NOAA Technical Report NMFS 77

Illustrated Key to the Genera of

Free-Living Marine Nematodes

of the Order Enoplida

Edwin J. Keppner

Armen C. Tarjan

July 1989

U.S. Department of Commerce
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34. Additions to a revision of the shark genus Carcharhinus: Synonymy of Aprionodon and Hypoprion, and description of a new species ofCarcharhinus (Carcharhinidae),

by J. A. F. Garrick. November 1985,26 p.

35. Synoptic review ofthe literature on the Southern oyster drill Thais haemas/oma

floridona, by Philip A. Butler. November 1985, 9 p.

36. An egg production method for estimating spawning biomass of pelagic fish:

Application to the northern anchovy, Engraulis mordax, by Reuben Lasker (editor).

December 1985, 99 p.

37. A histopathologic evaluation of gross lesions excised from commercially important North Atlantic marine fishes, by Robert A. Murchelano, Linda DespresPatanjo, and John Ziskowski. March 1986, 14 p.

38. Fishery atlas of the northwestern Hawaiian Islands, by Richard N. Uchida and

James H. Uchiyama (editors). September 1986, 142 p.

39. Survey of fish protective facilities at water withdrawal sites on the Snake and

Columbia Rivers, by George A. Swan, Tommy G. Withrow, and Donn L. Park. April

1986, 34 p.

40. Potential impact of ocean thermal energy conversion (OTEC) on fisheries, by

Edward P. Myers, Donald E. Hoss, Walter M. Matsumoto, David S. Peters, Michael

P. Seki, Richard N. Uchida, John D. Ditmars, and Robert A. Paddock. June 1986,

33 p.

41. A stationary visual census technique for quantitatively assessing community structure of coral reef fishes, by James A. Bohnsack and Scon P. Bannerot. July 1986, 15 p.

42. Effects oftemperature on the biology ofthe northern shrimp, Pa:uJalus borealis,

in the Gulf of Maine, by Spencer Apollonio, David K. Stevenson, and Earl E. Dunton,

Jr. September 1986, 22 p.

43. Environment and resources of seamounts in the North Pacific, by Richard N.

Uchida, Sigeiti Hayasi, and George W. Boehlert (editors). September 1986, 105 p.

44. Synopsis of biological data on the porgies, Calamus arc/ifrons and C. proridens

(Pisces: Sparidae), by George H. Darcy. September 1986, 19 p.

45. Meristic variation in Sebas/es (Scorpaenidae), with an analysis of character

association and bilateral panern and their significance in species separation, by Lochai Chen. September 1986, 17 p.

46. Distribution and relative abundance of pelagic nonsalmonid nekton off Oregon

and Washington 1979-84, by Richard D. Brodeur and William G. Pearcy. December

1986, 85 p.

47. Reproduction, maturation, and seed production of cultured species: Proceedings

of the twelfth U.S.-Japan meeting on aquaculture, Baton Rouge, Louisiana, October

25-29, 1983, by Carl J. Sindermann (editor). February 1987, 73 p.

48. Widow rockfish: Proceedings of a workshop, Tiburon, California, December

11-12, 1980, by William H. Lenan and Donald R. Gunderson (editors). January 1987,

57 p.

49. Reproduction, movements, and population dynamics of the southern kingfish,

Mentici"hus americanus, in the northwestern Gulfof Mexico, by Stephen M. Harding

and Mark E. Chinenden, Jr. March 1987, 21 p.

50. Preparation of acetate peels of valves from the ocean quahog, Arc/ica islandica,

for age determinations, by John W. Ropes. March 1987, 5 p.

51. Status, biology, and ecology of fur seals. Proceedings of an international

workshop, Cambridge, England, 23-27 April 1984, by John P. Croxall and Roger

L. Gentry (editors). June 1987,212 p.

52. Limited access alternatives for the Pacific groundfish fishery, by Daniel D.

Huppert (editor). May 1987, 45 p.

53. Ecology of east Florida sea turtles: Proceedings of the Cape Canaveral, Florida,

sea turtle workshop, Miami, Florida, February 26-27, 1985, by Wayne N. Witzell

(convener and editor). May 1987, 80 p.

54. Proximate and fany acid composition of 40 southeastern U.S. fmfish species,

by Janet A. Gooch, Malcolm B. Hale, Thomas Brown, Jr., James C. Bonnet, Cheryl

G. Brand, and Lloyd W. Reiger. June 1987,23 p.

55. Proximate composition, energy, fany acid, sodium, and cholesterol content of

fmfish, shellfish, and their products, by Judith Krzynowek and Jenny Murphy. July

1987, 53 p.

56. Some aspects of the ecology of the leatherback turtle Dennochelys coriacea at

Laguna Jolova, Costa Rica, by Harold F. Hirth and Larry H. Ogren. July 1987, 14 p.

57. Food habits and dietary vari~bility of pelagic nekton off Oregon and Washington,

1979-1984, by Richard D. Brodeur, Harriet V. Lorz, and William G. Pearcy. July

1987, 32 p.

58. Stock assessment of the Gulf of Menhaden, Brevoonia pa/ronus, fishery, by

Douglas S. Vaugh~n. September 1987, 18 p.

59. Atlantic Menhaden, Brevoonia tyrannus, purse seine fishery, 1972-84, with a

brief discussion of age and size composition of the landings, by Joseph W. Smith,

William R. Nicholson, Douglas S. Vaughan, Donnie L. Dudley, and Ethel A. Hall.

September 1987, 23 p.

60. Gulf Menhaden, Brevoortia po/ronus, purse seine fishery, 1974-85, with a brief

discussion of age and size composition of the landings, by Joseph W. Smith, Eldon

J. Levi, Douglas S. Vaughan, and Ethen A. Hall. December 1987, 8 p.

61. Manual for starch gel electrophoresis: A method for the detection of genetic variation, by Paul B. Aebersold, Gary A. Winans, David J. Teel, George B. Milner, and

Fred M. Uner. December 1987, 19 p.

62. Fishery publication index, 1980-85; Technical memoradum index, 1972-85, by

Cynthia S. Martin, Shelley E. Arenas, Jacki A. Guffey, and Joni M. Packard.

December 1987, 149 p.

63. Stock assessment of the Atlantic Menhaden, Brevoonia tyrannus, fishery, by

Douglas S. Vaughan and Joseph W. Smith. January 1988, 18 p.

64. illustrated key to penaeoid shrimps of commerce in the Americas, by Isabel Perez

Farfante. April 1988, 32 p.

65. History of whaling in and near North Carolina, by RandaI\ R. Reeves and Edward

Mitchell. March 1988, 28 p.

66. Atlas and zoogeography of common fishes in the Bering Sea and Northeastern

Pacific, by M. James Allen and Gary B. Smith. April 1988, 151 p.
NOAA Technical Report NMFS 77

Illustrated Key to the Genera of

Free-Living Marine Nematodes

of the Order Enoplida

Edwin J. Keppner

Armen C. Tarjan

July 1989

u.s. DEPARTMENT OF COMMERCE

Robert Mosbacher, Secretary

National Oceanic and Atmospheric Administration

William E. Evans, Under Secretary for Oceans and Atmosphere

National Marine Fisheries Service

James Brennan, Assistant Administrator for Fisheries
The National Marine Fisheries Service (NMFS) does not approve. recommend

or endorse any proprietary product or proprietary material mentioned in this

publication. No reference shall be made to NMFS. or to this publication furnished

by NMFS, in any advertising or sales promotion which would indicate or imply

that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause

directly or indirectly the advertised product to be used or purchased because of

this NMFS publication.

11
CONTENTS

Introduction 1

Acknowledgments 2

Citations 2

Use of the key 2

Flow chart of taxa included in key 3

Key to genera of the Enoplida 6

Illustration sources 24

References for illustration sources 25

COVER: Drawing of Metoncholaimus pristiurus by N.A. Cobb, 1. Wash. Acad.

