Nematodes of Puerto Rico: Actinolaimoidea New Superfamily with a Revision of Its Gener a and Species with Addenda to Belondiroidea (Nemata, Adenophorea, Dorylaimida)

TECHNICAL PAPER 43 OCTOBER 1967

NEMATODES OF PUERTO RICO: ACTINOLAIMOIDEA NEW SUPERFAMILY WITH A REVISION OF ITS GENER A AND SPECIES WITH ADDENDA TO BELONDIROIDEA (NEMATA, ADENOPHOREA, DORYLAIMIDA)

GERALD THOHNE

University of Puerto Rico Mayaguez Campus AGRICULTURAL EXPERIMENT STATION Rio Piedras, Puerto Rico 00928 Table of Contents

Introduction 5 Historical 6 Species of Actinolaimus through 1964 and their present position 7 Important Generic and Specific Diagnostic Characters 8 Diagrammatic Outline of the Actinolaimoidea 10 Key to Families, Subfamilies and Genera of the Actinolaimoidea 10 Actinolaimoidea, New Superfamily 12 Family Actinolaimidae (Thorne, 1939) Meyl, 1960 12 Key to Species of Actinolaimus 12 Subfamily Brittonematinae, n.subfam. 13 Key to Genera of Brittonematinae 13 Genus Brittonema, n.g. 13 Key to Species of Brittonema 14 Genus Actinocephalus, n.g. 18 Family Neoactinolaimidae, n.fam. 19 Genus Neoactinolaimus, n.g. 21 Key to Species of Neoactinolaimus 21 Genus Metactinolaimus Meyl, 1957 23 Genus Egtitus, n.g. 24 Key to Species of Egtitus 24 Family Paractinolaimidae, n.fam. 26 Genus Paractinolaimus Meyl, 1957 27 Key to Species of Paractinolaimus 27 Genus Westindicus, n.g. 32 Family Carcharolaimidae, n.fam. 33 Genus Carcharolaimus Thorne, 1939 34 Key to Species of Carcharolaimus 35 Genus Carcharoides, n.g. 39 Subfamily Caribenematinae, n.subfam. 39 Genus Cal'ibenema, n.g. 40 Family Trachypleurosidae, n.fam. 43 Family Mylodiscidae, n.fam. 43 Addenda to Belondiroidea 44 Genus Axonchoides, n.g. 44 Summary 47 Resumen 47 Literature Cited 47 NEMATODES OF PUERTO RICO: ACTINOLAIMOIDEA NEW SUPERFAMILY WITH A REVISION OF ITS GENERA AND SPECIES WITH ADDENDA TO BELONDIROIDEA

(NEMATA, ADENOPHOREA, DORYLAIMIDA) 1 Gerald Thorne2 INTRODUCTION This work continues the production of group papers begun by the writer with the publication on nematodes of the superfamily Belon- diroidea (21).3 It is based principally on specimens now in the perma- nent collection of the Puerto Rico Agricultural Experiment Station representing new species and genera which have made possible a complete revision of the forms previously described. Many old descriptions and illustrations are inadequate in important details and assignments to new genera may sometimes be found in error when collections are again made in type localities. In some instances it has been necessary to leave the species as inquirendae. For descriptions and illustrations of previously described species the reader is referred to: Monograph on the Nematodes of the Super- family Dorylaimoidea (20). As in the previous work (21) much of this material originated in the Luquillo Experimental Forest (EI Yunque area), where conditions are especially favorable for species of Actinolaimus, Brittonema, and other groups in which tails are elongated or filiform. From cultivated and dryer parts of the Island these genera are seldom collected and members of blunt-tailed Carcharolaimidae are more prevalent.

1J. B. Goodey in T. Goodey (13) abandoned the Phylum NEMATA and Classes SECERNENTEA and ADENOPHOREA, but this act is not herein accepted. . 2Consulting Nematologist, Agricultural Experiment Station, Mayaguez,Campus, University of Puerto Rico, Rio Piedras, P.R. The writer is indebted to Mrs. Carmen T. Ramirez Pagan who prepared the Cobb metal slide microscopic mounts, and to Mr. Jesse Roman who gave invaluable assistance in collecting specimens and pre- paring and handling the manuscript; and to Drs. Alejandro Ayala Almod6var and Luis F. Martorell, al'ld Miss Lucy Franqui Mendez for their kind assistance in check- ing and correcting the original publication and proofs, and indexing and typing of the same. Mr. Roy O. Woodbury, Plant Taxonomist of the Experiment Station, kindly allowed me to accompany him on several trips to the Rain Forest and other areas on the Luquillo Experimental Forest. Acknowledgment is due Dr. J. M. Good for library privileges of the Nematology Investigations Office, Beltsville, Md., and especially to Mrs. Grace S. Cobb for locating many references. Dr. and Mrs. John Ferris, Purdue University, also secured copies of certain essential papers. Dr. A. D. Baker's Check List of the Nematode Superfamilies Dorylaimoidea, Rhabditoidea, Tylenchoidea, and Aphelenchoidea has been invaluable as a reference source in nomenclature. 3Italic numbers in parentheses refer to Literature Cited, p. 47. 6 Gerald Thorne

Examination of intestinal tracts sometimes gave evidence of food habits and these are recorded for certain species. However, observa- tions on intestinal contents have not been very rewarding. Even when masses of food material are present it has been impossible to identify the particles. Usually we have regarded the actinolaims as predators but in not a single instance have remains of nematodes, oligochaetes or other animal life been identified. Only in Cal'charolaimus were oligocheate setae observed.

One of the most interesting discoveries was the presence of two larvae of what apparently is a mermithid which were found living in the body cavity of Paractinolaimus robustus.

HISTORICAL Nematodes of the Superfamily ACTINOLAIMOIDEA were first mono graphed under subfamily rank in 1939 by the writer (20). It was then noted that the group comprised species representing several types and that eventually higher rank would be justified. Meyl (18) elevated the group to family rank and erected the genera Paractinol- aimus, Actinolaimoides, and Metactinolaimus, giving the group a more substantial status. Goodey (13) suppressed Actinolaimidae and reestab- lished Actinolaiminae, but this act is not recognized herein. Numerous other species described by various workers are listed in the keys to species. Outstanding collections in Puerto Rico revealed new genera and family groups, making it imperative that superfamily rank be established. Complete references to species prior to 1939 are listed in Thorne, 1939 (20). Baker (8) and Goodey (13) should also be consulted for more recent citations.

Assignment of previously described species to other genera is difficult in many instances because of inadequate diagnosis. Frequently, absence of males complicates the problem because supplement arrange- ments are important diagnostic characters in certain instances. Paractinolaimus striatus (Thorne, 1939) Meyl, 1957, is an interesting species combining the strong spear and numerous denticles of Paractinolaimus with deep striae similar to those of BTittonema. Acti- nolaimus l'uwenzori has longitudinal striae but supplements are in a ventral series, not in fascicles as in true Actinolaimus. Bridging fOlms like these are to be expected among extensive collections and eventually these two species will probably be found to represent groups which merit generic rank. For the present it is considered best to allow them to remain in their present status. Brasilaimus subaquilus Lordello and Zamith (16) probably represents a valid genus although Clark (10) made it a synonym of Actino- laimus. More critical examination of the specimen is desil:able. Nematodes of Puerto Rico 7

SPECIESOF Actinolaimus THROUGH1964 ANDTHEm PRESENTPOSITION A. afl'icanus Filipjev, 1929-Neoactinolaimus afl'icanus (Filipjev, 1929) n.comb. A. bil'keti Altheer, 1960-position undetermined A. chappuisi Meyl, 1955-remains in Actinolaimus A. chitwoodi MOOlthy, 1937-Neoactinolaimus chitwoodi (Moorthy, 1937) n.comb. A. cinctus Cobb in Thorne, 1939-Westindicus cinctus (Cobb, 1939) n.comb. A. confol'mis W. Schneider, 1935-Trachypleurosum confol'mis (W. Schneider, 1935) Andrassy, 1959 A. costatus W. Schneider, 1935-remains in Actinolaimus A. cyatholaimus (Daday, 1905) Steiner, 1916-Egtitus cyatholaimus (Daday, 1905) n.comb. Synonym: DOl'ylaimus cyatholaimus Daday, 1905 A. elaboratus (Cobb, 1906) Cassidy, 1930-Egtitus elaboratus (Cobb, 1906) n.comb. Synonym: DOl'ylaimus elaboratus Cobb, 1906 A. filipjev W. Schneider, 1935-Paractinolaimus filipjev (Schneider, 1935) Meyl, 1957 A. frostae AUgen, 1952-Neoactinolaimus frostae (AUgen, 1952) n.comb. A. hintoni AUgen, 1952-Neoactinolaimus hintoni (AUgen, 1952) n.comb. A. hutchinsoni Filipjev, 1929-Neoactinolaimus hutchinsoni (Filipjev, 1929) n.comb. A. labyrinthstoma (Cobb, 1893) Cobb, 1913-Trachypleul'osum laby- l'inthstoma (Cobb, 1893) Andrassy, 1959 A. macl'olaimus (de Man, 1880) Steiner, 1916-Paractinolaimus macrolaimus (de Man, 1880) Andrassy, 1964 A. michaelseni Steiner, 1916-Neoactinolaimus michaelseni (Steiner, 1916) n.comb. A. micoletzkyi Schneider, 1935-Paractinolaimus micoletzki (Schneider, 1935) Meyl, 1957 A. microdentatus Thorne, 1939-Paractinolaimus micl'odentatus Thorne, 1939) Meyl, 1957 A. mpoumensis Altheer, 1916-position undetermined A. neocyatholaimus Kreis, 1936-Egtitus neocyatholaimus (Kreis, 1936) n.comb. A. nudus Wu & Hoeppli, 1929-Egtitus nudus (Wu & Hoeppli, 1929) n.comb. 8 Gerald Thorne

