
Bicirronema
Bitylenchus
Boleodorus
Brevibucca
Brittonema
Bunonema
Bursaphelenchus
Bursilla
Butlerius
Cacopaurus
Cactodera
Caenorhabditis

Feeding type
3
(6)
lb
3
3
3
3
3
3
5, 8
5, 8
5
1e
3
7 or 2
8
ld
3
ld
3
5a
la or b
2?
1c
lc
3
lc
2 or lb, le or if
2 or le
5, 8
8
5, 8
1b
2
5
1d
3
1, 8?
le
3
3
1d
3
ld
le or 2(?)
3
5, 8
3
2
3
3, 5
la
la
3

Family or literature citation

Rhabditidae
Achromodoridae
(66)
(111,126)
(71,104,126)
Cephalobidae
Cephalobidae
Cephalobidae
Diplogasteridae; (88)
Actinolaimidae
(56); (see 31,100)
Nygolaimidae
Tylenchidae; (122)
(see 77)
(119)
Dorylaimidae
Trichodoridae; (see Decraemer in 76)
Alaimidae
(14,36); (see Anderson & Potter in 76)
(122)
(18); (see 100)
(22); (see Krall in 76)
(see Nickle & Hooper in 76)
Hoplolaiminae
Hoplolaiminae
Halaphanolaimidae
(34); (see Fortuner in 76)
(68); (see 122)
(17,23,40,65); (see 122)
(117,122); (see 31)
Aporcelaimidae
(114,122); (see 31,100)
Pratylenchidae
Aphlenchoididae
(10); (see 100)
Tylenchidae
Cryptonchidae
(see 100)
Tylenchidae
Bastianiidae
Bathyodontidae; (see 12)
(15)
Chambersiellidae
Tylenchorhynchidae; (see Anderson & Potter in 76)
Tylenchidae
Brevibuccidae
Actinolaimidae
(77,128)
(32,63); (see Nickle & Hooper in 76)
Rhabditidae
(28,89,101); (see 100)
Paratylenchidae; (112); (see Raski in 76)
(19,98); (see Baldwin & Mundo-Ocampo in 76)
(25,78,97); (see 107)

320 Journal of Nematology, Volume 25, No. 3, September 1993

TABLE 1.

Continued

Genus

Caloosia
Campydora
Carcharolaimus
Cephalenchus
Cephalobus
Ceratoplectus
Cervidellus
Chambersiella
Cheilorhabditis
Chiloplacus
Choanolaimus
Chondronema
Chromadorina
Chromadorita
Chronogaster
Chrysonemoides
Clarkus
Coarctadera
Cobbonchus
Coomansus
Coslenchu~
Crozpedonema
Criconema
Crieonemoides
Crossonema
Cruznema
Cryphodera
Cryptonchus
Curviditis
Cuticonema
Cuticularia
Cylindrolaimus
Daptonema
Deladenus
Demaniella
Deontolaimus
Desmodora
Desmolaimus
Desmoscolex
Dichromadora
Diphtherophora
Diplenteron
Diplogaster
Diplogasteritus
Diplogasteroides
Diploscapter
Discolaimium
Discolaimus
Ditylenchus
Dolichodorus
Dolichorhabditis
Dolichorhynchus
Domorganus
Dorydorella
Dorylaimellus
Dorylaimoides
Dorylaimus
Doryllium
Drilocephalobus

Feeding type
1d
8?
5
ld
3
3
3
3
3
3
5a
7, 2?
3, 6?
6
3
?
5a
3
5a
5a
le
3
ld
ld
ld
3
la
3
3
3
3
3
3, 4, 5 or 6
2
3
3
3, 6
3
3
3, 6?
2
3?
3, 5a, 6 or 8
3
3
3
5?
5
2, or lb
ld
3
1d
3
8
1, 2?
8?
8
2?
3