Sci. 22(12):346.

iii

mustrated Key to the

Genera of Free-Living

Marine Nematodes of

the Order Enoplidal

EDWIN J. KEPPNER

Habitat Conservation Division

Southeast Regional Office

National Marine Fisheries Service, NOAA

3500 Delwood Beach Road

Panama City, FL 32407

ARMEN C. TARJAN

Department of Entomology and Nematology

University of Florida

Gainesville, FL 32611-0611

Abstract

A pictorial key to US genera of free-living marine nematodes

in the order Enoplida is presented. Specific morphological and

anatomical features are iUustrated to facilitate use of the key.

The purpose ofthis work is to provide a single key to the genera

of enoplid nematodes to facilitate identification ofthese organisms by nematologists and marine biologists working with

meiofauna.

'Florida Agricultural Experiment Station Journal Series No. 8673.

Introduction _

Free-living marine nematodes are a significant component

of estuarine and marine benthic communities and are considered a part ofthe meiofauna. Nematodes are often the most

numerically abundant organisms in estuarine and marine

sediments, and their species diversity within a given estuarine

or marine benthic habitat is usually quite large. Free-living

marine nematodes recently have been examined as possible

indicators of pollution in estuarine and marine environments.

In North America, studies to determine the ecological role

of nematodes and their response to pollution have been difficult. This is due to the instability of the taxonomy of these

organisms and the absence of keys to several of the genera

within each order.

Hope and Murphy (1972), Gerlach and Riemann (1974),

Andrassy (1976), Lorenzen (1981), and Inglis (1983) have

published classifications of free-living marine nematodes.

Major differences exist between these classifications which

can lead to confusion in the identification of these animals.

Much of the morphology of free-living marine nematodes

is not well understood. As a result, homologous structures

are difficult to identify. Existing classifications and keys to

the taxa of free-living marine nematodes reflect these

difficulties.

The systematics and general ecology of the free-living

marine nematodes have been summarized by Heip et al.

(1982, 1985). Tarjan (1980) provided an illustrated key to

a large number ofthe genera of free-living marine nematodes

in each order with numerous drawings of representative

species ofthese genera. Platt and Warwick (1983) developed

a pictorial key to the genera offree-living marine nematodes

and provided drawings and descriptions of the species of

Enoplida found in British waters. The use of these keys can

provide a sound basis for the identification of the genera of

marine nematodes. However, some ofthe genera of enoplid

nematodes are absent from both keys.

It was decided that the key provided by Tarjan (1980)

should be expanded to include the most commonly accepted

genera of the Enoplida. The goal was not to present a new

classification ofthe Enoplida but to provide a means ofidentification ofthe genera included in the key. The original key

of Tarjan (1980) was based on the classification given by

Andrassy (1976). The key presented herein is a compilation

of the published literature and relies heavily on classifications ofthe Enoplida given by Andrassy (1976) and Gerlach

and Riemann (1974). This key is not intended to reflect

phylogenetic relationships. The key includes the Tripyloidina

in the Enoplida simply so that those free-living marine

nematodes with segmented cephalic setae are all in one place.

Gerlach and Riemann (1974) placed them in the Enoplida

but Andrassy (1976) did not. The genus Polygastrophora

De Man, 1922 was removed from the Enchelidiidae and

placed with the Belbollidae for ease of identification because

both genera, Polygastrophora and Bolbellia (Cobb 1920)

Gerlach and Riemann 1974, possess a multibulbar esophagus,

and some species, such as Polygastrophora edax Wieser
and Hopper, 1967, do not have sexual dimorphism in the

stoma.

The key presented herein includes 118 genera of the

Enoplida and follows the pattern given in Tarjan (1980). The

decision as to which genera should be included in the key

was difficult and in some cases arbitrary. However, an attempt was made to include those genera which have been

reported primarily from estuarine and marine environments

rather than primarily from freshwater, those that contain

more than one species, and those that have been collected

by the authors. This key includes about 70% of the genera

listed for the Enoplida, and the genera included account for

about 90% of the known species of free-living marine

enoplids. Platt and Warwick (1983) provide a brief and concise discussion of the morphology of free-living marine

nematodes and enoplids that may be consulted as an aid to

using this key.

A key such as this inevitably perpetuates errors, misinterpretations, and inadequate descriptions of taxa that exist in

the literature. Accordingly, it is hoped that the present work

will stimulate others to correct such errors where they exist

and refine the definition of taxa where necessary.

Acknowledgments _

We express our sincere appreciation to those persons.who

reviewed the key and provided many helpful suggestIOns.

Comments and recommendations were received from Dr.

Istvan Andrassy, Eotvos Lorand University, Hungary; Dr.

W. Duane Hope, U.S. National Museum of Natural History;

Dr. Preben Jensen, Denmark; and Dr. Howard Platt, British

Museum (Natural History).

We are grateful to Lisa A. Hoover who devoted much time

to testing the key using specimens of marine nematodes co.l.

lected by her and the senior author. Her efforts resulted In

clarification of several potential problem areas.

Citations _

Andrassy, I.

1976 Evolution as the basis for the systematization of nematodes.

Akademiai Kaido, Budapest, 288 p.

Gerlach, S.A., and F. Riemann

1974 The Bremerhaven checklist of aquatic nematodes. Part 2.

Ver6eff. Inst. Meeresforsch. Bremerhaven, SuppI. 4, p. 405-736.

Heip, C., M. Vincx, N. Smol, and G. Vranken

1982 The systematics and ecology of free-living marine nematodes.

HelminthoI. Abstr. Ser. B. 51: 1-31.

Heip, C., M. Vincx, and G. Vranken .

1985 The ecology of marine nematodes. Oceanogr. Mar. BIOI. Ann.

Rev. 23:399-489.

Hope, W.D., and D.G. Murphy .

1972 A taxonomic heirarchy and checklist of the genera and higher

taxa of marine nematodes. Smithson. Contrib. Zool. 136: 1-101.

Inglis, W.G.

1983 An outline classification of the Phylum Nematoda. Aust. J.

Zool. 31:243-255.

2

Lorenzen, S.

1981 Entwurf eines phylogenetischen systems der freilebenden Nernatoden. Verafl'. Inst. Meeresforsch. Bremerhaven, Suppl. 7, p. 1-472.

Platt, H.M., and R.M. Warwick

1983 Free-living marine nematodes. Part 1. British Enoplids. Cambridge Univ. Press, Cambridge, 307 p.

Tarjan, A.C.

1980 An illustrated guide to the marine nematodes. Inst. Food Agric.

Sci., Univ. Florida, Gainesville, 135 p.

Use of the key _

Couplet numbers down the far-right margin of the page refer

to the location of related lower-ranked taxa. When one arrives at a family or subfamily name on the right side, rather

than another couplet, the genera within that taxon immediately follow in couplets designated by lower-case letters

on the left. For example, couplet 5(4)B ends with the family

Thoracostomatidae. The genera included in that family are

in couplets "a" and "b" immediately following. The parenthetical portion of the couplet number [e.g., (4) in 5(4)B]

indicates the referent couplet, allowing one to work backward, as well as forward, through the key. Numbers in parentheses within the descriptions refer to figures.

The following explanations of terms are provided to aid

in avoiding confusion, particularly at the beginning of the

key. The term stoma is used throughout to designate

modification of the anterior end for feeding. Stoma refers

to both the mouth opening that is present in all nematodes

and the buccal cavity which can be well developed or greatly

reduced. The esophagus of marine nematodes is, in general,

either muscular (outer border smooth and few nuclei and

vesicles visible within esophageal tissue) or vesiculatecellulate (outer border lobed or irregular and many nuclei

and vesicles visible). In some taxa such as the Phanodermatidae, the anterior portion of the esophagus is muscular

and the posterior part that comprises most of the esophagus

is vesiculate-cellulate. These taxa are considered to have a

vesiculate-cellulate esophagus. The anterior portion of the

esophagus in some free-living marine nematodes surrounds

the stoma and attaches to the cuticle to form a cephalic capsule. The thickness of the cephalic capsule and the extent

of contact with the cuticle varies. It is imperative to use the

illustrations cited in the couplets in deciding which statements

are appropriate.