A. occalescens W. Schneider, 1937- Egtitus occalescens (W. Schneider, 1937) n.comb. A. omel'coopel'i Filipjev, 1931-remains in Actinolaimus A. schuul'mansstekhoveni Meyl, 1957-remains in Actinolaimus Synonym: Actinolaimus omercoopel'i of Schuurmans Stekhoven & Teunissen, 1938 (not of Filipjev, 1931) A. palmael'i AUgen, 1953-position undetermined A. papillatus W. Schneider, 1935-remains in Actinolaimus A. propinquus Cobb, in M. V. Cobb, 1915-Egtitus propinquus (Cobb, 1915) n.comb. A. radiatus Cobb, 1913-remains as type species of Actinolaimus. Not a synonym of A. tl'ipapillatus as designated by Andrassy, 1954 A. l'otundicauda de Man, 1880-Cal'chal'Olaimus l'otundicauda (de Man, 1880) Thorne, 1939 A. l'uzwenzoTii de Coninck, 1935-position undetermined A. straeleni Meyl, 1957-remains in Actinolaimus A. striatus Thorne, 1939-Paractinolaimus stl'iatus (Thorne, 1939) Meyl, 1957 A. surinamensis Micoletzky, 1925-Egtitus sUl'inamensis (Micoletzky, 1925) n.comb. A. taylol'i Meyl, 1957-Bl'ittonema taylol'i (Meyl, 1957) n.comb. A. tenuiaculeatus (Kreis, 1924) Micoletzky, 1925-Bl'ittonema tenuiaculeatus n.comb. Synonym: DOl'ylaimus tenuiaculeatus Kreis, 1924. A. tenuis W. Schneider, 1935-Neoactinolaimus tenuis (W. Schneider, 1935) n.comb. A. toblel'i (Menzel and Micoletzky) in Micoletzky, 1925)-Actinolai- moides toblel'i (Menzel & Micoletzky, 1925) Meyl, 1957 A. tl'ipapillatus (Daday, 1905) Steiner, 1916-N eoactinolaimus tl'ipapillatus. n.comb. Synonym: Dorylaimus tripopillatus Daday, 1905 A. tripapillatus of Kreis, 1932-Metactinolaimus heisi Meyl, 1957 A. zeelandicus Clark, 1962-N eoactimolaimus zeelandicus (Clark, 1962) n.comb. IMPORTANT GENERIC AND SPECIFIC DIAGNOSTIC CHARACTERS Longitudinal striae are of two types: Elevated ridges as in Acti- nolaimus radiatus and deep grooves in the .cuticle as found on Bl'ittonema and Actinocephalus. Diagnosis of striated species usually do not include statements as to the type of striae present, thus making definite assignment to genera somewhat uncertain. Nematodes of Puerto Rico 9

Pharyngeal armature may vary from simple platelike pieces to elaborate ribbed basketlike structures, with or without strong onchia and numerous denticles. The spear guiding-ring may be a simple or massive single ring, or a slender double-type framework. A constriction of the esophagus just posterior to the spear extension frequently is an important generic character. Anterior portion of esophagus may be a slender hyaline tube or have radial muscles. Expansion to the enlarged basal portion may be gradual or abrupt. Esophageal gland nuclei vary greatly in visibiltiy, the dorsal one is always conspicuous while the four submedian are generally difficult to see. In some instances even their outlets in the esophageal lumen can- not be observed and it is even possible that they are not present. Lateral fields may have one or two lines of cells with pores which vary from conspicuous tubes to almost invisible fibers. A structure herein named the basal esophageal shield may be present or absent. These appear to be muscular pads varying from flat disks to cupped basins. Cardias usually are elongate conoid but may be hemispheroid. Nygolaimoid glands at esophagus base are known for Carcharolai- mus dentatus and C. aberrans. These species do not have basal shields. Vulva may be transverse or longitudinal slits or pOl'elike openings with tiny labia. Ovaries, two except in Actinolaimus radiatus, A. africanus, and Actinolaimoides tobleri. Supplements may be arranged in fascicles (bundles) or in ventral series. Ventrosubmedian papillae on males may be present or absent. Female tails conoid to subfiliform except blunt and rounded in Carcharolaimidae. Male tails blunt, rounded except Brittonematinae and Trachypleurosum. Generally face-view of dorylaims aids in identification, but the heavily sclerotized pharyngeal armature so obscures details that little can be seen that is of diagnostic value. Therefore they have not been illustrated. Details of pharyngeal armature vary when seen from different angles; a submedian view may be considerably different from a lateral or dorsal, and this fact must be remembered when comparing illustrations. 10 Gerald Thome

Nomenclature of the various parts of the stoma, or pharynx, has not been definitely determined. Chitwood (9) stated: Actinolaimus: the latter is a most unusual member of the group having a radially striated cheilorhabdion and four massive onchium-like stylet guides in addition to heavily sclerotized pro and mesorhabdions and stylet guiding ring. Since the latter structure is absent from such forms as Nygolaimus we may presume it to be the modified telorhabdion. However, its homology has not been determined.

DIAGRAMMATIC OUTLINE OF AcTINOLAIMOIDEA Family I Subfamily Genus

Actinolaiminac Actillolainms Actinolaimidae ) { Brittonema Brittonematinae Actinocephalus

{ Neoactinolaimus Neoactinolaimidae Metactinolaimus Egtitus

Superfamily ~ ACTINOLAIl\lOIDEA Paractinolaimidac { Paractinolaimus Westindicus

{ Carcharolaimus , Carcharolaiminae Carcharoides Carcharolaimidac Antholaimus

Caribenematinae Carib enema

{ Trachypleurosum Trachypleurosidac Actinolaimoides

Mylodiscidae Mylodiscus

KEY TO FAMILIES, SUBFAMILIES, AND GENERA OF ACTINOLAIMOIDEA 1. Cuticle marked by longitudinal striae Actinolaimidae 2 Cuticle with minute transverse striae only 4 2. Spear robust, aperture 2/5 - 1/2 its length Actinolaiminae A single genus, Actinolaimus Spear slender, aperture 1/3 or less its length Brittonematinae n.sf. 3 3. Head continuous with neck contour Brittonema n.g. Head broadly expanded Actinocephalus n.g. Nematodes of Puerto Rico 11

4. Lip-region with sclerotized disk, no teeth or denticles Mylodiscidae n.£. A single genus, Mylodiscus Lip-region not a sclerotized disk, teeth or denticles usually present 5 5. Tails·rounded or bluntly conoid 7 Tails of both sexes elongate conoid to filiform Trachypleurosidae n.£. 6 6. Pharynx with several rows of transverse denticles Trachypleurosum Pharynx with simple rodlike elements Actinolaimoides 7. Lateral fields with 2 lines of cells, pores usually obscure 8 Lateral fields with 1line of cells, pores prominent Carcharolaimidae n.£. 12 8. Pharynx with 4 large onchia or a fused spear-guide, few or no denticles Neoactinolaimidae n.£. 9 Pharynx with 4 large onchia, and many denticles Paractinolaimidae n.£. 11 9. Postextension constriction of esophagus absent Egtitus n.g. Postextension constriction of esophogus present 10 10. Onchia fused into ringlike spear-guide Metactinolaimus Onchia not fused into ringlike spear-guide Neoactinolaimus n.g. 11. Pharyngeal cavity massively ribbed Westindicus n.g. Pharyngeal cavity with obscure ribbing Paractinolaimus 12. Posterior pharynx narrow, small onchia, plates or denticles only Carcharolaiminae n.sf. 13 Posterior pharynx broad with large onchia, no denticles Caribenematinae n.sf. A single genus, Carib enema n.g. 13. Pharyngeal cavity with simple plates only Antholaimus Pharyngeal cavity strongly sclerotized bearing onchia or denticles 14 14. Tail blunt, rounded Carcharolaimus Tail bluntly conoid, slightly arcuate Carcharoides n.g.4

4The reader is referred to Thorne (20) for descriptions and illustrations of most of the genera and species not included in this paper. 12 Gerald Thorne

ACTINOLAIMOIDEA, NEW SUPERFAMILY Synonym: Actinolaimidae (Thorne, 1939) M eyl, 1960 Diagnosis: DORYLAIMIDA: Lip region with sclerotized pharyngeal walls varying from simple plates (Antholaimus) to strong, refractive basketlike or ribbed structures (Carcharolaimus), frequently accompanied by 4 large onchia which may be fused into a spear guide (Metactinolaimus) or accompanied by numerous other mural denticles (Pamctinolaimus). Cuticle with longitudinal striae in Actinolaimidae or plain with minute, transverse striae in other families. Esophagus enlarged near middle, the anterior portion either with well developed radial striae or a slender nonmuscular tube. A strong basal shield frequently surrounding base of esophagus. Lateral fields with one or two lines of cells from which rise 1, 2, or 3 series of spores which may be so reduced as to be almost invisible, or with conspicuous tubelike canals. Type family: Actinolaimidae (Thorne, 1939) Meyl, 1960 FAMILY ACTINOLAIMIDAE (THORNE, 1939) MEYL, 1960 Diagnosis emended: ACTINOLAIMOIDEA: Cuticle with longitudinal striae. Lateral field with 2 series of cells, their pores usually minute and difficult to see. Vestibule corrugated. Pharynx armed with 4 massive onchia, no denticles present. Basal shield present at esophagus base. Ovaries 1 or 2. Vulva longitudinal or porelike. Supplements in 2 or 3 fascicles (bundles), occasionally with one or more larger ventral supplements between the fascicles. Female tail conoid to filiform; male tail blunt, rounded with numerous lateral and ventrosubmedian papillae. Type subfamily: Actinolaiminae Thorne, 1939. Subfamily Actinolaiminae Thorne, 1939 Diagnosis emended: Actinolaimidae: Body cylindroid except at extremities. Cuticle striated. Pharynx with 4 massive onchia. Male tail blunt, rounded with supplements in fascicles. Female tail conoid to subfiliforrn. Spear with aperture occupying 1fz or more of its length. Constriction present at beginning of esophagus. Anterior portion of esophagus with radial muscles.

Type genus: Actinolaimus Cobb, 1913, the only genus. GENUS Actinolaimus COBB, 1913 Diagnosis emended: Actinolaiminae. Characters of the subfamily. Key to species of Actinolaimus 1. Number of body striae 16 papillatus Schneider, 1935 2.7 mm.; a - 45; b - 4.6; c - 19.1; V - 16 45 15 Nematodes of Puerto Rico 13

2.3 mm.; a-50; b - 5.2; c - 84; Number of body striae 24 - 32 2 2. Anterior ovary rudimentary, vulva 24 - 30 percent 2 Anterior ovary normal, vulva 40 - 46 percent 4 3. Number of body striae 24 straelini Meyl, 1957 5.0 mm.; a - 70; b - 5.6; c - 20; V - 7 24-30 ~8 4.1 mm.; a - 60; b - 5.0; c - 173 (145-208) Number of body striae 32 radiatus Cobb, 19135 5.0-7.0 mm.; a - 83; b - 6.6; c - 18; V - 4 25 42 5.5-6.5 mm.; a - 100; b - 6.6; c - 250; T - 78 4. Length 2.0 mm. schuurmansstekhoveni Meyl, 1957 2.0 mm.; a - 35; b - 4.0; c - 11.7; V - 8 39 9 Length 3.4 - 3.9 mm. costatus Schneider, 1935 3.9 mm.; a-54; b - 5.6; c - 17.7; V - 8 39 9 3.4 mm.; a-57; b - 4.9; c - 70-101; T - 68 Species of the genus Actinolaimus have not been found in Puerto Rico.

SUBFAMILY BRITTONEMATINAE, NEW SUBFAMILY

Diagnosis: Actinolaimidae. Body fusiform with very narrow head continuous with neck contour ( Brittonema ) or broadly expanded (Actinocephalus). Striae formed by deep furrows in the thick cuticle. Spear slender, 2 or 3 times as long as head, its aperture occupying 1/3 or less of length. Anterior portion of esophagus a slender hyaline tube. Basal shield of esophagus a prominent feature. Cardia elongate conoid. Female tails spicate to filiform. Male tail bluntly rounded but bearing a filiform extension. Supplements in fascicles. Type genus: Brittonema, n.g.