Family or literature citation

Hemicydiophorididae
Campydoridae
(28); (see 100)
(37,109); (see 122)
(81,88); (see 72,122)
Plectididae
(122)
Chambersiellidae
Rhabditidae
(118,122)
Choanolaimidae
(16)
Chromadoridae; (116)
(49)
Leptolaimidae
Chrysonematidae
(67,106); (see 101,130)
Rhabditidae; (see 107)
(see 100)
(31)
(as Aglenchus); (122)
Butonematidae
(see 26,93)
(see 26,93)
(see 26,93)
(108)
(see 8,60)
Cryptonchidae
Rhabditidae
Breviibuccidae
Rhabditidae
Diplopeltidae
(13)
(9,129); (see 62)
(88)
Leptolaimidae
Desmodoridae; (74)
Linhomoeidae
Desmoscolecidae
(13,120)
Diphtherophoridae; (see 4)
(64,125); (see 100)
(see 72,100)
(124)
Diplogasteroididae
(42,88)
(see 100)
(28,56); (see 31,100)
(55,122); (see Smrhan & Brezeski in 76)
(82,83,85); (see Smart & Nguyen in 76)
Rhabditidae
Dolichodoridae
Diplopehidae
Dorylaimidae
Belondiridae; (see 131)
Leptonchidae
(56); (see 77,100)
Leptonchidae
Osstellidae

Feeding Habits of Soil Nematodes: Yeates et al. 321

TABLE 1.

Continued

Genus

Durinema
Ecphyadophora
Ecumenicus
Elaphonema
Enchodelus
Epidorylaimus
Ereptonema
Ethmolaimus
Eucephalobus
Eudorylaimus
Eurystomina
Eumonhystera
Euteratocephalus
Fictor
Filenchus
Funaria
Fungiotonchium
Geocenamus
Geomonhystera
Glauxinema
Globodera
Goffartia
Goodeyus
Gracilacus
GranonchuIus
Haliplectus
Helicotylenchus
Hemicriconemoides
Hemicycliophora
Heterocephalobellus
Heterocephalobus
Heterodera
Heterorhabditis
Hexatylus
Hirschmanniella
Hoplolaimus
Hoplotylus
Howardula
Huntaphelenchoides
Iotonchus
Ironus
Isolaimium
Kirjanovia
Kochinema
Labronema
Laimaphelenchus
Laimydorus
Lelenchus
Leptolaimus
Leptonchus
Longidorella
Longidorus
Loofia
Macrotrophurus
Malenchus
Meloidodera
Meloidoderita
Meloidogyne
Merlinius

Feeding type
5
ld or 2(?)
8
3
8 or 6?
8
3
3, 6?
3
5, 8
(5)
3, 4
3
3, 5, 6
1e
2?
2
1d
3, 4
6
la
3
3
ld
5a
3
1c
ld
ld
3
3
la
7, 3
2
lb
1c
lb
7 or 2
2
5a
5a or 6
(3)
3
8
5, 8
5b, If, 2
8
1e
3
2
1d
1d
1d
Id
1e
la
la
la
ld

Family or literature citation

(see 77,100)
Ecphyadophoridae
Dorylaimidae
Elaphonematidae
Nordiidae
Dorylaimidae
Ereptonemidae
Ethmolaimidae
(122)
(29,44,117); (see 31,100)
Eurystominidae
Monhysteridae
Teratocephalidae
(88,103); (see 100)
Tylenchidae
Leptonchidae
Iontonchiidae (see 99)
Dolichodoridae
Monhysteridae
Neodiplogasteridae
(53); (see Baldwin & Mundo-Ocampo in 76)
Diplogasterididae
Cylindrocorporidae
Paratylenchidae; (see Raski in 76)
Mononchidae; (35)
Haliplectidae
(51); (see Fortuner in 76)
Criconematidae; (see 93)
Criconematidae; (see 93)
Cephalobidae
Cephalobidae
(see Baldwin & Mundo-Ocampo in 76)
(90); (see Wouts in 76)
(24); (see 72)
(7); (see Loof in 76)
Hoplolaimidae
Pratylenchidae
Allantonematidae
Aphelenchoididae
(67,100,101)
(47); (see 100)
Isolaimidae
Cephalobidae
Nordiidae
(28,30,88,122); (see 31,100)
(see 100)
(see 100)
Tylenchidae
(13)
(see 31)
Nordiidae
Longidoridae
Hemicycliophoridae
Dolichodoridae
Tylenchidae
(see Baldwin & Mundo-Ocampo in 76)
(see 76)
(see 27,96)
(see Anderson & Potter in 76)

3 2 2 Journal of Nematology, Volume 25, No. 3, September 1993

TABLE 1.