Illustrations used in the key were redrawn from published

references or are original drawings by the senior author. In

some cases, portions of the anatomy shown on the drawings

are labelled or emphasized with ink; others have been deleted

to better portray the structures of interest. Credit is given

for each illustration used. Since this key is proposed for nonprofit educational purposes, it is not considered to be an

infringement of the United States Revised Copywrite Law.
Flow chart of taxa included in key _

Subfamily

'"I---------1-----------.1

Enoplinae Enoploidinae Rhabdodemaniinae

I I I

Genus Genus Genus

I I I

1

Enoplina

I

I

Enoplidae

I

Phanodermatidae

I

Genus

I

1

Tripyloidina

Thoracostomopsis

I

Thoracostomopsidae

I

Genus

I

1

Leptosomatoidea

I

Tripylina

I

Superfamily

I

Phylum Nematoda

I

Class

Enoplia

I

Order

Enoplida

I

Suborder

I

Crenopharynx

Dayellus

Klugea

MicoletzkyiIJ

Phanoderma

Phanodermella

Phanodennopsis

1

Enoploidea

Family

I

I

Oncholaimina

Chaetonema

Rhabdodemania

Trileptium

Africanthion

Epacanthion

Enoploides

Enoplolaimus

FilipjeviIJ

Gairleanema

Mesacanthion

Mesacanthoides

Metenoploides

Oxyonchus

ParasaveljeviIJ

Paramesacanthion

Parenoplus

SaveljeviIJ

Enoplus

3
Leptosomatoidea

I

Family

1---------------------,1

Leptosomatidae Thoracostomatidae

I I

Subfamily Genus

'I-----------1------------" I Deontostoma

Antic1minae LePtosoratinae TriOdontrlaiminae Pseudocel/a

Thoracostoma

Genus Genus Genus

I I I

Anticoma Corythostoma Jaegerskioeldia

Anticomopsis Cylicolaimus Triodonto/aimus

Barbonema Leptosomatides

Odontanticoma Leptosomatina

Parabarbonema Leptosomatum

Paranticoma Leptosomel/a

Syringonomus Macronchus

TubolaimeUa Platycoma

Platycomopsis

Synonchus

1

Oncholaiminae

I

Genus

I

Adoncholaimus

Filoncholaimus

Kreisoncholaimus

Metaparoncholaimus

Metoncholaimoides

Metoncholaimus

Meyersia

Oncholaimium

Oncho/aimus

Pontonema

Prooncholaimus

Pseudoncholaimus

Wiesoncho/aimus

Pelagonematinae

I

Genus

I

Anoplostoma

Brodybucca

lilium

Pe/agonema

Pe/agonemeUa Subfamily

rl--------1------------,1

Thoonchinae Eurystomininae Pareurystomininae

I I I

Genus Genus Genus

I I I

DitlevseneUa Eurystomina Batheurystomina

Thoonchus Ger/achystomina Pareurystomina

Ledovitia

Oncholaimina

I

Superfamily

.....-------------.--- 1 --------,

1 I I

Pelagonematoidea Enchelidoidea Oncholaimoidea

I FallY FallY

FaInJ10IY I I , onChOI~1imidae

Bolbellidae Enchelidiidae Eurystominidae

I I

M ...·rd

Subfamily

.. .o:r E::C~ I o-:~~ 2~

I

Polygastrophora I I . Genus Genus Genus

I 1 I I

Curvolaiminae Calyptronema Octonchus Krampia

I ConistomeUa Mononcho/aimus

Genus Symplocostoma Oncholaimelloides I Symplocostomel/a Oncholaimellus

Curvolaimus Oncholaimoides

Viscosia

4
I

Halanonchinae

I

Genus

I

Halanonchus

1

Trefusiidae

I

Subfamily

I I

Trefusiinae

I

Genus

I

Cytolaimium

Rhabdocoma

Trefusia

Trefusiaiaimus

Tripyloidina

I

Family

I

I

Tripyloididae

I

Genus

I

Bathylaimus

Paratripyloides

Tripyloides

Lauratonema

Lauratonemoides

Tripylina

I

Superfamily

r-I-------1----------1

Ironoidea Oxystominoidea

I I

Family Family

I I II,----1------,[

Prismatolaimidae Tripylidae Paroxystominidae Oxystominidae

I I I

Genus Genus Genus

I I I

Onchulus Tripyla Maldivea

Paroxystomina

I

Lauratonematidae

I

Genus

I

I

Ironidae

Subfamily

I

Ironinae

I

Genus

I

Dolicholaimus

fronus

Syringolaimus

Trissonchulus

Subfamily

I

Thalassironinae

I

Genus

I

Conilia

fronella

Parironus

Pheronus

Thalassironus

Subfamily

I

Halalaiminae

I

Genus

I

Halalaimoides

Halalaimus

Subfamily

I

Oxystomininae

I

Genus

I

Litinium

Nemanema

Oxystomina

Thalassoalaimus

Wieseria

5
Abbreviations used in the key

a amphid d denticles I labia s spicules

ab apical band ds double seta lr longitudinal rods sp spear

an anus dt dorsal tooth Is labial seta spn spinneret

ap apophysis Isv left subventral (tooth) st stoma

e esophagus str striations

b bar ep excretory pore m mandibles sup supplement

bn band es esophageal structure ms solid mandibles svt subventral tooth

e chambers g gubernaculum oe ocellus t tooth

ee cephalic capsule tb transverse bar

cl claw h heavily cuticularized p exit pore tr tropis

cir circular structure walls

es cephalic seta rb refractive body v vulva

it inner thickening rsv right subventral (tooth)

w walls

Key to genera of the Enoplida _

Enoplida Filipjev, 1929

The genera in this key includes free-living marine nematodes with the following characteristics: Amphid variable, consisting

of slit-like opening and subcuticular pouch, may be enlarged, rounded, longitudinally oval or unispiral (Figs. 1,2,5,67,114);

integument smooth or annulated, rarely ornamented (punctate); lateral fields absent; stoma variable in shape, teeth and denticles sometimes present, mandibles present in some; esophagus cylindrical to clavate, rarely with bulb-like swellings or bulbs.

1 A Cephalic setae segmented (2es, 118). (Suborder Tripyloidina) 28

B Cephalic setae not segmented (5es) 2

2(1) A Esophagus only enveloping posterior end ofstoma (Ie); stoma spacious (1st); barrel-shaped with heavily cuticularized walls (lw) with or without teeth (It) and/or denticles; males of some genera with reduced stoma (76).

(Suborder Oneholaimina) 19

B Esophagus extending anteriorly to, or close to, lip region (2e); stoma narrow or large, with weakly cuticularized

walls (2st, 5w) 3

3(2) A Esophagus extending to lip region where it is attached to integument forming a cephalic capsule (3e, 6ce, 7ee);

precloacal supplements, when present, often cuticularized, tubular (4sup); amphids pocket-shaped (l2a). (Suborder

Enoplina) 4

B Esophagus extending to, or close to, lip region but not attached to integument (2e); precloacal supplements papilloid

or setose; amphids of variable shape. (Suborder Tripylina) 12

4(3) A Structure of esophagus muscular (2e), not vesiculate-cellulate (3es); stoma simple; teeth, if present, generally

of unequal size; mandibles (40m) absent. (Superfamily Leptosomatoidea) 5

B Esophagus with irregular outline and generally of vesiculate-cellulate structure; stoma generally with three teeth

of about equal size; mandibles present or absent (28m, 34m). (Superfamily Enoploidea) ,~

5(4) A Cephalic capsule inconspicuous with unmarked, weakly-lobed posterior margins. (Family Leptosomatidae)

......................................................................................... 6

B Cephalic capsule displaying conspicuous spots and indentations with strongly-lobed posterior margins (6ce)

......................................................................... Thoraeostomatidae

6
a

7

/'
a Ocelli with lens present (7oc); caudal glands extending anterior to anus b

Ocelli without lens; caudal glands confined to tail Pseudocella Filipjev, 1927

b(a) Tooth-like structure (tropis) present on ventral side of cephalic capsule (6tr) .