KEY TO GENERA OF BRITTONEMATINAE 1. Head continuous with neck contour Brittonema n.g. Head strikingly expanded Actinocephalus n.g. Brittonema, n.g. Diagnosis: Brittonematinae: Body fusiform. Head continuous with neck contour. Spear twice or more as long as head-width. Anterior portion of esophagus a slender tube. Female tails spicate to filiform. Male tails bluntly rounded with filiform extensions. Supplements in fascicles.

5Andhassy. 1954 made A. radiatlls a synoym of A. tripapillatlls, but this act is not here accepted. 14 Gerald Thome

Type species: Brittonema sulcatum, n.sp. Key to species of B1'ittonema 1. Striae often not adjacent, forming bands in cuticle fusifo1'mum n.sp. Striae adjacent, fonning sharp ridges in cuticle 2. Length 3.7 mm. taylori (Meyl, 1957) n.comb. a - 32; b - 4.0; c - 13.4; V - 44 Length well under 3.0 mm. 3 3. Head rounded, lips obscure 1.1-1.9mm.; a - 41; b - 3.9; c - 9.3; V - 45 tenuiaculeatus (Kreis, 1924) n.comb. Head truncate, angular 4 4. Female tail spicate spicatum n.sp. Female tail elongate filiform sulcatum n.sp. B1'ittonema sulcatum, n.sp. (Fig. 1, A-G) 2.1 mm.; a - 41; b - 4.1; c - 4.6; V - 9 41 11 1.9mm.; a - 45; b - 4; c - 9; T - 60 Body cylindroid from anterior third of neck to near anus. Neck tapering near anterior third for a short distance, then almost cylindroid for about 3 neck-widths when it gradually tapers to the lip-region which is only % as wide as neck-base. Tails of both sexes long, filifonn. Cuticle marked by deep longitudinal grooves with sharp ridges between them, about 40 being present at midbody. Minute transverse striae visible. Lateral fields about 1/6 as wide as body, pores not observed. Lip-region truncated, set off by slight depression, its papillae difficult of resolution. Amphid almost as wide as head, beaker-shaped, with sensillae about opposite spear base. Vestibule with minute corrugations. Pharynx with 4 strong onchia. Spear slender, 20 ft long, the aperture occupying 1/5 its length. Guiding-ring double. Spear extensions 23 ft long, heavily sclerotized. Esophagus a slender nonmuscular tube continuing back through nerve ring until it forms a fusiform hyaline enlargement, then expanding to the basal cylindroid portion with strong radial musculature and glandular tissues. Usually only the dorsal and two submedian gland nuclei visible. Nerve-ring conspicuous with many associated cells and ganglia similar to those of B1'ittonema spicatum (fig. 2,G). Cardia elongate conoid completely submerged into intestinal cells. Well-developed shield surrounding base of esophagus. Intestinal cells filled with rather unifonn light-brown granules. Ovaries reflexed half way ~ack to vulva. Vulva apparently a tiny elongate slit at bottom of a conoid depression in cuticle. Eggs 30 x 70 ft. Prerectum length 31f2times body width. Two pairs of lateral caudal pores. Nematodes of Puerto Sica 15

FIG. l.-Brittonema sulcatum: A, Head x 1,500; B, male tail x about 350; C, expansion region of esophagus; D,E, female posterior portion of body and terminus; F, esophagus base, basal shield, cardia, and anterior end of intestine; G, vulva region and vagina. All about 500 x. 16 Geralcl Thome

Male more cylindroid than female. Testes dorylaimoid. Posterior portion of body slightly bent ventrally. Tail dorsally convex conoid then elongate filiform to pointed terminus. This filiform terminus begins as a narrow isthmus, then expands slightly. Four pairs of caudal papillae seen. Supplements in 2 fascicles of 8 each, their ampullae extending almost halfway across body. Adanal pair of supplements unusually large, mammiform. Six pairs of ventrosubmedian papillae present. Spicula massive, arcuate, with slender lateral guides. Brittonema sulcatum is immediately distinguished by the peculiar tapering of the neck, fusiform hyaline bulb of esophageal tube just anteroir to basal enlargement, and unusually long, filiform tails of both sexes. Holotype female, allotype and other specimens as indexed under Brittonema 2. From forest and stream bank soil along Mount Britton Trail; EI Yunque Mountains, Luquillo Experimental Forest. Brittonema fusiformum, n.sp. (Fig. 2, A-C) 2.3 mm.; a - 35; b - 4.3; c - 11; V - ]3 42 1:1 Body fusiform, tapering rapidly to the lip-region which is only 1/6 as wide as neck-base. Posteriorly it ends in a subfiliform elongated tail. Cuticle near midbody marked by about 24 deep longitudinal striae, or grooves, which in this region produce an alternating series of bands and grooves, the bands being about twice as wide as the grooves. Anteriorly and posteriorly some of the bands become narrowed and disappear. Lateral pores not observed. The high, somewhat rec- tangular lip-region is a distinctive character. Cephalic papillae difficult of resolution. Pharyngeal cavity with four onchia, no denticles present. Guiding ring double. Spear unusually slender, 22 0 long its aperture occupying about 1/5 its length. Spear extensions 33 0 long. Amphids goblet shaped, with sensillae located just posterior to guiding ring. Esophagus a~ first a narrow, slender, nonmuscular tube which continues through the massive nerve-ring, finally joining the tapering posterior portion. Basal portion of esophagus strongly muscular, containing elaborate glandular network. Only the dorsal and 2 submedian gland nuclei observed. Cardia elongate conoid but this and the basal shield are compressed until accurate details are lost. Intestinal cells densely packed with tiny dark-brown granules. Ovaries reflexed half way to vulva. Eggs 45 x 90 0, 3 being present in the 1 gravid female seen. Uteri packed with spermatozoa. Prerectum length twice body width. Details of convex conoid, elongate tail as illustrated (fig. 2,B). Only 1 pair of caudal papillae observed. A portion of the filiform terminus may be missing. Male not collected. Holotype female as indexed under Brittonema 1. From moss on tree trunks, Mount Britton, EI Yunque Mountains, Luquillo Experi- mental Forest. Nematodes of Puerto Rico 17

F c

: : : . ·····'····· . • . , .' C : : . : :. mDJ· : .:

FIG. 2.-A-C, Brittonema fusiformum: A, Head x 1,500; B, posterior portion of female; C, section of cuticle showing grooves. Both about 500 x. D-G, Brittonema spicatum: D, Head x 1,500; E,F,G, posterior portion of female, vulva-vagina region, and nerve-ring and esophageal expansion region. 18 Gerald Thorne

Brittonema spicatum, n.sp. (Fig. 2, D-G) 1.8 mm.; a - 26; b - 4.6; c - 8.5; V - 12 42 15 Body fusiform, lip-region 1/5 width of neck-base. Cuticle marked by about 30 longitudinal grooves at midbody forming sharp ridges in cuticle. Lateral field 2 cells wide, occasional pores observed in 2 lines, the pores emerging at crest of ridges. Cephalic papillae very obscure, sometimes not seen. Amphids beaker-shaped, almost as wide as head. Vestibule with very minute corrugations, scarcely visible on some specimens. Pharynx armed with 4 strong onchia. Spear slender, 15 po long. Guiding-ring double. Esophagus a slender hyaline tube until it passes through nerve-ring, then becoming about twice as wide before joining basal enlarged portion. Only dorsal and 2 submedian gland nuclei usually visible. Nerve-ring unusually distinct with many cells surrounding it. Base of esophagus with shallow shield. Cardia elongate conoid, surrounded by intestinal cells. Intestine packed with fine light- brown granules. From lateral view, vulva assumed to be a short longitudinal slit. Vagina with massive musculature. Ovaries reflexed almost to vulva. Eggs 1h as wide as body, twice as long as wide. Prerectum length twice body-width. Tail conoid to elongate apicate. Males unknown and gravid females did not contain spermatozoa. This species is immediately distinguished by the rather obese fusiform body, apicate tail and monosexuality. Holotype female and other specimens as indexed under Brittonema 2. From moss and soil. Coca Falls, El Yunque Mountains, Luquillo Experimental Forest. Two males were found which resemble the above described females but they had blunt rounded tails, not filiform as in B. sulcatum. Supplements in 2 fascicles. It appears advisable to refrain from assign- ing them to B. spicatum until additional collections are made.

GENUS Actinocephalus, n.g. Diagnosis: Brittonematinae. Characters of the subfamily, but head greatly expanded and spear longer than in other members. Type species: Actinocephalus bizarrus, n.sp.

A.ctinocephaus bizarrus, n.sp. (Fig. 3, A-F) 2.8 mm.; a - 28; b - 3.8; c - 6.3; V - 10 43 10 Body fusiform, tapering to narrow head which is only % as wide as neck-base, posteriorly ending in a conoid filiform tail. Cuticle 5 fL thick, marked by about 100 longitudinal grooves at midbody, which Nematodes of Puerto Rico 19 decrease in numbers toward the extremities, about 50 present in the anal region. Lateral fields 115 body width, 2 cells wide but only 3 lateral pores seen rising from them. If others are present they are be- yond the limits of visibility. Lip-region set off by an unusual expansion and bearing a complicated assembly of sclerotized mouth parts (fig. 3,A). Usual circlets of 6 and 10 cephalic papillae easily visible. Amphids about 1fz as wide as head, their apertures obscure slits beneath contour of lateral lips. Vestibule corrugated, leading to a spacious pharynx with 4 strong onchia. Posterior extension of labial framework an unusual feature, extending back to guiding ring a distance of about 20 µ.. Spear 47 µ. long but only 2 µ. wide, its aperture occupying about VB its length. Extensions 80 µ. long, heavily sclerotized. Esophagus a slender tube with minute radial striae. After passing through nerve-ring it forms an elongate bulb which rather abruptly joins the basal enlargement which occupies 1fz the total length. Radial musculature of esophagus very fine interwoven through masses of glandular structures. Only 3 esophageal gland nuclei visible. Nerve-ring hyaline, obscure, with numerous cells grouped about it. Basal shield of esophagus moderately developed. Cardia elongate conoid, surrounded by intestinal cells. Intestine with minute granules. Vulva apparently longitudinal; vagina extending more than halfway across body. Uterine tract with many cells and conspicuous valvular apparatus. Posterior ovary illustrated (fig. 3,C) was in the process of producing syngonic sperms before beginning egg production. These were stored in an elongate sac at flexure of uterus. No males observed and no true spermatozoa present in uteri. Pre rectum length 4 times body-width. Rectum about twice as long as anal body diameter with conspicuous valvular apparatus at its junction with prerectum. Four pairs of caudal pores observed, all opening on crests.of longitudinal ridges. Tail bent dorsally, uniformly tapering to a minute rounded terminus which bears a tiny point. The species is immediately distinguished from all other actinolaims by its unusual expanded head, pharyngeal armature and long spear. Holotype female as indexed under Actinocephalus 1. From moss on trees by trail near summit of Mount Britton, El Yunque Mountains, Luquillo Experimental Forest. A male of this genus was collected in another locality on Mount Britton, but since A. bizarrus is syngonic additional collecting is necessary to determine if they reoresent the same or different species. Supplements in two fascicles, filiform terminus about half as long as that of Brittonema sulcatum.