Continued

Genus

Mesocriconema
Mesodiplogaster
Mesodorylaimus
Mesorhabditis
Metacrobeles
Metadiplogaster
Metateratocephalus
Miconchus
Microdorylaimus
Microlaimus
Monhystera
Monhystrella
Monobutlerius
Mononchoides
Mononchulus
Mononchus
Monotrichodorus
Morulaimus
Mylonchulus
Myolaimus
Nacobbus
Nagelus
Namibinema
Neoactinolaimus
Neoaplectana
Neodiplogaster
Neopsilenchus
Neothada
Nothacrobeles
Nothotylenchus
Nullonchus
Nygolaimium
Nygolaimoides
Nygolaimus
Odontolaimus
Odontopharynx
Odontorhabditis
Ogma
Oionchus
Onchulus
Opisthodorylaimus
Orrina
OssteUa
Oxydirus
Panagrellus
Panagrobelium
Panagrobelus
Panagrocephalus
Panagrolaimus
Paracrobeles
Paractinolaimus
Paracyatholaimus
Parahadronchus
Paralongidorus
Paramphidelus
Paraphanolaimus
Paraphelenchus
Paraplectonema
Parawtylenchus

Feeding type
ld
3, 5
8
3
3
3
3
5a
8
3
3, (4)
3, 4
3
3, 5a
3
5a
ld
ld
5a
3
la
1d
3
5, 8
7, 3
3, 4 or 5
1e
le or 2(?)
3
2
5a
5
5
5
3 or 6?
3, 5a
3
ld
3
5a or 6
8
la or b
3
1, 8?
3
3
3
3
3
3
5
6?
5a
ld
3
3
2
3
lc

Family or literature citation

(see 93)
(102,103); (see 100)
(29); (see 31,95,100)
( 107,122,126)
Cephalobidae
Diplogasteridae
Teratocephalidae
(see 100)
Nordiidae
Microlaimidae
(122); (see 77,115)
(13)
Diplogasteridae
(89); (see 100)
(see 31)
(38,88); (see 31,100)
Trichodoridae; (see Decraemer in 76)
Belonolaimidae; (see Smart & Nyguen in 76)
(48,67); (see 31,100)
Myolaimidae
(see Jatala in 76)
Dolichodoridae
Cephalobidae
(101); (see 100)
(see 76)
Neodiplogasteridae
Tylenchidae
Tylenchidae
Cephalobidae
Anguinidae
(see 100)
(see 100)
Nygolaimidae
(113,122); (see 31,100)
Odontolaimidae
Odontopharyngidae
Odontorhabditidae
(see 93)
Mononchulidae
Onchulidae
Dorylaimidae
(see Krall in 76)
Osstellidae
Belondiridae
(21)
Panagrolaimidae
Panagrolaimidae
Cephalobidae
(88,126)
Cephalobidae
(46) (see 100)
(13)
(see 100)
(see 11,54)
Alaimidae
Halaphanolaimidae
(110); (see Nickle & Hooper in 76)
Leptolaimidae
(see Fortuner in 76)

F e e d i n g H a b i t s o f Soil N e m a t o d e s : Yeates et al.

TABLE 1.