· Thoracostoma Marion, 1870

Tooth-like structure (tropis) absent Deontostoma Filipjev, 1916

6(5) A Cephalic capsule without noticeable inner thickening (12); teeth small if present (1St) Anticominae

a Cephalic setae about three head diameters long b

Cephalic setae not more than one head diameter long d

b(a) Stoma small, simple c

Stoma well developed Tubolaimella Cobb, 1933

c(b) Amphids posterior to longest postcephalic setae (9a); precloacal supplement absent .

· Barbonema Filipjev, 1927

Amphids anterior to longest postcephalic setae; precloacal supplement present (10sup) .

· Parabarbonema Inglis, 1964

d(a) Gubernaculum present e

Gubernaculum absent (11) Anticomopsis Micoletzky and Kreis, 1930

e(d) Excretory pore situated on cuticular projection (12) Paranticoma Micoletzky and Kreis, 1930

Excretory pore normal, not on cuticular projection f

f(e) Tubular precloacal supplement present or absent; setiform, subventral, precloacal supplements absent;

gubernaculum well developed (13) g

Tubular precloacal supplement absent; setiform, subventral supplements present; gubernaculum small,

simple (14g) Syringonomous Hope and Murphy, 1969

g(f) Stoma well developed with distinct, small teeth (1St) Odontanticoma Platonova, 1976

Stoma reduced, teeth absent Anticoma Bastian, 1865

B Cephalic capsule with noticable inner thickening (16it); teeth present or absent 7

7(6) A Stoma with three equally long mandibular teeth (17t) Triodontolaiminae

a Tail filiform with acute tip Triodontolaimus De Man, 1893

Tail short with clavate tip Jaegerskioeldia Filipjev, 1916

B Stoma with or without small teeth, of equal length if more than one Leptosomatinae

a Stoma with no more than one small tooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .b

Stoma with several teeth, plates, or dentated ribs f

b(a) Cervical setae present in two to three circles (18) Platycomopsis Ditlevsen, 1926

Cervical setae absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..... c

c(b) Tail clavate Leptosomella Filipjev, 1927

Tail cylindrical or conical, rounded or acute tip d

d(c) Cephalic sensillae papillose (7cs); ocelli present. e

Cephalic sensillae setose, well developed (19); ocelli absent Leptosomatina AUgen, 1951

e(d) Precloacal supplement present; gubernaculum large, with anteriorly directed projections (20g); sexual dimorphism in amphids absent Leptosomatides Filipjev, 1918

Precloacal supplement reduced or absent; gubernaculum small, without anteriorly directed projections

(21g); sexual dimorphism in amphids present Leptosomatum Bastian, 1865

8
"

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subventral section

vestibule &stoma

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9
f(a) Two flat setae situated near each amphid (22) Platycoma Cobb, 1894

Amphids not accompanied by two flat setae g

g(f) Stoma large, with apical dentated band (23ab) Cylicolaimus De Man, 1889

Stoma narrow (24st), with plates h

h(g) Three large teeth present in addition to small teeth on plates (24t); gubernaculum large with dentate distal

ends; precloacal region with long, stout ventrolateral setae in rows Macronchus Inglis, 1964

Three large teeth absent, small teeth present; gubernaculum small, without dentate distal ends; stout

precloacal setae absent i

i(h) Gubernaculum with two large apophyses (25ap) Corythostoma Hope and Murphy, 1972

Gubernaculum without two large apophyses (26g) Synonchus Cobb, 1894

8(4) A Stoma with a long, sharply pointed spear (27sp). (Family Thoracostomopsidae) .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thoracostomopsis Ditlevsen, 1918

B Spear absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9

9(8) A Stoma with three mandibles (28m). (Family Enoplidae) 10

B Stoma without three mandibles Phanodermatidae

a Cephalic capsule trilobed (29cc) b

Cephalic capsule not trilobed f

b(a) Cephalic capsule strongly developed, with or without striations (30cc) c

Cephalic capsule weakly developed d

c(b) Gubernaculum with dorsal apophysis (31ap) Dayellus Inglis, 1964

Gubernaculum without dorsal apophysis Phanoderma Bastian, 1865

aa Ocelli present Phanoderma Bastian, 1865

Ocelli absent Alyncoides Wieser, 1953

d(b) Cephalic capsule with three movable, labial lobes; labial papillae prominent; spicules elongate; precloacal

supplement present (32) Klugea Filipjev, 1927

aa Ocelli present Nasinema Filipjev, 1927

Ocelli absent Klugea Filipjev, 1927

Cephalic capsule without three movable, labial lobes; labial papillae not prominent; precloacal supplement present or absent e

e(d) Spicules long; tubular precloacal supplement absent Phanodermopsis Ditlevsen, 1926

Spicules short; tubular precloacal supplement present (33) Phanodermella Kreis, 1928

f(a) Precloacal supplement present in male Micoletzkyia Ditlevsen, 1926

Precloacal supplement absent in male Crenopharynx Filipjev, 1934

10(9) A Labial setae short, papilliform (28Is), teeth absent. (Subfamily Enoplinae, one genus) .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enoplus Dujardin, 1845

B Labial setae long, stout, setose (34Is); teeth present (34t) 11

11(10) A Lips relatively raised, conical (341) Enoploidinae

a Anterior end of subventral teeth long, extending to or beyond anterior end of mandibles (35) b

Anterior end of subventral teeth located far posterior to anterior end of mandibles f

b(a) Anterior end of subventral teeth at level of anterior end of mandibles c

Anterior end of subventral teeth extending beyond anterior end of mandibles d

10
®

11

® II
c(b) Mandibles arch-shaped, posterior apophyses absent (36) 0 0 0 0 0 0 •••• 0 0 0 0 0 .Oxyonchus Filipjev, 1927

Mandibles not arch-shaped, posterior apophyses present (37ap) 0 0 ••••••• 0 0 0 0 Filipjevia Kreis, 1928

d(b) Mandibles well developed (35m) 0 ••••••••••• 0 0 0 • 0 •• 0 0 0 0 •••• 0 0 0 0 0 0 Parasaveljevia Wieser, 1953

Mandibles reduced or vestigial 0 • 0 • 0 0 0 0 0 0 0 • 0 0 0 0 0 0 • 0 0 0 0 0 •••••• 0 0 0 0 0 ••••• 0 ••••••••••••• 0 •••e

e(d) Dorsal tooth reduced (38dt), subventral teeth larger (38svt); labial setae not bulbous. 0 0 • 0 ••• 0 0 0 •••

• •••••••••••••• 0 • 0 0 0 •••••••••••• 0 0 0 ••• 0 0 0 ••••• 0 •••••••• 0 0 •••••••Saveljevia Filipjev, 1927

Dorsal tooth not reduced, larger and more cuticularized than subventrals; labial setae bulbous (39)

• • 0 0 0 0 •••••••••••• 0 0 •••••• 0 ••••• 0 0 0 0 •• 0 0 0 0 ••• 0 0 0 •••Gairleanema Warwick and Platt, 1973

f(a) Teeth reduced, visible only in juveniles; anterior end of mandibles without claws (40) .. 0 ••••• 0 •••

• • 0 • 0 •••••••••••• 0 0 0 ••••• 0 0 0 •••••••••••• 0 0 0 0 0 ••• 0 0 ••••• 0 0 0 0 •••Parenoplus Filipjev, 1927