FA:MILY NEOACTINOLAIMIDAE, NEW FAMILY Diagnosis: ACTINOLAIMOIDEA: Cuticle with only minute transverse striae, not longitudinally striated. Lateral field with 2 lines of cells from which excessively minute fibers rise. Pharynx armed with

Nematodes of Puerto Rico 21

4 massive onchia but without numerous denticles. Postextension constriction of esophagus present or absent. Esophagus with or without basal shield. Female tails elongate, male bluntly rounded. Supplements in fascicles or a ventral series. Type genus: Neoactinolaimus, n.g. Diagnosis: Neoactinolaimidae. Cuticle without longitudinal striae. Pharynx armed with 4 large onchia, no denticles present. Postextension constriction of esophagus present. Anterior portion of esophagus with radial muscles. Ovaries, 2. Female tail elongated to a filiform terminus, male tail blunt, rounded. Supplements in fascicles. Type species: Neoactinolaimus agilis, n.sp. Members of Neoactinolaimus are immediately distinguished by the absence of longitudinal striae, presence of a postextension constriction of the esophagus and fascicled supplements. Occasionally a few ventral supplements are found between the fascicles.

Key to species Neoactinolaimus 1. Length between 5.0 and 6.0 mm. . .. ' 2 Length less than 4.0 mm. 3 2. Very slender species, a - 83 africanus (Filipjev, 1929) 5.8 mm.; a - 83; b - 5.8; c - 13.8; V - 8 27 28 More robust species, a-50 tripapillatus (Daday, 1905) 5.6 mm.; a-51; b - 4.8; c - 16; V - 27 5.4 mm. a-50; b - 4.5; c - 108 3. Spear twice as long as head width michaelsini (Steiner, 1916) 2.7 mm.; a - 60; b - 4.3; c - 124; T - 70 Spear distinctly less than twice head width 4 4. Small tooth back of each onchium agilis n.sp. Small tooth not present back of each onchium 5 5. Vulva at 69 percent ...... hutchinsoni (Filipjev, 1929) 2.0 mm.; a - 41; b - 3.3; c - 8.5; V - 25 69 25 2.3 mm.; a-56; b - 3.8; c - 113 Vulva at 38 to 50 percent 6 6. Length well over 3.0 mm. 7 Length well under 3.0 mm...... 8 7. Esophagus expansion a gradual tapering chitwoodi (MoOl·thy, 1937)

3.4 mm.; a-57; b - 5; c - 13.7; V - 17 40 2!l Esophagus with abrupt expansion. frostae (Allgen, 1952) 3.9 mm.; a-52; b - 4.3; c - 13.9; V - 47 3.6 mm.; a-50; b - 4.0; c - 125; T - ? 22 Gerald Thome NE

8. Robust species, a - 30 hintoni (Allgen, 1952) 2.0 mm.; a - 30; b - 4.3; c - 8; V - 45 1.5 mm.; a - 26; b - 3.3; c - 76; T - 73 More slender species, a-50 - 70 9 9. Vulva at 38 percent tenuis (W. Schneider, 1935) 2.8 mm.; a - 67; b - 5.3; c - 11.8; V _ 16 38 21 1.9 mm.; a-57; b - 4.0; c - 115; T _ ? Vulva at 50 percent zeelandicus (Clark, 1962) 2.6 mm.; a-50; b - 4.6; c - 16.5; V _ 14 51 17 2.6 mm.; a - 46; b - 4.6; c - 98; T - 27

Neoactinolaimus agilis, n.sp. (Fig. 4, A-G) 3.0 mm.; a - 67; b - 5.6; c - 17; V _ 16 4316 3.5 mm.; a - 78; b - 6.4; c - 140; T - 64 . ;1'::" Bodies of both sexes cylindroid throughout their length, except at the extremities. Cuticle without visible markings. Lateral fields % body- width, 2 cells wide, each cell with scattered variable sized granules and a nuclear body. Lateral pores not observed. Head set off by slight narrowing of neck. Lips low, rounded, their paillae not interfering with head contour. Amphids goblet-shaped, half as wide as head with sensillae just posterior to spear base. Vestibule a sclerotized ring without corrugations. Pharynx armed with 4 large onchia and 4 very small submedian teeth, unlike the numerous denticles of paJ'actinolaimus. Spear 22 µ.long, its aperture occupying 2/5 the length. Guiding- ring double. Spear extensions about as long as spear, surrounded by a hyaline muscular tissue set off from esophagus by a constriction. FIG. 4.-Neoactinolaimll Anterior portion of esophagus with strong radial striae, about % neck- plement fascicl field; F, vulv; width, slightly narrowing as it passes through the nerve-ring, then anterior end 0 gradually increasing in width to join the strongly muscled basal portion which is about half the neck-length. Five esophageal gland nuclei The small tooth b( usually visible. No basal shield present. Cardia a muscular disk with distinctive. an elongate tapering portion reaching deep into intestine, the discoid portion not surrounded by intestine. Intestine about 4 cells in circum- Holotype, allotype ference, the cells containing scattered granules of variable size. Vulva actinolaimus 1. From ' apparently porelike. Ovaries reflexed 3/5 distance to vulva, long tural Experiment Static slender with numerous oocytes. Prerectum 4 times body-width. Rectum GENDi length about twice anal body-width. Female tail uniformly conoid to filiform terminus. Head set off by sl Male with blunt rounded tail. Supplements in 2 fascicles, each with rugated. Onchia fused 6 to 8 innervations and traces of a third rudimentary group sometimes elements where postel visible as a slight elevation in the body contour. Tail with 5 pairs of guide. Spear 1% times lateral papillae. Ventromedian papillae and lateral pores not seen. 2/5 its length. Postext< Spicula and guides dorylaimoid. at 23-25 percent. Fern Male tail bluntly roune

24 Gerald Thome

Type species: Metactinolaimus kreisi Meyl, 1957. 5. Esophagus gr: Synonym: Actinolaimus tTipapillatus of Kreis, 1932. (Not Daday, 1905). 2.9mm.; a 2.6mm.; a 4.0-8.7 mm.; a - 58-73; b - 5 - 7; c - 17 - 25; V - 26 - 35 Esophagus wi 4.7-8.1 mm.; a - 52-78; b - 4 - 6; c - 125 - 195 6. Female tairta These measurements from Kreis who studied 13 females, 36 males and 53 young. The variations in size and proportions are unusual, if 2.2mm.; a all were mature specimens. Female tail de Goodey (13) suggested that the fused onchia forming a spear-guide 7. Maile tail hem may be an incorrect interpretation of Krei's illustrations. However 2.6mm.; a Meyl described and illustrated M. leloupi as having similar structures. 1.9 mm.; a Male tail blun GENUS Egtitus, N.C. Diagnosis: Neoactinolaimidae. Cuticle with only minute transverse striae. Pharynx armed with 4 large onchia but no denticles. Post- extension constriction of esophagus absent. Anterior portion of esopha- 2.0 mm.; a - 4L gus with fine radial muscles. Ovaries 2. Female tail elongated, male 1.9 mm.; a - 4~ tail blunt, rounded. Supplements in ventral series. Bodies practic: Members of Egtitus are immediately distinguished from Neo- actinolaimus by the absence of a postextension constriction of the minute transverse sl esophagus and ventral series of adjacent or closely spaced supplements. 2 lines of altematir Named in honor of Dr. E. G. Titus who advised the writer to enter which excessively n nematology. by slight depressiOl with 3 sclerotized e Type species: Egtitus bryophilus n.sp. tions. Pharynx anne and pharyngeal am Keys to species of Egtitus half as wide as hei length. Esophagus a 1. Lip-region conspicuous expanded (Daday, 1905) expanding to 1/3 n cyatholaimus ring, then suddenly 2.4. mm.; a - 33; b - 3.2; c - 46; T _ ? portion somewhat i Lip-region not conspicuous expanded 2 massive glandular 2. Spear-length 1% times head-width nuclei seen. A low propinquus (M. V. Cobb, 1915) a cylindroid isthml 2.4 mm.; a-50; b - 4.1; c - 11; V _ 1347 14 cells % its length. A Spear-length equal to or only slightly more than head-width 3 at base of esopha~ Vulva a short longit 3. Length under 2.0 mm. 4 Ovaries reflexed, bt; Length over 2.0 mm. 5 body-width. 4. Pharynx only slightly sclerotized surinamensis (Mico!., 1925) Male slightly 11 1.4 mm.; a - 38; b - 3.3; c - 63; T - 47 Supplements an ada Pharynx massively sclerotized elaboratus (Cobb, 1906) pairs of minute sub 1.8mm.; a - 36; b - 4.1; c - 14.3; V _ 1650 11 reaching about 1 bo arcuate with slende Nematodes of Puerto Rico 25

1957. 5. Esophagus gradually tapering posteriad ;, 1932. (Not Daday, nudus (Wu & Hoeppli, 1929) 2.9 mm.; a - 45; b - 4.4; c - 11; V _ 18 38 20 2.6 mm.; a - 37; b - 4.9; c - 129; T - ? ; V - 26 - 35 Esophagus with definite expansion near middle 6 95 6. Female tail tapering uniformly from in front of anus 13 females, 36 males neocyatholaimus (Kreis, 1936) tions are unusual, if 2.2 mm.; a - 32; b - 4.8; c - 9.4; V - 19 42 30 Female tail dorsally convex conoid, then filiform 7 )rming a spear-guide 7. Maile tail hemispheroid. occalescens (W. Schneider, 1937) ustrations. However 2.6 mm.; a - 37; b - 3.4; c - 10.4; V _ 10 51 7 19 similar structures. 1.9 mm.; a - 28; b - 3.7; c - 64; T - ? Male tail bluntly conoid bl'yophilus n.sp.