Continued

Genus

Paratrichodorus
Paratripyla
Paratrophurus
Parat2flenchus
Paravulvus
Paraxonchium
Pareudiplogaster
ParoigoIaimella
Paurodontus
Pellioditis
Pelodera
Peltamigratus
Phasmarhabditis
Placodira
Plectonchus
Plectus
Pleurotylenchus
Pratylenchoides
Pratylenchus
Prionchulus
Prismatolaimus
Pristionchus
Prochromadora
Prodesmodora
Prodorylaimium
Prodorylaimus
Proleptonchus
Protocylindrocorpus
Protorhabditis
Pseudac~vbeles
Pseudhalenchus
Pseudoaulolaimus
Psilenchus
Pterotylenchus
Pterygorhabditis
Punctodera
Punctodora
Pungentus
Quinisulcius
Radopholus
Rhabditis
Rhabditoides
Rhabditophanes
Rhabdolaimus
Rhabdontolaimus
Rhadinaphelenchus
Rhodolaimus
Rotylenchulus
Rotylenchus
Scottnema
ScuteUonema
Scutylenchus
Sectonema
Seinura
Seleborca
Sphaerolaimus
Sphaeronema
Sphae~.ularia
Sporonchulus

Feeding type
ld
5a
ld
ld
5?
5, 8
6, 3
3
2, 7
3
3
lc
3
3
3
3
1d
lb
lb
5a
3?
3, 5a
3, 6?
3
8
8
8?
3
3
3
2
3
1e
la or b
3
la
3, 6?
l d, 5, 8
ld
lb
3
3
3
3
3
lb
3
1a
lc
3
lc
ld
5, 8
5b
3
5a
la
7, 2
5a

Familyor literature citation

Trichodoridae; (see Decraemer in 76)
Tripylidae
Dolichodoridae
(94,122); (see Raski in 76)
Nygolaimidae
Aporcelaimidae
(13)
(88)
(as Neotylenchus); (40)
(107)
(88,107,122)
(see 76)
(107)
Cephalobidae
Brevibuccidae
(77,88,122)
Tyiodoridae
(see Loof in 76)
(see Loof in 76)
(5,61,69); (see I00)
(see 77)
Neodiplogasteridae
Chromadoridae
Desmodoridae
Dorylaimidae
Dorylaimidae
Leptonchidae
Cylindrocorporidae (88)
Rhabditidae
Cephalobidae
Anguinidae
Cryptonchidae
Psilenchidae
Anguinidae
Pterygorhabditidae
(see Baldwin & Mundo-Ocampo in 76)
Chromadoridae
(see 31,100)
(see 76)
(see Loof in 76)
(88,102,107,122)
Rhabditidae
Alloionematidae
(see 77)
(88)
(see Nickie & Hooper in 76)
Bunonematidae
(57)
Hoplolaiminae; (see Fortuner in 76)
Cephalobidae; (80)
Hoplolaiminae; (see Fortuner in 76)
Dolichodoridae
(see 100)
(28,41,43,101,123); (see 100)
Acrobelidae
Sphaerolaimidae
(see Raski in 76)
(91 )
(see 100)

323

324

Journal of Nematology, Volume 25, No. 3, September 1993

TABL~ 1.

Continued

Genus

StegeUeta
Stegelletina
Steinernema
Stomachoglossa
Subanguina
Sulphuretylenchus
Synonchium
Telotylenchus
Tenunemellus
Teratocephal~s
Teratolobus
Teratorhabditis
Theristus
Thonus
Thornenema
Thornia
Tobrilus
Torumanawa
Tricephalobus
Trichodorus
Tripius
T,~pyla
Trischistoma
Trophonema
Trophotylenchus
Trophurus
Turbatrix
Tylencholaimellus
Tylencholaimus
Tylenchorhynchus
Tyl~nchulus
Tylenchus
Tylocephalus
Tylodorus
Tylolaimophorus
Tylopharynx
Verutus
Westindicus
Wilsonema
Xiphinema
Ypsylonellus
Zeldia
Zygotylenchus

Feeding type
3
3
7, 3
(5)
la or b
7 or 2
5
1c
ld or 2(?)
3
3
3
3, 4 or 6
5, 8
8
8?
5a or 6
8
3
ld
7, 2
5a
5a
la
1a
1d
3
2
2
ld
la
If, 2?
3
1d
2
3
la
(5)
3
1d
3
3
lb