Teeth distinct at all stages (43t); mandibles well developed with claws (43c1) .. 0 0 0 ••••••••••• 0 ••g

g(f) Mandibles arch-shaped, with two longitudinal rods or plates (4Ur) connected by anterior bar (41b); if

anterior bar missing, rods separated by a narrow space (42) 0 0 0 0 0 0 0 ••••••••••• 0 0 ••••••h

Mandibular rods more solid, not as thin as above (34m) 0 0 0 0 0 • 0 ••••• 0 0 • 0 0 0 0 • 0 0 ••••••• 0 .1

h(g) Cephalic setae originate from base of cephalic capsule (43cc) 0 0 0 0 oEnoplolaimus De Man, 1893

Cephalic setae originate anterior to base of cephalic capsule (44) 0 0 0 0 0 • 0 0 0 •• 0 0 ••••• 0 0 0 •••••••• .i

i(h) Cephalic setae originate from middle or anterior end of cephalic capsule 0 0 0 0 ••••• 0 ••••••• 0 .j

Cephalic setae originate anterior to anterior end of cephalic capsule (45) 0 ••••••••

• •••• 0 0 0 0 •••••• 0 0 0 •••••• 0 ••••••••• 0 ••• 0 0 0 0 ••• 0 • 0 0 •••••• 0 0 Paramesacanthion Wieser, 1953

j(i) Longitudinal elements of mandibles rod-like, transverse anterior bar always present (41b) k

Longitudinal elements of mandibles plate-like, transverse anterior bar present or absent; if absent,

longitudinal elements separated by a narrow space (42) 0 ••••••• 0 •••••••Epacanthion Wieser, 1953

k(j) Precloacal supplement absent, in its place a row of short stout setae (46) .

• •••••••••••••• 0 • 0 0 •••••••••••••• 0 0 ••• 0 0 0 0 0 0 •• 0 0 0 0 ••••• 0 0 0 0 0 0 ••Africanthion Inglis, 1964

Precloacal supplement present or absent; if absent, midventral row of stout setae also absent 0 0 • 0 0 0 0

• •••• 0 • 0 ••••• 0 0 •••• 0 0 0 ••• 0 0 0 0 0 • 0 ••• 0 0 0 0 ••••••••••• 0 0 •••••• 0 0 Mesacanthion Filipjev, 1927

l(g) Lips unstriated (47) . 0 0 0 ••• 0 0 0 ••••••••• 0 0 0 •• 0 • 0 ••••• 0 0 0 ••• 0 • 0 0 oMesacanthoides Wieser, 1953

Lips striated (34str) .. 0 0 0 ••••• 0 0 0 0 0 0 0 • 0 0 0 0 0 ••••• 0 •••• 0 0 0 0 •••••• 0 0 0 0 • 0 0 0 • 0 ••••• 0 0 0 0 0 •••m

mO) Mandibles not extremely slender or elongate (ratio length-to-width at middle of shaft <10) .

• 0 • 0 0 0 • 0 0 •••• 0 • 0 0 0 0 •• 0 0 0 0 0 0 •••••• 0 0 ••• 0 0 0 0 •••••• 0 0 0 0 •••••• 0 0 0 • Enoploides Saveljev, 1912

Mandibles elongate, slender (ratio length-to-width ~1O) (48) . 0 0 0 0 • 0 0 • • Metenoploides Wieser, 1953

B Lips low, flattened (491) 0 0 0 ••••• 0 0 0 ••• 0 • 0 0 0 •••• 0 0 0 0 •••••• 0 ••••••••••• 0 ••••Rhabdodemaniinae

a Stoma with teeth 0 0 0 0 • 0 •••• 0 •• 0 0 • 0 0 0 0 0 •• 0 ••••••••• 0 0 0 ••••••••• 0 ••••••••••• 0 •••••••b

Stoma without teeth (50) 0 • 0 0 ••••• 0 0 0 0 0 • 0 0 0 0 0 ••••• 0 0 •••••• 0 0 0 0 0 •••Chaetonema Filipjev, 1927

b(a) Mandibles reduced to three rod-like structures or three paired teeth anteriorly; teeth also present posterior

in stoma (49t) 0 0 •••• 0 0 0 0 0 0 •••• 0 0 0 •• 0 0 0 0 0 0 •••• 0 •••Rhabdodemania Baylis and Daubney, 1926

Mandibles with two longitudinal rods (5Ur) connected by an anterior, transverse bar (51tb); teeth at level

of anterior bar (51t) ..... 0 0 ••••• 0 0 0 0 0 • 0 0 0 0 •••••••••••••••• 0 0 0 0 ••••••Trileptium Cobb, 1933

12(3) A Stoma narrow, amphids small, body not tapered anteriorly 0 ••• 0 ••• 0 0 0 0 ••••••••• 0 0 0 0 0 0 0 • 0 0 •••••• 0 013

B Stoma almost absent, amphids small to large, rounded to longitudinal; body tapered anteriorly (5). (Superfamily

Oxystominoidea) .. 0 0 •• 0 0 ••••• 0 0 ••••••• 0 0 0 •••• 0 0 0 0 0 0 • 0 0 0 0 0 0 •••••••••• 0 0 0 • 0 0 0 0 •••••• 0 0 0 0 •••17

13(12) A Stoma elongate-tubular with parallel walls; three claw-like teeth at anterior end of stoma (52t). (Family

Ironidae) ... 0 0 ••••••• 0 • 0 0 0 ••••• 0 0 0 0 • 0 ••••• 0 0 ••• 0 •• 0 ••••• 0 0 0 0 0 •••• 0 ••••• 0 •• 0 0 0 0 0 0 0 ••••••••14

B Stoma funnel- or barrel-shaped; teeth, when present, arising from walls of stoma .. 0 • 0 0 0 • 0 0 0 0 ••••••••15

12
13

@
14(13) A Cephalic sensillae papilliform (53cs) Ironinae

a Esophagus with distinct posterior bulb (54) Syringolaimus De Man, 1888

Esophagus without posterior bulb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . _. . . . . . . . . . . . . ..... b

b(a) Female gonad monodelphic, opisthodelphic Trissonchulus Cobb, 1920

Female gonad didelphic, amphidelphic c

c(b) Spinneret present, marine (55) Dolicholaimus De Man, 1888

Spinneret absent, freshwater Ironus Bastian,1865

B Cephalic sensillae setiform (S2cs) Thalassironinae

a Cephalic setae long, in two well-separated circles of 6+4 (56) Ironella Cobb, 1920

Cephalic setae in single circle of 10 b

b(a) Caudal glands and spinneret present d

Caudal glands and spinneret absent c

c(b) Cephalic setae short (57) Pheronus Inglis, 1966

Cephalic setae long (58) Parironus Micoletzky, 1930

d(b) Cephalic setae long or short; if long, cervical setae doubled (52ds); spicules short and equal in length

· Thalassironus De Man, 1889

Cephalic setae long, cervical setae not doubled; spicules unequal in length, one exceedingly long (59s)

· Conilia Gerlach, 1956

15(13) A Female genital duct opening (vulva) well separated from anus 16

B Female genital duct united with rectum forming a cloaca or vulva immediately anterior to anus .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lauratonematidae

a Cloaca present in females (60) Lauratonema Gerlach, 1953

Cloaca absent, vulva (6lv) immediately anterior to anus (61an) .