Egtitus bl'yophilus, n.sp. ly minute transverse no denticles. Post- (Fig. 5, A-E) '1' portion of esopha- 2.0 mm.; a - 44; b - 4.0; c - 13; V _ 11 50 11 tail elongated, male 1.9 mm.; a - 48; b - 4.2; c - 72; T - 59 Bodies practically cylindroid except at extremities. Cuticle with guished from Neo- minute transverse striae. Lateral fields % body-width at midbody, with constriction of the 2 lines of alternating cells. Each cell has a tiny glandular body from ;paced supplements. which excessively minute fibers arise to the surface. Lip-region set off the writer to enter by slight depression. Cephalic papillae minute but visible. Vestibule with 3 sclerotized elements, the posterior one bearing minute corrugations. Pharynx armed with 4 pointed onchia, no denticles. Guiding ring and pharyngeal armature as illustrated. Amphid goblet shaped, about half as wide as head. Spear 16 µ. long, the aperture occupying % its length. Esophagus at first about % neck width with fine radial muscles, 15 ) expanding to 1/3 neck-width for a short distance posterior to nerve- cyatholaimus ring, then suddenly expanded to more than % neck-width. Expanded portion somewhat irregular in width with strong radial muscles and 2 massive glandular tissues. Only the dorsal and 2 submedian gland nuclei seen. A low shield surrounds esophagus base. Cardia at first 1. Cobb, 1915) a cylindroid isthmus then elongate conoid, extending into intestinal 14 cells 1/2 its length. A conspicuous gland is located left dorsosubmedian at base of esophagus. Intestinal granules pale brown or colorless. 1 head-width 3 Vulva a short longitudinal slit at base of a shallow conelike depression. 4 Ovaries reflexed, but females immature. Prerectum length about twice 5 body-width. (MicoI., 1925) Male slightly more slender than female with conoid, blunt tail. Supplements an adanal pair and 7 ventrosubmedian, as illustrated. Four (Cobb, 1906) pairs of minute submedian, and 3 pairs of caudal papillae. Prerectum 111 reaching about 1 body-width anterior to supplements. Spicula strongly arcuate with slender guides. 26 Gerald Thotnt!

B c

FIC. 5.-Egtitus bryophillls: A, Head x 1,000; B, amphid; C, posterior portion of female; D, posterior portion of male; E, esophagus base, basal shield, cardia, dorsosubmedian gland, and intestine.

Holotype female, allotype and other specimens as indexed under Egtitus l. From moss on tree trunk beside trail near summit of Mount Britton, El Yunque Mts., Luquillo Experimental Forest.

FAMll..Y PARACTINOLAIMIDAE, NEW FAMll..Y Diagnosis: Actinolaimoidea. Cuticle with only fine transverse striae. Pharynx armed with 4 massive onchia which in Metactinolaimus are fused into a spear guide. Walls of vestibule with or without denticles. Postextension constriction of esophagus present. Anterior portion of esophagus with radial musculature. Ovaries 2, reflexed. Female tail elongated, male tail bluntly rounded with supplements in a ventral series. Type genus: Pamctinolaimus Meyl, 1957. Nematodes of Puerto Rico 27

Key to genera of Paractinolairnidae 1. Onchia fused into a massive spear-guide, no dentides present Metactinolaimus Onchia not fused into a ring, dentides present 2 2. Vestibule with massive basketlike ribbing Westindicus n.g. Vestibule without basketlike ribbing Paractinolaimus

GENUS Paractinolaimus MEYL, 1957 Diagnosis emended: Pharynx armed with 4 massive onchia which may appear to be fused into base of vestibule, sometimes appearing as an archlike structure. Walls of vestibule with numerous dentides rising from slightly ribbed plates. Postextension constriction of esophagus present. Ovaries 2. Supplements an adanal pair and a ventral series. Female tail elongated, male tail bluntly rounded. Type species: Paractinolaimus micoletzkyi (W. Schneider, 1935) Meyl, 1957.

Key to species of ParactinolaimusG 1. Female tail very long, filiform. filipjev (W. Schneider, 1935) 3.3 mm.; a - 43; b - 4.5; c - 6.8; V _ 9 45 9 Female tail elongated but c - 11 or less 2 2. Dentides 40 - 50, relatively sparse 3 Dentides very numerous, more than 100 4 3. Length about 4.0 mm. ormore, a-50 macrolaimus (de Man, 1880) 4.5 mm.; a-50; b - 5; c - 11.5; V _ 16 47 14 3.7 mm.; a-50; b - 5; c - 85; T - ? Length about 3.0 mm.; a - 37 microdentatus (Thorne, 1939) 3.2 mm.; a - 37; b - 4.3; c - 10.7; V _ 20 46 15 4. Vestibule almost as deep as wide baldus n.sp. Vestibule about twice as wide as deep 5 5. Onchia projecting into vestibule vigor n.sp. Onchia located at base of vestibule robustus n.sp.

6Pamctinolaimlls striatlls (Thorne, 1939) Meyl 1957 remains in this genus until additional collecting reveals its true relationships. 28 Gerald Thorne

Paractinolaimus vigor, n.sp. (Fig. 6, A-D)

2.0 mm.; a - 34; b - 3.7; c - 14; V _ 12 53 12 Lip-region set off by expansion, the lips angular and bearing the usual circlet of 6 and 10 papillae. Vestibule minutely corrugated. Pharynx with 4 massive onchia joined to a somewhat bowed basal section marked by minute spiral corrugations, reaching back to a massive, sclerotized guiding-ring. Spear about 28 p. long, its aperture occupying about % its length. Spear extensions almost as long as spear, heavy sclerotized. A conspicuous constriction in the anterior esophagus about 25 p. posterior to spear extensions. Esophagus at first a slender tube with radial muscles, then enlarged by irregular expansion to join the massive basal portion which occupies 3/5 of its length. This enlarged portion has strong radial muscles and conspicuous glandular tissues. Only the dorsal and 2 submedian gland nuclei usually visible. Two small glands sometimes visible in anterior tapering portion of esophagus. Cardiae region with a basinlike epithelial shield surrounding base of esophagus and an elongated cardia as illustrated (fig. 6-D). Intestinal cells packed with minute brown granules. Vulva a short longitudinal slit at bottom of a conical depression in cuticle. Ovaries reflexed about halfway to vulva. Males unknown and mature females contained no spennatozoa. This species is distinctive because of the broad, angular lip-region, many pharyngeal denticles, slightly ribbed pharyngeal chamber, spiral corrugations on posterior pharynx, and tail shorter than those of re- lated species. Holotype female and other specimens as indexed under Paracti- nolaimus 1.

Paractinolaimus baldus, n.sp. (Fig. 6, E-H)

2.6 mm.; a - 46; b - 4.2; c - 11; V - 14 52 14 2.9 mm.; a - 49; b - 4.3; c - 84; T - 53 Body cylindroid except at extremities. Lateral pores begin immediately behind the head, then shift to form a single line originating on the dorsal slide of the cells of the lateral cord. A short distance anterior to the supplements they also are found on the ventral side. However, these should not be confused with the ventrosubmedian papillae found on males of many groups of dorylaims. Lip-region rounded set off by shallow depression, papillae not interfering with head contour. Pharynx bearing 4 huge onchia and many minute denticles. Spear 27 p. long with aperture occupying about half its length. Guiding ring massive, sclerotized. A constriction in anterior esophagus as shown for Pamcti- Nematodes of Puerto Rico 29

FIG. 6.-A-D, Pamctinolaimus vigor: A, Head and antel'ior portion of esophagus x 1,000; B, expansion region of esophagus; C, posterior portion of female; D, base of esophagus, basal shield, cardia, and anterior intestine. E-H, Pamctinolaimus baldus: E, Head x 1,000; F, posterior portion of female; G, vulva-vagina region; H, posterior portion of male. 30 Gerald Thome nolaimus vigor (fig. 6,A). Anterior portion of esophagus with weak radial muscles, gradually expanding to the basal portion which is about 2/3 wide as body and bearing a basal shield. Cardia slender conoid, longer than body-width. Intestinal cells densely packed with minute brown granules. Vulva a short longitudinal slit or porelike, located in a conical depression. Ovaries reflexed 3/5 back to vulva, the uteri packed with spermatozoa. Prerectum length 2% times body- width. Tail elongate conoid, usually slightly bent dorsad. Male slightly more slender than female. Supplements 10-11, very low and obscure, arranged as illustrated. Six pairs of ventrosubmedian papillae. Prerectum extending almost to anterior supplements. Spiculae elongated and slender with simple lateral guides. Diagnosis: Paractinolaimus with above measurements and general description. Distinctive because of the rounded head set off by gradual depression, pharyngeal armature, form of tails, arrangement of supplements and slender elongate spicula. Holotype female and other specimens as indexed under Paracti- nolaimus 2. Stream bank soil from Coca Falls, El Yunque Mountains, Luquillo Experimental Forest.

Paractinolaimus robustus, n.sp. (Fig. 7, A-E)

3.0 mm.; a - 43; b - 5.4; c - 11; V - 11 50 11 2.8 mm.; a - 47; b - 4.8; c - 71; T - 62 Body cylindroid throughout most of its length, tapering gradually near extremities. Lateral fields about 1/3 body-width without definite cellular structures. Occasionally a tiny obscure ampulla can be seen rising to the surface from the dorsal side of the field. Lip-region set off by a shallow depression. Cephalic papillae rising slightly above head contour. Entrance to vestibule corrugated. Instead of the usual 4 large onchia, the pharynx contains a sclerotized arch which doubtless represents the 4 onchia fused together. Minute denticles are distributed throughout the lower half of the pharynx. Spear 23 µ, long, the aperture occupying about 2/5 its length. Pharyngeal walls marked by ridges forming obscure patterns. Guiding-ring massive, refractive. Postexten- sion constriction of esophagus present. Spear extensions well sclerotized. Esophagus at first a slender tube with distinct radial musculature until it expands by gradual enlargement leading to the basal portion which is made up of strong radial muscles and extensive glandular tissues. Basal shield of esophagus strongly developed. Cardia elongate conoid. Intestinal cells packed with small, uniform dark-brown granules. Vulva, a pore or short slit located at the bottom of a conelike depression. Vagina extending about halfway across body. Ovaries reflexed

32 Gerald Thorne about 3/5 distance back to vulva. Prerectum length 3 times body-width, rectum almost half as long as prerectum. Tail rather uniformly conoid to a fine-pointed terminus. A single pair of caudal pores seen. Male with typical spicula and simple guides. Supplements 10, very low, obscure, with conspicuous ampullae extending back into body. A series of 7 ventrosubmedian papillae are easily seen. Anterior to these are numerous other obscure ventral papillae extending as far forward as the testes region, their presence usually marked by a small depression in the body tissues. Caudal papillae 6 pairs, a very distinguishing feature of the species. Holotype female, allotype, and other specimens as indexed under Paractinolaimus 3. From dieback area on EI Yunque Mountain and moss along trail to Mount Britton, Luquillo Experimental Forest. The position of this species may.be questioned since the onchia are at least partly fused into an archlike structure which is best seen from a lateral view. When viewed dorsally the arch is not so complete. Because of the numerous dentides it certainly does not belong in Metactinolaimus and may actually represent a distinct new genus. One female contained two larvae within the body cavitv, apparently a mermithid or a similar group of parasites. The parasitized indi- vidual was only 2.0 mm. long and the ovaries had failed to develop so apparently the parasites had both a stunting and a sterilizing effect. A description of this parasite appears at the end of the Actinolai- moidea section. (See Nannomeris infestans, n.sp.).

GENUS Westindicus, N.C.