Order M onhysterida
Linhomoeidae: Desmolaimus
(3) bacterial feeding
Monhysteridae: Monhystera, Geomonhystera
3 bacterial feeding
4 substrate ingestion
Sphaerolaimidae: Sphae~vlaimus
5 predacious
Xyalidae: Theristus, Daptonema
3 bacterial feeding
4 substrate ingestion

Family or literature citation

Cephalobidae
Cephalobidae
(see 76)
(see 100)
(see 76)
(see 76)
(see 100)
Dolichodoridae
Ecphyadorphoridae
(see 77)
Cephalobidae
Rhabditidae
(see 77)
(31)
Dorylaimidae
(see 100)
(see 100)
Aporcelaimidae
Panagrolaimidae
Trichodoridae; (see Decraemer in 76)
(see 76)
(77,100)
Tripylidae; (see 100)
Tylenchulidae; (see Raski in 76)
Tylenchulidae
Dolichodoridae
(70,89)
(122)
(29,105,122); (see 31)
(see Anderson & Potter in 76)
(see Raski in 76)
Tylenchidae; (122)
(see 77)
Tylodoridae
Diphtherophoridae
Tylopharyngidae
(see Baldwin & Mundo-Ocarnpo in 76)
(see 100)
(77,122)
Longidoridae
Acrobelidae
(126)
(see Loof in 76)

5 predacious
6 algal feeding
Order Desmoscolecida
Desmoscolecidae: Desmoscolex
(3) bacterial feeding
Order Araeolaimida
All terrestrial forms are apparently
type 3.
Diplopeltidae: Cylindrolaimus, Domorganus

Feeding Habits of Soil Nematodes: Yeates et al. 325

Craspedonema
3 bacterial feeding
Cephalobidae: Cephalobus, Heterocephalobus, Eucephalobus
3 bacterial feeding
Chambersiellidae: Chambersiella, Bicirronema
3 bacterial feeding
Cylindrocorporidae: Protocylindrocorpus,
Goodeyus
(3) bacterial feeding
Diploscapteridae: Diploscapter
3 bacterial feeding
Elaphonematidae: Elaphonema
3 bacterial feeding
Myolaimidae: Myolaimus
3 bacterial feeding
Order Chromadorida
Odontopharyngidae: Odontopharynx
3 bacterial feeding
Achromadoridae: Achromadora
5 predatory
6 algal feeding
O d o n t o r h a b d i t i d a e : Odontorhabditis,
Choanolaimidae: Choanolaimus, Synonchium
Cheilorhabditis
5 predacious
3 bacterial feeding
Cyatholaimidae: P aracyatholaimus
Osstellidae: Osstella
6 algal feeding
3 bacterial feeding
Desmodoridae: Desmodora
Panagrolaimidae: Panagrolaimus, Panagrel(3) bacterial feeding
lus, Turbatrix
6 algal feeding
3 bacterial feeding (many are insect asEthmolaimidae: Ethmolaimus
sociates)
3 bacterial feeding
Pterygorhabditidae: Pterygorhabditis
6 algal feeding
3 bacterial feeding
Hypodontolaimidae: Chromadorita
Rhabditidae: Rhabditis, Mesorhabditis, Pelo6 algal feeding
dera
Microlaimidae: M icrolaimus
3 bacterial feeding (many are associated
3 bacterial feeding
with earthworms, arthropods, or ver6 algal feeding
tebrates)
Steinernematidae:
Steinernema (= NeoOrder Rhabditida
aplectana), Heterorhabditis
7 entomogenous
Members of this order that occur in soil
3 bacterial feeding in free-living stages
are basically type 3 (bacterial feeding).
Acrobelidae: Acrobeles, Cervidellus, Acrobe- Teratocephalidae: Teratocephalus, Euteratocephalus
loides
3 bacterial feeding
3 bacterial feeding
Alloionematidae: AUoionema, Rhabditophanes
3 bacterial feeding (soil-dwelling stages Order Diplogasterida
only)
The basic habit of these is apparently
7 insect parasites
type 3 (bacterial feeding).
Brevibuccidae: Brevibucca, Cuticonema
Diplogasteridae: Diplogaster, Butlerius
(3) bacterial feeding; insect associates
Bunonematidae: Bunonema, Rhodolaimus,
These may use insects for dispersal.