· Lauratonemoides De Coninck, 1965

16(15) A Stoma large, barrel-shaped or prismatic; esophagus muscular, areolated; stoma with large teeth (62). (Family

Prismatolaimidae) Onchulus Cobb, 1920

B Stoma reduced (63), funnel-shaped or straight; esophageal tissue more or less homogenous; freshwater forms

(except one genus). (Family Tripylidae) Tripyla Bastian, 1865

17(12) A Arnphids small, rounded (64); two ovaries; precloacal supplements large, knob-like or absent .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Paroxystominidae

a Supplements large, knob-like (65) Paroxystomina Micoletzky, 1924

Supplements absent Maldivea Gerlach, 1962

B Amphids large to very large, of diverse shapes; one or two ovaries; precloacal supplements minute, setose, or

absent. (Family Oxystominidae) 18

18(17) A Amphids elongate, longitudinally slit (5a); precloacal supplements absent Halalaiminae

a Caudal glands absent Halalaimoides Cobb, 1933

b Caudal glands present Halalaimus De Man, 1888

aa Distinct circle of labial setae (5Is) present Nualaimus Juario, 1974

Distinct circle of labial setae absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... bb

14
@

tb

®

15
bb(aa) Cephalic setae in two well-separated circles cc

Cephalic setae in two circles close together Nuada Southern, 1914

(= Tycnodora Cobb, 1920)

cc(bb) Amphids narrow (width <40% of corresponding diameter) Halalaimus De Man, 1888

Amphids wide (width ~40% of corresponding diameter) Pachydora Wieser, 1953

B Amphids rounded (66), not longitudinally slit; if present, precloacal supplements setose ....Oxystomininae

a Amphids very large, almost circular, orifice narrow; cephalic setae in two circles of six or single circle

ofl00r12 b

Amphids never very large; 10 cephalic setae in two circles c

b(a) Cephalic setae in two circles of six each, circles close together Litinium Cobb, 1920

Cephalic setae in single circle of 10 or 12 Thalassoalaimus De Man, 1893

c(a) Cephalic setae in two circles close together (67cs); circle of long cervical setae at level of amphids (67a)

· Wieseria Gerlach, 1956

Cephalic setae in two well-separated circles; without circle of long cervical setae at level of amphids

(68cs) d

d(c) Tail short, rounded (69) Nemanema Cobb, 1920

Tail long, attenuated (70) Oxystomina Filipjev, 1921

19(2) A Esophagus may be significantly enlarged posteriorly, occasionally with posterior bulb or multibulbar; sexual

dimorphism in stoma present or absent; if present, male stoma reduced; stoma with single large tooth (71t) or

three unequal teeth (8Ot), denticles may be present (80d); if so, male supplements may be knob-like, cuticularized. (Superfamily Enchelidoidea) 20

B Esophagus cylindrical or clavate, not significantly enlarged posteriorly, never multibulbar; sexual dimorphism

in stoma absent; stoma unarmed or with three teeth (72), denticles may be present (one genus); male supplements

papilloid, not cuticularized if present 24

20(19) A Esophagus with a row of moniliform bulbs (73) Bolbellidae

a Male supplements papilloid; sexual dimorphism in stoma (males reduced) present in some species ...

......................................................... .Polygastrophora De ~an, 1922

Male supplements cup-like, large (74); sexual dimorphism in stoma absent .

· Bolbellia Gerlach and Riemann, 1974

(= Belbolla Andrassy, 1976 = Bolbella Cobb, 1920)

B Esophagus without row of bulbs 21

21(20) A Stoma in both sexes spacious, about as long as wide; one to three teeth present; tooth, if single, originating

from base of stoma (75). (Family Eurystominidae) 22

B Stoma of female longer than wide; single tooth lies on wall of stoma (77t); stoma of males rudimentary (76).

(Family Enchelidiidae) Enchelidiinae

a Female stoma narrow, almost cylindrical; solid transverse stomatal rings present giving appearance of

more than two chambers (77c) b

Female stoma broader, divided into two unequal chambers (78c) by a transverse band, a ring or row of

denticles (78d); other rings, if present, faint and located posteriorly c

b(a) Transverse row of denticles in anterior portion of stoma; refractive body absent .

· Symplocostomella Micoletzky and Kreis, 1930

Anterior transverse stomatal ring solid rather than denticular; refractive body present (77rb) .

· Symplocostoma Bastian, 1865

16
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® @

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17
c(a) Posterior part of stoma longer than anterior part, asymmetrical; refractive body present (78rb) .....

· Calyptronema Marion, 1870

Posterior portion of stoma conical, symmetrical; refractive body absent .

· , Conistomella Stekhoven, 1942

22(21) A Stoma with single, large subventral tooth originating from base of stoma; male precloacal supplements present

(79sup) 23

B Stoma with three unequal teeth (8Ot); male precloacal supplements absent Thoonchinae

a Stoma with transverse rows of denticles (80d) Thoonchus Cobb, 1920

Stoma without transverse rows of denticles (81) Ditlevsenella Filijev, 1925

23(22) A Caudal glands and spinneret (79spn) present Eurystomininae

a Cervical setae extemely long (82), longer than corresponding body diameter .

· Ledovitia Filipjev, 1927

Cervical setae less than corresponding body diameter (75) b

b(a) Precloacal supplements (83sup) and gubernaculum (83g) without apophyses .

· Gerlachystomina Inglis, 1962

Precloacal supplements and gubernaculum with apophyses Eurystomina Filipjev, 1921

B Caudal glands and spinneret absent (84) Pareurystomininae

a Tail conical; precloacal supplements cup-shaped with apophyses (84ap) .

· Pareurystomina Micoletzky and Kreis, 1930

Tail flagellate (85); precloacal supplements tubular, without apophyses (86sup) .

· , Batheurystomina Lambshead and Platt, 1979

24(19) A Stoma unarmed or with minute teeth (87). (Superfamily Pelagonematoidea, Family Pelagonematidae) .

................................. , , 25

B Stoma armed with one to three large teeth (It). (Superfamily Oncholaimoidea, Family Oncholaimidae)

........................................................................................ 26

25(24) A Female gonad monodelphic, opisthodelphic; tail filiform. (Subfamily Curvolaiminae) .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . " Curvolaimus Wieser, 1953

B Female gonads didelphic, amphidelphic; tail not filiform Pelagonematinae

a Stoma small, short, cup-shaped, nearly as wide as long .

· Bradybucca Stekhoven, 1956

(= Anoncholaimus Stekhoven, 1950)

Stoma large, longer than wide b

b(a) Head with three lips; stoma cylindrical (88); males with bursa Anoplostoma Biitschli, 1874

Head with six lips; stoma cup-shaped or with curved or bipartite walls; males without bursa c

c(b) Stoma in two tandem parts (89) , Illium Cobb, 1920

(= Pseudopelagonema Kreis, 1932)

Stoma not in two tandem parts d

d(c) Esophagus with posterior bulb . ' Pelagonemella Kreis, 1932

Esophagus without posterior bulb Pelagonema Cobb, 1894

18
® ®

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I

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=r=

19
26(24) A Stoma with one large tooth (9Ot) and numerous smaller, well-developed denticles (9Od). (Subfamily Octonchinae)

o000000000000000000000000000000000000000000000000000000000000000000000Octonchus Clark, 1961

(= Polydontus Schulz, 1932)

B Stoma with one to three well-developed teeth, denticles absent 00000000000000000000. 0000000000. 000027

27(26) A Stoma with three teeth; right subventral very large (91rsv), left subventral (911sv) and dorsal (91dt) reduced

or even greatly reduced; males may have copulatory bursa; gubernaculum absent; demanian system (92a,b) present

or absent; if present, without exit pores (92a) 000000. 00000. 0. 00000000.. 00000000000oOncholaimellinae

a Females with one ovary; if present, bursa of males supported by long, knobbed papillae (93) 00000b

Females with two ovaries; if present, bursa ofmales not supported by long, knobbed papillae; circumcloacal

setae present 0000000000. 00000000000. . 00000000000. 0000000000000000000000000000. 000. 0000c

b(a) Stoma large, not cup-shaped; bursa in males supported by long, knobbed papillae (93) 00... 0. 00000

o 00000000000000000000000000000000000000000000000. 0000000. .Oncholaimelloides Timm, 1969

Stoma small, cup-shaped (94st); bursa absent 00000000000000.0000000.0 oKrampia Ditlevsen, 1921

c(a) Cuticle with distinct annules (95), longitudinal striations distinct or indistinct 00000000000000000000

o. 00000000000000000000000000000000.. 000000000000000000000Oncholaimoides Chitwood, 1937