Diagnosis: Paractinolaimidae. Lip-region 2/3 as wide as neck- base, set off by slight depression. Vestibule with 36 sderotized ribs forming a basketlike structure. Walls of vestibule bearing scores of minutely iagged den tides. Four massive, pointed onchia projecting into vestibule. Spear guide massive. Esophagus with postextension constriction and deeply cupped basal shield. Ventromedian supplements very obscure, spaced in a series. Ventrosubmedian papillae present. Type species: Westindicus brachycephalus, n.sp. Westindicus is immediately recognized by the unusually wide head, heavily ribbed vestibule, massive, forward pointing onchia, numerous jagged dentides, and obscure, well spaced supplements. Only 1 species known. Nematodes of Puerto Rico 33

Westindicus bmchycephalus, n.sp. (Fig. 8, A-G)

2.3 mm.; a - 41; b - 3.8; c - 9-14; V - 14 53 14 2.9 mm.; a - 44; b - 3.8; c - 82; T - 62 Lip-region about 2/3 as wide as neck-base. Lateral pores very obscure and difficult to observe, rising from dorsal side of lateral fields except in rare instances when they originate on ventral side. Cephalic papillae low, rounded, not interfering with head contour. Vestibule with minute corrugations. Pharynx elaborately ribbed, its lower walls bearing great numbers of minute denticles. Four massive onchia have their basal elements attached directly to the strong, refractive guiding ring. Spear 15 /-'-long, its aperture occupying slightly more than half its length. Spear extensions and esophageal lumen heavily sclerotized. Postextension constriction of esophagus present. Esophagus at first 1/4 as wide as neck, narrowing slightly as it passes through nerve-ring, then gradually becoming wider until it rather abruptly enlarged to join the basal portion which occupies almost 2/3 the total length. Only the dorsal and 2 submedian gland pores seen because of the strong radial musculature and dense glandular structures. Basal shield of esophagus strongly developed. Cardia conoid to elongate conoid. Intestine thin-walled, its cells filled with minute brown granules, completely surrounding cardia. Vulva transverse, ovaries reflexed about half their length. Pre rectum length twice body diameter. Rectum somewhat longer than anal body diameter. Female tail dorsally convex conoid to an elongated finely rounded terminus. Sometimes more gradually tapering and longer than illustrated. Male with the usual adanal pair and 5 to 7 ventromedian supplements which are very small and scarcely break the body contour. Four pairs of ventrosubmedian papillae present. Tail with 4 pairs of lateral pores. Spermatozoa about 3 x 11 /-'-,elongate drop shaped and minutely spinose, arranged in uterus with blunt end toward ovary. Holotype female, allotype, and other specimens as indexed under Westindicus 1. From moss and soil, Coca Falls and dieback area, El Yunque Tropical Mountains, Luquillo Experimental Forest.

FAMILY CARCHAROLAIMIDAE, NEW FAMILY Diagnosis: Actinolaimoidea. Cephalic region with framework which usually is a massively sclerotized basketlike structure which, in some instances, may be reduced to platelike elements. Anterior portion of esophagus a slender hyaline tube, posterior portion broad and muscular with strong radial musculature and dense glandular tissues which often obscure details of the esophageal gland nuclei. Basal shield often present. Nygolaimoid glands at base of esophagus known in 2 species. Ovaries 2, reflexed. Vulva transverse. Vagina unusually 34 Gerald Thome

D I;:: \;~ ,.,b l:! !1 " ff

FIG. 8.-Westindicus brachycephalus: A, Head and anterior end of esophagus x 1,000; B, ribbed framework of pharyngeal vestibule; C, postel'ior portion of female; D, vulva-vagina region; E, posterior portion of male; F, base of esophagus, basal shield, cardia, and anterior intestine; G, spermatozoa x 1,000.

short and small. Lateral field usually conspicuous with large closely spaced cells from which a single row of prominent lateral pores rise and sometimes additional lines of very minute, fiberlike pores. Tails usually blunt, rounded, rarely conoid arcuate. Males known only for C. teres. Females apparently syngonic hermaphrodites. Type subfamily: Carcharolaiminae, new subfamily. Subfamily Carcharolaiminae, new subfamily Diagnosis: Carcharolaimidae. Vestibule with or without denticles. Small onchia grouped about base of vestibule and pharynx. Spear- r length less the width of lip-region. Cardia elongate conoid or cylindroid. f Type genus: Carcha1'OlaimusThome, 1939. d

- Nematodes of Puerto Rico 35

GENUS Carchal'Olaimus THORNE, 1939 Diagnosis: Carcharolaiminae. Characters of the subfamily. Cephalic framework basketlike with ribs, dentides or small basal anchia. Type species: Carcharolaimus teres Thorne, 1939.

Key to species of Carcharolaimus 1. Pharynx without dentides 2 Pharynx with numerous dentides 5 2. Tail conoid, arcuate ..... aberrans n.sp. Tail blunt, rounded . . . . 3 3. Neck short, b - 5 teres Thorne, 1939 1.4-2.0 mm.; a - 37; b - 5.0; c - 50; V - s 48 s 1.8 mm.; a - 40; b - 5.0; c - 60; T - 50 Neck longer, b - 3.7 4 4. Spear of normal form 1'0tundicauda (deMan, 1880) 1.7 mm.; a - 40; b - 3.7; c - 67; V - 9 49 10 Spear somewhat sickle-shaped dl'epanodon Loof, 19647 5. Lip-region discoid or angular 6 Lip-region only angular, not discoid 7 6. Vulva near 45 percent discus n.sp. Vulva near 67 percent formosus Lordello, 1957 2.3 mm.; a - 47; b - 4; c - 81; V - 10 57 6 7. Nygolaimoid glands at esophagus base dentatus Thorne, 1939 2.2 mm.; a - 40; b - 5; c - 83; V - ]3 48 15 Basal shield on esophagus present ramirezi n.sp. Caracharolaimus pizae Lordello, 1953 is not included since the description is not available. Carcharolaimus ramirezi, n.sp. (Fig. 9, A-E) 2.2 mm.; a - 38; b - 4.0; c - 67; V - 10 49 10 Body cylindroid, tapering to head which is 2/3 width of neck- base. Lateral fields with about 125 cells from which conspicuous pores rise in an irregular line, usually from the dorsal side of the field. Numerous ventral pores rise from small glands, especially in esophageal region. Amphids 1/3 head-width, their apertures obscure slits beneath

7The position of C. drepanodon is questionable. Perhaps it should be trans- ferred to Caribenema. A more detailed study of the pharyngeal armature is desirable. Nematodes of Puerto F

contour of lips. Sensillae close to chamber. 1 striction, heavily sclerotized. Numerous jag A D pharynx and many minute denticles on wall~ 12 refractive ribs forming a basketlike cham pillars extend back to guiding-ring. Spear SOl side, 20 jJ, long with aperture occupying abou1 22 I-' long. Esophagus a slender nonmuscul massive basal portion which occupies 2/3 ( basal portion finely ribbed. Dorsal gland CI very obscure. Basal shield cup-shaped, en esophagus. Cardia elongate conoid, the ant surrounded by intestinal cells. Intestinal CE sized brown granules. Vulva transverse, ' across body. Eggs fill body cavity and 4 Ovaries reflexed, unusually short, only abol Prerectum scarcely differentiated from inte width. Males unknown and gravid females I This species is readily differentiated fn relative by the larger spear with long apert armature and longer, conoid tail. Holotype and other specimens as indexe Carcharolaimus mmil'ezi is named in honor 0 Pagan who collected these specimens from Lajas Experiment Substation, Lajas, P.R., ar much toward establishing the Puerto Rico J

H Carcharolaimus discus, (Fig. 9, F-I)

2.7 mm.; a - 43; b - 4; c - 86; V - 7 45 J Body closely resembling that of a Dis seen under the lower powers. Transverse st fields very slender near head, becoming al Lateral pores conspicuous, arranged in a fail discoid, half as wide as base of neck, reseml with submedian papillae on separate secto similar to that illustrated for D. major (~ great numbers of very fine refractive ribs. F onchia in base of pharynx, extending back tc about 16 I-' long, aperture occupying abou1 at first a slender nonmuscular tube, sudde FIG. 9.-A-E, Cm'charolaimus mmirezi: A, Head x 1,000; B, amphid two-thirds, irregular cylindroid with expande region; C, expansion region of esophagus; D, posterior I?ortion of and posterior ends. Only dorsal esophageal female; E, base of esophagus, basal shield, cardia, and anterior shield slightly cup-shaped. Cardia conoid, 1 intestine. F -I, Carchal'Olaimus discus: F, Head, lateral view x 1,000; G, head, sagittal section; H, ribbed framework of pharyn- surrounded by i~testinal cells. Intestinal cell~ geal vestibule. I, posterior portion of female. J,K, Carcharolaimus aberrans: Head x 1,000 and female tail. Nematodes of Puerto Rico 37 contour of lips. Sensillae close to chamber. Lip-region set off by constriction, heavily sclerotized. Numerous jagged teeth about base of pharynx and many minute denticles on walls. Pharynx hexagonal with 12 refractive ribs forming a basketlike chamber. Six strong sclerotized pillars extend back to guiding-ring. Spear somewhat concave on dorsal side, 20 µ.long with aperture occupying about half its length, extensions 22 µ. long. Esophagus a slender nonmuscular tube until it joins the massive basal portion which occupies 2/3 of total length. Lumen of basal portion finely ribbed. Dorsal gland conspicuous but remainder very obscure. Basal shield cup-shaped, enveloping tapering end of esophagus. Cardia elongate conoid, the anterior, disklike portion not surrounded by intestinal cells. Intestinal cells packed with medium- sized brown granules. Vulva transverse, vagina extending only 1/4 across body. Eggs fill body cavity and 4 times as long as wide. Ovaries reflexed, unusually short, only about as long as body-width. Prerectum scarcely differentiated from intestine, 1/2 as long as body- width. Males unknown and gravid females do not contain sperm. This species is readily differentiated from C. dentatus, its closest relative by the larger spear with long aperture, details of pharyngeal armature and longer, conoid tail. Holotype and other specimens as indexed under Carcharolaimus 4. Carcharolaimus ramirezi is named in honor of Mrs. Carmen T. Ramirez Pagan who collected these specimens from a sugarcane field near the Lajas Experiment Substation, Lajas, P.R., and who has contributed so much toward establishing the Puerto Rico nematology collection.

Carcharolaimus discus, n.sp. (Fig. 9, F-I)

2.7 mm.; a - 43; b - 4; c - 86; V - 7 45 10 Body closely resembling that of a Discolaimus, especially when seen under the lower powers. Transverse striae easily visible. Lateral fields very slender near head, becoming about ljg of midbody-width. Lateral pores conspicuous, arranged in a fairly regular line. Lip-region discoid, half as wide as base of neck, resembling that of a Discolaimus with submedian papillae on separate sectors of the labial assembly, similar to that illustrated for D. maior (20). Pharyngeal wall with great numbers of very fine refractive ribs. Four blunt outward hooked onchia in base of pharynx, extending back to spear guiding ring. Spear about 16 µ. long, aperture occupying about % its length. Esophagus at first a slender nonmuscular tube, suddenly expanded in posterior two-thirds, irregular cylindroid with expanded portions at both anterior and posterior ends. Only dorsal esophageal gland nucleus seen. Basal- shield slightly cup-shaped. Cardia conoid, l/3 body-width, completely surrounded by intestinal cells. Intestinal cells packed with small brown 38 Gerald Thome granules. Vulva transverse, vagina 1/3 body-width. Three pairs caudal pores apparently not associated with lateral field series. Carcharolaimus discus is immediately distinguished by its discoid tip region and four blunt pharyngeal onchia. Holotype and other specimens as indexed under Carcharolaimus 2. From soil about Hibiscus, Gurabo Experiment Substation, and declining sugarcane, Isabela, P.R.