(3) bacterial feeding

7 Domorganus (sometimes associated
with earthworms)
Halaphanolaimidae: Aphanolaimus
(3) bacterial feeding
Haliplectidae: Haliplectus
(3) bacterial feeding
Leptolaimidae: Leptolaimus, Chronogaster
3 bacterial feeding
Plectidae: Plectus, Anaplectus
3 bacterial feeding (Plectus)
Rhabdolaimidae: Rhabdolaimus
3 bacterial feeding
Wilsonematidae: Wilsonema, Tylocephalus,
Ereptonema
3 bacterial feeding

326 Journal of Nematology, Volume 25, No. 3, September 1993

3 bacterial feeding
5 predacious
Diplogasteroididae: Diplogasteroides, Gof-

fartia
3 bacterial feeding
Neodiplogasteridae: Neodiplogaster, Diplen-

teron, Fictor, Pristionchus

These may use insects for dispersal.
3 bacterial feeding
4 substrate ingestion
5 predacious
6 in Pareudiplogaster
Tylopharyngidae: Tylopharynx
3 bacterial feeding

Order Tylenchida
Suborder Tylenchina:
Allantonematidae: AUantonema, Howardula, Sulphuretylenchus
7 insect parasites
? alternate generation possibly feeds in
soil
Anguinidae: Anguina, Ditylenchus, Nothoty-

lenchus
1 parasites of aerial parts of plants
2 fungal feeders
Atylenchidae: Atylenchus, Eu~ylenchus
(1) root feeding
Caloosiidae: Caloosia
1 ectoparasites of roots
Chondronematidae: Chondronema
7 juveniles parasitic in insects
? adults free-living, possible nonfeeding
Criconematidae:
Criconema, Ogma,

Hoplolaimidae: Hoplolaimus, Rotylenchus,

Helicotylenchus
1 ectoparasitic or semi-endoparasitic on
roots
Iotonchiidae: Iotonchium
2 hyphal feeding in some stages
7 insect parasites
Meloidogynidae: Meloidogyne
1 obligate endoparasites; female saccate
Nacobbidae: Nacobbus
1 gall-forming root endoparasites; saccate females
Neotylenchidae: Fergusobia, Deladenus
7 insect parasites with alternate generation being
2 hyphal-feeding or
1 plant feeding
Paratylenchidae: Paratylenchus, Gracilacus
1 ectoparasites of roots; male often
without stylet
Paurodontidae: Paurodontus
2 hyphal-feeding generation and
7 probable insect-parasitic generation
Pratylenchidae: Pratylenchus, Radopholus
1 migratory endoparasites of roots
Psilenchidae: Psilenchus
1 associates of lower plants (other than
fungi); feed on root hairs and epidermal cells
Rotylenchulidae: Rotylenchulus
1 e c t o p a r a s i t i c on roots o f h i g h e r
plants; female saccate
Sphaeronematidae: Sphaeronema, Meloido-

derita

1 females sedentary ecto- or endoparasites of roots

1 ectoparasites of roots; males often Sphaerularidae: Sphaerularia, Tripius
2 hyphal-feeding generation may occur
without stylets
7 females parasitic in insect haemocoel
Dolichodoridae: Dolichodorus, TylenchorhynSychnotylenchidae: Sychnotylenchus, Neodichus, Merlinius, Belonolaimus
tylenchus
1 migratory ectoparasites of roots
7 associates of bark beetles
E c p h y a d o p h o r i d a e : Ecphyadophora, TeT y l e n c h i d a e : Tylenchus, Filenchus, Bonunemellus
leodorus
(1) root feeding
1 associates of algae, mosses, lichens,
Hemicycliophoridae: Hemicycliophora
and plant roots
1 ectoparasites of roots; males often
Tylenchulidae:
Tylenchulus, Trophonema
without stylet
1
ectoparasites
of roots; female saccate
Heteroderidae: Heterodera, Globodera, Crybut
usually
outside
root
phodera
1 obligate endoparasites; female cuticle Tylodoridae: Tylodorus, Cephalenchus
1 root feeding
produces resistant cyst