Cuticle without distinct transverse annulations or longitudinal striations 000000000000000000000000d

d(c) Right subventral tooth large, almost fIlls stomatal cavity (91rsv); left subventral (911sv) and dorsal teeth

(91dt) small, not elongate; bursa present or absent o. 0000... 0. 000000000. 0000000. 000000. 00000e

Right subventral tooth not as large as above (96rsv); left subventral (96lsv) and dorsal teeth (96dt) reduced

but elongate, distinct; small bursa present (97) or absent 000000. 0000000000Viscosia De Man, 1890

e(d) Right subventral tooth heavily cuticularized; stoma divided transversely by cuticularized band (91bn);

spicules moderate to very long; bursa present or absent 000000.. 000oOncholaimellus De Man, 1890

Right subventral tooth (981'sv) not as heavily cuticularized; dorsal (98dt) and left subventral (98Isv) teeth

greatly reduced; stoma not divided transversely by cuticularized band; bursa absent; circumcloacal setae

present; spicules short, about one anal diameter long, straight . 00000oMononcholaimus Kreis, 1924

B Stoma with three teeth, equal or unequal in size; if subventrals unequal, left largest; right subventral and dorsal

never greatly reduced; bursa absent; gubernaculum present or absent; demanian system present (92a,b) or absent,

exit pores present (92b,p) or absent 000.000000000000000000000000000000000.0000. 000oOncholaiminae

a Subcuticular trabeculae present (99) 00000000000000000. 0000000Prooncholaimus Micoletzky, 1924

Subcuticular trabeculae absent 00. 0000. 00. 0000000000. 0. 0000. 00000000000000000000000000000b

b(a) Females with two ovaries 000000. 0.. 0000000000000000000. 00. 0. 00000000000000000. 000000. 00c

Females with one ovary 0000000000000000000000. 0. 000. 00000000000000000000000. 0. 00000000h

c(b) Demanian system absent 000000000000000000000000000000000000000000000000000000. 00. 00000d

Demanian system present 00000000. 000. 000. 00000000000000... 00. 00000000. 00000. 00000. 00000e

d(c) Tail short (100); gubernaculum present 0000000000.0000.000000.00000 .. oPontonema Leidy, 1885

Tail long, filiform; gubernaculum absent 0000. 0000000000000. 000000Filoncholaimus Filipjev, 1927

e(c) Demanian system exit pore single, at level of vulva; subventral teeth equal (lOlsvt), larger and anterior

to dorsal (101dt); gubernaculum present (102) 00000000000 .. 0. 0000000000.Meyersia Hopper, 1967

Demanian system exit pores two or more (92b,p), posterior to vulva; teeth variable; gubernaculum present

or absent 00000000000000000.0 .. 00.0000000000000000000000000000 00000000000000000000000of

f(e) Subventral teeth equal, smaller than dorsal; spicules very long and thin (103); gubernaculum absent 0

. 00000000000000000000. 0000000. 00000000. 00000. 0. 0000000. oMetoncholaimoides VVieser, 1953

Subventral teeth equal or unequal, larger or equal to dorsal; gubernaculum present 0000000000. 00og

20
® a b

®

21
g(f) Stoma about three times as long as wide; gubernaculum small .... .Kreisoncholaimus Rachor, 1969

Stoma about two times as long as wide (104); gubernaculum larger (105) .

· Adoncholaimus Filipjev, 1918

h(b) Demanian system absent; spicules short; gubernaculum absent (106) .

· Pseudoncholaimus Kreis, 1932

Demanian system present; spicules variable; gubernaculum present or absent i

i(h) Spicules short; gubernaculum absent j

Spicules moderate to very long; gubernaculum present or absent k

jO) Subventral teeth equal (l07svt) or unequal; males with well-developed, fleshy or papilloid precloa"al organ

(108); demanian system with exit pores (92b,p) Oncholaimium Cobb, 1930

Subventral teeth equal or unequal; males without well-developed precloacal organ; demanian system without

exit pores Oncholaimus Dujardm, 1845

kO) Subventral teeth equal and larger than dorsal; spicules moderate to long; gubernaculum present or

absent 1

Subventral teeth unequal; spicules moderate to greatly elongate (setaceous) (109); gubernaculum present

or absent Metoncholaimus Filipjev, 1918

l(k) Spicules medium length; gubernaculum small (110) .

· Metaparoncholaimus De Coninck and Stekhoven, 1933

Spicules long, slender; gubernaculum large (111) Wiesoncholaimus Inglis, 1966

28(1) A Stoma with two chambers (112c); teeth present (112t) or absent; gubernaculum (113g) very large, equal to or

larger than spicules (113s); cephalic setae in single circle of 10 setae Tripyloididae

a Cephalic setae short, thick, distinctly segmented; stoma funnel-shaped (112c); teeth present or absent

.................................................................................. b

Cephalic setae long, thin, indistinctly segmented; anterior portion of stoma large, globular (114); teeth

present or absent Bathylaimus Cobb, 1893

b(a) Posterior chamber of stoma with teeth (112t) Tripyloides De Man, 1886

Posterior chamber of stoma without teeth Paratripyloides Stekhoven, 1950

B Stoma not divided into two chambers (115); teeth absent; gubernaculum smaller than spicules; cephalic setae

in single circle of six or in two circles of 6+4. (Family Trefusiidae) 29

29(28) A Stoma large, barrel-shaped; lips large, with three large circular structures (l16cir); cephalic setae very short

. (116cs). (Subfamily Halanonchinae) Halanonchus Cobb, 1920

B Stoma narrow, not barrel-shaped (115st); lips small, without circular structures; cephalic setae longer (115cs)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Trefusiinae

a Amphids circular; gubernaculum present b

Amphids oblong with posteriorly directed pouch (115a); gubernaculum absent c

b(a) Amphids closed circles (117); females with one ovary Rhabdocoma Cobb, 1920

Amphids open circles (118); females with two ovaries Cytolaimium Cobb, 1920

c(a) Male with two testes Trefusia De Man, 1893

Male with one testis Trefusialaimus Riemann, 1974

22
,

,

I

I

, @rr~"4i'\ ,

I

I \

I y' f \ , I (I __ "\, lll.'" \' ( , ( \ { .. " :' \ I '/ \ " 1." ,'4 I I

It" ~: \ I,"'\JI I," \ I I, ' :' I

23
Illustration sources _

Fig. no.