Carcharolaimus aberrans, n.sp. (Fig. 9, J-K) 1.7 mm.; a - 35; b - 4; c - 35; V _ 12 54 12 Lip-region almost 3/5 as wide as neck base set off by deep constriction. Lateral fields with usual series of conspicuous pores arranged in a somewhat irregular line. Cephalic papillae 6 and 10 in usual arrangement. Amphid about 1/3 as wide as head, sensillae easily observed, slightly back of a point opposite spear-base. Pharynx hexagonal with 12 refractive supporting ribs and 4 sharp, small sickle-shaped onchia about its base. These onchia extend back to guiding-ring. Spear 20 µ. long, its aperture occupying about 3/5 its length, extensions 38 µ.. Anterior esophagus with fusiform bulb just in front of nerve-ring in which spears are formed during molting. Orifices of 2 or 3 tiny glands are visible near middle of this bulb. Basal portion of esophagus suddenly enlarged until 2/3 neck-width, its muscular and glandular tissues obscuring location of all glands except dorsal one. Cardia elongate spheroid, 1/3 neck-width, almost entirely submerged in intestinal cells. Three conspicuous nygolaimoid glands at base of esophagus instead of usually basal shield, similar to those described for Carcharolaimus dentatus. Intestinal cells with scattered fine brown granules. Vulva transverse, vagina extending 2/5 across body. Ovaries of young female about as long as body-width, observed producing syngonic spermatozoa. Prerectum and rectum length each about equal to body-width. Carcharolaimus aberrans is immediately distinguished from other species of the genus by its enlongated, arcuate tail. Holotype and other specimens as indexed under Carcharolaimus 3. From soil about declining sugarcane, Isabela, P.R. The presence of nygolaimoid glands in this species is most interesting because among the actinolaims thev have previously been reported for only Carcharolaimus dentatus (20). Carcharolaimus teres Thorne, 1939 A young specimen of what arpears to be this species was found in soil about declining Hibiscus, Gurabo Experiment Substation, and a similar specimen from a sugarcane field near Santa Isabel, P.R. N~matodes of Puerto Rico 39

GENUS Carcharoides, N.C. Carcharolaimidae: Characters of the family with cephalic region massivelysclerotized and armed with 6 strong forward pointing onchia grouped just below vestibule. Esophagus with strong basal shield. Cardia flattened pear shape. Tail conoid, ventrally arcuate to subacute terminus. Type species: Carcharoides tigrodon, n.sp. Carcharoides combines the pharyngeal armature of CaJ'charolaimus with the hemispheroid cardia of Caribenema and its position is somewhat uncertain. Carcharoides tigrodon, n.sp. (Fig. 10, A-E) 1.8 mm.; a - 21; b - 4.4; c - 35; V _ 14 EV l4 Body cylindroid throughout most of its length. Cuticle with minute transverse striae. Lateral fields about 1f4 body-width at midbody with about 80 cells from each of which a single pore rises from the dorsal side of the cell, forming a single line. Lip-region set off by deep constriction. Anterior circlet of papillae located on prominent conoid ele- vations at the extreme margin of the head. Posterior circlet emerging through openings in the heavily sclerotized hexaradiate framework. Sixstrong sharp teeth arranged about entrance to cephalic framework. Spear 15 µ long, its aperture occupying almost 2/3 its length. Spear extensions about 30 µ long. Nerve-ring obscure. Anterior portion of esophagus a slender hyaline tube, posterior portion suddenly expanded and occupying 3/5 the total length. Strong basal-shield present. Cardia flattened, hemispheroid, about 1hsubmerged into intestinal cells and scarcely extending into lumen. Intestinal cells densely packed with dark brown granules. Vulva transverse, vagina extending 1/5 across body. Ovaries reflexed almost to vulva when oocytes develop. Pre- rectum length about 1 body-width, rectum length equal to anal body diameter. Lateral cells and pores especially prominent on tail and posterior portion of body. Males not known and mature egg-producing females do not contain spermatozoa. Holotype and 1 paratype specimen as indexed under Carcharoides 1. From dying areas in lawn, Commonwealth Capitol grounds, San Juan, P.R. Subfamily Caribenematinae, new subfamily Diagnosis: Carcharolaimidae. Posterior portion of pharynx broad and heavily sclerotized, bearing 11arched, inward-pointing onchia. Spear-guide massive, attached to 4 spheroid sclerotized structures to which the onchia are also joined. A minor ring located posterior to principal guide. Bands of conspicuous extrudor muscles extend from 40 Gerald Thome

((I j:

FIG. 10.-Carcharoides ti.grodon: A, Head x 1,000; B, cephalic framework; C, posterior portion of female; D, expansion region of esophagus; E, esophagus base, basal shield, cardia, and anterior end of intestine. base of pharynx to spear extensions. Lumen of basal portion of esophagus minutely corrugated. Basal shield discoid. Cardia hemispheroid, surrounded by intestinal cells. Type genus: Caribenema, n.g.

GENUS Cal'ibenema, N.C. Diagnosis: Caribenematinae: Body cylindroid, lip-region about '/2 width of neck base, set off by deep constriction. A conspicuous series of pores rise from lateral fields and minute fibers which may be Nematodes of Puerto Rico 41

rudimentary pores rise from the ventral side of lateral fields with occasionally one on dorsal side. A dorsal series of pores extend back as far as base of esophagus and a ventral series throughout the body. Head with complicated armature as illustrated and described above in diagnosis of Carcharolaimidae (fig. 11). Spear extremely pointed, slightly dorsally arcuate with aperture occupying more than half its length. Vagina extending halfway across body. Ovaries 2, observed producing syngonic sperms in very young females. Males unknown. Type species: Caribenema fe1'Ox, n.sp. Carib enema is distinctive because of unusual pharyngeal armature, spear with its extremely long aperture and vagina extending halfway across body.

Cal'ibenema feTox, n.sp. (Fig. 11, A-G)

2.4 mm.; a - 48; b - 4.2; c - 80; V - 7 47 7 (juvenile females) Characters of the genus: Cuticle marked by minute transverse striae. Lateral fields at first about 1/10 body-width, increasing to % at midbody and 1/3 near tail, bearing about 120-125 very conspicuous pores usually somewhat dorsal. An additional series of very fine pores rise from the dorsal and another from the ventral side of the field. Lip-region set off by deep constriction. Amphids below contour of lateral lips with obscure sensillae. Pharynx with 12 strong ribs bearing minute jagged denticles and great numbers of exceedingly fine denticles on plates between ribs. Four strong onchia project forward forming a domelike spear guide. Guiding ring heavily sclerotized with four refractive spheroid bodies attached. Spear 24 I-'- long, its aperture occupying 3/5 the length. Strong muscle bands extend from guiding ring to spear extensions. Esophagus enlarged in posterior 3/5, set off from slender anterior portion by a distinct constriction. Thick discoid basal shield present. Only dorsal esophageal gland usually visible. Cardia somewhat spheroid with intestinal cells completely surrounding it. Intestinal cells filled with scattered fine granules. Vulva transverse, vagina extending halfway across body. Prerectum less than 1 body-width long. Ovaries 2, producing syngonic hermaphroditic sperms in young females. Rectum equal to anal body- l, width. Tail dorsally conoid to blunt terminus. Two pairs of lateral caudal papillae in addition to 2 pores from lateral field. Males unknown. Caribenema ferox is immediately distinguished from C. longidens by shorter spear (24: 31 1-'-) and longer spear extensions (30: 16 1-'-). Holotype female and other specimens as indexed under Caribe- nema 1, from soil about roots of palm, Agricultural Experiment Station, Rio Piedras. 42 Gerald Thome

F I

FIG. ll.-A-G, CaTibenema fel'ox: A,B, Head lateral and dorsal views x 1,000; C, amphid; D, expansion region of esophagus; E, ovary showing syngonic production of spermatozoa; F, esophagus base, basal shield, cardia, and anterior end of intestine; G, posterior portion of female. H,I, Caribenema longidens: H, Head x 1,000; I, posterior portion of female. Nematodes of Puerto Rico 43

Caribenema longidens, n.sp. (Fig. 11, H-I)

2.5 mm.; a - 41; b - 4; c - 83; V - iJ 54 G Body cylindroid, tapering to head which is 2/5 width of neck- base. Entire body tissues very dense, obscuring many details. Cuticle marked by fine transverse striae. Lateral fields with about 125 pores, arranged in a rather uniform line. Additional obscure pores arise from both dorsal and ventral sides of the field. Numerous dorsal pores are present, as far back as esophagus base, ventral pores throughout most of body-length. Lip-region angular with prominent papillae which appear to be infested with some organism causing them to be blunt and corroded. Amphids 1/3 head, similar to those of C. ferox. Pharynx with strong refractive ribs bearing minute teeth and with tiny denticles over portion of the areas between ribs. Four sharp teeth arch over a dome-shaped spear guide. Viewed laterally this structure appears to be dome-shaped but from a dorsoventral view the massive onchia are visible (fig. 11, B). Remainder of head armature as illustrated. Spear dorsally arcuate, 28 µ. long, with aperture occupying 3/5 its length and extensions only 15 µ. long. Esophagus enlarged in posterior 3/5, set off from slender hyaline anterior portion by contraction. Basal shield discoid or slightly cupped. Cardia somewhat spheroid, submerged in dense glandular cells of intestine. Vulva transverse. Vagina extending almost halfway across body. An egg was 42 x 132 µ., filling body cavity. Prerectum about as long as body-width. Tail dorsally convex conoid to blunt, rounded terminus. Holotype and other specimens as indexed under Caribenema 2. From soil about roots of sugarcane, San German, P.R. Caribenema longidens is immediately distinguished from C. ferox by its longer spear, short extensions, and more bluntly rounded tail.

FAMILY TRACHYPLEUROSIDAE NEW FA:MILY Diagnosis: ACTINOLAIMOIDEA. Lip-region set off by expansion. Pharynx armed with several rows of transverse denticles. Esoph- agus enlarged near middle by gradual expansion. Ovaries 2. Tails of both sexes elongate conoid to filiform. Supplements in ventral series. Type genus and species: TmchypleuTosum confoTmis (Schneider, 1905) Andrassy, 1959.