CriconemeUa, Hemicriconemoides

Feeding Habits of Soil Nematodes: Yeates et al. 327

Suborder Aphelenchina:
Aphelenchidae: Aphelenchus
2 hyphal feeding; sometimes feed on
fungi in diseased plants
Aphelenchoididae: Aphelenchoides,
Bursaphelenchus, Anomyctus,
Rhadinaphelenchus
1 plant feeding in a minority of forms
(Aphelenchoides may feed on fungi but
also on aerial parts of plants
including ferns)
2 hyphal feeding in terrestrial species;
may be insect associated
5 predacious (Laimaphelenchus)
6 algal feeding (Laimaphelenchus)
Paraphelenchidae: P araphelenchus
2 hyphal feeders
Seinuridae: Seinura
5 predacious

Order Enoplida
Alaimidae: Alaimus, Amphidelus
3 bacterial feeding
Cryptonchidae: Cryptonchus, Aulolahnus
(3) bacterial feeding (Aulolaimus)
Ironidae: Ironus
5 predacious
Oncholaimidae: Bastiania
(3) bacterial feeding
Onchulidae: Onchulus, Stenonchulus
(3) bacterial feeding
(5) predacious
(8) omnivorous
Prismatolaimidae: Prismatolaimus
3 bacterial feeding
Tobrilidae: T obrilus
(5) predacious
(6) algal feeding
Tripylidae: Tripyla, Trischistoma
(5) predacious

Order Mononchida
Most common forms are type 5 (predacious) but some are type 3 (bacterial feeding).
Anatonchidae: Anatonchus
5 predacious; prey may be ingested
whole

Bathyodontidae: Bathyodontus, Mirolaimus

(3) bacterial feeding
Mononchidae: Mononchus
5 predacious; prey not ingested whole
(3) may also feed on bacteria
Mononchulidae: Mononchulus, Oionchus
(3) bacterial feeding

Order Dorylaimida
Actinolaimidae: Actinolaimus, Brittonema,
Actinca
5 predacious
(8) omnivorous
Aporcelaimidae: Aporcelaimus, Paraxonchium, Sectonema
5 predacious
(8) omnivorous
Belondiridae: Axonchium, Dorylaimellus, Oxydirus
(1) plant feeding
Diphtherophoridae: Diphtherophora, Triolaimophorus
(2) hyphal feeding
Discolaimidae: Discolaimus, Discolaimium
(5) predacious
Dorylaimidae: Dorylaimus, Laimydorus, Mesodorylaimus, Prodorylaimus, Thornenema,
Thor~?~s

(8) omnivorous
5 predacious (Labronema)
Leptonchidae: Leptonchus, Tylencholaimellus, Do*~yllium
(2) hyphal feeding
Longidoridae: Longidorus, Xiphinema
1 plant feeding; may transmit plant viruses
Nordiidae: Eudorylaimus
(8) omnivorous (possibly all genera are
plant feeding)
(1) plant feeding
Nygolaimidae: Nygolaimus
5 predacious, especially on enchytraeids
Trichodoridae: Trichodorus, Paratrichodorus
1 plant feeding; may transmit plant viruses

Order lsolaimida
Isolaimidae: Isolaimium
(3) bacterial feeding

328 Journal of Nematology, Volume 25, No. 3, September 1993

Order Stichosomida
Mermithidae: Mermis, Agamomermis
7 parasites of arthropods and other invertebrates; postparasitic stages may
occur in soil
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