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

21

22

23

24

25

26

27

28

29

30

31

32

33

34

35

36

37

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

Reference

Wieser 1959, fig. 26d

Keppner 1986, fig. 35, p. 333

Wieser 1953, fig. 35c, p. 69

Andrassy 1968, fig. c, p. 213

Wieser and Hopper 1967, fig. 3a, p. 309

Original drawing

Bongers 1983, fig. lOA, p. 827

Wieser 1954, fig. 2b, p. 176

Gerlach 1956b, fig. d, tafe1 26

Inglis 1964, fig. 134, p. 361

Micoletzky 1930, fig. 5c, p. 268

Mico1etzky 1930, fig. 4a, p. 262

Wieser 1953, fig. 3d, p. 18

Hope and Murphy 1969, fig. 2B, p. 514

Platonova and Gal'tsova 1976, fig. 53, p. 145

Inglis 1964, fig. 2, p. 274

Vitiello 1970, fig. 2a, p. 141

Vitiello 1970, fig. 6b, p. 145

Vitiello 1970, fig. 3b, p. 141

Bongers 1984, fig. 2B, p. 19

Bongers 1983, fig. 15B, p. 835

Inglis 1966, fig. 1, p. 84

Schuurrnans-Stekhoven 1935, fig. 72A, p. 11

Inglis 1964, fig. 141, p. 362

Hope 1967, fig. 3D, p. 9

Platt and Warwick 1983, fig. 83c, p. 187

Inglis 1964, fig. 104, p. 357

Inglis 1964, fig. 110, p. 358

Inglis 1964, fig. 38, p. 346

Steiner 1921, fig. 6a, tafe1 12

Inglis 1964, fig. 42, p. 346

Wieser 1953, fig. 29b, p. 57

Vitiello 1970, fig. 31e, p. 179

Wieser and Hopper 1967, fig. lOa, p. 312

Wieser 1959, fig. 35c, p. 69

Bress1au and Schuurrnans-Stekhoven 1940, fig. 17C,

tafel IV

Kreis 1928, fig. 13b, p. 195

Gerlach 1956b, fig. i, tafe! 28

Warwick and Platt 1973, fig. 11a, p. 152

Wieser 1953, fig. 37a, p. 71

Inglis 1966, fig. 10, p. 86

Inglis 1966, fig. 13, p. 90

Original drawing

Inglis 1966, fig. 13, p. 90

Vitiello 1970, fig. 34a, p. 184

Inglis 1964, fig. 94, p. 355

Wieser and Hopper 1967, fig. 13a, p. 313

Wieser 1953, fig. 52b, p. 93

Wieser 1959, fig. 2a, plate 1

Wieser 1953, fig. 54a, p. 93

Original drawing

Original drawing

Van der Heiden 1974, fig. 3A, p. 426

Chitwood 1951, fig. 91, p. 647

Wieser 1953, fig. 57d, p. 97

Original drawing

Inglis 1966, fig. 17, p. 94

24

58 Boucher 1971, fig. lA, p. 976

59 Gerlach 1956a, fig. 51, p. 38

60 Gerlach 1953, fig. 5, p. 46

61 Gerlach 1956b, fig. b, tafel 26

62 Andrassy 1964, fig. a, p. 31

63 Brzeski 1964, fig. I, p. 160

64 Wieser 1953, fig. 22a, p. 45

65 Wieser 1953, fig. 22b, p. 45

66 Gerlach 1962, fig. i, tafel 4

67 Gerlach 1956b, fig. i, tafel 26

68 Platt and Warwick 1983, fig. 88A, p. 199

69 Platt and Warwick 1983, fig. 88E, p. 199

70 Gerlach 1962, fig. f, tafel 4

71 Wieser and Hopper 1967, fig. 27A, p. 321

72 Schuurrnans-Stekhoven and Adam 1931, fig. 3, plate 5

73 Timm 1952, fig. 25, p. 17

74 Wieser 1953, fig. 83d, p. 137

75 Timm 1961, fig. 16A, p. 41

76 Jensen 1986, fig. 6a, p. 98

77 Micoletzky 1930, fig. 17, p. 310

78 Hopper and Meyers 1967, fig. 9, p. 492

79 Wieser 1959, fig. 87d, p. 141

80 Gerlach 1954a, fig. 3A, tafel 89

81 Riemann 1966, fig. 52f, p. 191

82 Wieser 1953, fig. 83b, p. 137

83 Gerlach 1954b, fig. 1b, p. 98

84 Original drawing

85 Lambshead and Platt 1979, fig. 2c, p. 375

86 Lambshead and Platt 1979, fig. 2f, p. 375

87 Vitiello 1970, fig. 46a, p. 198

88 Original drawing

89 Wieser and Hopper 1967, fig. 25a, p. 320

90 DeConinck 1965, fig. 472, p. 661

91 Original drawing

92 Hope 1974, fig. 11, p. 417

93 Murphy 1966, fig. 1d, p. 30

94 Bresslau and Schuurmans-Stekhoven 1940, fig. 21A,

tafel 5

95 Original drawing

96 Wieser and Hopper 1967, fig. 23a, p. 319

97 Wieser and Hopper 1967, fig. 22f, p. 319

98 Original drawing

99 Original drawing

100 Hopper and Meyers 1967, fig. 8, p. 492

101 Hopper 1967, fig. 12, p. 148

102 Original drawing

103 Wieser 1959, fig. 74a, p. 125

104 Hopper 1969, fig. 1, p. 677

105 Hopper 1969, fig. 2, p. 677

106 Belogurov et al. 1972, fig. I, p. 1451

107 Wieser and Hopper 1967, fig. 16a, p. 315

108 Wieser and Hopper 1967, fig. 15c, p. 315

109 Original drawing

110 Mawson 1958, fig. 38c, p. 352

III Inglis 1966, fig. 34, p. 99

112 Gerlach 1965, fig. 31a, tafel 155

113 Original drawing

114 Original drawing

115 Keppner 1986, fig. 29, p. 330

116 Wieser and Hopper 1967, fig. 5a, p. 310

117 Platt and Warwick 1983, fig. 133A, p. 289

118 Cobb 1920, fig. 31, p. 251
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Enoplida (Nematoda). Opusc. Zool. (Budap.) 5(1): 25-4:1.

1968 Fauna Paraguayensis 2. Nematoden aus den Galenewaldem des Acaray-Flusses. Opusc. Zool. (Budap.) 8(2):

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Belogurov, 1.0., L.S. Belogurova, and A.V. Leonova

1972 The review of the genus Pseudoncholaimus (Nematoda:

Oncholaimidae) with reference of monodelphness of nematodes and heterotopy of the female genital pore. Zool. Zh.

51(10): 1450-1456.

Bongers, T.

1983 Revision of the genus Leptosomatum Bastian, 1865

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1984 Revision of the genus Leptosomatides Filipjev, 1918.

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Boucher, G.

1971 Deux espeees nouvelles de nematodes de la vase terrigene cotiere de Banyul-sur-Mer. Bull. Mus. Nat!. d'Hist.

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Bresslau, E., and J.H. Schuurmans-Stekhoven

1940 Marine freilebende Nematoden aus der Nordsee. Brux.

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Brzeski, M.

1964 Revision der gattung Tripyla Bastian and Paratripyla

gen. n. (Nematoda, Tripylidae). Ann. Zool. (Warsaw)

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Chitwood, B.G.

1951 North American marine nematodes. Texas J. Sci.

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Cobb, N.A.

1920 One hundred new nemas. Contrib. Sci. Nematol.

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De Coninck, L.A.P.

1965 Classe de nematodes. In Grasse, P. (ed.), Traite de

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Gerlach, S.A.

1953 Lauratonema nov. gen., Vertreter einer neuen Familie

mariner Nematoden aus dem Kustengrundwasser. Zool.

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1954a Les nematodes marins libres des eaux souterraines

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1954b Nouveaux nematodes libres des eaux souterraines

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19568 Brasilianische Meeres-Nematoden 1. (Ergebnisse eines

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von Spitzbergen. Veroeft. Inst. Meeresforsch. Bremerhaven 16:109-172.

25

Hope, W.D.

1967 A review of the genus Pseudocella Filipjev, 1927

(Nematoda: Leptosomatidae) with a description of Pseudocella triaulolaimus n. sp. Proc. Helminthol. Soc. Wash.

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1974 Nematodes. In Giese, A.C., and 1.S. Pearse (eds.),

Reproduction of marine invertebrates, p. 391-469. Acad.

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Hope, W.D., and D.G. Murphy

1969 Syringonomous typicus new genus, new species (Enoplida: Leptosomatidae) a marine nematode inhabiting arenaceous tubes. Proc. BioI. Soc. Wash. 82:511-518.

Hopper, B.E.

1967 Free-living marine nematodes from Biscayne Bay,

Florida, II. Oncholaimidae; Description offive new species

and one new genus (Meyersia). Mar. BioI. 1:145-151.

1969 Marine nematodes of Canada. II. Marine nematodes

from the Minas Basin-Scots Bay area of the Bay of Fundy,

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Hopper, B.E., and S.P. Meyers

1967 Foliicolous marine nematodes on turtlegrass, Thalassia testudinum Konig, in Biscayne Bay, Florida. Bull. Mar.

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Inglis, W.G.

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26