FAMILY MYLODlSCIDAE NEW FAMILY Diagnosis: ACTINOLAIMOIDEA. Lip-region disklike, bearing a shallow, minutely dentate bowl-like plate. Lining of pharynx muscular, not sclerotized as in other actinolaims. Spear-guiding a ring a double sclerotized framework. Anterior esophagus slender with conspicuous lumen, suddenly expanding near middle to form strongly muscular 44 Gerald Thome basal portion. Ovaries 2. Lateral fields narrow with pores in single line. Tail of female blunt, rounded. Males unknown. Type genus and species: Mylodiscus nanus Thorne, 1939. Nannomermis infestans, n.sp. (Fig. 7, F-G) Larva: 0.24 mm.; a - 43; genital primordium at 59 percent. In process of moulting. Body contents withdrawn from extremities. Granules have a tendency to be grouped in bands as if beginning to form cells. A single prominent nucleus, genital primordium?, present. Probably the female of Paractinolaimus robustus had devoured the eggs and they had hatched in the intestinal tract. From there the larvae had penetrated the intestinal wall and moved to the body cavity where they were undergoing their first moult. The larval spear was not visible but one must have been present to enable them to puncture the intestinal wall. The fact that ovaries had not developed in the host indicates that the two larvae observed had a sterilizing effect as mermithids usually do when parasitizing insects. Dr. N. A. Cobb (11) reported numerous specimens of both male and female Mononchus subtenuis-Mylonchulus subtenuis (Cobb, 1917) Altheer, 1958, parasitized by very small, 0.14 mm.; mermithid larvae which he named Nannomel'mis nemicola.s These larvae pos- sessed well-developed spears and other anatomical characters suf- ficient to enable accurate identification as mermithids. The specimens here recorded constitute the second record of nemic parasitism in other nemas. It is assumed that this mermithid may belong in the same genus and, since it is so much larger, even at this early stage, it is considered to be a different and undescribed species. ADDENDA TO BELONDIROIDEA GENUS Axonchoides, N.C. Diagnosis: Axonchinae. Body massive, lip-region narrow, blunt rounded tails in both sexes. Lip-region rounded with conspicuous semisclerotized framework and mammiform papillae. Spear extensions in 2 sections. Anterior % of esophagus a slender, nonmuscular tube, not set off from enlarged basal portion by constriction. Basal % of esophagus with massive muscular and glandular tissues which obscure all gland nuclei except the dorsal one. Supplements few, widely spaced, not rising above body contour.

8The writer is indebted to Dr. J. R. Christie and Miss Edna M. Buhrer for their aid in locating the obscure referenC'e containing the description of Nanno- mermis nemicola. Nematodes of Puerto Rico 45

Axonchoides is immediately distinguished from Axonchium, its nearest relative, by the semisclerotized labial framework, mammiform cephalic papillae, nonmuscular anterior portion of esophagus, and supplements not rising above body contour. Type species: Axonchoides crassus, n.sp.

Axonchoides crassus, n.sp. (Fig. 12, A-G)

2.8 mm.; a - 31; b - 2.7; c - 94; V - 8 48 17 2.8 mm.; a - 33; b - 3.2; c - 95; T - 63

Body cylindroid except anteriorly where it tapers to a lip-region only about 1/6 as wide as neck-base. Cuticle 8 po thick near head, 4 po on body, marked by excessively minute transverse and radial striae which are best observed on the tails. Lateral fields only about 2 to 4 po wide with tiny glands from which rise a single row of easily observed refractive pores. Lip-region rounded with papillae on mammiform ele- vations, very different from the usual conoid type. Amphid apertures almost encompassing head, goblet-shaped, their sensillae located about opposite base of spear extensions. Two lateral papillae rise from the slender lateral cord above the amphidial tube. Head with 6 semisclerotized ribs forming the cephalic framework. Spear somewhat fusiform, 12 po long, its aperture occupying about 1/3 its length. Guid- ing-ring single. Posterior portion of spear extensions unusual in that they are divided into 2 sections, indicating how the tripartite spears of Dorylaimellus and Xiphinema are formed. Anterior to nerve-ring the nonmuscular esophageal tube becomes an elongate-fusiform swel- ling in which spear-producing glands are located and which function during moulting. Lumen of this portion exhibits broken walls which indicate that pores of 2 or 3 glands enter here. Basal bulb of esophagus beginning rather abruptly with lumen at first somewhat sinuous where the dorsal gland orifice is located. Massive basal portion occupying % the esophagus length, with strong radial musculature and dense glandular tissues which obscure details of lumen and gland nuclei. Base of esophagus tapering to join cardia which at first is a discoid section, then becoming cylindroid and extending deep into intestine. Cardia, except its terminus, surrounded by a sheath of intestinal tissues densely packed with minute, dark brown granules. Intestinal cells also densely packed with similar granules. Prerectum length twice body-width. Rectum about half as long as prerectum. Vulva a transverse slit located less than 4 body-widths back of esophagus. Vagina extending in and slightly back about halfway across body. Anterior rudimentary uterine branch extending almost to cardia, forming a roomy pouch in which are hundreds of spermatozoa 1.5 x 4 po in size. Posterior ovary reflexed with about 6 rows of oocytes rising 46 Gerald Thome

FIG. 12.-Axonchoides crassus: A, Head x 1,000; B, posterior portion of male; C, amphid; D, spear development section of esophagus; E, base of esophagus, cardia, anterior rudimentary uterus, and vulva-vagina region; F, female tail; G, expansion region of esophagus. from the cap cell, with finally about 6 oocytes arranged in tandem as they proceed toward uterus. Eggs 32 x 90 ft. Male similar to female in general body form. Posterior portion slightly arcuate. Testes dorylaimoid. Spicula massive with elongated narrow shaft. Lateral guides slender, slightly curved. Supplements an adanal pair and 5 ventromedian which do not rise above body contour, with long ducts leading in and forward. Submedian papillae not present. Prerectum extending about as far as supplement series. Tails of both sexes bluntly rounded, bearing two pairs of papillae. Holotype female and allotype as indexed under Axonchoides l. From moss and soil by stream, Coca Falls, El Yunque Mountains, Luquillo Experimental Forest, March 28, 1965. Nematodes of Puerto Rico 47

SUMMARY This paper is a continuation of the nematological investigations in taxonomy conducted in Puerto Rico. A brief historical account on the nematodes of the superfamily ACTINOLAIMOIDEA is given. This is followed by a list of the species of Actinolaimus known through 1964 giving their present position in taxonomical nomenclature. The new superfamily ACTINOLAIMOIDEA is erected and with it three new families and subfamilies. The paper also includes the description of nine new genera and eighteen new species. An addenda to the BELONDIROIDEA is included. RESUMEN Esta publicaci6n es una continuaci6n de las investigaciones que se han llevado a cabo en Puerto Rico sobre la taxonomia de los nematodos. Se incluye en el estudio 4n breve historial de los nematodos de la superfamilia ACTINOLAIMOIDEA, seguido de una lista de las especies del genero Actinolaimus conocidas hasta el 1964, con su posiciontaxonomica actual. Se establecen la nueva superfamilia ACTI- NOLAIMOIDEA y tres familias y subfamilias nuevas. En el estudio tambien se describen nueve generos y diez y ocho especies nuevas. Se incluye ademas una nota adicional en relacion a la superfamilia BELONDIROIDEA.

LITERATURE CITED 1. AUgen, C., Uber elmge freilebende Siisswasser-Nematoden von Mt. Kenya (Brit. Ostafrika); Ein kleiner Beitrag zur Kenntnis tropischer Nematoden, Ark. Zool., Stockholm, Ser. 2, 3 (1-2) 139-57, 1952. 2. , Terrestrial nematodes from Jan. Mayen. Ann. & Mag. Nat. Hist. 12. s. (69), 6: 665-688, 1953. 3. Altheer, E., Results from the Danish Expedition to the French Cameroons 1949-1950) XXVIII. Nematodes Limncoles; 8 figs. Bttll. deL'Institut Francais D' Afriqtte Noire 22: (3) 770-787, 1960. 4. Andrassy, L., Neubenenungen einiger Homonymen Nematoden-Gattungen. Nematologica 4: 223-6, 1959. 5. , Taxonomische iibersicht der Dorylaimen (Nematoda) II, Acta Zool. Acad. Sci. Httng. 6: (1-2) 1-28, 1960. 6. , Freilebende Nematoden aus Angola, I, Einige moosbewohnende Nematoden, Das Pttblicacoes Cultttris De Companhia de Diamentes de Angola 66: 57-78, 1963. 7. , Einige nematodes aus der umbegung des toten meeres, Israel, I. Zool. 13: 89-97, 1964. 8. Baker, A. D., Check lists of the nematode superfamilies Dorylaimoidea, Rhabditoidea, Tylenchoidea, and Aphelenchoidea, E. J. Brill, Leiden, 1962. 9. Chitwood, B. G. and Chitwood, M. B., An Introduction to Nematology. Section 1, Anatomy; Washington, D.C., B. G. Chitwood, revised ed., 1950. 10. Clark, W. C .. New species of dorylaimoid nematodes belonging to the genus Pungenttts Thorne & Swanger, Actinolaimus Cobb and DOTylaimellHs Cobb, New Zealand Jour. Sci. 6: (4) 565-76, 1962. TE( 48 Gerald Thorne

1l. Cobb, N. A., A new mermithid infesting other nemas, Nannomermis, nemicola, n.g. n.sp., Proc. H elmith. Soc. Washington in ]. Pal'S. 11: 217-29, 1925. (First reported without name in ]. Pars. 11: 120. Sept. 1924). 12. Gadea, E., Nota sobre un nuevo Actinolaimus de Fernando Po. Inst., BioI. Api. 7: 77-82, 1950. 13. Goodey, T., Soil and Freshwater Nematodes (2nd ed. revised by J. B. Goodey). Methuen & Co. Ltd., London, pp. 1-544, 1963. 14. Loof, P. A. A., Free-living and plant parasitic nematodes from Venezuela, Nematologica 10: 201-300, 1964. 15. , A note on Paractinolaimlls macrolaimtts (De Man, 1880) Andrassy, 1964, Nematologica 10: 647, 1965. 16. Lordello, L. G. E., and Zamith, A. P. L., Brasilaimlls, a new genus of free- living nematode, Rev. Bras. BioI. 17: (4) 455-7, 1957. 17. Meyl, A. B., Nematoden aus dem Psammon des Tanganyika-Sees, Zool. Anz., 155: (5-6) 135-8, 1955. 18. , Free-living Nematodes; in: Exploration Bydrobiologique du Lac Tanganika (1946-47), [nst. Roy. Sci. Nat. Belg., 3: (7) 27-51, 1957. 19. , Die freilebenden Stisswassernematoden (Fadenwtirmer) in: Die Tierwelt mitteleuropas (Bohmer. Ehrmann & Ulmer), Quelle & Meyer. Leipzig, pp. 1-273, 54 pIs. figs. 1-889, 1961. 20. Thorne, G., A monograph of the nematodes of the superfamily Dorylaimoic1ea. Capita Zoo1., 8 (5) 1-261, pIs. 1-29. figs. 1-214, 1939. 21. , Nematodes of Puerto Rico: Belondiroidea new Superfamily, Lep- tonchidae, Thorne, 1935. and Belonenchidae new Family. (Nemata. Ac1eno- phorea, Dorylaimida). Univ. P.R., Tech. Paper 39, pp. 1-51, Sept. 1964.

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